Advances in Ant System A Tics (Hymenoptera Formicidae) - 2007

Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O.
Wilson – 50 years of
contributions
Roy. R. Snelling, Brian L. Fisher, and Philip S. Ward (Editors.)
M emoirs of the American Entomological Institute, Volume 80
American Entomological Institute.
Gainesville, FL, September 2007
Slightly more than fifty years ago a significant advance in the systematics of ants was published - E. O. Wilson's doctoral thesis:
A monographic revision of the ant genus Lasius (Wilson, 1955). This was not quite Ed's first paper on formicid systematics
(Wilson, 1951, 1952), but it was the first comprehensive revision of an ant genus based on modern or Mayrian taxonomic
principles. Ant systematics had truly entered the modern era only five years earlier with W. S. Creighton's revolutionary Ants of
North America (Creighton, 1950). Creighton emphasized the need for species and subspecies based on solid morphological
features and sound field experience. Creighton's work clearly influenced Borgmeier's massive revision of the New World army
ants (1955). Enlarging upon Creighton's philosophical understanding that taxa must be based on secure morphological
differentiation and that species must be understood as populations subject to (often) considerable geographic variation,
Wilson's revision, following on Creighton's monumental accomplishment, effectively ended the era of such giants as Forel,
Emery, and Wheeler. Each of these workers had described numerous subspecies and varieties, often from inadequate
samples; these new taxa were all too frequently ascribed to species that were, themselves, poorly known.
Creighton had dismissed the varietal form, the backbone of the quadrinomial system then applied to ants. Some of the
"varieties" he did continue to recognize as geographic variants or subspecies, usually based on morphological characteristics;
most, however, were consigned to oblivion as nothing more than trivial sporadic variations within a population. Only five years
later, Wilson took the next step and dispensed with subspecies, having determined that most ceased to exist as taxonomically
meaningful entities when entire populations were critically examined.
I first began studying ants, as part of my larger interest in the aculeate Hymenoptera, in the years just prior to the appearance of
Creighton's book. Suffice to say that when I first realized just how chaotic ant taxonomy was, I promptly determined not to work
on ants. Some years later I acquired a copy of Ants of North America, but by then I was deeply involved with bees and social
wasps, and had only a casual interest in ants. It was not until about 1960 that I became more involved with ants. Ants of North
America was supplemented by reprints of more recent works, including Ed's Lasius monograph. Suddenly, ant systematics, a
previously arcane welter of confusing subspecies and varieties and wholly unworkable keys, had become comprehensible!
Those individuals entering ant systematics over the past half century cannot begin to comprehend the profound significance of
these two publications. But now, fifty years later, we can look back and appreciate Ed's contributions, examined more fully in the
following paper, to our field of endeavor.
The present volume, containing contributions from ant taxonomists the world over, began as a chance email interchange
between Jack Longino and myself in the spring of 2003. Basically, it was something on the order of (from Jack): "Maybe we
should put together a Festschrift for the Old Boy (i.e., Ed)." My response was, "Not a bad idea." We left it at that and signed off.
The idea didn't die there. Phil Ward, who happened to be visiting the LACM at the time, and I started talking about it and shortly
agreed in principle to explore the possibility. I immediately began contacting colleagues and potential contributors; most were
enthusiastic and agreed to provide papers. A lot of hard work by Ed's colleagues from around the world has resulted in this
volume. There are a couple of thoughtful essays, one from Barry Bolton, another from Bob Taylor. Another paper examines the
great diversity of ant species in arid lands in Australia. But the papers submitted here are primarily taxonomic/systematic in
nature.
All of the papers included herein have been reviewed by the editors and, sometimes also by the usual battery of "anonymous
reviewers." I wish to thank two of my colleagues at the Natural History Museum of Los Angeles County, Brian V. Brown and Fred
S. Truxal, for their encouragement. Particular thanks are tendered to my son and grandson, Gordon and James, respectively,
and to Weiping Xie and Bill Mertz who came to my assistance many times when my ineptitude with computer technology became
all too evident. And finally, I and my co-editors, Phil Ward and Brian Fisher, wish to express our profound gratitude to all the
authors whose contributions made this volume possible.
Roy. R. Snelling
Snelling, R. R. 2007. Preface, pp 1-2. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant
systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the
American Entomological Institute, 80.(pdf) [plazi]
Ward, P. S. 2007. Edward O. Wilson and his contributions to ant systematics, pp 3-7. In Snelling, R. R., B. L. Fisher,
and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years
of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Alpert, G. D. 2007. A review of the ant genus Metapone Forel from Madagascar, pp 8-18. In Snelling, R. R., B. L.
Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50
years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Andersen, A. N. 2007. Ant diversity in arid Australia: a systematic overview, pp 19-51. In Snelling, R. R., B. L. Fisher,
Bolton, B. 2007. How to conduct large-scale taxonomic revisions in Formicidae, pp 52-71. In Snelling, R. R., B. L.
Brandão, C. R. F. and A. J. Mayhé-Nunes. 2007. A phylogenetic hypothesis for the Trachymyrmex species groups,
Brandão, C. R. F. and A. J. Mayhé-Nunes. 2007. A phylogenetic hypothesis for the Trachymyrmex species groups,
and the transition from fungus-growing to leaf-cutting in the Attini, pp 72-88. In Snelling, R. R., B. L. Fisher, and P. S.
Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Cover, S. P. and M. Deyrup. 2007. A new ant genus from the southwestern United States, pp 89-99. In Snelling, R.
R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O.
Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Deyrup, M. and S. P. Cover. 2007. A new species of Crematogaster from the pinelands of the southeastern United
States, pp 100-112. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera:
Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute,
80.(pdf) [plazi]
Espadaler, X. 2007. The ants of El Hierro (Canary Islands), pp 113-127. In Snelling, R. R., B. L. Fisher, and P. S.
Fernández, F. 2007. Two new South American species of Monomorium Mayr with taxonomic notes on the genus, pp
128-145. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera:
80.(pdf) [plazi]
Fisher, B. L. 2007. A new species of Probolomyrmex from Madagascar, pp 146-152. In Snelling, R. R., B. L. Fisher,
Francoeur, A. 2007. The Myrmica punctiventris and M. crassirugis species groups in the Nearctic region, pp
153-145. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera:
80.(pdf) [plazi]
Kohout, R. J. 2007. Revision of the subgenus Aulacomyrma Emery of the genus Polyrhachis F. Smith, with
descriptions of new species, pp 186-253. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant
Lattke, J. E., F. Fernández and E. E. Palacio G.. 2007. Identification of the species of Gnamptogenys Roger in the
Americas, pp 254-270. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics
(Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American
Entomological Institute, 80.(pdf) [plazi]
Longino, J. T. and F. Fernández. 2007. Taxonomic review of the genus Wasmannia, pp 271-289. In Snelling, R. R.,
B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson
– 50 years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
McArthur, A. J. 2007. A key to Camponotus Mayr of Australia, pp 290-351. In Snelling, R. R., B. L. Fisher, and P. S.
Ogata, K. and H. Okido. 2007. Revision of the ant genus Perissomyrmex with notes on the phylogeny of the tribe
Myrmecinini, pp 352-369. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics
Schoedl, 2007. Revision of Australian Meranoplus: the Meranoplus diversus group, pp 370-424. In Snelling, R. R., B.
L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson –
50 years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Schultz, T. R. 2007. The fungus-growing ant genus Apterostigma in Dominican amber, pp 425-436. In Snelling, R. R.,
B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson
– 50 years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Shattuck, S. O. and N. J. Barnett. 2007. Revision of the ant genus Mayriella, pp 437-458. In Snelling, R. R., B. L.
Snelling, G. C. and R. R. Snelling. 2007. New synonymy, new species, new keys to Neivamyrmex army ants of the
United States, pp 459-550. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics
Snelling, R. R. 2007. A review of the arboreal Afrotropical ant genus Axinidris, pp 551-579. In Snelling, R. R., B. L.
Taylor, R. W. 2007. Bloody funny wasps! Speculations on the evolution of eusociality in ants, pp 580-609. In Snelling,
R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O.
R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O.
Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80.(pdf) [plazi]
Trager, J. C., J. A. MacGown and M. D. Trager. 2007. Revision of the Nearctic endemic Formica pallidefulva
group, pp 610-636. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera:
80.(pdf) [plazi]
Ward, P. S. 2007. The ant genus Leptanilloides: discovery of the male and evaluation of phylogenetic relationships
based on DNA sequence data, pp 637-649. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant
Yamane, S. 2007. Pachycondyla nigrita and related species in Southeast Asia, pp 650-663. In Snelling, R. R., B. L.
Yoshimura, M. and K. Onoyama. 2007. A new sibling species of the genus Strumigenys, with a redefinition of S.
lewisi Cameron, pp 664-690. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds) Advances in ant systematics
Snelling, R. R., B. L. Fisher, and P. S. Ward. (eds). 2007. Advances in ant systematics (Hymenoptera: Formicidae): homage
to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80: 1-670.
PREFACE
Roy R. Snelling
Natural History Museum of Los Angeles County
900 Exposition Boulevard, Los Angeles, California, U.S.A.
antmanrs@nhm.org
Slightly more than fifty years ago a significant advance in the systematics of ants was published —
E. O. Wilson's doctoral thesis: A monographic revision of the ant genus Lasius (Wilson, 1955). This
was not quite Ed's first paper on formicid systematics (Wilson, 1951, 1952), but it was the first
comprehensive revision of an ant genus based on modern or Mayrian taxonomic principles. Ant
systematics had truly entered the modern era only five years earlier with W. S. Creighton's
revolutionary Ants of North America (Creighton, 1950). Creighton emphasized the need for species
and subspecies based on solid morphological features and sound field experience. Creighton's work
clearly influenced Borgmeier's massive revision of the New World army ants (1955). Enlarging
upon Creighton's philosophical understanding that taxa must be based on secure morphological
differentiation and that species must be understood as populations subject to (often) considerable
geographic variation, Wilson's revision, following on Creighton's monumental accomplishment,
effectively ended the era of such giants as Forel, Emery, and Wheeler. Each of these workers had
described numerous subspecies and varieties, often from inadequate samples; these new taxa were
all too frequently ascribed to species that were, themselves, poorly known.
Creighton had dismissed the varietal form, the backbone of the quadrinomial system then
applied to ants. Some of the "varieties" he did continue to recognize as geographic variants or
subspecies, usually based on morphological characteristics; most, however, were consigned to
oblivion as nothing more than trivial sporadic variations within a population. Only five years later,
Wilson took the next step and dispensed with subspecies, having determined that most ceased to
exist as taxonomically meaningful entities when entire populations were critically examined.
I first began studying ants, as part of my larger interest in the aculeate Hymenoptera, in the
years just prior to the appearance of Creighton's book. Suffice to say that when I first realized just
how chaotic ant taxonomy was, I promptly determined not to work on ants. Some years later I
acquired a copy of Ants of North America, but by then I was deeply involved with bees and social
wasps, and had only a casual interest in ants. It was not until about 1960 that I became more
involved with ants. Ants of North America was supplemented by reprints of more recent works,
including Ed's Lasius monograph. Suddenly, ant systematics, a previously arcane welter of
confusing subspecies and varieties and wholly unworkable keys, had become comprehensible!
Those individuals entering ant systematics over the past half century cannot begin to comprehend the
profound significance of these two publications. But now, fifty years later, we can look back and
appreciate Ed's contributions, examined more fully in the following paper, to our field of endeavor.
The present volume, containing contributions from ant taxonomists the world over, began as a
chance email interchange between Jack Longino and myself in the spring of 2003. Basically, it was
something on the order of (from Jack): "Maybe we should put together a Festschrift for the Old Boy
(i.e., Ed)." My response was, "Not a bad idea." We left it at that and signed off. The idea didn't die
there. Phil Ward, who happened to be visiting the LACM at the time, and I started talking about it
and shortly agreed in principle to explore the possibility. I immediately began contacting colleagues
and potential contributors; most were enthusiastic and agreed to provide papers. A lot of hard work
by Ed's colleagues from around the world has resulted in this volume. There are a couple of
2 Memoirs of the American Entomological Institute, Volume 80
thoughtful essays, one from Barry Bolton, another from Bob Taylor. Another paper examines the
great diversity of ant species in arid lands in Australia. But the papers submitted here are primarily
taxonomic/systematic in nature.
All of the papers included herein have been reviewed by the editors and, sometimes also by the
usual battery of "anonymous reviewers." I wish to thank two of my colleagues at the Natural
History Museum of Los Angeles County, Brian V. Brown and Fred S. Truxal, for their
encouragement. Particular thanks are tendered to my son and grandson, Gordon and James,
respectively, and to Weiping Xie and Bill Mertz who came to my assistance many times when my
ineptitude with computer technology became all too evident. And finally, I and my co-editors, Phil
Ward and Brian Fisher, wish to express our profound gratitude to all the authors whose contributions
made this volume possible.
LITERATURE CITED
Borgmeier, T. 1955. Die Wanderameisen der Neotropischen Region. Studia Entomologica 3: 1-720.
Creighton, W.S. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology
104: 1-585.
Wilson, E.O. 1951. A new Leptothorax from Alabama. Psyche 57: 128-130.
Wilson, E. O. 1952. O complexo Solenopsis saevissima na America do sul. Memórias do Instituto
Oswaldo Cruz 50: 49-68.
Wilson, E. O. 1955. A monographic revision of the ant genus Lasius. Bulletin of the Museum of
Comparative Zoology at Harvard College 113: 1-201.
Ward, P. S. 2007. Edward O. Wilson and his contributions to ant systematics, pp. 3-7. In
Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics (Hymenoptera:
Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of the American
Entomological Institute, 80.
EDWARD O. WILSON AND HIS CONTRIBUTIONS TO ANT SYSTEMATICS
Philip S. Ward
Department of Entomology,
University of California, Davis
Davis, CA 95616, USA
psward@ucdavis.edu
This Festschrift celebrates the fiftieth anniversary of the publication of the Ph.D. thesis of the
world’s most prominent myrmecologist. Edward O. Wilson is a towering figure in ant biology,
and his influence extends well beyond myrmecology to broader aspects of science and society.
Yet many of his research accomplishments, especially from the earlier years, reflect a particular
concern with ant systematics and evolution. Among Wilson’s many contributions to this field,
his doctoral dissertation—a monographic revision of the ant genus Lasius—is a particularly
significant milestone. Published in March 1955, it represents one of the first serious attempts to
incorporate into ant systematics the principles of population biology and evolution. Wilson and
his associate, William L. Brown, Jr., pioneered the introduction of these ideas into ant taxonomy,
often in the face of resistance from more traditional workers.
The “new systematics”, as it was called by Julian Huxley (1940), drew inspiration from the
writings of Ernst Mayr, Bernard Rensch, Theodosius Dobzhansky, Sewall Wright and other
architects of the neodarwinian synthesis. Mayr’s (1942) Systematics and the origin of species
had an especially influential role. Species were envisaged as groups of interbreeding
populations, reproductively isolated from other such groups, and frequently containing a
substantial amount of variation. This biological species concept stood in contrast to the
typological view of species that was implicit in much earlier taxonomic work. Ant taxonomy
itself was burdened with a peculiar and unwieldy pentanomial nomenclature (Brown, 1955), and
was associated with the profligate naming of “species”, “subspecies”, and “varieties”, often
poorly diagnosed and with little thought directed to the nature of these forms or their
relationships to one another.
An early assault on infraspecific names in ant taxonomy was carried out by William S.
Creighton, most notably in his monumental Ants of North America (1950). Buhs (2000) credits
Creighton with introducing the “new systematics” to myrmecology, but Wilson and Brown went
further than Creighton and they had a more lasting impact. They argued compellingly against
the use of subspecies names in systematics (Wilson & Brown, 1953; Brown & Wilson, 1954);
they advocated a global approach to revisionary taxonomy; they were concerned with the higher
classification and phylogeny of ants; and they paid greater attention than other workers to the
quantification of intra- and interspecific variation. Wilson’s (1955) Lasius monograph
exemplified these ideals. It dealt with the world fauna, offered a provisional phylogeny,
abandoned the use of subspecies names, and attempted to delineate species while allowing for a
measure of intraspecific variation. It made extensive use of metric measurements and indices,
based on large population samples, and it introduced standardized terms—still in use today—for
describing the appearance of pilosity in ants.
These progressive features were not adopted by all workers. Creighton and other North
American-centered ant taxonomists became quite critical of the work of the “Happy Harvard
Team”, as they referred to Wilson and Brown. In retrospect, most of the criticisms appear to be
rather defensive and to miss their mark. One of the most persistent was that the Harvard
researchers lacked sufficient field experience to make prudent judgments about species
boundaries (Buhs, 2000). This arose specifically in the context of Wilson’s Lasius revision.
The criticism is difficult to sustain since there is abundant evidence in the monograph that
Wilson had direct experience in the field with most of the North American species. His
taxonomic studies of Melanesian ants were similarly preceded by an intensive collecting
expedition to that region of the world. And no ant systematist can—or perhaps ever will—match
Bill Brown for global field experience of ants in their natural habitats.
If the taxonomic disputes are dissected more carefully one sees that they often center on the
issue of how much variation one can expect to occur in single biological species. Wilson and
Brown tended to be “lumpers”, accepting more intraspecific variation, both ecological and
phenotypic, than Creighton, Cole, Gregg, Buren, and others, who were prone to split biological
diversity more finely. What is the verdict 50 years later? For many of these differences of
taxonomic opinion, involving such difficult genera as Formica and Lasius, it is sobering to
realize that we simply do not have adequate evidence to judge. In some instances we are dealing
with closely related allopatric populations for which assessment of species status is often
somewhat arbitrary. For Lasius it is noteworthy, however, that half a century later and after the
accumulation of much more material, Wilson’s treatment of the Nearctic species holds up quite
well. In the Palearctic region—from which Wilson had less material and no direct field
experience—the picture has undergone greater modification, with multiple species of Lasius
now being recognized under what Wilson considered to be single polytypic species (Seifert,
1988, 1992; Schlick-Steiner et al., 2003). There are some genetic data to support recognition of
these cryptic species (Steiner et al., 2004; Janda et al., 2004), but taxon sampling remains
incomplete and information from molecular markers other than mitochondrial DNA—which can
be quite misleading about species boundaries—is badly needed. If the situation in North
America is comparable to that in the Old World, then it is possible that some of the Nearctic
Lasius “species” will prove to be composed of multiple sibling species. There is ample scope
here for additional research in both North America and Eurasia to determine if we are dealing
with different biological species, i.e., reproductively isolated entities, or with geographical,
ecological and behavioral variants.
Another notable feature of Wilson’s (1955) Lasius revision was his exploration of the
processes responsible for the genesis of taxonomic diversity. For the first time in ant taxonomy
we find discussion of the patterns of species distributions and intraspecific geographical
variation, and the implications of these patterns for the mode of speciation. Considerable
attention was also given to biological traits. Under the accounts of individual ant species there
are summaries of what is known about nesting behavior, foraging habits, and habitat preferences.
The role of interspecific competition and divergent selection was of particular interest to Wilson,
and led to a co-authored paper with Brown on character displacement that appeared one year
later (see below).
Over a ten-year period between 1955 and 1965 Wilson published a series of additional
papers on ant taxonomy that reflected the thinking of the “new systematics”. This included
revisionary treatments of selected groups of Melanesian ants, mostly poneromorphs (Wilson,
1957, 1958a, 1958b, 1959b, 1959c); rediscovery and description of the enigmatic Sri Lankan ant
Aneuretus simoni (Wilson et al., 1956); taxonomic notes on certain Formica species (Wilson &
Brown, 1955); a review of dacetine ant evolution (Brown & Wilson, 1959); and a revision of the
Indo-Australian army ants (Wilson, 1964). The taxonomic work on Melanesian ants provided a
foundation for Wilson’s analysis of ecological and evolutionary trends in the ants of that region,
especially a pattern of range expansion and habitat shifts that Wilson (1959a, 1961) termed the
taxon cycle. Another influential idea, which arose in part from taxonomic patterns that Wilson
observed in Lasius, was the concept of character displacement (Brown & Wilson, 1956), the
notion that under some circumstances closely related species can be expected to show greater
divergence in sympatry than in allopatry, as a consequence of interspecific competition, a pattern
opposite to that which is predicted by occasional interspecific hybridization. Thus, not only was
Ward: Wilson - contributions to ant systematics 5
Wilson’s taxonomic work informed by the principles of evolutionary biology, but there was also
reciprocity: taxonomic findings themselves illuminated previously undisclosed phenomena,
emerging, as Wilson himself has noted, “as unplanned products of pedestrian daily research”
(Wilson, 1994: 205).
After 1965 there was less taxonomic output from Wilson as his inquiries focused on other
aspects of ant biology, and on broader questions about biogeography, the evolution of social
behavior, and the conservation of biodiversity. Yet there continued to be periodic contributions
to ant systematics, including the discovery and description of the first fossil ant from the
Cretaceous period (Wilson et al., 1967); a monograph on the ants of Polynesia (Wilson &
Taylor, 1967); a magisterial overview of social insect biology (Wilson, 1971) that included a
lucid discussion of the origin of ants and the contradictions inherent in treating both Amblyopone
and Sphecomyrma as models for the common ancestor of all ants; a series of taxonomic papers
on fossil ants, primarily from Dominican amber (Wilson, 1985a, 1985b, 1985c, 1985d, 1986);
and a useful taxonomic summary and set of keys to ant subfamilies and genera (appearing in
Hölldobler & Wilson, 1990). Although their primary focus is not systematics, the much
acclaimed books The insect societies (Wilson, 1971) and The ants (Hölldobler & Wilson, 1990)
have had an important indirect influence on ant systematics by luring acolytes into the field with
engaging and informative prose.
In 2003 Wilson published a prodigious tome on the New World Pheidole. Recognizing 624
species, of which more than half are new to science, this work provides the first comprehensive
treatment of Pheidole on such a large geographic scale. The monograph is copiously illustrated,
and identification of an unknown specimen is most easily accomplished by browsing the detailed
line drawings within the appropriate species group, paying attention to highlighted features and
lists of similar taxa. This “field guide” approach produces a more reliable result than resorting
to the species identification keys, which are cumbersome and often lead the user astray.
It may be difficult for non-systematists to appreciate what an enormous improvement
Wilson’s (2003) revision brings to the state of Pheidole taxonomy. There had been no earlier
synthetic work on the genus at such a scale—merely an accumulated morass of hundreds of
names, often associated with ill-defined or unrecognizable taxa, and a backlog of thousands of
Pheidole specimens in collections that could not be confidently identified to species. We now
have a robust framework for the classification and identification of the New World fauna,
available for further refinement and testing. Of course, this is certainly not the final word on the
taxonomy of New World Pheidole. Numerous additional species await discovery and
delineation. Longino’s fine-scaled analysis of the Costa Rican fauna (Longino, 2004) yields
species delimitations somewhat at variance with those of Wilson. Rather ironically—in view of
earlier taxonomic controversies—Longino interprets a number of Wilson’s “species” as
representing intraspecific variation, thus advocating a more polytypic view of species than that
adopted by Wilson.
While most of Wilson’s work in ant systematics has been concerned with species-level
issues, he has also contributed incisively to our understanding of the origin of ants (as noted
above), especially through the discovery and analysis of key fossil taxa (Wilson, 1971, 1987;
Wilson et al., 1967). His proposed phylogeny of Lasius (Wilson, 1955: 15) anticipated correctly
the inclusion of Acanthomyops within that genus, as now confirmed by molecular data (Janda et
al., 2004). A provisional phylogeny for Formicidae as a whole, based on internal and external
anatomy, that appeared in The ants (Hölldobler & Wilson, 1990: 26) provides an interesting
point of contrast with the picture that is emerging from molecular phylogenetic analyses. From
Lasius to Pheidole, from fossil ants to extant species, what underpins all of Wilson’s systematic
papers is an intense interest in understanding biological diversity—its origins, causes, and
consequences. His writings also convey a passion for scientific discovery and an ability to apply
critical evolutionary thinking to taxonomic problems. The result is an inspiring body of work
and an intellectual legacy that will continue to stimulate future generations of ant systematists.
LITERATURE CITED
Brown, W.L., Jr. 1955. Ant taxonomy. Pp. 569-572 in: Kessel, E. W. (ed.) A century of
progress in the natural sciences, 1853-1953. San Francisco: California Academy of Science,
x + 807 pp.
Brown, W.L., Jr. & Wilson, E.O. 1954. The case against the trinomen. Systematic Zoology 3:
174-176.
Brown, W.L., Jr. & Wilson, E.O. 1956. Character displacement. Systematic Zoology 5: 49-64.
Brown, W.L., Jr. & Wilson, E.O. 1959. The evolution of dacetine ants. Quarterly Review of
Biology 34: 278-294.
Buhs, J.B. 2000. Building on bedrock: William Steel Creighton and the reformation of ant
systematics, 1925-1970. Journal of the History of Biology 33: 27-70.
Creighton, W.S. 1950. The ants of North America. Bulletin of the Museum of Comparative
Zoology 104: 1-585.
Hölldobler, B. & Wilson, E.O. 1990. The ants. Cambridge, Massachusetts: Harvard University
Press, xii + 732 pp.
Huxley, J. (ed.) 1940. The new systematics. Oxford: Clarendon Press, viii + 583 pp.
Janda, M., Folková, D. & Zrzavý, J. 2004. Phylogeny of Lasius ants based on mitochondrial
DNA and morphology, and the evolution of social parasitism in the Lasiini (Hymenoptera:
Formicidae). Molecular Phylogenetics and Evolution 33: 595-614.
Longino, J. T. 2004. Ants of Costa Rica: Pheidole. http://www.evergreen.edu/ants/genera/
pheidole/specieslist.html (version dated 6 July 2004)
Mayr, E. 1942. Systematics and the origin of species from the viewpoint of a zoologist. New
York: Columbia University Press, xiv + 334 pp.
Schlick-Steiner, B.C., Steiner, F.M., Schödl, S. & Seifert, B. 2003. Lasius austriacus sp. n., a
central European ant related to the invasive species Lasius neglectus. Sociobiology 41: 725-
736.
Seifert, B. 1988. A revision of the European species of the ant subgenus Chthonolasius (Insecta,
Hymenoptera, Formicidae). Entomologische Abhandlungen. Staatliches Museum für
Tierkunde Dresden 51: 143-180.
Seifert, B. 1992. A taxonomic revision of the Palaearctic members of the ant subgenus Lasius
s.str. (Hymenoptera: Formicidae). Abhandlungen und Berichte des Naturkundemuseums
Görlitz 66 (5): 1-67.
Steiner, F.M., Schlick-Steiner, B.C., Schödl, S., Espadaler, X., Seifert, B., Christian, E. &
Stauffer, C. 2004. Phylogeny and bionomics of Lasius austriacus (Hymenoptera,
Formicidae). Insectes Sociaux 51: 24-29.
Wilson, E.O. 1955. A monographic revision of the ant genus Lasius. Bulletin of the Museum of
Comparative Zoology 113: 1-201.
Wilson, E.O. 1957. The tenuis and selenophora groups of the ant genus Ponera (Hymenoptera:
Formicidae). Bulletin of the Museum of Comparative Zoology 116: 355-386.
Wilson, E.O. 1958a. Studies on the ant fauna of Melanesia. I. The tribe Leptogenyini. II. The
tribes Amblyoponini and Platythyreini. Bulletin of the Museum of Comparative Zoology
118: 101-153.
Wilson, E.O. 1958b. Studies on the ant fauna of Melanesia III. Rhytidoponera in western
Melanesia and the Moluccas. IV. The tribe Ponerini. Bulletin of the Museum of Comparative
Zoology 119: 303-371.
Wilson, E.O. 1959a. Adaptive shift and dispersal in a tropical ant fauna. Evolution 13: 122-144.
Wilson, E.O. 1959b. Studies on the ant fauna of Melanesia V. The tribe Odontomachini.
Bulletin of the Museum of Comparative Zoology 120: 483-510.
Ward: Wilson - contributions to ant systematics 7
Wilson, E.O. 1959c. Studies on the ant fauna of Melanesia. VI. The tribe Cerapachyini. Pacific
Insects 1: 39-57.
Wilson, E.O. 1961. The nature of the taxon cycle in the Melanesian ant fauna. American
Naturalist 95: 169-193.
Wilson, E.O. 1964. The true army ants of the Indo-Australian area (Hymenoptera: Formicidae:
Dorylinae). Pacific Insects 6: 427-483.
Wilson, E.O. 1971. The insect societies. Cambridge, Massachusetts: Harvard University Press, x
+ 548 pp.
Wilson, E.O. 1985a. Ants of the Dominican amber (Hymenoptera: Formicidae). 1. Two new
myrmicine genera and an aberrant Pheidole. Psyche (Cambridge) 92: 1-9.
Wilson, E.O. 1985b. Ants of the Dominican amber (Hymenoptera: Formicidae). 2. The first
fossil army ants. Psyche (Cambridge) 92: 11-16.
Wilson, E.O. 1985c. Ants of the Dominican amber (Hymenoptera: Formicidae). 3. The
subfamily Dolichoderinae. Psyche (Cambridge) 92: 17-37.
Wilson, E.O. 1985d. Ants from the Cretaceous and Eocene amber of North America. Psyche
(Cambridge) 92:205-216.
Wilson, E.O. 1986. Ants of the Dominican amber (Hymenoptera: Formicidae). 4. A giant
ponerine in the genus Paraponera. Israel Journal of Entomology 19: 197-200.
Wilson, E.O. 1987. The earliest known ants: an analysis of the Cretaceous species and an
inference concerning their social organization. Paleobiology 13: 44-53.
Wilson, E.O. 1994. Naturalist. Washington, D.C.: Island Press, xii + 380 pp.
Wilson, E.O. 2003. Pheidole in the New World. A dominant, hyperdiverse ant genus.
Cambridge, Massachusetts: Harvard University Press, [ix] + 794 pp.
Wilson, E.O. & Brown, W.L., Jr. 1953. The subspecies concept and its taxonomic application.
Systematic Zoology 2: 97-111.
Wilson, E.O. & Brown, W.L., Jr. 1955. Revisionary notes on the sanguinea and neogagates
groups of the ant genus Formica. Psyche (Cambridge) 62: 108-129.
Wilson, E.O., Carpenter, F.M. & Brown, W.L., Jr. 1967. The first Mesozoic ants, with the
description of a new subfamily. Psyche (Cambridge) 74: 1-19.
Wilson, E.O., Eisner, T., Wheeler, G.C. & Wheeler, J. 1956. Aneuretus simoni Emery, a
major link in ant evolution. Bulletin of the Museum of Comparative Zoology 115: 81-99.
Wilson, E.O. & Taylor, R.W. 1967. The ants of Polynesia (Hymenoptera: Formicidae). Pacific
Insects Monograph 14: 1-109.
Alpert, G. D. 2007. A review of the ant genus Metapone Forel from Madagascar, pp. 8-18. In
A REVIEW OF THE ANT GENUS METAPONE FOREL

FROM MADAGASCAR
Gary D. Alpert
Entomology Department
Museum of Comparative Zoology
Harvard University, 26 Oxford Street, Cambridge, MA 02138 U.S.A.
gary_alpert@post.harvard.edu
ABSTRACT
The pattern of obligatory association between the Old World ant genus Metapone and termites is
confirmed. Metapone vincimus, new species, is described based upon workers and a queen
collected from northeastern Madagascar. Several nest series of M. vincimus were collected from
within a log in association with the dry wood termite, Cryptotermes kirbyi (Kalotermitidae). An
illustrated key is provided to separate the three species of Metapone known from Madagascar. A
remarkable ergatoid male caste is noted for Metapone madagascarica.
Key words: Hymenoptera, Formicidae, Metapone, new species, Madagascar, taxonomy,

ergatoid male, termites
Alpert: Metapone of Madagascar 9
INTRODUCTION
The genus Metapone Forel remains one of the most unusual and enigmatic ant genera in the Old
World. Most of the 17 described taxa are known from very few specimens, often single queens
or males are collected during dispersal flights and occasionally workers are collected from dead
wood. Forel (1911) established the genus Metapone based upon a series of workers, larvae and
reproductive pupae of the type species M. greeni Forel, collected from Peradenyia, Sri Lanka. In
the same paper, Forel designated Metapone as the type genus of a new tribe Metaponini and
provisionally placed this tribe into a new special section among the Ponerinae, which he called
the Promyrmicinae.
The genus Metapone was properly transferred to the subfamily Myrmicinae a year later by
Emery (1912) when he realized that the larvae were characteristic of the Myrmicinae and not the
Ponerinae. Unfortunately, Emery misinterpreted certain ponerine-like morphological traits of
Metapone as primitive rather than as derived adaptations to a highly specialized habit of foraging
in galleries of wood. As a result, Emery retained Forel’s section Promyrmicinae, transferred it to
the Myrmicinae, and expanded it to include the tribes Metaponini and Pseudomyrmicini.
Wheeler (1919) was able to clear up much of the earlier confusion after additional species of
Metapone became available for study. Wheeler rejected the term Promyrmicinae even though he
could not determine the position of the Metaponini among the other tribes within the
Myrmicinae. The genera Metapone, Liomyrmex, Vollenhovia and Xenomyrmex were included
until recently in the tribe Metaponini (Bolton, 1994, 1995). Metapone is presently the only genus
assigned to the tribe Metaponini (Bolton, 2003). The placement of the Metaponini within the
Myrmicinae remains problematic and awaits a comprehensive revision of the myrmicine tribes.
The genus Metapone is widely distributed throughout the Indo-Australian, Oriental and
Malagasy Regions (Taylor, 1991; Bolton, 1995; Eguchi, 1998). Even though specimens are rare,
new species continue to be discovered. Eguchi (1998) described a new species from Borneo, an
undescribed Metapone has been found in Gabon (B. Bolton, B. Fisher, personal communication)
and another undescribed Metapone has been found recently in New Caledonia (C. Burwell,
personal communication). There are several series of undescribed Metapone from Papua New
Guinea and Australia in the Australian National Insect Collection in Canberra, Australia. The
new species described in this paper, M. vincimus, is the third endemic Metapone species known
from Madagascar.
From the very first collections, Metapone workers have been found in association with
termites. The type species of the genus, M. greeni, was found in “galleries in a decayed branch,
which was also infested by two species of termites” (Forel, 1911, quoting E.E. Green, the
collector). Subsequently, Wheeler (1919, 1936) listed M. greeni as an inquiline of termites and
suggested that all species of Metapone probably form small colonies and live in or near the
galleries of termites in dead wood. Taylor (1991) has found several undescribed species of
Metapone in association with termites in Australia. In Papua New Guinea, Metapone were found
in rotten wood with termites of the genus Prorhinotermes (Leigh Miller, personal
communication). Eguchi (1998) reported that M. quadridentata Eguchi from Borneo “was
collected from the galleries of a termite nest in rotten wood”. Gregg (1958) obtained from Alfred
E. Emerson, the noted termite specialist, two new species of Metapone (M. emersoni and M.
madagascarica) from a series of termites collected from Madagascar by Harold Kirby in 1935.
Additional specimens of M. madagascarica were collected in association with Coptotermes
truncatus Wasmann and Cryptotermes sp. (Brian Fisher, written communication). Though the
evidence is anecdotal, the pattern of obligatory association with termites is clear. The new
species of Metapone described in this paper, M. vincimus, was found in association with a dry
wood termite, Cryptotermes kirbyi (Paul Eggleton, written communication). Live colonies of M.
madagascarica and M. vincimus along with their termite associates were transported to the
laboratory of Bert Hölldobler in Würzburg, Germany. The resulting studies (Hölldobler et al.,
2002a, 2002b) are the first in-depth investigations of the Metapone-termite relationship.
METHODS
Specimens were examined using a Leica MZ 16 binocular microscope. Digital images were
taken with a JVC digital camera and processed using Auto-Montage (Syncroscopy, Divison of
Synoptics, LTD) software. Morphological measurements follow the terminology of Eguchi
(1998) and include: head length, measured from the midpoint of a line drawn across from the
posterior margin of the head to the anterior clypeal margin (HL), head width (HW), cephalic
index (CI), scape length (SL), dorsal length of mesosoma excluding posterior declivities of the
propodeum (ML), promesonotal length in dorsal view (PML), maximum promesonotal width
(PMW), petiolar node length measured from the centers of the anterior and posterior faces
(PNL), maximum petiolar width in dorsal view (PW) and maximum postpetiole width in dorsal
view (PPW).
Material was examined from the Museum of Comparative Zoology, Harvard University,
Cambridge, MA, USA, (MCZC), California Academy of Sciences, San Francisco, California,
USA (CASC) and the Field Museum of Natural History, Chicago, Illinois, USA, (FMNH).
KEY TO WORKERS OF METAPONE FROM MADAGASCAR
1 Abdominal segment 2 from above twice as wide as long (Figs. 11, 12), third segment in
profile with a stout, ventrally directed spine (Fig. 6)................................ M. emersoni Gregg
-- Abdominal segment 2 from above quadrate to subquadrate (Figs. 8, 10), third segment with
an anteroventral transverse ridge that appears as a blunt tooth (Figs. 2, 4) ............................ 2
2 Abdominal segment 2 from above with narrow anterior margin, sides strongly diverging
posteriorly to form a wide, strongly concave posterior margin (Fig. 8) ..... M. vincimus, n. sp.
-- Abdominal segment 2 from above with sides weakly diverging posteriorly to form a slightly
wider, somewhat less concave posterior margin (Fig. 10)............... M. madagascarica Gregg
Metapone emersoni Gregg

Figures 5, 6, 11, 12, 17, 18
Gregg (1958) described this species and provided an illustration of the petiole and
postpetiole in lateral view. Additional figures are provided here of the entire ant for comparison
purposes. Gregg based his description on two workers found within a series of termites collected
12 miles from Perinet, Madagascar on June 28, 1935 by Harold Kirby. Kirby attached a locality
code T-4503, his record label T specifying a collection of termites. Many of Kirby’s termite
collections were later identified to species in a series of papers including Moszkowski’s 1955
paper on Malagasy termites. T-4503 however has never been listed as a locality code for
Cryptotermes kirbyi, or any other termite, leaving in question the identification of the host
termite for M. emersoni. One worker of M. emersoni has been collected subsequently from a
more northern location, (Toamasina, P.N. Mantadia, 18˚47’30”S, 48˚25’36”E, 895m, rainforest,
25 Nov-1Dec 1998, coll. H.J. Ratsirarson, HJR020, CASENT0003116, CASC).
A single queen assigned to M. emersoni was collected from the Andohahela Special Reserve
(24˚46’35”S, 45˚42’19”E), in southern Madagascar (P. Rabeson, 4.II. 1993, MCZC). When M.
emersoni is rediscovered and queens are collected in association with workers, then the status of
this single queen can be confirmed.
Metapone madagascarica Gregg

Figures 3, 4, 9, 10, 15, 16, 19-24
Gregg (1958) described this species and included an illustration of a worker in dorsal view.
Additional figures are provided here. M. madagascarica was described from a small series of
workers collected along with termites from a stump with a field label T–4403. Moszkowski
(1955, p. 34) described the associated drywood termite as a new species, Cryptotermes kirbyi
Moszkowski, gave the field label as T-4403, and described the collection locality in more detail
[14 km. East (23˚ 20’S, 43˚48’E) of Tulear, along Fiheranana River, Madagascar, coll. H. Kirby,
7.VI.1935, in large dead stump]. On February 10, 1993, Phil Ward, Emile Rajeriarison and the
author discovered a series of M. madagascarica from Berenty Reserve, 15m, 25˚01’ 3.9”S,
46˚18’ 21.8”E; spiny forest, in an Alluaudia sp. log in association with termites. On July 11,
2000, the author returned to this locality with Pascal Rabeson and Emile Rajeriarison and
collected several nest series of M. madagascarica in a dead hardwood tree (local Malagasy name
“Kelegnogne”) in association with the termite Cryptotermes kirbyi. These nest series included
larvae, pupae, workers, queens, males and ergatoid males. Additional material examined from
Madagascar: Toliara, Reserve Privé Berenty, Foret d’Anjapolo, 21.4 km 325˚ NW Ambosary,
24˚55’47”S, 46˚12’35”E, 65m, spiny forest/thicket, 7 Feb 2002 (coll. Fisher-Griswold
Arthropod Team) CASENT0004524, 1 ergatoid male, CASC; Reserve Privé Berenty, Foret de
Malaza, Mandrare River, 8.6 km 314˚ NW Ambosary, 25˚0’28”S, 46˚18’22”E, 40 m, gallery
forest, 6 Feb 2002 (coll. Fisher-Griswold Arthropod Team BLF5474) CASENT0004525, 1
worker, CASC; Parc National d’Andohahela, Foret de Manantalinjio, 33.6 km 63˚ENE
Ambosary, 7.6 km 99˚E Hazofotsy, 24˚49’1”S, 46˚36’36”E, 150 m, spiny forest/thicket, 12-16
Jan 2002 (coll. Fisher-Griswold Arthropod Team BLF4837) CASENT0004526, BLF4840,
CASENT0004529, 2 workers, CASC.
Worker-queen intermorphs were present in low numbers among the series of workers. Two
intermorphs exhibiting rudimentary wing articulations were present among 51 workers. Other
intermediate stages of intermorph development were also present and this is consistent with the
variation in development of ovarioles from six to two (Hölldobler et al., 2002b).
Several ergatoid males (Figs. 19, 21, 23) were collected along with typical winged males
(Figs. 20, 22, 24) within the same colony at more than one location. These ergatoid males are
almost identical to workers with the following exceptions, presence of male genitalia, large well-
developed eyes and typical male antennae. The presence of both winged males and an ergatoid
male caste in the same colony is exceptional in ants and warrants further study.
Metapone vincimus, new species

(Figs. 1, 2, 7, 8, 13, 14)
Diagnosis - Similar to M. madagascarica but readily distinguished by the following

characters: Transverse metanotal groove prominent and continuous across the mesosomal
dorsum. Petiole in dorsal view with strongly divergent sides, so that the flat anterior margin is
much shorter than the concave posterior margin. Median anterior lobe of clypeus blunt, not
forming teeth.
Description - Worker. Measurement for holotype given first, followed in brackets by range
for 15 paratype workers examined. HL 1.20 mm [1.20-1.44]; HW 0.90 mm [0.90-1.06]; ML
1.60 mm [1.58-1.90]; PML 0.90 mm [0.90-1.24]; PMW .64 mm [0.60-0.76]; PNL 0.34 mm
[0.32-0.46]; PW 0.44 mm [0.44-0.68]; PPW 0.46 mm [0.44-0.62].
Head in full-face view distinctly longer than broad (CI 0.75 [0.74-0.81]), lateral margins
sub-parallel, posterior margin broadly concave. Compound eyes small, reduced to about 6
weakly defined ommatidia, located near the mid-line of the head. Ocellar pits and ocelli absent.
Mandibles stout, with 5 rounded teeth, increasing in size progressively towards the mandibular
apices. Anterior clypeal margin with a quadrate median lobe formed into a blunt anterior edge
(0.10 mm wide). The vertex with a short median suture that extends from the base to the
posterior margin. Frontal carinae, widely separated, parallel and forming deep antennal scrobes.
The frontal carinae originating from the posterior border of the clypeus, diverging transversely
and extending posteriorly to behind the eye. Lateral clypeal lobes narrow, oblique, and separated
from the genae by distinct lines continuous with the posterior border of the clypeus. A series of
parallel striae begin just below the eye and extend across the posterior border of the lateral
clypeal margin.
Antennae 11-segmented, the scape short and flattened dorso-ventrally, SL 0.34 mm, dorsal
surface of scape with scattered erect hairs. First segment of funiculus elbowed, segments 2-7
gradually expanding before developing into a flattened, 3-segmented club. Numerous hairs on all
surfaces of the funicular segments.
Mesosoma long and narrow, the promesonotum separated from the propodeum by a well-
developed, complete, transverse metanotal groove. Entire mesosoma marginate to submarginate
laterally, margin becoming more distinct at the propodeal corners. Propodeal dorsum nearly
horizontal, passing through an abrupt concave angle to the vertical, posterior face. Petiole
weakly convex dorsally. In dorsal view, the anterior margin of the petiolar node straight, the
posterior margin deeply concave. Petiole ventrally with a thin, translucent median rounded keel.
In profile, the anterior and posterior face of the petiole strongly concave. In profile, postpetiole
almost flat dorsally. In dorsal view, anterior margin of postpetiole straight, posterior margin
slightly concave, the anterodorsal corners diverging to rounded posterodorsal corners. The
postpetiole joined to the gaster by a wide face, although a deep constriction exists between the
two segments. Anteroventral surface of postpetiole produced into a pronounced, triangular
rounded lobe or blunt tooth in side view.
The first gastric segment almost as long as the remaining gastral segments combined. Gaster
with numerous hairs arising from foveae, surface between foveae smooth and shiny. Coxae
stout and bulbous. Femora swollen and laterally compressed, the ventral surface longitudinally
grooved for the reception of the tibia. Tibia also stout and partly compressed. Protibia armed
apically with 1 small spine and a large pectinate spur. Mesotibia with a small, barely pectinate
spur, and 3 stout apical teeth. Apical tarsomere on all legs with simple claws.
Sculpture: Clypeus, frons, genae and antennal scrobes covered with fine, longitudinal striae,
essentially parallel, but which fade posteriorly, leaving the vertex, occiput and posterior part of
the genae smooth and shining, interrupted only by prominent punctures. Centrally two parallel
striae forming a groove that extends from the clypeal border to just beyond the limits of the
frontal carinae. Entire dorsum of mesosoma longitudinally striate. Dorsal surface of petiole with
hair-bearing punctures. Postpetiole and gaster with similar but finer punctures. All areas of the
body, including the legs and antennae, smooth and shiny.
Pilosity: Short, scattered, erect yellow hairs on all surfaces of head and mesosoma, many
hairs arising from punctures on the vertex, occiput and petiole. Hairs longer on mandibles,
anterior margin of the clypeus, lateral surfaces of legs and lower surfaces of petiole and gaster.
Pilosity most abundant on postpetiole and gaster.
Color: Head, mandibles and postpetiole dark reddish brown, the mesosoma and gaster a
lighter brown. Petiole, legs and antennae light yellowish brown.
Type Material. – Holotype worker from MADAGASCAR, 30km N of Antalaha, 3km W

to a hill, near Amboangy, 50 m, 14˚39’53.3”N, 50˚11’26.5”E, 29.VII.2000, secondary rainforest,
in log with termite Cryptotermes kirbyi, G. Alpert, P. Rabeson and E. Rajeriarison, #2278.
Deposited in MCZC. Paratype workers. 15 workers and 1 queen-worker intermorph, collected
from the same nest series as the holotype. Alate queen found in same locality on 24.I.1991.
Paratypes have been deposited in the following collections; Los Angeles County Museum of
Natural History, Los Angeles, California, USA, (LACM); California Academy of Sciences, San
Francisco, California, USA, (CASC); Bohart Museum of Entomology, University of California,
Davis, California, USA, (UCDC); The Natural History Museum, London, England (BMNH);
Australian National Insect Collection, Canberra, Australia (ANIC); and the National Museum of
Natural History, Smithsonian Institution, Washington, D.C., USA, (USNM). Paratypes and
voucher specimens including larvae are deposited in the Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts, USA, (MCZC). Cryptotermes termite voucher
specimens have been deposited in the American Museum of Natural History, New York, USA,
(AMNH); the Natural History Museum, London, England (BMNH); and the Museum of
Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA, (MCZC).
Queen-worker intermorph. Same characters as worker apart from being larger in most
dimensions, darker and having relatively larger postpetiole. Ocelli absent.
Queen. Same as worker with the following exceptions: Larger size, additional thoracic
sclerites, HL 1.20 mm; HW 0.98 mm; AL 2.20 mm; PW 0.46 mm; PNL 0.44 mm, dealate and
uniformly black in color. Three ocelli present and large compound eyes.
Male. Unknown.
Distribution. - Known only from the type locality, a secondary lowland rainforest habitat.
Etymology. - The specific name, vincimus = “we succeed”, is in recognition of the joint
effort expended in discovering this species. Extremely hard logs were searched by axe for weeks
before finally locating the ants and associated termites.
Remarks. - At least two separate M. vincimus colonies were present in the same fallen tree
(local Malagasy name “Ompa”). The colonies were separated by over 5 meters along the log and
workers attacked and killed members of the other colony when put together into a common
foraging arena. Workers and brood were found in chambers just below the log surface in close
proximity to Cryptotermes termite chambers. Colony size was small, one colony containing less
than 20 workers and about 40 larvae. A queen mesosoma along with worker remains was found
in a refuse chamber of this colony. Since queens are known for this species it is probable that
after the death of the primary reproductive, the colony is still able to produce workers via the
presence of gamergates (Hölldobler et al., 2000b).
ACKNOWLEDGEMENTS
This paper is dedicated to the recognition and appreciation of Edward O. Wilson, who has
supported the author for more than two decades with encouragement, advice and funding.
“Vincimus” (success through perseverance) was the operative word not only for the Shackelton
Expedition but also for the “Wilson Expedition” to explore myrmecological life on earth. I
acknowledge all who are undertaking this enterprise for their efforts and for their
encouragement.
Emile Rajeriarison and Pascal Rabeson were invaluable as field assistants and as
collaborators in the effort to discover colonies of Metapone. I thank Stefan Cover, Steve
Shattuck, Brian Fisher and Phil Ward for their review of this manuscript. I also thank the
Madagascar Institute pour le Conservation de l’Environment Tropicaux (MICET) for collection
permits and logistical support in Madagascar.
LITERATURE CITED
Bolton, B. 1994. Identification Guide to the Ant Genera of the World:Harvard University Press,
Cambridge, Massachusetts. 222pp.
Bolton, B. 1995. A New General Catalogue of the Ants of the World: Harvard University Press,
Cambridge, Massachusetts. 504 pp.
Bolton, B. 2003. Synopsis and classification of Formicidae. Memoirs of the American
Entomological Institute 71: 1-370.
Emery, C. 1912. Études sur les Myrmicinae. Annales de la Société Entomologique de Belgique
56: 94-105.
Eguchi, K. 1998. A new ant of the genus Metapone (Hymenoptera: Formicidae) from Sabah,
Borneo. Entomological Science 1: 605-609.
Forel, A. 1911. Sur le genre Metapone n.g. nouveau groupe des formicides et sur quelques
autres formes nouvelles. Revue Suisse de Zoologie 19: 445-459.
Gregg, R.F. 1958. Two new species of Metapone from Madagascar. Proceedings of the
Entomological Society of Washington 60: 111-121.
Hölldobler, B., Oldham, N.J., Alpert, G.D. & Liebig, J. 2002a. Predatory behavior and
chemical communication two Metapone species (Hymenoptera: Formicidae). Chemoecology
12: 147-151.
Hölldobler, B., Liebig, J. & Alpert, G.D. 2002b. Gamergates in the myrmicine genus
Metapone (Hymenoptera: Formicidae). Naturwissenschaften 89: 305-307.
Moszkowski, L. I. 1955. Cryptotermes kirbyi, new species from Madagascar and C. havilandi
(Sjöstedt) from Africa and introduced into Madagascar, India and South America (Isoptera:
Kalotermitidae). Memoires de L’Institut Scientifique de Madagascar. Série E. 6: 15-41.
Taylor, R.W. 1991. The nomenclature and distribution of some Australasian ants of the
Myrmicinae (Hymenoptera: Formicidae). Memoirs of the Queensland Museum 30: 599-614.
Wheeler, W.M. 1919. The ants of the genus Metapone Forel. Annals of the Entomological
Society of America 12: 173-191.
Wheeler, W.M. 1936. Ecological relations of Ponerine and other ants to termites. Proceedings
of the American Academy of Arts and Sciences 71(3): 159-243.
Figures 1-6. Metapone vincimus holotype worker (1, 2); M. madagascarica holotype worker (3,
4); M. emersoni holotype worker (5, 6).
Figures 7-12. Metapone vincimus holotype worker (7, 8); M. madagascarica holotype worker
(9, 10); M. emersoni holotype worker (11, 12).
Figures 13-18. Metapone vincimus queen (13, 14); M. madagascarica queen (15, 16); M.
emersoni queen (17, 18).
Figures 19-24. Metapone madagascarica ergatoid male (19-23); M. madagascarica male (20-
24).
Andersen, A. N. 2007. Ant diversity in arid Australia: a systematic overview, pp. 19-51. In
ANT DIVERSITY IN ARID AUSTRALIA: A SYSTEMATIC OVERVIEW
Alan N. Andersen
CSIRO Sustainable Ecosystems
Tropical Ecosystems Research Centre
PMB 44 Winnellie
NT 0822, Australia
Alan.Andersen@csiro.au
ABSTRACT
Australia is one of the world’s hotspots for ant diversity, but diversity patterns within the
Australian ant fauna remain poorly documented. The key to understanding patterns of ant
diversity in Australia is an understanding of its arid fauna, as this is primarily where most of
Australia’s highly diverse genera have radiated. Based on the 3,750 Australian ant species held
at the CSIRO Tropical Ecosystems Research Centre in Darwin, I estimate that the entire
Australian fauna comprises about 6,500 species, only about one-fifth of which have been
described. Species richness is heavily concentrated into 11 genera, each with over 100
Australian species. Most of these occur primarily in arid and semi-arid regions, and a systematic
overview is presented for five of the most important: Iridomyrmex, Melophorus, Camponotus,
Monomorium and Rhytidoponera. For each genus, a comprehensive species-group framework is
proposed as a basis for documenting patterns of diversity and distribution. Many of the taxa are
highly generalised morphologically, and much remains to be done to resolve difficult complexes
at the species level, and to resolve relationships between species-groups. An answer to the
fundamental question of why ant diversity is so remarkably high in arid Australia remains
elusive, but may be related to unusually high ant productivity and behavioural dominance over
evolutionary time.
Key words: ant diversity, arid, Australia, Camponotus, Formicidae, Hymenoptera, Iridomyrmex,
Melophorus, Monomorium, Rhytidoponera
20 Memoirs of the American Entomological Society, Volume 80
INTRODUCTION
Australia is one of the world’s hotspots for ant diversity, but little systematic information is
available on diversity patterns within the Australian ant fauna. The exceptional richness of local
ant communities in Australia has been commonly reported (Andersen, 1995), but species
richness within entire regional faunas have been poorly documented. Exactly how rich is the
Australian ant fauna, and how is the richness distributed both from systematic and
biogeographical perspectives? These questions can only be answered by a detailed
understanding of diversity patterns within the major Australian ant genera, most of which have
many more undescribed than described species, and lack comprehensive species-group
frameworks.
It is clear that the key to understanding patterns of ant diversity in Australia is an
understanding of the arid fauna. Most of Australia’s ‘megadiverse’ (>100 species) ant genera
have radiated primarily in the arid zone, and whichever way you look at it ant diversity in arid
Australia is extraordinary by world standards (Table 1). Inland Australia routinely packs more
than a hundred ant species per hectare, and in semi-arid regions one can find a hundred species
within 0.1 ha (Andersen, 2003). These are the richest local ant communities on Earth. As many
or more species can occur within a comparable area of lowland tropical rainforest, but here there
is extreme vertical stratification of species (Brühl et al., 1998), so that we are really dealing with
multiple communities stacked on top of each other. Ironically, Australia’s own lowland tropical
rainforests are relatively poor in species, supporting a depauperate subset of the rich Indo-
Malayan fauna (Taylor, 1972).
Table 1. Ant diversity at local and regional scales in arid Australia compared with arid North
America and southern Africa. Data are maximum numbers of species recorded in local (within 1
ha) communities (see Andersen, 1997, 2003), and broad estimates of total arid faunas.
Australia North America Southern

Africa
Max no. species/ha >100 35 30-40
Total no. species 4,500 300 400
Here I provide a systematic overview of diversity within the ant fauna of arid Australia. I
use the term ‘arid’ very broadly to include semi-arid regions - up to about 350 mm annual
rainfall in the southern semi-arid zone, and about 800 mm in the northern semi-arid tropics
(where rain falls almost exclusively during a summer wet season). I begin by providing an
overview of the diversity of the entire Australian fauna, before examining in detail the five major
genera of the arid zone. For each genus, a comprehensive species-group framework is proposed
as a basis for documenting patterns of diversity and distribution.
My analyses are based on specimens held at the CSIRO Tropical Ecosystems Research
Centre (TERC) in Darwin that my collaborators and I have accumulated over the past 25 years.
These holdings represent by far the largest collection of ants from arid Australia. As of January
2004 they comprise over 3,750 native Australian species, and have been accumulating at a
relatively constant 200 additional species per year over the past decade, without any sign of
levelling off (Fig. 1).
Andersen: Ant biodiversity in arid Australia 21
4000
3500
3000
2500
No. species
2000
1500
1000
500
0
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004
Fig. 1. Accumulation of Australian ant species in the CSIRO Tropical Ecosystems Research
Centre collection over the past decade.
DIVERSITY OF THE AUSTRALIAN ANT FAUNA
Shattuck (1999) lists 1275 described species and subspecies of ants from Australia. Based on
Taylor’s (1991) opinion that “Only about a quarter of the Australian fauna has been named”, this
gives an estimated 5,000 Australian species. My own estimate based on the TERC collection is
somewhat higher. About 40% (38 out of 103) of Australian genera contain twenty or more
species, and I estimate that together they contribute about 6,000 species (Table 2). This would
put the total Australian fauna at about 6,500 species.
A very large proportion (collectively about 80% of total species) of the fauna is contributed
by 11 megadiverse (each with over 100 Australian species) genera (Table 2). Chief among these
is Melophorus, with over 500 sorted species in the TERC collection and possibly as many more
yet to be collected. Next is Camponotus, likely to have well over 500 Australian species.
Monomorium, Pheidole, Meranoplus, Rhytidoponera, Polyrhachis, Iridomyrmex and
Tetramorium each are likely to have more than 300 Australian species. Cerapachys and
Myrmecia complete the 11 ‘centurion’ Australian genera. Such an array of megadiverse genera
is unparalleled anywhere in the world, with other continents having at most two or three genera
containing one hundred or more local species. Interestingly, the frequency distribution of species
richness in Australian ant genera is bimodal, with about twice as many genera containing more
than a hundred species as those containing between 50 and 100 (Fig. 2).
The ratio of undescribed to described Australian species is very uneven among genera
(Table 2). Myrmecia is by far the best known taxonomically of Australia’s megadiverse genera
in terms of proportion of species that are named (Ogata & Taylor, 1991), but even then a
significant proportion remains undescribed. However, it is likely that most of these undescribed
species are already represented in collections, awaiting the resolution of difficult species
complexes, such that the total species count is unlikely to be boosted markedly by the collection
Table 2. Diversity within Australia’s richest (those with 20 or more species) genera. Data are
provided for the number of described species (including subspecies; following Shattuck, 1999
and Bolton, 2000), the number of sorted species in the CSIRO Tropical Ecosystems Research
Centre (TERC) collection in Darwin (as of the end of 2004), and an estimate of total number of
Australian species. The ratio of estimated total: number of described species is also provided.
Estimated Total:
Described TERC Total Described
Myrmeciinae
Myrmecia 89 90 130 1.5
Amblyoponinae
Amblyopone 17 7 25 1.5
Ponerinae
Anochetus 5 25 30 5.0
Hypoponera 10 24 35 3.5
Leptogenys 25 35 55 2.2
Odontomachus 3 25 35 11.7
Bothroponera* 20 42 60 3.0
Ectatomminae
Rhytidoponera 76 247 350 3.25
Cerapachyinae
Cerapachys 44 95 150 3.4
Sphinctomyrmex 15 15 25 1.7
Myrmicinae
Crematogaster 34 60 80 2.4
Colobostruma 16 16 25 1.6
Epopostruma 18 12 30 1.7
Meranoplus 59 217 400 6.8
Monomorium 53 335 500 9.4
Orectognathus 18 3 20 1.1
Pheidole 53 297 500 9.4
Podomyrma 49 25 75 1.5
Solenopsis 7 26 40 5.7
Strumigenys 49 18 60 1.2
Tetramorium 24 192 350 14.6
Dolichoderinae
Anonychomyrma 14 25 35 2.5
Bothriomyrmex 5 13 20 4.0
Dolichoderus 22 28 40 1.8
Iridomyrmex 63 223 350 5.6
Table 2. (continued)
Estimated Total:
Described TERC Total Described
Leptomyrmex 27 16 30 1.1
Ochetellus 4 18 30 7.5
Papyrius 4 16 25 6.3
Tapinoma 5 22 30 6.0
Formicinae
Calomyrmex 11 19 25 2.3
Camponotus 128 485 750 5.9
Melophorus 29 505 1000 34.5
Notoncus 6 30 40 6.7
Opisthopsis 15 19 25 1.7
Paratrechina 10 55 80 8.0
Polyrhachis 115 241 350 3.0
Prolasius 18 34 45 2.5
Stigmacros 48 58 80 1.7
TOTAL 1119 3606 5920 5.3

*The synonymy of Bothroponera with Pachycondyla listed in Bolton (1994) is not recognized
here as it is based on an unpublished manuscript.
of significantly new material. Species of Myrmecia are very large and extremely conspicuous
ants, and, like most Australians, occur primarily in the temperate south. In other words, the
genus is unusually well collected. Indeed, Australia’s four richest genera in terms of described
species (Camponotus, Polyrhachis, Myrmecia and Rhytidoponera) are all among the country’s
largest and most conspicuous ants, and therefore have been subject to relatively high collecting
effort.
Polyrhachis stands out as being in a far better taxonomic state than indicated by the
relatively small proportion (about a third) of Australian species that are described, thanks to
decades of work by Rudy Kohout from the Queensland Museum. All known Australian species
outside the subgenera Chariomyrma and Campomyrma have been catalogued and assigned
manuscript names. In each of the rich sub-genera Hagiomyrma, Hedomyrma and Cyrtomyrma
there are more manuscript names than described species (R. Kohout, personal communication).
All these subgenera have their centres of Australian distribution in relatively well-collected
Queensland. In comparison, very large numbers of species of the far richer Chariomyrma and
Campomyrma occur in poorly collected central and northwestern Australia; the proportion of
undescribed species is far higher in these subgenera.
The most dramatic mismatch between described and actual species undoubtedly occurs in
Melophorus, where only 29 of the possibly thousand or more species have been named. The
majority of species are small, occur in remote regions, and forage during the hot parts of the day
when most ants and ant collectors alike are inactive. The genus has never attracted serious
attention by ant taxonomists. The true extent of their diversity has only been revealed during
extensive pitfall trapping over the past decade, involving literally thousands of sites across
central and northern Australia. The mismatch between described and actual species is also
especially pronounced for Tetramorium (Table 2). As in Melophorus, most Australian species of
Tetramorium are relatively small and inconspicuous, and occur in remote regions of the arid
zone.
25
20
No. genera
15
10
0
1 2-5 6-20 21-50 50-100 >100
No. species/genus
Fig. 2. Frequency distribution of species richness in Australian ant genera. Numbers of species
per genus have been estimated based on Shattuck (1999) and holdings in the TERC collection.
DIVERSITY WITHIN MAJOR ARID GENERA
Of Australia’s eleven megadiverse genera, all but Polyrhachis and Myrmecia have radiated
primarily in the arid zone. In Meranoplus, Tetramorium, Iridomyrmex, Cerapachys and
especially Melophorus the vast majority of species occur in arid habitats, whereas
Rhytidoponera, Monomorium, and especially Pheidole and Camponotus have a greater
proportion of species in mesic habitats. Here I examine in detail diversity within the five most
important of these arid-adapted genera: Iridomyrmex, Melophorus, Camponotus, Monomorium
and Rhytidoponera, building on preliminary species-group frameworks provided in Andersen
(1991, 2000).
Iridomyrmex
Iridomyrmex might not be the richest ant genus in Australia, but is easily the most important
ecologically, especially in the arid zone (Greenslade, 1979). Species of Iridomyrmex are highly
abundant and aggressive ants that exert a level of behavioural dominance over other ants that is
unparalleled in arid regions elsewhere in the world (Andersen, 2003). All species of
Iridomyrmex are sun-loving ants, with the vast majority occurring in arid and semi-arid habitats.
The genus can be reasonably well-represented in more humid regions, but tends to be restricted
to open habitats, or to open patches within more forested habitats.
A comprehensive species-group framework for Iridomyrmex is presented in Table 3. It
needs to be noted that the species- and even higher-level taxonomy of the genus is extremely
challenging, given that morphology is highly generalised and conservative throughout the genus.
There are many complexes of unresolved sibling species, and in many cases morphological
similarity between species might well be a result of convergence. The species-group framework
proposed here is therefore very preliminary.
Table 3. Classification, distribution and diversity of major species-groups and complexes of

Iridomyrmex in Australia. Figures are numbers of sorted species in the TERC collection as of the
end of 2004. Distribution is described as predominantly northern (N; occurring primarily within
the northern arid zone), southern (S; occurring primarily in the southern arid zone), monsoonal
(M; occurring primarily in the monsoonal tropics), widespread (W; well-represented throughout
arid Australia), or Bassian (B; occurring primarily in humid habitats of southern Australia).
calvus radiation
calvus group (3, B)
rufoinclinus group (6, N)
viridigaster group (3, S)
agilis radiation
agilis group (9, S)
anceps group (14, M)
bicknelli group (10, S)
gracilis group (25, W)
pallidus group (41, W)
purpureus group (12, W)
discors complex (1, S)
purpureus complex (11, W)
rufoniger radiation
conifer group (7, B)
mattiroloi group (44, W)
rufoniger group (21, W)
suchieri group (15, W)
vicinus group (10, B)
Group A (3, M)
calvus radiation
This is a relatively small but clearly defined radiation, characterised by a uniformly curved
configuration of the frontal carinae, and anteriorly inclined and asymmetrical (anterior face
markedly shorter than posterior face) petiolar node. The frontal carinae are sinuate in all other
groups of Iridomyrmex, suggesting a basis for a primary higher-level classification of the genus.
Species of the calvus radiation have been revised by Shattuck (1993a), who referred to them as a
single species-group. However, he recognised three distinct complexes, which I have elevated
to species-group status because of their diversity and morphological and biogeographical
distinctiveness.
The calvus group (ss) comprises eight smallish species from moister habitats of southern
Australia, Norfolk Island and New Caledonia. They superficially resemble many other small and
robust species of Iridomyrmex from unrelated groups. All species have restricted distributions,
with the notable exception of I. notialis, which occurs throughout subhumid and semi-arid
southeastern and southwestern Australia, as well in the McDonnell Ranges of central Australia.
The viridigaster and rufoinclinus groups contain larger, reddish or red and black species, all
of which are uncommon. Species of the viridigaster group have a prominently rounded
promesonotum, and scale-like petiole, and are restricted to the southern arid zone. In the
rufoinclinus group the mesosoma is more gracile, and the petiolar node is relatively long, low
and broadly rounded dorsally. Iridomyrmex cappoinclinus occurs in the central arid zone, I.
cephaloinclinus throughout the northern arid zone, I. rufoinclinus throughout the monsoonal
tropics, and I. anteroinclinus is restricted to the northern Kimberley region of far northern
Western Australia (WA). There are at least two additional, undescribed species in the group,
including a uniformly blackish species from Cape York Peninsula.
agilis radiation
This is an extremely rich radiation of arid-adapted groups of long-legged, often extremely
gracile species. In all cases, the antennal scapes exceed the vertexal margin by at least one-fifth
their length, and usually much more. The agilis group comprises relatively large, extremely
gracile and exceptionally fast-moving species. Iridomyrmex agilis is a red species with
contrasting black legs and gaster that occurs throughout arid Australia. Iridomyrmex rubriceps is
an unusual member of the group in that it occurs in humid coastal eastern Australia, where it is
patchily distributed in open habitats. The bicknelli group seems to be closely related to the agilis
group, with both occurring primarily in the southern arid zone. Species of the bicknelli group are
smaller and uniformly blackish, often with feeble iridescence, and have a convex vertexal
margin. Iridomyrmex bicknelli occurs throughout southern Australia, most abundantly in the arid
zone, but also in open habitats of mesic regions. An undescribed species has a similarly wide
distribution across northern Australia. Species of both the agilis and bicknelli groups do not
appear to be as aggressive as other species of Iridomyrmex.
As the name suggests, species of the gracilis group are also extremely gracile. They can
have particularly long scapes, exceeding the vertexal margin by up to half their length. They are
hairier than species of the agilis and bicknelli groups, and are more aggressive. Iridomyrmex
fusciventris is a relatively small species with hairy scapes that occurs throughout the southern
semi-arid zone. A more robust species with glabrous scapes has a similar distribution, but
extends into the central arid zone. Several particularly gracile species are locally dominant ants
throughout the northern arid zone and semi-arid tropics. The group includes a small number of
larger, reddish species (I. mayri and close allies), one of which is extremely abundant and
widespread in the northern arid zone. The group (apparently) also includes a very distinctive
mid-sized blackish species with bicolored black and brown head, which occurs in association
with salt lakes throughout the southern arid zone. Iridomyrmex gracilis itself is a medium-sized
blackish species with hairy scapes that is atypically distributed in coastal and subcoastal eastern
Australia. There are several similar-looking species, but with shorter antennal scapes, that also
occur in subcoastal southeastern Australia; it is possible that these (including I. gracilis) are
actually related to species of the temperate vicinus group discussed below, rather than to the arid
and northern species described above.
The tropical anceps group seems to be very closely related to the gracilis group, and indeed
it is difficult to find clear characters that separate them. Compared to the gracilis group, species
of the anceps group tend to be less gracile and less hairy, without, for example, erect hairs on the
scape or hind femur. One species of the anceps group (possibly I. anceps itself) is one of the
most abundant ants throughout the semi-arid tropics and northern monsoonal zone. It (or
possibly a sibling species) also occurs extensively in South-East Asia.
I have included the purpureus group in this radiation, with the yet-to-be-substantiated
proposition that it is most closely allied to the gracilis group. The group is most distinctive, with
very broad heads, relatively short scapes, and densely hairy scapes and legs. It includes the
famous meat ants of the purpureus group in the more narrow sense, which have been
comprehensively revised by Shattuck (1993b) following extensive ecological and molecular
work by John Greenslade and Bruce Halliday (e.g. Greenslade & Halliday, 1982). However, I
also include in the group as a small complex of species described by Shattuck (1996) as
belonging to the discors group (Iridomyrmex discors and I. obscurior).
The final group that I have included in the agilis radiation is the pallidus group of pale,
large-eyed, nocturnal species, which is represented throughout arid Australia. It would not be
surprising if this group proved to be polyphyletic. Iridomyrmex pallidus (originally described as
a subspecies of I. rufoniger, which belongs to an unrelated group; see below) is extremely

abundant throughout tropical Australia. Like most other species in the group, its mesosoma has
numerous short, silvery hairs. However, the group includes several glabrous species. One of
these is I. hartmeyeri, a relatively large species with a prominently rounded propodeum, which
occurs throughout most of arid Australia.
rufoniger radiation
This radiation comprises relatively small and stout brownish or blackish species with short
antennal scapes (only slightly exceeding the vertexal margin). Iridomyrmex rufoniger and allies
represent the most distinctive group within this radiation. The head is cordate, with a
conspicuously concave vertexal margin, and the mesosoma features a steeply rising, prominently
rounded pronotum and short propodeum. The pronotum sports a distinctive tuft of long hairs.
The group includes a complex of small, uniformly blackish species with a flattened propodeum.
One such species with a mostly glabrous gaster occurs throughout most of arid Australia. There
are several similar species with hairy gasters, mostly in the northern arid zone and semi-arid
tropics. A closely related complex contains larger reddish species. One of these, possibly I.
rufoniger itself, occurs throughout the southern arid zone. A similar species, but with hairy hind
tibiae, occurs in central WA. A third complex consists of slightly larger species with a
prominently rounded propodeum. I have included I. septentrionalis,a uniformly brownish
species that is extremely common throughout subcoastal eastern Australia, in the rufoniger
group, but it may actually be related to I. gracilis and close allies described above.
Species of the mattiroloi group superficially resemble those of the rufoniger group, sharing
a stout mesosoma with steeply rising pronotum and short propodeum. However, they lack the
cordate head and tuft of long hairs on the pronotum, and I suspect the two groups are not so
closely related. Whereas species of the rufoniger group occur almost exclusively in the arid
zone, the mattiroloi group occurs in a wide range of habitats. In many species the mesosoma is
hairy throughout, and in one of these from the northern arid zone the scapes are also hairy. In
other species the mesosoma is glabrous except for several hairs on the pronotum. One such
species is extremely common throughout the semi-arid tropics and northern monsoonal zone,
and the same or a similar species occurs throughout eastern Indonesia. Numerous other species
are entirely glabrous. A complex of glabrous species has a particularly deep metanotal groove,
and one such species from the southern arid zone has striking pink and blue iridescence.
Iridomyrmex mattiroloi itself is a relatively hairy species from southeastern forests, and recent
molecular analysis (Rod Eastwood, personal communication) suggests that it might in fact be
quite unrelated to the arid species with similar morphology.
The suchieri group is closely related to the mattiroloi group (at least to the arid species), and
indeed the boundary between them is not so clear cut. Species of the suchieri group tend to be
larger, the pronotum is not so steeply rising and prominently rounded, and the propodeum is
longer. Some species are densely hairy. Two of these, one with hairy and the other with glabrous
scapes, occur from central northern Queensland to central western New South Wales. A more
sparsely hairy species is the dominant species of Iridomyrmex in a range of relatively moist and
shady habitats throughout the semi-arid tropics and northern monsoonal zone. An entirely
glabrous species is distributed throughout most of the arid zone.
Species of the vicinus group resemble those of the suchieri group, but have longer scapes
(exceeding the vertexal margin by about one-fifth of their length) and a prominently rounded
propodeum. The group has a Bassian distribution, occurring primarily in mesic and sub-humid
habitats of southern Australia. Iridomyrmex vicinus is restricted to mesic southern Victoria and
northern Tasmania, and a similar species occurs in high rainfall areas of far southwestern
Australia. Several other species extend into the southern semi-arid zone. One particularly
distinctive species (possibly I. emeryi) occurs in montane and alpine areas from southeastern
New South Wales to central Tasmania. It is a conspicuous ant of alpine grasslands, building
nests of fragmented plant material around the bases of tussock grasses, presumably to avoid
waterlogged soils. The conifer group comprises relatively large species from southwestern
Australia that have a distinctively conical propodeum (Shattuck, 1998). Despite their distinctive
appearance, I consider them to be closely related to species of the vicinus group (especially the
alpine species discussed above).
Species of the final group (referred to as Group A in Table 3) in the rufoniger radiation also
resemble those of the suchieri group, with which they are presumably allied, but have larger
eyes and blue iridescence. The group is referred to as the cynaeus group in Andersen (2000), but
I am now not so sure that I. cyaneus in fact belongs to this group. One species is common
throughout most of tropical Australia.
Melophorus
Species of Melophorus are highly thermophilic and polymorphic formicines, and therefore
parallel (but are not related to) species of Myrmecocystus in the New World and Cataglyphis in
the Old World. Their foraging activity is typically restricted to hot parts of the day, where they
can tolerate temperatures that are lethal to most other ants (Fig. 3). As for Iridomyrmex, the
overwhelming impression for such a speciose genus is one of morphological generalisation and
conservatism, with many species-groups appearing superficially similar to each other. However,
such morphological conservatism belies remarkable biological diversity. Although the vast
majority of species seem to be generalist predators and scavengers, the genus includes specialist
predators of ant brood and termites, specialist granivores, and at least one ‘honeypot’ with
specialized repletes (M. bagoti). A comprehensive key to provisional species-groups is provided
as Appendix 1. A number of distinct radiations can be recognised within the genus (Table 4), as
outlined below.
47
46
Body temperature (oC)
45
44
43
42
41
40
10 11 12 13 14 15 16
Time (hrs)
Fig. 3. Diel patterns of body temperature of foraging Melophorus bagoti, one of the most
thermally tolerant insects known. Above-ground activity commences only when soil surface
temperature exceeds 50oC, and continues with temperatures exceeding 70oC. Data from
Christian & Morton (1992).
Table 4. Classification, distribution and diversity of species-groups of Melophorus. Figures are

numbers of sorted species in the TERC collection. Distribution is described as predominantly
northern (N; occurring primarily within the northern arid zone), southern (S; occurring primarily
in the southern arid zone) or widespread (W; well-represented throughout arid Australia).
fulvihirtus radiation
anderseni group (6, N)
fulvihurtis group (2, S)
Group G (2, N)
Group I (3, W)
majeri radiation
majeri group (1, S)
potteri radiation
potteri group (12,S)
aeneovirens radiation
aeneovirens group (16, N)
froggatti group (42, W)
Group A (24, N)
Group B (10, N)
Group L (4, N)
wheeleri radiation
wheeleri group (43, W)
Group F (17, N)
Group H (2, N)
Group K (1, N)
mjobergi radiation
hirsutus group (2, mesic SE)
mjobergi group (91, W)
Group C (18, W)
Group D (14, W)
Group J (9, N)
fieldi radiation
fieldi group (110, W)
perthensis group (5, W)
pillipes group (16, N)
Unclear affinities
iridescens group (9, W)
bruneus group (37, S)
Group E (9, N)
fulvihurtis radiation.
These relatively few species lack tibial spurs, and characteristically have a thick and
sculptured integument. Species of the fulvihirtus (southern arid zone) and anderseni (northern
arid zone and seasonal tropics) groups raid brood from the nests of highly aggressive species of
Iridomyrmex, without apparent interference from their unwitting hosts (Clark, 1941; Agosti,
1997). Species of Groups G and I are presumably also specialist brood-raiders. Species of the
fulvihirtus group, anderseni group and Group I all have reduced maxillary palps, with the final
segment peculiarly pointed apically. The majeri group, represented by a single (and rare) known
species from southwestern WA. (Agosti, 1997) also lacks tibial spurs and its integument is
densely sculptured. However, it is otherwise so morphologically distinct that I have considered it
to represent a separate radiation (indeed, it is arguably distinctive enough to warrant generic
rank).
potteri radiation
The bulldozer ants of the potteri group from the southern arid zone are characterised by
stout bodies, short legs, and massive mandibles, all adaptations for hunting their termite prey
(McAreavey, 1947). Two major complexes can be recognized. In one, the anterior clypeus is
acutely angled and projects over the base of the mandibles, and the mandibles have five or six
distinct teeth. In the other, the anterior clypeus is not acutely angled, and the mandibles are
particularly large and blade-like, with just a single apical tooth (Fig. 15f in Greenslade, 1979).
aeneovirens radiation
This comprises long-legged species with an acutely angled clypeus that projects over the
base of the mandibles, and a dome-shaped occiput. The froggatti group, characterised by its
short and high propodeum, occurs throughout inland Australia and is particularly rich in species.
The closely related aeneovirens group and Group A consist of relatively large and gracile
species that occur primarily in the northern arid zone; they are strongly reminiscent of species of
Myrmecocystus and Cataglyphis. Groups B and L also occur primarily in the northern arid zone.
The latter appears closely allied to Group A, whereas the former is very distinctive, with a thick
and often conspicuously sculptured integument, short funicular segments, and is hairy
throughout. In one complex the hairs are particularly long and abundant, and the head of minor
workers is peculiarly hour glass-shaped (Fig. 15d in Greenslade, 1979).
wheeleri radiation
These species have characteristically short maxillary palps, and often a bulging clypeus. The
radiation features the seed harvesters of the extremely rich wheeleri group, the only formicines
known to be specialist granivores (Andersen, 1991). The major workers have very large heads,
strangely reminiscent of those of the myrmicine genus Messor. Several distinct complexes can
be recognised within the group. One comprises relatively large and gracile, reddish species in
which the two posterior clypeal setae are located slightly behind the anterior margin of the
antennal sockets, rather than slightly in front as is typical for Melophorus. Another comprises
relatively large and gracile, blackish species with conspicuously sculptured heads, and
mandibles with six or seven (rather than five) teeth.
Species of Group F look very similar to those from a complex of less gracile species within
the wheeleri group, but major workers do not have such massive heads, and the species are
apparently not granivorous. Group K is represented by a single known species from the Northern
Territory’s (NT) Victoria River District. It is large, black, and heavily sculptured, with
mandibles that are remarkably armed with very numerous (15 or more), long, sharp teeth. It
presumably has a specialist diet unrelated to granivory. Group H is represented by two known
species (one from the northern Top End of the NT, and the other from the nearby Victoria River
District) that appear to be associated with nests of species of the rothsteini group of
Monomorium, which are of similar size and color (reddish head and mesosoma, with contrasting
black gaster). Is this another group of specialist brood-raiders?
mjobergi radiation
These relatively stout-bodied species have a short and high propodeum, head that is slightly-
to-markedly compressed dorso-ventrally, and typically a short, scale-like petiole. Species of the
mjobergi group are the smallest (total length of minor workers about 1.5 mm) of the genus, and
resemble plagiolepidines. The group is exceptionally rich, with an estimated 200 species. It
includes several very distinctive complexes. One of these has very long and flattened heads, and
is characteristic of clay soils across the semi-arid north. Another, more widespread, complex is
very gracile compared with typical species, with antennal scapes exceeding the vertexal margin
by half or more their length. In the vast majority of species the mesosoma is entirely glabrous,
but a complex of hairy species with very large eyes occurs in the eastern arid zone. Representing
another complex, two species from southern Western Australia have a bulbous metanotal
process, strongly reminiscent of the enormis group of Notoncus.
Group C appears to be closely allied to the mjobergi group, but its species tend to be larger
and more conspicuously sculptured, and are covered with adpressed pubescence in addition to
erect hairs. The group contains two distinct complexes: smaller, more robust, darkly colored
species with hairy scapes; and larger, more gracile and usually yellowish species with glabrous
scapes.
In Group D, the mesosoma is very stout, with a particularly short and high propodeum. The
group appears to be restricted to higher rainfall areas, being absent from the central arid zone
and penetrating wetter habitats where no other Melophorus occurs. For example, one species is
restricted to sub-coastal Top End of the NT and the far northern Kimberley region of WA, where
it occurs in relatively shady habitats, and another species occurs in mesic habitats of
southeastern Australia. I suspect that Group D is allied to the highly distinctive hirsutus group,
whose dark reddish, heavily sculptured and densely hairy mesosoma is remarkably barrel-
shaped, with dorsally protruding metanotal spiracles. The group is represented by a small
number of species restricted to relatively mesic habitats of southeastern Australia. The final
group that I place in this radiation, Group J, consists of densely hairy species with a biconvex
mesosoma, and a relatively long and narrow, rather than scale-like, petiolar node. The group
occurs primarily in central Queensland (Qld) and semi-arid NT.
fieldi radiation
This comprises morphologically generalised taxa that are among the most common ants
throughout arid Australia. The radiation includes two very distinctive groups, with
complimentary distributions in semi-arid Australia. One of these, the Bottle-brush ants of the
pillipes group, is characterised by a dense covering of long, erect hairs on the tibiae. The group
occurs throughout semi-arid Australia, but is best represented in central Qld. The other, the
perthensis group, is characterised by a short, low and prominently rounded propodeum. It is
found primarily in the southwestern semi-arid zone, but one species occurs from the southern
Top End of the NT to the Gulf region of northwestern Qld. Both groups appear to be absent from
the central arid zone.
I have placed all other species in the radiation into the fieldi group (Plate 17, Andersen,
2000), which, as constituted, is a grab-bag of very morphologically generalised species that may
prove to represent several distinct species-groups. In the strict sense, species of the fieldi group
are largely glabrous (minors at most with a pair of pronotal hairs), gracile ants. In some, antennal
scapes are entirely glabrous, whereas in others the scapes have a single row of sparse erect hairs.
In one complex the glabrous scapes are particularly long, exceeding the vertexal margin by
about half their total length, and the species are often yellow. In the broader sense, the group
includes relatively hairy species, as well as the turneri complex of relatively stout species
(scapes exceeding vertexal margin by only about one-third their total length) that are very
similar in appearance to species of Group F within the wheeleri radiation.
Other species-groups
There are some additional species-groups whose relationships to other species-groups are
unclear. The most important of these is the bruneus group, whose dozens of species occur
primarily in the southern arid zone. One radiation (including M. bruneus itself) is very
distinctive, with short funicular segments and a relatively thick and sculptured integument. The
scapes and mesosoma are covered with short, erect hairs, and the head is conspicuously
bicolored (dark brown posteriorly, yellowish anteriorly). One species is very widely distributed,
occurring in all mainland States and the NT. In other species of the group, antennal scapes are
more sparsely hairy and funicular segments are not so short. Some of these species otherwise
look similar to those described above, but sometimes with longer and stouter setae. Others are
clothed with long silvery hairs, often also with dense silvery pubescence. In another radiation the
mesosoma is very sparsely hairy, and the ants closely resemble species of the fieldi group.
Another group with unclear affinities is the extremely gracile iridescens group, consisting of
two distinct complexes. The iridescens complex consists of medium-sized species that are
reminiscent of species from Group A within the aeneovirens radiation. Major workers are
orange-red with contrasting black gaster, but minor workers are often uniformly brownish. The
species appear to be restricted to the southern semi-arid zone, except for one known from far
northwestern Australia. One species with a peculiarly flattened head (Fig. 15a in Greenslade,
1979) occurs in mallee habitats throughout the southern semi-arid zone. The bagoti complex
consists of at least three very large species that are reminiscent of species from the aeneovirens
group. Melophorus bagoti is the largest species of the genus, and has specialised repletes. It is
uniformly orange, and occurs throughout the central and northern arid zones. A closely related
but beautifully shiny species with iridescent black gaster occurs in the Great Sandy Desert of
northern WA. Another species with orange foreparts and contrasting black gaster occurs in
central WA. The very strong similarities with species from the aeneovirens radiation would
suggest that the iridescens group belongs there; however the latter lacks the projecting clypeus
that is so characteristic of the aeneovirens radiation.
The final group with unclear affinities is Group E, comprising densely hairy species with a
thick and heavily sculptured integument. All species have orange or red foreparts with
contrasting black gaster, and occur primarily in the northern arid zone.
Although the species richness of Melophorus in arid Australia is remarkable, it is not unique
among the Australian biota. Arid Australia supports many hundreds of species of the plant genus
Acacia (Maslin & Hopper, 1982). Acacia is absent from many habitats in the region, and at most
a handful of species co-occur at any particular site. In comparison, Melophorus is ubiquitous,
more than ten species frequently co-occur, and spatial turnover is particularly high, much more
so than in other ant genera. For example, ant surveys of two nearby stations in the Victoria River
District of the NT (Fisher et al., 2004) yielded a total of 32 species of Melophorus, only four
(13%) of which were shared; this compares with 44% shared species among the remaining 90
ant species collected. These observations suggest that the total number of Melophorus species in
arid Australia might be truly staggering.
Camponotus
As previously mentioned, Camponotus is well-represented in all Australian biomes, but is
particularly rich in the arid zone. The arid fauna includes several radiations that occur primarily
or exclusively in arid regions (Table 5), as well as representatives of virtually all other
Australian species-groups within the genus. The major exceptions are the groups of arboreal,
phragmotic species that are restricted to wetter forests (Andersen, 2000; McArthur & Shattuck,
2001).

Camponotus that occur primarily or exclusively in arid or seasonally arid Australia. Figures are
numbers of species in the TERC collection. Distribution is described as predominantly northern
(N; occurring primarily within the northern arid zone), southern (S; occurring primarily in the
southern arid zone), monsoonal (M; occurring primarily in the monsoonal tropics) or widespread
(W; well-represented throughout arid Australia).
aurocinctus radiation
aurocinctus group (5, S)
ceriseipes group (8, S)
denticulatus group (24, W)
denticulatus complex (13, N)
terebrans complex (10, S)
postcornutus complex (1, S)
perjurus group (2, S)
setosus group (4, M)
subnitidus radiation
subnitidus group (19, W)
rufus complex (1, S)
subnitidus complex (7, N)
tricoloratus complex (11, S)
Group A (10, W)
Group B (2, S)
whitei radiation
intrepidus group (3 arid species, S)
intrepidus complex (mesic habitats only)
suffusus complex (3, S)
whitei group (9, S)
leae complex (7, S)
whitei complex (2, S)
Group C (3, S)
Group D (3, M)
discors radiation
discors group (40, W)
evae group (9, S)
minimus group (18, S)
sponsorum group (13, S)
Group E (10, S)
maculatus radiation
ephippium group (46, W)
capito complex (11, S)
ephippium complex (19, W)
tasmani complex (16, S)
nigriceps group (27, S)
consobrinus complex (14, S)

nigriceps complex (13, S)
nigroaeneus group (32, W)
pellax group (30, M)
bigenus complex (19, M)
pellax complex (11, M)
aurocinctus radiation
The most conspicuous species of Camponotus in arid Australia belong to a radiation of fast-
moving, diurnal, reddish and/or black species that have a characteristic cluster of long, J-shaped
hairs on the base of the mentum (leading to the description ‘Bearded sugar ants’; Andersen,
2002). This is referred to as the wiederkehri group by Shattuck & McArthur (2002), but I
consider it to include a number of distinct species-groups (see aurocinctus radiation in Table 5),
all of which occur exclusively (or nearly so) in the arid zone or seasonal tropics. The most
common of these is the denticulatus group, whose minor workers have an undifferentiated
propodeum such that the mesosomal profile is a simple curve. The group includes the
denticulatus complex of the central and northern arid zones, and the terebrans complex of the
southern arid zone.
The denticulatus complex is considered a single species by Shattuck & McArthur (2002),
but I recognise at least 13 species in the TERC collection. The complex occurs throughout
central and inland northern Australia, building distinctive nests that have conspicuous mounds
and often slit-shaped entrances. Camponotus denticulatus itself is a bicolored black and red
species occurring throughout central Australia. Most other species are uniformly reddish, and
occur in the northern arid zone or monsoonal tropics. In the field, minor workers strongly
resemble meat ants (highly aggressive species of the purpureus group of Iridomyrmex, as
outlined above). Interestingly, C. denticulatus is primarily black, as is the dominant meat ant (I.
viridiaeneus) where it occurs, whereas most species in northern Australia are uniformly reddish,
as is the locally dominant meat ant (I. sanguineus). This suggests that species of the complex
might be meat ant mimics.
Unlike McArthur et al. (1998), I similarly recognise a number of species in the terebrans
complex. One of these (possibly the species described by Santschi as C. latrunculus victoriensis)
is restricted to mesic habitats of southern Victoria and Tasmania. In contrast, an undescribed
pale species with large, bulbous eyes extends into the central arid zone. It is sympatric with C.
terebrans s.s. in semi-arid northwestern Victoria. The complex seems to be particularly speciose
in southwestern WA. In one such species, the vertexal corners of major workers form angular
projections as in C. postcornutus, which I consider belongs to a monospecific complex within
the denticulatus group (Table 5). I also include C. gouldianus, occurring throughout the southern
semi-arid zone, in the terebrans complex.
Other groups within the aurocinctus radiation have a saddle-shaped mesosoma. In the
aurocinctus group, the mesosoma has a conspicuous metanotal groove and the petiolar node is
very long and low. Four species have been described in this group: C. arenatus, C. aurocinctus,
C. owensae and C. versicolor (Shattuck & McArthur, 2002), but I consider C. ‘aurocinctus’ to
represent a complex of at least three species. In C. aurocinctus ss, the propodeum has a concave
anterior face that forms an obtuse angle with the dorsal surface, and the gaster has conspicuous
golden pubescence. The species is widely distributed in arid Australia, occurring in most
mainland States and in the NT. It is morphologically uniform from southern South Australia
(SA), through central Australia, to its northern limit in northern central WA. In what I consider
to be an undescribed sibling species from the southeastern arid zone, the propodeum is variably
rounded (never obtusely angled), the gaster totally lacks golden pubescence, and the petiole is
shorter and higher. Another apparent sibling species without gastric pubescence occurs in WA; it
has a very broad and unusually hairy head.
In the remaining three groups of the aurocinctus radiation, a metanotal groove is more
weakly developed (often absent), and the petiolar node is relatively short, high and rounded
dorsally. The ceriseipes group consists of about ten species from the southern arid zone, four of
which are described (C. cerseipes, C. donnellani, C. prosseri and C. rufonigrus; Shattuck &
McArthur, 2002). The setosus group comprises at least four species from the northwestern
monsoonal zone, only one of which is described (as a member of the aurocinctus complex;
Shattuck & McArthur, 2002). Most species of the setosus group are covered with whitish
pubescence (more golden on the gaster) and are densely clothed with long hairs. Camponotus
setosus is a red and black species distributed from the Kimberley to the southern Top End of the
NT, occurring on sandy soils associated with sandstone outcrops. The three other species are
uniformly black; one of these from the Top End is unusual in that it is very sparsely hairy.
Finally, the perjurus group consists of two known species in which the head is peculiarly
attached well below the occipital margin (Shattuck & McArthur, 2002). In C. perjurus the
foreparts are red and only weakly iridescent. An undescribed species from SA is uniformly
blackish, with strong blue-green iridescence throughout.
subnitidus radiation
A second arid-adapted radiation of Camponotus comprises species of the subnitidus and
related groups, which have extremely long and glabrous antennal scapes (hence the name Aerial
sugar ants; Andersen, 2002). The clypeus of major workers projects as a simple, rectangular
lobe, distinguishing these ants from superficially similar species belonging to the maculatus
radiation (see below). The clypeal region of major workers is often heavily pitted and somewhat
flattened, suggestive of partial phragmosis. The ants are primarily nocturnal, so are somewhat
inconspicuous despite including some of the largest species of Australian Camponotus.
The subnitidus group comprises very large species in which the sides of the heads of minor
workers are markedly divergent anteriorly. Two major complexes can be recognised: the
subnitidus complex of northern Australia, in which the petiolar node is very long and low, with a
nipple-like projection dorsally, and the tricoloratus complex of the southern arid zone, where the
node is short and erect. A second group (Group A in Table 5) comprises smaller, often pale
species that look superficially like members of the claripes group from the maculatus radiation.
They occur throughout inland Australia, but are not commonly collected. The final group (Group
B in Table 5) consists of a small number of dark reddish brown species from the southern semi-
arid zone, in which the mesonotum of minor workers has a posterior hump.
whitei radiation
A third major arid-adapted radiation of Australian Camponotus, comprising the whitei and
related groups, consists of mostly dark-colored, diurnal species with thick and often coarsely
sculptured integuments. The whitei group consists of medium-sized species that occur primarily
in southern semi-arid regions. Two complexes can be recognised, one in which the mesosoma is
biconvex (whitei complex) and the other in which the mesosoma is saddle-shaped (leae
complex). Camponotus whitei is by far the most common and widespread species of the group,
occurring from central northern Qld, through western NSW to northwestern Victoria, and across
to southern WA. All other species of the group have been rarely collected. A related group
(Group C in Table 5) consists of small, robust and brownish species in which the mesosoma is
straight and uninterrupted in profile, with the propodeum often projecting backwards over the
petiole. The group appears to be endemic to central and southern WA. In another, possibly
closely-related group from northwestern Australia (Group D in Table 5) the mesosoma is
likewise straight and uninterrupted in profile, but the species are larger and more gracile. All
species of Groups C and D appear to be rare. Finally, the intrepidus group seems to belong to the
whitei radiation. Most species of this group occur in mesic habitats, but at least two species of
the suffusus complex occur in semi-arid southeastern Australia.
discors radiation
I have perhaps artificially clustered together a range of morphologically generalised species-
groups from arid Australia as the discors radiation, a fourth major radiation of arid-adapted
Australian Camponotus. Some of these species-groups closely resemble species-groups from the
maculatus radiation (see below), and very possibly belong there. However, in all cases major
workers lack the conspicuously notched or otherwise indented anterior clypeal margin that is so
characteristic of Australian groups within the maculatus radiation (except for the
novaehollandiae group). It is very possible that this clypeal character has been secondarily lost
in some or all of these species-groups, rather than them belonging to a separate radiation.
In the discors, minimus and evae groups the mesosoma is disc-shaped and laterally
compressed posteriorly. I suspect that such morphology is convergent, given that it occurs in
unrelated species-groups both in Australia and overseas. The discors group is particularly rich in
species, and occurs throughout inland Australia. Minor workers of all species are yellowish,
either uniformly so or with infuscated patches on the mesosoma and head. Species of the
apparently closely related minimus group, which are restricted to the southern arid zone, are
among the smallest of all terrestrial Camponotus, with minor workers only 3-4 mm in total
length. They are either blackish with pale legs (as in C. minimus) or uniformly yellowish (e.g. C.
scratius), and strongly resemble miniature species from the claripes group (C. minimus was in
fact described as a subspecies of C. claripes). In the evae group the mesosoma is especially
strongly compressed (the propodeal dorsum is ridge-like), and the pronotum is markedly
flattened dorsally. All species are black, often with pale legs, and the integument is rather
conspicuously sculptured and dull. Two complexes can be recognised, one in which the anterior
clypeal margin is a simple convexity, and the other in which the margin is concave medially. It
is possible that the Australian honeypot ant Camponotus inflatus also belongs here as a third
complex, rather than where it has been traditionally placed as a member of the nigroaeneus
group of the maculatus radiation.
I have also included the sponsorum group and apparently closely related Group E in the
discors radiation. Both groups comprise relatively small, diurnal species from the southern arid
zone. The petiole is relatively long and broadly rounded, and the gaster is covered with silvery
pubescence. Species of the sponsorum group have a saddle-shaped mesosoma where the
propodeum is markedly concave in profile. Most species are bicolored yellowish and black. One
such species (possibly C. sponsorum itself) is extremely widely distributed, occurring
throughout the southern arid zone and extending into central Qld. Species from Group E have
longer and narrower heads and longer scapes (exceeding vertexal margin by at least half their
total length), and the mesosoma has a simple profile (propodeum at most feeble concave in
profile). They have the appearance of small species of the nigroaeneus group, to which they are
in fact possibly related.
maculatus radiation
In addition to the four major arid-adapted radiations outlined above, several arid-adapted
species-groups occur within the cosmopolitan maculatus radiation, which is ubiquitous in
Australia (Table 5). As previously mentioned, this radiation comprises a range of species-groups
in which the anterior clypeal margin of major workers is conspicuously notched or otherwise
indented medially (see Greenslade, 1979). The nigriceps group consists of large, yellowish (head
and often also gaster contrasting black) nocturnal species from southern Australia, in which the
anterior clypeal margin of all workers projects as a pair of moustache-shaped angular lobes
(McArthur and Adams, 1996). Two complexes can be recognised, one (nigriceps complex) in
which the gula has numerous erect hairs, and the other (consobrinus complex) where the gula is
glabrous. Most species of the consobrinus complex have a bicolored gaster, with the first
segment yellowish anteriorly; exceptions include C. loweryi from eastern semi-arid Australia
(gaster uniformly yellowish) and an undescribed species with a uniformly black gaster from
northern NSW and southeastern Qld. Camponotus consobrinus itself is restricted to mesic
habitats of southeastern Australia (including Tasmania), but most other species occur in semi-
arid regions. The nigriceps complex extends deeper into the central arid zone.
The remaining arid-adapted groups within the maculatus radiation consist of fast-moving,
diurnal, reddish and/or black species, all with very many species. The most common of these are
species of the morphologically generalised nigroaeneus group. Camponotus nigroaeneus itself is
restricted to mesic habitats of southeastern Australia, and the closely related C. aeneopilosus is
distributed along the eastern seaboard from northern Victoria to North Queensland, but
otherwise the group occurs exclusively in the arid zone. In most species the gaster is covered in
silvery or golden pubescence, but there are many exceptions. One of these is C. amperi, which
occurs throughout the southern semi-arid zone. A similar, undescribed species is widespread on
cracking clay soils of the semi-arid tropics. A least two species of the nigroaeneus group are
strongly iridescent throughout.
The final two arid-adapted groups that I place within the maculatus radiation are the closely
related ephippium and pellax groups. In both cases the major workers are partly phragmotic, and
their occipital corners extend backwards as rounded lobes. Species of the ephippium group have
a characteristically concave mesosomal profile. Three distinct complexes can be recognised
(Table 5). In the ephippium complex the vertexal corners are distinctly angled laterally.
Camponotus ephippium occurs throughout the eastern arid zone, up to central Qld where it co-
occurs with the closely-related C. dromas, which is distributed throughout the semi-arid tropics.
Several uniformly black species with golden gasters occur in North Queensland, and another
such species occurs in the southern Kimberley. Species of the tasmani complex have rounded
rather than angled heads laterally, and occur primarily in the southern arid zone. In the capito
complex, the mesosoma is highly polished, very markedly concave in profile, and only sparsely
hairy. Most species are reddish with contrasting black gaster, and also occur in the southern arid
zone. Lastly, the pellax group consists of numerous relatively small species from the monsoonal
tropics. Species of the pellax complex have red foreparts and black gaster, and occur primarily in
the western monsoonal zone. At least two species have a strongly iridescent gaster. Species of
the bigenus complex are uniformly black, and occur primarily in the eastern monsoonal zone.
Several non-arid species groups of the maculatus radiation also occur in arid Australia,
including the tropical novaehollandiae group (Andersen, 2000) and cosmopolitan claripes and
rubiginosus groups. However, these are not considered here.
Monomorium
The exceptional diversity of the Monomorium fauna from arid Australia is not at all
reflected in the recent revision of Australian Monomorium by Heterick (2001), who recognised a
total of only 59 species throughout Australia. Many of Heterick’s ‘species’ were described as
being extremely widespread and highly variable, and most ant taxonomists would consider them
each to represent several to many valid species. The higher-level classification of species that I
propose here (Table 6) also differs significantly from that presented by Heterick (2001).
nigrium radiation
This is an extremely rich radiation of very small, mostly generalist species with 11-
segmented antennae, and smooth and shiny mandibles having three or four teeth. All other
Australian species groups of Monomorium have 12-segmented antennae, and were previously
referred to the now synonymised genus Chelaner. Species of the nigrium radiation are
ubiquitous in Australian ant communities outside the coolest and wettest habitats, but are
particularly diverse and abundant in the arid zone and seasonal tropics. The nigrium group
(considered by Heterick (2001) to represent a single variable species, M. fieldi) consists of very
many blackish species that are predominantly smooth and shiny (Plate 9, Andersen, 2000), with
most occurring in the tropical north. They are extremely challenging taxonomically, but several
complexes can be recognised. The nigrium complex comprises tiny species with short antennal
scapes, glabrous hind femora, and often a short and obliquely angled propodeum. They occur
throughout higher rainfall areas of northern Australia. The fieldi complex contains larger species
with longer scapes (approximately reaching the vertexal margin), and have at least a few erect
hairs on the hind femora. A third complex (with no described species) comprises a small number
of species from the Top End of the NT with broad heads, short scapes, and small petiolar nodes.
One species is weakly polymorphic, and two others have conspicuous sculpturing on the
propodeum.
Table 6. Classification, distribution and diversity of major species-groups of Monomorium that

occur primarily or exclusively in arid or seasonally arid Australia. Figures are numbers of
species in the TERC collection. Distribution is described as predominantly northern (N;
occurring primarily within the northern arid zone), southern (S; occurring primarily in the
southern arid zone), monsoonal (M; occurring primarily in the monsoonal tropics) or widespread
(W; well-represented throughout arid Australia).
nigrium radiation
carinatum group (23, N/M)
carinatum complex (15, M)
complex A (8, N)
eremophilum group (30, S)
laeve group (64, W)
nigrium group (24, M)
sydneyense group (7, W)
Group A (7, E)
rothsteini radiation
rothsteini group (47, W)
sordidum group (25, W)
antarcticum radiation
centrale group (4, W)
flavipes group (8, S)
flavipes complex (6, S)
euryodon complex (2, S)
insolescens group (30, M)
lacunosum group (5, S)
longiceps group (8, S)
legulus complex (3, S)
longiceps complex (5, S)
longinode radiation
capito group (5, N)
longinode group (2, S)
whitei radiation
falcatum group (4, S)
whitei group (18, S)
The carinatum group also has a tropical distribution, but occurs primarily in drier habitats.
The mesosoma and petiole are conspicuously sculptured, and at most sparsely hairy. The group
contains two complexes (Andersen, 2000). In species of the carinatum complex the
promesonotal dorsum is markedly flattened and broadly trapezoidal, often with distinct lateral
margins. There are at least 15 species, distributed throughout the northern arid zone and semi-
arid tropics. Species from the second complex (Fig. 16, Andersen, 2000; Complex A in Table 6)
are smaller and the mesosoma is rounded dorsally and always glabrous. Heterick (2001)
considered them all to be variants of M. sydneyense, which I believe belongs to a separate group.
Monomorium sydneyense has a finely reticulate propodeum but is otherwise smooth and shiny,
and is restricted to humid and sub-humid habitats of southeastern Australia (except for a recent
introduction into New Zealand). It is glabrous, and in southern Victoria co-occurs with a very
similar but hairy species that extends into Tasmania. Superficially similar species occur
throughout Australia outside the central arid zone. I have provisionally classed them together as
the sydneyense group, but they are very possibly polyphyletic. It is possible that M. sydneyense
and allies are closely related to the eremophilum group, which comprises very many granivorous
species from the southern arid zone. In all cases the median clypeal carinae project as a pair of
sharp teeth. Many species, including M. nanum, have very large, reniform or tear-shaped eyes.
Another group, with apparently no described species (Group A in Table 6), also has large,
reniform eyes. However, the anterior clypeal margin of these species is not laterally denticulate,
suggesting they are not granivorous. The final group in the nigrium radiation is the laeve group,
whose numerous yellowish, mostly nocturnal species are ubiquitous in arid Australia.
rothsteini radiation
This is an exclusively arid-adapted radiation that seems to be allied to the nigrium radiation.
The species have a saddle-shaped mesosoma (incorporating a deep metanotal groove), large and
erect petiolar node, smooth and shiny head and promesonotum, and mandibles with three teeth.
The radiation comprises two very distinct groups. Species of the sordidum group are highly
generalised ants that strongly resemble species of the nigrium group. In most species the
propodeum is densely sculptured, but in some from the southern arid zone it is almost entirely
smooth and shiny. Monomorium megalops is a species from the complex with very large eyes,
and occurs throughout the eastern arid zone. At least one other species has similarly large eyes.
The rothsteini group comprises granivorous species that are the most abundant harvester
ants in arid Australia (Andersen, 2000). The group is very conservative morphologically, but
there is undoubtedly very many species – apparently at least 50 and possibly many more. Several
complexes can be recognised. One of these has particularly long striae on the sides of the head
anteriorly (the striae extend to the lower margin of the eye), and includes three very common
and widespread species. One is a particularly large and robust species that occurs throughout
most of far northern Australia (it is the species studied in Andersen et al., 1991, and
Monomorium sp. 1 in Andersen et al., 2000). Another has a very deeply incised anterior clypeal
margin, and occurs throughout the semi-arid tropics. The third has a relatively short, narrow and
dorsally tapered petiolar node, and is widespread in central Australia. In a second complex, the
mesonotum is conspicuously sculptured dorsally. One such species is common throughout arid
NT, and others occur in northern WA.
Most species of the rothsteini group have reddish foreparts with contrasting black gaster.
However, in many species the foreparts are conspicuously infuscated, and some are uniformly
blackish. Several such uniformly blackish species occur in northern WA. In several otherwise
reddish species, peculiar black individuals can be observed foraging with normal workers. It is
not clear if these represent an apterous reproductive caste from the host colony, or are parasitic
species.
antarcticum radiation
This is a Gondwanan radiation of variously granivorous species that includes groups
restricted to wet forests (such as the leae group of southern Australia and the closely related
antarcticum group of New Zealand), as well as five major groups centred on the Australian arid
zone and monsoonal tropics. In all these arid groups the mesosomal profile is a simple curve,
with the propodeum at most feebly differentiated.
The flavipes group is closely related to the leae group; in both cases the mandibles have five
teeth, the clypeus projects over the mandibles as a broadly rounded or blunt lobe, and the species
are seed-eating omnivores. However, species of the flavipes group lack the differentiated
propodeum of the leae group, and occur in drier habitats. Two complexes can be recognised, the
flavipes complex comprising monomorphic species (as is typical for Monomorium) and the
euryodon complex containing polymorphic species. Several species of the flavipes complex are
known from subhumid habitats of southeastern Australia, another occurs throughout the
southern semi-arid zone, and another is restricted to semi-arid southwestern WA. The euryodon
complex includes at least two species, one occurring from semi-arid central Qld to western
NSW, and another in semi-arid southwestern WA.
The centrale group appears to be closely related to the flavipes group, from which it can be
distinguished by its anterior clypeal margin that is conspicuously concave medially, and longer
and more cuboid petiolar node. The basal tooth of the mandibles is reduced to a small denticle or
is absent altogether. As in the flavipes group, all species are yellowish or reddish brown. One is
widely distributed in the central arid zone, from SA to northwestern Qld, with most other species
occurring in southern semi-arid regions. However, at least one species is known from North
Queensland and another from the Kimberley region of far northern WA
The lacunosum group consists of a small number of uncommon, heavily sculptured species
that otherwise resemble species of the centrale group. Two complexes can be recognised. In one,
the propodeum is armed with teeth or short, broad spines, and the metapleural lobes are likewise
produced into broad spines. This complex includes M. lacunosum from semi-arid southwestern
WA, and an undescribed species from semi-arid northwestern Victoria. In the second complex,
the propodeum is unarmed and the metapleural lobes are more flange-like than spinose. The
complex comprises a small number of species known only from central Qld and central NSW.
The longiceps group comprises up to a dozen reddish brown, conspicuously sculptured
species occurring in eucalypt (particularly mallee) woodlands throughout the southern semi-arid
zone. They are major predators of eucalypt seeds (e.g. the species referred to as Chelaner sp. 3
in Andersen, 1982). The group contains two distinctive complexes. In the longiceps complex the
propodeum is armed with teeth, and the petiolar node is relatively short and erect. The species
resemble those of the lacunosum group, and presumably the two groups are closely related. The
legulus complex contains larger species with an unarmed propodeum, a long, broadly rounded
petiolar node, and a clypeus armed with a large pair of acutely angled projections.
The insolescens group appears to parallel the longiceps group in northern Australia, and is
the only arid-adapted group within the antarcticum radiation that has a northern distribution. The
group occurs throughout the monsoonal tropics, but primarily in the western monsoonal region,
and is particularly diverse in the Kimberley. The group as a whole is remarkable for both its
diversity and the limited distributions of its species. Of the 30 species in the TERC collection, 19
are known only from relatively small areas in the Kimberley, and most of the others appear to be
restricted to similarly small areas of the northern half of the NT. Three of the latter commonly
co-occur in subcoastal habitats of the Top End. Only one species within the group could be
considered widely distributed – a heavily sculptured species with very large eyes occurs across
semi-arid NT into northwestern Qld. This species, as well as several others that are also heavily
sculptured, bears a strong resemblance to species of the longiceps group. However, other species
of the insolescens group are markedly divergent. The most extreme belong to a complex of
large, smooth and shiny species, with long petiolar peduncles and high nodes. These species
have angular vertexal corners with a feebly to markedly concave vertexal margin, very
prominent rounded or acutely angled clypeal projections, and relatively long funicular segments
(see Fig. 18 in Andersen, 2000).
longinode radiation
This is a small radiation of granivorous species that is presumably allied to the antarcticum
radiation, and possibly belongs there. The species are heavily sculptured and typically have a
long and low petiolar node. Some species of the insolescens group also share these features, so
there is possibly a close relationship, particularly with the capito group. The capito group
comprises a small number of species from the northern arid zone, characterised by a very large,
longitudinally striate head, massive clypeal teeth, and reduced numbers (only 2 or 3) of
mandibular teeth. The most widespread of these is M. bifidum, which occurs in both the Top End
and the Kimberley. There are at least three other sibling species, one in the central arid zone,
another in the Victoria River District, and another in the East Kimberley. Monomorium capito
has a particularly long and low petiolar node, and is known from only a single specimen
collected from remote central Western Australia (Heterick, 2001). The only other known
member of the capito group is a remarkable species in which the head has an exceptionally low
attachment to the mesosoma, the propodeum is armed with a pair of teeth, and the petiole is
feebly sculptured and relatively short and high. It appears to be restricted to the Katherine region
of the southern Top End.
The longinode group consists of two or three species from southern WA. These species have
a similar mesosoma and petiole to M. capito, but have smaller, more coarsely and irregularly
sculptured heads, and the anterior clypeal margin has short, bluntly rounded lobes rather than
prominent teeth.
whitei radiation
This radiation comprises specialist granivores of the southern arid zone, and its relationship
to other radiations of Australian Monomorium is unclear. The whitei group consists of large-
headed species with a prominent promesonotum that is markedly higher than the propodeum in
profile. Monomorium whitei is a very large, polymorphic species that occurs throughout the
southern arid zone, from southwestern Qld to southern WA. It has a reddish brown head and
mesosoma, with contrasting yellowish brown gaster. There are several other very similar taxa in
the region that vary in coloration and sculpturing, indicating a complex of unresolved sibling
species. Monomorium bicorne is a closely related mid-size, uniformly black species occurring
throughout southeastern semi-arid Australia. A sibling species with a longitudinally striate head
and contrasting yellow-brown gaster occurs in southern WA. Most of the diversity within the
whitei group resides in several complexes of small, monomorphic species that have radiated
primarily in southern WA. One of these complexes is characterised by very long and elongate
eyes, and includes M. anthracinum.
The falcatum group consists of a handful of small, stout species with foveolate heads and
the presence of propodeal denticles or spines. Two distinct complexes can be recognised.
Species of the falcatum complex (including M. falcatum and M. elegantulum) have 12-
segmented antennae and prominent propodeal spines, and are represented throughout the
southern arid zone. Species of the decuria complex have 10-segmented antennae and the
propodeum is armed with small denticles only; they are restricted to southwestern WA.
Rhytidoponera
Rhytidoponera is widespread in the Australian region, and includes several species-groups
occurring primarily or exclusively in wet forests. However, it is particularly diverse in arid
Australia, where it is virtually ubiquitous. Species of Rhytidoponera are mid-sized to very large
ants with highly generalised habits, and are most abundant at relatively unproductive sites for
ants where behavioural dominance (especially by Iridomyrmex) is relatively low. In the arid
zone they are particularly important components of local ant communities on poorly drained
soils or at rocky sites (Andersen, 2003). Most species in arid Australia belong to exclusively
arid-adapted species-groups (and indeed radiations; Table 7), and there is very little penetration
into arid Australia from forest species-groups.

Rhytidoponera that occur primarily or exclusively in arid or seasonally arid Australia. Figures
are numbers of species in the TERC collection. Distribution is described as predominantly
northern (N; occurring primarily within the northern arid zone), southern (S; occurring primarily
in the southern arid zone), monsoonal (M; occurring primarily in the monsoonal tropics) or
widespread (W; well-represented throughout arid Australia).
tyloxys radiation
tenuis group (16, M)
tyloxys group (12, N)
dubia complex (5, N)
tyloxys complex (7, N)
metallica radiation
anceps group (1, S)
metallica group (37, S)
metallica complex (33, S)
borealis complex (4, M)
flavipes group (2, S)
mayri radiation
aurata group (12, M)
convexa group (62, W)
foveolata group (13, W)
foveolata complex (9, S)
micans complex (4, N)
mayri group (18, W)
taurus group (21, M)
reticulata radiation
reticulata group (20, M)
Group A (6, N)
tyloxys radiation
This radiation consists of two species-groups with highly reduced (often absent) hind tibial
spurs. The tyloxys group is arguably the most distinctive of all groups within Rhytidoponera, and
seems to be an outlier within the genus both morphologically and biologically. The long, narrow
head, very large eyes and massive mandibles suggest a high degree of specialisation, but details
of their biology are not known. Two clearly defined complexes can be recognised. In the tyloxys
complex the petiolar node is armed with a dorsal spine. In R. tyloxys the spine is reduced to a
short, angular projection; this species occurs throughout the northern arid zone and semi-arid
tropics. A species with a more prominent spine is extremely widespread (but uncommon)
throughout the central arid zone. A very similar species occurs in the Kimberley region, but
differs in having an elaborate rather than simple anterior clypeal margin. In two other species
from the Kimberley the petiolar spine is extremely long and thin.
In the dubia complex the petiole is broadly rounded dorsally (Fig. 9b, Andersen, 2000).
Rhytidoponera dubia is restricted to the Top End and northern Victoria River District of the NT,
where it is usually found foraging on low vegetation. A similar but larger species occurs
throughout the northern half of the NT and into the Gulf region of northwestern Qld. A larger
species again occurs throughout the central arid zone, and is the only species of the tyloxys group
known from SA.
The tenuis group seems much closer to mainstream Rhytidoponera both morphologically
and biologically. It consists of a large number of relatively small and gracile species from the
monsoonal zone (Fig. 9c, Andersen, 2000), apparently only one of which is described. Most
have rather limited distributions. A species with a densely punctate first gastric tergite is
common throughout the Top End of the NT, and a similar species occurs in central Qld. In
Rhytidoponera tenuis the first gastric tergite has feeble striations over dense background
punctation; it is restricted to the broader Townsville/Cairns region of North Queensland. In
several species the first gastric tergite is coarsely striate but otherwise shiny anteriorly, and
densely punctate posteriorly. In several other species the first gastric tergite is striate throughout.
metallica radiation
This is a radiation of smaller species occurring primarily in southern Australia. Most of the
species belong to the extremely rich but morphologically conservative metallica group.
Rhytidoponera metallica as I understand the species is a metallic green, highly opportunistic
species occurring in a wide range of open habitats throughout southeastern Australia. It is an
extremely common urban ant throughout the region, and is familiar to many people because of
its relatively painful sting. Very many similar species occur in the southern arid zone. In some
the mesosoma has bulging pronotal shoulders and an extended ‘neck’. A relatively smooth and
shiny species with sparse hairs occurs throughout the western semi-arid zone. Most species have
prominent, backward projecting occipital lobes, but in others these lobes are reduced or absent.
A species with coppery pink iridescence is widespread in the central arid zone, and other species
are brilliantly blue. A smaller number of species, including R. tasmaniensis from mesic habitats
of the southeast, and R. inornata from the southwest, lack iridescence altogether.
All the above species appear to be seed-eating omnivores. A separate complex of non-
granivorous, usually non-iridescent species occurs in the northern monsoonal tropics. The most
common of these are R. borealis (Fig. 9a, Andersen, 2000), R. brunnea and R. trachypyx, which
co-occurr along with an uncommon undescribed species in the the Top End of the NT.
Rhytidoponera brunnea was synonymised with R. borealis by Brown (1958), but can be
distinguished by its short and high petiolar node, and finer sculpturing on the occipital lobes.
Both species also occur in the northern Kimberley, along with another undescribed species with
small eyes and reduced occipital lobes. At least two undescribed species with bluish iridescence
and an unusually projecting clypeus occur in Cape York Peninsula.
The flavipes group consists of a small number of species from the southwestern arid zone.
They are more gracile, less coarsely sculptured ants with relatively narrow heads and very large
eyes, giving them the general appearance of species of the tyloxys group.
The final group within the metallica complex considered here is the anceps group.
Rhytidoponera anceps is a relatively small species with a cuboid petiolar node and large eyes. It
has a remarkably disjunct distribution, with distant populations in southern WA and central Qld.
Another complex (victoriae complex) in the anceps group is extremely common in mesic
habitats throughout the eastern seaboard, and is not considered here. There is a small number of
other groups within the metallica radiation that occur exclusively in eastern forests (e.g.
impressa group, Ward, 1980), and these are likewise not considered here.
mayri radiation
This is the major arid-adapted radiation within the genus, and includes all the larger species
of arid Australia. The most common representatives are species of the extremely rich but
morphologically uniform convexa group, which occurs throughout most of inland Australia. The
species are mid-sized for Rhytidoponera, and have a characteristically short, high and dorsally
flattened node, and a relatively simple occipital vertex. Within the group, there are one or more
complexes of somewhat larger, less coarsely sculptured species that have rectangular heads with
a distinctly concave vertexal margin. The head is variously punctate and foveolate, but never
coarsely rugose. One of these is R. maniae, a very common ant throughout semi-arid
southeastern Australia. In more typical species, the head is coarsely reticulate-rugose, and not so
broad behind the eyes (Fig. 9g, Andersen, 2000). One complex of species (R. violacea and allies)
from central and southern WA is conspicuously iridescent. Rhytidoponera rufithorax is a
distinctive reddish brown species with contrasting dark brown gaster that occurs throughout the
semi-arid tropics. It co-occurs throughout its range with an extremely similar but uniformly
blackish (gaster sometimes paler brown) species. Rhytidoponera convexa is a relatively hairy
species from central and North Queensland. Rhytidoponera pilosula is a particularly hairy,
uniformly blackish species from southwestern Qld and northwestern NSW. A very similar
species is highly abundant in Mitchell grasslands of northwestern Qld and semi-arid NT.
The foveolata group appears to be closely allied to the convexa group, especially to the first
complexes described above. They are larger, more robust species, with broadly rectangular
heads, usually with a concave vertexal margin. Species of the micans complex are finely but
densely punctate throughout, with scattered, shallow foveolae, and occur in the central and
northern arid zones. Species of the foveolata complex are more coarsely sculptured, and occur in
the southern arid zone.
The mayri group consists of numerous, very large species with prominently crested heads.
They typically construct ‘volcano’ nests, often at the bases of shrubs. Rhytidoponera mayri is
widespread in the southeastern arid zone, but most other species have more limited distributions.
In the monsoonal tropics the group is largely replaced by species of the closely related aurata
and taurus groups (Andersen, 2000). In the latter the vertexal lobes are produced as prominent
horn-like projections (Fig. 9h, Andersen, 2000). Rhytidoponera taurus is widely distributed in
the central and northern arid zones, but virtually all other species are restricted to the monsoonal
tropics. The group is particularly rich in northern WA, where apparently at least a dozen species
occur. In the aurata group (Plate 5, Andersen, 2000) the horns are reduced to acute vertexal
angles. With two exceptions, all species appear to be restricted to either the Top End or
Kimberley. One of the exceptions is a uniformly reddish brown species that occurs throughout
the monsoonal zone, and the other is a uniformly blackish brown species occurring in the Top
End, Victoria River District and Kimberley.
reticulata radiation
This is a tropical radiation containing two arid-adapted species-groups, each with a cuboid
petiolar node and long, spinose sub-petiolar process. In the reticulata group the head is at most
slightly crested and the vertexal corners are not prominently lobed. Rhytidoponera reticulata
(Fig. 9e in Andersen, 2000) is a slightly crested species that is very common throughout the Top
End. A very similar but more feebly crested, more densely hairy and less coarsely sculptured
species occurs in semi-arid NT and northwestern Qld. Most other species occur in northern WA.
The other species-group within this radiation (Group A in Table 7) comprises species with
the general appearance of those of the reticulata group, but with conspicuous vertexal crests or
lobes. There are six known species, none of which are described. All occur in the semi-arid
tropics, mostly in the NT.
CONCLUSION
This chapter has presented a systematic overview of the five major genera contributing to the
remarkable ant diversity in arid Australia. For some genera (Melophorus and Iridomyrmex) this
represents their first comprehensive treatment at the species-group level. For others
(Monomorium, Camponotus) the analyses significantly challenge recent taxonomic work,
particularly at the species level. All the genera contain very difficult species complexes whose
final resolution remains a long way off. Many species-groups and indeed entire genera are
highly conservative morphologically, often making it difficult to determine relationships
between groups, and sometimes to assign species to species-groups. The application of
molecular techniques would be highly informative here, but unfortunately has had very limited
use in Australian ant taxonomy. The use of molecular techniques would also be important for
investigating relationships between Australian and other species-groups in genera such as
Monomorium and Rhytidoponera.
The five major genera considered here exhibit highly divergent biogeographical profiles
(Fig. 4). The most extreme is shown by Iridomyrmex, where the overwhelming majority (85%)
of species belong to groups that are widely distributed throughout arid Australia (Fig. 4a). This
is perhaps explained by the genus being an ancient autocthonous element of the Australian ant
fauna that has radiated within Australia over a very long period of geological time. In contrast,
Monomorium has a far more even biogeographical profile, with substantial proportions of
species from both southern and northern species-groups (Fig. 4d). Monomorium has a complex
biogeographical history within Australia, apparently including relatively recent incursions of
tropical elements from South-East Asia (nigrium radiation), as well as Gondwanan stock (e.g.
antarcticum radiation).
Melophorus and Rhytidoponera both belong to entirely Gondwanan taxa (Melophorini and
Ectatomminae respectively), but both contain a substantial proportion of species from northern
groups within Australia (Figs 4b, e). For Rhytidoponera, this can be explained by the
predominantly tropical distribution of ectatommines generally. However, all other melophorines
occur primarily in cool-temperate climates, so in Melophorus the prominence of northern over
southern Australian elements is surprising.
The biogeographical history of Camponotus in Australia, where the arid zone supports a
high proportion of species from southern species-groups (Fig. 4c), is unclear. The aurocinctus,
whitei and discors radiations appear to be endemic to Australia. The subnitidus radiation occurs
in South-East Asia and possibly also southern Africa. The maculatus radiation (which possibly
includes groups assigned to the discors radiation in Table 5) occurs throughout the Old World.
Within this radiation, the novaehollandiae group would seem to be a geologically recent
incursion into Australia from South-East Asia, given its prevalence throughout that region and
its exclusively tropical distribution within Australia. As previously noted, its major workers lack
the notched anterior clypeal margin that is characteristic of other Australian groups within the
maculatus radiation, which all appear to be endemic to Australia. It is therefore possible that the
latter evolved within Australia from Gondwanan stock.
It is one thing to describe the remarkable ant diversity in arid Australia; it is another to
explain it. An answer to the fundamental question of why ant diversity is so remarkably high in
arid Australia remains elusive. However, it is possibly related to unusually high ant productivity
and behavioural dominance over evolutionary time, perhaps driven by an unusually high supply
of plant exudates, ultimately in response to low soil fertility and erratically pulsed rainfall
(Andersen, 2003). Highly productive communities are also typically highly diverse, due to
interaction selection producing numerous highly specialised taxa (Greenslade, 1983). It is clear
that interaction selection has played a major role in shaping the arid Australian ant fauna at
higher taxonomic levels, with the extent of morphological, physiological and behavioural
specialisation exhibited by genera such as Meranoplus, Melophorus and Cerapachys being
unparalleled in other arid ant faunas (Andersen, 2003). However, the extent to which this might
explain exceptional diversity at the species level remains unclear.
Fig. 4. Proportions (%) of species belonging to northern (N), southern (S) and widespread (W)
species-groups in terms of their distributions in arid Australia. Data are from Tables 3-7 (data for
northern species-groups also include those occurring primarily in monsoonal Australia).
LITERATURE CITED
Agosti, D. 1997. Two new engimatic Melophorus species (Hymenoptera: Formicidae) from
Australia. Journal of the New York Entomological Society 105: 161-169.
Andersen, A.N. 1982. Seed removal by ants in the mallee of northwestern Victoria. In: R.C.
Buckley (ed.) Ant-Plant Interactions in Australia. Junk Press, The Hague, pp. 31-43.
Andersen, A.N. 1991. The Ants of Southern Australia: A Guide to the Bassian Fauna. CSIRO
Publishing, Melbourne.
Andersen, A.N. 1997. Functional groups and patterns of organization in North American ant
communities: a comparison with Australia. Journal of Biogeography 24: 433-460.
Andersen, A.N. 2000. The Ants of Northern Australia: A Guide to the Monsoonal Fauna.
CSIRO Publishing, Collingwood, Australia.
Andersen, A.N. 2002. Common names for Australian ants. Australian Journal of Entomology
41: 285-293.
Andersen, A.N. 2003. Ant biodiversity in arid Australia: productivity, species richness and
community organization. Records of the South Australian Museum Monograph Series 7: 9-92.
Andersen, A.N., Azcárate, F.M. & Cowie, I.D. 2000. Seed selection by an exceptionally rich
community of harvester ants in the Australian seasonal tropics. Journal of Animal Ecology 69:
975-984.
Andersen, A.N., Blum, M.S. & Jones, T.M. 1991. Venom alkaloids in Monomorium
'rothsteini' Forel repel other ants: is this the secret to success by Monomorium in Australian
ant communities? Oecologia 88: 157-160.
Bolton, B. 1994. Identification Guide to the Ant Genera of the World. Harvard University Press,
Cambridge, Massechussetts.
Bolton, B. 2000. The ant tribe Dacetini. Memoirs of the American Entomological Institute 65: 1-
1028.
Brown, W.L., Jr. 1958. Contributions toward a reclassification of the Formicidae. II. Tribe
Ectatommini (Hymenoptera). Bulletin of the Museum of Comparative Zoology 118: 173-362.
Brühl, C.A., Gunsalam, G. & Linsenmair, K.E. 1998. Stratification of ants (Hymenoptera,
Formicidae) in a primary rain forest in Sabah, Borneo. Journal of Tropical Ecology 14: 285-
297.
Christian, K.A. & Morton, S.R. 1992. Extreme thermophilia in a central Australian ant,
Melophorus bagoti. Physiological Zoology 65: 885-905.
Clark, J. 1941. Australian Formicidae. Notes and new species. Memoirs of the National
Museum of Victoria 12: 71-94.
Fisher, A., Kutt, A., Andersen, A., Karfs, R., Milne, D. & Risler, J. 2004. “Land condition”
as a surrogate for biodiversity health in tropical savanna rangelands. Proceedings of the
Australian Rangeland Society 13th Biennial Conference, Alice Springs, 5th-8th July 2004:183-
188.
Greenslade, P.J.M. 1979. A Guide to Ants of South Australia. South Australiam Museum,
Adelaide.
Greenslade, P.J.M. 1982. Selection processes in arid Australia. In: W. R. Barker & P. J. M.
Greenslade (eds.) Evolution of the Flora and Fauna of Arid Australia, pp. 125-130. Peacock
Publishing, Frewville, South Australia.
Greenslade, P.J.M. & Halliday, B. 1982. Distribution and speciation in meat ants,
Iridomyrmex purpureus and related species (Hymenoptera: Formicidae). In: W. R. Barker &
P. J. M. Greenslade (eds.) Evolution of the Flora and Fauna of Arid Australia, pp. 249-255.
Peacock Publishing, Frewville, South Australia.
Heterick, B.E. 2001. Revision of the Australian ants of the genus Monomorium (Hymenoptera:
Formicidae). Invertebrate Taxonomy 15: 352-459.
McAreavey J.J. 1947. New species of the genera Prolasius Forel and Melophorus Lubbock
(Hymenoptera: Formicidae). Memoirs of the National Museum of Victoria 15: 7-27.
McArthur, A.J. & Adams, M. 1996. A morphological and molecular revision of the
Camponotus nigriceps group (Hymenoptera: Formicidae) from Australia. Invertebrate
Taxonomy 10: 1-46.
McArthur, A.J. & Shattuck, S.O. 2001. A taxonomic revision of the Camponotus
macrocephalus species group (Hymenoptera: Formicidae) in Australia. Transactions of the
Royal Society of South Australia 125: 25-43.
Ogata, K. & Taylor, R.W. 1991. Ants of the genus Myrmecia Fabricius: a preliminary review
and key to the named species (Hymenoptera: Formicidae: Myrmeciinae). Journal of Natural
History 25: 1623-1673.
Shattuck, S.O. 1993a. Revision of the Iridomyrmex calvus species-group (Hymenoptera:
Shattuck, S.O. 1993b. Revision of the Iridomyrmex purpureus species-group (Hymenoptera:
Shattuck, S.O. 1996. Revision of the Iridomyrmex discors species-group (Hymenoptera:
Formicidae). Australian Journal of Entomology 35: 37-42.
Shattuck, S.O. & McArthur, A.J. 2002. A taxonomic revision of the Camponotus wiederkehri
and perjurus species-groups (Hymenoptera: Formicidae). Transactions of the Royal Society of
South Australia 126: 63-90.
Shattuck, S.O. & McMillan P. 1998. Revision of the species of the Iridomyrmex conifer group
(Hymenoptera: Formicidae), with notes on their biology. Australian Journal of Zoology 46:
301-315.
Taylor, R.W. 1972. Biogeography of insects of New Guinea and Cape York Peninsula. In: D.
Walker (ed.) Bridge and Barrier: The Natural and Cultural History of Torres Strait, pp. 213-
230. Australian National University Press, Canberra.
Taylor, R.W. 1991. Formicidae. In: The Insects of Australia: A textbook for Students and
Research Workers, Second Edition, pp. 980-989. Melbourne University Press, Melbourne.
Ward, P.S. 1980. A systematic revision of the Rhytidoponera impressa group (Hymenoptera:
Formicidae) in Australia and New Guinea. Australian Journal of Zoology 28: 475-498.
Appendix 1. Key to species-groups of Melophorus, based on minor workers unless otherwise

stated.
1. Hind tibia without a conspicuous apical spur ....................................................................... 2

Hind tibia with a conspicuous apical spur ............................................................................ 6
2. Antennal scapes relatively short, exceeding vertexal margin by at most half their total length
............................................................................................................................................... 3
Antennal scapes very long, exceeding vertexal margin by well over half their total length . 5
3. Mesosoma glabrous .................................................................................................. Group G

Mesosoma covered with short erect hairs .............................................................................. 4
4. Head and mesosoma densely punctate and dull throughout; gaster covered with short,
silvery pubescence ...................................................................................... fulvihirtus group
Head and mesosoma smooth and shiny throughout; gaster likewise shiny, without silvery
pubescence .................................................................................................................Group I
5. Dorsum of mesosoma markedly flattened, with distinct lateral carinae, and glabrous; distal
half of antennal scapes conspicuously swollen; frontal carinae closely approximated,
separated by a distance no greater than the maximum scape diameter (southwestern WA)....
............................................................................................................................majeri group
Dorsum of mesosoma not markedly flattened, without distinct lateral carinae, and covered
with short erect hairs; distal half of antennal scapes not conspicuously swollen; frontal
carinae separated by a distance greater than twice the maximum scape diameter (northern
Australia) ......................................................................................................anderseni group
6. Antennal scapes very short, at most exceeding vertexal margin by a distance equal to their
maximum diameter; propodeal spiracle very large, as long as at least half the maximum
height of the propodeum .................................................................................... potteri group
Antennal scapes exceeding vertexal margin by a distance equal to at least twice their
maximum diameter; propodeal spiracle not so large, usually much shorter than half the
maximum height of the propodeum ...................................................................................... 7
7. Anterior clypeus projecting over mandibles and acutely angled in profile, such that there is a
V-shaped notch between the anterior clypeus and mandibles; in most cases, antennal scapes
exceeding vertexal margin by well over half their total length.............................................. 8
Anterior clypeus at most feebly projecting over mandibles, and never acutely angled in
profile; in most cases, scapes exceeding vertexal margin by half or less their total length . 12
8. Mesosoma entirely glabrous .................................................................................................. 9

Mesosoma with erect hairs (sometimes confined to the pronotum)..................................... 11
9. Mesosoma longitudinally compressed, with a truncate propodeum that has a very short
dorsal face........................................................................................................froggatti group
Mesosoma elongate and saddle-shaped, with propodeum of normal length........................ 10
10. Total length >4 mm; head and mesosoma uniformly red, contrasting with black gaster.........
..................................................................................................................aeneovirens group
Total length <4 mm; mesosoma at least partly brownish, not uniformly reddish .... Group A
11. Antennal scapes glabrous, and usually exceeding vertexal margin by about two-thirds their
total length; second funicular segment about twice as long as wide......................... Group L
Antennal scapes with erect hairs, exceeding vertexal margin by about half or less their total
length; second funicular segment less than twice as long as wide............................ Group B
12. Maxillary palps relatively short, length of fourth (of six) segment about half or less that of
third; mandible sometimes with more than five teeth; clypeus often bulging such that it has a
distinct anterior face above the mandibles........................................................................... 13
Maxillary palps relatively long, length of fourth (of six) segment about two-thirds or more
that of third; mandibles not with more than five teeth, and clypeus not bulging such that it
has a distinct anterior face above the mandibles.................................................................. 16
13. Dorsal surface of head covered with numerous short hairs; head and mesosoma heavily
sculptured, at most feebly shiny........................................................................................... 14
Dorsal surface of head with scattered setae only; integument usually smooth and shiny.... 15
14. Mandibles with five teeth; foreparts brick-red, contrasting with black gaster .......... Group H
Mandibles with >10 teeth; color uniformly blackish ................................................ Group K
15. Antennal scapes exceeding vertexal margin by about one-third their total length.....Group F
Antennal scapes exceeding vertexal margin by at most one-quarter their total length ............
........................................................................................................................ wheeleri group
16. Antennal scapes exceeding vertexal margin by well over half their total length.....................
...................................................................................................................... iridescens group
Antennal scapes exceeding vertexal margin by at most half their total length .................... 17
17. Propodeum shorter than high (rarely, propodeal suture absent); head often dorso-ventrally
compressed .......................................................................................................................... 18
Propodeum as long or longer than high; head not dorso-ventrally compressed................... 22
18. Petiolar node short and scale-like, in profile very thin and acutely angled dorsally............ 19
Petiolar node longer, broadly rounded dorsally in profile ......................................... Group J
19. Mesosoma and gaster covered with whitish or silvery adpressed pubescence in addition to
erect hairs.................................................................................................................. Group C
Mesosoma with or without erect hairs, but not covered with adpressed pubescence .......... 20
20. Very small species, total length about 1.5 mm; mesosoma almost always glabrous ...............
........................................................................................................................mjobergi group
Larger species, total length about 2 mm or longer; mesosoma usually hairy ...................... 21
21. Mesosoma barrel-shaped, without even a propodeal suture ............................hirsutus group

Mesosoma with conspicuous metanotal groove or depression ................................. Group D
22. Head conspicuously sculptured, dull or feebly shiny........................................................... 23

Head very finely and feebly scalloped, effectively smooth and shiny................................. 24
23. Legs and antennal scapes clothed in long white hairs; foreparts brick-red, contrasting with
black gaster ............................................................................................................... Group E
Legs and scapes with at most scattered hairs; mesosoma brownish or black ..bruneus group
24. Tibiae clothed with long (longer than maximum tibial diameter) fine, erect hairs, giving
them a bottle-brush appearance .........................................................................pillipes group
Tibiae with very sparse or no long, erect hairs .................................................................... 25
25. Propodeum very short (about one-quarter total mesosoma length), prominently rounded, and
often markedly lower than promesonotum in profile, giving the mesosoma a strongly
biconvex profile .......................................................................................... perthensis group
Propodeum not so short (about one-third total mesosoma length) and prominently rounded,
only slightly lower than promesonotum in profile................................................fieldi group
Bolton, B. 2007. How to conduct large-scale taxonomic revisions in Formicidae, pp. 52-71. In Snelling, R. R., B. L. Fisher,
and P. S. Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
contributions. Memoirs of the American Entomological Institute, 80.
HOW TO CONDUCT LARGE-SCALE TAXONOMIC REVISIONS IN FORMICIDAE
Barry Bolton
Department of Entomology, The Natural History Museum,
Cromwell Road, London SW7 5BD, U.K.
ABSTRACT
An essay is presented that illustrates the author’s personal system, developed and expanded over
many years, for the production of taxonomic revisionary work in Formicidae starting from first
principles. It indicates what revisionary taxonomy means, stresses the main points essential to good
taxonomic procedure, indicates the snares and pitfalls that may be encountered when conducting
such work, and shows how to avoid commonly committed errors. Emphasis is laid on features such
as general procedure, production of good keys, gradual accumulation of relevant data and proper
layout and reproducibility of results in all stages of the work from initiation to final publication.
Key words: Hymenoptera, Formicidae, taxonomy, revisions, methodology.

Bolton: Large-scale Taxonomic Revisions 53
INTRODUCTION
The purpose of this contribution is to outline a functional procedure for the production of large-scale
taxonomic revisions in the ants. The term “large-scale” is of course subjective and open to various
interpretations. For instance, a student undertaking his or her first attempt at such a project may
consider 5 - 6 species to be quite a major undertaking, whereas an experienced student would pitch
the number of taxa involved considerably higher. Some would not consider the task to be large-scale
if under 50 species were involved. So the methodology outlined here must satisfy all the conditions
necessary to carry out a revisionary survey where more than a mere two or three taxa are involved.
What follows is written in the form of an essay, rather than a formal publication. The main
reason for adopting this procedure is that its various sections attempt to point out good techniques
but also to illustrate poor methodology together with errors of omission and commission. The
sections of the essay are written in a sequence which implies a smoothly continuous process from
first considerations to completion of the task. In reality, of course, things never go that regularly or
that smoothly. Factors over which the researcher has no control mean that the treatment of some taxa
will progress at a different rate from that of others. For instance, some institutions will respond to
requests for loans much more quickly than others, some journals may be difficult to obtain, and
resolving the taxonomy of some species will always be more difficult than others. There is no quick
cure for these complaints, but if the taxonomic work is structured as laid out here they should not
impede the overall progress of the study. Each problem, as it is resolved, can be slotted into its
appropriate place in the sequence without causing disruption to the work as a whole.
Finally, because this is a personal system, presented as an informal essay rather than a standard
scientific publication, there are no formal references given in the text. In part this is to save the
blushes of good taxonomists who make the odd mistake (we have all done it) who would thus find
themselves thrown into the company of serial incompetents. This is not to say that the work exists in
a vacuum; much of the basis of the taxonomic work discussed below can be found in: Mayr, E. and
Ashlock, P.D. 1991. Principles of Systematic Zoology (second edition): 475 pp. McGraw-Hill.
The aims of this essay
What I hope to set out and explain is a methodology for conducting a taxonomic study or
revision; in particular:
i how to set about studying numbers of related species, some of which are probably or certainly
already described and some of which are probably or certainly undescribed, and the
establishment of the unique identity of each,
ii how to build up a formal taxonomic study or revision of a given group, regardless of its size or
rank,
iii how to present the findings in such a way that others can understand the taxonomy and use it to
identify their own specimens.
What the essay is not concerned with is piecemeal alpha-taxonomy. Taxonomy of this nature is
basically concerned with the detection, discrimination, description and publication of isolated new
taxa (species, genera, etc.). Each taxon is known or assumed to be new to science and is known or
assumed to be undescribed. The author therefore issues a description and names the taxon. This may
be described as the “one by one” technique and is usually acceptable in groups that are very well
known (birds, primates, butterflies), but in most insect groups the numbers of undescribed species
are vast and this “one by one” technique only serves to increase bafflement and frustration among
those wishing to identify taxa; and this applies to other taxonomists as well as non-taxonomic
biologists.
The main problem with the “one by one” approach is that anyone faced with the task of
identifying a species is confronted by numerous descriptions of single species with no way of
knowing where to begin. The specimen must be compared with each description in turn, in the hope
that eventually a match will be found. This is an incredibly frustrating and extremely
time-consuming exercise. The frustration is often compounded because exponents of the “one by
one” technique frequently fail to draw attention to uniquely diagnostic characters and fail to provide
any notes by which the new species may be compared with those that are already known. There are
even cases where piecemeal alpha-taxonomic authors have described new species in well known
genera, where keys to species already exist, and have failed to indicate where the new species fit into
the key or how they modify the pre-existing system. This is inexcusable: it is not just poor
taxonomic procedure, it is bad scientific methodology.
Finally, there are people who, with the best will in the world, cannot fulfil the four factors listed
below, try as they may. Their discriminatory faculties seem insufficient to cope with the huge
numbers of characters available, or they are unable to sort or categorize the observed characters in
any meaningful way. There are others who cynically disregard the basic factors in a rush to have
their names associated with supposedly new taxa, as if such a trivial aim was important. Such people
should avoid descriptive taxonomy and comparative morphology as their efforts, whether
unfortunately inept or cynically divisive, only add to the problems rather than contribute to solutions.
Good taxonomy ultimately relies upon:
i the detection of unique characters that diagnose a taxon,

ii the comparison of characters that differentiate related taxa,
iii the recognition of characters that unite related taxa,
iv the description and ordering of those characters in such a way that each taxon can be
recognized by other workers
In short: the function of taxonomy is to establish identity.
INITIATION PHASE OF THE PROJECT
Select a group to study
Choice of a study group may depend upon:
i economic importance,
ii value in biodiversity work,
iii understanding evolution,
iv fixing identities to investigate phylogeny,
v interpreting zoogeography,
vi interest in morphological adaptations,
vii establishing patterns of speciation,
viii forming a basis for ecological or ethological studies,
ix attempting to establish a natural classification,
x learning more about the inhabitants of the planet,
xi your supervisor says this is what you are going to do.
Depending on circumstances of time or funds available, and the degree of experience and
determination of the researcher, the group selected for study may consist of a species group, a genus,
a defined group of several genera, or a higher taxon such as a tribe or subfamily.
Ideally the area covered by a revision should encompass the entire range of the group in
question, or at least the species of a group from an entire Zoogeographical Region if the group is of
very wide distribution. However, political or financial considerations often restrict taxonomic
surveys to a single country, usually the one providing the finances and facilities. The danger here is
that instead of solving taxonomic problems a survey of a relatively small area, especially if the area
is treated as isolated, may add confusion. There is a real risk of the unwitting creation of junior
synonyms or homonyms because the researcher is insufficiently acquainted with the extralimital
fauna. Thus, if circumstances dictate that a project has to be restricted to a relatively small area it is
important that the researcher obtains a working knowledge of the entire group. It is useful to
remember that insect species do not recognize red lines on political maps, despite the fact that
bureaucrats tend to think they should.
Preliminaries
Once a group for study has been decided upon the next step is to acquire material so that work
can commence. For most researchers a small to moderate collection will usually be available in the
institution where the work is taking place. Additional material can be obtained from:
i museums and other state or local institutions,

ii private collections,
iii universities,
iv field collecting.
Collections held in categories i - iii can usually be borrowed through the mail but sometimes it
is necessary to visit the appropriate institution. Material held in these collections sometimes includes
types or other specimens of historical importance, and is always useful for obtaining data on
distribution and variation of the species concerned. Always treat borrowed material with respect and
return it in good condition within the terms of the loan agreement. When returning specimens
through the mail always remember to pack them safely; protect the specimens against mechanical
damage as most postal services are unaware that insects tend to be fragile.
Field collecting is the best way to obtain fresh specimens and to get to know the appearance and
behaviour of the group in its natural habitat. Take time to discover the best method(s) for collecting,
transporting and storing specimens of the group in question. If unaware of the techniques involved
then consult with other people who have experience in the same or related fields and ask their
opinion; they are usually very happy to advise. There are also some museum publications that deal
with the acquisition and storage of entomological specimens. When fresh material is obtained always
ensure that it is properly mounted and safely stored and has appropriate data labels. When collecting
abroad, always ensure that the correct permission to do so has been obtained. Many countries now
have strict regulations concerning the collecting of insects, especially the larger and showier groups,
and they tend to become irate if their regulations are flouted.
Note that the acquisition of material and preliminary work on it precedes searching the
literature for names, precedes amassing and reading vast photocopied files of original descriptions,
and precedes the examination of type-material.
This approach is important because at this stage the critical aspect is to have a reasonable
representation of the group from its entire range and to develop a personal insight into its
morphology and variation that is not unduly influenced by what has previously been published.
Initial dependence on previous publications has a strong tendency to restrict the scope of a new
investigation. There is real psychological pressure for the researcher to make an unwarranted
assumption that previous authors have already investigated all the useful characters. This is
especially true in groups where there are many isolated descriptions of single species, carried out by
multiple authors from several countries, over many years. In fact, what has usually happened in such
cases is that an initial author has described, say, the first species in a genus. He has stressed certain
characters that he considers most important, out of the enormous number available, merely because a
description must have a finite length. A second author, a few years later, adds a couple more species
to the genus. To contrast his species from the original he stresses differences from those characters
cited by the first author, especially if he has not bothered to see the original author’s type-specimens.
The third worker in the chair then reads these two works. He either again concentrates on variation
in the same characters, because he has no inkling of the state of characters that have not already been
mentioned; or he inserts characters that have not been discussed previously, assuming that because
they were not mentioned they were not present. You can see what is happening - the descriptions,
despite becoming less trustworthy, are assuming more importance than the specimens they purport to
represent. This procedure unfortunately becomes self-perpetuating and as it continues the
descriptions become more and more trivial and have less and less value. As a corollary, as the
number of supposedly different species, really just differing descriptions, increases, the ease with
which a serious taxonomist can compare all the type-material decreases.
Contrary to what many seem to believe, the objective of a taxonomic revision is to establish the
unique identities of the species involved and to transmit that information to a reader in a reasonably
compact, concise manner. It is not, and should not be reduced to, a mere review of previous
type-material with the express aim of finding new species that can be described.
The discovery and description of new species is incidental, it is not the prime objective of the
exercise.
The first thing that needs to be achieved when commencing work on a group is a reasonable
understanding of the limits of all the species that constitute the group, in a way that is not unduly
influenced by the opinions of earlier workers. It is therefore best to leave the examination of the
type-material of previously described species until a later stage, after a good understanding of the
characters and relationships of all the species has been independently acquired.
All students of taxonomy should be acquainted with the International Code of Zoological
Nomenclature (Fourth Edition, 1999) before embarking on a project. This volume sets out the rules
and regulations that govern the naming of animal taxa. It should be born in mind that the rules of
nomenclature are tools for ordering and understanding taxonomy and classification; nomenclature
should not be regarded as an end in itself.
Before undertaking a taxonomic revision all students should become familiar with, and
understand the function of, the type-concept in taxonomy, the rules governing synonymy and
homonymy, and those governing the status and availability of names; these aspects are not discussed
here.
Initiation of taxonomy
When a sufficient number of specimens of the target group has been accumulated, that is, when
representatives of several apparently distinct forms with consistently differing morphological
features have been amassed, the taxonomic investigation can begin. It is useful, initially, to adopt a
null hypothesis.
Assume that all previous work has no value, and that determination labels on all specimens
(except genuine type-material that happens to be at hand) are incorrect or are misidentifications.
This will certainly be untrue for a proportion of the already-named specimens, but is a good way
to begin as it puts every specimen on the same philosophical level.
Get to know the group intimately; take time over it. Examine closely all the material that is
available and compare and contrast individuals and groups of individuals. When a good overall
knowledge of the group’s morphology and variation has been acquired, and when the researcher is
conversant with any specialized morphological terminology that the group may require, search for
the following:
i unique characters that discriminate individual species,

ii characters that unite pairs or groups of species,
iii universal characters that encompass all species.
Polarity of these characters, that is whether they are plesiomorphic (ancestral) or apomorphic
(derived) characters, is not important at this stage; what the characters are, and their presence or
absence through the various species of the group, is the important feature as the aim is to establish
unique identity, not phylogeny. If, later in the investigation, a phylogeny is to be constructed, the
characters can quickly be reassessed for phylogenetic utility and fed into an appropriate data matrix.
As characters in groups i - ii are detected, arrange the specimens in collection drawers or good
quality specimen boxes, so that individuals considered to belong to a single species are grouped
together, and species that appear to be closely related because of shared characters are in proximity.
Separate these from species or groups of species with different shared characters.
During this process, or when all the species are considered to be adequately segregated, give
each a numerical or alphabetical recognition code; something simple like “species A”, or “species
1”. Do not worry about names at this stage, that will come later.
Also during this initial sorting of material take preliminary notes of the main characters of each
taxon. Make a separate heading for each of the taxa, using the code-number or code-letter heading of
each, and under it write down the critical characters so far discovered and which characters are also
shared with other taxa. Add observations on variation, if any, and list the locality data of the material
examined. For example:
Species A
Characters:
i) Antenna with 11 segments.
ii) Apical antennal segment with scale-like sensory hairs.
iii) Mandible with 5 teeth; third tooth from apex the longest.
iv) Second abdominal tergite smooth and shining everywhere.
v) Hind femur with spines present on dorsal surface.
Character comparisons:
i) also in sp. C and sp. E.
ii) and iii) also in sp. B.
iv) also in sp. C.
v) not in any other species.
Material examined
ENGLAND: Surrey, Kew Green, 12.x.1975, on oak tree trunk (collector’s name); BELGIUM: 5
km east of Ronse, 23.v.1906, on soil at base of oak tree (collector’s name); HOLLAND: Bunnk,
near Utrecht, no further data.
Using those characters that have been found to unite some species and separate them from
others, and those characters that are uniquely applicable to single species, and using the species
codes as end-points, the first draft of an identification key can be constructed.
First draft key
The key is the most important part of the study. It should be built up, developed and
continually revised and updated as the study progresses, so that it acts as a continuing bench-mark
for the progress of the work, and allows you and others to test the reproducibility of the results.
At risk of boring because of repetition, I stress: key construction and modification should be a
continuous process through the course of the study. The key should be modified and expanded as the
work progresses and as more taxa and their characters are discovered or differentiated and added in.
It is the most important part of the study, the part that reflects your hypotheses of what constitute
species in the group. It is also the part of the work that allows your conclusions to be tested by
others. Without an accurate key there is no easy way for any other worker to gain access to your
species concepts. It should not be the last task undertaken. In some taxonomic works it is obvious
that construction of the identification key has been the last act of the study. In other words, after the
researcher has examined the material, made decisions about species differentiation and written all
the descriptions, only then has the key been constructed, but from the descriptions and not from the
actual specimens. The key is thus being treated as a mere adjunct of the descriptions, built with
characters abstracted from the descriptive work and implying that the descriptions provide more
information than the specimens from which they were derived.
There are many ways of constructing keys for identification purposes but the simplest to
understand, and usually the most universally functional, is the dichotomous key. In this system the
reader is presented with a series of pairs of alternatives, each of which leads either to a second pair
of alternatives or an identification. Each pair of alternatives is called a couplet, and each half-couplet
a lug. The specimen to be identified is compared with the character(s) listed in each lug of the first
couplet, and the one which corresponds is followed through the remainder of the key.
Imagine that the “species A” mentioned above belongs in genus Xus, and let us say that after the
preliminary work described above five species have been differentiated and are represented by the
recognition codes “species A, B, C, D and E” respectively. Let us also say that these five can
apparently be grouped and segregated by different expressions and combinations of the following
characters: number of antennal segments, form of sensory hairs on the apical antennal segment,
number of teeth on the mandible and their relative lengths, form of sculpture on the second
abdominal tergite, and presence or absence of spines on the hind femur. A dichotomous key such as
the following could be drawn up.
1 Antenna with 11 segments............................................................................................... 2

- Antenna with 12 segments............................................................................................... 4
2 Mandible with 5 teeth. Hind femur with spines present on dorsal surface .........species A
- Mandible with 8 teeth. Hind femur without spines on dorsal surface.............................. 3
3 Second abdominal tergite smooth and shining. Counting from apex of mandible tooth 3 is
longer than teeth 4 and 5..................................................................................... species C
- Second abdominal tergite densely reticulate-punctate. Counting from apex of mandible
tooth 4 is longer than teeth 3 and 5 ..................................................................... species E
4 Principal sensory hairs on apical antennal segment scale-like. Mandible with 5 teeth.
Second abdominal tergite finely and densely longitudinally rugose................... species B
Principal sensory hairs on apical antennal segment filiform. Mandible with 7 teeth. Second
abdominal tergite transversely striate on basal half, smooth apically .................species D
Thus a specimen of Xus species E would track through the first lug of couplet 1 (which leads the
reader to couplet 2), the second lug of couplet 2 (which leads to couplet 3), and the second lug of
couplet 3, which gives the identity.
The sole function of a key is to allow taxa to be identified. It is concerned only with the
establishment of identity and to allow the recognition of taxa based on an ordering of characters. It
does not need to reflect classification or phylogeny, nor should it be a vehicle for reflecting the
ability of the author to impress the reader.
A few guidelines for key construction:
i Chose obvious and clear-cut characters, especially for main couplets that segregate groups of
species. Describe them accurately (illustrate if necessary). If any special condition, or a
particular orientation or viewing angle, is necessary then note it in the couplet; do not leave the
reader to guess.
ii Avoid hanging comparatives that are incomprehensible to a reader (e.g., legs longer versus legs
shorter; which begs the question “than what?”). Always give some indication of what the
comparison means. For example by direct comparison with some other aspect of the same
specimen (legs longer than head width versus legs shorter than head width), or use comparative
measurements (legs > 1.5 times head width versus legs < 1.5 times head width), or use absolute
measurements (legs 3.0 mm or more versus legs 2.0 mm or less).
iii Avoid purely subjective nonsense such as, “ head more reddish versus head not so reddish”;
“known from queens and workers versus known from workers alone”. A reader cannot possibly
interpret such statements unless all the author’s original specimens are to hand, which is
extremely unlikely. Experience teaches that even on the rare occasion when the specimens are
all available the task frequently remains impossible.
iv Ensure that lugs of couplets contrast directly. A couplet such as “legs red versus thorax black”,
is an absurdity as the two lugs are not directly comparable.
v Avoid the “not as above” lug. This involves a couplet that gives many characters, or even a
short description, in the first lug but merely says in the second lug “not as above”, or
“characters otherwise”, or words to that effect. Such couplets are incredibly frustrating as the
second lug is not transmitting any useful information. It leaves the reader in limbo, with no
direct alternatives between the two lugs to which the specimen under examination can be
compared.
Culmination of initial phase
By this stage the following results have been achieved:
i all the available material has undergone a careful sorting to species and the species have
been allocated recognition codes,
ii characters diagnostic of the species have been noted under corresponding coded headings,
iii characters that unite groups of species, or that isolate groups of species, have been noted,
iv a first-draft key, using the species-codes as end-points and the characters detected, has been
constructed.
INTERMEDIATE PHASE OF THE PROJECT

Consultation of literature
It now becomes necessary to search out and consult earlier literature on the group. The function
of the literature search at this stage is:
i to build up a complete list of previously established names,

ii to obtain the references that give the original descriptions of the names,
iii to establish what the type-material consists of and where it is deposited,
iv to discover the type-locality of each nominal taxon,
v to establish the current status of the name by reference to usage in the literature, between
its original description and the present.
All these data will be used later in the construction of the taxonomic synopsis of each species,
discussed below. Sometimes there is a recent check-list available for the group in question, or even a
full catalogue, which is better as it usually provides much more information than a check-list. Either
of these will provide a compendium of the required names and the latter will also give the essential
references in full. Sometimes however it is necessary to consult abstracting journals such as
Zoological Record or on-line compendia and go through them searching for appropriate entries of
original descriptions or later references involving names in the group. If a check-list or catalogue is
available ensure that the abstracting journals are checked from the catalogue’s date of publication to
the present, to catch any contributions published after the appearance of the catalogue. As references
are found, look them up in the library or reprint collection, or search for them on-line; photocopy
them if necessary. Be aware of the Inter-Library loan system, which can be extremely valuable in
obtaining copies of more obscure publications. If the authors of any previous contributions are still
alive, write and ask if a reprint or photocopy of the publication is still available. Most authors are
happy to distribute free copies of their papers to enthusiastic researchers.
The species in these publications will of course bear names, and these names will not
correspond to your species-codes at this time, nor do they need to. Open a separate entry or file for
each published name, regardless of its current status or rank. Under each name, as it is tracked down,
note the taxonomic name, author and bibliographical reference, what the type-material is and how
much there is of it, the type-locality and any other relevant data that are given, the name of the
collector and the institute in which the type-material is deposited. Also note any data concerning
material examined but excluded from the type-series and other comments (if any). Build up the data
thus obtained into a taxon file and a references file.
The relevant data for some species of our imaginary genus Xus (Xus aus, Xus bus, Xus cus and
Xus dus), together with their equally imaginary authors and other information, would appear as
follows.
TAXON FILE
Xus aus Green

Xus aus Green, 1883: 207, fig. 1. Holotype queen, FRANCE: near Paris, 15.vi.1880, under stone (M.
Brun) (Paris Museum).
Xus bus Green

Xus bus Green, 1885: 12. Syntype workers, BELGIUM: near French border, ix.1884, on tree trunk (U.
Blanc) (Paris Museum). [Junior synonym of aus: Blanc, 1890: 31 (Blanc maintains that queen aus
and worker bus belong to same species).]
Xus cus (Schwarz)

Gus cus Schwarz, 1888: 89. Syntype workers, GERMANY: no accurate locality given, 1886, swept from
plants (H. Grün) (Berlin Museum).
Xus cus (Schwarz); Blanc, 1890: 35 [combination in genus Xus].
Xus dus Blanc

Xus dus Blanc, 1890: 33, figs 1 - 4. Syntype workers, queens and males, FRANCE: near Belgian border,
21.ix.1885 (U. Blanc) (Paris Museum and Brussels Museum).
Note that proper names of species are always written in italics, and that the initial letter of the
genus-rank name only is a CAPITAL. Note also the brackets (parentheses) round the author’s name
in the case of Xus cus (Schwarz). These indicate that the species name has been transferred away
from the genus name with which it was originally combined (Gus) because a later author, in this case
Blanc, has judged that the name cus is properly ascribed to genus Xus, not genus Gus. Thus in the
original combination Gus cus Schwarz, the author’s name is not in brackets; in the changed
combination Xus cus (Schwarz) it is. This is the only circumstance in which brackets are placed
around the name of an author. Should the name cus revert to genus Gus at a later date (i.e. Blanc got
it wrong) then the brackets will again disappear. Note also that species described from series of
specimens at these relatively early dates usually leave them as a syntypic series, without nomination
of a holotype. During revision of such series it may be necessary to nominate one specimen as a
lectotype: see the appropriate sections of the International Code of Zoological Nomenclature (Fourth
Edition, 1999) for the correct procedure.
It is preferable to include the complete data of the type-material for each taxon in these
synopses; see the discussion of taxonomic synopses, below. Some authors include only the name and
bibliographical reference here (for instance Gus cus Schwarz, 1888: 89), and relegate all the other
information to other places in the text. I consider it important for any reader to have the full data that
applies to the type-material immediately visible, not to have to go searching for it in several different
places in the text or to have it mixed in with the list of other material that has been examined. This is
especially important in species with extensive synonymies as it allows the reader access to all the
relevant data pertaining to types and taxonomic history merely by running an eye down the list of
entries.
At the same time that the above data are being amassed, open a references file (bibliography)
and list all references found from which data have been abstracted. List them in alphabetical order
by author and in publication date order for each author who has provided more than one contribution
to the group. It is best practice to write out the name of the periodical, journal or book in full. In the
past it was common for authors to present the names of periodicals in abbreviated form, but these
days there are so many journals with very similar titles that full writing of their names is necessary to
avoid confusion. As with our imaginary taxa, so the authors, dates, titles and journals mentioned
here are also imaginary.
REFERENCES FILE
Blanc, U. 1890. Xus bus est un synonyme de Xus aus, et autres choses nouveaux. Journal
d’Entomologie Incroyable 12: 30-35.
Green, A. 1883. The genus Xus described for science. Bulletin of Imaginary Taxonomy 3:
200-250.
Green, A. 1885. Yet another species of the genus Xus in the West Palaearctic region. Bulletin
of Imaginary Taxonomy 5: 1-18.
Schwarz, E. 1888. Beschreibungen nicht-existierender Insekten Deutschlands: 1000 pp. Berlin.

Examination of types
At this stage it is possible to begin replacing the species-codes in the key and the taxon file with
genuine names. By reference to the data compiled in the taxon file, the various museums and other
institutions where type-material is deposited can be contacted and loans of the appropriate specimens
arranged. Or, if funds are available, the institution can be visited, which is usually pleasant and may
even broaden the mind. When borrowing specimens always ensure that the loan conditions stated by
the lending institution are complied with. Any specimen has considerable scientific value, as well as
historical value, but in the case of type-material this is particularly true because:
It is the type-material that establishes and carries the name of the species; not its original
description, not any later interpretation of the species, not any later opinion, not any later mention
in the literature.
Treat materials borrowed from other institutions with great care because any specimen entrusted
to you for study remains the property of its home institution and it must be retained in good
condition for future researchers.
On receipt of a purported type-specimen it should immediately be checked against the taxon file
and the original description, to ensure that it is genuine. This is important as sometimes material
becomes mislabelled at some point in its history, or labels may have been inadvertently switched at
some time in the past, or the original labels may have been misunderstood by a later curator so that
the specimen has acquired a spurious type-status when it is not actually part of the original
type-material. Its data label(s) should match the data noted for the specimen in the original
description, and the specimen itself should fit the description. On occasion, and especially with older
material, the data label(s) on the specimen may have information additional to that given in the
original description. Additional data from the label should be added to the taxon file, but only
provided the identity of the specimen is assured.
When a type-specimen has been confirmed as valid it should then be run through the
already-prepared draft identification key. There are four possible results.
i The type-specimen runs directly to one of the species-codes in the draft key.
It can then be compared directly to specimens bearing that code in the study collection. If the
type-specimen and coded collection material are judged identical and conspecific within the known
limits of variation of the species then the code reference can be abandoned throughout the
manuscript and replaced by the name.
For example, the type-material of Xus cus is found to run to species A of the draft key. The key
end-point (at first lug of couplet 2) is changed to reflect this.

2 Mandible with 5 teeth. Hind femur with spines present on dorsal surface ...................cus
- Second abdominal tergite densely reticulate................................................................ etc.
Also the file covering the taxon is changed from
Species A
Characters:
iii) Mandible with 5 teeth; third tooth from apex the longest....etc.
to
Xus cus (Schwarz)

Gus cus Schwarz, 1888: 89. Syntype workers, GERMANY: no accurate locality given, 1886,
swept from plants (H. Grün) (Berlin Museum).
Xus cus (Schwarz); Blanc, 1890: 35 [combination in genus Xus].
Characters:
iii) Mandible with 5 teeth; third tooth from apex the longest....etc.
iv) The type-specimen fails to run to any keyed species code.
The key must therefore be modified to accommodate this extra taxon and also a new taxon file
must be created. Write a new couplet and insert it at the appropriate place. Remember to adjust
couplet numbers, and lead-on numbers at the ends of lugs, where necessary throughout the
remainder of the key.
For example, the holotype of Xus fus has been obtained and fails to run to any terminal lug in
the key because its diagnostic characters exclude it from all previously incorporated taxa. Let us say
that Xus fus has 11 antennal segments and 8 teeth, which would bring it out in couplet 3 of the
original key, but that its head is longer than broad and its second abdominal tergite is densely rugose,
whereas the previous occupants of couplet 3 (species C and E) have heads that are broader than long
and different abdominal sculpture. The key can be modified to accommodate the extra species by
writing a new couplet 3 and making the original couplet 3 become couplet 4; but note how this
affects couplet numbers throughout.

- Antenna with 12 segments .............................................................................................. 5
2 Mandible with 5 teeth. Hind femur with spines present on dorsal surface ...................cus
3 Head longer than broad. Second abdominal tergite densely rugose.............................. fus
– Head broader than long. Second abdominal tergite smooth or reticulate-punctate, but not
densely rugose ................................................................................................................. 4
5 Principal sensory hairs on apical antennal segment scale-like. Mandible with 5 teeth.
Second abdominal tergite finely and densely longitudinally rugose................... species B
- Principal sensory hairs on apical antennal segment filiform. Mandible with 7 teeth. Second
abdominal tergite transversely striate on basal half, smooth apically .................species D
It is not compulsory to add the new couplet where it is placed in the above example, although in
this case it is reasonable because other characters of Xus fus seem to imply that it is closely related to
species C and E (same antennal segment count, same teeth). But suppose that fus is the only species
in the entire group which has the head longer than broad and that this feature is very obvious and
unmistakable. It could therefore be justified to make fus run out first, as the subject of a new couplet
1.
1 Head four times longer than broad................................................................................ fus

- Head two times broader than long ................................................................................... 2

Similarly, because it shares the same number of antennal segments with Xus cus, species C and
species E, it could be inserted as a new couplet 2.

2 Head four times longer than broad................................................................................ fus

- Head two times broader than long ................................................................................... 3
3 Mandible with 5 teeth. Hind femur with with spines present on dorsal surface ...........cus
For best results always organize a key so that the characters used, and the order in which they
are used, impart maximum clarity to the key and maximize the chance of a reader obtaining a
correct identification.
iii The type-specimen terminates at a lug already occupied by a name, that name being in
place as the result of keying in a previous type-specimen of a differently named taxon.
This event usually implies that synonymy, a situation in which a single taxon has been described
more than once, and has therefore been invested with more than one name, may be present.
However, it sometimes implies that further analysis may be required because two species really are
present but they are distinguished by characters that have not previously been included in the
analysis, so both of them will therefore run to the same end-point in the key.
Compare the type-specimens of the two nominal species directly and with any other material
standing under the original name in the collection. Review all previously used characters and their
variation (if any) and analyze for other discriminating characters, to ascertain if both type-specimens
represent one species or if two species are really present. If, after this re-analysis, all the material is
judged to represent one species, then there is synonymy and the appropriate rules come into force
because no single species is allowed to bear two different proper names. If two species can be
discriminated after the re-analysis, insert a new couplet in the key (as in ii., above) that
accommodates the new findings.
In the case of a confirmed synonymy combine the taxon files of both names under the senior
name (the older, first-described name is senior except when it is a junior homonym in need of
replacement). Replace the junior synonym (the younger, second-described name) with the senior
synonym if the junior happens to appear anywhere in the text, except where it appears in the taxon
file of the senior name, of course (see taxonomic synopses, below); and ensure that the correct name
occupies the end-point of the appropriate lug in the key.
iv The result of running the type-specimen through the key is ambivalent.
Re-examine all taxa that are grouped around the point where the type-specimen fails in the key.
Compare the type-specimen with all apparently related species and re-assess the validity of the
species themselves. It may be necessary to re-analyze and re-order a group of species and find
alternative characters, or it may be that the ambivalence is the result of previously unsuspected
variation. After re-examination has been concluded re-run the results through the revised key and
proceed through the above sections i, ii or iii again, if necessary.
When the study has reached this stage, or when convenient prior to this stage, enquiries should
be made to institutions and other entomologists that have not previously been approached, to
ascertain if any more material of the group under study is available. If the response is positive then
ask to examine it, by loan or visit. This will ensure that the study will have the maximum
representation of species and reasonable numbers of specimens of each species, and will furnish
extra information on variation and distribution. Any extra material acquired should be sorted and run
through the key. If previously unseen species are represented then new taxon files can be created and
new couplets inserted in the key. It is very useful to obtain some previously unseen material at this
time and feed it into the revision as it provides a test of the efficiency of the key and gives valuable
information about the validity of the characters used and the stability of the taxonomic conclusions
that have been postulated.
Finalizing the key
When the type-material of all previously described taxa has been keyed successfully then those
lugs that still terminate in a species-code number or code letter automatically represent undescribed
taxa (new species). Names must be selected for these and formal descriptions of them will need to be
made. It is worth bearing in mind that in the key the sum of all lugs leading to an identity constitutes
a definition of that species.
Additions to the key can also be made at this stage to include the known distributions of the taxa
and the figure numbers that illustrate the characters mentioned. Distributions need only be
generalized (countries or states/counties within larger countries), to give the reader an indication of
the range of the taxon, because more detailed geographical information will be available under the
heading of “Material examined” for each taxon.

2 Head four times longer than broad (Fig. 1). (Britain, Germany, Poland, Denmark, Norway,
Sweden) …… ............................................................................................................... fus
- Head two times broader than long (Fig. 2) ...................................................................... 3
3 Mandible with 5 teeth. Hind femur with spines present on dorsal surface (Fig. 3).
(Germany, Austria, Belgium, France, Spain)................................................................cus
- Mandible with 8 teeth. Hind femur without spines on dorsal surface (Fig. 4) ................. 4
CONCLUDING PHASE OF THE PROJECT
Taxonomic synopses
i Taxonomic synopses of previously described taxa
At completion of the study each previously established name should be subtended by a formally
laid out taxonomic synopsis that enables a reader to see at a glance the current status of the name and
a history of its use and how it was interpreted. The synopsis should show an initial entry giving the
reference of the original description, type-data, type-locality, type-depository, and an indication of
whether type-material has been examined during the course of the study. Subsequent entries should
indicate any synonymy or homonymy in which the name has been involved, any changes of status or
combination that the name may have been subjected to, and any other references that are felt to be
critical, such as those describing castes or sexes different to that of the original description.
What need not be included here is a compendium of papers that mention the name in a context
that is not critical to understanding its taxonomic history. References to ecology, ethology,
zoogeography and so on, can usually be relegated to the general discussion of the species following
its formal diagnosis or description. However, if such references are deemed necessary for any
reason, then they should be incorporated in the taxonomic synopsis in a way that makes it clear they
are not an essential part of the taxonomic history. For instance, they can be enclosed in square
brackets, or indented from the main text, or both. They should not interfere with the understanding of
the taxonomic synopsis, which should be clear and concise.
For example, imagine that all the presumed species listed in the initial taxon file above (Xus aus,
bus, cus, dus) had been examined, all their type-material and much other material had been analyzed
and compared, and the conclusion reached that only a single real species was represented. That is to
say, all the apparent differences invoked by the original describers of the names were merely the
result of normal variation within a single species. The taxonomic synopsis covering all these events
would appear something like this:
Xus aus Green
Xus aus Green, 1883: 207, fig. 1. Holotype queen, FRANCE: near Paris, 15.vi.1880, under stone (M.
Brun) (Paris Museum) [examined].
Xus bus Green, 1885: 12. Syntype workers, BELGIUM: near French border, ix.1884, on tree trunk
(U. Blanc) (Paris Museum) [examined]. [Junior synonym of aus: Blanc, 1890: 31 (aus and bus
are queen and worker of a single species); synonymy confirmed.]
Gus cus Schwarz, 1888: 89. Syntype workers, GERMANY: no accurate locality given, 1886, swept
from plants (H. Grün) (Berlin Museum) [examined]. Syn. n.
[Xus cus (Schwarz); Blanc, 1890: 35 (combination in genus Xus).]
Xus dus Blanc, 1890: 33, figs 1 - 4. Syntype workers, queens and males, FRANCE: near Belgian
border, 21.ix.1885 (U. Blanc) (Paris Museum and Brussels Museum) [examined]. Syn. n.
All the data necessary to understand the history and current status of this species, and the
specimens that were involved, are immediately obvious. They are presented in temporal order,
earliest event first, as far as is possible. In this example the only reference that is not in date order is
that indicating the synonymy of aus and bus, which immediately follows the entry for bus.
Note that in the case of cus the indication of synonymy follows the entry for the original
description, not the later entry indicating its change of generic combination. This is because the
original reference contains the type-data and it is the type-material that carries the name. Note also
that there are no punctuation marks allowed between the name of a taxon and the name of its original
author.
ii Taxonomic synopses of new taxa
New species do not have a formal synopsis such as that given above, for the simple reason that
they have no former history. It is, however, necessary to give each new species a name, and that
name must be different from any other formerly used in the genus. This applies to names that are
currently in synonymy as well as those still considered as valid species. The reason for this is that
names in synonymy are still taxonomically available because a later researcher may produce
evidence that a previous synonymy was incorrect. When this occurs the later researcher is at liberty
to resurrect the name from synonymy. The name of a new species must also be different from any
name that was originally proposed in the genus but which has since been transferred to a different
genus. When a name is proposed for the first time it must be followed by an indication that it is new.
The term “new species” may be written immediately after the name, or a contraction of the Latinized
version of this (species novus) may be used: sp. n.
Data explaining the type-specimens and provenance of new species must be included at some
point following the introduction of the new species. There is no fixed place to do this but it is usually
accomplished directly after the title of the species or immediately following the description. Suppose
that a new species was detected in our imaginary genus Xus, which was described and published by
Dr Black in 1952, who nominated the species as Xus nus. Black decided to propose the name, then
give the description, and then the type-data, thus:
Xus nus sp. n.
Holotype worker: definition (see diagnoses and descriptions, below).

Paratype workers, queens and males: further definition if necessary and notes on variation if
any.
Holotype worker, GERMANY: 2 km. NW of Riesa, on bank of Riv. Elbe, 21.x.1950, on trunk
of alder tree, no. 602 (H. Brun) (Humboldt University Museum).
Paratypes. 10 workers, 2 queens and 4 males with the same data; 5 workers and 1 queen with
same data but 1 km. NW Riesa, 22.x.1950, no. 605, under stone (H.Brun) (Humboldt University
Museum, BMNH, MNHN).
The type-data indicate what the type-material is, how much of it there is, where and when it was
collected, other data referring to the specimens (on alder tree and collection number), the name of
the collector (in italics in parentheses), and the institutions in which the type-material is deposited,
either by name or by abbreviation, also in parentheses.
At a later date, if this species is referred to again in a revision or other formal taxonomic
publication, the above data will form the information of the taxonomic synopsis and will appear like
this:
Xus nus Black

Xus nus Black, 1952: 64. Holotype worker, GERMANY: 2 km. NW of Riesa, on bank of Riv. Elbe,
21.x.1950, on trunk of alder tree, no. 602 (H. Brun); paratype workers, queens and males with
same data and paratype workers and queen, 1 km. NW Riesa, 22.x.1950, no. 605, under stone
(H.Brun) (Humboldt University Museum, BMNH, MNHN) [examined].
Diagnoses and descriptions
The definition of a species may consist of a diagnosis, a description, or both. Some authors
present diagnosis and description as separate entities but often the former is included within the
latter.
In a formal sense a diagnosis is a short statement that absolutely isolates a species (or a taxon of
any rank). It consists usually of the autapomorphy or autapomorphies that are unique properties of
the taxon and of none other in the group under investigation.
A description is more general and attempts to give an outline of the salient features of the
species, to give the reader an overview of its characterisation, regardless of whether the individual
characters are apomorphic or plesiomorphic. Having said that, a description within a large taxonomic
study should be concise and accurate. There is no need to repeat characters of significance at genus
rank as these are common to all species in the genus. Those characters that diagnose the genus are
usually presented before the part of the work that deals with the species, so that the reader can
quickly see what characters the researcher considers common to all the species in the study.
Similarly, if the various species can be arranged into groups which share one or a number of critical
characters (species groups), then those characters can be abstracted and listed separately as common
to the whole group and need not be repeated in each species description. As well as being
informative of how species fall into groups this is also a useful space saving device.
For instance, in our imaginary genus Xus only two species have 12-segmented antennae, all the
rest have 11. There happen to be other consistent characters in our genus, associated with antennal
segment count, that can be used to split the genus into two distinct groups.
Xus aus group

Antenna with 11 segments.
Maxillary palp with 6 segments.
Mandibles unsculptured.
Setae on vertex of head simple.
Included species: aus, cus, dus, fus.
Xus bus group

Antenna with 12 segments.
Maxillary palp with 4 segments.
Mandibles sculptured.
Setae on vertex of head plumose.
Included species: bus, eus.
The groups as stated convey only taxonomic information relating to identity; they do not give
any data with direct phylogenetic value. However, if the polarity of some or all of these characters
can be ascertained, then monophyly of the groups can be established and the apomorphies
incorporated into any phylogenetic analysis deemed necessary.
After characters common to the genus, and those common to various species groups have been
abstracted, what remains in each taxon file constitutes the basic skeleton of the species description.
This information is often sufficient by itself, but if not the author should be prepared to add whatever
features and comments are deemed necessary to give a good representation of the species.
When the formal description of a species has been completed it is always useful to add a few
lines or a short paragraph in which the species is compared to and contrasted with its closest
relatives. The reader can use this as a quick reference to what the author considers to be the most
important points for differentiation of the species.
Discussions
Beyond the description it is up to the author to decide what other information should be
included, but any doubts that remain about the sole identity of any species should be expressed. For
instance, it may sometimes be suspected that a taxon consists of two sibling species that cannot be
differentiated absolutely in the current treatment. Suspicions of this sort usually arise when there is
greater than normal variation in one or more characters but where insufficient material is available to
decide the significance of the variation. Note when this happens and discuss the reasons so that the
potential pair can be investigated later by molecular or other techniques, if necessary. Also, it may
be deemed necessary to discuss any new synonymy that has occurred, there may be publications
dealing with aspects of behaviour, ecology, etc., that the author judges necessary for discussion, or
there may be reasons to debate aspects of the species as it was treated by previous authors.
Material examined
At the conclusion of the description and discussion of each species a list of all the material seen
during the course of the study should be appended. The amount of information and degree of
accuracy of geography depends upon how important the author considers this aspect to be. At the
very least it should give a fair indication of the known range of the species, but geographical overkill
is not recommended. It is not necessary to list all the localities of the specimens as well as recording
their latitudes and longitudes and also drawing up a detailed distribution map as well. This is too
much information; or rather, it is repeating the same information too often.
Illustrations
The illustrations prepared, whether drawings, light photographs, or electron microscope

pictures, should be sufficient to encompass all the important aspects of the group in question. They
may be produced gradually during the course of the study or done as a single exercise when the text
is more or less complete, or by a combination of both. A gradual accretion of illustrations as the
study progresses allows the selection of the most essential and best illustrations at the end of the
survey. On the other hand doing all the illustrations after the text is complete allows for more
accurate selection of what is essential, but depends upon the necessary specimens still being to hand.
There is usually no need to illustrate every character or every taxon, though sometimes some
species will require more illustration than others. The number of illustrations needed per species or
per group depends roughly upon the difficulty of describing the characters of the group and the
degree to which characters are shared among the various species. In species where variation in shape
of body parts is critical to recognition it will obviously be more valuable to include illustrations of
the different shapes rather than to attempt their descriptions in words. On the other hand in taxa
where there are very conspicuous differentiating characters that can easily be described in a few
words, large numbers of illustrations are unnecessary.
Illustrations should be grouped to provide best access for a reader. Sometimes it is best to group
all illustrations in a single place (at the end of the key, at the end of the text, etc.) but other times it
may be deemed best to space them out (at the end of each species group, at the end of the fauna of a
single zoogeographical region, etc.). In some instances they may be included in the key as marginal
illustrations, but they should never interrupt key couplets or the flow of the text. Illustrations in the
margins of keys may be useful but they have the disadvantage of considerably narrowing the page
and thus reducing the width available for the key couplets.
Final points
When everything has been completed, review the manuscript for content, layout and spelling,
then apply these questions:
1 Is the revision an advance over any previous system?

2 Can the new system be followed and understood by other taxonomists?
3 Are the taxa adequately defined and differentiated so that their identities can be discerned
by others, who may not necessarily be taxonomists?
4 Is the system adequately presented, so that its findings can be duplicated, expanded or
altered by later taxonomists?
5 Does it satisfy normal taxonomic and nomenclatural procedure?
If the answer to all these questions is in the affirmative then the final step is publication of the
results.
APPENDIX: TAXONOMIC AND SCIENTIFIC COMMON SENSE
Taxon inflation
Taxon inflation is a possible pitfall in all taxonomic studies but is particularly one that afflicts
the inexperienced, or those working on higher taxa for the first time. It occurs when the student
spends all available time intensively studying a single group so that its characters, particularly its
differentiating characters, assume an importance that is out of proportion to those of the remainder of
its group (tribe, subfamily, family, etc.). The result is usually that the student promotes all his or her
taxa by one or more ranks. Thus subgenera become genera, genera become tribes, subfamilies
become families, and so on. The cause of this is generally lack of broader knowledge and experience
of groups that are closely related to that being studied. The result of it is that all taxa within the
inflated group end up out of proportion with their relatives in equivalent taxa.
To illustrate this, imagine that the ant fauna is being studied in a very small area, for example
the garden of an ordinary house. Two species are found in the soil, one of which is yellow and has
small eyes, the other of which is black and has large eyes. In the context of the garden alone these
two taxa can be assigned to any rank as they are being compared only to each other, not to the fauna
of the rest of the world. When viewed with a whole world perspective the characters that seemed so
important in the restricted area are discovered to be of little relevance whilst other characters,
apparently inconsequential in the small area, are constant and universal through all the species of all
the world: the two apparently so different taxa turn out to be two species that belong in the same
genus.
“Common names”
Back before 1758 Linnaeus had a remarkable insight, a flash of genius. He realized that any
living thing could be named accurately by the use of just two words. The two words, the first
representing a genus and the second a species, represent the unique proper name of a species that is
universally recognizable. And as they are pitched in a language that is extinct, no living language is
given priority over any other. It is a marvelous thing to contemplate: the name of any insect species
in the whole world, and there are now about a million of them, can be recognized by anyone. A
native English speaker can say the name of a species to a native speaker of Japanese and both
immediately understand what the species is. Set against this incredible achievement and its universal
application is the misplaced effort of formulating “common” names for species that do not need them
and are often anything but common. These “common” names are often formulated by a single
individual and are only understood in one language, and sometimes only in part of the range of one
language. Obviously, this condemnation does not apply to names that have evolved naturally for
large species in any language because such species feature in the lives of the whole population, for
instance oak, cow, dog in English. But deliberately to sit down and make up “common” names for
ant species is an exercise in pointlessness and futility.
ACKNOWLEDGEMENTS
My thanks and gratitude to Roy R. Snelling, Philip S. Ward and an unknown number of anonymous
reviewers, who suggested useful modifications to the first few drafts of this essay.
Brandão, C. R. F., Mayhé-Nunes, A. J. 2007. A phylogenetic hypothesis for the Trachymyrmex species groups, and the
transition from fungus-growing to leaf-cutting in the Attini, pp. 72-88. In Snelling, R. R., B. L. Fisher, and P. S. Ward
(eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions.
Memoirs of the American Entomological Institute, 80.
A PHYLOGENETIC HYPOTHESIS FOR THE TRACHYMYRMEX SPECIES GROUPS,

AND THE TRANSITION FROM FUNGUS-GROWING TO LEAF-CUTTING IN THE
ATTINI
C. Roberto F. Brandão
Museu de Zoologia,
Universidade de São Paulo,
Av. Nazaré 481, São Paulo, SP, 04263-000, Brazil
crfbrand@usp.br
and
Antonio J. Mayhé-Nunes
Departamento de Biologia Animal,
Instituto de Biologia, Universidade Federal Rural do Rio de Janeiro,
Seropédica, RJ, 23890-000, Brazil
amayhe@ufrrj.br
Abstract
As part of a series of papers dealing with an ongoing revision of Trachymyrmex (Myrmicinae:

Attini) ant species, we study here the relationships of its species. The main objective of the series
is to reevaluate the taxonomic status of the described taxa and to describe new species. A
revision of the Opulentus group is the first published paper in the series, followed by a revision
of the Iheringi group. At the beginning of our work, we accepted the species groups as proposed
by authors, but now decided to test this proposal, to avoid major rearrangements in the future.
Our parsimony analysis using 50 characters from external worker morphology and 32 taxa
(Mycetarotes parallelus as outgroup) indicates congruence with unpublished notions of
relationships within Trachymyrmex, but some important differences as well, and may help to
understand the transition from fungus-growing to leaf-cutting in Attini ants.
Key words: Hymenoptera, Formicidae, Attini, Trachymyrmex, fungus-growing ants, leaf-cutting

ants, Neotropics, phylogeny
Brandão and Mayhé-Nunes: Phylogeny of Trachymyrmex 73
Introduction
Reliance on fungi as the dominant food source evolved only once in ants, in the fungus-growing
Attini (Myrmicinae), that includes species necessarily dependent on the cultivation of a
mutualistic fungus to feed adults and larvae (Mueller et al., 2001). Members of the genus
Megalomyrmex of the silvestrii group, by contrast, are facultative social parasites of attines,
consuming fungus grown by their hosts or the hosts’ larvae when living inside attine nests
(Brandão, 2003).
Until recently, the monophyly of the Attini was supported by their exclusive primary ability
to cultivate a basidiomycete fungus in nearly axenic gardens (Mayhé-Nunes, 1995) that,
depending on the ant species, are manured with a variety of fresh and dead plant matter, and by
three morphological characters of the prepupae (Schultz & Meier, 1995): short, narrow labrum;
fleshy, nearly straight, subconical mandibles; and presence of leg vestiges as open slits in the
integument. Brandão & Mayhé Nunes (2001) described the first adult female autapomorphy for
the tribe: the median clypeal seta in all attines originates from the anteclypeus or at the meeting
of the anteclypeus and the clypeal border itself. Thirteen genera with over 210 species are now
included in Attini. The Sri Lankan genus Proatta, suggested by Moffett (1987) to be possibly an
attine, is now placed in Stenammini (Bolton, 2003).
The origin of the attine ant-fungus mutualism has received much attention (see for instance
North et al., 1997). Mueller (2002) suggested that the Attini arose at the beginning of the
Tertiary, some 45-65 Mya, bracketing the estimate of 50 Mya by Wilson (1971). Mueller et al.
(2001) summarized the hypotheses for the successive evolution of behavioral elements of attine
fungiculture in two main models: 1. the traditional “consumption first” model, in which initial
stage fungi growing accidentally in ant nests became part of their diet; subsequently the ants
evolved the ability to promote fungal growth, and finally the ants evolved a mechanism to
transmit fungi between parent and offspring nests, or 2. the alternative “transmission first”
model, where in the initial stage the ants did not feed on fungi, but instead were used by the
fungi for dispersion; subsequently the ants incorporate the fungus in their diet, and finally
evolved the ability to cultivate and transmit the symbiotic fungus. The exact nature of the
ancestral attine-fungus association is difficult to infer, given the long time elapsed and intensive
taxon diversification since the origin of the Attini (Mueller, 2002). Also there exist no extant
facultative attine-fungus stages of ant-fungus interaction, suggesting transience and instability of
non-obligate ant fungus associations and possibly a rapid evolutionary transition from a hunter-
gatherer ancestral ant to the derived fungicultural ant.
Most attine fungal cultivars belong to Leucoagaricus and Leucocoprinus (Agaricales:
Basidiomycota: Lepiotaceae: Leucocoprinae), except for some Apterostigma ants that
secondarily switched to Tricholomataceae (although basal Apterostigma cultivate lepiotaceous
fungi), and for a subgroup of the rimosus species group of Cyphomyrmex that secondarily
culture their leucocoprineous fungus as yeasts and not as mycelia. Leucocoprineous fungi are
common litter decomposers in the moist Neotropics, and have been probably domesticated by
the ants repeatedly and independently from free-living populations. Attine fungi are also
transferred laterally across nests of even different genera (Adams et al., 2000a; Adams et al.,
2000b). Moreover, attine ants combat outbreaks of the potentially devastating microfungal
parasite Escovopsis using antimicrobial pesticides produced by Streptomyces bacteria they grow
on specialized regions of their own bodies (Currie et al., 2003). Escovopsis phylogeny provides
evidence for lateral transfer of the parasite between closely related species. Currie et al. (2003)
concluded that Escovopsis parasitism of the ant-microbe symbiosis likely originated with a
parasite of free-living leucocoprineous fungi that invaded the symbiosis along with the
domestication of these free-living fungi.
The Attini genera were informally divided by Kusnezov (1964) into two groups: the
Paleoattini (also called “apterostigmoid” clade by Schultz & Meier, 1995), encompassing
Apterostigma, Myrmicocrypta, and Mycocepurus (but see Schultz & Meier, 1995 for apparent
paraphyly of Myrmicocrypta in relation to these lineages), and the Neoattini (correspondingly,
the “attoid” clade), for all other genera, including Sericomyrmex, Trachymyrmex and the
polymorphic leaf-cutter ants Acromyrmex and Atta.
Lower attines and Trachymyrmex and Sericomyrmex in the higher attines culture their
fungus with the type of substrate for which free-living leucocoprineous fungi are specialized:
plant detritus, flower parts, seeds and arthropod feces. Atta and Acromyrmex manure their
gardens with fresh leaf material. The fungi cultivated by the lower attines are polyphyletic,
probably because of the common horizontal transmission and repeated domestication from free
living cultivars. Chapela et al. (1994) showed that the fungus cultivated by higher attines is
monophyletic, probably because it is vertically transferred, hence clonally, to new colonies.
Also, mature cultivars of the higher attines (although literature is not clear on cultivars manured
by basal Trachymyrmex and Sericomyrmex species) produce conspicuous nodules called
staphylae, that are clusters of vacuolized hyphal-tip swellings rich in glycogen (gongylidia)
(Hölldobler & Wilson, 1990). Stradling and Powell (1986) demonstrated that the fungal
symbiont of Atta and Acromyrmex grows faster than those of Sericomyrmex and Trachymyrmex
and is therefore able to support larger colonies.
Trachymyrmex is a key taxon to the understanding of the phylogenetic relationships among
the derived Attini. Contrary to the lower attine colonies that number only a few dozen workers,
some Trachymyrmex and Sericomyrmex species live in colonies that can have up to a few
thousand workers; in the higher attines Acromyrmex colony sizes may reach several hundred
thousands or even 5-10 million workers in Atta. Also, a sound phylogenetic framework for
Trachymyrmex may help us to understand the most important transition in attine evolution once
their ancestral forms learned how to cultivate fungus, that is, the acquisition of a further
specialization of leaf-cutter ants to manure a derived monophyletic staphylae producing cultivar,
called Group 1 fungus group by Chapela et al. (1994).
Trachymyrmex is probably the most derived monomorphic attine ant, with some species
showing weak polymorphism (in the sense commonly applied to social insects). Trachymyrmex
species are widespread in the Neotropical region, with some forms exclusively found in the
Nearctic (Kusnezov, 1964; Chapela et al., 1994; Schultz & Meier, 1995; Mayhé-Nunes & Jaffé,
1998). Although abundant in most habitats, most Trachymyrmex species are small and
inconspicuous foragers on the floor of forests, harvesting opportunistically a wide variety of
items used to cultivate symbiotic fungus, including insect frass, plant debris and occasionally
freshly cut parts of plants (but see Waller, 1989 for T. turrifex). Trachymyrmex diversus actively
cuts and carries fresh leaves, at least in the laboratory (Schultz & Meier, 1995). Most
Trachymyrmex workers feign death when disturbed, with the known exception of T. diversus
(Schultz & Meier, 1995), and are, therefore, easily overlooked on the litter; yet their small to
medium sized nests built in the soil are usually conspicuous, because their entrance is normally
adorned by tower- or crater-shaped earth mounds (Bhatkar, 1974; Tschinkel & Bhatkar, 1974).
Trachymyrmex species can collect and disperse seeds, as is the case with different Brazilian
Atlantic Forest species that gather seeds of the common tree Miconia cabucu (Medeiros &
Morretes, 1994). The territorial and nocturnal foragers of T. urichi employ chemical
communication, laying a trail pheromone released from the poison gland whose concentration
regulates the number of workers recruited; nests have a specific odor used in nestmate
recognition; and workers secrete a defensive secretion that induces nestmates to attack any
object with it, even their gyne (Jaffé & Villegas, 1985). Trachymyrmex turrifex individuals show
little or no fidelity to food type or foraging routes (Waller, 1989). According to Gallardo (1916),
T. pruinosus used pieces of orange fruit as substrate to fungus garden in an artificial nest
maintained in laboratory. Weber (1945) also has observed workers of T. cornetzi gathering
orange rind and carrying it to the nest in the field. Torres et al. (1999) observed T. jamaicensis
utilizing a variety of fruits as the substrate for their fungal gardens. T. holmgreni collects fresh
leaves of the grass Paspalum ancylocarpum to such a degree that it has been considered an
agricultural pest (Gonçalves, 1975). Unidentified species of Trachymyrmex, and T. fuscus, have
also been considered agricultural pests in Eucalyptus plantations in Brazil, due to their relatively
large populations and relatively high density of nests in these agroecosystems (Oliveira et al.,
1998). More recently, however, Araujo et al. (2002) have observed that workers of T. fuscus
transport mainly dry vegetation to the nest. Beshers & Traniello (1994) studied the adaptiveness
of worker demography in two populations of T. septentrionalis. Army ants are important
predators of Trachymyrmex brood, and probably also of most attine ant species (LaPolla et al.,
2002).
Weber (1972) reported that Trachymyrmex zeteki accepted and ate (in laboratory conditions)
pieces of fungus taken from nests of other higher attine species, but these ants did not eat the T.
zeteki fungus. Sericomyrmex amabilis ate the fungus of most of these ants, but only two Atta
species tested ate the fungus of S. amabilis. This may indicate that Acromyrmex and Atta ants
and cultivars share characters still more derived in relation to other higher attines and their fungi.
Hinkle et al. (1994) suggest moreover that cultivars of Sericomyrmex and Trachymyrmex are
different from those cultivated by Atta and Acromyrmex, and that they are monophyletic. Based
on nuclear DNA sequence data, Escovopsis parasites living in Trachymyrmex nests also
represent a specific lineage (Currie et al. 2003).
The series of revisionary studies we are publishing aim to reevaluate the taxonomic status of
described infraspecies of Trachymyrmex and to describe new species. Existing revisionary
studies are partial and outdated. When we started the series of revisionary works on
Trachymyrmex, we decided to accept the proposal of species groups in drafts we found in the
Walter W. Kempf personal library, reserving a new classification, based on cladistic approach,
to the conclusion the series, when we would be ideally more acquainted with the limits of the
taxa within Trachymyrmex. However, as our studies proceeded, it became clear that testing
Kempf’s proposal is now necessary to avoid large taxonomic rearrangements in the future.
Prior to the present work, relationship proposals for Trachymyrmex species were mostly
based on opinions of describers, and had not been subjected to any formal analysis. However,
most authors recognize roughly the same species groups (judging from the comments that follow
the species descriptions): Opulentus, Iheringi, Urichi and Septentrionalis groups, with some
dispute over the Cornetzi and Farinosus groups, either as independent groups or subgroups of,
respectively, Septentrionalis and Urichi groups. Figure 1 summarizes Kempf’s ideas on the
relationships among Trachymyrmex species groups as interpreted from his drafts. Interestingly,
if this proposal proves to be right, the most derived clade includes species in Urichi and
Septentrionalis groups, that also show the most conspicuous differences in nestmate gyne and
workers, the largest size range among worker nestmates, and larger colonies in relation to other
Trachymyrmex.
The most dramatic evolutionary step in Attini history, after the origin of the fungus-ant
symbiosis, was the acquisition of staphylae rich monophyletic cultivars by the higher Attini (the
“attoid” clade), in particular Atta and Acromyrmex that collect only fresh vegetation as a novel
form of fungal substrate. Converting vegetation to substrate requires a series of operations and
specializations. Vegetation is cut by workers with head width 1.6 mm or greater (other tasks,
such as defense, involve workers of even greater size), while the care of the fungus requires very
small workers; intervening steps in gardening are conducted by workers of graded intermediate
size. We know very little about division of labor in the lower or basal higher attines. Atta and
Acromyrmex, however, show one of the most complicated caste systems found in ants
(Hölldobler & Wilson, 1990), although their division of labor is based on an enormous and
continuous size variation and alloethism, instead of allometry. Beshers & Traniello (1994) have
shown how variation in size can evolve in the context of life history strategies and how this
variation may serve as a preadaptation for division of labor.
On the ant side, this major evolutionary change seems to have put the higher Attini on a
course resulting in much larger colonies, by the increase in size differences between gynes and
daughter workers, and hence increase in ovariole counts, allowing rapid colony growth, and by
the increase in size variation among worker nestmates, able to execute all activities involved in
fungus cultivation. Atta species represent an extreme in this tendency because their single queen
per nest is huge in relation to the workers (even the extremely large majors), with ovaries
composed of hundreds of ovarioles that individually produce relatively more eggs than do
ovarioles of lower attine gynes (Weber, 1972), resulting in mature colonies that can attain
millions of individuals. The question is when in attine evolution these characters started to
change. There are no Atta or Acromyrmex species showing incipient stages in worker size
variation, while Trachymyrmex species commonly show a variation in these features. A study on
the evolution of polymorphism in basal “attoids” within a phylogenetic framework may be
therefore an interesting way to understand the origin of the world’s most powerful herbivores.
Wilson (1987) studied the then known Cretaceous ant species, concluding that females
appear to have differentiated as queen and worker castes belonging to the same colonial species,
instead of winged and wingless solitary females belonging to different species. He based his
study on the fact that gasters of workers of modern and extinct Miocene ant species are smaller
relative to the rest of the body than is the case for modern wingless solitary wasps. The wingless
Cretaceous formicoids conform to the proportion of ant workers rather than to those of wasps,
and hence are reasonably interpreted to have lived in colonies.
Inspired by Wilson’s (1987) work, we studied size differences between reproductive and
sterile females and size ranges among workers of different Trachymyrmex species and species
groups, superimposed on a new phylogenetic hypothesis based on a parsimony analysis of
worker external morphological characters.
SUMMARY OF TAXONOMIC HISTORY
Trachymyrmex was proposed by Forel (1893) as a subgenus of Atta Fabricius, when he

described and compared T. urichi with two species originally described in Atta (Acromyrmex)
and Oecodoma, respectively T. saussurei (Forel) and T. tardigrada (Buckley); the latter now is
considered unrecognizable and incertae sedis in Atta (Bolton, 1995). Two other species now
included in Trachymyrmex, T. jheringi (Emery) and T. jamaicensis (André), were also described
as Atta (Acromyrmex). After 1893, the subgenus Atta (Trachymyrmex) was adopted almost
universally, but surprisingly Forel (1914a, b) described two Atta species in Acromyrmex
(Trachymyrmex) and Bruch (1921) proposed Acromyrmex (T.) tucumanus cordovanus, ignoring
the raising of Trachymyrmex to generic rank by Wheeler (1916). In the same year, Santschi
(1916) described T. fiebrigi, considering Trachymyrmex as a subgenus of Cyphomyrmex,
following Emery´s (1913) ideas. Mann (1922) proposed Myrmicocrypta cucumis, now a junior
synonym of T. bugnioni, and T. opulentus in Sericomyrmex; Weber (1937) also described a
species in the latter genus, later recognized as Trachymyrmex. From 1930 on, a relatively large
number of taxa were described in Trachymyrmex, making it today the second richest genus in the
Attini; exceeded only by Acromyrmex. The striking morphological resemblance between some
Trachymyrmex and other attines can explain, however, descriptions of species now accepted in
Trachymyrmex in different taxa, supporting also the impression shared by several
myrmecologists that Trachymyrmex may not be a natural group (see comments by Schultz &
Meier, 1995 and references). Notwithstanding, reproductive and sterile females of all
Trachymyrmex share the presence of microtuberculated mesosomal projections, an exclusive and
hence apomorphic trait of the genus (Mayhé-Nunes & Brandão, 2002)—a character already
noticed by Forel when he choose the name (from Greek, trachis, coarse), although overlooked in
the recent literature (but see Gallardo, 1916 comments on T. pruinosus Emery).
MATERIALS AND METHODS
We measured head width in frontal view and the largest diameter of gasters in dorsal view of
workers and dealated gynes of all Trachymyrmex species for which we found gynes and workers
on the same pin in the collection of the Museu de Zoologia, Universidade de São Paulo, that is
collected at the same time, hence probably from the same colony, and probably representing
gynes not founding new colonies at the time of their collection, but instead from mature
colonies. In so doing, we could also study size distributions among workers of different
Trachymyrmex species.
We found 41 pairs of Trachymyrmex gynes and workers on the same pin. We list below the
species from which we took measurements and to which group we consider them to belong:
Septentrionalis group: T. septentrionalis (6 pairs), T. diversus (one pair), T. smithi (4 pairs),

and T. arizonensis (2 pairs);
Cornetzi (as subgroup of Septentrionalis): T. cornetzi (one pair), T. bugnioni (one pair), and
T. levis (one pair);
Iheringi group: T. holmgreni (one pair), and T. kempfi (7 pairs);
Opulentus group: T. relictus (5 pairs), and T. opulentus (one pair);
Urichi group (includes the Farinosus subgroup): T. urichi (one pair), T. farinosus (one pair),
T. ruthae (one pair), T. oetkeri (one pair), and T. turrifex (8 pairs).
The measurements were submitted to regression analysis by caste, by species, and by

species groups.
PHYLOGENETIC ANALYSIS
The character matrix (Table 2, Appendix) for 50 external morphological worker characters
compiled from 32 attine species, including Mycetarotes parallelus (Emery) as the outgroup, was
subjected to unweighted parsimony analysis using the computer programs NONA (Goloboff,
1993) and WinClada version 1.00.08 (Nixon, 2002). Characters were treated as unordered. The
data matrix was generated with the NEXUS Data Editor version 0.5.0 (Page, 2001). We searched
for trees using the following NONA parameters: hold 1000 (maximum trees to keep), mult*N
100 (number of replications), hold/ 5 (starting trees per rep), with mult*max* (multiple
TBR+TBR) as search strategy; the strict consensus was calculated for the obtained trees.
Table 1. List of worker morphological characters and character coding for the genus
Trachymyrmex.
1. Pilosity of the gaster and femora: (0) only hairs, without fine pubescence; (1) dense, long
dark hairs mixed with an extremely low, fine and abundant light pubescence.
2. Discal area of mandibles: (0) smooth; (1) finely striated; (2) coarsely striated.
3. Lateral borders of frontal lobes: (0) semicircular; (1) sub-triangular; (2) triangular.
4. Accessory lobes or teeth at the base of frontal lobes: (0) absent; (1) present.
5. Anterior border of frontal lobes: (0) straight; (1) convex; (2) concave.
6. Posterior border of frontal lobes: (0) straight; (1) convex; (2) concave.
7. Lateral borders of frontal lobes: (0) smooth; (1) crenulated.
8. Frontal carina: (0) not reaching the preoccipital (posterior) margin of head; (1) reaching the
preoccipital margin.
9. Frontal and preocular carina: (0) ending separated; (1) ending together.
10. Preocular carina: (0) strongly curved; (1) vertical.
11. Preocular carina: (0) unique; (1) double.
12. Scrobe: (0) absent; (1) present.

13. Scrobe: (0) opened; (1) closed.
14. Apical tubercle of the antennal scrobes: (0) absent; (1) present.
15. Basal lobes of the antennal scapes: (0) absent; (1) present.
16. Basal lobes of antennal scapes: (0) not transversely broadened; (1) transversely broadened
pointing mesad (2) transversely broadened pointing to both sides.
17. Anterior surface of antennal scapes: (0) smooth; (1) weakly microtuberculate; (2) notably
microtuberculate.
18. Length of antennal scapes: (0) surpassing weakly the posterolateral corners of head; (1)
surpassing notably the posterolateral corners; (2) not surpassing the posterolateral corners.
19. Preoccipital (posterior) margin of head: (0) distinctly notched; (1) almost straight.
20. Preoccipital spines: (0) larger than the tubercles of the preoccipital lobes; (1) nearly the
length of the tubercles of the preoccipital lobes; (2) absent or vestigial.
21. Supraocular projections: (0) absent or vestigial; (1) present.
22. Supraocular projections: (0) tooth or spine-like; (1) microtuberculate swelling.
23. Dorsal projections of mesosoma: (0) smooth spine- or tooth-like; (1) microscopically
multituberculate swelling, tooth- or spine-like.
24. Median pronotal projections: (0) present; (1) absent or vestigial.
25. Number of median pronotal projections: (0) two; (1) one.
26. Lateral pronotal projections: (0) spine-like; (1) tooth-like; (2) microscopically
multituberculate or multidentate swelling.
27. Lateral pronotal projections in frontal view: (0) directed upwards and outwards; (1) laterally
directed.
28. Inferior pronotal corner: (0) unarmed, obtusely angulate or rounded; (1) armed with a tooth
or spine.
29. Size of projection on the inferior pronotal corner: (0) weakly projected; (1) notably
projected.
30. Shape of projection on the inferior pronotal corner: (0) triangular; (1) spine-like; (2)
rounded.
31. Anterior mesonotal projections: (0) nearly of the length of the pronotal lateral ones; (1)
notably shorter than pronotal lateral ones; (2) notably longer than pronotal lateral ones.
32. Apex of projection on the inferior pronotal corner: (0) blunt; (1) acute.
33. Anterior mesonotal projections: (0) spine-like; (1) microscopically multituberculate or
multidentate swelling; (2) semicircular multidentate ridge from above.
34. Shape of median pronotal projections: (0) spine-like; (1) ridge or multituberculate swelling.
35. Posterior mesonotal projections: (0) present; (1) absent or vestigial.
36. Shape of posterior mesonotal projections: (0) spine-like; (1) ridge or multituberculate
tumulus.
37. Pilosity of mesopleura: (0) vestigial or absent; (1) present.
38. Projection on the inferior margin of mesopleura: (0) absent; (1) present.
39. Projection on the superior margin of mesopleura: (0) absent; (1) present.
40. Shape of projection on the superior margin of mesopleura: (0) small tooth or triangular
spine; (1) large lobe.
41. Projections at the meeting of basal and declivous faces of propodeum: (0) as long as the
larger projections of promesonotum; (1) shorter than promesonotal projections; (2) longer
than promesonotal projections.
42. Projections at the meeting of basal and declivous faces of propodeum: (0) as long as the
projections of basal face; (1) longer than the projections of basal face; (2) tooth-like, nearly
of the length of pronotal lateral ones.
43. Petiolar node: (0) unarmed; (1) with a pair of teeth; (2) with two pairs of teeth.
44. Petiolar node from above: (0) longer than broad; (1) as long as broad.
45. Postpetiole from above: (0) distinctly transverse; (1) as long as broad.
46. Posterior border of postpetiole: (0) straight; (1) superficially excised; (2) notably excised.
47. Hairs of the first gastric tergite: (0) strongly curved, hook-like; (1) in two shapes: straight or
weakly curved medially and hook-like; (2) straight.
48. Tubercles on the basal third of the first gastric tergite: (0) clearly in four longitudinal rows;
(1) more or less in four longitudinal rows; (2) randomly distributed; (3) inconspicuous, but
with notable ridge on each side.
49. Large welt on each side of lateral posterior region of the first gastric tergite: (0) absent; (1)
present.
50. First gastric sternite: (0) without hook-like hairs; (1) with hook-like hairs; (2) in two shapes:
straight or weakly curved medially and hook-like.
RESULTS
The tree presented in Fig. 3 is the strict consensus (L 323; CI 21; RI 38) of the 68 equally
parsimonious trees (L 259; CI 27; RI 54) resulting from the analysis, showing 15 unresolved
terminal branches, but keeping the Iheringi and “Cornetzi” species groups, and the clades
jamaicensis-zeteki and arizonensis-nogalensis. One of the 68 trees is presented in Fig. 2, as it
shows the species groups in Trachymyrmex proposed by Kempf in his manuscripts (Fig. 1), with
the following differences:
1. Urichi group: In our analysis it includes T. phaleratus, that Kempf included in the
Septentrionalis species group. By contrast, T. oetkeri appeared in our analysis out of this group,
where it has been traditionally included. Two character states are at the base of the Urichi clade:
8.1. frontal carinae reaching the preoccipital margin and 17.1. anterior scape surface weakly
microtuberculate. However, 8.1 reverts to 8.0 in T. ruthae and in the clade T. agudensis – T.
isthmicus; in T. jamaicensis character 17 appears in state 2. In this particular tree, these
characters are thus not synapomorphic for the group and generate homoplasies within the group.
2. Septentrionalis group: The major problem here is the lack of sound characters at the base
of the largest clade in the T. septentrionalis – T. levis branch (with nine species) and the fact that
T. mandibularis and T. saussurei appeared outside the species group. As said before, in our
analysis T. phaleratus is nested within the Urichi group.
3. We based our analysis on worker characters only, because we did not have enough
information on sexuals and/or immatures. However, Schultz & Meier (1995) studied larval
characters and found an apparent apomorphy shared by members of the Cornetzi group they
studied, T. bugnioni and T. cornetzi, supporting this clade. Only larvae of these species are
entirely devoid of hairs on the ventral region of the body. These two species are also the only
ones among Trachymyrmex larvae Schultz & Meier (1995) studied having only two subantennal
(genal) setae.
4. Trachymyrmex relictus does not belong to the Opulentus species group clade in the
consensus, but in 60 of the 68 trees (88%) it is nested within the species group we assigned.
DISCUSSION
As far as informed by external characters of workers, Trachymyrmex seems to be a

monophyletic taxon, including species that share unique microtuberculate mesosomal
projections.
We found only four characters that can clearly differentiate monophyletic groups within
Trachymyrmex. The Opulentus group females present pilosity interspaced with fine decumbent
pubescence, more clearly visible at the hind femora (Mayhé-Nunes & Brandão, 2002). Female
specimens in the Iheringi group (Mayhé-Nunes & Brandão, 2005) are easily recognized by the
presence of a lobe at the base of the antennal scapes.
We were not able to find synapomorphies to support the two larger species groups, Urichi
and Septentrionalis, and have also doubts of an Arizonensis subgroup, but we did not include all
species of these groups in our analysis. Apparently, females in the Urichi group have the
posterolateral corners of the head angulate, while in the Septentrionalis group these corners are
rounded. In each of these two large groups, we identified two possible subgroups: Farinosus
with exclusive and very conspicuous posterolateral gastral protuberances and Cornetzi with low
promesonotal projections. However, our study supports a Cornetzi group, as it consistently
showed a clade formed by T. cornetzi, T. bugnioni, T. imgardae and T. levis. Also, all trees show
a clade formed by T. arizonensis and T. nogalensis, traditionally assigned, however, to the
Septentrionalis group. The same is true for the clade T. jamaicensis and T. zeteki, assigned by
most authors to the Urichi group.
When we superimpose the data on size differences between gynes and workers of
Trachymyrmex on our preferred tree (Fig. 4), members of the Septentrionalis group show the
greatest differences. Trachymyrmex species reported as agricultural pests also belong to this
species group, possibly meaning that mated queens can produce more workers in a shorter time
and that their relatively larger gasters support more and/or more productive ovarioles.
Members of the Septentrionalis and Urichi groups show also the largest size range among
workers. Our proposal suggests the Septentrionalis group as the basal Trachymyrmex group.
However, our morphological analysis indicates a derived position for species in this group. In
any topology, Urichi and Septentrionalis groups never came up together.
Analysis of measurements taken from pairs of nestmate gynes and workers of different
Trachymyrmex species indicates that worker and gynes appear clearly differentiated, with very
little overlap. Although the differences among species, within groups or even between species
(workers and gynes taken separately) are not statistically significant, our results suggest
secondary departures from the strict monomorphic state of basal Trachymyrmex and lower
attines, and moreover, that the increase in size differences between worker and gynes and the
increase in size range among workers occurred independently in the main Trachymyrmex clades.
The two major groups of Trachymyrmex species, Urichi and Sepentrionalis, are mostly
distributed in southern and central-North America, respectively. Of the 12 extant species that
seem to belong to the Septentrionalis group, six are endemic to the Nearctic region (Mayhé-
Nunes, 1995); the others are widespread in northern South America, above parallel 10ºS. The
Dominican amber fossil T. primaevus seems to belong to this group.
The majority of the species of the Urichi group are also found in northern South America,
but two of them occur in Central America (T. isthmicus and T. zeteki) and other two are found
between the parallels 10º-25ºS (T. agudensis and T. fuscus); there is also an endemic Nearctic
species, T. turrifex. However, T. jamaicensis, a species widely distributed in the Caribbean, also
occurs in Florida. The Opulentus group has three species widespread in northern South America,
with a unique species (T. opulentus) found in Central America, and another (T. dichrous) that
seems to be a typical inhabitant of the vast savannas of Central Brazil. Contrary to other groups,
Iheringi is restricted to the southern Neotropical region, although some species are widespread in
South America (as T. holmgreni and T. kempfi), the distribution of the group does not cross the
parallel 10ºS.
The topology of the species groups cladogram is partially congruent with a pattern
explained by vicariant events of speciation. Particularly, by a first split in Nearctic and
Neotropical clades, probably due to repeated breaks between North and South America during
the Cenozoic Era, until the definitive formation of the Isthmus of Panama (3-5 Mya). The close
proximity among the clades T. arizonensis - T. nogalensis and T. cornetzi - T. levis (see also the
consensus in Fig. 3) can indicate shared ancestral lineages from the north of South America that
migrated to Central America, with some of them later established in the Nearctic region. Another
example of probable vicariance is the clade T. jamaicensis - T. zeteki. In spite of our insufficient
knowledge on the geographic distribution of T. zeteki (known only from type-locality, Panama),
it possibly shared an ancestor with T. jamaicensis, that in the past was able to cross the passage
between Yucatan and Cuba. On the other hand, the limited distribution of the Iheringi group is
sufficient to postulate that its component species shared an ancestor that lived in the southern
Neotropical region. In some cladograms T. papulatus is the nearest species to the clade T.
pruinosus - T. tucumanus, although it does not belong to the Iheringi group because it lacks
lobated antennal scapes; it is found only in southern South America (Argentina).
Baroni Urbani (1980) said that some questions could be solved with a reasonable
phylogenetic proposal for Trachymyrmex, and we can, hopefully start to answer them now: The
Miocene-Oligocene Dominican amber fossil T. primaevus does not seem to be the sister species
of recent Antillean Trachymyrmex, so we may assume that the Antilles were colonized more
than once by Trachymyrmex. T. primaevus belongs to a primarily Nearctic clade, but other
Septentrionalis group species occur in the Caribbean islands, strongly suggesting multiple
arrivals of Trachymyrmex in the islands.
Our studies indicate independent origins of polymorphism and rapid colony growth in the
higher attines, or “attoids”, and may help to devise the evolutionary steps and their sequence in
the attine evolution.
CONCLUSIONS
Our phylogenetic analysis of Trachymyrmex is congruent in many ways with relationships

proposed informally by other authors, especially ideas presented in Walter W. Kempf’s
unpublished drafts. Contrary to some opinions, however, Trachymyrex is accepted as
monophyletic, composed of six recognizable species groups. Size differences between nestmate
gynes and workers and size range among nestmate workers of different Trachymyrmex species
depart from monomorphism independently in the two main Trachymyrmex clades, suggesting at
least two origins for rapid colony growth, increased polymorphism, and large colony populations
in the genus.
ACKNOWLEDGMENTS
It is a pleasure and an honor for us to participate in this homage to Edward Wilson. Ed has been
a continuous source of inspiration for all of us, and instrumental in bringing myrmecology to the
scientific center stage. We heartily thank the organizers of the book for the invitation. Rodrigo
M. Feitosa helped with measurements, Rogerio Rosa da Silva with the statistics, and Mirian D.
Marques and Alexandre P. Aguiar with the text.
LITERATURE CITED
Adams, R.M.M., Mueller, U.G., Green, A.M. & Narozniak, J.M. 2000a. Garden sharing and
garden stealing in fungus-growing ants. Naturwissenschaften 87: 491-493.
Adams, R.M.M., Mueller, U.G., Schultz, T.R. & Norden, B. 2000b. Agropredation:
usurpation of attine gardens by Megalomyrmex ants. Naturwissenschaften 87: 549-554.
Araujo, M.S., Della Lucia, T.M.C. & Mayhé-Nunes, A.J. 2002. Caracterização de ninhos e
atividade forrageadora de Trachymyrmex fuscus Emery (Hymenoptera, Formicidae) em
plantio de eucalipto. Revista Brasileira de Zoologia 19: 419-427.
Baroni Urbani, C. 1980. First description of fossil gardening ants. (Amber collection Stuttgart
and Natural History Museum Basel; Hymenoptera: Formicidae. I: Attini). Stuttgarter
Beiträge fur Naturkunde. Series B (Geologie und Paläontologie) 54: 1-13.
Beshers, S.N. & Traniello, J.F.A. 1994. The adaptiveness of worker demography in the attine
ant Trachymyrmex septentrionalis. Ecology 75: 763-775.
Bhatkar, A.P. 1974. Oriented crescentic mounds of a gardening ant Trachymyrmex

septentrionalis. Florida Entomologist 57: 96.
Bolton, B. 1995. A new general catalogue of the ants of the world. Cambridge, Massachussetts:
Harvard University Press, 504 pp.
Brandão, C.R.F. 2003. Further revisionary studies on the ant genus Megalomyrmex Forel
(Hymenoptera: Formicidae: Myrmicinae: Solenopsidini). Papéis Avulsos de Zoologia 43:
145-159.
Brandão, C.R.F. & Mayhé Nunes, A.J. 2001. A new fungus-growing ant genus,
Mycetagroicus, gen. n., with the description of three new species and comments on the
monophyly of the Attini (Hymenoptera: Formicidae). Sociobiology 38: 639-665.
Bruch, C. 1921. Estudios mirmecológicos. Revista del Museo de La Plata 26: 175-211.
Chapela, I.H., Rehner, S.A., Schultz, T.R. & Mueller, U.G. 1994. Evolutionary history of the
symbiosis between fungus-growing ants and their fungi. Science (Washington, D.C.) 266:
1691-1694.
Currie, C.R., Wong, B., Stuart, A.E., Schultz, T.R., Rehner, S.A., Mueller, U.G., Sung, G.-
H., Spatafora, J.W. & Straus, N.A. 2003. Ancient tripartite coevolution in the attine ant-
microbe symbiosis. Science (Washington, D.C.) 299: 386-388.
Emery, C. 1913. Études sur les Myrmicinae. [V-VII]. Annales de la Société Entomologique de
Belgique 57: 250-262.
Forel, A. 1893. Note sur les Attini. Annales de la Société Entomologique de Belgique 37: 586-
607.
Forel, A. 1914a. Einige amerikanische Ameisen. Deutsche Entomologische Zeitschrift 1914:
615-620.
Forel, A. 1914b. Formicides d’Afrique et d’Amérique nouveaux ou peu connus. Bulletin de la
Société Vaudoise des Sciences Naturales 50: 211-288.
Gallardo, A. 1916. Notas acerca de la hormiga Trachymymrex pruinosus Emery. Anales del
Museo Nacional de Historia Natural de Buenos Aires 28: 241-252.
Goloboff, P.A. 1993. NONA version 1.0. San Miguel de Tucumán, Argentina: published by the
author.
Gonçalves, C.R.G. 1975. Formigas dos gêneros Sericomyrmex e Trachymyrmex cortando folhas
verdes de plantas. Abstracts. 27a. Reunião da SBPC.
Hinkle, G., Wetterer, J.K., Schultz, T.R. & Sogin, M.L. 1994. Phylogeny of the attine ant
fungi based on analysis of the small subunit ribosomal RNA gene sequences. Science
(Washington, D.C.) 266: 1695-1697.
Hölldobler, B. & Wilson, E.O. 1990. The ants. Cambridge, Massachusetts: Harvard University
Press, xii + 732 pp.
Jaffe, K. & Villegas, G. 1985. On the communication system of the fungus-growing ant
Trachymyrmex urichi. Insectes Sociaux 32: 257-274.
Kempf, W.W. 1972. Catálogo abreviado das formigas da região Neotropical (Hymenoptera:
Formicidae). Studia Entomologica 15: 3-344.
Kusnezov, N. 1964(“1963”). Zoogeografía de las hormigas de Sudamérica. Acta Zoologica
Lilloana 21: 123-251.
LaPolla, J.S., Mueller, U.G., Seid, M. & Cover, S.P. 2002. Predation by the army ant
Neivamyrmex rugulosus on the fungus-growing ant Trachymyrmex arizonensis. Insectes
Sociaux 49: 241-256.
Mann, W.M. 1922. Ants from Honduras and Guatemala. Proceedings of the United States
Natural Museum 61: 1-54.
Mayhé-Nunes, A.J. 1995. Filogenia de los Attini (Hym., Formicidae): un aporte al

conocimiento de las hormigas fungívoras. Unpublished thesis. Universidad Simón Bolivar,
Caracas, Venezuela.
Mayhé-Nunes, A.J. & Brandão, C.R.F. 2002. Revisionary studies on the attine ant genus
Trachymyrmex Forel. Part 1: Definition of the genus and the Opulentus group
(Hymenoptera: Formicidae). Sociobiology 40: 667-698.
Mayhé-Nunes, A.J. & Brandão, C.R.F. 2005. Revisionary studies on the attine ant genus
Trachymyrmex Forel. Part 2: the Iheringi group (Hymenoptera: Formicidae). Sociobiology
45: 271-305.
Mayhé-Nunes, A.J. & Jaffé, K. 1998. On the biogeography of Attini (Hymenoptera:
Formicidae). Ecotropicos 11: 45-54.
Medeiros, J.D. & Morretes, B.L. 1994. Dispersal of Miconia cabucu seeds by ant
Trachymyrmex spp. Biotemas 7: 124-126.
Mueller, U.G. 2002. Ant versus fungus versus mutualism: ant-cultivar conflict and the
deconstruction of the attine ant-fungus symbiosis. American Naturalist 160(suppl.): 67-98.
Mueller, U.G., Schultz, T.R., Currie, C.R., Adams, R.M.M. & Malloch, D. 2001. The origin
of the attine ant-fungus mutualism. Quarterly Review of Biology 76: 169-197.
Nixon, K.C. 2002. NEXUS Data Editor version 0.5.0. http://taxonomy.zoology.gla.ac.uk/rod/
rod.html.
North, R.D., Jackson, C.W. & Howse, P.E. 1997. Evolutionary aspects of ant-fungus
interactions in leaf-cutting ants. Trends in Ecology and Evolution 12: 386-389.
Oliveira, M.A., Della-Lucia, T.M.C. & Anjos, N. 1998. Ocorrência e densidade de ninhos de
formigas cortadeiras em plantios de eucalipto no sul da Bahia. Revista Brasileira de
Entomologia 42: 17-21.
Page, R.D.M. 2001. WinClada version 1.00.08. http://www.cladistics.com.
Santschi, F. 1916. Formicides sudaméricains nouveaux ou peu connus. Physis 2: 365-399.
Schultz, T.R. & Meier, R. 1995. A phylogenetic analysis of the fungus-growing ants
(Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae.
Systematic Entomology 20: 337-370.
Stradling, D.J. & Powell, R.J. 1986. The cloning of more highly productive fungal strains: a
factor in the speciation of attine ants. Pp. 625-626 in: Eder, J. & Rembold, H. (eds)
Chemistry and biology of social insects. München: Verlag & J. Peperny, xxxv + 757 pp.
Tschinkel, W.R. & Bhatkar, A. 1974. Oriented mound building in the ant, Trachymyrmex
septentrionalis. Environmental Entomology 3: 667-673.
Torres, J.A., Santiago, M. & Salgado, M. 1999. The effects of the fungus-growing ant,
Trachymyrmex jamaicensis, on soil fertility and seed germination in a subtropical dry forest.
Tropical Ecology 40: 237-245.
Waller, D.A. 1989. Foraging behavior of Trachymyrmex turrifex Wheeler (Formicidae: Attini).
Southwestern Naturalist 34: 271-275.
Weber, N.A. 1937. The biology of the fungus-growing ants. Part I. New forms. Revista de
Entomologia 7: 378-409.
Weber, N.A. 1945. The biology of the fungus-growing ants. Part VIII. The Trinidad, B. W. I.,
species. Revista de Entomologia 16: 1-87.
Weber, N.A. 1972. Gardening ants. The attines. Philadelphia: American Philosophical Society.
Wheeler, W.M. 1916. Ants collected in British Guiana by the expedition of the American
Museum of Natural History during 1911. Bulletin of the American Museum of Natural
History 35: 1-14.
Wilson, E.O. 1971. The insect societies. Cambridge, Massachusetts: Harvard University Press, x
+ 548 pp.
Wilson, E.O. 1987. The earliest known ants: an analysis of the Cretaceous species and an
inference concerning their social organization. Paleobiology 13: 44-53.
Figure 1. Sketch of Kempf’s ideas on the relationships among main species groups of
Trachymyrmex Forel. Numbers between parentheses represent respectively described and
undescribed species.
Figure 2. One of the 68 equally parsimonious trees for the data matrix (L 259; CI 27; RI 54).
Numbers in branches correspond to synapomorphies: characters above and state below,
respectively (see list of characters in table 1). CA, Central America; CI, Caribbean islands; NR,
Nearctic region; NS, northern part of South America (above parallel 10ºS); SS, southern part of
South America (below parallel 10ºS).
Figure 3. Strict consensus cladogram for Trachymyrmex species relationships obtained from the
68 equally parsimonious trees, using the data matrix presented in Table 2 in the appendix. L 323;
CI 21; RI 38.
90
80 Septentrionalis
Cornetzi
70 Iheringi
Gaster width
Relictus
60
Urichi
Septentrionalis
50
Cornetzi
Iheringi
40
Relictus
30
Urichi
20
30 35 40 45 50 55 60 65 70 75 80
Head width
Figure 4. Head and gaster widths of nestmate workers (open symbols) and dealated gynes (filled
symbols) of several Trachymyrmex species groups (see Material and Methods for explanation).
Measurements were not transformed into mm.
APPENDIX
Table 2. Data matrix of thirty-one taxa of Trachymyrmex (outgroup Mycetarotes parallelus) and
fifty morphological characters of workers.
1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2
1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5
Mycetarotes 0 1 0 0 0 0 0 1 0 0 0 0 - 1 0 - 0 0 0 0 0 - 0 0 0
T. agudensis 0 0 2 0 0 0 1 0 1 1 1 1 0 1 0 - 1 0 0 0 1 1 1 0 1
T.arizonenensis 0 2 1 0 2 2 0 0 0 0 0 0 - 0 0 - 0 1 1 1 0 - 1 1 0
T. bugnioni 0 2 0 0 2 0 0 0 0 1 0 1 0 0 0 - 0 0 0 1 0 - 1 0 0
T. compactus 1 0 0 0 1 1 1 0 1 1 0 1 0 1 0 - 0 0 0 2 1 1 1 1 -
T. cornetzi 0 0 1 0 0 0 0 0 0 1 0 0 - 0 0 - 0 0 1 1 0 - 1 0 0
T. dichrous 1 0 1 0 0 0 0 0 0 1 0 0 - 0 0 - 0 0 0 2 1 1 1 1 -
T. diversus 0 1 1 0 0 0 0 0 0 0 0 0 - 0 0 - 0 1 1 0 1 1 1 0 0
T. farinosus 0 0 0 1 1 1 1 1 1 1 0 1 1 1 0 - 1 2 0 0 1 0 1 0 1
T. fuscus 0 0 0 1 1 1 1 1 1 1 0 1 1 1 0 - 1 0 0 0 1 0 1 0 1
T. holmgreni 0 1 1 0 0 2 0 0 0 1 0 1 0 0 1 1 2 1 0 0 1 1 1 0 0
T. iheringi 0 1 2 0 2 0 0 0 0 1 0 1 0 0 1 2 2 1 1 0 1 1 1 0 1
T. irmgardae 0 2 1 0 0 0 0 0 0 1 0 1 0 0 0 - 0 0 0 2 0 - 1 1 -
T. isthmicus 0 1 0 1 1 1 1 0 0 1 0 1 0 1 0 - 0 0 0 0 1 0 1 0 0
T. jamaicensis 0 0 0 0 2 1 1 1 0 1 0 1 0 1 0 - 2 1 0 0 1 0 1 0 1
T. levis 0 2 0 0 0 0 0 0 0 1 0 1 0 0 0 - 0 0 1 1 0 - 1 0 0
T. mandibularis 0 2 2 0 0 0 0 0 0 0 0 0 - 0 0 - 0 1 0 0 0 - 1 0 0
T. nogalensis 0 2 2 0 2 0 0 0 0 0 0 0 - 0 0 - 0 1 1 1 0 - 1 1 -
T. oetkeri 0 0 0 0 1 1 0 0 0 1 0 1 0 1 0 - 0 1 1 0 1 0 1 0 1
T. opulentus 1 2 0 0 0 1 1 0 1 1 0 1 0 1 0 - 0 0 0 2 1 0 1 0 1
T. papulatus 0 0 1 0 0 2 0 0 1 1 0 1 0 0 0 - 1 1 0 1 1 0 1 0 1
T. phaleratus 0 1 0 0 1 1 1 1 1 1 0 1 1 0 0 - 1 0 1 0 1 1 1 0 0
T. pruinosus 0 0 2 0 0 0 0 0 0 1 0 1 0 0 1 0 2 1 0 0 1 0 1 0 0
T. relictus 1 1 0 0 1 1 1 0 0 1 0 1 0 1 0 - 0 0 1 2 1 0 1 1 -
T. ruthae 0 0 0 1 1 1 1 0 1 1 0 1 0 1 0 - 1 2 0 0 1 0 1 0 0
T. saussurei 0 2 2 0 0 0 0 0 0 0 0 0 - 0 0 - 0 1 1 1 1 0 1 0 0
T.septentrionalis 0 2 1 0 0 0 0 0 0 0 0 0 - 0 0 - 0 1 1 0 1 0 1 0 0
T. smithi 0 2 0 0 0 0 0 0 0 0 0 0 - 0 0 - 0 0 1 1 0 - 1 0 0
T. tucumanus 0 0 1 0 2 2 0 0 0 1 0 1 0 0 1 2 1 1 0 0 1 1 1 0 0
T. turrifex 0 0 0 0 1 1 1 1 1 1 0 1 1 1 0 - 1 2 1 0 1 0 1 0 1
T. urichi 0 0 0 0 1 1 1 1 1 1 0 1 1 1 0 - 1 1 1 0 1 0 1 0 1
T. zeteki 0 0 0 1 1 1 1 1 0 1 0 1 0 1 0 - 1 2 0 0 1 0 1 0 1
Table 2 (continued)
2 2 2 2 3 3 3 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 4 5
6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0
Mycetarotes 0 0 1 0 0 0 0 0 0 0 0 0 0 0 - 0 1 1 0 1 1 2 3 0 0
T. agudensis 0 0 1 1 1 0 0 0 0 1 0 0 0 1 1 1 1 0 0 1 0 1 0 0 1
T.arizonenensis 0 0 1 1 1 0 0 0 0 1 - 0 0 0 - 1 1 2 1 0 0 1 2 0 1
T. bugnioni 1 1 1 1 1 0 0 1 0 1 - 0 0 0 - 0 0 1 1 1 0 0 2 0 1
T. compactus 0 0 0 - - 1 - 1 1 1 - 1 0 0 - 1 1 0 0 0 2 2 2 0 1
T. cornetzi 1 1 1 1 1 0 1 1 0 1 - 0 1 0 - 1 1 1 1 1 1 1 0 0 1
T. dichrous 2 0 0 - - 2 - 0 0 0 0 1 0 0 - 1 2 0 0 0 2 2 3 0 0
T. diversus 0 1 1 1 2 1 0 1 0 1 - 0 0 1 0 1 0 1 1 0 1 0 2 0 1
T. farinosus 0 0 1 1 1 0 1 0 0 0 0 0 0 0 - 0 0 2 1 0 0 1 2 1 1
T. fuscus 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 2 0 2 0 1 0 1 0 0 1
T. holmgreni 0 1 1 1 1 0 1 0 1 1 - 0 0 1 0 2 1 1 1 1 0 1 1 0 1
T. iheringi 0 0 1 1 0 1 0 1 0 1 - 0 0 1 0 0 1 2 1 0 0 1 3 0 1
T. irmgardae 1 1 1 1 1 1 0 1 1 1 - 0 0 0 - 1 0 1 1 1 0 0 2 0 1
T. isthmicus 0 0 0 - - 0 - 0 0 0 0 1 0 0 - 1 1 2 0 1 0 1 0 0 1
T. jamaicensis 0 0 1 1 1 0 0 0 0 1 - 0 0 0 - 0 0 1 1 1 0 0 1 0 1
T. levis 1 1 1 0 0 1 0 1 1 1 - 0 0 1 0 1 1 1 1 1 0 0 2 0 1
T. mandibularis 1 1 1 0 0 2 1 0 0 0 0 0 0 0 - 0 0 2 0 1 0 0 2 0 1
T. nogalensis 0 0 1 1 1 0 1 0 0 0 1 0 0 0 - 1 1 2 0 0 1 1 2 0 1
T. oetkeri 0 0 1 0 0 0 1 0 0 0 0 0 1 1 0 0 0 2 1 0 0 0 2 0 1
T. opulentus 1 1 1 1 0 2 1 0 0 0 1 1 0 0 - 1 0 0 0 0 0 2 2 0 0
T. papulatus 0 1 1 1 0 1 1 1 1 1 - 0 0 1 0 0 0 2 1 0 0 1 2 0 1
T. phaleratus 0 0 1 0 0 1 1 1 0 0 0 0 1 1 0 0 0 2 0 1 0 1 1 0 1
T. pruinosus 0 1 1 0 0 1 1 1 0 1 - 1 0 1 0 0 1 2 1 0 0 1 1 0 1
T. relictus 0 0 1 1 0 1 1 1 0 0 0 0 0 0 - 0 0 2 1 0 0 2 0 0 0
T. ruthae 0 0 1 1 0 1 1 2 0 1 - 0 0 0 - 0 0 2 0 0 1 2 2 1 2
T. saussurei 0 0 1 1 1 0 1 0 0 0 0 0 0 1 0 0 0 2 1 1 1 0 2 0 1
T.septentrionalis 0 0 1 0 0 0 1 0 0 1 - 0 0 1 0 0 0 2 1 0 1 0 2 0 1
T. smithi 0 1 1 1 0 1 1 1 0 1 - 0 0 1 0 0 0 2 1 0 1 0 3 0 1
T. tucumanus 0 0 1 1 1 1 0 1 0 1 - 1 0 1 0 0 1 2 1 0 0 1 3 0 1
T. turrifex 0 0 1 1 0 1 1 1 0 1 - 0 0 0 - 0 0 2 1 0 0 1 2 0 1
T. urichi 0 1 1 1 0 0 1 0 0 0 0 0 0 1 0 2 0 2 1 0 0 1 0 0 1
T. zeteki 0 0 1 1 1 0 1 0 0 1 - 0 0 0 - 0 0 2 0 0 0 0 1 0 1
Cover, S. P., Deyrup, M. 2007. A new ant genus from the southwestern United States, pp. 89-99.
In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics
(Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs of
the American Entomological Institute, 80.
A NEW ANT GENUS FROM THE SOUTHWESTERN UNITED STATES
Stefan Cover
Department of Entomology
26 Oxford Street, Cambridge
MA 02138. USA
cover@oeb.harvard.edu
and
Mark Deyrup
Archbold Biological Station,
P.O. Box 2057
Lake Placid, FL 33852. USA
MDeyrup@archbold-station.org
ABSTRACT
A new ant, Dolopomyrmex pilatus (new genus, new species), is described from the southwestern
United States. The ant belongs to the subfamily Myrmicinae and is placed in the tribe
Solenopsidini, as defined by Bolton (1987, 2003). Within the tribe it seems closer to the Old
World genera Anillomyrma and Bondroitia, rather than the New World genus Tranopelta, as
previously suggested by some. D. pilatus appears to be almost exclusively subterranean, which
may account for the rarity with which it has been collected. Brief observations on a living colony
fragment revealed that the workers fed only on termite nymphs and the brood of other ants.
Key words: Formicidae, Myrmicinae, Solenopsidini, Dolopomyrmex, Tranopelta, Anillomyrma,

Bondroitia, desert, new genus, new species, United States, Mexico.
INTRODUCTION
Although the ant fauna of North America north of the Mexican border is diverse, consisting of
an estimated 800-900 species, no new ant genus has been described from material collected in
the United States or Canada since Mycetosoritis, described by William Morton Wheeler in 1907.
The new genus described herein was first discovered in the southwestern United States, in the
Madrean Archipelago region of southern Arizona, known as the “Sky Islands.” The exuberant
ant fauna of this area is but a northern extension of the enormous, yet still poorly understood
fauna of the Sierra Madre Occidentale of Mexico, and we expect other discoveries of
comparable importance will be made as the ants of this region become better known. Additional
collections of the new genus have been made in New Mexico and California from community
types that, like the type locality, are clearly derived from the mountain and desert ecosystems of
northern Mexico.
Specimens were examined and measured using a Wild (now Leica) M-5A with micrometer.
Measurement conventions and indices are as defined in Bolton (1987), with the exception of AL
(alitrunk length), here recorded as ML (mesosoma length).
Dolopomyrmex, new genus

Figures 1-3
Diagnosis of Worker. Monomorphic subterranean myrmicine ants with the following

combination of characters:
1) Palp formula 3,2

2) Mandible with 4 teeth and strongly oblique cutting margin, the largest gap between
mandibular teeth separating the basal tooth from the third subapical tooth.
3) Clypeus ecarinate. Median clypeal seta absent, instead anterior margin with pair of long,
ventrally curved setae, one on each side of the midpoint, flanked laterally by fringe of 5-6
long, forward-projecting setae on each side of the clypeal margin.
4) Median portion of clypeus narrow, strongly elevated, antennal insertions closely
approximated.
5) Frontal lobes small, subtriangular, fully covering condylar bulbs of antennae. Frontal
carinae absent.
6) Eyes vestigial or entirely absent, when present, consisting only of the remnant of a single
ommatidium.
7) Antennae 11 segmented, with an enormous 3 segmented apical club longer than the
remainder of the funiculus.
8) Mesosoma elongate, in profile, compressed laterally at promesonotal juncture with
propodeum giving it a distinctive hourglass shape in dorsal view.
9) Metanotal impression present, propodeum lacking teeth or spines, in profile dorsally
convex, with flat posterior face in side view.
10) Propodeal spiracle circular, of moderate size. Metapleural gland well-developed.
Metapleural lobes small, rounded and inconspicuous.
11) Petiole with short, indistinct peduncle, node low, rounded, suborbicular in dorsal view.
Subpetiolar process absent. Spiracle circular, located laterally on the anterior slope of the
node.
Cover & Deyrup: New Ant Genus 91
12) Postpetiole low and rounded in profile, lacking a ventral projection or tooth, suborbicular in
dorsal view.
13) Tergite and sternite of the fourth abdominal segment meeting to produce an angulate corner
on each side of the postpetiole seen in dorsal view, as in the unrelated myrmicine genus
Pheidole.
14) Sting simple, robust.
15) Outer surfaces of middle and hind tibiae and tarsomeres with abundant stiff, spine-like, erect
to suberect setae.
Diagnosis of Queen. As in worker except:
1) Enormous in comparison with worker caste. ML always > TL of largest workers.

2) Mandible with strongly oblique cutting margin and 4 teeth that decrease progressively in
size from apical to basal.
3) Submedian ventrally curved setae on anterior clypeal margin not clearly separable from
other setae present.
4) Ventral surface of head with abundant, curved, delicate erect setae.
5) Compound eyes large, with scattered very short, stubble-like, erect setae. Ocelli well-
developed but small.
6) Antennae 11 segmented as in worker, 3 segmented apical club present, but less pronounced.
7) Body in general somewhat flattened, probably as an adaptation for subterranean life.
8) Mesosoma massive, with full complement of sclerites associated with the presence of
wings.
9) Petiolar node subrectangular, wider than long, in dorsal view. Postpetiole subelliptic in
dorsal view.
10) Spine-like erect setae abundant on outer surface of hind tibia, especially abundant on the
outer tibial surface of the middle leg.
Diagnosis of Male.
1) Palpal formula 2,1. Palps very short.

2) Mandible strap-like, with strongly oblique cutting margin and 4 teeth, decreasing
progressively in size from the apex. Rarely, 1-2 teeth may be absent, leaving only the apical
and subapical teeth present.
3) As in worker caste, clypeus ecarinate, median clypeal seta absent, paired submedian
ventrally curved setae present.
4) Median portion of clypeus narrow, elevated, antennal insertions closely approximated.
5) Frontal lobes small, only partly covering the condylar bulbs, frontal carinae absent.
6) Compound eyes large, with short erect setae as in queen. Ocelli well-developed but small.
7) Antennae 11 segmented, with large 3 segmented club. First antennal segment greatly
enlarged to form distinct scape longer than the apical club.
8) Mesosoma massive, alate, wing venation as in Fig. 3. Propodeum lacking teeth or spines,
spiracle lenticular, metapleural lobes very small.
9) Petiolar node low and rounded in profile, peduncle absent. Petiolar spiracles located at the
tips of large, laterally projecting connules.
10) Postpetiole unique, as in Fig. 3. Attachment to fourth abdominal segment extraordinarily
broad, spiracles as in petiole, located at the tips of large, laterally projecting connules.
11) Erect or suberect setae present on outer surfaces of middle and hind tibiae, but neither stiff
and spine-like, nor as abundant as in the female castes.
Type species: Dolopomyrmex pilatus Cover & Deyrup, new species.

Etymology: Dolops, Dolopos (ancient Greek, “lurker in ambush”) + myrmex, myrmekos

(Greek, “ant”).
Dolopomyrmex pilatus, new species

Figures 1-3
Holotype worker: TL 2.0, HL 0.54, HW 0.49, CI 91, SL 0.33, SI 67, PW 0.33, ML 0.70.
With characters of the generic diagnosis and as illustrated in Fig. 1. Head broadly
rectangular, posterior corners rounded, posterior margin flat, entire dorsal surface covered with
coarse setigerous foveolae, generally separated by 1-2× their diameters. Spaces between
foveolae smooth and shining. Eye remnant barely visible; unpigmented. Numerous short,
decumbent to appressed setae present on dorsal surface of head, sparse appressed setae present
on ventral cephalic surface. Antennal scapes in repose extending about 3/4 the distance towards
the posterior corners of the head, with numerous erect or suberect hairs on outer surfaces. Body
surfaces generally smooth and shining on mesosoma, petiole, postpetiole, and remainder of
abdomen. Numerous, short, curved erect hairs present on promesonotum, longer erect hairs
present on propodeal dorsum, petiolar node, and postpetiole, respectively. Remainder of
abdomen (= gaster) with evenly spaced erect hairs as in Fig. 1. Color pale yellow.
Paratype workers: TL 1.6-2.4, HL 0.47-0.56, HW 0.42-0.51, CI 89-91, SL 0.27-0.34, SI
67-68, PW 0.26-0.33, ML 0.59-0.74.
Paratype queen: TL 9.5, HL 1.27, HW 1.49, CI 117, SL 0.79, SI 53, ML 3.2. With
characters in generic diagnosis and as illustrated in Fig. 2. Head broader than long, posterior
margin flat, dorsal surface foveolate, foveolae larger and much sparser on posterior half of head.
Spaces between foveolae smooth and shining, except on anterior half, where weak striae are
present surrounding the antennal sockets and near the mandibular insertions. Clypeus foveolate,
except for smooth median strip, erect setae numerous. Antennal scape in repose against head
with many erect setae on outer surfaces. Mesosoma largely smooth and shining, sparsely and
finely foveolate on most surfaces, coarser foveolae present on mesopleuron. Petiole and
postpetiole smooth and shining with scattered fine foveolae, each with a tuft of long, erect setae
on ventral surface. Remainder of abdomen smooth and shining with sparse, very fine foveolae
and moderately abundant, short erect hairs. Outer surfaces of middle and hind tibiae with
abundant, coarse, setigerous foveolae. Color: body generally light to medium yellowish-brown,
appendages a lighter brownish yellow. Posterior dorsal surface of head with large, dark brown
spot centering on the ocelli.
Paratype male: TL 5.0, HL 0.68, HW 0.72, CI 106, SL 0.45, SI 62, ML 1.85. With
characters in generic diagnosis and as illustrated in Fig. 3. Head with fine striate sculpture over
most of dorsal surface, except for smooth median clypeal strip and two lateral smooth, shining
patches between the ocelli and compound eyes. Mesosoma generally unsculptured and shining
with widely spaced very fine setigerous foveolae, except for some coarser foveolae on the
mesopleuron, and fine striate sculpture on the mesopleuron and the propodeal sides. Petiole and
postpetiole with small tuft of long, erect setae on ventral surface. Remainder of abdomen with
widely-spaced, very fine setigerous foveolae, erect hairs sparse and short. Color: head and
mesoscutum blackish brown, Mesosoma, petiole, and postpetiole variably light to medium
yellowish brown. Legs light yellowish brown, antennae pale yellow.
Etymology: “pilum” is Latin for the short, but powerful throwing spear of the Roman
legionaries. So Dolopomyrmex pilatus is the “spear-bearing ambush ant.”
Type Locality: USA, Arizona: Cochise Co. Chiricahua Mtns. 0.8mi NW Jct. Forest
Service Road 42 on FSR 42B (Paradise Road). 31°55.24’N, 109°09.67’W. elevation 4700’
(1432 m.). Broad, grazed wash dominated by mesquite and acacia to 3 m. tall. Fine-textured,
densely compacted, sandy soil with some clay. Note: this is privately owned land adjacent to the
Coronado National Forest.
Type Series: holotype worker (SP Cover 1441) and the following paratypes: 22 workers,
13 males, 15 queens [3-VIII-1988, SPC 1441], 9 workers, 12 queens [4-VIII-1988, SPC 1454], 6
workers [17-VIII-1989, SPC 2173], 13 workers [17-VIII-1989, SPC 2174], 4 workers [3-VIII-
1990, SPC 2530]. Holotype and paratypes deposited in the Museum of Comparative Zoology
(Cambridge, Mass., USA). Additional paratypes will be deposited in the Natural History
Museum (London, U.K), the Natural History Museum of Los Angeles County (Los Angeles,
California), the National Museum of Natural History (Washington, DC), the California Academy
of Sciences (San Francisco, CA), the Museu de Zoologia (Sao Paolo, Brazil), the Australian
National Insect Collection (Canberra, Australia), the University of California (Davis, California)
and in the collection of William P. Mackay. Note: images of the holotype will be available on
the online MCZ Type Database [http://mcz-28168.oeb.harvard.edu/default.htm] and images of
paratypes on AntWeb [http://www.antweb.org].
Additional specimens examined: USA, California: Los Angeles Co. Saddleback Butte.
23-IX-1989. coll. G.C. Snelling (2 males, 2 dealate females) [LACM]. New Mexico: Dona Ana
Co., 45km NE Las Cruces. Jornada LTER. 3-X-1970. colls. G. Richardson and C.W. O’Brien (3
alate females, 1 dealate) [LACM]; Socorro Co. Servilleta National Wildlife Refuge. Date
unknown (not seen by the authors, but cited in Mackay and Mackay (2002).
DISCUSSION
It is possible the California collections may represent a second species of Dolopomyrmex. The
two queens are somewhat larger than those from Arizona and New Mexico and exhibit some
differences in morphology and pilosity. The two males taken with them, however, closely
resemble males from the type series. These specimens are tentatively assigned to D. pilatus
pending the collection of workers, upon which their status should be re-evaluated.
The type locality is a desert wash with highly compacted, sandy soil located at the border
between relatively flat desert and the foothills of the Chiricahua Mountains. It is a very arid,
sunbaked habitat for most of the year. Ant species found at the site include: Crematogaster
opuntiae, Aphaenogaster albisetosa, Pogonomyrmex barbatus, P. imberbiculus, Pheidole
desertorum, P. rugulosa, Dorymyrmex insanus, Forelius maccooki, and Camponotus fragilis, all
desert or desert foothills ants common throughout much of the American Southwest. The
following notes summarize the circumstances of each collection at the type locality. SPC 1441
was collected when a single worker was seen entering a minute, nearly invisible hole with a
trace of excavated soil around it, in bare soil between mesquite clumps a day after rain.
Chambers containing winged queens, males, some brood, and workers were found between 30-
50 cm. deep. SPC 1454 was found in soil “while excavating a nest of another species” and
included winged queens as well as workers. SPC 2173, 2174 and 2530 are all small series of
workers taken 15-25 cm deep in soil when excavating nests of Pogonomyrmex imberbiculus
after rain at the type locality. The Pogonomyrmex nests were all located in bare soil between
mesquite clumps. SPC 1441 was maintained alive for about a week in a large plastic petri dish
with a plaster bottom and moist cotton in it. The workers fed intermittently on fresh ant brood
and termite nymphs, but ignored other dead insects and sweet substances. Mackay & Mackay
(2002) report this ant (as Tranopelta sp.) from “Creosotebush scrub and mesquite dominant
zones, often near desert playas or arroyos, sometimes in open desert or in forest meadows.
Occasionally they are found in salt flats or saltbush communities.”
Dolopomyrmex fits comfortably within the myrmicine tribe Solenopsidini, as newly
expanded by Bolton (2003). Its affinities within the tribe are the subject of some disagreement.
Since the discovery of Dolopomyrmex in 1988, several myrmecologists have suggested
informally that it is most likely an undescribed species of the exclusively New World myrmicine
genus Tranopelta. Indeed, Mackay & Mackay (2002) place the ant in that genus (without
formally describing it) and summarize the reasons for doing so. The hypothesis makes some
sense. Dolopomyrmex and Tranopelta share important character states. The workers of both lack
a bicarinate clypeus and a median clypeal seta, and instead possess a pair of subparallel setae
that straddle the clypeal midpoint. The antennae of both are 11 segmented with a prominent 3
segmented apical club. The queens and males also share some notable similarities. Both
Dolopomyrmex and Tranopelta have fully subterranean life-histories, workers yellow in color
and with highly reduced eyes, and sexual forms far larger than the corresponding workers. In
addition, the geographic range of Tranopelta extends north into southern Mexico, so it is not
impossible that a species might occur in the extreme southwestern United States.
In our opinion, however, a closer examination reveals differences that not only preclude the
inclusion of D. pilatus in Tranopelta, but also in fact, reveal that the two genera are not closely
related. For example, in the worker caste of Dolopomyrmex the clypeus is narrow, the median
portion strongly elevated, and the antennal sockets closely approximated (clypeus broad, median
portion weakly elevated, antennal sockets well-separated in Tranopelta), the cutting edge of the
mandible is strongly oblique (more or less transverse in Tranopelta), the petiolar node is low,
rounded, and nearly circular in dorsal view (strongly upright and wider than long in dorsal view
in Tranopelta), the petiolar spiracle is located on the anterior side of the node (on the well-
developed peduncle in Tranopelta), and a subpetiolar process is lacking (one, sometimes two
present in Tranopelta). In addition, compound eyes are vestigial or lacking in Dolopomyrmex
(present, with at least several well-defined and pigmented ommatidia in Tranopelta), and the
workers are monomorphic (weakly polymorphic in Tranopelta). Lastly, the males are strikingly
different. Those of Dolopomyrmex possess a true antennal scape nearly as long as the funiculus,
and the petiolar and postpetiolar spiracles are located at the tips of unique lateral connules. In
Tranopelta males, the first antennal segment is very short, no longer than the following two
funicular segments taken together, and the petiolar and postpetiolar spiracles are normal in
configuration.
Instead, within the Solenopsidini Dolopomyrmex appears to be closer to two exclusively Old
World solenopsidine genera, Anillomyrma and Bondroitia. A comparison of the workers is
instructive and striking. All three genera share a closely similar structure of the clypeus, frontal
lobes, and antennae, strongly oblique mandibular cutting margins with 4 teeth (3 teeth in one
Anillomyrma species), and similar petiolar structures, most notably having the spiracle located
anterolaterally on the side of the node, reduced palpal counts, and vestigial or absent compound
eyes. The only truly discordant note is the absence of a median clypeal seta in Dolopomyrmex,
present in Anillomyrma, Bondroitia, and most other Solenopsidine genera. The presence of a
median clypeal seta was once thought to be a diagnostic character for the Solenopsidini (Bolton,
1987), but is presently seen to be more variable and less important than previously thought.
Bolton’s (2003) recent redefinition of the tribe reflects this change. We look forward to future
discoveries that will help further clarify the phylogeny within this important and interesting
group of genera. With respect to Dolopomyrmex in particular, we need better collections of the
sexual forms of both Anillomyrma and Bondroitia. Two damaged males and a fragmentary
queen of Bondroitia lujae at the MCZ collection show important similarities, as well as some
notable differences, with the comparable castes of Dolopomyrmex. See also the figures of
Bondroitia lujae in Bolton (1987).
To facilitate recognition of the new genus we present a modified version of the portion of
Bolton’s (1987) key to the workers of the Solenopsidini that separates genera in which the
antennae have eleven segments. In addition to Dolopomyrmex, Tranopelta is added to the key.
7 Propodeal spiracle notably enlarged ..................................................................................... 8

-- Propodeal spiracle normal in size ......................................................................................... 9
8 Median clypeal seta present. Eyes absent .........................................Bondroitia (Afrotropical)

-- Median clypeal seta absent. Eyes present, consisting of at least several distinct, pigmented
ommatidia ......................................................................................... Tranopelta (Neotropical)
9 Antennal club 2 segmented. Head almost circular in full-face view. Eyes posterior to
cephalic midlength.......................................................................................... Phacota (Spain)
-- Antennal club 3 segmented. Head not almost circular in full-face view. Eyes, when present,
at or anterior to cephalic midline ........................................................................................ 10
10 Propodeum in profile sharply angulate to bidentate ............................................................ 11

-- Propodeum in profile evenly rounded .................................................................................. 12
11 [Couplet as in Bolton, which leads to Oxyepoecus and Allomerus (in part), both Neotropical
genera]
12 Median portion of clypeus distinctly elevated. In profile, postpetiole less voluminous than
petiolar node, postpetiole narrowly attached to abdominal segment IV in dorsal view ....... 13
-- Median portion of clypeus evenly transversely convex, not distinctly elevated. In profile,
postpetiole more voluminous than petiolar node, postpetiole broadly attached to abdominal
segment IV in dorsal view ........................................................... Diplomorium (Afrotropical)
13 Clypeus strongly or weakly bicarinate. Median clypeal seta present ....Monomorium (in part)
-- Clypeus lacking carinae. Median clypeal seta absent, instead clypeus with pair of long,
ventrally curving setae that straddle the midpoint ..........................Dolopomyrmex (Nearctic)
In conclusion, Ed Wilson once remarked (context and exact wording now scrambled in
declining memory of first author) that the soil was perhaps the final frontier of biodiversity
exploration. Ants like Dolopomyrmex are just a hint of the diversity that remains to be
discovered if we can improve methods for finding the invertebrates that apparently abound in the
middle and lower soil horizons, at present safely hidden from our sight.
DEDICATION
The authors dedicate this paper to Edward O. Wilson on the occasion of his “golden jubilee”
year as a publishing ant systematist. Much will rightly be said concerning the importance of his
many scientific accomplishments. Here we wish to honor his service to myrmecology as a friend
to all interested in ants, as an encourager and supporter of myrmecological research and
education, and as a mentor, teacher, companion, and guide to all of us called to the honorable
study of the little creatures that rule the world.
ACKNOWLEDGMENTS
We wish to thank Barry Bolton, Phil Ward, Bill Mackay, and Roy Snelling for thoughtful and
valuable discussions and for the loan of important specimens, and Phil Ward and Roy Snelling
for much appreciated comments on the manuscript. We also thank Wade and Emily Sherbrooke,
then at the Southwestern Research Station (Portal, Arizona), for abundant kindness, good cheer,
and logistical support. This research was made possible by funding from the Museum of
Comparative Zoology, and the Theodore Roosevelt Fund at the American Museum of Natural
History.
LITERATURE CITED
Bolton, B. 1987. A review of the Solenopsis genus-group and revision of Afrotropical

Monomorium Mayr (Hymenoptera: Formicidae). Bulletin of the British Museum of Natural
History. Entomology Series 54: 263-452.
Mackay, W. & Mackay, E. 2002. The Ants of New Mexico (Hymenoptera: Formicidae). The
Edwin Mellen Press, Ltd., Lewiston, New York, 398 pp.
Cover & Deyrup: New Ant Genus
Figure 1. Dolopomyrmex pilatus, Worker habitus.

97
Figure 2. Dolopomyrmex pilatus, Queen habitus.

Figure 3. Dolopomyrmex pilatus, Male habitus.

Deyrup, M., Cover, S. P. 2007. A new species of Crematogaster from the pinelands of the
southeastern United States, pp. 100-112. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds).
Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
A NEW SPECIES OF CREMATOGASTER FROM THE PINELANDS OF THE

SOUTHEASTERN UNITED STATES
Mark Deyrup
Archbold Biological Station
P.O. Box 2057
Lake Placid, FL 33862
MDeyrup@archbold-station.org
and
Stefan Cover
Harvard University
26 Oxford Street, Cambridge
MA 02138. USA
cover@oeb.harvard.edu
ABSTRACT
A new species of arboreal Crematogaster, C. pinicola, is described from the southeastern United
States. It is a cryptic species closely related to the common C. ashmeadi Mayr, from which it can
be distinguished primarily by its red and black coloration (uniform dark brown to black in C.
ashmeadi), and by its narrow ecological specialization: it nests only in open-grown pines with
thick twigs (C. ashmeadi occurs in many forest types and nests in a wide range of hardwoods and
conifers). Crematogaster pinicola is believed to be a relic of the fire-maintained pine ecosystems
that once dominated much of southeastern North America. A recent study shows that C. pinicola
is a major part of the diet of the endangered red-cockaded woodpecker.
Key words: Hymenoptera, Formicidae, Myrmicinae, Crematogaster, new species, taxonomy, fire
ecology.
Deyrup & Cover: A new pine-inhabiting Crematogaster 101
INTRODUCTION
Like many other large ant genera, the genus Crematogaster presents ant taxonomists with an
unending series of challenging taxonomic puzzles. These problems are not inconveniences: to the
thoughtful myrmecologist they are further evidence that biodiversity is as subtle as it is
spectacular. The solving of these problems is at the heart of what Wilson (1992) calls “the great
Linnean enterprise,” the inventory of the biosphere.
The first useful account of the North American Crematogaster was Creighton’s (1950)
treatment of the genus in the “Ants of North America.” From the past Creighton inherited a
confusing list of ill-defined taxa, the customary legacy to modern taxonomists from the “Bronze
Age” of Ant Taxonomy (1750 - 1950), an era characterized by much honest descriptive effort that
was critically undermined by the lack of a biologically realistic conceptual framework. Creighton
began the process of bringing order to the North American ant fauna, and to the Crematogaster in
particular, by employing the newly popularized biological species concept to elicit meaning from
what seemed to be an intractable morass of names and morphological variation (see Buhs, 2000
for a discussion of this development). Expanding on this pioneering work, Buren (1958, 1968)
revised the North American Crematogaster again, incorporating much newly available
information. Like Creighton, Buren made extensive use of distributional data and natural history
observations in making taxonomic decisions and, as a result, his work still forms the basis for our
modern understanding of the genus in North America. Not much has been added since Buren’s
studies. Johnson (1988) reviews the eastern species and presents a key to them. In an excellent
new revision of the Crematogaster of Costa Rica, Longino (2003) makes several taxonomic
changes that affect the North American fauna.
The new species of Crematogaster described here was first recognized as an undescribed
species separate from C. ashmeadi Mayr by William Buren himself. After retiring, Buren moved
to Florida, where he retained his interest in Crematogaster. Hand-labeled specimens in Buren’s
collection show he believed that the common southeastern C. ashmeadi included a second,
previously unrecognized species distinguished by its red and black coloration. His student James
Trager also knew of this species, as evidenced by specimens of C. pinicola collected in 1981 at the
Archbold Biological Station and labeled by him “Crematogaster n. sp.” Buren probably would
have described this species, but he became ill and died in 1983. The same ant is also the
“undescribed species” of Crematogaster referred to by Deyrup and Trager (1986). In his review of
eastern Crematogaster, Johnson (1988) was the first to address the problem of separate color
forms in C. ashmeadi in print, but he did not arrive at a definite conclusion concerning their
biological basis. We believe the accumulated evidence supports the hypothesis that the red and
black form of Crematogaster ashmeadi is, in fact, a valid sibling species.
METHODS AND MATERIALS
Specimens were examined and measured using an ocular micrometer scale in a Leitz
stereomicroscope at 40x. Measurement conventions and indices follow those used in Bolton
(1994).
Crematogaster pinicola, new species

Figures 1-3
DIAGNOSIS: Workers morphologically indistinguishable from those of C. ashmeadi, except

for a distinctive color difference in freshly collected material. In C. pinicola workers, the head,
mesosoma, petiole, postpetiole, and appendages are ferrugineous red, and the gaster is black. In C.
ashmeadi, mature specimens are always a uniform dark brown to black. A similar color distinction
is seen in alate queens; those of C. pinicola are notably bicolored, those of C. ashmeadi are
uniformly brown or black. Males of C. pinicola are generally somewhat lighter in color than those
of C. ashmeadi, but are harder to distinguish reliably than the corresponding female castes. Note:
detailed morphometric studies might possibly reveal the existence of minute, but consistent
morphological differences between the two species, but a detailed examination of all three castes
in both species has not provided hints that such differences exist.
Worker: Measurements (mm) (holotype in parenthesis): total length: 2.31-3.64 (3.22); head
width at eyes: 0.67-0.90 (0.87); length of antennal scape: 0.44-0.60 (0.60); distance from
mesothoracic spiracle to propodeal spiracle: 0.23-0.33 (0.31); distance from lower edge of
propodeal spiracle to tip of propodeal spine: 0.12-0.17 (0.16). Head: in frontal view, posterior half
smooth, shining, covered with sparse appressed silvery hairs separated at their bases by a distance
slightly shorter than length of a hair; orientation of hairs convergent toward lower midline of
frons; frons with a series of erect hairs in a line just mesad of imaginary vertical lines extending up
from frontal carinae, 4 hairs on right side, 3 on left (in holotype); fine striae covering malar area,
extending up about 1/3 of way along eye on inner side. Antennal scape with appressed hairs only.
mandible with 4 teeth. Mesosoma: pronotum with one standing curved humeral hair on each side;
pronotum and mesonotum with sparse appressed silvery hairs, with bases separated by more than
half length of a hair and less than twice length of a hair; pronotum and mesonotum shining, with
very faint shagreening; mesopleuron finely, evenly reticulate up to level of mesothoracic spiracle;
metapleuron with longitudinal carinae covering its upper 3/4, fine reticulations between more
widely spaced carinae; propodeal spine in lateral view wedge-shaped, sharply pointed, with a fine
dorsal carina; dorsal areas of propodeum with sparse, appressed silvery hairs divergent from the
midline; all legs with sparse, appressed, silvery hairs. Gaster: First tergite sparsely covered with
longitudinally oriented, appressed, silvery hairs whose bases are slightly closer together than the
length of a hair; submarginal bands of similar, but longer hairs on tergites 1 - 3; a sparse
submarginal band of erect hairs on tergites and sternites 1 - 4. Color: Body and appendages except
for gaster ferruginous; gaster black.
Queen from nest of holotype (Fig. 3). Measurements (mm): total length: 7.48-7.90; head
width at eyes: 1.44-1.54; length of mesosoma (lateral view): 2.20-2.45; length of forewing: 6.46-
6.84. Head, legs, body reddish brown, except mesonotum, scutellum blackish brown, gaster black;
wings hyaline, major veins pale testaceous. Mandible with 5 teeth, mandibular striae with sparse,
coarse punctures; median ocellus separated from lateral ocelli by about 1.8 times diameter of
lateral ocellus; anterior half of dorsum of head finely striate, including clypeus, except for lower
median area of frons; posterior half of dorsum of head shining, finely punctate, with appressed
hairs. Mesonotum strongly shining, no reticulate areas; fine striations on lateral margins of
mesonotum and posterior quarter of mesopleuron; metapleuron coarsely but evenly striate,
propodeum coarsely, unevenly striate; gaster shining, first gastral tergite with appressed hairs
slightly longer than distance between their bases, and a few scattered, suberect longer hairs.
Male from nest of holotype (Fig. 2). Measurements (mm): total length: 3.02-3.24; head width
at eyes: 0.62-0.66; length of mesosoma (lateral view): 1.06-1.30; length of forewing: 2.87-3.13.
Head and body blackish brown; femora medium brown, lighter than head and body; tibiae, tarsi,
mandibles testaceous; wings hyaline with no infuscation, heavier veins pale testaceous. Mandible
with 3 subequal teeth; median ocellus separated from lateral ocelli by twice diameter of lateral
ocellus; head with sparse sub-appressed hairs, those on occipital area procumbent, about as long as
distance between their bases; hairs on frons convergent toward midline; frons with a few
conspicuous large punctures on each side; malar space weakly striate; area between eye and
antennal sockets not striate; antennal scape shorter than last antennal segment. Mesosoma shining,
without reticulate areas, smooth except for weak, fine striations on lateral areas of mesonotum and
posterior fourth of mesopleuron, metapleuron more coarsely striate; mesonotum with sparse, short
hairs embedded in elongate punctures, usually farther apart than length of a hair; wing venation as
in figure 2; gaster smooth, shining, first gastral tergite with short, appressed, embedded hairs.
TYPE MATERIAL
USA: Florida: Highlands County, Archbold Biological Station, 24-VI-1996, M. Deyrup.

Florida Scrub habitat. Nest in 6 cm diameter branch of Pinus elliottii in firelane. The entire type
series is from a single colony. Holotype and 37 paratypes deposited in the Museum of
Comparative Zoology, Harvard University, Cambridge, Massachusetts. Deposition of additional
paratypes: 24 workers, 2 alate queens, 2 males: Natural History Museum of Los Angeles County,
Los Angeles, California; 17 workers, 2 alate queens, 1 male: Florida State Collection of
Arthropods, Gainesville; 17 workers, 2 alate queens, 1 male: National Museum of Natural History,
Smithsonian Institution, Washington, D.C.; 14 workers, 1 alate queen, 1 male: The Natural
History Museum, London; 12 workers: collection of William Mackay, El Paso, Tex.; remaining
type material: Archbold Biological Station, Lake Placid, Florida.
ETYMOLOGY
The specific epithet is derived from Latin: “pine dweller.”
DISCUSSION
Geographic Range and Variation in Florida.
The distribution map provided in Fig. 4 shows localities for collections documented by
voucher specimens only. The real distribution of C. pinicola is undoubtedly much more extensive
and continuous. Without mentioning specific collections, Johnson (1988) says that the species
occurs in Georgia, South Carolina, and Alabama also. We have not seen Johnson’s specimens, but
expect that C. pinicola will be found to occur widely on the southern coastal plain where suitable
habitats are found. Material collected throughout Florida shows a remarkable consistency in color,
size range, and morphology. Crematogaster pinicola is clearly less variable than its sister species,
C. ashmeadi, which shows more obvious variation in size and color over its much wider
geographic distribution.
In describing a species of ant that can be recognized visually only by its color we are aware
that we a treading on the myrmecological equivalent of “thin ice.” Early ant systematists were
notorious for naming new taxa (both specific and infraspecific) based on minute (and indeed,
sometimes entirely imaginary) differences in color, sculpture, or pilosity. Creighton (1950) was
withering in his criticism of these practices, and subsequent generations of ant taxonomists have
been strongly conditioned to believe that color alone is unreliable as a separatory character on the
species level. Much cumulative experience with the genus Crematogaster is an additional reason
for caution. Color variation is not uncommon in Crematogaster species. For example, dark and
bicolored variants have been observed in other Crematogaster of the eastern United States,
notably C. atkinsoni and C. pilosa. In both species, northern specimens tend to be uniformly black
or brown in color, while bicolored specimens are sometimes found in Florida and elsewhere along
the Gulf Coast. With precisely this in mind, Johnson (1988) treated C. pinicola as a color variant
form of C. ashmeadi.
We believe that ecological evidence clearly indicates that C. ashmeadi and C. pinicola are
reproductively isolated, and thus C. pinicola is a good species, not merely a color variant. While
broadly sympatric with C. ashmeadi across most of Florida, C. pinicola is distinctively different in
its ecology. It nests exclusively in pine trees, particularly slash pine (Pinus elliotii) and longleaf
pine (P. palustris), which have relatively robust twigs. Pines occupied by C. pinicola are usually
open-grown (i.e., well-separated from other trees) and surrounded by low brush or low perennial
herbs and grasses. These pines may be in dry sandhill habitats or in wetter flatwoods habitats.
Both habitat types are maintained by frequent fires that retard the invasion of other woody plant
species. In sharp contrast, C. ashmeadi shows much wider ecological tolerances in nest site
selection, and in habitat preferences. C. ashmeadi nests in pines, in many hardwoods including
oaks (Quercus spp.), hickories (Carya spp.), ash (Fraxinus caroliniana), red maple (Acer rubrum),
shrubs such as winged sumac (Rhus copallina L.), vines such as greenbriar (Smilax spp.) and
grape (Vitis spp.) and in the hollow stems of large herbs such as dog fennel (Eupatorium
capillifolium). Trees, shrubs, vines and herbs occupied by C. ashmeadi may be in open sites or
under a dense canopy, and may be in dry or wet habitats. Pines growing in mixed hardwood
stands, surrounded by high brush, or thickly covered with vines, are much more likely to be
occupied by C. ashmeadi than C. pinicola. Our extensive collecting experience has shown that the
two species show considerable segregation by habitat across much of Florida, but that zones of
overlap, where pines mix with hardwoods, occur as well. Within these zones it is not uncommon
to find C. pinicola in a large pine tree only a few meters from an oak inhabited by C. ashmeadi.
What is especially noteworthy is that these forms maintain their integrity where they co-occur; we
have no found color intergrades in these overlap areas, and we have not found dark queens with
bicolored workers or bicolored queens with dark workers. Based on these observations, we
conclude that the two forms are reproductively isolated and thus constitute separate, if closely
related species.
Given the natural history described above, it is difficult to imagine a biologically convincing
scenario for regarding these ants as two forms of a single, panmictic species. It would be necessary
to hypothesize that something about the environment of open-grown pines usually (but not
always) causes a developmental shift that makes all members of a colony bicolored. Furthermore,
this shift must not occur in colonies in relatively shaded pines, in open-grown hardwoods or in
open-grown shrubs, vines and weed stems. While not utterly inconceivable, this scenario is
complex and inherently improbable, especially given that there are no known examples of
anything comparable occurring in the Formicidae. Likewise, we find the observations reported by
Johnson (1988) unconvincing as possible objections to regarding the two forms as good species.
Johnson claims to have seen mixed foraging columns in overlap zones and says that colonies
containing both color morphs have been found. In all our extensive collecting experience, we have
never seen a single instance of either phenomenon. The rare occurrence of mixed colonies,
however, is no serious impediment. Mixed colonies containing two non-parasitic ant species are
not unknown, and they can be an accidental byproduct of territorial interactions in which nests are
raided and brood stolen. Johnson (1988) also notes (correctly) that callow workers of C. ashmeadi
are bicolored, with the gaster darker than the head and mesosoma, as in workers of C. pinicola.
We observe that the head and mesosoma of C. ashmeadi callows are grayish in color because they
lack the strong yellow to reddish undertones always present on the head and mesosoma of C.
pinicola workers, regardless of age. As such, they do not offer convincing evidence of
intergradation between the two forms. Note: it is sometimes difficult to assign names to museum
specimens of the C. ashmeadi - C. pinicola complex. Specimens of C. ashmeadi, which appear

blackish in the field, often appear brownish in collections, especially if the specimens have been
stored for several years in alcohol prior to mounting. The gaster may fade somewhat less than the
head and mesosoma, so specimens may appear obscurely bicolored. Specimens of C. pinicola can
fade from bright mahogany-red to a dull brown, converging in color with some specimens of C.
ashmeadi.. The authors cannot at present confidently identify all specimens in collections.
Associations with alates are sometimes useful, as queens seem to show less color change than
workers as they age. We have not found differences in male genitalic structures, so males are no
help in resolving this problem.
Crematogaster pinicola provides evidence that speciation may occur in Crematogaster with
negligible structural divergence. While morphometric studies could possibly reveal small
morphological differences between C. pinicola and C.. ashmeadi, the existence of such
differentiation should not to be presumed. Morphological differentiation is not necessarily an
immediate or even an eventual consequence of speciation, and taxonomists must be careful not to
make finding such differentiation a requirement for assigning species status. In this regard,
Umphrey’s (1996) work on the Aphaenogaster rudis complex is illuminating. He found that some
cryptic species (defined as such by karyotypic differences) could be distinguished from congeners
morphometrically, whereas other karyotypically distinct cryptic species were morphologially
indistinguishable. No doubt, many cases similar to that of C. pinicola and C.ashmeadi will come
to the attention of myrmecologists as our knowledge of the world ant fauna improves.
BIOLOGY
Prior to European settlement, much of the southeastern United States was covered with pine
forests that were structured and maintained by frequent fires (Frost 1993). These forests might be
on wet sites (flatwoods) or dry sites (sandhill) but they were similar in general structure, with a
dense herbaceous ground layer and large, usually widely scattered pines, such as slash pine, P.
elliottii, and longleaf pine, Pinus palustris. Shrubs and woody vines were suppressed by fire, but
persisted in patches that were protected by natural fire breaks. The pines survived the fires by a
series of adaptations, the most important of which (from an ant’s point of view) were the thick,
loose layers of insulating bark at the base of the tree, thick twigs (which are less flammable than
fine twigs), and the tendency for the lower branches and twigs to die and drop off, even when they
receive plenty of light (thus reducing the chance that fire will be carried into the crown of the
tree). Crematogaster pinicola appears to be specialized to endure frequent fires by taking
advantage of the fire adaptations of pines. When the lower twigs and branches begin to senesce,
they are quickly attacked by scavenging insects. At the Archbold Biological Station in central
Florida for example, these are scolytids, such as Pityoborus comatus and Pityophthorus pulicarius
and cerambycids, such as Eupogonius pauper. The activities of the beetle larvae provide a
succession of thick hollow twigs where C. pinicola can become established, even in young trees.
Once established colonies gradually relocate up the trunk as branches die. As a tree matures, there
are occasional dead twigs and branches up in the crown, and the buildup of thick layers of bark
along the trunk provides another nesting area, especially where the outer bark is riddled with the
abandoned galleries of moths. The creation of cavities in the bark by moths has been quantified by
Tschinkel (2002). The thickest bark is at the base of the tree, and in large trees larvae and pupae of
males and queens may be found near the root crown, sometimes below ground level. This brings
part of the colony in contact with the diverse subterranean fauna of southeastern pinelands,
including a large number of potential predators, such as army ants. On the other hand, the
sequestration of these larvae, especially the large queen larvae, at or below ground level may
provide some protection from woodpeckers.
Walter Tschinkel and several associates have published detailed studies of the ecology of a
pine-inhabiting Crematogaster (referred to as C. ashmeadi) in the Appalachicola National Forest
in northern Florida. (Hahn and Tschinkel, 1997, Hess and James, 1998, Baldacci and Tschinkel,
1999, Tschinkel and Hess, 1999, and Tschinkel (2002). Tschinkel (pers. comm.) says the ants
were, “were red and black, and considering their rather particular life cycle and nesting habits,
quite distinct from the all-black species on hardwoods.” Based on this comment, and our own
collections on pines in the same area, we are confident that the ant studied by Tschinkel is C.
pinicola, not C. ashmeadi.
In the Apalachicola National Forest, mating flights of C. pinicola occur in June and July
(Tschinkel 2002). Nest-founding queens regularly occur in abandoned beetle galleries in small
dead branches on pine saplings (Hahn and Tschinkel 1995, Baldacci and Tschinkel 1999). Small
trees, under 7 m tall, are preferred, and trees with more than two dead branches are also preferred
(Baldacci and Tschinkel, 1999). Such trees are far too small to support a mature colony of C.
pinicola, which contains several tens of thousands of individuals (Tschinkel 2002). Workers from
one colony are hostile to workers from other colonies (Tschinkel 2002), and it is possible that it is
safest course of action for founding queens is to found a colony in a tree that is yet unsuitable for
large colonies. Once established, these small founding colonies could then send scouts to find a
large tree that is not already well defended by another colony. It appears that there is never more
than one colony in a tree, and few colonies occupy more than one tree (Tschinkel 2002). Surveys,
using baits and other methods, of trees in class sizes suitable for mature colonies revealed 55 to
almost 90% of the trees were occupied by C. pinicola (Tschinkel and Hess 1999, Tschinkel 2002).
The higher percentage was obtained by more diverse and intensive survey techniques, and is
probably the more accurate figure(Tschinkel 2002). There are no co-dominant arboreal ants in
these pine forests, although several other arboreal ant species may co-occur with C. pinicola
(Tschinkel and Hess 1999).
Crematogaster pinicola may be an important part of the diet of southeastern pine
woodpeckers, especially the endangered red-cockaded woodpecker, whose diet was studied by
Hess and James (1998). In the Appalachicola National Forest they found that C. pinicola
comprised about 43% of the woodpecker’s arthropod diet, a degree of specialization on a single
prey species that may be unique for insectivorous birds in the United States. Many birds show
ephemeral specialization on single species of insects that are at a high point in a population cycle,
but the red-cockaded woodpecker can afford persistent specialization because ant colonies
themselves are abundant, long-lived, and available at all seasons. This ant revises the moral of
Aesop’s fable of the grasshopper and the ant: the improvident grasshopper may vanish in the
winter, but the thrifty and industrious ant can be eaten all year long! The original distribution of C.
pinicola, like that of the red-cockaded woodpecker, was probably centered in the distribution of
the longleaf pine ecosystem. This ecosystem, which once covered approximately 92 million acres,
from the southeastern tip of Virginia to eastern Texas, has been almost completely destroyed (97%
of the old growth forest is gone), and much of what remains is highly fragmented and difficult to
manage with fire (Frost 1993). Crematogaster pinicola may have undergone a major decline with
the reduction of its habitat, but it is definitely not an endangered species. The gross inequalities
imposed by size scale are all in favor of C. pinicola: while a single family group of the red-
cockaded woodpecker requires about 40 hectares of foraging habitat (Hooper 1996), a single
hectare of large pines could support many colonies and thousands of individuals of C. pinicola.
The red-cockaded woodpecker is not threatened by loss of its food supply, but by the lack of
suitable nesting trees, and by its tendency to leave small, remnant patches of pine forest
(Wilson 1992).
ACKNOWLEDGMENTS
This paper is dedicated to Edward O. Wilson, with special appreciation for his participatory
enthusiasm for faunistics and basic taxonomy, an enthusiasm that adds considerable luster to these
important endeavors. This research was supported by the Archbold Biological Station and by the
Museum of Comparative Zoology, Harvard University. The collecting trip to the Apalachicola
National Forest was supported by the Wilson Ant Collection Fund.
LITERATURE CITED
Baldacci, J. & Tschinkel, W.R. 1999. An experimental study of colony-founding in pine saplings
by queens of the arboreal ant, Crematogaster ashmeadi. Insectes Sociaux 46: 41-44.
Bolton, B. 1994. Identification Guide to the Ant Genera of the World. Harvard University Press,
Cambridge, MA. 222 pp.
Buhs, J.B. 2000. Building on bedrock: William Steel Creighton and the reformation of ant
systematics, 1925-1970. Journal of the History of Biology 33: 27-70.
Buren, W.F. 1958. A review of the species of Crematogaster, sensu stricto, in North America
(Hymenoptera: Formicidae). Part I. Journal of the New York Entomological Society 66: 119-
134.
Buren, W.F. 1968. A review of the species of Crematogaster, sensu stricto, in North America
(Hymenoptera, Formicidae). Part II. Descriptions of new species. Journal of the Georgia
Entomological Society 3: 91-121.
Zoology at Harvard College 104: 585 pp.
Deyrup, M. & Trager, J. 1986. Ants of the Archbold Biological Station, Highlands County,
Florida (Hymenoptera: Formicidae). Florida Entomologist 69: 206-228.
Frost, C.B. 1993. Four centuries of changing landscape patterns in the longleaf pine ecosystem.
Proceedings of the Tall Timbers Fire Ecology Conference 18: 17-43.
Hahn, D.A. & Tschinkel, W.R. 1997. Settlement and distribution of colony-founding queens of
the arboreal ant, Crematogaster ashmeadi, in a longleaf pine forest. Insectes Sociaux 44: 323-
336.
Hess, C.A. & James, F.C. 1998. Diet of the red-cockaded woodpecke4r in the Apalachicola
National Forest. Journal of Wildlife Management 62: 509-517.
Hölldobler, B. & Wilson, E.O. 1990. The Ants. Belknap Press of Harvard University Press,
Cambridge, Mass. 732 pp.
Hooper, R.G. 1996. Arthropod biomass in winter and the age of longleaf pines. Forest Ecology
Management 82: 115-131.
Johnson, C. 1988. Species identification in the eastern Crematogaster (Hymenoptera:
Formicidae). Journal of Entomological Science 23: 314-332.
Longino, J.T. 2003. The Crematogaster (Hymenoptera, Formicidae, Myrmicinae) of Costa Rica.
Zootaxa 151:1-150.
Tschinkel, W.R. 2002. The natural history of the arboreal ant, Crematogaster ashmeadi. Journal
of Insect Science 2: 1-15.
Tschinkel, W.R. & Hess, C.A. 1999. Arboreal ant community of a pine forest in northern Florida.
Annals of the Entomological Society of America 92: 63-70.
Umphrey, G.J. 1996. Morphometric discrimination among sibling species in the fulva-rudis-
texana complex of the ant genus Aphaenogaster (Hymenoptera: Formicidae). Canadian
Journal of Zoology 74: 528-559.
Wilson, E.O. 1992. The diversity of life. The Belknap Press of Harvard University Press,
Deyrup & Cover: A new pine-inhabiting Crematogaster
Figure 1. Crematogaster pinicola, worker; length of specimen 3.2 mm.

109
Figure 2. Crematogaster pinicola, male; length of specimen 3.2 mm.

Figure 3. Crematogaster pinicola, alate queen; length of specimen 7.8 mm.

Figure 4. Collection localities for C. pinicola.

Espadaler, X. 2007. The ants of El Hierro (Canary Islands), pp. 113-127. In Snelling, R. R., B. L.
Fisher, and P. S. Ward (eds) Advances in ant systematics (Hymenoptera: Formicidae): homage to
E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute, 80.
THE ANTS OF EL HIERRO (CANARY ISLANDS)
Xavier Espadaler
CREAF and Unit of Ecology
Autonomous University of Barcelona
08193 Bellaterra, Spain.
Xavier.espadaler@uab.es
ABSTRACT
The ants of El Hierro, the smallest and youngest of the Canary Islands, are updated. Twenty-one
species are listed, with seven considered as exotics. Twelve species and six genera are added to the
myrmecofauna of El Hierro. Two endemic new species are described, Temnothorax bimbache sp.n.,
and Monomorium wilsoni sp.n., and Tetramorium depressum Forel is elevated to specific status
(stat. nov.). The population of Plagiolepis (provisionally identified as P. maura) has the interesting
characteristic of having apterous sexuals. Distribution data are presented.
Key words: Hymenoptera, Formicidae, Temnothorax bimbache n. sp., Monomorium wilsoni n. sp.,
taxonomy, distribution.
“…why does a biologist do research at all? To discover, of course” (Wilson 1989)
INTRODUCTION
The Canary Islands consist of seven main islands and six islets, which are the independent tips of an
enormous volcanic mountain range lying under the Atlantic Ocean. Their nearest neighbor is
Morocco, about 95 km (59 mi) east of Fuerteventura. The islands include a huge variety of
landscapes, with cloud forests surrounded by mist, volcanic plateaus, cliffs hit by Atlantic storms,
green pastures and desert volcanic landscapes. The fertile volcanic soils and varied altitude have
combined to create several biological treasures in the Canaries. Roughly a fourth of the islands' 2000
plant species are endemic, including the Canary Island palm, the Canary pine and the recently
described dragon tree, Dracaena tamaranae, an ancient survivor from the last ice age.
Microclimates in the islands allow for great variation in vegetation, from the laurisilva, with lichen-
covered laurels, holly and other broadleaves, to the dry scrublands and semi-desert areas where
saltbush, cactus-like plants and palms grow. In short, the Canary Islands have a high level of
endemicity that is currently receiving much attention (Juan et al., 2000).
The Canaries have a permanent subtropical climate, with mean temperatures ranging from 18°C
(64°F) in winter to 24°C (75°F) in summer. Apart from Lanzarote and Fuerteventura, the northern
side of the islands is subtropical, while the south, including those two islands, is somewhat warmer
and drier. Rain is scarce except on parts of the exposed northern coasts, especially on the northern
side of the more mountainous islands. Lanzarote and Fuerteventura, with no mountains to trap rain
clouds, receive a mean of 150 mm of rain per year. Northeast trade-winds are predominant. Located
27°38' to 27°51'N, and 17°53' to 18°09'W, El Hierro is the smallest (278 km2), the youngest
(1.2×106 years; Guillou et al. 1996) and, jointly with La Palma, the most Atlantic of the Canary
Islands. Its large scale topography, with three large embayments separated by three ridges, is the
result of recent—between 15,000 and 200,000 years ago—giant landslides (Gee et al., 2001). In
spite of those conditions El Hierro has a great and well preserved biological diversity that has
recently (2000) granted it the status of a Biosphere Reserve. More than 58% of the surface is under
protection.
The island's vegetation, constituted from mesic areas of laurel forest to arid areas, may be
summarized as follows, from sea level upwards: A) Halophile coastal zone, dominated by Limonium
pectinatum and Schizogyne sericea and falaises by species from the genera Aeonium. B) The dry
coastal matorral with Euphorbia balsamifera (tabaiba dulce), E. broussonettii (tabaiba amarga), E.
canariensis (cardón) and Rumex lunaria. C) The thermophile juniper forest at El Sabinar with
Juniperus turbinata ssp. canariensis (sabina) as the dominant species, Olea europaea (acebuche),
Maytenus canariensis and Visnea mocanera (mocán). D) The most interesting botanical zone, the
monteverde or laurisilva, on the Northern cliffs hanging above El Golfo, dominated by Myrica faya,
Erica arborea (brezo) and Ilex canariensis, with the more rare Laurus azorica and Picconia excelsa.
E) Huge pine forests on the southern slopes, with Pinus canariensis, Chamaecytisus proliferus and
Adenocarpus foliolosus. Lastly, agricultural land and high mountain pastures are to be found on
rural areas. From its rich flora, new taxa in higher plants are still being described (Scholz et al.,
2000; Gaisberg & Wagenitz, 2002). Endemics are abundant (Juan et al., 2000; Izquierdo et al.,
2001). Among vertebrates, the very rare giant (up to 60 cm) lizard (Gallotia simonyi machadoi), the
“lagarto del Salmor”, is in danger of extinction. An ongoing captive breeding program near
Frontera is allowing its reintroduction.
Ants from El Hierro have not received any special attention and only nine species have been
noted: Crematogaster alluaudi (Wellenius, 1955: Frontera); Temnothorax gracilicornis nivarianus
(Wellenius, 1955: Frontera); Tetramorium semilaeve depressum (Wellenius, 1955: Guarazoca, El
Pinar, Frontera); Cardiocondyla mauritanica Forel (Seifert, 2003: without locality); Linepithema
humile (Wellenius, 1955: Valverde); Plagiolepis barbara canariensis (Wellenius, 1955: Guarazoca,
El Brezal, Frontera); Camponotus hesperius (Wellenius, 1955: Valverde, Guarazoca, El Pinar, El
Espadaler: Ants of El Hierro Island 115
Brezal, Frontera); Camponotus feai (Wellenius, 1955: Valverde; Barquín, 1981: La Lajura, La
Restinga); Camponotus guanchus (Wellenius, 1955: Guarazoca, Frontera; Cagniant & Espadaler,
1993: El Sabinal).
I have had the opportunity to visit twice this interesting island, one week in spring 1998 and one
week in winter 2003. As a result, 12 species and 6 genera are added to its myrmecofauna. Two
species are proposed as new and the status of the remaining taxa is discussed. A third of the species
are ant exotics.
I enjoyed ant searching by a classical entomological survey, when visiting all vegetation zones.
Nests were located under stones and by checking rock crevices when possible with a small pickax—
volcanic rocks are extremely hard!—tree stumps and breaking dead wood remains, fallen logs or
small branches. At urban localities, I looked at irrigated gardens, flower pots or next to accumulated
organic debris. Recent surveys of ants from the other Macaronesian archipelagos of Azores
(Wetterer et al., 2004), Madeira and Cape Verde (Wetterer et al.; in preparation) indicate that the
non-native component of the myrmecofauna is very abundant in urban habitats. Complete societies
of interesting species were collected by careful digging and maintained in artificial nests. Ants were
fed artificial diet (Bhatkar & Whitcomb, 1970), supplemented with cut Tenebrio larvae, until sexuals
were eventually produced. Vouchers will be deposited at the following institutions: Museum of
Comparative Zoology (MCZC) (Cambridge, USA), Natural History Museum of Los Angeles
County (LACM) (Los Angeles, USA) and Museo de Ciencias Naturales (MCNC) (Tenerife, Spain).
A check list, taxonomic comments, nomenclatural changes and descriptions of new species follow.
Generic names adhere to recent taxonomic arrangements by Bolton (2003). Measurement and
indices are as in Bolton (1982), except that ML (mesosomal length) is used rather than AL (alitrunk
length). Propodeal spine development (Buschinger’s index; Buschinger, 1966) is measured as the
maximum length from center of propodeal spiracle to the tip of spines/minimum length from center
of propodeal spiracle to the vertical, declivitous face of propodeum as seen in profile.
The holotypes of the two new species described below are currently deposited in my personal
collection. It is my intention that the entire collection will ultimately be deposited in a public
institutional collection.
Localities. Names are from the Kompass map #242 El Hierro, Edition 01-02 (1:30000). Data
from 2003 indicate geographical coordinates by a Garmin Summit eTrex GPS; geographical
coordinates from 1989 are back transformed from UTM coordinates, using GeoTrans 2.2.5.
1. Arenas Blancas (N27º46.051’ W18º7.339’). 20 m. Halophile vegetation. 30.03.1989;

17.12.2003.
2. Between Echedo and Charco Manso (N27º48.98’ W17º35.69’). 200 m. Dry coastal matorral,
with Euphorbia obtusifolia. 28.03.1989.
3. Caldereta del Tabaibal Manso (N27º43.14’ W18º8.02’). 480 m. Succulent scrub, with
Euphorbia regis-jubae. 30.3.1989.
4. Binto (N27º43.20’ W18º5.29’). 950 m. Pine forest. 30.3.1989.
5. El Brezal (N27º43.856’ W17º59.709’). 1275 m. Laurisilva. 31.3.1989; 18.12.2003.
6. El Fayal (N27º43.85’ W18º0.44’). 1300 m. Myrica, Erica, Urtica, very humid. 31.3.1989.
7. El Julán (N27º42.77’ W18º2.54’). 1000 m, transition to cultivated fields. Pines and fig trees.
18.12.2004.
8. El Julán (N27º42.80’ W18º1.26’). 1100 m. Very dry pine forest. 29.3.1989.
9. El Morcillo (N27º43.59’ W17º59.95’). 1250 m. Pine forest with some very old trees. 29.3.1989.
10. El Sabinal (N27º44.356’ W18º7.366’). 695 m. Degraded juniper forest. 17.12.2003.
11. El Sabinal (N27º44.917’ W18º7.729’). 570 m. Very well preserved juniper forest. 2.4.1989;
17.12.2003.
12. Ermita de la Peña (N27º48.091’ W17º58.779’). 760 m. Rural, mountain pastures. 16.12.2003.
13. Hoya de Fileba (N27º44.812’ W 17º59.425’). 1280 m. Laurisilva dominated by Erica.
19.12.2003.
14. La Caleta (N27º48.040’ W17º53.184’). 20 m. Dry coastal matorral next to the small town.
16.12.2003.
15. La Frontera (N27º45.170’ W18º0.727’). 275 m. Town, streets and public gardens. 20.12.2003.
16. Ladera Cabello (N27º44.397’ W18º4.727’). 215 m. Ruderal vegetation. 17.12.2003.
17. Llano de Guillén (N27º43.29’ W18º0.97’). 1200 m. Open pine forest, humid, many grasses.
18.12.2003.
18. Los Jables (N27º41.93’ W18º1.60’). 600 m. Track. 29.3.1989.
19. Los Letreros (N27º42.147’ W18º2.141’). 630 m. Ruderal vegetation with scattered pine trees.
18.12.2003.
20. Los Llanillos (N27º45.170’ W18º1.318’). 305 m. Town streets. 20.12.2003.
21. Mercadel (N27º42.642’ W18º1.568’). 1010 m. Pine forest, humid. 18.12.2003.
22. Mirador de Isora (N27º44.35’ W17º57.11’). 850 m. Ruderal vegetation with Hypericum
canariensis, Ficus, Rumex lunaria. 1.4.1989.
23. Mirador de las Playas (N27º43.801’ W17º58.515’). 1080 m. Pine forest. 16.12.2003.
24. Mocanal (N27º50.16’ W17º56.35’). 200 m. Rumex lunaria, Aeonium sp. “picón”. 28.3.1989.
25. Montaña de la Fuente (N27º44.805’ W17º59.430’). 1310 m. Laurisilva with Erica. 20.12.2003.
26. Montaña de La Limera (N27º48.80’ W17º56.62’). 780 m. Degraded fayal-brezal with Apollonia
canariensis (barbusano) and ruderal vegetation. 28.3.1989.
27. Montaña de las Cuevas (N27º43.97’ W 18º7.14’). 720 m. Pastures with megaforbs. 30.3.1989.
28. Montaña de los Charcos (N27º45.733’ W18º8.367’). Dry coastal matorral. 17.12.2003.
29. Montaña del Gajo (N27º43.812’ W17º59.489’). 1280 m. Transition from laurisilva to pine
forest. 19.12.2003.
30. Montaña Hoya de la Vaca (N27º46.021’ W 17º58.511’). 1180 m. Rural, mountain pastures.
16.12.2003.
31. Montaña la Empalizada (N27º41.953’ W18º1.175’). 770 m. Ruderal vegetation. 18.12.2003.
32. Montaña Tembárgena (N27º43.44’ W 18º6.51’). 750 m. 30.3.1989.
33. Pico del Malpaso (N27º44.33’ W 18º2.89’). 950 m. Laurisilva. 2.4.1989.
34. Pie del Risco, Frontera (N27º45.91’ W17º59.70’). 150 m. Ruderal, Opuntia ficus-indica.,
Nicotiana glauca. 31.3.1989.
35. Playa de las Almorranas (N27º43.858’ W17º56.588). 2 m. Rocky beach, halophile vegetation.
1.4.1989; 19.12.2003.
36. Pozo de las Calcosas (N27º50.355’ W17º56.870’). 125 m. 16.12.2003. Highly degraded coastal
matorral near the town.
37. Puerto de la Estaca (N27º47.169’ W17º54.031’). 2 m. Harbor. 19.12.2003.
38. Punta de la Dehesa (N27º46.150’ W18º7.621’). 25 m. Halophile vegetation. 30.3.1989;
17.12.2003.
39. Punta del Bajío (N27º46.73’ W17º54.30’). 20 m. Periploca laevigata, Senecio kleinia.
1.4.1989.
40. Punta del Guincho (N27º44.22’ W17º55.46’). 50 m. Abandoned fields with Euphorbia
canariensis, Limonium, Astydamia, Periploca. 1.4.1989.
41. Punta Grande (N27º47.859’ W17º59.434’). 20 m. Halophile and ruderal vegetation and
bungalow gardens. 31.3.1989; 20.12.2003.
42. La Restinga (N27º38’ W17º59.425’). 5 to 20 m. Fishermen small town, streets. Rocky coastal.
18.12.2003.
43. Tacorón (N27º38.234’ W17º 59.428’). 250 m. Dry coastal matorral. 18.12.2003.
44. Tamaduste (N27º48.131’ W 17º53.178’). 30 m. Town streets. 15.12.2003.

45. Valverde (N27º49.120’ W17 º54.950’). 600 m. Town streets and gardens. 15.12.2003.
CHECK LIST
Twenty-one ant species were found, with the Myrmicinae being the dominant subfamily with
11 species, followed by the Formicinae with 6 species and Ponerinae and Dolichoderinae with two
species each. Numbers after the species name indicate localities where found and collected castes: w
(workers), q (queens), m (males). (*) indicates a first record for El Hierro.
SUBFAM. PONERINAE
1. Hypoponera ragusai (Emery). (*) (35, w). A nest with 13 workers was recovered in the humid
spot under a stone, a few meters from the sea, during the 1989 visit. No nest was found in 2003. The
morphology matches the drawings of the mesosoma profile of H. ragusai var. santschii Emery
(Emery, 1909, without a mesopropodeal depression, although the petiole is more akin to the petiole
of H. ragusai. The whole genus is in need of revision.
2. Hypoponera eduardi (Forel). (*) (26, w). A few workers of this widespread Mediterranean
species were collected under a stone on a slope of volcanic charcoal (“picón”).
SUBFAM. DOLICHODERINAE
3. Tapinoma melanocephalum (Fabricius). (*) (41, w). Running workers were detected in irrigated
gardens next to recently built bungalows. This is the first record for El Hierro of this well known
tramp species.
4. Linepithema humile (Mayr). (2, w; 8, w, m; 15, w; 23, w; 26, w; 31, w; 41, w; 44, w; 45, w). The
Argentine ant is known from all the Canary Islands (Espadaler & Bernal, 2003). At El Hierro it
occupies habitats from next to sea level to one thousand meters, in pine forests. Confronted with the
two populations known to exist in North Mediterranean Europe (Giraud et al., 2002), the Argentine
ants from El Hierro showed aggressiveness towards the “Catalan” population and reacted peacefully
towards the “Main” population from mainland Europe. Aggression tests (one to one worker; five
replicates) were run with two samples from El Hierro (La Frontera; Mirador de las Playas). I
conclude that both samples from El Hierro belong to the genotypic profile of the “Main” population,
the more abundant in Western Mediterranean Europe.
SUBFAM. MYRMICINAE
5. Cardiocondyla emeryi Forel (*) (37, w; 41, w). This widespread tramp species was detected only
at two seaside small towns, on poorly attended gardens.
6. Cardiocondyla mauritanica Forel (1, w; 3, w, q; 11, w; 20, w; 38, w; 41, w, q). From 20 to 570
m, open dry habitats.
7. Crematogaster alluaudi Emery (1, w; 3, w; 11, w; 12, w; 16, w; 28, w; 34, w; 36, w; 39, w; 40,
w). From 20 to 760 m. The specimens from El Hierro are somewhat more pilose and the pubescence
on the head, gaster and legs is more detached, than in populations from Tenerife. The enhanced
pilosity and pubescence is also present in samples from the island of La Palma. This species nests in
dead shrubs and is rather inconspicuous unless one happens to touch the vegetation or breaks their
nest: then they rush out and come to any available surface, fiercely biting the skin. On examining the
surface of leaves of Aeonium sp. (Crassulaceae) plants at Ladera Cabello, I found dead ants attached
to the viscous surface. Those crassulaceous plants are a trap for flying insects: seven males of C.
alluaudi, one of Solenopsis canariensis, one possibily of Temnothorax bimbache and one queen of
the big Camponotus hesperius were recovered from a few plants, in addition to many small flies.
Crawling insects may also be trapped, as shown by the capture of two workers of Camponotus
guanchus and one small carabid beetle.
8. Monomorium wilsoni, new species Figures. 4, 5, 6
Holotype queen: TL 4.7, HL 0.98, HW 1.05, CI 106, SL 0.82, SI 78, PW 0.66, ML 1.49.
Queen description (Figures 5, 6). Clypeus with the anterior margin feebly concave, without
differentiated teeth. Clypeal carinae very weak. Frontal lobes with longitudinal superficial striae,
present also between the base of mandibles and the eyes. Mandibles with four teeth. Head broader
than long. Sides of head rounded. Antennae with 12 segments and the scape reaching the occiput.
The posterior margin is concave, depressed in dorsal view. Eye maximum diameter 0.21-0.25, about
0.20-0.23 x HW, with 15-18 ommatidia in the longest row. Eyes with curved anteriorly pilosity of
0.04-0.05 mm. Mesosoma apterous. Sides of pronotum visible in dorsal view. Promesonotal dorsum
in profile see fig. Scutellum saddle-shaped, rounded in profile. No traces of tegulae in the > 25
queens from 10 nests examined. Seen from above, petiole width 0.40-0.46, in profile with a
posterior dorsal node two-faced. Postpetiole width 0.44-0.52, in profile with a vaulted node and a
short ventral point. Whole body with a long pilosity. Whole body shining, with alutaceous
microsculpture. Sides of propodeum and katepisternum with a fine reticulum. Declivitous face of
propodeum transversely striate. General color brown. Mandibles, pronotum, anepisternum and
metanotum reddish, contrasting with the brownish propodeum. Gaster blackish brown.
Worker description (Fig. 4). Clypeus as in the queen. Mandibles with 2-4 poorly developed
longitudinal striae and four teeth, decreasing in size from apex to base. Palp formula 2, 2. Head
longer than wide. Sides of head very slightly curved; vertex margin straight or very feebly concave.
Scape surpassing the posterior border. Eye maximum diameter 0.12-0.14, about 0.23-0.27 x HW,
with 9-11 ommatidia in the longest row. Promesonotal dorsum in profile with a well marked
metanotal groove. Declivitous face of propodeum shallowly concave. Petiole higher than
postpetiole. Anterior border of clypeus with a middle seta and 3-4 pairs of diminishing size towards
the lateral margins. Median portion of clypeus with one pair directed anteriad and two pairs directed
upwards. Dorsum of head with two pairs of standing hairs: one pair situated at the distal end of
frontal lobes and one pair in midline of the head. Sometimes, a pair of shorter hairs is present
between this last pair and the posterior margin. Posterior margin and dorsal mesosoma without
standing hairs. Underside of head with 6-10 hairs of variable length. Petiole without hairs;
postpetiole with 2-4 hairs directed posteriad. First gastral tergite with 0-2 hairs in front of the apical
row. Dorsum of head shining, with piligerous pits and a very superficial alutaceous sculpture.
Mesosoma shining. Sides and dorsum of propodeum, entire mesopleura and posterior half of
propleura with visible reticulation that becomes increasingly superficial towards the anterior part of
mesosoma. Body color brown with mandibles, antennomeres 3-12, lower half of petiole and
postpetiole distinctly brownish red.
Additional paratype measurements, Workers: TL 2.4-2.7, HL 0.63-0.70, HW 0.50-0.57, CI 79-
82, SL 0.57-0-62, SI 107-113, PW 0.32-0.36, ML 0.74-0.82. Queens: TL 4.7-5.1, HL 0.97-1.01, HW
1.01-1.06, CI 104-108, SL 0.82-0.84, SI 77-81, PW 0.64-0.68, ML 1.43-1.51.
Holotype queen: Spain, Canary Islands, El Hierro, Ermita de la Peña (N27º48.091’ W17º58.779’).
760 m. Rural, mountain pasture. 16.12.2003. (X. Espadaler), presently deposited in my personal
collection (XEPC). Paratypes. 16 workers, 8 queens, all same data as holotype. In eight pins, each
pin with 2 workers, one female (queens), deposited in LACM, MCNC, MCZC and XEPC.
Additional specimens examined: 1, w; 3, w, q; 4, w, q; 8, w, q; 9, w; 10, w; 11, w, q; 12, w, q; 17,

w; 18, w; 19, w, q; 21, w; 22, w; 28, w, q; 29, w, q; 32, w; 39, w, q; 41, w, q
Etymology. This species is named in honor of, and gratefulness to, Prof. Edward Osborne Wilson.
Remarks. The species is a member of the salomonis-group as defined by Bolton (1987). The closest
relatives of this species belong to the whole group of endemic Monomorium from the Canary
Islands, which have differentiated into a minimum of five species (unpublished observations).
Specific differences are best appreciated in the apterous queen caste, in the density and length of
pilosity, shape of mesosoma, petiole and postpetiole, and biometry. In the apterous queens of the
species from the Canary Islands a cline can be established in which the more dense the pilosity, the
shorter is, and the more advanced are queens in the stage towards an ergatoid condition. Variation in
pilosity is best appreciated in head profile, petiole and postpetiole in dorsal view. The most modified
species is M. hesperium Emery, in which queens are the smallest among the group in the Canary
Islands and are next to completely smooth and shining and hairs are very long and comparatively
sparse. M. medinae Forel, instead, is the most densely pilose but hairs are very short and head
sculpture is more developed, as is also in the mesopleurae. M. wilsoni is intermediate in pilosity
density and length. The head is wider than in M. medinae or M. hesperium. The mesosomal profile,
especially the propodeum, the development of scutum and scutellum (side view) and the form of the
strange petiole and postpetiole are also characteristic for each species. This peculiar modified
thoracic structure is known in other Monomorium species and has been related to nest founding by
budding (Bolton, 1986). Material is being collected for a revision of the entire group of endemic
Macaronesian Monomorium.
Natural history. This is one of the most abundant ants at El Hierro. I collected it from sea level and
up to 1300 m, under stones in open, dry, coastal habitats and up to the humid pine and laurel forests.
This wide ecological plasticity is found also in its congener M. carbonarium in Madeira (unpub.
observ.) and the Azores (Wetterer et al., 2004). Highly distinctive, the huge numbers of dark and
shining workers attack and sting fiercely. No special skin reaction follows. The species is
polygynous.
9. Monomorium subopacum (F.Smith) (*) (22, w q; 34, w; 35, w q m; 37, w; 40, w). From sea level
to a single outpost at 850 m, the Mirador de Isora, a highly degraded site.
10. Pheidole teneriffana Forel (*) (37, w). A nest with winged males was detected on cracks in the
pavement. Ants rushed out after a small air blow with the aspirator.
11. Solenopsis canariensis Forel (*) (2, w; 6, w q; 10, w; 11, w; 16, w; 18, w; 39, w). From 20 to
1300 m, cryptic and underground nesting species. The whole group of small yellow species of
Solenopsis is in need of a revisionary study. The small, smooth, shining and morphologically
convergent workers are extremely similar and sexuals seem to be the proper phenotype on which to
base a sound taxonomy.
12. Temnothorax gracilicornis (Emery) (5, w q; males eclosed in the laboratory). Three colonies
from this Canarian endemic were collected: two were nesting under mosses on a stone; the third was
nesting under a small stone. The morphology and coloration of workers and queens fits the
description of L. gracilicornis, not of the darker and shinier var. nivarianus Santschi, that Wellenius
(1955) collected at Frontera. As I have not seen type material of both taxa and samples from
Tenerife show varying coloration—from yellow to brownish, thus including both names—and body
surface shininess, I refer the samples of El Hierro to the nominal species. This is not a formal
proposal of synonymy.
13. Temnothorax bimbache new species Figures 1, 2, 3
Holotype worker: TL 3.7, HL 0.89, HW 0.73, CI 82 , SL 0.78, SI 106, PW 0.52, ML 1.14.
Worker description (Figs. 1, 2). Mandibles with fine longitudinal sculpture. Palp formula 5, 3.
Clypeal margin convex and with a broad medial band smooth and shining. Median clypeal carina
absent. Eyes large, maximum diameter 0.20-0.26, about 0.29-0.33 x HW, with 16-19 ommatidia in
the longest row. Micropilosity present. Head in full-face view oval-shaped. Vertex margin slightly
convex, with a slightly projecting rim visible in full-face view; posterolateral corners rounded. With
mesosoma in profile the promesonotum evenly convex, the metanotal area with a shallowly
impressed groove; propodeum with two long spines. Buschinger’s index 3.6. Femora strongly
inflated. Petiole in profile with a slender neck and a broadly rounded and big node. An acute and
long, directed anteriorly, subpetiolar process. Postpetiole in profile rounded and with a short
posterior neck. In dorsal view the petiole node is oval-shaped, longer than wide. Postpetiole wider
than long. Head sculpture: widely spaced longitudinal rugulae, with spaces smooth and shining or
with vestiges of ground-sculpture. Rugulae reaching the vertex. Seven rugulae between frontal
carinae. The area behind the eyes shows some irregular cross-meshes. Dorsum of promesonotum
with a few longitudinal rugae, the space between them smooth and shining. Sides of mesosoma with
irregular longitudinal rugae and a fine reticulum between them, but otherwise shining. Metanotal
and propodeal dorsum without rugae, smooth and shining. Petiole with a pair of short rugae in the
declivity to the neck and with a fine superficial shagreening, also present in the postpetiole. Gaster
smooth and shining but for the hair pits. All dorsal surfaces of head and body with slightly curved or
straight hairs wich are shorter on the head. Appendages without such long hairs. Longest pronotal
hair 0.12. Bicolored, with the head and gaster deep brown. Legs brownish, contrasting with the
orange-reddish mesosoma. The mesopleurae vary from pure red to brownish. Mandibles, antennae,
orange. Petiole orange to brownish. Pospetiole brownish.
Queens (Fig. 3) follow the usual differences from workers. A main variation is the petiole shape,
rather triangular in profile. The anterior medial zone of the scutum is infuscated, as are the two
zones proximal to the tegulae.
Additional paratype measurements. Workers: TL 3.4-4.1, HL 0.88-0.96 , HW 0.70-0.80, CI 78-

85, SL 0.70-0.76 , SI 94-103, PW 0.50-0.56, ML 1.09-11.19. Queens: TL 4.9-5.5, HL 0.96-1.02,
HW 0.84-0.92, CI 87-90, SL 0.76-0.80, SI 85-90, scutum width 0.92-1.02, ML 1.53-1.69.
Holotype worker: Spain, Canary Islands, El Hierro, Punta de la Dehesa, 50m, N27º46.150’ W
18º7.621’, 17 December 2003, under small volcanic stone (X. Espadaler), presently deposited in my
personal collection. Paratypes. 32 workers, 16 queens, all same data as holotype. On sixteen pins,
each pin with 2 workers, one female (queens), deposited in LACM, MCNC, MCZC, XEPC.
Additional specimens examined: 16, w; 38, w q.
Etymology. The species is named after the “bimbaches”, as were named the former aboriginal
people, cave-inhabitants of El Hierro.
Remarks. The closest relative of L. bimbache, which shows the same bicolored pattern, is L. risi
Forel, found in Tenerife and Gran Canaria. This last species is larger and has much shorter spines, a
stouter petiole node and less shining head surface sculpture. Another related species is L. neminan
Espadaler, from the Anti-Atlas mountains in Morocco, that has similarly long spines but is smaller,
and a dull surface sculpture and distinctly lower petiole node.
Natural history. Four nests were found under blackish, porous, volcanic, small stones. Nest
situation showed a peculiar orientation: the stones were protected by vegetation and placed in the
northern face. The ants occupied the first two cm of soil but also the holes and voids of the volcanic
stone. Winged females eclosed at the laboratory in April and May 2004.
14. Tetramorium caldarium (Roger) (*) (37, w; 41, w). This tramp ant was collected only from two
urban sites.
15. Tetramorium depressum Forel (new status) (3, w; 4, w; 9, w; 11, w; 13, w; 14, w; 22, w; 26, w;
27, w; 30, w; 33, w; 39, w; 40, w; 41, w). Forel ((1892) described this ant as a subspecies of T.
semilaeve André. In addition to the head surface differences already noted by Forel, both taxa have
different male genitalia and head worker sculpture and are here formally proposed as distinct
species. From 20 to 1300m although big, populous nests are found at the higher altitudes, usually
under stones. The few nests located in the drier coast seemed to be in a miserable and suffering state,
with a small number of workers. But for a smaller size the morphology of lowland populations was
indistinguishable from other populations.
SUBFAM. FORMICINAE
16. Camponotus feai Emery (3, w; 8, w q; 14, w, q; 18, w; 22, w q; 28, w q; 35, w q; 36, w; 38, w;
39, w; 40, w; 41, w; 42, w; 43, w). Low to mid altitude species: sea level to 1000m (a southern, very
dry slope). All populations from El Hierro belong to the homogeneous black phenotype; some
populations from other islands show a remarkable variation in color of head and thorax, from pure
black to a splendid red.
17. Camponotus hesperius Emery (2, w q; 5, w; 6, w q; 9, w; 10, w q; 11, w; 12, w; 13, w q; 16, w;
24, w q; 25, w; 27, w). This big and beautiful shining species is usually found from mid to high
altitude, up to 1300 m, the lowest point at 200 m. Usually soil nesting, it can be found under stones
also in very humid spots on volcanic charcoal.
18. Camponotus guanchus Santschi (11, w q m; 16, w m). Nests were found in dead wood and rock
crevices at the thermophile juniper forest. In December 2003, winged queens and males were
present inside the nests. This Canarian species is known from the islands of Tenerife, La Palma and
El Hierro.
19. Paratrechina longicornis (Latreille) (*) (14, w; 20, w; 34, w q; 37, w; 41, w; 42, w; 44, w). This
lowland species is found only in urban parks or suburban dirty habitats.
20. Paratrechina jaegerskioeldi (Mayr) (*) (41, w). Workers were collected running amid the steps
in the recently built bungalows where Tapinoma melanocephalum was also recovered. It was
previously known from the island of Tenerife (Espadaler & Bernal, 2003).
21. Plagiolepis maura Santschi cf. (1, w q m; 18, w; 22, w q; 28, w q; 31, w q; 34, w m; 35, w; 36,
w q; 37, w; 38, w; 39, w; 41, w q; 44, w). This is a frequent ant at low altitudes although it was
collected also at 600m and 850m. Nests are in soil, usually under stones or small wood debris. This
identification is provisional. I have not used the name P. barbara canariensis, used by Wellenius
(1955) for his samples from El Hierro, because males and females of this species are much larger; in
addition, the conspecificity of this last taxon with P. schmitzii is highly probable (unpublished
observations). This species has been exceedingly difficult to deal with. The West Mediterranean
Plagiolepis sorely need a complete revision, as what follows attest. The key characteristic is aptery
in sexuals: in early spring, small males were present in a nest at Arenas Blancas, running very
rapidly. Its apterous aspect, suggested at first a social parasite. Thereafter, as all queens from El
Hierro showed also to be apterous, their true nature was evident. Sexuals are not ergatoid: but for the
lack of wings they have the characteristic male and gyne morphology. At Pié del Risco, one fully
winged male was collected. This is the first free-living formicine ant in which wingless males have
been noted. After checking material from other islands, it was soon clear that this is best interpreted
as a case of wing polymorphism: in queens from the islands of Gran Canaria and La Gomera there is
a variable degree of thoracic simplification, from a complete winged thorax to an apterous
morphology, lacking tegulae and with a fused scutum and scutellum. An enhanced pilosity in
pronotum and scutellum seems to be correlated with thorax simplification.
Aptery in ant males and queens has appeared independently several times under distinct
selective pressures and its functional significance is also diverse (Heinze & Ysuji, 1995; Espadaler
1997). Two other species in the genus Plagiolepis are known to have wingless males (P. xene
Stärcke and P. ampeloni (Faber) or wingless females (P. ampeloni) although this is probably related
with the parasitic life of both species. It is legitimate to speculate on wing loss though none of the
hypotheses exposed in Wagner & Liebherr (1992) seems to apply in this case. Habitat stability is not
applicable because of the recent volcanic eruptions and large-scale landslides at El Hierro (Gee et
al., 2001). Habitat isolation neither seems to be appropriate as the numerous sites where this species
has been collected attest. The energetic cost of flight is also to be rejected: El Hierro has not a cold
climate and, even if the prevailing trade winds, from the Southwest are high, this admittedly ad-hoc
explanation is invalidated by the fact that the similarly sized males and females of Solenopsis
canariensis have retained full wings. The possibility of winged queens at El Hierro cannot be
rejected.
DISCUSSION
El Hierro is the youngest of the Canary Islands. Thus, one would expect it to be the island with
a lesser degree of endemic species. Instead, as Izquierdo et al. (2001: 23) show, it has the highest
percentage of endemics, whether considering all terrestrial species of the whole Canary Islands
(24.6%) or only the Insecta (40.6%). It has a similar level of endemic vascular flora (18.4%) as La
Palma, Gomera or Tenerife. On the contrary, if we consider only the subset of Canarian endemics,
and the endemics strictly found in a single island, El Hierro has the lowest percentage (15.7) of this
last selective group. This seemingly contradictory pattern is probably caused by its young age and
by the paucity of colonization events given the main direction of the southwest blowing trade winds.
Natives and exotics

Seven ant species I consider as exotics on El Hierro:,Cardiocondyla emeryi, Pheidole
teneriffana, Tetramorium caldarium, Paratrechina longicornis, P. jaegerskioeldi, Tapinoma
melanocephalum, and Linepithema humile. But for the last species, the other are new records for the
island; not a good prospect. Fortunately, those exotics are restricted to heavily disturbed habitats, the
streets and gardens, with the exception of the Argentine ant that, in addition, can occupy habitats
rather well preserved, as the extensive southern pine forests in the surroundings of El Pinar.
Probably only this last species is of some concern in conservation terms.
Future research
It is difficult to ascertain the completeness of any inventory if, as mine, it is based on a non-
systematic and non-quantitative sampling scheme. The ant species number of El Hierro, given the
small island size, is probably rather complete in the native complement; exotic species such as
Monomorium pharaonis or Pheidole megacephala, already known from other Canary Islands
(Espadaler & Bernal 2003), the Azores (Wetterer et al., 2004) or Madeira (unpublished
observations), can arrive haphazardly and establish in urbanized habitats. A few taxonomic
problems remain: the specific identity of Plagiolepis, with the aptery problem at its center, deserves
an all Canary Islands sampling program, in concert with a revision of the West-Mediterranean
Plagiolepis. The population of Hypoponera ragusai is a taxon still to be settled, pending a revision
of Hypoponera. The proper evaluation of the morphological differences of Crematogaster alluaudi
merits intensive collecting as El Hierro, in spite of being the youngest island, may have already
produced some genetic segregation. Pestano et al., (2003) found substantial molecular
differentiation in populations of some bat species from El Hierro as compared with bats from other
Canary Islands. Lastly, the finding of the male caste in Monomorium wilsoni would allow a
complete specific description.
The laurisilva, a most interesting but poorly studied habitat is somehow deceptive in its species-
poor myrmecofauna. I have had the same deceiving impression when visiting other Macaronesian
islands. This scarcity is probably due to the dense canopy that intercepts nearly all incoming
radiation (Aschan et al., 1994) thus rendering difficult ant nesting.
The endemic Monomorium from the Canary Islands offer an interesting specific diversification
on which to establish morphological and molecular phylogenies, and deduce sequential colonization
of the islands, as did Juan et al., (1996) with the beetle genus Hegeter.
ACKNOWLEDGEMENTS
Rafael Rodríguez and the late Cristina Vicente were kind companions during the first field excursion
in 1989. Thanks are due to Pere Oromí (La Laguna, Tenerife) for a critical reading of a first draft.
Partial funding was provided by the National Geographic Society and private funds.
LITERATURE CITED
Aschan, G., Jiménez, M.S., Morales, D. & Lösch, R. 1994. Aspectos microclimáticos de un
bosque de laurisilva en Tenerife. Vieraea 23: 125-141.
Barquín, J. 1981. Las hormigas de Canarias. Taxonomía, ecología y distribución de los Formicidae.
Colección Monografías (Secr. Publ. Univ. La Laguna) 3: 1-584.
Bhatkar, A.P. & Whitcomb, W.H. 1970. Artificial diet for rearing various species of ants. Florida
Entomologist 53: 229-232.
Bolton, B. 1982. Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax,
Melissotarsus, Messor and Cataulacus (Formicidae). Bulletin of the British Museum (Natural
History) Entomology 45: 307-370.
Bolton, B. 1986. Apterous females and shift of dispersal strategy in the Monomorium salomonis-
group. Journal of Natural History 20: 267.272.
Bolton, B. 1987. A review of the Solenopsis-genus group and a revision of the Afrotropical
Monomorium Mayr (Hymenoptera: Forrmicidae). Bulletin of the British Museum (Natural
History) Entomology 54: 263-452.
Buschinger, A. 1966. Leptothorax (Mychothorax) muscorum Nylander und Leptothorax (M.)
gredleri Mayr, zwei gute arten. Insectes Sociaux 13: 165-162.
Cagniant, H. & Espadaler, X. 1993. Camponotus guanchus Santschi, 1908, stat. nov. et études des
populations de Camponotus sicheli Mayr, 1866 (Hymenoptera, Formicidae). Journal of African
Zoology 107: 419-438
Espadaler, X. 1982. Xenhyboma mystes Santschi, 1919 = Monomorium medinae Forel, 1892.
Évidence biologique de la synonymie (Hymenoptera, Formicidae). Nouvelle Revue
d’Entomologie (N.S.) 12: 111-113.
Espadaler, X. 1997. Leptothorax caesari sp. n. (Insecta: Hymenoptera: Formicidae), a granivore
with apterous males. Annalen der Naturhistorisches Museum Wien B. Bot. Zool. 99: 145-150.
Espadaler, X. & Bernal, V. 2003. Exotic ants in the Canary Islands (Hymenoptera, Formicidae).
Vieraea 31: 1-7.
Emery, C. 1908. Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes.
(Hym.) Teil VIII. Deutsche Entomologische Zeitschrift 1909: 355-376.
Gaisberg, M. v. & Wagenitz, G. 2002: Carduus baeocephalus subsp. microstigma Gaisberg &
Wagenitz, a new subspecies from the Canary Islands (Cardueae, Carduinae) – a facultative
autogamous descendant of Carduus baeocephalus Webb subsp. baeocephalus. Candollea 57:
271-282.
Gee, M.J.R., Watts, A.B., Masson, D.G. & Mitchell, N.C. 2001. Landslides and the evolution of
El Hierro in the Canary Islands. Marine Geology 177: 271-293.
Giraud, T., Pedersen, J.S. & Keller, L. 2002. Evolution of supercolonies: the Argentine ants of
southern Europe. Proceedings of the National Academy of Sciences of the United States of
America 99: 6075–6079.
Guillou, H., Carracedo, J.C., Torrado, F.P., & Badiola, E.R. 1996. K-Ar ages and magnetic
stratigraphy of a hotspot-induced, fase grown oceanic island-El Hierro, Canary Islands. Journal
of Volcanology and Geothermal Research 73: 141-155.
Heinze, J. & Tsuji, K. 1995. Ant reproductive strategies. Research in Population Ecology 37: 135-
149.
Izquierdo, I., Martin, J.L., Zurita, M. & Arechavaleta, M. (eds.). 2001. Lista de especies
silvestres de Canarias (hongos, plantas y animales terrestres). Consejería de Política Territorial
y Medio Ambiente. Gobierno de Canarias. 437 pp.
Juan, C., Oromí, P. & Hewitt, M. 1996. Phylogeny of the genus Hegeter (Tenebrionidae,
Coleoptera) and its colonization of the Canary Islands deduced from cytochrome oxidase I
mithocondrial DNA squences. Heredity 76: 392-403.
Juan, C., Emerson, B., Oromí, P., & Hewitt, G.M. 2000. Colonization and diversification:
towards a phylogeographic synthesis fort he Canary Islands. Trends in Ecology and Evolution
15: 104-109.
Pestano, J., Brown, R.P., Suárez, N.M. & Fajardo, S. 2003. Phylogeography of pipistrelle-like
bats within the Canary Islands, based on mtDNA sequences. Molecular Phylogeny and
Evolution 26: 56-63.
Scholz, H., Stierstorfer, Ch. & Gaisberg, M. v. 2000: Lolium edwardii sp. nova (Gramineae) and
its relationship with Schedonorus sect. Plantynia Dumort. Feddes Repertorium 111: 561-565
Wagner, D.L. & Liebherr, J.K. 1992. Flightlessness in insects. Trends in Ecology and Evolution
7: 216-220.
Wellenius, O. 1955. Formicidae insularum Canariensium. Societas Scientiarum Fennica,
Commentationes Biologicae 15(8): 1-20.
Wetterer, J., Espadaler, X., Wetterer, A. & Cabral, S.. 2004. Native and exotic ants of the
Azores (Hymenoptera, Formicidae). Sociobiology 44: 1-19.
Wheeler, W.M. 1927. The ants of the Canary Islands. Proceedings of the American Academy of
Arts and Sciences 62: 93-120.
Wilson, E.O. 1989. The coming pluralization of biology and the stewardship of systematics.
BioScience 39: 242-245.
Figure 1. Temnothorax bimbache. Worker profile.
Figure 2. Temnothorax bimbache, worker, profile of petiole and postpetiole.

Figure 3. Temnothorax bimbache. Queen profile.
Figure 4. Monomorium wilsoni. Worker profile.

Figure 5. Monomorium wilsoni. Queen, profile.
Figure 6. Monomorium wilsoni. Queen, profile of propodeum, petiole and postpetiole

Addendum
September 26, 2007
Corrections to the captions for Figures 4 - 6:
Figure 4. Monomorium wilsoni. Queen profile.
Figure 5. Monomorium wilsoni. Queen, profile of propodeum, petiole and postpetiole.
Figure 6. Monomorium wilsoni. Worker, profile.
Correspondingly, on page 118, in the queen description paragraph, “(Figures 5, 6)” should be “(Figures 4, 5)”.
In the worker description paragraph, “(Figure 4)” should be “(Figure 6)”.
Fernández, F. 2007. Two new South American species of Monomorium Mayr with taxonomic notes
on the genus, pp. 128-145. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant
systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions.
Memoirs of the American Entomological Institute, 80.
TWO NEW SOUTH AMERICAN SPECIES OF MONOMORIUM MAYR

WITH TAXONOMIC NOTES ON THE GENUS
Fernando Fernández
Instituto de Ciencias Naturales
Universidad Nacional de Colombia
Apartado 7495
Bogotá D.C., Colombia
ffernandezca@unal.edu.co
ABSTRACT
This article describes two new species of Monomorium from Brazil. One, M. delabiei n. sp. is
reminiscent of the genus Megalomyrmex, although it lacks the transverse propodeal carina
characteristic of that genus. The other, M. inusuale n.sp., is even more interesting: given the current
concepts in Solenopsidini, it could be treated as a new genus within the tribe due to its distinctive
morphological characteristics (mandibular configuration, vestibulate propodeal spiracle, propodeal
carinae, micropegs on the last tergum). Broadening the limits of Monomorium requires including
Nothidris, Phacota and Epelysidris as junior synonyms (syn. nov.). A working key to Neotropical
species of Monomorium is provided along with taxonomic notes.
Key words: Hymenoptera, Formicidae, Myrmicinae, Solenopsidini, Monomorium, Nothidris,

Phacota, Eplysidris, Neotropical, key, new species.
Fernández: South American Monomorium 129
INTRODUCTION
Monomorium is one of the most diverse ant genera, with more than 300 described species (Bolton,
1995; Heterick, 2001), the majority of which are Old World, especially Afrotropical. For a long
time there were various generic or subgeneric names associated with this genus and related groups
(Ettershank, 1966), without critical study of their limits or validity. Ettershank (1966) made the first
attempt to classify Monomorium and related groups; Bolton (1987) subsequently modified this
arrangement in his study of the Afrotropical fauna of the genus. Since then, the only regional studies
have been those of DuBois (1986) and Heterick (2001), and our understanding of the monophyly and
relationships of the group remains far from adequate. The present work is intended as a modest
contribution to this end; it consists of the description of two interesting species attributable to
Monomorium, with taxonomic notes, synonyms, and an operational key for the Neotropical species.
The species described herein require extending the generic limits of Monomorium, or forming
bridges among neighboring genera. Among the consequences is that Nothidris and Epelysidris
cannot be sustained as distinct genera, that Megalomyrmex is left in a precarious state, and that
Phacota becomes, once again, a synonym of Monomorium.
Heterick (2003) observed that : “Momomorium sublamellatum ... pushes back the diagnostic
boundaries of the genus Monomorium, and cannot be placed at present in existing Australian
species-groups of Monomorium” (Heterick, 2003). The description of the two new species is fraught
with analogous interpretational dilemmas with reference to the Neotropical Monomorium fauna,
within which perhaps neither is rightly included. Both species may eventually prove to lie outside
Monomorium. For further analysis of the issue, readers are referred to the discussion.
Depositories of Material
BMNH Natural History Museum, London, England.

CEPLAC Centro de Pesquisas do Cacau, Itabuna, Bahia, Brazil.
ICN Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C.,
Colombia.
LACM Los Angeles County Museum of Natural History, Los Angeles, USA.
MZSP Museu de Zoologia, Universidad de São Paulo, SP, Brazil.
Terminology
CI Cephalic index: HW/HL × 100.

EL Eye length: maximum diameter of the eye.
GL Gaster length: the maximum length, in lateral view, from the anteriormost point of the
gaster to gastral apex.
HL Head Length: maximum length, in full face view, from the midpoint apex of anterior
clypeal margin to midpoint of preoccipital (posterior) margin of head.
HW Head width: maximum width in full face view.
PL Petiole length: in dorsal view, from anterior to posterior articulation.
PPL Postpetiole length: in dorsal view, from anterior to posterior articulation.
SL Scape length: maximum length of scape, excluding basal condyle, in straight line distance.
TL Total length: total length of the ant from anterior clypeal margin to gastral apex.
WL Weber's length: the diagonal length of mesosoma from anterior pronotal margin to basal
angle of metapleuron.
Illustrations
Scanning Electron Microscope (SEM) photographs were taken at the University of Alberta in
Edmonton. Observations and measurements of pinned specimens were made using a Nikon SMZ 2T
stereomicroscope at 80X magnification and a fiber ring lamp. All measurements are in millimeters.
Monomorium Mayr
Monomorium Mayr, 1855: 452. Type-species: Monomorium monomorium Bolton, 1987: 287.
Phacota Roger, 1862: 260. Type species: Phacota sichelli Roger, 1862: 262. Synonym of Monomorium:
Ettershank, 1966: 82. Genus revalidated: Bolton, 1987: 281. Syn. rev.
Nothidris Ettershank, 1966: 105. Type-species: Monomorium latastei Emery, 1895: 10. Syn. n.
Antichthonidris Snelling, 1975: 5. Type-species: Monomorium denticulatum Mayr, 1887: 614. Junior
synonym of Monomorium: Heterick, 2001: 361.
Epelysidris Bolton, 1987: 279. Type-species: Epelysidris brocha Bolton, 1987: 280. Syn. n.
For a full list of synonymies before Heterick (2001) see Bolton (1987: 287-288).
WORKER DIAGNOSIS (after Bolton, 1987: 289; Heterick, 2001: 363-364).
Monomorphic to polymorphic. Minute to moderately large in total length. Mandibles with 4 to

5 teeth. Maxillary palps with 2 to 4 segments. Median clypeal seta present, sometimes displaced or
absent. Median portion of clypeus raised, longitudinally bicarinate, the carinae rarely effaced.
Frontal carinae absent past frontal lobes. Antennal scrobes absent. Antennae with 11-12 segments
and with club of 3 (rarely 4) segments. Eyes present, sometimes reduced. Metapleural glands never
bulging or hypertrophied. Metapleural lobes usually small, rounded. Propodeum normally unarmed,
sometimes angulated to dentate, rarely with lamelliform process. Propodeal spiracle usually circular
and at about midlength of the sclerite, rarely in another position. Petiole pedunculated, the petiolar
spiracle usually close to or at node. Sting functional.
Nothidris was created by Ettershank (1966) and further delimited by Snelling (1975), who
created Antichthonidris to accomodate some species. Bolton (1987:284-285) discussed the traits
proposed for the latter, demonstrating their weakness and dubious value as generic-level characters:
a vestibulated propodeal spiracle appears to be present in some Australian Monomorium species
(Bolton, 1987), for instance, as well as in M. delabiei. Moreover, the inclusion of Antichthonidris in
Monomorium, as proposed by Heterick (2001), leaves no justification for maintaining Nothidris as a
separate genus.
Phacota has been a taxonomic problem in the myrmicines, due to its poor description, the
disappearance of the type specimen, and the lack of collected material referable to P. sichelii
(Bolton, 1987), all of which have impeded an evaluation of its taxonomic status. Ettershank (1966)
considered this name a junior synonym of Monomorium. Bolton (1987) subsequently revived the
genus, citing the few attributes that can be retrieved from Roger's (1862) original description;
nevertheless, he made explicit his strong suspicion that the putative species is based on a wingless,
ergatoid Monomorium female, perhaps from the M. salomonis group. Both the meager description
(e.g., that the gaster is bigger than the head) and the important fact that the species has not been
rediscovered in Spain or any other nearby location, are consistent with this interpretation. Given that
the European ant fauna can be considered acceptably collected and studied, and in light of the
group's importance, samples assignable to Phacota would surely have been detected and described
by now. According to its description, Phacota is characterized by 11-segmented antennae with a 2-
segmented club. Some Neotropical Solenopsis females possess this combination, but it is an
antennal configuration unknown in Monomorium, and it is highly probable that the description of the
number of flagellomeres in the antenna and club is erroneous. It would not be the only inadvertent
mistake of this type in the history of ant systematics, especially given the size of the ants and the
optical resolution possible in the 19th century. It seems of little practical use to maintain a badly-
described genus, with no associated type material, and no other collected material, and I recommend
that Phacota once again be demoted as proposed by Ettershank (1966) until and unless more
material is discovered, or the type specimen (in good condition) reappears.
Epelysidris is a monotypic genus of eastern Malaysia, easily separable by the distinctive pair of
lobules on the basal border of each mandible, mandibular and clypeal structure, and palpal formula
(Bolton, 1987). Although this taxon is undoubtedly monophyletic, its continued recognition as a
separate genus would create the same dilemmas that characterize Antichthonidris, Nothidris, and
some others. It is preferable to leave brocha as one additional (although highly apomorphic) species
within Monomorium; I propose here that Epelysidris thus be considered a junior synonym of
Monomorium.
Neotropical species of Monomorium (includes recent introductions*).
M. bidentatum Mayr, 1887 comb. rev. – Chile, Argentina

M. brasiliense Forel, 1908 - Brazil
M. carbonarium Fr. Smith, 1858 – Azores
M. cekalovici (Snelling, 1975) comb. nov. - Chile
M. chilensis, n.name for N. bicolor Ettershank, 1965:55, preoccupied by M. bicolor Emery,
1877:368 - Chile
M. cyaneum Wheeler, 1914 - Mexico
M. compressum Wheeler, 1914 – Mexico
M. delabiei sp.n. - Brazil
M. denticulatum Mayr, 1887 comb. rev. – Chile, Argentina
M. destructor (Jerdon, 1852)* - Widespread
M. ebeninum Forel, 1891 – Caribbean and coastal Mesoamerica
M. floricola (Jerdon, 1852)* - Widespread
M. inquilinum DuBois, 1980 - Mexico
M. inusuale sp.n. - Brazil
M. latastei Emery, 1895 comb. rev. - Chile
M. marjoriae DuBois, 1986 – Mexico
M. minimum (Buckley, 1867) – Mexico (?) Paraguay (?)
M. monomorium Bolton, 1987* – Barbados
M. pharaonis (Linnaeus, 1758)* - Widespread
M. salomonis (Linnaeus, 1758)* - Widespread
M. subcoecum Emery, 1894 – Caribbean (St. Thomas and Puerto Rico)
M. subopacum Fr. Smith, 1858* – Antigua
Outside the Neotropical fauna the following changes are proposed:
Monomorium sichelii (Roger, 1862) comb. rev.

Monomorium brocha (Bolton, 1987) comb. n.
SPECIES DESCRIPTIONS AND NOTES
Monomorium bidentatum Mayr comb. rev.
Monomorium bidentatum Mayr, 1887: 616 (w, q).

Monomorium (Notomyrmex) bidentatum: Emery, 1922: 169.
Monomorium (Notomyrmex) bidentatum piceonigrum Borgmeier, 1948: 468 (w); Kusnezov, 1960: 345
(as junior synonym of bidentata).
Notomyrmex bidentatum: Kusnezov, 1960: 345.
Nothidris bidentatus: Ettershank, 1966: 107.
Antichthonidris bidentatus: Snelling, 1975: 5-6 (w, q, m); Wheeler & Wheeler, 1980: 533 (larvae).
Antichthonidris bidentata: Bolton, 1995: 67.
Heterick (2001) synonymized Antichthonidris under Monomorium, but did not make the
appropriate nomenclatural changes in the two species assigned to that genus.
Monomorium brasiliense Forel
Monomorium minutum subsp. brasiliense Forel, 1908: 361 (w).

Monomorium monomorium subsp. brasiliense: Brandao, 1991: 357.
Monomorium brasiliense: Bolton, 1995: 260.
Bolton (1987) did not study the type of this species, nor the possibly corresponding native
Monomorium material. The Monomorium native to South America are moderate to large in size; M.
inusuale and M. delabiei are small but with HW greater than 0.60 mm. There is a possibility that M.
brasiliense might in fact be a tramp species, erroneously described by Forel as a different
(subspecific) taxon.
Monomorium cekalovici (Snelling) new combination
Nothidris cekalovici Snelling, 1975:3 (w); Bolton, 1995: 292.
Monomorium chilensis, new name.
Megalomyrmex bicolor Ettershank, 1965:55 (w, larva); Kempf, 1970: 359.

Nothidris bicolor: Snelling, 1975: 3; Bolton, 1995: 292.
M. chilensis is the new name proposed for bicolor, a name preoccupied by Monomorium bicolor
Emery, 1877: 368.
Monomorium cyaneum Wheeler
It is not easy to separate M. cyaneum from M. ebeninum and M. depressum. In the syntypes of
M. cyaneum examined (LACM) the mesopleuron is smooth, whereas the DuBois (1986) diagnosis
and keys indicate punctate mesopleura for the species. There are Mexican workers (LACM) with
punctate mesopleura identified as M. cyaneum, but other material referable to this species (Mexico,
LACM) lacks this attribute. Perhaps there is variability in the character, but the limited material is
insufficient to evaluate this possibility. In M. cyaneum the propodeum is more weakly angled than
in the other two species and supposedly there are 8 or fewer hairs on the mesosoma (DuBois, 1986).
The pilosity varies somewhat in samples referable to these three species. M. cyaneum and M.
depressum share a petiole that is slightly curved in frontal view, as well as a petiolar node that is
thicker in lateral view. M. ebeninum has a petiole that is flat or weakly concave in frontal view.
Thus the problem is how to separate M. cyaneum and M. depressum. As pointed out above, the
punctate sculpturing of the mesopleura does not seem to be universal in cyaneum (three syntypes
examined, LACM, do not have appreciable sculpturing on the mesopleura). A characteristic of the
syntypes and other specimens (from Hatillo, Mexico, LACM) is the presence of a light metallic blue
sheen on the entire body (from which the species name derives). A specimen labeled as cyaneum
(LACM) by DuBois has mesopleural sculpturing but does not have the blue sheen.
Monomorium delabiei, new species
Description
Worker measurements (mm): HW 0.65; HL 0.73; SL 0.53; EL 0.16; WL 1.03; PL 0.36; PPL
0.24; GL 1.00; TL 3.50. Indices: CI 89; SI 82.
Head slightly longer than wide. Sides of head slightly convex, evenly rounded into the posterior
border, latter more or less straight with a small medial concavity. Mandibles with four teeth: the
apical larger than the others, the subapical separated by a space from the basal teeth. Maxillary
palps 2 segmented. Clypeus convex with no trace of carinae, extended posteriorly as smooth and
shiny frontal area. Clypeus anteriorly convex. Antenna 12-segmented with club 3-segmented.
Frontal carinae short.
Eyes large, well developed, with about 11 ommatidia in the maximum diameter, situated in the
middle of the head.
In lateral view mesosoma with promesonotum convex. Metanotal groove deep, well marked.
Propodeum unarmed. Declivity of propodeum devoid of transverse carinae. Propodeal spiracle with
a thin walled vestibule anteriorly. Metapleural lobes small, rounded. Petiole with short peduncle,
node rounded above. Petiolar spiracle at beginning of node. Petiolar ventral process consisting of
anterior median keel. In dorsal view postpetiole wider than petiole. In side view postpetiole
subcampaniform, with a ventral strong median tubercle, pointed anteriorly. Sting developed.
Entire body smooth and shining. Moderate erect pilosity on head, promesonotum, petiole,
postpetiole and gaster, very few on propodeum. Larger hairs about 0.15 mm, those of head shorter.
Clypeus with two series of transverse setae, those of anterior margin increasing in size toward the
somewhat larger median apical setae. Body brown, most of gaster dark brown.
Queen and male unknown.
Type data: Holotype worker, Brazil, Bahia, Fazenda Amarillina, Guaratinga, 28.vii.1993
(Pula/Michelli No. 4675-D) (CEPLAC).
Distribution: Eastern Brazil.
Etymology: This species is named for my friend and colleague Dr. Jacques Delabie, for his
continuous supply of interesting ants.
Comments
In M. delabiei there are two transverse rows of hairs on the clypeus. In the row on the anterior
border, the hairs increase in length from the lateral ones towards the midpoint, where they are
longest (although not very much longer than the rest); in other words, there is no single medial hair
that stands out, a characteristic typical of the tribe. This is also typical of some Megalomyrmex, and
in at least some “Antichthonidris” workers, there is no apical medial seta. The clypeus forms a
medial convexity with only a very weak trace of carinae. The tentorial pit is more than halfway
between antennal receptacles and mandibular base (as in some “Antichthonidris”), rather than near
the antennal alveolus (as in some Neotropical Monomorium). The form of the propodeal spiracle is
somewhat similar to that of Nothidris (Bolton, 1994: 104), with a visible vestibule preceding
anteriorly the spiracle proper: in typical Monomorium and Megalomyrmex the spiracle is round and
without anterior vestibules. The postpetiole has a robust, pointed ventral tubercle, a characteristic
apparently not shared with other members of the genus or with Megalomyrmex. The absence of the
transverse propodeal carina excludes, by definition, the inclusion of this species in the typical
Megalomyrmex, although the character is not universal within the genus, and can be present in some
Monomorium.
Monomorium denticulatum Mayr revised combination
Monomorium denticulatum Mayr, 1887: 614 (w, q); Wheeler & Wheeler, 1980: 532 (larvae).
Monomorium denticulatum navarinensis Forel, 1904: 7 (q); Kusnezov, 1960: 347 (as junior synonym of
denticulata).
Monomorium denticulatum picea Emery, 1906: 120 (w); Kusnezov, 1960: 347 (as junior synonym of
denticulata).
Monomorium (Notomyrmex) denticulatum: Emery, 1922: 169.
Monomorium (Notomyrmex) denticulatum navarinensis: Emery, 1922: 169.
Monomorium (Notomyrmex) denticulatum inerme Borgmeier, 1948: 469 (worker); Kusnezov, 1949:432
(as junior synonym of denticulata).
Notomyrmex denticulatum: Kusnezov, 1960: 347.
Nothidris denticulatus: Ettershank, 1966: 107.
Antichthonidris denticulatus: Snelling, 1975: 6; Heterick, 2001: 353.
Antichthonidris denticulata: Bolton, 1995: 67.
*Monomorium destructor (Jerdon)
A widespread tramp species, M. destructor is readily recognized by the fine transverse striae on
the vertex, as well as the deep metanotal groove.
Monomorium ebeninum Forel
This species is similar to M. compressum and, to a lesser extent, to M. cyaneum. A

characteristic that appears to separate it from similar species is the petiolar profile, which in frontal
view is flat to weakly concave.
*Monomorium floricola (Jerdon)
This is a small adventive Old World species, as is M. monomorium, with HW less than 0.35
mm. In M. floricola the pale mesosoma is in sharp contrast to the darker head and gaster.
Monomorium inquilinum DuBois
Known only from queen and male castes. DuBois (1986) suggests that this species is a social
parasite of M. cyaneum. Known only from the type locality, between Mexico City and Querétaro,
Mexico.
Monomorium inusuale, new species

Figures 1-2
Description
Worker, measurements (mm), holotype (paratypes, n=3): HW 0.48 (0.46-0.48); HL 0.57 (0.57-
0.62); SL 0.37 (0.37-0-41); EL 0.03 (0.03); WL 0.72 (0.70-0.72); PL 0.25 (0.25); PPL 0.18 (0.18);
GL 0.73 (0.73-0.80); TL 2.58 (2.58-2.61). Indices: CI 84 (74-84); SI 77 (77-89).
Head longer than wide. Sides of head parallel, very slightly wider anteriorly, broadly rounded
into the posterior border, which is more or less straight. Mandibles with four teeth: the three apical
separated from isolated basal by a large diastema. Maxillary palps 2 segmented. Median lobe of
clypeus strong, bicarinate, narrowing posteriorly to narrow strip between frontal lobes. Clypeus with
apical median seta and several paracarinal setae. Antenna 12 segmented with strong 3 segmented
club; scapes fail to reach the vertexal border. Antennal insertions close together. Frontal carinae
and antennal scrobes absent. Eyes reduced to indistinct ommatidia, placed in front of head midline.
In lateral view promesonotum slightly convex. Metanotal groove very deep, well marked.
Propodeum unarmed, with a faint cuticular crest, from the meeting point of dorsal and declivity
faces, extended and slightly higher, continuous with the small metapleural lobes. Propodeal spiracle
circular, opened posteriorly. Bullae of metapleural glands large. Petiole with peduncle and node
well differentiated, the node more or less with the sides parallel, dorsum convex. Petiolar spiracle at
beginning of node. Petiolar ventral process tooth-like. Postpetiole subcampaniform, with a ventral
strong transverse carina. Apical portion of penultimate tergite with four small pegs or teeth, hairs
arising from outermost.
Mandibles, most of promesonotum, dorsum of petiole and postpetiole and gaster smooth and
shining. Head with longitudinal rugulae mixed with dense foveae. Posterior promesonotum with
feeble short rugulae. Most of mesopleura with irregular short longitudinal striae, mixed with fine
reticulation. Most of propodeum and sides of petiole and postpetiole densely reticulated. Declivity
of propodeum with several fine transverse carinae, the most posterior more marked. Moderate erect
pilosity on head, promesonotum, petiole, postpetiole and gaster, very few on propodeum. Scapes
with several erect hairs. Larger hairs about 0.15 mm, those of head shorter. Body brown,
appendages lighter.
Type data: Holotype worker, Brazil, Bahia, Barrolandia, 16-23.07.1994 (S. Lacau) (CEPLAC).
Paratypes, 1 worker, same data as holotype, deposited in ICN; 3 workers, Brazil, Amazonas,
Benjamin Constant, 21.ix.1962 (W.L. Brown Jr.) (MZSP); 2 workers, Ecuador, Napo, Limoncocha,
1973 (M. Rettenmeyer) (MZSP, BMNH).
Distribution: Ecuador and Brazil.
Etymology: The name refers to the unusual traits of the ant, and the taxonomical difficulties for
their generic placement.
Comments
This species presents a series of unusual characters that are interesting in the context of the
Solenopsidini, and especially Monomorium and its allied genera. The long diastema between the 3
apical teeth and the basal tooth does not appear to be common in the genus or any other close genera.
The same can be said for the much reduced eyes, since large, multifaceted eyes are common in the
genus, although some Afrotropical species have reduced eyes (Snelling, personal communication).
The narrow carina or propodeal crest that extends down to join the metapleural lobes is reminiscent
of Epelysidris. The anterior and posterior sides of the petiolar node are more or less straight; in
other species in this and neighboring genera, they tend to be rounded. The postpetiole has a strong
transverse keel in the ventral part, which is reminiscent of Adelomyrmex. The head sculpturing is
also unusual, with longitudinal rugules mixed with obvious foveae, a trait which is absent in other
Neotropical Monomorium. The most outstanding feature is the series of minute teeth or pegs in the
last abdominal tergum, from at least the outermost of which arise hairs. This appears to be an
autapomorphic structure, absent in other Solenopsidini and probably in Myrmicinae; it is somewhat
like the pygidial pegs of Cerapachyinae and Pachycondyla crassinoda workers.
The Brazilian specimens are uniform in size and general aspect. Nevertheless, the two
Ecuadorian specimens differ somewhat in size and some measurements (slightly shorter dorsal side
of the propodeum, for instance) and the visible micropegs of the last tergum are less conspicuous
than those of Benjamin Constant, Brazil. These structures are well-developed in the type specimen,
from Bahia, but the propodeum is intermediate between the other Brazilian specimens and those
from Ecuador. Given that in the other essential characteristics all samples are similar, I prefer to
consider these differences as within-species variability, instead of creating a new taxon that might
result in a paraphyletic species.
Is M. inusuale truly a Monomorium? I recently was loaned two interesting myrmicines from
CEPLAC. The first of these, a worker from Guaramiranga (Brazil, Ceará) is reminiscent in general
form of the Australian M. sublamellatum, although without a single apical clypeal seta and clypeus
so conspicuously bicarinate. The Brazilian specimen could pass as an unusual Monomorium.
However, the discovery of a preoccipital carina—supposedly apomorphic in Rogeria—could require
the inclusion of the species in the latter genus, in spite of the fact that other characteristics do not
coincide (Kugler, 1994). Or, alternatively, the Australian species possibly does not belong in
Solenopsidini; Heterick (2003) places sublamellatum in Monomorium by default, since it does not fit
in any other genus of the tribe. The paraphyletic nature of Monomorium renders it a “basket” or
“catch-all” genus. Maybe sublamellatum, like this Ceará worker, belongs to Stenammini (Bolton,
personal communication).
The second CEPLAC specimen from Bahia (Brazil, S. Jose Victoria, No. 2139, provisionally
determined as Monomorium sp.) has a general aspect typical of Monomorium, the genus to which the
Bolton (1994) key leads, if one ignores the absence of the apical clypeal seta (which is variable in
Solenopsidini). Nevertheless, as in the above case, this specimen might better be associated with
some Stenammini, possibly Rogeria, although it is impossible to confirm the preoccipital carina in
the unique dry-mounted specimen, and the total absence of color and other traits make it difficult to
place in Rogeria (as defined by Kugler, 1994).
Bolton (personal communication) suggests that M. inusuale might be a member of Stenammini,
which was my first suspicion due to the overall resemblance with members of this tribe. The traits
of frontal lobes and toruli, as well as clypeal posterior border, places inusuale in the Stenammini as
diagnosed in Bolton (2003:58). However, the first gastral tergite clearly overlapping the first sternite
on ventral surface, and the single medial clypeal seta puts the taxon in the Solenopsidini, again sensu
Bolton (2003:59-60). Although the solenopsidine tribe group (Bolton 2003:57) may be a
monophyletic group, the tribes proposed by Bolton (2003) as Stenammini and Solenopsidini
currently lack synapomorphies. Perhaps the mixed traits of inusuale, the Australian species, and the
Brazilian specimens referred to above may justify merging the two tribes. On the other hand,
moving these taxa from Monomorium to Stenammini simply transfers the problem of generic
allocation from one tribe to the other.
Bolton (2003) has clarified some of the uncertainties in the systematics of the myrmicines allied
to Solenopsidini and Stenammini. However, there are many problems to be resolved only when the
limits of the Myrmicine tribes are clearly defined. For these reasons, I think that the only option at
the moment is to leave inusuale as a member of Monomorium.
Monomorium latastei Emery revised combination
Monomorium latastei Emery, 1895: 10 (w); Wheeler & Wheeler 1980: 533 (larva).
Monomorium (Notomyrmex) latastei: Emery, 1922: 169.
Nothidris latastei: Ettershank, 1966: 106.
Monomorium marjoriae DuBois
The worker caste of this species is indistinguishable from those of M. minimum and M.
wheelorum. However, M. marjoriae is the only species (of these three) recorded for México
(DuBois, 1986).
Monomorium minimum (Buckley)
Kempf (1972) recorded this species from Paraguay, following Forel (1917). However, I know
of no confirmed records of this species from Paraguay or any other Neotropical country and it is not
included in the key below.
*Monomorium monomorium Bolton
Wilson and Taylor (1967) mention the presence of this Old World species (as M. minutum
Mayr) from Brazil. However, I know of no confirmed records in the literature. It has also been
recorded from Barbados (Kempf, 1972).
*Monomorium pharaonis (Linnaeus)
Monomorium pharaonis, M. salomonis and M. subopacum are the only species in the
Neotropics with head and mesosoma with fine reticulate-punctate sculpturation and mandibular
dorsum with coarse longitudinal rugulae. M. pharaonis has two rows of hairs on the head between
the vertex and carinae, which distinguishes this species from the other two. M. pharaonis is one of
the best-known Old World tramp species.
*Monomorium salomonis (Linnaeus)
The body is sculptured as in M. pharaonis and M. subopacum. It differs from these by the lack
of hairs on the dorsum, and presence of a medial notch in the anterior clypeal margin. It is a
widespread species, introduced in the Neotropics.
Monomorium subcoecum Emery
This is a minute, pale ant, with propodeum angles well marked. Mesopleuron sculptured. Body
with abundant pilosity. It was described from St. Thomas in the Lesser Antilles, but is found also in
Puerto Rico (R. R. Snelling, pers. comm.).
Monomorium subopacum (Fr. Smith)
Similar in general form as M. pharaonis and the body is without erect hairs; clypeus without the
medial notch as in M. salomonis. Originally described from the Azores, M. subopacum has been
recorded from Antigua in the Lesser Antilles (Kempf, 1972).
THE LIMITS OF MONOMORIUM AND RELATED GENERA
Bolton (1987) characterized the genus and listed its extensive associated synonymy, including
new synonymies. This author proposed species groups for the Afrotropical fauna, some of which
might be applicable to the fauna outside of Africa. Since then, DuBois (1986) has studied the M.
minimum species group native in America and Heterick (2001) has revised the Australian fauna,
placing Antichthonidris Snelling as junior synonym of Monomorium.
Monomorium was included within the tribe Solenopsidini by Bolton (1987), citing several traits
and giving emphasis to the possession of a single medial apical seta or hair on the anterior margin of
the clypeus. This characteristic at times is variable (the seta maybe displaced to one side, or a pair of
setae more or less arising from the apical part of the medial clypeus), a feature that is also present in
Adelomyrmex, Cardiocondyla, Tyrannomyrmex and some Stenammini (Bolton, 2003; Fernández,
2003).
Later, Bolton (2003) redefined Solenopsidini and proposed two groups in the tribe: Solenopsis
genus group (Solenopsidini s.str., as in Bolton, 1987) and Carebara genus group, containing most of
the former tribe Pheidologetonini. Although it is possible to recognize several monophyletic
groupings within the Solenopsidini s.str. (e.g., Allomerus, Solenopsis), the monophyly of
Monomorium and allied groups (Antichthonidris, Megalomyrmex, Nothidris, Phacota) is not clear.
In fact, Heterick (2001) in his revision of the Australia Monomorium formally synonymized
Antichthonidris with Monomorium. Bolton (1987) had already warned of the precariousness of this
genus and of Nothidris Ettershank, names whose validity become feeble upon examination of the
group in a worldwide context. Phacota Roger was revalidated by Bolton (1987) as a synonym of
Monomorium (Ettershank, 1966). Unfortunately the type of this group has disappeared and Bolton
1987) suspected that the description might have been based on a wingless ergatoid female.
What are the limits of Monomorium?
There has been no phylogenetic study of the Solenopsidini. Kusnezov (1957) discussed some
problems in the taxonomy of Solenopsis and neighboring groups and Bolton (1987) ventured some
possible relationships within the tribe (now Solenopsis genus group: see Bolton, 2003). Some
groups, e.g., Anillomyrma + Bondroitia, or Diplomorium + Allomerus, as well as Solenopsis +
Oxyepoecus, possess distinctive shared morphological traits. Notwithstanding that these groups
might be thus be considered monophyletic, the arrangement would leave Monomorium and other
close genera as paraphyletic. No morphological trait exists, either in workers or sexuals, which
would robustly separate Monomorium from other solenopsidines. The existence of certain
characteristics or sets of characteristics that were seen from a local rather than global perspective
favored the creation of many weakly-delimited genera, whose fate has been synonymy. In fact, strict
application of cladistic norms would result in synonymy of all the genera in the tribe, leaving a giant
inclusive genus, Solenopsis, which would be rejected by the majority of myrmecologists. Even so,
the monophyly of this grouping would not be clear, since Solenopsidini still lacks a strong
synapomorphy: the apical central clypeal seta is not universal.
Perhaps a moderate solution might be to maintain as valid those groups that can be clearly
distinguished from Monomorium (Allomerus, Anillomyrma, Bondroitia, Diplomorium, Oxyepoecus
and Solenopsis), leaving the rest in a single genus, Monomorium. This group is paraphyletic, but can
be justified on practical grounds as a provisional arrangement, pending molecular systematics and

chemotaxonomy that will probably illuminate relationships not apparent solely on the basis of
external morphology. Some studies, for instance, suggest that the defensive chemicals in Solenopsis
and neighboring groups will be phylogenetically useful traits (Van der Meer, 1986). Although
molecular phylogenetic studies are not nearly as compelling as the discipline's followers contend, the
joint examination of molecular, chemical and morphological characters will probably be
enlightening. This problem is not exclusive to Monomorium and its neighbors; it is highly likely that
the same situation exists in genera such as Camponotus and its neighbors in the tribe Camponotini
(Brady et al., 1999) and in the taxa created around Leptothorax (Bolton, 2003).
The species M. inusuale described here creates an interesting dichotomy. On one hand, it
possesses unique features (mandibles with diastema, greatly reduced eyes, propodeal crest, pegs on
the penultimate abdominal segment) whose nature might justify the creation of a new genus, as was
the case with Nothidris, Antichthonidris and Epelysidris. Nevertheless, given the morphological and
behavioral richness of Monomorium (Bolton, 1987; Heterick, 2001), it is difficult to justify such
strict generic limits. If Nothidris or Epelysidris are accepted as valid genera, it requires that M.
inusuale also be considered a separate genus. Nevertheless, by so doing, it would favor the
indiscriminate creation of many monotypic genera (or with just a few species) around Monomorium.
Although such genera might be monophyletic, characterizing them as separate genera might
confound the taxonomy of the group. It seems thus more advisable to regard these as monophyletic
lineages that have evolved separately, each with unique traits, within Monomorium. This is the
decision taken recently by Heterick (2003) in placing the highly distinctive M. sublamellatum in
Monomorium, instead of erecting a new monotypic genus to accommodate this species.
To summarize, this work proposes the following: To accept as valid genera within the tribe
Solenopsidini those that can be unambiguously distinguished from Monomorium: Anillomyrma,
Solenopsis and others. To consider as congeners those genera that cannot be clearly separated from
Monomorium: Nothidris, Phacota, Epelysidris and perhaps Megalomyrmex. This is a practical
position, but there are other considerations as well: the phylogeny of the Myrmicinae is barely in its
first stages. This means that some genera placed in Solenopsidini (Anillomyrma, for instance) might
eventually belong to a different lineage, which may restrict the size of the tribe. New morphological,
molecular, and chemical characters, as well as the description of the males and females of many of
these lineages will perhaps in the future create a firm foundation for understanding the limits and
relationships of the components of this interesting tribe.
KEY TO NEOTROPICAL MONOMORIUM WORKERS
This key should be considered a working draft, since it has been constructed based more on the
literature than on specimens: Snelling & Hunt, 1976; DuBois, 1986; Bolton, 1994. I have not been
able to examine various specimens assignable to some of the species reported for the Neotropical
region. It will be of special interest in this context to study the type of M. brasiliense and establish
its actual status. Observations by users will surely be useful for improving future versions.
1 Head and dorsal mesosomal sculpture with longitudinal striations or fine reticulation .......... 2
– Head and mesosomal dorsum smooth and shining; any sculpture, if present, reduced........... 5
2 Last abdominal tergite with a series of small pegs (Fig. 2); mandibles with four teeth, the apical
ones separated from basal by diastema (Fig. 1); Brazil and Ecuador..................... M. inusuale
– Last abdominal tergite without small pegs; mandibles with different configuration .............. 3
3 Front of head with several erect hairs forming two rows of about 4 hairs each; mesosoma with
several hairs .........................................................................................................M. pharaonis
– Front of head devoid of erect hairs; mesosoma without erect hairs ........................................ 4
4 Clypeal margin with medial notch .......................................................................M. salomonis

– Clypeus without medial notch; Antigua.............................................................M. subopacum
5 Maxillary palps with 4 segments; Chile.................................................................................. 6

– Maxillary palps with fewer than 3 segments........................................................................... 8
6 Head concolorous with mesosoma; gaster darker; propodeum distinctly angulate in profile
orwith short sharp teeth .......................................................................................................... 7
– Head concolorous with gaster; mesosoma lighter; propodeum rounded in profile ...................
.............................................................................................................................M. cekalovici
7 Propodeum angulate in profile, but without sharp teeth; penultimate antennomere 1.25 – 1.45×
longer than wide .................................................................................................... M. chilensis
– Propodeum at least sharply angulate, usually with distinct triangular teeth; penultimate
antennomere 0.77 - 1.20× longer than wide............................................................. M. latastei
8 Head with fine transverse rugulae on vertex........................................................M. destructor

– Vertex without obvious rugulae.............................................................................................. 9
9 Propodeum armed with well developed spines; Chile and Argentina................ M. bidentatum
– Propodeum unarmed or with small spines or angles ............................................................. 10
10 Head width more than 0.60 mm............................................................................................ 11

– Head width less than 0.60 mm .............................................................................................. 12
11 Clypeus bicarinate; mandibles with 5 teeth; Chile and Argentina .................. M. denticulatum
– Clypeus feebly bicarinate; mandibles with 4 teeth; Brazil...................................... M. delabiei
12 Mesosoma and petiole clear yellow or at most light brownish, contrasting with dark reddish
brown head and gaster ........................................................................................... M. floricola
– Mesosoma concolorous at least with either head or gaster ................................................... 13
13 Propodeum abruptly subangulate in profile; several short parallel, longitudinal rugulae on

clypeus; head and body shiny black..................................................................... M. ebeninum
– Propodeum distinctly rounded in profile or with a pair of short sharp spines at propodeal angles
.............................................................................................................................................. 14
14 Scape surpassing posterior margin of head by slightly less than length of following segment.
............................................................................................................................. M. marjoriae
– Scape never surpassing posterior margin of head ................................................................. 15
15 Propodeum with small spines; clypeus without teeth; eyes reduced to one or two ommatidia;
yellowish species; Puerto Rico, Virgin Islands.................................................. M. subcoecum
– Propodeum unarmed, rounded in profile; clypeus with two submedian teeth; eyes multifaceted;
body brown to black ............................................................................................................. 16
16 Scape with reclinate hairs; body brown ................................................. M. compressum (part)

– Scape with suberect hairs; body blackish brown to black..................................................... 17
17 HW equal to or more than 0.38mm; Mexico ........................................................................ 18

– HW less than 0.36mm; introduced in Brazil (?)..............................................M. monomorium
18 Body with metallic bluish reflections; clypeal teeth as long as wide at base .........M. cyaneum
– Body without bluish reflections; clypeal teeth stout, wider at base than long ..........................
…………………………………………………………………….……..M. compressum (part)
ACKNOWLEDGMENTS
I am pleased to offer my thanks to many people who helped to produce this contribution. Beto
Brandão (MZSP), Roy R. Snelling and Brian Brown (LACM), Charles Delabie (CEPLAC) and
James Wetterer loaned or provided ants. Sara Bennet made useful translation and/or suggestions in
the English. Partial support was received from NSF Grant DEB Nos. 9972024 and 0205982 to
Michael Sharkey and Brian Brown. SEM pictures were taken by George D. Braybrook (Department
of Earth and Atmospheric Sciences, University of Alberta) and prepared by Danny Shepley
Department of Biological Sciences, University of Alberta); costs incurred were met through Grant
1399 of National Research Council of Canada to George E. Ball (Department of Biological
Sciences). This paper is a partial product of my PhD Thesis at the Universidad Nacional de
Colombia under the directorship of Dr. Rodulfo Ospina. Special thanks to Roy R. Snelling and Phil
Ward for the useful corrections and comments to the manuscript.
LITERATURE CITED
Bolton, B. 1987. A review of the Solenopsis genus-group and revision of Afrotropical Monomorium
Mayr (Hymenoptera: Formicidae). Bulletin of the British Museum (Natural History)
Entomology 54: 263-452.
Bolton, B. 1994. Identification guide to the ant genera of the world. Cambridge, Massachusetts:
Bolton, B. 1995. A new general catalogue of the ants of the world. Cambridge, Massachusetts:
Borgmeier, T. 1948. Einige ameisen aus Argentinien. Revista de Entomologia 19: 459-471.
Brady, S.G., Gadau, J. & Ward, P.S. 1999. Is the ant genus Camponotus paraphyletic? Pp. 27-28
in: 4th International Hymenopterists Conference, 6-11th January 1999, Canberra, Australia,
Abstracts.
Brandão, C.R.F. 1991. Adendos ao catálogo abreviado das formigas da região Neotropical. Revista
Brasileira de Entomologia 35: 319-412.
DuBois, M. 1986. A revision of the native New World species of the ant genus Monomorium
(minimum group) (Hymenoptera: Formicidae). University of Kansas Science Bulletin 53: 65-
119.
Emery, C. 1877. Catalogo delle formiche esistenti nelle collezioni del Mueso Civico di Genova.
Parte prima. Formiche provenienti dal Viaggio dei signori Antinori, Beccari e Issel nel Mar
Rosso e nel paese dei Bogos. [part]. Annali del Museo Civico di Storia Naturale di Genova 9:
363-368.
Emery, C. 1894. Studi sulle formiche della fauna Neotropica. Bulletino della Societá Entomologica
Italiana 26: 137-241.
Emery, C. 1895. Deuxiéme note sur les fourmis du Chili. Actes de la Societé Scientifique du Chili 5:
10-18.
Emery, C. 1906. Studi sulle formiche della fauna Neotropica. Bulletino della Societá Entomologica
Italiana 37: 107-194.
Emery, C. 1922. In Wytsman, P. Genera Insectorum. Hymenoptera, Fam. Formicidae, subfam.
Myrmicinae. Fasc. 174B & 174C: 95 -206, 207-397. Bruxelles.
Ettershank, G. 1965. A new species of Megalomyrmex from the Chilean Andes. Psyche
(Cambridge) 72: 55-58.
Ettershank, G. 1966. A generic revision of the world Myrmicinae related to Solenopsis and
Pheidologeton. Australian Journal of Zoology 14: 73-171.
Fernández, F. 2003. A new myrmicine ant genus from Malaysia with uncertain affinities
(Hymenoptera: Formicidae). Zootaxa 341: 1-6.
Forel, A. 1891. In Grandidier, A. Histoire Physique, Naturelle et Politique de Madagascar 20.2.
Les Formicides: 1-231.
Forel, A. 1904. Formicidae. Ergebnisse der Hamburger Magalhaensischen Sammelreise 7(8): 1-7.
Forel, A. 1908. Ameisen aus Sao Paulo (Brasilien), Paraguay, etc. Gesammelt von Prof. Herm.v.
Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der k.k. Zoologisch-Botanischen
Gessellschaft in Wien 58: 340-418.
Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bulletin de la Societé
Vaudoise des Sciences Naturelles 51: 229-253.
Heterick, B.E. 2001. Revision of the Australian ants of the genus Monomorium (Hymenoptera:
Heterick, B.E. 2003. Two new Australian Monomorium Mayr (Hymenoptera: Formicidae)
including a highly distinctive species. Australian Journal of Entomology 42: 249-253.
Kempf, W.W. 1970. Taxonomic notes on ants of the genus Megalomyrmex Forel, with the
description of new species. Studia Entomologica (N.S.) 13: 353-364.
Kempf, W.W. 1972, Catálogo abreviado das formigas da região Neotropical. Studia Entomologica
(N.S.) 15: 3-344.
Kugler, C. 1994. A revision of the ant genus Rogeria with descriptions of the sting apparatus
(Hyemnoptera: Formicidae). Journal of Hymenoptera Research 3: 17-89.
Kusnezov, N. 1949. El género Monomorium en la Argentina. Acta Zoologica Lilloana 7: 423-448.
Kusnezov, N. 1957. Die Solenopsidinen-Gattungen von Südamerika. Zoologischer Anzeiger 158:
266-280.
Kusnezov, N. 1960. La fauna de hormigas del oeste de la Patagonia. Acta Zoologica Lilloana 17:
321-401.
Mayr, G. 1855. Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate
aufgefundenen Ameisen nebst Hinzufügung jener in Deutschland, in der Schweitz und in Italien
vorkommenden Ameisen. Verhandlungen des Zoologisch- Botanischen Vereins in Wien 5: 271-
478.
Mayr, G. 1887. Südamerikanische Formiciden. Verhandlungen der k.k. Zoologisch-Botanischen
Gesellschaft in Wien 37: 511-632.
Roger, J. 1862 Beiträge zur Kenntniss der Ameisen-Fauna der Mittelmeerländler. Berliner
Entomologische Zeitschrift 6: 255-262.
Snelling, R.R. 1975. Descriptions of new Chilean ant taxa. Contributions in Science 274: 1-19.
Snelling, R.R. & Hunt, J.H. 1976. The ants of Chile. Revista Chilena de Entomologia 9: 63-129.
Vander Meer, R.K. 1986. Chemical taxonomy as a tool for separating Solenopsis spp. Pp. 316-326
in: Lofgren, C.S. & Vander Meer, R.K. (eds.). Fire ants and leaf-cutting ants. Biology and
management. Boulder: Westview Press, xv + 435 pp.
Wheeler, G.C. & Wheeler, J. 1980. Supplementary studies on ant larvae: Ponerinae, Myrmicinae
and Formicinae. Transactions of the American Entomological Society 106: 527-545.
Wheeler, W.M. 1914. Ants collected by W.M. Mann in the state of Hidalgo, Mexico. Journal of
the New York Entomological Society 22: 37-61.
Wilson, E.O. & Taylor, R.W. 1967. The ants of Polynesia. Pacific Insects Monograph 14: 1-109.
Figure 1. Monomorium inusuale. A, head in full face view; B, clypeus in oblique view; C, lateral
view of worker; D & E, two lateral views of mesosoma.
Figure 2. Lateral (A) and dorsal (B-D) views of the last abdominal tergum in M. inusuale (Brazil).
Fisher, B. L. 2007. A new species of Probolomyrmex (Hymenoptera: Formicidae) from Madagascar, pp. 146-152. In
Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to
E. O. Wilson – 50 Years of contributions. Memoirs of the American Entomological Institute, 80.
A NEW SPECIES OF PROBOLOMYRMEX FROM MADAGASCAR
Brian L. Fisher
California Academy of Sciences
875 Howard Street
San Francisco, California 94103 USA
bfisher@calacademy.org
ABSTRACT
The worker and queen of Probolomyrmex tani (sp. nov.) from Madagascar are described. This is
the first record of the genus from the Malagasy region. P. tani is widespread throughout western
Madagascar but rare in collections. Collection data indicate that this species nests and forages in
soil or litter in a range of habitats from montane forest to dry spiny bush. P. tani is most similar
in size and general shape to P. guineensis found in western and central Africa.
Key words: Hymenoptera, Formicidae, Probolomyrmex tani, Madagascar, taxonomy, new

species, littoral forest.
Fisher: Probolomyrmex of Madagascar 147
INTRODUCTION
Probolomyrmex was revised globally by Taylor (1965). Additional species were subsequently
described from the Neotropics (Agosti, 1994; O’Keefe & Agosti, 1997) and the Oriental region
(Tanaka, 1974; Brown, 1975; Terayama & Ogata, 1988). This paper describes a new species and
the first record of the genus from the island of Madagascar. The genus Probolomyrmex was
placed in the subfamily Proceratiinae by Bolton (2003). This placement has been confirmed by
molecular work (Ouellette et al., 2006). As in other regions, Probolomyrmex in Madagascar is
cryptic and rarely collected using the traditional arsenal of ant collecting methods (sifted litter,
hand collecting, pitfall, Malaise, light trap, and beating low vegetation). Over 6,000 leaf litter
samples, 4,000 pitfalls and 8,000 additional hand collecting events throughout Madagascar from
1992-2004 turned up only 25 specimens from nine localities.
MATERIAL AND METHODS
The following species of Probolomyrmex were examined from Africa and Asia for comparison
with specimens from Madagascar: bidens (specimens from India), filiformis (specimen from
South Africa), guineensis (specimens from Guinea, Ivory Coast, Cameroon, Gabon, Central
African Republic, Democratic Republic of Congo), and ponce (specimens from India). All
species and type material examined in this study have been imaged and are available on AntWeb
(www.antweb.org). Material was deposited at California Academy of Sciences, San Francisco
(CASC) and Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts (MCZC).
Digital images (Fig. 1-6) were created using a JVC KY-F75 digital camera and Syncroscopy
Auto-Montage (v 5.0) software. All metric measurements were taken at 80× power with a Leica
MZ APO microscope using an orthogonal pair of micrometers and recorded to the nearest 0.001
mm and rounded to two decimal places for presentation. Measurement indices and their
abbreviations used in the paper are based on those used by Taylor (1965). Size and the shape of
the petiolar node are the most important characters for the identification and delimitation of
Probolomyrmex species. Both Brown (1975) and Agosti (1994) found the node measurements
petiolar height (PH) and petiolar length (PNL), as defined by Taylor, somewhat ambiguous (see
definitions below). I include these measurements used by Taylor for comparison with species in
other studies, but I also provided additional measurements (PNH and DPNL, see below). PNH
does not include the subpetiolar process which is variable in species with a thin projecting
cuticular tooth or lamella. DPNL measures the length of the petiolar node in dorsal view and
includes only the posterior margin of the node and not the posteriormost extension of the petiolar
tergum where it surrounds the gastric articulation.
HL Head length: maximum longitudinal length from the anteriormost portion of the projecting
clypeus to the midpoint of a line across the back of the head.
HW Head width: maximum width of head, including the eyes in queens.
LHT Length of the hind tibia, measured in dorsolateral view, from the articulation with the femur,
excluding the proximomedial condyle, to the distal extremity of the tibia.
CI Cephalic index: HW/HL × 100.
SL Scape length: maximum chord length excluding basal condyle and neck.
SI Scape index: SL/HW × 100.
WL Weber’s length: in lateral view of the mesosoma, diagonal length from posteroventral corner of
mesosoma to the farthest point on anterior face of pronotum, excluding the neck.
PW Pronotum width (workers only): maximum width of pronotum in dorsal view.
MW Mesonotal width (queen only): maximum width of mesoscutum in dorsal view.
DPW Dorsal petiolar width: maximum width of the petiole in dorsal view.
PNI Petiolar node index (workers only): DPW/PW × 100.
PH Petiole height: maximum height of petiolar segments in side view, measured vertically from the
posteroventral corner of the subpetiolar process to the level of the petiolar apex.
PNH Petiole node height: maximum height of petiolar node in side view, measured vertically from a
line tangent to the posterior and anteriormost points of the tergosternal suture to the level of the
highest point on the node.
PNL Petiole node length: maximum length of the node, measured longitudinally from the level of the
spiracular process to that of the posteriormost extension of the petiolar tergum, where it
surrounds the gastric articulation.
DPNL Petiole node length: in dorsal view, maximum length of the node, measured from the anterior
margin of the node (excluding articulation with propodeum) to the posteriormost dorsal margin of
node. If anterior or posterior margin is concave, the length is measured from the midpoint of a
line drawn across the margin.
LPI Lateral petiolar index: PNL/PH × 100.
LPNI Lateral petiolar node index: DPNL/PNH × 100.
TL Total length: sum of HL + WL + PNL + length of gaster in lateral view (abdominal segment III
to the apex of last visible abdominal segment, usually pygidium)
Probolomyrmex tani, new species

Figures 1-7
TYPE MATERIAL
Holotype worker, Madagascar: Prov. Antsiranana, Forêt d'Analabe, 30.0 km 72° ENE
Daraina, 13°05'00"S, 049°54'30"E, 30m, 27 Nov 2003, littoral rainforest, (coll. B.L. Fisher et
al.) Collection code: BLF9426, specimen code: CASENT0041505, (CASC) Paratypes: 1
worker and 1 dealate queen with same data as holotype but with specimen codes
CASENT0041506 (1dQ) (CASC) and CASENT0041507 (1w) (MCZC).
ETYMOLOGY
The specific name is an arbitrary combination, to be treated as a noun in apposition.
DESCRIPTION
Worker Measurements (mm): maximum and minimum based on all specimens, n = 22,
(holotype), [paratype].
TL 2.3-2.76 (2.5) [2.5], HL 0.55-0.64 (0.60) [0.59], HW 0.37-0.43 (0.40) [0.40], LHT 0.34-
0.46 (0.39) [0.39], CI 65-75 (67) [67], SL 0.36-0.44 (0.37) [0.37], SI 85-104 (91) [94], WL 0.69-
0.87 (0.77) [0.75], PW (w) 0.28-0.36 (0.31) [0.31], DPW 0.17-0.22 (0.19) [0.19], PNI (w) 56-69
(62) [61], PH 0.25-0.32 (0.28) [0.27], PNL 0.24-0.32 (0.27) [0.28], LPI 86-104 (96) [104], PNH
0.20-0.23 [0.22], DPNL 0.20-0.25 [0.25], LPNI 96-117 [117].
Characters of Probolomyrmex worker as described by Brown (1975: 7). In full face view,
posterior margin of head straight, lateral margins convex, posterolateral corners rounded.
Mandible with apical tooth followed by a series of 6 small denticles. Palp formula 4,2. Eyes
absent. Antennae 12 segmented.
General form of mesosoma and petiole as in figure 5. Dorsal outline of mesosoma in profile
more or less straight, posterolateral corners of propodeum with blunt angles; posterior declivity
of propodeum weakly concave in dorsal view. Dorsal sutures of mesosoma absent. Posterior face
of petiolar node in lateral view more or less straight, with only a slight concave impression and
enclosed laterally and dorsally by a low carina. Subpetiolar process developed as in figure 2;
posteroventral portion projected into an acute angle (shape of posteroventral lamella variable in
shape in specimens examined), without an acute tooth.
Cephalic dorsum with distinct punctures that overlay a surface of fine superficial punctures;
both sizes of punctures separated by distances about equal to their average diameters. Superficial
punctures also present on mesosoma and metasoma but less distinct. Mesosomal dorsum
overlain with larger punctures similar in size to those on dorsum of head. Sides of mesosoma,
petiole, and abdominal segments III and IV with shallow foveolate punctures. Head, mesosoma,
metasoma finely shagreened and moderately shining. Pilosity typical for the genus and limited to
opening of metapleural gland and mandibles. Pubescence whitish, extremely fine, adpressed and
present over entire body, most abundant on abdominal sternites III and IV, and least developed
on dorsum of mesosoma and underside of head. Color reddish brown
Queen measurements (mm): maximum and minimum based on all specimens, n = 3,

[paratype].
TL 2.4-2.6 [2.6], HL 0.56-0.60 [0.60], HW 0.40-0.47 [0.42], LHT 0.32-0.41 [0.41], CI 67-
83 [67], SL 0.33-0.39 [0.39], SI 80-99 [99], WL 0.71-0.80 [0.80], MW 0.32-0.36 [0.34], DPW
0.16-0.23 [0.2], PH 0.27-0.31 [0.28], PNL 0.24-0.29 [0.29], LPI 82-91 [91] PNH 0.19-0.24
(0.20) [0.20], DPNL 0.21-0.29 (0.24) [0.25], LPNI 101-121 (125) [122].
Characters of Probolomyrmex queen as described by Brown (1975: 7). General features as
in figures 2, 4, and 6. Eyes large (maximum length of compound eye: 0.10-0.12), not convex,
situated anterior to the midline of the head. Form of head, mandibles, and petiolar node as in
worker. Surface sculpture, pubescence and color developed as in worker. General form of
mesosoma as in figure 2. Based on specimen CASENT0102226 from Berenty, wing venation
highly reduced, as in previously described species (Taylor 1965).]
DISTRIBUTION AND BIOLOGY
P. tani is infrequently collected but widespread throughout western Madagascar (Fig. 7).
This species has been collected most often in lowland (>180 m) xeric habitats (littoral rainforest,
tropical dry forest, and spiny bush). There are two examples, however, where the species has
been collected in mesic forest: rainforest in Manongarivo (780m) and mountain rainforest in
Binara (1100 m). These two localities are nested within the western dry forest ecoregion and are
adjacent to dry forest habitats. There are no records of Probolomyrmex from the large block of
mesic forest in eastern Madagascar.
The type locality is a littoral forest in northeastern Madagascar. This was chosen as the type
locality to draw attention to the importance of littoral forest in Madagascar. The remnant patches
of littoral forest habitat along the northeastern coast north of Antalaha, such as Analabe and
Ambondrobe, are rich in ant species and contain many endemics, but are highly threatened and
without protective status. These forest patches do not harbor important vertebrate taxa and thus
have not received priority status for conservation in Madagascar. Their rich arthropod fauna,
however, should be recognized as an important component of diversity for reserve design
(Simons et al., 2004).
The majority of specimens in this study were collected from sifted litter, but one specimen
was obtained from under a stone, and one from a pitfall trap. These data suggest that this species
nests and forages in soil or litter in a range of habitats from montane forest to dry spiny bush and
rarely forages above ground. Unlike the other leaf litter nesting ants such as Strumigenys that
demonstrate high species turnover across the island (Fisher 1999, 2000), Probolomyrmex tani
has a wide distribution across many habitats. The subterranean habitat of this species may allow
it to survive in a wide range of ecoregions.
DISCUSSION
Two specimens from the southwest of Madagascar (Tsimanampetsotsa: CASENT0004401

(1w) and Berenty CASENT0102226 (1aQ)) differ from other specimens from the north in
lacking distinct shallow punctures on the mesosoma and metasoma. The queen from Berenty is
also notably smaller HL 0.56 HW 0.40, LHT 0.32, and SL 0.33. Though the size and sculpture
differences are notable, I consider the variation to be intraspecific.
P. tani is most similar to P. guineensis Taylor from Africa in size, and general shape.
Workers of P. tani may be easily distinguished from those of P. guineensis by the shape of the
posterior margin of the petiolar node in lateral view. In P. tani, the posterior margin is more or
less straight, with only a slight concave impression, while in most other species including, P.
guineensis, it is distinctly concave. In addition, the lateral and dorsal carina that encloses the
posterior face of the petiolar node is more developed in P. guineensis than in P. tani. P. tani is
easily distinguished from other ant genera and species in Madagascar by the absence of eyes in
worker, and the presence of frontal lobes reduced to a narrow, sharp ridge between antennal
sockets with antennal insertions clearly visible when viewed from the front.
OTHER MATERIAL EXAMINED
MADAGASCAR: Prov. Antsiranana, Forêt d'Anabohazo, 21.6 km 247° WSW

Maromandia, 14°18'32"S, 047°54'52"E, 120m, tropical dry forest, 11-16 Mar 2001 (coll. Fisher
et al.) - BLF3338: CASENT0458322 (1dQ), CASENT0458323 (1w); Forêt de Binara, 9.4km
235° SW Daraina, 13°15'48"S, 049°36'00"E, 1100m, montane rainforest, 5 Dec 2003 (coll.
B.L.Fisher et al.) - BLF98000: CASENT0043467 (1w), CASENT0043468 (1w),
CASENT0043471 (1w); Montagne des Français, 7.2 km 142° SE Antsiranana (=Diego Suarez),
12°19'22"S, 049°20'17"E, 180m, tropical dry forest, 22-28 Feb 2001 (coll. Fisher et al.) -
BLF3123: CASENT0004400 (1w); R.S. Manongarivo, 12.8 km 228° SW Antanambao,
13°58'36"S, 048°25'24"E, 780m, rainforest, 11 Oct 1998 (coll. B.L.Fisher et al.) - BLF1862(15)-
5 (1w); Mahajanga, Parc National d'Ankarafantsika, Ampijoroa Station Forestière, 40 km 306°
NW Andranofasika, 16°19'15"S, 046°48'38"E, 130m, tropical dry forest, 26 Mar-1 Apr 2001
(coll. Fisher et al.) - BLF3522: CASENT0465467 (1dQ), CASENT0465863 (1w); Parc National
d'Ankarafantsika, Ampijoroa Station Forestière, 5.4 km 331° NW Andranofasika, 16°17'56"S,
046°48'47"E, 70m (coll. Fisher et al.) - BLF3571: CASENT0469570 (1w), CASENT0469571
(1w), CASENT0469572 (1w), CASENT0469573 (1w), CASENT0469574 (1w),
CASENT0469575 (1w), CASENT0469576 (1w), CASENT0469577 (1w), CASENT0469578
(1w), CASENT0469579 (1w); Parc National Tsingy de Bemaraha, 3.4 km 93° E Bekopaka,
Tombeau Vazimba, 19°08'31"S, 044°49'41"E, 50m, 6-10 Nov 2001 (coll. Fisher et. al.) -
BLF4232: CASENT0477984 (1w), CASENT0477985 (1w), CASENT0477986 (1w); Toliara,
Parc National de Tsimanampetsotsa, Forêt de Bemanateza, 20.7 km 81° E Efoetse, 23.0 km 131°
SE Beheloka, 23°59'32"S, 043°52'50"E, 90m, spiny forest/thicket, 22-26 Mar 2002 (coll. Fisher
et. al.) - BLF6263: CASENT0004401 (1w); Toliara, Berenty 12 km N.W. Amboasary, 5-
15.v.1983. (col. J.S.Noyes, M.C.Day) - BM.1983-201: CASENT0102226 (1aQ).
ACKNOWLEDGMENTS
This work was supported in part by the National Science Foundation under Grant No. DEB-
0072713 to B. L. Fisher and C. E. Griswold and DEB-0344731 to B. L. Fisher and P. S. Ward.
P. S. Ward and R. Snelling made valuable comments on the manuscript. Fieldwork that provided
the basis for this work could not have been completed without the gracious support of the
Malagasy people, and the Arthropod Inventory Team (Balsama Rajemison, Helian Ratsirarson,
Jean Claude Rakotonirina, Jean-Jacques Rafanomezantsoa, Chrislain Ranaivo, Coco
Randriambololona, Hanitriniana Rasoazanamavo, Nicole Rasoamanana, Clavier

Randrianandrasana, Pascal Rabeson, Valerie Rakotomalala, Dimby Raharinjanahary and Dylan
Burge)
LITERATURE CITED
Agosti, D. 1994. A revision of the South American species of the ant genus Probolomyrmex
(Hymenoptera: Formicidae). Journal of the New York Entomological Society 102: 429-
434.
Bolton, B. 2003. Synopsis and classification of Formicidae: Memoirs of the American
Brown, W.L., Jr. 1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae,
tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and
Aenictogitini Search. Agriculture (Ithaca, New York) 5: 1-115.
Fisher, B. L. 2005. A model for a global inventory of ants: a case study in Madagascar. Pp. 78-
89 in Jablonski, N.G. (ed.), Biodiversity: A Symposium Held on the Occasion of the
150th Anniversary of the California Academy of Sciences June 17-18, 2003.
Proceedings of the California Academy of Sciences (4) 56, Suppl. I.
Fisher, B.L. 1999. Improving inventory efficiency: a case study of leaf litter ant diversity in
Madagascar. Ecological Applications 9: 714-731.
Fisher, B.L. 2000. The Malagasy fauna of Strumigenys. Pp. 612-710 in Bolton, B. The ant tribe
Dacetini. Memoirs of the American Entomological Institute 65: 1-1028.
O'Keefe, S. & Agosti, D. 1997. A new species of Probolomyrmex (Hymenoptera: Formicidae)
from Guanacaste, Costa Rica. Journal of the New York Entomological Society 105:
190-192.
Ouellette, G. D., Fisher, B. L. & Girman, D. J. 2006. Molecular systematics of basal
subfamilies of ants using 28S rRNA (Hymenoptera: Formicidae). Molecular
Phylogenetics and Evolution 40: 359–369.
Simon F., Powell, G.V.N., Richardson, K.S., Manion, G., Overton, J.M., Allnutt, T.F.,
Cameron, S.E., Mantle, K., Burgess, N.D., Faith, D.P., Lamoreux, J.F., Kier, J.,
Hijmans, R.J., Funk, V.A., Cassis, G.A., Fisher, B.L., Flemons, P., Lees, D., Lovett,
J.C. & Van Rompaey, R.S.A.R. 2004. Mapping more of terrestrial biodiversity for
global conservation assessment. BioScience. 54: 1101-1109.
Tanaka, M. 1974. Description of a new species of the ant of the genus Probolomyrmex Mayr
from Malaysia (Hymenoptera, Formicidae). Entomological Review of Japan 26: 35-37.
Taylor, R.W. 1965. A monographic revision of the rare tropicopolitan ant genus
Probolomyrmex Mayr (Hymenoptera: Formicidae). Transactions of the Royal
Entomological Society of London 117: 345-365.
Terayama, M. & Ogata, K. 1988. Two new species of the ant genus Probolomyrmex
(Hymenoptera, Formicidae) from Japan. Kontyû 56: 590-594.
Figures 1, 3, 5: Probolomyrmex tani worker: holotype CASENT0041505. Figures 2, 4, 6:

Probolomyrmex tani queen CASENT0041506. Figure 7: Collection localities of
Probolomyrmex tani in Madagascar. Map shows major ecoregions: east (white): rainforest,
central (gray): montane forest; central (black): montane shrubland; west (hatch); southwest (gray
hatch): desert spiny bush thicket.
Francoeur, A. 2007. The Myrmica punctiventris and M. crassirugis species groups in the
Nearctic region, pp. 153-185. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in
ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
THE MYRMICA PUNCTIVENTRIS AND M. CRASSIRUGIS SPECIES GROUPS

IN THE NEARCTIC REGION
André Francoeur
Département de biologie
Université du Québec à Chicoutimi
555 boulevard de l'Université
Saguenay, Québec, Canada G7H 2B1
andre_francoeur@uqac.ca
ABSTRACT
This paper deals with two Nearctic species groups of the ant genus Myrmica. The punctiventris
group includes four species: M. pinetorum Wheeler, M. punctiventris Roger, M. semiparasitica
n. sp. and M. punctinops n. sp. The first three occur in the eastern deciduous forest biome, while
M. punctinops is presently known only from California. A key to species for the group is
provided. The crassirugis group is established to include two new species: M. crassirugis and M.
wheelerorum. These ant species are associated with warm climates of western North America.
General ecology and distribution data based on this study are presented. All species are
illustrated with photographs.
Key words: Hymenoptera, Formicidae, Myrmicinae, taxonomy, new species, type designations,
ecology, distribution, Nearctic.
INTRODUCTION
The last taxonomic revision of the Nearctic species of the ant genus Myrmica dates back to
Creighton's book The ants of North America (1950). Subsequently a few species were added and
some nomenclatural changes were proposed (Bolton 1995), but it has become progressively
difficult, often even impossible, to identify with confidence many encountered forms of this
widespread genus, a basic component of the temperate myrmecofauna.
The accumulated material over subsequent years shows a much more diverse genus, richer
in species than previously known. Years ago I undertook revision of Nearctic Myrmica as a part-
time project. Except for one name all nominal taxa, active or synonymised, were examined. Two
fundamental conclusions became evident. Type series of old names often include more than one
species and species concepts for commonly used names in the literature have shifted from the
original concept. Both of these problems plague the name Myrmica emeryana Forel for example.
As a result it quite difficult to use or interpret published data without voucher specimens. The
recognition of species groups based on new sets of morphological characters is one key to
resolve this taxonomic tangle. Ecological biogeography is another.
The results presented in this paper are offered to honor a great and inspiring Naturalist.
Source of material
This taxonomic study is based on personal examination of the type series of the involved
described Myrmica species and infra-specific forms for the Nearctic region and other specimens
(about 800) from institutional collections: Museum of Comparative Zoology, Harvard University
(MCZC); Los Angeles County Museum of Natural History (LACM); Department of
Entomology, United States National Museum of Natural History (USNM); American Museum
of Natural History, New York (AMNH); University of California at Davis (UCDC); W.F Barr
Entomological Collection, University of Idaho (DEUI); Museum of Natural History, Albertson
College of Idaho (ACIC); State Biological Survey, University of Kansas (SBSK); Academy of
Natural Sciences of Philadelphia (ANSP); Department of Entomology, Cornell University
(DECU); Department of Entomology, University of Minnesota (DEUM); Collection des insectes
du Québec, ministère des Resources Naturelles (CIQ); A. Francoeur Collection, Université du
Québec à Chicoutimi (CAFR); Entomology Department, Royal Ontario Museum (ROM);
Muséum d'Histoire Naturelle, Genève (MHNG); and Naturhistorisches Museum Wien
(NHMW). Also a number of private collectors provided material as gifts or voucher depositions;
see acknowledgments for list. Detailed collection data are not given in this paper, but can be
provided upon request.
Measurements and indices

All specimens were measured using a reticule accurate to 0.01 mm coupled to an electronic
instrument displaying absolute values (Wild MMS 235). Biometric standards are adapted from
Seifert (1988), Radchenko & Elmes (1998) and from my previous publications on Myrmica
species except that indices are not expressed here in percentages (Francoeur, 1981, 1984). The
following abbreviations for morphometrics and indices are used.
Measurements. HL: length of head in dorsal view, measured in a straight line from the
anterior point of median clypeal margin to mid-point of the preoccipital margin. HW: maximum
width of head in dorsal view behind the eyes. FMW: minimum width of frons between the
frontal lobe margins. FXW: maximum width of frons between the frontal lobe margins. FL:
length of frontal lobes measured in a straight line from the anterior point of external border (near
base of frontal triangle) to the posterior end (near head surface). SL: maximum straight-line
length of antennal scape shaft (exclusive of the condylar bulb) in dorsal view (perpendicularly to
Francoeur: Myrmica punctiventris and M. crassirugis groups 155
flagellar articulation plane). ML: diagonal length of mesosoma seen in profile, from anterior
margin of pronotal collar to posterior margin of propodeal lobe. PNW: maximum width of
pronotum from above, measured in front of tegulae in queens and males. SPL: maximum length
of propodeal spine in lateral view. SPD: distance between tips of propodeal spine from above.
PL: length of petiole in profile from anterior margin of subpetiolar process to posterior end of
node. PH: maximum height of petiole in profile. PW: maximum width of petiole from above.
PPL: maximum length of postpetiole in profile exclusive of the helcium. PPH: maximum height
of postpetiole in profile. PPW: maximum width of postpetiole from above.
Indices. Cephalic (Cl) = HW/HL. Frontal (Fl) = FXW/FL. Frontal lobe FLI = FMW/FXW.
Scape (Sl) = SL/HW. Mesosomal (MI) = PNW/ML. Spine (SPI) = SPD/SPL. Petiole (PI) =
PW/PL. Petiole node PNI = PW/PH. Postpetiole PPI = PPW/PPL. Postpetiole node PPNI =
PPW/PPH. Postpetiole lateral PPLI = PPH/PPL
MYRMICA PUNCTIVENTRIS GROUP
This group includes three eastern species sharing the following exclusive combination of
morphological characters for females. Clypeus elongate, with anterior margin angulo-convex.
Frontal lobes laterally developed over the antennal socket as triangular or angulo-convex
surfaces; frontal area transverse-rectangular. Eyes oval, of medium size. Antennal scapes evenly
bent basally, smooth and shining, shorter than head length; base flattened dorsally. Mesosomal
profile typically with a rather flattened promesonotum, higher than the propodeum; in dorsal
view promesonotum pear-shaped, posterior end of mesonotum narrower and angulo-convex. In
lateral view metasternal flanges shaped as inverted triangular lamellae. Petiole and pospetiole
short. First gastric (fourth abdominal) segment with large, rounded piliferous punctures. Wings
tinted brown. In addition their distribution occurs within the range of the same biogeographic
biome: the northeastern deciduous forest.
Taken alone a character may not be exclusive to this group. Similar characteristics of the
antennal scapes and the triangular shape of the frontal lobes also occur in the M incompleta
group. This latter group includes M. incompleta Provancher, M. alaskensis Wheeler, M. lampra
Francoeur and M. quebecensis Francoeur. The last two species are workerless parasites
(Francoeur 1984) while M. semiparasitica, described below has retained the worker caste. Large
gastral punctures exist also in M. striolagaster Cole.
Although M. punctinops is a Pacific coast species, inhabiting a very different eco-
biogeographical range, it is provisionally included in the punctiventris group with which it
shares the cephalic features. However, the mesosomal profile is quite different, similar to that of
the crassirugis group, the first gastral segment lacks enlarged punctures, and the erect hairs of
the gastral dorsum are longer than in the eastern species. The frontal lobes are similar to those of
M. alaskensis. Based on general examination of Palearctic species I believe that M. punctinops
(together with another species) belongs to what I named a “Pacific ant fauna” in my Formica
fusca group revison (Formica subelongata Francoeur is an example) (Francoeur, 1972). This
term is used to mean that there exist Nearctic species limited to the Pacific Coast of which
equivalent forms occur in the Asiatic side of the Pacific ocean. Such a faunal trend is also
observed for the genera Formicoxenus (Francoeur et al., 1985) and Temnothorax (unpublished).
Such a phenomenon can be expected for ants as well in the history of the Holarctic region. A
comparative study of this myrmecofauna should be most significative not only in terms of
phylogeny, but also of evolutionary eco-biogeography.
KEY TO WORKERS OF THE MYRMICA PUNCTIVENTRIS GROUP
M. punctiventris group: Antennal scapes seen in profile evenly downcurved basally, base
without any outgrowth. Clypeus entire and prominent, median anterior margin angulo-convex.
Frontal lobes laterally developed over head dorsum into an angular surface with rounded angle.
Eyes oval. In dorsal view promesonotum pear shaped, posteriorly angulo-convex, distinctly
higher than propodeum in profile. First gastric segment with coarse, round punctures. If without
such punctures, mesosomal profile is convex.
1 First gastric segment with coarse, large piliferous punctures; mesosomal dorsum profile
weakly convex to almost flat (eastern North America) ......................................................... 2
– First gastric segment without coarse, large punctures; mesosomal dorsum profile convex;
length of longest erect hairs on gaster dorsum equal to or longer than greatest diameter of
eye (California).............................................................................................. punctinops n. sp.
2 Propodeal spines elongated with a narrow, thin base; spine index < 1.5; subpetiolar process
small and dentiform (larger species with normal queens........................................................ 3
– Propodeal spines short and dentiform with a large, thick base; spine index > 1.5; petiole
sternal process lamelliform (smaller species with microgynes; parasitic species associated
with M. punctiventris)............................................................................. semiparasitica n. sp.
3 Dorsal surface of head coarsely striated; rugae high and thick with flattened top in cross
section; frontal lobes smaller, difference between maximum and minimum width usually <
0.12 mm; propodeal spines distinctly longer than the distance separating their tips, usually
deflected.....................................................................................................punctiventris Roger
– Dorsal surface of head delicately striated, rugulae low and rounded in cross section; frontal
lobes larger, difference between maximum and minimum width usually > 0.12 mm;
propodeal spines shorter, about equal to the distance separating their tips, usually straight ....
.................................................................................................................. pinetorum Wheeler
Myrmica pinetorum Wheeler

Figures 1-7
Myrmica punctiventris subsp. pinetorum Wheeler, 1905: 348 (w, q). (examined)
Myrmica punctiventris pinetorum: Weber, 1950: 217 (m).
Myrmica pinetorum: Creighton, 1950: 102.
Myrmica punctiventris var. isfahani Forel, 1922: 92 (w, q). New synonymy.
Types. M. pinetorum: Lakehurst, New Jersey, USA; lectotype worker, here designated, in
AMNH; paralectotype workers and queens in AMNH, MCZC. M. punctiventris var. isfahani:
Mt. Mitchell, Tyson, North Carolina, USA; lectotype worker, here designated, in MHNG;
paralectotype workers and queens in MHNG.
The syntype series (typus and cotypus) of Forel’s isfahani in MHNG includes specimens of
both M. pinetorum and M. punctiventris collected by him on Mt. Mitchell, Tyson, North
Carolina, at different altitudes, and also workers of M. punctiventris from Virginia. The workers
and alate queens labeled as typus belong to M. pinetorum; all others are workers of M.
punctiventris. The “typus” series is here selected as lectotype and paralectotypes, and this
secures the synonymy of isfahani under M. pinetorum.
Worker. Habitus, figures 1-2, 4. Measurements and indices in tables 1-2. Head in full face
view subrectangular with convex sides; preoccipital (posterior) margin straight, corners largely
rounded. Eyes convex and suboval, located slightly anteriorly of the mid point of the head sides.
Anterior margin of clypeus angulo-convex; lateral wings thin and flat, with 1-3 short rugae. In
dorsal view frontal lamellae laterally developed over the antennal articulation, triangular in
shape with a rounded angle; posterior margin distinctly arrower and ending as a carinae merging
into the head dorsum. Antennae: fossae rather shallow; scapes shorter than head length and
width; in profile base evenly bent, dorsoventrally flattened with feeble dorsal concavity; in
dorsal view shaft width regular along its axis. Funicular segments 3-5 as wide as long, remaining
segments longer than broad; apical club of 4 segments.
Mesosoma, in profile, with mesometasternal external margin horizontally aligned,
promesonotum very feebly convex, almost straight in larger specimens, distinctly higher than
propodeum, both joining through an angle at the mesopropodeal furrow which remains shallow.
In dorsal view promesonotum typically pear-like, posterior end of mesonotum narrower and
anguloconvex. Strigil of protibia with a basal tooth; meso and metatibiae with delicate spurs,
finely and pectinate on distal half. Propodeal lobes small, with a posterodorsal angle. Propodeal
spines straight and acuminate, rather short and thin with a narrow base, shorter than the distance
separating their tips, projecting backwards and upwards at 45°, usually parallel; sometimes a
very feeble curve after the base. Petiole short, about as high as long but narrower; peduncle
hidden by propodeal lobes; node seen in profile with anterior face slightly concave, forming a
right angle with the dorsal surface which is rather flattened, followed by another angle with the
concave posterior face, inclined down to the posterior margin. Postpetiole shorter than high and
wide, height and width about equal; node profile typically with very short anterior and posterior
vertical surfaces, united by a large convex one; sternal process strongly convex and globular,
making 1/3 of the postpetiole height.
Mandibles striate with piligerous punctures. Frons and clypeus with parallel, acute and thin
carinae, widely separated by subopaque, faintly microsculptured surface; reminder of head with
reticulations. Mesosoma generally striatorugulose; rugulae thicker on pleurae and somewhat
sinuous on promesonotum. Antennal fossae with parallel and convex carinae. Petiole and
postpetiole rugose. Gaster smooth and shining; first segment with large round punctures. Long
erect hairs moderately abundant on body; suberect on scapes. Gastric dorsum without distinct
pubescence. General body color light to dark reddish brown; gaster darker; appendages lighter or
more yellowish.
Queen. Habitus, figures 3, 5. Measurements and indices in tables 1-2. Basically similar to
worker in shape of head, characters of sculpture, color and pilosity of body except the following.
Head with three ocelli. Usual distinct mesosomal development of a queen and body size larger.
Sculpture coarser on mesonotum, petiole and postpetiole. Mesopleurae with more delicate
rugulae; transverse groove narrow and deeply impressed. Surface between spines smooth and
shining; sometimes vestigial shagreening on lateral borders. Wings feebly tinted; submarginal
cell of anterior wings partly subdivided posteriorly.
Male. Habitus, figures 6-7. Measurements and indices in tables 1-2. Smaller than queen. In
full face view head slightly longer than broad, narrower before eyes, with shallow elongated
antennal fossae, posterior half evenly rounded. Mandibles elongate, blade subtriangular;
masticatory margin with three apical teeth followed by 2-3 teeth or denticles. Clypeus convex,
anterior margin angulate, finely lamellar mesially. Malar space short. Frontal triangle shallow
and weakly delimited. Frontal lobes poorly developed, but distinct as thin carinae with straight
lateral margins that diverge posteriorly, originating from toruli. Antennae 13-merous; scapes
very short, shorter than or equal to combined length of next 2 segments; in profile scape base
with a very faint dorsal flattening; length of second funicular segment equal to next two;
funicular club of 5 segments. Eyes large and globular. Ocelli small, the antero-median 0.06-0.07
mm in diameter; distance between the posterior ocelli equal to 4-5 x diameter of anterior ocellus.
In lateral view, mesosoma rather long; mesonotum high. Mayrian furrows not impressed,
weakened or absent posteriorly. Spurs of meso- and metatibia weakly pectinate. Metapleural
lamellae small. Wings as in queen, but usually darker (figure 7). Propodeum with two small,
dentiform spines or two more or less developed protuberences marked by carinae, surface
between them smooth and shining; spiracles round and well marked. In profile petiole rather
short, with an anterior peduncle mostly hidden by propodeal lobes; ventral margin straight or
very weakly concave with an anterior denticle; node with an anterior face concave, summit
convex with longitudinal rugulae running to posterior margin. Postpetiole shorter than high and
wide, about as large as wide; anterior and dorsal surfaces of dorsum forming a convex slope with
summit ending posteriorly by a short declivity; sternum longer than high, ventral margin convex.
Head sculpture very fine, mainly shagreened; faint and short rugulae present on front and
malar space, anastomosed on temples, surface punctulate. Mandibles faintly sculptured. Clypeus
very faintly microsculptured, with or without short rugulae. Frontal triangle punctulate. Front
with few rugulae, some reaching the ocellar triangle. Antenna with suberect to subdecumbent
fine hairs longer than the width of segments, but shorter on funicular club. Pronotum densely
shagreened; anterior and lateral areas of mesoscutum smooth and shining, Mayrian furrow as a
thin line from which originate short rugulae, the medial ones longest. Meso- and metapleurae
with parallel rugulae obliquely oriented; transverse grooves feebly impressed. Propodeal
protuberences or spines with a row of fine erect hairs. Body pilosity moderately abundant,
delicate, erect to decumbent. Lateral sides of petiole and postpetiole faintly sculptured, median
area of dorsum smooth and shining. Gaster smooth and shining with some appressed hairs; first
segment with large rounded, piliferous punctures. Body color black to blackish brown;
appendages lighter.
Specimens examined. CANADA. Ontario (ROM), Québec (CAFR, CIQ). USA.

Connecticut (CAFR, LACM), Illinois (MCZC), Indiana (MCZC), Massachusetts (MCZC,
MHNG, USNM), Michigan (AMNH, CAFR, LACM), New-Jersey (AMNH, CAFR, NHMG,
NHMW), New-York (AMNH, LACM, MCZC), North Carolina (LACM, MCZC, MHNG), Ohio
(LACM), Pennsylvania (CAFR, USNM), Tennessee (LACM, MCZC), Virginia (CAFR, LACM,
MCZC, USNM).
Also mentioned in the literature from Oklahoma, Missisipi and South Carolina (D.R. Smith
in Krombein et al., 1979).
Notes. M. pinetorum closely resembles punctiventris but averages smaller in size, with
larger frontal lobes (figure 33), shorter scapes and spines (figure 32 and 34), more delicate body
sculpture. According to material examined, the presence of propodeal spines in males is
observed in the northern part of the species range.
Range. Eastern North America, from southern Canada south probably to the US Gulf states.
Ecology. A forest dweller associated with the eastern deciduous forest biome. Based on
collection data M. pinetorum inhabits not only coniferous forest, but also mixed and rather open
deciduous woods, from dry to more humid conditions. In Pennsylvania I found this ant in a stand
of Populus with Acer and Betula trees. Colonies are small and their nests in different types of
soil: sandy, earthy or rocky, under denuded surface or under leaf litter, lichens, mosses and
rocks. Dennis (1938) reported a nest in a cavity in dead wood. Wesson and Wesson (1940)
observed carton turrets as nest entrances.
Myrmica punctiventris Roger

Figures 8-13
Myrmica punctiventris Roger, 1863: 190 (w).

Myrmica punctiventris: Mayr, 1886: 450 (q)
Myrmica punctiventris: Emery, 1895: 312 (m).
Myrmica punctiventris: Creighton, 1950: 102.
Types. North America (according to Creighton, 1950). Based on known range: eastern N.
A. Type material, if still in existence, should be in the Berlin Museum.
Worker. Habitus, figures 8, 11. Measurements and indices in tables 1-2. Head in full face
view subrectangular with convex sides; preoccipital margin straight and corners broadly
rounded. Eyes small, convex and suboval, located slightly anteriorly of the mid point of the head
sides. Anterior margin of clypeus anguloconvex; lateral wings thin and flat, with 1-3 short rugae.
In dorsal view frontal lamellae laterally feebly developed over the antennal articulation,
approximately triangular or anguloconvex in shape; posterior margin weakly narrower and
ending as a carina merging into the head dorsum. Antennae: fossae rather shallow; scapes
shorter than head length; in profile base evently bent, dorsoventrally flattened with a feeble
dorsal concavity; in dorsal view shaft width regular along its axis. Funiculus segments 3-5 as
large as long, others longer than broad; apical club of 4 segments.
Mesosoma in profile, mesometasternum external margin horizontally aligned,
In dorsal view promesonotum typically pear-shaped, posterior end of mesonotum narrower and
anguloconvex. Strigil of protibia with basal tooth; meso- and metatibiae with delicate spurs,
finely pectinate on the distal half. Propodeal lobes small, with a posterodorsal angle. Propodeal
spines long and acuminate with a narrow base, longer than the distance separating their tips,
projecting backwards and upwards, either almost straight or most often deflected, usually
subparallel. Petiole short, about as high as long but narrower; peduncle hidden by propodeal
lobes; anterior face of node seen in profile slightly concave, forming a right angle with the
somewhat flattened dorsal surface, concave posterior face inclined down to posterior margin.
Postpetiole shorter than high and wide, height and width about equal; node profile typically with
very short anterior and posterior vertical surfaces, united by a large convex one; sternal process
strongly convex and globular, making 1/3 of the postpetiole height.
Mandibles striate with piligerous punctures. Frons and clypeus with parallel, coarse rugae,
separated by subopaque, faintly microsculptured surface; reminder of head with reticulation.
Mesosoma generally striatorugulose; rugae thicker on pleurae and somewhat sinuous on
promesonotum. Antennal fossae with parallel and convex rugae. Petiole and postpetiole rugose.
Gaster smooth and shining; first segment with large round punctures. Long erect hairs
moderately abundant on body; suberect on scapes. Gastric dorsum without distinct pubescence.
General body color light to dark reddish brown; gaster darker; appendages lighter or more
yellowish.
workers in shape of head, characters of sculpture, color and pilosity of body but with the
following usual caste differences: three ocelli present; mesosoma modified for flight; body size
larger. Sculpture coarser on posterior half of dorsum of head, on petiole and postpetiole.
Mesosoma coarsely rugose; surface between rugae faintly microsculptured. Mesopleural
transverse groove rather large and shallow, impressed; katepisternum with widely spaced,
oblique, parallel rugae. Surface between spines smooth and shining. Wings tinted brownish.
Submarginal cell of anterior wing partly subdivided.
Male. Habitus, figures 10-13. Measurements and indices in tables 1-2. Smaller than queen.
In full face view head slightly longer than broad, narrower before eyes, with shallow elongated
antennal fossae, posterior half evenly rounded. Mandibles elongate, blade subtriangular;
masticatory margin with three apical teeth followed by 2-3 denticles. Clypeus convex, anterior
margin angulate. Malar space short. Frontal triangle shallow and weakly delimited. Frontal lobes
poorly developed, but distinct, as thin carinae with straight lateral margins that diverge
posteriorly, originating from toruli. Antennae 13-merous; scapes very long, equal to first 6-7
funicular segments; in profile scape base with faint dorsal flattening; length of second funicular
segment equal to the length of next two; club 5-merous. Eyes large and globular. Ocelli rather
large, 0.07-0.09 mm in diameter; distance between posterior ocelli 3-4 x diameter of anterior
ocellus.
In lateral view, mesosoma elongate; mesonotum high. Mayrian furrows not impressed,
weakened or absent posteriorly. Mesoscutellum anguloconvex posteriorly in dorsal view. Spurs
of meso- and metatibiae pectinate. Metapleural lamellae small. Wings as in queen, usually
darker. Propodeum with more or less developed prominences marked by carinae, sometimes
spiniform, surface between them smooth and shining; spiracles rounded and well marked. In
profile petiole short, with anterior peduncle mostly hidden by propodeal lobes; ventral margin
straight or very weakly concave with an anterior denticule; node with anterior slope straight,
summit rounded with horizontal rugulae. Postpetiole shorter, slightly wider than long; in profile
higher than long; anterior and dorsal surfaces of dorsum forming a convex slope with apex
posterior to center; sternum longer than high, ventral margin straight or convex.
Head sculpture generally fine; rugulae present on front, shorter and stronger around eyes
and malar space, anastomosed on temples, surface punctulate. Mandibles subopaque, faintly
sculptured. Clypeus faintly sculptured, often with short median ruga extending back anterior
margin. Frontal triangle punctulate. Frons with rugulae reaching the ocellar triangle, median part
often only punctulate and shining; lateral lobes reduced to feebly lamellar parallel carinae
originating from toruli. Temples punctate, with a varying abundance of short, partly
anastomosed rugulae. Antennal scapes with suberect hairs over pubescence, most shorter than
width of scape; funiculi with few short, suberect fine hairs on segments over the pubescence,
sparse on club. Mesosoma generally rugulose. Pronotum partly shagreened and mesoscutum
partly smooth and shining; mesoscutellum with longitudinal rugulae; mesopleuron with stronger
rugulae on dorsoposterior corner of katepisternum; anepisternum with an anterior smooth area;
transverse grooves feebly impressed, dark. Propodeal protuberences with a row of fine erect
hairs. Petiole node with rugulae. Postpetiole node smooth and shining with lateral shagreening;
sternum with rugulae. Body pilosity moderately abundant, fine, erect to decumbent; denser on
legs. Gaster smooth and shining; first segment with very faint punctures. Body color black to
blackish brown; appendages lighter.
Specimens examined. BERMUDA: Bermuda Island (DECU, MCZC). CANADA: Ontario

(CAFR, LACM), Québec (CAFR, CIQ). USA. Connecticut (CAFR), District of Columbia
(DECU, MHNG, USNM), Georgia (DECU,MCZC, USNM), Kansas (SBSK), Illinois (AMNH,
MCZC, USNM), Massachusetts (AMNH, LACM, MCZC), Minnesota (DEUM), Michigan
(AMNH, CAFR, LACM), New-Hampshire (CAFR), New Jersey (AMNH, ANSP, CAFR,
LACM, USNM), New-York (LACM, MCZC), North Carolina (MCZC, MHNG), Ohio
(LACM,USNM), Pennsylvania (DECU, DEUM, LACM), Tennessee (LACM), Texas (MCZC),
Virginia (CAFR, MCZC, MHNG, USNM), West Virginia (CAFR).
Also mentioned in literature from Arkansas, Iowa and Nebraska (D.R. Smith in Krombein et
al., 1979).
Notes. M. punctiventris differs from M. pinetorum as follows: averaging larger, coarser

sculpture, frontal lobes less developed and less contrast between maximum and minimum width
(figure 33), longer scapes and spines (figures 32, 34).
Range. Eastern North America, from southern Canada south to US Gulf states.
Ecology. The most frequently collected of the species in this group, this ant appears
primarely as a forest dweller associated with the eastern deciduous forest biome. The species has
been found in a variety of wooded habitats: Laurentian maple, mixed hardwoods (Culvert 1974),
mixed pines, oak, oak-hickory, and mixed stands. Kannowski (1959) reported it from bogs in
Michigan. From dry to humid conditions M. punctiventris seems most prosperous in mesic partly
open woods. Nests are built usually in soil under litter, mosses or rocks, in acorns, occasionally
in wood fragments. Wesson and Wesson (1940) report nest entrances surmounted with turrets of
crude carton. Colony structure, reproduction and genetics were recently documented (Snyder &
Herbers, 1991; Herbers & Mouser, 1997).
Myrmica semiparasitica, new species

Figures 14-19
Types. Holotype worker and 14 paratypes: USA, Long Island, Suffolk County, New York,
U.S.A., 2.v.1982 (S. P. Cover), a nest series collected including 9 workers, 1 ergatogyne, 2
queens and 3 males. Additional paratypes: 11 alate queens, 23 males, CANADA, Mont Rigaud,
Rigaud, Québec, (2 series: 14.viii.1981, 30.vii & 16.viii.1982, captured among swarmings of
Myrmica on the mountain top (Leprince and Francoeur 1986). Holotype worker and most
paratypes in MCZC; paratype workers, queens, and males also in LACM, USNM, and CAFR;
paratype ergatogyne in MCZC.
Etymology. Meaning partly parasitic, since this species is probably a temporary social
parasite.
Worker. Habitus, figures 14, 17. Measurements and indices in tables 1-2. Head in full face
view subquadrate with almost straight sides; preoccipital margin straight and corners rounded.
Eyes rather large, convex and suboval, located on the mid point of the head sides. Anterior
margin of clypeus angulo-convex; lateral wings thin and flat, apex crossed by genal carinae. In
dorsal view frontal lamellae small. Extending partly over the antennal articulation, loosely
triangular or angulo-convex in shape; posterior margin feebly narrower and ending as a carinae
merging into the head dorsum. Antennae: fossae rather shallow; scapes shorter than head length
and width; in profile base evently bent, dorso-ventrally flattened with a feeble dorsal concavity;
in dorsal view shaft width regular along its axis. Funiculus segments 3-5 as large as long, others
longer than broad; apical club of 4 segments.
promesonotum flattened, distinctly higher than propodeum, both joining through an angle at the
mesopropodeal furrow which remains shallow. In dorsal view promesonotum typically pear-like,
posterior end of mesonotum narrower and angulo-convex. Strigil of protibia with basal tooth;
meso and metatibiae with minute to vestigial spurs, very finely and shortly pectinated on the
distal third or only barbulate at apex. Propodeal lobes small, with a posterodorsal angle.
Propodeal spines dentiform and feebly pointed, subparallel, shorter than the distance separating
their tips, projecting backwards and upwards at 45°. Petiole short, higher than wide; peduncle
hidden by propodeal lobes; node seen in profile anterior face slightly concave, forming a right
angle with the dorsal surface which is rather flattened, meeting the posterior face through a
rounded angle; sternum with a lamellar process varying in size. Postpetiole shorter than high and
wide, the latters about equal; node profile typically with very short anterior and posterior vertical
surfaces, united by a large convex one; sternal process strongly convex or subglobular, making
1/3 of the postpetiole height.
Mandibles faintly striated with ciliated punctures. Frons and clypeus with parallel, acute and
thin carinae, spreading fan-wise towards the occiput, separated by faintly microsculptured
surface; reticulations on temples and the preoccipital margin. Antennal fossae with parallel and
convex carinae. Mesosoma generally striato-rugulose; rugulae acute and largely reticulated on
on promesonotum, more rounded and parallel on mesopleurae and dorsum of propodeum. Space
between spines with transverse and faint lines, remaining ventral area smooth. Petiole and
postpetiole rugose. Gaster smooth and shining; first segment with large rounded punctures. Long
and erect hairs moderately abundant on body; suberect on scapes. Gastric dorsum without
distinct pubescence. General body color light to dark reddish brown; appendages lighter or more
yellowish.
workers in shape of head, characters of sculpture, color and pilosity of body except the
following. Head with 3 proportionatly large ocelli. Usual distinct mesosoma development of a
queen and body size larger. Mesosoma rugulose; reticulations on lateral areas of promesonotum;
central part of mesonotum with diverging rugulae. Ventral lamella of petiole digitiform or
angulate, varying in size. Sculpture coarser on posterior half dorsum of head, on petiole and
postpetiole, reduced on anteromedian third of pronotum, behind the collar. Transverse groove of
mesopleurae narrow and impressed; katepisternum with thinner rugulae, separated, obliquely
parallel. Wings translucid. Submarginal cell of anterior wing not always partly subdivided.
Male. Habitus, figures 16-19. Measurements and indices in tables 1-2. Smaller than queen.
In full face view head slightly longer than broad, narrower before eyes, with shallow elongated
antennal fossae, posterior half evenly convex. Mandibles elongated, blade subtriangular;
masticatory margin quite varying, with 1-4 apical teeth followed by 1-2 denticles. Clypeus
convex, anterior margin angulo-convex. Malar space short. Frontal triangle wide and shallow
and more or less delimited. Frontal lobes reduced to minute but distinct carinae, diverging
posteriorly. Antennae with 13 segments; scapes short, equal to less than the first 3 funiculus
segments; in profile scape base with a faint dorsal flattening; length of second funicular segment
equal to next two; funicular club of 5 segments. Eyes large and globular. Ocelli propotionately
large, 0.07-0.08 mm in diameter; distance between the posterior two equal to diameter of 3
anterior ocelli.
In lateral view, mesosoma rather long; mesonotum high. Mayrian furrows not or feebly
impressed. Spurs of meso- and metalegs varying in size and development from minutely
pectinated to normal. Metapleural lamellae small and largely convex. Wings as in queen.
Propodeum with two more or less developed protuberences marked by carinae, surface beteen
them smooth and shining; spiracles rounded and well marked. In profile petiole short, with an
anterior peduncle hidden by propodeal lobes; ventral margin with a lamellar process rectangular
in shape, sometimes angulate anteriorly; summit of node convex with few longitudinal rugulae.
Postpetiole shorter than high and wide; about as large high; in profile anterior and dorsal
surfaces of dorsum forming a convex slope with summit anding posteriorly by a very short
declivity; sternum longer than high, ventral margin more or less convex.
Head sculpture generaly fine; rugulae present or not on frons, shorter and malar spaces,
reticulated on temples. Mandibles very faintly sculptured. Clypeus very faintly microsculptured,
with a short median carina originating from the median angle of anterior margin. Frontal triangle
partly punctulate and smooth. Antennae with suberect fine hairs longer than the width of
segments, but shorter on funiculus club. Pronotum densely shagreened; mesoscutum mostly
smooth and shining. Mayrian furrow of mesonotum as a thin line from which originate short
rugulae. Mesometapleurae with parallel rugulae obliquly oriented, transverse grooves feebly
impressed. Propodeal protuberences with a row of fine erect hairs. Lateral sides of petiole faintly
shagreened with some rugulae; median area of dorsum smooth and shining. Sides of postpetiole
shagreened, smooth elsewhere Body pilosity moderately abundant, fine, erect to decumbent;
denser and longer on head and legs. Gaster smooth and shining; first segment without large
punctures. Body color black to blackish brown; appendages lighter.
Specimens examined. Presently known only from type series.
Notes. This is the smallest species of the group. The reduced size is more obvious in the
queens and males. The sternal process of the petiole comports with the parasitic syndrome of
Wilson (1971). The frontal lobes are similar to the ones of M. punctiventris while the general
size of worker, shorter scapes and spines (figures 32-33), the relative proportions of the waist,
the delicate rugulate sculpture are closer to M. pinetorum.
Range. Presently known from southern Québec to New England.
Ecology. The fact that the species was found in a colony of M. punctiventris with the two
female forms and males, and that it performed nuptial flights support the probability of a
temporary parasitic ant.in nature. S. Cover collected his specimens from a hickory nut in a rich
oak woods of the East Farm Preserve. The host species, as well as M. pinetorum, are found in
red oak stands on and around Mont Rigaud, Québec.
Myrmica punctinops, new species

Figures 20-27
Types. Holotype worker and 17 worker paratypes: USA, Charlton Flat, San Gabriel
Mountains, Los Angeles Co., California, U.S.A., 25.v.2002 (R. R. & G. C. Snelling). Additional
paratypes: 1 worker, same locality, 18.vi.1944 (collector unknown); 2 workers, Brown’s Flat,
San Gabriel Mountains, Los Angeles Co., 3.xi.1960 (E.L. Sleeper); 2 workers, Fort Ord, UCSC
Reserve, Monterey Co., 5.xi.1998 (L. DiGirolamo). Holotype and most paratypes in LACM;
paratypes also in CAFR, MCZC, and UCDC.
Worker. Habitus, figures 20-21. Measurements and indices in tables 1-2. Head in full face
view subrectangular with convex sides; preoccipital margin straight and corners largely rounded.
Eyes small, convex and suboval, located slightly anteriorly of the mid point of the head sides.
Anterior margin of clypeus angulo-convex; lateral wings thin and flat, with 1-3 starting genal
carinae. In dorsal view frontal lamellae large, laterally developed over the antennal articulation,
triangular in shape with a rounded angle; posterior margin narrower and ending as a carinae
merging into the head dorsum. Antennae: fossae rather shallow; scapes shorter than head length
and width; in profile base evently bent, dorso-ventrally flattened with a feeble dorsal concavity;
in dorsal view shaft width regular along its axis. Funiculus segments 3-5 as large as long, other
longer than broad; apical club of 4 segments.
In dorsal view promesonotum typically pear-like, posterior end of narrower and angulo-convex.
Strigil of protibia with a basal tooth; meso and metatibiae with delicate spurs, finely and shortly
pectinate on the distal half. Propodeal lobes small, with a posterodorsal angle. Propodeal spines
straight and acuminate, rather short and thin, with a narrow base, shorter than the distance
separating their tips, projecting backwards and upwards at 45°, usually parallel; in dorsal view a
feeble recurvation after the base often present. Petiole short, about as high as long but narrower;
peduncle hidden by propodeal lobes; node seen in profile anterior face slightly concave, forming
a right angle with the dorsal surface which is rather flattened, follow another angle with the
concave posterior face, inclined down to the posterior margin. Postpetiole shorter than high and
wide, the latters about equal; node profile typically with very short anterior and posterior vertical
surfaces, united by a large convex one; sternal process strongly convex and globular, making 1/3
of the postpetiole height.
Mandibles striated with ciliated punctures. Frons and clypeus with parallel, acute and thin
carinae, widely separated by subopaque, faintly microsculptured surface; reminder of head with
reticulation. Mesosoma generally striato-rugulose; carinae thicker on pleurae and somewhat
sinuous on promesonotum. Antennal fossae with parallel and convex carinae. Petiole and
postpetiole rugose. Gaster smooth and shining; first segment with large rounded punctures. Long
and erect hairs moderately abundant on body; suberect on scape. Gastric dorsum without distinct
pubescence. General body color light to dark reddish brown; gaster darker; appendages lighter or
more yellowish.
Queen and male. Unknown.
Specimens examined. Presently known only from the types.
Range. Presently known only from California. It probably occurs in Oregon.
Ecology. The species was found at 44 m and 1615 m altitude. The Charlton Flat specimens
were sifted from oak and pine forest litter. Those from Fort Ord were found in a kind of
maritime chaparral on sand, dominated by the rare species Arctostaphylos pumila, with a
mixture of thermophilic and cold-tolerant ant species (P.S. Ward, pers. comm.).
MYRMICA CRASSIRUGIS GROUP
This group contains two sister species. The females are characterized by the following
combination of morphological characters. Head rounded. Clypeus not elongated, with anterior
margin evenly convex. Eyes oval. Frontal lobes relatively wide, upwardly produced, with a
strongly convex margin on nearly half of the length, follow carinae curving outwards to merge
with front rugae; front area sagitally elongated. Scapes slightly shorter than head length;
angularly bent basally, bend surrounded by a lamina small to spoonlike. Mesosoma profile
convex with a promesonotal furrow very shallow; body coarsely striato-rugulose. Spines of
medium length, shorter than the distance separating their tips, subdiverging. In lateral view
metasternal flanges bilobate or bidentate from a thin rectangular base. Wings translucid. Western
species occuring in dry and warm habitats.
Myrmica crassirugis, new species

Figures 28-31
Types. Holotype worker and paratypes (all castes): USA, Idaho National Engineering and
Environmental Laboratory (INEEL), Bingham and Clark Counties, Idaho, U.S.A., 2 nest series,
31.viii.1991 (#9172, with males) and 22.ix.1989 (#8929, with alate queens) (W. H. Clark).
Holotype worker and paratypes (all castes) in MCZC; paratypes also in ACIC, BMNH, CAFR,
CASC, LACM, MNHG, and USNM. Topotypes in ACIC.
Etymology. From latin crassus = thick and ruga = wrinkle. The name is based on a feature
of the mesosoma sculpture.
Worker. Habitus, figures 22, 25. Head in full face view, typically rounded, eyes centered
on mid-line cutting the lateral margin, malar margin convex. Head slightly longer than broad
(suboval) with convex sides, a convex or slightly straight (larger specimens) preoccipital margin,
and very broadly rounded preoccipital corner. Median half of anterior margin of clypeus convex,
border of lateral wings thick and sculptured so as to close the antennal fossae. Frontal lamellae
rounded, reduced posterioly to short and narrowly subparallel carinae, merging to dorsum
sculpture. Antennal scape rather long, surpassing the preoccipital corner by the length of the first
two funiculus segments; sharply bent at base (almost right angle), with a reduced to moderately
developed dorsal carina, either thin or lobate and surounding the shaft angle, shortly developing
laterally and associated to a vertical thin and narrow ridge along basal shaft. In dorsal view shaft
narrower along anterior half, about 1/3 the maximal width, particularly just after the bend;
funicular segments 3-6 less than 1.5 times longer than broad; club 4-segmented.
In profile, mesometasternum external margin horizontally aligned, mesosoma evenly
convex, lower posteriorly; mesopropodeal suture not or very shallowly impressed; propodeal
lobes angulate posterodorsally, but not pointed. Strigil of protibia with a basal tooth; middle and
hind tibiae with large pectinate spurs. Propodeal spines of medium length, subparallel,
surpassing slightly the propodeal lobes, usually straight and acuminate, sometimes slighly
deflected, projecting backwards and upwards at just less than 45°. Petiole higher than long and
wide with a short but distinct peduncle; seen in profile the anterior face concave, forming a
rounded right angle with the dorsal surface which is almost straight, while the posterior surface
forms an arch. Postpetiole relatively short and low, very slightly larger than high in profile, node
rounded, sternum subglobular.
Mandibles finely rugulose with ciliated punctures. Frons and clypeus longitudinally rugose;
remainder of head dorsum with reticulated rugae, surface between rugae subopaque and
punctulate. Antennal fossae with faint rugulae. Mesosoma strongly and deeply striated; rugae
with flattened summit, longitudinally aligned, sometimes sinuous on pronotum. Petiole and
pospetiole rugose. Gaster smooth and shining. General body color light to dark brownish-red;
mesosoma often more reddish. Erect body hairs moderately abundant and long; gastric
pubescence very dilute.
Queen. Habitus, figures 23, 26. Basically similar to workers in shape of head, characters of
sculpture, color and pilosity of body except the following. Usual distinct mesosoma
development of a queen and body size larger. Sculpture coarser on posterior half dorsum of
head, on petiole and postpetiole, reduced on anteromedian third of pronotum, behind the collar.
Surface between rugae of mesopleurae distinctly punctate; transverse groove narrow and deeply
impressed; katepisternum with thinner rugae, largely separated, obliquely straight. Surface
between spines subshining, faintly sculptured in upper third, then shagreened. Wings not tinted;
submarginal cell partly divided. Gaster smooth and shining.
Male. Habitus, figures 24, 27. About the size of queen. In full face view head slightly longer
than broad, narrower before eyes, without distinct antennal fossae, posterior half evenly rounded.
Mandibles elongate, blade large and subtriangular; masticatory margin with three apical teeth
followed by 2-3 teeth or denticles. Clypeus more or less bulging, anterior margin weakly
convex. Malar space short. Frontal triangle depressed; lateral carinae paralled, weak, originating
from toruli. Antennae with 13 segments; scapes rather thick and long, equal to first 3-4 funiculus
segments; shaft with a short base, equal before the weak angle to 1/4-1/3 the scape length,
flattened dorsally; funiculus with weakly enlarged club of 4 segments; second funicular segment
longer than first, about twice the length of third. Eyes large and globular, with some erect,
minute hairs. Ocelli small, 0.06-0.08 mm in diameter; distance between the posterior two equal
to diameter of 4 anterior ocelli.
In lateral view, mesosoma rather long. Mayrian furrows not impressed, weakened or absent
posteriorly. Spurs of meso and metalegs well pectinated. Metapleural lamellae small, not
surpassing the petiole peduncle height. Wings as in queen. Propodeum without spines, just a
more or less developed protuberences; spiracles rounded and well marked; surface with long,
well separated, dark rugae. In profile petiole elongated with an anterior peduncle; ventral margin
straight or very weakly concave; node with an anterior slope straight, summit rounded with
horizontal rugulae. Postpetiole shorter, slightly larger than long; in profile higher than long;
anterior and dorsal surfaces of dorsum forming a convex slope with apex posterior to center;
sternum longer than high, ventral margin straight or convex.
Head sculpture generally fine; rugulae present on front, shorter and stronger around eyes
and malar space, anastomosed on temples, surface punctulate. Mandibles subopaque, faintly
sculptured. Clypeus very faintly sculptured, partly smooth, subshining mesially, often with a
short median ruga from anterior margin. Frontal triangle punctulate. Front area with rugulae
reaching the ocellar triangle, median part often only punctulate and shining; lateral carinae
paralled, weak, originating from toruli. Temples punctate, with a varying abundance of short,
partly anastomosed rugulae. Antennal scape with long suberect hairs over pubescence; funiculus
with short, dense and erect pubescence, in addition a second layer of longer, erect dilute fine
hairs. Pronotum and mesoscutum faintly sculptured, partly shagreened and smooth;
mesoscutellum with longitudinal rugulae; mesoplurae shagreened or punctulate with few rugae
or rugulae mostly associated to borders, transverse grooves strongly impressed, dark, base of
some rugae origin. Propodeal protuberances with a row of fine erect hairs. Body pilosity
moderately abundant, fine, erect to decumbent; denser and longer on head and legs. Postpetiole
and gaster smooth and shining. Body color black to blackish brown; appendages lighter.
Specimens examined. All nest series available were recently collected (≥ 1980).
CANADA. British Columbia (CAFR). USA. Idaho (ALBRCIDA, CAFR, DEUI, LACM,
UCDC), Montana (CAFR, USNM), Nevada (CAFR, LACM), Oregon (CAFR), Utah (CAFR,
LACM), Washington (CAFR, LACM).
Notes. Rarely found in collections. Then either not identified or under the names M.
americana, M. emeryana and M. lobifrons. Based upon specimens deposited in CAFR, the
species was reported from the INEEL by Allred and Cole (1971) as Myrmica lobicornis.
The only literature concerning this species is Jackson et al. (1991) which was a
chemotaxonomic study of the species based on specimens from INEEL. It showed that M.
crassirugis is distinctive from 14 described Nearctic Myrmica and that it shares some similarities
with European species of Myrmica.
Range. A western species presently known from southern Canada down to southern USA.
This ant should be found in Arizona, California, New Mexico, Wyoming and southern Alberta in
addition to states and province mentioned hereafter.
Ecology. Collection records indicate this ant lives in semi-xerophilous prairie like sagebrush
habitats and open rather dry coniferous forest of the upper Sonoran zone in mountains. Foragers
have been collected on Atriplex confertifolia, Agropyron cristatum, Pterysia terebinthina,
Balsamorhiza sagitta and Pinus sp. Collecting altitudes include 880 m in Oregon, 1465 - 2380 m
in Idaho, 1870 m in Montana, 1935 - 2129 m in Utah, 1935 - 3134 m in Nevada. Nests are dug
in mineral soils, uncovered or under rock and vegetation (herbs, shrubs). W.H. Clark and P Blom
(personal communication) found M. crassirugis in all of the major soil and vegetation types of
the INEEL site, despite its very cryptic habits. A detailed account of the natural history of this
ant will appear in a report on the ant fauna of that site to be published in Sociobiology.
Myrmica wheelerorum, new species

Figures 32-35
Types. Holotype worker, paratype workers and alate queens: USA, 4 mi NNE Vya, 5600 ft.
elev., Washoe Co., Nevada, U.S.A., 16.vi.1971 (G. C. & J. Wheeler, NEV-2076 & NEV-2078);
additional paratypes (3 workers, 3 alate queens), Florissant, Colorado, no date (W.M. Wheeler).
Holotype and paratypes in LACM; paratypes also in CAFR, MCZC, and USNM.
Etymology. Named after George C. and Jeanette N. Wheeler who collected this ant in
Nevada.
Worker. Habitus, figures 32, 34. Measurements and indices in Table 3-4. Head in full face
view, typically large and rounded, with big oval eyes centered on mid-line cutting the lateral
margin. Head suboval, slightly longer than broad with convex sides, a convex or slightly straight
(larger specimens) preoccipital margin, and very broadly rounded preoccipital corner, malar
margin convex. Mandibles large, with 6-7 teeth and 1-2 denticles. Median half of clypeus some
what bulging, lateral wings flattened; anterior margin of clypeus convex, lateral wings flattened.
Frontal lamellae rounded, reduced posteriorly to short and outcurved carinae merging to dorsum
sculpture. Antennal scape rather long, surpassing the preoccipital corner by the length of the first
two funiculus segments; bent basally at right angle, with a moderately to strongly developed,
then spoonlike, dorsal lamina surounding the shaft angle, shortly developing laterally and
associated with a vertical ridge along basal shaft. In dorsal view shaft narrower along anterior
half, about half the maximal width, particularly just after the bend; funiculus segments 3-6 less
than 1.5 times longer than broad; club of 4 segments.
In profile, when mesometasternum external margin horizontally aligned, mesosoma evenly
convex, lower posteriorly; mesopropodeal suture not or very shallowly impressed; propodeal
lobes angulate posterodorsally. Strigil of protibia with a basal tooth; middle and hind tibiae with
finely pectinated spurs. Propodeal spines rather short and diverging, surpassing slightly the
propodeal lobes, usually straight and acuminate, projecting backwards and upwards at about 45°
to the horizontal. Petiole longer than high and higher than wide, with a short peduncle; seen in
profile the anterior face concave, forming a rounded right angle with the dorsal surface which is
short, often straight or participating in the arch formed by the posterior surface. Postpetiole
distinctly higher and larger than long, about as high than wide, its sternum convex.
Mandibles finely rugulose with ciliated punctures. Sculpture of head delicate and thin; frons
and clypeus longitudinally rugulose; reminder of head dorsum with loosely reticulated rugulae,
surface between rugulae large, subshining and distinctly punctulate. Antennal fossae with faint
rugulae; area of lateral wings of clypeus smooth and shining but rugulose at apex. Mesosoma
strongly and deeply striated; rugae with flattened summit, longitudinally aligned, sometimes
sinuate on pronotum. Petiole and pospetiole rugose. Gaster smooth and shining. General body
color light to dark yellowish-red; gaster somewhat darker and appendages lighter. Erect body
hairs moderately abundant and long; gastric pubescence very dilute.
Queen. Habitus, figures 33, 35. Measurements and indices in Table 3-4. Basically similar to
workers in shape of head, characters of sculpture, color and pilosity of body except the
following. Usual distinct mesosomal development of a queen and body size larger. Head with
three rather large ocelli. Sculpture of head more developed, anastomoses more abundant except
on front and clypeus where carinae remain parallel. Sculpture coarser on lateral parts of
pronotum, mesonotum, petiole and postpetiole. Rugulae of mesopleurae obliquely aligned,
intersurface distinctly punctate; transverse groove narrow and deeply impressed. Surface
between spines subshining, faintly sculptured in upper half, ventral half smooth and shining.
Male. Unknown.
Notes. This is a sister species of Myrmica crassirugis, differing by its striking yellowish
color, exceptional for the genus, a more sturdy head and frontal lamellae, but with more delicate
and reticulated sculpture, a more developed lamina on scape bend, and diverging spines. Its
ecological distribution is probably more southern.
The Wheelers in The ants of Nevada (1986) mentioned in their key to Myrmica species a
concolorous yellowish red form as sp. nov., after my evaluation of their material. It was this new
species, but their series was labelled M. tahoensis Wheeler.
Range. Presently known only from the type localities in Nevada and Colorado.
Ecology. This attractive ant was collected at an altitude of 1680 m (5600 feet) in Nevada.
There are no data for the Florissant specimens. It is probably associated mainly with the upper
Sonoran zone.
ACKNOWLEDGEMENTS
Thanks are due to institutional collection curators, active or retired, for their helpful
cooperation: E. O. Wilson and S. Cover (MCZC), R. R. Snelling (LACM), D. R. Smith
(USNM), P. Ward (UCDC), the late W. L. Brown (DECU), M. Favreau (AMNH), M. B. DuBois
(SBSK), P. J. Clausen (DEUM), R. Béïque and C. Piché (CIQ), C. Bésuchet and B. Merz
(MHNG), M. Fischer and S. Schödl (NHMW). Also collectors kindly provided gifts of
specimens from different states and provinces, particularly W. H. Clark in Idaho, G. C. & J.
Wheeler in Nevada, Paul B. Kannowski and Mary Talbot in Michigan, D. M. Allred and G. F.
Knowlton in Idaho and Utah, S.O. Shattuck in Oregon, J. Heron in Washington, and R. Higgins
in British Columbia. I am also indebted to many students and collaborators, especially Robert
Loiselle and Myriam Tremblay. Financial support for the Myrmica revision was provided by the
National Research Council of Canada and the Museum of Comparative Zoology of Harvard
University.
LITERATURE CITED
Allred, D.M. & Cole, A.C. 1971. Ants of the National Reactor Testing Station. Great Basin
Zoology 104: 1-585, 57 plates.
Culvert, D.C. 1974. Species packing in Caribbean and North temperate ant communities.
Ecology 55: 974-988.
Dennis, C.A. 1938. The distribution of ant species in Tennessee, with reference to ecological
factors. Annals of the Entomological Society of America 31: 267-308.
Emery, C. 1895. Beiträge zur Kenntniss der nordamericanischen Ameisenfauna. (Schluss.)
Zoologische Jarhrbücher. Abtheilung für Systematik, Geographie und Biologie der Tiere
8: 257-360.
Forel, A. 1922. Glanures myrmécologiques en 1922. Revue Suisse de Zoologie 30: 87-102.
Francoeur, A. 1973. Révision taxonomique des espèces néarctiques du groupe fusca, genre
Formica (Formicidae, Hymenoptera). Mémoires de la Société Entomologique du Québec
3: 1-316.
Francoeur, A. 1981. Le groupe néarctique Myrmica lampra (Formicidae, Hymenoptera).
Canadian Entomologist 113: 755-759.
Francoeur, A. & Loiselle, R. 1984. Description du mâle et notice sur la biologie de la fourmi
parasite Myrmica quebecensis (Formicidae, Hymenoptera). Revue d’Entomologie du
Québec 29: 3-11.
Francoeur, A., Loiselle, R. & Buschinger, A. 1985. Biosystématique de la tribu Leptothoracini

(Formicidae, Hymenoptera). 1. Le genre Formicoxenus dans la région holarctique.
Naturaliste Canadien 112: 343-403.
Herbers, J.M. 1997. Microsatellite DNA markers reveal details of social structure in forest ants.
Molecular Ecology 7: 299-306.
Jackson, B.D., Keegans, S.J., Morgan, E.D., Clark, W.H. & Blom, P.E. 1991.
Chemotaxonomic study of undescribed species of Myrmica ant from Idaho. Journal of
Chemical Ecology 17: 335-342.
Leprince, D.J. and Francoeur, A. 1986. Hilltop swarming by ants (Hymenoptera: Formicidae)
in southwestern Quebec and interspecific competition at the swarm marker. Annals of the
Entomological Society of America 79: 865-869.
Mayr, G. 1886. Die Formiciden der Vereinigten Staaten von Nordamerica. Verhandlungen der
k. k . Zoologisch-Botanischen Gesellschaft in Wien 36: 419-464.
Radchenko, A.G. & Elmes, G.W. 1998. Taxonomic revision of the ritae species-group of the
genus Myrmica (Hymenoptera, Formicidae). Vestnik Zoologii 32(4): 3-27.
Roger, J. 1863. Die neu aufgeführten Gattungen und Arten meines Formiciden-Verzeichnisses,
nebst Ergänzung einiger früher gegeben Beschreibungen. Berliner Entomologische
Zeitschift 7: 131-214.
Seifert, B. 1988. A taxonomic revision of the Myrmica species of Europe, Asia Minor, and
Caucasia (Hymenoptera, Formicidae). Abhandlungen und Berichte der
Naturkundemuseums Görlitz 62(3): 1-75.
Smith, D.R. 1979. Superfamily Formicoidea. Pp. 1323-1467 in Krombein, K.V., Hurd, P.D.,
Smith, D.R. & Burks, B.D. (eds.) Catalog of Hymenoptera in America North of Mexico.
Volume 2. Apocrita (Aculeata). Washington, D.C.: Smithsonian Institution Press, pp.
1199-2209.
Snyder, L.A. & Herbers, J.M. 1991. Polydomy and sexual allocation ratios in the ant Myrmica
punctiventris. Behavioral Ecology and Sociobiology 28: 409-415.
Weber, N.A. 1950. A revision of the North American ants of the genus Myrmica Latreille with a
synopsis of the Palearctic species. Annals of the Entomological Society of America 43:
189-226.
Wesson, L.G. & Wesson, R.G. 1940. A collection of ants from southcentral Ohio. American
Midland Naturalist 24: 89-103.
Wheeler, G.C. & Wheeler, J. 1986. The ants of Nevada. Los Angeles: Natural History Museum
of Los Angeles County, 138 pp.
Wheeler, W.M. 1905. An annotated list of the ants of New Jersey. Bulletin of the American
Museum of Natural History 21: 371-403.
Wilson, E.O. 1971. The insect societies. Cambridge, Massachusetts: Harvard University Press,
548 pp.
Table 1. Morphometrics (mm) for the Myrmica punctiventris group.
Measures * pinetorum punctiventris semiparasitica punctinops

Range Mean Range Mean Range Mean Range Mean
Workers (17) ** (25) (5) (7)
HW 0.77 - 0.94 0.84 0.81 - 1.01 0.93 0.76 - 0.84 0.80 0.82 - 1.05 0.89
HL 0.97 - 1.14 1.04 0.98 - 1.18 1.11 0.91 - 0.98 0.95 1.02 - 1.27 1.08
SL 0.74 - 0.87 0.79 0.83 - 0.98 0.93 0.68 - 0.75 0.72 0.87 - 1.06 0.94
FMW 0.37 - 0.44 0.40 0.38 - 0.50 0.46 0.38 - 0.40 0.39 0.36 - 0.45 0.40
FXW 0.50 - 0.62 0.56 0.46 - 0.62 0.55 0.46 - 0.49 0.47 0.56 - 0.73 0.60
FL 0.25 - 0.35 0.30 0.25 - 0.33 0.30 0.24 - 0.28 0.26 0.34 - 0.43 0.38
YD 0.18 - 0.24 0.21 0.20 - 0.26 0.23 0.19 - 0.22 0.21 0.20 - 0.24 0.23
PNW 0.57 - 0.68 0.62 0.59 - 0.74 0.67 0.53 - 0.60 0.56 0.59 - 0.74 0.66
ML 1.30 - 1.48 1.40 1.39 - 1.67 1.56 1.20 - 1.29 1.26 1.37 - 1.73 1.54
SPL 0.23 - 0.35 0.29 0.35 - 0.45 0.41 0.16 - 0.19 0.17 0.22 - 0.34 0.30
SPD 0.30 - 0.40 0.35 0.43 - 0.47 0.48 0.28 - 0.33 0.30 0.37 - 0.52 0.45
PL 0.32 - 0.43 0.37 0.36 - 0.49 0.43 0.27 - 0.35 0.32 0.39 - 0.48 0.43
PH 0.31 - 0.41 0.35 0.31 - 0.40 0.37 0.30 - 0.34 0.33 0.33 - 0.40 0.36
PW 0.23 - 0.33 0.28 0.25 - 0.34 0.31 0.24 - 0.27 0.25 0.27 - 0.37 0.30
PPL 0.26 - 0.40 0.32 0.29 - 0.37 0.34 0.23 - 0.27 0.25 0.25 - 0.37 0.29
PPH 0.37 - 0.51 0.43 0.43 - 0.51 0.47 0.36 - 0.43 0.39 0.37 - 0.47 0.41
PPW 0.35 - 0.48 0.42 0.41 - 0.50 0.45 0.34 - 0.39 0.36 0.36 - 0.47 0.41
Queens (7) (10) (5)

HW 0.81 - 1.01 0.94 0.96 - 1.07 1.03 0.77 - 0.82 0.80
HL 1.01 - 1.21 1.16 1.17 - 1.25 1.22 0.91 - 0.93 0.93
SL 0.75 - 0.89 0.85 0.95 - 1.00 0.97 0.66 - 0.72 0.69
FMW 0.41 - 0.50 0.46 0.48 - 0.54 0.51 0.37 - 0.39 0.38
FXW 0.55 - 0.69 0.64 0.59 - 0.64 0.62 0.44 - 0.47 0.45
FL 0.30 - 0.34 0.32 0.30 - 0.37 0.33 0.22 - 0.27 0.25
YD 0.27 - 0.30 0.29 0.25 - 0.31 0.29 0.23 - 0.26 0.24
PNW 0.74 - 0.92 0.87 0.88 - 0.95 0.92 0.62 - 0.67 0.67
ML 1.53 - 1.86 1.78 1.85 - 2.03 1.95 1.32 - 1.36 1.34
SPL 0.32 - 0.36 0.33 0.36 - 0.43 0.41 0.15 - 0.20 0.17
SPD 0.35 - 0.49 0.45 0.47 - 0.52 0.51 0.28 - 0.33 0.31
PL 0.37 - 0.59 0.51 0.48 - 0.58 0.53 0.33 - 0.35 0.34
PH 0.42 - 0.51 0.47 0.44 - 0.49 0.46 0.31 - 0.34 0.32
PW 0.30 - 0.41 0.36 0.35 - 0.40 0.37 0.24 - 0.27 0.26
PPL 0.32 - 0.47 0.41 0.35 - 0.45 0.40 0.23 - 0.26 0.24
PPH 0.48 - 0.63 0.57 0.51 - 0.63 0.57 0.38 - 0.41 0.39
PPW 0.45 - 0.60 0.54 0.51 - 0.61 0.55 0.34 - 0.39 0.37
Males (4) (10) (5)

HW 0.62 - 0.72 0.67 0.68 - 0.82 0.75 0.59 - 0.65 0.61
HL 0.70 - 0.80 0.76 0.78 - 0.89 0.84 0.65 - 0.71 0.67
SL 0.21 - 0.27 0.24 0.72 - 0.79 0.75 0.20 - 0.24 0.23
FXW 0.26 - 0.33 0.30 0.31 - 0.37 0.34 0.27 - 0.30 0.28
FL 0.16 - 0.19 0.18 0.21 - 0.29 0.25 0.15 - 0.19 0.17
YD 0.27 - 0.34 0.31 0.29 - 0.33 0.31 0.26 - 0.29 0.28
PNW 0.67 - 0.94 0.83 0.71 - 0.91 0.83 0.69 - 0.79 0.73
ML 1.40 - 1.79 1.58 1.49 - 1.84 1.67 1.28 - 1.46 1.34
PL 0.34 - 0.45 0.40 0.41 - 0.50 0.45 0.33 - 0.40 0.34
PH 0.27 - 0.35 0.31 0.31 - 0.39 0.34 0.27 - 0.30 0.28
PW 0.21 - 0.28 0.25 0.26 - 0.35 0.30 0.20 - 0.23 0.21
PPL 0.26 - 0.33 0.29 0.23 - 0.31 0.27 0.22 - 0.25 0.23
PPH 0.36 - 0.43 0.40 0.39 - 0.48 0.43 0.32 - 0.35 0.34
PPW 0.35 - 0.43 0.39 0.39 - 0.46 0.42 0.31 - 0.36 0.32
* See text for measure abbreviations ** Number of specimens

Table 2. Indices for the Myrmica punctiventris group.
Index pinetorum punctiventris semiparasitica punctinops

Range Mean Range Mean Range Mean Range Mean
Workers (17) (25) (5) (7)
CI 0.79 - 0.82 0.79 0.81- 0.86 0.84 0.83 - 0.86 0.84 0.79 - 0.83 0.81
FI 1.68 - 2.15 1.94 1.73 - 2.03 1.85 1.75 - 1.92 1.90 1.50 - 1.80 1.66
FLI 0.68 - 0.78 0.72 0.80 - 0.87 0.83 0.82 - 0.83 0.83 0.60 - 0.67 0.64
SI 0.93 - 0.99 0.95 0.95 - 1.05 1.00 0.89 - 0.91 0.90 1.01 - 1.08 1.05
OI 0.23 - 0.27 0.25 0.23 - 0.26 0.24 0.25 - 0.26 0.26 0.22 - 0.26 0.25
AI 0.43 - 0.45 0.44 0.41 - 0.46 0.43 0.44 - 0.47 0.45 0.42 - 0.44 0.43
SPI 1.09 - 1.43 1.23 1.07 - 1.30 1.17 1.67 - 1.83 1.74 1.33 - 1.68 1.51
PI 0.71 - 0.83 0.78 0.83 - 0.97 0.91 0.73 - 0.93 0.81 0.66 - 0.70 0.69
PNI 0.74 - 0.86 0.81 1.03 - 1.22 1.10 0.77 - 0.83 0.80 0.78 - 0.93 0.83
PLI 0.92 - 1.00 0.96 0.79 - 0.85 0.83 0.94 - 1.11 1.02 0.83 - 0.85 0.84
PPI 1.15 - 1.41 1.35 1.27 - 1.47 1.34 1.37 - 1.54 1.48 1.23 - 1.48 1.34
PPNI 0.93 - 1.00 0.98 0.89 - 0.98 0.96 0.89 - 0.94 0.93 0.95 - 1.05 1.00
PPLI 1.24 - 1.44 1.38 1.31 - 1.53 1.39 1.52 - 1.65 1.59 1.27 - 1.48 1.34
Queens (7) (10) (5)

CI 0.79 - 0.83 0.81 0.82 - 0.87 0.84 0.85 - 0.87 0.86
FI 1.67 - 2.20 1.99 1.76 - 2.03 1.88 1.74 - 2.05 1.82
FLI 0.68 - 0.75 0.72 0.80 - 0.83 0.82 0.81 - 086 0.84
SI 0.88 - 0.96 0.91 0.92 - 0.99 0.94 0.86 - 0.88 0.87
OI 0.29 - 0.33 0.31 0.25 - 029 0.28 0.29 - 0.32 0.30
AI 0.47 - 0.51 0.49 0.45 - 0.49 0.47 0.48 - 50.0 0.49
SPI 1.09 - 1.57 1.36 1.16 - 1.36 1.25 1.50 - 1.87 1.94
PI 0.68 - 0.81 0.71 0.80 - 0.93 0.88 0.69 - 0.82 0.76
PNI 0.71 - 0.80 0.77 1.00 - 1.14 1.09 0.75 - 0.84 0.80
PLI 0.86 - 1.14 0.92 0.77 - 0.85 0.81 0.91 - 0.97 0.95
PPI 1.27 - 1.41 1.33 1.29 - 1.46 1.37 1.42 - 1.58 1.51
PPNI 0.93 - 1.00 0.95 0.93 - 1.00 0.97 0.93 - 0.95 0.93
PPLI 1.35 - 1.50 1.40 1.38 - 1.46 1.41 1.54 - 1.67 1.62
Males (4) (10) (5)

CI 0.87 - 0.90 0.89 0.83 - 0.92 0.88 0.88 - 0.97 0.92
FI 1.63 - 1.83 1.73 1.28 - 1.50 1.39 1.42 - 2.00 1.69
SI 0.33 - 0.38 0.35 0.93 - 1.06 0.99 0.32 - 0.39 0.37
OI 0.44 - 0.47 0.45 0.38 - 0.43 0.42 0.42 - 0.47 0.44
AI 0.48 - 0.57 0.52 0.46 - 0.55 0.50 0.51 - 0.57 0.55
PI 0.54 - 0.71 0.62 0.72 - 0.84 0.79 0.58 - 0.64 0.60
PNI 0.68 - 0.89 0.79 0.82 - 0.96 0.90 0.71 - 0.78 0.75
PLI 0.78 - 0.79 0.79 0.81- 0.94 0.88 0.75 - 0.83 0.80
PPI 1.27 - 1.43 1.34 1.39 - 1.83 1.56 1.28 - 1.44 1.40
PPNI 0.95 - 1.05 0.99 0.84 - 1.08 0.98 0.91 - 1.09 1.00
PPLI 1.21 - 1.43 1.36 1.52 - 1.70 1.60 1.32 - 1.48 7.00
Table 3. Morphometrics (mm) for the Myrmica crassirugis group.
Measures crassirugis wheelerorum

Range Mean Range Mean
Workers (35) (9)
HW 0.81 - 1.09 0.94 0.93 - 1.08 1.01
HL 1.01 - 1.20 1.10 1.09 - 1.20 1.15
SL 0.83 - 1.05 0.93 0.93 - 1.05 0.99
FMW 0.27 - 0.38 0.32 0.28 - 0.35 0.32
FXW 0.38 - 0.51 0.44 0.44 - 0.48 0.46
FL 0.26 - 0.38 0.31 0.30 - 0.43 0.31
YD 0.20 - 0.27 0.23 0.24 - 0.28 0.26
PNW 0.64 - 0.80 0.71 0.68 - 0.75 0.71
ML 1.44 -1.82 1.64 1.60 - 1.72 1.68
SPL 0.23 - 0.38 0.31 0.27 - 0.36 0.31
SPD 0.36 - 0.58 0.43 0.42 - 0.52 0.48
PL 0.43 - 0.54 0.47 0.44 - 0.53 0.49
PH 0.32 - 0.43 0.37 0.37 - 0.39 0.38
PW 0.25 - 0.36 0.30 0.26 - 0.33 0.30
PPL 0.28 - 0.37 0.33 0.30 - 0.36 0.34
PPH 0.37 - 0.50 0.44 0.41 - 0.49 0.45
PPW 0.38 - 0.50 0.45 0.39 - 0.48 0.46
Queens (15) (2)

HW 1.00 - 1.16 1.07 1.05 - 1.08 1.07
HL 1.13 - 1.30 1.22 1.24 - 1.28 1.26
SL 0.88 - 1.05 0.97 0.98 - 1.02 1.00
FMW 0.36 - 0.44 0.39 0.33 - 0.35 0.34
FXW 0.46 - 0.56 0.51 0.51 - 0.53 0.52
FL 0.28 - 0.39 0.34 0.37 - 0.40 0.39
YD 0.28 - 0.33 0.30 0.31 - 0.32 0.32
PNW 0.93 - 1.02 0.98 1.00 - 1.01 1.01
ML 1.99 - 2.19 2.10 2.12 - 2.14 2.13
SPL 0.32 - 0.40 0.36 0.33 - 0.35 0.34
SPD 0.52 - 0.62 0.56 0.50 - 0.54 0.52
PL 0.57 - 0.64 0.60 0.58 - 0.59 0.59
PH 0.47 - 0.51 0.48 0.48 - 0.49 0.49
PW 0.37 - 0.45 0.39 0.36 - 0.38 0.37
PPL 0.37 - 0.43 0.40 0.37 - 0.38 0.38
PPH 0.53 - 0.60 0.56 0.51 - 0.52 0.52
PPW 0.54 - 0.63 0.58 0.56 - 0.57 0.57
Males (11)
HW 0.73 - 0.90 0.82
HL 0.80 - 0.97 0.91
SL 0.40 - 0.63 0.49
FXW 0.29 - 0.36 0.32
FL 0.20 - 0.28 0.24
YD 0.31 - 0.35 0.33
PNW 0.86 - 1.03 0.95
ML 1.75 - 2.09 1.96
PL 0.48 - 0.57 0.52
PH 0.30 - 0.40 0.36
PW 0.28 - 0.32 0.30
PPL 0.32 - 0.38 0.35
PPH 0.40 - 0.46 0.43
PPW 0.40 - 0.49 0.46
Table 4. Indices for the Myrmica crassirugis group.
Index crassirugis wheelerorum

Range Mean Range Mean
Workers (35) (9)
CI 0.77 - 0.91 0.86 1.11 - 1.17 1.14
FI 1.24 - 1.71 1.43 1.38 - 1.55 1.48
FLI 0.88 - 1.21 1.04 0.63 - 0.74 0.70
SI 0.94 - 1.15 0.98 0.95 - 1.00 0.98
OI 0.22 - 0.28 0.25 0.23 - 0.27 0.25
AI 0.41 - 0.47 0.43 0.41 - 0.45 0.42
SPI 1.25 - 1.59 1.39 1.26 - 1.85 1.57
PI 0.55 - 0.71 0.64 0.59 - 0.69 0.63
PNI 0.74 - 1.03 0.81 0.68 - 0.86 0.79
PLI 0.67 - 0.85 0.78 0.75 - 0.86 0.79
PPI 1.17 - 1.48 1.35 1.30 - 1.45 1.36
PPNI 0.95 - 1.11 1.02 0.95 - 1.07 1.02
PPLI 1.17 - 1.48 1.32 1.29 - 1.42 1.33
Queens (15) (2)

CI 0.85 - 0.91 0.88 1.18 - 1.19 1.18
FI 1.28 - 1.69 1.52 1.28 - 1.43 1.35
FLI 0.97 - 1.29 1.16 0.65 - 0.66 0.65
SI 0.88 - 0.94 0.90 0.93 - 0.94 0.94
OI 0.26 - 0.30 0.28 0.30 - 0.30 0.30
AI 0.44 - 0.48 0.46 0.47 - 0.47 0.47
SPI 1.25 - 1.84 1.55 1.43 - 1.64 1.53
PI 0.57 - 0.73 0.64 0.62 - 0.64 0.63
PNI 0.71 - 0.88 0.80 0.75 - 0.78 0.76
PLI 0.77 - 0.86 0.81 0.83 - 0.83 0.83
PPI 1.35 - 1.50 1.44 1.47 - 1.50 1.49
PPNI 1.02 - 1.09 1.04 1.08 - 1.12 1.10
PPLI 1.33 - 1.45 1.38 1.34 - 1.37 1.36
Males (11)
CI 0.87 - 0.94 0.90
FI 1.04 - 1.60 1.32
SI 0.50 - 0.73 0.60
OI 0.39 - 0.44 0.41
AI 0.45 - 0.53 0.49
PI 0.54 - 0.60 0.58
PNI 0.77 - 1.07 0.83
PLI 0.56 - 0.73 0.69
PPI 1.18 - 1.50 1.33
PPNI 1.00 - 1.10 1.06
PPLI 1.16 - 1.44 1.25
Figures 1-3. Myrmica pinetorum. 1. Worker profile without legs. 2. Dorsal view of worker. 3.
Profile of queen.
Figures 4-7. Myrmica pinetorum. Frontal view of head. 4. Worker. 5. Queen. 6. Male. 7. Profile
of alate male body without legs.
Figures 8-10. Myrmica punctiventris. Profile of body. 8. Worker. 9. Queen without wings. 10.
Male.
Figures 11-13. Myrmica punctiventris. Frontal view of head. 11. Worker. 12. Queen. 13. Male.
Figures 14-16. Myrmica semiparasitica. Profile of body without legs. 14. Worker. 15. Queen.
16. Male.
Figures 17-19. Myrmica semiparasitica. Frontal view of head. 17. Worker. 18. Queen. 19. Male.
Figures 20-21. Myrmica punctinops. Worker. 20. Profile of body. 21. Dorsal view of head.
Figures 22-24. Myrmica crassirugis. Profile of body without legs. 22. Worker. 23. Queen
without wings. 24. Male.
Figures 25-27. Myrmica crassirugis. Frontal view of head. 25. Worker. 26. Queen. 27. Male.
Figures 28-31. Myrmica wheelerorum. Profile of body. 28. Worker without legs. 29. Queen
without left antenna and wings. Frontal view of head. 30. Worker. 31. Queen.
Figure 32. Variations of the scape length in the Myrmica punctiventris group.
Figure 33 (top). Variations of frontal lobe development in the Myrmica punctiventris group.
Figure 34 (bottom). Variations in spine length in the Myrmica punctiventris group.
Kohout, R. J. 2007. Revision of the subgenus Aulacomyrma Emery of the genus Polyrhachis F.
Smith, with descriptions of new species, pp. 186-253. In Snelling, R. R., B. L. Fisher, and P. S.
Ward (eds.) Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson –
50 years of contributions. Memoirs of the American Entomological Institute, 80.
REVISION OF THE SUBGENUS AULACOMYRMA EMERY

OF THE GENUS POLYRHACHIS FR. SMITH,
WITH DESCRIPTIONS OF NEW SPECIES
Rudolf J. Kohout
Higher Entomology Section, Queensland Museum
PO Box 3300, South Brisbane, Queensland, 4101, Australia
rudyk@qm.qld.gov.au
ABSTRACT
The subgenus Aulacomyrma Emery of the genus Polyrhachis Fr. Smith is revised. Thirty-six
species are recognized, including seven previously described: P. dohrni Forel, exarata Emery,
excellens Viehmeyer, geometrica Fr. Smith, pallipes Donisthorpe, porcata Emery and sculpta
Emery, and twenty-nine species described as new: P. abdita, annulata, bedeloweryi, breviata,
decora, gentilis, gressitti, impressa, kokoda, lumi, maai, mamba, minima, monticola, orokana,
parva, pelecta, planata, retusa, roberti, roomi, simulans, snellingi, sulcifera, umboi, variegata,
wamuki, wardi and wilsoni. A neotype for P. dohrni Forel is designated. P. sculpta Emery,
previously placed in the subgenus Campomyrma Wheeler, is now recognized as a member of the
subgenus Aulacomyrma. A key based on the worker caste is provided and known distributions of
individual species are given.
Key words: Hymenoptera, Formicidae, Formicinae, Polyrhachis, Aulacomyrma, New Guinea,

Bismarck Archipelago, Indonesia, systematics, new species, distribution.
Kohout: Revision of subgenus Aulacomyrma 187
INTRODUCTION
The subgenus Aulacomyrma was erected by Emery (1921) as a subgenus of Polyrhachis Fr. Smith
with P. porcata Emery as the type species. He characterized the subgenus by the following: first
gastral segment occupying a large portion of the gaster, anteriorly elevated and projecting above
the petiolar scale; scale biconvex with arched dorsal margin terminating on each side in acute
teeth; mesosoma stout, partly marginate; humeri armed with teeth or spines; body sculpture
consisting of regular, more-or-less coarse furrows. Emery (1921) also included another four
species in the subgenus, P. dohrni Forel, exarata Emery, excellens Viehmeyer and serrata Fr.
Smith. Evidently he had doubts about including serrata, but continued to list this species as
Aulacomyrma with a question mark (Emery, 1925), until Donisthorpe (1932) placed it ultimately
under Echinopla Fr. Smith. Chapman & Capco (1951) misspelt the name as Aulocomyrma and
included six species, P. exarata, excellens, mystica Karavaiev, pallipes Donisthorpe, porcata and
serrata, with the last species listed twice (under Polyrhachis and Echinopla). Bolton (1995) listed
nine species of Aulacomyrma, including P. carbonaria Fr. Smith, dohrni, exarata, excellens,
geometrica Fr. Smith, mystica, pallipes, porcata and schizospina Karavaiev. Dorow (1995)
transferred P. carbonaria to the subgenus P. (Myrma) and synonymized P. mystica with P.
cryptoceroides Emery within the subgenus P. (Myrmhopla). Kohout (1998) considered the type of
P. schizospina to be a queen of P. numeria Fr. Smith a species within the subgenus P. (Myrma). P.
sculpta Emery, listed by Emery (1925), Chapman & Capco (1951), Bolton (1995) and Dorow
(1995) in the subgenus P. (Campomyrma), is now recognised as a member of the subgenus
Aulacomyrma, bringing the number of previously described species in the subgenus to seven.
Illustrations
Photographs of some specimens were prepared courtesy of Brian L. Fisher (CASC,

Department of Entomology) and photographed by April Nobile with a JVC KY-F70B 3CCD
digital camera attached to a Leica M420 stereomicroscope. The remaining photographs were
prepared, courtesy of Steve O. Shattuck (ANIC, CSIRO Division of Entomology), by Natalie
Barnett using a ProgRes 3012 scanning digital camera (Jenoptik) attached to a Leica MZ16A
stereomicroscope. All digital images were processed using Auto-Montage (Syncroscopy, Division
of Synoptics Ltd, USA) software. Images of the specimens will also be available in full color on
the ant the following websites: www.calacademy.org/research/entomology/types
(www.antweb.org) and www.ento.csiro.au/science/ants (www.ozants.com.au). All illustrations
depict the primary types, except those of P. pallipes (Figures 19, 22 & 25) representing a
type-compared specimen. Digital images of the holotype of pallipes are available on CASC ant
website (CASTYPE06955).
Methods
References and synonyms of individual species are listed only where relevant to the context
of this paper. For full synonymy citations see Bolton (1995) and Dorow (1995). Publication dates
and the spelling of species epithets and authors’ names follow Bolton (1995). Where a holotype
specimen is mentioned as ‘unique’, this infers that this was the only specimen available for that
species description and no syntypes or paratype specimens are known to exist.
Morphological terms used in this paper closely follow those defined by Bolton (1994), except
for the alitrunk, which in all my previous taxonomic papers has consistently been referred to as the
mesosoma. However, following a specific request by the editors, the morphological terms
referring to the occiput, occipital margin and the occipital corners were, on this occasion, replaced
by the terms vertex, preoccipital margin and the posterolateral corners, respectively, with their
meanings remaining the same.
The present study is based on the worker caste only, as the queens of the majority of species
are unknown. Where queens associated with workers are known, notes are added under the
appropriate species heading. Males are extremely rare and in fact only a single associated male is
known for only one species. As such, it has no taxonomic value for the revision of the group and
has been omitted from this work.
The localities at which ants were collected by the Bishop Museum’s collectors were checked
against that institution’s list of New Guinean localities (BPBM, 1966, unpublished). In some cases
the latitude and longitude co-ordinates, or altitude, are only approximate. The use of the words
“New Guinea” or “Bismarck Archipelago” alone indicate the geographic delimitation of these
regions regardless of the current political boundaries.
Standard Measurements and Indices
Measurements and indices follow those of Kohout (1990: 499): TL – Total length (the
necessarily composite measurement of the entire ant when measured in profile); HL – Head length
(the maximum measurable length of the head in perfect full face view, measured from the
anterior-most point of the clypeal border or teeth, to the posterior-most point of the preoccipital
margin); HW – Head width (width of the head in perfect full face view, measured immediately in
front of eyes); CI – Cephalic index (HW × 100/HL); SL – Scape length (length of the antennal
scape, excluding the condyla); SI – Scape index (SL × 100/HW); PW – Pronotum width (width of
the pronotal dorsum measured at the bases of pronotal spines or teeth); MTL – Metathoracic tibial
length (maximum measurable length of the tibia of the hind leg).
Dimensions for the holotype of each species are in parentheses following the range for all
specimens.
All measurements were taken using a Zeiss (Oberkochen) SR stereomicroscope with an
eyepiece graticule calibrated against a stage micrometer. All measurements are expressed in
millimeters.
Abbreviations
Abbreviations used for specimen data are: Arch. = Archipelago; c. = about (L. circa); I. =
Island; Is = Islands; Mt = Mount; Mtn = Mountain; Mts = Mountains; nr = near; Pen. = Peninsula;
Pltn = Plantation; Prov. = Province; R. = River; Ra. = Range; Rd = Road; rf. = rainforest; w =
worker/s.
Institutions and depositories (with the names of cooperating curators)
ANIC – Australian National Insect Collection, CSIRO, Division of Entomology, Canberra,

Australia ( S.O. Shattuck, R.W. Taylor); BMNH – The Natural History Museum, London, U.K.
(B. Bolton); BPBM – Bernice P. Bishop Museum, Honolulu, HI, U.S.A. (K.T. Arakaki); CASC –
California Academy of Sciences, San Francisco, CA., U.S.A. (B.L. Fisher); MCZC – Museum of
Comparative Zoology, Harvard University, Cambridge, MA., U.S.A. (S.P. Cover); LACM –
Natural History Museum of Los Angeles County, Los Angeles, CA, U.S.A. (R.R. Snelling);
MNHU – Museum für Naturkunde, Humboldt-Universität, Berlin, Germany (Dr F. Koch, A.
Kleine-Möllhoff ); OXUM – Oxford University Museum, Oxford, U.K. (C. O’Toole); QMBA –
Queensland Museum, Brisbane, Australia (C. Burwell, G.B. Monteith); SNSD – Stätliche
Naturhistorische Sammlungen, Museum für Tierkunde, Dresden, Germany (R. Emmrich, U.
Kallweit); UCDC – University of California, Davis, Department of Entomology, CA., U.S.A.
(P.S. Ward).
SYSTEMATICS
Genus Polyrhachis Fr. Smith, 1857
Polyrhachis Fr. Smith, 1857: 58. Type species: Formica bihamata Drury, 1773: 73, pl. 38, figs 7, 8, worker;
by original designation.
Subgenus Aulacomyrma Emery, 1921

Aulacomyrma Emery, 1921: 17 (as subgenus of Polyrhachis Fr. Smith). Type species: Polyrhachis porcata
Emery, 1921: 20, worker; by original designation.
Aulacomyrma Emery; Emery, 1925: 197. (as subgenus of Polyrhachis Fr. Smith). Diagnosis revised.
Aulocomyrma Emery; Chapman & Capco, 1951: 256 (misspelling). Subgenus of Polyrhachis Fr. Smith,
1857.
Aulacomyrma Emery; Hung, 1967: 402. Senior synonym of Johnia Karavaiev, 1927.
Aulacomyrma Emery; Dorow, 1995: 12. As subgenus of Polyrhachis Fr. Smith, 1857.
Aulacomyrma Emery; Bolton, 1995: 21. As subgenus of Polyrhachis Fr. Smith, 1857.
Diagnosis of the subgenus
Worker. Small ants (HL 1.00-1.70) with general characteristics of the genus. Anterior clypeal
margin arcuate, majority of species with small notch or shallow emargination medially. Frontal
carinae sinuate with laminate lobes; frontal triangle usually indistinct or visible only in certain
illuminations and views. Eyes ranging from flattened to strongly protuberant. Preoccipital margin
with a distinct ridge posteriorly and laterally in all but one species (decora), where it is suppressed
by overlying cephalic sculpture. Mesosoma with dorsum showing all stages from fully marginate
to completely immarginate. Pronotal humeri armed with spines, teeth or simply angular.
Promesonotal suture present. Mesonotal and propodeal dorsa virtually fused. Metanotal groove
completely lacking or only very weakly indicated by a hairline break in dorsal sculpture and/or by
shallow incisions of lateral margins in species with marginate mesosoma. Propodeum terminating
posteriorly in a pair of short teeth, tubercles or simply rounded, except in one species (wilsoni),
where it is armed with long, mostly outwardly directed spines. Petiole scale-like; dorsal margin
acute, usually entire, but occasionally with intercalary teeth or short spines, medially emarginate
or unevenly jagged, laterally armed with short spines or teeth. In several species (decora and its
allies) petiole deeply transversely sculptured with dorsal margin more-or-less blunt. Base of first
gastral segment usually concave medially, accomodating posterior face of petiole; dorsal margin
of concavity acute or blunt and often raised above dorsum of segment; first gastral segment
sometimes truncate or simply convex. Sculpture of head and body mostly consisting of regularly
spaced striae that are either longitudinal or convoluted producing a characteristic “geometrical”
appearance; gastral sculpture more modest, usually consisting of longitudinal striae; most distinct
on sides of first gastral segment.
Queen. Very much like worker with usual characters identifying full sexuality, including
three ocelli and complete thoracic structure with wings. Armament of pronotal humeri somewhat
reduced, lateral petiolar spines distinctly shorter. Sculpture of head and body similar to that of
worker, pattern of striation following structural characteristics of fully developed mesosoma.
Male. A single associated male is known for only one species (wardi) and as such has been
omitted from this work. Hung (1967) studied the male genitalia of Polyrhachis and noted that they
proved to be quite similar throughout the genus.
Distribution and biology. The known distribution of the subgenus ranges from the islands of
eastern Indonesia (Aru, Misool), across the New Guinean mainland and adjacent islands (Umboi)
to the Bismarck Archipelago, including New Britain and New Ireland. Little is known about their
biology and nesting habits, however, a small colony of P. dohrni was collected by the author from
a dry hollow twig on a living tree at the edge of lowland rainforest. The internal walls of the twig
cavity were lined with a little silk. The only other record of a nest is of P. wardi, collected by Dr
Phil Ward, also from a dry twig of a rainforest tree. The colonies of both species were rather small,
with only a few workers (5 and 11 respectively, including 2 and 3 alate queens and a single male).
If such a nesting pattern is the norm for other species of the subgenus, that might explain the
general scarcity of Aulacomyrma material even in the best collections. This might also account for
the fact that all previously described species were based on unique specimens. In spite of the
exemplary cooperation of the listed institutions, this situation has only marginally improved in this
revision with almost half the new species described from single specimens.
KEY TO AULACOMYRMA SPECIES

(Based on worker caste)
1 Anterior face of first gastral segment distinctly concave medially, accomodating posterior
face of petiole; concavity delimited dorso-medially by acute or blunt margin that is often
raised above dorsal surface of segment .................................................................................. 2
- Anterior face of first gastral segment medially truncate or convex; if weakly concave then
concavity too shallow to accommodate posterior face of petiole and its dorsal border rounded,
without distinct margin ........................................................................................................... 3
2 Mesosomal dorsum distinctly marginate laterally along its entire length; margins interrupted
only at sutures ....................................................................................................................... 10
- Mesosomal dorsum with ill-defined margins, formed in part by outermost dorsal striae or with
margins completely lacking and dorsum rounding onto the sides in laterally uninterrupted
curve ..................................................................................................................................... 27
3 Sculpture of mesosomal dorsum consisting of mostly longitudinal striae, that weakly

converge anteriorly on pronotal dorsum ............................................................... bedeloweryi
- Sculpture of mesosomal dorsum with striae forming variously curved, transverse, V-, U- or
semicircular-shaped pattern .................................................................................................... 4
4 Eyes convex or protuberant, clearly breaking cephalic outline in full face view.................... 5
- Eyes flattened and somewhat sunk into shallow concavity in cephalic sculpture, at most
touching but not extending beyond lateral cephalic outline in full face view......................... 9
5 Petiole with both faces distinctly striate ................................................................................. 6

- Petiole with both faces finely reticulate-punctate or shagreened............................................ 8
6 Mesosoma distinctly antero-posteriorly compressed; propodeal declivity very high, vertical

to main axis of body .................................................................................................... gressitti
- Mesosoma relatively long, not antero-posteriorly compressed; propodeal declivity relatively
low, oblique to main axis of body........................................................................................... 7
7 Pronotal dorsum rather massive, particularly anteriorly, distinctly convex in lateral view;
petiole with lateral spines curved outwards and backwards; first gastral segment uniformly
finely reticulate-punctate ...............................................................................................kokoda
- Pronotal dorsum in lateral view only moderately convex anteriorly, rather flat towards
promesonotal suture; petiole with short, upturned lateral teeth; first gastral segment distinctly
longitudinaly striate laterally, finely reticulate-punctate dorsally.................................. decora
8 Pronotal humeri with short, acute spines; dorsal petiolar margin with pair of short intercalary
teeth and pair of blunt angular projections situated between them and lateral petiolar teeth....
....................................................................................................................................breviata
- Pronotal humeri with broadly-based, angular teeth and lateral margins distinctly raised and
continuous to midlength of the pronotum; dorsal petiolar margin jagged, without distinct
intercalary teeth .............................................................................................................pelecta
9 Larger species (HL > 1.56); petiole finely shagreened with sharp dorsal margin bearing pair
of short, acute, intercalary teeth.......................................................................................roomi
- Smaller species (HL <1.40); petiole with both faces distinctly transversely striate; dorsal
margin blunt, intercalary teeth lacking ......................................................................... mamba
10 Clypeus unsculptured, smooth and shiny.............................................................................. 11

- Clypeus sculptured as rest of head........................................................................................ 13
11 Petiole with dorsal margin entire; head smooth and very shiny ....................................... maai
- Petiole with dorsal margin medially emarginate with a pair of intercalary teeth flanking
emargination; head distinctly, mostly longitudinally, striate ................................................ 12
12 Petiole strongly transverse, with rather distinct, acute intercalary teeth; propodeal dorsum
terminating in very prominent, somewhat upturned, tooth-like prominences, with blunt
apices; mandibles distinctly reddish-brown; pilosity and pubescence very sparse to absent....
...................................................................................................................................... roberti
- Petiole about as wide as high, with rather blunt, short intercalary teeth; propodeal dorsum
terminating in rounded angles; mandibles, except narrow masticatory border, black; pilosity
and pubescence moderately abundant but not concealing underlying sculpture......... simulans
13 Eyes moderately convex, in full face view exceeding, even if only marginally, lateral outline
of head .................................................................................................................................. 14
- Eyes rather flat, in full face view sometimes almost level with, but not exceeding lateral
outline of head ...................................................................................................................... 25
14 Propodeal dorsum separated from declivity by distinct, transverse ridge that may be straight,
curved or arched and somewhat weakened medially ............................................................ 15
- Propodeal dorsum not separated from declivity by transverse ridge or margin; both surfaces
meeting in medially uninterrupted, although abrupt, curve .................................................. 18
15 Lateral margins of mesonotal-propodeal dorsum confluent, converging posteriorly and

terminating in blunt angles with their posterior margins continued medially, forming almost
straight, transverse ridge that may be somewhat weakened medially..............................parva
- Lateral margins of mesonotal-propodeal dorsum not confluent; margins of mesonotum
converging posteriorly, those of propodeum subparallel or diverging, terminating posteriorly
in pair of distinct spines, laterally directed teeth or rounded prominences with their posterior
margins continued medially, forming more-or-less inward bowed, somewhat arched,
transverse ridge..................................................................................................................... 16
16 Propodeal dorsum armed posteriorly with very distinct, long, outwards, backwards and
weakly downwards directed spines; their posterior margins forming a deeply, inwardly
bowed and distinctly medially arched, transverse ridge ................................................wilsoni
- Propodeal dorsum armed posteriorly with short teeth or rounded prominences; their posterior
margins forming a weakly inwardly bowed ridge, that may be somewhat weakened medially
............................................................................................................................................. 17
17 Propodeal teeth acute, broad-based and directed outwards; first gastral segment only very
finely longitudinally striate laterally; numerous hairs and appressed pubescence present on all
body surfaces ...................................................................................................................wardi
- Propodeal teeth reduced to small, rounded prominences in dorsal view; first gastral segment
distinctly longitudinally striate laterally; hairs and appressed pubescence virtually absent
from most body surfaces............................................................................................... exarata
18 Sculpture of mesosomal dorsum consisting of longitudinal, widely-spaced striae, with only

about 13 striae spread across width of pronotal dorsum; mesonotal-propodeal dorsum
distinctly transverse, almost twice as wide as long..................................................... sulcifera
- Sculpture of mesosomal dorsum consisting of mostly longitudinal, narrowly-spaced striae,
with about 20 or more striae spread across width of pronotal dorsum; mesonotal-propodeal
dorsum only marginally wider than long .............................................................................. 19
19 Smaller species (HL <1.10) .................................................................................................. 20

- Larger species (HL >1.20) .................................................................................................... 21
20 Sides of mesosomal dorsum subparallel; propodeal dorsum terminating posteriorly in

distinct, rounded angles with their posterior margins weakly curved inwards, continuing
medially for some distance but failing to meet ............................................................. minima
- Sides of mesosomal dorsum distinctly converging posteriorly; propodeal dorsum completely
unarmed, descending into declivity across its full width .............................................. gentilis
21 Dorsum of pronotum transversely convex between lateral margins ..................................... 22

- Dorsum of pronotum virtually flat................................................................................ planata
22 Propodeal dorsum terminating in dentiform angles, that are not distinctly raised above its
surface; striae on pronotal dorsum converging anteriorly..................................................... 23
- Propodeal dorsum terminating in distinct teeth; their lateral margins raised, leaving the
posterior margin of propodeum weakly concave medially; striae on pronotal dorsum
longitudinal, more-or-less parallel, not converging anteriorly........................................... lumi
23 Vertex with transverse, strongly bowed, almost semicircular striae; lateral margins of
mesonotal-propodeal dorsum confluent, converging posteriorly, not distinctly incised at
metanotal groove (New Guinea)....................................................................................sculpta
- Vertex longitudinally striate; lateral margins of mesonotal-propodeal dorsum distinctly
incised at metanotal groove; mesonotal margins converging posteriorly, propodeal margins
more-or-less parallel-sided ................................................................................................... 24
24 Femora distinctly bicolored; light reddish-brown with very dark brown apices; eyes only
moderately convex, in full face view just marginally breaking lateral outline of head
(Bismarck Archipelago)................................................................................................. dohrni
- Femora unicolored, very dark brown or black; eyes convex, clearly extending beyond lateral
outline of head (Umboi Island) ....................................................................................... umboi
25 Anterior face of first gastral segment with acute margin that is moderately raised above face
of segment; legs distinctly light, mostly yellow or reddish-yellow ..............................pallipes
- Anterior face of first gastral segment with blunt margin that is not raised above face of
segment; legs generally dark reddish-brown or black........................................................... 26
26 Eyes situated close to lateral outline of head in full face view; pronotal dorsum with greatest
width at about midlength of segment (New Ireland) ...................................................... abdita
- Eyes situated well away from lateral outline of head in full face view; pronotal dorsum with
greatest width across humeri (Irian Jaya) ..................................................................variegata
27 Propodeal dorsum separated from declivity by distinct, rather acute, transverse ridge ........ 28
- Propodeal dorsum not separated from declivity by transverse ridge; both surfaces
medially confluent ................................................................................................................ 29
28 Eyes strongly protuberant, almost fully extending beyond lateral outline of head in full face
view; mesonotal-propodeal dorsum strongly transverse, distinctly wider than long, sides
subparallel....................................................................................................................orokana
- Eyes only moderately convex, situated in shallow depression in cephalic sculpture and only
partly extending beyond lateral outline of head in full face view; mesonotal-propodeal
dorsum about as long as width across posterior transverse ridge; sides of mesonotal-propodeal
dorsum converging posteriorly ...................................................................................impressa
29 Petiole with dorsal margin entire .......................................................................................... 30

- Petiole with dorsal margin more-or-less emarginate medially.............................................. 33
30 Smaller species (HL <1.15) .................................................................................................. 31

- Larger species (HL >1.34) .................................................................................................... 32
31 Petiole strongly transverse, both faces finely shagreened; propodeal dorsum terminating in a
pair of small rounded prominences...............................................................................wamuki
- Petiole about as wide as high, both faces distinctly striate; propodeal dorsum terminating in
downward and weakly inward curved angles ............................................................. snellingi
32 Pronotal dorsum only weakly transversely convex, almost flat, with spines directed
anteriorly; mesosomal dorsum with sides subparallel; mesosoma with prominent, regularly
spaced, longitudinal striae, those on pronotal dorsum anteriorly converging; anterior face of
first gastral segment with acute margin that is strongly raised above dorsal face of segment ..
...................................................................................................................................excellens
- Pronotal dorsum strongly transversely convex with spines directed more-or-less laterally;
mesosomal dorsum with sides converging posteriorly so that pronotum is notably broader
than propodeum; pronotal dorsum with inverse, narrowly U-shaped to semicircular striae;
mesonotal-propodeal dorsum with V-shaped striae; anterior face of first gastral segment with
acute margin that is only moderately raised above face of segment .............................porcata
33 First gastral segment finely shagreened ...................................................................... annulata

- First gastral segment distinctly longitudinally striate laterally ............................................. 34
34 Propodeal declivity oblique to main axis of body; femora of front legs very light,
yellowish-brown, those of mid- and hind legs very dark, reddish-brown or black...................
............................................................................................................................... geometrica
- Propodeal declivity abrupt, vertical to main axis of body; femora of all three pairs of legs
uniformly dark reddish-brown or black ................................................................................ 35
35 Dorsal petiolar margin medially emarginate with distinct acute teeth flanking emargination;
larger species (HL 1.37)............................................................................................ monticola
- Dorsal petiolar margin only narrowly and shallowly emarginate without distinct teeth
flanking emargination; smaller species (HL 1.12).......................................................... retusa
The species-groups
The thirty-six species of Aulacomyrma can be divided, on the basis of external morphological
characters, into two, mostly intergrading, species-groups:
The dohrni – group, including abdita sp. nov., dohrni Forel, exarata Emery, gentilis sp. nov.,
lumi sp. nov., minima sp. nov., pallipes Donisthorpe, parva sp. nov., planata sp. nov., roberti sp.
nov., sculpta Emery, simulans sp. nov., sulcifera sp. nov., umboi sp. nov., variegata sp. nov.,
wardi sp. nov. and wilsoni sp. nov.
The porcata – group, including annulata sp. nov., bedeloweryi sp. nov., breviata sp. nov.,
decora sp. nov., excellens Viehmeyer, geometrica Fr. Smith, gressitti sp. nov., impressa sp. nov.,
kokoda sp. nov., maai sp. nov., mamba sp. nov., monticola sp. nov., orokana sp. nov., pelecta sp.
nov., porcata Emery, retusa sp. nov., roomi sp. nov., snellingi sp. nov. and wamuki sp. nov.
The margination of the mesosoma or its partial or complete loss is an important character,
evident in a number of species that are intermediate between the dohrni-group, which includes
species with a laterally marginate mesosomal dorsum and the porcata-group with a partly or
completely immarginate condition. The transition is illustrated by the following series of species:
exarata → planata →sculpta → excellens → porcata → orokana →geometrica →decora. In the
first species, exarata, a member of the dohrni-group, the mesosoma is fully laterally and
posteriorly, marginate. The second species, planata, still has complete lateral margins, but the
posterior propodeal margins extend only partly towards the midline as short ridges. They fail to
meet medially and there is a gap through which the dorsum meets the declivity. In sculpta, which
could be considered intermediate between the dohrni- and porcata-group, the lateral margins of
the mesosoma are less distinct posteriorly, leaving the propodeum only weakly marginate laterally
and completely immarginate posteriorly. The margins of the mesosoma are incomplete in
excellens and related species, with the pronotal margins weak and extending from the pronotal
spines for only part of the length of the pronotum. The margins merge with the outermost dorsal
striae and curve onto the sides of pronotum, failing to reach the promesonotal suture. The
mesonotal-propodeal dorsum is usually only partly marginate, leaving one or the other laterally
immarginate. In porcata there is only an indication of weak lateral margins of the mesosoma, with
the propodeum terminating in barely distinct tubercles. The propodeum in orokana has a distinct
transverse posterior margin, but the lateral margins of mesosoma are completely lost. A complex
of species, represented in the series by geometrica, has the mesosoma virtually immarginate,
although in some the outermost striae of the mesonotal dorsum form ill-defined lateral margins.
Finally decora and its allies have the mesosoma completely immarginate.
Within the series, changes in other characters are also evident, notably in the convexity of the
mesosomal dorsum and the degree of concavity of the anterior face of the first gastral segment. In
species related to dohrni and excellens, the mesosomal dorsum between the lateral margins is
almost flat or only moderately transversely convex, and the first gastral segment is more-or-less
distinctly concave anteriorly. By contrast, in species related to porcata and geometrica, the
dorsum of mesosoma is distinctly transversely convex and rounds onto the sides in an
uninterrupted curve. Also, as the series progresses, the first gastral segment tends to be somewhat
less concave and finally culminates in species having the anterior face only truncate or even
convex, as in decora. In most species the surface of the petiole is shagreened with the dorsal
margin distinctly acute and armed with rather well developed lateral spines. In species allied to
decora, the petiolar node in lateral view is blunt with both faces deeply transversely striate and
armed with rather short lateral teeth. The affinities of these species are not immediately apparent,
but porcata and snellingi, both possessing a transversely striate petiole with a rather acute dorsal
margin, evidently represent intermediate forms.
Parallel to the characters discussed above is a change in the form of the sculpture of the
mesosoma which, in dohrni-group species, consists mostly of simple, longitudinal striation. The
sculpture tends to become more complicated as the series progresses, with the dorsal striation in
porcata-group species curved into simple V- or U-shapes, notably on the pronotal dorsum. The
sculpture of species at the end of the series is even more complex, featuring variously curved
patterns, giving them a characteristic “geometrical” appearance.
Polyrhachis dohrni species-group
This group is characterized by the following: mesosoma distinctly and fully laterally
marginate with margins interrupted only at sutures. Pronotal humeri armed with broad-based
teeth, short spines or acute angles which are laterally continuous with the pronotal margins.
Propodeal dorsum terminating in dentiform angles or tubercles, but also in distinct, posteriorly
directed tooth-like prominences as in roberti and lumi, or in laterally directed blunt teeth as in
wardi. In one species, wilsoni, the lateral margins are greatly extended and terminate in long,
outwardly curved, acute spines. Posterior margination of propodeum varying from complete
transverse ridge separating dorsum from declivity as in exarata and parva, where ridge is straight,
and wilsoni, where it is deeply curved inwards, to a situation where posterior margin is lacking and
both surfaces are confluent, as in dohrni or pallipes. Petiole scale-like with anterior and posterior
faces convex and dorsal margin acute, with rather distinct lateral spines or teeth. Species related to
dohrni have dorsal petiolar margin entire, or at most weakly jagged, as in lumi; in roberti and
simulans dorsal petiolar margin medially emarginate with short teeth or denticles flanking
emargination. Anterior face of first gastral segment in all but one species (roberti) distinctly
concave, accommodating posterior face of petiole; anterodorsal margin of concavity acute and
weakly elevated, as in pallipes and planata, or blunt and not at all raised above dorsal surface of
segment, as in abdita and simulans. Sculpture of head and mesosomal dorsum consisting of
mostly longitudinal striation, that in most species tends to be anteriorly convergent on pronotum.
In two species, roberti and simulans, longitudinal sculpture is virtually absent from clypeus that is
virtually smooth, only very finely reticulate-punctate.
Polyrhachis (Aulacomyrma) abdita, new species

Figures 1, 4, 7
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, New Ireland Prov., Lelet Plateau, c. 03º20’S, 151º56’E,
800-1000m, 19-24.vii.1984, R.J. Kohout acc. 84.95 (worker). PARATYPES: data as for holotype (12
workers). Type distribution: holotype and 2 paratypes in ANIC; 2 paratypes each in BMNH, CASC, LACM,
MCZC and QMBA.
WORKER
Dimensions: TL c. 4.64-5.44 (5.19); HL 1.15-1.37 (1.31); HW 1.03-1.22 (1.18); CI 88-90

(90); SL 1.22-1.40 (1.40); SI 114-119 (119); PW 1.00-1.18 (1.18); MTL 1.28, 1.09-1.34 (1.28) (13
measured).
Anterior clypeal margin arcuate with shallow emargination medially; in profile clypeus
virtually straight with very weakly impressed basal margin, indicated laterally by a faint line that
barely breaks cephalic sculpture. Frontal carinae sinuate, with laminate lobes. Sides of head very
weakly convex, almost straight in front of eyes before converging towards bases of mandibles.
Eyes rather flat, with anterior margins somewhat sunk into shallow depressions in cephalic
sculpture; in full face view almost level with, but failing to break lateral cephalic outline.
Mesosoma marginate along entire length. Pronotal humeri armed with broad-based, angulate,
acute teeth, that are laterally continuous with pronotal margin. Dorsum of pronotum widest at
midlength. Promesonotal suture distinct; metanotal groove lacking dorsally, position marked by
shallow emarginations in lateral margin. Propodeum with lateral margins only narrowly rounded
posteriorly; dorsum descending into virtually vertical declivity in an unobstructed curve. Petiole
with sharp dorsal margin, its apex higher than anterodorsal margin of first gastral segment; lateral
petiolar spines directed outwards and slightly curved backwards from bases. First gastral segment
concave medially, accomodating posterior face of petiole; anterodorsal margin of concavity rather
blunt, not elevated above dorsal face of segment.
Mandibles finely, mostly longitudinally, striate-rugose, with shallow pits towards masticatory
border. Sculpture of body consisting of more-or-less regularly spaced striae, that are mostly
longitudinal on head, distinctly anteriorly converging on pronotal dorsum and posteriorly
converging on mesonotal-propodeal dorsum; striae oblique on sides of mesosoma and
longitudinal on sides of gaster. Petiole and dorsum of gaster finely, microscopically, shagreened.
Entire body with numerous semierect to erect, off-white to silvery hairs, that do not exceed
greatest diameter of eye in length. Silvery to silvery-grey appressed pubescence fairly abundant,
partly concealing underlying sculpture, notably on propodeal dorsum, where it curves towards
midline. Gastral pubescence rather abundant, silvery at sides and along posterior margins of
segments; becoming distinctly silvery-grey on dorsal surface of first gastral segment.
Black; mandibular masticatory border, apices of antennal scapes, funiculi, femora and tarsi
medium to dark reddish brown, with funiculi and tarsi becoming lighter apically. Tibiae distinctly
lighter, except their proximal and distal ends, which are very narrowly dark, reddish brown.
Sexuals and immature stages unknown.
ETYMOLOGY
Name derived from the Latin word abditus, meaning hidden, concealed.
REMARKS
The known distribution of P. abdita is limited to Lelet Plateau on New Ireland. P. abdita is
very similar to P. dohrni, but can be distinquished by the characters listed under the latter. Of
these, the most important are the more flattened eyes and complete absence of an anterodorsal
process on the first gastral segment in abdita. Because of their similarity and adjoining
distributions, abdita could be considered a higher altitude population of dohrni. However, with
most of the distinguishing characters constant and no intermediate forms detected, I believe they
represent distinct species.
Polyrhachis (Aulacomyrma) dohrni Forel, 1901

Figures 2, 5, 8
Polyrhachis dohrni Forel, 1901: 34. Holotype queen. Type locality: BISMARCK ARCHIPELAGO, Credner
I. (F. Dahl), ?MNHU (holotype apparently lost).
Polyrhachis (Aulacomyrma) dohrni Forel; Emery, 1921: 17. Combination in P. (Aulacomyrma).
Polyrhachis (Aulacomyrma) dohrni Forel; Emery, 1925: 198.
NEOTYPE DESIGNATION
The holotype of this species should be lodged in the MNHU, together with other specimens
collected by Friedrich Dahl from the Bismarck Archipelago. Extensive searches by Dr Frank
Koch and the author failed locate it there or in any other collections examined and it is considered
lost. Forel (1901: 35) stated that the specimen was taken from the stomach of a bird Pachycephala
melanura on Credner I., 7.viii.1896. Because of considerable damage to the specimen (“Pro- und
Mesothorax stark defekt”), Forel was apparently uncertain about the caste and in the original
description listed the specimen as a worker with a question mark. However, further on he wrote:
“... drei deutliche Punktaugen (♀ ??)”. Because dohrni workers do not possess ocelli, the
specimen Forel described was undoubtedly a queen.
To establish the nomenclatural stability of the species, I have designated, in accordance with
Article 75.3.5. of the International Code of Zoological Nomenclature (Fourth Edition), a worker
specimen as the neotype of Polyrhachis dohrni Forel. The specimen was selected from a colony of
5 workers and 2 alate queens collected from a nest inside a dry hollow twig on a living tree in the
grounds of the Lowland Agricultural Experimental Station, Kerevat, East New Britain Province,
04º20’S, 152º02’E, c. 50m, rf. edge, 14.vii.1984, R.J. Kohout acc. 84.67. The locality is situated
only about 35km SW of Credner Islands, the type locality of P. dohrni. The neotype has been
deposited in ANIC.
WORKER

(92); SL 1.25-1.43 (1.40); SI 113-121 (114); PW 1.06-1.25 (1.18); MTL 1.15-1.37 (1.37) (9
measured).
Anterior clypeal margin arcuate with shallow notch medially. In profile clypeus virtually
straight with very weakly impressed basal margin that is laterally indicated by a hairline break in
cephalic sculpture. Frontal carinae sinuate, with laminate lobes. Frontal triangle indistinct. Sides
of head weakly convex. Eyes moderately convex, in full face view usually breaking lateral
cephalic outline. Mesosoma marginate along entire length; pronotal humeri armed with
broadly-based, acute teeth that are laterally continuous with posteriorly converging, uninterrupted,
pronotal margin. Promesonotal suture distinct; metanotal groove lacking dorsally, position
marked by shallow emargination of lateral margins. Mesonotal margins posteriorly converging;
propodeal margins subparallel, terminating in blunt processes that appear as minute teeth in
profile. Propodeal dorsum descending abruptly into weakly concave propodeal declivity. Petiole
scale-like, biconvex, with sharp dorsal margin, its apex higher than anterodorsal margin of first
gastral segment; lateral petiolar spines directed outwards and slightly curved backwards from
bases. First gastral segment concave medially, accommodating posterior face of petiole;
anterodorsal margin not elevated above dorsal face of segment in lateral view, however, when
viewed from behind, appearing somewhat medially produced.
Mandibles finely, mostly longitudinally, striate-rugose. Sculpture of body consisting of
more-or-less regularly spaced striae, mostly longitudinal on head, pronotal dorsum and sides of
gaster; V-shaped on mesonotal and propodeal dorsa, oblique on sides of mesosoma. Petiole and
dorsum of gaster finely, microscopically, shagreened.
Entire body with numerous semierect to erect, off-white to pale-yellow hairs, that do not
exceed greatest diameter of eye in length. Silvery appressed pubescence fairly abundant, but not
concealing underlying sculpture, except on propodeal dorsum, where it curves towards midline.
Gastral pubescence rather abundant, silvery at sides and along posterior borders of segments,
becoming distinctly golden on dorsal aspect of first gastral segment.
Black; mandibular masticatory border, apices of antennal scapes, funiculi, distal parts of
femora and tarsi medium to dark reddish brown, with funiculi and tarsi becoming lighter apically.
Tibiae distinctly lighter, except proximal and distal ends, that are very narrowly dark, reddish
brown.
QUEEN
Dimensions: TL c. 6.50-6.95; HL 1.53-1.56; HW 1.34-1.40; CI 87-90; SL 1.47-1.56; SI

109-112; PW 1.37-1.47; MTL 1.56-1.59 (4 measured).
Very much like worker with usual characters identifying full sexuality, including three ocelli
and complete thoracic structure with wings. Sculpture, pilosity and color as in worker except eyes
more prominent, clearly breaking cephalic outline in full face view. Lateral petiolar spines
distinctly shorter.
Male and immature stages unknown
MATERIAL EXAMINED
PAPUA NEW GUINEA: East New Britain Prov., Gazelle Pen., LAES Kerevat, 04°20’S, 152°02’E,
10-14.vii.1984 (RJK accs 84.27, 67, 71) (w,♀); ditto, 60m, 31.viii-1.ix.1955 (J.L. Gressitt) (w); ditto,
28.viii.1955 (J.L. Gressitt) ( ♀ ); ditto, 135m, 20-25.xi.1959 (T.C. Maa) (w); Malmaluan-Vunakanau,
8.v.1956 (J.L. Gressitt) (w); Baining Mts, St Paul Mission, 04º15’S, 151º47’E, 350m, 5.ix.1955 (J.L.
Gressitt) (w); ditto, 7.viii.1955 (J.L. Gressitt) (♀); ditto, Gaulim, 04º26’S, 152º05’E, 140m, 19-28.x.1962 (J.
Sedláček) (w). New Ireland Prov., Kableman, 8km ESE of Kavieng, 02º37’S, 150º51’E, c. 5m, 25.vii.1984
(RJK acc. 84.120) (w); East Coast, c. 3km S of Konos, 03º09’S, 151º43’E, c. 100m, 22.vii.1984 (RJK acc.
84.113) (w); West Coast, Gilingil Pltn, c. 04º28’S, 152º40’E, 2m, 6.vii.1956 (E.J. Ford Jr) (w); Ridge above
“Camp Bishop”, 15km up Kait R., 250-500m, 11.vi.1956 (J.L. Gressitt) (w).
REMARKS
P. dohrni is apparently limited to the Bismarck Archipelago: New Britain, neighboring

islands and low altitude localities along the coast of New Ireland. It is similar to P. abdita from
Lelet Plateau, but differs in several characters. The eyes of dohrni are moderately convex and in
full face view break the lateral outline of the head, if only marginally. The mesosomal dorsum is
more slender and the pronotal humeri armed with broad-based, more anteriorly directed, acute
teeth. The propodeal margins terminate posteriorly in blunt distinct processes that appear as
minute teeth in profile. The anterodorsal margin of the first gastral segment is not raised above its
dorsal face in lateral view but, viewed from behind, the margin appears medially produced. The
legs, notably the proximal halves of the femora, and apices of antennal scapes are distinctly lighter
reddish-yellow and the dorsal gastral pubescence is more golden. In contrast, P. abdita has
distinctly flattened eyes that do not reach the lateral outline of the head in full face view. The
mesosoma is somewhat wider posteriorly. The pronotal teeth are less produced, more angular and
the greatest pronotal width is at its midlength. The lateral propodeal margins are only narrowly
rounded posteriorly, not dentiform. The anterodorsal margin of the first gastral segment, when
viewed from behind, is smoothly rounded, without any indication of a median process. The
antennal scapes and femora are more-or-less uniformly medium to dark reddish-brown and the
relatively abundant pubescence, including that of the gaster, is silvery or silvery-gray.
Polyrhachis (Aulacomyrma) exarata Emery, 1887

Figures 3, 6, 9
Polyrhachis exarata Emery, 1887: 226. Holotype worker. Type locality: INDONESIA, Misool I.,
Waigamma (O. Beccari), MCSN (examined).
Polyrhachis (Aulacomyrma) exarata Emery; Emery, 1921: 17. Combination in P. (Aulacomyrma).
WORKER
Dimensions of holotype: TL c. 4.84; HL 1.18; HW 1.06; CI 90; SL 1.31; SI 123; PW 1.00 ;

MTL 1.21.
Clypeus with anterior margin shallowly truncate in middle; median longitudinal carina
distinct. Sides of head in front of eyes only weakly convex. Eyes convex, in full face view clearly
breaking lateral cephalic outline. Mesosoma marginate along entire length. Pronotal humeri with
rather strong, forward directed, broad-based teeth; propodeal margins terminating posteriorly in
short, rounded processes, that appear in profile as minute, weakly upturned teeth. Propodeal
dorsum separated from declivity by a distinct, slightly inward bowed, transverse ridge. Petiole
with dorsal margin sharp, somewhat sinuate in dorsal view; lateral spines acute, bent outwards and
backwards. First gastral segment only shallowly concave; dorso-medial margin of concavity
blunt, not raised above dorsal face of segment.
Dorsum of head and pronotum with shiny, regularly spaced, striae, longitudinal on pronotum
and posteriorly converging towards midline on mesonotal-propodeal dorsum. Petiole with both
faces finely shagreened. Sides of first gastral segment distinctly, longitudinally striate with
sculpture becoming more reticulate-puncate on dorsum.
Mandibles and sides of head with a few short semi-erect to erect hairs that are virtually absent
from dorsum of head, vertex, mesosoma and petiole, except for a few scattered hairs at base of
pronotal teeth. Sides and apex of gaster with numerous semi-erect to erect, medium length hairs
and with somewhat dilute, silvery, appressed pubescence lining posterior margin of first gastral
segment. Antennae and legs with yellow, appressed pubescence and numerous relatively short
erect hairs.
Black, with mandibular masticatory border and antennal scapes reddish brown. Funiculus a
shade lighter with extreme tip of last segment narrowly black. Legs mostly yellow, except distal
portion of femora, proximal ends of tibiae and first tarsal segment, that are medium to dark reddish
brown.
REMARKS
In general appearance, P. exarata resembles P. parva. They share rather strong, broad-based
pronotal teeth that are somewhat weakly emarginate at their bases; a distinct, weakly inward
bowed, transverse ridge separating the propodeal dorsum from the declivity and an almost
identical color scheme. However, they differ in several characters, including the longitudinal
striation of the pronotal dorsum. In P. exarata the striation is parallel with the pronotal margins,
while in parva the striae distinctly converge anteriorly. The propodeal margins in exarata
terminate in narrowly rounded processes that are weakly widened before rounding into the
posterior margin. In contrast, the propodeal margins in parva terminate in minute blunt angles.
The anterior face of first gastral segment is medially concave in exarata with the anterodorsal
margin of the concavity bluntly rounding onto the dorsum of the segment. In parva the concavity
is strongly and acutely marginate, with its anterodorsal margin produced dorso-medially and
raised above the dorsal face of the gaster.
Polyrhachis (Aulacomyrma) gentilis, new species

Figures 10, 13, 16
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Morobe Prov., Huon Pen., Mongi Watershed,
Zingzingu, 1200m, 9-10.iv.1955, E.O. Wilson #761 (worker). Type distribution: unique holotype
in MCZC.
WORKER
Dimensions of holotype: TL c. 4.84; HL 1.28; HW 1.09; CI 85; SL 1.31; SI 120; PW 0.87;

MTL 1.22.
Anterior clypeal margin with shallow notch medially. In profile clypeus almost straight
anteriorly, weakly convex posteriorly with shallow basal margin. Frontal carinae sinuate with
raised laminate lobes; frontal triangle indistinct. Sides of head almost straight in front of eyes,
converging anteriorly towards bases of mandibles; gently rounding behind eyes into preoccipital
margin. Eyes moderately convex, in full face view breaking lateral cephalic outline. Mesosoma
laterally marginate, virtually parallel-sided. Pronotal humeri armed with broad-based teeth that are
laterally continuous with pronotal margins; margins weakened posteriorly before reaching distinct
promesonotal suture. Metanotal groove absent. Propodeal dorsum terminating in rather distinct,
rounded prominences, their posterior margins continued medially but failing to meet, leaving a
very narrow gap through which propodeal dorsum descends into declivity; viewed from behind,
posterior margin forming an open inverted ‘V’; propodeal declivity concave in lateral view.
Petiole with dorsal margin acute, entire; lateral spines relatively short, curved backwards, sinuate
in dorsal view. Anterior face of first gastral segment concave, anterodorsal margin of concavity
rather acute, but not raised above dorsal face of segment.
Mandibles with rather distinct striae radiating from bases towards masticatory borders. Head
mostly longitudinally striate, striae on sides curving inwards from posterolateral corners and
continuing anteriorly towards mandibular bases. Striae on vertex mostly longitudinal, a few
irregular, mostly transversely bowed striae, medially along preoccipital margin. Pronotal dorsum
with striae converging anteriorly towards pronotal collar; striae on mesonotal-propodeal dorsum
longitudinal for most of their length, curving strongly inwards before reaching posterior margin.
Petiole with both faces and first gastral segment laterally and dorsally shagreened.
Head with numerous, erect and semi-erect, relatively short, hairs that do not exceed the
greatest diameter of the eyes in length; hairs distinctly less abundant on mesosomal dorsum and
first gastral segment, where they are posteriorly curved. Posterior and ventral gaster with more
abundant hairs, some as long as greatest diameter of eyes. Mostly medium length, pale yellow to
golden, diluted appressed pubescence on all body surfaces, most abundant and golden on gastral
dorsum, almost hiding underlying sculpture; pale yellow or silvery on metapleuron, propodeal
declivity and posterior margins of gastral segments.
Black, with legs dark to medium reddish brown. Tibiae, except proximal ends, apical
segments of tarsi and funiculi distinctly light yellowish brown.
ETYMOLOGY
Name derived from the Latin word gentilis, meaning gentle.

REMARKS
With its small size and parallel sided mesosoma, P. gentilis superficially resembles P.
wamuki. However in wamuki the pronotal margins are lacking and the lateral margins of the
pronotal spines curve directly onto the sides of the segment without continuing towards the
promesonotal suture. The sculpture of the pronotal dorsum also differs with the striae in gentilis
only weakly converging, while they are strongly inversely V-shaped and merge along the midline
in wamuki.
Polyrhachis (Aulacomyrma) lumi, new species

Figures 11, 14, 17
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA: West Sepik Prov., Torricelli Mts, Lumi, 03º28’S,
142º02’E, 400-550m, 4-13.viii.1984, R.J. Kohout acc. 84.273) (worker). PARATYPES: data as
for holotype (2 workers). Type distribution: holotype in ANIC, 1 paratype each in QMBA and
MCZC.
WORKER
Dimensions: TL c. 5.69-5.90 (5.69); HL 1.50-1.53 (1.53); HW 1.31-1.37(1.37); CI 89, 87-89

(89); SL 1.53-1.56 (1.56); SI 114-117 (114); PW 1.15-1.18 (1.18); MTL 1.43-1,47 (1.47) (3
measured).
Clypeus arcuate, anterior margin medially truncate; median carina blunt, but distinct; in
profile clypeus straight with base rather flat. Frontal triangle well defined. Frontal carinae sinuate
with raised laminate lobes; central area relatively wide with weak, blunt, longitudinal carina. Sides
of head in front of eyes weakly converging towards mandibular bases. Eyes moderately convex, in
full face view just reaching lateral cephalic outline. Mesosoma with lateral margins converging
posteriorly. Pronotal humeri armed with broad-based, acute teeth, their anterior and lateral
margins sharp and distinctly raised. Mesonotum and propodeum with hairline break in dorsal
sculpture and emarginations of lateral margins indicating position of metanotal groove. Propodeal
dorsum with lateral margins emarginate, terminating posteriorly in blunt, tooth-like prominences
that are distinctly raised, posterior margins continued for some distance, forming short, transverse,
medially unconnected ridges. Dorsal margin of petiole sharp, with some irregularities along
outline, lateral petiolar spines acute, posteriorly curved. Anterior face of first gastral segment
medially concave with anterodorsal margin of concavity blunt.
Sculpture of body consisting of more-or-less regular striae of various intensities, striae rather
flat and anteriorly converging on clypeus and vertex, where they are strongly impressed, meeting
along midline; striae mostly longitudinal on sides of head. Mesosomal dorsum with striae
regularly spaced, those along pronotal margin somewhat converging anteriorly towards pronotal
collar; striae converging posteriorly on propodeum and oblique on sides of mesosoma. Both faces
of petiole finely shagreened. Sides of first gastral segment with regular longitudinal striae, less
distinct on dorsum.
Erect to semi-erect, medium length hairs on mandibular masticatory border, median portion
of anterior clypeal margin, along frontal carinae, antennal scapes and vertex. In full face view,
hairs completely absent from sides of head between eyes and mandibular bases. Dorsum of
mesosoma and gaster with numerous erect hairs that are directed posteriorly on gastral dorsum and
distinctly longer towards apex of gaster. Petiole with a few, short, erect hairs along dorsal margin.
Mostly silvery, appressed pubescence scattered in various densities over body; most dense on
pronotal humeri and propodeal dorsum, almost obscuring underlying sculpture. Pubescence
off-white or yellowish on gaster, densest along gastral margins and curving towards midline.
Black; distal funicular segments and legs medium to dark reddish-brown.
ETYMOLOGY
Named after the type locality, Lumi village in the Torricelli Mountains, Papua New Guinea.
REMARKS
P. lumi is close to exarata, but has the head, in full face view, distinctly larger and the eyes
only reaching and not exceeding the lateral cephalic outline; the mesosoma in lateral view strongly
arched, terminating in rather distinct, posteriorly and dorsally directed, blunt teeth and the legs
dark brown. In contrast, the eyes in exarata clearly break the lateral cephalic outline, the
mesosomal dorsum is almost flat with the propodeum terminating in narrowly rounded processes
that are weakly widened laterally before rounding inwards and forming a transverse posterior
margin and the legs are mostly yellow.
Polyrhachis (Aulacomyrma) minima, new species

Figures 12, 15, 18
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Morobe Prov., Huon Pen, lower Busu R., 5.v.1955,
lowland rf., E.O. Wilson #954 (worker). Type distribution: unique holotype in MCZC.
WORKER

MTL 1.06.
Anterior clypeal margin arcuate, with very shallow notch medially; clypeus in profile
virtually straight anteriorly, with basal margin shallowly impressed. Frontal carinae strongly
sinuate with laminate lobes. Sides of head in front of eyes only weakly converging anteriorly.
Eyes convex, in full face view clearly breaking lateral cephalic outline. Mesosoma marginate,
margins of pronotal dorsum rather weak, but reaching promesonotal suture. Pronotal dorsum
virtually parallel sided; humeri armed with acute, laterally directed, broad-based teeth.
Promesonotal suture distinct; metanotal groove indicated by distinct incisions in lateral margins.
Propodeum with lateral margins weakly widened posteriorly, terminating in rounded prominences
and continued inwards, forming medially discontinued ridges that partly separate propodeal
dorsum from relatively high and concave declivity. Petiole transverse, dorsal margin acute and
weakly notched medially, strongly sinuate in dorsal view; lateral spines backwardly curved.
Anterior face of first gastral segment concave; anterodorsal margin of concavity acute but only
very weakly elevated above dorsal face of segment.
Mandibles finely striate. Head mostly longitudinally, somewhat irregularly, striate; striae on
sides of head more regular and weakly curved inwards. Sculpture of mesosomal dorsum
consisting of longitudinal striae, somewhat anteriorly converging on pronotal dorsum, posteriorly
converging on mesonotal-propodeal dorsum; oblique on sides. Propodeal declivity finely
shagreened; somewhat tranversely wrinkled medially, very smooth and glossy ventrally. Petiole
and dorsum of gaster shagreened, sides of first gastral segment longitudinally striate.
Relatively short, yellowish or off-white, erect or curved hairs on head and mesosoma, most
dense and longest on mandibles and clypeus, only a few short, variously curved hairs breaking the
outline of vertex and mesosoma. Dorsum of first gastral segment with only a few short hairs, more
abundant, longer and more golden towards apex. Somewhat untidy, relatively long, appressed or
decumbent, greyish or silvery pubescence present in various density on all body surfaces, densest
along preoccipital margin, pronotal humeri, propodeal dorsum, meso- and metapleura and coxae;
fairly dense on gastral dorsum, ranging from silvery to golden.
Black. Mandibles, antennae and legs light to medium yellow, only basal portion of scapes,
distal ends of femora and first tarsal segments a shade darker.
ETYMOLOGY
Name derived from the Greek word minys, meaning little or small, referring to its small size.
REMARKS
P. minima is one of the smallest members of the subgenus. It is somewhat similar to P. wardi
but, besides its smaller size (HL 1.09 in minima versus HL 1.15-1.34 in wardi), it differs in having
a more convex mesosoma, posteriorly weakened pronotal margins, propodeal margins that are not
produced into laterally directed teeth and a higher and more concave propodeal declivity.
Polyrhachis (Aulacomyrma) pallipes Donisthorpe, 1948

Figures 19, 22, 25
Polyrhachis (Aulacomyrma) pallipes Donisthorpe, 1948: 603. Holotype worker. Type locality: NEW
GUINEA, Finschhafen (E.S. Ross), CASC (examined).
WORKER

(90); SL 1.22-1.31 (1.31); SI 114-118 (117); PW 0.94-1.06 (1.03); MTL 1.12-1.22 (1.18) (6
measured).
Anterior clypeal margin arcuate, with shallow notch medially; in lateral view basal margin
only weakly impressed, indicated by hairline break in cephalic sculpture. Frontal triangle
indistinct. Frontal carinae with strongly sinuate, laminate lobes. Sides of head weakly converging
anteriorly; convex and more strongly converging posteriorly into preoccipital margin. Eyes
weakly convex, in full face view not or only just reaching lateral cephalic outline. Mesosoma
marginate along entire length. Pronotal dorsum armed with broad-based, anteriorly pointed teeth,
their lateral margins shallowly emarginate at bases. Promesonotal suture distinct; metanotal
groove lacking, position indicated by shallow emarginations in lateral margins. Propodeal dorsum
descending into declivity in medially uninterrupted curve; lateral margins terminating posteriorly
in short, blunt, processes resembling teeth in lateral view. Petiole with dorsal margin entire, acute;
lateral spines relatively long, curved backwards and upwards. First gastral segment concave
anteriorly, dorsal margin of concavity sharp and produced dorso-medially above dorsal surface of
segment.
Sculpture of head consisting of somewhat irregularly spaced, mostly longitudinal striae,
rather flat on clypeus and preoccipital margin, becoming stronger laterally. Mesosomal dorsum
with mostly longitudinal striae, anteriorly converging on pronotal dorsum, V-shaped on propodeal
dorsum. Anterior and posterior face of petiole shagreened. Gaster finely longitudinally striate;
concavity of first gastral segment smooth and shiny.
Medium length, mostly erect, yellow to golden hairs scattered over entire body surfaces,
longest on head and apically and ventrally on gaster, hairs shorter on mesosomal dorsum; only a
few, short hairs on dorsum of first gastral segment. Appressed, mostly silvery pubescence on head,
mesosoma and petiole; rich golden and dense pubescence on dorsum of first gastral segment,
largely obscuring underlying sculpture.
Black; mandibular masticatory margin, distal ends of scapes and basal segments of funiculi
light reddish brown; funicular segments becoming gradually lighter towards apex of antennae,
apical segments light yellow. Trochanters, femora, tibiae and tarsi light yellow, except distal ends
of femora and basal ends of tibiae and tarsi narrowly reddish-brown.
ADDITIONAL MATERIAL EXAMINED
PAPUA NEW GUINEA: Bubia, Markham V., 50m, 20.ix.1955 (J.L. Gressitt) (w); Huon Pen., lower
Busu R., 27.iv.1955, lowl. rf. (E.O. Wilson #705) (w); Lae, Busu R. area, 8.i.1968, rf. (B.B. Lowery) (w);
40km W of Madang, 05º13’S, 145º25’E, 140m, 30.i.1989, rf. (P.S. Ward #10088-14) (w). Northern Prov.,
Kokoda, 25.v.1972 (P.M. Room #364) (w); ditto, 400m, 22.iii.1956 (J.L. Gressitt) (w).
REMARKS
The head and mesosoma of the holotype have almost completely lost the silvery or pale
golden pubescence, that is still evident on the propodeal declivity and gastral dorsum. The
anterodorsal process of the first gastral segment is well developed, almost as prominent as in
porcata. Specimens from Kokoda differ in several aspects, notably in the color of the antennal
scapes, joints of the femora, tibiae and basal tarsal segments which are very dark brown.
Polyrhachis (Aulacomyrma) parva, new species

Figures 20, 23, 26
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Central Prov., nr Eilogo, 21.xii.1980, J.W. Ismai
(worker). PARATYPES: data as for holotype (4 workers). Type distribution: holotype in BMNH;
1 paratype each in, ANIC, CASC, MCZC and QMBA.
WORKER

(88); SL 1.12-1.28 (1.28); SI 114-125 (123); PW 0.87-1.03 (0.94); MTL 1.18, 1.06-1.22 (1.18) (6
measured).
Anterior clypeal margin with shallow notch medially; clypeus in profile straight anteriorly,
with a weakly impressed basal margin. Frontal carinae sinuate, with laminate lobes. Sides of head
in front of eyes gently converging towards bases of mandibles. Eyes convex, distinctly breaking
lateral cephalic outline. Mesosomal dorsum marginate along entire length, margins distinctly
converging posteriorly. Pronotal humeri armed with broad-based, forward projecting, triangular
teeth. Promesonotal suture distinct; metanotal groove lacking. Propodeal dorsum terminating
posteriorly in minute blunt angles, their posterior margins continuous and meeting medially,
forming transverse, somewhat medially weakened ridge separating propodeal dorsum from
declivity. Dorsal margin of petiole acute, entire, terminating laterally in acute, outwards and
backwards curved spines. First gastral segment concave anteriorly; anterodorsal margin of
concavity acute, produced dorso-medially and weakly raised above dorsal face of segment.
Mandibles finely longitudinally striate, becoming finely rugose-punctate towards masticatory
borders. Head, including clypeus, regularly, longitudinally striate. Mesosomal dorsum
longitudinally striate, interspaces wider than those on head; striae mostly oblique on lateral
mesosoma. Petiole with anterior and posterior faces shagreened, wrinkled on sides. First gastral
segment finely, longitudinally, striate-rugose laterally, shagreened dorsally.
Entire body with relatively short, erect or curved, silvery hairs. Appressed, mostly silvery
pubescence abundant everywhere, notably on dorsum of mesosoma, where it fills interspaces
between striae; dense on dorsum of gaster, partly obscuring underlying sculpture.
Black. Antennal scapes, joints of femora, tibiae and basal tarsal segments very dark brown,
almost black. Mandibular masticatory border reddish brown. Funiculi reddish-brown at bases,
each subsequent segment progressively lighter.
PAPUA NEW GUINEA: Gulf Prov., Ivimka camp, Lakekamu Basin, 7.7°S 146.8°E, 120m, lowland
wet forest, 23.xi.1996 (R.R. Snelling #96-357) (w); Central Prov., Karema, Brown R., lowland rf.,
8-11.iii.1955 (E.O. Wilson #534, 564) (w).
ETYMOLOGY
Derived from the Latin word parvus, meaning little, in reference to its small size.
REMARKS
Superficially similar to exarata, with the differences listed under the remarks section of that
species.
Polyrhachis (Aulacomyrma) planata, new species

Figures 21, 24, 27
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, East Sepik Prov., 8km SSW of Dagua, 03º29’S,
143º16’E, 800m, lowland rf., 12-14.vii.1980, P.S. Ward #4747-10 (worker). Type distribution:
unique holotype in ANIC.
WORKER

MTL 1.18.
Clypeus with anterior margin arcuate, entire. Sides of head weakly convex and gently
converging in front of eyes towards bases of mandibles. Eyes convex, clearly breaking cephalic
outline in full face view. Mesosomal dorsum distinctly marginate along entire length, pronotal and
mesonotal margins converging posteriorly. Pronotal dorsum rather flat; humeri armed with
broad-based, blunt teeth. Propodeal margins divergent, terminating posteriorly in blunt processes,
that appear as minute teeth in profile and form short, transverse, medially unconnected ridges, so
that propodeal dorsum descends into concave declivity in medially uninterrupted curve. Petiole
strongly transverse, dorsal margin entire; lateral spines very distinct, strongly curved backwards
and upwards. Anterior face of first gastral segment concave medially; anterodorsal border of
concavity sharply margined, but not distinctly raised above dorsal surface of segment.
Mandibles very finely longitudinally striate. Head and dorsum of mesosoma more-or-less
regularly striate; striae mostly longitudinal on head and dorsum of mesosoma, anteriorly
converging on clypeus and oblique on sides of mesosoma. First gastral segment shagreened
dorsally, longitudinally striate laterally, striae much finer than those on rest of body.
Relatively short, mostly erect, off-white or yellowish hairs, scattered over all dorsal body
surfaces; hairs longer on apical segments and venter of gaster, some almost as long as greatest
diameter of eye. Rather sparse, appressed, silvery pubescence scattered over dorsum of head and
mesosoma dense on pronotal humeri, propodeal dorsum, along lateral and posterior borders of
gastral segments and especially on metapleuron and propodeal declivity; midline of declivity
glabrous.
Black. Mandibular masticatory border widely, inner mandibular border narrowly, light
yellowish brown. Antennal scapes dark reddish brown; funiculi brown, segments becoming
progressively lighter towards apex, apical segments yellowish brown. Legs very light, yellowish
white (unique holotype may not be fully pigmented individual), only basal portion of trochanters,
distal ends of femora, proximal ends of tibiae and basal tarsal segments dark, reddish-brown.
ETYMOLOGY
Name derived from the Latin word planus, meaning flat, alluding to its virtually flat pronotal
dorsum.
REMARKS
In general appearance and color, P. planata is similar to P. pallipes. However, it differs from
that and other Aulacomyrma species by its virtually flat pronotal dorsum and distinctly transverse
petiolar node with its lateral spines strongly bent backwards.
Polyrhachis (Aulacomyrma) roberti, new species

Figures 28, 31, 34
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Tatupiti nr Tapini, 1200m, viii.1962, rf., R.W.
Taylor acc. 2293 (worker). PARATYPES: data as for holotype (2 workers). Type deposition:
holotype in ANIC; 1 paratype each in MCZC and QMBA.
WORKER

(88); SL 1.62-1.65 (1.65); SI 114-116 (115); PW 1.18-1.25 (1.25); MTL 1.56-1.65 (1.65) (3
measured).
Median portion of anterior clypeal margin projecting anteriorly as narrow, medially
emarginate lobe. Clypeus transversely convex, straight in profile with rather distinct basal margin
and frontal triangle. Frontal carinae sinuate with laminate lobes. Central area with short but
distinct longitudinal carina. Sides of head in front of eyes moderately convex, behind eyes
rounding into convex preoccipital margin. Mesosoma marginate along entire length, margins
broken at promesonotal suture and distinctly notched at metanotal groove. Pronotal dorsum
moderately convex; humeri armed with broad-based, acute teeth. Promesonotal suture distinct;
metanotal groove indicated dorsally by weakly incised line breaking sculpture. Propodeum armed
posteriorly with pair of strong, broad-based, triangular, dorsally curved teeth, their posterior
margins continued medially but failing to meet, leaving a small gap through which propodeal
dorsum meets declivity. Petiole strongly transverse, dorsal margin medially emarginate with pair
of distinct, acute, intercalary teeth flanking emargination; lateral spines long and straight with tips
weakly curved upwards; dorsal margin between intercalary teeth and lateral spines jagged.
Anterior face of first gastral segment very shallowly concave with dorso-medial margin blunt.
Mandibles with very fine shallow punctures. Clypeus very finely, microscopically
shagreened, contrasting with rest of head that is finely, but distinctly and regularly striate; striae
mostly longitudinal on sides of head, converging medially on vertex. Mesosoma longitudinally
striate, striae converging posteriorly on mesonotum and oblique on sides. Both faces of petiole
microscopically shagreened. First gastral segment shagreened, a few longitudinal striae on sides.
Dorsum of head with very few erect hairs on anterior clypeal margin, antennal scapes, frontal
carinae and area between eyes and posterolateral corners. Mesosomal dorsum with a few erect or
semierect, medium length hairs on pronotal humeri, mesonotal and propodeal dorsa and dorsal
surfaces of propodeal teeth. Petiole with several short hairs arising from posterior face just below
dorsal margin. Dorsum and apex of gaster with more numerous, distinctly longer, golden hairs.
Silvery, rather diluted, appressed pubescence on pronotal humeri, dorsum and sides of
propodeum, coxae and petiole, more dense and distinctly golden on dorsum of first gastral
segment.
Black and glossy. Mandibles and eyes distinctly light reddish-brown. Appendages very dark
brown, almost black, inner surfaces of front femora, front and middle tibiae and distal segments of
tarsi medium to light reddish brown. Apex of gaster reddish brown.
ETYMOLOGY
Named in honor of my former mentor, Dr Robert W. Taylor, who introduced me to and

guided my first steps in my studies on Polyrhachis ants.
REMARKS
This species is characterized by well developed pronotal and propodeal teeth and notably by
its remarkable petiole, which is strongly transverse and furnished with a pair of acute dorsal teeth
with a medial emargination between them and relatively long, lateral spines. It stands near P.
simulans with both sharing similar sculpture of the head and mesosoma, including the absence of
any striae on their mandibles and clypeus. However, they differ in a number of characters, of
which the most distinct are discussed under simulans.
Polyrhachis (Aulacomyrma) sculpta Emery, 1887

Figures 29, 32, 35
Polyrhachis sculpta Emery, 1887: 226. Holotype worker. Type locality: NEW GUINEA, Sorong (L.M.
D’Albertis), MCSN (examined).
Polyrhachis (Campomyrma) sculpta Emery; Emery, 1925: 180. Combination in P. (Campomyrma).
WORKER

MTL 1.34.
Clypeus arcuate, medially shallowly emarginate with distinct longitudinal median carina; in
profile clypeus virtually straight with basal margin weakly impressed medially. Frontal triangle
indistinct. Frontal carinae strongly sinuate with raised laminate lobes. Eyes relatively large,
convex, in full face view clearly breaking lateral cephalic outline. Sides of head only weakly
converging anteriorly, convex and converging behind eyes. Pronotal and mesonotal dorsa
marginate, lateral margins of propodeal dorsum ill-defined. Pronotal humeri armed with triangular
teeth, lateral margins shallowly emarginate at bases. Promesonotal suture distinct; metanotal
groove lacking, position indicated by interruption of lateral mesosomal margin. Propodeal dorsum
descending into virtually vertical declivity in rather abrupt, medially unbroken curve; lateral
propodeal margins terminating in small, blunt, tubercles. Petiole with sharp dorsal margin,
shallowly emarginated medially; lateral teeth short, very weakly curved backwards. First gastral
segment concave anteriorly, anterodorsal margin of concavity elevated above dorsal surface of
segment.
Mandibles finely, longitudinally striate. Sides and front of head regularly, longitudinally
striate, striae on clypeus converging anteriorly and meeting along midline; striae on vertex
distinctly transversely bowed, almost semicircular. Mesosoma with striae converging anteriorly
on pronotal dorsum, somewhat V-shaped on mesonotal and propodeal dorsa and oblique on sides.
Petiole and dorsum of gastral segments shagreened.
Silvery or off-white hairs present on all body surfaces and appendages. Numerous, semi-erect
to erect, very short to medium length hairs on front and sides of head in front of eyes; hairs very
short and strongly anteriorly inclined on vertex. Antennal scapes with numerous hairs along
leading edges and at distal ends. Hairs on pronotal dorsum very uneven in length and direction,
ranging from short to medium, erect to semi-erect or undulated; hairs mostly posteriorly directed
on mesonotal and propodeal dorsa. Petiole with a few hairs along dorsal margin. Gaster with
mostly short, posteriorly directed hairs on dorsum of first gastral segment, increasing in length and
density towards gastral apex where they are relatively long and more golden. Silvery appressed
pubescence scattered over body; most dense on dorsum and sides of propodeum, where it almost
completely hides underlying sculpture, and along posterior margins of gastral segments.
Black, with antennal scapes, basal segments of funiculi and femora very dark reddish brown.
Apical segments of funiculi, tibiae and tarsi light to very light yellowish brown.
REMARKS
P. sculpta is characterized by the posteriorly less distinct lateral margins of the mesosoma,
leaving the propodeum only weakly margined laterally and completely immarginate posteriorly.
The sculpture of the vertex is transversely bowed in contrast to the longitudinal striation of the rest
of the head. With its rather untidy silvery pubescence sculpta superficially resembles P. variegata,
but they differ in many aspects, especially the eyes, that are convex in sculpta, and distinctly flat in
variegata. The propodeal declivity is virtually vertical in sculpta, while it is distinctly oblique in
variegata. The anterodorsal margin of first gastral segment is elevated above the dorsal surface of
the segment in sculpta, while it is only weakly concave in variegata, with the dorso-medial
margin blunt.
Polyrhachis (Aulacomyrma) simulans, new species

Figures 30, 33, 36
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Morobe Prov., Huon Penins., lower Busu Riv.,
5.v.1955, lowland rf. (E.O. Wilson #950) (worker). Type distribution: unique holotype in MCZC.
WORKER

MTL 1.37.
Anterior clypeal margin arcuate, entire; in profile clypeus weakly convex with rather flat
basal margin. Frontal triangle indicated by change in cephalic sculpture. Frontal carinae sinuate,
with raised laminate lobes. Sides of head gently convex in front of eyes. Eyes only moderately
convex, not reaching lateral cephalic outline in full face view. Mesosoma laterally marginate
along entire length. Pronotal humeri armed with acute teeth, their margins raised and continuous
with pronotal margins. Promesonotal suture distinct; metanotal groove indicated by shallow
incisions in lateral margins. Propodeal dorsum with lateral margins terminating posteriorly in
narowly rounded angles with weakly raised margins, appearing as small teeth in lateral view;
medially propodeal dorsum rounding into declivity in an uninterrupted curve. Dorsal petiolar
margin acute, with medial emargination flanked by pair of intercalary teeth; lateral margin
somewhat jagged and terminates in outwards and weakly backwards directed, slender spines.
Anterior face of first gastral segment concave, dorso-medial margin of concavity rather blunt.
Mandibles, clypeus and frontal triangle rather smooth, very finely, superficially shagreened
with scattered minute punctures, strongly contrasting with rest of head, that is longitudinally
striate with striae weakly curved on sides; striae less regular on vertex and somewhat curved along
midline. Striation on pronotal dorsum distinctly converging anteriorly and medially; striae on
mesonotal- propodeal dorsum converging towards propodeal declivity and oblique on sides.
Petiole with both faces finely shagreened. First gastral segment longitudinally striate, striae on
dorsum distinctly finer and converging medially towards anterior margin.
Head, body and appendages with numerous erect to semi-erect, whitish or silvery hairs; no
hairs on mandibles and clypeus, except a few along anterior margin, and sides of head between
eyes and mandibular bases. Gaster with numerous, short, backward-curved hairs, those closer to
apex distinctly longer. Mostly appressed, greyish or silvery pubescence on all body surfaces, most
diluted on head and pronotum, with denser patches on meso- and metapleura, lateral borders of
propodeal declivity and propodeal dorsum, where it curves towards midline. Pubescence
relatively abundant on gastral dorsum, but not hiding underlying sculpture.
Black or very dark brown, with funiculi and tibiae of hind legs a shade lighter. Mandibular
masticatory borders, tibiae, except their basal margin and apical tarsal segments light, almost
yellow with reddish tint.
ETYMOLOGY
Name derived from the Latin word simulo, meaning imitate or copy, in referrence to its
similarity to P. roberti.
REMARKS
P. simulans closely resembles P. roberti and they share similar sculpture of the head and
mesosoma, including the absence of striae on their mandibles and clypeus. However, they differ in
a number of characters, of which the most distinct is the anterior clypeal margin which is entire in
simulans, while in roberti the median portion of the margin projects as a narrow, medially
emarginate lobe. In full face view the eyes do not reach the cephalic outline in simulans, while
they clearly exceed it in roberti. In simulans the lateral margins of the propodeal dorsum terminate
in blunt rounded angles, that do not project posteriorly. In contrast, the margins in roberti
terminate in a pair of strong, broad-based, triangular, dorsally curved teeth, that project well
beyond the posterior border of the propodeum. The petiole in simulans is about as wide as high,
while in roberti it is clearly transverse, distinctly wider than high. Most of the body surfaces in
simulans have abundant, erect hairs, while only a few hairs break the outline of the head and
mesosoma in roberti.
Polyrhachis (Aulacomyrma) sulcifera, new species

Figures 37, 40, 43
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Wum, Upper Jimmi Valley, 05º25’S, 144º23’E,
840m, 17.vii.1955, J.L. Gressitt (worker). Type distribution: unique holotype in MCZC.
WORKER

MTL 1.28.
Anterior clypeal margin arcuate, entire; in profile clypeus very weakly convex with rather
shallow basal margin indicated by hairline break in cephalic sculpture. Sides of head in front of
eyes only weakly convex. Eyes strongly convex, in full face view clearly breaking cephalic
outline. Mesosomal dorsum laterally marginate along entire length. Pronotal humeri armed with
strong, broad-based, acute teeth, with distinctly raised margins. Promesonotal suture distinct;
metanotal groove lacking, position indicated laterally by shallow emargination in the mesosomal
margin. Propodeal dorsum with laterally widened margin, forming distinct rounded prominences,
posteriorly continued as transverse, inwardly bowed ridges that almost completely separate
dorsum from declivity, except for very narrow medial gap. Petiole distinctly transverse, with
acute, medially jagged dorsal margin and slender, acute, strongly upturned lateral spines. Anterior
face of first gastral segment concave with anterodorsal margin acute, but not distinctly raised
above dorsal face of segment.
Mandibles finely longitudinally striate. Sculpture of head and body consisting of widely
spaced, mostly regular, longitudinal striae, giving dorsal surfaces a “ploughed” appearance. Sides
of mesosoma with mostly oblique striae, propodeal declivity rather smooth. Petiole shagreened.
Gaster with fine, mostly longitudinal striation, striae on dorsum converging towards anterodorsal
margin.
Rather short, mostly erect, silvery and yellowish hairs present on all body surfaces, except
inferior edges of antennal scapes, dorsal petiolar margin and extensor surfaces of femora and
tibiae. Hairs on gastral dorsum more golden and posteriorly curved, those on apex distinctly
longer. Appressed pubescence generally very sparse or absent, somewhat more abundant and
yellowish on pronotal and propodeal dorsa, notably on humeral teeth and along posterior
propodeal margin, silvery on metapleuron and lateral borders of propodeal declivity. Gastral
pubescence more abundant and distinctly golden.
Black, with striae and interspaces rather shiny. Antennal scapes, joints of femora and tibiae
and proximal portion of basal tarsal segments black or very dark brown. Funiculi and rest of legs
light or medium yellow.
ETYMOLOGY
Derived from the Latin word sulcus, meaning furrow or goove in reference to the “ploughed”
appearance of the dorsal sculpture.
REMARKS
A remarkable species easily recognized by its cephalic and mesosomal striation, that has only
about 13 widely spaced striae present across the pronotal dorsum.
Polyrhachis (Aulacomyrma) umboi, new species

Figures 38, 41, 44
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Umboi I., 1km N of Awelkom, 600m,

21-28.ii.1967, G.A. Samuelson (worker). Type distribution: unique holotype in MCZC.
WORKER

MTL 1.28.
Anterior clypeal margin with distinct notch medially; in profile clypeus only very weakly
convex with shallow impression just behind anterior margin and moderately impressed basal
margin, indicated laterally by break in cephalic sculpture. Frontal triangle indistinct. Frontal
carinae sinuate with moderately raised laminate lobes. Head widest just behind eyes, sides
converging anteriorly towards mandibular bases. Eyes convex, in full face view clearly breaking
cephalic outline. Mesosoma marginate; pronotal and mesonotal dorsa with lateral margins
converging posteriorly, propodeal margins parallel. Pronotal humeri armed with angle-like teeth
with moderately raised margins, bases weakly emarginate laterally. Promesonotal suture distinct;
metanotal groove lacking dorsally, position indicated by slight notch and change in direction of
lateral margins. Propodeal dorsum terminating in minute blunt angles, posterior margins
continued inwards for only a short distance with dorsum descending into declivity in medially
uninterrupted curve. Petiole with dorsal margin acute, weakly jagged; lateral spines curved
slightly backwards and upwards. Anterior face of first gastral segment concave, anterodorsal
margin distinct, but not acute or raised above dorsal face of segment.
Sculpture of head and mesosoma consisting of more-or-less regularly spaced striae, mostly
longitudinal on sides of head, slightly converging on clypeus; striae on vertex curving from
posterolateral corners towards central area; a few short, transverse striae along medial preoccipital
margin. Pronotal dorsum with striae weakly converging anteriorly, striae on mesonotal-propodeal
dorsum distinctly converging posteriorly. Petiole shagreened and somewhat wrinkled. Sides of
first gastral segment longitudinally striate, dorsum shagreened.
Head with numerous, erect, rather short, pale yellow hairs not longer than greatest diameter of
eye, hairs much less abundant on mesosoma, only few breaking mesosomal outline in lateral view.
Gaster with a few short hairs on dorsum of first segment; hairs denser and distinctly longer on
ventral surfaces and towards gastral apex. White or silvery, rather diluted, appressed pubescence
on head; denser patches on pronotal humeri, propodeal dorsum, metapleuron, coxae and petiole.
Propodeal declivity bordered dorsally and laterally by longer and semierect silvery pubescence.
Gastral pubescence rather diluted and silvery along posterior borders of tergites; more abundant
and pale golden on dorsum of first gastral segment.
Black, tibiae and apical tarsal segments medium to light reddish-brown; funiculi progressively
lighter towards apices, terminal segments light reddish-brown.
ETYMOLOGY
Named after the locality Umboi Island, situated just off the Huon Peninsula along the
northeastern coast of Papua New Guinea.
REMARKS
P. umboi is somewhat similar to P. pallipes, but lacks an acute and raised anterodorsal margin
of first gastral segment and having less acute pronotal teeth and generally darker appendages.
Polyrhachis (Aulacomyrma) variegata, new species

Figures 39, 42, 45
TYPE MATERIAL
HOLOTYPE: INDONESIA, IRIAN JAYA, Guega, W of Swart Valley, 03º36’S, 138º25’E,

1200m, 15.xi.1958, J.L. Gressitt (worker). Type distribution: unique holotype in MCZC.
WORKER

MTL 1.40.
Anterior clypeal margin distinctly medially emarginate; in profile clypeus very weakly
convex, almost straight with rather flat but distinct basal margin. Frontal carinae sinuate, rather
short, only moderately raised; frontal triangle indistinct. Sides of head convex, distinctly
converging anteriorly towards mandibular bases. Eyes distinctly flat, not reaching cephalic outline
in full face view. Mesosoma marginate with propodeal margins somewhat weakly defined.
Pronotal humeri armed with laterally directed, broad-based teeth, margins raised and laterally
continuous with pronotal margins. Promesonotal suture distinct; metanotal groove lacking.
Propodeum terminating in small but distinct tuberculae. Propodeal dorsum descending into
oblique declivity in uninterrupted curve. Petiole with acute, finely jagged dorsal margin and short,
weakly backwards and upwards curved lateral spines; in dorsal view, dorsal petiolar margin
weakly anteriorly bowed. Anterior face of first gastral segment very weakly concave with
anterodorsal margin blunt, not raised above dorsum of segment.
Mandibles finely longitudinally striate at bases, becoming more reticulate with several shallow
pits towards masticatory borders. Clypeus with very fine, weakly V-shaped striae at base, merging
into irregular rugae anteriorly. Vertex and sides of head with fine, rather flat, mostly longitudinal
striae. Pronotal dorsum with striae diverging, more-or-less parallel with lateral pronotal margins;
striae posteriorly convering on mesonotal-propodeal dorsum, mostly oblique on sides of
mesosoma, propodeal declivity only finely shagreened. Petiole with both faces finely shagreened.
Gaster with sides longitudinally striate, striae distinctly finer dorsally, first gastral tergite finely
reticulate anteromedially.
A few, rather short, erect hairs along outer mandibular edges, anterior clypeal margin, leading
edge of antennal scapes, frontal carinae and outer surfaces of hind tibiae; a solitary hair arising
from vertex. Hairs completely absent from dorsum of mesosoma, petiole and first gastral segment;
only a few hairs, shorter than greatest diameter of eye, concentrated on apex and venter of gaster.
Silvery, appressed and semierect, somewhat untidy pubescence in various densities on vertex,
pronotal humeri, propodeal dorsum, metapleura, coxae and anterior face of petiole; pubescence
virtually absent from front of head; gaster with pubescence more abundant dorsally but not
obscuring underlying sculpture.
Black, mandibular masticatory border dark red. Funiculi apically and legs generally dark to
medium reddish-brown, tibiae and apical tarsal segments a shade lighter.
ETYMOLOGY
Named for its somewhat different appearance from that of other members of the subgenus
(Latin varius, variegatus = different, different sort).
REMARKS
A rather distinct species easily separated from most others by the posteriorly wide head and
distinctly flat eyes, that are situtated well away from the lateral margins of the head. The rather flat
and fine sculpture, untidy silvery pubescence and notably the obliquely descending propodeal
declivity are also quite distinctive.
Polyrhachis (Aulacomyrma) wardi, new species

Figures 46, 49, 52
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, East Sepik Prov., 3km S of Wewak, 03º37’S,
143º37’E, 400m, 15.ii.1989, P.S. Ward #10199 (worker). PARATYPES: data as for holotype (10
workers, 3 alate ♀, with some of the specimens not entirely hardened and fully pigmented). Type
distribution: holotype, 1 nidoparatype worker and 1 nidoparatype queen in ANIC; 2 nidoparatype
workers each in BMNH, CASC, MCZC and QMBA.
WORKER

(85); SL 1.31, 1.22-1.37; SI 114-126 (120); PW 0.94-1.06 (0.97); MTL 1.12-1.25 (1.18) (11
measured).
Anterior clypeal margin shallowly emarginate medially; clypeus in profile weakly convex with
shallow basal margin. Frontal carinae sinuate, with laminate lobes. Sides of head weakly convex,
anteriorly converging. Eyes strongly convex, in full face view clearly breaking cephalic outline.
Mesosomal dorsum marginate along entire length; weakly convex in profile. Pronotal humeri
virtually right-angled, with margins raised. Promesonotal suture distinct. Metanotal groove
lacking dorsally, position indicated by emarginations of lateral margins. Propodeal dorsum with
lateral margins diverging and extended into rather short, blunt, laterally directed teeth, posterior
margins continued as transverse, somewhat incurved ridge, dividing propodeal dorsum from
declivity. Petiole with dorsal margin acute, entire, terminating laterally in distinct, backwardly
curved spines with weakly upturned tips. First gastral segment concave anteriorly; concavity
sharply margined dorsally, but not elevated above dorsal face of segment.
Mandibles irregularly longitudinally striate. Head mostly longitudinally, somewhat irregularly
striate-rugose; sculpture more regular on sides; striae originating from below posterolateral
corners bowed inwards and then outwards, terminating at bases of mandibles. Striation of
mesosomal dorsum longitudinal, anteriorly diverging on pronotum, posteriorly converging on
mesonotal-propodeal dorsum, mostly oblique on sides of mesosoma. Petiole with both faces
shagreened, some rugosity evident towards sides. First gastral segment mostly shagreened
dorsally, sides finely, longitudinally striate.
Rather short, mostly erect, silvery and yellowish hairs on all body surfaces and appendages.
Appressed, silvery or greyish pubescence very sparse on head, more abundant on dorsum of
mesosoma, curving towards midline, not concealing underlying sculpture; patches of denser
pubescence on meso- and metapleuron, coxae and petiole; pubescence fairly abundant on first
gastral segment, silvery on sides and richly golden on dorsum, almost hiding underlying sculpture.
Black, including coxae and most of antennal scapes. Apices of scapes, funiculi, mandibular
masticatory border, distal portion of femora, proximal ends of tibiae and basal tarsal segments
very dark to dark reddish brown. Rest of legs light yellow, almost white in some specimens.
QUEEN
Dimensions: TL c. 5.84-6.10; HL 1.43-1.47; HW 1.25-1.26; CI 85-87; SL 1.40-1.43; SI

112-113; PW 1.25-1.28; MTL 1.40 (3 measured).
Similar to worker with usual differences associated with caste. Sculpture similar, pattern of
striation following fully developed thoracic structure. Pronotal humeri less acute, propodeal teeth
less distinct and petiolar spines shorter. Color of body and appendages identical to worker.
Male and immature stages (larvae and a pupa) present in ANIC.
INDONESIA, IRIAN JAYA (New Guinea: Neth. on data label), Waris, S of Hollandia, 450-500m,
1-7.viii.1959, sweeping, T.C. Maa (w). PAPUA NEW GUINEA: East Sepik Prov., Amok, N of Maprik,
03º35’S, 142º57’E, 165m, 6.i.1960 (T.C. Maa) (w).
ETYMOLOGY
Named in honor of the collector of the type series, Dr Phil S. Ward.
REMARKS
P. wardi is a very distinct species separated from other Aulacomyrma by the distinctly
laterally widened margins of the propodeal dorsum that extend into short, bluntly terminated,
laterally directed teeth.
Polyrhachis (Aulacomyrma) wilsoni, new species

Figures 47, 50, 53
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Huon Pen., lower Busu R., 5.v.1955, lowland rf.,
E.O. Wilson #950 (worker). PARATYPES: data as for holotype, but 3.v.1955, E.O. Wilson #895,
969 (3 workers) and 9.v.1955, E.O. Wilson #984) (worker); Bulolo, 3000ft, 19.xii.1970, rf., B.B.
Lowery (4 workers); W. Bulolo, 2500ft, 22.xii.1970, rf., B.B. Lowery (3 workers). Type
deposition: holotype and 2 paratypes in MCZC; 2 paratypes each in QMBA, ANIC, BMNH
and CASC.
WORKER

(87); SL 1.22-1.31 (1.28); SI 114-122 (117); PW 0.94-1.09 (1.00); MTL 1.09-1.22 (1.22)
(12 measured).
Anterior margin of clypeus arcuate with very shallow notch medially; in profile clypeus
weakly convex. Sides of head in front of eyes gently converging anteriorly. Eyes convex, clearly
breaking cephalic outline in full face view. Frontal carinae distinctly raised, sinuate; frontal area
relatively wide, with short median carina that merges posteriorly into cephalic sculpture.
Mesosoma marginate along entire length. Pronotal humeri armed with broad-based, triangular,
blunt, teeth. Mesonotal and propodeal dorsa fused, with lateral margins deeply emarginate,
terminating in very long, outwardly curved spines, their posterior margins continued as tranverse,
deeply inwardly bowed ridges, that meet medially and separate propodeal dorsum from declivity.
Petiole with sharp, dorsal margin, shallowly indented medially; lateral spines acute and curved
backwards, their tips slightly upturned. Anterior face of first gastral segment concave,
accommodating posterior face of petiole; anterodorsal margin of concavity distinct medially, but
not raised above dorsal face of segment.
Sculpture of head, mesosoma and sides of gaster consisting of regularly spaced striae, mostly
longitudinal on dorsum of head and sides of gaster, anteriorly converging on pronotal dorsum,
converging posteriorly and joining along midline on mesonotal and propodeal dorsa. Propodeal
declivity, petiole and dorsum of first gastral segment shagreened.
Abundant, yellowish, semi-erect to erect hairs present on all body surfaces and appendages,
longest hairs almost as long as greatest diameter of eye. Hairs absent from inferior edges of scapes
and dorsal surfaces of femora. Body with mostly off-white or yellowish, appressed pubescence,
denser and silvery on metapleuron and reddish on gastral dorsum.
Black, with antennal scapes, distal portions of femora, proximal and distal ends of tibiae and
basal segments of tarsi dark to medium brown; funiculi and rest of legs distinctly lighter, medium
to light yellowish brown or reddish-yellow.
PAPUA NEW GUINEA: Adelbert Mts, Wanuma, 800-1000m, 26.x.1958 (J.L. Gressitt) (w); Wau
Gorge, 3000’, rf., 30.xii.1970 (B.B. Lowery) (w); ditto, 7.i.1971 (B.B. Lowery) (w).
ETYMOLOGY
Named in honor of Prof. Edward O. Wilson of Harvard University, who collected many new
Aulacomyrma species during his extensive field work throughout Papua New Guinea.
REMARKS
A remarkable species, easily distinguished from all other known members of the subgenus by
the very elongate, strongly divergent, propodeal spines.
.
Polyrhachis porcata species-group
This group is characterized by the following: mesosomal margins poorly defined or

completely absent. Pronotal humeri armed with broad-based teeth or short spines; pronotal lateral
margins absent or extending posteriorly for only a short distance before merging with outermost
dorsal striae and curving onto sides of pronotum without reaching promesonotal suture.
Mesonotal-propodeal dorsum usually with only one or other marginate, or completely
immarginate. Petiole scale-like with anterior and posterior faces convex and dorsal margin acute
or blunt. Anterior face of first gastral segment distinctly concave in all species, except decora and
its allies, where it is truncate or even convex. Within the group the species tend to polarise into
four, rather intergrading, complexes, centering on excellens, porcata, geometrica and orokana.
Species in the excellens complex are more similar to the dohrni-group species, featuring short and
weak pronotal margins and only a weakly convex mesosomal dorsum. The sculpture of mesosoma
consists of mostly longitudinal, relatively widely spaced striae of a “ploughed” appearance, as in
excellens or maai. Species of the porcata complex tend to have the mesosomal dorsum, distinctly
transversely convex, the pronotal dorsum completely immarginate, and the mesonotal-propodeal
dorsum with one or the other marginate, as in porcata or monticola. The sculpture of the
mesosomal dorsum consists mostly of longitudinal striae curved into various V- or U-shaped
patterns, notably on the pronotum. The geometrica complex include species with a virtually
immarginate mesosoma, both laterally and posteriorly, although in some species closely related to
geometrica the outermost striae of the mesonotal dorsum form an ill-defined lateral margin. The
pronotal humeri are armed with distinct, more-or-less isolated teeth or spines. The anterior face of
the first gastral segment is only shallowly concave, as in geometrica or roomi, or simply convex as
in decora or kokoda. The sculpture of the mesosomal dorsum is intricate with the striae forming
transverse, V-, U- or semicircle-shaped patterns, giving the surface a characteristic “geometrical”
appearance.
The two species comprising the orokana complex are in most aspects very similar to species
of the geometrica complex. However, they differ by the presence of a very prominent transverse
ridge on propodeum, separating the propodeal dorsum from the declivity and by the complete lack
of lateral mesosomal margins.
Polyrhachis (Aulacomyrma) annulata, new species

Figures 48, 51, 54
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Mt Tafa, 8km SE of Mondo, 08º38’S, 147º07’E,

8500ft, ii.1934, L.E. Cheesman (B.M.1934-321) (worker). Type distribution: unique holotype in
BMNH.
WORKER
Dimensions of holotype: TL 5.54; HL 1.47; HW 1.47; CI 100; SL 1.43; SI 97; PW 1.12; MTL
1.40.
Anterior clypeal margin arcuate, emarginate medially. In profile clypeus moderately convex
with basal margin weakly impressed, indicated by hairline break in cephalic sculpture. Frontal
carinae sinuate with moderately raised laminate lobes. Sides of head in front of eyes converging
anteriorly; rounding behind eyes into relatively shallow preoccipital margin. Eyes moderately
convex, just reaching lateral cephalic outline in full face view. Mesosoma only partially
marginate, somewhat antero-posteriorly compressed; pronotal humeri armed with broad-based
acute spines with lateral margins continued posteriorly for a short distance before merging with
outermost dorsal striae and curving onto sides. Promesonotal suture distinct; metanotal groove
lacking. Mesonotal dorsum with distinct lateral margins formed by outermost lateral striae.
Propodeum immarginate laterally; propodeal dorsum separated from relatively high, shallowly
concave declivity by posterior-most stria that forms a weakly defined posterior margin. Petiole
strongly transverse, dorsal margin acute, medially emarginate, strongly sinuate in dorsal view;
lateral spines rather short, weakly bent backwards. First gastral segment concave with the
anterodorsal margin of concavity rather blunt.
Mandibles finely longitudinally striate-rugose. Head mostly regularly striate, striae V-shaped
on clypeus, longitudinal along sides of head and somewhat converging anteriorly from vertex
towards frontal carinae. Mesosomal dorsum striate; pronotal striae longitudinal, anteriorly
converging on pronotum, oblique on sides; mesonotal-propodeal dorsum with striae strongly
bowed, more-or-less semicircular, oblique on sides. Petiole shagreened, becoming more
transversely reticulate towards base. Gaster uniformly, finely reticulate.
Mandibles with a few, anteriorly curved hairs along outer and masticatory borders. Antennal
scape with a few erect hairs along leading edge and at apex. A few scattered, rather short,
variously directed hairs breaking outline of head and mesosoma. Gaster with some longer erect
hairs, bordering segments. Appressed pubescence virtually absent from dorsum of head and
mesosoma; pale golden pubescence on gastral dorsum, almost obscuring underlying sculpture.
Black. Antennae, femora and first tarsal segments very dark brown, rest of legs medium to
light reddish brown; mandibles with anterior half reddish brown; apex of last funicular segment
distinctly yellow.
ETYMOLOGY
Name derived from the Latin word annulatus, meaning ringed or circular, with reference to
the form of the sculpture of the mesonotal-propodeal dorsum.
REMARKS
With the antero-posteriorly compressed mesosoma, dorsal petiolar margin medially

emarginate and striae of the mesonotal-propodeal dorsum more-or-less semicircular, annulata is
somewhat similar to breviata. However, both species are easily separated by their size (HL 1.47 in
annulata versus 1.15 in breviata) and shape of the petiole which, in annulata, is strongly
transverse, while in breviata the petiole is almost as high as wide. The sculpture of the pronotal
dorsum in annulata consists of longitudinal, anteriorly converging striae, while in breviata the
striae are tranverse and strongly anteriorly bowed medially.
Polyrhachis (Aulacomyrma) bedeloweryi, new species

Figures 55, 58, 61
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, E. Highlands Prov., Mingende, 05º58’S, 144º53’E,

5200ft, 11.i.1968, B.B. Lowery #NG507 (worker). PARATYPES: data as for holotype (3
workers). Type deposition: holotype in ANIC; 1 paratype each in BMNH, MCZC and QMBA.
WORKER

(91); SL 1.53; SI 122-125 (1.22); PW 0.94-0.97 (0.97); MTL 1.56-1.59 (1.59) (4 measured).
Anterior clypeal margin arcuate with very shallow notch medially; basal margin flat, indicated
only by a faint hairline; clypeus with shallow medial depression; very weakly convex in profile.
Frontal carinae sinuate with laminate lobes; central area with a weak, short, longitudinal carina.
Sides of head in front of eyes converging anteriorly, weakly convex; behind eyes rounding into
convex preoccipital margin. Eyes convex, in full face view clearly breaking lateral cephalic
outline. Mesosomal dorsum laterally marginate except pronotal margin posteriorly incomplete.
Pronotal humeri armed with short, distinct teeth, their lateral margins continued posteriorly before
merging with the outermost dorsal striae and curving onto sides. Promesonotal suture distinct;
metanotal groove indicated by a shallow emargination in lateral outline of segment. Mesonotal
and propodeal dorsa fused, posteriorly unarmed; descending into declivity in an abrupt,
uninterrupted curve. Petiole with dorsal margin entire; lateral spines weakly curved backwards
with tips upturned. First gastral segment with base medially truncate, anterodorsally immarginate
and evenly curved onto dorsum of segment.
Mandibles finely longitudinally striate. Clypeus and sides of head shagreened, overlaid by
weak irregular rugosity; rest of head with rather weak and shallow, mostly longitudinal striation,
most distinct on vertex with striae converging between and behind frontal carinae, fading before
reaching basal clypeal margin. Mesosomal dorsum with rather distinct, more-or-less regular
striae; anteriorly converging on pronotal dorsum with outermost striae continued obliquely along
sides. Striae on mesonotal and propodeal dorsa mostly longitudinal with outermost striae forming
rounded posterior corners; median striae rounding into propodeal declivity, terminating just below
level of dorsal face of segment. Sides of propodeum with mostly horizontally aligned striae.
Petiole with both faces rather smooth, finely shagreened. Sides of first gastral segment distinctly
longitudinally striate, striae becoming much finer towards dorsal and anterior faces of segment
that are shagreened.
Off-white, sparse, rather short, erect or semierect hairs on all body surfaces, including leading
edge of antennal scape. Greyish appressed pubescence present as scattered short hairs on dorsum
of head; more concentrated on pronotal humeri, propodeal dorsum, petiole and coxae. Sides of
propodeum and dorso-lateral borders of propodeal declivity with patches of relatively dense
pubescence. Gastral dorsum anteriorly and laterally with light cover of off-white or golden
pubescence, not masking underlying sculpture.
Black and rather shiny with appendages almost uniformly very dark reddish brown or black.
Mandibular masticatory border reddish brown. Apex of last funicular segment and apical tarsal
segments, distinctly light yellowish brown.
PAPUA NEW GUINEA: E. Highlands Prov., Chuave, 06º07’S, 145º08’E, 22km SE, 2300m,
24-28.xi.1967 (P. Colman) (w).
ETYMOLOGY
Named after the late B.B. (Bede) Lowery, S.J., whose significant collecting skills and activities
across Australia, New Guinea and elsewhere in Southeast Asia, brought to light many new ant
species, including a number of those described here.
REMARKS
The specimens of the type series were collected at the base of a Casuarina tree on the edge of
a coffee plantation. P. bedeloweryi is relatively close to P. porcata. They share almost identical
sculpture of the mesosomal dorsum. However, bedeloweryi differs in having the striation upon the
head much finer, the eyes distinctly more convex, the petiole finely shagreened and the first
gastral segment with the base only shallowly truncate, lacking the anterodorsal process found in
porcata.
Polyrhachis (Aulacomyrma) breviata, new species

Figures 56, 59, 62
TYPE MATERIAL
HOLOTYPE: INDONESIA, IRIAN JAYA (New Guinea: Neth. on data label), Wisselmeren
(= Wissel Lakes), Urapura, Kamo Valley, 03º55’S, 136º15’E, 1530m, 11.viii.1955, J.L. Gressitt
(worker). PARATYPE: data as for holotype (worker). Type distribution: holotype in MCZC;
paratype in QMBA.
WORKER

(95); SL 1.22-1.25 (1.22); SI 112; PW 0.78-0.81 (0.78); MTL 1.12 (2 measured).
Anterior clypeal margin arcuate, shallowly emarginate medially; in profile clypeus weakly
sinuate with shallowly impressed basal margin. Frontal carinae relatively short with laminate
lobes. Sides of head convex in front of eyes, rounding behind into rather shallowly convex
preoccipital margin. Eyes moderately convex, in full face view breaking lateral cephalic outline.
Mesosoma rather short, somewhat antero-posteriorly compressed, dorsum completely
immarginate. Pronotal dorsum widest at midlength; humeri armed with short, acute teeth.
Promesonotal suture distinct; metanotal groove lacking. Mesonotal-propodeal dorsum rounding
into sides and declivity in uninterrupted curve. Dorsal margin of petiole medially emarginate with
a pair of acute intercalary teeth flanking emargination; additional pair of blunt, tooth-like
projections, half way between intercalary teeth and rather short lateral teeth. Anterior face of first
gastral segment truncate medially, rounding onto dorsal face of segment, without a distinct
anterodorsal margin.
Mandibles finely longitudinally striate. Sculpture of head and mesosoma consisting of
relatively fine striation; striae on head mostly longitudinal along moderately convex outline of
head; V-shaped on clypeus and vertex, meeting along midline. Pronotal dorsum with transverse,
strongly anteriorly bowed striae continued obliquely on sides. Striae on mesonotal-propodeal
dorsum forming virtual semicircles or U-shaped patterns, continued obliquely along sides;
posterior-most striae medially interrupted by rather smooth surface at base of propodeal declivity.
Petiole with anterior face weakly transversely striate, medially overlaid by superficial, irregular,
mostly vertical, reticulation; posterior face shagreened. First gastral segment more finely
longitudinally striate, striae on dorsum converging anteriorly and merging into rather smooth,
finely shagreened base.
A few erect or semi-erect, rather short, off-white or silvery hairs scattered on dorsum of head
and mesosoma, a few hairs arising from leading edge of antennal scapes and dorsal margin of
petiole. Propodeal declivity bordered laterally with short, semi-erect, silvery-gray hairs, that
extend onto propodeal lobes. Somewhat longer, yellowish to golden hairs around apex and venter
of gaster, almost completely absent from dorsum. Appressed pubescence virtually absent, except
on gastral dorsum where it is silvery to golden and partly obscures underlying sculpture.
Black, including antennal scapes and coxae. Funiculi medium reddish brown with proximal
portion of basal segments bordered black. Legs mostly light to medium yellowish brown; distal
half of femora, proximal end of tibiae and basal segment of tarsi dark reddish brown.
ETYMOLOGY
Name derived from the Latin word brevior, meaning shorter, referring to the distinctly short
body.
REMARKS
P. breviata is rather similar to gressitti, with both species featuring a distinctly shortened
body and similar configuration of the petiole, including an emarginate dorsal margin bordered by
intercalary teeth. However, gressitti is easily recognized by its very high propodeal declivity and
the direction of the striae that are mostly longitudinal on the vertex. In breviata the striae on the
vertex are distinctly V-shaped and meet along the midline. This species also somewhat resembles
annulata with distinguishing characters listed in the remarks section under that species.
Polyrhachis (Aulacomyrma) decora, new species

Figures 57, 60, 63
TYPE MATERIAL
HOLOTYPE: INDONESIA, IRIAN JAYA (New Guinea: Neth. on data label), Vogelkop,
Fak Fak, S. coast of Bomberai Pen., 02º55’S, 132º18’E, 10-100m, 3.vi.1959, T.C. Maa (worker).
PARATYPES: data as for holotype (3 workers); Vogelkop, Bomberi, Bomberai Pen., 02º 48’S,
132º50’E, 700-900m, 6.vi.1959, T.C. Maa (2 workers); ditto, 10.vi.1959, J.L. Gressitt (worker).
Type distribution: holotype and 2 paratypes in MCZC; 1 paratype each in ANIC, BMNH, CASC
and QMBA.
WORKER

(83); SL 1.37-1.47 (1.43); SI 126-131 (131); PW 0.90-0.94 (0.90); MTL 1.43-1.47 (1.43) (7
measured).
Clypeus with anterior margin arcuate, shallowly medially emarginate; in profile clypeus
weakly convex with distinct notch anteriorly formed by furrow between clypeal striae; basal
margin very weakly impressed. Frontal triangle indistinct. Frontal carinae very short, strongly
raised with laminate lobes. Eyes very convex, protuberant, somewhat protracted posteriorly. Sides
of head in front of eyes weakly converging anteriorly; strongly converging behind into laterally
and posteriorly immarginate posterolateral corners; in lateral view relatively high vertex descends
abruptly into narrow margin. Mesosomal dorsum laterally and posteriorly immaginate. Pronotal
humeri armed with very strong, laterally and anteriorly directed spines with raised margins,
dorsum of spines shallowly concave. Promesonotal suture distinct; metanotal groove lacking.
Combined length of mesosomal and propodeal dorsa distinctly longer than pronotum. Propodeal
dorsum descending uninterrupted into rather short propodeal declivity. Petiole scale-like with
sharp dorsal margin terminating laterally in short, acute teeth. Base of first gastral segment
anteriorly convex.
Mandibles finely longitudinally striate. Sculpture of head, mesosoma and petiole consisting of
strongly raised, regularly spaced striae with deep interspaces resulting in a “ploughed”
appearance; striae anteriorly converging on clypeus, U-shaped on apex of vertex, strongly curved
medially from posterolateral corners towards central area between frontal carinae. Pronotal
dorsum with inverse, open V-shaped striae, descending obliquely and uninterrupted onto sides.
Mesonotal and propodeal dorsa with regular, U-shaped striae that are continued along sides and
interrupted medially by smooth surface of declivity. Petiole with dorsally bowed striae on both
faces. Gaster with distinctly finer striae, longitudinal on sides, less distinct and anteriorly
converging on dorsum.
Off-white, erect, medium length hairs on dorsum of head and mesosoma, a few distinctly
shorter hairs along dorsal margin of petiole. Hairs on gaster golden, somewhat longer, curved
posteriorly on dorsum. Silvery, appressed pubescence virtually absent from head and mesosoma;
small, thin patches on propodeal lobes, along declivity, coxae and subpetiolar process. Gaster with
fairly dense, reddish-golden pubescence mostly restricted to dorsum of first segment.
Black with very smooth and glossy striae on head, mesosoma and petiole. Antennal scapes
very dark, funiculi dark reddish brown with only apical funicular segment distinctly lighter. Legs
dark reddish brown, except trochanters and femora, blending from light yellow over proximal
halves to very dark brown at distal ends.
PAPUA NEW GUINEA: Huon Pen., Mongi Watershed, Wamuki, 800m, 19-20.iv.1955 (E.O. Wilson
#847 (w).
ETYMOLOGY
Named for its highly decorative sculpture pattern.
REMARKS
P. decora is rather similar to gressitti, kokoda and mamba which all have virtually identical
sculpture of the mesosoma and a rather convex base of the first gastral segment. It differs from all
of them in having a highly convex vertex, a feature notably distinct in lateral view. Additionally,
decora differs from gressitti in having the mesosomal dorsum distinctly longer, from kokoda by
the longitudinally striate first gastral segment and from mamba by the strongly convex,
protuberant eyes.
Polyrhachis (Aulacomyrma) excellens Viehmeyer, 1912

Figures 64, 67, 70
Polyrhachis excellens Viehmeyer, 1912: 14, fig. 19, 19a, 19b. Holotype worker. Type locality: NEW
GUINEA (Kais. Wilhelmsland on data label), Torricelli Mts (Schlaginhaufen), SNSD (examined).
Polyrhachis (Aulacomyrma) excellens Viehmeyer; Emery, 1921: 17. Combination in P. (Aulacomyrma).
Worker

MTL 1.40.
Anterior clypeal margin arcuate with shallow notch medially; basal margin indicated by
hairline break in cephalic sculpture. Frontal triangle indistinct. Frontal carinae strongly raised and
sinuate with laminate lobes; central area relatively wide with median stria forming short,
longitudinal carina. Sides of head in front of eyes weakly converging anteriorly. Eyes strongly
convex, in full face view clearly extending beyond lateral cephalic outline. Pronotal dorsum
immarginate, rather flat in profile with greatest width at about midlength. Pronotal humeri armed
with very prominent, acute spines. Promesonotal suture distinct, well impressed. Mesonotal and
propodeal dorsa fused, with ill-defined lateral margins formed by outermost dorsal striae,
terminating posteriorly in short, dorsally flattened, blunt teeth. Propodeal dorsum descending into
declivity in even, uninterrupted curve. Petiole with dorsal margin sharp, entire; lateral spines
relatively short, curved backwards. First gastral segment concave anteriorly; dorsal margin of
concavity produced dorso-medially into an extremely high, lip-like prominence that appears very
thin in lateral view and almost reaches height of apex of petiolar margin.
Mandibles finely longitudinally striate at bases, more reticulate-punctate at masticatory
borders. Sculpture of head and mesosoma very prominent, consisting of strongly raised,
regularly-spaced striae that are longitudinal on sides of head and mesonotal-propodeal dorsum;
anteriorly converging on clypeus, vertex and pronotal dorsum; oblique on sides of mesosoma.
Dorsal surface of pronotal spines and both faces of petiole smooth, microscopically shagreened.
First gastral segment very finely striate, striae converging anteriorly and terminating before
anterodorsal process that is finely microscopically shagreened.
Semi-erect to erect, short, off-white hairs fringing outer mandibular margins and antennal
scapes, only a few hairs arising from clypeus, sides of head near mandibular bases, frontal carinae,
vertex and pronotal dorsum. Hairs completely absent from mesonotal-propodeal dorsum. Gaster
with a few hairs dorsally and numerous, medium length, golden hairs along posterior margins of
apical gastral segments and at apex. Appressed pubescence virtually absent from body, except
small diffused patch bordering propodeal declivity.
Body black with striae smooth and shiny. Antennal scapes and femora dark to very dark
reddish-brown. Rest of legs medium to light yellowish-brown, except proximal and distal ends of
segments bordered dark reddish-brown. Funiculi missing.
REMARKS
The holotype is the only known specimen of this spectacular species. The very strongly
produced anterodorsal margin of the first gastral segment in P. excellens represents the most
extreme development of this remarkable character.
Polyrhachis (Aulacomyrma) geometrica Fr. Smith, 1859

Figures 65, 68, 71
Polyrhachis geometricus Fr. Smith, 1859: 141. Holotype worker. Type locality: INDONESIA, Aru I. (A.R.
Wallace), OXUM (examined).
Polyrhachis (Aulacomyrma) geometrica Fr. Smith: Donisthorpe, 1932: 453. Combination in (Aulacomyrma).
WORKER
Dimensions: TL c. 4.89-5.14 (4.89); HL 1.25-1.34( 1.25); HW 1.09-1.18 (1.09); CI 87-88

(87); SL 1.31-1.37 (1.31); SI 116-122 (120); PW 0.90-0.94 (0.94); MTL 1.25- 1.28 (1.25) (3
measured).
Anterior clypeal margin arcuate, only very shallowly truncate medially; clypeus virtually
straight in profile. Frontal carinae strongly sinuate, with raised margins; central area relatively
wide with distinct, longitudinal carina running towards anterior clypeal margin. Frontal triangle
indistinct. Eyes convex, clearly breaking lateral cephalic outline in full face view. Sides of head in
front of eyes only weakly converging anteriorly. Pronotum immarginate, widest just below humeri
that are armed with small, acute, dorso-ventrally flattened, triangular teeth. Promesonotal suture
distinct; metanotal grove lacking, position indicated by slight change in direction of dorsal
sculpture and slight depression in lateral outline. Propodeum unarmed. Petiole with dorsal margin
emarginate between distinct intercalary teeth; lateral spines relatively long, curved backwards.
Anterior face of first gastral segment very shallowly concave medially at base.
Head in front of eyes and on sides more-or-less regularly, longitudinally striate; preoccipital
margin with median striae transversely bowed and lateral striae converging anteriorly towards
central area between frontal carinae. Sculpture of pronotal dorsum consisting of inverse, wide U-
and V-shaped striae, that are laterally uninterrupted and continuous along sides. Mesonotal
dorsum with more regular, semicircular striae, outermost forming an ill-defined lateral margin;
propodeal dorsum with striae continued obliquely along sides. First gastral segment distinctly, and
more densely, longitudinally striate; striae converging anteriorly towards base.
Holotype with pilosity almost completely absent from dorsum of mesosoma and gaster, a few
erect to semierect, short hairs on dorsum and sides of head, vertex, along antennal scapes and on
tibia of middle and hind legs. Rather sparse, Appressed, silvery pubescence scattered over dorsum
of head and mesosoma; more abundant along dorso-posterior borders of gastral segments.
Black, including antennal scapes, coxae, trochanters (except narrow light band at distal ends)
and femora of middle and hind legs. Femora of front legs, tibiae and tarsi distinctly light yellowish
or reddish brown, except proximal and distal ends of segments very dark brown or black. Basal
funicular segments very dark brown with paler, reddish brown bands at their apices; lighter bands
becoming progressively wider in distal funicular segments, that are mostly pale, reddish-brown.
PAPUA NEW GUINEA: Huon Pen., lower Busu R., 12.v.1955, lowl. rf., E.O. Wilson #1018 (w). Milne
Bay Prov., Opanabru, 12mi W of Raba Raba, 2600ft, 1.viii.1969, rf, R. Pullen (worker).
REMARKS
P. geometrica is similar to the other species of the complex, notably to annulata, breviata and
monticola, and in addition to the characteristic “geometrical” sculpture they share the medially
emarginate dorsal margin of the petiole and rather shallowly concave base of the first gastral
segment. However, geometrica is easily recognizable by the color of the front femora which are
very light, yellowish brown in contrast to the entirely black femora of the middle and hind legs. In
the other allied species the femora of all legs are more-or-less uniform in color.
Polyrhachis (Aulacomyrma) gressitti, new species

Figures 66, 69, 72
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA: Adelbert Mts, Wanuma, 04º36’S, 145º06’E,

800-1000m, 25.x.1958, J.L. Gressitt (worker). Type deposition: unique holotype in MCZC.
WORKER

MTL 1.25.
Anterior clypeal margin with minute notch medially; clypeus in profile virtually straight with
weakly impressed basal margin. Frontal triangle indistinct. Frontal carinae sinuate, with rather
short, laminate, anteriorly truncate lobes. Sides of head in front of eyes converging anteriorly;
rounding behind eyes into convex preoccipital margin. Eyes strongly convex, protuberant, in full
face view clearly breaking lateral cephalic outline. Mesosoma laterally and posteriorly
immarginate, distinctly antero-posteriorly compressed, notably when viewed from side. Pronotal
humeri armed with rather strong, horizontal, anteriorly directed spines; pronotal dorsum evenly
curved in profile. Promesonotal suture strongly impressed; metanotal groove lacking.
Mesonotal-propodeal dorsum distinctly wider than long; propodeal declivity exceptionally high.
Petiole with dorsal margin acute, sinuate in dorsal view, lateral spines relatively short, curved
backwards. Anterior face of first gastral segment convex.
Mandibles finely longitudinally striate. Sculpture of head and mesosoma consisting of
regularly spaced, smooth and glossy striae; mostly longitudinal and somewhat anteriorly
converging on head, transverse and anteriorly bowed on pronotum; striae on mesonotal-propodeal
dorsum forming imperfect semicircular pattern with outmost striae continuing obliquely onto
sides and posteriorly extending into propodeal declivity. Petiole with both faces transversely
striate, striae less distinct dorsally, becoming wrinkles just below dorsal margin. First gastral
segment distinctly longitudinally striate, dorsum with striae finer and distinctly converging
towards base.
Mostly medium length, erect or curved, golden hairs on dorsum of head, mesosoma and
gaster; distinctly shorter hairs along leading edge of antennal scapes, dorsal margin of petiole and
legs. Appressed pubescence virtually absent from most body surfaces, except dorsum of first
gastral segment, where it is rather diluted and reddish.
Black; mandibular masticatory border, antennal scapes, most of femora, basal segment of
tarsi and apical segments of gaster reddish brown to dark rusty red. Funiculi, trochanters, proximal
ends of femora and apical tarsal segments, light to very light reddish brown.
ETYMOLOGY
Named in honor of the collector, the late J.L. Gressit, Bernice P. Bishop Museum, Honolulu,
Hawai’i.
REMARKS
Another remarkable species very similar to P. decora, but with a distinctly longer and
exceptionally high propodeum with its outline forming an almost perfect semicircle in dorsal
view.
Polyrhachis (Aulacomyrma) impressa, new species

Figures 73, 76, 79
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Huon Pen., lower Busu R., 10.v.1955, lowl. rf., E.O.
Wilson #1003 (worker). PARATYPES: data as for holotype, but 27.iv.1955, E.O. Wilson #706
(worker); 10.v.1955, E.O. Wilson #988 (2 workers); 12.v.1955, E.O. Wilson #1018 (worker) and
17.v.1955, E.O. Wilson #1054 (7 workers, 1 dealate ♀). Type distribution: holotype, 1 paratype
worker and paratype ♀ in MCZC; 2 paratype workers each in ANIC, BMNH, CASC and QMBA.
WORKER
Dimensions: TL c.5.09-5.69 (5.19); HL 1.31-1.47 (1.34); HW 1.12-1.28 (1.12); CI 83-89

(83); SL 1.34-1.47 (1.34); SI 120-122 (120); PW 0.94-1.09 0.94); MTL 11.25-1.34 (1.28) (12
measured).
Anterior clypeal margin arcuate, narrowly truncate medially; basal clypeal margin flat,
indicated only by hairline break in cephalic sculpture. Frontal triangle indistinct. Frontal carinae
relatively short, strongly raised. Sides of head in front of eyes weakly converging towards
mandibular bases; strongly rounded behind into rather short preoccipital margin. Eyes convex,
somewhat sunk into shallow concavity in cephalic sculpture on sides of head; in full face view
only moderately extending beyond lateral cephalic outline. Mesosoma laterally immarginate with
dorsum rounding onto sides in unbroken curve. Pronotal humeri armed with short, acute spines.
Promesonotal suture deeply impressed; metanotal groove lacking. Propodeal dorsum delimited
posteriorly by distinct transverse ridge that separates dorsum from declivity, posterolaterally with
minute, somewhat dorsally raised, rounded prominences. Petiole with sharp dorsal margin; lateral
spines acute, weakly curved backwards and upwards. First gastral segment shallowly concave
anteriorly, anterodorsal margin of concavity blunt.
Sculpture of head and mesosoma mostly consisting of regularly spaced striae; longitudinal on
sides of head, converging anteriorly on clypeus and vertex; somewhat inversely U-shaped on
pronotal dorsum and obliquely rounding onto sides. Striae on mesosomal and propodeal dorsa
forming incomplete semicircles, outermost striae continued obliquely on sides of propodeum.
Both faces of petiole shagreened. First gastral segment with dorsum finely striate; intensity of
sculpture increasing laterally with sides of gaster very distinctly longitudinally striate.
Relatively short, off-white and yellowish hairs scattered over all dorsal surfaces of head,
mesosoma and gaster; hairs silvery and distinctly shorter along dorsal and lateral borders of
propodeal declivity and dorsal margin of petiole. Off-white or silvery appressed pubescence,
generally directed towards midline, present over all dorsal surfaces of body; most dense on
pronotal humeri and dorsum and sides of propodeum. Gastral pubescence golden, rather dense,
almost completely obscuring underlying sculpture dorsally.
Body black. Antennae, femora, proximal ends of tibiae and basal tarsal segments black or very
dark brown; extreme tip of apical funicular segment, distal end of trochanters, tibiae and apical
tarsal segments medium to light yellowish to reddish brown.
QUEEN
Dimensions: TL c. 6.20; HL 1.56; HW 1.31; CI 84; SL 1.53; SI 117; PW 1.25; MTL 150 (1
measured).
Queen essentially as worker, with usual characters identifying full sexuality, including three
ocelli and complete thoracic structure with wings. Sculpture of head and body similar to worker,
direction of striae following structural characteristics of fully developed mesosoma.
Male and immature stages unknown.
ETYMOLOGY
The name impressa is derived from a self explanatory term ‘impressive’ relating to the rather
striking appearance of this species.
REMARKS
P. impressa is very similar to P. orokana described below and both species share the laterally
immarginate mesosomal dorsum, strong transverse ridge separating the propodeal dorsum from
declivity and virtually identical sculpture of the mesosomal dorsum. They can be distinguished by
the shape of the fused mesonotal-propodeal dorsum which is about as wide as long in impressa
and is strongly transverse, distinctly wider than long in orokana. The eyes in impressa are only
moderately convex and situated in a shallow depression in the cephalic sculpture, only partly
extending beyond the lateral outline of the head in full face view. In contrast, the eyes in orokana
are strongly convex, protuberant and in full face view they fully extend beyond the lateral cephalic
outline. The antennal scapes in impressa bear numerous hairs over all their surfaces, while in
orokana the hairs are confined to their leading edges. The petiole in impressa has the lateral spines
relatively long and weakly bent backwards, while they are rather short and virtually straight in
orokana.
Polyrhachis (Aulacomyrma) kokoda, new species

Figures 74, 77, 80
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Northern Prov., Kokoda, 9.iii.1972, P.M. Room
#250 (worker). PARATYPES: data as for holotype (1 worker); Oivi Ridge, Kokoda Rd, 1200ft,
rf., 18.i.1971, B.B. Lowery (1 worker). Type distribution: holotype in ANIC; 1 paratype each in
MCZC and QMBA.
WORKER

(79); SL 1.36-1.43 (1.43); SI 135-139 (139); PW 0.97-1.03 (1.00); MTL 1.31-1.40 (1.37) (3
measured).
Clypeus with anterior margin medially truncate or very shallowly emarginate; median
longitudinal carina weakly elevated posteriorly before descending towards shallowly impressed
basal margin. Frontal carinae sinuate, with rather short, laminate, anteriorly truncate lobes. Sides
of head in front of eyes convex; narrowed behind into convex preoccipital margin. Eyes strongly
convex, clearly breaking cephalic outline in full face view. Mesosoma immarginate, dorsum
sloping onto sides and propodeal declivity in unbroken curve. Pronotal dorsum very strongly
raised anteriorly, convex in profile. Pronotal humeri armed with acute, rather long, downward
curved spines; bases broad, margins raised. Promesonotal suture distinct; metanotal groove
lacking. Petiole with both faces convex and dorsal margin somewhat angular; lateral spines short,
curved backwards. Anterior face of first gastral segment convex.
Mandibles distinctly, longitudinally striate. Sculpture of head, mesosoma and petiole
consisting of regularly spaced, rather smooth and glossy striae. Head with longitudinal, mostly
anteriorly converging striae; clypeal striae terminating just short of anterior margin. Striae on
mesosomal dorsum forming an imperfect elliptical pattern, divided in half by promesonotal
suture; outermost striae continued obliquely onto sides and posteriorly extending onto propodeal
declivity. Petiole with tranverse, dorsally arched striae on anterior face; inversely V-shaped on
posterior face and converging upwards towards dorsal margin. Gaster finely, microscopically,
shagreened.
Abundant, long, mostly erect or curved, golden hairs on most of body and appendages, hairs
reaching greatest diameter of eyes in length, distinctly shorter on antennal scapes and along dorsal
margin of petiole. Propodeal declivity with patch of very short, somewhat dorsally curved,
off-white hairs. Appressed, golden pubescence very sparse on dorsum of head and mesosoma;
more distinct on pronotal dorsum, forming V-shaped patch running from pronotal spines across
the segment, and posteriorly towards propodeal declivity. Thin covering of mostly white or silvery
pubescence on sides of mesosoma, coxae, appendages and posterior margins of gastral segments;
pubescence most abundant and distinctly reddish on gastral dorsum, somewhat obscuring
underlying sculpture.
Black, masticatory margins of mandibles, antennal scapes and most of legs medium to dark
reddish-brown. Trochanters and basal femora distinctly yellow.
PAPUA NEW GUINEA: Catalina Estate, 48km N of Port Moresby, 09º27’S, 147º09’E, 500m,
3.ix.1959 (T.C. Maa) (w).
ETYMOLOGY
Named after the type locality, Kokoda village, situated on the northern rim of the Owen
Stanley Ranges in the Northern Province of Papua New Guinea.
REMARKS
P. kokoda is superficially similar to P. mamba described below, with both species evidently
sympatric. The paratype of kokoda was collected together with a mamba queen at the same
locality at Oivi Ridge and they were subsequently mounted on the same pin. Both species share
the somewhat elliptical striation of the mesosomal dorsum but there the similarity ends. They
differ in numerous characters, including the shape of the eyes, which in kokoda are strongly
convex and clearly project beyond the lateral outline of the head. In contrast the eyes in mamba are
virtually flat and sunk into a shallow concavity in the cephalic sculpture and they do not break the
cephalic outline in full face view. In lateral view the outline of the mesosoma in kokoda features a
strongly convex pronotal dorsum and the propodeal dorsum descends into the declivity in an
uninterrupted curve. In mamba the pronotum is flatter and the propodeal dorsum descends
abruptly into a rather short, vertical declivity. The dorsum of the petiole has an acute margin in
kokoda, while it is bluntly rounded in mamba. The first gastral segment of kokoda is finely
shagreened in contrast to mamba, where its sides are distinctly, longitudinally striate. The
pubescence on the head and body is golden in kokoda, with a distinct reddish tint on the gastral
dorsum, while in mamba the pubescence is uniformly greyish or white.
Polyrhachis (Aulacomyrma) maai, new species

Figures 75, 78, 81
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, W. Highlands, Minj, 05º51’S, 144º40’E,

8-13.ix.1959, T.C. Maa (worker). PARATYPE: PAPUA NEW GUINEA, W. Highlands,
Nondugl, 05º52’S, 144º43’E, 1600m, 9.vii.1955 (J.L. Gressitt) (worker). Type distribution:
holotype in MCZC, paratype in QMBA.
WORKER

(92); SL 1.40-1.50 (1.50); SI 119-125 (119); PW 0.90-1.00 (1.00); MTL 1.34-1.53 (2 measured).
Anterior clypeal margin arcuate, narrowly and shallowly notched medially; clypeus weakly
convex in lateral view. Sides of head in front of eyes gently converging anteriorly; curving rather
abruptly behind eyes into relatively short preoccipital margin. Eyes moderately convex, only
marginally extending beyond cephalic outline. Frontal carinae only moderately raised; central
area with short, weakly defined, median carina. Mesosomal dorsum with ill-defined margins.
Pronotal dorsum with sides parallel; humeri armed with broad-based, triangular teeth with slightly
raised anterior and lateral margins, with latter continued posteriorly towards well impressed
promesonotal suture. Mesonotal-propodeal dorsa fused, sides weakly converging posteriorly
before widening and again converging and terminating in acute teeth. Petiole with dorsal margin
sharp, entire; lateral spines acute and curved backwards and slightly upwards. Anterior face of
first gastral segment concave, accommodating posterior face of petiole, anterodorsal margin of
concavity weakly medially produced above dorsal face of segment.
Head very smooth and shiny. Mesosomal dorsum smooth between widely spaced, longitudinal
striae, that are broken at promesosotal suture and terminate at point where propodeal dorsum
descends into declivity that is medially smooth, lacking any sculpture. Sides of mesosoma
similarly sculptured, with striae mostly oblique. Anterior face of petiole smooth and shiny,
posterior face and dorsum of first gastral segment shagreened. Sides of gaster with very fine,
closely spaced, longitudinal striae.
Off-white or yellowish hairs present on mandibles, anterior portion of clypeus and leading
edges of antennal scapes; a few hairs arising along frontal carinae and on vertex. In frontal view,
no hairs projecting from sides of head between eyes and mandibular bases. A few short, semierect
hairs, scattered over mesosomal and gastral dorsa, density and length increasing towards apex and
venter of gaster. Pubescence virtually lacking, except for patches of fine, silvery short hair, at
sides of propodeal declivity, anterior face of petiole, and dorsum of first gastral segment where it
has distinct reddish tint.
Body, including antennal scapes and legs, virtually black, only mandibular masticatory border
and apical segments of funiculi and tarsi dark to medium reddish-brown. Color of legs a shade
lighter in paratype.
ETYMOLOGY
Named in honor of the late T.C. (Tsing-Chao) Maa of the Tunghai University in Taiwan, who
was a longtime associate of the Bernice P. Bishop Museum, Honolulu, Hawai’i. Maa was a field
worker supreme who collected many of the types lodged in that museum, including a number of
the new species described in this paper.
REMARKS
P. maai is a very distinctive species that somewhat resembles excellens. Both species share
the similar, almost parallel-sided mesosomal dorsum with mostly longitudinal striation and a
rather glossy appearance. However, besides the very prominent, anterodorsal process of first
gastral segment in excellens, they differ in the sculpture of the head, that is very smooth and shiny
in maai and distinctly longitudinally striate in excellens. Also the interspaces between the
mesosomal striae are rather shallow in maai, while they are much deeper in excellens.
Polyrhachis (Aulacomyrma) mamba. new species

Figures 82, 85, 88
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Northern Prov., Owen Stanley Ra., Mamba
Plantation, c. 7km WNW of Kokoda, 08º51’S, 147º41’E, 500m, 31.viii-1.ix.1984, R.J. Kohout
acc. 403) (worker). PARATYPES: data as for holotype (60 workers, 1 dealate ♀). Type
distribution: holotype, most paratype workers and paratype ♀in ANIC; 2 paratype workers in
each BMNH, CASC, MCZC, LACM and QMBA.
WORKER
Dimensions: TL c. 6.00, 5.04-5.54; HL 1.47, 1.31-1.40; HW 1.28, 1.12-1.18; CI 87, 84-85;

SL 1.47, 1.37-1.40; SI 115, 119-122; PW 1.06, 0.90-0.94; MTL 1.50, 1.34-1.40 (4 measured).
Anterior clypeal margin arcuate, entire to narrowly truncate or very shallowly emarginate
medially. Longitudinal median carina weakly raised, terminating posteriorly in a shallow basal
margin. Frontal carinae strongly sinuate with rather short, laminate, anteriorly sharply truncate
lobes. Side of the head in front of eyes very weakly convex, rounded behind into preoccipital
margin. Eyes flat, situated well forward on sides of head, virtually identical to those in P. impressa
and P. roomi, not breaking lateral cephalic outline in full face view. Mesosoma immarginate
laterally and posteriorly. Pronotal humeri armed with rather broad-based, laterally and anteriorly
directed, more-or-less blunt spines. Promesonotal suture shallow; metanotal groove lacking.
Petiole in profile with dorsum rounded, lateral teeth minute, upturned. Anterior face of first gastral
segment convex.
Mandibles finely and densely, longitudinally striate. Dorsum of head with mostly regular
striae, similar to those in P. roomi in orientation; longitudinal and anteriorly converging on
clypeus; obliquely curving from sides of head towards basal clypeal margin; transverse and
anteriorly bowed on vertex, outermost striae converging forwards between frontal carinae.
Mesosoma with striation resembling that of P. kokoda, notably on mesonotum and propodeum.
Pronotal striation in mamba somewhat different, due to shape of dorsum between pronotal spines;
curved inwards in kokoda, straight in mamba. Anterior and posterior faces of petiole with
tranverse, slightly dorsally bowed striae, that are connected along sides. First gastral segment with
sides finely longitudinally striate; dorsally striae becoming less distinct, dorsum finely
shagreened.
Short to medium length, erect, curved or sinuate, off-white to silvery hairs present on dorsum
of head, mesosoma, petiole and first gastral segment; somewhat longer, yellowish to golden and
more erect hairs on legs and gaster. Appressed, mostly greyish pubescence in various densities on
most of body, notably on pronotal dorsum, including spines, meso- and metapleuron, lateral
borders of propodeal declivity and coxae; somewhat longer on dorsum of gaster and almost
obscuring underlying sculpture.
Black; mandibular masticatory border, antennal scapes, joints of femora and tibiae, basal
tarsal segments and apical segments of gaster dark reddish-brown. Funiculi, except base of first
segment, and most of legs light to very light reddish-yellow.
QUEEN
Dimensions: TL c. 6.35; HL 1.53; HW 1.37; CI 89; SL 1.56; SI 114; PW 1.34; MTL 1.59 (1
measured).
Similar to worker with usual differences indicating caste. Sculpture of head and body similar
to worker, direction of striae following structural characteristics of fully developed mesosoma.
Colour as in worker except funiculi distinctly darker and femora and tibiae with more extensive
dark patches around joints.
PAPUA NEW GUINEA: Northern Prov., Kokoda, 25.v.1972 (P.M. Room #360) (w); ditto, 1.vi.1972
(P.M. Room #360) (w); Oivi Ridge, Kokoda Rd, 1200ft, rf., 18.i.1971, B.B. Lowery (dealate♀). Morobe
Prov., Bulolo, 2500-3500ft, rf., 2.i.1968, B.B. Lowery (w).
ETYMOLOGY
Species named after the type locality, Mamba Plantation nr Kokoda, Papua New Guinea.
REMARKS
P. mamba is similar to P. kokoda, with distinguishing characters given in the remarks section
of the latter species. Among these, the most important are the flat eyes in mamba that do not reach
the lateral cephalic outline in full face view. Also, the striae on the vertex in mamba are
tranversely bowed, while they are longitudinal in kokoda. One dealate queen was collected
foraging with workers on the trunks and branches of freshly felled trees on the edge of a recent
rainforest clearing. The second queen was collected with a kokoda worker at Oivi Ridge nr
Kokoda.
Polyrhachis (Aulacomyrma) monticola, new species

Figures 83, 86, 89
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Morobe Prov., Sarawaget Ra., 3km E of Gain,
06º25’S, 146º46’E, 1000-1200m, rf., 26.viii.1984, R.J. Kohout acc. 84.363 (worker). Type
deposition: unique holotype in ANIC.
WORKER

MTL 1.31.
Anterior clypeal border arcuate, emarginate medially; clypeus straight in profile, basal
margin weakly impressed, laterally indicated by distinct line breaking cephalic sculpture. Frontal
carinae with lamellate lobes. Sides of head weakly convex in front of eyes, rounding behind into
weakly convex preoccipital margin. Eyes moderately convex, only marginally exceeding lateral
cephalic outline. Mesosomal dorsum immarginate for most of length, distinctly converging
posteriorly. Pronotal humeri armed with broad-based, triangular, dorso-ventrally flattened teeth.
Promesonotal suture relatively shallow, distinctly incised laterally; metanotal groove only weakly
indicated by faint break in sculpture. Mesonotal dorsum with outermost dorsal striae forming an
ill-defined lateral margin. Propodeal dorsum immarginate, terminating posteriorly in small
rounded prominences formed by inwardly curved lateral striae; medially dorsum sloping into
declivity in rather abrupt, uninterrupted curve. Dorsal margin of petiole with acute intercalary
teeth, bordering medial emargination, margin jagged laterally, terminating in relatively long,
strongly backward- curved spines with weakly upturned tips. First gastral segment concave
anteriorly; concavity with blunt dorsal margin that is not elevated above dorsal surface of
segment.
Mandibles finely, somewhat irregularly, longitudinally striate-rugose. Sculpture of head
consisting of regularly spaced striae; V-shaped on clypeus and mostly longitudinal on sides; striae
on vertex converging anteriorly between and along frontal carinae. Striae on pronotal dorsum
strongly converging anteriorly; striae on mesonotal-propodeal dorsum converging posteriorly,
forming V-shaped pattern; laterally striae continued obliquely onto sides. Petiole with both faces
microscopically shagreened. First gastral segment distinctly, longitudinally striate laterally;
dorsally striae becoming finer and less distinct, medially and anteriorly finely shagreened.
A few short, erect or curved, yellowish hairs on head, including mandibular masticatory
border, anterior clypeal margin, sides of head between eyes and posterolateral corners and along
leading edges of antennal scapes; hairs absent between eyes and bases of mandibles or along
inferior edges of scapes. Mesosomal dorsum and first gastral segment with a few short erect hairs,
variously curved. Petiole with several very short hairs fringing dorsal margin. Apex of gaster with
numerous, relatively long, golden hairs. Mostly silvery, very sparse appressed pubescence on
head; more abundant on pronotal humeri, propodeal dorsum, sides and upper portion of declivity,
coxae, petiole and posterior margins of gastral segments; first gastral segment with distinctly
golden pubescence almost obscuring sculpture.
Black; antennae, femora, bases of tibiae and basal tarsal segments very dark reddish-brown.
Tip of last funicular segment, distal ends of trochanters, most of tibiae and tarsi, light to very light
reddish-brown.
ETYMOLOGY
Name derived from the Latin word montis, meaning mountain.
REMARKS
Known from a single worker collected foraging on low vegetation. P. monticola is rather
similar to annulata and breviata and shares with them the somewhat antero-posteriorly
compressed body and the medially emarginate dorsal petiolar margin. However, monticola is
easily separated by the sculpture of the mesonotal-propodeal dorsum that consists of V-shaped,
posteriorly converging striae, while in annulata and breviata the striae form virtual semicircles or
U-shaped patterns.
Polyrhachis (Aulacomyrma) orokana, new species

Figures 84, 87, 90
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, E. Highlands, Orokana, 27.xi.1967, under bark of

tree, B.B. Lowery (worker). PARATYPES: data as for holotype (3 workers). Type distribution:
holotype in ANIC; 1 paratype each in BMNH, MCZC and QMBA.
WORKER
Dimensions TL c. 4.94-5.69 (c. 4.94); HL 1.25-1.31 (1.25); HW 1.00-1.06 (1.03); CI 76-82

(82); SL 1.25-1.28 (1.25); SI 121-125 (1.21); PW 0.87-0.90 (0.870; MTL 1.18-1.22 (1.18) (4
measured).
Anterior clypeal margin arcuate, entire; basal margin very weakly impressed, indicated by
hairline break in cephalic sculpture. Frontal triangle indistinct. Frontal carinae relatively short,
strongly raised with laminate lobes. Sides of head slightly convex in front of eyes, rounding
almost immediately into preoccipital margin behind. Eyes strongly convex, situated well back on
sides of head, fully protruding beyond lateral cephalic outline. Mesosoma laterally immarginate
with dorsum rounding onto sides in unbroken curve; evenly convex in profile. Pronotal humeri
armed with somewhat dorso-ventrally flattened, acute teeth; anterior margins dorsally raised;
lateral margins continued obliquely downwards and merging on sides with lateral striae.
Promesonotal suture distinct, metanotal groove lacking. Mesonotal and propodeal dorsa fused,
distinctly transverse, separated from declivity by distinct transverse ridge terminating laterally in
rounded prominences. Petiole with acute dorsal margin terminating laterally in very small,
upturned teeth. First gastral segment shallowly concave anteriorly, anterodorsal margin of
concavity obtuse, not raised above dorsal face of segment.
Sculpture of head and mesosoma consisting of regularly spaced striae, mostly longitudinal on
head, converging anteriorly on clypeus. Striation on pronotal dorsum inversely U-shaped
anteriorly, becoming widely V-shaped towards promesonotal suture and continuing obliquely
onto sides. Mesonotal and propodeal dorsa with almost semicircular striae, extended obliquely
along sides. Petiole shagreened. First gastral segment longitudinally striate laterally and dorsally.
Rather short, erect or curved, silvery and off-white hairs on most dorsal surfaces of body,
shortest lining dorsal margin of petiole, longest and yellowish on dorsum and apical gaster.
Silvery appressed pubescence on pronotal dorsum forming distinct V-pattern, running diagonally
from pronotal humeri towards middle and dispersing before reaching promesonotal suture.
Pubescence on mesonotal and propodeal dorsa becoming rather dense and somewhat directed
medially; sparse on sides of propodeum, dense posteriorly on propodeal lobes and coxae.
Propodeal declivity bordered laterally and dorsally with very fine, short, upward curved
pubescence. Petiole and dorsum of gaster with fairy abundant pubescence, not obscuring
underlying sculpture.
Black. Antennae, femora, proximal ends of tibiae and first tarsal segment very dark to dark
reddish-brown, tibiae and tarsi distinctly light yellow.
Queen and male unknown. Larvae of various stages of development and a pupa in ANIC.
ETYMOLOGY
Named after the locality Orokana in the Southern Highlands Province of Papua New Guinea.
REMARKS
P. orokana is very similar to P. impressa, both featuring almost the same sculpture pattern of
the mesosomal dorsum, somewhat posteriorly protracted eyes and the concave anterior face of
first gastral segment. However they differ in a number of characters, including the shape of the
mesosomal-propodeal dorsum which, in orokana is transverse, distinctly wider than long, while in
impressa it is as long as wide. The eyes in orokana are convex and protuberant and fully extend
beyond the lateral cephalic outline in full face view. In comparison the eyes in impressa are less
convex, somewhat sunk into a shallow concavity in the cephalic sculpture and extend only
moderately beyond the cephalic outline. The lateral petiolar teeth in orokana are very small and
upturned, while in impressa they are relatively long and curved backwards and upwards. The
lateral gaster has distinct, horizontally directed, longitudinal striae in orokana; while in impressa
the striae are oblique and directed towards the dorso-anterior margin of the first gastral segment.
Polyrhachis (Aulacomyrma) pelecta, new species

Figures 91, 94, 97
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Karubaka, Swart Valley, 03º35’S, 138º30’E,

1500m, 11.xi.1958, J.L. Gressitt (worker). PARATYPE: data as for holotype (1 worker). Type
distribution: holotype in MCZC; paratype in QMBA.
WORKER
Dimensions (holotype cited first): TL c. 5.49; HL 1.47, 1.43; HW 1.28, 1.28; CI 87, 89; SL
1.50, 1.50; SI 117, 117; PW 1.12, 1.12; MTL 1.43, 1.40 (2 measured).
Anterior clypeal margin arcuate, entire; slightly sinuate in profile. Frontal triangle indistinct.
Frontal carinae strongly sinuate, raised, with laminate lobes; central area relatively wide with
rather distinct longitudinal carina formed by strongly raised median stria. Sides of head gently
convex, only weakly converging anteriorly. Eyes convex, in full face view breaking lateral
cephalic outline. Pronotal humeri armed with broad-based triangular teeth, that are shallowly
concave dorsally between distinctly raised margins; lateral margins continued for some distance
before merging with dorsal striae along sides. Promesosonal suture distinct; metanotal groove
lacking. Mesonotal dorsum only partially marginate. Propodeal dorsum immarginate, with
posterior angles weakly indicated. Petiole relatively narrow, as wide as high, with dorsal margin
somewhat jagged; lateral spines short, slender, directed outwards and only slightly curved
backwards. Anterior face of first gastral segment only shallowly concave medially at base.
Sculpture of head consisting of regularly spaced striae; mostly longitudinal, converging
medially and anteriorly on clypeus and between frontal carinae. Pronotal dorsum with broadly
open, inverse V-shaped striae. Sculpture of mesonotal-propodeal dorsa differing between
holotype and paratype; striae in holotype forming incomplete semicircles, open inversely
U-shaped posteriorly; in paratype striae strongly converging posteriorly and meeting along
midline in V-shaped pattern. Dorsum of first gastral segment microscopically shagreened,
becoming finely longitudinally striate towards sides.
Short, erect or semierect, off-white or yellowish hairs present in various densities on most
body parts, except inferior edge of antennal scapes. Rather sparse, appressed pubescence
everywhere; more dense on coxae and anterior face of petiole and propodeal declivity that is
bordered laterally by very short, greyish, semierect fine hairs. Gastral dorsum with light cover of
silvery pubescence not concealing underlying sculpture.
Black; antennae and legs dark to very dark reddish-brown. Mandibular masticatory borders,
funiculi, tibiae, except proximal ends, and distal tarsal segments distinctly lighter.
ETYMOLOGY
Name derived from the Latin pellax, meaning deceitful or decoy, in reference to its close
resemblance to P. geometrica.
REMARKS
P. pelecta is rather close to P. geometrica but differs in having the pronotal teeth broad-based,
the lateral propodeal margins feebly dentate posteriorly, the dorsal margin of the petiolar node
more-or-less entire and the first gastral segment with very fine striations confined to its sides. In
contrast, the pronotal teeth in geometrica are distinctly smaller and more acute, the propodeum
smoothly rounds into the declivity, the dorsal margin of petiolar node is distinctly medially
emarginate and the whole surface of first gastral segment is distinctly and finely, striate.
Polyrhachis (Aulacomyrma) porcata Emery, 1921

Figures 92, 95, 98
Polyrhachis (Aulacomyrma) porcata Emery, 1921: 20. Holotype worker. Type locality: NEW GUINEA
(NE), (Staudinger et Bang-Hass), MCSN (examined).
Polyrhachis (Aulacomyrma) porcata Emery, 1925: 198, Pl. 3, figs 29, 29a.
WORKER
Dimensions (holotype cited first): TL c. 5.74, 6.35; HL 1.47, 1.68; HW 1.28, 1.47; CI 87, 87;
SL 1.56, 1.75; SI 122, 119; PW 1.06, 1.18; MTL 1.62, 1.75 (2 measured).
Anterior clypeal margin arcuate, with very narrow medial notch; clypeus weakly convex in
profile with basal margin slightly impressed medially and marked laterally by a hairline break in
cephalic sculpture. Frontal triangle indistinct; frontal carinae strongly sinuate with laminate lobes.
Eyes only weakly convex, not breaking cephalic outline in full face view. Sides of head weakly
converging anteriorly; rounding behind eyes into convex preoccipital margin. Pronotal dorsum
immarginate; humeri armed with short, laterally and anteriorly directed, rather acute spines,
margins raised with dorsal surface between them weakly concave. Promesosonal suture strongly
impressed; metanotal groove lacking. Mesonotal and propodeal dorsa with outermost striae
stronger than others, forming ill-defined and incomplete lateral margins. Propodeal dorsum
posteriorly immarginate, unarmed, descending into declivity in sudden, unbroken curve. Petiole
with dorsal margin acute, entire, somewhat medially and dorsally compressed and sinuate in
dorsal view; lateral teeth rather short, curved posteriorly. Anterior face of first gastral segment
concave medially, dorsal margin of concavity produced dorso-medially into prominent ridge,
distinctly raised above dorsal surface of segment.
Mandibles finely, longitudinally striate. Head mostly longitudinally striate with striae on
vertex converging medially between frontal carinae. Pronotal dorsum with outermost striae almost
semicircular and continuing uninterrupted in oblique direction on sides; innermost striae more
narrow and inverse U-shaped. Striae on mesonotal and propodeal dorsa converging posteriorly,
terminating just short of declivity. Anterior and posterior faces of petiole transversely striate,
striae somewhat bowed dorsally. (These are well marked laterally, but somewhat obscure
medially in specimen from Gemeheng.) Dorsum of first gastral segment finely, longitudinally
striate, central striae converging anteriorly and terminating before reaching almost smooth, finely
shagreened, dorso-medial prominence.
Medium length, yellowish or golden hairs present on all body surfaces; erect or semierect on
dorsum and sides of head in front of eyes, between frontal carinae and on vertex. Antennal scape
with numerous short erect hairs along leading edge. Numerous erect, suberect and variously
curved hairs on dorsum of mesosoma, but virtually absent from sides. Petiole with pair of short
erect hairs arising on each side along dorsal margin. Gaster with numerous, relatively long hairs
along posterior margins of segments; hairs on dorsum of first gastral segment shorter and
distinctly curved posteriorly. Appressed pubescence virtually absent, with only small, silvery
patches on metapleuron, lateral and dorsal borders of propodeal declivity, sides of petiole and
coxae. Pubescence on dorsum of gaster confined to narrow, silvery band along posterior edge of
first gastral segment; anterior portion of dorsal face, with rather diluted golden pubescence.
Black. Appendages very dark reddish-brown, with only tip of apical funicular segment and
apical tarsal segments very light reddish-brown.
NEW GUINEA: Huon Penins., Mongi Watershed, Gemeheng, 1300m, 11-13.iv.1955 (E.O. Wilson
#788) (w).
REMARKS
A very distinct, virtually black species that is more-or-less regularly longitudinally striate
with the petiole transversely striate. Pubescence is virtually lacking from the body, with only a few
scattered, erect, hairs on the head, dorsum of mesonotum, propodeum and the gaster.
Polyrhachis (Aulacomyrma) retusa, new species

Figures 93, 96, 99
TYPE MATERIAL
HOLOTYPE: INDONESIA, IRIAN JAYA (New Guinea: Neth. on data label), Vogelkop,
Fak Fak, S. coast of Bomberai Pen., 02º55’S, 132º18’E, 10-100m, 3.vi.1959, T.C. Maa (worker).
Type distribution: unique holotype in MCZC.
WORKER

MTL 1.12.
Clypeus with anterior margin arcuate, weakly truncate; in profile clypeus straight anteriorly
with outline raised towards basal margin due to a short carina formed by median stria; basal
margin impressed medially, laterally indicated by break in cephalic sculpture. Frontal triangle
indistinct. Frontal carinae with raised laminate lobes. Sides of head in front of eyes converging
anteriorly towards mandibular bases; posteriorly rounding into weakly convex preoccipital
margin. Eyes strongly convex, clearly breaking cephalic outline in full face view. Pronotal humeri
armed with acute, broad-based teeth, lateral margins curving from bases onto sides, pronotal
dorsum immarginate. Promesonotal suture distinct. Metanotal groove lacking. Lateral margin of
mesonotal dorsum formed by outermost dorsal striae. Propodeum immarginate laterally and
posteriorly with dorsum descending into declivity in uninterrupted curve. Dorsal margin of petiole
with shallow emargination medially, weakly jagged laterally, lateral spines relatively long, curved
outwards, backwards and upwards. Anterior face of first gastral segment concave, with dorsal
margin of concavity acute, not elevated above dorsal face of segment.
Sculpture of head and body consisting of more-or-less regularly spaced striae; mostly
longitudinal on dorsum of head, those on vertex curving medially towards frontal carinae and
central area. Pronotal striation distinctly converging anteriorly, several median striae curving
tranversely onto pronotal collar. Mesonotal-propodeal dorsum with striae V-shaped, converging
posteriorly and terminating just before propodeal dorsum descends into declivity. Petiole with
anterior face wrinkled, transversely at base of segment and vertically on sides; posterior face of
petiole finely shagreened. First gastral segment distinctly striate laterally, striae less distinct
dorsally and becoming reticulate-punctate centrally.
Head and mesosoma with numerous, yellow, erect hairs, notably on front of head; distinctly
shorter hairs arising from vertex and mesosomal dorsum. Petiole with a few short, erect hairs on
dorsal margin. First gastral segment with a few, short, posteriorly directed hairs, more erect and
distinctly longer on apex and venter. Very sparse, pale yellow, appressed pubescence on dorsum
of head, mesosoma and gastral dorsum.
Black, including antennal scapes, femora and proximal ends of hind tibiae. Funiculi, except
apices and basal tarsal segments medium to dark reddish-brown. Tibiae, except their proximal
ends, apical segments of tarsi and last funicular segments distinctly light reddish-brown.
ETYMOLOGY
From the Latin word retusus, meaning blunted, rounded, refering to the bluntly terminated
propodeal dorsum of this species.
REMARKS
P. retusa is relatively close to porcata, however, they differ in a number of characters,

including their relative size (HL 1.12 in retusa versus 1.50-1.68 in porcata). The pronotal dorsum
in lateral view is much less convex in retusa with the pronotal teeth short and broad-based and the
petiole is narrow in profile with its dorsal margin distinctly acute. The first gastral segment of
retusa is concave, but its dorsal margin is not elevated dorsally. In contrast, the pronotal dorsum in
porcata is distinctly convex in lateral view with the humeri armed with laterally and anteriorly
directed, acute spines. The petiole, in lateral view, is scale-like, with both faces distinctly convex
and the dorsal margin less acute. The first gastral segment is concave with the dorsal margin of the
concavity produced above the face of the segment.
Polyrhachis (Aulacomyrma) roomi, new species

Figures 100, 103, 106
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA, Northern Prov., Sangara, 18.xii.1971, P.M. Room
(worker). PARATYPES: data as for holotype (4 workers). Type distribution: holotype in ANIC, 1
paratype each in BMNH, CASC, MCZC and QMBA.
WORKER

(86); SL 1.56-1.72 (1.59); SI 115-126 (119); PW 1.03-1.18 (1.06); MTL 1.53-1.68 (1.59) (5
measured).
Anterior clypeal margin distinctly emarginate medially, dentate on each side of emargination;
clypeus with weakly raised longitudinal carina, almost straight in profile; basal margin weakly
impressed and indicated laterally by faint line breaking cephalic sculpture. Frontal carinae with
strongly sinuate and distinctly raised laminate lobes. Sides of head convex. Eyes virtually flat,
situated well forward on sides of head; inner margins somewhat sunk into shallow concavity in
cephalic sculpture; not or only just reaching lateral cephalic outline in full face view. Mesosoma
with pronotal and propodeal dorsa laterally immarginate; mesonotal dorsum with outermost striae
forming short margin. Pronotal humeri armed with short, laterally and anteriorly directed,
horizontal spines. Promesonotal suture rather shallow, indicated by change in direction of
sculpture; metanotal groove lacking. Propodeal dorsum separated from declivity by faint posterior
border that, viewed from behind, forms inverse ‘V’ terminating laterally in small denticles. Dorsal
petiolar margin with pair of acute intercalary teeth medially, shallowly emarginate between them;
laterally margin somewhat jagged, lateral teeth acute, weakly upturned. Anterior face of first
gastral segment shallowly concave, narrowly rounding onto dorsum of segment.
Mandibles densely, rather finely, longitudinally striate. Dorsum of head with mostly regular
striae; longitudinal and anteriorly converging on clypeus, obliquely curving from sides of head
towards basal clypeal margin, transverse and anteriorly bowed on vertex with outermost striae
converging towards central area between frontal carinae. Pronotal dorsum mostly regularly,
transversely striate; striae curving vertically onto sides. Striae on mesonotal-propodeal dorsum
transverse, bowed posteriorly into open U-shapes; sides with striae more-or-less horizontal.
Petiole and gaster finely shagreened.
Dorsum of head and mesosoma with medium length, mostly erect, off-white or yellowish
hairs; hairs somewhat posteriorly curved and golden on gastral dorsum, distinctly longer along
sternites and on gastral apex. Appressed pubescence much reduced on head and mesosomal
dorsum, only patches of sparse, off-white or silvery pubescence on sides of propodeum, along
propodeal declivity, petiole and coxae. Golden and rather dense gastral pubecence mostly
confined to dorsum of first segment, not completely concealing underlying sculpture.
Black; most of antennal scapes, basal half of first funicular segment, joints of femora and
tibiae, bases of basal tarsal segments and gaster dark to very dark reddish-brown. Mandibular
masticatory border, basal and distal ends of antennal scapes, funiculi and most of legs light to very
light reddish-yellow.
ETYMOLOGY
Named in honour of the collector, Dr Peter M. Room of CSIRO, Division of Entomology,

Indooroopilly, Queensland, Australia, who collected scores of ants, many new to science, during
the years of his appointment as an agricultural entomologist in Papua New Guinea.
REMARKS
This species is easily recognisable by its relatively large size.
Polyrhachis (Aulacomyrma) snellingi, new species

Figures 101, 104, 107
TYPE MATERIAL
HOLOTYPE: INDONESIA, IRIAN JAYA, PT. Freeport Concession, Siewa camp, 03.04ºS,
136.38ºE, 200ft, 18-30.iv.1998, lowland secondary rf., Malaise trap, R.R. Snelling #16 (worker).
Type distribution: unique holotype in LACM.
WORKER

MTL 1.15.
Anterior clypeal margin with shallow notch medially; in profile clypeus virtually straight with
only very weakly impressed basal margin. Frontal carinae sinuate with laminate lobes. Sides of
head weakly convex, gently converging anteriorly, curving behind eyes into relatively shallow
preoccipital margin. Eyes convex, in full face view clearly breaking lateral cephalic outline.
Mesosomal dorsum only partially marginate. Pronotal humeri armed with very strong, anteriorly
directed, broad-based spines, with raised margins, central areas distinctly concave; lateral margins
of spines extended posteriorly for some distance, merging with outermost dorsal striae and
curving onto sides, not reaching distinct promesonotal suture. Mesonotal dorsum marginate;
propodeum poorly margined, outermost striae delimiting lateral edges of dorsum and terminating
in downward and weakly inward curved angles. Propodeal dorsum descending into declivity in
even, uninterrupted curve. Petiole with both faces convex, dorsal margin acute; lateral spines
relatively short, acute, curved weakly backwards and upwards. First gastral segment with anterior
face concave; dorsal margin of concavity sharp, not elevated above dorsal face of segment.
Mandibles finely, mostly longitudinally, striate. Head, including clypeus, distinctly striate;
striae mostly longitudinal on front of head and vertex, somewhat oblique on sides. Dorsum of
mesosoma distinctly longitudinally striate, striae converging anteriorly; dorsal surfaces of
pronotal spines not striate, finely shagreened. Mesonotal-propodeal dorsum longitudinally striate,
striae converging posteriorly towards declivity. Petiole with anterior face transversely striate;
posterior face with fan-shaped striae running from apex of dorsal margin towards sides and base of
segment. Anterior face of first gastral segment with transverse striae that are continued posteriorly
along sides and towards dorsum; dorsal surface more finely, longitudinally, striate.
Numerous erect or semierect, yellowish or off-white hairs, shorter than greatest diameter of
eye, present on most body surfaces. Mostly greyish or silvery, appressed or semierect pubescence
present in various densities on most body surfaces, sparse on head, rather abundant on pronotal
dorsum and mostly confined to interspaces between striae and pronotal spines. Propodeal dorsum
with pubescence relatively long and distincly curved towards midline. Gastral dorsum with rather
dense pubescence partly obscuring sculpture.
Black, mandibular masticatory border very narrowly and funiculi towards apices, very dark
reddish-brown. Legs generally medium reddish-brown, apices of mid and hind femora, tibiae and
first tarsal segments, very dark reddish-brown.
ETYMOLOGY
Named in honor of its collector, Roy R. Snelling of the Natural History Museum of Los
Angeles County.
REMARKS
P. snellingi is a very distinctive species. It is the only known member of the subgenus having
the lateral striation of the first gastral segment extending transversely across its anterior face.
Polyrhachis (Aulacomyrma) wamuki, new species

Figures 102, 105, 108
TYPE MATERIAL
HOLOTYPE: PAPUA NEW GUINEA: Morobe Prov., Huon Penins., Mongi Watershed,
Wamuki (= Wamuri), 06º32’S, 147º30’E, 800m, 19-20.iv.1955, E.O. Wilson #847 (worker).
PARATYPE: data as for holotype (1 worker). Type distribution: holotype in MCZC;

paratype in QMBA.
WORKER
Dimensions (holotype cited first): TL c. 4.38, 4.23; HL 1.15, 1.12; HW 1.04, 1.00; CI 90, 89;
SL 1.22, 1.15; SI 117, 115; PW 0.84, 0.81; MTL 1.09, 1.06 (2 measured).
Anterior clypeal margin arcuate, shallowly emarginate medially; in profile clypeus virtually
straight anteriorly, with very weakly impressed basal margin. Frontal carinae sinuate, with
laminate lobes. Sides of head in front of eyes almost straight, anteriorly converging. Eyes
moderately convex, breaking cephalic outline in full face view. Mesosomal dorsum virtually
parallel-sided. Pronotal dorsum immarginate; humeri armed with acute, broad-based teeth with
weakly raised margins. Promesonotal suture distinct; metanotal groove lacking.
Mesosomal-propodeal dorsum with ill-defined lateral margins formed by outer-most dorsal striae.
Propodeum terminating in distinct, rounded prominences, their posterior margins continued
inwards for some distance, forming short, medially unconnected ridges, partly separating dorsum
from shallowly concave declivity. Dorsal petiolar margin sinuate in dorsal view, somewhat jagged
medially and towards sides, with relatively long, acute lateral spines, directed laterally and weakly
curved backwards. First gastral segment with anterior face concave; anterodorsal margin of
concavity rather sharp and moderately elevated above dorsal face of segment.
Mandibles finely longitudinally striate; striae overlaid by shallow punctures towards
masticatory margin. Head, including clypeus, distinctly, mostly regularly, longitudinally striate;
striae on sides of head and vertex weakly curved and converging between frontal carinae.
Mesosomal dorsum with regular striae; inversely V-shaped on pronotal dorsum, converging
anteriorly and medially and continued obliquely onto sides; mesonotal-propodeal dorsum with
striae curving posteriorly and inwards towards declivity, where they terminate and form an
ill-defined blunt margin. Petiole shagreened. First gastral segment very distinctly longitudinally
striate laterally; striae much less distinct dorsally, median area behind anterodorsal margin
shagreened.
Mostly golden, rather short hairs dispersed over most body surfaces. Appressed, silvery
pubescence rather sparse on head and mesosoma; more abundant, mostly golden pubescence on
gastral dorsum.
Black, with antennal scapes and legs at their joints, dark reddish-brown. Mandibular
masticatory border, funiculi and most of legs yellow or relatively light, yellowish-brown with
reddish tint.
ETYMOLOGY
Named after the type locality, Wamuki village on Huon Peninsula, Papua New Guinea.
REMARKS
P. wamuki is characterised by its small size, virtually parallel-sided mesosomal dorsum and
its wide, strongly transverse, petiolar node.
ACKNOWLEDGEMENTS
I am very grateful to Steve O. Shattuck (ANIC) and Stefan P. Cover (MCZC), for unlimited access
to the collections in their care. My deepest appreciation goes to the Ernst Mayr Grant committee of
Harvard University for this prestitious award that was bestowed upon me twice. Without their
support which allowed me to study the Polyrhachis material in the collection of the Museum of
Comparative Zoology, this study could not have been accomplished. I am also very much indebted
to Brian L. Fisher (CASC), Steve O. Shattuck, Natalie Barnett (ANIC) and April Nobile (CASC)
for their help in obtaining digital images of the specimens. My sincere thanks are due to the
curators, researchers and other persons in charge of the collections in numerous museums and
institutions named above, for the loan of types and other valuable specimens. My gratitude must
finally be extended to Chris Burwell (QMBA), for his support during preparation of this paper and
for reading and commenting on a draft of the manuscript.
LITERATURE CITED
Bolton, B.. 1994. Identification guide to the ant genera of the world. Harvard University Press,
Bolton, B. 1995. A new general catalogue of the ants of the world. Harvard University Press,
Chapman, J.W. & Capco, S.R. 1951. Check list of the ants (Hymenoptera: Formicidae) of Asia.
Monographs of the Institute of Science and Technology, Manila 1: 327pp.
Donisthorpe, H. 1932. On the identity of Smith’s types of Formicidae collected by Alfred Russell
Wallace in the Malay Archipelago, with descriptions of two new species. Annals and
Magazine of Natural History (10) 10: 441-476.
Donisthorpe, H. 1948. A third instalment of the Ross Collection of ants from New Guinea.
Annals and Magazine of Natural History (11) 14 (1947): 589-604.
Dorow, W.H.O. 1995. Revision of the ant genus Polyrhachis Smith, 1857 (Hymenoptera:
Formicidae: Formicinae) on subgenus level with keys, checklist of species and bibliography.
Courier Forschungsinstitut Senckenberg 185: 1-113.
Emery, C. 1887. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova.
Parte terza. Formiche della regione Indo-Malese e dell’ Australia. Annali del Museo Civico di
Storia Naturale di Genova (2)4[24] (1886): 209-258.
Emery, C. 1921. Le genre Polyrhachis. Classification; espèces nouvelles ou critiques. Bulletin de
la Société Vaudoise des Sciences Naturelles 54: 17-25.
Formicinae. Fasc. 183:302 pp. Bruxelles.
Forel, A. 1901. Formiciden aus dem Bismarck-Archipel, auf Grundlage des von Prof. Dr. F. Dahl
gesammelten Materials bearbeitet. Mitteilungen aus dem Zoologischen Museum in Berlin 2:
1-38.
Hung, A.C.F. 1967. A revision of the ant genus Polyrhachis at subgeneric level. Transactions of
the American Entomological Society 93: 395-422.
International Commission on Zoological Nomenclature 1999. International Code of
Zoological Nomenclature (Fourth Edition). London. 306pp.
Karavaiev, W. 1927. Ameisen aus dem Indo-Australischen Gebiet. III. Académie des Sciences
de l’Ukraïne. Mémoires de la Classe des Sciences Physiques et Mathématiques (Travaux du
Musée Zoologique 3). Kiev 7(1): 3-52.
Kohout, R.J. 1990. A review of the Polyrhachis viehmeyeri species-group (Hymenoptera:
Formicidae: Formicinae). Memoirs of the Queensland Museum 28(2): 499-508.
Kohout, R.J. 1998. New synonyms and nomenclatural changes in the ant genus Polyrhachis Fr.
Smith (Hymenoptera: Formicidae: Formicinae). Memoirs of the Queensland Museum 42(2):
505-531.
Smith, F. 1857. Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount
Ophir, Malacca; and at Singapore, by R.A. Wallace. Journal of the Proceedingsof the
Linnean Society of London, Zoology 2:42-88.
Smith, F. 1859. Catalogue of Hymenopterous insects collected by Mr. A.R. Wallace at the Islands
of Aru and Key. Journal of the Proceedings of the Linnean Society of London, Zoology 3:
132-158.
Viehmeyer, H. 1912. Ameisen aus Deutsch Neuguinea gesammelt von Dr. O. Schlaginhaufen.
Nebst einem Verzeichnisse der papuanischen Arten. Abhandlungen und Berichte des Königl.
Zoologischen und Anthropologisch-Ethnographischen Museums zu Dresden 14 :1-26.
Figures 1-3, head in full face view: 1, P. abdita; 2, dohrni; 3, exarata. Figures 4-6, dorsal view of
mesosoma and petiole: 4, P. abdita; 5, dohrni; 6, exarata. Figures 7-9, Lateral view of mesosoma
and petiole: 7, P. abdita; 8, dohrni; 9, exarata.
Figures 10-12, Head in full face view: 10, P. gentilis; 11, lumi; 12, minima. Figures 13-15, Dorsal
view of mesosoma and petiole: 13, P. gentilis; 14, lumi; 15, minima. Figures 16-18, Lateral view
of mesosoma and petiole: 16, P. gentilis; 17, lumi; 18, minima.
Figures 19-21, Head in full face view: 19, P. pallipes; 20,parva; 21, planata. Figures 22-24,
Dorsal view of mesosoma and petiole: 22, P. pallipes; 23, parva; 24, planata. Figures 25-27,
Lateral view of mesosoma and petiole: 25, P. pallipes; 26, parva; 27, planata.
Figures 28-30, Head in full face view: 28, P. roberti; 29, sculpta; 30, simulans. Figures 31-33,
Dorsal view of mesosoma and petiole: 31, P. roberti; 32, sculpta; 33, simulans. Figures 34-36,
Lateral view of mesosoma and petiole: 34, P. roberti; 35, sculpta; 36, simulans.
Figures 37-39, Head in full face view: 37, P. sulcifera; 38, umboi; 39, variegata. Figures 40-42,
Dorsal view of mesosoma and petiole: 40, P. sulcifera; 41,umboi; 42, variegata. Figures 43-45,
Lateral view of mesosoma and petiole: 43, P. sulcifera; 44, umboi; 45, variegata.
Figures 46-48, Head in full face view: 46, P. wardi; 47, wilsoni; 48, annulata. Figures 49-51.
Dorsal view of mesosoma and petiole: 49, P. wardi; 50, wilsoni; 51, annulata. Figures 52-54,
Lateral view of mesosoma and petiole: 52, P. wardi; 53, wilsoni; 54, annulata.
Figures 55-57, Head in full face view: 55, P. bedeloweryi; 56, breviata; 57, decora. Figures 58-
60, Dorsal view of mesosoma and petiole: 58, P. bedeloweryi; 59, breviata; 60, decora. Figures
61-63, Lateral view of mesosoma and petiole: 61, P. bedeloweryi; 62, breviata; 63, decora.
Figures 54-66, Head in full face view: 64, P. excellens; 65, geometrica; 66, gressitti. Figures 67-
69, Dorsal view of mesosoma and petiole: 67, P. excellens; 68, geometrica; 69, gressitti. Figures
70-72, Lateral view of mesosoma and petiole: 70, P. excellens; 71, geometrica; 72, gressitti.
Figures 73-75, Head in full face view: 73, P. impressa; 74, kokoda; 75, maai. Figures 76-78,
Dorsal view of mesosoma and petiole: 76, P. impressa; 77, kokoda; 78, maai. Figures 79-81,
Lateral view of mesosoma and petiole: 79, P. impressa; 80, kokoda; 81, maai.
Figures 82-84, Head in full face view: 82, P. mamba; 83, monticola; 84, orokana. Figures 85-87,
Dorsal view of mesosoma and petiole: 85, P. mamba; 86, monticola; 87, orokana. Figures 88-90,
Lateral view of mesosoma and petiole: 88, P. mamba; 89, monticola; 90, orokana.
Figures 91-93, Head in full face view: 91, P. pelecta; 92, porcata; 93, retusa. Figures 94-96,
Dorsal view of mesosoma and petiole: 94, P. pelecta; 95, porcata; 96, retusa. Figures 97-99,
Lateral view of mesosoma and petiole: 97, P. pelecta; 98, porcata; 99, retusa.
Figures 100-102, Head in full face view: 100, P. roomi; 101, snellingi; 102, wamuki. Figures
103-105, Dorsal view of mesosoma and petiole: 103, P. roomi; 104, snellingi; 105, wamuki.
Figures 106-108, Lateral view of mesosoma and petiole: 106, P. roomi; 107, snellingi; 108,
wamuki.
Lattke, J. E., Fernández, F., Palacio, E. E. 2007. Identification of the species of Gnamptogenys
Roger in the Americas, pp. 254-270. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances
in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
IDENTIFICATION OF THE SPECIES OF GNAMPTOGENYS ROGER

IN THE AMERICAS
John E. Lattke
Instituto de Zoología Agrícola
Universidad Central de Venezuela
Apartado 4579
Maracay 2101-A, Venezuela
piquihuye@hotmail.com
Fernando Fernández
Apartado 7495
ffernandez@unal.edu.co
and
Edgard E. Palacio
Fundación Nova Hylaea
Apartado Aéreo 7954
Santafé de Bogotá D.C., Colombia
parasitoideus@yahoo.com
ABSTRACT
A dichotomous key is presented for identifying workers of recent species of the genus
Gnamptogenys in the Americas. Included are 81 species with their known distribution. G. wilsoni
Lattke, n. sp., is described.
Key words: Hymenoptera, Formicidae, Gnamptogenys, taxonomy, identification key, Americas.
Resumen: Se presenta una clave dicotómica para identificar a las especies recientes de obreras del
género Gnamptogenys en América. Se incluyen 81 especies con su distribución conocida. Se
describe G. wilsoni Lattke, n. sp.
Palabras claves: Hymenoptera, Formicidae, Gnamptogenys, taxonomía, clave para identificación,

América
Lattke, et al.: Identification of American Gnamptogenys 255
INTRODUCTION
Ants, in accordance with their ecological importance, numerical dominance, and considerable
diversity are becoming ever more present in studies as biological indicators (Alonso, 2000). The
genus Gnamptogenys is one of the larger groups with more than 120 species worldwide, of which
over 80 are known from the New World. It ranks fifth place amongst the poneromorph subfamilies
in number of species (Bolton, 1995) and tenth place amongst the ant genera recovered in Neotropical
leaf litter samples (Ward, 2000). Most species inhabit mesic forested areas, nesting in decomposing
wood on the ground and hunting their prey on the soil and amongst the leaf litter. The New World
fauna was recently revised by Lattke (1995) but subsequent work has increased the number of
species, and some new synonyms have been determined (Lattke, 2002; Lattke et al., 2004; Pacheco
et al., 2004). Given these changes, a new version of the key for identifying the species is presented.
The results should be confronted with descriptions in order to support the conclusions. The unnamed
taxa ¨species A, B, and C¨ are included in the key but not in the species list, and will be described in
a future paper.
LIST OF RECENT NEW WORLD GNAMPTOGENYS SPECIES
G. acuminata Emery: Tropical South America

G. acuta (Brown): Colombia to Bolivia
G. alfaroi Emery: Costa Rica to Ecuador
G. ammophila Lattke: southeastern Venezuela
G. andina Lattke: Colombia - Ecuador
G. annulata Mayr: Costa Rica to Tropical South America
G. banksi (Wheeler): Panama to Ecuador
G. biquetra Lattke: Panama
G. bispinosa (Emery): Costa Rica to Colombia
G. bisulca Kempf and Brown: Costa Rica to Ecuador, Andes
G. boliviensis Lattke: northern Bolivia
G. bruchi (Santschi): northern Argentina
G. brunnea Lattke: western Colombia
G. bufonis (Mann): southern Mexico - Nicaragua
G. caelata Kempf: Colombia - southern Brazil
G. concinna (F. Smith): Mexico to Tropical South America
G. continua Mayr: Mexico to Tropical South America
G. cuneiforma Lattke: Panama
G. curvoclypeata Lattke: Venezuela to Colombia
G. ejuncida Lattke: Colombia
G. enodis Fernández Colombia
G. ericae Forel: northern South America
G. extra Lattke: western Colombia - Ecuador
G. falcaria Lattke: Hispaniola
G. falcifera Kempf: northern South America
G. fernandezi Lattke: Venezuela to Ecuador
G. fieldi Lattke: north-central Venezuela
G. flava Pacheco, MacKay & Morgan: Bolivia
G. gentryii Lattke: southwestern Colombia
G. gracilis (Santschi): Guyana Shield
G. haenschi Emery: Costa Rica - Tropical South America
G. hartmanni (Wheeler): southern U.S. - northern South America
G. haytiana (Wheeler & Mann): Hispaniola

G. horni Santschi: Panama to Bolivia
G. ilimani Lattke: Bolivian Andes
G. ingeborgae Brown: Colombia
G. insularis Lattke: Hispaniola
G. interrupta Mayr: Mexico to Tropical South America, Jamaica
G. kempfi Lenko: Brazil & Peru
G. lanei Kempf: lower Amazon watershed
G. laticephala Lattke: Ecuador
G. lavra Lattke: SE Brazil
G. lineolata Brown: Hispaniola
G. lucaris Kempf: southern Brazil
G. mecotyle Brown: Panama - Tropical South America
G. mediatrix Brown: Cuenca Amazónica
G. menozzii Borgmeier: southern Brazil
G. mina (Brown): northern South America
G. minuta (Emery): Belize - Brazil
G. moelleri (Forel): Tropical South America
G. mordax (F. Smith): Mexico to Tropical South America
G. nana Kempf: Brazil & northern Argentina
G. nigrivitrea Lattke: Colombia
G. perspicax Kempf & Brown: Colombia to Ecuador
G. petiscapa Lattke: northwestern Venezuela
G. pilosa Lattke: western Colombia
G. pittieri Lattke: north central Venezuela
G. pleurodon (Emery): Tropical South America
G. porcata (Emery): Honduras to Bolivia
G. rastrata (Mayr): southern Brazil
G. regularis Mayr: Mexico to Paraguay
G. reichenspergeri (Santschi): Orinoco - Amazon watershed
G. relicta (Mann): Orinoco - Amazon watershed
G. rimulosa (Roger): southern Brazil
G. rumba Lattke: Cuba
G. schmitti (Forel): Hispaniola
G. semiferox Brown: Hispaniola
G. siapensis Lattke: southern Venezuela
G. simulans (Emery): Costa Rica
G. stellae Lattke: Costa Rica, Colombia
G. striatula Mayr: Mexico to northern Argentina, Caribbean
G. strigata (Norton): Mexico to Colombia
G. striolata (Borgmeier): southwestern Brazil
G. sulcata (F. Smith): Mexico to tropical South America
G. tortuolosa (F. Smith): Amazon - Orinoco watershed
G. transversa Lattke: Panama
G. triangularis Mayr: Florida, USA.; Costa Rica to Argentina
G. vriesi Brandão & Lattke: Ecuador
G. wilsoni Lattke, n. sp: southeastern Brazil
KEY TO THE NEW WORLD SPECIES OF GNAMPTOGENYS (workers)

1 Frontal carina not covering all of base of scape, at least neck visible; petiolar spiracle
facing ventrolaterally, not in depression; promesonotal suture well marked, totally
interrupting dorsal mesosomal sculpture ......................................................................... 2
-- Frontal carina not totally covering base of scape, at least neck visible; petiolar spiracle
facing ventrolaterally, not in depression; promesonotal suture absent to well-impressed but
never interrupting dorsal mesosomal sculpture.............................................................. 25
-- Frontal carina in dorsal view covering all of scape base, including condyle and neck;
petiolar spiracle facing ventrally, located in depression; promesonotal suture absent to
lightly impressed, never interrupting dorsal mesosomal sculpture ................................ 75
2(1) Petiolar node with semiparallel anterior and posterior margins in lateral view, both forming
approximate right angles with the dorsal margin (Fig. 1); cephalic vertex usually smooth
and shining, sometimes with weak transverse striae........................................................ 3
-- Petiolar node with a single curved anterodorsal margin or anterior and dorsal margins
separated by a blunt angle (Figs. 2-4); cephalic vertex usually with sculpturing ........... 6
3(2) Mesosomal dorsum with large smooth and shining areas................................................ 4

-- Mesosomal dorsum totally sculptured, usually costulate or rugulose .............................. 5
4(3) Metanotal groove well-impressed, interrupting dorsal mesosomal sculpture; metacoxal

dorsum unarmed; propodeal declivity lacking lobes................................................relicta
-- Metanotal groove absent; metacoxal dorsum with a lobe or denticle; propodeal declivity
with anterolateral lobes............................................................................. reichenspergeri
5(3) Postpetiolar dorsum transversely rugulose; body yellowish ................................ haytiana

-- Postpetiolar dorsum longitudinally costulate; body black to dark brown .................. mina
6(2) Propodeal spiracle separated from declivitous margin by more than one diameter in lateral
view; metacoxal dorsum unarmed .............................................................................lavra
-- Propodeal spiracle separated from declivitous margin by less than one diameter; metacoxal
dorsum with a lobe or denticles ....................................................................................... 7
7(6) Mesosomal dorsum with a well defined metanotal groove .............................................. 8

-- Metanotal groove absent................................................................................................ 12
8(7) Dorsum of abdominal tergites III and IV smooth and shining ................................gentryi
Dorsum of abdominal tergites III and IV longitudinally costulate.................................. 9
9(8) Subpetiolar process subquadrate in lateral view, with a posterior angle........................ 10

-- Subpetiolar process forming an anterior projecting lobe, without a posterior angle...... 11
10(9) Area between promesonotal suture and metanotal groove narrow and of uniform width;
anterior mesonotal margin convex..........................................................................bisulca
-- Area between promesonotal suture and metanotal groove wide and ovaloid; anterior
mesonotal margin forms a blunt angle ............................................................... species A
11(9) Anterior margin of clypeal lamella bluntly angular; cephalic vertex longitudinally costulate
............................................................................................................................. brunnea
-- Anterior margin of clypeal lamella evenly convex; cephalic vertex with less than five
transverse costulae ............................................................................................. species B
12(7) Petiolar node erect in lateral view, the dorsal margin separated from the anterior and
posterior margins by blunt angles (Fig. 2) ..................................................................... 13
-- Petiolar node posteriorly inclined in lateral view, with a broadly curved anterodorsal
margin contrasting with the angle separating the dorsal margin from the posterior margin
(Figs. 3-4) ...................................................................................................................... 15
13(12) SL < 1.1 mm .................................................................................................................. 14

-- SL> 1.1 mm .......................................................................................................... moelleri
14(13) Propodeal declivity with 5-11, sometimes fewer, longitudinal costulae between spiracles
and 1-3 transverse costulae between each spiracle and the longitudinal costulae (Fig. 6);
scapes with sparse pilosity, less than six decumbent and semi-erect hairs ...........striatula
-- Propodeal declivity with 13-16 longitudinal costulae between spiracles and no transverse
costulae between each spiracle and the longitudinal costulae (Fig. 5); scapes with dense,
uniformly decumbent pilosity, more than six decumbent and semi-erect hairs
......................................................................................................................... ammophila
15(12) Petiolar node with blunt or pointed apex overhanging posterior margin in lateral view;
subpetiolar process in ventral view with edge of mostly uniform width, widening only at
union with sternite (Fig. 7) ............................................................................................ 16
-- Petiolar node with apex barely overhanging posterior margin; subpetiolar process in ventral
view cuneiform, anteriorly of uniform width, widening posteriorly into a bifurcate process
(Fig. 8) ........................................................................................................................... 22
16(15) Petiolar node with a blunt apex in lateral view .............................................................. 17

-- Petiolar node with an acute apex .............................................................................. acuta
17(16) Postpetiolar sternite with a median smooth and shining area; vertex usually with 4 - 5
transverse costulae next to vertexal carina (Fig.9)................................................. gracilis
-- Postpetiolar sternite totally costulate or striate; cephalic vertex usually with a single,
occasionally more than three, transverse costulae next to vertexal carina (Fig. 10) ...... 18
18(17) Scape in frontal view with more than 10 erect hairs along its posterior margin, not
including basal pilosity (Fig. 12) ................................................................................... 19
-- Scape in frontal view with fewer than 10 erect hairs along its posterior margin (Fig. 11)
....................................................................................................................................... 20
19(18) Body and legs brown; body striate or costulate, with abundant decumbent pubescence;
subpetiolar process forming a rounded lobe, without sharp angles in lateral view ...........
............................................................................................................................. ejuncida
-- Body black and legs ferruginous; body usually costate, with sparse decumbent pubescence;
subpetiolar process usually with a sharp anterior angle, blunt in Central American
specimens ..............................................................................................................porcata
20(18) Tibiae and scapes with 5 or more erect hairs ................................................................. 21

-- Tibiae and scapes with 1 or no semi-erect or semi-decumbent hairs ......................... extra
21(20) Propodeal declivity with transverse costulae; petiolar node slightly pedunculate (Fig. 4)
...........................................................................................................................pleurodon
-- Propodeal declivity with longitudinal costulae; petiolar node sessile (Fig. 13) .................
.......................................................................................................................... nigrivitrea
22(15) Postpetiolar sternite with well-defined costulae or striae; propodeal dorsum not depressed
below level of mesonotum in lateral view; dorsal and declivitous propodeal faces confluent
in lateral view ................................................................................................................ 23
-- Postpetiolar sternite with weakly-defined rugulae; propodeal dorsum depressed below level
of mesonotum in lateral view; dorsal and declivitous propodeal faces separated by ridge in
lateral view ............................................................................................................ pittieri
23(22) Scape with moderate to scarce pubescence, with 5 or more erect hairs........................ 24
-- Scape with dense white pubescence, with 0 - 2 erect hairs.....................................ilimani
24(23) HW > 0.84 mm, WL > 0.35 mm; OI < 0.16 ........................................................... andina
-- HW < 0.84 mm, WL < 0.35 mm; OI > 0.16 ..........................................................strigata
25(1) Scape not reaching the preoccipital margin in dorsal view............................................ 26

-- Scape surpassing preoccipital margin by at least one apical width................................ 42
26(26) Mandibular dorsum with striae, costulae or rugulae on at least one-fourth of its length ...
....................................................................................................................................... 27
-- Mandibular dorsum totally smooth and shining, with sparse punctures ........................ 29
27(26) Body with regular striae or costulae; mandibles triangular or semi-triangular; not found in
Antilles .......................................................................................................................... 28
-- Body with irregular costulae; mandibles falcate; Antillean (Hispaniola) ............. falcaria
28(27) Metacoxal dorsum unarmed; eye small (OI < 0.1); mandibular dorsal surface densely striate
............................................................................................................... haenschi (in part)
-- Metacoxal dorsum with lobe; eye large (OI > 0.2); mandibular dorsal surface costulate on
basal fourth or less, the rest smooth and shining .................................hartmanni (in part)
29(26) Propodeal dorsum mostly transversely striate................................................................ 30

-- Propodeal dorsum mostly longitudinally striate or costulate ......................................... 32
30(29) Head subquadrate in dorsal view (CI > 0.78); body brown to dark brown .................... 31
-- Head rectangular, (CI < 0.78); body black ............................................................. alfaroi
31(30) Mesosoma and petiole costulate; both propodeum and metacoxal dorsum edentate .........
.................................................................................................................................kempfi
-- Mesosoma and petiole finely striate; both propodeum and metacoxal dorsum dentate .....
............................................................................................................................. annulata
32(29) Clypeal lamella laterally rounded in dorsal view, never angular; subpetiolar process usually
rounded in lateral view, rarely with a posterior tooth .................................................... 33
-- Clypeal lamella with sharp lateral angles; subpetiolar process with a posterior tooth in
lateral view .................................................................................................................... 39
33(32) Mandible semi-triangular to semi-falcate in dorsal view; metanotal groove well-defined

....................................................................................................................................... 34
-- Mandible triangular; metanotal groove absent........................................................... nana
34(33) Metacoxal dorsum unarmed, at most with a low tubercle or swelling ........................... 35
-- Metacoxal dorsum with a lobe or tooth ......................................................................... 36
35(34) Body striate; clypeal lamella laterally with blunt angles in dorsal view, medially anteriorly
projecting and with a small median concavity .........................................................stellae
-- Body costulate; clypeal lamella with a broad median concavity and laterally rounded in
dorsal view........................................................................................................... simulans
36(34) Metacoxal dorsum with a low triangular lobe; HW < 1.12 mm, WL < 1.81 mm .......... 37
-- Metacoxal dorsum with a slender and parallel-sided lobe or tooth; HW > 1.12,
WL > 1.81 mm....................................................................................................... mordax
37(36) Propodeal spiracle separated from declivity by less than its diameter in lateral view .......
................................................................................................................................wilsoni
-- Propodeal spiracle separated from declivity by more than its diameter......................... 38
38(37) Cephalic dorsum striate; propodeal declivity inclined and relatively straight in lateral view;
with weakly developed lateral lobes ................................................................. boliviensis
-- Cephalic dorsum costulate; propodeal declivity with a posterior concavity formed by well-
developed anterolateral lobes............................................................................... continua
39(32) Metanotal groove well-defined, visible in any angle; subpetiolar process without a posterior
tooth in lateral view ....................................................................................................... 40
-- Metanotal groove weakly defined, visible only in limited angles; subpetiolar process with
posterior tooth in lateral view ........................................................................................ 41
40(39) Anterior margin of clypeal lamella straight in dorsal view; metacoxal dorsum with a lobe
............................................................................................................................. rimulosa
-- Anterior margin of clypeal lamella with a bluntly angular concavity; metacoxal dorsum
unarmed ................................................................................................................biquetra
41(39) Propodeal declivity with longitudinal costulae that converge posteriorly; anterior face of
petiolar node transversely costulate; mesometapleural suture absent or weakly impressed
...................................................................................................................................horni
-- Propodeal declivity with parallel longitudinal costulae; most of anterior face of petiolar
node longitudinally costulate, but frequently with a basal strip of transverse costulae that
occasionally covers all of anterior face; mesometapleural suture well-impressed.............
.............................................................................................................................regularis
42(25) Mandible elongate, falcate or semi-falcate in dorsal view, with just the apex touching or
overlapping the opposite mandible when totally closed ................................................ 43
-- Mandible triangular or semi-triangular in dorsal view, with a fourth or more of the
masticatory border overlapping the opposite mandible when closed............................. 48
43(42) Propodeal declivity with teeth or denticles; metacoxal dorsum with tooth; mostly
continental species ......................................................................................................... 44
-- Propodeal declivity rounded, without teeth or denticles; metacoxal dorsum unarmed;
endemic to Hispaniola ................................................................................................... 47
-- Propodeal declivity with teeth or denticles; metacoxal dorsum unarmed; Cuban endemic
.................................................................................................................................rumba
44(43) Mandible with short triangular tooth on basal internal margin ...................................... 45
-- Mandible with a lobe or broad convexity on basal internal margin .................... mediatrix
45(44) Clypeus with a lobe anterior to each antennal fossa; each lobe partially covers the clypeal
lamella in dorsal view; mesonotal spiracle in depression below level of surrounding
integument ....................................................................................................... laticephala
-- Clypeus without lobes between antennal fossae and lamella; mesonotal spiracles at same
level as surrounding integument .................................................................................... 46
46(45) Antennal scape surpasses posterior cephalic margin by at least twice its maximum width;
preoccipital lamella visible, with a median convexity in cephalic dorsal view..................
............................................................................................................................species C
-- Antennal scape surpasses posterior cephalic margin by no more than one maximum
diameter ...................................................................................................................banksi
47(43) Mandible elongate and slender; propodeal dorsal surface with smooth transverse costulate
...............................................................................................................................schmitti
-- Mandible shorter and thicker; propodeal dorsal surface with vermiculate longitudinal
costulae ...............................................................................................................semiferox
48(42) Mandibular dorsal surface mostly smooth and shining, sometimes with striae or costulae
extending not beyond the basal third; mesosomal dorsum usually without transverse
sutures, if otherwise, then the ant is ferruginous and finely striate ................................ 49
-- Mandibular dorsal surface mostly striate or rugulose; metanotal groove well-impressed .
....................................................................................................................................... 62
49(48) HW > 1.4 mm; WL > 2.3 mm........................................................................................ 50

-- HW < 1.4 mm; WL < 2.3 mm........................................................................................ 51
50(49) Mesosoma costulate; metacoxal dorsum with dorsal tooth; apex of petiolar node ending in a
blunt angle in lateral view; body black ..............................................................tortuolosa
-- Mesosoma striate; metacoxal dorsum unarmed; apex of petiolar node ending in a blunt
point; body ferruginous........................................................................................concinna
51(49) Mesosoma striate; propodeal declivity with lateral longitudinal crests distinct from
mesosomal striae............................................................................................................ 52
-- Mesosoma costulate; propodeal declivity without crests distinct from mesosomal striae
....................................................................................................................................... 53
52(51) Posterior face of petiolar node longitudinally costulate.......................hartmanni (in part)

-- Posterior face of petiolar node transversely costulate..............................................bruchi
53(51) Posterior face of petiolar node longitudinally costulate................................................. 54

-- Posterior face of petiolar node transversely costulate.................................................... 58
54(53) Clypeal lamella laterally angular; mandible semi-triangular, the internal and masticatory
margins united by a broad convexity ............................................................................. 55
-- Clypeal lamella laterally rounded; mandible triangular, the internal and masticatory
margins separated by an angle ....................................................................................... 57
55(54) Petiolar node with a pointed apex in lateral view; propodeal declivity longitudinally
costulate; body uniform brown ......................................................................... acuminata
-- Petiolar node sometimes with a posterior projection but never pointed; propodeal declivity
usually transversely costulate, rarely longitudinal; color variable, usually with both brown
and ferruginous parts, rarely totally brown .................................................................... 56
56(55) Metacoxal tooth absent or vestigial; propodeal declivity without lateral lobes; body dark
brown to bicolored..................................................................................................sulcata
-- Small metacoxal tooth present; propodeum with small lateral lobes; body yellow ... flava
57(54) Metacoxal dorsum with a lobe; clypeal lamella with two convexities; petiolar node with
dorsal and posterior margins forming approximately a right angle in lateral view ............
......................................................................................................................... transversa
-- Metacoxal dorsum unarmed; clypeal lamella with a single convexity; petiolar node with an
apical point that overhangs the posterior margin .............................................. fernandezi
58(53) Clypeal lamella laterally sharply angular....................................................................... 59

-- Clypeal lamella laterally rounded or obtusely angular................................................... 60
59(58) HW < 1.1 mm; WL < 1.7 mm; clypeal lamella with a straight anterior margin ..... ericae
-- HW > 1.1 mm; WL > 1.7 mm; clypeal lamella medially convex, laterally concave .........
.....................................................................................................................curvoclypeata
60(58) Propodeal declivity longitudinally costulate; clypeal lamella straight medially and rounded
laterally in frontal view; mandible semi-triangular, the internal and masticatory margins
united by a broad convexity........................................................................................... 61
-- Propodeal declivity transversely costulate; clypeal lamella medially concave, laterally
convex laterally; mandible triangular, the internal and masticatory margins separated by an
angle ......................................................................................................................volcano
61(60) Petiolar node with a more or less convex dorsal margin in lateral view; metacoxal dorsum
with an acute tooth; subpetiolar process with sharp angles ................................... lucaris
-- Petiolar node with a straight dorsal margin; metacoxal dorsum with a triangular tooth;
subpetiolar process with blunt angles ..................................................................siapensis
62(48) Scape usually smooth and shining with sparse punctulae.............................................. 63

-- Scape rugulose or striate (sculpture sometimes attenuated)........................................... 70
63(62) Mandibular dorsal surface striate or rugulose; metacoxal dorsum toothed.................... 64

-- Mandibular dorsal surface smooth and shining; metacoxal dorsum unarmed...... insularis
64(63) Masticatory margin of mandible with distinct denticles ................................................ 65

-- Masticatory margin of mandible edentate or with indistinct denticulation .................... 66
65(64) Propodeum unarmed; body striate ....................................................................... lineolata

-- Propodeum with denticles; body costulate....................................................... ingeborgae
66(64) Scape surpasses posterior cephalic margin by at least twice its apical width; HW > 1.1 mm;
WL > 1.6 mm................................................................................................................. 67
-- Scape surpasses posterior cephalic margin by not more than its apical width; HW < 1.1
mm; WL < 1.6 mm ............................................................................................... rastrata
67(66) Petiolar node shorter than wide in dorsal view .......................................................... lanei
-- Petiolar node longer than wide in dorsal view ............................................................... 68
68(67) Propodeal tooth longer than diameter of propodeal spiracle; subpetiolar process projecting
anteriorly as a triangular lobe in lateral view............................................. enodis (in part)
-- Propodeal tooth shorter than diameter of propodeal spiracle; subpetiolar process
subquadrate.......................................................................................................... menozzii
69(62) Propodeal teeth lacking.......................................................................... haenschi (in part)

-- Propodeal teeth present.................................................................................................. 70
70(69) Anterior pronotal margin with at least 5 rugulae or transverse costulae; eyes prominent and
protruding ...................................................................................................................... 71
-- Anterior pronotal margin with longitudinal costulae; eyes not prominent nor protruding,
relatively flat................................................................................................... triangularis
71(70) Cephalic vertex with 4 - 5 transverse costulae; anterior half of postpetiolar tergite rugose
....................................................................................................................................... 72
-- Vertex longitudinally costulate; anterior half of postpetiolar tergite longitudinally costulate
....................................................................................................................................... 73
72(71) Propodeal teeth at least as long as the distance between their bases; body ferruginous ....
........................................................................................................................... bispinosa
-- Propodeal teeth shorter; body black....................................................................perspicax
73(71) Transverse costulae from the metanotum to the propodeum............................ cuneiforma

-- Longitudinal costulae from the metanotum to the propodeum, partially effaced on the
propodeal declivity ........................................................................................................ 75
74(73) Pronotum longitudinally costulate; subpetiolar process rectangular in lateral view……….

............................................................................................................................. mecotyle
-- Pronotum with anterior strip of transverse costulae; subpetiolar process shaped a modest
lobe, without angles ................................................................................... enodis (in part)
75(1) Body sculpture mostly granulose and opaque; clypeus without anteromedian denticles
....................................................................................................................................... 76
-- Body finely striate, silky and shining; clypeus without anteromedian denticles............ 78
-- Body rugulose or roughly costulate; clypeus with anteromedian denticles ................... 81
76(75) Mandible falcate, the masticatory margin edentate and concave ............................... fieldi
-- Mandible triangular, the masticatory margin fairly straight and denticulate.................. 77
77(76) Metanotal groove well-impressed; spiracles of abdominal segments I and II shining and
conspicuous ........................................................................................................ petiscapa
-- Metanotal groove absent; spiracles of abdominal segments I and II opaque and not
conspicuous ............................................................................................................ minuta
78(75) Metanotal groove well-impressed.................................................................................. 79

-- Metanotal groove absent or weakly impressed .............................................................. 80
79(78) Clypeal lamella medially concave and with 2 angular lateral lobes; eyes prominent and
globulose; WL > 3.0 mm .......................................................................................... vriesi
-- Clypeal lamella medially straight to slightly convex and laterally rounded; eyes not
prominent; WL < 3.0 mm ...................................................................................... bufonis
80(78) Mandible triangular, masticatory margin with distinct teeth.................................falcifera

-- Mandible falcate, masticatory margin edentate ................................................. interrupta
81(75) Postpetiolar tergite mostly with arched punctate-rugulae, longitudinally costulate

posteromedially; meso- and metatibial spurs present............................................striolata
-- Postpetiolar tergite with rough longitudinal costulae; meso- and metatibial spurs absent
............................................................................................................................... caelata
Gnamptogenys wilsoni Lattke, new species

Figures 14, 15
TYPE MATERIAL
Holotype worker. Brazil, Bahia, CEPC 6. Ilheus, 1986, J. Delabie 32. Deposited in CEPEC,
Ilheus, Brazil.
DIAGNOSIS
Small (TL: 3 mm) with elongate and subfalcate mandible, internal margin and base of
masticatory margin with convex lobe, mandibular apex broadly curved and tapering; propodeal
spiracle slightly protruding posterad beyond propodeal declivity in lateral view.
WORKER
Measurements: HL 0.59, HW 0.52, ML 0.46, SL 0.43, ED 0.09, WL 0.94 mm; CI 0.88, SI 0.83,
MI 0.89, OI 0.17. Total Length: 3 mm.
Head subquadrate in dorsal view, posterior cephalic margin straight, lateral margin broadly
convex; clypeal lamella laterally curved, anterior margin broadly concave. Cephalic dorsum, and
vertex longitudinally costulate, individual costula slightly rugulose, not smooth. Scape failing to
reach posterior cephalic border by one apical width, dorsum smooth and shining with sparse
punctulae. Mandible elongate, semifalcate, with internal and masticatory margins joined by
convexity followed by brief concavity then straight to apical tooth, masticatory margin with series of
low blunt denticles, better observed in oblique ventral view of mandible; low strigulae present along
lateral basal third of mandible, the rest mostly smooth. Cephalic vertexal face flat, meeting dorsum at
blunt angle; eye small, set just anterad of cephalic mid-length in dorsal view; ventral cephalic face
longitudinally costulate.
Mesosoma with mostly flat dorsal margin in lateral view, propodeal dorsum curving onto
declivity, declivitous margin slightly interrupted by spiracle; propodeal spiracle on low tubercle,
situated less than one diameter from propodeal dorsum. Mesosomal side longitudinally costate;
mesopleuron roughly triangular with small dorsal lobe; mesometapleural suture well-impressed;
metapeural B propodeal suture indistinct or absent. Anterior pronotal margin with 3 transverse
costae; mesosomal dorsum and propodeal declivity longitudinally costate; promesonotal suture
absent, metanotal groove shallow but distinct.
Petiole in lateral view low, subquadrate, anterior margin brief and irregular, dorsal margin
broadly convex, anterior face with 3 transverse costae, laterally and dorsally with longitudinal
costae; posterior margin bound by single transverse costa in dorsal view, posterior face narrow and
smooth. Subpetiolar process in lateral view subquadrate with acute angles anterad and posterad,
ventral margin concave. Gaster longitudinally costate, constriction of abdominal pretergite I
longitudinally costae, postpetiolar anterior face with narrow smooth strip; postpetiolar sternite with
costae slightly weakened posteromedially. Protarsus opposite protibial strigil with single stout seta;
procoxa smooth and shining in lateral view; metacoxa with low dorsal tooth. Antennae, mandibles,
legs ferruginous; head brown, thorax and abdomen ferruginous brown.
Queen, male. Unknown.
ETYMOLOGY
The species is named in honor of Dr. Edward O. Wilson, in recognition of his outstanding
professional career in myrmecology, tropical biology and biological diversity issues.
COMMENTS
Based upon several morphological features (e.g., head shape, mandibular shape, laterally rounded
clypeal lamella, absent promesonotal suture) this species seems closest the the mordax-group
(Lattke, 1995) but differs due to the posterior position of the propodeal spiracle, in close proximity
to the propodeal declivity, as in the striatula-group. It falls closest to G. boliviensis and G. continua
in the key. G. wilsoni can be separated from G. boliviensis by the position of the propodeal spiracle,
which is separated by several times its diameter from both the propodeal dorsum and declivity. G.
boliviensis is very finely costulate, the mandible lacks a convex lobe at the interface of the internal
and masticatory margins, the apex remaining subparallel in dorsal view and ending abruptly, without
an apical tooth. G. continua has a more narrow clypeal lamella than in G. wilsoni, with a deeper
median concavity; its mandible has more developed denticles and no convex lobe along the internal
margin; the cephalic vertex is smooth; and the propodeal spiracle is separated from the declivity by
at least its diameter, and from the dorsal margin by several diameters. The mesopleuron in G. wilsoni
seems to be constituted mostly by the katepisternum, with the anepisternum reduced to a small
dorsal lobe.
LITERATURE CITED
Alonso, L.E. 2000. Ants as indicators of diversity. Pp. 80-88; in Agosti, D., Majer, J., Alonso, L. &
Schulz, T. (eds.). Ants, Standard methods for measuring and monitoring biodiversity: 280pp.
Smithsonian Institution Press, Washington, D.C.
Bolton, B. 1994. Identification guide to the ant genera of the world: 222pp. Harvard University
Press, Cambridge, Mass.
Bolton, B. 1995. A taxonomic and zoogeographical census of the extant ant taxa (Hymenoptera:
Formicidae). Journal of Natural History 29: 1037-1056.
Brown, W.L., Jr. 1992. Two new species of Gnamptogenys, and an account of millipede predation
by one of them. Psyche 99: 275-289.
Lattke, J.E. 1995 Revision of the ant genus Gnamptogenys in the New World (Hymenoptera:
Formicidae). Journal of Hymenoptera Research 4: 137-193.
Lattke, J.E. 2002. Nuevas especies de Gnamptogenys Roger, 1863 de América (Hymenoptera:
Formicidae: Ponerinae). Entomotropica 17(2): 135-144
Lattke, J.E., Fernández, F., & Palacio, E.E. 2004. Una nueva especie de Gnamptogenys
(Hymenoptera: Formicidae) con lista comentada de las especies de Colombia y Ecuador.
Iheringia (Series Zoológica) 43: 341-349.
Pacheco, J., MacKay, W. & Morgan, C. 2004. A new species of Gnamptogenys Roger of the
sulcata group (Hymenoptera: Formicidae) from Bolivia. Proceedings of the Entomological
Society of Washington 106: 434-437
Ward, P.S. 2000. Broad scale patterns of diversity in leaf litter communities, pp. 99-121; in Agosti,
D., Majer, J., Alonso, L. & Schulz, T. (eds.). Ants, Standard methods for measuring and
monitoring biodiversity: 280 pp. Smithsonian Institution Press, Washington, D.C.
Figures 1-4. Lateral view of petiole, (1) G. relicta, (2) G. moelleri, (3) G. acuta, (4) G. pleurodon.
Figures 5-6. View of declivitous propodeal face, (5) G. ammophila, (6) G. striatula.
Figures 7-8. Ventral view of subpetiolar process, (7) G. gracilis, (8) G. andina.
Figures 9-10. Cephalic vertex (9) G. gracilis, (10) G. porcata.
Figures 11-12. Frontal view of scape, (11) G. ammophila, (12) G. striatula.

Figure 13. Lateral view of mesosoma of G. nigrivitrea.
Figures 14-15. Gnamptogenys wilsoni, n. sp. Scale bar = 0.5 mm, (14) Lateral view of body, (15)
Dorsal view of head
Addendum
September 14, 2007
The following couplet:
53(51) Posterior face of petiolar node longitudinally costulate................................................. 54

-- Posterior face of petiolar node transversely costulate.................................................... 58
Should read:
53(51) Posterior face of petiolar node transversely costulate.................................................... 54

-- Posterior face of petiolar node longitudinally costulate................................................. 58
Longino, J. T., Fernández, F. 2007. Taxonomic review of the genus Wasmannia, pp. 271-289. In
TAXONOMIC REVIEW OF THE GENUS WASMANNIA
John T. Longino
The Evergreen State College
Olympia WA 98505 USA
longinoj@evergreen.edu
and
Fernando Fernández
Apartado 7495
ffernandezca@unal.edu.co
ABSTRACT
The Neotropical ant genus Wasmannia is reviewed and a key is provided to all Wasmannia
species. By far the most common species is W. auropunctata, native to the Neotropics and
becoming a serious pest ant when introduced elsewhere. Although W. auropunctata is variable,
there is no evidence that it is composed of multiple cryptic species. Queen size is dimorphic in
Costa Rica, but the taxonomic and biological significance of this is unknown. Other species of
Wasmannia are rare and inconspicuous. The following taxonomic changes are made: W.
australis Emery 1894, laevifrons Emery 1894, obscura Forel 1912, pulla Santschi 1931,
nigricans Emery 1906, and rugosa (Forel 1886) are synonymized under auropunctata (Roger
1863); W. weiseri Forel 1912 is synonymized under sulcaticeps Emery 1894. There is a negative
relationship between queen size and worker size among Wasmannia species. Wasmannia
auropunctata has the greatest difference, which may contribute to its ecological success.
Key words: Hymenoptera, Formicidae, Myrmicinae, Wasmannia auropunctata, invasive ants,

taxonomy, Neotropics.
INTRODUCTION
The genus Wasmannia is endemic to the Neotropics. Its most famous member is W.
auropunctata, the “little fire ant”. This species does very well in synanthropic habitats
throughout the Neotropics and has been introduced to tropical locales throughout the world,
where it often becomes a severe pest ant. Other species in the genus are few, and all of them are
uncommon relative to their famous relative. Twenty-one available names have accumulated in
the taxonomic literature, but there has never been a synthetic review of the genus. Kusnezov
(1952) characterized the Wasmannia species of Argentina, but no other work has focused on the
taxonomy of the genus. The purpose of this report is to provide a revised key, taxonomic
commentary, and natural history notes on all species in the genus. Particular emphasis is placed
on the occurrence of the genus in Costa Rica, where extensive inventory work has been carried
out.
Wasmannia was established in 1893 by Forel. The most recent ant classification (Bolton,
2003) places Wasmannia in the tribe Blepharidattini, within the attine tribe group. Bolton
proposes as an autapomorphy for the attine tribe group the “anterior clypeal margin with a broad
anteclypeal apron or flange that fits tightly over basal margins of mandibles and is at an angle to
outline of clypeus proper (not a direct continuation of median clypeus). Anteclypeal apron of
different sculpture/texture from median portion of clypeus.” Within this tribe group there are two
tribes: Attini (the fungus-growing ants) and Blepharidattini. Workers of Attini are differentiated
from the Blepharidattini by (1) the fungus-growing habit; (2) masticatory margin of mandibles
longer than basal margin, usually with 7 or more teeth (secondarily reduced to 5 teeth in some
groups); and (3) the 11-segmented antennae gradually incrassate. In contrast, the Blepharidattini
(1) do not culture fungi; (2) have relatively shorter mandibles, the masticatory margin with 5 or
fewer teeth and subequal in length to the basal margin; and (3) the 11-segmented antennae have
a discrete 2-segmented club. Additional blepharidattine characters described by Bolton are
clypeus broadly inserted between the frontal lobes and the propodeal spiracle low on the side of
the propodeum. The Blepharidattini contains two genera, Blepharidatta and Wasmannia.
Blepharidatta have very strongly developed antennal scrobes, the upper margin of which is
prolonged posteriorly so that the posterolateral margins of the vertex are drawn out as posteriorly
directed teeth or lobes, the anterior margins of the frontal lobes project forward almost to the
anterior margin of the clypeus, and the petiolar node is long, low, and without differentiated
anterior face. In Wasmannia, the antennal scrobes are shallow and never so strongly developed;
the posterolateral margins of the vertex are not produced as lobes or teeth; the frontal lobes do
not project forward; and the petiolar node is always well-developed, with a distinct anterior face.
In practice, Wasmannia may be confused with some species of Ochetomyrmex. For
example, the face of O. semipolita is very similar to Wasmannia (Fernandez, 2003).
Ochetomyrmex have less developed antennal scrobes, the clypeal apron is lacking, and there is a
slightly impressed mesonotal suture which is never present in Wasmannia. In addition,
Wasmannia lacks a bifurcated carina on the ventral surface of the petiole, a character present in
Ochetomyrmex, Tranopelta, and other myrmicines (Fernandez, 2003).
METHODS
The following terminology and abbreviations are used:
HL: head length; perpendicular distance from line tangent to rearmost points of vertex
margin to line tangent to anteriormost projections of clypeus, including anterior flange,
in full face view.
HW: head width; maximum width of head in face view, including eyes if they project beyond
the sides of the head.
Longino & Fernández: Review of Wasmannia 273
EL: eye length, measured along maximum diameter.

WL: Weber’s length; viewing mesosoma in lateral profile, distance from approximate
inflection point, where downward sloping pronotum curves into anteriorly projecting
neck, to posteroventral propodeal lobes.
CI: cephalic index; HW/HL.
OI: ocular index; EL/HL.
Collections are referred to by the following acronyms:
MHNG: Muséum d’Histoire Naturelle, Geneva, Switzerland.

MZSP: Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.
NHMB: Naturhistorisches Museum, Basel, Switzerland.
NMW: Naturhistorisches Museum, Vienna, Austria.
TAXONOMIC SYNOPSIS
Wasmannia affinis Santschi 1929. Brazil (Paraná).

Wasmannia auropunctata (Roger 1863). Throughout Neotropics, introduced elsewhere.
= atomum (Santschi 1914).
= australis Emery 1894. NEW SYNONYMY
= glabra Santschi 1931.
= laevifrons Emery 1894. NEW SYNONYMY
= obscura Forel 1912. NEW SYNONYMY
= panamana (Enzmann 1947).
= pulla Santschi 1931. NEW SYNONYMY
= nigricans Emery 1906.. NEW SYNONYMY
= rugosa (Forel 1886). NEW SYNONYMY
Wasmannia iheringi Forel 1908. Brazil (Espirito Santo), Peru, Costa Rica.
Wasmannia lutzi Forel 1908. southeastern Brazil.
Wasmannia rochai Forel 1912. Brazil to Costa Rica.
Wasmannia scrobifera Kempf 1961. Surinam, Costa Rica.
Wasmannia sigmoidea (Mayr 1884). Guianas, Antilles (St. Vincent; Puerto Rico), Costa Rica.
Wasmannia sulcaticeps Emery 1894. Argentina.
= bruchi Santschi 1919. TL: Argentina.
= weiseri Forel 1912. TL: Argentina. NEW SYNONYMY.
Wasmannia villosa Emery 1894. Brazil (Rio Grande do Sul).
Wasmannia williamsoni Kusnezov 1952. Argentina.
KEY TO WORKERS
(The worker of Wasmannia villosa is unknown.)
1a. Petiolar node strongly quadrate in lateral view, with nearly vertical anterior face and forming
sharp right angle with dorsal face (Fig. 1); petiolar peduncle about as long as node; setae on
mesosomal dorsum erect, long, about 0.1 mm or longer; antennal scrobe narrow, not
extending to side of head, ventral margin defined by preocular carina that runs from dorsal
margin of eye to margin of vertex; side of head posterior to eye rounded; color red brown to
orange .................................................................................................................auropunctata
1b. Petiolar node more rounded, with sloping anterior and posterior faces, and without sharply
differentiated dorsal face or clypeus strongly projecting and box-like and dorsal pilosity
composed of a very short stubble; petiolar peduncle longer or shorter than node; antennal
scrobe various; color various .................................................................................................. 2
2a. Scape strongly flattened; clypeus strongly projecting and box-like, sharply divided into equal
dorsal and anterior faces that meet at a right angle; setae on mesosomal dorsum abundant,
straight, filiform, short (about 0.04 mm long), forming a stubble; propodeal spines very
short, about same length as setae on mesosomal dorsum .........................................scrobifera
2b. Scape more terete, less flattened; clypeus not strongly box-like, rounded from posterior to
anterior margins; setae on mesosomal dorsum longer and/or curved, clavate; propodeal
spines various ......................................................................................................................... 3
3a. First gastral tergite lacking erect setae; petiolar peduncle longer than node (Fig. 1); dorsal
setae on mesosoma and face thin, flexuous; color yellow orange ................................iheringi
3b. First gastral tergite with abundant erect setae; petiolar peduncle equal to node or shorter;
dorsal setae stiff to slightly clavate; color various .................................................................. 4
4a. Antennal scrobe narrow, not extending to side of head, ventral margin defined by preocular
carina that runs from dorsal margin of eye to margin of vertex; side of head posterior to eye
rounded ................................................................................................................................... 5
4b. Antennal scrobe broad and flat, extending to side of head, ventral margin formed by angular
side of head posterior to eye ................................................................................................... 8
5a. Face between frontal carinae with about 12 distinct longitudinal striae overlaying strong
punctate sculpture; propodeal spiracle small, diameter less than width of base of propodeal
spine (Argentina) .................................................................................................................... 6
5b. Face between frontal carinae with fewer and more irregular longitudinal rugae, overlaying
opaque but not as strongly punctate sculpture; propodeal spiracle large and conspicuous,
diameter about equal to width of base of propodeal spine...................................................... 7
6a. Head width less than 0.50 mm; longitudinal striae on face very regular to margin of vertex;
color yellow red; posterior face of petiole rounded, anterior face slightly angulate .................
................................................................................................................................ sulcaticeps
6b. Head width greater than 0.54 mm; striae on face somewhat less parallel, becoming irregular
near margin of vertex; color uniformly dark maroon, abdomen black; anterior and posterior
faces of petiolar node similarly rounded.................................................................williamsoni
7a. Dorsal setae on mesosoma and gaster straight to weakly curved, thin, not clavate; head
relatively long, CI about 0.90; propodeal spines relatively long and upturned (Fig. 1); eyes
relatively larger (OI 0.26) ........................................................................................ sigmoidea
7b. Dorsal setae on mesosoma curved and clavate; head relatively short, CI 0.95-1.00; propodeal
spines short and directed posteriorly (Fig. 1); eyes shorter (OI 0.21-0.24) ....................rochai
8a. Postpetiole in dorsal view subquadrate to slightly trapezoidal, with widest portion anterior to
midlength; postpetiolar dorsum strongly punctate and opaque; propodeal spines relatively
long and robust, in dorsal view about as long as distance between their tips .................... lutzi
8b. Postpetiole in dorsal view elliptical, widest portion at or posterior to midlength; postpetiolar
dorsum feebly punctate, sublucid medially; propodeal spines shorter, in dorsal view shorter
than distance between tips .............................................................................................. affinis
KEY TO QUEENS
1a. Body covered with abundant, long, appressed pilosity; propodeal spine robust, short, forming
nearly a right angle; length about 4 mm ........................................................................ villosa
1b. Pilosity erect; propodeal spine long or short but usually more spiniform, acute; length
various .................................................................................................................................... 2
2a. Clypeus strongly projecting and box-like, sharply divided into equal dorsal and anterior faces
that meet at a right angle...........................................................................................scrobifera
2b. Clypeus not strongly box-like, rounded from posterior to anterior margins ............................ 3
3a. Head width less than 0.65 mm ................................................................................................. 4

3b. Head width greater than 0.65 mm ............................................................................................ 7
4a. Petiolar peduncle longer than node ................................................................................iheringi

4b. Petiolar peduncle about as long as node .................................................................................. 5
5a. Mesosomal dorsum almost uniformly punctate, with faint longitudinal striae on mesonotum;
striae on face weak....................................................................................................sigmoidea
5b. Mesosomal dorsum strongly longitudinally striate, on both mesonotum and scutellum; striae
on face strong.......................................................................................................................... 6
6a. Cephalic striae continuous, not mixed with other sculpture; color yellow red; posterior face
of petiole rounded, anterior face slightly angulate.................................................. sulcaticeps
6b. Cephalic striae mixed, appearing rugo-reticulated at least near vertex margin; color
uniformly dark maroon, abdomen black; anterior and posterior faces of petiolar node
similarly rounded ....................................................................................................williamsoni
7a. Antennal scrobe broad and flat, extending to side of head, ventral margin formed by angular
side of head posterior to eye; outer margin of frontal carinae not strongly convex, diverging
at level of ocelli; propodeal spines spiniform, longer than width at base; anteroventral
postpetiolar teeth short, inconspicuous .................................................................................. 8
7b. Antennal scrobe narrow, side of head rounded, ventral margin of scrobe defined by preocular
carina that runs from dorsal margin of eye to margin of vertex or ventral margin not
defined; frontal carinae, propodeal spines, and ventral postpetiolar teeth variable ................ 9
8a. Head in face view strongly trapezoidal, with widest portion of head behind eyes; rugae on
face fading posteriorly, weak on cephalic dorsum near margin of vertex ......................... lutzi
8b. Head in face view less trapezoidal, widest across eyes; rugae on face continuing to margin of
vertex .............................................................................................................................. affinis
9a. Antennal scrobe a distinct longitudinal trough, with defined ventral margin; propodeal spines
subtriangular, about as long as wide at base; head width more than 0.75 mm ....auropunctata
9b. Antennal scrobe weakly impressed, ventral margin not defined; propodeal spines spiniform,
longer than width at base; head width about 0.75 mm or less ........................................rochai
SPECIES ACCOUNTS
Wasmannia affinis Santschi

Table 1
Wasmannia affinis Santschi, 1929: 300, fig. 25, 26. Holotype worker: Brazil, Paraná, Rio
Negro (Reichensperger) [NHMB] (examined).
Wasmannia sigmoidea Mayr (part): Mayr, 1887: 622.
Taxonomic comments
Wasmannia affinis and lutzi are two related species from southeastern Brazil. They share a
unique development of the antennal scrobe. The scrobe is very broad, forming a flat surface that
extends from the frontal carinae to the side of the head. The side of the head is somewhat
angular posterior to the eye. The preocular carina is faint and does not form the ventral border of
the scrobe. In contrast, all other species of Wasmannia have a more narrow scrobe that does not
reach the side of the head in full face view. The ventral margin of the scrobe is limited by the
preocular carina or, in cases where the carina is faint or absent, where it would be if it extended
posterior to the eye. The side of the head behind the eye is rounded. The expanded scrobe is also
present in the queen of lutzi, resulting in a strongly trapezoidal head shape, such that the head is
broader behind the eyes than across them. Wasmannia affinis differs from lutzi in (1) the
propodeal spines are shorter, and (2) in dorsal view, the postpetiole is elliptical with rounded
sides, and the widest point is at or behind the midlength.
Mayr described W. sigmoidea in 1884, based on specimens from Cayenne (see below).
Later (Mayr, 1887) he identified a series of specimens from Santa Catarina state in Brazil as W.
sigmoidea. One of us (JTL) examined these Santa Catarina specimens (they have a variety of
labels, some indicating they are from Santa Catarina and collected by Hechko, some just saying
“Brazil 188,” and some erroneously labeled as types of sigmoidea). The workers in the series are
all very uniform and we suspect they are from a single original collection. These workers are not
sigmoidea, but instead match the holotype of Santschi’s affinis.
In addition to the material above, we have examined three collections from Santa Catarina
and São Paulo states in Brazil.
Wasmannia auropunctata (Roger)

Fig. 1, Table 1
Tetramorium auropunctatum Roger, 1863: 182. Syntype worker, queen, male: Cuba (not
examined). Combination in Ochetomyrmex: Forel, 1886; in Wasmannia: Forel, 1893.
Description of larva: Wheeler, G.C. & Wheeler, J., 1954: 444.
Ochetomyrmex auropunctatus var. rugosus Forel, 1886: xlix. Syntype worker: Guatemala,
Retalhuleu (Stoll) [MHNG] (examined). Combination in Wasmannia: Forel, 1901: 128.
Description of queen, male: Santschi, 1929: 299. NEW SYNONYMY.
Wasmannia auropunctata var. australis Emery, 1894: 193. Syntype worker: Brazil, Rio
Grande do Sul (v. Ihering). NEW SYNONYMY.
Wasmannia auropunctata var. laevifrons Emery, 1894: 193. Syntype worker: Bolivia,
Coroico, Chulumani Yungas (Balzan). NEW SYNONYMY.
Wasmannia auropunctata var. nigricans Emery, 1906: 160. Syntype worker: Paraguay,
Tacurú Pucú. NEW SYNONYMY.
Wasmannia auropunctata var. obscura Forel, 1912: 1. Syntype worker: Dibulla, Burithaka
and Don Diego, Sierra Nevada de Santa Marta, Colombia (Forel); Ceará, Brazil (Diaz da
Rocha); Santos, Brazil (Dr. Ris) [MHNG] (Dibulla and Ceará syntypes examined, Ceará
worker here designated LECTOTYPE). NEW SYNONYMY.
Xiphomyrmex atomum Santschi, 1914: 370. Syntype worker: Gabon, Libreville (Silvestri)
[NHMB] (examined). Combination in Wasmannia, and variety of auropunctata: Santschi,
1916:504. Junior synonym of auropunctata: Wheeler, 1922: 912.
Wasmannia glabra Santschi, 1931: 272. Syntype queen: French Guiana, St. Laurent du
Moroni (Le Moult) [NHMB] (examined). Junior synonym of auropunctata: Kempf,
1964:66.
Wasmannia auropunctata st. pulla Santschi, 1931: 272. Syntype worker: Panama, France
Field, 9 May 1930 (A. Bierig) [NHMB] (examined). NEW SYNONYMY.
Hercynia panamana Enzmann, J. 1947: 44, pl. 5. Syntype worker, queen: Panama. Junior
synonym of auropunctata: Brown, 1948: 102.
Taxonomic comments
Workers of W. auropunctata have a strongly quadrate petiolar node. The anterior face of the
node is sharply differentiated from both the peduncle and the dorsal face of the node, meeting
both at nearly right angles, and forming a strongly step-like profile. This is a highly distinctive
feature that easily distinguishes auropunctata workers from all other Wasmannia species. Within
the species there is abundant variation in the strength of sculpturing and coloration, and this has
engendered the naming of nine infraspecific forms in addition to the nominotypical. We have
never been able to discover evidence of discrete forms among the workers of auropunctata,
either in sympatry or allopatry; the variation appears continuous. Three of the forms have
already been synonymized by others: atomum, glabra, and panamana. We have been able to
examine types of three additional forms—obscura, pulla, and rugosa—and they fall well within
our concept of auropunctata. The remaining forms—australis, laevifrons, nigricans—we have
not examined, but the published descriptions give no indication that they are beyond the range of
variation of auropunctata. We have synonymized them, following the philosophy that taxa
should be synonymized unless evidence of distinctness is obtained.
The above synonymy does not assure that there is no genetic structuring, and perhaps
species-level differences, within the broad concept of auropunctata. There may even be distinct
sympatric species. In Costa Rica, there appear to be two size classes of queens (Fig. 2). Queens
with smaller heads include ten from various sites in Costa Rica, including La Selva Biological
Station and the Peñas Blancas Valley, one from Jamaica, and one from Venezuela. Queens with
large heads are all from the Atlantic slope of Costa Rica. Three are from La Selva (two from
different Winkler samples of sifted leaf litter from the forest floor, one from a small nest under
an epiphyte mat in an old treefall) and one is an alate queen found in a Cecropia sapling near
Volcan Arenal. Thus the small-headed and big-headed forms are broadly sympatric in Costa
Rica. Among the small-headed queens four are definitively associated with workers from the
same colony, and among the big-headed queens one is associated with workers. Others have
workers doubtfully associated (together in the same Winkler sample). We can discern no
differences in workers associated with the two types of queens. The cause of the two size classes
of queens is unknown, but could reflect either differences between cryptic species or
intraspecific polymorphism.
An aberrant worker form is frequently encountered in large samples of W. auropunctata.
These aberrant workers have the head grossly swollen. The entire head is more spherical than
normal, as if the head were inflated like a balloon. The rest of the body is little different from a
normal worker. These aberrant workers are occasionally encountered in Winkler samples that
contain hundreds or thousands of auropunctata workers.
Natural History
Wasmannia auropunctata is a widespread pest ant (Clark et al., 1982; De Souza et al., 1998;
Fabres & Brown, 1978; Jourdan, 1997; Lubin, 1984; Ulloa Chacón & Cherix, 1990; Williams,
1994; Wetterer & Porter, 2003). In its presumed native range it occurs from Argentina to Mexico
(Kempf, 1972; Wetterer & Porter, 2003). Its introduced range includes the Galapagos Islands,
West Africa (Gabon, Cameroon, and possibly the Republic of Congo and the Democratic
RepubIic of Congo), Melanesia (New Caledonia, Solomon Islands, Vanuatu, and possibly
Tuvalu), Polynesia (Wallis and Futuna and Hawaii), parts of the US (Florida and possibly
California), and subtropical Atlantic islands (the Bahamas and Bermuda) (Wetterer & Porter,
2003). It is widespread on Caribbean islands, but it is unclear whether these are long-term native
populations or recent introductions (Wetterer & Porter, 2003).
The species is remarkably catholic in its habitat preference. It is common in habitats ranging
from wet to dry and from early successional to mature. In an elevational gradient of mature wet
forest on the Atlantic slope of Costa Rica (the Barva Transect, from La Selva Biological Station
to 2000m elevation on the slope of Volcan Barva) it is abundant at 50m and 500m elevations,
but nearly absent at 1070m (Table 2). In the lowland habitats where it is abundant, it occurs in
leaf litter on the forest floor and at all levels in the vegetation.
Although it occurs frequently in samples from mature forest habitats in Costa Rica, it is
never so abundant in those habitats that it is noticeable as a pest or appears to be displacing other
native species (Tennant, 1994; McGlynn & Kirksey, 2000; pers. obs.). In contrast, in certain
agricultural habitats (banana plantations) and in parts of the tropics where it has been introduced
it becomes super-abundant, with negative impacts on native species and human comfort (Clark
et al., 1982; Wetterer & Porter, 2003). In dry-forest fragments in Colombia there is a negative
correlation between W. auropunctata abundance and overall ant diversity (Armbrecht & Ulloa
Chacón, 2003). Where introduced in New Caledonia it invades dense native forest and displaces
native ants (Le Breton et al., 2003). Behavioral tests and cuticular hydrocarbon analysis show
that W. auropunctata is multicolonial in its native range in Brazil, unicolonial where introduced
in New Caledonia (Errard et al., 2002).
The sting of Wasmannia is noteworthy. These are extremely tiny ants, barely visible in the
field. When the senior author first began studying ants in Costa Rica, he was at first puzzled
about Wasmannia. By literature accounts Wasmannia was reputed to have a terrible sting, but he
had been collecting them for months in Corcovado National Park without ever experiencing the
famous sting. One day he was collecting from a populous nest and some workers made it up to
the soft skin of his inner forearm and began to sting. The sting was definitely noticeable, about
as severe as a fire ant (i.e., Solenopsis geminata) but inordinately strong for an ant that could
barely be seen! Workers are so small they cannot sting through the thicker skin of the hands.
Surprisingly, the chemical and toxicological nature of the venom of W. auropunctata has
not been investigated. Howard et al. (1982) discovered an alkylpyrazine compound in the
mandibular glands, which acted as an attractant to conspecifics and a repellent to heterospecifics.
They speculated that the workers might apply the mandibular gland product as an irritating
secretion, augmenting the defensive properties of the venomous sting. It would be interesting to
investigate whether the venom alone is the powerful agent in this small ant, or if the strong
burning sensation is a synergetic effect of venom plus mandibular gland product.
Nests can be almost anywhere: in rolled leaves or dead sticks in the leaf litter, under stones,
in rotten wood, in hollow stems suspended above the ground, in ant-plant domatia, and under
epiphytes. Workers are omnivorous scavengers and predators and can rapidly recruit to food.
Colonies are polygynous and it is never clear where colony boundaries are. Dozens of dealate
queens may be found together in nests. Males are rare but do occasionally occur.
Wasmannia iheringi Forel

Fig. 1, Table 1
Wasmannia iheringi Forel, 1908: 359. Syntype worker, queen: Brazil, São Paulo (v.
Ihering) [MHNG] (examined).
Comments
Prior to this report W. iheringi was known only from the type specimens. Forel’s description
and specimen labels indicate the specimens were collected in São Paulo by von Ihering. The
types also bear a pencil label with “2265.” Luederwaldt (1926) referred to W. iheringi, stating
“Mr. E. Garbe found in Espirito Santo a small nest, constructed of fine carton [“serragem fina”],
on a leaf of Cecropia. N. 2.265.” Given the match of the collection numbers We assume these
represent a single collection. We also presume Luederwaldt’s data are more accurate than
Forel’s. Kempf (1972) lists only the type locality for the range of W. iheringi, suggesting it
remained known only from the types at the time of his Neotropical catalogue.
Surprisingly, this species has been discovered in Costa Rica, where it inhabits the canopy of
lowland rainforest on the Atlantic slope. Morphologically the Costa Rican specimens are
identical to the types. It has been collected at La Selva Biological Station, at 500m elevation on
the Barva Transect, and at the old Carrillo station at 600m in Braulio Carrillo National Park. The
quantitative sampling by the ALAS project shows it to be a moderately abundant component of
the arboreal fauna (Table 2).
One nest has been observed, at La Selva Biological Station in Costa Rica. The nest was on
the undersurface of a single leaf of a fuzzy-leaved Guarea growing along a stream. A 2cm long
felt-like carton roof covered a portion of the leaf between two veins. The nest contained 27 adult
workers, one dealate queen, four adult males, and brood of various sizes. This observation is
similar to Garbe’s collection from southeastern Brazil, suggesting a specialized nesting behavior
that is the same in both Costa Rica and southeastern Brazil.
On another occasion at La Selva the species was observed in a patch of old second growth
forest comprised of medium to large trees over abandoned cacao. A small patch was being felled
for an experiment at La Selva and collecting was carried out in these newly-felled trees. A dense
aggregation of workers and two dealate queens were found on a branch of a felled Coussapoa,
suggesting that colonies can be polygynous.
Alate queens were collected at blacklights at La Selva in October 1991.
We have also examined workers in Erwin’s fogging samples from Tambopata, Peru. The
species is now known from three widely separated localities, but it is unknown whether these are
disjunct populations or a result of undersampling in intervening regions.
Wasmannia lutzi Forel

Table 1
Wasmannia lutzi Forel, 1908: 357. Syntype worker, queen, male: Brazil, São Paulo (Lutz)
[MHNG] (examined).
See under W. affinis.
Wasmannia rochai Forel

Fig. 1, Table 1
Wasmannia rochai Forel, 1912: 1. Syntype worker: Ceará, Brazil (Diaz da Rocha) [MHNG]
(examined, one worker here designated LECTOTYPE).
Comments
Wasmannia rochai occurs widely in the mainland Neotropics, from Guatemala south to São
Paulo state in Brazil. The syntype workers from Ceará are slightly larger than workers from
Panama and Costa Rica, but with similar proportions. The differences between rochai and
sigmoidea are subtle but consistent. Compared to sigmoidea, rochai is smaller, with a relatively
shorter and broader head. The propodeal spines are shorter and are directed posteriorly, instead
of upturned in sigmoidea. The setae on the face, mesosoma, and gaster are more curved,
appearing shorter than the setae of sigmoidea because of the greater curvature. They are also
more clavate, swelling noticeably at the tips.
In some specimens of rochai the outer margin of the antennal scrobe is weakly defined,
approaching the condition seen in affinis and lutzi. However, the face sculpture of rochai is
always much more feeble than the coarse reticulate rugose sculpture on affinis and lutzi.
Kempf (1972) recorded the range of rochai as Panama, the Guianas, Trinidad, and six
Brazilian states from Ceará south to São Paulo. Given the prior uncertainty of species
differences among affinis, lutzi, rochai, and sigmoidea, earlier determinations should be treated
cautiously pending reexamination of existing material. We have examined material from
Guatemala, Costa Rica, Venezuela, and Brazil (Amazonas, Bahia, Ceará, and São Paulo states).
This species appears to be rare in Costa Rica, although its superficial similarity to W.
auropunctata may result in its being overlooked in a sea of the latter species. Only two Costa
Rican collections are known: Phil Ward collected workers and a dealate queen (PSW#7628) in a
recent treefall at Carara Biological Reserve. The senior author collected a lone dealate queen in a
canopy tree at Sirena in Corcovado National Park. The species has been collected multiple times
on Barro Colorado Island in Panama, where it is a relatively common part of the canopy ant
fauna (Mike Kaspari pers. comm.).
Wasmannia scrobifera Kempf

Fig. 1, Table 1
Wasmannia scrobifera Kempf, 1961: 512, figs. 14, 15. Holotype worker: Poeroe man
kemisa, Surinam (van der Drift).
Comments
Kempf described this species from a single worker. Numerous collections are now known
from Costa Rica, and we have also seen material from Colombia and Brazil (Bahia and Matto
Grosso states).
In Costa Rica, this species is infrequently encountered. It inhabits mature lowland rainforest
in the Atlantic lowlands. It has been collected in Winkler samples of sifted litter from the forest
floor at Hitoy Cerere Biological Reserve, Casa Plastico near Rara Avis, the 500m site on the
Barva Transect, and La Selva Biological Station. Quantitative sampling by the ALAS project
reveals it to be a low density species occurring most often in fogging samples, and less often in
Winkler and Berlese samples of forest floor litter (Table 2). Dinah Davidson collected it in a
Piper plant at La Selva. Mary Cornelius, an OTS student, found a nest in a leaf domatium of an
ant-plant (Tococa, Melastomataceae) at Tortuguero, but it contained only workers and brood.
Grant Gentry found small carton nests under leaves at La Selva (pers. comm.). Workers have
been collected from the stomachs of dendrobatid frogs in the Chocó region of Colombia. These
results suggest that the species may nest in the low arboreal zone yet forage in the leaf litter on
the forest floor.
Wasmannia sigmoidea (Mayr)

Fig. 1, Table 1
Tetramorium sigmoideum Mayr (in Radoszkowsky), 1884: 33. Syntype worker: Cayenne,
French Guiana (Jelski) [NMW] (examined, one worker here designated LECTOTYPE).
Combination in Wasmannia and description of queen and male: Forel, 1893:386.
Comments
Wasmannia sigmoidea is the second oldest name in the genus and to date has been poorly
characterized. It has never been satisfactorily differentiated from other species in the genus,
particularly rochai. Forel (1884) identified material from St. Vincent Island in the Antilles as W.
sigmoidea, and described the queen and male. We have not been able to examine the workers of
this collection, but the queens match queens of sigmoidea from Puerto Rico and Costa Rica.
Kempf (1972) gives the range of sigmoidea as Guianas, Antilles St. Vincent, Grenada, and
Santa Catarina state of Brazil. The Guianas are listed because of the type locality in French
Guiana and some Surinam specimens he tentatively identified as sigmoidea (Kempf 1961). The
St. Vincent record is based on Forel’s publication. We do not know the basis of the Grenada
record. The Santa Catarina record is based on a published record by Mayr (1887), which is a
misidentification of specimens of affinis (see under affinis). We have examined abundant
material from Puerto Rico, multiple collections from Costa Rica, and a collection from Guarico
state in Venezuela. Thus the current known range of sigmoidea is circumcaribbean.
The few Costa Rican records are as follows. David Olson collected workers during his study
of Winkler and pitfall trap sampling methods at La Selva Biological Station (Olson 1991, as
Wasmannia sp.1). This was the only known collection from La Selva, in spite of intensive
inventory effort there (Longino et al., 2002), until an August, 2004 collection of workers and
alate queens from the rootball of a palm tree in the laboratory clearing. Workers occurred in four
different samples from the Project ALAS expeditions to the 500m site on the Barva Transect:
two Malaise trap samples, one flight-intercept sample, and one sweep net sample (Table 2). It is
likely that sigmoidea prefers open and synanthropic habitats, hence its undersampling in Costa
Rica, where sampling emphasis has been in forested habitats.
Wasmannia sulcaticeps Emery

Table 1
Wasmannia sulcaticeps Emery, 1894: 195. Holotype worker: Argentina, Buenos Aires.
Description of queen, male: Forel, 1914: 280.
Wasmannia sulcaticeps var. weiseri Forel, 1914: 281. Syntype queen, male: Argentina,
Cordoba, Canals (Weiser) [MHNG] (examined). NEW SYNONYMY.
Wasmannia bruchi Santschi, 1919: 43. Syntype worker: Argentina, La Plata (Bruch)
[NHMB] (examined). Junior synonym of sulcaticeps: Kusnezov, 1952: 174.
Comments
Wasmannia sulcaticeps and williamsoni are two related species that occur at the far southern
limit of the genus, in Argentina. They are both distinguished from other members of the genus
by the heavy striate sculpture on the face and by the very small propodeal spiracle.
Kusnezov (1952) described the Wasmannia fauna of Argentina, and sulcaticeps and
williamsoni were described as having an allopatric distribution. Wasmannia sulcaticeps occurred
in more humid environments near Buenos Aires and in the northern provinces of Tucumán,
Salta, and Jujuy. Wasmannia williamsoni occurred in the more arid habitats west of Buenos
Aires. Kusnezov did not mention var. weiseri, and its type locality is between the ranges of
sulcaticeps and williamsoni shown on his distribution map. Forel’s description of weiseri noted
only minor differences from sulcaticeps.
Wasmannia villosa Emery
Wasmannia villosa Emery, 1894: 196. Holotype queen: Brazil, Rio Grande do Sul (v.
Ihering).
Comments
Emery described this single queen from Rio Grande do Sul. He distinguished it from other
species by the long, abundant, appressed pilosity; robust dentiform propodeal spine; and
transverse petiolar node. We have not examined this specimen, but the description sets it apart
from any Wasmannia queen with which we are familiar. However, its status as a species of
Wasmannia is not assured. It would not surprise us if it were revealed to be an attine queen or a
member of some other myrmicine genus.
Wasmannia williamsoni Kusnezov

Table 1
Wasmannia williamsoni Kusnezov, 1952: 181, figs. 1, 4–7. Syntype worker, queen:
Argentina, La Pampa, Winifreda, Castex (Kusnezov). [MZSP] (worker examined).
Remarks
Wasmannia williamsoni has the largest workers in the genus, based on the one worker we
have examined. It is far larger than any other Wasmannia worker we have seen (Table 1). See
further information under sulcaticeps.
GENERAL DISCUSSION
Kusnezov (1952) observed that Wasmannia auropunctata was the dominant Wasmannia species
in Argentina and that it had the greatest difference between worker size and queen size. The
Argentinean species sulcaticeps and williamsoni were restricted to the subtropical and semiarid
limits of the genus, and the queens were much smaller and closer to the workers in size. He
suggested that the species such as sulcaticeps and williamsoni were the more primitive members
of the genus, and auropunctata was more recent and more advanced. We examined the
relationship between worker size and queen size for the eight species of Wasmannia for which
we had data, and there is a negative relationship (Fig. 3). Species with the smallest workers have
the largest queens. The species fall roughly into three groups. Wasmannia sigmoidea, iheringi,
sulcaticeps, and scrobifera have very small queens, little larger than the workers. Wasmannia
rochai, affinis, and lutzi are intermediate, with relatively smaller workers and larger queens.
Wasmannia auropunctata is at the top, with the smallest workers and largest queens.
These results support Kusnezov’s conclusions. A general trend in ant evolution is the
development of increasing caste differences, and it follows that a greater difference in size
between worker and queen is often apomorphic. If that is the case with Wasmannia, then
auropunctata, lutzi, affinis, and rochai may form a clade within Wasmannia based on larger
queen size. If increasing caste differentiation is associated with greater ecological success, the
ecological dominance of auropunctata could be explained by it having the largest queens and
smallest workers in the genus.
A case can also be made for iheringi being the sister taxon to all other Wasmannia. It has
the most plesiomorphic mesosoma and petiole shape, resembling other generalized myrmicines.
Other Wasmannia show a trend toward shorter, more compact mesosoma and shorter petiolar
peduncle and/or more quadrate node. A biogeographic scenario would thus have iheringi as the
oldest lineage in the genus, with disjunct populations at scattered localities over a large area,
having been largely displaced from a formerly extensive range. Other rare or localized species in
the genus — sulcaticeps, williamsoni, sigmoidea, scrobifera, affinis, lutzi — could also be relicts
from an early radiation of the genus. Wasmannia rochai is the second most common and
widespread species. Its small workers and large queens could have been an initial step in the
direction of ecological dominance. Wasmannia auropunctata, with its even smaller workers and
larger queens, is the most recent manifestation of this trend. It has swept the Neotropical field
and is now conquering the world.
ACKNOWLEDGMENTS
We thank museum curators B. Merz (MHNG), D. Burckhardt (NHMB) and S. Schödl (NMW)
for loans of type material, and the ALAS staff for help with field and lab work. This work was
supported by National Science Foundation grants DEB-0072702 and DBI-0215820, and
National Geographic Society grant 7331-02.
LITERATURE CITED
Armbrecht, I. & Ulloa Chacón, P. 2003. The little fire ant Wasmannia auropunctata (Roger)
(Hymenoptera: Formicidae) as a diversity indicator of ants in tropical dry forest fragments
of Colombia. Environmental Entomology 32: 542-547.
Brown, W.L., Jr. 1948. The status of the genus Hercynia J. Enzmann (Hymenoptera:
Formicidae). Entomological News 59: 102.
Clark, D.B., Guayasamín, C., Pazmiño, O., Donoso, C. & Páez de Villacís, Y. 1982. The
tramp ant Wasmannia auropunctata: autecology and effects on ant diversity and distribution
on Santa Cruz Island, Galapagos. Biotropica 14: 196-207.
De Souza, A.L.B., Delabie, J.H.C. & Fowler, H. G. 1998. Wasmannia spp. (Hym. Formicidae)
and insect damages to cocoa in Brazilian farms. Journal of Applied Entomology 122: 339-
341.
Emery, C. 1894. Studi sulle formiche della fauna neotropica. VI-XVI. Bullettino della Società
Entomologica Italiana 26: 137-241.
Emery, C. 1906 (“1905”). Studi sulle formiche della fauna neotropica. XXVI. Bullettino della
Società Entomologica Italiana 37: 107-194.
Enzmann, J. 1947. Hercynia, a new genus of myrmicine ants. Journal of the New York
Errard, C., Jourdan, H., Glaude, C., Delabie, J.H. & Hefetz, A. 2002. Intraspecific and
interspecific discrimination in the tramp ant Wasmannia auropunctata (Hym. Formicidae).
Actes des Colloques Insectes Sociaux 15: 82-89.
Fabres, G. & Brown, W.L., Jr. 1978. The recent introduction of the pest ant Wasmannia
auropunctata into New Caledonia. Journal of the Australian Entomological Society 17:
139-142.
Fernández, F. 2003. The myrmicine ant genera Ochetomyrmex Mayr and Tranopelta Mayr
(Hymenoptera: Formicidae). Sociobiology 41: 633-661.
Forel, A. 1886. Espèces nouvelles de fourmis américaines. Annales de la Société Entomologique
de Belgique 30: xxxviii-xlix.
Forel, A. 1893. Formicides de l’Antille St. Vincent, récoltées par Mons. H. H. Smith.
Transactions of the Entomological Society of London 1893: 333-418.
Forel, A. 1901. I. Fourmis mexicaines récoltées par M. le professeur W.-M. Wheeler. II. A
propos de la classification des fourmis. Annales de la Société Entomologique de Belgique
45: 123-141.
Forel, A. 1908. Ameisen aus São Paulo (Brasilien), Paraguay etc. gesammelt von Prof. Herm. v.
Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandlungen der Kaiserlich-Königlichen Zoologisch-
Botanischen Gesellschaft in Wien 58: 340-418.
Forel, A. 1912. Formicides néotropiques. Part IV. 3me sous-famille Myrmicinae Lep. (suite).
Mémoires de la Société Entomologique de Belgique 20: 1-32.
Forel, A. 1914. Formicides d’Afrique et d’Amérique nouveaux ou peu connus. Bulletin de la
Société Vaudoise des Sciences Naturelles 50: 211-288.
Howard, D.F., Blum, M.S., Jones, T.H. & Tomalski, M.D. 1982. Behavioral responses to an
alkylpyrazine from the mandibular gland of the ant Wasmannia auropunctata. Insectes
Sociaux 29: 369-374.
Jourdan, H. 1997. Threats on Pacific islands: the spread of the tramp ant Wasmannia
auropunctata (Hymenoptera: Formicidae). Pacific Conservation Biology 3: 61-64.
Kempf, W.W. 1961. A survey of the ants of the soil fauna in Surinam (Hymenoptera:
Kempf, W.W. 1964. Miscellaneous studies on Neotropical ants. III. (Hymenoptera:
Kempf, W.W. 1972. Catálogo abreviado das formigas da Região Neotropical. Studia
Entomologica 15: 3-344.
Kusnezov, N. 1952 (“1951”). El género Wasmannia en la Argentina (Hymenoptera,
Formicidae). Acta Zoologica Lilloana 10: 173-182.
Le Breton, J., Chazeau, J. & Jourdan, H. 2003. Immediate impacts of invasion by Wasmannia
auropunctata (Hymenoptera: Formicidae) on native litter ant fauna in a New Caledonian
rainforest. Austral Ecology 28: 204-209.
Longino, J.T., Colwell, R.K. & Coddington, J.A. 2002. The ant fauna of a tropical rainforest:
estimating species richness three different ways. Ecology 83: 689-702.
Lubin, Y.D. 1984. Changes in the native fauna of the Galápagos Islands following invasion by
the little red fire ant, Wasmannia auropunctata. Biological Journal of the Linnean Society
21: 229-242.
Luederwaldt, H. 1926. Observações biologicas sobre formigas brasileiras especialmente do
estado de São Paulo. Revista do Museu Paulista 14: 185-303.
Mayr, G. 1887. Südamerikanische Formiciden. Verhandlungen der Kaiserlich-Königlichen
Zoologisch-Botanischen Gesellschaft in Wien 37: 511-632.
McGlynn, T.P. & Kirksey, S.E. 2000. The effects of food presentation and microhabitat upon
resource monopoly in a ground-foraging ant (Hymenoptera: Formicidae) community.
Revista de Biologia Tropical 48: 629-642.
Olson, D.M. 1991. A comparison of the efficacy of litter sifting and pitfall traps for sampling
leaf litter ants (Hymenoptera, Formicidae) in a tropical wet forest. Biotropica 23: 166-172.
Radoszkowsky, O. 1884. Fourmis de Cayenne Française. Trudy Russkago Entomologicheskago
Obshchestva 18: 30-39.
Roger, J. 1863. Die neu aufgeführten Gattungen und Arten meines Formiciden-Verzeichnisses
nebst Ergänzung einiger früher gegebenen Beschreibungen. Berliner Entomologische
Zeitschrift 7: 131-214.
Santschi, F. 1914. Formicides de l’Afrique occidentale et australe du voyage de Mr. le
Professeur F. Silvestri. Bollettino del Laboratorio di Zoologia Generale e Agraria della
Reale Scuola Superiore d’Agricoltura. Portici 8: 309-385.
Santschi, F. 1916 (“1915”). Descriptions de fourmis nouvelles d’Afrique et d’Amérique.

Annales de la Société Entomologique de France 84: 497-513.
Santschi, F. 1919. Nouveaux formicides de la République Argentine. Anales de la Sociedad
Cientifica Argentina 87: 37-57.
Santschi, F. 1929. Nouvelles fourmis de la République Argentine et du Brésil. Anales de la
Sociedad Cientifica Argentina 107: 273-316.
Santschi, F. 1931. Fourmis de Cuba et de Panama. Revista de Entomologia (Rio de Janeiro) 1:
265-282.
Tennant, L.E. 1994. The ecology of Wasmannia auropunctata in primary tropical rainforest in
Costa Rica and Panama. Pp. 80-90 in: Williams, D. F. (ed). Exotic ants: biology, impact,
and control of introduced species: 332 pp. Westview Press, Boulder, Colorado, USA.
Ulloa Chacón, D. & Cherix, D. 1990. The little fire ant Wasmannia auropunctata (Roger)
(Hymenoptera: Formicidae). Pp. 281-289 in: Vander Meer, R. K., Jaffe, K. & Cedeno, A.
(eds). Applied myrmecology: a world perspective: 741 pp. Westview Press, Boulder,
Colorado, USA.
Wetterer, J.K. & Porter, S.D. 2003. The little fire ant, Wasmannia auropunctata: distribution,
impact, and control. Sociobiology 42: 1-41.
Wheeler, G.C. & Wheeler, J. 1954. The ant larvae of the myrmicine tribes Meranoplini,
Ochetomyrmicini and Tetramoriini. American Midland Naturalist 52: 443-452.
Wheeler, W.M. 1922. Ants of the American Museum Congo expedition. A contribution to the
myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bulletin
of the American Museum of Natural History 45: 711-1004.
Williams, D.F. (ed). 1994. Exotic ants. Biology, impact, and control of introduced species: 332
pp. Westview Press, Boulder, Colorado, USA.
Table 1. Measurement data for Wasmannia queens and workers. The additional set of
measurements for W. auropunctata queen head sizes are shown in Fig. 2.
Species specimen1 WL HW HL EL CI OI
Queens
affinis Brazil (SC), MZSP2594 1.16 0.74 0.72 0.16 1.01 0.22
auropunctata INBIOCRI002280087 1.13 0.77 0.69 0.22 1.13 0.33
auropunctata INBIOCRI002280107 1.19 0.80 0.71 0.21 1.13 0.30
auropunctata holotype glabra - 0.84 0.74 0.26 1.13 0.35
iheringi INBIOCRI001271987 0.86 0.60 0.61 0.18 0.99 0.29
iheringi syntype 0.88 0.62 0.63 0.16 0.99 0.26
lutzi syntype 1.15 0.76 0.69 0.19 1.10 0.27
rochai LACM ENT 141402 1.08 0.68 0.64 0.22 1.06 0.34
rochai INBIOCRI002280101 1.08 0.68 0.65 0.20 1.05 0.31
rochai Brazil (SP), MZSP1843 1.20 0.75 0.70 0.24 1.07 0.34
scrobifera INBIOCRI002280114 0.67 0.56 0.60 0.17 0.94 0.28
scrobifera Brazil (BA), ICN 0.70 0.54 0.61 0.16 0.88 0.27
sigmoidea St. Vincent 0.82 0.61 0.62 0.20 0.97 0.33
sigmoidea Puerto Rico LACM 0.86 0.58 0.65 0.21 0.89 0.32
sulcaticeps syntype weiseri 0.80 0.59 0.63 0.18 0.93 0.28
Workers
affinis holotype 0.58 0.51 0.55 0.11 0.93 0.20
affinis Brazil (SC), MZSP2594 0.62 0.55 0.61 0.13 0.91 0.21
auropunctata lectotype obscura 0.46 0.42 0.46 0.11 0.92 0.23
auropunctata syntype rugosa 0.48 0.45 0.48 0.10 0.95 0.20
auropunctata Colombia ICN 0.46 0.42 0.46 0.11 0.92 0.23
iheringi syntype MHNG 0.65 0.57 0.53 0.11 1.06 0.21
iheringi syntype MZSP 0.63 0.49 0.56 0.11 0.87 0.20
iheringi INBIOCRI001271987 0.65 0.52 0.54 0.13 0.97 0.24
lutzi syntype 0.56 0.49 0.55 0.10 0.89 0.19
rochai LACM ENT 140078 0.46 0.43 0.45 0.11 0.97 0.24
rochai LACM ENT 141402 0.45 0.44 0.46 0.10 0.96 0.21
rochai lectotype 0.54 0.50 0.50 0.11 0.99 0.22
rochai Brazil (SP), MZSP 0.50 0.47 0.48 0.11 0.97 0.23
scrobifera INBIOCRI002280114 0.56 0.52 0.52 0.15 0.99 0.29
sigmoidea LACM ENT 141401 0.56 0.49 0.55 0.14 0.89 0.26
sigmoidea lectotype 0.58 0.51 0.56 0.14 0.91 0.26
sigmoidea INB0003235900 0.60 0.52 0.57 0.15 0.90 0.26
sigmoidea INB0003231652 0.61 0.52 0.58 0.15 0.90 0.26
sulcaticeps syntype bruchi 0.57 0.48 0.55 0.12 0.88 0.22
sulcaticeps syntype bruchi 0.58 0.49 0.57 0.12 0.85 0.21
williamsoni syntype MSZP 0.71 0.59 0.69 0.15 0.85 0.22
1. Specimen barcodes or type data. Barcoded specimens are all from Costa Rica; additional
specimen data available from senior author.
Table 2. Abundance of Wasmannia species along an elevational gradient on the Atlantic slope
of Costa Rica. Table values are percent of samples in which a species occurred. Details of
sampling methods are in Longino et al. (2002) and http://viceroy.eeb.uconn.edu/ALAS/
ALAS.html.
No. Samples Wasmannia Wasmannia Wasmannia Wasmannia
auropunctata iheringi sigmoidea scrobifera
La Selva
Baits 40 0% 0% 0% 0%
Berlese 217 11% 0% 0% <1%
Fogging 52 40% 25% 0% 23%
Malaise 62 11% 15% 0% 3%
Litter nests 222 8% 0% 0% 0%
Winkler 41 83% 2% 0% 7%
500m
Berlese 94 14% 0% 0% 0%
Fogging1 3 0% 0% 0% 0%
Malaise2 20 65% 5% 10% 0%
Sweepnet 58 33% 0% 2% 0%
Flt intercept3 10 70% 0% 10% 0%
MiniWinkler 250 20% 0% 0% <1%
1070m
Berlese 104 0% 0% 0% 0%
Malaise 20 5% 0% 0% 0%
Sweepnet 34 0% 0% 0% 0%
Flt intercept 10 0% 0% 0% 0%
MiniWinkler 150 0% 0% 0% 0%
2000m Sampling effort similar to 1070m; Wasmannia absent

1. Each fogging sample is one fogged tree with 40 1m2 funnels.
2. Each sample is 10-week catch of one Malaise trap.
3. Each sample is 6-weeks catch of one flight-intercept trap.
Figure 1. The five most widely distributed Wasmannia species. Views from left to right are face,
lateral mesosoma, and lateral petiole. Scale bars in face and mesosoma views are 0.5 mm, in
petiole views 0.1 mm.
Figure 2. Queens of Wasmannia auropunctata occur in two size classes. In cluster of queens
with smaller head size, one is from Jamaica and one from Venezuela. All others (n=14) are from
Costa Rica.
Figure 3. Relationship of queen head width to worker head width among Wasmannia species.
Average values are shown, based on data in Table 1 and Figure 2. Wasmannia auropunctata is
plotted twice, with different values for the dimorphic queens. A regression of queen head width
on worker head width has a negative slope, r2=0.50, p<0.05 (removing the large-headed W.
auropunctata queens renders the regression nonsignificant).
McArthur, A. J. 2007. A key to Camponotus Mayr of Australia, pp. 290-351. In Snelling, R. R.,
B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae):
homage to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological
Institute, 80.
A KEY TO CAMPONOTUS MAYR OF AUSTRALIA
Archie J. McArthur
Honorary Research Associate
South Australian Museum
North Terrace
Adelaide, SA 5000, Australia
mcarthur.archie@saugov.sa.gov.au
ABSTRACT
This key has been constructed to allow identification of 91 species and 10 subspecies of
Camponotus workers found in Australia. Species recognized in material examined have been
divided into 23 groups and are recorded in a checklist of valid and invalid names, authors, year
described and locations of type specimens. Abbreviations are listed for type locations and
measurements. A key to the 23 groups is provided followed by a key to each species. Brief
diagnoses of the species and distribution maps of material examined are located throughout the
key. Diagnoses and keys to the species are placed on adjacent pages for user benefit. A table of
relevant character states is provided in the Appendix. Described species not included in the key
are recorded in the Appendix. This key is based on characters selected for user convenience and
no attempt has been made to correlate the groups with their phylogeny.
Key words: Taxonomy, Hymenoptera, Formicidae, Formicinae, Camponotus, distribution

McArthur: Key to Australian Camponotus 291
INTRODUCTION
In this paper, a key is presented to allow identification of workers from the species and
subspecies of the ant genus Camponotus found in Australia. Literature is now available for
classifying Australian ants to subfamily and genus (Bolton, 1995; Shattuck, 1999). Little is
known of the biology of Camponotus ants even though species inhabit all parts of Australia
occurring in variable size populations. Species of the macrocephalus group nest in vegetation
where major workers with their flat truncated heads guard the entrance to their nest (McArthur
& Shattuck, 2001), while other species are known to nest in the ground. Many are nocturnal
foragers but some forage diurnally. Some are known as “sugar ants” (Froggatt, 1905), some are
known to have mutual relationships with butterflies (Braby, 2000), to assist in revegetating
disturbed sites by transporting seed (McArthur, 2003b), and to have lived for 23 years (Haskins,
1992). Some notes on biology have been recorded (Greenslade, 1979; Shattuck, 1999). The
taxonomy of the genus Camponotus is complicated by the wide variation in color, profile and
integument of the worker castes within a species. However pilosity is a good character for
matching collections of major, medium and minor worker castes because the number and
placement of setae (seta, singular; a sclerotized hair like projection, not to be confused with
spines on the inner margins of tibiae) is relatively consistent across workers in most Australian
species.
Depositories of Type Material
AMS = Australian Museum, Sydney, Australia; ANIC = Australian National Insect

Collection, Canberra, Australia; BMNH = The Natural History Museum, London, U.K.; MCG =
Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy; MCZ = Museum of
Comparative Zoology, Harvard, Cambridge, Massachusetts, U.S.A.; MHNG = Muséum
d'Histoire Naturelle, Geneva, Switzerland; NMHW = Naturhistorisches Museum, Vienna,
Austria; NHMB = Naturhistorisches Museum, Basel, Switzerland; NMV = Museum of Victoria,
Melbourne, Australia; OUM = Oxford University Museum, Oxford, U.K.; SAMA = South
Australian Museum, Adelaide, Australia; ZMB = Museum fur Naturkunde an der Universität
Humboldt zu Berlin, Germany.
Measurements.
Terms are illustrated in Fig. 1. All measurements are expressed in millimetres.
Declivity (D). Length from the center of angle to junction of propodeum with pedicel.
Eye length (EL). Maximum longitudinal length, in dorsal view.
Frontal carinae width (FCW). Maximum distance between carinae, in dorsal view.
Head length (HL). Maximum length measured along center-line of head from anterior margin of
clypeus to posterior margin of head, in dorsal view.
Head width (HW). Maximum head width in dorsal view.
Propodeal dorsum (PD). Length from center of angle to anterior margin of the propodeum.
Pronotal width (PW). Maximum width of pronotum in dorsal view.
Scape Index (SI) Ratio scape length / head width.
Scape length (SL).The straight-line length from base, exclusive of basal condyle, to apex.
Fig. 1. Camponotus worker. Morphology of the head in dorsal view, leg in front view, body in
lateral view and some terms used in the text.
TAXONOMIC HISTORY AND CLASSIFICATION
The genus Camponotus has been placed in the subfamily FORMICINAE whose members
possess a distinctive acidopore (appearing as a cone fringed with fine setae in Fig. 2), a single
node on the pedicel and lacking a sting. Camponotus can be confused with members of the
subfamily DOLICHODERINAE where the acidipore is replaced by a slit as shown in Fig.3.
Fig. 2. Side view of C. loweryi showing Fig. 3. Posterior view of Dolichoderus

acidopore typical of subfamily sp. typical of subfamily
FORMICINAE. DOLICHODERINAE.
Camponotus and Colobopsis were described as separate genera (Mayr, 1861). Colobopsis
was proposed as a subgenus of Camponotus (Emery, 1889) and that union has been accepted.
Species of Camponotus of the world have been divided into numerous groups of species named
subgenera (Emery, 1925) and in particular for ‘Ancien Continent et Australie’ where 21 groups
were described. Only two of Emery’s groups, viz. Colobopsis (as macrocephalus) and nigriceps
have been continued herein as groups 2 and 3. Characters defining groups 2, 3 and 4 have been
established previously (McArthur & Adams, 1996; McArthur & Shattuck, 2001; Shattuck &
McArthur, 2002) and incorporated into this key, while 20 other groups are proposed for the first
time.
Over the last 185 years, 163 species and subspecies of Camponotus have been described
from Australia. Of these, 28 are invalid names through synonymy or unavailability of a name,
leaving a total of 135 valid species and subspecies (Bolton, 1995). Based on the material
examined by the author from over 5000 localities, 101 species and subspecies are recognized. Of
the 34 taxa not recognized in this study, 6 have types which are believed to be lost (Appendix 1).
Twenty nine subspecies have been described from Australia although the concept is
generally considered undesirable. Already 12 of these subspecies have been raised to species
rank; three of which resulted from morphological and molecular studies (McArthur & Adams,
1996; McArthur, Adams & Shattuck, 1997) and seven from rigorous morphological examination
(McArthur & Shattuck, 2001; Shattuck & McArthur, 2002). It is outside the scope of this paper
to revise the status of the 10 subspecies included in this key. A lectotype has been designated for
C. darlingtoni (McArthur, Adams & Shattuck, 1997).
This key has been constructed to allow identification of 91 species and 10 subspecies of
Camponotus workers found in Australia. Species recognized in material examined have been
divided into 23 groups and are recorded in a checklist of valid and invalid names, authors, year
described and locations of type specimens. Abbreviations are listed for type locations and
measurements. A key to the groups is provided followed by a key to each species. Brief
diagnoses of the species and distribution maps of material examined are located throughout the
key. Diagnoses and keys to the species are placed on adjacent pages for user benefit. A table of
characters is provided as an appendix. This key is based on characters selected for the user’s
convenience and no attempt has been made to correlate the groups with their phylogeny. This
key should be considered temporary as it will require amendment following the anticipated
finding of new species. Twenty one new species have been described during the last eight years.
Previously, definitions of the genus Camponotus (Bolton, 1995; Shattuck, 1999) in Australia
stipulated the absence of a visible metapleural gland (a source of anti-bacterial secretions).
Camponotus thadeus Shattuck 2004 possesses a metapleural gland and its inclusion in this key is
justified since Camponotus gigas (a common ant of South East Asia) does possess such a gland
and is classified as Camponotus. This emphasises the need for a global review of the characters
defining the genus Camponotus.
Because minor workers comprise the caste most likely to be collected in the field, this key is
based on characters of minor workers where possible. However in a few pairs of species, minor
workers appear so morphologically similar, it is impossible to find characters to separate them.
In these cases it is fortunate that the major workers have distinctive characters for determining
identification. On the other hand there are a few pairs of morphologically similar species with
workers which can only be separated by color so far. In these and many other cases, it would be
advantageous to determine genetic distance by molecular techniques, one of which was
employed in the study by McArthur & Adams (1996).
Characters in the key refer to minor workers, unless otherwise stated.
CHECK LIST OF CAMPONOTUS SPECIES INCLUDED IN THE KEY
The 101 species and subspecies have been recognized in material examined in this study and are
assigned to groups numbered 1 to 23. Valid species names are recorded in bold type (invalid in
not-bold).
1, aureopilus group
thadeus Shattuck, 2004 (ANIC)
2, macrocephalus group
anderseni McArthur & Shattuck, 2001 (SAMA)
annetteae McArthur & Shattuck, 2001 (ANIC)

conithorax Emery, 1914 (MCG)
gasseri Forel, 1894 (MHNG)
= gasseri coloratus Wheeler, 1934
= gasseri lysias Forel, 1913
= gasseri obtusitruncatus Forel, 1902
howensis Wheeler, 1927 (MCZ)
janeti Forel, 1895b (MHNG)
janforrestae McArthur & Shattuck, 2001 (ANIC)
mackayensis Forel, 1902 (MHNG)
= reticulatus mackayensis Forel, 1902
macrocephalus Erichson, 1842 (ZMB)
= fictor Forel, 1902
= factor augustulus Viehmeyer, 1925
= semicarinatus Forel, 1895b
sanguinifrons Viehmeyer, 1925 (ZMB)
vitreus Smith, 1860 (OUM)
= nitida Smith, 1859
3, nigriceps group
clarior Forel, 1902 (MHNG)
= nigriceps clarior Forel, 1902
consobrinus Erichson, 1842 (ZMB)
= consobrinus dimidiatus Roger, 1863
= consobrinus obniger Forel, 1902
dryandrae McArthur & Adams, 1996 (SAMA)
eastwoodi McArthur & Adams, 1996 (SAMA)
longideclivis McArthur & Adams, 1996 (SAMA)
loweryi McArthur & Adams, 1996 (SAMA)
nigriceps Smith, 1858 (BMNH)
= nigriceps perthiana Wheeler, 1933
pallidiceps Emery, 1887 (MCG)
= nigriceps pallidiceps Emery, 1887
prostans Forel, 1910 (MHNG)
4, wiederkehri group
arenatus Shattuck & McArthur, 2002 (ANIC)
aurocinctus Smith, 1858 (BMNH)
= midas Froggatt, 1896
ceriseipes Clark, 1938 (NMV)
donnellani McArthur & Shattuck, 2002 (ANIC)
gouldianus Forel, 1922 (MHNG)
owensae Shattuck & McArthur, 2002 (SAMA)
perjurus Shattuck & McArthur, 2002 (ANIC)
postcornutus Clark, 1930a (NMV)
prosseri Shattuck & McArthur, 2002 (ANIC)
rufonigrus Shattuck & McArthur, 2002 (ANIC)
setosus Shattuck & McArthur, 2002 (ANIC)
terebrans Lowne, 1865 (BMNH)
= latrunculus victoriensis Santschi, 1928
= myoporus Clark, 1938
= testaceipes Smith, 1858
versicolor Clark, 1930a (NMV)
wiederkehri Forel, 1894 (MHNG)

= denticulatus Kirby, 1896; Clark 1930b (worker redescribed)
= latrunculus lucidor Forel, 1910
= latrunculus Wheeler, 1915
5, ephippium group
afflatus Viehmeyer, 1925 (ZMB)
capito Mayr, 1876 (NHMW)
capito ebinithorax Forel, 1915 (MHNG)
dromas Santschi, 1919 (NHMB)
ephippium narses Forel, 1915 (MHNG)
ephippium Smith, 1858 (BMNH)
fieldellus Forel, 1910 (MHNG)
tasmani Forel, 1902 (MHNG)
6, whitei group
leae Wheeler, 1915 (SAMA)
whitei Wheeler, 1915 (SAMA)
= scutellus Clark, 1930a
7, johnclarki group
johnclarki Taylor, 1992 (NMV)
= Notostigma sanguinea Clark, 1930a
8, intrepidus group
intrepidus Kirby, 1819 (BMNH)
= agilis Smith, 1858
= magnus Mayr, 1862
intrepidus bellicosus Forel, 1902 (MHNG)
piliventris Smith, 1858 (BMNH)
suffusus Smith, 1858 (BMNH)
= schenki Mayr, 1862
9, chalceus group
chalceus Crawley, 1915b (BMNH)
fergusoni McArthur, 2003a (SAMA)
hartogi Forel, 1902 (MHNG)
= ferruginipes Poulton & Crawley, 1922
innexus Forel, 1902 (MHNG)
pawseyi McArthur, 2003a (SAMA)
10, sponsorum group
froggatti Forel, 1902 (MHNG)
sponsorum Forel, 1910 (MHNG)
11, aeneopilosus group
aeneopilosus Mayr, 1862 (NHMW)
nigroaeneus Smith, 1858 (BMNH)
12, subnitidus group
dorycus confusus Emery, 1887 (MCG)
rufus Crawley, 1925 (OUM)
subnitidus Mayr, 1876 (NHMW)
tricoloratus Clark, 1941 (NMV)
13, maculatus group
extensus Mayr, 1876 (NHMW)
fieldeae Forel, 1902 (MHNG)
maculatus humilior Forel, 1902 (ANIC)
novaehollandiae Mayr, 1870 (NHMW)
oxleyi Forel, 1902 (MHNG)

spenseri Clark, 1930b (NMV)
= reticulatus Kirby, 1896
villosus Crawley, 1915a (BMNH)
14, darlingtoni group
cinereus amperei Forel, 1913 (MHNG)
claripes elegans Forel, 1902 (MHNG)
darlingtoni Wheeler, 1934 (MCZ)
= rottnesti Donisthorpe, 1941
inflatus Lubbock, 1880 (not known)
= aurofasciatus Wheeler, 1915
pitjantjatarae McArthur, 2003a (SAMA)
15, scotti group
cinereus Mayr, 1876 (NHMW)
claripes marcens Forel, 1907b (MHNG)
longifacies McArthur, 2003a (ANIC)
scotti McArthur, 2003a (SAMA)
scratius Forel, 1907b (MHNG)
16, esau group
arcuatus Mayr, 1876 (NHMW)
esau Forel, 1915 (MHNG)
17, gibbinotus group
cowlei Froggatt, 1896 (AMS)
discors Forel, 1902 (MHNG)
gibbinotus Forel, 1902 (MHNG)
18, walkeri group
adami Forel, 1910 (MHNG)
walkeri Forel, 1893 (MHNG)
19, armstrongi group
armstrongi McAreavey, 1949 (NMV)
macareaveyi Taylor, 1992 (NMV)
= sanguinea McAreavey, 1949
20, evae group
evae Forel, 1910 (MHNG)
rudis McArthur, 2003a (ANIC)
tristis Clark, 1930a (NMV)
21rubiginosus group
evae zeuxis Forel, 1915 (MHNG)
lownei Forel, 1895a (MHNG)
rubiginosus Mayr, 1876 (NHMW)
simpsoni McArthur, 2003a (SAMA)
22, oetkeri group
michaelseni Forel, 1907b (MHNG)
oetkeri Forel, 1910 (MHNG)
23, claripes group
claripes Mayr, 1876 (NHMW)
claripes minimus Crawley, 1922 (OUM)
eremicus Wheeler, 1915 (SAMA)
KEY TO GROUPS OF SPECIES
All characters in the key refer to minor workers, unless otherwise stated.
KEY TO SPECIES
1, aureopilus group
Camponotus thadeus Shattuck

Sole species in group. Worker. HW 2.00 - 3.20; HL 2.15 - 2.80;
PW 1.85 - 2.15. Black; mesonotum, propodeum and anterior gaster
with dense bright yellow setae up to 1 mm long, scapes and tibiae
with short setae raised to 20°, no erect setae on underside of head;
mesosoma finely reticulate, head finely punctate; metapleural gland
opening 0.25 - 0.27; head sides straight tapering forward, widest
behind eyes. Major worker. Vertex feebly concave; eyes distant from
corners by more than EL; anterior clypeal margin feebly projecting,
median section concave between two rounded convexities. Minor worker. Anterior clypeal
margin feebly projecting, mostly straight; vertex straight; eyes near corners.
2, macrocephalus group
E
Camponotus annetteae McArthur & Shattuck

Worker. HW 1.40; HL 1.50; PW 1.40. Major worker not yet
described. Minor worker. Entirely red; mesosoma, under head, scapes
and tibiae covered with dense, erect, short setae < EL; propodeal dorsum
nearly hemispherical; node summit sharp, pointing forward in lateral
view, concave in rear view; vertex slightly convex in front view, angular
in lateral view; clypeus convex in all directions, anterior margin convex.
Camponotus janeti Forel

Worker. HW 0.95 - 1.70; HL 1.00 - 2.00; PW 0.70 - 1.50. Head,
mesosoma, legs dark brown; mesosoma, under head covered with dense
erect setae nearly EL; scapes and tibiae with short setae raised to 45°;
eyes looking sideways. Major worker. Head sides straight; vertex
convex; anterior clypeal margin, feebly concave; front of head coarsely
punctate, straight anterior separated from posterior by a curve; propodeal
dorsum straight, angle nearly 135°; node summit blunt. Minor worker.
Head sides straight, tapering forward; vertex convex; anterior clypeal margin projecting feebly
convex; front of head finely punctate, not truncate; propodeal dorsum straight, angle well
rounded nearly 150°; metanotum distinct.
Camponotus mackayensis Forel

Worker. HW 1.00 - 1.15; HL 1.00 - 1.70; PW 0.80 - 1.05. Red-
brown, pronotum lighter than mesonotum; propodeal dorsum a shallow
concavity; long erect setae on most surfaces including scape. Major
worker. Head sides straight; vertex straight, anterior half coarsely
punctate, clypeus oval, widest at truncation, anterior margin narrow;
node summit blunt; metanotum a wide shallow groove. Minor worker.
Head sides anterior half tapering forward; vertex slightly convex
between widely rounded corners; anterior clypeal margin projecting, convex; propodeal dorsum
straight, angle well rounded nearly 150°; metanotum a distinct vee; propodeal dorsum anterior
inclined upward to a ridge then shallowly concave, then a wide angle; node summit rounded.
Camponotus janforrestae McArthur & Shattuck

Worker. HW 1.6; HL 1.8; PW 1.2. Major worker not yet described.
Minor worker. Black covered in long and short erect white setae
including under head, scapes and tibiae; propodeum a hemispherical
hump; node summit rounded; head sides straight tapering forward;
vertex rounded; clypeus wide, glossy, anterior margin feebly projecting,
in center a narrow concavity bounded on each side by a tooth.
Camponotus vitreus Smith

Worker. HW 0.85 - 1.55; HL 0.85 - 1.55; PW 0.60 - 1.05. Black to
dark brown; glossy; propodeum hemispherically domed; all surfaces
excepting under head covered with erect long setae; node summit sharp;
dimorphic. Major worker. Head sides straight, feebly tapering forward;
clypeus tapering to a narrow convex anterior margin, in lateral view
anterior clypeus straight, smoothly truncated from convex posterior
clypeus. Minor worker. Head sides straight tapering forward; clypeus wide, anterior margin
convex projecting.
Camponotus conithorax Emery

Worker. HW 1.5 - 2.1; HL 1.6 - 2.3; PW 1.1 - 1.3. Dark brown;
glossy; metanotal spiracles well below dorsum; erect setae mostly
confined to gaster. Major worker. Head sides straight, nearly parallel;
clypeal median portion wide, anterior margin projecting, bounded
laterally with angles. Minor worker. Head sides straight, tapering
forward, clypeal median portion wide, sides tapering rearward, bounded
laterally with angles, feebly concave between.
Camponotus gasseri Forel

Worker. HW 0.9 - 1.6; HL 1.0 - 1.7; PW 0.65 - 1.2. Black to brown;
glossy; anterior head often red; metanotum with spiracles above dorsum;
propodeum hemispherically domed; erect setae confined to gaster and
front of head; node summit rounded; dimorphic. Major worker. Head
sides straight, feebly tapering forward; anterior head striate, truncate;
clypeus tapering to narrow convex anterior margin. Minor worker. Head
sides straight, tapering forward; vertex flattish; clypeus wide, anterior
margin convex, projecting; anterior head finely punctate.
Camponotus sanguinifrons Viehmeyer

Worker. HW 0.75 - 1.10; HL 0.85 - 1.40; PW 0.45 - 0.55. Major
worker. Dark brown, mandibles and legs lighter; mesonotum a gentle
curve; head sides straight, parallel; clypeus, mandibles and cheeks
forming flat circular area, coarsely punctate, with few coarse longitudinal
striations; plentiful, short, erect, club-like setae; clypeal lateral margins
widest at centre of circular area; anterior margin short, straight. Minor
worker. A few long erect setae under head and few more on gaster,
sparse elsewhere; propodeal dorsum mostly straight and sloping downward; head sides straight,
tapering slightly to front; anterior margin projecting, evenly convex, wide.
Camponotus anderseni McArthur & Shattuck

Worker. HW 0.80 - 1.20; HL 0.90 - 1.30; PW 0.60 - 0.90. Light
brown; glossy; dorsum of mesosoma flattish, metanotum distinct;
mesosoma, under head lacking erect setae, slightly raised on tibiae; eyes
elongate looking outward, nearer to mandibles than vertex; dimorphic.
Major worker. Clypeus depressed, finely punctate, anterior margin
convex. Minor worker. Clypeus wide, anterior margin convex,
projecting.
Camponotus macrocephalus Erichson

Worker. HW 0.90 - 1.70; HL 1.10 - 1.90; PW 0.70 - 1.20. Yellow brown; glossy; dorsum of
mesosoma evenly flatly convex; erect setae confined to front of head and
gaster; eyes placed near vertex; dimorphic. Major worker. Head sides
straight, parallel; vertex straight; anterior head sharply truncated, deeply
striate lengthwise and at 45°; clypeal median section clearly edged,
parallel, anterior margin convex and posterior to mandible insertions.
Minor worker. Head sides mostly straight, slightly tapering forward;
vertex rounded; clypeus not truncate, anterior clypeal margin convex,
projecting past mandibular insertions.
Camponotus howensis Wheeler

Black, otherwise similar to macrocephalus, confined to Lord
Howe Is.
3, nigriceps group
Camponotus loweryi McArthur & Adams

Worker. HW 1.60 - 4.05; HL 2.20 - 4.25; PW 1.50 - 2.50.
Whole of gaster uniform in color; under head lacking erect setae,
mesosoma with golden erect more plentiful on pronotum than
propodeum, all pointing forward, tibiae and scapes with short
setae raised to < 5°; mesosoma dorsum evenly convex; node
sharp; finely reticulate; short flat-lying setae sparse; polymorphic.
Major worker. Head sides convex, tapering forward; metanotum
distinct, narrow; eyes flat. Minor worker. Head sides straight,
parallel; vertex convex; metanotum absent; eyes bulbous.
Camponotus consobrinus Erichson

Worker. HW 1.30 - 3.30; HL 1.75 - 3.40; PW 1.4 - 2.30.
Under head lacking erect setae, mesosoma with a few scattered
golden erect setae, all pointing forward; tibiae and scapes with
short setae raised to < 5°; anterior gaster lighter color than
posterior; mesosoma dorsum evenly convex; node sharp; finely
reticulate; short flat-lying setae sparse; polymorphic. Major
worker. PD/D about 1.5; head sides convex, tapering forward;
metanotum distinct, narrow; eyes flat. Minor worker. PD/D about
2.5; head sides straight parallel; vertex convex; metanotum
absent; eyes bulbous.
Camponotus longideclevis McArthur & Adams

Worker. HW 1.70 - 3.50; HL 2.20 - 3.30; PW 1.5 - 1.90.
Under head lacking erect setae, mesosoma with a few scattered
golden erect setae, all pointing forward, short setae raised to > 5°
on tibiae and to < 5 ° on scapes; mid-tibia inside with > 8 spikes;
anterior gaster lighter color than posterior; mesosoma dorsum
evenly convex; node sharp; finely reticulate; short flat-lying setae
sparse; polymorphic. Major worker. PD/D about 1.2; head sides
convex, tapering forward; metanotum distinct, narrow; eyes flat.
Minor worker. PD/D about 1.5; Head sides straight parallel;
vertex convex; metanotum absent; eyes bulbous.
Camponotus dryandrae McArthur & Adams

Worker. HW 1.70 - 3.90; HL 2.40 - 3.50; PW 1.60 - 2.5.
Under head with a few long erect setae, mesosoma with scattered
golden long fine erect setae, on propodeum confined to near
angle; mesosoma with plentiful flat-lying short setae; tibiae with
short setae raised to 30°; scapes raised to 10°; mesosoma dorsum
evenly convex; node summit blunt; finely reticulate; polymorphic.
Major worker. PD/D about 1.5; head sides convex, tapering
forward; metanotum marked by fine lines; eyes flat. Minor
worker. PD/D about 3; Head sides straight, parallel; vertex
convex; metanotum absent; eyes bulbous.
Camponotus clarior Forel

Worker. HW 1.90 - 3.90; HL 2.50 - 3.50; PW 1.6 - 2.2.
Mostly yellow; under head with plentiful long erect fine setae,
mesosoma with scattered golden long fine erect setae (more
plentiful anteriorly) interspersed with plentiful flat-lying short
setae; tibiae and scapes with short setae raised to 30°; mesosoma
dorsum evenly convex; node summit blunt; finely reticulate;
polymorphic. Major worker. PD/D about 1.5; head sides convex,
tapering forward; eyes flat; metanotum feeble. Minor worker.
PD/D 3; Head sides straight parallel; vertex convex; metanotum
absent; eyes bulbous.
Camponotus nigriceps Smith

Worker. HW 1.60 - 4.10; HL 2.20 - 4.10; PW 1.4 - 2.6. Black
head; under head with plentiful long erect fine setae, mesosoma
with scattered golden long fine erect setae, plentiful flat-lying
short setae; tibiae and scapes with short setae raised to 30°;
mesosoma dorsum evenly convex; node summit sharp; finely
reticulate; polymorphic. Major worker. PD/D about 1.5; head
sides convex, tapering forward; metanotum marked by fine lines;
eyes flat. Minor worker. PD/D about 3; head sides straight
parallel; vertex convex; metanotum absent; eyes bulbous.
Camponotus eastwoodi McArthur & Adams

Worker. HW 1.60 - 4.10; HL 2.20 - 4.10; PW 1.4 - 2.6.
Under head with a few long erect fine setae, more in minor
workers; mesosoma with scattered golden long fine erect setae
and plentiful flat-lying short setae; tibiae and scapes with short
setae raised to 30°; mesosoma dorsum evenly convex; node
summit sharp; finely reticulate; polymorphic; head and mesosoma
black or dark brown. Major worker. PD/D about 1.5; head sides
convex, tapering forward; metanotum marked by fine lines; eyes
flat. Minor worker. PD/D 3; head sides straight, parallel; vertex convex; metanotum absent; eyes
bulbous.
Camponotus prostans Forel

Worker. HW 1.90 - 2.00; HL 2.60 - 3.60; PW 1.5 - 2.2. Black
head; under head with a few long erect setae, more with minor
workers; mesosoma with a few scattered golden long fine erect
setae; mesosoma with plentiful flat-lying short setae; tibiae and
scapes with short setae raised to 30°; mesosoma dorsum evenly
convex; node summit sharp; finely reticulate; polymorphic. Major
worker. PD/D about 1.5; head sides convex, tapering forward;
metanotum marked by fine lines; eyes flat. Minor worker. PD/D
3; head sides straight, parallel; vertex convex; metanotum absent;
eyes bulbous.
Camponotus pallidiceps Emery

Worker. HW 1.60 - 4.10; HL 2.20 - 4.10; PW 1.4 - 2.6. Light
brown head; under head with a few long erect fine setae, more
with minor workers; mesosoma with scattered golden long fine
erect setae; mesosoma with plentiful flat-lying short setae; tibiae
and scapes with short setae raised to 30°; mesosoma dorsum
evenly convex; node summit sharp; finely reticulate;
polymorphic. Major worker. PD/D about 1.5; head sides convex,
tapering forward; metanotum marked by fine lines; eyes flat.
Minor worker. PD/D 3; head sides straight, parallel; vertex
convex; metanotum absent; eyes bulbous.
4, wiederkehri group
Camponotus perjurus Shattuck & McArthur

worker not yet described. Minor worker. Purplish, tending iridescent;
unique head, pronotum attached well below vertex, resembling
Iridomyrmex purpureus in attachment; propodeal dorsum concave; a
few scattered erect setae, none under head, on tibiae nor on scapes;
head sides straight, tapering forward; vertex nearly semicircular;
anterior clypeal margin projecting, convex; color mostly red-brown
with gaster darker.
Camponotus setosus Shattuck & McArthur

worker not yet described. Minor worker. Gaster black, most of head,
mesosoma and node black, remainder with red patches; antennae
dark brown; pronotum and mesonotum form together an even, raised
convexity followed by angular trough of metanotum and feebly
convex dorsal surface of propodeum; entire body covered with dense
flat-lying setae, erect setae absent from antennae.
Camponotus gouldianus Forel

Worker. HW 1.6 - 4.7; HL 1.8 - 4.3; PW 1.1 - 2.7. Brown, legs
lighter; propodeum with more than 40 erect short and long setae;
abundant, overlapping, short, flat-lying setae on head and gaster;
mesosoma flat and long; propodeal angle rounded. Major worker.
Anterior clypeal margin nearly straight with angular corners; node
summit blunt. Minor worker. Anterior clypeal margin feebly convex,
strongly projecting, crenulate, anterior corners with wide angles;
PD/D 3; node summit bluntly rounded.
Camponotus terebrans Lowne

Worker. HW 1.1 - 3.6; HL 1.3 - 3.28; PW 0.9 - 2.0. Head red-
brown to black, mesosoma and node yellow to brown, gaster darker
than mesosoma, legs lighter; propodeum with 10 to 25 erect setae;
sparse short flat-lying setae on head and gaster; node summit
angular. Major worker. Medial section of anterior clypeal margin
straight, with rectangular corners; propodeum straight, angle well
rounded; node summit sharp. Minor worker. Anterior clypeal
margin medial section convex, strongly projecting; dorsal
propodeum surface straight; node summit blunt.
Camponotus postcornutus Clark

Worker. HW 2.0 - 4.9; HL 1.9 - 4.2; PW 1.7 - 3.2. Red brown,
gaster darker; glossy; posterior corners of head in larger workers
taper rearward into blunt protuberances; mesosoma dorsum evenly
convex; angle rounded; front and under head, mesosoma, petiole,
gaster and coxae with sparse reddish, long erect setae; polymorphic.
Major worker. Medial section of anterior clypeal margin mostly
straight, feebly projecting anteriorly with broad lateral angles, carina
distinct; node summit moderately sharp. Minor worker. Anterior
clypeal margin projecting feebly, clypeal carina sharp; PD/D about 2; node summit bluntly
rounded.
Camponotus arenatus Shattuck & McArthur

Worker. HW 1.5 - 1.8; HL 1.9 - 2.2. Major worker not yet
described. Minor worker. Pronotum anterior regions dark red to
black, distinctly darker than mesonotum and propodeum; metanotal
groove depressed below level of anterior region of propodeum; node
anterior face much shorter than posterior face; tibiae and scapes
lacking erect setae; anterior clypeal margin broadly convex;
propodeum lacking a distinct angle, PD/D about 1.5; node summit
broadly convex; erect setae on all surfaces of head and mesosoma,
node and gaster, absent from scapes and tibiae.
Camponotus owensae Shattuck & McArthur

Worker. HW 1.6 - 2.0; HL 2.0 - 2.4; PW 1.4 - 1.60. Major worker not yet described. Minor
worker. Entirely black; propodeum with a wide concavity and a
posterior hump; metanotal groove depressed below level of anterior
region of propodeum; node summit long and flat, its anterior face
much shorter than posterior; setae on tibiae raised to 20°, none visible
on scapes, otherwise overall plentiful white flat-lying, with a few
erect; anterior clypeal margin projecting bounded by rounded angles.
Camponotus aurocinctus Smith

Worker. HW 1.6 - 4.9; HL 2.0 - 4.1; PW 1.5 - 2.6. Mesosoma
uniform in color, red to black, first gastral tergite anterior similar in
color to propodeum; metanotal groove depressed below level of
anterior region of propodeum; propodeum with a gentle angle; erect
setae absent from scapes, tibiae, node, sparse on head, mesosoma and
gaster, 0 to 30 under head; finely and closely punctate; polymorphic.
Major worker. Anterior clypeal margin feebly convex; node summit
varying from a broad, blunt angle to uniformly convex. Minor worker. Anterior clypeal margin
convex; node summit elongate.
Camponotus versicolor Clark

Worker. HW 1.8 - 3.4; HL 2.2 - 3.2; PW 1.4 - 2.5. First two
gastral tergites distinctly lighter in color than propodeum, gastral
tergites never with golden bands, otherwise similar to aurocinctus.
Camponotus wiederkehri Forel

Worker HW 1.2 - 3.6; HL 1.5 - 3.3; PW 0.9 - 2.2. Yellow-red to dark
brown, gaster darker; posterior section of mesonotum flat (or nearly
so) immediately anterior of metanotal groove; erect setae plentiful,
scattered, less on propodeal angle and declivity, absent from scapes,
flat-lying on tibiae. Major worker. Anterior clypeal margin strongly
projecting, straight, corners broadly angular; pronotum and
mesonotum slightly raised, evenly convex; anterior section of
propodeal dorsum feebly concave anteriorly and feebly convex
posteriorly, angle widely rounded. Minor worker. Clypeus convex;
anterior section of mesonotum feebly convex, remainder joins with propodeal dorsum to form a
long flat surface ending in a widely rounded angle; PD/D about 3; entire body clothed with fine
flat-lying short setae.
Camponotus donnellani Shattuck & McArthur

Worker. HW 1.40; HL 1.58. Major worker not yet described.
Minor worker. Head, mesosoma and node red with upper surfaces of
head, pronotum and sometimes mesonotum with blotches of darker
color; propodeum with at most 4 elongate erect setae near angle;
anterior propodeal dorsum feebly concave, posterior straight; node
summit broadly rounded; head sides nearly parallel; vertex rounded;
anterior clypeal margin feebly projecting, broadly convex; long setae
scattered on all surfaces, absent from scapes and tibiae; glossy.
Camponotus rufonigrus Shattuck & McArthur

Worker. HW 1.1 - 1.4; HL 1.3 - 1.6; PW 0.9 - 1.2. Major worker
not yet described. Minor worker. Black head contrasting with red
mesonotum; propodeum with more than 10 erect setae scattered;
pronotum and mesonotum evenly convex; metanotum indistinct;
propodeum concave anteriorly, flat posteriorly, angle rounded, PD/D
about 1.5; anterior clypeal margin evenly convex, carina
conspicuous; dorsal and under surfaces of head, mesosoma, petiole,
gaster and coxa with sparse long erect setae; entire body with short
indistinct flat-lying short setae; tibiae and scapes lacking erect setae.
Camponotus ceriseipes Clark

Worker. HW 1.2 - 4.1; HL 1.4 - 3.4; PW 1.0 - 2.5. Head same
color as mesonotum (red to black); posterior mesonotum feebly
convex immediately anterior of metanotal groove (more so in
minors), node broadly rounded above; tibiae and scapes lacking erect
setae, propodeum with > 10 erect setae, scattered on dorsum;
(generally node is higher and narrower, integument shiny in
ceriseipes compared with prosseri.) Scapes relatively short in minors,
SI < 1.5. Major worker. Metanotum distinct, propodeal dorsum
feebly convex, sometimes slightly stronger near metanotum; angle well rounded; node summit
rounded, posterior surface feebly concave near summit; anterior clypeal margin broadly convex
across entire width, scarcely projecting, with a weak carina; head, mesosoma, node and gaster
with scattered long setae. Minor worker. Anterior propodeal region feebly concave, posterior
region straight, angle distinct and widely rounded, PD/D nearly 2; node summit rounded; head,
mesosoma, petiole and gaster with scattered long setae; anterior clypeal margin convex, carina
distinct. (This species can be confused with prosseri; in larger minor workers of ceriseipes scape
is relatively short, but this difference is minimal in smaller workers.)
Camponotus prosseri Shattuck & McArthur

Worker. HW 1.0 - 3.9; HL 1.5 - 3.2; PW 1.0 - 2.4. Similar to ceriseipes except in scape
length of larger minor workers. In ceriseipes, SI < 1.5; whereas in
prosseri, SI > 1.4.
5, ephippium group
Camponotus ephippium narses Forel

Worker. HW 1.1 - 2.8; HL 1.5 - 2.8; PW 1.2 - 1.8. Black, legs
brown; setae on tibiae flat-lying, none visible on scapes, no erect
setae under head, plentiful elsewhere. Major worker. Head front
feebly truncate, coarsely punctate; clypeus scarcely projecting. Minor
worker. Dorsum of mesosoma evenly convex; PD/D > 3; golden,
short flat-lying setae on gaster; median section of clypeal anterior
margin wide, strongly projecting, mostly straight; about 8 teeth.
Camponotus tasmani Forel

Worker. HW 1.5 - 3.5; HL 1.8 - 3.5; PW 1.3 - 2.4. Entirely
clothed in erect whitish setae about EL; red and black; propodeum
feebly concave. Major worker. Head sides feebly convex not
tapering; vertex straight swollen; frontal carinae wide (about 1/3
HW); anterior clypeal margin with two blunt teeth separated by a
shallow concavity; node summit blunt. Minor worker. Front of head
clothed in short white setae hiding integument; anterior clypeal
margin strongly projecting convex; node summit convex; PD/D about
3.5.
Camponotus ephippium Smith

Worker. HW 1.6 - 3.8; HL 1.9 - 3.6; PW 1.55 - 2.8. Variations of
red and black; mostly glossy; setae on tibiae distinct, short,
overlapping raised to 20°, none on scapes, long erect setae scattered
everywhere including under head. Major worker. Head sides mostly
straight, parallel; cheeks and vertex swollen, corners projecting
rearward; eyes small, flat, placed > 2 EL distant from corners;
metanotum a distinct trough; propodeal dorsum straight, angle 135°
rounded; anterior clypeal margin feebly projecting, median section
concave, bounded by two blunt teeth. Minor worker. Head sides nearly straight, parallel, corners
abrupt; eyes bulbous, large, < 0.5 EL from corners; propodeal dorsum concave; PD/D about 3;
node summit convex; most of ant covered with fine short white overlapping setae; about 8 teeth.
Camponotus afflatus Viehmeyer

worker not described. Minor worker. Red, gaster darker; glossy; head
sides straight, feebly tapering to front; no erect setae under head,
sparse elsewhere, fine short flat-lying setae scarcely visible on tibiae
and scapes; propodeal dorsum long, concave, PD/D about 3; angle
about 150°; clypeal anterior margin convex projecting; 6 teeth.
Camponotus fieldellus Forel

Worker. HW 1.6 - 3.2; HL 1.7 - 3.2; PW 1.3 - 2.2. Dark brown;
setae on tibiae flat-lying or at most raised to 5°, none on scapes, no
erect setae under head. Major worker. Head corners rounded, clypeus
scarcely projecting. Minor worker. Head sides straight, distinctly
tapering forward; head and mesosoma covered with white short flat-
lying setae, more golden on gaster; node summit convex.
Camponotus capito Mayr

Worker. HW 1.65 - 3.35; HL 1.85 - 3.20; PW 1.45 - 2.4. Red-
brown, gaster darker; sparse erect setae to 0.5 mm long, scattered on
mesosoma, mostly near angle, none under head, on scapes nor tibiae.
Major worker. Head sides straight, tapering to front; anterior clypeal
margin middle third strongly projecting, narrow, bounded by wide
angles, crenulate between; forehead swollen; metanotum distinct.
Minor worker. Anterior clypeal margin middle half strongly
projecting, bounded by right angles; pronotum and mesonotum uniformly humped, high.
Camponotus dromas Santschi

Worker. HW 1.4 - 3.3; HL 1.7 - 3.25; PW 1.35 - 2.4. Mesosoma
contrasting red and black, red legs; tibiae with flat-lying short
overlapping setae, a few long erect setae under head. Major worker.
Front of head feebly truncate; median section of anterior clypeal
margin projecting, narrow, concave, bounded by two blunt teeth.
Minor worker. Anterior mesosoma black contrasting with red
posterior; node summit convex, sometimes flattish; median section of
anterior clypeal margin wide, convex; about 8 teeth.
Camponotus capito ebinithorax Forel

Worker. HW 1.5 - 3.00; HL 1.6 - 2.95; PW 1.5 - 2.3. Red to
brown; sparse long setae scattered along mesosoma, plentiful under
head, flat-lying short and indistinct on scapes and tibiae. Major
worker. Median third is bounded by two projecting teeth with a
feebly crenulated concavity between; head sides posterior half
straight parallel, otherwise convex; vertex straight, node summit
sharp; eyes small, elongate. Minor worker. Head rectangular; anterior
clypeal margin projecting, convex; pro and mesonotum strongly
humped; propodeum deeply concave; plentiful white short flat-lying setae; eyes small, round.
6, whitei group
Camponotus whitei Wheeler

Worker. HW 1.40 - 2.70; HL 1.50 - 2.70; FCW 0.50 - 1.10; PW
1.15 - 1.75. Predominantly brown; entirely covered with plentiful
erect setae including scapes, tibiae and under head; frontal carinae
wide. Major worker. Head sides posterior straight, anterior tapering
forward; vertex straight; pronotum convex; mesonotum slightly
convex; metanotum distinct; propodeal dorsum humped; PD/D about
1; anterior clypeal margin median portion projecting, bounded by two blunt teeth with a deep
concavity between; node summit sharp. Minor worker. Head sides straight, tapering to front;
vertex straight; pronotum convex; mesonotum straight; metanotum a wide V; propodeal dorsum
domed; PD/D about 1; anterior clypeal margin median portion projecting, wide, slightly convex,
crenulate; node summit convex, higher than long; integument coarsely punctate.
Camponotus leae Wheeler

Worker. HW 1.05 - 2.90; HL 1.30 - 2.90; FCW 0.34 - 1.00;
PW 0.95 - 2.00. Red brown and brown combination; entirely
covered with plentiful erect setae (more on major than minor)
including scapes, tibiae and under head. Major worker (not
previously described). Head sides mostly straight, tapering
forward; vertex straight; forehead swollen; anterior clypeal
margin median portion not projecting, narrow (about 1/3 anterior
head width), bounded by two blunt teeth with a deep concavity
between; frontal carinae scarcely diverging, wide; integument of head finely reticulate;
pronotum convex; mesonotum straight; metanotum distinct; propodeal dorsum straight; PD/D
about 1; node summit sharp. Minor worker. Head sides straight, parallel; vertex convex;
pronotum convex; mesonotum straight; metanotum a wide V; propodeal dorsum straight; PD/D
about 2; anterior clypeal margin median portion projecting, wide, slightly convex, crenulate;
node summit flat, longer than high; integument of head finely and coarsely punctate.
7, johnclarki group
Camponotus johnclarki Taylor
Sole species in group. Worker. HW 1.8; HL 2.2; PW 1.3.

Reddish. Major worker. Head sides convex; vertex straight to
concave; frontal carinae narrow; anterior clypeal margin median
section feebly projecting, straight, crenulate, bounded by two
rounded angles; node leaning forward, summit convex. Minor
worker. Head sides straight, parallel; vertex convex; anterior
clypeal margin median section projecting, wide, convex;
pronotum, mesonotum, propodeum and under head each have
more than 20 short erect setae; scapes and tibiae with plentiful
setae raised to 30°; pronotum and mesonotum evenly convex; metanotum a wide groove;
propodeal dorsum nearly straight; angle well rounded; PD/D about 3; node summit long, flat,
inclined forward, highest in front; 8 or 9 teeth.
8, intrepidus group
Camponotus piliventris Smith

Worker. HW 1.6 - 3.5; HL 2.2 - 3.3; FCW 0.5 - 1.0; PW 1.3 - 2.0. Black to dark brown, legs
much lighter red-brown; gaster with golden short flat-lying
overlapping setae; anterior propodeum raised above metanotum;
pronotum and mesonotum evenly convex; scapes and tibiae with
plentiful erect setae; anterior clypeal margin median section
convex; head finely distinctly punctate, mesonotum more
distinctly reticulate; metanotum spiracle directed upward; node
anterior and posterior faces feebly convex, summit blunt. Major
worker. Head sides posterior half straight, parallel, anterior
tapering to front; vertex straight; propodeal dorsum slightly
convex, angle well rounded; PD/D about 1; front of head coarsely punctate. Minor worker. Head
sides feebly convex, tapering forward; vertex strongly convex; pronotum and mesonotum evenly
convex; propodeal dorsum slightly convex; angle well rounded; PD/D about 2; node summit
sharp.
Camponotus suffusus Smith

Worker. HW 2.2 - 3.8; HL 2.4 - 3.4; FCW 0.6 - 1.0; PW 1.6 -
2.3. Red-brown, legs nearly similar color; gaster with golden
short flat-lying, overlapping setae; scapes and tibiae with plentiful
erect setae; anterior clypeal margin median section projecting,
straight, bounded by angles; head finely punctate; node, anterior
and posterior faces straight, summit sharp. Major worker. Head
sides tapering forward; vertex straight; metanotum spiracle
directed upward; propodeal dorsum flat not raised above level of
metanotum; angle distinct about 135°; PD/D about 1.5. Minor
worker. Head sides feebly convex tapering forward, anterior clypeal margin median section
straight; vertex mostly straight; pronotum and mesonotum evenly convex; propodeal dorsum
flat, angle distinct, about 135°; PD/D about 1.8; node summit sharp.
Camponotus intrepidus Kirby

Worker. HW 2.3 - 4.6; HL 2.9 - 4.5; PW 1.8 - 2.6. Head and
gaster dark brown, mesosoma lighter reddish brown; anterior
mesosoma uniformly convex; scapes and tibiae with plentiful
erect setae; head finely punctate; gaster glossy with fine
striations, metanotum flat. Major worker. Head sides convex
tapering forward, head widest at eye level; vertex straight;
anterior clypeal margin median section convex; mandibles
striate; head finely punctate; propodeum predominantly flat;
angle well rounded; PD/D about 1.5, node thick, anterior face
convex, summit blunt, posterior face straight. Minor worker.
Head sides convex tapering forward; vertex flattish; anterior clypeal margin projecting, median
section convex; propodeum mostly convex; PD/D about 1.5.
Camponotus intrepidus bellicosus Forel

Very similar to intrepidus except head of major worker is widest
behind eyes, head of minor worker tapers rearward; vertex convex.
9, chalceus group
Camponotus chalceus Crawley

Worker. HW 1.5 - 2.4; HL 1.7 - 2.35; PW 1.2 - 1.6. Black and
red; finely punctate; pronotum and mesonotum evenly convex;
declivity feebly concave; erect setae about EL scattered over
mesosoma, more under and on front of head, absent on scapes, tibiae;
node summit convex. Major worker. Head sides strongly convex;
anterior clypeal margin middle half narrow, concave between two
teeth. Minor worker. Head sides tapering forward; middle half of
anterior clypeal margin nearly straight, projecting forward, bounded
by wide angles.
Camponotus fergusoni McArthur

Worker. HW 1.2 - 2.8; HL 1.4 - 2.8; PW 1.15 - 2.0. Mostly
black, propodeum with some red; finely punctate; propodeal dorsum
concave; mesosoma and node covered with white fine short flat-lying
short flat-lying setae. Major worker. Head sides straight, feebly
tapering forward; vertex straight; clypeus with a few fine long setae,
feebly projecting forward, anterior margin evenly concave; node
summit blunt. Minor worker. Head sides straight, tapering forward;
vertex convex; anterior clypeal margin median section wide, nearly straight, projecting forward;
dense, short flat-lying setae nearly hiding integument; pronotum, mesonotum and half
propodeum evenly convex; node with a long, feebly convex summit.
Camponotus pawseyi McArthur

Worker. HW 1.5 - 3.0; HL 1.6 - 3.0; PW 1.3 - 2.1. Color varies
between black and red; entirely covered with dense erect setae. Major
worker. Forehead swollen; anterior head (lateral view) feebly
truncated; clypeus coarsely punctate with a distinct carina, anterior
margin median section projects with a central concavity; mandibles
striate; propodeal dorsal surface flat; node summit bluntly pointed.
Minor worker. Head sides straight, eyes near corners of head (very
similar to tasmani) except vertex convex, anterior clypeal margin projecting convex; crenulate; 7
or 8 teeth; entirely covered with plentiful erect setae, flat-lying setae hiding integument of
clypeus; propodeal dorsum deeply concave; node summit convex.
Camponotus hartogi Forel

Worker. HW 1.3 - 2.1; HL 1.55 - 2.2; PW 1.1 –1.5.
Combinations of red, brown and black; anterior pronotum margined;
erect setae about EL sparsely scattered on scapes, indistinct flat-lying
on tibiae. Major worker. Head sides convex strongly tapering
forward; vertex straight; pronotum and mesonotum slightly convex;
propodeal dorsum straight; angle rounded; anterior clypeal margin
median section feebly projecting, narrow, bounded by 2 teeth with a
deep concavity between. Minor worker. Head sides convex; vertex
mostly straight; propodeal dorsum feebly concave.
Camponotus innexus Forel

Worker. HW 2 - 2.4; HL 2 - 2.3; PW 1.8 - 1.9. Very similar to
hartogi except in innexus, where vertex of minor worker is convex, in
hartogi it is mostly flat.
10, sponsorum group
Camponotus froggatti Forel

Worker. HW 0.9 - 1.8; HL 1.1 - 1.8; PW 0.8 - 1.2. Head,
mesosoma, node and gaster black, legs lighter; glossy; propodeal
dorsum mostly concave, angle 90°; long erect setae sparse
everywhere, none on scapes, tibiae; anterior clypeal margin
projecting; node summit convex. Major worker. Head sides mostly
straight, tapering to front, widest behind eyes; anterior clypeal margin
narrow, feebly concave, bounded by two small teeth; central clypeus
widest posteriorly, raised above lateral margins. Minor worker. Head
sides long, straight, tapering to front; anterior clypeal margin wide, convex; most of ant with
white non-overlapping flat-lying setae; a few erect setae under head.
Camponotus sponsorum Forel

Worker. HW 0.85 - 2.0; HL 1.05 –2.05; PW 0.8 - 1.4. Yellow
brown, sometimes mottled, gaster darker; sparse long erect setae
everywhere except scapes and tibiae; anterior clypeal margin
projecting; node summit sharp. Major worker. Head sides straight,
parallel, forehead swollen, anterior clypeal margin concave bounded
by two teeth; propodeal dorsum straight or feebly concave. Minor
worker. Head sides, straight, tapering to front; anterior clypeal
margin wide, strongly convex; most of ant with white non-
overlapping flat-lying setae.
11, aeneopilosus group
Camponotus aeneopilosus Mayr

Worker. HW 1.3 - 2.8; HL 1.5 - 2.5; PW 1.2 - 1.75. Black,
legs brown; pronotum feebly margined in front, more distinct in
minors; sparse yellow, erect setae scattered on mesosoma, none
under head; sparse short, white, flat-lying setae on mesosoma
scapes and tibiae; frontal carinae wide; node summit blunt; finely
punctate. Major worker. Anterior clypeal margin median section
concave bounded by two blunt teeth; head sides convex, tapering
to front; propodeal angle rounded; node summit blunt. Minor
worker. In smallest minors anterior clypeal margin median
section convex, in larger minors concave; head sides tapering forward; vertex slightly convex;
PD/D about 3.
Camponotus nigroaeneus Smith

legs brown; pronotum feebly margined in front, more distinct in
minors; mesosoma with sparse scattered erect setae and short
sparse flat-lying setae; under head with a few erect setae, scape
and tibia short setae raised to about 15°; plentiful short erect setae
on sides of head; finely punctate. Major worker. Anterior clypeal
margin median section concave, bounded by two blunt teeth;
head sides mostly straight; propodeal angle rounded; node
summit blunt. Minor worker. Anterior clypeal margin median
section concave; head sides parallel, straight; vertex convex; PD/D about 3.
12, subnitidus group

Camponotus dorycus confusus Emery

Worker. HW 1.7 - 4.4; HL 2.5 - 4.9; PW 1.5 - 2.4. Brown; with
dense erect setae; node summit sharp, front and back convex; 6 teeth.
Major worker. Head sides mostly straight, tapering to front; vertex
concave; anterior clypeal margin projecting, wide, straight, bounded
by two blunt angles; propodeal dorsum convex; PD/D about 1.5;
angle wide; eyes flattish. Minor worker. Head sides mostly straight,
tapering into a collar at rear; eyes bulbous; head finely reticulate,
darker than mesosoma; anterior clypeus margin wide, convex,
projecting; propodeal dorsum straight, PD/D about 2.5, angle wide; frontal carinae narrow; node
summit sharp, sides convex.
Camponotus subnitidus Mayr

Worker. HW 2.0 - 4.8; HL 3.2 - 5.5; PW 1.5 - 2.7. Brown; erect
setae sparse, more plentiful under head, tibiae and scapes lacking
erect setae. Major worker. Head sides straight, nearly parallel; vertex
straight; anterior head feebly truncate; eyes flattish; anterior clypeal
margin straight, narrow, bounded by 135° angles, projecting, in
lateral view concave near margin; propodeum uniformly convex, an
angle. Minor worker. Head sides mostly straight, tapering into a
collar at rear; head finely reticulate; anterior clypeal margin wide,
convex, projecting, with a strong carina; propodeal dorsum straight, PD/D about 4, angle
indistinct; eyes bulbous; frontal carinae narrow; node long, anterior face short, summit rising
upward to a transverse ridge, posterior face concave above, straight below.
Camponotus tricoloratus Clark

Worker. HW 1.9 - 5.1; HL 3.2 - 5.0; PW 1.5 - 2.8. Yellow to
brown; glossy; plentiful erect setae under head, none on tibia and
scape, sparse otherwise. Major worker. Head sides mostly straight;
vertex straight; in profile feebly truncate; clypeus in profile concave
near anterior margin, in front view straight, crenulate, projecting with
135° corners; propodeal dorsum and declivity straight, angle well
rounded; node summit sharp, front and back convex. Minor worker.
Head sides straight tapering to a collar behind; frontal carinae narrow
in front, slightly wider behind; clypeus wide, anterior margin convex,
projecting; node long, summit blunt, front and back convex.
Camponotus rufus Crawley

Worker. HW 2.0 - 5.6; HL 3.1 - 5.1; PW 1.7 - 2.9. Red, gaster
black; glossy; erect setae sparse, none on tibia, on scape, on
propodeum nor under head, flat-lying on tibia. Major worker. Head
sides convex; vertex straight; in profile feebly truncate; clypeus
finely punctate, in profile concave, in front view anterior margin
nearly straight, crenulate, projecting with 135° corners; propodeal
dorsum straight, angle well rounded; node summit sharp, front and
back convex. Minor worker. Head sides straight tapering to a collar
behind; frontal carinae nearly parallel and narrow; anterior clypeal margin convex, crenulate,
projecting; propodeal dorsum evenly convex; node summit blunt, front and back convex.
13, maculatus group
Camponotus oxleyi Forel

Worker. HW 1.93 - 2.97; HL 1.97 - 2.74; PW 1.4 - 2.0.
Brown; finely punctate when viewed from front; scapes and tibiae
with sparse, flat-lying, short setae, a few raised to about 30°;
pronotum, mesonotum and propodeum each with 2 or 3 erect
setae about EL; shorter and more plentiful under head. Major
worker. Head is longer than wide; anterior clypeal margin median
section bounded by square corners with a wide concavity
between. Minor worker. Head sides convex, tapering forward,
much wider behind than in front; vertex convex; anterior clypeal
margin median section widely convex; node summit convex; propodeal dorsum feebly convex,
angle distinct, well rounded; PD/D about 2.5.
Camponotus extensus Mayr

Worker. HW 1.1 - 2.2; HL 1.6 - 2.7; PW 0.95 - 1.25. Brown;
finely punctate, more so in front view, glossy; scapes and tibiae
with plentiful short setae raised to about 45°; mesosoma long
(particularly propodeum) with about 20 erect setae varying in
length from half EL to EL, plentiful short erect setae under head;
legs slightly lighter. Major worker. Head is much longer than
wide, sides convex and tapering strongly to front; vertex straight
in front view; anterior clypeal margin median section bounded by
square corners widely concave between; node summit blunt;
PD/D about 3.5. Minor worker. Head sides convex, parallel, tapering feebly to rear (more so in
smallest workers); vertex convex; anterior clypeal margin median section bounded by rounded
corners, straight between; node summit convex; PD/D about 4.
Camponotus villosus Crawley

Worker. HW 1.4 - 3.0; HL 2.1 - 3.2; PW 1.2 - 1.70. Yellow;
glossy; scapes and tibiae with plentiful short setae raised to about
45°; mesosoma with about 20 erect setae about EL mostly on
pronotum and near angle, plentiful under head; propodeal dorsum
slightly convex, angle well rounded. Major worker. Head much
wider behind than in front; node summit blunt; anterior clypeal
margin median section projecting, bounded by square corners
with a wide concavity between. Minor worker. Head sides
convex, parallel, tapering feebly to rear (more so in smallest
workers); vertex convex; anterior clypeal margin median section bounded by rounded corners,
straight between; node summit convex; PD/D about 4.
Camponotus maculatus humilior Forel

Worker. HW 1.0 - 2.1; HL 1.45 - 2.3; PW .85 - 1.2.
Yellowish, head slightly darker than mesosoma; tibiae and scapes
with distinct setae raised up to 30°, a few scattered long setae
overall including under head. Major worker. Head sides posterior
half parallel, anterior tapering to front; anterior margin of clypeus,
strongly projecting with 90° corners, feebly concave between;
node summit blunt; eye centers at or just anterior to line joining
posterior tips of frontal carinae. Minor worker. Head sides mostly
parallel, posterior rounded; anterior margin of clypeus, strongly
projecting with square corners, wide, straight; PD/D about 3.
Camponotus novaehollandiae Mayr

Color, a combination of browns and yellows, otherwise similar to
villosus.
Camponotus spenseri Clark

Worker. HW 1.4 - 2.5; HL 1.8 - 2.7; PW 1.1 - 1.50. Brown;
finely punctate; scapes and tibiae with plentiful short setae raised to
about 45°; mesosoma with about 20 erect setae longer than EL,
plentiful short erect setae under head. Major worker. Head sides
convex, tapering strongly to front, anterior clypeal margin median
section bounded by square corners with a wide shallow concavity
between; node summit blunt; PD/D about 2.5. Minor worker. Head
sides evenly convex; vertex convex; anterior clypeal margin median
section bounded by rounded corners, straight between; node summit blunt; propodeum dorsum
straight, angle distinct, about 150°; PD/D about 3.
Camponotus fieldeae Forel

Worker. HW 1.0 - 2.25; HL 1.4 - 2.4; PW 0.8 - 1.30. Propodeum
dorsum is mostly feebly convex (in spenseri straight), angle well
rounded; otherwise similar to spenseri.
14, darlingtoni group
Camponotus claripes elegans Forel

legs yellow; metanotum distinct, propodeum dorsum straight, higher
than mesonotum (more so in minors); scattered setae on all surfaces,
raised to 10° on tibiae and scapes otherwise erect; frontal carinae
wide. Major worker. Anterior clypeal margin median section narrow,
deeply concave; node summit sharp. Minor worker. Anterior clypeal
margin median section widely concave; head sides straight, tapering
rearward in smallest minors, otherwise parallel; head thin in lateral view.
Camponotus inflatus Lubbock

Worker. HW 1.3 - 2.4; HL 1.5 - 2.3 ; PW 1.25 - 1.6. Black;
matte; plentiful erect setae on all surfaces; anterior clypeal margin
median section mostly straight. Major worker. Head sides convex;
vertex concave. Minor worker. Head sides straight tapering forward;
vertex mostly straight; propodeum dorsum slightly convex.
Camponotus cinereus amperei Forel

Worker. HW 1.3 - 2.2; HL 1.4 - 2.25; PW 1.3 - 1.6. Black, legs slightly lighter, mandibles
partly red; pronotum margined in front; a few erect setae on
mesosoma, none under head, flat-lying on scape and tibia. Major
worker. Head sides convex; anterior clypeal margin median section
feebly concave. Minor worker. Head sides straight, nearly parallel,
slightly tapering to front; vertex nearly flat; anterior clypeal margin
median section feebly convex crenulated; node summit convex.
Camponotus pitjantjatarae McArthur

Worker. HW 1.9 - 2.6; HL 1.8 - 2.3; PW 1.6 - 2.0. Black; similar to
inflatus but with less pilosity, erect setae scattered on front of head,
mesosoma, node, gaster, none under head, on scapes nor tibiae. Major
worker. Head sides mostly convex, anterior clypeal margin projecting;
node summit blunt. Minor worker. Head triangular, sides straight.
Camponotus darlingtoni Wheeler

Worker. HW 0.95 - 2.65; HL 1.3 - 2.6; PW 0.9 - 1.65. Black to dark
brown, legs lighter; a few erect setae scattered over mesosoma, none
under head, on scapes nor tibiae. Major worker. Anterior clypeal margin
projecting, median section straight, bounded by rounded corners. Minor
worker. Anterior clypeal margin evenly convex with rounded corners.
15, scotti group

Camponotus claripes marcens Forel

Worker. HW 1.1 - 2.85; HL 1.6 - 3.0; PW 1.0 - 1.9. Brown to
yellow; plentiful erect setae on mesosoma, under head, mostly flat-lying
on scapes with a few raised to 30°, tibiae to 5°. Major worker. Anterior
clypeal margin shallowly concave. Minor worker. Head sides tapering to
rear; anterior clypeal margin feebly concave, bounded by angles.
Camponotus scotti McArthur

Worker. HW 1.2 - 2.4; HL 1.4 - 2.3; PW 1.0 - 1.5. Red to black
combinations; short white curved setae distinct on mesosoma in lateral
view; erect setae, sparse on head, mesosoma, node and gaster, none
under head on scapes nor tibiae; front of head finely and feebly punctate
otherwise finely reticulate, frontal carinae wide. Major worker. Head
wider at back; vertex straight, anterior clypeal margin feebly projecting
with two teeth on either side of a shallow concavity; clypeus coarsely
punctate. Minor worker. Head sides nearly parallel; vertex convex, anterior clypeus margin
projecting, evenly convex.
Camponotus longifacies McArthur

Worker. HW 1.2 - 1.7; HL 1.0 - 1.6; PW 0.7 - 1.4. Combination of
yellow and brown; small. Major worker. Square head; mesosoma evenly
convex; node summit pointed; clypeus mid section raised into a flattish
plane above cheeks, anterior margin concave, bounded by two small
teeth. Minor worker. Head sides straight, tapering forward; node summit
elongate.
Camponotus cinereus Mayr

Worker. HW 1.25 - 2.1; HL 1.65 - 2.15; PW 0.7 - 1.55. Mostly red
to red brown, otherwise similar to cinereus amperei.
Camponotus scratius Forel

Worker. HW 0.80 - 1.5; HL 1.0 - 1.6; PW 0.75 - 1.1. Yellowish
brown, legs lighter; glossy; long erect setae sparse, scattered, flat-lying
on scapes, raised to 10° on tibiae; node summit sharp. Major worker.
Head sides convex, widest at eye level; anterior clypeal margin median
section projecting, slightly convex, bounded by angles. Minor worker.
Eyes large; head sides feebly tapering forward; vertex convex; anterior
clypeal margin median section projecting, straight, bounded by angles.
16, esau group
Camponotus arcuatus Mayr

Worker. HW 0.9 - 1.8; HL 1.1 - 1.7; PW 0.9 - 1.3. Black to very
dark brown; glossy; finely reticulate; long erect setae on all surfaces
sparse, < 10 on mesosoma, white flat-lying short overall, distinct on
scapes and on tibiae; frontal carinae wide. Major worker. Head sides
widest behind eyes, nearly straight, tapering to front; vertex straight;
anterior clypeal margin concave between 2 teeth. Minor worker.
Head sides nearly straight, tapering to front; vertex convex; HW near
PW; anterior clypeal margin broadly convex with a weak central
concavity; pronotum margined in front; node columnar, summit convex.
Camponotus esau Forel

Worker. HW 1.05; HL 1.2; PW 1.04. Major worker not yet
described. Black. Minor worker. Matte, finely reticulate; widespread
white flat-lying short setae, overlapping on gaster, anterior clypeal
margin broadly convex, nearly straight; otherwise similar to
arcuatus.
17, gibbinotus group
Camponotus cowlei Froggatt

Worker. HW 1.30 - 3.5; HL 1.77 - 3.4; PW 1.19 - 2.1; CW 0.38 -
1.0. Yellowish brown; glossy; erect setae sparse on mesosoma,
plentiful on head, slightly raised on scapes and tibiae; node sharp;
frontal carinae narrow. Major worker. Head sides convex, widest just
behind eye level; vertex concave; anterior clypeal margin median
section projecting, nearly straight, bounded by angles. Minor worker.
Head sides straight tapering feebly to front; vertex convex; anterior
clypeal margin median section projecting, convex.
Camponotus discors Forel

Worker. HW 1.34 - 3.02; HL 1.74 - 2.97; PW 1.19 - 1.83; CW
0.42 - 0.80. Yellow to brownish-yellow; glossy; sparse pilosity; erect
setae sparse, a few scattered under head (more so with major
workers), flat-lying, sparse but distinct; node summit sharp, thicker in
minor worker; frontal carinae narrow. Major worker. Head sides
mostly straight; vertex straight; anterior clypeal margin median
section projecting, nearly straight, crenulate, bounded by angles; eyes
small. Minor worker. Head sides nearly straight, feebly tapering
forward; eyes situated < EL/2 from corners; anterior clypeal margin median section projecting,
convex.
Camponotus gibbinotus Forel

Worker. HW 1.1 - 3.1; HL 1.5 - 3.05; PW 1.0 - 1.85. Yellow,
legs lighter in color; glossy; erect setae sparse, a few inconspicuous
short in depression of head where pronotum attaches, flat-lying
sparse and indistinct, none raised on scapes nor tibiae; node summit
sharp; frontal carinae narrow. Major worker. Head sides convex,
tapering forward, widest at eye level; vertex straight to concave;
anterior clypeal margin median section projecting nearly straight,
bounded by angles; eyes small. Minor worker. Head sides slightly
convex; eyes large (EL/HW about 0.25) situated < EL/2 from corner; anterior clypeal margin
median section projecting, convex; propodeum narrow (rear view).
18, walkeri group
Camponotus adami Forel

Worker. HW 1.00 - 2.10; HL 1.30 - 2.20; PW 0.80 - 1.30.
Mostly black, legs, antennae yellowish; glossy; sparse long erect
setae scattered on mesosoma, two or three under head, flat-lying
on scapes and tibiae; anterior clypeal margin median section
projecting, straight, bounded by angles. Major worker. Propodeal
angle about 150°; node summit sharp; head sides convex, widest
at eye level; vertex mostly straight. Minor worker. Propodeal
dorsum and declivity uniformly convex lacking a distinct angle;
head sides straight, nearly parallel, feebly tapering forward;
vertex, median third nearly flat, corners widely rounded; eyes large and bulbous; node summit
blunt; finely punctate, each punctation reflecting light.
Camponotus walkeri Forel

Worker. HW 1.45 - 2.9; HL 1.84 - 2.7; PW 1.2 - 1.75.
Mesosoma red-brown, head, gaster darker, legs lighter; glossy;
propodeal dorsum and declivity uniformly convex lacking a
distinct angle; a few long erect setae scattered on mesosoma, four
or five confined to angle, four or five under head, raised to 5° on
scapes and tibiae; vertex straight; node summit sharp. Major
worker. Head sides mostly straight; anterior clypeal margin
median section projecting, concave, bounded by teeth. Minor
worker. Head sides slightly convex; anterior clypeal margin
median section projecting, straight, bounded by angles; propodeal dorsum mostly straight.
19, armstrongi group
Camponotus armstrongi McAreavey

Worker. HW 1.2 - 2.0; HL 1.30 - 2.0; PW 1.05 - 1.40. Mesosoma bright red, legs lighter,
head and gaster black or dark brown; glossy; dorsum of
mesosoma evenly convex; declivity mostly concave; plentiful
erect long setae, > 20 on mesosoma (more for minor workers),
flat-lying on scapes and tibiae; pronotum margined in front; node
summit sharp. Major worker. Head sides mostly straight, parallel;
vertex straight; forehead swollen; anterior clypeal margin
projecting, median section concave, bounded by two teeth. Minor
worker. Head sides feebly convex, tapering forward, widest at eye
level; vertex, straight to concave; anterior clypeal margin wide,
projecting, convex.
Camponotus macareaveyi Taylor

Worker. HW 1.5 - 2.5; HL 1.4 - 2.4; PW 1.10 - 1.75. Dull
red, sometimes in part brown; pronotum and mesonotum slightly
convex; propodeal dorsum mostly straight; plentiful erect long
setae (> 20 on mesosoma), raised to 5° on scapes and tibiae;
pronotum margined in front; node summit blunt; declivity mostly
straight. Major worker. Head sides straight, parallel; vertex
straight; forehead swollen; anterior clypeal margin projecting,
median section concave, bounded by two teeth; darker than minor
worker. Minor worker. Head sides feebly convex, tapering forward, widest at eye level; vertex,
straight; anterior clypeal margin wide, projecting, convex.
20, evae group
Camponotus tristis Clark

and antennae dark brown; finely punctate; matte; < 6 erect setae on
mesosoma, none under head, on scapes nor tibiae; pronotum
margined front and sides; mesosoma dorsum evenly flatly convex.
Major worker. Head sides convex tapering forward, widest behind
eyes, feebly truncate in lateral view; vertex feebly concave; anterior
clypeal margin projecting, deeply concave in middle, bounded by
teeth; node summit sharp. Minor worker. Head sides convex, widest
at eye level; vertex flat; eyes small; anterior clypeal margin projecting, crenulate in middle;
frontal carinae narrow.
Camponotus rudis McArthur

legs, antennae and mandibles lighter; matte, finely punctate; < 10
erect setae on mesosoma, a few under head, flat-lying on scapes and
tibiae; pronotum margined front and sides; mesosoma dorsum evenly
slightly convex; declivity feebly concave; node summit sharp. Major
worker. Head sides convex, tapering forward, widest at eye level;
vertex straight; anterior clypeal margin projecting, convex with a
shallow concavity in middle. Minor worker. Head sides straight, tapering forward; vertex
convex; eyes small; anterior clypeal margin projecting, convex, crenulate in middle.
Camponotus evae Forel

legs lighter; matte, finely punctate; > 30 erect setae on mesosoma,
few under head, raised to 40° on scapes and tibiae, distinct; pronotum
margined front and sides; node summit sharp. Major worker. Head
sides mostly straight, parallel; vertex straight; anterior clypeal margin
projecting, widely indented in middle. Minor worker. Head sides
convex tapering forward; vertex, slightly convex; anterior clypeal
margin projecting, convex in middle.
21, rubiginosus group
Camponotus simpsoni McArthur

lighter; glossy; erect setae absent under head, mesosoma with < 40;
otherwise similar to evae zeuxis especially in major worker head
sides.
Camponotus rubiginosus Mayr

Worker. HW 1.4 - 2.5; HL 1.35 - 2.7; PW 1.2 - 1.8. Red and
black, bi-coloration more distinct on gaster; long erect setae > 50 on
mesosoma some to 0.4 mm long, short setae raised to 10° on scapes
and to 30° on tibiae, otherwise sparse, flat, fine; pronotum margined
in front and on side. Major worker. Head swollen, sides mostly
straight feebly tapering to front, widest behind eyes; vertex straight;
anterior clypeal margin median third marked by sharp teeth, strongly
concave between; propodeal declivity (rear view) wide, flat, bare,
fringed with long setae. Minor worker. Head sides slightly convex, strongly tapering; vertex
straight; anterior clypeal margin wide, convex.
Camponotus lownei Forel

Worker. HW 1.4 - 1.8; HL 1.2 - 2.1; PW 0.9 - 1.3. Dark brown
to black, legs lighter; glossy; mesosoma with > 30 erect setae,
plentiful under head, scape and tibia with short setae raised to 45º.
Major worker. Head sides straight and parallel for 4/5 of their length,
then slightly tapering forward; vertex tending concave; frontal
carinae wide, anterior clypeal margin median third bounded by two
pointed teeth, with a deep semicircular notch between; node summit
sharp. Minor worker. Head sides evenly slightly convex, tapering
slightly to front; vertex feebly convex, angles rounded; frontal carinae wide, anterior clypeal
margin median third nearly straight, bounded by obtuse angles; node summit blunt.
Camponotus evae zeuxis Forel

Worker. HW 1.2 - 2.5; HL 1.4 - 2.6; PW 1.2 - 1.7. Black to dark
brown; glossy; scape and tibia with plentiful setae up to 0.2 mm long,
raised to 45°, just overlapping; > 50 erect setae up to 0.4 mm long
spread along mesosoma extending down declivity to near pedicel, a
few on front of and under head; pronotum feebly margined; frontal
carinae wide; node summit sharp. Major worker. Head sides convex
tapering forward; anterior clypeal margin median third projecting,
deeply concave in middle, bounded by two teeth; vertex flat. Minor worker. Head sides mostly
uniformly slightly convex tapering to front; vertex slightly convex; anterior clypeal margin
median two thirds projecting, evenly convex.
22, oetkeri group
Camponotus michaelseni Forel

Worker. HW 1.5 - 3.7; HL 1.7 - 3.45; PW 1.3 - 2.1. Brown; reticulate striate; erect setae
sparse, < 6 on mesosoma, a few under head and propodeum,
indistinct flat-lying on scapes and tibiae; propodeal declivity mostly
straight, longer than dorsum; node summit sharp; pronotum margined
at front and side. Major worker. Head sides convex, widest at eye
level; vertex straight; anterior clypeal margin projecting, median third
bounded by teeth on each side with one in middle, shallowly concave
between outer and middle teeth. Minor worker. Propodeal angle 120°
abrupt; head sides straight, tapering forward; vertex slightly convex;
anterior clypeal margin projecting, median half bounded by angles each side, crenulate between.
Camponotus oetkeri Forel

front of head lighter; glossy, finely reticulate; no erect setae on
mesosoma nor under head, indistinct flat-lying on scapes and tibiae;
propodeal angle wide; node summit sharp; pronotum margined at
front and side. Major worker. Head sides convex, tapering forward;
vertex straight with well rounded corners; anterior clypeal margin
projecting, median third shallowly concave. Minor worker. Propodeal
angle 150° widely rounded; head sides convex, tapering forward;
vertex slightly convex; anterior clypeal margin feebly projecting, median two thirds widely
concave.
23, claripes group
Camponotus eremicus Wheeler

Worker. HW 1.95; HL 1.92; PW 1.28; FCW 0.47. Brown,
antennae lighter, legs yellow. Major worker. No erect setae on
mesosoma or under head; head sides convex, tapering forward;
mesosoma dorsum in profile evenly convex, sharp at angle (blade-like
in rear view); pronotum margined at front and sides; anterior clypeal
margin median two thirds feebly projecting, straight, bounded by wide
angles; frontal carinae narrow. Minor worker. Sparse erect setae on
mesosoma and under head, none on scapes and tibiae; head sides straight, feebly tapering
forward; vertex convex; frontal carinae narrow; mesosoma dorsum evenly convex; pronotum not
margined; pronotum convex on top (rear view).
Camponotus claripes Mayr

Worker. HW 0.9 - 2.7; HL 1.5 - 2.85; PW 0.85 - 1.65. Yellow
brown, legs lighter; glossy; dense long erect setae on all surfaces of
head, less on mesosoma, short raised to 20° on scapes and tibiae, flat-
lying, short, widespread not overlapping on mesosoma. Major worker.
Head sides mostly slightly convex; vertex straight; anterior clypeal
margin bounded by angles with a concavity between; mesosoma
dorsum slightly convex; propodeal angle well rounded. Minor worker.
Head sides straight, parallel, much longer than wide; head thin (lateral view); vertex convex;
anterior clypeal margin median third projecting, convex, sometimes with a wide central
concavity; mesosoma dorsum slightly convex; propodeal angle well rounded; large eyes.
Camponotus claripes minimus Crawley

Worker. HW 0.95 - 2.44; HL 1.22 - 2.40; PW 0.80 - 1.45. Brown,
legs lighter; glossy; sparse erect setae on head, < 10 on mesosoma,
indistinct flat-lying on scapes and tibiae; node summit sharp. Major
worker. Head sides posterior half straight, parallel then tapering
forward; vertex straight; anterior clypeal margin median third
projecting, bounded by wide angles, crenulate between; mesosoma
dorsum slightly convex; metanotum distinct, depressed front and back,
elevated between; propodeal angle well rounded; frontal carinae wide. Minor worker. No erect
setae under head; head sides straight, parallel; vertex straight; anterior clypeal margin projecting,
wide, convex; mesosoma dorsum slightly convex; propodeal angle 160° well rounded; large
eyes.
ACKNOWLEDGEMENTS
This work has been made possible through the support of the South Australian Museum, CSIRO
Entomology and grants from Australian Biological Resources Study and the Mark Mitchell
Trust. The South Australian Department for Environment and Heritage, Biodiversity and
Monitoring Section provided many of the specimens for study. Special thanks are due to S.
Barker (SAMA), B. Bolton (BMNH), J. A. Forrest OAM (SAMA), R. Hutchinson (SAMA), H.
Landsberg (ZMB), F. Koch (ZMB), R. J Lavigne (SAMA), E. G. Matthews (SAMA), B. Merz
(MHNG), R. Poggi (MCG), S. O. Shattuck (ANIC), S. Schödl (NHMW), R. R. Snelling
(LACM), C. H. Watts (SAMA) and to P. J. Fargher and J. J. Weyland for help with the
manuscript
LITERATURE CITED
Bolton, B. 1994. Identification guide to the ant genera of the world: 222 pp. Harvard University
Bolton, B. 1995. A new general catalogue of the ants of the world: 504 pp. Harvard University
Braby, M.F. 2000. Butterflies of Australia. Their identification, biology and distribution: 976
pp. CSIRO Publishing, Australia.
Clark, J. 1930a. Some new Australian Formicidae. Proceedings of the Royal Society of Victoria
42: 116-128.
Clark, J. 1930b. New Formicidae, with notes on some little known species. Proceedings of the
Royal Society of Victoria 43: 2-25.
Clark, J. 1938. Reports of the McCoy Society for Field Investigation and Research. No. 2. Sir
Joseph Banks Islands.10. Formicidae. Proceedings of the Royal Society of Victoria 50: 356-
382.
Clark, J. 1941. Australian Formicidae. Notes and new species. Memoirs of the National
Museum of Victoria 12: 71-93.
Crawley, W.C. 1915a. Ants from north and central Australia, collected by G. F. Hill. Part 1.
Annals and Magazine of Natural History (8) 15: 130-136.
Crawley, W.C. 1915b. Ants from north and south west Australia (G. F. Hill, Rowland Turner)
and Christmas Island, Straights Settlements. Part 2. Annals and Magazine of Natural
History (8) 15: 232 - 239.
Crawley, W.C. 1922. New ants from Australia. Annals and Magazine of Natural History (9) 10:
16-36.
Crawley, W.C. 1925. New ants from Australia. II Annals and Magazine of Natural History (9)
16: 577-598.
Donisthorpe, H. 1941. Synonymical notes etc., on Formicidae. Entomologist’s Monthly
Magazine 77: 37-240.
Emery, C. 1887. Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova.
Parte terza. Formiche della regione Indo-Malese e dell'Australia. Annali del Museo Civico di
Storia Naturale "Giacomo Doria" (2) 4 [24] (1886): 209-258.
Emery, C. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. 20. Formiche di
Birmania e del Tenasserim raccolte da Leonardo Fea (1885-1887 Annali del Museo Civico
di Storia Naturale "Giacomo Doria" 7: 485-520.
Emery, C. 1914. Les Fourmis de la Nouvelle-Calédonie et des îles Loyalty. In Sarasin, F. &
Roux, J. (eds) Nova Caledonie Zoologie 1: 393-437 Wiesbaden.
Emery, C. 1920. Studi sui Camponotus. Bullettino della Società Entomologica Italiana 52: 3-
49.
Emery, C. 1925. Hymenoptera. Fam. Formicidae. Subfam. Formicinae. In Wytsman, P.(ed.)
Genera Insectorum: 1-302. Louis Desmet-Verteneuil: Bruxelles- Fascicule 183.
Erichson, W.F. 1842. Beitrag zur Insecten-Fauna von Vandiemensland. Arkiv fur
Naturgeschichte 8: 83-287.
Forel, A. 1893. Nouvelles fourmis d'Australie et des Canaries. Annales de la Société
Entomologique de Belgique 37: 454-466.
Forel, A. 1894. Quelques fourmis de Madagascar (recoltees par M. le Dr Völtzkow); de
Nouvelle Zelande (recoltees par M W.W.Smith); de Nouvelle Caledonie (recoltees par
M.Sommer); de Queensland (Australie) (recoltees par M. Wiederkehr); et de Perth
(Australie occidentale) (recoltees par M. Chase). Annales de la Société Entomologique de
Belgique 38: 226-237.
Forel, A. 1895a. Nouvelles fourmis de diverses provenances, surtout d’Australie. Annales de la
Société Entomologique de Belgique 39: 41-49.
Forel, A. 1895b. Nouvelles fourmis d'Australie, récoltées à The Ridge, Mackay, Queensland par
M. Gilbert Turner. Annales de la Société Entomologique de Belgique 39: 417-428.
Forel, A. 1902. Fourmis nouvelles d'Australie. Revue Suisse de Zoologie 10: 405-548.
Forel, A. 1907a. Formicides du Musée National Hongrois. Annales Historico-Naturales Musei
Nationalis Hungarici 5: 1-42.
Forel, A. 1907b. Formicidae. In Michaelsen, W.& Hartmeyer, R. (eds). Die Fauna Südwest
Australiens 1: 263-310 Jena.
Forel, A. 1910. Formicides australiens recus de MM.Froggatt et Rowland Turner. Revue Suisse
de Zoologie 18: 1-94.
Forel, A. 1913. Fourmis de Tasmanie et d'Australie récoltées par MM.Lea, Froggatt etc. Bulletin
de la Société Vaudoise des Sciences Naturelles 9: 173-196.
Forel, A. 1915. Results of Dr. E. Mjöberg’s Swedish Scientific Expeditions to Australia 1910-
1913. 2 Ameisen. Arkiv fur Zoologi 16: 1-119.
Forel, A. 1922. Glanures myrmécologiques en 1922. Revue Suisse de Zoologie 30: 87-102.
Froggatt, W.W. 1896. Honey ants. In Spencer, B. (ed.). Report on the work of the Horn
Scientific Expedition to Central Australia Pt. 2 Zoology: 385-392. Melville, Mullen &
Slade, Melbourne.
Froggatt, W.W. 1905. Domestic insects: ants. Agricultural Gazette of New South Wales 1905:
861-6.
Greenslade, P.J.M. 1979. A guide to the ants of South Australia. 44 pp. South Australian
Museum.
Haskins, C.P. & Haskins, E. 1992. Note on extraordinary longevity in a queen of the formicine
ant genus Camponotus. Psyche 99: 31-33.
Kirby, W. 1819. A description of several new species of insects collected in New Holland by
Robert Brown, Esq., F. R. S. Transactions of the Linnean Society of London 12 1818: 454-
478.
Kirby, W.F. 1896. Hymenoptera. In Spencer, B. (ed.). Report on the work of the Horn Scientific
Expedition to Central Australia Pt. 1 Supplement: 203-209. Melville, Mullen & Slade,
Melbourne.
Lowne, B.T. 1865. Contributions to the natural history of Australian ants. Entomologist 2: 273-
280.
Lubbock, J. 1880. Observations on ants, bees and wasps with a description of a new species of
honey-ant. Part vii. Ants. Journal of the Linnaean Society of London. Zoology 15: 41-52.
McAreavey, J.J. 1949. Australian Formicidae. New genera and species. Proceedings of the
Linnean Society of New South Wales 74: 1-25.
McArthur, A.J. 2003a. New species of Camponotus (Hymenoptera: Formicidae) from

Australia. Transactions of the Royal Society South Australia 127: 5-14.
McArthur, A.J. 2003b. Ants and acacias. The South Australian Naturalist 78 (3/4): 3-7.
McArthur, A.J. & Adams, M. 1996. A morphological and molecular revision of the
Camponotus nigriceps group (Hymenoptera: Formicidae) from Australia. Invertebrate
Taxonomy 10: 1-46.
McArthur, A.J., Adams, M. & Shattuck, S.O. 1997. A morphological and molecular review
of Camponotus terebrans (Lowne) (Hymenoptera: Formicidae). Australian Journal of
Zoology 45: 579-598.
McArthur, A.J. & Shattuck, S.O. 2001. A taxonomic revision of the Camponotus
macrocephalus species group (Hymenoptera: Formicidae) in Australia. Transactions of the
Royal Society South Australia 125: 25-43.
Mayr, G.L. 1861. Die Europäischen Formiciden. (Ameisen.). (Carl Gerolds Sohn, Vienna).
Mayr, G.L. 1862. Myrmecologische Studien. Verhandlungen der Zoologisch-Botanischen
Mayr, G.L. 1870. Neue Formiciden. Verhandlungen der k.k. Zoologisch-Botanischen
Mayr, G.L. 1876. Der Australischen Formiciden. Journal des Museum Goddefroy (4) 12: 56-
115.
Menozzi, C. 1925. Qualche formica nuova od interessante del Deutsch. Entomol. Institut di
Dahlem. Entomologische Mitteilungen 14: 368-371.
Poulton, E.B. & Crawley, W.C. 1922. Notes on some Australian ants. Entomologist’s Monthly
Magazine (3) 8: 118-126.
Roger, J. 1863.Verzeichniss der Formiciden-Gattungen und Arten. Berliner Entomologische
Mitteilungen 7 (Beilage): 1-65.
Santschi, F. 1919. Cinq notes myrmécologiques. Bulletin de la Société Vaudoise des Sciences
Naturelles 52: 325-350.
Santschi, F. 1928. Nouvelles fourmis d’Australie. Bulletin de la Société Vaudoise des Sciences
Naturelles 56: 465-483.
Shattuck, S.O. & McArthur, A.J. 1995. Generic placement of Australian ants described by
W.F. Erichson (Hymenoptera: Formicidae). Journal of the Australian Entomological
Society 34: 121-123.
Shattuck, S.O. 1999. Australian ants: their biology and Identification. Monographs on
Invertebrate Taxonomy 3: 226 pp.
Shattuck, S.O. & McArthur, A.J. (2002). A taxonomic revision of the Camponotus
wiederkehri and perjurus species groups (Hymenoptera: Formicidae). Transactions of the
Royal Society South Australia, 126: 63-90.
Shattuck, S.O. 2004. Review of the Camponotus aureopilus species group (Hymenoptera,
Formicidae) including a second Camponotus with a metapleural gland. Zootaxa 903: 1-20.
Smith, F. 1858. Catalogue of Hymenopterous Insects in the collection of the British Museum.
Part 6. Formicidae: 216 pp. London.
Smith, F. 1859. Catalogue of hymenopterous insects collected by Mr A. R. Wallace in the
islands of Aru and Key. Journal of the Proceedings of the Linnaean Society of London.
Zoology 3: 132-158.
Smith, F. 1860. Catalogue of hymenopterous insects collected by Mr A. R. Wallace in the
islands of Bachian, Kaisaa, Amboyna, Gilolo and at Dory in New Guinea. Journal of the
Proceedings of the Linnaean Society of London. Zoology 5 (supplement to volume 4): 93-
143.
Taylor, R.W. 1992. Nomenclature and distribution of some Australian and New Guinean ants of
the subfamily Formicinae. Journal of the Australian Entomological Society 31: 57-69.
Viehmeyer, H. 1925. Formiciden der australischen Faunenregion.(Schluss.) Entomologische

Mitteilungen 14: 139-149.
Wheeler, W.M. 1908. Honey ants, with a revision of the American Myrmecocysti. Bulletin of
the American Museum of Natural History 24: 345-397.
Wheeler, W.M. 1915. Hymenoptera. Transactions of the Royal Society South Australia 39: 805-
823.
Wheeler, W.M. 1927. The ants of Lord Howe Island and Norfolk Island. Proceedings of the
American Academy of Arts and Sciences 62: 121-153.
Wheeler, W.M. 1933. Mermis parasitism in some Australian and Mexican ants. Psyche 40: 20-
31.
Wheeler, W.M. 1934 Contributions to the fauna of Rottnest Island, Western Australia. No. IX.
The ants. Journals of the Royal Society Western Australia 20: 137-163.
APPENDIX
TABLE 1. Camponotus taxa not included in the key
Species Author Type Location

aeneopilosus flavidopubescens Forel, 1902 MHNG
arcuatus aesopus Forel, 1907b not found
bigenus Santschi, 1919 NHMB
cameratus Viehmeyer, 1925 ZMB
chalceoides Clark, 1938 NMV
cinereus notterae Forel, 1907b not found
claripes inverallensis Forel, 1910 MHNG
claripes nudimalis Forel, 1913 MHNG
claripes orbiculatopunctatus Viehmeyer, 1925 ZMB
claripes piperatus Wheeler, 1933 MCZ
consectator Smith, 1858 BMNH
crenatus Mayr, 1876 NHMW
cruentatus aspera Menozzi, 1925 not found
discors yarrabahensis Forel, 1915 MHNG
erythropus Viehmeyer, 1925 ZMB
horni Kirby, 1896 not found
insipidus Forel, 1893 MHNG
lividicoxis Viehmeyer, 1925 not found
longinodis Forel, 1915 NMV
molossus Forel, 1907b MHNG
nigriceps lividipes Emery, 1887 MCG
nigroaeneus divus Forel, 1907a MHNG
nigroaeneus xuthus Forel, 1915 MHNG
nitidiceps Viehmeyer, 1925 ZMB
oetkeri voltai Forel, 1913 MHNG
pellax Santschi, 1919 NHMB
punctiventris Emery, 1920 MCG
scratius nunctius Forel, 1907b not found
simulator Forel, 1915 MHNG
spinitarsus Emery, 1920 MCG
subnitidus famelicus Emery, 1887 MCG
suffusus bendigensis Forel, 1902 MHNG
tumidus Crawley, 1922 OUM
walkeri bardus Forel, 1910 MHNG
TABLE 2. Character states and legend for species in the key.

TABLE 3. Character states and legend for species in the key
Worker Major Minor

Character A B C D E F GH I J KLMN ABCDE F GH I JKLMN
adami not available 1 2 2 2 3 3.1 4 2 3 1 1 2 1 2
aeneopilosus 2 1 2 1 5 3.3 3 2 1 1 2 2 2 1 2 2 2 1 3 2.9 3 2 1 1 2 2 1 1
afflatus 2 2 2 2 5 2.4 1 2 3 1 1 2 1 2 1 2 2 2 4 3.3 1 2 1 1 1 2 2 2
anderseni 1 1 1 2 4 1.3 3 2 1 1 1 2 1 1 2 2 4 2.0 1 2 1 1 1 1 1 2
annetteae not described 1 2 1 1 3 1.5 1 2 3 1 3 3 3 2
arcuatus 2 2 2 1 5 2.6 4 2 3 1 2 2 2 2 1 2 2 1 4 2.3 4 2 3 1 2 2 2 2
arenatus not described 2 2 1 2 4 3.2 1 2 2 1 3 2 1 2
armstrongi 1 2 1 1 5 2.6 3 1 3 2 3 2 1 2 1 2 2 1 4 2.8 5 1 3 2 3 2 1 2
aurocinctus 1 1 2 1 3 4.0 1 4 2 1 1 2 1 2 2 1 1 2 3 3.5 1 4 3 1 1 2 1 2
capito 1 1 2 1 5 2.9 1 2 1 1 2 2 2 2 1 1 2 2 4 3.0 1 2 1 1 2 2 1 2
c. ebinithorax 1 2 1 1 5 3.3 3 2 3 2 2 2 1 2 1 2 2 1 4 2.8 1 2 3 1 1 2 1 2
ceriseipes 1 1 2 1 3 3.9 1 2 2 1 1 2 1 2 1 1 2 2 4 4.1 1 4 2 1 2 2 1 2
chalceus 2 1 2 1 5 3.1 2 2 3 1 2 2 2 2 2 1 2 2 4 2.6 2 2 3 1 2 2 1 2
cinereus 1 1 2 1 5 3.3 3 2 1 1 1 2 1 2 2 2 2 1 4 2.9 3 2 1 1 2 2 1 2
c. amperi 1 1 2 1 5 3.3 3 2 1 1 1 2 1 2 2 2 2 1 4 2.9 3 2 1 1 2 2 1 2
clarior 1 1 2 1 1 4.0 3 2 3 2 3 2 1 2 1 1 1 1 1 3.7 3 2 3 2 3 2 1 2
claripes 1 2 2 1 5 3.0 4 2 3 2 3 2 1 2 1 2 1 1 4 3.1 3 2 3 2 2 2 1 2
c. elegans 1 1 2 1 5 2.8 3 2 3 2 2 2 1 2 1 2 1 1 4 2.6 3 2 3 2 2 2 1 2
c. -marcens 2 2 1 2 5 2.8 3 1 3 1 3 2 1 2 1 2 3 2 2 2.9 3 2 3 1 2 2 1 2
c. minimus 1 1 2 1 5 3.3 3 2 3 1 2 2 2 2 1 2 2 1 4 2.3 3 2 1 1 1 2 1 2
conithorax 1 2 1 2 3 1.8 1 2 1 1 1 1 1 2 1 2 2 2 4 2.0 2 2 1 1 1 1 1 2
consobrinus 1 1 2 1 1 3.5 3 2 1 2 2 2 1 2 1 2 1 2 2 3.0 3 2 1 2 2 2 1 2
cowlei 1 1 2 1 3 3.5 3 2 3 2 2 2 1 2 1 2 1 1 3 3.3 4 2 3 2 1 2 1 2
darlingtoni 1 1 2 1 3 3.0 3 2 1 1 2 2 1 2 1 2 2 1 4 2.5 3 2 1 1 2 2 1 2
discors 1 1 2 1 3 3.7 3 2 3 1 1 2 1 2 1 2 1 1 4 3.3 3 2 1 1 1 2 1 2
donnellani not described 1 2 2 2 4 3.7 1 2 2 1 2 2 1 2
d. confusus 2 2 1 1 3 3.8 3 2 3 3 3 3 1 2 1 2 3 3 4 2.9 3 2 3 3 3 3 1 2
dromas 1 2 1 1 5 2.4 1 2 3 1 3 2 1 2 1 2 2 1 4 2.8 1 2 3 1 3 2 1 2
dryandrae 1 1 2 1 1 3.3 3 2 3 2 3 3 1 2 1 2 1 1 2 3.3 3 2 3 2 3 3 1 2
eastwoodi 1 1 2 1 1 3.9 3 2 3 1 3 3 1 2 1 2 3 1 2 3.1 3 2 3 1 3 3 1 2
ephippium 1 2 1 1 5 3.0 1 2 3 2 3 2 1 2 1 2 1 1 4 2.8 1 2 3 2 3 2 1 1
e. narses 1 2 2 2 5 2.7 3 2 1 1 3 2 1 2 4 2 1 1 3 2.9 3 2 1 1 3 2 1 1
eremicus not available 1 1 2 1 4 3.9 4 1 3 1 1 2 1 2
esau 2 1 2 1 5 2.9 3 2 3 2 2 2 1 1 2 2 2 1 4 2.6 3 2 3 2 2 2 1 1
evae not available 2 1 2 1 4 2.9 3 2 3 2 3 2 1 2
e. zeuxis 1 1 2 1 5 3.0 3 1 3 3 3 2 1 2 1 2 2 1 4 2.8 3 2 2 3 3 2 1 2
extensus 2 1 2 1 3 3.3 3 2 3 2 2 3 3 2 2 2 3 3 3 3.1 3 2 3 2 2 3 3 2
TABLE 3. continued
Worker Major Minor

Character A B C D E F GH I JKLMN ABCDE F GH I J KLMN
fergusoni 1 1 2 1 5 3.2 3 2 3 1 2 2 3 2 4 2 1 1 4 3.4 1 2 3 1 2 2 1 1
fieldeae 2 1 2 1 3 3.4 3 2 3 3 2 2 1 2 2 2 1 1 3 3.1 3 2 3 3 2 2 1 2
fieldellus 1 2 1 1 5 2.6 3 2 1 1 2 2 1 2 4 2 2 2 4 3.0 3 2 1 1 2 2 1 1
froggatti 1 2 2 1 5 2.7 2 2 1 1 2 2 1 2 1 2 2 1 4 2.5 2 2 3 1 1 2 1 2
gasseri 1 2 1 1 4 1.5 1 2 1 1 1 2 1 2 2 2 4 1.8 1 2 1 1 1 2
gibbinotus 1 1 2 1 3 3.7 3 2 1 1 1 2 1 2 1 2 1 1 4 3.3 3 2 1 1 1 2 1 2
gouldianus 1 1 2 1 3 4.0 3 2 2 3 3 3 1 2 4 1 1 2 4 3.2 3 2 2 3 3 3 1 1
hartogi 2 1 2 1 5 2.8 3 2 3 1 3 2 2 2 2 1 2 1 5 2.6 3 2 3 1 3 2 1 2
howensis 1 2 1 1 3 1.8 3 2 1 1 1 2 1 1 1 2 4 2.2 3 2 1 1 1 2
inflatus 2 2 2 2 3.0 3 2 3 2 3 2 1 2 2 1 2 1 4 2.9 3 2 3 2 3 2 1 2
innexus 2 1 2 1 5 2.9 3 2 3 1 2 2 1 2 2 2 2 2 5 2.6 3 2 3 1 3 2 1 2
intrepidus 2 1 2 1 6 3.9 1 2 3 3 3 1 1 2 2 2 1 2 4 3.4 1 2 3 3 3 1 1 2
i. bellicosus 2 1 2 1 6 4.1 1 2 3 3 3 3 3 2 2 2 1 2 4 3.7 1 2 3 3 3 1 1 2
janeti 1 2 1 1 4 1.8 3 2 3 3 3 2 1 2 1 2 1 1 4 1.6 3 2 3 3 3 2 1 2
janforrestae not described 1 2 2 2 5 2.1 1 2 3 3 3 2 1 2
johnclarki not available 1 2 1 3 4 4.2 1 5 3 3 3 2 1 2
leae 3 2 2 1 5 2.8 1 1 3 3 3 2 1 2 3 1 1 2 6 2.9 1 4 1 3 2 2 1 1
longideclevis 1 1 2 1 1 3.5 3 2 1 2 2 3 3 2 1 2 1 1 2 3.3 3 2 1 2 2 2 3 2
longifacies 1 1 2 1 3 3.3 3 2 3 1 2 3 1 2 1 2 2 2 4 3.4 1 2 3 1 2 2 1 1
loweryi 1 1 2 1 1 3.7 3 2 1 1 3 2 3 2 1 2 1 1 2 3.1 3 2 1 1 3 2 3 2
lownei 1 2 1 1 5 2.8 3 2 3 3 3 2 1 2 1 1 2 1 4 2.9 3 2 3 3 3 2 1 2
macareaveyi 1 2 1 1 5 3.0 3 2 3 3 3 2 1 2 1 2 2 1 4 2.9 3 2 3 3 3 2 1 2
mackayensis 2 2 2 1 4 1.9 3 2 3 1 2 3 1 2 1 1 2 2 4 2.4 2 2 1 1 2 2 1 2
macroceph. 1 2 1 1 4 1.7 3 2 1 1 1 2 1 1 1 2 4 2.0 3 2 1 1 1 2
m. humilor 1 1 2 1 3 2.9 3 2 3 2 2 2 3 2 1 2 1 2 3 2.5 3 2 3 2 2 2 3 2
michaelseni 1 1 2 1 6 3.5 3 1 1 1 1 2 1 2 1 1 2 1 4 3.0 3 2 3 1 1 2 1 2
nigriceps 1 1 2 1 1 3.6 3 2 3 2 3 2 1 2 1 2 1 2 2 2.9 3 2 3 2 3 2 1 2
nigroaeneus 2 2 1 1 5 3.0 3 2 3 2 2 2 1 2 1 2 1 2 5 2.6 3 2 3 2 2 2 1
novaeholland. 2 2 2 1 4 3.3 3 2 3 2 3 2 3 2 1 2 3 3 3 2.7 3 2 3 2 3 2 3 2
oetkeri 1 1 2 1 5 4.1 3 1 1 1 1 2 1 1 2 1 4 4.4 3 1 1 1 1 2 1 2
owensae not described 1 2 1 1 4 3.1 1 4 2 2 2 3 1 2
oxleyi 2 1 2 1 5 2.9 3 2 3 1 3 2 1 2 2 2 1 2 5 2.8 3 2 3 1 3 2 1 1
pallidiceps 1 1 2 1 1 3.7 3 2 3 2 2 3 1 2 1 2 1 2 2 3.0 3 2 3 2 2 2 1 1
pawseyi 1 2 1 1 5 2.9 1 2 3 3 3 3 1 2 4 2 2 2 4 2.9 1 2 3 2 3 2 1 1
perjurus not described 1 2 2 2 4 3.1 1 2 1 1 2 2 1 2
piliventris not available 2 1 2 2 4 3.7 1 2 3 3 3 1 1 1
pitjantjat. 1 1 2 1 5 3.4 3 2 1 1 2 2 1 2 2 1 2 1 5 3.2 3 2 1 1 2 2 1 2
postcornutus 1 1 2 1 3 4.1 3 2 2 2 2 3 1 2 1 1 2 2 4 2.9 3 2 2 1 2 2 1 2
TABLE 3. continued
Worker Major Minor

Character A B C D E F GH I JKLMN ABCDE F GH I JKLMN
prosseri 1 1 2 1 3 3.8 3 2 2 1 2 2 1 2 1 1 1 2 4 3.3 1 2 2 1 2 2 1 2
prostans 1 1 2 2 1 3.6 3 2 3 2 2 2 3 2 1 2 1 1 2 3.1 3 2 3 2 2 2 3 2
rubiginosus 1 2 2 1 5 3.2 3 2 3 3 3 2 1 2 1 2 2 1 4 3.0 3 2 3 3 3 2 1 2
rudis 2 1 2 1 5 3.6 3 2 3 2 2 2 1 2 2 2 2 2 4 3.3 3 2 3 1 2 2 1 2
rufonigris 1 1 2 1 4 3.8 1 2 2 1 2 2 1 2 1 1 2 2 4 3.3 1 2 2 1 2 2 3 1
rufus 1 1 2 1 4 3.6 3 2 1 1 1 2 1 2 1 2 3 3 3 3.5 3 3 1 1 1 2
sanguinifrons 1 2 1 1 3 1.8 1 2 3 1 2 3 1 2 1 2 2 2 4 2.4 1 2 3 1 1 2 1 2
scotti 1 1 2 1 5 3.1 3 2 1 1 2 2 1 2 1 1 2 1 4 2.7 3 2 1 1 2 2 1 2
scratius 1 1 2 1 5 3.0 3 1 3 2 2 2 1 2 1 2 2 1 4 2.7 3 2 3 2 1 2 1 2
setosus not described 4 1 1 2 4 3.4 1 2 2 3 3 2 1 1
simpsoni 1 2 1 1 5 3.0 3 1 1 1 2 2 1 2 1 1 2 1 5 2.7 3 1 1 1 3 2 1 2
spenseri 2 1 2 1 3 3.3 3 2 3 3 2 2 3 2 2 1 1 1 3 3.0 3 2 3 3 2 2 3 2
sponsorum 1 2 1 2 5 2.8 1 2 3 1 2 2 1 2 1 2 2 2 4 2.9 1 2 3 1 2 2 1 2
subnitidus 2 2 2 1 3 3.8 3 2 3 1 2 3 2 2 2 3 3 4 3.8 3 3 3 1 1 3 1 2
suffusus 2 1 2 1 3 3.7 1 2 3 3 3 1 1 1 2 1 1 2 4 3.5 1 2 3 3 3 1 1 1
tasmani 1 2 1 1 5 2.6 1 2 3 3 3 2 1 2 1 2 2 1 4 3.4 1 2 3 3 3 2 1 2
terebrans 1 1 2 1 3 4.2 3 2 2 3 2 2 1 2 1 1 1 2 4 3.2 3 2 2 3 2 2 1 2
thadeus 2 2 2 1 5 3.4 3 2 1 1 3 2 1 1 2 2 2 1 4 2.8 3 2 1 1 3 2 1 1
tricoloratus 1 2 2 1 3 3.6 3 2 3 1 2 3 1 2 1 2 3 3 4 3.5 3 3 3 1 2 3 1 2
tristis 2 1 2 1 5 3.7 3 1 1 1 1 2 1 2 2 1 2 3 5 3.6 3 1 1 1 1 2
versicolor 1 1 2 1 3 4.0 1 4 2 1 1 2 1 2 2 1 1 2 3 4.0 1 4 3 1 1 2 1 2
villosus 2 2 2 1 4 3.3 3 2 3 2 3 2 3 2 1 2 3 3 3 2.3 3 2 3 2 3 2 3 2
vitreus 1 2 1 1 4 1.8 1 2 1 3 3 2 1 2 1 1 2 2 4 1.8 1 2 1 3 3 2 1 2
walkeri 1 1 2 1 3 3.4 3 1 3 1 2 2 1 2 1 2 2 1 4 2.9 3 2 3 1 2 2 1 2
whitei 1 1 2 1 5 2.3 1 2 3 3 3 2 1 2 3 1 1 1 4 2.7 1 2 3 3 2 2 1 2
wiederkehri 1 1 2 1 3 3.4 1 1 2 1 2 2 1 2 1 1 1 2 4 2.9 3 2 2 1 2 2 1 1
Ogata, K., Okido, H. 2007. Revision of the ant genus Perissomyrmex with notes on the
phylogeny of the tribe Myrmecinini, pp. 352-369. In Snelling, R. R., B. L. Fisher, and P. S.
Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson
– 50 years of contributions. Memoirs of the American Entomological Institute, 80.
REVISION OF THE ANT GENUS PERISSOMYRMEX,

WITH NOTES ON THE PHYLOGENY OF THE TRIBE MYRMECININI
Kazuo Ogata
and
Hirofumi Okido 1
Institute of Tropical Agriculture, Kyushu University,
Fukuoka 812-8581, Japan
kogata@agr.kyushu-u.ac.jp
ABSTRACT
The ant genus Perissomyrmex is one of the four extant genera of the tribe Myrmecinini. The
present study revises the genus including a definition of the taxon, a description of a new species
from China, discussions on the phylogenetic position in the tribe and biogeography. Our
phylogenetic analysis based on morphological characters shows that (1) the tribe Myrmecinini is
justified as a monophyletic taxon by the structures of the antennal base and the labrum, (2) the
genera Perissomyrmex and Pristomyrmex are sister groups, sharing the short masticatory margin
of the mandible nearly vertical to the long axis of the head and more or less raised structure on
the basal margin of the mandibles, and (3) each of the genera is recognized as a monophyletic
taxon. Our provisional conclusion of the phylogeny is (Myrmecina + (Acanthomyrmex +
(Pristomyrmex + Perissomyrmex))).
A key to the four species of Perissomyrmex is given. P. snyderi from Central America is
distinctive in having a subpetiolar process and three other Asian species are distinguishable by
the dentition on the anterior clypeal margin and the shape of the ventral margin of the petiole.
The disjunct distribution of the genus is also discussed. The discovery of a fourth species from
SW China, Perissomyrmex emarginatus, n. sp., suggests that the genus is an old temperate ant
living in the boundary to northern zoogeographical regions. Judging from the sister group
relation to Pristomyrmex that occurs in the Old World tropics and from higher species diversity
in Asia, Perissomyrmex would have originated in Asia. The fossil record of Ennaemerus in
Baltic Amber suggests that the Proto-Perissomyrmex might have Arcto-Tertiary distribution, and
the present distribution would be a relic of the old temperate range in Paleogene.
Key words: Hymenoptera, Formicidae, Perissomyrmex, new species, phylogeny, biogeography
1
Present address: Kyushu Environmental Evaluation Association, 1-10-1 Matsukadai, Higashi-
ku, Fukuoka 813-0004 Japan
Ogata & Okido: Revision of Perissomyrmex 353
INTRODUCTION
The ant genus Perissomyrmex is a small, rare taxon including only three known species
occurring in the Oriental and Neotropical regions. The ant was first described from a quarantine
specimen in the USA and the original locality of the specimen was supposed to be from
Guatemala (Smith, 1947). This presumed original locality had been questioned because the
genus shows a close affinity to the genus Pristomyrmex, found exclusively in the Old World
(Bolton, 1981). The discovery of the second species from Bhutan (Baroni Urbani & de Andrade,
1993) encouraged speculation that Perissomyrmex might not be native in Central America.
However, the rediscovery of Perissomyrmex in Mexico and Guatemala (Longino & Hartley,
1994) confirmed the disjunct distribution of the genus, although the relationships of the genus
remained unclear.
Perissomyrmex is easily distinguished from the genus Pristomyrmex in having 9-segmented
antennae and a small but distinct tooth on the basal margin of the mandible, all of which are
apomorphic, while for Pristomyrmex there have been no clear synapomorphies although Baroni
Urbani & Andrade (1993) cited the absence of the metanotal groove. Once Perissomyrmex is
recognized as a separate genus, Pristomyrmex would become a paraphyletic taxon. This might
lead to a case of a possible junior synonym of Perissomyrmex under Pristomyrmex. Practically,
since there have been only two known species of the Perissomyrmex until 2000, the generic
characters were not clear. Bolton (2003) diagnosed the tribal characters, but the generic
relationship among the members of the Myrmecinini to which the Perissomyrmex and
Pristomyrmex belong is still unclear.
Thus, to summarize the taxonomic situation concerning Perissomyrmex, the following
questions exist: (1) the identities of Perissomyrmex and Pristomyrmex, (2) phylogenetic
relationships among the genera of the tribe Myrmecinini, and (3) disjunct distribution of
Perissomyrmex. We have already given a brief discussion of this matter (Ogata & Okido, 2002),
with a report of an undescribed species from China.
Recently Radchenko (2003) described a new species of Perissomyrmex from Nepal and
India, which is different from our Chinese species. Now the genus includes four species and
these allow insights into the taxonomic status of the genus and the tribe. The present study
analyzes the phylogeny of the genera of the tribe Myrmecinini based on morphological
characters. Finally, we clarify the taxonomic position of Perissomyrmex in relation to
Pristomyrmex, giving a key to the species, redescriptions of three known species and a
description of a new species of Perissomyrmex from southwestern China. Finally the
biogeography of Perissomyrmex is discussed.
Measurements and indices
Total Length (TL). The total outstretched length of the individual, from mandibular apex to
gastral apex.
Head Length (HL). The length of the head capsule, measured in a straight line from the mid-
point of a line connecting the median paired teeth of the anterior clypeal margin to the mid-
point of the preoccipital margin, in full-face view.
Head Width (HW). The maximum width of the head excluding eyes in full-face view; in
Perissomyrmex broadest in the anterior part of head.
Cephalic Index (CI). HW/HL × 100
Scape Length (SL). The maximum straight-line length of the antennal scape, excluding the
basal condylar bulb.
Scape Index (SI). SL/HW × 100
Eye Length (EL). The maximum diameter of the eye.
Pronotal Width (PW). The maximum width of the pronotal in dorsal view.
Mesosomal Length (ML). The diagonal length of the mesosoma (= alitrunk) in lateral view
from the point at which the pronotum meets the cervical shield to the posterior base of the
metapleural lobes or teeth.
Gastral Length (GL). The maximum straight-line length of the gaster in lateral view.
Gastral Width (GW). The maximum width of the gaster.
All measurements are expressed in millimeters.
Abbreviations of museums
BMNH The Natural History Museum, London, U.K.

NMB Naturhistorisches Museum, Basel, Switzerland
KU Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan
UCD University of California, Davis, California, U.S.A.
PHYLOGENY AND TAXONOMIC POSITION OF PERISSOMYRMEX
The tribe Myrmecinini and its monophyly

Presently the tribe Myrmecinini comprises 4 extant genera: Myrmecina, Acanthomyrmex,
Pristomyrmex and Perissomyrmex (Bolton, 2003). When Ashmead (1905) established the tribe,
it included two genera: Myrmecina and Podomyrma. Emery (1922) considered the tribe to
include 10 genera, divided into two subgroups: subtribe Myrmecinini including Myrmecina,
Acanthomyrmex, Pristomyrmex and Dacryon; subtribe Podomyrmini including Podomyrma,
Lordomyrma, Atopomyrmex, Dilobocondyla, Atopula and Terataner. Although Wheeler (1922)
did not recognize the subdivisions and did not include the genus Atopula, his concept of the tribe
was almost identical to Emery’s. Later Perissomyrmex was described and assigned to the tribe
(Smith, 1947), Dacryon was synonymized with Podomyrma (Brown, 1973; Taylor & Brown,
1985), and the latest concept of the tribe treats the subtribe Podomyrmini sensu Emery as an
independent tribe (Dlussky & Fedoseeva, 1987: Lordomyrma was removed to the tribe
Pheidolini) or placed under the tribe Formicoxenini (Bolton 1994: Lordomyrma was removed to
the tribe Stenammini). In any case Podomyrma and related genera are removed from the
Myrmecinini and the tribe is presently diagnosed as follows (Bolton, 2003):
(1) Anterior margin of labrum angled abruptly downward preapically so that margin appears
very thick in anterior view
(2) Labrum mediodorsally, at line of down curvature (apparent anterior margin), with a
transverse cuticular raised ridge, or series of 2 - 3 small teeth, or both of these.
(3) Antennal sockets relatively close to anterior margin of head (except in major workers of
Acanthomyrmex).
(4) Median portion of clypeus broad, broadly inserted between antennal sockets, the latter
relatively widely separated.
(5) Lateral portion of clypeus usually raised into a ridge in front of toruli.
(6) Stridulum present on pretergite of abdominal segment IV.
(7) Antennae with 9, 11, or 12 segments, with a 3-segmented club (12-13 segments in male).
Among the characters listed above, Bolton considered the first two to be synapomorphies
but the phylogenetic relationship among the genera of the tribe has been left unchallenged.
The characters and the phylogenetic relationships among the genera

Taxonomic and biogeographic overview of the members of tribe Myrmecinini is as follows.
Myrmecina includes 35 described species, but there are almost as many undescribed species in
the Oriental and Australopapuan faunas, so we are estimating 60-70 species in the genus.
Acanthomyrmex is a relatively small genus including 16 described species from the Oriental and
Indo-Australian regions. Pristomyrmex contains 52 species from the Afrotropical, Oriental,
Australian and the Pacific regions (Wang, 2003). Perissomyrmex comprises three known and
one new species described in this paper, showing disjunct distribution in Central America and
Himalayan region.
We have examined the morphology of above genera and found the following characters as
useful for constructing the phylogeny:
1) Frontal lobe: In most of the genera of the subfamily Myrmicinae, the frontal lobe
covering antennal insertion is developed, as in Myrmecina (Fig. 1), but in the genera
Acanthomyrmex, Pristomyrmex, and Perissomyrmex, the structure is reduced, so that the toruli
are exposed (as in Fig. 2). The character is a synapomorphy of those three genera.
2) Genal carina: The genus Myrmecina has unique carinae originated from mandibular
insertion, running parallel in the genal region, and connecting to the preoccipital carina (Fig. 3).
This structure would be a reinforcement of the head capsule. A similar structure is also observed
in Vombisidris, but since the position is different, this would be homoplasous. In any case, the
genal carina of Myrmecina is an autapomorphy of the genus.
3) Antennomere count: The number of antennal segments in worker and queen varies in
genera: 12 in Myrmecina and Acanthomyrmex, 11 in Pristomyrmex, and 9 in Perissomyrmex.
Bolton (2003) showed the count 11 to 12 in Myrmecina, possibly the result of a miscount of
Myrmecina cacabau (see Brown, 1971), in which Taylor (1980) found the 3rd antennal segment
to be distinctly small and hairless. We have confirmed that this is true of all the species of the
genus Myrmecina and unique to the genus (Fig. 4). The reduction in size of the 3rd antennal
segment would be a synapomorphy of the genus. Judging from this situation, the reduction of the
antennomere count in Pristomyrmex would be a loss of the original 3rd antennal segment. And if
the further trait on the reduction in size of an antennal segment were observed in Pristomyrmex,
we could conclude a successive reduction of antennomere count in Perissomyrmex. But there is
no such structure in Pristomyrmex, so we suppose the reduction occurred independently in
Pristomyrmex and Perissomyrmex.
4) Antennal scape base: Moffett (1986) showed that the basal end of the antennal scape in
Acanthomyrmex, Pristomyrmex and Myrmecina has a characteristic structure which forms a
flange encircled by a groove (as in Fig. 5). We found the same structure in Perissomyrmex. The
situation in Myrmecinini is unique and thus the flange is a synapomorphy of the tribe. The
structure might be correlated with the reduction of the frontal lobe, since the flange sometimes
develops to form a lamella covering the antennal insertion.
5) Labrum: The labrum of the tribe is also unique in having a ridge or teeth on the dorsal
surface. In most cases the labrum is a simple flap-like structure covering the labio-maxillary
complex. But in the tribe Myrmecinini as Bolton (2003) described, the anterior portion is sharply
bent inward forming angled preapical margin. Along with transverse preapical margin a ridge or
teeth are developed (Figs. 6). The presence of dorsal teeth on the labrum in Myrmecina and
Pristomyrmex was pointed out by Ogata (1991), and the transverse ridge in Perissomyrmex by
Longino & Hartley (1994) (‘ledge’). In Acanthomyrmex, the portion bears transverse ridge. The
structure suggests a peculiar function, that is, they use the labrum not only for a cover of labio-
maxillary complex but also for grasping or fixing an object with mandibles.
6) Mandible: The mandible of ants is typically subtriangular in shape, with basal shaft,
basal margin, masticatory margin and outer margin in dorsal view. The terminology of the
mandible and dentition is sometimes inconsistent. For example, in Perissomyrmex, Smith (1947)
used “superior border” in describing the basal margin, Baroni Urbani & de Andrade (1993)
interpreted the tooth on the basal margin as “basal tooth”. Here we use the names of teeth only
on the masticatory margin. In the genera Pristomyrmex and Perissomyrmex, the mandibles are
sometimes referred to as elongate (e.g. Smith, 1947). This is due to the extension of the basal
margin of the mandible and the masticatory margin becomes short and somewhat vertical to the
long axis of the head. Figs. 7 and 8, where the left mandible is removed, show contrasting
examples of the angle of masticatory margin in Acanthomyrmex and Pristomyrmex. In
Acanthomyrmex (Fig. 7), like in most of the genera of Myrmicinae, the masticatory margin
inclines from dorsal point that fits for anterior margin of the clypeus to ventral point, so that the
masticatory margin is elongate, while in Pristomyrmex (Fig. 8) and Perissomyrmex the axis of
the masticatory margin turns outward, which probably correlates with elongate basal shaft and
basal margin. Although the elongation of the basal shaft of mandible is also observed in
Myrmecina, the vertical masticatory margin is distinct and would be a synapomorphy of
Pristomyrmex and Perissomyrmex. In both genera, the basal margin has a protuberance in the
middle; broadly produced and sometimes weak in Pristomyrmex (Wang, 2003) but forming a
distinct tooth in Perissomyrmex.
7) Mesopleuron: Anterior free margin of lower part of mesopleuron sometimes expands to

form a lamella partly covering the procoxa. This situation is observed in Myrmecina and
Pristomyrmex (Fig. 9). The expansion of the mesopleuron would be an apomorphy, but not a
synapomorphy in both genera, because the situation occurs in many genera of different tribes
such as Hylomyrma, Mayriella, Paedalgus, Tetramorium. In his terminology of Bolton (1994),
the mesopleuron was divided into upper anepisternum and lower katepisternum by a transverse
groove. But this usage is not correct, because the groove is a secondary structure and not
homologous with the paracoxal suture which separates the anepisternum from the katepisternum
(Matsuda, 1970). The mesopleural part of ants is largely occupied by the anepisternum.
8) Spines on mesosoma: Spines are often developed on the pronotum in most

Pristomyrmex species, all the Acanthomyrmex species and some Myrmecina. It is unclear
whether this structure in the different genera is homologous or not. One of the unique characters
of Myrmecina is paired metanotal teeth in front of the base of the propodeal spines, although not
clearly developed in all the species.
9) Metanotal groove: The ‘propodeal suture’ mentioned by Baroni Urbani & de Andrade
(1993) is synonymous with the metanotal groove. They inferred that the loss of the suture would
be an autapomorphy of Pristomyrmex. We confirmed their statement, but the loss also occurs in
Myrmecina which would be a homoplasy.
10) Tibial spur on middle and hind legs: Smith (1947) mentioned that the absence of
spurs on the middle and hind legs of Perissomyrmex is one of the characters to distinguish it
from Pristomyrmex. But actually Pristomyrmex has no spurs and only Acanthomyrmex has long
and distinct spurs in the tribe Myrmecinini. The absence of the tibial spurs on middle and hind
legs is found in many genera of Myrmicinae.
11) Petiole: The petiole of Myrmecina is unique in shape. Most of the keys say "sessile" for
describing the form (e.g. Bolton, 1994), but it would be interpreted that the node is depressed
because the anterior oblique face corresponds to peduncle of the petiole (Fig. 11). This
characteristic shape of the petiole is an autapomorphy of Myrmecina. In Acanthomyrmex the
petiolar node is consistently bispinose, which is an independant synapomorphy of
the genus.
12) First gastral tergum: Acanthomyrmex has a unique ventral margin of the 1st gastral
tergum, forming a M-shaped line (Fig. 12), which is due to a characteristic constriction of the
anterior part of the 1st tergum. This is a synapomorphy of the genus.
Those characters discussed above are summarized in Table 1. Based on these, applying a
classical Hennigian algorithm, the provisional phylogenetic relationships among four genera are
shown in Fig. 13. Now it is clear that the genera Perissomyrmex and Pristomyrmex are each a
monophyletic taxon justified by synapomorphies, and they together form a sister group within
the tribe Myrmecinini.
Perissomyrmex M. R. Smith
Perissomyrmex M. R. Smith 1947: 281. Type-species: Perissomyrmex snyderi, by original

designation.
DIAGNOSIS
Worker and Female

Monomorphic or polymorphic terrestrial myrmicine ants with the following combination of
characters:
1. Head almost as broad as long in minor worker, or slightly broader in major worker,
divergent anteriorly.
2. Eyes protruding, situated at about the midlength of the side of head, or slightly
posteriorly.
3. Clypeus with a few paired teeth on the anterior margin; lateral portions reduced to a
thin raised strip in front of the antennal insertions.
4. Frontal lobe absent; frontal carina poorly developed.
5. Antennal toruli exposed.
6. Antennae 9-segmented, scape extending over posterior border of head, apical 3
segments forming a club.
7. Mandibles with long basal margin and short masticatory margin; basal margin with a
distinct tooth; masticatory margin with one basal and two apical teeth separated by a
diastema.
8. Labrum with transverse ridge at the preapical portion.
9. Palp formula 4,2.
10. Pronotum unarmed, promesonotal suture weakly impressed.
11. Lower part of mesopleuron not expanding to cover on procoxa.
12. Metanotal groove deeply impressed.
13. Propodeal spiracle rounded situated in the middle of lateral surface of propodeum.
14. Propodeal spine long and acute.
15. Propodeal lobe low and obtuse angled.
16. Petiole pedunculate with lateral projection, petiolar node rounded.
17. Postpetiole with rounded node.
Male Unknown.
Of the four species of the genus, P. snyderi is easily diagnosed by the presence of a
subpetiolar process in the workers, the dentition of the anterior margin of the clypeus and the
coarse and irregular sculpturation on the head. This does not mean that the three Asian species
comprise a monophyletic group. Because the loss of the subpetiolar process is frequently
observed in Myrmecina and Pristomyrmex, we do not confirm the polarity of the character state
of the dentition and the sculpture. There is no strong evidence, however, that Central American
snyderi has a close relationship with any of the Asian species. The detailed phylogeny of the
four species is not clear.
The polymorphism of the genus was reported by Longino & Hartley (1994) in P. snyderi
from Mexico. While no Asian species are known to be polymorphic, it is possible. Because the
available materials were mostly from litter extractions, whole colony composition has not been
clear.
It should be noted that the individuals of the Perissomyrmex are highly variable in the
morphology. In particular, projected structures, such as teeth on the mandibles, propodeal spines,
lateral projections on the petiolar peduncle, are varying due to wearing down or polymorphism.
Perissomyrmex species
emarginatus n. sp.
monticola de Andrade
nepalensis Radchenko
snyderi Smith, M.R.
KEY TO PERISSOMYRMEX SPECIES (workers)
1. Subpetiolar process small but distinct (Fig. 17d); median paired teeth of anterior margin
of clypeus distinct but base not forming differentiated tubercle so that outline of
anterior clypeal margin is evenly convex (Fig. 16g, h); body almost blackish; Central
America .............................................................................................................................
............................................................................................................snyderi M.R. Smith
-- Subpetiolar process absent (Fig. 17a-c); median paired teeth on anterior clypeal margin
strongly protuberant so that there is a distinct median projection (Fig. 16a-f); body color
brownish; Asia................................................................................................................. 2
2. Lateral paired teeth on anterior clypeal margin close to median pair, bases fused
(Figs.16a-c); median notch of anterior margin of clypeus inverted U-shape (Fig. 16a);
SW China .......................................................................................... .emarginatus n. sp.
-- Lateral paired teeth on anterior clypeal margin well isolated from median paired teeth
(Figs. 16d-f); median notch of anterior margin of clypeus inverted V-shape.................. 3
3 Ventral margin of propodeum mostly straight, curved in front of the area below spiracle
(Fig. 17b); gaster reddish brown, contrasting with blackish mesosoma; Bhutan...............
....................................................................................................... monticola de Andrade
-- Ventral margin of propodeum curved behind the area below the spiracle (Fig. 17c);
gaster concolorous with mesosoma, but lighter; Nepal & NE India..................................
...................................................................................................... nepalensis Radchenko
SPECIES ACCOUNTS
Perissomyrmex emarginatus new species

Figures 14, 16a-c, 17a
DESCRIPTION
Worker: Holotype measurements (mm): TL 3.73, HL 0.86, HW 0.90, , SL 0.78, EL 0.10,

PW 0.57, ML 0.98, GL 1.07, GW 0.86. Indices: CI 104.65, SI 86.67.
Paratype measurements (4 workers, 3 measured): TL 3.97-4.14, HL 0.86-0.96, HW 0.92-
0.94 , SL 0.74-0.78, PW 0.57-0.59, ML 0.90-1.11, GL 1.07-1.39, GW 0.86-0.90). Indices: CI
95.74-106.98, SI 82.22-86.67.
Head subrectangular, with slightly concave posterior border and rounded posterior corners;
sides slightly diverging anteriorly. Eyes situated in the midlength of sides, comprising 15-18
ommatidia. Anterior margin of clypeus with four teeth; median paired teeth large and distinct,
lateral paired teeth smaller and lower; the median paired ones separated by median notch, the
lateral tooth located just in front of antennal insertion, and the median and lateral ones closely
situated each other, sometimes fused at the base in each side; median notch distinct and deep,
inverted U-shape, the bottom of notch reaching or exceeding level of anterior margin of lateral
edge in front of antennal insertion. Frontal lobes almost reduced, frontal carina low, diverging
posteriorly, reaching the level of eyes. Antennal toruli exposed, antennal insertion forming deep
hollow forming distinct anterior edge. Mandible elongate with long basal margin and short
masticatory margin; basal margin with one small but distinct tooth in the midlength; masticatory
margin equipped with one apical and one subapical teeth followed by a diastema and one basal
tooth. Antennae 9 segmented, scape exceeding posterior corner of head by its width; apical three
segments forming club but obscure. Labrum with transverse preapical ridge; two small
projections present on the dorsal surface in the middle.
Promesonotum raised, but the dorsal surface somewhat depressed; promesonotal suture
weakly impressed. Anteroventral free margin of mesopleuron carinated but not forming
expansion covering procoxa. Metanotal groove complete, deeply impressed. Dorsal outline of
propodeum slightly raised. Spiracle circular and large, situated on lateral surface of propodeum
in the middle. Propodeal spine straight, long and acute, directed backward and upward.
Propodeal lobe low and rounded.
Petiole with anterior declivity, rounded tip and posterior face; ventral margin weakly sinuate
but no subpetiolar process; anterior peduncle with distinct paired lateral projections in dorsal
view, situated anterodorsal to the spiracle in lateral view. Postpetiole, in profile, with raised
dorsal outline, anterior face convex, posterior face nearly vertical and in frontal view the crest
rounded but pointed. Ventral margin to 1st gastral segment not sinuate.
Head, mandible and mesosoma costate, lower part of mesopleuron and posterior part of
propodeal spiracle unsculptured. Petiole mostly unsculptured. Anterior face of postpetiole
smooth and shining, the rest finely punctate. Body dark yellowish brown, the appendages lighter.
Queen. TL 4.63-4.92, HL 1.07-1.19, HW 1.11-1.23, CI 103.36-110.81, SL 0.78-0.90, SI
68.91-73.17, PW 0.82-0.90, ML 1.31-1.39, GL 1.35-1.48, GW 1.11-1.19 (5 measured).
Similar to the worker, but the head wider than long with three ocelli. The dentition on
mandibles and anterior clypeal margin basically same as in worker but tending to be worn down,
so that the teeth are often low and obscure; the median portion of anterior clypeal margin
produced with less distinct median notch and four teeth which are sometimes fused together.
Mesosoma robust short and high, mesonotum thick, mesoscutellum overhanging metanotum,
metanotum more of less convex in profile, propodeal spine robust directed backward. Petiole
basically as in worker, but the ventral surface with distinct paired longitudinal keel originated
from posterior end of the petiole running anteriorly and convergent at anterior 1/4 of petiole,
lateral surface with longitudinal ridge from the portion above the spiracle to posterior end, and
lateral projection more distinct. Apex of postpetiolar node pointed. Sculpture more distinct than
that of worker. Coloration same as worker.
TYPE MATERIAL
Holotype, Worker, Lianwangpo, Mt. Emeishan, Emeishan Shi, Sichuan Prov., CHINA ,
2310m alt., 4.x.1996 (S. Nomura & Zhao) (KU No. 3149 ). Paratypes (All from CHINA): 4
workers, 2 queens, same as holotype; 2 queens, Mt. Jiudingshan, Chapingshan Mts., Mao Xian,
Sichuan Prov., 3670 m alt., 22.ix.1996 (S. Nomura & Zhao); 1 queen, Leidongping, Mt.
Emeishan, Emeishan Shi, Sichuan Prov., 2310-2350 m alt., 4.x.1996 (S. Nomura & Zhao).
Remarks
All the material was collected by Berlese funnel. Collecting site of Mt Emeishan is a road
side of mixed coniferous forest, Dabei site is more like evergreen forest, and Mt Jiudingshan site
is somewhat shrubby. The altitudinal range of collecting sites is from 2300 m to 3700 m elev-
ation. The lateral projections of the petiolar peduncle are sometimes asymmetrically developed.
In the worker caste, P. emarginatus is distinguished from two other Asian species in having
the deeper median notch on the anterior clypeal margin. The bottom of the notch reaches or even
exceeds the level of anterior ridges in front of antennal insertions. The body of P. emarginatus
has more sculptured surface and the color is more brownish than the two other Asian species.
The propodeal spines, if they are fully developed, are also different from those of P. monticola
and P. nepalensis, being straight without upper curve. In the queen the median notch is
shallower than that of the nonspecific workers, so that the depth and the shape cannot be used in
separating species. Although we did not observe the queen of P. nepalensis, the dentition of the
anterior clypeal margin is still useful; the 4 teeth are arranged closer together forming a blunt
median projection in P. emarginatus, while the lateral paired teeth are small and separate from
the median pair in P. monticola.
Perissomyrmex monticola de Andrade

Figures 16d,e, 17b
Perissomyrmex monticola de Andrade, in Baroni Urbani & de Andrade 1993: 90. Holotype worker.
Bhutan.
Diagnosis
Worker, measurements (mm): TL 4.6, HL 1.12, HW 1.08, SL 0.92, PW 0.64, AL 1.24, GL
1.35, GW 0.96. Indices: CI 96.4, , SI 85.2,
Four teeth of anterior margin of clypeus well isolated; median paired teeth large and
distinct; lateral paired teeth small and their tip not reaching the level of the bottom of median
notch; median notch converted V shape; the bottom of notch not reaching the level of anterior
margin of lateral ridge in front of antennal insertion; distance between a median teeth closer than
that between a median tooth and a lateral tooth; position of lateral tooth just below in the middle
of toruli. Promesonotum forming single raised convex. Propodeal spines long and acute,
directing backward and curved upward. Propodeal lobe low and rounded. Following the
restriction on anterior articulator portion to mesosoma, sides of petiole almost parallel in dorsal
view.
Posterior face of postpetiolar node declined. Head capsule and mesosoma heavily striate.
Mandibles very lightly striate. Mesopleural area, propodeum smooth and shining, petiole smooth
and shining.
Body bicolored, with dark brown head and mesosoma and reddish gaster, antennae and legs
yellowish.
Queen: TL 5.32, HL 1.20, HW 1.20; CI 100, SL 1.00, SI 75, PW 0.84, AL 1.52, GW 1.20,
GL 1.56.
Dentition of anterior margin of clypeus and mandible as in worker, but robust and lower.
Mesoscutellum overhanging metanotum, metanotum less convex in profile. Petiole as in worker
but with low ventral keel and more distinct lateral projection.
Remarks
Seemingly P. monticola is distinct in having a bicolored body with blackish head and
mesosoma and reddish gaster. The species is similar to P. nepalensis in having an inverted V
shaped median notch on the anterior clypeal margin. The notch in P. monticola and P.
nepalensis is shallower than in P. emarginatus. The bottom of the notch in both species does not
reach the level of anterior ridges in front of the antennal insertions. The teeth arrangement of the
anterior clypeal margin is similar to that of P. nepalensis as well, showing that the lateral paired
teeth are more spaced to the median paired teeth. The shape of the propodeal spines, upward
curved profile, also show the similarity in P. monticola and P. nepalensis. According to the key
provided by Radochenko (2003) the species is distinct in having abundant short standing hairs
on the antenna and legs, but the character is somewhat delicate and P. emarginatus also shows
the same pilosity. It seems that the ventral profile of the petiole is more reliable.
Among the three Asian species, the sculpture of P. monticola is the most distinct, and the
unsculptured areas on the mesopleuron and propodeum are broadest. The species is known only
from the type locality.
Specimens examined: 1 worker, 1 queen, Nobding 41 km O, Wangdi Ph. 2800 m, Bhutan

(NMB).
Perissomyrmex nepalensis Radchenko

(Figs. 16f, 17c)
Perissomyrmex nepalensis Radchenko 2003:12. Holotype worker. Nepal.
Diagnosis
Worker, measurements (mm): TL 3.74-4.38, HL 0.90-1.10, HW 0.92-1.10, SL 0.8-0.9, PW
0.56-0.64, ML 1.00-1.10, GL 1.02-1.30, GW 0.80-0.96. Indices: CI 100-102, SI 82-87.
Four teeth on anterior margin of clypeus well isolated; median paired teeth large and
distinct, lateral paired teeth small and spaced from the median teeth; median notch converted V
shape; the bottom of notch not reaching anterior margin of lateral ridge in front of antennal
insertion. Promesonotum raised but less convex. Propodeal spine directed backward and curved
upward. Ventral margin of petiole nearly straight. Subpetiolar process absent. Costulation on
head and pronotum low and spaced, unsculptured area on mesosoma and propodeum wider.
Body color light reddish brown, antennae and legs yellowish.
Specimens examined: 2 workers, W. Bengal, Darjeering, Tiger Hill, 2450 m, 28.viii.1997,

India (BMNH).
Remarks
The dorsal outline of the promesonotal area varies in the size of workers. In minor worker
the area is less raised but in major worker weakly convex in profile. P. nepalensis is
distinguishable from P. monticola by the ventral profile of the petiole: nearly straight in the
former, but widely convex in the latter. Sculpturation of P. nepalensis is weaker and more
spaced than that of P. montiocola. Radchenko (2003) noticed the standing hairs on the antennae
and legs in distinguishing P. nepalensis from P. monticola; short and abundant in the former,
long and sparse in the latter.
The species was found in Himalayan region (Nepal and northeastern India) at the altitudinal
range of 2450 to 3000m, collected by pitfall traps (Radochenko, 2003).
Perissomyrmex snyderi Smith, M.R.

Figures 16g, h, 17d
Perissomyrmex snyderi Smith, M.R. 1947:282. Holotype worker. Guatemala.
Diagnosis.
Worker, TL 3.74-4.94, HL 0.88-1.24, HW 0.88-1.34, CI 100-108, SL 0.82-0.94, SI 73-93,
PW 0.56-0.72, ML 0.98-1.20, GL 1.00-1.40, GW 0.80-1.06.
Polymorphic. Labrum with lamellate flange developed at basal part of curvature. Anterior
clypeal margin with 3 pairs of projection, the shape and size varying to reduce or fused;
basically, the median pair broad and robust, inner lateral pair smaller; outer lateral pair smallest,
sometimes reduced or lost; the median notch roundly concave; in some case the median paired
teeth fused without median notch in major worker. Propodeal spine straight or slightly curved
upward. Subpetiolar process present anteriorly; ventral margin of petiole almost straight in
profile; anterior portion of petiole diverging toward midlength of petiole in dorsal view.
Sculpture on head and mesosoma irregularly costate. Body color blackish.
Specimens examined: 9 workers (including 1 major), 4 km N Union Juarez, Volcan

Tacann, Lower slopes 1950 m, Chiapas, Mexico (4 in UCD, 5 in BMNH).
Remarks
The species shows great variation in worker caste body size, the shape of spines and the
teeth on the anterior clypeal margin, but the subpetiolar process is consistently present, which is
unique among the four Perissomyrmex species. In addition, P. snyderi is also unique in having
coarser and more irregular sculptures on the head and mesosoma and dark color of the body. The
degree of the protuberance of the anterior clypeal margin is less distinct than that in Asian
species, even though variation exists.
The rediscovered individuals were collected from Berlese samples of a cloud forest floor of
1700 to 2000 m altitude in Chiapas, Mexico (Longino & Hartley, 1995).
Biogeography of Perissomyrmex
Perissomyrmex has a unique biogeographic pattern showing a disjunct Oriental-Neotropical
distribution (Fig. 15). According to Bolton’s census data (1994) there are no genera showing
such disjunct distribution in ants. Seemingly the genus is a tropical element, but the actual
distribution records are restricted to higher elevation at moderate northern latitude; 1700 to 2000
m in Central America and over 2000 m in Asia. The species of Asia are never found in the
tropical mountains nor in lowlands of temperate zones. The distribution range of the genus in
Asia is shown in Fig. 18 with altitude and latitude forest zones. Their habitat includes Lauro-
Fagaceae forest, evergreen forest, temperate conifer & deciduous forest. The genus cannot be a
tropical element but might be a relic of an old temperate taxon.
Baroni Urbani & de Andrade (1993) and Radchenko (2003) supposed the origin of the
genera in Asia. This is supported by the present study as follows: (1) the sister group of
Perissomyrmex is Pristomyrmex, distributed in the Old World Tropics; (2) a clade of
Perissomyrmex + Pristomyrmex is closely related to Acanthomyrmex, also distributed in tropical
Asia (Fig. 18), and (3) species diversity of Perissomyrmex is highest in Asia. Considering the
present ecological distribution of Perissomyrmex, the speciation of the ancestor of
Perissomyrmex + Pristomyrmex might have occurred in temperate-tropical regions.
Longino & Hartley (1994) discussed the origin of the disjunct distribution and mentioned
that P. snyderi is not an accidental introduction to Central America but a relic of widely
distributed taxon. If their hypothesis is correct, the Asian and Central American species would
be different monophyletic species groups. Morphologically, the Asian species share several
characters, e.g. the loss of the subpetiolar process, the distinct protuberance of the median
portion of the anterior clypeal margin, and more or less parallel sculpturation on the head. But
because of the difficulties of their polarities as mentioned above, we have not confirmed
monophyly.
The evidence of vicariance might be in fossil records. There are two more extinct genera of
the tribe Myrmecinini: Stiphromyrmex Wheeler and Ennaemerus Mayr of the Baltic Amber
inclusions. Among them, the genus Ennaemerus has a 9 segmented antenna, as in
Perissomyrmex. It should be noted that when Wheeler (1915) referred to the resemblance of
Ennaemerus to Pristomyrmex, Perissomyrmex was not discovered yet. If the tribal assignment of
them is correct, Ennaemerus would be a close relative of Perissomyrmex. And the ‘Proto-
Perissomyrmex’ (Ennaemerus + Perissomyrmex) mights have a wide range of distribution. For
reference of this pattern, Myrmecina, the early derivative of the tribe, shows wide range of
distribution (Fig. 19). The difference is that Myrmecina would be basically a tropical origin,
while Perissomyrmex is a temperate one.
Tiffney (1985) suggested that there are five major periods of migration between eastern
Asia and eastern North America: pre-Tertiary, Early Eocene, Late Eocene-Oligocene, Miocene
and Late Tertiary-Quaternary. Recent molecular analysis of Magnoliaceae (Azuma et al., 2001)
showed that tropical disjunction occurred during the middle to late Eocene and that the
temperate disjunction occurred in the Oligocene. General resemblance of the distribution pattern
of that plant might be referential in considering the disjunction of Perissomyrmex. Since
geological time of the Baltic Amber is thought to be the Late Eocene (e.g. Dlussky, 1999;
Bolton, 2003), the ‘Proto-Perissomyrmex’ (Ennaemerus + Perissomyrmex) might have Arcto-
Tertiary distribution, expanding their distribution from Eurasia to North America during one of
the three possibilities of Tiffney’s five periods of migrations: Late Eocene-Oligocene, Miocene
and Late Tertiary-Quaternary. The present distribution would be a relic of the old temperate
range in Paleogene. Molecular analysis would be needed to estimate the detailed phylogeny and
diverging time of disjunction.
ACKNOWLEDGEMENTS
We thank Shuhei Nomura for providing samples from China; P. S. Ward, M.L. de Andrade, G.
R. Else, and S. Ryder for their kindness in arranging loans of specimens; and R. Snelling for
arranging publication. The study was partly supported by a Grant-in-Aid for an Overseas
Research Project (No 14255016) from the Ministry of Education, Science, Technology, Sports
and Culture, Japan.
LITERATURE CITED
Ashmead, W.H. 1905. A skeleton of a new arrangement of the families, subfamilies, tribes and
genera of the ants, or the superfamily Formicidae. Canadian Entomologist 37: 381-384.
Azuma, H. Garcia-Franco, J.G., Rico-Gray, V. & Thien, L.B. 2001. Molecular phylogeny of
the Magnoliaceae: The biogeography of tropical and Temperate disjunctions. American
Journal of Botany 88: 2275-2285.
Baroni Urbani, C. & de Andrade, M.L. 1993. Perissomyrmex monticola n. sp., from Bhutan:
the first natural record for a presumed Neotropical genus with a discussion on its taxonomic
status. Tropical Zoology 6:89-95.
Bingham, C.T. 1903. The fauna of British India, including Ceylon and Burma. Hymenoptera 2.
Ants and Cuckoo Wasps, 506 pp. London.
Bolton, B. 1981. A revision of six minor genera of Myrmicinae in the Ethiopian
zoogeographical region. Bulletin of the British Museum (Natural History) (Entomology) 43:
245-307.
Bolton, B. 1994. Identification Guide to the Ant Genera of the World: 222 pp. Cambridge, Mass.
Brown, W.L., Jr. 1971. Characters and synonymies among the genera of ants. Part 4. Some
genera of subfamily Myrmicinae. Breviora 365: 1-5.
Brown, W.L., Jr. 1973. A comparison of the Hylean and Congo-West African rain forest ant
faunas. Pp. 161-185. in: Meggers, B.J., Ayensu, E.S. & Duckworth, W.D. (eds). Tropical
Forest Ecosystem in Africa and South America: a Comparative Review. 350 pp. Washington
D.C.
Dlussky, G.M. 1997. Genera of ants from Baltic Amber. Paleontological Journal 31: 616-627.
Myrmicinae. Fasc. 174C: 207-397. Bruxelles.
Longino, J.T. & Hartley, D.A. 1994. Perissomyrmex snyderi (Hymenoptera: Formicidae) is
native to Central America and exhibits worker polymorphism. Psyche 101: 195-202.
Matsuda, R. 1970. Morphology and evolution of the insect thorax. Memoirs of the
Entomological Society of Canada 76: 1-431.
Moffett, M.W. 1986. Revision of the myrmicine genus Acanthomyrmex. Bulletin of the Museum
of Comparative Zoology 151: 55-89.
Ogata, K. 1991. A generic synopsis of the poneroid complex of the family Formicidae
(Hymenoptera). Part II. Subfamily Myrmicinae. Bulletin of the Institute of Tropical
Agriculture, Kyushu University 14: 61-149.
Ogata, K. & Okido, H. 2002. Taxonomy of the ant tribe Myrmecinini with special references to
the enigmatic genus Perissomyrmex. Proceedings of the XIV International Congress of
IUSSI, 27 July - 3 August 2002, Hokkaido University, Sapporo, Japan, p. 157.
Ohsawa, M. 1990. An interpretation of latitudinal patterns of forest limits in South and East
Asian mountains. Journal of Ecology 78: 326-339.
Radchenko, A. 2003. Perissomyrmex nepalensis sp. nov. - new evidence of Old World origin
for the genus. Entomologica Basiliensia 25: 13-22.
Smith, M.R. 1947. A new genus and species of ant from Guatemala. Journal of the New York
Taylor, R.W. 1980. The rare Fijian ant Myrmecina (=Archaeomyrmex) cacabau (Mann)
rediscovered. New Zealand Entomologist 7: 122-123.
Taylor, R.W. & Brown, D.R. 1985. Zoological Catalogue of Australia 2. Hymenoptera:
Formicoidea, Vespoidea and Sphecoidea: 381 pp. Canberra
Tiffney, B.H. 1985 Perspectives on the origin of the floristic similarity between eastern Asia and
eastern North America. Journal of the Arnold Arboretum 66: 243-273.
Wang, M. 2003 A monographic revision of the ant genus Pristomyrmex (Hymenoptera:
Formicidae). Bulletin of the Museum of Comparative Zoology 157:383-542.
Wheeler, W.M. 1915. The ants of the Baltic Amber. Schriften der Physikalisch-Ökonomischen
Gesellschaft zu Königsberg 55 (1914): 1-142.
Wheeler, W.M. 1922. The ants of the Belgian Congo. Bulletin of the American Museum of
Natural History 45: 1-1139.
Figures 1-12. Characters of Myrmecinini: antennal insertion of Myrmecina (1), Pristomyrmex (2);
genal carina of Myrmecina (3); base of funiculus of Myrmecina (4), base of scape of Myrmecina (5);
labrum of Pristomyrmex (6); masticatory margin of Acanthomyrmex (7), Pristomyrmex (8);
mesosoma of Pristomyrmex (9); propodeal spine of Myrmecina queen (10); petiole of Myrmecina
(11); 1st gastral tergum of Acanthomyrmex (12).
Table 1. Characters in the tribe Myrmecinini
# Apomorphic Characters
1 Preapical portion of labrum sharply bent inward
2 Transverse ridge or teeth on anterodorsal part of labrum
3 Basal fringe at antennal scape
4 Ventral carina on the head
5 Reduction of 3rd antennal segment
6 Reduction of petiolar node
7 Loss of frontal lobe
8 M-shaped ventral margin of 1st gastral tergum
9 Petiolar node bispinose
Short masticatory margin of mandible nearly vertical to long axis of
10
head
11 Protuberance on basal margin of mandible in the midway
12 11-segmented antenna
13 Expansion of mesopleuron partly covering procoxa
14 Loss of metanotal groove
15 9-segmented antenna
16 Distinct tooth on basal margin of mandible
17 Lateral projection on petiolar peduncle
Figure 13. Phylogenetic relationships of the genera of the tribe Myrmecinini. Numbers on each
clade show synapomorphies listed in Table 1.
Fig. 14. Perissomyrmex emarginatus n. sp.
Figure 15. Distribution of Perissomyrmex
Figure 16. Anterior clypeal margin of Perissomyrmex spp., worker of P. emarginatus (a),
female of P. emarginatus (b-c), worker of P. monticola (d), female of P. monticola (e), worker
of P. nepalensis (f), minor worker of P. snyderi(g), major worker of P. snyderi (h).
Figure 17. Petiole of Perissomyrmex workers in profile, P. emarginatus (a), P. monticola (b), P.
nepalensis (c), P. snyderi (d).
Figure 18. Distribution range of Perissomyrmex spp. in Asian forest zones (altitudinal and
latitudinal vegetation in SE & E Asia after Ohsawa, 1990).
Figure 19. Distribution of the genera of the tribe Myrmecinini.

Schödl, S. 2007. Revision of Australian Meranoplus: the Meranoplus diversus group, pp. 370-
424. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics
(Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of contributions. Memoirs
of the American Entomological Institute, 80.
REVISION OF AUSTRALIAN MERANOPLUS:

THE MERANOPLUS DIVERSUS GROUP
Stefan Schödl1
2. Zoologische Abteilung
Naturhistorisches Museum
Burgring 7, A-1014 Vienna, Austria
ABSTRACT
The Meranoplus diversus group of species is taxonomically revised. Twenty-five species are
recognized, nineteen of which are described as new herein: arcuatus sp.n., angustinodis sp.n.,
berrimah sp.n., christinae sp.n., convexius sp.n., crassispina sp.n, deserticola sp.n., digitatus
sp.n., discalis sp.n., diversoides sp.n., mcarthuri sp.n., naitsabes sp.n., occidentalis sp.n.,
orientalis sp.n., snellingi sp.n., taurus sp.n., tricuspidatus sp.n., variabilis sp.n. and wilsoni sp.n.
Lectotypes are designated for M. duyfkeni Forel, 1915, M. mars Forel, 1902, M. oxleyi Forel,
1915 and M. unicolor Forel, 1902. The status of M. dichrous Forel, 1907 remains doubtful.
Morphological details are figured and distribution data based on this study are presented.
Key words: Hymenoptera, Formicidae, Myrmicinae, taxonomy, lectotypes, new species, key.
1
Deceased 20 April 2005
Schödl: Revision of Australian Meranoplus 371
INTRODUCTION
Meranoplus is an Old World genus, occuring in the Ethiopian, Oriental and Australian regions.
Fewer than 30 Australian taxa are presently known (Bolton, 1995; Shattuck, 1999; Andersen,
2000), and the taxonomy of several still needs to be resolved (Taylor, 1990; Schödl, 2004).
However, these relatively few species by no means reflect the actual number of taxa found
throughout Australia with an overwhelming diversity and with countless still undescribed
species. Amongst others a group of apparently seed-harvesting species, the M. diversus group
(Andersen, 2000, and citations therein), has evolved in Australia. This group alone comprises
more species than found in the combined Ethiopian and Oriental realms (Bolton, 1981; Schödl,
1998). The members of this group, very distinct within the genus and for the Australian fauna,
are herein treated taxonomically for the first time. The type of Meranoplus dichrous Forel, 1907,
known from a single queen, was destroyed during World War II (Taylor, 1990). From the
original description it remains doubtful whether the taxon is conspecific with any of those herein
treated, in particular the smaller, bicolored species.
Material and methods
Dry and point-mounted specimens were examined with an Olympus SZH10 Research
Stereo binocular microscope. Measurements were taken with an ocular grid at magnifications
from 70× - 140×. Line drawings were prepared with a camera lucida apparatus, attached to the
binocular. Distribution maps were generated with the use of BioLink®.
Gynes have not been considered in this study. Although they frequently occur in samples,
they are still missing in too many species. In general they look very much alike workers in head
morphology and thus can be assigned to one or another species.
Morphometric Measurements and Indices
CI Cephalic Index: HW × 100 / HL

CS Cephalic Size: arithmetic mean of HL + HW
EL Eye Length: maximum length of compound eye, measured in lateral view
EW Eye Width: maximum width of compound eye, measured in lateral view
EYE (EL + EW) / CS
FC Frontal Carinae: minimum distance between frontal carinae, measured in full face view; if
frontal carinae are evenly narrowed, measured in front of eyes
FI HW × 100 / FC
HL Head Length: length of head, measured from mid-point of preoccipital margin to mid-point of
anterior clypeal margin
HW Head Width: maximum width of head behind compound eyes, measured in full face view
ML Mesosomal Length: length of mesosoma, measured in lateral view from anterior pronotal
tooth to caudalmost part of basal propodeal lobe
PMD Promesonotal Diameter, measured from apex of pronotal anterolateral projection to apex of
mesonotal posterolateral projection of opposite side.
PMI Promesonotal Index: PW × 100 / PML
PMI2 Promesonotal Index 2: PW × 100 / PMD
PML Promesonotal Length: length of promesonotal shield, measured from anterior mid-point of
pronotum behind collar, that is the mid-point of a virtual line, where the anterior pronotal
margins meet, to mid-point of hind margin of mesonotum (lamella included) above propodeal
declivity
PPI Postpetiolar Lateral Index: PPLL × 100 / PPLH
PPLH Postpetiolar Lateral Height: height of postpetiole, measured in lateral view
PPLL Postpetiolar Lateral Length: length of postpetiole, measured in lateral view
PSL Propodeal Spine Length: length of propodeal spine in lateral view, measured from mid-point
of propodeal spiracle to spinal apex (specimens need to be tilted in case of diverging spines)
PTI Petiolar Lateral Index: PTLL × 100/ PTLH
PTLH Petiolar Lateral Height: height of petiole, measured in lateral view
PTLL Petiolar Lateral Length: length of petiole, measured in lateral view
PW Pronotal Width: maximum width of pronotum, measured posterior to base of anterolateral
pronotal projections (angles) in dorsal view
REL Relative Eye Length: EL / HL
SL Scape Length: length of antennal scape, excluding basal condylar bulb, measured in caudal
view
SI1 Scape Index 1: SL × 100 / HW
SI2 Scape Index 2: SL / CS
TL Total Length: length of out-streched individual from mandibular apex to apex of gaster,
usually measured in lateral view as a sum of HL, ML and nodes + gaster length
Depositories of Studied Material
ANIC Australian National Insect Collection, Canberra, Australia

BMNH The Natural History Museum, London, U.K. [= British Museum of Natural History]
JDMP J.D. Majer collection, Curtin University, Perth, Western Australia, Australia
LACM Natural History Museum of Los Angeles County, Los Angeles, California, USA
MCZC Museum of Comparative Zoology, Cambridge, Massachusetts, USA
MHNG Muséum d'Histoire naturelle, Geneva, Switzerland
MVMA Museum Victoria, Melbourne, Victoria, Australia
NHMB Naturhistorisches Museum, Basle, Switzerland
NHMW Naturhistorisches Museum, Vienna, Austria
NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden
PSWC P.S. Ward Collection, University of California, Davis, California, USA
QMBA Queensland Museum, Brisbane, Queensland, Australia
SAMA South Australian Museum, Adelaide, South Australia, Australia
UQIC University of Queensland Insect Collection, Brisbane, Queensland, Australia
USNM National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
WAMP Western Australian Museum, Perth, Western Australia, Australia
DIAGNOSIS OF M. DIVERSUS GROUP (WORKERS)
Although several of the species found in this group are among the largest in Meranoplus, the
diversus group comprises medium sized to smaller species as well. Species are readily separated
from members of other groups by the distinctive clypeal morphology. The posterior (dorsal) part
of the clypeus is constructed as a massive plate fit in between the anterolateral frontal corners; it
may show various forms of elaborations such as keels or plates or, it may be distintly excavated
at its apparent anterior margin which frequently surpasses the anterolateral frontal corners in
length. The true anterior clypeal section is strongly concave in lateral view and hidden below the
posterior section. Most species have large, massive heads in relation to their body size.
Mandibles are short, stout and armed with three to five teeth and are mostly overhung by the
clypeal plate. These are frequently worn down completely so that the masticatory margin may
secondarily be edentate. The compound eyes generally are of moderate size (except in
occidentalis and taurus). The frontal carinae are markedly narrower than the head, so that the
genae, eyes and antennal scrobes are partly visible in dorsal view. The promesonotal shield
either has very reduced blunt (to almost absent) projections or prominent spines, frequently
laterally flanged and translucently margined. The petiole is triangular to box-shaped, the
postpetiole nodular with an antero-ventral tooth of variable size. Species are unicolorous or
bicolored with the gaster and appendages usually brighter than the remainder of body. Both
color forms may occur in one and the same species.
In general only the larger species with a large, bulbous head capsule and reduced
projections on the promesonotal shield are assigned to the M. diversus group. However the
majority of the species are considerably smaller (see also Andersen, 2000). Due to various
characteristics such as shape of the dorsal shield, clypeal structure and sculpturation of the
dorsal surface species cluster to subgroups. These are not recognized in detail herein, since these
subgroups are not clearly defined.
Species either have a limited distribution, a circumstance that may be due to climatic and
ecological factors, while others are widely distributed without significant restriction to their
radiation (none of the species herein treated is known to have crossed the Torres Strait). Some
species show a higher degree of morphological plasticity than do others. For this reason, some
material has only been tentatively assigned to species. Further material will be needed to clarify
the taxonomic position of these 'left overs'.
Members of the M. diversus group nest in the ground, as evidenced by the large number of
pitfall specimens, label data and observations of collectors. They are granivores and thus play an
important role in seed dispersal (Taylor & Brown, 1987; Andersen, 2000, and citations therein),
comparable to that of of Messor, a genus missing in Australia.
KEY TO WORKERS OF M. DIVERSUS GROUP
The key herein presented does not reflect phylogeny. The worn-down condition of the dentition
in many specimens renders more difficult to decide how many mandibular teeth are structurally
present.
1 Promesonotal shield with posterolateral and posterior projections short and at most bluntly
rounded or triangular, lateral margins of the shield not or only little overhanging lateral
mesosomal sides and propodeal declivity (Figs. 2, 3, 5, 7, 9, 11, 13, 41) .............................. 2
-- Promesonotal shield with posterolateral and posterior projections well developed as acute or
narrowly rounded projections, lateral margins distinctly overhanging lateral mesosomal
sides and propodeal declivity (Figs. 15, 17, 19, 21, 23, 25, 27, 29, 31, 33, 35, 37, 39, 43, 45,
47, 49, 51)............................................................................................................................. 8
2 Paramedian projections on mesonotal hind margin absent (Fig. 41). Eyes very large (EYE
0.37 - 0.42; REL 0.23 - 0.26). (Western Australia) ...............................................occidentalis
-- Paramedian projections on mesonotal hind margin at least present as minute denticles. Eyes
distinctly smaller (EYE < 0.35; REL < 0.22) ......................................................................... 3
3 Clypeus anteriorly bluntly bidentate. Posterior face of petiole costate or rugose. With four
mandibular teeth .................................................................................................................... 4
-- Clypeal projection different. Posterior face of petiole rugose to rugulose. With three
mandibular teeth ..................................................................................................................... 5
4 Posterior petiolar face rugose. Promesonotal shield translucently margined, with mesonotal
projections narrowly rounded (Fig. 11). Dorsal ocular margin well separated from ventral
scrobal margin. (north-western Australia) ..................................................................duyfkeni
-- Posterior petiolar face evenly and densely costate. Promesonotal shield not translucently
margined, with mesonotal projections reduced or seen as minute denticles (Fig. 13). Dorsal
ocular margin reaching ventral scrobal margin. (Queensland) ..................................orientalis
5 Clypeus with conspicuous dorsal median keel-like carina, accompanied by lateral more or
less developed carinate projections (Figs. 52, 53). (Widespread throughout most of arid to
monsoonal Australia, absent in southernmost parts) ......................................................... ajax
-- Dorsal clypeus at most with weak median keel or with flat elaboration ................................ 6
6 Anterior clypeal margin with upward flexion, somewhat lamellate and laterally denticulate
and broadly rounded, frequently with weak median carina (Fig. 55; plate I, A). (Top End,
Northern Territory) ..................................................................................................... snellingi
-- Anterior clypeal margin not lamellate, concave, with dorsal flat anteriorly rectangular or
excised elaboration that may surpass anterior clypeal margin................................................ 7
7 Larger species, HW 1.78 - 2.37. Dorsal clypeal elaboration seldom surpassing anterior
margin (Fig. 54). First gastral tergite entirely coarsely striate to striatopunctate. (Kimberley
throughout central Australia to northeastern Queensland) ......................................... unicolor
-- Smaller species with HW 1.58 - 1.80. Dorsal clypeal elaboration usually distinctly
surpassing anterior clypeal margin (Fg. 56). First gastral tergite microreticulate, without
striation. (Top End, Northern Territory)....................................................................berrimah
8 Posterior petiolar face evenly and conspicuously costate. Concolorous or bicolored species 9
-- Posterior petiolar face reticulate rugulose. Distinctly bicolored species ............................. 21
9 Clypeus bidentate or anteriorly deeply excised, with or without additional medial carina or
bulge ..................................................................................................................................... 10
-- Clypeus not bidentate, either transversally vaulted with the anterolateral clypeal corners
more or less acutely extended, or the clypeal projection flat with an occasional medial
elevation and the anterior margin sinuate (Figs. 65 - 68) ..................................................... 18
10 Clypeus dorsally with additional medial bulge or carina that may surpass the anterior
clypeal margin ...................................................................................................................... 11
-- Clypeus without such structure............................................................................................. 12
11 Clypeus medially with triangular bulge-like projection (Fig. 59). Dorsal ocular margin
confluent with ventral scrobal margin. (south-central to north-east coastal Australia) ....mars
-- Clypeus medially with acute tooth merging into a narrow carina (Fig. 61). Dorsal ocular
margin well separated from lower scrobal margin. (coastal and south-central Queensland;
Alice Springs) ..................................................................................................... tricuspidatus
12 Dorsal ocular margin well separated from lower scrobal margin......................................... 13

-- Dorsal ocular margin very close to or confluent with lower scrobal margin........................ 15
13 Promesonotal shield in overall impression not square, with apically acute posterolateral and
posterior projections. Petiole in profile with posterior face angularly convex. Frontal carinae
in anterior half abruptly broader (Fig. 71; plate I, B). (south-central Queensland; New South
Wales).......................................................................................................................christinae
-- Promesonotal shield in overall impression somewhat square with apically rounded posterior
and posterolateral projections. Petiole in profile acutely triangular. Frontral carinae
anteriorly not abruptly broader ............................................................................................. 14
14 Clypeus medially deeply excised, with long, acute anterolateral projections that almost
reach anterolateral frontal lobes (Fig. 69) (w. coastal to central Australia)................ diversus
-- Clypeus not as deeply excised, clypeal teeth short and bluntly triangular with upward
flexion, distinctly shorter than extended frontal lobes (Fig. 70). (Queensland; southern-most
Northern Territory) ................................................................................................. diversoides
15 Larger species with HW 1.65 - 1.80. Frontal carinae distinctly narrower than head width (FI
124 - 129). (central Australia) ................................................................................ deserticola
-- Smaller species with HW 1.15 - 1.50. Frontal carinae less narrower than head width (FI 115
- 122) .................................................................................................................................... 16
16 Propodeal spines long and massive (PSL 0.58 - 0.63) (northern South Australia) ..................
............................................................................................................................... crassispina
-- Propodeal spines shorter (PSL 0.45 - 0.56) .......................................................................... 17
17 Eyes relatively large (REL 0.21 - 0.24) with 15 - 18 ommatidia. Promesonotal shield
posteriorly not distinctly narrower (Fig. 21), with shorter pilosity. (Kimberley to Top End) ..
........................................................................................................................................ oxleyi
- Eyes relatively smaller (REL 0.18 - 0.20) with 14 - 15 ommatidia. Promesonotal shield
posteriorly distinctly narrower (Fig. 17), with longer pilosity. (north-eastern NSW, south-
central and costal Queensland) ..................................................................................... wilsoni
18 Clypeus a flat disc with an occasional medial elevation, anterior clypeal margin sinuate
(Fig. 68). Propodeal spines short (PSL 45 - 50). (Cape York) ..................................... discalis
-- Clypeus transversely vaulted with lateral clypeal corners extended (Figs. 65 - 67).
Propodeal spines longer (PSL 54 - 70) ................................................................................. 19
19 Eyes small (REL 0.15 - 0.17) with 14 - 17 ommatidia in the longest row. (south-east
Australia) ................................................................................................................. convexius
-- Eyes larger (REL 0.16 - 0.20) with 17 - 19 ommatidia in the longest row........................... 20
20 Promesonotal shield distinctly wider than long (PMI 130 - 138), more irregularly reticulate-
rugulose. Posterior medial projections digitate, with reduced or without lamellate material
inbetween (Fig. 31). (NSW, south-central Queensland)............................................. digitatus
-- Promesonotal shield not as distinctly wider than long (PMI 119 - 127), regularly reticulate.
Posterior medial projections with lamellate material between. (Northern Territory) naitsabes
21 Anterior clypeal margin broadly concave (Fig. 75). Mandibles with five teeth. (central and
southern Australia; Western Australia) ....................................................................mcarthuri
-- Anterior clypeal concavity much narrower. Mandibles with four teeth ............................... 22
22 Anterolateral clypeal corners acutely pointing forward (Fig. 51). Posterior and
posterolateral projections of promesonotal shield short and apically bluntly rounded,
translucently flanged (Fig. 76). With short scapes (SI1 47 - 53), markedly narrow frontal
carinae (FI 131 - 146) and relatively large eyes (REL 0.26 - 0.29). (central and south-
eastern Australia) ........................................................................................................... taurus
-- Anterolateral clypeal corners less acutely pointing forward. Posterior and postero-lateral
projections of promesonotal shield more strongly developed. Scapes longer (SI > 56),
frontal carinae broader (FI > 113 - 129) and eyes relatively smaller (REL 0.22 - 0.27) ...... 23
23 Petiole and postpetiole in lateral view narrow and high (PTI 55 - 61; PPI 48 - 58). south-
eastern Australia) ................................................................................................. angustinodis
-- Petiole and postpetiole in lateral view not as narrow and not as high (PTI > 64; PPI 53 - 72)
.............................................................................................................................................. 24
24 Petiole in lateral view dorsally truncated with distinct dorsal face (Fig. 44). (central to
south-east Australia) ..................................................................................................variabilis
-- Petiole in lateral view triangular without or with narrow oblique dorsal face (Fig. 46).
(Western and South Australia) ...................................................................................arcuatus
SPECIES OF M. DIVERSUS GROUP
Meranoplus ajax Forel, 1915

(Figs. 1, 2, 3, 52, 53, 78, 90)
Meranoplus mars r. ajax Forel, 1915: 44 (Kimberley district, Western Australia; worker). - Taylor &
Brown, 1985: 68 (catalogue), Taylor, 1987: 39 (listed) Taylor, 1990: 34 (Raised to species), Bolton,
1995: 250 (catalogue), Shattuck, 1999: 142 (listed). Holotype worker (NHRS; examined), 'Kimberley
district [printed] \ N. V. Austr. Mjöberg [printed] \ mars [printed] \ Meranoplus Mars For. r. Ajax
Forel.♀ typus unicus. [handwritten] \ 388 [printed] 88 [handwritten; both on red label] \ Riksmuseum
Stockholm [printed on light-blue label]'.
WORKERS (n = 43). TL 6.50 - 9.25, HL 1.65 - 2.40, HW 1.70 - 2.60, FC 1.38 - 1.95, CI
100 - 117, CS 1.68 - 2.50, SL 0.89 - 1.20, SI1 43 - 52, SI2 45 - 53, PML 1.07 - 1.58, PW 1.38 -
2.05, PMD 1.35 - 2.08, PMI2 91 - 104, ML 1.50 - 2.0, PSL 0.53 - 0.84, PTLL 0.50 - 0.70,
PTLH 0.63 - 0.90, PTI 72 - 82, PTDW 0.63 - 0.90, PPLL 0.43 - 0.58, PPLH 0.58 - 0.93, PPI 59
- 78, PPDW 0.68 - 0.95, PT/PP 80 - 100.
Mandible with three teeth. Clypeus in full face view longitudinally carinulate, with
conspicuous somewhat tricuspidate anteriorly projecting elaboration, consisting of strong medial
carina and lateral carinae, that merge into lateral angulate to rounded lobes, anterior projections
occasionally reduced. Head distinctly wider than long, lateral sides almost straight, feebly
narrowing anteriorly, preoccipital corners bulbously rounded, rear margin emarginate. Frontal
carinae markedly narrower than head (FI 124 - 143), sinuately and evenly narrowed from about
middle of length of head towards clypeus, anterolaterally extended into narrowly rounded
projections which together with clypeus distinctly overhang mandibular section. Antennal
scrobe in lateral view reaching middle of length of head. Distinctly transversely carinulate at
rear, occasionally with additional shagreening. Genae and ventrolateral sides of head carinate to
rugose, preoccipital lobes reticulate. Compound eyes relatively small (REL 0.15 - 0.19, EL 0.31
- 0.41, with 15 - 21 ommatidia in the longest row), situated distinctly in front of middle of lateral
sides of head, dorsal ocular margin never reaching ventral scrobal margin.
Promesonotum wider than long (PMI 120 - 140), concealing lateral sides of mesosoma only
anteriorly. Propodeal declivity and propodeal spines visible from above; lateral sides of
promesonotum occasionally translucently margined, anterolateral corners rectangular, at most
with stout anterolaterally projecting teeth; promesonotal suture absent or occasionally seen as
faint transverse line; at its level laterally with deep excision on each side of the shield;
posteriorly to excision with acute posteriorly directed lateral projection; posterolateral and
posterior mesonotal projections reduced to triangular stout teeth. Propodeal spines long and
slender, in dorsal view markedly diverging and slightly curved, emerging distinctly above
middle of propodeal length.
Petiole in profile broadly triangular, sometimes obliquely truncated, anterior face straight
and delicately rugulose, posterior face distinctly rugose. Dorsal postpetiole nodiform, tapering
towards base, occasionally wedge-shaped, with ventral medium sized tooth, rugose throughout.
First gastral tergite conspicuously striate to mictroreticulate. Dorsum of head longituinally

costulate, additionally with few transverse meshes, at rear reticulate, with microsculpture which
varies from nothing to a dense reticulum. Promesonotal shield irregularly to very regularly
rugose or rugoreticulate. Pilosity consisting of short and longer hairs reaching to 600 µm.
Concolorous brown to dark-brown, frequently with the gaster somewhat brighter.
MATERIAL EXAMINED
South Australia: 9 km / 9.4 km SE Maryinna Hill, 14.-18.iii.1995 (Hirst & Lands); 7.5 km
WNW, 8 km NW, 7.5 km NNW Mt. Kintore, 5.-10.v.1994 (coll. unknown); Musgrave Ranges,
5 km E Mitchell Knob, 20.-21.x.1994; 19.9 km W Indulkana, 25.-31.x.1998 (coll. unknown);
18.5 km WNW Ungarinna Rockhole, 14.-18.iii.1995 (coll. unknown). Queensland:
'Queensland', 21.vi.1902 (F.P. Dodd); ca. 100 km NW Mt Isa, Buckley R., 25.vii.1981 (B.B.
Lowery); Weipa, vii.1983 (J.D. Majer); 6 km NE Ebagoola, 5.viii.1983 (P.S. Ward);
Townsville, 5./6.xi.1914 (Wheeler), ibid., 19.xi.1901 & 23.iv.1902 (F.P. Dodd); Porcupine
Gorge NP, Hughenden, 13.v.1980 (B.B. Lowery); 30 km N Giru, 20.iii.1980 (B.B. Lowery); 10
km S Bowen, 24.v.1981 (B.B. Lowery); Torrens Ck., E Hughenden, 14.v.1980 (B.B. Lowery);
18 km W Paluma, 21.i.1995 (J. Bugeja); Cape York, Lockerbie, 10.vi.1969 (G.B. Monteith),
ibid., i.1958 (Darlington); 20 km W Mount Surprise, 12.viii.1975 (B.B. Lowery); 10 km E
Mareeba, 3.viii.1975 (B.B. Lowery); N of Mareeba, ii.1958 (Darlington); Coen, C. York,
vii.1932 (Darlington); Cooktown, 20.vi.1955 (J. Beauglehole). Northern Territory: Tanami
Desert, 20.v.1986 (P.J.M. Greenslade); Alice Springs, vi.1951 (Lowe), ibid., 28.vi.1951 (W.L.
Brown); 27.8 km NE Alice Springs, 13.vii.2003 (R. R. Snelling); Tennant Creek, 2.vii.1951
(W.L. Brown); Powell Ck, 1933 (C. Barrett); 40 km W Wave Hill, 14.ix.1981 (D. Davidson &
S. Morton); Daly R. (Nesselmann); Darwin, 11.ii.1945 (Malkin), ibid., 31.vii.1961 (B.B.
Lowery); Berrimah, 8.vii.1951 (W.L. Brown), ibid., 28-29.vii. 2003 (R.R. Snelling); Darwin, 30
km SE, Stuart Highway, 4.viii.1972 (W. L. Brown); Stuart Hwy., 132 mi S Darwin, 5.vii.1951
(W.L. Brown); 9 km NW Adelaide River, 15.viii.1983 (P.S. Ward); Kununurra Dam,
11.vii.1967 (G. Campbell); Katherine, 18.vii.1981 & 1.vii.1985 (B.B. Lowery); Katherine
Gorge, 21.x.1977 (P.J.M Greenslade); Katherine, 40 / 45 km NW, 7.iv.1978, 27 km SE,
8.iv.1978, 35 km NW, 11.iv.1978 (P.J.M. Greenslade); Marbullo SW Katherine, 25.x.1977
(P.J.M. Greenslade); Arnhem H'way, 15 km E Adelaide, 20.vii.1981 (B.B. Lowery); Mataranka,
30.viii.1985 (B.B. Lowery); Edith Falls, 6.vii.1985 (B.B. Lowery); Larrimah, 16.vii.1981 &
10.vii.1985 (B.B. Lowery); Victoria River, 1.vii.1985 (B.B. Lowery); Alligator Rivers area,
Kapalga, 6.ix.1983 (P.J.M Greenslade), ibid., 1933, (C. Barret), ibid., Thorack Res., 1980 (M.
Andrew); Doyles Ridge nr. Birdum, 24.vi.1936 (coll. unknown). Western Australia: Mount
Fanny, 28 km NE of Mt. Aloysius, 16.xi.1977 (J.E. Feehan); Meekatharra / Billiluna, Pool
Canning Stock Rte, iv.1930-viii.1931 (coll. unknown); 42 km S Derby, nr. Willare Rd. House,
9.viii.1966 (B.B. Lowery); Kununurra, 8.viii.1966(B.B. Lowery), ibid., 11.vii./1.viii.1967
(Campbell); Kimberley region nr Kalumburu mission, viii.1967 (coll. unknown); Surveyors Pool
Camp, Mitchell Plateau, 3.v.1992 (S.O. Shattuck); Derby (W.D. Dodd). (412 workers, 24 gynes
in ANIC, BMNH, JDMP, LACM, MCZC, NHMB, NHMW, PSWC, QMBA, SAMA, USNM,
WAMP).
DISCUSSION
M. ajax not only is one of the more widely distributed taxa, it is by far also one of the most
variable species within the diversus-group. However, certain trends within the variability are
displayed by different populations: Southern and central Australian samples show a distinct
gastral striation and long thin whitish pilosity, whereas the striation vanishes and is reduced to
almost entirely absent in northern populations, where the pilosity is shorter and stouter. In
addition, the shape of the clypeal prominence varies markedly as does the petiolar shape and the
outline of the promesonotal shield. Since transition between these populations seems to occur, I
here refrain from describing new taxa. Genetic information may help to elucidate whether there
are well defined subspecies or there is even more than one species hidden in this taxon.
However, despite the high degree of variability, M. ajax as here defined is easily separated from
others by the clypeal prominence. This is a widespread species covering most of the arid to
monsoonal areas of Australia, absent in the southernmost parts.
Meranoplus unicolor Forel, 1902

(Figs. 4, 5, 54, 79, 91)
Meranoplus diversus Sm. r. unicolor Forel, 1902: 455 (King Sound, Western Australia; worker) - Taylor &
Brown, 1985: 67 (catalogue), Taylor, 1987: 38 (listed), Taylor, 1990: 34 (raised to species), Bolton,
1995: 252 (catalogue), Shattuck, 1999: 143 (listed). Lectotype worker (MHNG, here designated),
'Typus [printed on red label] \ M. diversus ♀ Sm r. unicolor For Kings Sound N. W. Austr Frogatt
[handwritten] \ 8 \ r. M. unicolor Forel [handwritten] \ Coll. Forel'; 4 paralectotypes in ANIC, MHNG.
According to Taylor (1990) one further syntype in Mus Victoria (not examined).
WORKERS (n = 22). TL 6.15 - 8.40, HL 1.66 - 2.15, HW 1.78 - 2.37, FC 1.27 - 1.73, CS
1.72 - 2.26, SL 0.88 - 1.1, SI1 44 - 52, SI2 46 - 54, PML 1.0 - 1.4, PW 1.38 - 1.85, PMI 118 -
138, PMD 1.28 - 1.80, PMI2 89 - 100,ML 1.45 - 1.97, PSL 0.48 - 0.85, PTLL 0.46 - 0.63,
PTLH 0.60 - 0.83, PTDW 0.6 - 0.85, PPLL 0.4 - 0.64, PPLH 0.55 - 0.83, PPI 64 - 77, PPDW
0.55 - 0.88, PT/PP 93 - 109.
Mandibles with three teeth. In full face view clypeal projection distinctly exceeding
anterolateral frontal corners, with a median bicarinate flat elaboration, that may considerably
surpass the anterior clypeal margin. Frontal carinae sinuately narrowed towards clypeus,
distinctly narrower than head width (FI 134 - 148). Head weakly to distinctly wider than long
(CI 104 - 118), preoccipital margin distinctly concave. Antennal scrobe in lateral view
posteriorly surpassing middle of length of head, merging into lateral sides far before posterior
corners; distinctly transversely carinulate at rear, occasionally with additional shagreen. Genae
and ventrolateral sides of head carinate to rugose, preoccipital lobes reticulate. Eyes of moderate
size (EL 0.30 - 0.36, REL 0.16 - 0.20, with 16 - 19 ommatidia in the longest row), in lateral
view situated in anterior half of lateral sides of head, not reaching ventral scrobal margin. Dorsal
surface of head with additional microreticulum between ridges of rugo-reticulation.
Promesonotal shield only very narrowly translucently margined. Mesonotum posteriorly
markedly narrowing with the posterolateral projections reduced and bluntly rounded,
occasionally as more or less well developed acute denticles. Propodeal declivity visible from
above. Propodeal spines due to variable body size varying in length (0.48 - 0.85), situated above
middle of length of declivity, acute and considerably diverging when seen from above.
Petiole in lateral view higher than long (PTI 70 - 86), box-shaped to roughly triangular with
anterior and posterior faces meeting in an angle. Postpetiole nodiform with short to medium-
sized antero-basal tooth.
First gastral tergite entirely and distinctly striate to punctate-striate, with lateral striation on
corresponding ventrite as well. Surface covered with moderately dense regular pilosity
consisting of shorter decumbent hairs and longer erect ones, pilosity on dorsal head shorter than
that on mesosoma and gaster.
Concolorous brown to dark-brown.
MATERIAL EXAMINED
South Australia: 6 km SE Box Creek, 24.ix./ 24.x.1993 (McArthur & Adams); 19.5 km
WNW Anta Hill, 16.-19.ix.1998 (coll. unknown); 20 km NE Macumba Station, 6.x.1981 (D.
Davidson & S. Morton). Western Australia: Kununurra, 8.viii.1986 (B.B. Lowery); Broome,
11.viii.1986 (B.B. Lowery); 24 mi SE Broome, 17.iv.1963 (McInnes & Dowse); King's Sound,
'Forel No 8'. Northern Territory: 25 km W Tempe Downs, 20.v.1997 (J. & J. Schapel);
Larrimah, 16.vii.1981 & 28.vi.1985 (B.B. Lowery); Tanami Desert, 18.v.1986 (P.J.M.
Greenslade); Alligator Rivers area, Jabiru, 10.v.1983 (P.J.M. Greenslade); 60 km E Three
Ways, 24.vii.1981(B.B. Lowery); Manbulloo SW Katherine, 25.x.1977 (P.J.M. Greenslade);
Elliot, Stuart H'way, 16.vii.1981 (B.B. Lowery); 40 km N Wave Hill, 14.ix.1981 (D. Davidson &
S. Morton); Davenport Ra. Rd. at Bonney Creek, 6.vii.2003 (R.R. Snelling); Kidman Springs,
23-30.iv.1997 (A. Salvarani). Queensland: Camooweal, 25.vii.1981 (B.B. Lowery); 19 km N
Ebagoola, 12.viii.1983 (P.S. Ward); Moreton Teleg. Stn., 11.viii.1958 (E.M. Exley); Cooktown
(Staudinger); 10 km E Mt.Isa Gorge Ck., 12.v.1980 (B.B. Lowery); SW Sandringham, 4.vi.1980
(P.J.M. Greenslade); Alice River (Mjöberg); Gubberamunda, 7.-9.v.2002 (S.G. Wright);
Gumbardo, iv.2001 (T. Beutel). (104 workers, 3 gynes in ANIC, LACM, MCZC, NHMW,
QMBA, SAMA, UQIC).
DISCUSSION
M. unicolor is a variable taxon, in particular with regard to the clypeal projection and the
petiolar shape. Nonetheless, the clypeal structure in addition to the distinct gastral striation
should leave no doubt as to the identity of the taxon. Additional material is needed to show
whether more than a single taxon is involved. Occuring from Kimberley over central Australia
to the north-east coast of Queensland; missing in southern Australia.
Meranoplus berrimah Schödl sp.n.

(Figs. 8, 9, 56, 89)
HOLOTYPE WORKER. TL 5.15, HL 1.50, HW 1.58, FC 1.25, CS 1.54, SL 0.75, SI1 48,
SI2 49, PML 1.02, PW 1.25, PMD 1.30, PMI2 104, ML 1.25, PTLL 0.38, PTLH 0.53, PTDW
0.55, PPLL 0.35, PPLH 0.50, PPI 70, PPDW 0.54, PT/PP 102.33.
Mandible with three teeth. In full face view the clypeal elaboration distinctly exceeding
anterolateral frontal corners, with a dorsal median bicarinate, anteriorly concave flat projection
and lateral even longer arcuate acute denticles, the latter situated ventrally to the median
projection. Head only moderately wider than long (CI 105), almost square, preoccipital margin
distinctly concave. Frontal carinae sinuately narrowed towards clypeus, not as narrow as in
former species (FI 126). Antennal scrobe in lateral view posteriorly surpassing middle of length
of head, distinctly transversely carinulate at rear, without additional shagreening. Genae and
ventrolateral sides of head carinate to rugose, preoccipital lobes reticulate. Eyes small (EL 0.26,
REL 0.17, with 15 ommatidia in the longest row), in lateral view situated in anterior half of
lateral sides of head, dorsal ocular margin never reaching ventral scrobal margin. Surface of
head between ridges of rugo-reticulation with microsculpture.
Promesonotal shield wider than long (PMI 123), only narrowly translucently margined.
Mesonotum posteriorly markedly converging, with the posterolateral projections acute and
stout, the posterior medial projections bluntly rounded. Propodeal declivity partly visible from
above. Propodeal spines rather short (0.44) situated above middle of length of declivity, when
seen from above acute, straight and considerably divering. Petiole in lateral view higher than
long (PTI 71), somewhat tectiform, anterior and posterior faces meeting in an acute angle with
anterior petiolar face angulate, posterior face convex.
Postpetiole elongately nodiform to drop-shaped with a medium-sized anterio-basal tooth.

Gaster entirely microreticulate, with faint basal striation. Surface covered with moderately dense
regular pilosity consisting of shorter decumbent hairs and longer erect ones, in general pilosity
on dorsal head shorter than on mesosoma and gaster.
Concolorous brown to dark-brown.
WORKERS (n = 7). TL 5.15 - 6.35, HL 1.48 - 1.70, HW 1.58 - 1.80, FC 1.25 - 1.35, FI
126 - 133, CI 103 - 110, CS 1.53 - 1.74, SL 0.75 - 0.88, SI1 47 - 50, SI2 49 - 52, PML 1.02 -
1.15, PW 1.25 - 1.5, PMI 119 - 136, PMD 1.3 - 1.5, PMI2 98 - 106, ML 1.25 - 1.50, PSL 0.43 -
0.50, PTLL 0.38 - 0.45, PTLH 0.53 - 0.64, PTI 67 - 74, PTDW 0.55 - 0.65, PPLL 0.35 - 0.43,
PPLH 0.50 - 0.61, PPI 61 - 77, PPDW 0.53 - 0.64, PT/PP 100 - 110, EL 0.25 - 0.29, REL 0.16 -
0.18, with 15 - 17 ommatidia in the longest row.
ETYMOLOGY
The name refers to the type locality.
TYPE MATERIAL
Holotype worker, Northern Territory: 'AUSTRALIA. N.Terr.: CSIRO-TERC. Berrimah

12.42°S 130.92°E 17 Dec. 2000 #00-163 \ #00-163. Secondary tropical savannah. Ex nest in
soil. coll. R. R. Snelling' (ANIC). Paratypes. 3 workers, same data as holotype; 37 workers,
same locality but '14-16 Dec. 2000 #00-163' and 'Foragers in litter', '28-29 June 2003', '21 June
2003 #03-157', '21 June 2003 #03-158' and '22 June 2003 #03-163' (LACM, NHMW, ANIC).
Northern Territory: Groote Eylandt, vii.1983 (J.D. Majer); Nourlangie Rock, 17.viii.1983
(P.S.Ward); Mudginberri, 18.vii.1976 (R. Mercer); Koolpinyah, 1933 (C. Barrett); Larrimah,
16.vii.1981 (B.B. Lowery); Mataranka Homestead, 22.viii.1981 (B.B. Lowery); 20 km W
Katherine, 1.vii.1985 (B.B. Lowery); 23 km SW Katherine, 24.x.1977, 27 km SW Katherine,
25.x.1977, 30 km SW Katherine, 10.iv.1978 (P.J.M. Greenslade); Berrimah, Darwin distr.,
8.vii.1951 (W.L. Brown). (43 workers in ANIC, JDMP, MCZC, NHMW, PSWC, USNM).
DISCUSSION
M. berrimah is a quite uniform taxon within its limited range, but shows variation in the
clypeal elaboration to some extent, concerning both size and occasional additional lateral
carinae. However, its structural distinctness in combination with the relatively small size (HW
1.60 - 1.80), the non-striate gaster and the petiolar shape separate M. berrimah easily from other
related group members. Small M. unicolor workers, that might show a similar clypeal structure
are readily separated by their striate gaster. Hitherto known from Top End, Northern Territory.
Meranoplus snellingi Schödl sp.n.

(Figs. 6, 7, 55, 87; Plate 1, A)
0.78, PPLL 0.58, PPLH 0.75, PPI 77, PPDW 0.85, PT/PP 91.
Mandible with three teeth. Clypeal projection distinctly exceeding anterolateral frontal
corners, with weak median carina and additional lateral carinulae, the anterior clypeal margin
concave with upward flexion, the lateral corners rounded, denticulate. Head wider than long (CI
112), preoccipital margin deeply concave. Frontal carinae distinctly sinuately narrowing towards
clypeus, markedly narrower than width of head (FI 136). Antennal scrobe in lateral view
scarcely surpassing middle of length of head, merging into lateral sides of head distinctly
anteriorly to rear corners; distinctly transversely carinulate at rear with additional shagreening.
Genae and ventrolateral sides of head carinate to rugose, preoccipital lobes reticulate. Eyes
relatively small (EL 0.35, REL 0.16, with 18 ommatidia in the longest row), situated well in
front of middle of lateral sides of head, the dorsal ocular margin far from the ventral scrobal
margin. Surface of head between ridges of rugo-reticulation with microreticulum.
Promesonotal shield distinctly wider than long (PMI 130), only very narrowly translucently
margined. Mesonotum with posterolateral projections triangular, the posterior medial
projections rounded. Propodeal declivity visible from above. Propodeal spines of medium length
(PSL 0.73) situated above middle of length of declivity, acute and slightly arcuate when seen
from above.
Petiole in lateral view box shaped, distinctly higher than long (PTI 75), with the anterior
face straight, meeting oblique dorsum in a more or less right angle, the latter merging roundly
into posterior face. Postpetiole nodiform with small anterio-basal tooth.
Gaster entirely and roughly striate (elongately carinulate, with microsculpture between
carinulae). Dorsal surface covered with scattered regular pilosity consisting of short and longer
more or less erect stiff hairs, in general pilosity on head dorsum shorter than on mesosoma
and gaster.
WORKERS (n = 8). TL 7.20 - 8.75, HL 1.88 - 2.19, HW 2.0 - 2.43, FC 1.48 - 1.75, FI 134
- 142, CI 103 - 112, CS 1.94 - 2.31, SL 1.0 -1.15, SI1 44 - 51, SI2 46 - 53, PML 1.23 - 1.45, PW
1.50 - 1.83, PMI 120 - 130, PMD 1.45 - 1.83, PMI2 94 - 103, ML 1.70 - 2.0, PSL 0.65 - 0.73,
PTLL 0.58 - 0.68, PTLH 0.73 - 0.84, PTI 74 - 81, PTDW 0.66 - 0.90, PPLL 0.45 - 0.60, PPLH
0.63 - 0.83, PPI 70 - 77, PPDW 0.71 - 0.95, PT/PP 91 - 102, EL 30 - 0.37, REL 0.15 - 0.19,
with 16 - 18 ommatidia in the longest row.
ETYMOLOGY
Named for Roy R. Snelling, myrmecologist at the Natural History Museum of Los Angeles
County. He provided me with numerous important specimens for this study.
TYPE MATERIAL
Holotype worker, Northern Territory: 'AUSTRALIA. N.Terr.: CSIRO-TERC. Berrimah

12.42°S 130.92°E 14-16 Dec. 2000 #00-145 \ #00-145. Secondary tropical savannah. Ex nest in
soil coll. R. R. Snelling' (ANIC). Paratypes. 23 workers, same data as holotype; 72 workers
with same locality data but '21 Dec 2000, #00-190', '19 June 2003 #03-143', '22 June 2003 #03-
162', '28-29 June 2003' (LACM, NHMW, ANIC).
Northern Territory: Maningrida, xii.1975 - i.1976 (J. Grigg); Nourlangie Rock,

17.viii.1983 (P.S. Ward); Bathurst Island (G.F. Hill); Howard Springs near Darwin, 19.vii.1981
(B.B. Lowery); Darwin, 13-17 km E, viii.1972 (W.L. Brown); Alligator Rivers area, Kapalga,
8.ix.1983 (P.J.M. Greenslade); Alligator Rivers area, Jabiru, 11.v./10.ix.1983 (P.J.M.
Greenslade); Katherine, 4.-5.vii.1951 (W.L. Brown); 21/22 km SE Katherine, 8.iv.1977 (P.J.M.

Greenslade); Finniss Range, 11.xi.1952 (Bateman); Arnhem Highway, 15 km E Adelaide (B.B.
Lowery). (47 workers, 1 gyne in ANIC, MCZC, NHMW, PSWC, USNM).
DISCUSSION
M. snellingi is a medium sized to large brown to blackish-brown species which apart from
differences in size shows only very little variation within its range. It is obviously most closely
related to the two preceding taxa but easily separable by the unique clypeal structure. Restricted
to Top End, Northern Territory.
Meranoplus convexius Schödl sp.n.

(Figs. 26, 27, 65, 84)
Mandible with three teeth. In full face view the clypeus anteriorly a strongly vaulted,
rugulose to carinulate projection, which markedly exceeds the anterolateral frontal corners; with
its anterior margin concave and the anterolateral corners acutely directed antero-ventrad. Head
only moderately wider than long (CI 107), preoccipital margin shallowly though markedly
concave. Frontal carinae sinuately narrowed towards clypeus (FI 124). Antennal scrobe
distinctly surpassing middle of lateral sides of head, anteriorly glossy, distinctly transversely
carinulate at rear, posteriorly well demarcated from remainder of head. Genae and ventrolateral
sides of head rugose, preoccipital lobes reticulate. Eyes small (EL 0.24, REL 0.16, with 15
ommatidia in the longest row), situated in front of middle of lateral sides of head, dorsal ocular
margin not reaching ventral scrobal margin. Head in posterior half distinctly reticulate,
inbetween ridges of rugo-reticulation shiny.
Promesonotal shield wider than long (PMI 122), rather flat, distinctly translucently
margined, provided with well developed projections concealing lateral sides of mesosoma and
propodeal declivity, invisible from above. Propodeal spines long in relation to body size (PSL
0.61) situated above middle of length of declivity, robust and acute, moderately diverging and
slightly arcuate when seen from above.
Petiole in lateral view higher than long (PTI 77), with anterior face straight, meeting
strongly convex posterior face in an acute angle. Postpetiole nodiform with a small antero-basal
tooth.
Gaster entirely microreticulate, basally with additional carinulae. Surface covered with
evenly distributed pilosity consisting of shorter decumbent and longer more or less erect arcuate
stiff hairs.
Distinctly bicolored with the gaster and appendages brown and remainder of body dark-
brown.
WORKERS (n = 10). TL 5.78 5.35 - 6.35, HL 1.38 - 1.60, HW 1.48 - 1.83, FC 1.20 - 1.38,
FI 123 - 135, CI 106 - 115, CS 1.43 - 1.71, SL 0.75 - 0.84, SI1 44 - 52, SI2 47 - 54, PML 1.05 -
1.25, PW 1.30 - 1.53, PMI 120 - 129, PMD 1.40 - 1.63, PMI2 107 - 111, ML 1.25 - 1.43, PSL
0.58 - 0.68, PTLL 0.40 - 0.48, PTLH 0.55 - 0.64, PTI 65 - 77, PTDW 0.53 - 0.63, PPLL 0.40 -
0.48, PPLH 0.55 - 0.64, PPI 69 - 77, PPDW 0.51 - 0.63, PT/PP 95 - 104, EL 0.22 - 0.26, REL
0.15 - 0.17, with 14 - 17 ommatidia in the longest row.
ETYMOLOGY
The name refers to the strongly vaulted clypeus.
TYPE MATERIAL
Holotype worker, Queensland: 'SEQ:25°34'Sx151°42'E Wetheron, 3 km SW, O/F 27 Jan -

2 Jun 1999 Monteith & Thompson 150m. pitfall 7718' (QMBA). Paratypes. 2 workers, same
data as holotype (QMBA, NHMW).
Queensland: Mount Coot-tha, Brisbane, 16.xii.1956 (B.B. Lowery). New South Wales: 9
km W Rankin Springs, 9.i.1967 (coll. unknown); 25 km N Dubbo, Eumungerie SF, 9.ix.1979
(B.B. Lowery); Urana golf links, 12.vi.1979 (B.B. Lowery); Tallimba, 21.vii.1979 (B.B. Lowery);
4 mi E Inverell, 22.viii.1968 (B.B. Lowery); Trundle, 30.xii.1963 (B.B. Lowery). (57 workers in
ANIC, NHMW, QMBA).
DISCUSSION
M. convexius together with the two following taxa forms a cluster within the diversus-group.
They all show a similar clypeal structure but may be separated by the different shape and surface
of the promesonotal shield, in addition with the size of the eyes. Distributed from the Murray-
Darling Basin to SE Queensland.
Meranoplus naitsabes Schödl sp.n.

(Figs. 28, 29, 66, 84)
SI2 47, PML 1.02, PW 1.30, PMD 1.38, PMI2 106, ML 1.30, PSL 0.58, PTLL 0.40, PTLH
0.58, PTDW 0.52, PPLL 0.40, PPLH 0.54, PPI 74, PPDW 0.52, PT/PP 100.
Mandible with three teeth. Clypeal projection very similar to that of preceding species, with
lateral teeth less produced. Head moderately wider than long (CI 108), preoccipital margin
shallowly though still markedly concave. Frontal carinae more evenly and less sinuately
narrowed towards clypeus (FI 129). Antennal scrobe not surpassing middle of lateral sides of
head as far as in convexius, anteriorly glossy, distinctly transversely carinulate at rear,
posteriorly indistinctly demarcated from remainder of head. Genae and ventrolateral sides of
head rugose, preoccipital lobes reticulate. Eyes relatively larger (EL 0.29, REL 0.19, with 17
ommatidia in the longest row), situated in front of middle of lateral sides of head, dorsal ocular
margin not reaching ventral scrobal margin. Head in posterior half distinctly reticulate, between
ridges of rugo-reticulation with faint microsculpture.
Promesonotal shield wider than long (PMI 127) with regular reticulation, lateral sides only
very narrowly translucently margined, projections less developed as in preceding species.
Propodeal declivity overhung by posterior mesonotal margin, only partly visible from above.
Propodeal spines long in relation to body size (PSL 0.58) situated above middle of length of
declivity, acute and moderately diverging when seen from above.
Petiole in lateral view more or less triangular, higher than long (PTI 70), with anterior face
straight, meeting convex to sinuate posterior face in a crest. Postpetiole nodiform with a small
anterio-basal tooth.
First gastral tergite entirely microreticulate, basally with additional carinulae. Pilosity
similar to that of preceding species.
Most workers are slightly bicolored with the gaster and appendages usually ferrugineous
and the remainder of body brown.
WORKER (n = 6). TL 5.05 - 5.70, HL 1.35 - 1.53, HW 1.48 - 1.65, FC 1.13 - 1.28, FI 125
- 131, CI 107 - 110, CS 1.41 - 1.6, SL 0.74 - 0.78, SI1 45 - 50, SI2 47 - 52, PML 1.0 - 1.08, PW
1.20 - 1.33, PMI 119 - 127, PMD 1.29 - 1.45, PMI2 106 - 109, ML 1.20 - 1.35, PSL 0.54 - 0.63,
PTLL 0.35 - 0.40, PTLH 0.51 - 0.65, PTI 62 - 71, PTDW 0.48 - 0.55, PPLL 0.33 - 0.40, PPLH
0.50 - 0.56, PPI 65 - 76, PPDW 0.48 - 0.55, PT/PP 100 - 105, EL 0.25 - 0.29, REL 0.18 - 0.19,
ETYMOLOGY
An anagram based on the name of my son Sebastian, who occasionally joins me on

entomological trips and does some real good collecting—in particular crickets and flies.
TYPE MATERIAL
Holotype worker, Northern Territory: 'NT Barrow Creek 5 km NE 21.29S 133.35E

10/10/81 135b D. Davidson / S. Morton' (ANIC). Paratypes. 5 workers, same data as holotype;
12 workers with same locality data but '135a' and '135c' (ANIC, NHMW).
Northern Territory: Tanami Desert, 20.v.1986 (P.J.M. Greenslade); 40 km W Wave Hill,

14.ix.1981 (D. Davidson / S. Morton). (25 workers in ANIC, NHMW).
DISCUSSION
This species is known only from arid portions of Northern Territory.
Meranoplus digitatus Schödl sp.n.

(Figs. 30, 31, 67, 85)
0.63, PTI 72, PTDW 0.61, PPLL 0.43, PPLH 0.65, PPI 66, PPDW 0.61, PT/PP 100.
Mandible with three teeth. Clypeal projection similar to that of preceding species. Frontal
carinae broad, distinctly sinuately narrowed towards clypeus (FI 123). Head wider than long (CI
110), preoccipital margin similar to that of preceding species. Antennal scrobe scarcely
surpassing middle of lateral sides of head and posterior ocular margin, anteriorly glossy,
transversely carinulate at rear, posteriorly indistinctly demarcated from remainder of head.
Genae and ventrolateral sides of head rugose, preoccipital lobes reticulate-foveolate. Eyes
relatively larger (EL 0.31, REL 0.18, with 19 ommatidia in the longest row), situated anteriorly
to middle of lateral sides of head, dorsal ocular margin not reaching ventral scrobal margin.
Head in posterior half distinctly reticulate to reticulate-foveolate, in between ridges of rugo-
reticulation with very faint microsculpture.
Promesonotal shield vaulted, distinctly wider than long (PMI 130), only pronotal lateral
sides translucently margined, mesonotal posterolateral and posterior projections digitate,
dorsally irregularly rugoreticulate, with the meshes elongate and longitudinally orientated.
Propodeal declivity partly overhung by posterior mesonotal margin. Propodeal spines relatively
long (PSL 0.70) situated above middle of length of declivity, straight, acute and moderately
diverging when seen from above.
Petiole in lateral view higher than long (PTI 72), with anterior face straight, meeting convex
posterior face in a crest. Postpetiole nodiform with a small anterio-basal tooth.
Gaster entirely microreticulate, with occasional glossy spots inbetween. Surface covered
with evenly distributed pilosity consisting of shorter decumbent and longer more or less erect
arcuate stiff hairs.
128 - 137, CI 107 - 112, CS 1.48 - 1.79, SL 0.71 - 0.80, SI1 41 - 47, SI2 43 - 48, PML 0.95 -
ETYMOLOGY
The name refers to the digitate projectons on the mesonotal hind margin.
TYPE MATERIAL
Holotype worker, New South Wales: 'N.S.W. Goolgowi 18.vii.1979 BBLowery Red soil \
ANIC ANTS VIAL 68.123' (ANIC). Paratypes. 8 workers, same data as holotype (ANIC,
NHMW).
New South Wales: Rata, 16.xii.1965 (B.B. Lowery); CSIRO Lake Mere field stn., nr.
Louth, i.1995 (M. Bryannah); Goolgowi, 18.vii.1979 (B.B. Lowery); Condobolin, 11.i.1967
(B.B. Lowery); 5 km W Merriwagga, 17.iv.1978 (B.B. Lowery); Bogan R. (J. Armstrong).
Queensland: Merigol, various sites and pitfalls, iv.2001 (T. Beutel). (113 workers, 7 gynes in
ANIC, NHMW, QMBA).
DISCUSSION
Distinct from both preceding taxa by the more or less digitate posterior mesonotal
projections which only seldomly are fused by lamellate material. Rather uniformly ferrugineous
with gaster and appendages usually only feebly brighter than remainder of body. Known from
the Murray-Darling basin.
Meranoplus discalis Schödl sp.n.

(Figs. 32, 33, 68, 86)
Mandible with four blunt teeth. In full face view the clypeal projection a truncated flat to
feebly vaulted shelve, not exceeding anterolateral frontal corners, with a sinuate anterior margin.
Head only moderately wider than long (CI 108), preoccipital margin shallowly concave. Frontal
carinae, distinctly sinuately narrowed towards clypeus (FI 124). Antennal scrobe in lateral view
posteriorly surpassing middle of length of head, ending far before posterior corners, transversely
carinulate at rear, posteriorly well defined from preoccipital section. Genae and ventrolateral
sides of head carinate, preoccipital lobes reticulate to foveo-reticulate. Eyes of moderate size
(EL 0.25, REL 0.19, with 14 ommatidia in the longest row), situated in front of middle of lateral
sides of head. Head in posterior half distinctly reticulate, in between ridges of rugo-reticulation
smooth and glossy.
Promesonotal shield wider than long (PMI 121), moderately vaulted; its surface coarsely
and irregularly rugose to rugoreticulate, the meshes somewhat longitudinally orientated; both
lateral sides and rear broadly translucently margined, posterolateral projections well developed,
arcuately directed outwards, posterior projections short and triangular. Propodeal declivity
overhung by mesonotal hind margin. Propodeal spines moderate in length related to body size
(PSL 0.50) situated above middle of length of declivity, massive and straight and moderately
diverging in dorsal view.
Petiole in lateral view distinctly higher than long (PTI 65), narrowly triangular with anterior
face straight, meeting posterior face in an acute angle. Postpetiole elongately nodiform with a
small anterio-basal tooth.
Gaster entirely microreticulate with glossy spots around hair pits. Surface covered with
evenly distributed pilosity exsiting of shorter decumbent and longer more or less erect arcuate
stiff hairs.
Distinctly bicolored with gaster and appendages ferrugineous and remainder of body
piceous.
PARATYPE WORKERS (n = 2). TL 4.95, - 5.25, HL 1.28 - 1.33, HW 1.30 - 1.40, FC

1.08 - 1.13, FI 121 - 124, CI 102 - 106, CS 1.29 - 1.36, SL 0.68 - 0.72, SI1 51 - 52, SI2 52 - 53,
PML 0.95 - 1.05, PW 1.18 - 1.28, PMI 121 - 124, PMD 1.35 - 1.48, PMI2 115 - 116, ML 1.15 -
1.20, PSL 0.45 - 0.48, PTLL 0.35 - 0.38, PTLH 0.54 - 0.58, PTI 65, PTDW 0.46 - 0.49, PPLL
0.33 - 0.35, PPLH 0.53 - 0.55, PPI 62 - 64, PPDW 0.48 - 0.50, PT/PP 97 - 98, EL 0.23 - 0.24,
REL 0.18, with 11 - 13 ommatidia in the longest row.
ETYMOLOGY
The name refers to the flat clypeal projection.
TYPE MATERIAL
Holotype worker, Queensland: 8-18 mi N.W. of \ Silver Plains Cape York, Q. May-June
'58 Darlingtons\gum forest (MCZC). Paratypes. 2 workers, same data as holotype (MCZC,
NHMW).
DISCUSSION
The paratypes resemble the holotype in all features. The characteristic clypeal structure
separates M. discalis from all other related forms. Only known from the type locality, Cape
York, Queensland.
Meranoplus diversus Smith, 1867

(Figs. 34, 35, 69, 83, 95)
Meranoplus diversus F. Smith, 1867: 527, pl.26, fig.2 (Champion Bay [= Geraldton], Western Australia;
worker), Forel, 1915: 44 (male). Taylor & Brown, 1985: 67 (catalogue), Taylor, 1987: 38 (listed),
Bolton, 1995: 251 (catalogue), Shattuck, 1999: 142 (listed). Holotype worker (BML, examined),
'Smith coll. pres. by Mrs. Farren White. 99-303. [printed] \ Meranoplus diversus. Sm. Trans. Ent. Soc.
[handwriten on bright violet label] \ Brit. Mus [handwritten on bright violet label] \ Holotype [round
printed red circled label]'.
1.63 - 1.87, SL 0.8 - 0.9, SI1 44 - 46, SI2 46 - 50, PML 1.05 - 1.27, PW 1.37 - 1.55, PMI 118 -
133, PMD 1.50 - 1.77, PMI2 107 - 120, ML 1.30 - 1.75, PTLL 0.40 - 0.50, PTLH 0.60 - 0.75,
PTDW 0.48 - 0.64, PPLL 0.38 - 0.50, PPLH 0.48 - 0.78, PPI 63 - 79, PPDW 0.48 - 0.60, PT/PP
100 - 109.
Mandible with three teeth. Clypeus medially deeply excavated, distinctly acutely bidentate,
longitudinally carinulate, sunk into prolonged anterolateral frontal projections. Head distinctly
wider than long (CI 112 - 120), lateral sides almost straight, feebly narrowing anteriorly,
preoccipital corners bulbously rounded, the rear margin emarginate. Frontal carinae evenly
sinuately narrowed towards clypeus, distinctly narrower than head width (FI 132 - 145).
Antennal scrobes in lateral view surpassing middle of length of head posteriorly, distinctly
transversely carinulate in posterior half, occasionally with additional shagreening, posteriorly
rather distinctly demarcated from remainder of head. Genae and ventrolateral sides of head
carinate to rugose, preoccipital lobes reticulate. Compound eyes moderately large (EL 0.25 -
0.32, REL 0.17 - 0.20, with 15 - 20 ommatidia in the longest row) situated distinctly in front of
middle of lateral sides of head, dorsal ocular margin not reaching ventral scrobal margin.
Promesonotum wider than long (PMI 118 - 133), somewhat flanged and broadly
translucently margined, concealing lateral sides of mesosoma and propodeal declivity.
Propodeal spines of medium length (PSL 0.58 - 0.78) situated above middle of length of
declivity, acute and slightly arcuate when seen from above.
Petiole distinctly higher than long (PTI 66 - 73), in profile wedge-shaped with anterior face
straight and unsculptured, posterior face convex, distinctly and regularly costate. Postpetiole
elongately nodiform distinctly tapering towards base, with ventral medium sized tooth, rugose
throughout.
First gastral tergite with dense microreticulum, basal half with additional distinct striation.
Dorsum of head longitudinally costulate, additionally with rugulae and only few transverse
meshes, at very rear reticulate, interspaces with microsculpture; with scattered, suberect to erect
hairs. Promesonotal shield coarsely rugoreticulate, with pilosity consisting of short decumbent
and longer more or less erect hairs.
Concolorous brown to fuscous, frequently with the gaster somewhat brighter.
MATERIAL EXAMINED
Western Australia: Winburn Rocks, 95 km E by N of Warburton, 16.xi.1977 (J.E.

Feehan). South Australia: 26.1 km / 26.3 km ENE Mimili, 25.-31.x.1998 (coll. unknown); 1.9
km WNW Mt. Lindsay, 16.-20.x.1996 (coll. unknown); 18.5 km WNW Ungarinna Rockhole
(coll. unknown); 14.2 km ESE Maryinna Hills, 14.-18.iii.1995 (coll. unknown); Womikata Bore,
21.x.1994 (coll. unknown); 0.5 km WSW Cheesman Peak, 25.x.1996 (coll. unknown) (38
workers, 2 gynes in ANIC, NHMW, SAMA).
DISCUSSION
Distributed from the west coast to central Australia. Although a wide gap is present between
the type locality (Geraldton) and most material studied herein, the species should be found
throughout the arid and semi-arid zones of central and Western Australia.
M. diversus is unique by the bifurcate clypeus in combination with the flanged

promesonotal shield and the costate triangular petiole.
Meranoplus diversoides Schödl sp.n.

(Figs. 36, 37, 70, 83)
Mandible with three teeth. Clypeus in full face view not exceeding anterolateral frontal
corners, medially bidentate, medial dentate section with slight upward flexion, sunk into
prolonged lateral frontal projections, with additional wrinkles or carinulae. Head distinctly wider
than long (CI 115), preoccipital lobes bulbously rounded, rear margin of head concave. Frontal
carinae distinctly sinuate and translucently margined anteriorly, distinctly narrower than head
width (FI 141), anterolaterally extended into narrowly rounded projections exceeding clypeal
fork. Antennal scrobe in lateral view surpassing middle of length of head, irregularly
transversely carinulate in posterior half, with additional shagreening, ill defined near posterior
section of head. Genae and ventrolateral sides of head rugose, posterior lateral corners of head
reticulate. Compound eyes relatively small (EL 0.29, REL 0.17, with 17 ommatidia in the
longest row), situated distinctly in front of middle of lateral sides of head, dorsal ocular margin
not reaching ventral margin of antennal scrobe.
Promesonotum markedly wider than long (PMI 130), concealing lateral sides of mesosoma
and propodeal declivity, translucently margined to a greater extent. Propodeal declivity above
middle of its length with two long slender, in dorsal view feebly diverging slightly arcuated
lateral spines.
Petiole higher than long (PTI 72) in profile broadly triangular, with anterior face straight
and unsculptured, posterior face convex, distinctly and regularly costate. Postpetiole in profile
dorsally nodiform, distinctly tapering towards base, with ventral medium sized tooth, rugose
throughout.
First gastral tergite entirely microreticulate, basally with fine longitudinal costulae. Dorsum
of head longitudinally costulate with additional rugulae and only few transverse meshes, behind
level of eyes reticulate, interspaces with faint microsculpture. Promesonotal shield coarsely
rugoreticulate. All dorsal surfaces with pilosity of decumbent curved hairs and longer more or
less erect ones.
Color dark brown to brown, with the gaster and appendages brighter.
128 - 142, CI 111 - 120, CS 1.51 - 1.90, SL 0.79 - 0.93, SI1 45 - 49, SI2 49 - 52, PML 1.07 -
ETYMOLOGY
The name refers to the close relationship to M. diversus.
TYPE MATERIAL
Holotype worker, Queensland: 'QLD Tambo airport 24 Oct 1979 RH Mew 1 [handwritten]
\ Royce H. Mew collection [printed] \ AUSTRALIA Qld. Tambo 24°53'S 146°15'E 24. Okt 79;
R.H.Mew [printed; subsequetly added locality label]' (SAMA). Paratypes. 10 workers, same
data as holotype (ANIC, SAMA, NHMW).
Queensland: Collinsville, 3 km N Johnson-Johnson Farm, 28.ii.2000 (Buschinger);

Toowoomba, 4.v.1951 (F.A. Perkins), ibid., picnic point & Prince Henry Dr., 11./13.iv.1962,
13.xii.1973 (B.B. Lowery); Gatton, 12.ii.51 (S.G. Grimmett); 8.5 km SW Mt. Hutton, 6.iii.2002
(S.G. Wright); Gumbardo, mulga, iv. 2001 (T. Beutel); Merigol, iv.2001, mulga (T. Beutel); St
George's, nr. Balonne R., 18.i.1966 (B.B. Lowery), ibid., reddish soil plain, box pine scrub,
6.i.1966 (B.B. Lowery); 15 km W Cloncurry, 12.v.1980 (B.B. Lowery); 80-100 mi S. Sarina
xii.1956-vi.1958 (Darlingtons); (Sarina to) Rockhampton, iii.1958 (Darlingtons). Northern
Territory: 3 miles SW Alice Springs, 28.vi.1951, mulga (W.L. Brown); Alice Springs,
26.xii.2003 (A. Narendra). (94 workers, 3 gynes in ANIC, MCZC, NHMW, QMBA, UQIC).
DISCUSSION
M. diversoides is closest to M. diversus but easily separated by the less prominently

developed clypeal structure. The species is found throughout Queensland and southernmost
Northern Territory.
Meranoplus christinae Schödl sp.n.

(Figs. 38, 39, 71, 82; Plate 1, B)
Mandible with three large acute teeth. Clypeus medially excavated, distinctly bidentate and
longitudinally carinulate, with denticles laterally sinuately merging into anterolateral frontal
projections. Head distinctly wider than long (CI 122), preoccipital corners bulbously rounded,
the rear margin emarginate. Frontal carinae in posterior half almost straight, anteriorly abruptly
becoming broader and sinuately narrowed towards clypeus, distinctly narrower than head width
(FI 132). Antennal scrobe in lateral view surpassing middle of length of head posteriorly,
transversely carinulate in posterior half, occasionally with additional microsculpture, posteriorly
weakely demarcated from remainder of head. Genae and ventrolateral sides of head carinate to
rugoreticulate, preoccipital lobes reticulate. Compound eyes of moderate size (EL 0.30, REL
0.18, with 17 ommatidia in the longest row) situated well in front of middle of lateral sides of
head, dorsal ocular margin distinctly separated from ventral scrobal margin.
Promesonotum markedly wider than long (PMI 135), translucently margined laterally as
well as posteriorly, concealing lateral sides of mesosoma and propodeal declivity. Mesosomal
section markedly narrower than pronotum, with acute well developed projections. Propodeal
spines rather long (PSL 0.69) situated above middle of length of declivity, acute and straight,
moderately diverging when seen from above.
Petiole higher than long (PTI 78), in profile with anterior face straight, posterior face
convex, distinctly and regularly costate. Postpetiole elongately nodiform with ventral medium
sized tooth, rugose throughout.
First gastral tergite elongately and irregularly carinulate in basal half, posteriorly with
microreticulum and interspersed glossy spots, with pilosity of differently sized setose hairs.
Dorsum of head anteriorly rugose, with additional rugulae and few transverse ridges, posteriorly
reticulate, interspaces with microsculpture; with scattered arcuate thin hairs and fewer setose
ones. Promesonotal shield coarsely rugoreticulate, with similar, though longer pilosity.
Concolorous brown to fuscous, frequently with the gaster somewhat brighter.
129 - 134, CI 120 - 123, CS 1.78 - 1.91, SL 0.91 - 1.0, SI1 46 - 49, SI2 50 - 54, PML 1.20 -
1.43, PW 1.65 - 1.83, PMI 125 - 140, PMD 1.70 - 1.90, PMI2 101 - 106, ML 1.5 - 1.7, PSL 0.69
- 0.75, PTLL 0.45 - 0.53, PTLH 0.61 - 0.75, PTI 70 - 78, PTDW 0.56 - 0.70, PPLL 0.45 - 0.50,
ETYMOLOGY
Named for Christine, my partner. I simply adore her.
TYPE MATERIAL
Holotype worker, Queensland: 'Qld:26°47.8'Sx145°49.3'E "Merigol", site 4, mulga. Apr

2001. T.Beutel. pitfall trap #4. 10858' (QMBA). Paratypes. 6 workers, same data as holotype;
62 workers, 1 gyne with same locality data but various site and pitfall-trap numbers (ANIC,
NHMW, QMBA).
Queensland: Gumbardo, iv.2001 (T. Beutel). New South Wales: Rata, 16.xii.1965, Sparse
sclerophyll (B.B. Lowery); Condobolin, 11.i.1967, 650ft., Red soil (B.B. Lowery). (30 workers,
1 gyne in ANIC, NHMW, QMBA).
DISCUSSION
One of the larger species, M. christinae may be distinguished from all others by the clypeal
projection and in the almost flanged frontal carinae. M. occidentalis, which has a similar clypeus
is readily separated by the distinctly larger eyes, by the different promesosomal shield with
missing posterior projections and by the different distribution.
Known from south-central Queensland and New South Wales.
Meranoplus mars Forel, 1902

(Figs. 14, 15, 59, 78, 94)
Meranoplus mars Forel, 1902: 454 (Charters Towers, Queensland; worker). - Taylor & Brown, 1985: 68
(catalogue), Taylor, 1987: 39 (listed), Bolton, 1995: 251 (catalogue), Shattuck, 1999: 143 (listed).
Lectotype worker (MHNG, here designated), 'Typus [printed on red label] \ M. Mars [?'type';
illegible] Forel Chartesr Towers [?'ou' or 'ost'] Queensland (Wiederkehr) [handwritten] \ sp. M. Mars
Forel [handwritten] \ Coll. Forel [printed]'; 3 paralectotypes in ANIC, MHNG, NHMB.
1.43 - 1.69, SL 0.80 - 0.95, SI1 52 - 59, SI2 55 - 61, PML 1.0 - 1.3, PW 1.25 - 1.55, PMD 1.32 -
1.63, PMI2 102 - 111, ML 1.35 - 1.65, PTLL 0.40 - 0.48, PTLH 0.55 - 0.65, PTDW 0.45 - 0.59,
PPLL 0.33 - 0.43, PPLH 0.55 - 0.68, PPI 57 - 68, PPDW 0.45 - 0.60, PT/PP 93 - 105.
Mandible with three teeth. Clypeus in full face view anteriorly tridentate, the median tooth
frequently reduced to a blunt bulge, or all three teeth reduced to short blunt dents, moderately
surpassing anterolateral frontal corners. Head moderately wider than long (CI 104 - 113),
preoccipital corners evenly rounded, rear margin with shallow depression. Frontal carinae
posteriorly almost parallel-sided, anteriorly broadly rounded and evenly narrowed towards
clypeus, not distinctly narrower than head width (FI 115 - 122). Antennal scrobe in lateral view
posteriorly reaching far beyond middle of head, feebly to distinctly transversely carinulate in
posterior half, occasionally with additional shagreening, posteriorly distinctly demarcated from
remainder of head. Genae and ventrolateral sides of head carinate to rugose, preoccipital lobes
reticulate. Compound eyes rather large (EL 0.26 - 0.33, REL 0.18 - 0.21, with 16 - 19
ommatidia in the longest row) situated at about middle of lateral sides of head, dorsal ocular
margin confluent with ventral scrobal margin.
Promesonotum moderately wider than long (PMI 112 - 129), only pronotum narrowly
translucently margined, propodeal declivity partly visible from above. Anterior mesonotal
projections hook-like, posterolateral and postrior teeth more or less acute and narrow,
mesonotum posteriorly distinctly indented. Propodeal spines of moderate length (PSL 0.48 -
0.65), slightly arcuate and acute, distinctly diverging when seen from above.
Petiole higher than long (PTI 71 - 80), in profile triangular with anterior face straight and
unsculptured, posterior face convex, distinctly and regularly costate. Postpetiole elongately
nodiform distinctly tapering towards base, with ventral medium sized tooth, rugose throughout.
First gastral tergite with dense microreticulum, basally with additional scattered carinulae.
Dorsum of head regularly costate, with few oblique transverse meshes, at very rear
reticulate; interspaces with microsculpture. Promesonotal shield elongately rugose to
rugoreticulate. All dorsal surfaces with short decumbent arcuate and long outstanding (> 500
µm) thin hairs.
Concolorous brown.
MATERIAL EXAMINED
Northern Territory: Kunoth Polk nr Alice Spings, 14-17.ii.1975 (P.J.M. Greenslade); 17

mi NW Hamilton Downs H.S., 9.iv.1963 (McInnes & Dowse). Queensland: Tingoora,
Chinchilla Rd., 4.vi.1959 (coll. unknown); Townsville, 1952 (S. Couleth); St George's, nr.
Balonne River, 18.i.1966 (B.B. Lowery); 45 km N Bowen, Bruce H'way, 7.iv.1981 (B.B.
Lowery). New South Wales: Mungindi, 21.i.1966 (B.B. Lowery). South Australia:
Innamincka, 8.3 km SSW Candradecka Dam, 10.-14.xi.1996 (coll. unknown); 2.8 km NNW
Four Hills Trig, 1.-5.iii.1996 (coll. unknown); 2.9 km SW Big Blyth Bore, 1.-5.iii.1996 (coll.
unknown); Allendale Stn., 10 km NNW Ucutanna Hill, 13.-17.xi.1995 (coll. unknown); 6.3 km
E Unducurra Hill, Eringa Stn., 14.-17.xi.1995 (coll. unknown); Innamincka, 6.6 km SW Table
Hill, 4.-9.xi.1997 (coll. unknown); Cowarie, 5 km NE Witchinna WH, 29.iv.1995 (Brandle); 3
km NW Apollo Bore, iv.1995 (coll. unknown); Murnpeowie, 1 km WSW Mt Playford,
11.xi.1994 (J. Reid); Mulga View Stn., 6.3 km SSE Mulga View H.S. (coll. unknown) (59
workers, 4 gynes in ANIC, NHMW, SAMA, UQIC).
DISCUSSION
M. mars is distinct by the clypeal structure, by the hook-like posteriorly directed mesonotal
spines in additon with the strongly diverging propodeal spines. M. wilsoni, with which it occurs
sympatrically partly has the clypeus bidentate and never with an additional medial bulge and
less diverging propodeal spines. Distributed from south-central to north-east coastal Australia.
Meranoplus deserticola Schödl sp.n.

(Figs. 22, 23, 63, 81)
Mandible with four teeth. Clypeus carinulate, in full face view bluntly bidentate, scarcely
surpassing anterolateral frontal projections. Frontal carinae rather broadly and distinctly
sinuately narrowed towards clypeus (FI 127). Head wider than long (CI 115), posterior
preoccipital margin very shallowly concave only. Antennal scrobe surpassing middle of lateral
sides of head, glossy with additional distinct transverse carinulae at rear, merging weakly
defined into posterior section of head. Genae and ventrolateral sides of head carinate,
preoccipital corners reticulate. Eyes relatively larger (EL 0.32, REL 0.21, with 19 ommatidia in
the longest row), situated at about middle of lateral sides of head, dorsal ocular margin very
close to ventral scrobal margin. Frons rather evenly longitudinally carinate, head only
posteriorly reticulate. Promesonotal shield vaulted, with somewhat angulate profile, moderately
wider than long (PMI 118), lateral sides narrowly translucently margined.
Pronotum anteriorly reticulate, remainder of shield longitudinally rugose, with few cross
meshes. Propodeal declivity partly overhung by posterior mesonotal margin. Propodeal spines
moderate long (PSL 0.50) situated above middle of length of declivity, straight, acute and
moderately diverging when seen from above.
Petiole in lateral view triangular (PTI 67), with anterior face straight, meeting broadly
convex posterior face in a crest. Postpetiole elongately nodiform with a rather large antero-basal
tooth.
Gaster entirely microreticulate, with occasional glossy spots inbetween. Surface covered
with evenly distributed pilosity consisting of short decumbent and long, more or less erect
outstanding stiff hairs.
125 - 129, CI 113 - 118, CS 1.54 - 1.68, SL 0.90 - 0.98, SI1 53 - 55, SI2 56 - 59, PML 1.20 -
0.48, PPLH 0.68 - 0.75], PPI 57 - 63, PPDW 0.51 - 0.61, PT/PP 95 - 107, EL 0.29 - 0.34, REL
ETYMOLOGY
The name refers to the preferred habitat.
TYPE MATERIAL
Holotype worker, Northern Territory: '25.18S 130.44E NT. Valley of Winds The Olgas,
18 Nov. 1977 J.E. Feehan \ associated collector T.A. Weir \ ANIC ANTS VIAL 42.37' (ANIC).
Paratypes. 7 workers, same data as holotype; 7 workers, same data as holotype but 'ANIC
ANTS VIAL 44.232' (ANIC, NHMW).
South Australia: 38 km ESE Amata, 21.-24.x.1998 (coll. unknown); Musgrave Ra., 2.5 km
SW Womikata Bore HmL., 18.-21.x.1994 (coll. unknown); 7 mi W of Mt Davis, Tomkinson
Ra., 19.x.1960 (McInnes & Dowse). Western Australia: 28 km ESE Warburton, 15.xi.1977
(J.E. Feehan); 1 mi S Agnew, 12.x.1960 (McInnes & Dowse) (40 workers, 1 gyne in ANIC,
NHMW, SAMA).
DISCUSSION
Variability in this species becomes apparent in the rugosity of the shield. Occasionally the
dorsal ocular margin may be confluent with the ventral scrobal margin. Specimens from South
Australia are somewhat more shiny. Closest related to crassispina but different by the features
given below under that taxon. Occuring in arid central Australia.
Meranoplus crassispina Schödl sp.n.

(Figs. 24, 25, 64, 85)
0.50, PPLL 0.35, PPLH 0.65, PPI 54, PPDW 0.48, PT/PP 105.26.
Mandible with four teeth. Clypeus carinulate, in full face view bluntly bidentate, scarcely
surpassing anterolateral frontal projections. Frontal carinae broad and distinctly sinuately
narrowed towards clypeus (FI 120). Head wider than long (CI 113), preoccipital margin very
shallowly concave only. Antennal scrobe distinctly surpassing middle of lateral sides of head,
glossy with additional distinct transverse carinulae at rear, merging very weakly defined into
posterior section of head. Genae and ventrolateral sides of head carinate, preoccipital corners
reticulate. Eyes moderately sized (EL 0.28, REL 0.21, with 16. ommatidia in the longest row),
situated slightly in front of lateral sides of head, dorsal ocular margin touching ventral scrobal
margin. Frons rather evenly longitudinally costulate, head only posteriorly reticulate.
Promesonotal shield vaulted, with evenly convex profile, moderately wider than long (PMI
118), lateral sides narrowly translucently margined.
Pronotal margins almost parallel-sided, with distinct indentation at level of promesonotal
suture, with the anterior mesonotal projections distinctly defined. Promesonotal shield coarsely
rugose, with occasional cross meshes only. Propodeal declivity partly overhung by posterior
mesonotal margin. Propodeal spines in relation to body size very long (PSL 0.63), situated high
above middle of length of declivity, massively developed, straight, apically acute and distinctly
diverging when seen from above.
Petiole in lateral view broadly triangular (PTI 71), with anterior face straight, meeting
convex posterior face in a crest. Postpetiole elongately nodiform with a rather large
anteroventral tooth.
Gaster entirely microreticulate. Surface covered with evenly distributed pilosity exsiting of
short decumbent and long, more or less erect outstanding stiff hairs.
120 - 122, CI 113 - 118, CS 1.39 - 1.40, SL 0.78 - 0.80, SI1 53, SI2 55 - 58, PML 1.05 - 1.25,
PW 1.25 - 1.38, PMI 110 - 119, PMD 1.43 - 1.53, PMI2 111 - 114, ML 1.30 - 1.50, PSL 0.58 -
ETYMOLOGY
The name refers to the shape of the propodeal spines.

TYPE MATERIAL
Holotype worker, South Australia: 'S. AUST., 5.6 km SSE Mosquito Camp Dam, Crown
Point Stn 26°09'28"S 134°30'49"E \ pitfalls 20.-23.Nov 1995 Stony Desert Survey LC05'
(SAMA). Paratypes. 12 workers, same data as holotype; 1 worker, '6.8 km WSW Mosquito
Camp Dam 26°08'44"S 134°25'56"E \ pitfalls 20.-23.Nov 1995 Stony Desert Survey LC02'
(ANIC, SAMA, NHMW).
Western Australia: Kimberley district, N.W. Austr (Mjöberg) (1 worker in NHRS).
DISCUSSION
This taxon in many respects resembles deserticola and obviously is a sibling. However, the
broader frontal carinae, the outline of the dorsal shield and, in particular the distinctly differently
structured propodeal spines separate crassispina well from the former. Between the known
localities in Central and north-western Australia a wide gap is present, which most likely is due
to insufficient collecting.
Meranoplus wilsoni Schödl sp.n.

(Figs. 16, 17, 60, 88)
SI2 57, PML 1.03, PW 1.30, PMD 1.44, PMI2 111 ML 1.28, PTLL 0.38, PTLH 0.55, PTDW
Mandible with four teeth. Anterior margin of clypeus in full face view medially excavated,
acutely bidentate, particularly laterally carinulate, moderately exceeding anterolateral frontal
margins. Head moderately wider than long (CI 115), with preoccipital margin shallowly
concave. Frontal carinae posteriorly almost parallel-sided, anteriorly broadly translucent and
evenly narrowed towards clypeus, moderately narrower than head width (FI 118). Antennal
scrobe reaching far beyond middle of lateral side of head, distinctly transversely carinulate in
posterior half, only weakly demarcated from remainder of head at the rear. Genae and
ventrolateral sides of head carinate to rugose, postocular section reticulate. Compound eyes
moderately large (EL 0.25, REL 0.20, with 15 ommatidia in the longest row) situated at about
middle of lateral sides of head, dorsal ocular margin almost touching ventral scrobal margin.
Promesonotum wider than long (PMI 127) and rather flat, outline distincly translucently
margined, concealing lateral sides of mesosoma and propodeal declivity. Propodeal spines of
medium length (PSL 0.53) situated above middle of length of declivity, acute and slightly
arcuate when seen from above.
Petiole higher than long (PTI 68), in profile triangular with anterior face straight and
nodular, tapering towards base, with ventral medium sized tooth, rugose throughout.
First gastral tergite with irregular microreticulum, with interspersed glossy spots, basally
with additional carinulae. Dorsum of head only anteriorly rugose, from about level of eyes on
rugo-reticulate. Promesonotal shield with coarse rugoreticlum. Dorsal surfaces covered with
evenly distributed pilosity consisting of short decumbent and long, more or less erect
outstanding stiff hairs, similar to that of preceding species.
Concolorous brown.
114 - 121, CI 112 - 118, CS 1.20 - 1.39, SL 0.71 - 0.83, SI1 53 - 57, SI2 57 - 62, PML 0.9 - 1.2,
ETYMOLOGY
Named for Prof. Edward O. Wilson, outstanding scientist and one of the pioneers of modern
myrmecology.
TYPE MATERIAL
Holotype worker, Queensland: 'Qld. 10 km S Bowen 24.v.1981 BBLowery Casuarina

scrub \ ANIC ANTS VIAL 68.187' (ANIC). Paratypes. 8 workers, same data as holotype; 12
workers, '8 km W Bowen 10.viii.1981 BBLowery, woodland \ ANIC ANTS VIAL 68.16'
(ANIC, MCZC, NHMW).
Queensland: 'Queensland', 1902-319. / [overleaf]: 22.4.02 (F.P. Dodd); Townsville,

13.i.1997 (D. Grund); Allinga, Chinchilla, Charleys Ck., 18.i.1986 (G. Lithgow); Woodstock,
1.iv.1980 (B.B. Lowery); 4 mi WNW Yelarbon, 1.xii.1949 (T. Greaves); 40 km W Warwick,
Gore Rail Res., 4.i.1966 (B.B. Lowery); St Georges, nr. Balonne River, 7.i./18.i.1966 (B.B.
Lowery); Gladstone, ii.1915 (F.H. Taylor); MacKay, ii.1915 (F.H. Taylor); Mount Elliot,
Lingum Townsville, 2.iii.1958 (P.F. Darling). New South Wales: Mungindi, 21.i.1966 (B.B.
Lowery); Legume, 20.ix.1974 (P.J.M. Greenslade). (62 workers, 3 gynes in ANIC, BMNH,
MCZC, NHMW, QMBA, SAMA).
DISCUSSION
This is a uniform taxon that shows only similarities with mars (see under that species).
Distributed in the Northern Murray-Darling Basin to NE coastal Queensland.
Meranoplus tricuspidatus Schödl sp.n.

(Figs. 18, 19, 61, 87)
Mandible with four teeth. Clypeus in full face view surpassing anterolateral frontal corners,
acutely bidentate, with an additional short acute medial denticle, accompanied by a low crest.
Head only moderately wider than long (CI 106), rear margin shallowly concave. Frontal carinae
evenly and rather broadly narrowing towards clypeus (FI 130). Antennal scrobe in lateral view
surpassing middle of length of head posteriorly, distinctly transversely carinulate in posterior
half, with additional microreticulation, posteriorly distinctly demarcated from remainder of
head. Genae rugose, ventrolateral sides of head rugoreticulate, preoccipital lobes reticulate.
Compound eyes of moderate size (EL 0.27, REL 0.18, with 16 ommatidia in the longest row) in
lateral view situated in front of middle of lateral sides of head, dorsal ocular margin not reaching
ventral scrobal margin.
Promesonotum wider than long (PMI 119), outline broadly translucently flanged,
concealing lateral sides of mesosoma and propodeal declivity. Propodeal spines relatively short
(PSL 0.51), acute and slightly arcuate and diverging, when seen in dorsal view.
Petiole distinctly higher than long (PTI 68), in profile with anterior face straight and
nodiform, with ventral medium sized tooth, rugose throughout.
First gastral tergite with dense microreticulum, basally with additional carinulae, with
relatively short stiff decumbent and additional longer erect hairs. Dorsum of head longitudinally
costulate anteriorly, posteriorly distinctly reticulate, interspaces with additional distinct
microreticulum, with short decumbent hairs and longer stiff erect differently sized pilosity.
Promesonotal shield rugoreticulate, pilosity similar to that of head.
Brown to fuscous with the gaster, legs and antennae slightly brighter.
129 - 137], CI 106 - 109, CS 1.28 - 1.58, SL 0.67 - 0.83, SI1 50 - 52, SI2 52 - 55, PML 0.93 -
0.16 - 0.18, with 13 - 17 ommatidia in the longest row
ETYMOLOGY
The name refers to the tridentate clypeal projection.
TYPE MATERIAL
Holotype worker, Queensland: 'Qld. Cape Upstart 21.xi.1980 BBLowery Cleared ground \
ANIC ANTS VIAL 68.198' (ANIC). Paratypes. 8 workers, same data as holotype; 12 workers,
'Qld. Cape Upstart Mt Ayr 21.xi.1980 BBLowery flat sea level \ ANIC ANTS VIAL 67'.
(ANIC, NHMW).
Queensland: Emerald, 15.xii.1972 (B.B. Lowery); Moonie, 60 mls N Goondiwindi, 4.i.1965

(B.B. Lowery); N of Bowen, xi.1957 (Darlington); Allinga, Chinchilla, 31.vii.1986 (G.
Lithgow). Northern Territory: Kunnoth Polk nr. Alice Springs, x.1974 & 24.-26.ix.1980
(P.J.M. Greenslade). (13 workers, 1 gyne in ANIC, MCZC, NHMW, QMBA).
DISCUSSION
Superficially this taxon somewhat resembles M. wilsoni and M. mars, but is readily
separated by the clypeus and, in addition by the translucently margined promesonotal shield.
Paratypes and additional other material from Queensland resemble the type. Specimens
from Northern Territory show a slightly longer and thinner pilosity and the dorsal surface of the
head is shiny with the microsculpture less developed. Known from coastal and south-central
Queensland, with a single sample taken at Alice Springs. The disjunct occurrence of M.
tricuspidatus most likely is only due to the lack of additional samples. This gap might be closed
as soon as new material shows up.
Meranoplus duyfkeni Forel, 1915

(Figs. 10, 11, 57, 80, 92)
Meranoplus diversus r. Duyfkeni [sic] Forel, 1915: 45 (Kimberley District, Western Australia; worker,
gyne) - Taylor & Brown, 1985: 67 (catalogue), Taylor, 1987: 38 (listed), Taylor, 1990: 34 (Raised to
species), Bolton, 1995: 251 (catalogue), Shattuck, 1999: 142 (listed). Lectotype worker (MHNG, here
designated), 'Typus [printed on red label] \ Meranoplus diversus Sm. r. Duyfkeni For. type
[handwritten] \ Australien Mjöberg No 33 [handwritten] \ Coll. A. Forel [printed]'. Fifteen
paralectotypes (workers, 1 gyne in MHNG, NHMB, ANIC, MCZC and NHRS.
Remarks: Out of 14 topotypical workers and two gynes, housed in NHRS, only one worker and one gyne
are labeled as types. Since it is subsequently impossible to state which of the specimens Forel had before
him, only the two specimens labeled as types are here considered to belong to the original syntype series.
1.63 - 1.89, SL 0.90 - 1.03, SI1 50 - 52, SI2 53 - 56, PML 1.05 - 1.20, PW 1.32 - 1.53, PMD
1.35 - 1.60, PMI2 100 - 105, ML 1.45 - 1.65, PTLL 0.36 - 0.45, PTLH 0.58 - 0.70, PTDW 0.49
- 0.63, PPLL 0.38 - 0.45, PPLH 0.51 - 0.70, PPI 60 - 78, PPDW 0.46 - 0.58, PT/PP 95 - 109.
Mandible with four teeth. Clypeus in full face view bluntly bidentate, additionally
carinulate, dents not or only weakly surpassing anterolateral corners of frontal carinae. Head
wider than long (CI 113 - 119) with the preoccipital corners evenly rounded and the rear margin
concave. Frontal carinae posteriorly narrow, anteriorly broadly and translucently narrowing
towards clypeus, distinctly narrower than head width (FI 131 - 139). Antennal scrobe in lateral
view surpassing middle of length of head posteriorly, finely carinulate in whole length, with
additional distinct microreticulum, posteriorly rather well defined from remainder of head.
Genae below scrobal margin evenly carinulate to carinate, ventrolateral sides of head carinate to
rugose, preoccipital lobes reticulate. Compound eyes moderate in size (EL 0.29 - 0.34, REL
0.19 - 0.20, with 17 - 18 ommatidia in the longest row) in lateral view situated slightly in front
of middle of lateral sides of the head, dorsal ocular margin well separated from ventral scrobal
margin.
Promesonotum trapezoid, wider than long (PMI 120 - 129), narrowly translucently
margined, concealing lateral sides of mesosoma, propodeal declivity visible from above.
Propodeal spines of medium length (PSL 0.50 - 0.63) situated above middle of length of
declivity, acute and straight and distinctly diverging in dorsal view.
Petiole distinctly higher than long (PTI 59 - 70), in profile triangular with anterior face
straight and unsculptured, the posterior face convex to angulately rounded, rugose. Postpetiole
nodiform, with large ventral tooth.
First gastral tergite entirely microreticulate, with occasional interspersed glossy spots,
basally an indistinct fine striation may be apparent. Dorsum of head costulate to rugose with
more or less developed microsculpture, frontal margins irregularly sculptured, with pilosity
consisting of decumbent arcuate short and scattered outstanding long hairs. Promesonotal shield
coarsely irregularly rugose to rugoreticulate.
Concolorous brown to fuscous.
MATERIAL EXAMINED
Northern Territory: Keep River National Park E Kununurra, 15.vii.1990 (R.P. McMillan).
Western Australia: Derby, vi.1984 (G. McKenzie), ibid. (W.D. Dodd); 'Kimberly R.S.', iii.1950
(B. Rudeforth) (19 workers in ANIC, JDMP, MCZC, NHMW, WAMP).
DISCUSSION
This medium sized species is readily separated from others by the unique translucently
flanged promesonotal shield with projections on mesonotal hind margin directed posteriorly.
Restricted to north-western Australia.
Meranoplus orientalis Schödl sp.n.

(Figs. 12, 13, 58, 80)
SI2 58, PML 1.20, PW 1.48, PMD 1.48, PMI2 100, ML 1.70, PTLL 0.50 PTLH 0.65, PTDW
Mandible with four teeth. Clypeus in full face view bluntly bidentate additionally
carinulate, denticles not or only weakly surpassing anterolateral corners of frontal carinae. Head
wider than long (CI 115) with the preoccipital corners evenly rounded and the rear margin
shallowly concave. Frontal carinae posteriorly parallel-sided, more evenly and not as broadly
and translucently narrowing towards clypeus, still distinctly narrower than head width (FI 125).
Antennal scrobe in lateral view reaching far beyond middle of length of head posteriorly,
distinctly carinulate in whole length, with additional distinct microreticulum, posteriorly ill
defined from remainder of head. Genae below scrobal margin and ventrolateral sides of head
carinate, preoccipital lobes reticulate. Compound eyes moderately large (EL 0.34, REL 0.21,
with 19 ommatidia in the longest row) in lateral view situated slightly behind middle of lateral
sides of the head, dorsal ocular margin touching ventral scrobal margin.
Promesonotum trapezoidal, wider than long (PMI 123) without developed shield, propodeal
declivity visible from above. Promesonotal projections almost completely reduced with
posterolateral spines bluntly triangular and the posterior projections as short denticles.
Propodeal spines of medium length (PSL 0.53) situated above middle of length of declivity,
massively built, acute and straight and distinctly diverging in dorsal view. Petiole higher than
long (PTI 77), in profile triangular with anterior face straight and unsculptured, the posterior
face convex, regularly costate. Postpetiole elongately nodiform, with medium sized ventral
tooth. First gastral tergite entirely microreticulate, with interspersed glossy spots, basally an
indistinct fine striation may be apparent. Dorsum of head costate, with indistinctly developed
microsculpture, frontal margins irregularly sculptured. Promesonotal shield regularly elongately
rugose. With pilosity similar to that of preceding species.
Concolorous brown to fuscous.
121 - 126, CI 113 - 116, CS 1.48 - 1.70, SL 0.90 - 0.98, SI1 53 - 57, SI2 57 - 61, PML 1.13 -
ETYMOLOGY
Named for its easterly distribution.
TYPE MATERIAL
Holotype worker, Queensland: '6 m. W. of Blackwater Qld. 26.III.62. J.E.Dowse \ Series

A95' (ANIC). Paratypes. 5 workers (one coated), same data as holotype (ANIC, NHMW).
Queensland: 30 mls E of Morven S.CQ., 16.v.1963 (Dahms) (4 workers in QMBA,

NHMW).
DISCUSSION
Doubtless M. orientalis is closely related to duyfkeni. However, apart from the completely
different distribution there are distinct characteristics that clearly separate the two taxa. The
clypeus in orientalis occasionally shows an additional anterio-median blunt bulge, the
promesonotum is never translucently flanged and its dentition is either absolutely missing or, in
specimens from Morven, slightly more clearly developed but still very similar to that of the type.
Restricted to southern Queensland.
Meranoplus oxleyi Forel, 1915

(Figs. 20, 21, 62, 86, 93)
Meranoplus diversus r. oxleyi Forel, 1915: 45 (Kimberley district, Western Australia; worker) - Taylor &
Brown, 1985: 67 (catalogue), Taylor, 1987: 38 (listed), Taylor, 1990: 34 (raised to species), Bolton,
1995: 251 (catalogue), Shattuck, 1999: 143 (listed). Lectotype worker (MHNG, here designated),
'Typus [red label, printed] \ jan [printed] \ Kimberley district [printed] \ N. V. Austr. Mjöberg [printed]
\ Meranoplus diversus Sm. oxleyi For. % type \ Coll. A. Forel'; 2 paralectotypes in ANIC and NHRS.
Remarks: three specimens with identical locality labels, one additionally labeled as type (NHRS)
[whether all three are syntypes cannot be stated; however, the specimen with the type label is here
regarded to be a paralectotype].
1.10 - 1.34, SL 0.65 - 0.78, SI1 53 - 57, SI2 55 - 60, PML 0.88 - 1.0, PW 1.05 - 1.25, PMD 1.20
- 1.43, PMI2 113 - 119, ML 1.0 - 1.25, PTLL 0.33 - 0.38, PTLH 0.48 - 0.60, PTDW 0.43 - 0.54,
PPLL 0.30 - 0.40, PPLH 0.45 - 0.55, PPI 63 - 73, PPDW 0.40 - 0.49, PT/PP 103 - 111.
Mandible with four teeth. Clypeus in full face view acutely bidentate, medially excavated,
not or only weakly surpassing anterolateral frontal angles, glossy except carinulae posterior to
dentition. Head moderately wider than long (CI 108 - 114), preoccipital margin with shallow
concavity. Frontal carinae markedly broadened before narrowing towards clypeus, moderately
narrower than head width (FI 116 - 121). Antennal scrobe in lateral view surpassing middle of
length of head posteriorly, transversely carinulate in posterior half with additional shagreen,
distinctly demarcated from remainder of head. Genae and ventrolateral sides of head carinate to
rugose, preoccipital lobes reticulate. Compound eyes medium sized (EL 0.23 - 0.26, REL 0.21 -
0.24, with 15 - 18 ommatidia in the longest row) situated at or slightly behind middle of lateral
sides of head in lateral view, dorsal ocular margin confluent with ventral scrobal margin.
Promesonotum moderately wider than long (PMI 113 - 125), outline partly translucently
margined, concealing lateral sides of mesosoma and propodeal declivity. Propodeal spines of
medium length (PSL 0.45 - 0.50) situated above middle of length of declivity, considerably
diverging acutely arcuate apically when seen from above.
Petiole distinctly higher than long (PTI 58 - 68), in profile wedge-shaped with anterior face
straight and unsculptured, posterior face convex, distinctly and regularly costate. Postpetiole
dorsally nodiform, distinctly tapering towards base, with a well developed tooth, rugose
throughout.
First gastral tergite entirely microreticulate with variable amount of glossy spots inbetween,
with setose variably sized decumbent to erect pilosity. Most of dorsum of head longitudinally
costate to rugose, posteriorly reticulate, interspaces with microsculpture, with rather dense layer
of short arcuate decumbent and longer stiff suberect to erect hairs. Promesonotal shield
irregularly rugoreticulate with predominantly elongate meshes with pilosity similar to that of
head.
Concolorous brown to fuscous, the gaster and appendages brighter.
MATERIAL EXAMINED
Northern Territory: Larrimah, 16.vii.1981 (B. B. Lowery); Katherine, 5 km NW,

19.x.1977; 7.5 km SW, 21.x.1977; 34 km NW, 22.x.1977; 55 km SW, 24.x.1977; 41 km SW,
6.iv.1978; 12 km SW, 6.iv.1978; 23 km SW, 9.iv.1978 (P.J.M. Greenslade). (20 workers,
ANIC, NHMW).
DISCUSSION
A small species with little variability, except specimens from Larrimah having a more
distinct frontal microreticulation. M. oxleyi differs from its closest relatives by the relatively
large eyes (REL 0.21 - 0.24) and by the posteriorly not distinctly converging promesonotal
shield. Hitherto known from Kimberley and Top End.
Meranoplus occidentalis Schödl sp.n.

(Figs. 40, 41, 72, 79; Plate 1, C)
Mandible with four teeth. In full face view clypeus anteriorly deeply excavated, distinctly
exceeding anterolateral frontal corners as a bilobed carinulate projection. Head somewhat wider
than long (CI 109), preoccipital margin moderately concave. Frontal carinae distinctly narrower
than head width (FI 139), evenly narrowing towards clypeus. With head in profile antennal
scrobe reaching beyond middle of lateral side of head, merging into lateral sides posteriorly,
glossy except transverse carinulae at rear. Genae and ventrolateral sides of head evenly carinate,
preoccipital lobes reticulate. Eyes exceptionally large (EL 0.43, REL 0.26, with 18 ommatidia in
the longest row), situated at about middle of lateral sides of head, the dorsal ocular margin
confluent with the ventral scrobal margin. Dorsal surface of head anteriorly costulate with
oblique anastomoses, posterior section reticulate, with additional faint microsculpture.
Promesonotal shield distinctly wider than long (PMI 138), lateral projections translucently
fused, posterior projections absent, mesonotal hind margin with transverse ridge, thus entire
propodeal declivity visible from above. Propodeal spines short (PSL 0.45) and in profile broad-
based, in dorsal view acute and arcuate.
Petiole in lateral view elongately trapezoid, markedly higher than long (PTI 59), with
anterior face more or less straight, roundly meeting oblique reticulate dorsum, the latter merging
into rugose posterior face, ventrally strongly sinuate. Postpetiole dorsally nodiform with large
anterio-basal tooth.
Gaster shiny, entirely microreticulate, striate at very base. Dorsal surfaces covered with
scattered pilosity of shorter decumbent and longer more or less erect soft hairs.
131 - 144, CI 109 - 121, CS 1.70 - 1.94, SL 0.90 - 1.08, SI1 48 - 53, SI2 52 - 56, PML 1.0 -
0.48, PPLH 0.74 - 0.88, PPI 50 - 56, PPDW 0.53 - 0.66, PT/PP 90 - 104, EL 0.37 - 0.44, REL,
ETYMOLOGY
Named for its restricted westerly distribution.
TYPE MATERIAL
Holotype worker, Western Australia: 'TAMBREY WESTERN AUSTRALIA 7 AUG

1987 R.P. MCMILLAN \ ON GROUND \ Western Australian Museum Entomology Reg no.
23155' (WAMP). Paratypes. 8 workers, 3 alate gynes, 4 males, same data as holotype (WAMP,
ANIC, NHMW).
Western Australia: Lyndon Stn [several localities possible] (Snell); Ashburton River,
22.vi.1967 (G. Campbell); Jigalong (J. Hickmer); Ethel Creek, iv.1993 (P.A. Varris); Mt.
Whaleback, Newman, vii.1984 (K.J. Walker); Manilya Stn., vi.1955 (A. Douglas); 80 km S
Barradale, 22.vi.1975 (P.J.M. Greenslade); 30 km S Minilya, 15.viii.1984 (B.B. Lowery); S.
bank of Gascoyne River, 10 km E of H'way, 10.viii.1983 (B.B. Lowery); 60 km N of Carnarvon
H'way, 12.viii.1984 (B.B. Lowery); Carnarvon, Ellanella Rd., 14.vii.1969 (L.E. Koch). (63
workers in ANIC, JDMP, MCZC, NHMW, SAMA, USNM, WAMP).
DISCUSSION
M. occidentalis is found in two distinct color varieties. Workers in the type series and
several additional samples are light-brown with the gaster and appendages ferrugineous. Other
samples have the gaster and appendages brown with the remainder of the body piceous.
Intermediate forms are so far unknown. Gynes in the the type sample are concolorous, males
strictly bicolored. The species is not to be mixed up with any other by the large eyes and the
unique promesonotal shield. Known from south-western and western coastal Australia.
Meranoplus angustinodis Schödl sp.n.

(Figs. 42, 43, 73, 81)
SI2 61, PML 1.12 PW 1.40, PMD 1.62, PMI2 116, ML 1.33, PTLL 0.35, PTLH 0.63, PTDW
Mandible with four teeth. In full face view clypeus anteriorly with deep oblique concavity,
markedly exceeding anterolateral frontal corners and distinctly carinulate. Head wider than long
(CI 115), preoccipital margin moderately concave. Frontal carinae broad, only moderately
narrower than head width (FI 118), evenly narrowing towards clypeus. With head in profile
antennal scrobe reaching distinctly beyond middle of lateral side of head, transversely carinulae
at rear, well defined posteriorly against remainder of head. Genae evenly carinate, ventrolateral
sides of head and posterolateral lobes reticulate. Eyes medium sized (EL 0.33, REL 0.26, with
18 ommatidia in the longest row), situated at about middle of lateral sides of head, the dorsal
ocular margin reaching the ventral scrobal margin. Dorsal surface of head coarsely
rugoreticulate, interspaces glossy.
Promesonotal shield moderately wider than long (PMI 127), with a somewhat overall
square impression, with mesonotal projections well developed, posterolateral spines curved and
with distinct translucent fusion between projections. Propodeal spines not very long (PSL 0.45),
narrow and acute, in dorsal view moderately arcuate.
Petiole in lateral view narrowly triangular, distinctly higher than long (PTI 56), with
anterior face more or less straight, meeting convex rugose posterior face in a crest, ventral face
sinuate. Postpetiole elongately nodiform with large antero-basal tooth.
First gastral tergite entirely microreticulate. Dorsal surfaces covered with shorter decumbent
and longer erect and more stiff hairs.
Distinctly bicolored with the gaster and appendages yellow, the femora basally fuscous.
Remainder of body piceous, except frontal carinae anterolaterally and translucent areas on
promesonotal shield brighter.
113 - 123, CI 111 - 119, CS 1.20 - 1.43, SL 0.75 - 0.88, SI1 56 - 60, SI2 60 - 64, PML 0.93 -
ETYMOLOGY
The name refers to the narrow shape of the nodes.
TYPE MATERIAL
Holotype worker, South Australia: 'S.Aust.-Collinsville, 4.5 km SW 33°22'S 139°17'E,

19-23.x.1992 South Olary Plains Survey' (SAMA). Paratypes. 10 workers, same data as
holotype (ANIC, NHMW, SAMA).
South Australia: Middleback Stn. HS, 20.-25.vii.1995, (McAllister); 11.5 km SE Wares

Peak, 5.x.1995 (coll. unknown); 8 km SE Paralena Hill, Woltana Stn., 26.-31.viii.1996 (coll.
unknown); 16 km SSE Mount Coffin, 9.-12.xii.1997 (coll. unknown); 14.7 km ESE Mt. Coffin,
Nth Moolooloo, 9.-12.xii.1997 (coll. unknown); 9 km SSE Oakbank, 7.-10.x.1992 (coll.
unknown); Murnpeowie, 17.xi.1992 (R. Brandle); Murnpeowie, Mary's Pool, 17.xi.1992 (R.
Brandle); Mount Hopeless, 15.xi.1994 (R. Tynan); Screech-Owl Creek, 27.iv.1995 (coll.
unknown); Willow Spring Stn., 13.5 km NNE Willow Spring HS, 15.-25.iii.1995 (coll.
unknown); 4.5 km SW Waraweena, 4.-7.vii.1999 (coll. unknown); 3 km SSW Waraweena, 1.-
10.x.1999 (coll. unknown); Coober Pedy, 26.v.1969 (P.J.M. Greenslade); Halbury, 10.x.1984
(B.B. Lowery); Flinders Ranges, Oraparinna, 15.-17.ix.1971 (P.J.M. Greenslade); Moockra
Tower, 6.x.1974 & 13.x.1980 (P.J.M. Greenslade); Tailem Bend, 21.vi.1969 (B.B. Lowery);
Hawker, 18.vi.1972 (B.B. Lowery).Victoria: Patho (H.A. Petter); N Ouyen, ix.1913 (coll.
unknown). New South Wales: 15 km N Coombah, 31.x.1975 (P. Ward); Fowlers Gap Stn., 110
km N Broken Hill, 11.x.1980 (S.R. Morton); Hillston, 8.xii.1965 (B.B. Lowery). (84 workers, 6
gynes, 1 male in ANIC, MVMA, NHMW, SAMA).
DISCUSSION
M. angustinodis is a very distinctive species, which is easily separated from others by the
narrowly triangular petiole and the elongate large-toothed postpetiole. Widely distributed
throughout south-eastern Australia.
Meranoplus variabilis Schödl sp.n.

(Figs. 44, 45, 74, 89)
Mandible with four teeth. In full face view lateral sides of clypeal projection distinctly
converging, anteriorly narrowly excavated with anterolateral corners produced into triangular
blunt teeth, surpassing anterolateral frontal corners, the clypeal surface coarsely rugose,
medially less structured. Head wider than long (CI 114), preoccipital margin shallowly concave.
Frontal carinae moderately narrower than head width (FI 126), evenly sinuately narrowing
towards clypeus. With head in profile antennal scrobe reaching beyond middle of lateral side of
head, with distinct transverse carinulae at the rear, posteriorly ill defined against preoccipital
section. Genae and ventrolateral sides of head rugoreticulate, preoccipital lobes reticulate. Eyes
moderate in size (EL 0.31, REL 0.25, with 18 ommatidia in the longest row), situated in front of
lateral sides of head, the dorsal ocular margin touching the ventral scrobal margin. Dorsal
surface of head almost entirely reticulate, only frons anteriorly rugose, without any further
microsculpture.
Promesonotal shield somewhat square, distinctly wider than long (PMI 138), projections
except posterior bluntly rounded ones well developed and more or less acute, margins
overhanging lateral mesosomal sides and propodeal declivity. Propodeal spines moderate in
length (PSL 0.44), narrowly and acutely triangular, in dorsal view more or less straight and
distinctly diverging.
Petiole in lateral view broadly trapezoid, higher than long (PTI 73), with straight anterior
and posterior faces roundly meeting feebly oblique reticulate dorsum. Postpetiole nodiform with
small antero-basal tooth.
Gaster shiny, entire first gastral tergite densely and evenly microreticulate, at base with
irregular carinulae. Dorsal surfaces covered with a layer of short decumbent and longer more or
less erect soft hairs.
Strictly bicolored with similar patterns as in preceding species.
126 - 129, CI 107 - 115, CS 1.19 - 1.33, SL 0.70 - 0.80, SI1 56 - 60, SI2 59 - 63, PML 0.85 -
1.0, PW 1.13 -1.38, PMI 125 - 147, PMD 1.20 - 1.45, PMI2 104 - 113, ML 1.08 - 1.28, PSL
ETYMOLOGY
The name refers to the variable overall shape of the species.
TYPE MATERIAL
Holotype worker, South Australia: 'S.Aust.- Cullyamurra WH Coopers Creek 140°50'E

27°42'S 21-24.ix. 1990 J. Shaw ANZSES Surv.' (SAMA). Paratypes. 6 workers, same data as
holotype (ANIC, SAMA, NHMW).
South Australia: Mount Barry Stn., 4.3 km SSE Jimmy WH, 18.-21.ix.1996 (coll.
unknown); 4 km N Halifax Hill, 29.ix.-5.x.1995 (coll. unknown); Mount Aroona, 2.-5.xii.1997
(coll. unknown); 19.9 km WNW Indulkana, 25.-31.x.1998 (coll. unknown); Murnpeowie,
Weathered Hill, 3 km ENE 18.xi.1992 (R. Brandle); Murnpeowie, Mount Hopeless,
15./16.xi.1994 (R. Tynan); Flinders Ranges, Oraparinna, 15.ix.1971 (P.J.M. Greenslade);
Moockra Tower, 7.x.1974 / 13.x.1980 (P.J.M Greenslade); Monarto Zoological Park, c 50 km E
Adelaide, 2.xii.2003 (coll. unknown); 26 km ENE Mimilli, 25.-31.x.1998 (coll. unknown); 12.5
km W Mimilli, pitfalls, 20.-24.iii.1993 (coll. unknown); 2 km S Wares Peak, 5.x.1995 (coll.
unknown). New South Wales: Mungindi, 21.i.1966 (B.B. Lowery); Moulamein, 29.viii.1970
(B.B. Lowery); N Nyngan 'Kapunda', 5.-6.v.1987 (P.J.M. Greenslade). Northern Territorry: 9
km E Curtin Springs, 20.xi.1977 (J.E. Feehan); Tanami Rd., 35 km W Stuart Hwy., 5.vii.2003
(Snelling). (60 workers, 2 gynes in ANIC, LACM, NHMW, SAMA).
DISCUSSION
A variable taxon regarding the outline of the promesonotal shield as well as the shape of the
petiole, the length and shape of the propodeal spines and pilosity. For this reason numerous of
the material, namely from Indulkana, Murnpeowie, Monarto, Mimilli and Wares Peak is here
only provisionally added. Further samples are needed to elucidate whether more than one
species is involved. Nonetheless, typical material is well defined by the forward oriented clypeal
projection together with the box-shaped petiole. Distributed from central to south-east Australia,
partly sympatrically occuring with M. angustinodis, reaching further to the north and east.
Meranoplus arcuatus Schödl sp.n.

(Figs. 46, 47, 77, 82)
Mandible with four teeth. In full face view the clypeal projection very similar to that of
preceding species, with the anterior concavity somewhat broader and with anterolateral corners
more blunt. Head moderately wider than long (CI 115), preoccipital margin almost straight.
Frontal carinae markedly narrower than head width (FI 125), posteriorly almost parallel-sided,
evenly curved towards clypeus. With head in profile antennal scrobe distinctly surpassing
middle of lateral side of head, distinctly transversely carinulae at the rear, anteriorly with
additional oblique carinulae, rather well defined posteriorly against remainder of head. Genae
and ventrolateral sides of head rugoreticulate, preoccipital lobes coarsely reticulate. Eyes
moderately large (EL 0.31, REL 0.26, with 15 ommatidia in the longest row), situated at about
middle of lateral sides of head, the dorsal ocular margin confluent with the ventral scrobal
margin. Frons anteriorly rugose, remainder of head coarsely rugoreticulate, interspaces glossy.
Promesonotal shield little wider than long (PMI 115), overhanging lateral sides of mesosoma
and propodeal declivity, near lateral margin with translucent fenestra at level of promesonotal
boarder. Mesonotum indented and projections well developed, translucently lamellate between
projections. Propodeal spines in profile short (PSL 0.43), situated at about middle of lateral sides
of the propodeal declivity, broad-based and acute, in dorsal view distinctly curved towards
petiole.
Petiole higher than long (PTI 68), in lateral view an equilateral triangle with anterior and
slightly convex posterior faces meeting in a crest, posterior face rugose. Postpetiole nodular with
well developed anterio-basal tooth.
First gastral tergite entirely microreticulate. Dorsal surfaces covered with shorter decumbent
and longer erect more or less setose hairs.
Distinctly bicolored as in preceding species.
117 - 125, CI 113 - 118, CS 1.15 - 1.31, SL 0.73 - 0.80, SI1 56 - 59, SI2 60 - 64, PML 1.03 -
ETYMOLOGY
The name refers to the incurved propodeal spines.
TYPE MATERIAL
Holotype worker, Western Australia: 'Lake Marmion 29°43'S 121°31'E pitfall 12-Mar-96
J A Forrest' (SAMA). Paratypes. 11 workers, same data as holotype; 6 workers, 'Nr. Lake
Marmion 29°47'S 121°35'E pitfalls, 12.ii.96 Forrest' (ANIC, NHMW, SAMA).
South Australia: Cowarie, 1.2 km SW Kalladeina Bore, 27.iv.1995 (T. Robinson); 19.5 km
WNW Anta Hill [= ?Antakatanya Hill], 16.-19.ix.1998 (coll. unknown); 22.5 km WSW Haines
Hill (coll. unknown). 25.8 km N Miandana WH, 9.-14.x.1998 (coll. unknown) (7 workers in
NHMW, SAMA).
DISCUSSION
M. arcuatus differs from other species by the distinctly incurved propodeal spines in
addition to the triangular petiole and the outline of the promesonotal shield. Only known from
the type locality and few samples in northeastern South Australia, with a wide gap in between.
Meranoplus mcarthuri Schödl sp.n.

(Figs. 48, 49, 75, 84)
Mandible with five teeth. In full face view clypeus distinctly carinulate, oblique anterior
margin broadly and deeply excavated, anterolateral corners produced into acute teeth, weakly
surpassing anterolateral frontal corners. Head wider than long (CI 121), preoccipital margin only
shallowly concave. Frontal carinae markedly narrower than head width (FI 126), weakly
sinuately narrowing towards clypeus. With head in profile antennal scrobe reaching far beyond
middle of lateral side of head, distinctly transversely carinulate with additional shagreening, well
defined posteriorly by a carina against remainder of head. Genae rugose, ventrolateral sides of
head and preoccipital lobes coarsely reticulate. Eyes moderately large (EL 0.35, REL 0.25, with
19 ommatidia in the longest row), situated slightly in front of middle of lateral sides of head, the
dorsal ocular margin confluent with the ventral scrobal margin. Dorsal surface of head coarsely
rugoreticulate, interspaces glossy.
Promesonotal shield moderately wider than long (PMI 126), mesonotum indented with
mesonotal projections well developed and rather acute, translucently fused. Propodeal spines
moderately long (PSL 0.55), situated at about middle of lateral sides of the propodeal declivity,
narrow and acute, in dorsal view markedly arcuate and diverging.
Petiole in lateral view narrowly trapezoid, higher than long (PTI 70), dorsum convex and
reticulate, posterior face rugose. Postpetiole nodular with small anterio-basal tooth.
First gastral tergite entirely microreticulate. Dorsal surfaces covered with short decumbent
and longer erect thin whitish hairs.
Distinctly bicolored as in preceding species.
WORKER (n = 8). TL 4.90 - 6.50, HL 1.25 - 1.45, HW 1.43 - 1.73, FC 1.15 - 1.45, FI 123
- 128, CI 114 - 125, CS 1.34 - 1.59, SL 0.85 - 1.0, SI1 57 - 61, SI2 61 - 66, PML 0.95 - 1.25,
ETYMOLOGY
Named for Archie McArthur, great (old) man from Adelaide, who I had the honor and
pleasure to meet in Vienna during his trip to Europe in early spring 2004. He sent hundreds of
pitfall samples which included most important material for this study and which certainly will
bring to light further interesting data.
TYPE MATERIAL
Holotype worker, Northern Territory: 'Goldunda Hstd. Alice Springs G. Campbell 26. 8.
63 Meranoplus sp. \ Royce H. Mew collection 132' (SAMA). Paratypes. 5 workers, same data
as holotype (ANIC, NHMW, SAMA).
Northern Territory: 9 km E Curtin Springs, 20.xi.1977 (J.E. Feehan). New South Wales:
2 mi. W Jacob's Well, 23.xi.1949 (T. Greaves). South Australia: 26.3 km ENE Mimili, 25.-
31.x.1998 (coll. unknown); 48.9 km SE Mount Lindsay, 26.x.1996 (coll. unknown); 8 km NW
Morgan, 15.xii.1976 (P.J.M. Greenslade); Victoria Desert, 10 km E Emu, 11.x.1976 (P.J.M.
Greenslade); 35 km S Arraby, Leap yr. Bore, viii.1975 (Forrest). Victoria: Linga (F.E.
Wilson); M. Western Australia: Morawa, 1961 (G.H. Lowe). State unknown: 'Cent Australia'
(coll. unknown) (37 workers in ANIC, NHMW, QMBA, SAMA, USNM, WAMP).
DISCUSSION
M. mcarthuri is well defined among related species by the mandibular dentition and by the
widely concave clypeal structure. Specimens from Morowa (WA) have the promesonotal shield
more distinctly indented between anterior and posterolateral mesonotal projections and the
cephalic index markedly lower (CI 114; 118 - 125 in others). Widely distributed throughout
central and southern Australia, with a single sample from Western Australia.
Meranoplus taurus Schödl sp.n.

(Figs. 50, 51, 76, 88)
0.36, PPLL 0.28, PPLH 0.43, PPI 64.71, PPDW 0.40, PT/PP 91.
Mandible with four teeth. In full face view anterior clypeal margin with few carinulae,
bifurcate with anterolateral corners produced into long acute teeth; the latter distinctly
surpassing the anterolateral frontal corners. Head almost square, only little wider than long (CI
110), preoccipital margin very shallowly concave. Frontal carinae markedly narrower than head
width (FI 144), sinuately converging towards clypeus. With head in profile antennal scrobe
reaching beyond middle of lateral side of head, weakly transversely carinulate with distinct
additional shagreening, distinctly defined posteriorly against remainder of head. Genae and
anterior ventrolateral section of head carinate, preoccipital lobes coarsely reticulate. Eyes
relatively large and rather flat (EL 0.27, REL 0.29, with 14 ommatidia in the longest row),
situated slightly in front of middle of lateral sides of head, the dorsal ocular margin touching the
ventral scrobal margin. Dorsal surface of head densely rugoreticulate, interspaces
microsculptured.
Promesonotal shield rugoreticulate, short and distinctly vaulted (PMI 142), somewhat
square with bluntly reduced projections. Propodeal spines medium sized in relation to small
body (PSL 0.35), situated rather high on lateral sides of the propodeal declivity, acute and in
dorsal view slightly arcuate and diverging.
Petiole in lateral view trapezoid, higher than long (PTI 66), dorsum convex and reticulate.
Postpetiole reticulate, nodular with triangular antero-basal tooth.
First gastral tergite regularly and densely microreticulate, basally with few additional
carinulae. Dorsal surfaces covered with short decumbent and longer erect thin whitish hairs.
Distinctly bicolored.
131 - 146, CI 107 - 115, CS 1.04 - 1.28, SL 0.59 - 0.65, SI1 48 - 53, SI2 51 - 56, PML 0.65 -
0.26 - 0.29. with 14 - 17 ommatidia in the longest row.
ETYMOLOGY
The name refers to the horn-shaped anterolateral clypeal projections ('taurus', Latin for
bull).
TYPE MATERIAL
Holotype worker, South Australia: 'S. Aust. William Creek 28°55'S 136°20'E 5-Sep-90, P
& I Gee.' (SAMA). Paratypes. 14 workers, same data as holotype; 9 workers, same locality data
but '28.55S 136.21E' (ANIC, NHMW, SAMA).
Northern Territory: 15 km SW Katherine, 7.iv.1978 (P.J.M. Greenslade); Tanami Rd 98

km W Stuart Hwy., 7.vii.2003 (R.R. Snelling); Tanami Rd 35 km W Stuart Hwy., 5.vii.2003
(R.R. Snelling). Queensland: Sandringham, 18.-25.ii.198 (P.J.M. Greenslade). New South
Wales: N. Nyngan 'Kapunda', 6.v.1987 (P.J.M. Greenslade); c. 45 km W by S of Wilcania,
11.xii.1973 (R.J. Kohout); 8 km S Conargo, 8.v.1979 (B.B. Lowery); Lightning Ridge,
10.v.1961 (B.B. Lowery). Victoria: Patho (H.A. Potter). South Australia: 2.5 km NW Ninety
Nine Dam, Anna Creek Stn., 1.-5.iii.1996 (coll. unknown); Top Camp Well, Hamilton Stn., 14.-
17.xi.1995 (coll. unknown); Innamincka, 9.5 km NW Patchawara bore, 10.-14.xi.1995 (coll.
unknown); 4.3 km NE Armistice bore, Anna Creek Stn., 1.-5.iii.1996 (coll. unknown); Cordillo
Downs Stn., 3.7 km W Frew well (coll. unknown); Murnpeowie, Mount Moolawatana, 16.-
17.xi.1994 (R. Tynan); Andamooka, 5.6 km WNW Netting Dam, 10.-14.xi.1996 (coll.
unknown); 3 km E Myra Mitta bore, 28.iv.1995 (T. Robinson); Nilpinna Stn., 4.1 km SSE,
Arckaringa WH, 15.-20.ix.1996 (coll. unknown); 5.2 km ESE Strangeways Springs, Stuart
Creek Stn., 25.-30.ix.1995 (coll. unknown); Stuart Creek Stn., 7.5 km E sheep camp WH (coll.
unknown); Anna Creek Stn., 28 km WNW Backadinna Dam, 2.-6.x.1995 (coll. unknown); Anna
Creek Stn., 15.6 km SW Backadinna Hill, 2.-6.x.1995 (coll. unknown); Peake Stn., 0.9 km NE
Four Hills Trig, 1.-5.iii.1996 (coll. unknown); 8 km NW Apollo Bore, iv.1995 (coll. unknown);
7.5 km NW Apollo Bore, iv.1995 (coll. unknown); Lake Eyre north, 20.vi.1991 (P. & I. Gee);
Screech-Owl Creek, 27.iv.1995 (coll. unknown); 15 km SE Balcanoona HS, 19.vii.1997 (coll.
unknown); 3.8 km ESE Coppertop Hill, Peak Stn., 1.-5.iii.1996 (coll. unknown); Woorana WH,
6.2 km S Macumba, 21.-25.ix.1996 (coll. unknown). Western Australia: 8 km E Gascoyne
junction, road turn-off, 4.vii.1981 (B. Heterick). (82 workers, 2 gynes, in ANIC, JDMP, LACM,
NHMW, SAMA).
DISCUSSION
M. taurus shows very little variability throughout its wide range. However, specimens from
a single sample taken at Katherine, Northern Territory are uniformly ferrugineous and have the
frontal striation more delicate. Southeastern-most samples, namely from Nyngan, Conargo,
Lightning Ridge and Patho differ by the clypeal bifurcation less acutely developed, by the
petiole more angulate in profile and by the markedly smaller eyes (REL 0.21 - 0.24), with 13 -
15 ommatidia in the longest row). Both forms are at least very closely related and are here
tentatively added to taurus. Further material will be needed to decide whether these samples
belong to taxa different from taurus. Occuring more or less sympatric with angustinodis and
variabilis in central and south-east Australia, except two remote collection sites in the far west
and one in the north, respectively. This small species is very distinctive by the large eyes, the
clypeal bifurcation and the square promesonotal shield with its reduced blunt projections.
ACKNOWLEDGEMENTS
I wish to thank all persons concerned with the loan and/or donation of material. These are
(in alphabetic order) Barry Bolton (BMNH), Michael Brancucci & Daniel Burckhardt (NHMB),
Chris Burwell (QMBA), Stefan P. Cover (MCZC), Greg Daniels (UQIC), Brian Hetterick
(JDMP), Terry Houston (WAMP), Archie McArthur (SAMA), Bernhard Merz (MHNG),
Thomas Pape & Bert Viklund (NHRS), Ted R. Schultz (USNM), Steve O. Shattuck (ANIC),
Roy R. Snelling (LACM), Ken Walker & Peter Lillywhite (MVMA), and Philip S. Ward
(University of California at Davis). Thanks are due to Alfred Buschingr (TU Darmstadt) and
Ajay Narendra, Macquarie University, Sydney for sending and donating additional material of
M. diversoides, Brian Hanich (WAMP) for information on uncertain locality data of M.
duyfkeni, and Alan N. Andersen (CSIRO, Darwin) for a constructive and helpful
correspondence. I am particularly indebted to R.R. Snelling (Los Angeles) and P.S. Ward
(University of California at Davis) for their invitation to publish this article herein. And, I thank
Matthias Buch, who prepared the magnificent water color illustrations of Meranoplus heads on
Plate 1.
LITERATURE CITED
Andersen, A.N. 2000. The Ants of Northern Australia. A guide to the monsoonal fauna: 106 pp,
CSIRO Publishing.
Bolton, B. 1981. A revision of the genera Meranoplus F. Smith, Dicroaspis Emery and
Calyptomyrmex Emery in the Ethiopian zoogeographical region. Bulletin of the British
Museum (Natural History). Entomology 42: 43-81.
Bolton, B. 1995. A New General Catalogue of the Ants of the World: 504 pp. - Harvard
University Press: Cambridge Massachusetts.
Forel, A. 1902. Fourmis nouvelles d`Australie. Revue Suisse de Zoologie 10: 405-548.
Forel, A. 1907. Formicidae. In Michaelsen, W & Hartmeyer, R. Die Fauna Südwest-
Australiens.Ergebnisse der Hamburger südwest-australischen Forschungsreise 1905 I: 263-
310.
Forel, A. 1910. Formicides Australiens. Revue Suisse de Zoologie 18: 1-94.
Forel, A. 1915. Results of Dr. E. Mjöberg's Swedish scientific expeditions to Australia, 1910-
1913. 2. Ameisen. Arkiv för Zoologie 9 (16): 1-119.
Schödl, S. 1998. Taxonomic revision of Oriental Meranoplus F. Smith, 1853 (Insecta:
Hymenoptera: Formicidae: Myrmicinae). Annalen des Naturhistorischen Museums in Wien
100B: 361-394.
Schödl, S. 2004. On the taxonomy of Meranoplus puryi Forel, 1902 and Meranoplus puryi
curvispina Forel, 1910 (Insecta: Hymenoptera: Formicidae). Annalen des Naturhistorischen
Museums in Wien 105B: 349-360.
Shattuck, S.O. 1999. Australian ants. Their biology and identification. Monographs on
Invertebrate Taxonomy 3: 226 pp.
Smith, F. 1867. Descriptions of new species of Cryptoceridae. Transactions of the
Entomological Society of London (3) 5: 523-528.
Taylor, R.W. 1987. A checklist of the ants of Australia, New Caledonia and New Zealand.
(Hymenoptera: Formicidae). CSIRO Division of Entomology Report 41: 1-92.
Taylor, R.W. 1990. The nomenclature and distribution of some Australian and New Caledonian
ants of the genus Meranoplus Fr. Smith (Hymenoptera: Formicidae: Myrmicinae). General
and Applied Entomology 22: 31-40.
Taylor, R.W. & Brown, D.R. 1985. Formicoidea, pp. 1-149. In: Walton, D.W. (ed.).
Zoological Catalogue of Australia 2 Hymenoptera: Formicoidea, Vespoidea and Sphecoidea:
381 pp. Canberra, Australian Government Publishing Service.
Figs. 1 - 7 Meranoplus workers, lateral aspects and dorsal views of promesonotal shield of (1, 2)
ajax, Maryinna Hill, SA, (3) ajax holotype, (4, 5) unicolor lectotype (pilosity omitted due to
abraded condition), (6, 7) snellingi.
Figs. 8 - 13 Meranoplus workers, lateral aspects and dorsal views of promesonotal shields of (8,
9) berrimah, (10, 11) duyfkeni lectotype, (12, 13) orientalis.
Figs. 14 - 19 Meranoplus workers, lateral aspects and dorsal views of promesonotal shields of
(14, 15) mars lectotype, (16, 17) wilsoni, (18, 19) tricuspidatus.
(20, 21) oxleyi lectotype, (22, 23) deserticola, (24, 25) crassispina.
(26, 27) convexius, (28, 29) naitsabes, (30, 31) digitatus, (32, 33) discalis, Silver Plains.
(34, 35) diversus, (36, 37) diversoides, (38, 39) christinae, Merigol.
(40, 41) occidentalis, (42, 43) angustinodis, (44, 45) variabilis.
(46, 47) arcuatus, (48, 49) mcarthuri, (50, 51) taurus
Figs. 52 - 64 Meranoplus workers, heads of (52) ajax, Ungarinna, (53) ajax holotype, (54)
unicolor lectotype, (55) snellingi, (56) berrimah, (57) duyfkeni lectotype, (58) orientalis, (59)
mars lectotype, (60) wilsoni, (61) tricuspidatus, (62) oxleyi, (63) deserticola, (64) crassispina.
Figs. 65 - 77 Meranoplus workers, heads of (65) convexius, (66) naitsabes, (67) digitatus, (68)
discalis, (69) diversus, (70) diversoides, (71) christinae, (72) occidentalis, (73) angustinodis,
(74) variabilis, (75) mcarthuri, (76) taurus, (77) arcuatus.
Figs. 78 - 83 Distribution of M. diversus-group species, as known from studied material.

Figs. 84 - 89 Distribution of M. diversus-group species, as known from studied material.

Figs. 90 - 95 Historic type label data of Meranoplus species (90) ajax, holotype, (91) unicolor,
lectotype, (92) duyfkeni, lectotype, (93) oxleyi, lectotype, (94) mars, lectotype, (95) diversus,
holotype.
Plate 1 Heads of Meranoplus workers, showing surface structure and pilosity (A) snellingi, (B)
christinae, (C) occidentalis.
Schultz, T. R. 2007. The fungus-growing ant genus Apterostigma in Dominican amber, pp. 425-
436. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics
THE FUNGUS-GROWING ANT GENUS APTEROSTIGMA IN DOMINICAN AMBER
Ted R. Schultz
Department of Entomology, MRC 188, P.O. Box 37012
National Museum of Natural History
Smithsonian Institution
Washington, DC 20013-7012, U.S.A.
schultzt@si.edu
ABSTRACT
The first fossil species of the fungus-growing ant genus Apterostigma (Myrmicinae: Attini) are
described from Dominican amber. Two new species are described, and the likely placement of
each within the genus is discussed. The phylogenetic position of Apterostigma within the Attini
is summarized, as is the evolutionary history of the transition from the cultivation of
leucocoprineaceous fungi (the ancestral condition) to the cultivation of pterulaceous fungi (the
derived condition) by different lineages of Apterostigma species. I conclude by speculating on
the possible implications of the fossil species for understanding this transition, which is unique
within the fungus-growing ants.
Key words: Hymenoptera, Formicidae, Attini, Apterostigma, fungus-growing ants, fossils,

Dominican amber.
INTRODUCTION
Fungus-growing ants in the tribe Attini are one of only a few animal groups that practice true
agriculture. Because, so far as is known, this fungicultural life-history strategy is entirely unique
in ants and, because it parallels human agriculture in many remarkable ways (Schultz et al.,
2005), the origin and evolution of the fungus-growing behavior has been the subject of recent
intensive study (e.g., Chapela et al., 1994; Hinkle et al., 1994; Schultz & Meier, 1995; Wetterer
et al., 1998; Mueller et al., 1998; Mueller et al., 2001). Various lines of evidence suggest that
the fungus-growing behavior arose a single time in an ancestral attine ant sometime between 45-
65 million years ago on the South American continent (Mueller et al., 2001). Perhaps the most
compelling pattern supporting this hypothesis is the geographic distribution, extant and
historical, of the Attini: fungus-growing ants occur only in North, Central, and South America,
and on some Caribbean islands (Kempf, 1972; Wilson, 1988; Brandão, 1991). They also occur
as fossils in Dominican amber (Oligo-Miocene, ~20 mya), but are unknown in Old World
fossils, including Baltic amber (Eocene, ~45 mya). To date, three attine ant species have been
described from Dominican amber: Trachymyrmex primaevus (Baroni Urbani, 1980),
Cyphomyrmex maya, and Cyphomyrmex taino (de Andrade, 2003). In addition, Brown (1973)
refers to possible Mycetosoritis males in Chiapas amber (Oligo-Miocene, ~20 mya), Wilson
(1985) refers to Cyphomyrmex in Dominican amber, and Baroni Urbani (1995) refers to
Apterostigma and Cyphomyrmex in Dominican amber.
Here I describe two fossil species of the fungus-growing ant genus Apterostigma preserved
in Dominican amber.
Amber specimens were studied and measured using a Leica Wild M10 stereomicroscope.
Illustrations were executed by USNM scientific illustrator V. Malikul, who directly examined
the specimens using a Nikon SMZ 1500 stereomicroscope. Specimens were photographed using
a JVC KY-F70B video camera mounted on a Leica M420 microscope and attached to an IBM
Intellistation M Pro computer, on which composite images were assembled using Auto-Montage
Version 3.04 software (Synoptics Ltd.). The following are abbreviated in the text: American
Museum of Natural History (AMNH), National Museum of Natural History (USNM), and the
Smithsonian Institution Ant Database (SIANT).
SPECIES DESCRIPTIONS
Apterostigma electropilosum, new species

(Figures 1 and 2)
HOLOTYPE (worker): AMBER: Oligo-Miocene; Dominican Republic; AMNH no. DR-

14-984; no other locality data. USNM SIANT database specimen reference number 00443127.
Deposited in AMNH.
Measurements: head length = 1.06 mm; head width (not including eyes) = 0.68 mm; scape
length = 0.99 mm; Weber’s length = 1.37 mm; metafemur length = 1.30 mm.
DESCRIPTION: In full-face view, head elongate, evenly rounded at the posterolateral

corners and slightly concave at the median posterior border. As in all Apterostigma species,
occiput drawn out posteriorly into a “neck” or “collar”; in this species, the collar short, not
expanded posteriorly, and lacking integumental rugosities. Frontal carinae strongly produced,
extending posterad to well past the level of the eyes. Vertexal carinae not visible, possibly
Schultz: Apterostigma in Amber 427
obscured by the amber matrix and by the unusual texture of the integument, discussed below.
Vestigial preocular carinae extending posterad to top of eye. Frontal lobes evenly rounded.
Antennae with the typical attine number of eleven segments, the apical antennal segment 2.5× as
long as the subapical segment, the subapical segment subequal in length to the pedicel. Clypeus
clearly with a smooth, shining anterior border, produced medially into a broad, V-shaped angle.
Mandibles with eleven teeth, all unusually sharp and showing no wear, teeth decreasing evenly
in size toward the mandibular base. Lateral corners of the hypostoma produced into blunt,
rounded “hypostomal teeth.” Eyes with six ommatidia across at narrowest circumference
(subparallel to the transverse plane) and ten ommatidia across at greatest circumference
(subparallel to the longitudinal axis). Eyes prominent and hemispherical, each hemisphere
interrupted from behind (i.e., ventrally) by a straight line of occluding integument.
Promesonotum with a pair of longitudinal, uniformly low carinae. Anterolateral mesonotal
carinae apparently absent and humeral prominences vestigial. Posterior mesonotum with a pair
of reduced, very short, almost vestigial carinae (“arista metanotal” of Lattke, 1997), best
described as vestigial tubercles. Ventral mesopleural carinae present and complete, but not
lamellate. Meso- and metacoxae of typical form, without lamellate or keel-like carinae.
Propodeum carinate on the basal and declivous faces, the carinae essentially interrupted
extensions of the promesonotal pair. Propodeal spines or tubercles absent, propodeal shoulder
evenly rounded.
Petiole short and thick, entirely lacking a node; anterior ventral tooth apparently absent.
Viewed dorsally, postpetiole roughly as long as broad, and about half as broad anteriorly as
posteriorly. First gastral (fourth abdominal) tergite strongly laterally carinate for at least three-
fourths of its length.
Body covered with long, fine, simple, erect setae with a maximum length of 0.15 mm.
Integument with a mottled, coarsely granulate texture that interferes with the study of
microsculpture. On close inspection, this texture apparently caused by shining bubbles, possibly
of an oily secretion, that arose from punctures in the integument while the amber was in a
semiliquid state. A similar phenomenon was observed in a modern Apterostigma specimen
embedded in Canada balsam by N. A. Weber.
The shining clypeal border places this species in the pilosum group (sensu Lattke, 1997),
one of two informal subgeneric groupings. The specimen keys out to A. wasmanni (southern
Brazil) in Lattke’s (1997) key, but differs from that species in being less strongly sculptured and
in having the median clypeal angle more pronounced. In habitus it is similar to many small
cryptic species that key out to Lattke’s unrevised “pilosum complex” of species, but is distinct
among them in combining evenly rounded frontal lobes, eleven mandibular teeth, and a medially
angulate clypeus.
ETYMOLOGY: The species name indicates that this is a species of the pilosum group, and
probably of the pilosum complex, in amber (electrum).
Apterostigma eowilsoni, new species

(Figures 3, 4, and 5)
HOLOTYPE (worker): AMBER: Oligo-Miocene; Dominican Republic; AMNH no. DR-

16-292; no other locality data. USNM SIANT database specimen reference number 00443150.
Deposited in AMNH.
Measurements: head length = 0.99 mm; head width (not including eyes) = 0.68 mm; scape
length = 0.99 mm; Weber’s length = 1.44 mm; metafemur length = 1.37 mm.
DESCRIPTION: In full-face view, head bluntly angled at posterolateral corners and again
at vertexal carinae, and slightly indented medially. Occipital “collar” short and not expanded
428
Memoirs of the American Entomological Institute, Volume 80
Fig. 1a. Apterostigma electropilosum, full-face view. Fig. 1b. Apterostigma electropilosum, lateral view.
Schultz: Apterostigma in Amber
Fig. 2a. Apterostigma electropilosum, full-face view. Fig. 2b. Apterostigma electropilosum, lateral view.
429
Fig. 3a. Apterostigma eowilsoni, lateral view.
Fig. 3b. Apterostigma eowilsoni, full-face view.

Fig. 4a. Apterostigma eowilsoni, lateral view.
Fig. 4b. Apterostigma eowilsoni, full-face view.

posteriorly; collar integument with a series of longitudinal rugae. Frontal carinae strongly
produced, extending posterad past the level of the eyes. Vertexal carinae strong, preocular
carinae weak. Frontal lobes evenly rounded. Antennae with the typical attine number of eleven
segments, the apical segment 2.25× as long as the subapical segment, the subapical segment
two-thirds the length of the pedicel. Clypeal border broadly convex, and, as far as can be seen,
with only an exceedingly thin strip of cuticle, forming the anterior edge of the clypeus, smooth
and of a darker color than the rest of the clypeal integument, the rest identical to the integument
of the rest of the head. Mandibles apparently with eleven sharp teeth, the teeth arranged in a
curious pattern heretofore unencountered in the genus: reckoning from the apex, the sixth and
eighth teeth on the left mandible and the fourth and seventh teeth on the right mandible distinctly
larger than the rest; the smaller teeth in between of various sizes, not decreasing in size toward
the mandibular base. Lateral corners of the hypostoma produced into blunt, rounded hypostomal
teeth. In frontal view the eyes typical, in lateral view the eyes half-hemispheres truncated from
behind by integument, but in dorsal view the eyes forward-directed subconical hemispheres
mounted on lobes or tubercles that project, perpendicular and earlike, from the sides of the head.
As far as can be determined given the complications of observing through the amber matrix, eye
width approximately nine ommatidia across the transverse circumference and approximately
twelve ommatidia across the longitudinal circumference.
Promesonotum with a pair of strong longitudinal carinae, in lateral view the carinae ending
anteriorly in an abrupt vertical wall above the promesonotal junction. Anterolateral mesonotal
carinae strong and humeral prominences apparently absent. Posterior mesonotum (“metanoto” of
Lattke, 1997) without sculpture. Ventral mesopleural carina present and complete, but not
lamellate. Meso- and metacoxae of typical form, without lamellate or keel-like carinae.
Propodeum carinate on the basal face, weakly carinate on the declivous face, and lacking
propodeal spines. Propodeal shoulder evenly rounded, the propodeal spiracles directed posterad
and mounted on tubercles. Ventral surfaces of the mesopleura covered with six small spherical
objects, apparently clusters of minute bubbles.
Petiole with an elongate peduncle bearing an anterior ventral tooth and a low but distinct
node. What appears to be a second, posterior ventral tooth is actually debris in the amber matrix.
Viewed dorsally, the postpetiole approximately 1.3X broader than long, subtriangular in shape,
and about half as broad anteriorly than posteriorly. Entire length of the first gastral (fourth
abdominal) tergite strongly laterally carinate. Body covered with long, fine, simple, erect setae
with a maximum length of 0.15 mm.
By Lattke’s (1997) primary criterion of the presence of a smooth and shining clypeal border,
A. eowilsoni belongs to the pilosum group. The anterior clypeal border in this species is,
however, extremely reduced and thus represents a credible intermediate in the morphocline
spanning the typical (and presumably plesiomorphic; Lattke, 1999) state in the pilosum group
and the derived state (clypeal border absent) in the auriculatum group. Of the species known to
me, the reduced clypeal border in A. eowilsoni most resembles that of an undescribed Costa
Rican species that keys out to the unrevised “pilosum complex” of species in Lattke’s (1997)
key. Certainly the most striking character of A. eowilsoni is the remarkably protruding eyes,
which are similar in form to—but far more developed than—the eyes found in the auriculatum-
group species A. pariense (Venezuela and Bolivia; specimens examined) and A. reburrum
(Colombia; specimens not seen), which also have their eyes mounted on markedly protruding
tubercles. Apterostigma eowilsoni clearly possessed excellent stereoscopic forward vision,
limited lateral vision, and no rearward vision.
ETYMOLOGY: It gives me great pleasure to name this striking and possibly

phylogenetically important fossil fungus-growing ant after E.O. Wilson, in celebration of his
long career of myrmecological discovery.
EVOLUTIONARY IMPLICATIONS
Current evidence suggests that the tribe Attini is divided into two major clades, the result of an
ancient, phylogenetically basal divergence: (i) Kusnezov’s (1963) “Paleoattini,” consisting of the
three genera Myrmicocrypta, Mycocepurus, and Apterostigma, and (ii) the “Neoattini,”
consisting of all other attine genera (Kusnezov, 1963; Schultz & Meier, 1995; Schultz,
unpublished). The Paleoattini are characterized by a number of compelling synapomorphies,
including: (i) short antennal pedicel in the male caste; (ii) the presence of a unique “fenestra”
(clear spot) in the wings of gynes (Emery, 1913); (iii) paired hypostomal teeth in workers and
gynes (secondarily lost in some species of Apterostigma and Mycocepurus); and (iv) the absence
of an inferior pronotal angle or tooth. (The monophyly of the Paleoattini does not preclude the
possibility that some attine species, known or as yet unknown and probably fitting our current
concept of Myrmicocrypta, may belong to earlier diverging lineages.) Within the Paleoattini,
ants of the genus Apterostigma are by far the most evolutionarily derived. This genus is
characterized by a number of synapomorphies, including: (i) a reduced palpal formula of 3, 2
(Kusnezov, 1953), in contrast to the formula of 4, 2 present in all other Attini except the
Argentinean social parasite Pseudoatta argentina; (ii) unusual wing venation, including loss of
the rsf1 vein; (iii) the wings of gynes with a distinctively positioned, reflective fenestrum, and of
both sexes with a smoky coloration; (iv) pilosity of fine, simple, erect setae, in some species
short and in most species long and flexuous; and (v) cytochrome oxidase mtDNA sequence with
a unique amino acid insertion.
Most remarkably, with regard to symbiont association a subset of Apterostigma species are
the most highly derived of all attine ants. All other attine ants, including the leaf-cutters in the
genera Atta and Acromyrmex, cultivate “parasol mushrooms” in the single tribe Leucocoprineae
(family Lepiotaceae), consisting of Leucocoprinus and Leucoagaricus, as well as a few species
currently assigned to Lepiota (Mueller et al. 1998; Johnson 1999). Apterostigma auriculatum
and, presumably, other species in the auriculatum group retain this ancestral attine association
with the Leucocoprineae. In contrast, all Apterostigma pilosum-group species (sensu Lattke,
1997) studied to date cultivate “coral fungi” closely related to the genera Pterula and Deflexula
(Pterulaceae), which are quite distantly related to the Leucocoprineae (Munkacsi and
McLaughlin 2001; Muncaksi et al., 2004). A molecular phylogeny for Apterostigma (Villesen et
al., 2004; summarized in Figure 6) indicates that the earliest Apterostigma species cultivated
leucocoprineaceous fungi. Sometime after the origin of Apterostigma, in an unparalleled event in
the history of the Attini, an Apterostigma species host-switched to a radically different fungal
cultivar in the Pterulaceae, which all of its descendants (the majority of extant Apterostigma
species, all derived) have been cultivating ever since. Moreover, it appears that the first
Pterulaceae-cultivating Apterostigma species cultivated the distinctive “G4” fungal clade,
whereas later in evolutionary time a pilosum-group Apterostigma species host-switched to the
“G2” fungal clade, a behavior inherited by all of its descendant species. Unlike G4-cultivating
species, G2-cultivating Apterostigma ants (including A. dentigerum, A. dorotheae, A. collare,
and a subset of Lattke’s [1997] “pilosum complex”) weave the elongate aerial hyphae of their
fungal cultivars into protective tent-like veils that entirely surround their gardens.
Fig. 5. Apterostigma eowilsoni, posterior view of head showing protruding eyes.
Fig. 6. A phylogeny of Apterostigma based on Villesen et al. (2004) indicating known cultivar
associations and the speculative positions of A. megacephala, A. eowilsoni, and A.
electropilosum.
Recently, Lattke (1999) described Apterostigma megacephala, a bizarre species that

combines the smooth and shining clypeal border that is the primary character of the pilosum
group with the strongly sculptured integument that is usually associated with the auriculatum
group. The first Apterostigma very probably also combined these traits. A phylogeny based on
morphological character data, incorporating A. megacephala as well as the two fossil species
recorded here, is necessary for accurately reconstructing the phylogenetic positions of the fossil
species, speculatively reconstructed in Figure 6. This information, combined with an expanded
molecular phylogeny, will permit the assignment of minimum dates to particular nodes, which
will in turn provide information critical for estimating the dates of origin of (i) the genus
Apterostigma; (ii) the auriculatum group; (iii) the shift from lepiotaceous to pterulaceous
cultivars; and (iv) the shift from “G4” to “G2” pterulaceous cultivars.
ACKNOWLEDGEMENTS
This work was supported by National Science Foundation grant IRCEB DEB 0110073. I am
grateful to D. Grimaldi for discovering and preparing these amber fossil specimens and for
bringing them to my attention; to V. Malikul for producing the illustrations; to C. Marshall for
helping to revise the illustrations; to M. Engel for instructing me in the fine art of amber study;
to R. Wilson for preparing the figures; to J. Sosa-Calvo for help with Auto-Montage and for
discovering Emery’s reference to paleoattine fenestrae; and to R.R. Snelling and P.S. Ward, both
for inviting me to contribute to this volume and for useful comments on the manuscript.
Most of all, I am grateful to E.O. Wilson for inspiring me to become a myrmecologist. In
particular, my life was transformed by his 1971 book The Insect Societies, in which appears the
following passage: “So long as the evolutionary origin of the Attini remains a mystery, we
cannot hope to gain any convincing insight into the evolutionary beginnings of fungus
gardening.”
LITERATURE CITED
Baroni Urbani, C. 1980. First description of fossil gardening ants (Amber Collection Stuttgart
and Natural History Museum Basel; Hymenoptera: Formicidae. I: Attini). Stuttgarter
Beiträge zur Naturkunde, B 54: 1-13.
Baroni Urbani, C. 1995. Invasion and extinction in the West Indian ant fauna revised: the
example of Pheidole (Amber Collection Stuttgart: Hymenoptera, Formicidae. VIII:
Myrmicinae, partim). Stuttgarter Beiträge zur Naturkunde, B 222: 1-29.
Brandão, C.R.F. 1991. Adendos ao Catálago abreviado das formigas da Região Neotropical
(Hymenoptera: Formicidae). Revista Brasileira de Entomologia 35: 319-412.
Brown, W.L., Jr. 1973. A comparison of the Hylean and Congo-West African rain forest ant
faunas. Pp. 161-185 in Tropical Forest Ecosystems in Africa and South America, a
Comparative Review, B.J. Meggers, E.S. Ayensu, and W.D. Duckworth, editors.
Washington, DC: Smithsonian Institution Press.
Chapela, I.H., Rehner, S., Schultz, T.R. & Mueller, U. 1994. Evolutionary history of the
symbiosis between fungus-growing ants and their fungi. Science 266: 1691-1694.
de Andrade, M.L. 2003. First descriptions of two new amber species of Cyphomyrmex from
Mexico and the Dominican Republic (Hymenoptera: Formicidae). Beiträge zur Entomologie
53: 131-139.
Emery, C. 1913. Études sur les Myrmicinae V. Les genres des Attini; descriptions de nouvelles
formes de Mycocepurus et de Myrmicocrypta. Annales de la Société Entomologique de
Belgique 57: 250-262.
Hinkle, G., Wetterer, J.K., Schultz, T.R. & Sogin, M.L. 1994. Phylogeny of the attine ant
fungi based on analysis of small subunit ribosomal RNA gene sequences. Science 266:
1695-1697.
Johnson, J. 1999. Phylogenetic relationships within Lepiota sensu lato based on morphological
and molecular data. Mycologia 91: 443-458.
Kempf, W.W. 1972. Catálago abreviado das formigas da Região Neotropical (Hymenoptera:
Formicidae). Studia Entomologia 15: 3-344.
Kusnezov, N. 1953. Phyletische Bedeutung der Maxillar- und Labialtaster der Ameisen.
Zoologischer Anzeiger 153: 28-38.
Kusnezov, N. 1963. Zoogeogafia de las hormigas in Sudamerica. Acta Zoologica Lilloana 19:
25-186.
Lattke, J.E. 1997. Revisión del género Apterostigma Mayr (Hymenoptera: Formicidae).
Arquivos de Zoologia 34(5): 121-221.
Lattke, J.E. 1999. A new species of fungus-growing ant and its implications for attine
phylogeny (Hymenoptera: Formicidae). Systematic Entomology 24: 1-6.
Mueller, U.G., Rehner, S.A. & Schultz, T.R. 1998. The evolution of agriculture in ants.
Science 281: 2034-2038.
Mueller, U.G., Schultz, T.R., Currie, C.R., Adams, R.M.M. & Malloch, D. 2001. The origin
of the attine ant-fungus mutualism. Quarterly Review of Biology 76: 169-197.
Munkacsi, A. & McLaughlin, D.J. 2001. Evolutionary relationships of Pterula and Deflexula
within Agaricales sensu stricto and their relationships with the tricholomataceous attine
fungi. Abstract, 2001 Mycological Society of America Meeting, August 25-29, 2001, Salt
Lake City, Utah.
Munkacsi, A.B., Pan, J.J., Villesen, P., Mueller, U.G., Blackwell, M. & McLaughlin, D.J.
2004. Convergent coevolution in the domestication of coral mushrooms by fungus-growing
ants. Proceedings of the Royal Society (London), Series B 271: 1777-1782.
Schultz, T.R. & Meier, R. 1995. A phylogenetic analysis of the fungus-growing ants
(Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae.
Schultz, T.R., Mueller, U.G., Currie, C.R. & Rehner, S.A. 2005. Reciprocal illumination: A
comparison of agriculture in humans and in fungus-growing ants. Pp. 149-190 in Insect-
Fungal Associations: Ecology and Evolution, F. E. Vega and M. Blackwell, editors. New
York: Oxford University Press.
Synoptics Ltd. 2003. Auto-Montage Version 3.04 (computer software). United Kingdom:
Cambridge.
Villesen, P., Mueller, U.G., Schultz, T.R., Adams, R.M.M. & Bouck, M.C. 2004. Evolution
of ant-cultivar specialization and cultivar switching in Apterostigma fungus-growing ants.
Evolution 58: 2252-2265.
Wetterer, J.K., Schultz, T.R. & Meier,R. 1998. Phylogeny of fungus-growing ants (tribe
Attini) based on mtDNA sequence and morphology. Molecular Phylogenetics and Evolution
9: 42-47.
Wilson, E.O. 1971. The Insect Societies. Cambridge, Massachusetts: Harvard University Press.
Wilson, E.O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 1. Two new
myrmicine genera and an aberrant Pheidole. Psyche 92: 1-9.
Wilson, E.O. 1988. The biogeography of the West Indian ants (Hymenoptera: Formicidae). Pp.
214-230 in Zoogeography of Caribbean Insects, J.K. Liebherr, editor. Ithaca, New York:
Cornell University Press.
Shattuck, S. O., Barnett, N. J. 2007. Revision of the ant genus Mayriella, pp. 437-458. In
REVISION OF THE ANT GENUS MAYRIELLA
Steven O. Shattuck and Natalie J. Barnett

CSIRO Entomology
P. O. Box 1700
Canberra, ACT 2601, Australia
Steve.Shattuck@csiro.au
ABSTRACT
The ant genus Mayriella is revised at the species level. The genus contains seven species: M.
abstinens Forel (north-east Australia and New Zealand), M. ebbei Shattuck & Barnett (new
species) (south-east Australia), M. granulata (Vietnam), M. overbecki Viehmeyer (removed
from synonymy with M. abstinens) (east-central Australia), M. sharpi Shattuck & Barnett (new
species) (eastern Papua New Guinea), M. spinosior Wheeler (north-east Australia) and M.
transfuga Baroni Urbani (Nepal and India east to Hong Kong and the Philippines and south to
Borneo and Java, Indonesia). M. hackeri Wheeler and M. venustula Wheeler are synonymized
with M. abstinens. All species occur in rainforest or wet sclerophyll forests and nest in soil,
often under rocks, or in rotten wood on the ground.
Key words: Hymenoptera, Formicidae, Myrmicinae, Mayriella, taxonomy, new species,

synonymy, Southeast Asia, Australia.
INTRODUCTION
Mayriella Forel is an Indo-Australian ant genus belonging to the subfamily Myrmicinae. The
genus was established by Forel (1902) for the newly described species M. abstinens from
Mackay, Queensland, Australia. Viehmeyer (1925) added a second species, M. overbecki, from
material collected at Trial Bay, New South Wales. Wheeler (1935) revised Mayriella based on
holdings in the Museum of Comparative Zoology, Harvard University. He synonymized M.
overbecki with M. abstinens and established 3 new taxa, M. abstinens hackeri, M. abstinens
venustula and M. spinosior. These taxa were based on material originating from Queensland.
Unfortunately, Wheeler had access to only limited material including four workers and six
queens of M. a. hackeri, single workers of M. a. venustula and M. spinosior, and apparently
only a single syntype worker of M. abstinens.
Mayriella has since been found outside Australia. Taylor (1961) reported an introduced
population of M. abstinens in New Zealand. He also gave the first detailed biological
observations on these ants and questioned the validity of the forms described by Wheeler
(1935). Taylor (1961) did not, however, formally synonymize Wheeler's names. Baroni Urbani
(1977) greatly expanded the known range of Mayriella when he described M. transfuga from
Nepal and Bhutan. This was the first published record of the genus outside the Australian region
(although material had been present in collections since the mid-1950s). More recently, Dlussky
and Radchenko (1990) added an additional species from South-east Asia, M. granulata, a
species very similar to the wide-spread M. transfuga.
Mayriella was re-examined by Taylor (1991). He found the genus to be "intractable at
species level" and containing anything from a single variable species to three or more sibling
species in Australia. Taylor (1991) did not propose any new species-level synonymies as he
found the available material showed extensive variation in size, propodeal spination, sculpturing
and color, and because some series contained dealate queens while others contained apparent
ergatoid queens. Taylor proposed the use of an "aggregate" name for the Australian species of
Mayriella, and raised M. abstinens hackeri and M. a. venustula to full species rank "for ease of
indexing." Finally, Taylor presented a detailed list of known collection localities for all species
of the genus based on material in the Australian National Insect Collection and the Queensland
Museum. More recently, Shattuck (1999) provided an overview of the genus in Australia,
providing notes on biology and distribution.
During the present study, Mayriella was found to contain seven valid species. Five of these
(abstinens, granulata, overbecki, spinosior, transfuga) were previously described, while two
(ebbei, sharpi) were undescribed. Two of the previously described taxa (hackeri, venustula)
were found to be conspecific with M. abstinens. Five of these species pose few taxonomic
problems as they are easily diagnosed and show minimal morphological variation. However, M.
abstinens (as defined here) possesses considerable variation in body color, propodeal spine
development and, to a lesser extend, sculpture. This variation is considered intraspecific because
(1) several nest series contain both color forms, (2) other nest series show intermediate color
patterns, (3) propodeal spine morphology varies independently of color and (4) no other
characters could be found to indicate two separate species are involved. See M. abstinens below
for further details. It should also be noted that Taylor’s (1961, 1991) suggestion that ergatoid
(worker-like) queens are present in this species could not be confirmed. Finally, M. granulata
shows only minor differences from M. transfuga and may prove to be conspecific with it.
Species of Mayriella show strong preference for moist, forested areas. Most specimens have
been collected from rainforest or wet sclerophyll woodlands with relatively few collections from
dry sclerophyll or scrub habitats (and these are restricted to Australia). Nests are normally found
in soil under stones or with a small mound of loose dirt at the entrance. They have also been
found nesting in rotten logs and occasionally arboreally. The number of workers in nests
Shattuck & Barnett: Revision of Mayriella 439
averages between 50 and 100 and about one-half of the nest series containing queens contain
more than one, suggesting that species are polygynous. Foraging workers are most commonly
collected from litter samples. Nests appear to be locally abundant although patchily distributed
on a larger scale. For example, in the Brindabella Ranges near Canberra, ACT, M. ebbei is
infrequently encountered and is known from only a few localities. However, near Baroomba
Rocks (located on the eastern slope of the Brindabella Ranges) M. ebbei occurs commonly
under rocks in an area covering several hundred square meters. This population appears limited
to this small area as nests could not be found in areas with similar geography and vegetation
located several kilometers away. However, observations are limited and additional data will be
required to determine if these habits are widespread and/or occur in other species. Additionally,
while Taylor (1961) made similar observations for M. abstinens, these were based on the
introduced population found in New Zealand and should be confirmed for naturally occurring
populations of the genus.
Placement of Mayriella in the subfamily Myrmicinae
Forel (1902) placed Mayriella in the myrmicine tribe Meranoplini. This treatment was
followed by Emery (1922) and Wheeler (1935). Wheeler (1935) supported this placement by
discussing numerous characters which Mayriella shared with Meranoplus, Willowsiella and
Romblonella. However, Wheeler also noted differences among these taxa which suggested they
may not be as closely related as previously thought. More recent treatments of the tribal
classification of the Myrmicinae have either followed Forel (Wheeler & Wheeler, 1973, 1985),
or not placed the genus within a specific tribe (Hölldobler & Wilson, 1990), or only tentatively
placed it (Bolton, 1994, in the tribe Stenammini). An examination of the sting apparatus of M.
ebbei (reported as M. abstinens) by Kugler (1997) found similarities with the genus Proatta,
suggesting a close relationship and thus a placement in the Stenammini, supporting the treatment
of the genus by Bolton (1994). Finally, Bolton’s (2003) recent review of the higher level
classification of the entire family has moved Mayriella to the Carebara genus group of the tribe
Solenopsidini and includes detailed lists of characters supporting this placement. Bolton also
provides a more detailed overview of the literature concerning this genus than offered here.
Measurements
The following measurements were recorded in millimeters. Indices were calculated using
the measurements indicated.
CI cephalic index: HW/HL

HL maximum head length in full face view, measured from the anterior clypeal margin
(excluding the projecting clypeal teeth) to the midpoint of a line drawn across the
posterior margin of the head
HTL maximum length of hind tibia, excluding the proximal part of the articulation which is
received into the distal end of the hind femur
HW maximum head width in full face view, excluding eyes.
ML mesosomal length measured from the point at which the pronotum meets the cervical
shield to the posterior base of the metapleuron
PW maximum pronotal width in dorsal view
SI scape index: SL/HW
SL length of the scape (first antennal segment) excluding the basal constriction and condylar
bulb
Depositories
ANIC – Australian National Insect Collection, Canberra, ACT, Australia

BMNH – The Natural History Museum, London, United Kingdom
MCZC – Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts, USA.
MHNG – Muséum d'Histoire Naturelle, Geneva, Switzerland
TERC CSIRO Tropical Ecology Research Centre (A. N. Andersen Collection),
Darwin, Northern Territory, Australia
NHMB – Naturhistorisches Museum, Basel, Switzerland
UASK – Institue of Zoology, Ukrainian Academy of Science, Ukraine
Mayriella Forel, 1902

Figures 1-25
Mayriella Forel, 1902: 425.
Type species. Mayriella abstinens Forel, 1902, by monotypy.
Diagnosis. Workers of Mayriella may be separated from other members of the Myrmicinae
by the presence of a 10-segmented antenna with a 2-segmented club; well developed antennal
scrobes; the clypeus with the mediolateral regions concave and the anterior margin with a
bidentate process (these structures combining to form an extension of the scrobes which receive
the terminal segments of the antennae when the antennae are retracted); and elongate compound
eyes which are narrowed anteroventrally. These characters, especially the shape of the eyes and
the configuration of the clypeus, are distinctive within the subfamily and it is unlikely these ants
will be confused with close relatives.
Key to Species of Mayriella (workers)
1 In dorsal view, anterior region of outer margin of postpetiole expanded laterally compared
to posterior regions so its overall outline is trapezoidal (Fig. 9) ............................................ 2
- In dorsal view, outer margins of postpetiole approximately the same width anteriorly as
posteriorly so its overall outline is cylindrical or slightly barrel-shaped (Fig. 5) ................... 3
2 Dorsum of postpetiole with more than four erect hairs and dorsum of gaster with numerous
erect hairs; dorsum of petiole in lateral profile uniformly convex and without distinct,
separate dorsal and posterior faces (Fig. 8) ......................................ebbei Shattuck & Barnett
- Dorsum of postpetiole with at most four erect hairs and dorsum of gaster generally lacking
erect hairs (rarely hairs present); dorsum of petiole in lateral profile angular and with
distinct, separate dorsal and posterior faces (Fig. 22)..................................spinosior Wheeler
3 Occurring in south-east Asia (Fig. 24) ................................................................................... 4

- Occurring in Papua New Guinea or Australia ........................................................................ 5
4 Propodeal spines relatively thin and appearing long, length greater than 1.5 times basal
width (Fig. 25) .................................................................................. transfuga Baroni Urbani
- Propodeal spines relatively broad and appearing short, length less than basal width (Fig. 12)
...........................................................................................granulata Dlussky and Radchenko
5 Ventral extension of compound eye terminating in a sharp point (Fig. 19); head
approximately square (CI ca. 1.0) (Papua New Guinea) ................ sharpi Shattuck & Barnett
- Ventral extension of compound eye rounded below (Fig. 15); head longer than wide (CI <
0.98) (Australia)...................................................................................................................... 6
6 Sculpturing in posterior section of antennal scrobe well developed and distinct; sculpturing
on dorsal surface of mesosoma consisting of large, closely spaced (separated by less than
their diameter) pits (Fig. 4); dorsum of petiole in lateral profile strongly angular, the
posterior face relatively short (Fig. 3) ..............................................................abstinens Forel
- Sculpturing in posterior section of antennal scrobe at most weakly developed and always
indistinct; sculpturing on dorsal mesosoma consisting of small, widely spaced (separated by
greater than their diameter) pits (Fig. 16); dorsum of petiole in lateral profile broadly
rounded, the posterior face relatively long (Fig. 15) .............................. overbecki Viehmeyer
Mayriella abstinens Forel, 1902

Figures 1-5
Mayriella abstinens Forel, 1902: 452.

Mayriella abstinens hackeri Wheeler, 1935: 157 syn. n.
Mayriella abstinens venustula Wheeler, 1935: 158 syn. n.
Mayriella hackeri; Taylor, 1991: 602.
Mayriella venustula; Taylor, 1991: 602.
TYPE MATERIAL
M. abstinens: Worker syntypes (1 in MCZC, examined, additional in MHNG, not examined)

from Mackay, Queensland. M. hackeri: Worker and queen syntypes (four workers and six
queens in MCZC, examined) from nr. Brisbane, Queensland. M. venustula: Holotype worker
(MCZC, examined) from Tambourine Mt., Queensland.
ADDITIONAL MATERIAL
(In ANIC unless otherwise noted). Australia, New South Wales: 10km S Mangrove Mt.
(Lowery, B.B.); 15km E Legume (Greenslade, P.J.M.); 15mi N Lismore, Whian Whian State
Forest (Lowery, B.B.); 1mi E Murwillumbah (Lowery, B.B.); 1mi E Murwillumbah Rail Station
(Lowery, B.B.); 20km W Urbenville (Greenslade, P.J.M.); Blue Knob Mt., Nightcap Range
(Lowery, B.B.); Brindle Creek, Border Ranges Natl. Pk, 28°22'S 153°05'E (Naumann, I.D.);
Dorrigo Natl Pk, E end Blackbutt Track (Newton, A. & Thayer, M.) (MCZC); Doyle's River
State Forest, Oxley Highway, W Wauchope (Lowery, B.B.); Foxground, 10mi S Kiama
(Lowery, B.B.); Foxground, 8mi S Kiama (Lowery, B.B.); Genoa (Lowery, B.B.); Jerusalem
Bay, Cowan, Sydney (Lowery, B.B.); Macksville (Lowery, B.B.); Mt. Keira, 34°24'S 150°51'E
(Ward, P.S.); Mt. Ousley, near Wollongong, 34°23'S 150°52'E (Ward, P.S.); Mt. Saddleback,
Kiama (Lowery, B.B.); Mt. Warning (Lowery, B.B.); Never Never Picnic area, Dorrigo Natl Pk
(Newton, A., Thayer, M.); Ourimbah (collector unknown) (MCZC); Ourimbah State Forest,
Wyong (Lowery, B.B.); Pymble (Lowery, B.B.); Royal Natl. Pk., 34°09'S 151°01'E (Ward,
P.S.); Seal Rocks, 32°26'S 152°32'E (Ward, P.S.); Sheepstation Creek, Wiangaree State Forest
(Newton, A., Thayer, M.) (MCZC); St. Ives, Sydney (Lowery, B.B.); Swampy forest behind 7
Mile Beach, Gerroa (Lowery, B.B.); Thone River, Oxley Highway, W Wouchope Tomewin,
10mi N Murwillumbah (Lowery, B.B.); Whian Whian State Forest, 28°39'S 153°20'E((Lowery,
B.B.); (Ward, P.S.); upper Allyn Valley, nr. Eccleston (Taylor & Brooks; Taylor, R.W.);
Queensland: 1.5km EbyN Mt. Sorrow, 16°05'S 145°27'E (Calder, A. & Weir, T.); 3 mi S
Blackall (Perkins, F.A.); 6km SSW North Tamborine, 27°56'S 153°11'E (Ward, P.S.);
Alexandra Bay, 16°12'S 145°26'E (Taylor, R.W. & Feehan, J.); Bald Knob, 5km ESE Maleny,
26°46'S 152°53'E (Hill, L.); Binna Burra, Lamington Natl Pk (Lawrence, J. & N.); Boombana
Natl. Pk., Mt. Glorious (Taylor, R.W.); Brisbane (Hacker,H.); Cairns District (Lea, A.M.);
Camp Mt., Brisbane (Lowery, B.B.); Cape Tribulation, 16°04'S 145°28'E (Calder, A. & Weir,
T.); Cedar Creek, Tamborine Mt. (Brown, W.L.) (MCZC); Clump Point (Woodward, T.E.);
Coopers Plains Br. (Perkins, F.A.); Crawfords Lookout, Palmerston Natl. Pk (Taylor, R.W.);
Cunningham's Gap (Lowery, B.B.); Dalsy Hill State Forest, 24km SE Brisbane (Lowery, B.B.);
Eacham Natl Pk, 17°18'S 145°37'E (Taylor, R.W.); Eungella Natl. Pk., 50km W Mackay
(Lowery, B.B.); Gadgaria (Greenslade, P.J.M.); Gayundah Creek, Hinchinbrook Island, 18°22'S
146°13'E (Davies, Thompson & Gallon; Monteith, Davies, Thompson & Gallon; Thompson,
G.); Goodna, Brisbane (Lowery, B.B.); Ithica Creek, Brisbane (Lowery, B.B.); Kirrama Range,
via Kennedy (Monteith, G.); Kondalilla Natl Pk, Blackall Ranges (Brown, W.L.) (MCZC);
Kroombit Tops, 65km SW Gladstone (Monteith, G. & Thompson, G.); Kroombit Tops, SSW
Calliope, Beauty Spot 98, 24°22'S 150°59'E (Monteith, G.); Kroombit Tops, SSW Calliope,
Three Moon Scrub, 24°25'S 151°03'E (Monteith, G.); Kuranda (Brown, W.L.) (MCZC);
Kuranda, Black Mt. Road, 16°45'S 145°33'E (Taylor, R.W. & Feehan, J.); Kweebank Cave
track, Binna Burra, Lamington Natl. Pk. (Taylor, R.W.); Landsborough (Perkins, F.A.); Mackay
(Ridy; Turner, G.); McNamee Creek, 17°40'S 145°48'E (Taylor, R.W. & Feehan, J.); Melita,
Cooloola Natl Pk (Greenslade, P.J.M.); Mt. Coot-tha, Brisbane (Lowery, B.B.); Mt. Mee State
Forest, 27°06'S 152°42'E (Monteith, G.B.); Mt. Nebo (Taylor, R.W.); Mt. Nebo, Brisbane
(Lowery, B.B.); Mt. Webb Natl. Pk., 15°04'S 145°07'E (Calder, A. & Feehan, J.); Mulgrave
River Road, 7km WbyS Bellenden Ker, 17°16'S 145°47'E (Calder, A. & Weir, T.); Noah Creek,
7km ENE Thornton Peak, 16°08'S 145°26'E (Calder, A. & Weir, T.); Noosa River, Cooloola
Natl Pk (Greenslade, P.J.M.); Pingin Hill (Holt, J.); Shipton's Flat (S of Cooktown) (Darlington,
P.F.) (MCZC); Spicers Gap, 28°05'S 152°25'E (Ward, P.S.); Tambourine Mt. (Lea, A.M.);
Thornton Range, 16°14'S 145°26'E (Taylor, R.W., Feehan, J.); Tully Falls Natl. Pk., 17°47'S
145°33'E (Taylor, R.W. & Feehan, J.); Warrawonga, Cooloola Natl Pk (Greenslade, P.J.M.);
Weipa (Andersen, A.N.) (TERC); Wongabel State Forest, 5km S Atherton (Monteith &
Thompson); nr. Brisbane (Hacker, H.); New Zealand: Grey Lynn, Auckland (Taylor, R.W.)
(MCZC); Mt. Alber, N side, Auckland (Taylor, R.W.); Mt. Eden, Auckland (Hammond, P.M.);
Te Atatu, Auckland (Keall, J.B.).
DIAGNOSIS
This taxon can be separated from other Australian species of this genus by the presence of
well developed sculpturing in the posterior section of the scrobe, the large, closely spaced pits
on the mesosomal dorsum, and the parallel lateral surfaces of the postpetiole. It can be separated
from the south-east Asian M. transfuga by the less angular petiolar node and more heavily
sculptured postpetiole.
WORKER DESCRIPTION
Sculpturing in posterior section of antennal scrobe well developed and distinct; sculpturing
on dorsal surface of mesosoma consisting of large, closely spaced pits (or rarely smaller and
more widely spaced); propodeal spines varying from short and triangular to elongate and thin;
dorsal surface of petiole in lateral profile uniformly convex, without distinct dorsal and posterior
faces and forming an obtuse angle with the anterior face; in dorsal view, postpetiole with the
anterior and posterior regions approximately the same width (the region connecting them either
flat or weakly convex); postpetiole and gaster lacking erect hairs dorsally.
Measurements. Worker (n = 10) - CI 0.89-0.97; HL 0.42-0.57; HTL 0.23-0.32; HW 0.39-
0.50; ML 0.42-0.57; PW 0.29-0.37; SI 0.58-0.73; SL 0.24-0.36.
COMMENTS
As conceived here, M. abstinens shows considerable variation in several morphological

traits. The most obvious of these include the overall body color and the length of the propodeal
spines. The body color varies from uniform light yellow to uniform dark brown, as well as
bicolored dark yellow and dark brown. This variation was noted by Wheeler (1935) during his
revision and was used when establishing his subspecies M. abstinens hackeri and M. a.
venustula. The propodeal spines vary in shape from short and triangular to elongate and thin.
During the present study, it was found that the majority of available material could be
sorted into yellow forms and dark brown forms and that most of the variation in color occurred
between nest series. However, a small number of nests were intermediate between the lightest
and darkest forms or were bicolored yellow and brown and could not be easily placed in either
form. To complicate matters further, a few series, apparently representing single nests, contained
workers of both color forms. Thus color could not be used to sort the available material without
arbitrarily placing some specimens.
Propodeal spine morphology shows similar variation to body color, except three classes
could be established which overlap minimally. These include very short triangular spines,
elongate triangular spines, and elongate thin spines. As with body color, the majority of
specimens can be sorted into these three classes with only a small number of specimens being
intermediate, and only a few nest series showing variation that spans more than a single class.
When body color and spine morphology are taken together, the same pattern as seen in the
characters individually emerges again. The majority of dark individuals have elongate triangular
spines with a few having short triangular or elongate thin spines. In contrast, most lightly
colored individuals have short triangular spines but a significant number have elongate
triangular or elongate thin spines. Thus the majority of the specimens show a consistent pattern
as would be expected for two distinct species, but a minority of specimens do not fit with this
pattern and suggest that a single species is involved.
Geographically, light body color ranges from the northern extreme of the species south to
south-east Queensland while the dark body color ranges from approximately Kennedy (north of
Ingham), Queensland south to the southern extreme of the range. All forms of the propodeal
spines occur in the Queensland populations while only the intermediate form occurs in New
South Wales. Thus body color shows a distinct geographic pattern with light individuals in the
north, both color forms throughout the central part of the range, and dark forms occurring in the
southern part of the range. Spinal morphology shows a similar pattern, but without the northern
differentiation seen in body color.
In addition to color, several other characters show variation within this species. The
sculpturing on the head and mesosoma is more pronounced in the darker form when compared
with the lighter form. However, it is difficult to determine if this difference is caused by the
underlying cuticular coloration accentuating the sculpturing differences, or if the difference is in
the cuticular structure. In a few rare cases (for example specimens from Seal Rocks, New South
Wales) the dorsum of the mesosoma is essentially smooth with small, widely spaced pits similar
to those in M. overbecki. These specimens are placed here because of the relatively well
developed sculpturing in the antennal scrobes and the angular petiole with a short posterior face.
In addition to the sculpture, there is a weak trend for the darker form to have a more angular
antero-dorsal petiolar face when compared with the lighter morph. However, this shows
considerable variation within both forms so that the entire range of variation can be found in
each.
Because all of the characters show considerable variation, no individual character could be
found to suggest that more than a single variable species is involved and because color and spine
morphology do not covary, all specimens are here placed in a single species.
This species has been collected in habitats ranging from dry sclerophyll woodlands (less
commonly) to rainforests (more commonly), and once in a garden. Nests are found mainly in
soil, either in the open or between rocks, and they have been found nesting within the mounds of
Myrmecia auriventris, M. brevinoda and M. flavicoma and Pachycondyla (Bothroponera)
mayri. They also nest arboreally under dead bark and in rotten wood on the ground. Workers
forage on the ground (where they are often found in leaf litter samples) as well as arboreally.
Mayriella ebbei Shattuck & Barnett, sp. nov.

Figures 6-9
TYPE MATERIAL
Holotype worker from Australia, Booroomba Rocks, 35°33'S 148°59'E, Australian Capital
Territory, 16 March 1992, S. O. Shattuck, eucalypt woodland (ANIC); ca. 60 worker and 1
dealate queen paratypes, same data as holotype (ANIC, BMNH, MCZC) (as well as numerous
additional larvae and worker pupae and 2 male pupae).
ADDITIONAL MATERIAL
(In ANIC unless otherwise noted). Australia, ACT: 5km SW Orroral Tracking Station
(Lowery, B.B.); Black Mt., S slope (lake road) (Taplin, I.C.); Blundells Creek Rd, 3.5km E
Piccadilly Circus (Lawerence, J.); Booroomba Rocks, 35°33'S 148°59'E (Shattuck, S.O.);
Brindabella Range (Lowery, B.B.); Wombat Creek, 6km NE Piccadilly Circus, 35°19'S
148°51'E (Weir, Lawerence & Johnson); foot of Mt. Majura (Lowery, B.B.); nr. Lees Spring,
Brindabella Range (Taylor, R.W.); New South Wales: 4.5km WNW Pigeon House Mt., 35°21'S
150°13'E (Hill, L.); Burns Bay, Lane Cove, Sydney (Lowery, B.B.); Castle Flat, Clyde River
floodplain, 32°21'S 150°13'E (Hill, L.); Gerroa, 10mi S Kiama (Lowery, B.B.); Kanangra Brook
and Rocky Spur, Kanangra-Boyd Natl Pk (Hill, L.); Kanangra-Boyd Natl Pk (Hill, L.); Lane
Cove, Sydney (Lowery, B.B.); Riverview College, Lane Cove, Sydney (Lowery, B.B.);
Tallaganda State Forest, Captain's Flat (Lowery, B.B.); Tamborine Bay, Lane Cove, Sydney
(Lowery, B.B.); Tamborine Bay, Sydney (Lowery, B.B.); South Australia: Belair (Greenslade,
P.J.M.); Bridgewater (Hutson); Bridgewater, Englebrook (Greenslade, P.J.M.); Christenson
Park, Sevenhill (Lowery, B.B.); Maclaren Flat (Kirkby, C.A.); Mt. Lofty, S Para (Hutson);
Sevenhill (Lowery, B.B.); West Bay, Kangaroo Island (Greenslade, P.J.M.); Tasmania:
Eddystone Point (Trueman, J. & Cranston, P.); Victoria: 12km E Warburton (Newton, A. &
Thayer, M.); Arthur's Seat (McCrae) (Brown, W.L.) (MCZC); Mt. Buffalo Natl Pk (Newton, A.
& Thayer, M.); Narbethong (McAreavy, J.); Oberon Creek, Summit Mt., Wilsons Prom.
(Ettershank, G.); One Tree Hill, Melbourne (Lowery, B.B.); Portland (Beauglehole, C.);
Queenstown, nr. Hurstbridge (Lowery, B.B.).
DIAGNOSIS
This species can be separated from others in this genus by the presence of numerous erect
hairs on the gaster. It shares the shape of the outer margins of the postpetiole with M. spinosior,
but differs from this species in the shape of the petiolar node and in having more than four erect
hairs on the postpetiole.
WORKER DESCRIPTION
on dorsal surface of mesosoma consisting of large, closely spaced pits; propodeal spines
relatively long and thin; dorsal surface of petiole in lateral profile uniformly convex, without
distinct dorsal and posterior faces and forming an obtuse angle with the anterior face; in dorsal
view, anterior region of postpetiole expanded relative to posterior region; dorsum of postpetiole
with more than four erect hairs; dorsum of gaster with numerous erect hairs.
0.54; ML 0.49-0.64; PW 0.30-0.40; SI 0.62-0.66; SL 0.27-0.35.
COMMENTS
This is the southernmost species of Mayriella and occurs in generally drier sites compared
to other species. Although widely distributed it shows minimal geographic variation in the
characters examined during this study. Biologically, this species is found primarily in drier
habitats such as sclerophyll woodlands, low scrub, ti-tree scrub and coastal heath and less
commonly in wet sclerophyll and rainforests. Nests occur in soil in the open or under stones or
other objects on the ground, and in rotten wood.
Mayriella granulata Dlussky & Radchenko, 1990

(Figs 10-12)
Mayriella granulata Dlussky & Radchenko, 1990: 123.
TYPE MATERIAL
Holotype worker and two paratype workers (UASK) from “Archipelago Baitylong, Isl.
Dongho“, Quang Ninh Prov., Vietnam (examined). This island is also known as Dảo Dông
Khoa, Ba Mùn Island and Cao Lô Island (and probably other names) and is located approx.
100km ENE of Hải Phòng at approx. 21º00’N 107º35’E. No additional material currently
available.
DIAGNOSIS
This species is similar to M. transfuga and can be separated from it by the presence of
relatively short and broad propodeal spines (the spines are at most about as long as the width of
their bases while they are at least 1.5 times the width of the base in M. transfuga). It can be
separated from the remaining species in the genus by the presence of well developed sculpturing
in the posterior section of the scrobe, the large, closely spaced pits on the mesosomal dorsum,
the parallel lateral surfaces of the postpetiole and the strongly angular petiolar node.
WORKER DESCRIPTION
relatively long and broad; dorsal surface of petiole in lateral profile uniformly convex, without
distinct dorsal and posterior faces and forming a sharp angle with the anterior face; in dorsal
view, postpetiole with the anterior and posterior regions approximately the same width (the
region connecting them either flat or weakly convex); postpetiole and gaster lacking erect hairs
dorsally.
Measurements. Holotype - CI 0.98; HL 0.43; HTL 0.22; HW 0.42; ML 0.45; PW 0.30; SI
0.57; SL 0.24.
COMMENTS
This species is essentially identical to M. transfuga in all characters examined during this
study with the exception of the broader and slightly shorter propodeal spines (as pointed out in
the original description) and the apparently longer hairs on the lateral margins of head,
especially the pair originating immediately posterior to the eyes. But even here, the spines in one
of the paratypes (the upper one) are noticeably narrower than those of the holotype and
remaining paratype and approach those found in some transfuga specimens (for example from
Kota Tinggi, E. Johore, Malaysia). Also, the lengths of the head hairs are difficult to assess
critically because they vary considerably in the degree of curvature, making precise
measurements difficult. While there is currently little support for treating this taxon as distinct
from transfuga, it would be helpful to have additional material before deciding on its true status.
Thus, this species is retained for the time being while noting that it may well represent little
more than a slight variation of transfuga.
Mayriella overbecki Viehmeyer, 1925, n. stat.

Figures 13-16
Mayriella overbecki Viehmeyer, 1925: 26 (removed from synonymy with M. abstinens).
TYPE MATERIAL
Worker (NHMB) (examined) and 1 queen (MCZC) (examined) syntypes from Australia,
Trial Bay, New South Wales.
ADDITIONAL MATERIAL
Australia, New South Wales: Brindle Creek, Wiangaree State Forest (collector unknown)
(ANIC, MCZC); Bruxner Park, via Coffs Harbour (Monteith, G.) (ANIC); Dorrigo Natl Pk, E
end Blackbutt Track (Newton, A. & Thayer, M.) (ANIC, MCZC); Mt. Warning (Lowery, B.B.)
(ANIC).
DIAGNOSIS
This species can be separated from others in the genus by the combination of small,
scattered pits on the mesosomal dorsum, weak sculpturing in the posterior sections of the
antennal scrobes and the petiole with a relatively long posterior face and acute angle.
WORKER DESCRIPTION
Sculpturing in posterior section of antennal scrobe at most weakly developed and always
indistinct; sculpturing on dorsal mesosoma consisting of small, widely spaced pits; propodeal
spines relatively short and thick; dorsal surface of petiole in lateral profile uniformly convex,
without distinct dorsal and posterior faces and forming a blunt angle with the anterior face; in
dorsal view, postpetiole with the anterior and posterior regions approximately the same width
(the region connecting them either flat or weakly convex); postpetiole and gaster lacking erect
hairs dorsally.

0.53; ML 0.54-0.62; PW 0.33-0.37; SI 0.62-0.68; SL 0.30-0.35.
COMMENTS
This is one of the rarer species of Mayriella, being known from only five collections. These
collections were from rainforest in north-eastern New South Wales, with nests being found in
rotten logs.
Mayriella overbecki was established for a number of workers and a queen collected at Trial
Bay, New South Wales. It was considered a junior synonym of M. abstinens by Wheeler (1935),
a treatment followed by subsequent authors. Wheeler, however, based his synonymy on
Viehmeyer's original description and did not examine type material. This synonymy was curious
given that Wheeler established two new species based on subtle characters and slight
differences. This taxon is here removed from synonymy and recognized as a distinct species.
Mayriella sharpi Shattuck & Barnett, sp. nov.

Figures 17-19
TYPE MATERIAL
Holotype worker from Bisianumu, nr. Sogeri (approx. 9º24’S, 147º24’E), Central Province,
Papua New Guinea, 15-20 March, 1955, E. O. Wilson, rainforest (ANIC).
DIAGNOSIS
M. sharpi is immediately recognizable by the sharply pointed anteroventral extensions of

the compound eyes. It is also the only species currently known from Papua New Guinea.
WORKER DESCRIPTION
Sculpturing in posterior section of antennal scrobe well developed and distinct; compound
eye forming a sharp point ventrally; sculpturing on dorsal mesosoma consisting of small, widely
spaced pits; propodeal spines relatively short and thick; dorsal surface of petiole in lateral profile
uniformly convex, without distinct dorsal and posterior faces and forming a blunt angle with the
anterior face; in dorsal view, postpetiole with the anterior and posterior regions approximately
the same width (the region connecting them either flat or weakly convex); postpetiole and gaster
lacking erect hairs dorsally.
Measurements. Holotype - CI 1.01; HL 0.45; HTL 0.25; HW 0.46; ML 0.48; PW 0.32; SI
0.63; SL 0.29.
COMMENTS
The only known collection of this species consists of a single worker found in rainforest.
The specimen is pale yellow and appears to be callow. The specimen is unusual in having the
sting placed forward along the ventral surface of the gaster and directed downward. Given the
possibly callow nature of the specimen it is difficult to assess whether this is its normal position
or if the gaster has been deformed during preservation in alcohol before being point mounted.
Additional specimens will be required to ascertain the true nature of the sting placement.
Mayriella spinosior Wheeler, 1935

Figures 20-22
Mayriella spinosior Wheeler, 1935: 159.
TYPE MATERIAL
Holotype worker from Australia, Cairns district, Queensland (MCZC) (examined).
ADDITIONAL MATERIAL
(In ANIC unless otherwise noted). Australia, Queensland: 14km WbyS Eungella, 21°10'S
148°22'E (Gillison, A.); 1km W Imbil (Hill, L.); 1km W Kuranda (Doyen, J.) (MCZC); 8.5km
WSW Seaforth, 20°56'S 148°53'E (Gillison, A.); Braemar State Forest, via Kogan, 27°13'S
150°50'E (Raven, P.); Burpengary, via Cabodture (Monteith, G.B.); Cape Tribulation, 16°04'S
145°28'E (Calder, A. & Weir, T.); Carnarvon Gorge (Woodward, T.E.); Lake Barrine, Atherton
Tableland (Darlington) (MCZC); Melita, Cooloola Natl Pk (Greenslade, P.J.M.); Mossman
Bluff Track, 5-10km W Mossman (Monteith & Thompson); Mt. Webb Natl. Pk., 15°04'S
145°07'E (Calder, A. & Feehan, J.); Plowman, Cooloola Natl Pk (Greenslade, P.J.M.); Reedy
Creek, via Somerset Dam (Monteith, G.B.); Thornton Range, 16°14'S 145°26'E (Taylor, R.W.
& Feehan, J.).
DIAGNOSIS
This species can be separated from others in this genus by the lateral margins of the
postpetiole being expanded anteriorly and forming a trapezoid when viewed from above (similar
to Fig. 9), combined with the lack of erect hairs on the gaster.
WORKER DESCRIPTION
relatively long and thin; dorsal surface of petiole in lateral profile angular and with distinct
dorsal and posterior faces; in dorsal view, anterior region of postpetiole expanded relative to
posterior region; dorsum of postpetiole with at most four erect hairs; gaster generally lacking
erect hairs dorsally, very rarely (only known in one specimen) erect hairs present.
0.55; ML 0.46-0.60; PW 0.29-0.40; SI 0.59-0.65; SL 0.26-0.33.
COMMENTS
This species has been found primarily in litter samples collected in rainforests with a few
additional records from Eucalyptus woodlands. It occurs along essentially the entire Queensland
coast with several inland records in the southern part of its range.
This species is sympatric with M. abstinens, to such an extent that they have been collected
together in the same litter samples. This co-occurrence is at least partly the cause of earlier
taxonomic confusion in the genus as several individual pins in the ANIC contained both species,
complicating initial species characterizations. Once this sympatry was recognized and the
species separated, diagnoses were easily developed.
Mayriella transfuga Baroni Urbani, 1977

Figures 23-25
Mayriella transfuga Baroni Urbani, 1977: 411.
TYPE MATERIAL
Holotype worker and 7 worker paratypes from Nepal, 6 km NW Narainghat (NHMB,

BMNH) (examined).
ADDITIONAL MATERIAL
(In ANIC unless otherwise noted). China, Hong Kong SAR: Tai Po Kau forest, New
Territories (Winney, R.) (BMNH); India, Uttar Pradesh: Lachiwala forest, Dehra Dun, Garwal
District (Lobl, I); Indonesia, South Kalimantan: 17-46km W Batulitjin (Brown, W.L.) (BMNH,
MCZC); West Java: Bogor (Imadate, G.) (BMNH); Lampung: Liwa, 5 °04'S 104°03'E (Harvey,
M.S.); Malaysia, Johor: Kota Tinggi (Murphy, D.H.); Pahang: Tanjong Bunga (Murphy,
D.H.); Perak: Sungei Simei Falls, Cameron Highlands (Jaccoud, T. & Marcuard, P.); Pulau
Pinang: George Town Botanic Gardens (Waterfall) (Taylor, R.W. & Barrett, R.A.); Sabah:
Poring Hot Springs (Burckhardt & Löbl) (BMNH); Sepilok Forest Reserve, nr. Sandakan
(Taylor, R.W.); Umas Umas nr. Tawau (Taylor, R.W.); Sarawak: Kampong Segu, 20mi SW
Kuching (Taylor, R.W.); Semengoh Forest Reserve, 11mi SW Kuching (Taylor, R.W); nr. Miri
(Hammond, P.) (BMNH); G. Mulu Natl Pk, RGS Expd Long Pala (Bolton, B.) (BMNH); Gn.
Mulu Natl Pk (Hammond, P. & Marshall, J.E.) (BMNH); Selangor: Gombak, vic. of Univ. of
Malaya Field Station, 9th Mile (Murphy, D.H.); Ulu Gombak Forest Reserve (Crozier, R.)
(MCZC); Ulu Gombak, nr. Kuala Lumpur (Taylor, R.W.); upper Gombak Valley (Murphy,
D.H.) (MCZC); upper Gombak Valley, nr. Kuala Lumpur (Taylor, R.W.); Nepal: 6km NW
Narainghat (Wittmer, W. & Baroni Urbani, C.) (BMNH); Philippines, Luzon: Mt. Makiling,
Lagunas (Dumont, K. & Morse, R.) (MCZC); Singapore: Nee Soon (Taylor, R.W.); Thailand:
Khao Sabap Natl Pk (Löbl & Burckhardt) (BMNH).
DIAGNOSIS
This taxon can be separated from other species in the genus by the presence of well
developed sculpturing in the posterior section of the scrobe, the large, closely spaced pits on the
mesosomal dorsum, the parallel lateral surfaces of the postpetiole, the strongly angular petiolar
node and the relatively long propodeal spines (length greater than 1.5 times the width of their
bases). It is most similar to M. granulata but can be separated based on the longer and narrower
propodeal spines.
WORKER DESCRIPTION
relatively long and thin; dorsal surface of petiole in lateral profile uniformly convex, without
distinct dorsal and posterior faces and forming a sharp angle with the anterior face; in dorsal
view, postpetiole with the anterior and posterior regions approximately the same width (the
region connecting them either flat or weakly convex); postpetiole and gaster lacking erect hairs
dorsally.
0.45; ML 0.41-0.47; PW 0.28-0.35; SI 0.49-0.61; SL 0.22-0.25.
COMMENTS
M. transfuga was described by Baroni Urbani (1977) from material collected in Bhutan and
Nepal. It is now known to occur much more widely, being found from Nepal and India east to
Hong Kong and the Philippines and south to Borneo and Java, Indonesia. While this is by far the
broadest geographic range of any species in the genus, this species shows minimal geographic
variation and there is no suggestion that more than one species is involved. The majority of
records are from primary and secondary rainforests where workers have been found foraging in
leaf litter or nesting in soil.
Recently, the species granulata was described from Vietnam by Dlussky and Radchenko
(1990). This species is morphologically similar to transfuga and may be conspecific with it. For
further discussion see under that species.
ACKNOWLEDGEMENTS
We would like to thank Stefan Cover (MCZC), Barry Bolton (BMNH), Roy Snelling (LACM)
and Alex Radchenko (UASK) for making material in their care freely available; Bob Taylor for
his extensive curation work in the ANIC; Frank Koch (Museum für Naturkunde, Berlin) for
examining the types of M. overbecki; Ewa Slipinska for translating the M. granulata description,
Alex Radchenko for information on the type locality of M. granulata and Charles Kugler for
detailed notes on the sting apparatus. Comments on this manuscript provided by Gary Alpert,
Roy Snelling, Phil Ward and an anonymous reviewer were most welcome.
LITERATURE CITED
Baroni Urbani, C. 1977. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen

Museums in Basel. Hymenoptera: Fam. Formicidae Genus Mayriella. Entomologica
Basiliensia 2: 411-414.
Bolton, B. 1994. Identification guide to the ant genera of the world. Harvard University Press,
Cambridge, Mass., 222 pp.
Dlussky, G.M. & Radchenko, A.G. 1990. The ants (Hymenoptera, Formicidae) of Vietnam.
Subfamilies Pseudomyrmicinae and Myrmicinae (tribes Calyptomyrmecini, Meranoplini
and Cataulacini). Pp. 119-125. In Akimov, I.A., Emelianov, I.G. & Zerova, M.D. (eds.).
News of Faunistics and Systematics: 184 pp. Kiev (Naukova Dumka).
Emery, C. 1922. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [concl.] In: Wytsman,
P. (Ed.) Genera Insectorum, Fasc. 174C. L. Desmet-Verteneuil, Brussels, pp. 207-397.
Forel, A. 1902. Fourmis nouvelles d'Australie. Revue Suisse de Zoologie 10: 405-548.
Hölldobler, B. & Wilson, E.O. 1990. The ants. Harvard University Press, Cambridge, Mass.,
xii + 732 pp.
Kugler, C. 1997. Stings of some species of Lordomyrma and Mayriella (Formicidae:
Myrmicinae). Insecta Mundi 11: 193-199.
Shattuck, S.O. 1999. Australian Ants: Their biology and identification. Monographs in
Invertebrate Taxonomy 3: xi + 1-226.
Taylor, R.W. 1961. Notes and new records of exotic ants introduced into New Zealand. New
Zealand Entomologist 2(6): 28-37.
Taylor, R.W. 1991. Nomenclature and distribution of some Australasian ants of the
Myrmicinae (Hymenoptera: Formicidae). Memoirs of the Queensland Museum 30: 599-614.
Viehmeyer, H. 1925. Formiciden der australischen Faunenregion. (Fortsetzung.)
Entomologische Mitteilungen 14: 25-39.
Wheeler, G.C. & Wheeler, J. 1973. The ant larvae of six tribes: second supplement
(Hymenoptera: Formicidae: Myrmicinae). Journal of the Georgia Entomological Society 8:
27-39.
Wheeler, G.C. & Wheeler, J. 1985. A simplified conspectus of the Formicidae. Transactions
of the American Entomological Society 111: 255-264.
Wheeler, W.M. 1935. The Australian ant genus Mayriella Forel. Psyche (Cambridge) 42: 151-
160.
Figs. 1-5. Mayriella abstinens Forel (Hinchinbrook Is., Gayundah Ck., Queensland; ANIC
Specimen Number 32-014526). 1, front of head; 2, distribution of material examined; 3, lateral
body; 4, dorsum of mesosoma; 5, postpetiole in dorsal view.
Figs. 6-9. Mayriella ebbei Shattuck & Barnett (Blundells Creek Rd, 3.5km E Piccadilly Circus,
ACT; ANIC Specimen Number 32-014469). 6, front of head; 7, distribution of material
examined; 8, lateral body; 9, postpetiole in dorsal view.
Figs. 10-12. Mayriella granulata Dlussky & Radchenko (holotype). 10, front of head;
11, distribution of material examined (type locality); 12, lateral body.
Figs. 13-16. Mayriella overbecki Viehmeyer (Brindle Creek, Wiangaree State Forest, New
South Wales; ANIC Specimen Number 32-014041). 13, front of head; 14, distribution of
material examined; 15, lateral body; 16, dorsum of mesosoma.
Figs. 17-19. Mayriella sharpi Shattuck & Barnett (holotype; ANIC Specimen Number
32-014459). 17, front of head; 18, distribution of material examined; 19, lateral body.
Figs. 20-22. Mayriella spinosior Wheeler (1km W Kuranda, Queensland; ANIC Specimen
Number 32-014058). 20, front of head; 21, distribution of material examined; 22, lateral body.
Figs. 23-25. Mayriella transfuga Baroni Urbani (Sungei Simei Falls, Cameron Highlands,
Malaysia; ANIC Specimen Number 32-001550). 23, front of head; 24, distribution of material
examined; 25, lateral body
Snelling, G. C., Snelling, R. R. 2007. New synonymy, new species, new keys to Neivamyrmex army
ants of the United States, pp. 459-550. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds).
Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
NEW SYNONYMY, NEW SPECIES, NEW KEYS TO NEIVAMYRMEX

ARMY ANTS OF THE UNITED STATES
Gordon C. Snelling
13161 Rancherias Road
Apple Valley, CA 92308, U.S.A.
myrmecophile@armyants.org
and
Roy R. Snelling
900 Exposition Boulevard
Los Angeles, CA 90007 U.S.A.
antmanrs@nhm.org
ABSTRACT
The United States species of Neivamyrmex are here reviewed. The following new synonymy is
proposed (senior synonym listed first): N. moseri Watkins = N. isodentatus Mackay; N. swainsoni
(Shuckard) = N. mexicanus (F. Smith) = N. fallax Borgmeier; N. halidaii (Shuckard) = N. mexicanus
(E. V. Enzmann), not N. mexicanus (F. Smith). N. mandibularis (M. R. Smith) is elevated to species
level, new status, having previously been regarded as a subspecies of N. pilosus (F. Smith). One
species, N. graciellae (Mann) is recorded from the United States (Arizona) for the first time. Four
new species are described: N. goyahkla (southern Arizona; workers only), N. kiowapache (Kansas to
southern Arizona; all castes), N. ndeh (southern Arizona; males only) and N. wilsoni (southern
California; workers). New keys are presented for the separation of worker and male castes of all
known United States species, and most species are illustrated.
Key words: Hymenoptera, Formicidae, Ecitoninae, Neivamyrmex, army ants, new species.
INTRODUCTION
The systematics of the New World army ants has been fairly stable since the massive revision by
Borgmeier (1955). The few species described since 1955 are mostly included in the keys to New
World army ants by Watkins (1972, 1976, 1982, 1990).
Borgmeier's classification of the New World army ants divided them into five genera:
Cheliomyrmex (the sole member of the tribe Cheliomyrmecini), Labidus, Eciton, Nomamyrmex, and
Neivamyrmex, all in the tribe Ecitonini, and all hitherto regarded as subgenera within Eciton. The
three genera Eciton, Neivamyrmex, and Nomamyrmex should possibly be recombined to form a
single genus, Eciton. As defined by Borgmeier the characters separating these genera are trivial and
the distinguishing feature for Neivamyrmex (the absence of a tooth on the pretarsal claw of the
worker) is equivocal. Genera should be based on characters of a more fundamental nature. Despite
numerous papers on the Ecitonini, Watkins never attempted a critical analysis of the status of the
various genera. A detailed study of character states in the numerous species of Ecitonini must be
made before any substantive conclusion can be reached; such a study is far beyond the scope of this
paper.
Borgmeier, as had Creighton (1950) before him, abandoned the quadrinomial system of
classification and instead recognized geographical subspecies for many of the species treated. These
were often based on trivial features of color and the proposed system has since proven untenable for
many species. Watkins, in his various contributions to the taxonomy of New World army ants
avoided entirely the matter of geographical variation. Strangely, while he included the infraspecific
names in his various regional lists, he never attempted to analyze populations in an effort to
determine whether or not these forms represented taxonomically significant entities, even though he
commonly used the subspecific names when identifying material. This short-coming had little impact
on those species of Neivamyrmex found in the United States, from which he listed only two
subspecies, both attributed to a single species.
Specimens utilized in the course of this study have been examined from the following institutional
and private collections:
American Museum of Natural History, New York, New York, U. S. A. (AMNH)

Arizona State University, Tempe, Arizona, U. S. A. (ARSU)
California Academy of Sciences, San Francisco, California, U. S. A. (CASC)
Clemson College, Clemson, South Carolina, U. S. A. (CLMC)
Cornell University, Ithaca, New York, U. S. A. (CUIC)
Essig Entomological Museum, University of California, Berkeley, California, U. S. A. (UCBC)
Gordon C. Snelling, personal collection, Apple Valley, California, U. S. A. (GCSC)
Jamie King, personal collection, San Diego, California, U. S. A. (JAKC)
Kenneth W. Cooper, personal collection, Riverside, California, U. S. A. (KWCC)
Natural History Museum of Los Angeles County, Los Angeles, California, U. S. A. (LACM)
Lloyd Davis, personal collection Gainesville, Florida, U.S.A. (LDPC)
University of Michigan, Ann Arbor, Michigan, U. S. A. (MICH)
University of Minnesota, Minneapolis, Minnesota, U. S. A. (MINN)
Museum of Comparative Zoology, Cambridge, Mass., U. S. A. (MCZC)
University of Pennsylvania, Philadelphia, Pennsylvania, U. S. A. (PENN)
Robert A. Johnson personal collection, Tempe, Arizona, U. S. A. (RAJC)
Snow Entomological Museum, University of Kansas, Lawrence, Kansas, U. S. A. (SEMC)
Texas A & M University, College Station, Texas, U. S. A. (TAMU)
Snelling and Snelling: United States Neivamyrmex 461
The Natural History Museum, London, United Kingdom (BMNH)

University of California, Davis, California, U. S. A. (UCDC)
University of California, Riverside, California, U. S. A. (UCRC)
University of Arizona, Tucson, Arizona, U. S. A. (UNAR)
William P. MacKay, personal collection, El Paso, Texas, U. S. A. (WPMC)
Other collections cited:
ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, U. S. A.

IBVB Instituto de Biologia Vegetal, Brazil
MCSN Museo Civico di Storia Naturale, Genoa, Italy
MHNG Muséum d'Histoire Naturelle, Geneva, Switzerland
NHMV Naturhistorisches Museum, Vienna, Austria
TERMINOLOGY
All measurements were made from mounted, fully dry, specimens under a binocular microscope with
10× oculars, and a Mitutoyo, x-y axis stage and Mitutoyo digital micrometers. Images were made
with a Hitachi S-3000 scanning electron microscope.
We do not use the term “occiput” when referring to the posterior margin of the head; that term is
morphologically incorrect. Instead we refer to the posterior margin as the preoccipital margin; in
Neivamyrmex species it is often marked by a more or less distinct preoccipital carina.
Traditionally, the lengths of the petiolar and postpetiolar nodes are measured in dorsal view,
beginning at the anterior margin of the node; this margin is often difficult to define and resulting
measurements may thus be inaccurate to an unacceptable degree. We have made all petiole and
postpetiole measurements in side view, starting at the anterior edge of the spiracle to the posterior
margin of the segment. Although the spiracle may not correspond exactly to the anterior edge of the
node it provides an easily recognized and consistent starting point with which to begin a
measurement.
Also brought into conformity with generally accepted usage elsewhere among the Hymenoptera
are two terms used in describing male genitalia of army ants: the “stipes” or “stipites” are here called
the parameres (= gonostylus); the sagitta is more correctly to be referred to as the aedeagus and that
is employed here.
In the keys and descriptions, the following acronyms are employed:
CI Cephalic index: (HW/HL) (100).

EL Male only: maximum length of compound eye in lateral view.
EW Male only: maximum width of compound eye in lateral view.
FI Femoral Index: (HFW/HFL) (100).
HFL Hind femur length, greatest length along dorsal margin, base to apex.
HFW Hind femur width, measured perpendicular to HFL
HL With head in full face view, the maximum length from anteriormost margin of clypeus
(the thin lamelliform clypeal apron) to vertexal margin.
HW Worker and queen: with head in full face view, the maximum width, exclusive of eyes;
male HW measured across eyes.
IOD Male only: The minimum distance between the inner margins of the posterior ocelli.
OD Male only: The transverse diameter of the anterior ocellus.
OI Ocular index: (EL/HL) (100).
OMD The distance between the lower margin of the compound eye and the base of the
mandible, measured in lateral view.
OOD Male only: The minimum distance between the outer margin of a posterior ocellus and
the adjacent inner margin of the compound eye.
OVD Male only: With the head in frontal view, the shortest distance between either posterior
ocellus and the preoccipital margin.
PL Petiole length as defined above.
PPL Postpetiole length, using the same criteria as for PL.
PPW Maximum width of postpetiole in dorsal view.
PW Petiole width in dorsal view.
SI Scape index: (SL/HL) (100).
SL Maximum length of scape, exclusive of basal condyle.
WL Diagonal length of mesosoma in profile, from anterior declivity of pronotum (exclusive
of pronotal “neck”) to apex of propodeal lobe
MAPS
An attempt has been made to produce range maps, which are as accurate as possible. All maps were
produced with the “shareware” mapping program Versamap, available at Versamap.com. For
localities that did not contain coordinate data but were a known distance from a known locality the
program Geocalc was used to determine the correct coordinates of the site. Geocalc is available from
Victor Fraenckel (victorf@windreader.com).
NEIVAMYRMEX: KEY TO MALES
1 Vertex concave in profile and preoccipital margin raised and collar-like (Figs. 101, 111)
........................................................................................................................................... 2
-- Vertex flat or low-convex and preoccipital margin not collar-like (Fig. 98, 107, 109) ... 3
2(1) Inner margin of mandible moderately convex at midlength (Fig.110); paramere narrowly
rounded at apex (Fig, 135)......................................................................................pilosus
-- Inner margin of mandible straight or weakly convex at midlength (Fig.100); paramere blunt
and broadly truncate at apex (Fig. 127) ........................................................ mandibularis
3(1) Mandible strongly curved, rapidly tapering to acute apex (Figs.94, 95, 102, 104).......... 4
-- Mandible blade straight or weakly curved, sometimes parallel-sided (Figs. 96, 97, 106,
108)................................................................................................................................ 12
4(3) Larger species, HW at least 2.0 mm ................................................................................ 5

-- Smaller species, HW less than 2.0 mm ........................................................................... 6
5(4) ML subequal to EL; OD less than 2.5 x OOD (Fig.94); paramere with rectangular dorsal
process and triangular ventral process (Fig. 124) ...................................................andrei
-- ML greater than EL; OD more than 2.7 x OOD; paramere broadly rounded with shallow
dorsal concavity (Fig. 136) .............................................................................. swainsonii
6(4) Apicodorsal process of aedeagus broad and beak-like, extending beyond apicoventral
process in lateral view (Fig. 114, 115) ............................................................................ 7
-- Apicodorsal process of aedeagus not beak-like, not extending beyond apicoventral process
(Fig.113, 116) ................................................................................................................. 8
7(6) Larger species, HW at least 1.5mm; pilosity of body appressed; volsella with apical fork
......................................................................................................................... fuscipennis
-- Smaller species, MHW not over 1.2mm; pilosity of body consisting of long sparse suberect
to erect hairs; volsella without apical fork (Fig. 139) ....................................melshaemeri
8(6) Ocelli small, OOD equaling or exceeding OD (Fig. 102, 103) ....................................... 9
-- Ocelli larger, OOD not more than 0.30× OD (Fig 104, 105) ...................................minor
9(8) Head in frontal view without prominent dorsolateral angles (Figs. 96, 97, 102. 104, 106,
108,112 109); volsella with apical fork (Figs 140, 141, 142, 143) ................................ 10
-- Head in frontal view with prominent dorsolateral angles (Fig 95); volsella without apical
fork .........................................................................................................................baylori
10(9) Head and mesosoma black, gaster reddish brown (Arizona, California, Baja California).
....................................................................................................................................... 11
-- Head and entire body dark charcoal gray (Arizona) ............................................ microps
11(10) Very small species, HW less than 1.0 mm; head and mesosoma blackish brown, gastral
segments 1-2 mostly brown, merging into more reddish distal segments; posterior margin
of head concave, preoccipital margin forming low upturned collar; first two gastral terga
brownish, following segments more distinctly reddish (Arizona) ............................ ndeh
-- Larger species, HW over 1.3 mm; posterior margin of head broadly convex; preoccipital
margin inconspicuous, forming a small ridge or lip; gaster uniformly reddish brown
(California, Baja California) ..................................................................................mojave
12(3) OOD exceeding OD, usually at least 2.0× OD ............................................................. 13

-- OOD no more than about 0.5× OD, usually less (Fig. 104, 105) ........................... harrisi
13(12) Distal 0.66 of mandible slightly convex along inner margin, usually wider than base, apex
narrow and slightly pointed (Figs. 106, 108, 112,) ....................................................... 14
-- Distal 0.66 of mandible no wider than base and apex usually blunt (Figs. 96, 97)........ 15
14(13) Frons, on each side, prominently swollen above antennal fossa (Figs. 106, 107); proximal
flange on ventral margin of paramere complete (Fig. 133)............................... nigrescens
-- Frons not swollen above antennal fossa; (Figs, 108, 109); proximal flange on ventral
margin of paramere incomplete (Fig.134)....................................................... opacithorax
15(13) Mandible evenly tapering to broadly rounded tip (Fig 96, 97); paramere without prominent
apicodorsal process (Fig 125, 126) ................................................................................ 16
-- Mandible about equally wide for most of its length, abruptly narrower near apex (Fig. 112);
paramere with prominent apicodorsal process, small sharp hooked tooth at tip of
apicodorsal process (Fig. 137) ............................................................................... texanus
16(15) Larger species maximum HW ca 1.75 mm; scape approximately parallel-sided, 3× longer
than broad (Fig. 96) .......................................................................................carolinensis
-- Smaller species, HW not over 1.5 mm; scape approximately pyriform, 2× longer than broad
(Fig. 97, 99) ................................................................................................... kiowapache
NEIVAMYRMEX: KEY TO WORKERS
1 Eye present as a distinct convex facet.............................................................................. 2

-- Eye absent (pigmented eye-spot may be present) .......................................................... 14
2(1) Suture between promesonotum and mesepisternum incomplete or absent (Fig.32-49) ... 3
-- Suture present and complete between promesonotum and mesepisternum (Fig. 31).........
.................................................................................................................................. agilis
3(2) Concave portion of preoccipital margin not distinctly wider than greatest width of
pronotum (dorsal view) and strongly tapered, neck-like in appearance and vertex without
prominent dorsolateral angles (Figs. 13a-c, 28)............................................................... 4
-- Concave portion of preoccipital margin as wide as or wider than greatest width of
pronotum; vertex strongly developed dorsolaterally, angulate or not (Figs. 27, 29, 30)....
......................................................................................................................................... 6
4(3) Subpetiolar process a prominent posteriorly directed spine; color variable black to brown or
bicolored .......................................................................................................................... 5
-- Subpetiolar process absent or present as a small anteriorly directed tooth; usually bicolored,
red and black, but if all black, then mesonotum distinctly granulose ...... melanocephalus
5(4) Mesonotum strongly convex in profile and posterior face of propodeum about as long as
dorsal face (Fig. 44); color uniformly brownish to blackish ...................................pilosus
-- Mesonotum weakly convex in profile and posterior face of propodeum distinctly shorter
than dorsal face (Fig. 35); color uniformly reddish ........................................... graciellae
6(3) Petiole node in dorsal view less than 1.2× longer than broad (Figs. 63, 66, 67).............. 7
-- Petiole node in dorsal view at least 1.2× longer than broad (Figs. 62, 69) ................... 10
7(6) Head and body dull, coarsely granulopunctate .................................................. rugulosus

-- Head and body smooth and shiny .................................................................................... 8
8(7) Dorsal and lateral surfaces of petiole and postpetiole smooth and shiny, coarse punctures
may be present but scattered; dorsolateral corners of head weakly developed, slightly
rounded ............................................................................................................................ 9
-- Dorsal and lateral surfaces of petiole and postpetiole strongly granulose; dorsolateral
corners of head well developed and angulate ......................................................... harrisi
9(8) Cephalic punctures in middle of frons fine, separated by 2 PD or more; HF 5.15× longer
than thick .............................................................................................. carolinensis (part)
-- Cephalic punctures coarser, 1.2-1.7 PD apart in middle of frons; HF 4.4× longer than thick
.............................................................................................................. kiowapache (part)
10(6) Head smooth and shiny between scattered fine piligerous punctures ............................ 11
-- Head distinctly sculptured and roughened, often appearing granulopunctate................ 13
11(10) Inner basal margin of mandible evenly curved into masticatory margin, without tooth or
sharp angle at juncture (Fig. 18); pronotal sculpture variable, but side of pronotum usually
at least weakly reticulate-punctate rather than exhibiting a smooth shiny surface that
contrasts with the sculptured mesosoma dorsum ........................................................... 12
-- Inner basal margin of mandible straight, with distinct tooth or sharp angle at juncture with
masticatory margin (Fig. 20); side of pronotum usually smooth and shiny, and contrasting
with the rugulose-punctate sculpture that covers at least part of the mesosoma dorsum
........................................................................................................................ opacithorax
12(10) Dorsal face of propodeum distinctly depressed below level of mesonotum (Fig. 41);
metafemur 6.4× longer than greatest thickness (Fig.92) ........................ nigrescens (part)
-- Dorsal face of propodeum not distinctly depressed (Fig. 32); metafemur 6.2× longer than
greatest thickness (Fig. 88) ............................................................................. californicus
13(10) Posterior face of propodeum slightly concave in profile and distinctly angulate at juncture
with dorsal face angulate (angle may be indistinct in small specimens) (Fig. 47); petiole
relatively short and high; surface of most of the metapleural gland bulla densely punctate
and opaque, sometimes with a very small, thin shiny strip above the flange of the
metapleural gland orifice ....................................................................................... texanus
-- Posterior face of propodeum not concave in profile and rounded into juncture with dorsal
face (Fig. 41); petiole height lower; sculpture on surface of metapleural bulla becoming
obsolete, lower half or more is smooth (or weakly reticulate) and conspicuously shiny
................................................................................................................ nigrescens (part)
14(1) Petiole with anteroventral tooth ..................................................................................... 15

-- Petiole without anteroventral tooth ................................................................................ 16
15(14) Petiole tooth well developed and with posteriorly directed spine; hind femur at least 4×
longer than thick; subantennal lamella absent or poorly developed (Fig. 25)... swainsonii
-- Petiole tooth small, lacking posteriorly directed spine (Fig. 77); hind femur very short and
robust, 3.4× longer than thick (Fig. 90); subantennal lamella distinct (Fig. 4) ....goyahkla
16(14) Basal (innermost) mandibular tooth not conspicuously enlarged relative to remaining teeth
(Figs. 23-25) .................................................................................................................. 17
-- Basal mandibular tooth greatly enlarged relative to remaining teeth (Fig. 22) ....... moseri
17(16) Petiole node no longer than broad or only slightly so (Fig. 71)..................................... 18
-- Petiole node distinctly longer than broad (Figs. 68, 69, 70) .......................................... 19
18(17) Broad lamella present below antennal fossa (Fig. 26); head shiny between scattered distinct
punctures...............................................................................................................leonardi
-- Without lamella below antennal fossa (Fig. 24, 25); head shiny, with only scattered minute
and indistinct piligerous punctures ......................................................................pauxillus
19(17) Antennal scape short and stout, not evenly tapered basad (Fig. 7, 8, 16) ...................... 20
-- Antennal scape long and slender (about 3× as long as broad) (Fig. 11), gradually tapered
basad ...................................................................................................................... nyensis
20(19) Subantennal lamella well developed (Fig. 8, 16, 26) .................................................... 21

-- Subanatennal lamella absent or greatly reduced (Fig. 3, 7) .......................................... 22
21(20) Mesosomal dorsum smooth and shiny (Fig. 54) ................................................... microps
-- Mesosomal dorsum distinctly sculptured (Fig. 61), slightly shiny .........................wilsoni
22(20) Cephalic punctures in middle of frons fine, separated by 2 PD or more; HF 5.15× longer
than thick .............................................................................................. carolinensis (part)
-- Cephalic punctures coarse 1.2-1.7 PD apart in middle of frons; HF 4.43× longer than thick
.............................................................................................................. kiowapache (part)
Neivamyrmex agilis Borgmeier

Figures 1a-b, 17, 31, 49
Eciton (Acamatus) cocula Mann, 1925: 77 (w). Nomen nudum.

Neivamyrmex agilis Borgmeier, 1953: 45 (w). MEXICO, Chihuahua, Río Santa María (MCZC).
Borgmeier, 1955: 406-407 (w). Watkins, 1976: 17. Watkins, 1982: 211; pl. 8, fig. 8 (w). Watkins,
1985: 482 (w). MacKay & MacKay, 2002: 47.
DISTRIBUTION (Map 2)
UNITED STATES: Arizona, New Mexico; MEXICO: Chihuahua, Jalisco (Watkins, 1982).
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Cochise Co.: 0.5 mi. S. Paradise 5500' (LACM); 13.84 mi. E
Coronado National Monument (31.36ºN 110.51ºW) (GCSC, LACM). Santa Cruz Co.: 1 mi. SE
Peña Blanca Lake, 3950', under rock (ARSU, GCSC). Gila Co.: Hwy. 77, 12.1 mi. N Globe @ 0.8
mi. W Hwy 77 (33.55ºN 110.68ºW), 4920' (GCSC, LACM, RAJC). NEW MEXICO, Grant Co.: 3
mi. S. I-10 3.8 mi. E. Separ I-10, in Pogonomyrmex rugosus nest (ARSU, GCSC, MCZC).
DISCUSSION
A seldom collected species, N. agilis is known from only a handful of records. Until now known
only from the area around Paradise Arizona, we take this opportunity to present new locality records
for this species.
The male and queen of this species are so far unknown, but we believe that N. andrei is the
likely male.
Neivamyrmex andrei (Emery)

Figure 94, 124
Eciton andrei Emery, 1901: 53 (m). MEXICO (no specific locality) (MCSN).
Eciton (Acamatus) andrei: Emery, 1910: 25.
Eciton (Acamatus) oslari Wheeler, 1908:4 15; pl. 26, fig. 8 (m). UNITED STATES, Arizona, Nogales
(lost ?).
Neivamyrmex andrei: Borgmeier, 1953: 7, 19. Borgmeier, 1955: 451-453 (m). Watkins, 1976: 24 (m).
Watkins, 1982: 213; (m). Watkins, 1985: 484. MacKay & MacKay, 2002: 47.
UNITED STATES: New Mexico and Arizona. MEXICO: Colima, Nayarit, Sinaloa and Vera
Cruz (Watkins, 1982).
SPECIMENS EXAMINED
We have 39 United States records of this rather poorly known species.

DISCUSSION
In addition to the characters cited in the key, N. andrei may be separated from all other known
United States species, except N. swainsonii, by the broad front coxae (in all our other species, they
are distinctly longer than broad). As noted above, we believe that N. agilis will prove to be the
female castes of this species.
Neivamyrmex baylori Watkins

Figures 95
Neivamyrmex baylori Watkins, 1973: 430-433 (m). USA, Texas, McLennan County, Waco (USNM)
examined. Watkins, 1976: 27 (m). Watkins, 1985: 485 (m).
UNITED STATES, Texas
SPECIMENS EXAMINED
UNITED STATES, TEXAS, McLennan Co.: Waco (USNM). Uvalde Co.: Rio Frio River,
Haven Camp, (LACM).
DISCUSSION
This species is known from only three specimens, all collected in Texas. The worker and queen
of this species is unknown, but is likely to be N. pauxillus or, less probably, N. moseri.
Neivamyrmex californicus (Mayr)

Figures 2, 18, 32, 62, 75, 88
Eciton californicum Mayr, 1870: 969 (w). USA, California, San Francisco (NHMV). Lectotype
designated by Ward, 1999: 76
Eciton (Labidus) californicum: Mayr, 1886b: 121 (w).
Eciton (Acamatus) californicum: Emery, 1894: 184 (w). Emery, 1900: 523 (w).
Eciton (Acamatus) californicum var. obscura Forel, 1914a: 265 (w). USA, California, Vista (MHNG).
Lectotype designated by Ward, 1999: 76
Eciton (Neivamyrmex) californicum: M. R. Smith 1942: 560 (w). Creighton, 1950: 70 (w).
Neivamyrmex californicus: Borgmeier 1953: 11 (w). Borgmeier, 1955: 517-519 (w). Watkins, 1976: 15
(w). Ward, 1999 (w).
UNITED STATES, California. MEXICO, Baja California.
SPECIMENS EXAMINED
UNITED STATES, CALIFORNIA, Contra Costa Co.: 9km ENE Danville, 490m (37°51'N
121°54'W) (UCDC); El Dorado Co.: 14km NW Shingle Springs, 340m (38º45'N 121º02'W) (UCDC,
GCSC, LACM). Los Angeles Co.: Arcadia (USNM); La Verne (USNM, MCZC); Los Angeles
(USNM). Orange Co.: Laguna Hills (LACM); Tonner Canyon (WPMC). Riverside Co.: Riverside,
“in house” (GCSC, KWCC, LACM); Lake Skinner (UCRC0. Santa Barbara Co.: T5N, R28W s.17,
500m, (JTLC). San Diego Co.: National City (USNM); Vista (USNM); Camp Pendleton (LACM);
5mi NE Poway, 600m (UCDC); Mt. Laguna, MSP Site, 6050 ft. (LACM); Mira Mar, Elliot Reserve
(32.84ºN 117.11ºW) (JAKC). Yolo Co.: Davis (LACM, PENN).
MEXICO, BAJA CALIFORNIA: 28 km E Ensenada, 750m (31º53'N 116º18'W) (UCDC); San
Carlos Canyon, 9.0 mi E Hwy 1,300 ft., mesic canyon bottom, sifted from litter (31º47.88'N 116º
30.05'W) (ARSU).
DISCUSSION
The male form of this ant is unknown. This species appears to be primarily an ant of montane
and foothill areas. Previously believed to occur only in California, recent extensive collection in Baja
California has located this species there. All previous out of state records believed to apply to this
species have proven to refer to the recently recognized shiny headed form of N. nigrescens (Ward
1999). Watkins (1972) described what he believed was the queen of N. californicus, however recent
reexamination of the specimen in question (Ward, 1999) has led to the conclusion that it too is a
representative of the shiny form of N. nigrescens.
Little is known about the feeding preferences of this species other than that it presumably
attacks exclusively, or nearly so, other ant species. Ward (1999) lists the following as prey species of
this ant: Messor andrei, Solenopsis molesta, Pheidole californica and P. hyatti. One of us (GCS)
observed a portion of a raid on a Solenopsis xyloni colony; the raid was well underway when it was
discovered. It was apparently a violent one with S. xyloni vigorously defending its nest. Whether or
not the raid was successful from the standpoint of N. californicus is unknown but the ground was
littered with dead and dying of both species.
Automontage images of N. californicus may be viewed at antweb.org.
The wingless phorid fly, Xanionotum hystrix Brues (Diptera: Phoridae), has been associated
with a colony of N. californicus collected in San Diego County, California (J. H. Hunt, pers. comm.).
Neivamyrmex carolinensis (Emery)

Figures 3, 19, 33, 63, 76, 89, 96, 113, 125
Eciton (Acamatus) carolinense Emery, 1894: 184 (w). USA, North Carolina, Belmont (MCSN); syntypes
in MCZC, AMNH; examined.
Eciton (Neivamyrmex) carolinense: M. R. Smith, 1942: 564 (w).
Neivamyrmex carolinensis: Borgmeier, 1953: 6 (w).
UNITED STATES; Virginia and North Carolina south to Florida, west to Tennessee and
Louisiana.
SPECIMENS EXAMINED
UNITED STATES, ALABAMA, Tuscaloosa Co.: Bryce Lake, University of Alabama

(MCZC); Hurricane Creek nr. Peterson (MCZC). FLORIDA, Alachua Co.: Gainesville (UMICH);
Gainesville, San Felasco Hammock State Preserve, sandhill/xeric Oak Hammock, at base of dead
Turkey Oak (GCSC, LACM). Lake Co.: Astatula, Jct S.R. 561 & 48, Sand Pine/Rosemary Scrub, at
base of dead sand pine (GCSC, LACM). Citrus Co.: Withlacoochee State Forest, 1/2 mi S. of S. R.
44, in sand next to Pheidole morrissi colony (GCSC, LACM). Marion Co.: 3.5 mi E Dunnellon,
Pine Oaks Estates, (CLMC). Volusia Co.: Orange City, Blus Spring St. Park, ex pitfall trap in Sand
Pine/Oak Scrub (GCSC, LACM). GEORGIA, Fulton Co.: Atlanta, ex oak stump (MCZC). Putnam
Co.: Oconee National Forest (SEMC, WPMC). Rabun Co.: Clayton, 2000-3700' (MCZC).
KENTUCKY, Edmonton Co.: Mammoth Cave Nat. Park, Bruce Hollow, ex long/stump litter
(MCZC). MISSISSIPPI, Hinds Co.: Taylorsville (LACM). Lowndes Co.: Columbus (CASC).
Monroe Co.: Aberdeen (CASC, CORN). Oktibbeha Co.: A & M College (MINN, PENN). ORTH
CAROLINA: Burk Co.: 4 - XI- 1957 (Scott; MCZC). SOUTH CAROLINA, Greenville Co.:
Greenville (LACM). Oconee Co.: Clemson College (MCZC). TENNESSEE, Blount Co.: Cades
Cove (LACM). Knox Co.: University of Tennessee Farm (LACM).
DISCUSSION
Neivamyrmex carolinensis is a wide ranging eastern species. Long considered to extend into the
central and western United States as far as Arizona, it is now known that the populations west of the
Mississippi River belong to a hitherto unrecognized sibling species. That species is described below
as N. kiowapache. We are unable to confirm the presence of this species in Nebraska and Ohio (M.
R. Smith, 1942) nor have we seen any specimens from Louisiana.
This species is unusual among our species of Neivamyrmex, in that the presence of a distinct eye
facet is variable, present in some individuals, absent in others. Generally, a facet is present in larger
individuals, but even this is subject to variation. This feature is shared with its sister species, N.
kiowapache.
Automontage images of this ant may be viewed at antweb.org
Neivamyrmex fuscipennis (M. R. Smith)

Figure 114
Eciton (Acamatus) spoliator: Wheeler, 1908: 416-417 (m). Misidentification

“Acamatus fuscipennis Cresson”: Wheeler, 1908: 417 (m). Nomen nudum.
Eciton (Neivamyrmex) fuscipennis M. R. Smith, 1942: 578 (m). USA: Texas (Belfrage) (USNM)
examined.
Neivamyrmex fuscipennis: Borgmeier, 1953: 18. Borgmeier, 1955: 642. Watkins, 1975: 85-90.
Neivamyrmex macropterus Borgmeier, 1953: 40 (m). MEXICO, Durango, Lerdo (USNM). Borgmeier,
1955: 645-646 (m). Watkins, 1975: 85-90. NEW SYNONYMY.
UNITED STATES: Texas and Kansas, west to Arizona, south to Mexico (Morelos and San
Luis Potosí).
MATERIAL EXAMINED
UNITED STATES, ARIZONA, Cochise Co.: SWRS. 5 mi. W. Portal, 5400' (LACM, UNAR,
USNM); Herb Martyr Dam, Chiricahua Mtns. (UNAR.); near Portal (SEMC). Santa Cruz Co.:
Madera Canyon, 4880' (LACM). KANSAS Stevens Co.: Hugoton (SEMC). TEXAS, Jeff Davis
Co.: Ft. Davis Resort, 5800' (LACM). Presidio Co.: 40 mi. ESE Presidio, at UV light (UNAR). Rusk
Co.: Henderson (TAMU).
MEXICO, MORELOS: 10 mi N Cuernavaca (UCDC).
DISCUSSION
As Creighton (1950) had pointed out, the authorship of this species has been confused. Wheeler
(1908) recorded two male specimens from an unspecified locality in Texas as Eciton (Acamatus)
spoliator Forel, 1899, a species originally described from Costa Rica. He provided a translation of
Forel's species and a figure that was clearly based on one of the Texas males. Wheeler further noted
that the specimens bore an unpublished Cresson name, Acamatus fuscipennis.
M. R. Smith (1942) correctly recognized that the Texas specimens were not the same as Forel's
species and redescribed the two males as Eciton (Neivamyrmex) fuscipennis, attributing the species
authorship to Wheeler (1908). Subsequent authors have followed Smith's lead in awarding
authorship to Wheeler. D. R. Smith (1979), however, determined that the correct author for N.
fuscipennis should be M. R. Smith, 1942. Watkins (1975) concurred and selected one of the Texas
males as lectotype.
Borgmeier (1953, 1955) noted the similarities between his new species, N. macropterus,
described from Mexico, and N. fuscipennis, but cited relatively larger ocelli and wings as
distinguishing features for N. macropterus. Watkins (1975) took up the matter and, after examining
more than 50 males from 9 localities found that the differences in eye length and wing length cited
by Borgmeier were unreliable. He did conclude that, in N. fuscipennis the distance between the
lateral ocellus and the margin of the compound eye was slightly greater than in N. macropterus. He
further noted that this difference could be determined in many specimens only by careful, precise
measurements. His final determination was that the two species are so closely related that they could
not be reliably separated, especially in specimens from intermediate localities.
According to Watkins (1985) in N. fuscipennis the “distance from the lateral ocellus to the
compound eye was usually about one-half diameter of median ocellus” vs. “about one-fourth (or
less)” in N. macropterus. We have found that when males are so positioned that the appropriate
measurements of anterior ocellus and of the distance between the compound eye and the nearest
lateral ocellus are uniform these differences between the two supposed taxa disappear. Accordingly
we here treat N. macropterus as a synonym of N. fuscipennis.
Neivamyrmex goyahkla, new species

Figures 4, 34, 50, 64, 77, 90
DIAGNOSIS
Worker: Eye spot present; head distinctly longer than broad, sides weakly convex in frontal
view and upper margin narrowly rounded onto vertex margin; front of head smooth and shiny
between scattered minute punctures; antennal scape 3.5 x longer than broad; mesosomal dorsum
smooth and shiny, with widely scattered small punctures; petiole longer than broad in dorsal view,
sides approximately parallel; node evenly rounded in profile and dorsum smooth and shiny;
subpetiolar tooth small but distinct; postpetiole broader than long, disc smooth and shiny. Queen and
male unknown.
DESCRIPTION
Worker measurements (mm) (n = 12). HL 0.45-0.72 (0.64); HW 0.34-0.61 (0.54); SL ).25-0.36

(0.33); SW 0.08-0.13 (0.095); PW 0.12-0.20 (0.196); PL 0.14-0.25 (0.24); Ppl 0.10-0.20) (0.19);
Ppw 0.15-0.28 (0.25); HFL (0.48); HFW (0.14). Indices. CI 72-85 (85); FI 26-33 (31); SI 45-57
(51.5).
Head: distinctly longer than broad; sides weakly convex in frontal view; upper margin narrowly
rounded onto vertex margin; preoccipital carina sharply angulate laterad. Small yellow eye spot
present. Front of head smooth and shiny and with scattered minute punctures. Subantennal lamina
well-developed. Mandible triangular, junction of upper and masticatory margins rounded, small peg-
like tooth present on basal margin; outer surface with fine longitudinal rugulae. Antennal scape 3.5 x
longer than broad, evenly tapering toward base.
Mesosoma: dorsum smooth and shiny, with widely scattered small punctures. Promesonotal
suture indistinct to absent; metanotal suture distinct across dorsum. Dorsal face of propodeum about
as long as declivitous face, shiny between scattered fine punctures; declivitous face straight to
slightly concave in profile.
Petiole longer than broad in dorsal view, sides approximately parallel; node evenly rounded in
profile; subpetiolar tooth small but distinct; dorsum smooth and shiny. Postpetiole broader than long,
disc smooth and shiny.
Gaster smooth and shiny between scattered fine punctures.
Pilosity about as usual for the genus, consisting of sparse hairs of highly variable length, from
short to long.
TYPE DATA
Holotype and numerous paratypes: U.S.A., Arizona, Santa Cruz Co., Ruby Road, 6.7 mi west
of I-19, 6 April 1998 (R. A. Johnson & G. C. Snelling). Holotype and most paratypes in LACM;
paratypes also in ARSU, BMNH, CASC, MCZC, UCDC, USNM.
ETYMOLOGY
This species is named for Goyahkla, the Bedonkohe Chiricahua Apache war leader more widely
known by his Spanish name, Geronimo. It is most easily pronounced “Goyák lay”.
Presently known only from south-central Arizona.
DISCUSSION
In addition to the type series, we have examined a few specimens from Arizona, Cochise Co.:
Chiricahua Mountains, 2 August 1954, collected by A. C. Cole, Jr. (LACM).
This small shiny yellow species is similar to N. leonardi and N. nyensis, but is easily separated
from those species by the presence of a small but distinct subpetiolar tooth. It is further distinguished
from N. leonardi by the mandibular structure.
The type series was discovered while overturning rocks in the late afternoon. A small number of
workers were present under the rock, and a more detailed search of the area located a fairly strong
column of ants trailing across the soil. Although they made use of cover as much as possible a good
percentage of the column was exposed on the surface. The column was followed for approximately
thirty feet before it was finally lost. It is unknown if this was a raiding or emigration column but
since no brood was observed our assumption is that this was the beginning of a raid.
Neivamyrmex graciellae (Mann)

Figures 5a-b, 35, 51, 65, 78
Eciton gracellae Mann, 1926: 97 (w). MEXICO, JALISCO, Ototonilco (USNM) examined.
Neivamyrmex graciellae: Borgmeier, 1953: 10. Emendation. Borgmeier, 1955: 388-389 (w). Watkins &
Coody, 1986: 257 (q).
UNITED STATES: Arizona. MEXICO: Jalisco, Oaxaca.

SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Santa Cruz Co., Florida Canyon, Santa Rita Mts.
(LACM, GCSC).
MEXICO, JALISCO, Chamela Biological Station (USNM); Ototonilco (USNM). OAXACA,
13 mi NW Oaxaca, 5500 ft. (LACM).
DISCUSSION
This species has not been previously recorded from the United States. Its discovery in southern
Arizona raises the possibility that this species, rather than N. melanocephalus, might be the worker
caste of N. mandibularis. The question cannot now be resolved.
Neivamyrmex halidaii (Shuckard)
Labidus latreillii: Haliday, 1836: 328 (m). Misidentification

Labidus halidaii Shuckard, 1840: 200. BRAZIL, São Paulo (BMNH).
Eciton halidayi: Dalla Torre, 1893: 3. Erroneous spelling by Dalla Torre and most subsequent authors.
Labidus gravenhorstii Westwood, 1842: 76 (m). BRAZIL, Goiás, Rio Vendhina.
Labidus amplipennis F. Smith, 1859: 6 (m). COLOMBIA, Bogotá (BMNH).
Eciton (Acamatus) Le Moulti Santschi, 1912: 524 (m). FRENCH GUYANA, St. Laurent de Maroni
(Basel)
Eciton (Acamatus) colombi Santschi, 1921: 94 (m). COLOMBIA, Bogotá (Basel)
Eciton (Labidus) frontalis Menozzi, 1924: 29 (m). BRAZIL,
Woitkowskia mexicana E. Enzmann, 1952: 445 (m). MEXICO, Chiapas, Tuxtla Gutierrez (type lost ?).
NEW SYNONYMY
Neivamyrmex halidaii: Borgmeier, 1953: 12, 15, 18 (m). Bolton, 1995: 289.
Neivamyrmex amplipennis: Borgmeier, 1953: 17.
Neivamyrmex halidaii lemoulti: Borgmeier, 1953: 14.
Neivamyrmex colombi: Borgmeier, 1953: 13.
Neivamyrmex frontalis: Borgmeier, 1953: 11.
Neivamyrmex halidayi: Borgmeier, 1955: 437-445 (m). Watkins, 1982: 212. Watkins, 1990:381.
DISTRIBUTION
MEXICO: Vera Cruz and Oaxaca to Chiapas (Watkins, 1982), south to BOLIVIA and
northern ARGENTINA.
DISCUSSION
The genus Woitkowskia was described by Enzmann (1952) for two neotropical species. It was
synonymized by Borgmeier (1955) with Neivamyrmex and its type species, W. connectens Enzmann,
synonymized with N. walkerii (Westwood, 1842); the status of W. mexicana Enzmann, 1952, was
left unresolved. As the name implies, W. mexicana was described from a single male from Tuxtla
Gutierrez, Chiapas, Mexico; the present location of the type is unknown. Despite its verbosity the
original description is mostly worthless and the accompanying figures are inept. It is not surprising
that subsequent workers have largely avoided dealing with this taxon. Watkins, in his several papers
on North American Neivamyrmex ignored this species altogether. Bolton (1995) merely noted that
Enzmann's name was an unresolved junior seconday homonym of N. mexicanus (F. Smith, 1859).
Worthless as Enzmann's description and figure of W. mexicana appear to be, it is possible to

match them to males of N. halidaii, a species that is common at the type locality of W. mexicana.
Accordingly we have synonymized that name here because there appears to be no reason not to do
so. Unless the type is some day located and proven not to be a synonym of N. hallidaii, this seems
the best way to deal with Enzmann's name.
Neivamyrmex harrisii (Haldemann)

Figures 6, 20, 36, 66, 79
Labidus harrisii Haldemann, 1852: 367 (m). USA: Texas, Fort Gates (no types known to exist).
Eciton (Labidus) harrisi: Mayr, 1886c: 441.
Eciton harrisi: Forel, 1899: 28 (m).
Eciton (Acamatus) harrisi: Emery, 1900: 515; fig. 18.
Eciton wheeleri Emery, 1901: 55, fig. 8 (w). USA: Texas, Hays Co. (MCSN).
Eciton (Acamatus) wheeleri: Wheeleri, 1908: 412.
Eciton (Acamatus) wheeleri subsp. dubia Creighton, 1932: 75-77 (w, q). U.S.A., Texas, 5 mi W Ft. Worth
(AMNH?).
Eciton (Neivamyrmex) wheeleri: M. R. Smith, 1942: 561. Creighton, 1950: 76.
Eciton (Neivamyrmex) harrisii: M. R. Smith, 1942: 572. Creighton, 1950: 72.
Neivamyrmex wheeleri: Borgmeier, 1953: 6. 1955: 511-514; (q, w).
Neivamyrmex harrisi: Borgmeier, 1955: 522-524. Watkins, 1968: 275.
UNITED STATES: Oklahoma and Texas, west to Arizona; MEXICO: border states south at
least to Colima in the east and Nayarit in the west.
SPECIMENS EXAMINED
We have studied 141 samples of this common species from the United States and Mexico.
DISCUSSION
M. R. Smith (1942) opined that N. wheeleri represented the female castes of N. harrisii. This
suspected synonymy was confirmed by Watkins (1968).
Males of this common species are regularly collected at lights. They are superficially similar to
those of N. andrei but the mandibular shape and the large ocelli will serve to separate them from
that species.
Neivamyrmex kiowapache, new species

Figures 7, 21, 37, 52, 67, 91, 93, 97-99, 126, 138
Neivamyrmex carolinensis: Borgmeier, 1955: 507-511. Misidentification, in part. Watkins, 1976: 16, 27.
Misidentification, in part. Watkins, 1985: 483, 485. Misidentification, in part. MacKay et al., 1985:
611 Misidentification MacKay & MacKay, 2002: 43, 45, 48, (w, m). Misidentification
DIAGNOSIS
Worker: Eye lens present or replaced by eye spot; head slightly longer than broad; antennal
scape barely exceeding level of eye spot; mesosomal dorsum subopaque to opaque; pronotal disc
coarsely punctate between weak longitudinal rugae; pronotal side smooth and shiny; petiole in dorsal
view slightly longer than broad, dorsal node weakly sculptured and shiny; postpetiole about as long
as broad in dorsal view, disc smooth and shiny.
Queen: Color pale yellow, Head about as broad as long; distinct yellow eyes spot present;
preoccipital lobes rounded; clypeal margin straight but slightly concave in middle; Scape thickened
distally, not reaching eye level; subantennal lamella absent. Dorsal surface of propodeum longer
than declining face, smooth and shining with widely scattered coarse punctures.. Petiole
approximately twice as broad as long, posterolateral corners of petiole well developed and angulate.
Male: Head less that 1.5mm wide measured across eyes; smooth and shiny with scattered small
punctures; OOD at least 1.5 x OD. Mandible blade-shaped, approximately parallel-sided and
tapering to blunt tip.
DESCRIPTION
Worker, measurements (mm) (n = 12): HW 0.45-0.81 (0.79); HL 0.55-0.83 (0.82); SL 0.28-

0.51 (0.51); SW 0.08-0.16 (0.16); PW 0.18-0.25 (0.25); PL 0.165-0.28 (0.28); PpL 0.19-0.29 (0.28);
PpW 0.21-0.29 (0.29); HFL 0.40-0.74 (0.71); HFW 0.11-0.21 (0.16). Indices: CI 87-100 (96); FI 21-
27 (23); SI 51-62 (63).
Head: slightly longer than broad, sides weakly convex in frontal view and narrowing above;
vertex corners rounded and preoccipital carina weakly developed at angle. Front of head shiny
between scattered fine punctures; yellow eye spot present. Antennal scape barely exceeding level of
eye spot, about 3.2 x longer than wide. Subantennal lamella prominent across median 0.33 of
antennal fossa. Mandible triangular, upper margin acutely angulate at juncture with masticatory
margin, latter with three or more small indistinct teeth, but tooth at juncture distinct.
Mesosoma: dorsum sub-opaque to opaque, pronotal disc coarsely puncture between weak
longitudinal rugae; pronotal side smooth and shiny. Promesonotal suture indistinct; promesonotum
slightly convex in profile, disc moderately shiny, with sparse coarse punctures, interspaces weakly to
moderately sculptured. Propodeum slightly depressed below mesonotum, disc and sides granulate;
dorsal face about as long as slightly concave declivitous face.
Petiole in dorsal view slightly longer than broad, dorsal face lightly granulate and shiny, sides
weakly longitudinally striate, shiny; subpetiolar tooth minute. Postpetiole about as long as broad in
dorsal view, smooth and shiny.
Gaster smooth and shiny, with abundant long semi-erect hairs.
Queen, measurements (mm) (n = 8): HW 1.21-1.26; HL 1.16-1.24; SL 0.47-0.54; SW 0.18-

0.22; PW 0.61-0.68; ML 0.57-0.64. Indices: CI 105; SI 46.
Head a little broader than long, sides convex in frontal view and dorsolateral corners rounded;
front of head smooth and shiny between numerous small punctures. Clypeal free margin mostly
straight, but slightly concave in middle. Mandible blade-shaped, about half as long as head, with
strong convexity at midpoint of inner margin, tapering to sharp apex. Scape thickened distally, not
reaching level of eye-spot. Frontal carinae low and rounded; frons with deep median groove from
base of clypeus to about level of eye-spot. Subantennal lamella absent.
Mesosoma. Pronotum rounded anteriorly in dorsal view; promesonotal suture indistinct;
promesonotum in side view arched, but mesonotum slightly concave; metanotal suture distinct; all
segments smooth and shiny. Dorsal face of propodeum longer than declivitous face, smooth and
shiny between scattered coarse punctures.
Petiole smooth and shiny, about twice as wide as long; posterolateral corners well developed
and angulate; subpetiolar process well-developed and triangular in profile.
Gaster smooth and shiny.

Pilosity of head consisting of scattered long erect hairs among more numerous shorter curved
hairs; anterior margin of pronotum with scattered curled hairs among more numerous shorter erect
hairs; gastral terga 1-4 with hairs along posterior margins, segment 5 with short hairs over entire
surface.
Color uniformly pale yellow.
Male, measurements (mm) (n = 2): HW 1.1-1.3; HL 0.79-0.81; ML 0.15-0.17; SL 0.42-0.45;

SW 0.18-0.22; EL 0.50-0.54 (0.54); EW 0.37-0.44; OD 0.16-0.18; OOD 0.26-0.29; IOD 0.34; OMD
0.07-0.12; PW 1.135. Indices: CI 150 SI 36
Head about twice as wide as long, dorsolateral corners rounded into preoccipital margin, not
projecting; frons smooth and shiny; surface between lateral ocelli flat. Free margin of clypeus
slightly concave. Frontal carinae sharp below, becoming rounded above, curving lateral to form
moderate swelling above antennal fossae; frontal area with deep elongate groove. Mandible blade-
shaped, approximately parallel-sided and tapering to blunt tip. Scape somewhat pyriform.
Mesosomal dorsum with dense fine punctures and widely scattered coarser punctures, only
slightly shiny; sides shiny between scattered large punctures.
Petiole about twice as wide as long, posterolateral corners broadly rounded; densely punctate
and slightly shiny.
Gaster: segments slightly shiny between dense fine punctures. Subgenital plate shovel-like, with
strongly convex sides, lateral teeth large, median tooth broadly triangular. Paramere blade-shaped in
profile, sides parallel before abruptly angling to sharp narrow tip, with broad shoulder at angle.
Aedeagus in profile with apically rounded posterodorsal lobe and slightly longer out-turned sharply
pointed posteroventral lobe. Volsella narrow basad, becoming broader at slightly angulate preapical
curve; outer face without apical tubercles; numerous hairs along ventral margin and outer face.
Color: Head, mesosoma and appendages black; gaster reddish brown; wings clear brown.
TYPE MATERIAL
Holotype and numerous paratypes, U.S.A., Colorado, Jefferson Co., Red Rock Park, 5 May
1999 (L. Davis). Holotype and most paratypes in MCZC; paratypes in BMNH, CASC, LACM,
LDPC, UCDC, USNM.
ADDITIONAL SPECIMENS EXAMINED (not types)
UNITED STATES, ARIZONA, Cochise Co.: Cochise Stronghold, Dragoon Mts. (WPMC); 1.8
mi S Southwest Research Station, Chiricahua Mts. (ASUC, LACM). Gila Co.: Hwy. 288 @ Honey
Creek Divide, Sierra Ancha (ASUC, LACM). Navajo Co.: Hwy. 60 @ Corduroy Creek (ASUC,
LACM). Santa Cruz Co.: Madera Cyn. (LACM). Yavapai Co.: Yarnell (ASUC). KANSAS,
Pottawatomie Co.: Little Gobi Desert (BPBM). Riley Co.: Kansas Hill (USNM); N of Tuttle Creek
Dam (Bill Clark, LACM); . Wallace Co.: 10 mi N, 14 mi W Sharon Spring (LACM, Bill Clark).
NEW MEXICO, Mora Co.: 12 km N Wagon Mound (TAMU). Taos Co.: 14 km S Tres Piedras
(WPMC). TEXAS, Sabine Co.: 14.5 km E Hemphill (WPMC).
ETYMOLOGY
The name is derived from a combination of the names of two great Native American nations, the
Kiowa of the midwestern states and the Apache of the southwestern states, a reflection of the broad
distribution of this species.
UNITED STATES, Kansas and Colorado, south to Texas, west to Arizona; MEXICO,
Chihuahua.
DISCUSSION
This is the western form that had been identified as N. carolinensis by previous authors. It
differs from that species in the distinctly smaller size in all castes, but most notably the sexual forms.
In addition to the smaller size, the males differ from those of N. carolinensis by the relatively shorter
and broader, more strongly pyriform, antennal scape. The queen differs from that of N. carolinensis
by its smaller size, the presence of a distinct yellow eye spot and a prominent subpetiolar process.
These differences are not great, but they are consistent. That workers of N. carolinensis and N.
kiowapache are very similar is without question. In view of this similarity we would be hesitant to
describe the western form based solely on the worker caste. We are fortunate to have the sexual
castes as well and the differences, particularly in size, are consistent and non-overlapping and non-
convergent. The westernmost samples that we have seen of N. carolinensis (from Mississippi and
Arkansas) and the easternmost samples of N. kiowapache (from Kansas and eastern Texas)
demonstrate no tendency toward intergradation in size or other features. Therefore, for the present
we recognize the western form, N. kiowapache, as distinct from the eastern ant, N. carolinensis in
the absence of any evidence that they are conspecific
Specimens from Nebraska and Louisiana were recorded by M. R. Smith (1942) as N.
carolinensis, but we have been unable to locate these specimens and cannot comment with certainty
as to their identity. Based solely on distribution, we assume that the Nebraska record will prove to be
referable to N. kiowapache. The Louisiana record remains problematic until the specimens can be
examined.
Rettenmeyer & Watkins (1978) noted the presence of multiple queens in a colony from Kansas
identified as N. carolinensis. We have examined samples from this colony and find that they belong
to N. kiowapache. In addition to the Kansas material cited in the paper, there were four samples from
Georgia and South Carolina. We presume these samples to be true N. carolinensis, but were unable
to examine the material during the course of this study, This phenomenon may be common, if not
consistent, in both N. carolinensis and N. kiowapache judging from the number of occurrences
reported.
Neivamyrmex leonardi (W. M. Wheeler)

Figures 27, 38, 53
Eciton (Acamatus) leonardi Wheeler, 1915: 392 (w). USA, California, Point Loma (MCZC) examined.
Eciton (Acamatus) peninsulare Mann, 1926: 98 (w). MEXICO: Baja California [Sur], La Palma Davila
(USNM) examined.
Eciton (Neivamyrmex) leonardi: M. R. Smith, 1942: 570 (w). Creighton, 1950: 72 (w).
Eciton (Neivamyrmex) peninsulare: Borgmeier, 1949: 101 (w).
Neivamyrmex leonardi: Borgmeier, 1955: 431-434 (w). Watkins, 1971: 101-103 (w). Wheeler & Wheeler,
1973: 38 (w). Watkins, 1976: 13, 18 (w). Cokendolpher & Francke, 1990: 11.
UNITED STATES: Oklahoma and Texas, west to California and Nevada; MEXICO, Baja
California Sur and Tamaulipas south to Guerrero.
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Pima Co.: Tucson, Prince Rd. (UNAR). Santa Cruz Co.:
Nogales (LACM). CALIFORNIA, Inyo Co.: Saline Valley Rd., 1.7 mi. W Grapevine Canyon Rd.
(36º30.7"N 117º33.3"W) (GCSC, CUIC, LACM, UCDC); 2 km ESE Furnace Creek Inn, Death
Valley Natl. Pk. (36º26'N 116º50'W), 100m,ex Solenopsis xyloni middens (UCDC); 9-mile Canyon
(35.85ºN 117.90ºW), 3400 ft. (LACM); 7 mi. S. Little Lake (LACM); 13.44 km SSE Lone Pine, ex
Messor pergandei midden (LACM). Los Angeles Co.: Santa Monica Mtns., Stunt Ranch (34º06'N
118º39'W), 410m (LACM, UCDC). Riverside Co.: Deep Canyon, 1300 ft. (UCRC); Deep Cyn.,
5200 ft. (UNAR). San Bernardino Co.: “Barstow Fossil Beds” (LACM)); 4 mi. NNW Adelanto
(LACM). San Diego Co.: Point Loma (USNM). NEVADA, Churchill Co.: Sand Springs Dune
1200m 39º17'N 118º25' W (UCDC). Lyon Co.: Weeks 1280m (39º18'N 119º15'W), 1280m (UCDC).
TEXAS, Mclennan Co. (SEMC). Bosque Co. (SEMC).
MEXICO, GUERRERO, 29 km NE Fila de Caballo (GCSC, WPMC).
DISCUSSION
This is a poorly known species. The synonymous form N. peninsulare was described from a few
workers taken under a stone. Watkins (1971) also reported finding it under a stone in Texas and
“...observed a few workers...in a weak nocturnal raiding column of N. opacithorax...” Our specimens
from 9-mile Canyon were found in soft sand at a depth of about 45 cm. This was in Sagebrush
Desert with some intermixed Larrea. This ant has also been collected from Creosote Bush Scrub-
Joshua Tree Woodland. In the chaparral biome of Deep Canyon, Wheeler and Wheeler (1973) found
a nest under a slightly buried stone.
Mann (1925) described the staphylinid beetle Pulicomorpha coecum from specimens collected
with a colony of N. leonardi (as Eciton peninsularis Mann) in Baja California Sur, Mexico.
The queen and male forms are unknown; however based on range data we surmise that N. minor
is probably the male form of N. leonardi.
Automontage images of N. leonardi may be viewed at antweb.org.
Neivamyrmex mandibularis (M. R. Smith) NEW STATUS

Figures 100, 101,127
Eciton (Neivamyrmex) pilosus subsp. mandibulare M. R. Smith, 1942: 548 (m). U.S.A., Arizona, 30 mi E
Quijotoa, Pima Co. (USNM) examined. Creighton, 1950: 68, 76.
Neivamyrmex pilosus subsp. mandibularis: Borgmeier, 1955: 375 (m).
UNITED STATES: Arizona and New Mexico; MEXICO: border states south to Chiapas
and Colima.
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Pima Co.: Santa Rita Mts., 4000 ft. (USNM); Forestry Cabin,
Brown Canyon, Baboquivari Mts., 3500 ft. (LACM); Baboquivari Mts. (LACM). Santa Cruz Co.:
Peña Blanca (LACM); Madera Canyon, Santa Rita Mts. (CASC). NEW MEXICO, Catron Co.:
Whitewater Creek, 5 mi NE Glenwood (SEMC). Grant Co.: 100 km NW Silver City (WPMC).
MEXICO, CHIAPAS, Tuxtla Gutierrez, 1000 ft. (CUIC). DURANGO, Nombre de Dios
(UCBC). JALISCO, Chamela, 3000 ft. (USNM). OAXACA, 1 mi SE Tamazulapám, 6200 ft.
(UCBC). SINALOA, Mazatlán (USNM); 5 mi N Mazatlán (UCBC). VERA CRUZ, Fortín de las
Flores, 3000 ft. (CUIC); 8 mi S Elota (UCDC).
DISCUSSION
When M. R. Smith (1942) described this species as a subspecies of N. pilosus he did so

primarily on similarities in the shape of the mandible. This difference is consistent throughout the
range of this species, a range that lies almost entirely within the much more extensive range of N.
pilosus. Were this the only difference between the two, there is no doubt that they would be regarded
as conspecific. However, there is a difference, too, in the shape of the paramere. Although quite
variable in shape, the paramere of N. pilosus (Figs. 128) is narrowly rounded at the apex; in N.
mandibularis the paramere is blunt and broadly rounded. Additionally, the hairs of the compound
eyes in N. mandibularis are generally longer and more flexuous than they are in most populations of
N. pilosus, but in areas where both forms are present, they are more consistent in this difference.
Workers of N. pilosus are relatively large and are conspicuous when that species is present. No
workers of N. pilosus have been found in southern Arizona, even though this is one of the most
heavily collected areas for ants in the United States. While absence of proof is not proof of absence,
we have allowed this consideration to influence our thinking. Additionally, workers of another
species, N. melanocephalus, are available as a possible match for N. mandibularis. At present, we
are reluctant to do more than suggest this match-up since recently examined material of N. graciellae
(Mann, 1926), has raised the possibility that this might be the worker of N. mandibularis. That
species was originally described from Ototonilco, Jalisco, Mexico, well within the range of N.
mandibularis, and this species is another member of the N. pilosus group. For the present, then, we
leave the question unresolved. But, in any case, we have concluded that N. mandibularis is best
regarded as a species apart from N. pilosus.
Neivamyrmex melanocephalus (Emery)

Figures 28, 80
Eciton (Acamatus) melanocephalum Emery, 1895: 260 (w). MEXICO, Tepic (MHNG).
Eciton (Neivamyrmex) melanocephalum: M. R. Smith, 1942: 549 (w) (part). Creighton, 1950: 72 (part).
Neivamyrmex melanocephalus: Borgmeier, 1953: 19. Borgmeier, 1955: 385-387 (w).
Eciton (Acamatus) melanocephalum subsp. xipe Wheeler, 1914: 41; worker. MEXICO, Hidalgo, San
Miguel (MCZC).
UNITED STATES: Arizona; MEXICO: south at least to Oaxaca and Morelos.

SPECIMENS EXAMINED
UNITED STATES, ARIZONA: Santa Cruz Co.: Upper White Rock Campground, Peña Blanca
Lake (31.39° N 111.08° W), 1200m (GCSC, LACM); Nogales, on sidewalk (USNM).
MEXICO, OAXACA: 14.5km S Ocotlán, 1500m (GCSC, WPMC). MORELOS: La Luz
Tetecala (WPMC).
DISCUSSION
This rather large distinctive species is only rarely encountered in the United States and its
principal range lies in Mexico and Guatemala. As discussed above, we believe that N. mandibularis
is the likely male of N. melanocephalus, but it is also possible that N. mandibularis could be the
opposite sex of N. graciellae.
One foraging raid of N. melanocephalus was observed. It occurred in full daylight and the
workers were taking a variety of small arthropods (B. V. Brown, pers. comm).
Neivamyrmex melshaemeri (Haldemann, 1852)

Figures 115, 128, 139
Labidus melshaemeri Haldemann, 1852: 368; pl. 9 fig. 7-9 (m). U.S.A, Texas, Fort Gates (type lost).
Eciton (Labidus) melshaemeri: Mayr, 1886c: 442. Emery, 1895: 261 (m). Emery, 1896: 33.
Eciton (Acamatus) melshaemeri: Emery, 1900: 516, 525 (m). Emery, 1910: 26.
Eciton melsheimeri: Forel, 1899: 28 (m).
Eciton (Acamatus) melshaemeri: Wheeler, 1908: 418 (m).
Eciton (Neivamyrmex) melshaemeri: M. R. Smith, 1942: 576 (m). M. R. Smith, 1951:780. Creighton,
1950: 73.
Neivamyrmex melsheimeri: Borgmeier, 1955: 647-649 (m). Watkins, 1972: 351 (m). Watkins, 1976: 23
(m); Watkins, 1982: 213 (m).
UNITED STATES: Louisiana and Oklahoma west to Texas; MEXICO: Tamaulipas and San
Luis Potosí south to Chiapas and Yucatán; SOUTH AMERICA.
SPECIMENS EXAMINED
UNITED STATES, LOUISIANA, Beauregard Parish: DeRidder (LACM). TEXAS, Bexar

Co.: San Antonio (CASC). Brazos Co.: College Station (TAMU, WPMC); 16 km W Bryan
(WPMC). Cameron Co.: nr. Southmost Sabal Palm Grove Sanctuary (TAMU, WPMC); Brownsville
(LACM, GCSC, UCBC, USNM). Dimmit Co.: Winter Haven (MINN). Grayson Co., Sherman
(TAMU). Liberty Co.: Devers (CUIC). Matagorda Co.: Blessing (TAMU). Morris Co.: Dangerfield
(TAMU). Tarrant Co.: Fort Worth (LACM). Travis Co.: Austin (TAMU). Val Verde Co.: Juno
(CUIC). Victoria Co.: Victoria (CUIC). Wharton Co.: Wharton (CUIC, LACM).
MEXICO, CHIAPAS: 12 mi N. Ocozocoautla (TAMU). TAMAULIPAS: Ciudad Victoria
(WPMC); Ciudad Victoria, Canon La Libertad (TAMU).
DISCUSSION
This common small species can be confused with no other United States species. The long
flexuous hairs on the body of this ant readily distinguish it from our other Neivamyrmex. Throughout
its range from the southern United States to South America this ant is morphologically quite
uniform.
Neivamyrmex microps Borgmeier, 1955

Figures 8, 39, 54, 68, 81, 102, 103, 129, 140
Neivamyrmex microps Borgmeier, 1955: 635 (m). U.S.A., Arizona, Phoenix (USNM). Watkins, 1972:
352. Watkins, 1976: 25.
The worker caste has not been previously described.
DIAGNOSIS
Worker. Eye absent; antennal scape not extending to midlength of head; head with numerous
suberect hairs, but no erect hairs; propodeal dorsum smooth and shiny, distinctly longer than
declivitous face, latter smooth and shiny; petiole node longer than broad in dorsal view, sides
slightly convex; postpetiole slightly wider than long, disc smooth and shiny, sides weakly
sculptured.
DESCRIPTION
Worker, measurements (mm) (n = 12): HW 0.59; HL 0.73; SL 0.34; SW 0.11; PW 0.22; ML

0.36; PL 0.27; PpW 0.29; PpL 0.22; HFL 0.50; HFW 0.15. Indices: CI 70-85 FI 28-39; SI 39-51
Head distinctly longer than broad; smooth and shiny with only scattered small punctures. Eye
absent. Dorsolateral corners slightly angulate, preoccipital carina weak. Scape not reaching middle
of head length, 3 x as long as greatest breadth, broad distad, abruptly narrowed at base. Subantennal
lamella well developed. Head without erect hairs, but with numerous suberect hairs. Mandible
triangular, upper margin straight and distinctly angulate at juncture with masticatory margin; upper
margin with distinct small distal tooth (Fig. 7).
Mesosomal dorsum smooth and shiny; promesonotal suture indistinct; metanotal groove distinct.
Propodeal dorsum smooth and shiny, distinctly longer than declivitous face, latter smooth and shiny.
Metafemur stout.
Petiole node longer than broad in dorsal view with sides slightly convex and narrowing
anteriorly; evenly convex in lateral view; sides slightly granulate; subpetiolar tooth minute.
Postpetiole slightly wider than long, dorsum smooth and shiny, sides lightly granulopunctate.
Gaster smooth and shiny on first tergum, second segment weakly sculptured and less shiny.
UNITED STATES: Known only from Arizona, but almost certainly extends into adjacent
eastern California and south into MEXICO.
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Maricopa Co.: Phoenix (USNM) (m); Scottsdale (GCSC,
LAAG, LACM) (m); 4 Peaks Rd. (Mazatzal Mts.), 5 mi E Hwy. 87 (RAJC, GCSC, LACM) (w).
Pinal Co.: Red Rock (34.30°N 110.19°W (ARSU) (m). Pima Co.: Tucson, in house (UNAR) (m).
Santa Cruz Co.: Madera Canyon, 4000', Santa Rita Mts. (LACM) (m). Yuma Co.: Hoodoo Well,
Kofa Mts. (GCSC) (m).
DISCUSSION
Neivamyrmex microps is one of a number of species, the males of which all have a distally
bifurcate volsella, that possess minute tubercles on the outer face of the volsella (much as in Fig.
140). The number and distribution of these tubercles may be consistent and characteristic from
species to species, but we have not investigated this matter in detail, nor have we examined all
species with bifurcate volsellae. At this stage we can only stipulate that they are not present in all
species with bifurcate volsellae. Among the United States species we can affirm their presence in
males of N. microps, N. minor, N. ndeh and N. swainsonii. They are absent in males of N. andrei and
N. fuscipennis. In the case of N. microps these tubercles are sufficiently numerous that three or four
are visible within the apical crotch as minute teeth.
In the original description of this species Borgmeier incorrectly cited the collector of the type
specimen as “R. H. Randall”; the correct name is R. H. Crandall. For many years known only from
the type specimen, this species has recently been collected again in a number of Arizona localities.
Although we now know that N. microps has a fairly extensive range within Arizona, it will likely
continue to be a rarely collected species. Current records indicate that this species inhabits a wide
range of habitats, having been found in areas ranging from Creosote scrub to Oak woodland, with an
elevational range between 1000 and 4000 feet.
Based on circumstantial evidence, we have associated the above described workers with this
species. They were collected in Maricopa County, along 4 Peaks Road by R. A. Johnson, 27 March
1993. These workers belong to the correct species group and and the known distribution makes this
association probable. Workers may be confused with those of N. leonardi, but may be differentiated
by the less well developed mandibular teeth, the longer, narrower postpetiole, and the less convex
mesosomal profile.
Nothing is known of the biology of this species. However, label data indicate that the activity
period for this species closely coincides with the summer/fall monsoon season in Arizona.
Neivamyrmex minor (Cresson)

Figures 104, 105, 116, 130, 141
Labidus minor Cresson, 1872: 195 (m). U.S.A., Texas (ANSP).

Eciton (Labidus) minor: Mayr, 1886: 441 (m). Emery, 1895: 261 (m).
Eciton (Acamatus) minus: Wheeler, 1908: 418 (m).
Eciton (Neivamyrmex) minus: M. R. Smith, 1942: 544 (m). Creighton, 1950: 73 (m).
Neivamyrmex minor: Borgmeier, 1955: 630-632 (m). Watkins, 1976: 25 (m). Wheeler & Wheeler, 1986:
20. Cokendolpher & Francke, 1990: 11.
UNITED STATES: Texas to Kansas, west to southern Nevada and California; MEXICO: Baja
California, Baja California Sur, Coahuila (Watkins, 1982).
SPECIMENS EXAMINED
We have 64 records from the United States.
DISCUSSION
Neivamyrmex minor is certainly one of the more widespread species. Occurring in a wide
variety of habitats it is one of the most frequently encountered Neivamyrmex species, and is
commonly collected at light. The female castes are currently unknown but, based on ranges, we posit
that N. leonardi will prove to be the worker.
Although this species has been collected in variety of habitats, it has only rarely been collected
in the desert areas of the west. Elevational amplitude ranges from near sea level to about 7000 feet.
Automontage images of this ant are available at antweb.org.
Neivamyrmex mojave (M. R. Smith)

Figures 117, 131, 142
Eciton (Neivamyrmex) mojave M. R. Smith, 1943: 196 (m). U.S.A., California, Mojave Desert (USNM)
examined. Creighton, 1950: 73 (m).
Neivamyrmex mojave: Borgmeier, 1955: 633-635 (m). Watkins, 1976: 25; map 47 (m).
DISTRIBUTION (Map. 5)
UNITED STATES: Southern California; MEXICO: Baja California Sur.
SPECIMENS EXAMINED
UNITED STATES, CALIFORNIA, Riverside Co.: Riverside (LACM). San Bernardino Co.:
Mojave Desert (USNM); 4 mi. NNW Adelanto 2900' (LACM).
MEXICO, BAJA CALIFORNIA SUR: 8.4 mi W on Ramal á Los Naranjo, at light (LACM).
DISCUSSION
This species has been previously known only from the type specimen, from an unknown locality
in the Mojave Desert. The specimen from north of Adelanto was picked up dead on the ground.
Neivamyrmex mojave is evidently nearest to N. microps and superficially looks much like that
ant. However it may be distinguished from that species by the non-toothed volsella, and differences
in coloration. This species might also be mistaken for N. minor but may be readily distinguished
from that species by the small size of the ocelli.
The recently described N. nyensis Watkins may be the worker form of this species.
Automontage images of this species are available at antweb.org
Neivamyrmex moseri Watkins

Figures 9, 22, 40, 55, 82
Neivamyrmex moseri Watkins, 1969: 529 (w). U.S.A., Louisiana, Rapides Parish, Kisatchue National Forest
(USNM) examined.
Neivamyrmex isodentatus MacKay, 1998: 333-335 (w). U.S.A., Texas, Kleberg Co., Kleberg Airport
(MCZC) examined. NEW SYNONYMY.
UNITED STATES: Louisiana and Texas.
SPECIMENS EXAMINED
In addition to the type material cited above, we have studied a small series from Texas, Bell Co.,
Bowmer Ranch 18 June 1970 (J. F. Watkins II; W164; LACM, WPMC).
DISCUSSION
Neivamyrmex moseri is easily separated from all other described United States species by its
unique mandibular structure: there are three large teeth that are usually approximately equal in size;
much smaller intercalary denticles may be present. MacKay (1995) described N. isodentatus as a
similar species that differed in details of mesosomal and petiolar sculpture, presence of a subantennal
lamella, fewer short bristly hairs on the funiculus, and a subtle difference in the direction of the
opening of the propodeal spiracle. None of these features is sufficiently distinctive to suggest a new
species, since all are features that are typically variable in any species of Neivamyrmex. The types of
N. isodentatus were compared with a few specimens from Bell County, Texas, identified by Watkins
as N. moseri, but not with the types of that species. Had the N. isodentatus material been compared
with N. moseri types, the conformity of the two would have been obvious. Both type series differ
from the Bell County specimens in the same manner and we conclude that N. isodentatus is a
synonym of N. moseri.
With the limited material available we are forced to agree with Watkins that the Bell County
specimens are also N. moseri, even though differing in a few subtle features that are variable and
somewhat illusory, depending upon the angle of view. We should note further that the statement by
Watkins (1969) that N. moseri workers lack a subantennal lamella is only partly correct; the lateral
extension of the frontal carina that runs below the socket is present, but is not elevated to form a
distinct lamella in most specimens examined. A low lamella is present in a few specimens, but not to
quite the same degree as seen in the types of N. isodentatus.
Neivamyrmex ndeh, new species

Figures 118, 132, 143
DIAGNOSIS
Male: head and mesosoma blackish brown, gaster reddish brown; head 1.5× broader than long
measured across eyes; smooth and shiny between scattered small punctures; mesosoma blackish
brown, entire surface smooth and shining with numerous scattered small punctures, pilosity suberect
and dense. Queen and worker unknown.
DESCRIPTION
Male, measurements (mm) (n = 2): HW 0.86-0.91 (0.91); HL 0.55-0.58 (0.58); SL 0.39-0.41

(0.41); SW 0.15-0.16 (0.15); EL 0.305-0.306 (0.305); EW 0.23-0.26 (0.23); OD 0.09-0.10 (0.09);
OOD 0.15-0.175 (0.175); OMD 0.07-0.12 (0.12); OVD 0.20; PW 0.70 (0.70); ML 0.40 (0.40); PL
0.62-0.70 (0.70). Indices: CI 156-158 (158); SI 49-57 (57).
Head almost 1.6× as broad as long. Dorsal margin, in frontal view, concave and rounded at
sides; preoccipital carina weak and not forming distinct collar. Free clypeal margin (ventral margin)
slightly concave; frontal carina sharp between antennal fossae, but becoming rounded dorsad and
curving laterad to form moderate swelling above antennal fossae. Mandible sickle-shaped and acute
at apex, about 0.40 mm long. EL 1.3× EW. OOD about 1.75× OD. Scape extending to level of upper
eye margin, about 3× longer than broad; subantennal lamella prominent.
Mesosoma smooth and shiny, with numerous scattered small punctures.
Petiole slightly longer than broad; posterolateral corners rounded; dorsal surface shiny, ventral
surface less so.
First three segments of gaster smooth and shiny, following segments duller, more distinctly,
finely punctate. Subgenital plate longer than broad, somewhat spatulate; apicolateral teeth
prominent, median tooth small. Dorsal margin of paramere broadly triangular, margins densely
clothed with long flexuous hairs. Volsella with apical fork, dorsal process large and sharp, ventral
process blunt and much shorter; inner margin each segment with a minute denticle near fork;
numerous long erect hairs along ventral margin. Aedeagus in profile with apically rounded
posterodorsal lobe and slightly longer, out-turned sharply pointed posteroventral lobe.
Color: head and mesosoma blackish brown, first two gastral segments similar, following
segments becoming reddish brown on posterior segments; wings clear brownish.
TYPE MATERIAL (Map 5)

Holotype: U. S. A., Arizona, Santa Cruz Co., Yanks Canyon (31.42°N 111.17°W), 12-15 Aug.
1993 (B.V. Brown and D. Feener). Paratype: U. S. A., Arizona, Cochise Co., Portal, 4800 ft., 4
Aug. 1959 (H. E. Evans). Both specimens in LACM.
ETYMOLOGY
This species is named for the Ndeh or Apache people of the southwestern United States and
adjacent Mexico; the name pronounced approximately “in déh”, accenting the second syllable.
DISCUSSION
This minute species, currently known only from the two specimens cited above, is close in size
only to N. baylori and should be readily identifiable on that basis alone. The holotype was taken in a
Malaise trap and the Portal specimen was presumably taken at black light. The worker caste of this
distinct species is unknown but is certainly likely to be another minute species; possibly it is N.
goyahkla, described above.
Neivamyrmex ndeh is superficially similar to N. microps, but is significantly smaller and with
distinctive genitalic features: the crotch of the apical fork of the volsella has only two barely
perceptible teeth that are well removed from each other (see discussion of N. microps).
Neivamyrmex nigrescens (Cresson, 1872)

Figures 10, 29, 41, 56, 69, 83, 92, 106, 107, 119, 133. 144
Labidus nigrescens Cresson, 1872: 194 (m). U.S.A., Texas (ANSP).

Eciton nigrescens: Dalla Torre, 1893: 5.
Eciton sumichrasti: Mayr, 1886a: 120. Mayr, 1886b: 440 (in part). Forel, 1899: 27 (in part). Wheeler,
1900: 563, fig. 1-3 (w, q). Wheeler & Long, 1901: 160, note 2. Mann, 1926: 99-100 (q).
Misidentification
Eciton (Acamatus) schmitti Emery, 1894: 183 (w). U.S.A. Missouri, Ripley Co., Doniphan (MCSN).
Emery, 1895: 258 (w). Forel, 1899: 28. Wheeler & Long, 1901: 161 (m). Wheeler, 1908c: 410 (w, m).
M. R. Smith, 1924: 84. M. R. Smith, 1927: 401-404. Borgmeier, 1936: 59. M. R. Smith, 1938: 160. G.
Wheeler, 1942: 331.
Eciton (Labidus) nigrescens: Emery, 1895: 261.
Eciton (Acamatus) nigrescens: Emery, 1900: 517, 525. Wheeler, 1908c: 417; pl. 26 fig. 2 (m). Emery,
1910b: 27. M. R. Smith, 1938: 157 (m).
Eciton (Neivamyrmex) nigrescens: M. R. Smith, 1942: 551; fig. 4, 23 (w, q, m) (part). Borgmeier, 1948:
193. Creighton, 1950: 66, 69, 73-74; pl. 12 (w, q, m).
Eciton (Neivamyrmex) californicum: Creighton, 1950: 70 (part).
Neivamyrmex nigrescens: Borgmeier, 1955: 494-501 (w, q, m) (part). Watkins, 1972: 358-363 (w, q, m).
Wheeler & Wheeler, 1973: 37, 38-40 (w, q, m). Watkins, 1976: 15, 22 (w, q, m). Cokendolpher &
Francke, 1990: 12. Allred, 1982: 492. Wheeler & Wheeler, 1986: 20. Ward, 1999: 74-97.
Neivamyrmex californicus: Watkins, 1972: 363 (part); Watkins, 1985: 482 (part).
UNITED STATES: transcontinental across southern states, north to West Virginia, Tennessee,
Colorado and Nebraska; MEXICO: Baja California, Nayarit, Oaxaca, Sonora (Watkins, 1982).
SPECIMENS EXAMINED
We have 133 records from the United States.

DISCUSSION
This common species is by far the most widespread species in the United States. As a result of
this wide range it is also by far the most studied and best known of the Neivamyrmex. Not
surprisingly, given such an extensive range, N. nigrescens also shows an extremely wide habitat
range. Rarely encountered in desert environments it is nonetheless present, apparently largely
confined to canyons and hillsides. Ward (1999) listed the elevational range from sea level to 1460m
in California, but we have records up to 2200m in areas outside of California.
Automontage images of the worker caste may be viewed at antweb.org.
Colonies studied by Wheeler (1900) consisted of “thousands” of individuals, while Schneirla
(1958) estimated 150,000 to 250,000 workers per nest.
Neivamyrmex nigrescens has a nomadic/statary cycle like Neotropical army ants such as Eciton.
The nomadic phase of the cycle begins when pupae eclose to workers. The whole colony then moves
along a trail, usually during night hours, capturing any insects they encounter and raiding the nests
of other ant species encountered. Columns may be 90m long and are headed by scouts. The colony
bivouacs before dawn, using natural cavities or nests of other species, which they have pillaged. The
following night they again move and raid. This nomadic cycle lasts for about three weeks or until the
larvae in the colony (which they transport each night) begin to pupate. The statary phase then begins
and the ants nest in subterranean cavities, either under stones or in abandoned ant nests for about 18
days (Schneirla, 1958). Raids continue but are less extensive than during the nomadic phase.
New colonies of N. nigrescens are formed when “a daughter queen leaves the parental nest,
accompanied by a number of workers. A mature colony is capable of producing a small number of
females, some of which may be fertilized in the nest by their brothers, but this does not preclude
mating outside the nest, or with males of other colonies. Since females are never winged, they can
make no nuptial flight.” (Smith 1965). Recent very preliminary data for this species suggest that N.
nigrescens may, at least at times, have more than one functional queen present in the colony.(D.
Kronauer, pers. comm.)
Other ants form an important part of the diet of N. nigrescens. Mallis (1938) observed this
species carrying larvae and pupae of Tetramorium caespitum (Linné), as well as click beetles,
mayflies, water boatmen and crickets. Wheeler & Long (1901) found larvae of Solenopsis geminata
(Fabr.) and three species of Pheidole, as well as dead carabid beetles, in nests they studied in Texas.
Ward (1999) further notes that in California Messor andrei (Mayr), Pheidole californica Mayr, P.
hyatti Emery, Solenopsis molesta (Say) and Formica moki Wheeler are also prey items of this
species. In Arizona N. nigrescens has been observed regularly raiding Pheidole obtusospinosa
Pergande (as P. subdentata) and Pheidole desertorum Wheeler. Neece & Bartell (1982) noted the
presence of unidentified mites of the family Trachyaropodidae in colonies of N. nigrescens.
The blind snake, Leptotyphlops dulcis, is able to follow the pheromone trails of N. nigrescens to
locate columns and feed on the ant brood (Watkins et al., 1967). When the army ants attack the
snake it forms a protective ball-like coil and smears a cloacal fluid on its body, which discourages
further ant attacks (Watkins et al., 1972).
Several species of scuttle flies (Diptera: Phoridae) are known to parasitize adults of N.
nigrescens. These include species in the genera Dacnophora and Cremersia (B. V. Brown, pers.
comm.), and Xanionotum (Rettenmeyer and Akre 1968). The diapriid wasp, Ecitovagus gibbus
Masner has been found as a parasitoid of N. nigrescens in southeastern Arizona (Masner 1977).
Myrmecophilous Staphylinidae (Coleoptera) associated with this ant in areas other than California
include: Microdonia laticollis Brues, M. nitidiventris Brues, M. occipitalis Casey, Ecitoxenidia
brevicornis Seevers, E. brevipes Brues, Dinocoryna carolinensis Seevers, and Ecitonidia wheeleri
Wasmann (Seevers 1965).
At least two species in the carabid beetle genus Helluomorphoides (H. ferrugineus Casey and H.
latitarsis LeConte) are specialized predators on both the booty and brood of N. nigrescens in
southeastern Arizona: “The beetles were observed running in army ant columns or standing off to the
sides of the columns, behind rocks or beneath clusters of leaf litter. During their predatory activities,
beetles ran along the trails in both directions, 'plowing' through the continuous two-way ant traffic.
When a beetle of either species contacted a worker ant bringing booty back to her bivouac, the ant
usually dropped the booty. On some occasions, if the booty was a larval or pupal individual of
another ant species, the beetle immediately ate it and continued on the trail. On other occasions the
beetle picked up the dropped booty, left the raiding column, and proceeded to a nearby rock. There,
the beetle quickly ate the larva or pupa, returned to the column, and resumed running along the trail”
(Topoff, 1969). Beetles were observed to forcibly take booty from the ants. The beetles were also
seen to feed on brood caches of the Neivamyrmex colony with which they became associated.
For further information on the biology and behavior of this species, see Ward (1999).
Neivamyrmex nyensis Watkins

Figures 11, 23, 42, 57, 70, 84
Neivamyrmex nyensis Watkins, 1977: 421-423 (w). U.S.A., Nevada, Nye Co., 5 1/2mi S, 3 mi E Beatty
(USNM) examined. Wheeler & Wheeler, 1986: 20.
UNITED STATES: Southern Nevada, Arizona, California; MEXICO: Baja California.
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Coconino Co.: Havasu Canyon (UCDC, LACM).

CALIFORNIA, Imperial Co.: 2.9 mi N Glamis (LACM). Riverside Co.: Chuckwalla Mts., Red
Cloud Cyn. (LACM). San Bernardino Co.: Lucerne Valley (GCSC). NEVADA, Nye Co.: 5 1/2 mi
S, 3 mi. E Beatty 3500 ft (LACM, USNM).
MEXICO, BAJA CALIFORNIA: 28 km E Ensenada (LACM, UCDC).
DISCUSSION
This small subterranean species is poorly known. Not surprisingly most of these collections
were from under partially buried stones; the Glamis sample consists of a single specimen taken from
a foraging worker of Forelius maccooki (Forel) and the Lucerne Valley record is based on a head
capsule recovered from a refuse midden of Solenopsis xyloni McCook. The Red Cloud Canyon site
is in a rocky streambed on the west side of the range, in Creosote/Cactus Scrub desert. The
specimens were collected in late afternoon beneath a partially buried stone. There had been rain
several days prior to the collection and the soil under the stone was slightly damp. The Havasu
Canyon collection was made under similar circumstances.
Neivamyrmex nyensis may eventually prove to be the worker form of N. mojave.
Automontage images are available at antweb.org.
Neivamyrmex opacithorax (Emery)

Figures 108, 109, 120, 134, 145
Eciton (Acamatus) californicum subsp. opacithorax Emery, 1894: 184 (w, in key). U.S.A., Missouri,
Doniphan (MHNG). Emery, 1895b: 259. Pergande, 1895: 874. Forel, 1899: 28.
Eciton (Acamatus) opacithorax: Emery, 1900a: 524; Emery, 1910: 25. Wheeler & Long, 1901: 163, 173
(w, q). Wheeler, 1908b: 411 (m). M. R. Smith, 1924: 84. Mallis, 1941: 62.
Eciton (Acamatus) carolinense: Wheeler, 1921: 314 (q). Misidentification
Eciton (Neivamyrmex) opacithorax: M. R. Smith, 1932: 555 (w, q, m). M. R. Smith, 1951 [in Muesebeck,
et al.]: 781. Buren, 1944: 180. Creighton, 1950: 74.
Eciton (Acamatus) opacithorax var. castaneum Borgmeier, 1939a: 416 (w). COSTA RICA, San Jose
(MCSN).
Eciton (Acamatus) californicum: Mallis, 1941: 62 (w). Misidentification
Eciton (Neivamyrmex) californicum: M. R. Smith, 1942: 560 (w). Misidentification
Eciton (Neivamyrmex) opacithorax subsp. castaneum: Borgmeier, 1948: 191 (w, q m). COSTA RICA,
San José.
Neivamyrmex opacithorax: Borgmeier, 1953: 6. Watkins, 1972: 349; Watkins, 1976: 16, 22. Watkins,
1985: 482, 484.
UNITED STATES: Virginia and Tennessee, south to Florida, west to California; MEXICO
(Baja California, Jalisco); GUATEMALA; COSTA RICA.
SPECIMENS EXAMINED
We have 51 records from within the United States.
DISCUSSION
Although N. opacithorax is a widespread species it is not as commonly encountered as other

members of the N. nigrescens group. Within the group it is readily recognized by the distinctively
shaped mandible and the shiny head. It is presumably a raider on other ant species. Workers of this
species were discovered during the processing of a number twig cuttings which contained a colony
of Pseudomyrmex championi (Forel) in Guatemala. It is unknown if the Neivamyrmex were actively
entering the colony while it was intact or if the raiding began as the twigs were cut and collected for
processing (P. S. Ward, pers. comm.).
Automontage images of N. opacithorax may be viewed at antweb.org.
Neivamyrmex pauxillus (W.M. Wheeler)

Figures 12, 24, 43, 58, 71, 85
Eciton (Acamatus) pauxillum Wheeler, 1903: 93 (w). U.S.A, Texas, Austin, Paisano Pass (AMNH,
MCZC).
Eciton (Neivamyrmex) pauxillum: M. R. Smith, 1942: 569
Neivamyrmex pauxillus: Borgmeier, 1953: 570-572. Watkins, 1982: 197-247. Watkins, 1985: 479-502.
UNITED STATES: Louisiana and Texas; MEXICO: Hidalgo (Watkins, 1982).
SPECIMENS EXAMINED
UNITED STATES, LOUISIANA, Rapides Parish: Kisatchue National Forest nr. Alexandria
(LACM). TEXAS, Bell Co.: Bowmer Ranch (LACM, WPMC). Brewster Co.: Paisano Pass, nr.
Alpine (MCZC). Travis Co.: Austin (AMNH, MCZC).
DISCUSSION
This is one of several small, nondescript yellow Neivamyrmex species. Because they are rarely
encountered, little is known of their habits. Some success has been had locating these minute species
using underground baiting, a practice that should be frequently employed. They will likely prove to
be subterranean predators of either other ants or of termites.
Neivamyrmex pilosus (F. Smith)

Figures 13a-b, 44, 59, 72, 86, 1110, 111, 121, 135, 146, 150, 151
Eciton pilosa F. Smith, 1858: 151 (w). BRAZIL, Amazonas, Vila Nova (BMNH).
Eciton clavicornis Norton, 1868: 46 (w). MEXICO, Vera Cruz, Hacienda Potrero (types lost?).
Eciton pilosum: Mayr, 1865:77; 1886: 120. Dalla Torre, 1893:5.
Eciton (Labidus) subsulcatum Mayr, 1887: 440 (m). U.S.A., Texas (NHMV).
Eciton mexicanus: Dalla Torre, 1893: 4. Forel, 1899: 27.
Eciton (Acamatus) pilosum: Emery, 1894a: 183. Emery, 1900a: 524; Emery, 1910: 25. Borgmeier, 1923,
48. Borgmeier, 1934: 94. Wheeler, G. C., 1943: 331 (larva).
Eciton (Labidus) mexicanum: Emery, 1895b: 260. Emery, 1896a: 33. Misidentification
Eciton pilosum: Forel, 1899: 27. Wheeler & Long, 1901: 165.
Eciton (Acamatus) mexicanum: Emery, 1900a: 515, 525. Emery, 1910: 26. Wheeler, 1908: 414. M. R.
Smith, 1931: 16.
Eciton aztecum Forel, 1901: 49 (m). GUATEMALA, San José (MHNG).
Eciton (Acamatus) pilosum: Wheeler, 1908: 412. Wheeler, 1921: 313. M. R. Smith, 1924: 85; M. R.
Smith, 1931: 295. Reichensperger, 1939: 297 (q).
Eciton (Acamatus) pilosum var. angustius Forel, 1909: 256 (w). PARAGUAY, San Bernardino (MHNG).
Emery, 1910: 25. Wheeler, 1921: 312.
Eciton (Acamatus) mexicanum var. azteca: Emery, 1910: 26.
Eciton (Acamatus) pilosum var. beebei Wheeler, 1921: 312 (w, m). GUYANA, Kartabo (MCZC). NEW
SYNONYMY.
Eciton (Acamatus) pilosum var. aztecum: Wheeler, 1921: 314.
Eciton (Labidus) spininodis subsp. militarium Santschi, 1929: 85 (w). PANAMA, Bocas del Toro,
Changuinola (BASEL). Borgmeier, 1953: 14.
Eciton (Acamatus) porrectognathum Borgmeier, 1933b: 167 (m). BRAZIL, Rio de Janeiro, Angra dos
Reis u. Itatiaia ( ? ).
Eciton (Acamatus) pilosum var. mexicanum: Borgmeier, 1936: 59. Borgmeier, 1939a: 416.
Misidentification
Eciton (Neivamyrmex) pilosum: M. R. Smith, 1942: 544. Creighton, 1950: 75.
Neivamyrmex pilosus porrectognathum: Borgmeier, 1953: 5. BRAZIL, Angra dos Reis and Itatiaia
(IBVB).
Neivamyrmex pilosus mexicanus: Borgmeier, 1953: 17. Misidentification
Neivamyrmex pilosus beebei: Borgmeier, 1953: 19.
Neivamyrmex pilosus: Borgmeier, 1955: 361-373 (w, q, m).
Neivamyrmex pilosus subsp. beebei: Borgmeier, 1955: 373 (w, m).
Neivamyrmex pilosus subsp. mexicanus: Borgmeier, 1955:374 (w, q, m). Watkins, 1976: 16. Watkins,
1982: 197-247. Watkins, 1985: 479-502. Misidentification
DISTRIBUTION (Map. 2)
UNITED STATES: Arkansas and Mississippi west to southern Arizona; MEXICO: border
states south to Chiapas; south to BRAZIL and PARAGUAY.
SPECIMENS EXAMINED
We have 56 records of this wide-ranging species in the United States.
DISCUSSION
The male type of Labidus mexicanus, described from Orizaba, Mexico by F. Smith (1859), has
been examined. It is a specimen of N. swainsonii (Shuckard), not a form of N. pilosus. Accordingly,
L. mexicanus is removed to the synonymy of that species.
We have examined both males and workers from throughout the extensive range of this species.
We have found that the features distinguishing the several recognized subspecies form a continuum
from north to south, with no clear demarcation for any of these features. Accordingly, all these are
here placed in synonymy.
Watkins (1985) indicated on his map of the distribution of N. pilosus mexicanus that this ant had
been collected from California. We have examined the specimens, purportedly collected at Exeter,
Tulare County, and concur that they are N. pilosus. However, it is our opinion that these males are
probably incorrectly labeled and we have not included California within the distribution of N.
pilosus.
Automontage images of the worker caste (as N. pilosus mexicanus) are located at antweb.org.
Neivamyrmex rugulosus Borgmeier

Figures 14, 45, 60
Neivamyrmex rugulosus Borgmeier, 1953: 49 (w). MEXICO, Jalisco, Zapotlán) (AMNH).

Eciton (Acamatus) schmitti Pergande, 1895: 874. Misidentification
Eciton (Acamatus) sumichrasti Wheeler, 1908: 410. Misidentification
UNITED STATES: Arizona; MEXICO: Jalisco, Nayarit, Sonora (Watkins, 1982).
SPECIMENS EXAMINED
UNITED STATES, ARIZONA, Cochise Co.: 0.3 mi W Southwest Research Station, 5500',
Chiricahua Mts. (LACM, MCZC); Cave Creek, Chiricahua Mts. (LACM, SEMC, USNM, WPMC);
Southwest Research Station, 5400’ (WPMC); 3.5 mi. NW Portal, mesquite foothills, 5450', raiding
Pheidole desertorum (GCSC, LACM, RAJC).
DISCUSSION
This is a poorly known species, which has been collected in the United States only a handful of
times. All of our records are from the Chiricahua Mountains in southeastern Arizona where it has
been collected above the 5000-foot level. Although records are few in the United States it is
probable that the species is more widespread than currently indicated, however this appears to be a
primarily Mexican species that is at the extreme northern limit of its range in the United States. Little
is known of the biology of this species, and the only prey records for it are other ant species:
Trachymyrmex arizonensis (LaPolla et al., 2002) and Pheidole desertorum (R. A. Johnson, pers.
comm.).
LaPolla et al. (2002) mention that a male of N. rugulosus had been collected near Portal,
Arizona. That male is in the LACM collections. We are unable to distinguish between this male and
those of N. harrisi. Since they note that it was attracted to a head lamp, the association of this male
with N. rugulosus appears to be accidental.
Neivamyrmex swainsonii (Shuckard)

Figures 15, 25, 30, 46, 73, 122, 136, 147, 149
Labidus swainsonii Shuckard, 1840: 201 (m). BRAZIL (BMNH). Westwood, 1842: 76. F. Smith, 1859: 8
(m).
Labidus mexicanus F. Smith, 1859: 7 (m). MEXICO, Orizaba (BMNH) (examined). NEW
SYNONYMY.
Eciton (Acamatus) nitens: Pergande, 1895: 874. Misidentification
Eciton (Acamatus) swainsonii: Emery, 1900: 515, 525. Emery, 1910: 27. Santschi, 1916: 370. Santschi,
1931: 74. Gallardo, 1920: 379. Borgmeier, 1923: 50.
Eciton (Acamatus) arizonense Wheeler, 1908c: 414 (m). U.S.A., Arizona, Nogales. (MCZC). Emery
1910:25. M. R. Smith, 1942: 581 (m)
Eciton (Acamatus) lieselaei: Gallardo, 1920: fig. 32. Misidentification
Eciton (Neivamyrmex) swainsoni: Borgmeier, 1948b: 462.
Eciton (Neivamyrmex) arizonense: Creighton, 1950: 69-70 (m).
Neivamyrmex swainsoni: Borgmeier, 1953: 16. Borgmeier, 1955: 454-458 (m). Watkins, 1972: 352 (m).
Watkins, 1976: 24 (m). Cokendolpher & Francke, 1990: 13.
Eciton (Neivamyrmex) commutatum: M. R. Smith, 1942: 568 (misidentification).
Neivamyrmex swainsoni arizonensis: Borgmeier, 1953: 19 (m).
Neivamyrmex fallax Borgmeier, 1953: 48 (w). U.S.A., Texas, Victoria (USNM). NEW SYNONYMY
UNITED STATES: Kansas, Louisiana and Texas, west to California; MEXICO: border states
south to Chiapas and Yucatán; south to ARGENTINA.
SPECIMENS EXAMINED
We have 123 records for this species in the United States.
DISCUSSION
F. Smith's Labidus mexicanus was made a variety of pilosus by Borgmeier (1936), a synonym of
pilosus by M. R. Smith (1942) and, finally, a subspecies of pilosus by Borgmeier (1953), where it
has remained. We have examined the type of L. mexicanus and determined that none of the above
decisions was correct. Labidus mexicanus F. Smith is, instead, a junior synonym of N. swainsonii
(New synonymy).
Due to its large size and relative abundance N. swainsonii is easily one of North America’s most
conspicuous army ants. Surprisingly, for many years the worker form of this common and wide-
ranging species was unknown. We have determined that N. fallax is the worker of N. swainsonii.
The evidence for this association is scanty: it is based on a worker of N. fallax found attached to the
leg of a male collected in Arizona. Although throughout the United States and Mexico the ranges of
these two taxa overlap nicely, N. fallax is unknown south of Guatemala.
Differences do exist between the North and South American populations; however after having
examining the type specimen from Brazil and several other specimens from Central and South
America we are forced to conclude that we are dealing with a single wide ranging and slightly
variable species.
Automontage images of the male caste may be viewed at antweb.org
Neivamyrmex texanus Watkins

Figures 47, 112, 123, 137, 148
Neivamyrmex texanus Watkins, 1972: 353 (w, q, m). U.S.A., Texas, Travis Co., Austin (USNM).
UNITED STATES: Virginia to Florida, west to Colorado and Arizona; MEXICO: San Luis
Potosí to Hidalgo and Jalisco.
SPECIMENS EXAMINED (U.S.A.)
In addition to numerous paratypes, we have 34 records from various states.
DISCUSSION
Small workers may be difficult to distinguish from those of N. nigrescens, but larger workers
are fairly easy to distinguish with a little practice. Males may be readily recognized by the distinctive
shape of the paramere.
This is a widespread ant, found in many different types of habitat. Although not commonly seen,
N. texanus is a large (for Neivamyrmex) and conspicuous ant when it is actively foraging.
Immigration columns are prominent and may often be seen from some distance away when crossing
a suitable background. One observed near Tucson stretched well over 100 yards across the desert
and could be readily seen as it meandered across the light colored sand. The blind snake,
Leptotyphlops dulcius, was observed in the column. Occasionally, when an ant displayed an interest
in the snake, it was quickly flicked off (pers. obs., GCS).
Plsek et al. (1969) observed Helluomorphoides texanus (LeConte) in raiding columns of N.
texanus and reported their observations on behavior of this species in the laboratory.
Automontage images of the worker are available at antweb.org.
Neivamyrmex wilsoni, new species

Figures 16, 26, 48, 61, 74, 87
DIAGNOSIS
Worker. Eye spot absent; front of head smooth and shiny, with scattered coarse punctures;
subantennal lamella well developed; petiole longer than broad in dorsal view, sides anteriorly
convergent; postpetiole distinctly wider than long, disc smooth and shiny, sides shiny and lightly
sculptured. Queen and male unknown.
DESCRIPTION
Worker measurements (mm) (n = 12). HL 0.48-0.77 (0.77); HW 0.36-0.70 (0.70); SL 0.24-

0.0.405 (0.405); SW 0.07- 0.14 (0.14); PW 0.14-0.24 (0.24); PL 0.16-0.27 (0.27); PpL 0.12-0.23
(0.23); PpW 0.19-0.30 (0.30); HFL 0.34-0.62 (62) ; HFW 0.09-0.17 (0.17). Indices: CI 75-93 (90);
FI 25-31 (28); SI 49-61 (52).
Head slightly longer than broad to distinctly longer than broad in minors, sides weakly convex;
front of head smooth and shiny, with scattered coarse punctures; dorsolateral angle slightly
developed, with strong preoccipital carina extending ventrad. Antennal scape about 2.9× longer than
broad, evenly tapering toward base. Subantennal lamella well-developed. Mandible triangular, outer
face weakly longitudinally striate; inner margin sharply angulate with masticatory margin, latter with
small indistinct teeth.
Mesosoma contiguously punctate, slightly shiny. Promesonotal suture indistinct. Mesonotum
smooth and shiny. Dorsum of propodeum opaque, in profile distinctly longer than slightly concave
declivitous face.
Petiole longer than broad in dorsal view and with sides convergent anteriorly, in profile dorsum
evenly arched and abruptly descending anteriorly, disc shiny between sparse fine punctures; side
punctate and opaque; subpetiolar process usually absent, but sometimes with small anteriorly
directed tooth. Postpetiole distinctly wider than long, disc smooth and shiny, sides shiny and lightly
sculptured.
Gaster smooth and shiny between small widely scattered punctures.
Pilosity about as usual in Neivamyrmex: a combination of sparse short to long, suberect to erect
simple hairs.
TYPE MATERIAL
Holotype and numerous paratypes, U.S.A., California, San Diego Co., Elliot Reserve, Mira
Mar (32.84° N 117.11° W), 28 March 1996 (A. Suarez). Holotype and most paratypes in LACM;
paratypes also in BMNH, CASC, MCZC, and UCDC.
ADDITIONAL SPECIMENS
CALIFORNIA, Orange Co.: Starr Ranch, 520m (33°36'N 117°33'W) (P.S. Ward, #14325-3;
UCDC), ex midden of Solenopsis xyloni. Riverside Co.: Lake Skinner, 462m (33°35'N 117°02'W)
(A.V. Suarez #1378.7; UCDC), ex midden of Forelius sp. San Bernardino Co.: Caruthers Canyon
(32.25°N 115.30°W), 1725m (P.S. Ward, #15175; UCDC), under stone.
ETYMOLOGY
We name this species in honor of Ed Wilson, a small recognition of his many achievements in
ant systematics, sociobiology and ecology.
Presently known only from southern California.
DISCUSSION
This small species is most similar to N. nyensis and may be confused with that species. It may
be separated by the shorter antennal scape (3.0× as long as wide vs. 4.0× as long as wide) and the
opaque mesosomal dorsum. The only other local species with which N. wilsoni might be confused is
N. leonardi, but in that species the mandibular teeth are better defined and the postpetiole is twice as
wide as long. N. wilsoni is also a more hairy ant than other similar species. Large workers are further
characterized by having the lower two-thirds or three-fourths of the head weakly longitudinally
strigulose.
Known habitats include oak-pine-juniper woodland, chamise chaparral and coastal sage scrub.
Automontage images of this species (as Neivamyrmex CA-01) are available at antweb.org.
ACKNOWLEDGEMENTS
We are grateful to the following for the opportunity to study specimens in their respective
institutions and/or personal collections: Cheryl Barr (UCBC), Bob Blinn (NCSU), Barry Bolton
(BMNH), Ken Cooper (KWCC), Stefan Cover (MCZC), Lloyd Davis (LDPC), Brian Fisher
(CASC), Phil Ward (UCDC), Jack Longino (JTLC), Bill MacKay (WPMC), Rob Brooks (SEMC),
Bob Johnson (ARSU, RAJC), James Carpenter (AMNH), Mickey Ladkin (TTUC), Ted Schultz
(USNM), Richard Hoebeke (CUIC), Steven Krauth (UWIS), John Morse (CLMC), Victoria Mosely
(LSAM), Carl Olson (UNAR), Steve Prchal (SASI), Zack Prusak (FSCA), David Richman (NMSU),
Ed Riley (TAMU), Mark O'Brien (MICH), Philip Clausen (MINN), Bob Byers (PENN), Dale Ward
(DKWC) and Doug Yanega (UCRC). We also thank Rosser Garrison (CDFA) and David C. Taylor
(Antimite Termite and Pest Control) for their comments and suggestions on very early incarnations
of this paper and James Snelling for his invaluable assistance with the illustrations, and his fine line
drawings of N. baylori, N. carolinensis and N. pilosus.
LITERATURE CITED
Allred, D. 1982. Ants of Utah. The Great Basin Naturalist 42: 415-511.
Cambridge, MA, 504 pp.
Borgmeier, T. 1949. Formigas novas ou pouco conhecidas de Costa Rica. Revista Brasileira de
Biologia 9: 201-210.
Borgmeier, T. 1953. Vorarbeiten zu einer Revision der Neotropischen Wanderameisen. Studia
Entomologica 2: 1-51.
Borgmeier, T. 1955. Wanderameisen der Neotropischen Region. Studia Entomologica 3: 1-720.
Cokendolpher, J.C. & Francke, O.F. 1990. The ants (Hymenoptera, Formicidae) of western
Texas. Part II. Subfamilies Ecitoninae, Ponerinae, Pseudomyrmecinae, Dolichoderinae and
Formicinae. Special Publications, The Museum, Texas Tech University 30: 76 pp.
Creighton, W.S. 1932. A new female Acamatus from Texas. Psyche 39: 73-78.
Creighton, W.S. 1950. The ants of North America. Bulletin of the Museum of Comparative Zoology
at Harvard College 104: 1-585.
Cresson, E.T. 1872. Hymenoptera Texana. Transactions of the Entomological Society of America 4:
153-292.
Dalla Torre, C.G. de. 1893. Catalogus Hymenopterorum, hucusque descriptorum systematicus et
synonymicus 7: 289 pp. Lipsiae.
Emery, C. 1894. Studi sulle formiche della fauna Neotropica. Bullettino della Società Entomologica
Italiana 26: 137-241.
Emery, C. 1896. Studi sulle formiche della fauna Neotropica. Bullettino della Società Entomologica
Italiana 28: 33-107.
Emery, C. 1900. Nuovi studi sul genere Eciton. Memorie della R. Accademia delle Scienze
dell'Istituto di Bologna (5) 8: 173-188 (pp. 511-526 in reprint).
Emery, C. 1895. Beiträge zur Kenntniss der nordamerikanischen Ameisen-fauna (Schluss.).
Zoologische Jahrbücher. Abtheilung für Systematik, Geographie und Biologie der Tiere 8: 257-
360.
Emery, C. 1901. Note sulle Doriline. Bullettino della Società Entomologica Italiana 33: 43-56.
Dorylinae. Fasc. 102: 34 pp. Bruxelles.
Enzmann, E.V. 1952. Woitkowskia, a new genus of army ants. Proceedings of the Iowa Academy of
Science 59: 442-447.
Forel, A. 1899. Biologia Centrali-Americana; or, contributions to the knowledge of the fauna and
flora of Mexico and Central America. Insecta: Hymenoptera. 3 (Formicidae): 169 pp. London.
Forel, A. 1914. Formicides d'Afrique et d'Amerique nouveaux ou peu connus. Bulletin de la Société
Vaudoise des Sciences Naturelles 50: 211-288.
Gallardo, A. 1920. Las hormigas de la República Argentina. Subfamilia dorilinas. Anales del Museo
Nacional de Historia Nautral de Buenos Aires 30: 281-410.
Gregg, R.E. 1963. The ants of Colorado. University of Colorado Press, Boulder, CO, 792 pp.
Haldeman, S.S. 1852. Appendix C.-Insects (pp. 366-378). In Stansbury, H. An expedition to the
Valley of the Great Salt Lake of Utah: including a description of its geography, natural history,
and minerals, and an analysis of its waters. 487 pp. London.
Haliday, A.H. 1836. Descriptions of the Hymenoptera. In: Curtis, J., Haliday, A. H., & Walker, F.
Descriptions etc. of the insects collected by Captain P. P. Ring R. N., F. R. S., in the survey of
the Straits of Magellan. Transactions of the Linnean Society of London. 17: 315-331.
LaPolla, J.S., Mueller, U.G., Seid, M., & Cover, S. 2002. Predation by the army ant Neivamyrmex
rugulosus on the fungus-growing ant Trachymyrmex arizonensis. Insectes Sociaux 49: 251-256.
MacKay, W.P. 1998. A new army ant from southern Texas. The Southwestern Naturalist 43: 333-
335.
MacKay, W.P., Dominguez, J.F.P. & Sanchez, L.I.V. 1985. The army ants of the State of
Chihuahua, Mexico (Hymenoptera: Formicidae: Ecitoninae). The Southwestern Naturalist 30:
611-612.
MacKay, W. & MacKay, E. 2002. The ants of New Mexico (Hymenoptera: Formicidae). Edwin
Mellen Press, Lewiston, New York, 398 pp.
Mallis, A. 1938. Army ants in California. Scientific Monthly 47: 220-226.
Mallis, A. 1941. A list of the ants of California with notes on their habits and distribution. Bulletin
of the Southern California Academy of Science 40: 61-100.
Mann, W.M. 1925. New beetle guests of army ants. Journal of the Washington Academy of
Sciences 15: 73-77.
Mann, W.M. 1926. Some new neotropical ants. Psyche 33: 97-107.
Mayr, G. 1870. Neue Formiciden. Verhandlungen der K.K. Zoologisch-Botanische Gesellschaft in
Wien 20: 939-996.
Mayr, G. 1886a. Ueber Eciton-Labidus. Ein myrmecologischer Beitrag. Wiener Entomologische
Zeitung 5: 33-36.
Mayr, G. 1886b. Ueber Eciton-Labidus (schluss). Wiener Entomologische Zeitung 5: 115-122.
Mayr, G. 1886c. Die Formiciden ver Vereinigten Staaten von Nord-Amerika. Verhandlungen der
k.k. Zoologisch-Botanishcen Gesellschaft in Wien 36: 419-464.
Mayr, G. 1887. Südamerikanische Formiciden. Verhandlungen der k.k. Zoologisch-Botanishcen
Menozzi, C. 1924. Una specie inedita di Eciton Latr. del sottogenere Labidus Jr. Bollettino della
Società Entomologica Italiana 56: 29-31.
Neece, K.C. & Bartell, D.P. 1982. A faunistic survey of the organisms associated with ants of
western Texas. Graduate Studies of Texas Tech University 25: 36 pp.
Norton, E. 1868. Notes on Mexican ants. American Naturalist 2: 57-72.
Pergande, T. 1895. Formicidae of Lower California, Mexico. Proceedings of the California
Academy of Sciences (ser. 2) 4: 161-165.
Rettenmeyer, C.W. & Watkins J.F. 1978. Polygyny and monogyny in army ants (Hymenoptera:
Formicidae). Journal of the Kansas Entomological Society 51: 581-591.
Santschi, F. 1912. Quelques fourmis de l'Amérique australe. Revue Suisse de Zoologie 20: 519-534.
Santschi, F. 1921. Ponerinae, Dorylinae et quelques autres formicides néotropiques. Bulletin de la
Société Vaudoise des Sciences Naturelles 54: 81-103.
Schneirla, T.C. 1956. The army ants. Smithsonian Report for the Year 1955: 379-406.
Schneirla, T.C. 1958. The behavior and biology of certain Nearctic army ants. Last part of the
functional season in Arizona. Insectes Sociaux 5: 215-255.
Schneirla, T.C. 1971. Army ants. W. H. Freeman & Co., San Francisco, 349 pp.
Shuckard, W.E. 1840. Monograph of the Dorylidae, a family of the Hymenoptera Heterogyna.
Annals of Natural History; or Magazine of Zoology, Botany and Geology 5: 188-201.
Smith, D.R. 1979. Formicoidea (pp. 1323-1467). In Krombein, K. V., Hurd, P. D. Jr., Smith, D. R.,
& Burks, B. D. (eds.) Catalog of Hymenoptera in America North of Mexico. 2 Apocrita
(Aculeata):1199-2209. Washington, D. C.
Smith, F. 1858. Catalogue of Hymenopterous Insects in the Collection of the British Museum. 6
Formicidae. British Museum, London, 216 pp.
Smith, F. 1859. Catalogue of Hymenopterous Insects in the Collection of the British Museum. 7
Dorylidae and Thynnidae. British Museum, London, 76 pp.
Smith, M.R. 1927. A contribution to the biology and distribution of one of the legionary ants,
Eciton schmitti Emery. Annals of the Entomological Society of America 28: 401-404.
Smith, M.R. 1942. The legionary ants of the United States belonging to Eciton subgenus
Neivamyrmex Borgmeier. American Midland Naturalist 27: 537-590.
Smith, M.R. 1943. A new male legionary ant from the Mojave Desert, California. Lloydia 6: 196-
197.
Topoff, H.R. 1969. A unique predatory association between carabid beetles of the genus
Helluomorphoides and colonies of the army ant Neivamyrmex nigrescens. Psyche 76: 375-381.
Ward, P.S. 1999. Deceptive similarity in army ants of the genus Neivamyrmex (Hymenoptera:
Formicidae): taxonomy, distribution and biology of N. californicus (Mayr) and N. nigrescens
(Cresson). Journal of Hymenoptera Research 8: 74-97.
Watkins, J.F., II. 1969. A new species of Neivamyrmex from Louisiana (Hymenoptera:
Watkins, J.F., II. 1971. A taxonomic review of Neivamyrmex moseri, N. pauxillus and N. leonardi,
including new distribution records and original descriptions of queens of the first two species.
Journal of the Kansas Entomological Society 44: 93-103.
Watkins, J.F., II. 1972. The taxonomy of Neivamyrmex texanus n. sp., N. nigrescens and N.
californicus (Formicidae: Dorylinae), with distribution map and keys to the species of
Neivamyrmex of the United States. Journal of the Kansas Entomological Society 45: 347-372.
Watkins, J.F., II. 1973. Neivamyrmex baylori n. sp. from Waco, Texas, U.S.A. (Hymenoptera:
Watkins, J.F., II. 1975. The relationship of Neivamyrmex fuscipennis to N. macropterus
(Dorylinae: Formicidae). Southwestern Naturalist 20: 85-90.
Watkins, J.F., II. 1976. The identification and distribution of New World army ants: Baylor
University Press, Waco, Texas, 102 pp.
Watkins, J.F., II. 1977. Neivamyrmex nyensis, n. sp. (Formicidae: Dorylinae) from Nye County,
Nevada, U. S. A. The Southwestern Naturalist 22: 421-425.
Watkins, J.F., II. 1982. The army ants of Mexico (Hymenoptera: Formicidae: Ecitoninae). Journal
of the Kansas Entomological Society 55: 197-247.
Watkins, J.F., II. 1985. The identification and distribution of the army ants of the United States of
America (Hymenoptera, Formicidae, Ecitoninae). Journal of the Kansas Entomological Society
58: 479-502.
Watkins, J.F., II. 1990. The army ants (Formicidae: Ecitoninae) of the Chamela Biological Station
in Jalisco, Mexico. Folia Entomológica Mexicana 77: 379-393.
Watkins, J.F., II. & Coody, C. J. 1986. The taxonomy of Neivamyrmex graciellae (Mann)
including an original description of the queen and field observations. Southwestern Naturalist
31: 256-259.
Watkins, J.F., II., Gehlbach, G.R. & Baldridge, R.S. 1967. Ability of the blind snake,
Leptotyphlops dulcis, to follow pheromone trails of army ants, Neivamyrmex nigrescens and N.
opacithorax. Southwestern Naturalist 12: 455-462.
Watkins, J.F., II., Gehlbach, G.R. & Plisek, R.W. 1972. Behavior of blind snake (Leptotyphlops
dulcis) in response to army ant (Neivamyrmex nigrescens) raiding columns. Texas Journal of
Science 23: 556-557.
Westwood, J.O. 1842. Monograph of the hymenopterous group, Dorylides (No. 5, pp. 73-80).
Arcana Entomologica; or illustrations of new, rare, and interesting insects 1:192 pp. London.
Wheeler, G.C. & Wheeler, J.N. 1973. The ants of Deep Canyon. University of California,
Riverside, California, 162 pp.
Wheeler, G.C. & Wheeler, J.N. 1986. The ants of Nevada. Natural History Museum of Los
Angeles County, Los Angeles, California, 138 pp.
Wheeler, W.M. 1900. The female of Eciton sumichrasti Norton, with some notes on the habits of
Texan Ecitons. American Naturalist 34: 563-574.
Wheeler, W.M. 1908. The ants of Texas, New Mexico and Arizona. Part I. Bulletin of the American
Wheeler, W.M. 1915. Some additions to the North American ant-fauna. Bulletin of the American
Wheeler, W.M. & Long, W.H. 1901. The males of some Texan Ecitons. American Naturalist 35:
157-173.
Map 1. Distribution of: N. andrei ( ); N. californicus ( ); N. carolinensis ( ).
Map 2. Distribution of: N. agilis( ); N. pilosus ( ); N. wilsoni ( ).

Map 3. Distribution of: N. baylori ( ); N. fuscipennis ( ); N. graciellae ( ); N. mandibularis ( ).
Map 4. Distribution of: N. microps ( ); N. moseri ( ); N. nyensis ( ); N. rugulosus ( ).

Map 5. Distribution of: N. mojave ( ); N. ndeh ( ); N. pauxillus ( ).
Map 6. Distribution of: N. kiowapache ( ); N. leonardi ( ); N. goyahkla ( ).

Map 7. Distribution of: N. melanocephalus ( ); N. melshaemeri ( ).
Map 8. Distribution of N. minor.

Map 9. Distribution of N. nigrescens.
Map 10. Distribution of N. opacithorax.

Map 11. Distribution of N. swainsonii.
Map 12. Distribution of N. texanus.

Map 13. Distribution of N. harrisi.

Figures 1-2. Frontal view of head of: 1a-b, N. agilis; 2, N. californicus.

Figures 3-5. Frontal view of head of: 3, N. carolinensis; 4, N. goyahkla; 5a,b N. graciellae
Figures 6-9. Frontal view of head of: 6, N. harrisi; 7, N. kiowapache; 8, N. microps; 9, N. moseri.
Figures 10-13a. Frontal view of head of: 10, N. nigrescens; 11, N. nyensis; 12, N. pauxillus; 13a, N.
pilosus.
Figures 13b. Side view of head: 13b, N. pilosus.

Figures 14-16. Frontal view of head of: 14, N. rugulosus; 15, N. swainsonii; 16 N. wilsoni.
Figures 17-19. Mandibles and subantennal lamella of: 17, N. agilis; 18, N. californicus; 19, N.
carolinensis.
Figures 20-22. Mandibles and subantennal lamella of: 20, N. harrisi; 21, N. kiowapache; 22, N.
moseri.
Figures 23-25. Mandibles of: 23, N. nyensis; 24, N. pauxillus; 25, N. swainsonii.
Figure 26. Mandible of N. wilsoni. Figures 27-28: Dorsal view of head and pronotum of: 27, N.
leonardi; 28, N. melanocephalus.
Figures 29-30. Dorsal view of head and pronotum of: 29, N. nigrescens; 30, N. swainsonii.
Figure 31. Lateral view of mesosoma of: 31, N. agilis.
Figures 32-34. Lateral view of mesosoma of: 32, N. californicus; 33, N. carolinensis; 34, N.
goyahkla.
Figures 35-37. Lateral view of mesosoma of: 35, N. graciellae; 36, N. harrisi; 37, N. kiowapache.
Figures 38-40. Lateral view of mesosoma of: 38, N. leonardi; 39, N. microps; 40, N. moseri.
Figures 41-43. Lateral view of mesosoma of: 41, N. nigrescens; 42, N. nyensis; 43, N. pauxillus.
Figures 44-46. Lateral view of mesosoma of: 44, N. pilosus; 45, N. rugulosus; 46, N. swainsonii.
Figures 47-48. Dorsal view of mesosoma of: 47, N. texanus; 48, N. wilsoni.
Figure 49, Dorsal view of mesosoma of N. agilis.
Figures 50-53. Dorsal view of mesosoma of: 50, N. goyahkla; 51, N. graciellae; 52, N. kiowapache;
53, N. leonardi.
Figures 54-57. Dorsal view of mesosoma of: 54, N. microps; 55, N. moseri; 56, N. nigrescens; 57,
N. nyensis.
Figures 58-61. Dorsal view of mesosoma of: 58, N. pauxillus; 59, N. pilosus; 60, N. rugulosus; 61,
N. wilsoni.
Figures 62-65. Dorsal view of petiole and postpetiole of: 62, N. californicus; 63, N. carolinensis; 64,
N. goyahkla; 65, N. graciellae.
Figures 66-69. Dorsal view of petiole and postpetiole of: 66, N. harrisi; 67, N. kiowapache; 68, N.
microps; 69, N. nigrescens.
Figures 70-73. Dorsal view of petiole and postpetiole of: 70, N. nyensis; 71, N. pauxillus; 72, N.
pilosus; 73, N. swainsonii.
Figure 74. Dorsal view of petiole and postpetiole of N. wilsoni. Figures 75-76, Lateral view of
petiole of: 75, N. californicus; 76, N. carolinensis.
Figures 77-79. Lateral view of petiole of: 77, N. goyahkla; 78, N. graciellae; 79, N. harrisi.
Figures 80-82. Lateral view of petiole of: 80, N. melancephalus; 81, N. microps; 82, N. moseri.
Figures 83-85. Lateral view of petiole of: 83, N. nigrescens; 84, N. nyensis; 85, N. pauxillus.
Figures 86-87. Lateral view of petiole of: 86, N. pilosus; 87, N. wilsonii.
Figure 88. Hind femur of N. californicus.
Figures 89-92. Hind femora of: 89, N. carolinensis; 90, N. goyahkla; 91, N. kiowapache; 92, N.
nigrescens.
Figure 93. Frontal view of head of queen N. kiowapache. Figures 94-95. Frontal view of head of
males: 94, N. andrei; 95, N. baylori. Figure 95 by James Snelling.
Figures 96-97. frontal view of head of male: 96, N. carolinensis; 97, N. kiowapache. Figure 98,
lateral view of head of male N. kiowapache. Figure 96 by James Snelling.
Fig. 99. Antenna of male N. kiowapache.

Figures 100-101: N. mandibularis: 100, mandibles; 101, lateral view of head.
Figures 102-104. Head of males of: 102-103. N. microps; 104, N. minor.

Figures 105-107. Head of males of: 105, N. minor; 106-107, N. nigrescens.

Figures 108-110. Head of males of: 108-109, N. opacithorax; 110, N. pilosus.

Figure 111-112. Head of male: 111, N. pilosus; 112, N. texanus.

Figures 113-116. Aedeagus of : 113, N. carolinensis; 114, N. fuscipennis; 115, N. melshaemeri; 116,
N. minor.
Figures 117-120. Aedeagus of: 117, N. mojave; 118 N. ndeh; 119, N. nigrescens; 120, N.
opacithorax.
Figures 121-123. Aedeagus of: 121, N. pilosus; 122, N. swainsonii; 123, N. texanus.
Figure 124. Paramere of: 124, N. andrei.
Figures 125-128. Paramere of: 125, N. carolinensis; 126, N. kiowapache; 127, N. mandibularis;
128, N. melshaemeri.
Figures 129-132. Paramere of: 129, N. microps; 130, N. minor; 131, N. mojave; 132, N. ndeh.
Figures 133-136. Paramere of: 133, N. nigrescens (after Ward 1999); 134, N. opacithorax (after
Ward 1999); 135. N. pilosus; 136, N. swainsonii.
Figure 137. Paramere of N. texanus. Figures 138-140. Volsella of: 138, N. kiowapache; 139, N.
melshaemeri; 140, N. microps.
Figures 141-144. Volsella of: 141, N. minor; 142, N. mojave; 143, N. ndeh; 144; N. nigrescens.
Figures 145-148. Volsella of: 145, N. opacithorax; 146, N. pilosus; 147, N. swainsonii; 148, N.
texanus.
Figure 149. Dorsal view of mesosoma of N. swainsonii.

Figure 150. Frontal view of head of N. pilosus (type). Figure by James Snelling.
Snelling and Snelling: United States Neivamyrmex
Figure 151. Lateral view of head of N. pilosus (type). Figure by James Snelling.
549
Snelling, R. R. 2007. A review of the arboreal Afrotropical ant genus Axinidris, pp. 551-579. In
A REVIEW OF THE ARBOREAL AFROTROPICAL ANT GENUS AXINIDRIS
Roy R. Snelling
900 Exposition Boulevard
Los Angeles, CA 90007, U.S.A.
antmanrs@nhm.org
ABSTRACT
The species of the Afrotropical genus Axinidris are here reviewed. The type species is
recharacterized following the discovery that the two type specimens have been artificially abraded of
almost all pilosity. Thirteen species were recognized by Shattuck (1991) in his revision of the genus.
New data extend the ranges of several previously described species and eight new species are
described: A. gabonica (Gabon), A. icipe (Kenya), A. lignicola (South Africa), A. luhya (Kenya), A.
mlalu (Central African Republic), A. namib (Namibia), A. okekai (Kenya) and A. stageri (Tanzania).
Technomyrmex hypoclinoides Santschi is here transferred to Axinidris (new combination) with A.
parvus Shattuck as a junior synonym (new synonymy). A key is given for the separation of the
workers of all known species and the new species are illustrated.
KEY WORDS: Hymenoptera, Formicidae, Dolichoderinae, systematics, Axinidris, Afrotropical.

INTRODUCTION
Axinidris is an exclusively Afrotropical genus of arboreal ants, first described by Weber (1941) for a
single bizarre Sudanese species, A. acholli Weber. The genus was recharacterized by Shattuck
(1991), two species were transferred from Engramma Forel, 1905 (subsequently treated as a
synonym of Technomyrmex Mayr, 1872, by Shattuck, 1992), and ten new species were described.
An additional eight species are described below and it thus seems likely that the actual diversity of
this genus is not yet fully realized, given that most species remain known either from type specimens
and/or from only one or two localities.
Species of Axinidris appear to nest exclusively within hollow plant stems, both living and dead,
and in rotten wood. They are found in forested areas throughout the Afrotropical region, but are
most abundant and diverse in the moist equatorial forests. Workers are primarily arboreal foragers,
but may occasionally forage in ground litter.
SPECIMENS EXAMINED
The material on which this study is based has come from the following institutional collections:
Alexander Koening Research Institute and Museum of Zoology, Bonn, Germany (AKRI)
California Academy of Sciences, San Francisco, USA (CASC)
Museum of Comparative Zoology, Harvard University, Cambridge, USA (MCZC)
Natural History Museum of Los Angeles County, Los Angeles, USA (LACM)
Naturhistorisches Museum, Basle, Switzerland (NHMB)
South African Museum, Cape Town, South Africa (SAMC)
The Natural History Museum, London, UK (BMNH)
Voucher specimens of my field work have been placed in most of these institutions, as well as:
International Centre of Insect Physiology and Ecology, Nairobi, Kenya (ICIPE)

National Museum of Kenya, Nairobi, Kenya (NMKC)
MEASUREMENTS AND DEFINITIONS
Measurements of morphological features were made at 50× using a Leitz stereo microscope, with a
dual-axis stage micrometer wired to Autometronics digital readouts. Measurements were recorded
in thousandths of a millimeter, bur are here expressed to the nearest hundredth. Head measurements
were made in full-face (frontal) view, as in Shattuck (1991).
The Scanning Electron Microscope (SEM) images were made using a Hitachi S-3000 SEM.
Terms for surface sculpture follow those of Harris (1979). Shattuck (1991) utilized the same
terminology and defined the varieties of surface sculpture as follows: colliculate: continuously
covered with low, rounded elevations; coriarious: leather-like, with minute cracks; imbricate:
appearing similar to shingles on a roof, i.e., slightly overlapping; punctate: with small pits;
scabriculous: with fine and regular, short sharp wrinkles and/or projections; strigulate: with
numerous short, fine raised lines. To these may be added: smooth: with no apparent surface
sculpture other than punctures; rugae: coarse, raised lines or ridges, sometimes parallel, sometimes
irregularly anastamosing, forming coarse reticulae or foveae.
Descriptive terminology employed here differs somewhat from that of Shattuck in the following
particulars. I consider the ant head to be hypognathous rather than prognathous, so the vertex (see
below) is dorsal and the mandibles are ventral. Mesosomal length (ML) is substituted for Shattuck's
"alitrunk length or AL". Ocular Index (OI) is an established term in ant systematics and is
Snelling: Review of Axinidris 553
substituted for REL (Relative Eye Length). Instead of the morphologically incorrect "occiput" (as
noted by Bolton, 1994) employed by Shattuck I use vertex. Appropriate changes are thus necessary
for several anagrams used by Shattuck (see below). In the descriptions below, the expression
"distance between [propodeal] spines" means the distance across the apices of the spines. Reference
to the profile of the medial carina of the propodeum is to its appearance at the summit of the
declivity.
Cephalic Index (CI): (HW/HL)(100). Note: This is the conventional definition of CI and expressed
as a whole number rather than as a fraction as in Shattuck's revision.
Clypeal Notch Depth (CND): measured from midpoint of a line drawn across ventral-most
projecting points of the clypeus to the dorsal-most point of the concavity.
Clypeal Notch Index (CNI): (CND/CNW)(100). Note: This is a modification of Shattuck's method,
allowing the expression as a whole, rather than fractional, number.
Clypeal Notch Width (CNW): measured between the ventral-most projecting points of the clypeal
notch (contact areas of a line drawn across notch opening.
Eye Length (EL): measured in full face (frontal) view.
Eye Width (EW): measured in full face view.
Head Length (HL): measured in full face view, from uppermost margin of vertex to free margin of
clypeus.
Mesosomal Length (ML): measured from anterior-most point of the pronotum (exclusive of neck) to
the posterior-most point of the propodeal processes (= AL of Shattuck).
Ocular Index (OI): (EL/HL)(100) (= REL of Shattuck); this is the conventional anagram.
Oculo-vertexal Distance (OVD): Distance between upper eye margin and a line drawn across the
dorsal margin of the vertex (= OOD of Shattuck).
Pronotal Width (PNW): maximum width of pronotum in dorsal view. Note: Most of the borrowed
types had been returned before I discovered that Shattuck's measurements did not include PNW;
in the measurements given below, in such cases I have indicated after PNW n/a (not available).
Propodeal Width (PPW): maximum width of propodeum in dorsal view. (Note: after I had returned
most of the type specimens borrowed, I found that Shattuck (1991) had not included this
measurement among those that he had made; in such cases PPW is annotated n/a)
Scape Index (SI): (SL/HW)(100) (= ScI of Shattuck); this is the conventional anagram.
Scape Length (SL): maximum length of scape, excluding basal condyle.
SYSTEMATICS
Workers and queens of Axinidris may be recognized among the Dolichoderinae by the following
suite of characters: clypeus with distinct median notch on free margin; metapleural gland opening
large, approximately circular and surrounded by long, stiff straight hairs; propodeum armed with
spines or tubercles located near the dorsal margin of the declivitous face, generally short and
rounded but may be reduced to mere angles, or more strongly developed as elongate spines that are
curved dorsad; propodeal spiracle near dorsal face of propodeum and with at least anterior margin
raised on a slight to strong projection, spiracular opening thus directed more or less distad;
propodeum usually with elongate median ridge or carina, sometimes sharply elevated and
compressed; petiolar node reclinate, narrowly rounded to subacute dorsally, strongly inclined
posteriorly; in worker and queen, gaster with only four exposed terga in dorsal view, tergum 5
situated beneath tergum 4. Males are similar to those of Technomyrmex, from which they differ in
the evenly arched profile of the petiolar node, which is not concealed by the first gastral segment as
seen in dorsal view.
KEY TO AXINIDRIS WORKERS
1 Propodeal ridge cariniform or absent; pronotal dorsum evenly rounded at sides ............ 2
-- Propodeal ridge compressed and strongly elevated, axiniform (Fig. 11); pronotal dorsum
submarginate at side (Sudan; Kenya).......................................................... acholli Weber
2(1) Pronotum and antennal scape shaft without erect hairs; gastral terga usually without erect
hairs ................................................................................................................................. 3
-- Pronotum with at least two erect hairs (caution: one or both, if only two are normally
present, may be broken off or lying on the surface); antennal scape shaft and gastral terga
with or without erect hairs ............................................................................................. 10
3(2) Medial carina present at least on posterior half of dorsal face & at least slightly elevated at
summit of declivity; propodeal spines various, but often more elongate and with surface
between them concave in dorsal view ............................................................................. 4
-- Medial carina absent from entire length of dorsal face; propodeal spines triangular in dorsal
view and surface between them straight ........................................ bidens Shattuck (part)
4(3) Pronotal dorsum conspicuously imbricate and only slightly shiny; mesonotum,
mesepisternum and propodeum duller, finely and contiguously punctate; dorsal surfaces
with abundant prostrate pubescence; distance between outer margins of propodeal spines
subequal to width of propodeum at spiracles.................................................................. 5
-- Pronotum and mesepisternum smooth and shiny, contrasting sharply with finely and
contiguously punctate propodeum; pronotum with scattered prostrate pubescence; distance
between outer margins of propodeal spines much less than width of propodeum at spiracles
………. ........................................................................................................................... 7
5(4) Vertex without erect hairs; gastral terga without erect hairs............................................ 6
-- Vertex with one pair of short erect hairs below dorsal margin; gastral terga 3 and 4 each
with several erect hairs (Liberia) ..................................................... occidentalis Shattuck
6(5) Head longer than broad; head and body dark brown, lower half of clypeus, mandibles, most
of scape yellowish red; meso- and metatarsi whitish (Kenya) .............. icipe, new species
-- Head broader than long; head and body black, hypostomal area, head below, mandibles and
legs reddish black; tarsi dark yellowish red (Kenya) .................... kakamegensis Shattuck
7(4) Propodeal carina low and right-angular to obtuse in profile (Figs. 17, 18)...................... 8
-- Propodeal carina elevated and acute in profile (Malawi; South Africa) .. tridens (Arnold)
8(7) At least fourth gastral tergum with at least 2 erect hairs; head and body light to dark brown.
……… ............................................................................................................................. 9
-- Third and fourth gastral terga without erect hairs; head and mesosoma light reddish, petiole
and first gastral segment yellowish (Central African Republic) .......... mlalu, new species
9(8) Hypostomal area without erect or suberect hairs; longest hair on fourth tergum about
0.10mm long and vertex hair about 0.66 x length of that on frontal lobe (Kenya;
Cameroun)………………………………………………………………okekai, new species
-- Hypostomal area with several short erect and suberect hairs; longest hair on fourth tergum
about 0.06mm long and longest vertexal hairs about 0.60 x length of that on frontal lobe
(Namibia)............................................................................................ namib, new species
10(2) Propodeum without medial carina and lateral spines virtually absent in dorsal view;
pronotum with one pair of long erect hairs and propodeal spines each with similar hair;
clypeal notch broad and shallow (CNI less than 75)...................................................... 11
-- Propodeum with distinct medial carinae which, at summit of declivity, is usually a low
obtuse lobe, less often sharply elevated or dentiform; pilosity various, but usually not as
above; clypeal notch relatively narrow, CNI usually over 100, but if as little as 80, then
medial carina sharply defined ........................................................................................ 12
11(10) Head and body medium to dark brown; gena distinctly finely imbricate; larger species, HW
consistently over 0.50mm (Ghana; Cameroun; Kenya) .................. bidens Shattuck (part)
-- Head and body light reddish yellow, gaster sometimes slightly darker; gena smooth and
shiny, sometimes weakly imbricate; smaller species, HW usually less than 0.50mm, but
may range up to 0.52mm (Liberia; Democratic Republic of Congo; Kenya) ………….
.................................................................................................... hypoclinoides (Santschi)
12(10) Distance between outer margins of propodeal spines, in dorsal view, equal to or exceeding
width of propodeum at spiracles; pronotum dull and closely pubescent, surface coarsely
imbricate or punctate ..................................................................................................... 13
-- Distance between outer margins of propodeal spines, in dorsal view, distinctly less than
width of propodeum at spiracles and/or pronotum shiny and sparsely pubescent, surface
weakly imbricate............................................................................................................ 14
13(12) Erect hairs of head and mesosoma brown, coarse and seta-like; propodeal spines short, no
longer than thick in profile (Ghana)....................................................... nigripes Shattuck
-- Erect hairs of head and mesosoma yellowish and fine, not seta-like; propodeal spines
distinctly longer than thick in profile (Kenya)..................................... luhya, new species
14(12) Mesonotum either smooth and shiny between several short longitudinal rugae or slightly
shiny and coarsely rugulose .......................................................................................... 15
-- Mesonotum moderately shiny, anteriorly finely imbricate, posteriorly duller and densely
punctulate ..................................................................................................................... 16
15(14) Mesonotum smooth and shiny between short longitudinal rugae; head and body light
reddish brown (Tanzania) .................................................................. stageri, new species
-- Mesonotum slightly shiny and coarsely rugulose; head and body dark brown (South Africa)
………………………………………………………………………lignicola, new species
16(14) Scape shaft with fine prostrate pubescence, but no suberect to erect hairs; front of head,
exclusive of clypeus, with six hairs (two along each frontal carina and two on upper frons)
....................................................................................................................................... 17
-- Scape shaft with at least one or two short erect hairs in addition to appressed pubescence;
front of head, exclusive of clypeus, with two hairs along each frontal carina and four or
more on frons................................................................................................................. 19
17(16) Distance between outer margins of propodeal spine apices distinctly less than width of
propodeum at level of spiracles; frons and vertex moderately shiny, weakly imbricate and
with scattered weak punctures; gaster concolorous with mesosoma.............................. 18
-- Distance between outer margins of propodeal spine apices at least equal to width of
propodeum at level of spiracles; frons and vertex weakly shiny, contiguously finely
punctate and strigulate; mesosoma brown, gaster yellowish red (Ghana) ........................
........................................................................................................palligastrion Shattuck
18(17) Medial carina broadly rounded in profile; second gastral tergum moderately shiny and
densely punctate (Republic of Congo)............................................ denticulata (Wheeler)
-- Medial carina subacute in profile; second gastral tergum shiny between sparse punctures
(Ghana) ............................................................................................... ghanensis Shattuck
19(16) Pronotum with at least two fully erect hairs in addition to humeral pair; frons slightly shiny
to shiny, varying from finely colliculate to weakly coriarious and scabriculous ........... 20
-- Pronotum with only two long fully erect hairs, one at each humerus; frons shiny, varying
from smooth to obscurely coriarious (Gabon) ............................... gabonica, new species
20(19) Scape shaft distinctly with six or more short erect hairs; frons slightly shiny, finely
colliculate to imbricate................................................................................................... 21
-- Scape shaft with only one or two very short erect hairs; frons shiny, weakly coriarious
(Nigeria) ................................................................................................... kinoin Shattuck
21(20) Body color reddish yellow; gaster yellow, similar in color to posterior portion of petiole
(Ghana) ............................................................................................... hylekoites Shattuck
-- Entire head and body reddish brown, gaster darker than posterior portion of petiole
(Cameroun; Ghana; Kenya) .................................................................. murielae Shattuck
Axinidris acholli Weber

Figures 1, 11, 21
Axinidris acholli Weber, 1941: 193 (w). SUDAN: Imatong Mountains, 4800 and 6200 ft. (N.A.
Weber) (MCZC) examined. Shattuck, 1991: 109-111; figs. 5-7, 35.
Worker diagnosis. Pronotal disc with 8-10 coarse rugae that more or less diverge behind;
mesepisternum with 4 or 5 coarse longitudinal to oblique rugae; medial propodeal carina compressed
and conspicuously higher than long and longer dorsally than at base; head and body with abundant
long, slender whitish hairs.
Worker measurements (mm) (n = 12). HW 0.79-0.90; HL 0.91-1.01; SL 0.79-0.88; EL 0.18-

0.23; EW 0.11-0.13; OVD 0.36-0.42; PNW 0.46-0.59; PPW 0.33-0.40; WL 1.08-1.29. Indices. CI
85-91; CNI 71-87; OI 23-26; SI 95-103.
Worker description. The worker caste has been adequately described by Shattuck (1991),
except for the presence of numerous long flexuous hairs as noted above in the diagnosis.
SPECIMENS EXAMINED
In addition to the lectotype and lectoparatype, I have collected numerous workers from
KENYA, Kakamega District, Isecheno, Kakamega Forest (00.24°N 034.85°E), 1550-1600m
(LACM), running on vines, except two in litter. All specimens were in dense forest as opposed to
being on trees at the edge of a clearing.
DISCUSSION
The bizarrely developed propodeal structures (Figs. 11, 21) are sufficient to separate A. acholli
from all the known remaining species. Additionally, no other species is known that has such an
abundance of long, flexuous white hairs. Only A. lignicola and A. stageri are almost as hairy, but in
both the propodeal structures are much less extreme, the hairs are shorter and straighter, and the
antennal scapes are proportionately much shorter.
I had originally regarded the Kenyan specimens as a previously undescribed species. When I
examined the two type specimens of A. acholli, however, I began to doubt that this was correct. The
only difference that I could discern was that the Kenyan specimens were abundantly hairy while the
A. acholli types were almost completely devoid of hairs. The type specimens, lectotype and
lectoparatype, consist of fragments mounted on points. The lectotype head has only a single
antenna. The lectoparatype is in even worse condition: the head lacks antennae, the mesosoma is
partly broken, and only a single detached hind leg is present. Both specimens appear to be severely
abraded, lacking hairs where all other species possess hairs (e.g., the mandibles, clypeus, frontal
carinae). Once it was clear that these poor specimens had been artificially denuded, it was obvious
that my fresh Kenyan samples were conspecific.
In addition to Sudan and Kenya, I expect that A. acholli will also be found in Uganda and the
Democratic Republic of Congo, at the very least.
Axinidris bidens Shattuck

Figures 2, 12, 22
Axinidris bidens Shattuck, 1991: 111-112; figs. 8-10, 35 (w). GHANA: Tafo (C. Campbell)
(BMNH) examined.
Worker diagnosis. Pronotal humeri and propodeal spines each with a single long fully erect
hair, mesosoma otherwise without erect hairs; antennal scape shaft without erect hairs; medial carina
absent and propodeal spines distinct.

0.19; OVD 0.24-0.29; PNW 0.33-0.38; PPW 0.18-0.26; WL 0.79-0.92. Indices. CI 80-89; CNI 50-
80; OI 26-34; SI 75-85.
Worker description. The worker caste has been adequately described by Shattuck (1991).
Queen measurements (mm) (n = 4). HW 0.62-0.63; HL 0.77-0.81; SL 0.50-0.51; EL 0.21-

0.23; OVD 0.29-0.32; PNW 0.51-0.58; PPW 0.37; WL 1.05-1.17. Indices. CI 76-82; CNI 80; OI
33-38; SI 80-83.
Queen description. Generally similar to worker but a little larger and with more elongate head;
mesosoma fully developed for flight; interocular distance 1.61-1.87× EL; interocellar distance 3×
diameter of median ocellus; ocellocular distance about 6× diameter of median ocellus; medial carina
absent; sculpture and pilosity similar to those of worker; mesonotum and metanotum each with a
single long erect hair on each side; gastral pilosity about as in worker, but appressed pubescence
sparser; total length about 2.5mm.
Male unknown.
SPECIMENS EXAMINED
In addition to the three type specimens I have examined samples from the following localities.
CAMEROUN, Southwest Prov., Big Ngwandi, Rumpi Hills (LACM). CENTRAL AFRICAN
REPUBLIC, Res. Dzanga-Sangha (CASC). KENYA, Kakamega Distr., Buyangu, Isecheno and
Kisere, all in Kakamega Forest, 1550-1600m (AKRI, LACM).
DISCUSSION
The samples from Cameroun were collected from living stems of Leonardoxa africana. At
Isecheno I collected workers running on a fallen tree trunk. One colony collected at Isecheno, on 30
April 2003 from a living stem of Acanthus eminens consisted of a single queen and 44 workers in
addition to 19 pupae (17 workers). Alate queens were collected from colony samples in Cameroun
during the month of November. This species has been collected by Thomas Wagner while fogging
Teclea nobilis, a common forest tree species. Other fogging samples, collected by W. Freund,
included some from Heinsenia diervillioides, in addition to T. nobilis. These data suggest that A.
bidens, and probably all species of Axinidris, are able to utilize stems or branches of a wide variety
of plant species as nesting sites.
Automontage images of this species (as Axinidris sp. 3) may be viewed at www.antweb.org.
Axinidris denticulata (Wheeler)
Engramma denticulatum Wheeler, 1922: 205; fig. 51 (w). DEMOCRATIC REPUBLIC OF

CONGO, between Lukolela and Basoko (H.O. Lang) (MCZC) examined.
Axinidris denticulatum: Shattuck, 1991: 112-114; figs. 11-13, 35.
Axinidris denticulata: Bolton, 1995: 77.
Worker diagnosis. Antennal scape without and pronotum with erect hairs; medial carina
present and distance between spines less than PPW; mesonotum without rugae; medial carina
rounded in profile; gastral tergum 2 densely punctate.
Worker measurements (mm) (n = 1). HW 0.71; HL 0.79; SL 0.62; EL 0.19; OVD 0.30; PNW
0.44; PPW 0.37; WL 0.91. Indices. CI 90; CNI 153; OI 27; SI 88.
Worker description. The worker caste has been adequately redescribed by Shattuck (1991).
DISCUSSION
This species is still known only from the three type specimens. Shattuck (1991) selected as
lectotype the only specimen that is in fair condition; the two lectoparatypes are so covered with mold
that they are nearly worthless for diagnostic purposes.
Axinidris gabonica, new species

Figures 3, 13, 23
Worker diagnosis. Antennal scape with sparse short erect hairs; upper frons and pronotum
each with two long erect hairs; gastral terga 3-4 each with one pair of long erect hairs; propodeal
spines short and thick; head and body mostly shiny.
Worker measurements (mm) (n = 3). HW 0.77-0.83; HL 0.83-0.90; SL 0.72; EL 0.21-0.22;

OVD 0.28-0.30; PNW 0.50-0.56; PPW 0.36-0.40; WL 1.01-1.10. Indices. CI 91-94; CNI 117-140;
OI 25-26; SI 86-93.
Worker description. Head shiny, frons weakly coriarious between sparse distinct punctures,
more strongly sculptured on frontal area; gena and malar area shinier and almost smooth. Antennal
scape shafts with a few widely spaces fine erect hairs; frons, below vertex, with 1 pair of long erect
hairs and a shorter pair near upper end of frontal carinae and a still shorter pair at lower end of
carinae; clypeus with about 6 long erect hairs.
Pronotum shiny and nearly smooth between distinct sparse punctures and with a single long
erect hair near each humerus (Fig. 13). Mesonotum similar to pronotum, but narrowly contiguously
punctate posteriorly. Metanotal spiracles elevated but appressed to base of propodeum, not
conspicuous in profile. Mesepisternum largely smooth and shiny anteriorly, becoming duller and
contiguously punctate posteriorly. Propodeum slightly shiny, contiguously to subcontiguously
punctate; spiracular prominence weak and spiracular opening directed distad; medial carina present
on posterior half of dorsal face, absent from declivitous face, slightly elevated at summit of declivity
and low and broadly rounded in profile; propodeal spines short and blunt, obtuse in dorsal view with
straight outer margins, distance between outer margins of apices distinctly less than distance
between spiracles.
Gastral terga shiny, smooth to weakly coriarious; T3-T4 each with a single long erect hair on
each side; appressed pubescence sparse.
Head blackish, body dark brown; distal portion of mandible yellowish, with reddish yellow
teeth; legs brown, distitarsi yellowish; petiole mostly whitish.
TYPE MATERIAL
Holotype worker, GABON, Prov. Ogoové-Maritime, Réserve des Mons Doudou, 25.2 km 304°
NW Doussala (2°13.63'S 10°23.67'E), 660m, 17 March 2000 (S. van Noort), sweeping in coastal
lowland rainforest. Paratypes: 2 workers, same data as holotype, 2 workers, same except 16 March
2000. Holotype and 2 paratypes in CASC; 1 paratype in LACM.
ETYMOLOGY
The name is derived from that of the country of origin.
DISCUSSION
This species will run to A. murielae Shattuck in the key by Shattuck (1991), but is immediately
separable by the scarcity of erect hairs on the vertex and frons. The general habitus is similar, but A.
gabonica is darker, and in both species the punctures of the frons and pronotum are relatively coarse.
Axinidris ghanensis Shattuck
Axinidris ghanensis Shattuck, 1991: 114; fig. 35 (w). GHANA, Mampong (P. Room) (BMNH)
examined.
Worker diagnosis. Antennal scape without and pronotum with erect hairs; medial carina
present and subacute in profile; distance between spines less than PPW; gastral tergum 2 shiny and
sparsely punctate.
n/a; PPW 0.34; WL 0.98. Indices. CI 91; CNI 150; OI 28; SI 86.
Worker description. The worker caste has been adequately described by Shattuck (1991).
DISCUSSION
This species is known only from the single worker holotype. Nothing is known of its biology,
although it is assumed to be arboreal. The most similar species is A. luhya, described below, from
which it differs as detailed under that species.
Axinidris hylekoites Shattuck
Axinidris hylekoites Shattuck, 1991: 114; figs. 14-16 (w, q). GHANA, Tafo (B. Bolton)
(BMNH) examined.
Worker diagnosis. Head and mesosoma reddish, gaster yellow; scape and pronotum each with
4 or more erect hairs; medial carina present; distance between spines less than PPW.
Worker measurements (mm) (n = 4). HW 0.69; HL 0.76-0.79; SL 0.57-0.61; OVD 0.25-0.27;

PNW n/a; PPW 0.32-0.33; WL 0.71-0.92. Indices. CI 88-91; CNI 77-119; OI 25-28; SI 83-88.
Worker and Queen have been adequately described by Shattuck; the male caste remains
unknown.
DISCUSSION
This species is known only from the several specimens comprising the type series. They were
collected from a nest in a rotten tree branch.
As Shattuck (1991) originally noted it is most similar to A. murielae but differs in the lighter
body color. The surface sculpture of the head is slightly coarser and the propodeal spines slightly
wider; this latter difference is almost certainly sufficiently variable as to be of little value.
Axinidris hypoclinoides (Santschi) new combination

Figures 9, 19, 29
Technomyrmex hypoclinoides Santschi, 1919: 89-90; (w). DEMOCRATIC REPUBLIC OF

CONGO (= Belgian Congo, Avakubi , 6 Jan. (J. Bequaert) (NHMB).
Axinidris parvus Shattuck, 1991: 118; fig. 35 (w). LIBERIA, Paiata (= Payeta) (J. Bequaert)
(MCZC) examined. NEW SYNONYMY (B. Bolton, pers. comm.).
Worker diagnosis. Antennal scape shaft without erect hairs and pronotum (Fig. 19) with 1 pair
of long erect hairs and each propodeal spine with 1 long erect hair; head (excluding clypeus) with 2
pairs of erect hairs; medial propodeal carina absent; first tergum with 2 long erect hairs; gena shiny
and finely imbricate; head and body yellow.

75; OI 28-32; SI 79-83.
Worker description. The worker has been adequately described by Shattuck (1991).
SPECIMENS EXAMINED
I have examined the two type specimens of A. parvus and others from the following localities.
GABON, Ogooue Prov., Maritime Res. Mons Doudou (CASC); KENYA, Kakamega Forest,
Isecheno and Isiukhu (AKRI, LACM).
DISCUSSION
The type of Technomyrmex hypoclinoides Santschi was examined by B. Bolton who informed
me that it is a senior synonym of A. parvus. Both the new combination and new synonymy should
be attributed to Mr. Bolton.
The few additional specimens available somewhat amplify the range of size variation, but
otherwise are in good agreement with the type specimens and with Shattuck's original description.
The presence of a single pair of relatively long erect hairs on the pronotal disc and a single such hair
at the apex of each propodeal spine is an unusual feature shared only with A. bidens, a larger and
darker species. Both species are present in the Kakamega forest of Kenya, but A. bidens is
encountered much more frequently.
Photomontage images are available at www.antweb.org as Axinidris sp. 1.
Axinidris icipe, new species

Figures 4, 14, 24
Worker diagnosis. Antenna scape shaft, pronotal disc and all gastral terga without erect hairs;
each frontal carina with a single erect hair and frons with none; medial carina subacute in profile;
pronotal disc slightly shiny and distinctly coriarious.
Worker measurements (n = 10). HW 0.60-0.68; HL 0.73-0.77; SL 0.44-0.49; EL 0.14-0.17;

OVD 0.32-0.33; PNW 0.37-0.42; PPW 0.26-0.28; WL 0.77-0.90. Indices. CI 84-90; CNI 60-80; OI
22-28; SI 67-76.
Worker description. Frons slightly shiny, weakly coriarious and densely micropunctate; gena
similar but becoming smooth and shiny toward mandible; lower frontal area and clypeus dull and
sharply sculptured. Frons and vertex without erect hairs; one erect hair on each frontal carina above
level of antennal insertion; clypeus with one pair of long erect hairs; scape shaft without erect hairs.
Mesosomal dorsum (Fig. 14) without erect hairs. Pronotum slightly shiny and sharply
coriarious. Mesonotum finely reticulopunctate; mesepisternum and side of propodeum similar but
more sparsely punctate and interspaces shinier. Metanotal spiracles low and barely visible in
profile.
Spiracular prominence of propodeum low and inconspicuous, spiracular opening directed distad;
medial carina present on posterior half of dorsal face, continuing onto declivitous face, carina right-
angular to obtuse in profile at summit; spines blunt in dorsal view, distance between their outer
apices greater than width of propodeum at spiracles.
Gastral terga shiny between close fine piligerous punctures and without erect hairs.
Head and body dark blackish brown, lower half of clypeus, mandibles, most of scape yellowish
red; meso- and metatarsi dirty whitish.
TYPE MATERIAL
Holotype worker and 20 worker paratypes, KENYA, Kakamega Dist., Isiukhu, Kakamega
Forest (0.27°N 34.88°E), January 2003 (W. Freund and C. Schmidt), fogging Teclea nobilis.
Holotype in LACM; paratypes in AKRI, BMNH, LACM, MCZC, NMKC, and United States
National Museum of Natural History (USNM).
ETYMOLOGY
The name is derived from the anagram for the International Centre of Insect Physiology and
Ecology, Nairobi; it here used as a noun in apposition.
DISCUSSION
This species may be confused with A. kakamegensis, a species still known only from the type
specimen. It shares with that species the extreme reduction in erect pilosity; i.e., the lack of erect
hairs on the antennal scapes, frons, mesosomal dorsum and gastral terga. It differs from A.
kakamegensis in the narrow head (longer than broad), the lack of strigulate sculpturing on the frons
and malar area, and the lack of erect hairs between the frontal lobes; the clypeus has only a single
pair of erect hairs (several present in A. kakamegensis). Axinidris icipe is also a notably smaller
species in which the head width is less than 0.70 mm versus 0.97 mm in the type of A.
kakamegensis.
Axinidris kakamegensis Shattuck
Axinidris kakamegensis Shattuck, 1991: 115; figs. 17-19, 35 (w). KENYA, Kakamega Forest
(A.E. Stubbs) (BMNH) examined.
Worker diagnosis. Scape and pronotum without erect hairs; head broader than long and vertex
without erect hairs; distance between spines greater than PPW.
DISCUSSION
Known only from the worker type specimen from an unspecified locality in the Kakamega
Forest.
Axinidris kinoin Shattuck
Axinidris kinoin Shattuck, 1991: 115-116; figs. 20-22, 35 (w). NIGERIA, "specific locality not
given, probably Gambari Experiment Station) (B. Taylor)” (BMNH) examined.
Worker diagnosis. Pronotum with erect hairs and scape with 1 or 2 short erect hairs;
mesonotum without rugae; medial carina present; distance between spines less than PPW; head and
body reddish brown.
Worker measurements (mm) (n = 2). HW 0.75-0.76; HL 0.83-0.86; SL 0.68-0.69; EL 0.22;

OVD 0.26-0.28; PNW n/a; PPW 0.37-0.38; WL 1.01. Indices. CI 88-91; CNI 128-131; OI 29; SI
91.
DISCUSSION
This species remains known only from the two type specimens from an unspecified locality in
Nigeria, but assumed to be the Gambari Experiment Station (Shattuck, 1991). It is most similar to A.
denticulata from which it differs as noted in the key.
Axinidris lignicola, new species

Figures 5, 15, 25
Worker diagnosis. Antennal scape shaft without erect hairs and pronotum with 12+ short
suberect hairs and 4-6 longer erect hairs; propodeal spines short, dentiform; propodeal dorsum
obliquely elevated distad; medial carina strongly elevated, obtuse in profile; first gastral tergum
with erect discal hairs and marginal row of suberect hairs.

170; OI 24-30; SI 86-100.
Worker description. Front of head smooth to weakly coriarious between fine subcontiguous
punctures; gena and malar area distinctly coriarious between well-spaced punctures, many of which
are smaller than on frons. Frons with 4-5 pairs of erect hairs, uppermost pair longest; vertex with 1
pair of suberect hairs subequal in length to longest frontal pair. Pubescence short, mostly appressed
but some on side of head, especially on malar area, subappressed to reclinate. Shaft of antennal
scape with appressed pubescence only.
Pronotum moderately shiny and coriarious between well-separated fine piligerous punctures.
Mesonotum irregularly rugulose and moderately shiny. Mesepisternum moderately shiny between
short, weak, more or less longitudinal rugae anteriorly, becoming duller and colliculate posteriorly
and ventrad. Propodeal dorsum so coarsely rugulose that medial carina is obscured; medial carina at
summit of declivity a short, strongly elevated obtuse lobe; spiracle on well developed prominence,
opening directed posterolaterad; propodeal spines short and nearly right-angular in dorsal view,
distance between their outer apices less than distance between spiracles. Pronotum (Fig. 15) with 4-
5 pairs of short erect hairs; mesonotum and dorsum of propodeum with variable number of suberect
to erect hairs of variable length.
Gastral tergum 1 moderately shiny and weakly coriarious, following segments less shiny and
more distinctly sculptured. Terga 1-2 with marginal band of suberect hairs; terga 2-4 with short
suberect discal hairs and all segments with abundant coarse subappressed pubescence.
Head and body dark brown; lateral lobes of clypeus and transverse stripe at base of mandibular
teeth yellowish.
TYPE MATERIAL
Holotype worker and numerous worker paratypes, SOUTH AFRICA, Western Cape,
Grootvadersbosch (38°59'S 20°49'E), 16 September 1995 (H.G. Robertson), "in centre of dead tree
trunk c 2.5cm dia., 1 m above ground", in indigenous forest. Holotype and most paratypes in
SAMC; paratypes also in BMNH, CASC, LACM and MCZC.
ETYMOLOGY
Wood-dweller, from L., lignum (wood) + L., colo (inhabit).
DISCUSSION
In addition to the type series I have seen one worker, also in SAMC, collected by A.J. Prins at
the same locality, October 1985 and provided with an unpublished name by Prins and Roux.
The strongly elevated medial carina at the summit of the propodeal declivity, as well as the
abundance of suberect to erect pilosity in combination with the lack of erect hairs on the scape shaft,
will serve to distinguish this species from all others known to me.
Axinidris luhya, new species

Figures 6, 16, 26
Worker diagnosis. Erect hairs present on pronotal disc but absent from scape shaft; propodeal
spines thick and directed posterolaterally, distance between their apices greater than propodeal
width; metanotal spiracles prominent in profile and base of propodeum abruptly elevated behind
metanotal suture; propodeum dull and contiguously minutely punctate.

0.28; OVD 0.37-0.42; PNW 0.60-0.68; PPW 0.38-0.46; WL 1.13-1.26. Indices. CI 92-100; CNI
12-15; OI 22-28; SI 69-74.
Worker description. Head slightly shiny, with fine strigulate sculpture, interspaces
contiguously finely punctate; clypeal disc and malar area with fine longitudinal rugae. Vertex with
0-3 pairs of short stiff hairs; frons with 6-8 widely spaced hairs of variable length in addition to
much longer pair along each frontal carina; entire head with abundant short coarse subappressed to
appressed pubescence. Shaft of antennal scape without erect or suberect hairs.
Entire mesosoma contiguously punctate and slightly shiny and with abundant coarse
subappressed to appressed pubescence; mesepisternum and side of propodeum also with weak
irregular longitudinal rugae. Pronotal disc (Fig. 16) with 2-3 pairs of long erect hairs and a variable
number (but fewer than 12) of much shorter erect hairs; 1-3 short erect hairs on each propodeal
spine. Mesonotum flat in profile; metanotal spiracles prominent. Propodeum abruptly elevated
behind metanotal suture; disc nearly flat; propodeal spiracle on well-developed prominence and
spiracular opening directed distad; medial carina low; propodeal spines thick, their apices subacute
in profile; spines broad in dorsal view and direct posterolaterally, their outer margins straight and
distance between their apices about equal to width of propodeum at spiracles; 4-10 fine erect hairs
present, including on spines.
Gastral terga slightly shiny and weakly imbricate, with abundant minute piligerous punctures;
terga with close coarse appressed pubescence and discs of two basal segments also usually with
scattered short erect hairs; T3-4 with 2-4 longer erect hairs.
Head and body blackish, antennae and legs dark brownish, tarsi reddish brown; yellowish spot
on head between eye and antennal socket; mandibles largely reddish, but blackish at base and apex.
TYPE MATERIAL
Holotype worker: KENYA, Kakamega Distr., Isecheno, 19 March 2002 (R. R. Snelling, #02-
156), on tree trunk. Numerous additional worker paratypes from Isecheno, collected by R. R.
Snelling, various dates, mostly on trunk of Solanum mauritiana. Holotype in LACM; paratypes in
AKRI, BMNH, CASC, ICIPE, LACM, MCZC, NMKC, SAMC, and USNM.
ETYMOLOGY
This species is named for the Luhya people who live in the Kakamega area and, in particular,
for those at Isecheno who made me feel welcome there; the name is a noun in apposition.
DISCUSSION
Additional specimens (not types) are from Yala and "Colobus Trail", Kakamega Forest, both
taken while fogging Teclea nobilis, by W. Freund. There is also a single worker collected in
GABON: Prov. Ogoové-Maritime, Réserve des Monts Doudou, 24.3 km 307° NW Doussala,
2°13.35'S 10°24;35'E, 370m, 7.iii.2000, S. van Noort (CASC), on vegetation in lowland rainforest.
The erect hairs of the antennal scape are pale, very fine and short, about 0.05mm long, and are
widely and irregularly distributed along the length of the shaft. The number varies from 2 to 6, but
one specimen was examined that lacked erect hairs on one scape, but had 4 on the other. Also
variable is the number of erect hairs on the mesosomal dorsum. Although the pronotum often has
but a single pair of long (up to 0.10mm) hairs, the number varies from 0 (2 specimens) to 8 within a
nest series. Occasionally, 1 or 2 short erect hairs may be present on the propodeal dorsum in the
vicinity of the spiracles; 1 or 2 similar hairs are present on each spine rising above the sometimes
suberect pubescence.
This appears to be an eastern sister species to A. occidentalis, currently known only from
Liberia. The two are similar is size, head shape, mesosomal profile and in most details of sculpture,
but A. luhya has the mesosoma more coarsely sculptured and there are long erect hairs on both the
upper frons (usually) and pronotal disc. In general habitus A. luhya might also be confused with A.
ghanensis, but in that species the distance between the outer margins of the propodeal spines is no
greater than that between the spiracles, the pronotum has only a single pair of erect hairs and none
are present on the propodeum and on gastral tergum 3.
Axinidris mlalu, new species

Worker diagnosis. Scape shaft, pronotum and gastral terga without erect hairs; distance
between apices of propodeal spines less than distance between propodeal spiracles; gastral tergum 1
yellow, segments 2-4 dark brown.
Worker measurements (mm) (n - 1). HW 0.74; HL 0.86; SL 0.68; EL 0.21; OVD 0.31; PNW
0.49; PPW 0.36; WL 1.03. Indices. CI 87; CNI 100; SI 91; OI 28.
Worker description. Front of head moderately shiny, weakly coriarious and strigulate between
sparse minute punctures; gena shinier and with few strigulae, malar area more strongly coriarious.
Antennal scape without erect hairs; each frontal carina with 1 pair of long erect hairs; upper frons
with a single submedian pair of long hairs below vertex margin.
Mesosomal dorsum without erect hairs. Pronotal disc smooth and shiny. Mesonotum shiny,
anteriorly smooth, posteriorly sharply punctate. Mesepisternum shiny, with faint longitudinal rugae.
Metepisternum densely and relatively coarsely punctate. Propodeum contiguously finely punctate;
dorsal face slightly convex in profile; medial carina absent from dorsal face except at summit of
declivity, where it is low and broadly rounded; medial carina present on declivitous face; spiracular
prominence well-developed, spiracular opening directly obliquely distad; propodeal teeth short and
distance between their bases less than distance between spiracles.
Gastral terga shiny and unsculptured except for close fine piligerous punctures; all terga without
erect hairs.
Head light brownish with yellowish clypeus and mandibles; mesosoma light brownish, paler on
pronotum and sides; gaster dark brown but tergum 1 yellow, concolorous with petiole.
TYPE MATERIAL
Holotype worker, CENTRAL AFRICAN REPUBLIC, Songha-Mbaéré Pref., Parc National

Dzanga-Ndoki, "37.9 km 169°S" Lidjombo, 2°22'N 16°10'E, 360m, 20-28 May 2000 (B.L. Fisher,
#4128), beating low vegetation, rainforest, in CASC.
ETYMOLOGY
The name is an arbitrary combination to be treated as a noun in apposition.
DISCUSSION
The sharply contrastingly bicolored gaster and lack of erect hairs on both the mesosomal and
gastral dorsa are distinctive for this species. Photomontage images are available at www.antweb.org
as Axinidris sp. 2.
Axinidris murielae Shattuck
Axinidris murielae Shattuck, 1991: 116-117; figs. 23-25, 35 (w, q). CAMEROUN, Nkoemvon
(D. Jackson) (BMNH) examined.
Worker diagnosis. Antennal scape with more than 5 and pronotum with 2 erect hairs;
pronotum mostly smooth and shiny; medial carina present and obtuse in profile; distance between
spines less than PPW; head and body reddish brown.

0.26; OVD 0.24-0.31; PNW n/a; PPW 0.31-0.46; WL 0.89-1.30. Indices. CI 87-89; CNI 90-129;
OI 26-34; SI 80-85.
DISCUSSION
Worker and queen have been adequately described by Shattuck (1991); the male remains
unknown.
The two species A. gabonica and A. murielae are similar and may ultimately prove to belong to
a single somewhat variable species. For the present the two are separable as noted in the key: A.
murielae has 4-6 erect hairs on the pronotum (only 2 in A. gabonica) and numerous erect hairs on
the front of the head (only 4 in A. gabonica) and also with erect hairs on gastral terga 2-4 (one in A.
gabonica).
The types of A. murielae are from Cameroun; Shattuck (1991) also recorded specimens from
Ghana. I have also seen a few specimens from the vicinity of Busumbuli in the Kakamega Forest of
Kenya. Both samples were taken by tree fogging, one from Teclea nobilis and one from Heinsenia
diervillioides.
Axinidris namib, new species

Figures 7, 17, 27
Worker diagnosis. Scape shaft and pronotum and gastral terga 1-3 without erect hairs;
hypostomal area with erect hairs; mesepisternum with distinct longitudinal rugae; medial carina
present on posterior half of dorsal face of propodeum and continued down declivitous face.
Worker measurements (mm) (n = 1). HW 0.81; HL 0.86; SL 0.64; El 0.24; OVD 0.31; PNW
0.51; PPW 0.36; WL 1.03. Indices. CI 94; CNI 100; SI 79; OI 30.
Worker description. Frons and vertex moderately shiny, finely longitudinally strigulate
between close minute punctures; gena shinier, finely coriarious between sparse shallow punctures;
malar area shiny, sharply strigulate and sparsely punctate. Each frontal carina with a single erect
hair at lower and upper ends; frons with a pair of very short submedian hairs below summit of
vertex; scape shaft without erect hairs; hypostomal area with several short erect hairs on each side.
Dorsum of mesosoma without erect hairs. Pronotal disc moderately shiny and coriarious to
strigulate between sparse minute punctures. Mesonotum smooth and shiny anteriorly, posterior
three-fourths moderately shiny and with sparse fine longitudinal rugae. Mesepisternum shiny
between relatively coarse more or less longitudinal rugae. Propodeal dorsum weakly shiny,
contiguously punctate and with irregular short, fine rugae, side (below spiracle) shiny between
sparse, short rugae; spiracular prominence distinct, spiracular opening directed obliquely distad;
medial carina present on posterior half of dorsal face and on declivitous face, low and broadly
rounded in profile at summit of declivity; propodeal spines short and nearly right-angular in dorsal
view, distance between their apices less than distance between spiracles.
Gastral terga moderately shiny, coriarious between close minute punctures; terga 1-3 without
erect hairs, 4 with one submedian pair near base.
Head and mesosoma reddish brown, mesosomal dorsum slightly darker; gaster darker reddish
brown; clypeus and mandibles more yellowish.
TYPE MATERIAL
Holotype worker, NAMIBIA ("S. W. AFRICA"), Keetmans Dist., 1 km W Mata Mata,

Welverdiend Farm No. 328 (25°47'S 19°59'E), 19 October 1972 (C.L. Hogue), in LACM.
ETYMOLOGY
The name is derived from that of the country of origin and is here a noun in apposition.
DISCUSSION
The presence of a number of short erect hairs on the hypostomal area is unusual among the
species of Axinidris, but does occur in several other species (acholli, lignicola, stageri). From A.
acholli, A. namib differs by its much smaller size and the less spectacularly developed medial carina
and propodeal spines. Both A. lignicola and A. stageri possess conspicuous erect pronotal hairs.
Axinidris nigripes Shattuck
Axinidris nigripes Shattuck, 1991: 117-118; fig. 35 (w). GHANA, Aburi (P. Room) (BMNH)
examined.
Worker diagnosis. Scape and pronotum with erect hairs; medial carina present; distance
between spines exceeding PPW; hairs of head and mesosoma brown, coarse and seta-like.
DISCUSSION
This species is similar to both A. luhya and A. palligastrion. It differs from the latter in its
larger size, deeper clypeal notch and differing color pattern, the gaster being darker than the head
and mesosoma, the reverse of the pattern in A. palligastrion. From A. luhya, A. nigripes differs by
the coarser, darker mesosomal hairs (fine and yellowish in A. luhya.
Axinidris occidentalis Shattuck
Axinidris occidentalis Shattuck, 1991: 118; figs. 26-28, 35 (w). LIBERIA, Paiata (= Payeta) (J.
Bequaert) (MCZC) examined.
Worker diagnosis. Scape and pronotum without erect hairs; pronotum weakly shiny, sharply
sculptured and densely pubescent; medial carina present; T3-T4 each with several erect hairs.

0.23; OVD 0.31-0.35; PNW 0.53-0.58; PPW 0.34-0.41; WL 1.02-1.10. Indices. CI 99-101; CNI
133-153; OI 23-25; SI 64-69.
DISCUSSION
In A. occidentalis the pronotum lacks erect hairs, but they are present on gastral terga 3 and 4,
and the distance across the propodeal spines is subequal to the propodeal width, a combination of
features unique to this western species.
Axinidris okekai, new species

Figures 8, 18, 28
Worker diagnosis. Antennal scape shaft and pronotal disc (Fig. 18) without erect hairs and
gastral terga 3-4 each with 2-4 moderately long erect hairs; upper frons with 2 hairs; propodeal
spines (dorsal view) short and right-angular, obtuse in profile; base of propodeum convex in profile,
medial carina obtusely angulate at declivity; pronotum polished between sparse piligerous punctures.

114; OI 25-30; SI 81-91.
Worker description. Frons moderately shiny, with sparse, fine irregular longitudinal strigulae
between contiguous minute punctures; vertex shiny between sparse piligerous punctures; gena shiny
and weakly imbricate to smooth and malar area closely strigulate. Upper frons with 1 pair of erect
hairs, each about as long as maximum width of first funicular segment; frontal area with 1 pair of
erect hairs and each frontal carina with 2 pairs. Front of head with sparse, short appressed brownish
hairs. Shaft of antennal scape without erect hairs, but with sparse, short appressed brownish hairs.
Pronotal disc polished and shiny between sparse minute piligerous punctures; anterior half of
mesonotum similar, posterior half reticulose; mesepisternum shiny and weakly sculptured anteriorly,
becoming contiguously, but shallowly, punctate behind; metanotal spiracle prominent in profile, but
set so close to base of propodeum that it is not readily visible. Propodeum slightly shiny and
contiguously finely punctate; base face convex in profile; spiracular prominence weak, opening
directed posterolaterally; medial carina present only at summit of declivity, low and obtusely
angulate in profile; propodeal spines short, in dorsal view right-angular and distance between their
outer margins much less than width of propodeum at spiracles; spines obtuse to right-angular in
profile.
Gastral terga shiny and weakly imbricate between sparse, minute piligerous punctures bearing
short, brownish appressed hairs; terga 1-2 without erect hairs, 3-4 each with 2-4 erect hairs.
Head and body dark brownish, mesosoma slightly paler than head; mandibles yellowish at base
of teeth.
TYPE MATERIAL
Holotype and 6 worker paratypes: KENYA, Kakamega Distr., Isecheno, 21 March 2002 (R. R.
Snelling, #02-162), on vines. Additional paratypes: 2 workers, Kakamega Forest, 7-11 February
1999 (T. Wagner), ex fogging Teclea nobilis; 2 workers, "Colobus Trail", Kakamega Forest, Sept.-
Oct. 2001 (W. Freund & C. Schmidt), ex fogging Heinsenia diernilliodes. Holotype and several
paratypes in LACM; paratypes also in AKRI, BMNH, MCZC and NMKC.
ETYMOLOGY
This species is dedicated to my friend, Wilberforce Okeka of KEEP (Kakamega Environmental

Education Programme, Isecheno), who was so helpful during my visits to Kakamega Forest.
DISCUSSION
In addition to the above material, I have seen a single worker that appears to belong to this
species: CAMEROUN, Mbalmayo, Nov. 1993 (N. Stark; BMNH). It differs from the types in being
more reddish, the mandibles largely yellowish; the head and pronotum are less closely sculptured.
The few specimens in the type series are consistent in their features. In stature and color it is
most like A. bidens but is immediately separable by the lack of erect hairs on the pronotal humeri
and propodeal spines.
Axinidris palligastrion Shattuck
Axinidris palligastrion Shattuck, 1991: 118; figs. 29-31; fig. 35 (w). GHANA, Nswam (=
Nsawam) (P. Room) (BMNH) examined.
Worker diagnosis. Scape without and pronotum with erect hairs; medial carina present;
distance between spines equal to PPW; pronotum smooth and shiny; mesonotum without rugae;
gaster yellowish red, not concolorous with mesosoma.

0.30; OVD 0.29-0.32; PNW n/a; PPW 0.42-0.45; WL 1.05-1.16. Indices. CI 91-94; CNI 107-134;
OI 31-35; SI 72-78.
DISCUSSION
Although similar to A. nigripes, this species is separable as noted in the discussion of that
species. The gaster is distinctly lighter in color than the head and mesosoma, but is not distinctly
bicolored as in the otherwise similar A. mlalu, a species that lacks erect pronotal hairs (present in A.
palligastrion).
Axinidris stageri, new species

Figures 10, 20, 30
Worker diagnosis. Apex of scape distinctly surpassing vertex margin; scape and pronotum
with numerous erect hairs; medial carina profile elevated and acute at apex, continued down
propodeal declivity.
Worker measurements (mm) (n = 3). HL 0.81-0.89; HW 0.20-0.22; SL 0.68-0.76; EL 0.11-

0.13; OVD 0.28-0.34; PNW 0.49-0.58; PPW 0.33-0.38; WL 0.97-1.11. Indices. CI 96-98; CNI 5-6;
OI 25-26; SI 87-88.
Worker description. Head shiny between dense minute punctures; gena weakly imbricate and
with scattered minute punctures; malar area polished and unsculptured. Apex of antennal scape
exceeding vertex by more than greatest width of scape. Numerous suberect hairs on head margins in
frontal view, especially above eyes; front of head and vertex with numerous short to moderately long
suberect and erect hairs; scape shaft with sparse long erect hairs and numerous short suberect hairs.
Pronotal disc shiny and weakly imbricate. Mesonotum shiny, with coarse irregular longitudinal
rugae evenly sloping in profile and metanotal spiracle clearly projecting. Mesepisternum shiny, with
finer longitudinal rugae. Propodeal dorsum less shiny than pronotum, coarsely imbricate; side shiny
and irregularly longitudinally rugose; spiracular tubercle prominent in dorsal view, anterior edge of
spiracle moderately elevated so that opening is directed obliquely distad; medial carina sharp,
strongly elevated behind and acute in profile, continued ventrad on posterior face; propodeal spines
reduced to minute teeth, distance between their apices much less than propodeal width at spiracles.
Pronotum (Fig. 20) with 20+ erect hairs; mesonotum and propodeum each with 10+ similar hairs.
Gastral terga subpolished and weakly imbricate; all terga with scattered erect and suberect hairs
in addition to sparse appressed pubescence.
Head and body uniformly yellowish red, appendages paler.
TYPE MATERIAL
Holotype and two paratype workers, TANZANIA, Bunduki, Uluguru Mts., 6-15 January 1964
(K.E. Stager), all in LACM.
ETYMOLOGY
This species is dedicated to Dr. Kenneth E. Stager, former Curator of Ornithology at the LACM,
in appreciation for the many samples of ants that he collected for me in various parts of the world.
DISCUSSION
This species is easily recognized by the abundance of erect pilosity on the body and appendages
and by its reddish color. Morphologically it is most similar to A. tridens, but the medial carina is
higher than in available material of that species. Additionally, A. tridens is much less hairy, with no
erect hairs on the antennal scapes and the gastral terga.
Axinidris tridens (Arnold)
Engramma tridens Arnold, 1946: 58 (w). MALAWI, Mlanje (= Mulanje) (SAMC).

Axinidris tridens: Shattuck, 1991: 119-120.
Worker diagnosis. Antennal scape shaft, pronotum and gastral terga without erect hairs;
pronotum densely pubescent; medial carina acute in profile.
Worker measurements (mm) (n = 2). HW 0.81-0.88; HL 0.86-0.95; SL 0.72-0.82; OVD 0.33;

PNW n/a; PPW 0.31-0.34; WL 1.07. Indices. CI 93-94; CNI 64-95; OI 24-28; SI 88-93.
Worker description. The worker has been adequately redescribed by Shattuck (1991).

DISCUSSION
Shattuck (1991) removed this species from Engramma to Axinidris. He also reported one
worker from St. Lucia Estuary, Natal, SOUTH AFRICA. This is the only specimen known, other
than the types.
ACKNOWLEDGEMENTS
For the loan of specimens in their care, I am deeply indebted to Barry Bolton (BMNH), Stefan Cover
(MCZC), Brian Fisher (CASC) and Hamish Robertson (SAMC). My field work in Kenya was
facilitated and encouraged by Scott Miller. Hans Herren, Susan Kimani, Nixon Onyimbo and Lucie
Rogo, all of ICIPE, greatly aided my field work in Kenya, with further assistance at Isecheno from
Wilberforce Okeka, Boniface Omolo, Andrew Chiapya and David Makonjio. Numerous canopy
fogging samples from Kakamega Forest that yielded Axinidris specimens were contributed by
Marcell Peters and Thomas Wagner (AKRI). Thanks are also extended to Giar-Ann Kung for her
patient assistance during my sessions with the scanning electron microscope.
DEDICATION
This paper is dedicated to Ed Wilson, not only in recognition of his contributions to ant taxonomy,
but also for the advances in understanding the biogeography and ecology of ants and of his many
stimulating contributions toward understanding social behavior, from insects to humans.
LITERATURE CITED
Arnold, G. 1946. New species of African Hymenoptera. No. 6. Occasional Papers of the National
Museum of Southern Rhodesia 2: 49-97.
Bolton, B. 1994. Identification guide to the ant genera of the world. Harvard University Press,
Forel, A. 1905. Miscellanea myrmécologiques, 2. (1905) Annales de la Société Entomologique de
Belgique 49: 155-185.
Harris, R.A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, California
Department of Food and Agriculture 28: 1-31.
Mayr, G. 1872. Formicidae Borneenses collectae a J. Doria et O. Beccari in territorio Sarawak annis
1865-1867. Annali del Museo Civico di Storia Naturale de Genova 2: 133-155.
Santschi, F. 1919. Nouvelles fourmis du Congo Belge du Musèe du Congo Belge, à Tervueren.
Revue Zoologique Africaine 7: 79-91.
Shattuck, S.O. 1991. Revision of the dolichoderine ant genus Axinidris (Hymenoptera: Formicidae).
SystematicEntomology 16: 105-120.
Shattuck S.O. 1992. Generic revision of the ant subfamily Dolichoderinae (Hymenoptera:
Formicidae). Sociobiology 21: 1-181.
Weber, N.A. 1941. Four new genera of Ethiopian and Neotropical Formicidae. Annals of the
Wheeler, W.M. 1922. Ants of the American Museum Congo Expedition. A contribution to the
myrmecology of Africa. II. The ants collected by the American Museum Congo Expedition.
Bulletin of the American Museum of Natural History 45: 39-269.
Figures 1-4. Frontal view of head of: 1, Axinidris acholli; 2, A. bidens; 3, A. gabonica; 4, A. icipe.
Figures 5-8. Frontal view of head of: 5, A. lignicola; 6, A. luhya; 7, A. namib; 8, A. okekai.
Figures 9-10. Frontal view of head of: 9, A. hypoclinoides; 10, A. stageri. Figures 11-13. Lateral
view of mesosoma of: 11, A. acholli; 12, A. bidens; 13, A. gabonica.
Figures 14-19. Lateral view of mesosoma of: 14, A. icipe; 15, A. lignicola; 16, A. luhya; 17, A.
namib; 18, A. okekai; 19, A. hypoclinoides.
Figure 20. Lateral view of mesosoma of A. stageri. Figures 21-24. Dorsal view of mesosoma of: 21,
A. acholli; 22, A. bidens; 23, A. gabonica; 24, A. icipe.
Figures 25-30. Dorsal view of mesosoma of: 25, A. lignicola; 26, A. luhya; 27, A. namib; 28, A.
okekai; 29, A. hypoclinoides; 30, A. stageri.
Taylor, R. W. 2007. Bloody funny wasps! Speculations on the evolution of eusociality in ants,
pp. 580-609. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances in ant systematics
BLOODY FUNNY WASPS!

SPECULATIONS ON THE EVOLUTION OF
EUSOCIALITY IN ANTS
Robert W. Taylor
Australian National Insect Collection,
CSIRO, Division of Entomology, Canberra, ACT, Australia.
bob.taylor@homemail.com.au
ABSTRACT
Following review of formicid characters a new model for evolution of ant eusociality is
presented. This “semisocial ancestry hypothesis” has the following steps: (1) Ants are descended
from a solitary wasp which accumulated and collectively reared its brood in acellular nests
(without individual brood cells). Metapleural glands delivering antibiotic compounds evolved in
adults as a correlated adaptation for nest sanitation. (2) Such wasps gathered to form cooperating
family groups of congenerational alate females and their offspring. (3) Female dealation
evolved, correlated with pedestrian foraging. Colonies now comprised coeval dealate females
rearing alate daughters and sons. Competition generated reproductive and non-reproductive
castes in the parental generation, resulting in semisocial colonies, eventually with one dealate
mated reproductive queen per nest, and a force of congenerational non-reproductive dealate
“workers”. These semisocial formicids progressively acquired petiolar nodes, elbowed antennae,
increased longevity and ant-like larvae. (4) Eusociality began when daughter females dealated
prematurely and joined the parental-generation worker force. (5) Daughter (eusocial) workers
adaptively superceded parental-generation (semisocial) workers and, with the evolution of
mechanisms for caste determination, lost mesosomal complexity to constitute eusocial colonies
like those of modern ants, with alate/dealate female reproductives and mesosomally-reduced
daughter workers. (6) Some of these gave rise to modern ants. (7) Competition between eusocial
and semisocial formicids resulted in extinction of the latter, leaving eusocial ants transcendent.
These events occurred early in formicid history. The fossil record indicates that eusocial
ants referable to several modern subfamilies were present by late Cretaceous times.
The semisocial formicids could have included derived species either with morphologically
undifferentiated apterous mesosomally-reduced females, or with alate/dealate queens and
congenerational apterous workers. The extinct upper Cretaceous Sphecomyrma freyi is possibly
the female or worker of one of these.
It is proposed that the worker caste in modern ants has evolved more than once, and that
eusociality arose several times in the semisocial ancestral group, producing separate lineages of
eusocial ants, several of which survived to constitute the modern fauna.
The unusual reproductive biology of the aberrant ponerine genus Diacamma is reviewed.
Key words: Diacamma, Formicidae, Hymenoptera, sociality, Sphecomyrma.

Taylor: Bloody Funny Wasps! 581
INTRODUCTION
Ants are classified in the family Formicidae assigned to suborder Apocrita of order
Hymenoptera. Within Apocrita they are allocated to superfamily Vespoidea. The several other
vespoid families comprise the “true” wasps. Three apocritan superfamilies, Vespoidea,
Chrysidoidea and Apoidea, are referred to collectively as aculeates. Superfamily Apoidea
includes bees and sphecid wasps. This classification represents ants as specialized vespoid
wasps.
This essay reviews salient characteristics of the Formicidae, especially their
synapomorphies. It considers the evolutionary origins and adaptive features of relevant
characters and the possible sequence of their phylogenetic acquisition, their possible
evolutionary interactions, the possible effects of each on options for future evolution, and their
roles in the evolution of formicid sociality. Some propositions are new to the scientific
discussion of ant evolution. It is hoped that they might encourage further discussion and
research.
Nomenclature here follows Bolton (2003). “Ponerinae” (with quotes) or “Ponerinae sens.
lat.” refer to that subfamily in the sense of the previous Emery/Wheeler classification. Bolton’s
terms “mymeciomorph”, “poneromorph” etc, refer to sets of putatively related subfamilies. The
words “formicid” and “formicids” refer to taxa which have contributed directly to ant
phylogeny. The term “standard reproductives” refers to alate females which later dealate (these
are also referred to as alate/dealate females).
Many general statements about ants are not been referenced below. They are well covered
by Wilson (1971) and Hölldobler and Wilson (1990).
Symplesiomorphies of the Formicidae
Several prominent symplesiomorphies of ants are synapomorphies of order Hymenoptera.

Thus (1) sex is genetically determined by haplodiploidy (males haploid and impaternate, females
diploid) as in all Hymenoptera (Crozier and Pamilo, 1996), (2) ant mouthparts, apart from the
labrum and mandibles, form a labiomaxillary complex with an associated infrabuccal pouch, and
(3) mature ant larvae, prior to pupation, enclose themselves in cocoons of silk spun from their
labial glands (the many formicid exceptions result from repeated secondary loss of this
character).
Other formicid characteristics are synapomorphies of suborder Apocrita: (4) The first true
abdominal segment is fused with the thorax to constitute the propodeum, configuring a novel 4-
segmented thoracic tagma, the mesosoma, and (5) a deep constriction separates the propodeum
and the second true abdominal segment forming a waist between the mesosoma and the
segmentally reduced abdomen (termed the metasoma). A small anterior exoskeletal section of
the first metasomal segment forms a ball-joint within the propodeum. This facilitates rotation of
the sting-bearing metasoma. The operative musculature is largely in the propodeum. Abdominal
articulation from the metathorax would be compromised by its leg-bearing function, hence the
propodeum has evolved.
Some symplesiomorphies of ants are synapomorphies of the aculeate Apocrita. These
include (6) modification of the ovipositor to function as a venom-injecting sting, (7) apodous
larvae, and (8)(probably) antennae primitively with 12 articles in females, 13 in males.
Further putative formicid symplesiomorphies relevant here are: (9) ocelli present, (10) a
univoltine life cycle with one new generation produced each year, (11) sperm in mated females
retained in the spermatheca and used to fertilise eggs for long periods following mating, (12)
adults are liquid feeders, (13) larvae are entomophagous, (14) females collect prey to provision
larvae, (15) prey is delivered to brood whole, without dissection, (16) males take no part in the
lives of females or offspring following copulation, and (17) navigation is visually mediated. The
points of phylogenetic origin of these attributes in formicid evolution are less obvious than for
those features mentioned previously. All are acceptable as putative formicid symplesiomorphies
(Carpenter, 1991).
Formicid synapomorphies
Synapomorphies are the particular features by which we recognize and characterize groups
of organisms. The truly homologous synapomorphic characters of any taxon are represented
nowhere outside that taxon. Also, where several synapomorphies are recognized, any one of
them, on its own (with due recognition of its tansformations), can distinguish a subject taxon
from its sister taxa.
Putative formicid synapomorphies relevant here include: (1) female antennae elbowed, with
an unusually elongate basal segment (the scape), (2) wing bases in reproductive females
structured to facilitate wing-shedding (the action “dealation”, the process “allectomy”), (3)
presence of a permanently apterous (wingless) non-reproductive worker caste, (4) metapleural
glands present in both adult sexes, (5) first metasomal segment (2nd true abdominal) forming a
posteriorly constricted waist node, the petiole, and (6) nests acellular, without individual brood
cells. (See Bolton, 2003, for other formicid synapomorphies).
Synapomorphic character states are uniquely and separately acquired in phylogeny. It
follows that when several synapomorphies are recognized within a taxon they would usually
have arisen in temporal evolutionary succession. The succession can, nonetheless, be so rapid
that sets of characters may appear almost simultaneously in phylogeny when organisms are
adapting rapidly to extreme selection pressures. Indeed, separate adaptively-related characters
probably arise in concert under some circumstances (this might have been the case with co-
evolution of acellular nesting and metapleural glands in ants – see below).
The first appearance of each eventual synapomorphy as a newly evolved character would
often represent some sort of adaptive break-through, contributing to relative success and
diversification of the newly equipped species. Acquisition of a further such character in the
accumulation of a set of successively derived future synapomorphies, being a unique event,
could occur in only one of the lineages within the subject clade, leaving the remainder as a
paraphyletic rump exhibiting the first acquired character but not the second.
Thus, the evolutionary history of acquisition of the formicid synapomorphies listed above
must have involved a chain of successive steps, in which taxa equipped with newly evolved
characters survived, while their sister groups lacking those characters did not. This process must
have been repeated at least often enough to match the number of character acquisitions, and the
loss of sister groups must have been complete at each step – how else could so many
characteristics of modern formicids be considered synapomorphies. In effect a group possessing
multiple synapomorphies could have not just one common ancestor, but a chain of common
ancestors, and its lineage might have historically experienced a sequence of extreme diversity
bottlenecks.
Characters homoplasious (evolved convergently, and analogous but not homologous) with
any of the synapomorphies recognized in a current natural taxon would occur only in separate
taxa. These would lack some or all of the remaining synapomorphic character set
Importantly, the most recently acquired apparently synapomorphic character observed in a
contemporary taxonomic group could in reality comprise a set of successfully deceptive
homoplasious characters separately evolved in the immediately ancestral stock. This is much
more likely than with previously evolved characters. We will revisit this proposition below.
Characters resembling most of the above formicid synapomorphies and symplesiomorphies
are present as homoplasies elsewhere in nature. Examples include haplodiploidy, production of
pupal cocoons, elbowed antennae, univoltinism, visual navigation etc. This is to be expected,
and does not diminish the arguments presented here.
Metapleural glands – the classic formicid synapomorphy
Metapleural glands are characteristic of family Formicidae. No antecedent or similar

structures are known in other taxa. Their absence in several genera, e.g. Oecophylla,
Dendromyrmex, Polyrhachis, Camponotus (except C. gigas (Taylor, 1992) and at least two other
species, S.O. Shattuck & G.D. Alpert, pers. coms.) and various males (perhaps those of the
majority of ants) is clearly due to secondary loss (Brown, 1968; Hölldobler & Engel-Siegel,
1985). Their cuticular components can often be clearly discerned in fossils. Wheeler (1928: 112)
first drew attention to the presence of metapleural glands as a character defining the Formicidae.
The glands are bilateral, situated posterolaterally on the mesosoma. Each has an accessory
cuticular chamber, usually marked by an external swelling, with a small opening to the exterior.
Internally a specialized section of the inner chamber wall receives unicellular ducts from a
cluster of relatively large cells lying in the body cavity immersed in haemolymph (Hölldobler &
Engel-Siegel, 1985). The structure is almost uniform throughout the Formicidae, with the
specialized section of chamber wall usually a flat sieve-like disc, but in Nothomyrmecia,
Myrmecia and some other ants a microperforate vermiform cuticular collecting sac receives the
cell ducts, the cell mass is relatively large and the cells relatively small (pers. obs.; Tullock et al.,
1962; Hölldobler & Engel-Siegel, 1985)). There seems no reason to doubt that metapleural
glands were uniquely evolved and are a genuine formicid synapomorphy.
The glandular fluid contains antibiotic molecules which are proof against various soil
bacteria and fungi (Beattie et al., 1986; Veal, Trimble & Beattie, 1992; Mackintosh et al., 1995).
Its function is clearly protection of ants, especially their larvae, from contamination or infection.
Adult ants groom more actively than other aculeates, and this spreads metapleural gland
secretion on their bodies and elsewhere in the nests. The gland does not manufacture the
chemicals it secretes, but extracts them from the hemolymph. Some of the compounds involved
are found in other insects as components of defensive arrangements equivalent to, but very
different from, the vertebrate immune system (Kaaya, 1993; Casteels, 1998).
Metapleural glands are important in the social biology of ants. Their adaptive significance is
obvious, and their first appearance was doubtless profoundly significant in ant evolution
The powerfully antibiotic metapleural gland secretions are lethal to pollen grains. This
explains why ants are barely involved in pollination of flowering plants, and why many flowers
are adapted to exclude or repel ants (Beattie et al., 1986). While the acquisition of metapleural
glands was a major step in ant evolution, the prospect of a formicid role in pollination was
simultaneously closed. Pollination is, of course, a prominent activity among other aculeates,
especially in bees
The acellular nest as a synapomorphy of family Formicidae
Various taxa of non-formicid aculeates form reproductive social groups ultimately

comprising colonies of one or several coeval females with their brood and adult offspring. There
is sometimes also a non-reproductive, exclusively female worker caste. Colonies are typically
housed in nests ranging from simple excavations to elaborate constructions. Many of the
between-group similarities seen in aculeate sociality and nesting behavior are homoplasious.
Comparative studies of living aculeates with varying levels of sociality and nest complexity
reveal that both features are derived ultimately from stocks in which solitary, winged, mated
non-social females excavated and provisioned simple nests, each accommodating a single
offspring. As more offspring came to be assembled in nests they were, apparently from the
beginning, typically housed in separate individual brood cells constructed by the parental
females, even when only two or three immatures were involved. Aculeate larvae are primarily
entomophagous, and the likelihood of cannibalism among ravenous larvae clustered in undivided
nest chambers with immobilized prey is thought to explain the initial adaptiveness of larval
segregation. Isolation of individual larvae in such “cellular” nests would likely also impede the
spread of infectious disease vectors and parasites, and reduce the risk that all of the brood would
be lost in a predatory attack.
Modern nests, no matter how elaborate, are indubitably cellular in virtually all non-formicid
aculeates – specifically constructed to segregate each developing individual from its siblings.
Cellular nests are remarkably persistent in relevant lineages, and the structural integrity of nests
with sections assembled from close-packed hexagonal cells providing strong architectural
elements has contributed to the evolution of large, intricate nests such as those of the honeybees
and vespine wasps.
On the other hand, the colonies of all ants reside in continuous nest spaces, which house the
various brood stages in company with resident adults. The nest space is often a simple chamber,
but it may be elaborately divided; either randomly, as when colonies nest in leaf litter; or
structurally, as when large nest mounds contain numerous interconnected chambers. Brood is
often sorted in stage-specific groups, but no ant is known to house individual immatures in
individual cells equivalent to those of other aculeates. Ant nests are “acellular”.
Accommodation of brood in continuous nest spaces is rare among aculeates. Reported
examples in solitary sphecid wasps (three species in subfamily Sphecinae, one each in Larrinae
and Crabroninae) (Krombein, 1967) are barely exceptions. Loosely partitioned cells are
constructed successively in restricted linear spaces by parental females, and chambers with
several larvae result when partitions are omitted or later break-down. Krombein reviewed three
related Isodontia species (Sphecini) “one of which makes individual cells, another with a single
brood chamber but sometimes with individual cells, and one which always makes a single brood
chamber”.
Females of the solitary bee Megachile policaris provision individual brood chambers (of
which there may be several) with a pollen mass in which eggs are laid, eventually producing a
congregation of larvae. According to Krombein this species “differs markedly from other known
megachilid bees and, in fact, from almost all aculeate Hymenoptera in that several larvae
develop amicably in a single large brood cell with no apparent cannibalism”. All species of
Lithurgus (Megachilidae) have similar behavior (C.D. Michener, pers. comm.). Bumble bees
(Apidae: Bombini) lay up to 16 eggs in distendable cells. The larvae congregate initially, but
later separate themselves with silk partitions (Michener, 1974).
Colonies of the incipiently eusocial allodapine bees (Xylocopinae: Allodapini) and their
brood are housed, like ants, in acellular nests. These frequently occupy the slender tubular pith
cavities of herbaceous plants or other similar spaces, where the accommodation of numerous
individual brood cells would be physically constrained. The adaptive rationale for these nesting
arrangements is clear. Allodapines are believed to be descended from ancestors which
constructed loosely partitioned cells not unlike the sphecids discussed above. Their acellular
nests are to that degree a secondary development. In evolutionary terms the loss of brood cells
occurred in correlation with the development of novel nesting arrangements whereby otherwise
inaccessible tubular spaces were able to provide accommodation for populous colonies
(Michener, 1974). Acellular nesting is an allodapine synapomorphy acknowledged to have
arisen uniquely from ancestral stock. Its origin establishes a single exception to the aculeate
norm (M.P. Schwarz, pers. comm.).
These putative exceptions all involve descent from stock with few larvae and little evident
commitment to cellular nesting. There is no evidence of reversals from cellular to non cellular
nesting among architecturally more advanced aculeates, suggesting that cellular nesting behavior
might be functionally irreversible, particularly when nests are substantially complex.
The above considerations imply that cellular nests might never have featured in the ant
lineage (or were never developed to a level precluding evolutionary reversal). Formicids are
therefore likely to have originated from a solitary flying wasp which accumulated its larvae in a
single nest space without cellular partitions. This behavioral acquisition marked the inception of
the formicid phylogenetic clade, and the acellular nest has persisted through millions of years to
stand eventually as a synapomorpy of the family Formicidae.
The existence of ants today could have depended on the absence of cannibalism in the
larvae of that ancestor. Evolution is like that – very chancy! There would, of course, be extreme
selective pressure against larval cannibalism once such nesting arrangements were established.
The earliest formicids
The first formicids would, by virtue of their ancestry, have displayed haplodiploidy; a
labiomaxillary complex; ocelli; a propodeum, constricted waist and venom-injecting sting.
Adults would have used liquid food and fertilized their eggs with spermathecally-stored sperm.
They would have practiced general brood care and foraged for insect or other similar prey
returned to the nest as larval food without dissection. Reproduction would have been univoltine,
the pupae enclosed in cocoons, and brood overwintered in the nests, probably as larvae. There
would be strong selective pressure for overwintering by adults in nests, which would provide
protection for hibernating brood. Navigation would have been visually mediated.
The proclivities of modern ants imply that nests would have been constructed in the soil,
and that mature larvae within the nests would have been banked with soil by workers prior to
spinning their pupal cocoons (as in Nothomyrmecia, Myrmecia, Amblyopone and other
“primitive” ants). Observations on living solitary wasps indicate that foraging would likely have
been epigaeic and diurnal, with the prey encountered above ground, on vegetation and other
exposed surfaces, but not retrieved from within the soil, leaf litter or fallen rotting wood.
The brood would be less protected from disease or fungal infection than in cellular nests.
For this reason the evolution of a precursor to the metapleural gland, capable of antibiotic
delivery, is likely to have occurred in close correlation with acellular nesting. The near
uniformity of metapleural gland accessory cuticular structures implies that the glands would
probably have rapidly developed to resemble the modern organs. Recognition of early gland
developmental stages might be difficult in fossils because the cells would probably have opened
minutely to the body surface prior to evolution of cuticular reservoir chambers.
Cooperation of two or more females in nest foundation, egg laying, brood care, nest defense
and foraging would improve brood protection, since the nest could be permanently guarded.
Communal aggregation would also support evolutionary increase in colony populations and
reproductive productivity, and have an additive effect on efficiency of biocidal protection from
metapleural gland secretions, supporting increased nest size. Aggregation would almost certainly
have been selectively favored. It would have assembled “primitively social” “polygynous family
groups”, as discussed by West-Eberhard (1978).
The evolution of progressive provisioning of the brood would occur as a consequence of the
association of adults and larvae in nests. This behavior is found elsewhere among vespoids only
in some highly social Vespidae – all other vespoids aside from ants are mass-provisioning (J.M.
Carpenter, pers. comm.).
Joint colony foundation (pleometrosis) by recently mated females is practiced today by
Nothomyrmecia, some Myrmecia species and other ants (Hölldobler & Taylor, 1983; Wheeler,
1933, Wilson, 1971). The characteristics of this behavior might be homologous with those of the
ancestral nesting arrangements hypothecated here.
The ancestral formicids would have generally resembled modern, flying solitary aculeate
wasps, except that they would have possessed metapleural glands or antecedent structures.
The univoltine life cycle, brood synchronicity and longevity
Aculeates are generally univoltine, producing one brood each year. Eggs are usually laid in
spring. Adults are short-lived, few surviving more than about a year. In most cases colonies die
out each winter, with only a few individuals surviving in hibernation to continue the cycle the
following spring. In advanced species these are mated females, often found claustral in
excavated or preexisting hibernacular cavities (Evans and West-Eberhard, 1973). Overwintering
as prepupae or cocoon-enclosed adults is practiced in solitary taxa or those with low levels of
social development (Krombein, 1967).
Constructed nests may be perennial but most species construct new nests each year. Reuse
of nests is not common, even in socially advanced groups like polistine wasps (Matsuura and
Yamane, 1984). In a few cases, mainly in the bee subfamily Apinae, nests are occupied at all
seasons by queens, workers, and brood, with in-nest hibernation in areas of severe climate. Even
in these, adults are not long-lived and few individuals apart from queens, that tend to live longer
than workers, survive more than one active season (Michener, 1974).
Extreme longevity has been frequently reported in queen ants, and workers may survive for
several years at least (Hölldobler and Wilson, 1990). Thus at some point in the ant lineage
selection favored increased longevity, probably among the semisocial formicids hypothecated
below.
There is a general misconception that egg-laying by queen ants is continuous and that all
brood stages are present in ant nests at all seasons. Even the writings of myrmecologists can
indicate a lack of understanding that ants are primitively univoltine and that the life-cycles of all
ants are derived from univoltine ancestry.
Univoltinism is a significant trait of ants. It is demonstrated in colonies of the Australian
bulldog ant Myrmecia gulosa (Myrmeciinae), in which queens lay eggs for a period of only a
few weeks each spring. As hatching proceeds eggs are progressively replaced by small hatchling
larvae until only larvae remain. They grow throughout summer and autumn and overwinter as
large, incompletely grown individuals. The following spring larval growth resumes and mature
larvae begin to spin cocoons. This occurs at about the time egg laying is again initiated to
produce the following generation. Emergence of callow adults from the cocoons is complete by
early to mid summer. Overwintering larvae do not feed (and might be diapaused, though this has
not been investigated), and the workers cease foraging for insect prey during winter, when they
seldom depart the nests (pers. obs.).
Transformation of this life history has been frequent among ants. Myrmecia tarsata does not
overwinter brood. Eggs are laid in spring and emergence complete before the following winter.
During winter larvae are absent and foraging for larval food unnecessary. The nest entrances are
then sealed with soil, enclosing the overwintering broodless colony. Other Myrmecia species
seem to be bivoltine, producing a “fast” summer brood maturing in late summer from spring-laid
eggs, and a “slow” overwintering brood maturing in spring from eggs laid the previous summer.
A survey of the brood contents of many preserved colony series of Australian Sphinctomyrmex
species (Cerapachyinae) has demonstrated that they overwinter no brood (pers. obs.).
There is evident seasonal synchronicity of brood in the nests of most ant colonies
encountered in the field. Winter nests of many species contain only large larvae, while in spring
eggs, hatchling larvae and pupae are present. This applies to a range of taxa in both temperate
and tropical habitats. Dacetinops species in Borneo and lowland New Guinea are typical
(Taylor, 1985). These observations imply that the univoltine life cycle with overwintering larvae
is common among modern ants. Despite literature search I am unable to say with certainty
whether the queens of any ants at all are continuous egg layers. Reporting of seasonal brood
contents in ant nests has been inadequate for such analysis
Ant taxa with unusual derived life cycles include the North American primitive formicine
Prenolepis imparis. It has a seasonally reversed univoltine life cycle, with actively foraging
workers and feeding larvae in winter and summer closure of the nests (Tschinkel, 1987). The
Australian formicine genus Myrmecorhynchus, judging from its winter foraging activity, could
be similar.
The so called “Schneirla cycle” of ecitononine and aenictine army ants involves a number of
brood cycles each year, and these ants are multivoltine. Brood development is highly
synchronized in each colony, and cycles of colony migration are correlated with developmental
status of the brood. If the brood events of a single Schneirla cycle are represented on a clock
diagram and compared with a similarly derived diagram of the annual cycle of a univoltine ant
like Myrmecia gulosa the two figures are almost identical, except for their time scales, one of
about 6 weeks duration, the other of 12 months. It seems likely that this multivoltine system
originated by temporal compression and repetition of the annual life cycle of a univoltine
ancestor. The Australian amblyoponine army ant Onychomyrmex hedleyi has three cycles per
year homoplasious with those of the Ecitoninae. The ancestral condition is probably that
exemplified by the related Amblyopone australis, which is univoltine, with a brood cycle almost
exactly resembling that of Myrmecia gulosa (pers. obs.).
Univoltine life cycles have almost certainly been the norm throughout ant phylogeny, and
the various derived life cycles among living ants probably all evolved following the acquisition
of eusociality.
Allectomy as a formicid synapomorphy
Allectomy is a synapomorphy of family Formicidae. Dealation is practiced by fertilized

winged female ants after mating and aerial dispersal prior to colony foundation. Wing retention
is then no longer adaptive so the wings are discarded and formerly “alate” females become
“dealate”. Dealation is universal in winged female ants.
Flight for mating and dispersal prior to dealation is clearly adaptive, since wings are
developed in the majority of formicid species. Many ants have evolved secondarily worker-like
apterous (ergatoid) queens, or the queens have been evolutionarily superceded by mated workers
(gamergates) (Peeters & Ito, 2001). These usually mate outside the nests using sexual calling
pheromones to attract males, as in Rhytidoponera metallica (Hölldobler & Haskins, 1977).
Nothomyrmecia queens have very reduced wings, barely longer than the mesosoma (Taylor,
1978), and cannot fly. These vestiges might facilitate recognition of nubile virgin queens by
males. It is likely that some foraging, mated dealate foundress queens and numerous workers
would be abroad at the time of mating activity. Mistakes by males in mating with either of these
would involve gamete wastage, and would be sociobiologically maladaptive. Also, if queens
dealated immediately after copulation this would forestall undesirable supplementary matings.
Some species of Myrmecia have similarly vestigial wings (Clark, 1951).
Allectomy is rare elsewhere among Hymenoptera. The only reported examples appear to be
in the parasitic genera Leaiopria and Mimopria (Proctroproidea: Diapriidae) which are parasites
of termites (Naumann & Masner, 1985). Females shed their wings upon entering established
termite nests after flight from parental termitaria. The wings are used for dispersal (and perhaps
in mating flight), but are no longer functional in the confines of host termite colonies, so they are
discarded. The adaptive rationale is clear.
Allectomy in the ant lineage could have evolved only in females of stock which, though
winged, had largely or fully dispensed with flight while foraging, and were therefore already
capable of successfully provisioning their nests on foot. This important truism seems seldom to
have impressed myrmecologists, who have usually assumed that dealation first evolved because
wings would be obstructive in confined nest galleries (as in the aforementioned diapriids). This
might be true, but established prior pedestrian foraging would be necessary in any case. The
primary adaptive function of allectomy is more likely related to evolutionary loss through wing
redundancy in foraging.
The fossil record and other considerations suggest that ants arose as part of the late
Mesozoic radiation of insects following the Gondwanic origin and early diversification of
flowering plants beginning in the late Jurassic, about 130 million years ago (Grimaldi, Agosti &
Carpenter, 1997; Schultz, 2000). Potential prey organisms, especially relatively sedentary
phytophagous insects in dense populations, would have been readily accessible as prey for
ancestral formicids during those times (as is the case today with Nothomyrmecia and various
species of Myrmecia).
The proposition that pedestrian foraging would be functional in such circumstances is
entirely plausible. If an ant ancestor with biology similar to that reviewed above was able to
forage successfully without flight, natural selection would act to favor stocks where wing-
shedding was present, and to improve its efficiency. That scenario is accepted here.
Pedestrian foraging, nocturnality and hypogaeic behavior
The model proposed here posits that pedestrian foraging originated in formicid phylogeny
as an inevitable correlate of allectomy by mated females, prior to appearance of the “true”
eusocial worker caste. This may be considered the single most influential event in ant
phylogeny, notwithstanding the acquisition of metapleural glands. Some of the reasons for that
assertion relate to the activities of modern worker ants, and will be deferred (since eusocial
workers have not yet entered this discussion). Others are appropriate for review now.
The development of pedestrianism positioned the ancestral formicids for two profoundly
important further developments in foraging behavior and other out-of-nest activities: (1)
nocturnality, and (2) hypogaeic foraging. Both of these assertions are justified by biological
comparison of modern-day ants with contemporary wasps, among which these activities are
barely represented.
Nocturnality
The first pedestrian formicids, perhaps alone among their aculeate competitors, had
incidentally acquired the capacity to forage nocturnally – a far from trivial event in their
evolutionary success. This would have facilitated activity at times when diurnally active
potential competitors, predators and parasites were inactive. Predatory success would be greatly
enhanced, since nocturnally torpid resting diurnal insects would be relatively easily captured.
Nocturnal activity is frequently observed among modern epigaeic ants. In many habitats
foragers are far more common and diverse at night than in the day. Diurnal activity levels can be
high in many habitats, involving an array of species. Suites of other species are abroad at night,
with diversity and biomass almost always exceeding those of the day. Ants of many genera like
Nothomyrmecia, Myrmecia, numerous Ponerinae sens. lat., Myrmicinae like Meranoplus and
Podomyrma, and many reported dacetines, are nocturnal epigaeic foragers. Numerous
formicines like Myrmecorhynchus, Notostigma and many Polyrhachis and Camponotus species
are also nocturnal. Colony migration and foraging in the various taxa of army ants is an almost
entirely nocturnal activity. In these characteristics ants stand in strong exception to other
aculeates.
Nocturnal foraging is prominent in only two vespid wasp genera, the neotropical Apoica
(Polistinae - with 9 species) and the S. E. Asian Provespa (Vespinae - with three species)
(Schremmer, 1972; Matsuura, 1991). These are proportionately large-eyed compared to
relatives, and their nocturnality is crepuscular and otherwise dependent on moon light. They
appear to have no special adaptations for this life style, apart from extended visual acuity, and
explosive departure of foragers from nests at dusk in Apoica (Hunt, Jeanne and Keeping, 1995)
(a phenomenon also observed in Nothomyrmecia). Both genera are sister to diurnal relatives.
Nocturnality is a synapomorphy in each, so only two evolutionary events were required to

originate their unusual behavior.
Nocturnality is also exceptional among bees. Species of Caupolicana (Zikanapis)
(Colletidae) have enlarged ocelli and fly when the moon is out. Lasioglossum (Sphecodogastra)
texanum is crepuscular but requires flight on moonlight nights to obtain sufficient pollen for cell
provisioning (C.D. Michener, pers. comm.). Species of Megalopta (Halictidae: Augochlorini)
and Perdita (Xerophasma) (Andrenidae: Panurginae) are night-flying, and the former genus
includes a subgenus of nocturnal cleptoparasites, Megalopta (Noctoraptor) (M.S. Engel, pers.
comm.).
The repeated evolution of nesting behavior among the aculeate Apocrita required the
correlated development of precise navigational capacities. Female aculeates have the ability to
depart from and navigate back to their nests. Research demonstrates that their navigation is
almost exclusively visual, and apparently for that reason overwhelmingly diurnal. Chemical
navigation using scattered scent marks is present in some winged species as an unusual
exception not necessarily correlated with nocturnality (it is used for example in tribe Epiponini
(swarm founding Polistinae) during colony foundation – J.M. Carpenter, pers. comm.).
Nocturnal visual navigation by winged aculeates must be sufficiently difficult that it has
seldom evolved in flying, nest-building species. Some parasitic wasps fly at night, but they have
no requirement for precisely navigated return flights, and within appropriate habitats their flights
and resting stations are essentially random.
Visual epigaeic navigation derived from wasp ancestry is clearly the ancestral condition in
ants. Much has been written on the subject. The classic study of Wehner, Harkness & Schmid-
Hempel (1983) is a well-referenced source. Such navigation by pedestrian epigaeic ants at night,
using local landmarks over relatively short distances, is clearly more easily effected than its
equivalent in flying visual navigators. It is the modus operandi of Nothomyrmecia macrops, of
several observed Myrmecia species, and many other ants (Taylor, 1978; Hölldobler and Taylor,
1983).
Unpublished field experiments involving canopy manipulation or screening, and the
removal or duplication of local tree trunks, demonstrate that Nothomyrmecia navigates visually,
using the overhead tree canopy silhouetted against the night sky as a map. Similar canopy
navigation occurs in the African ponerine Paltothyreus tarsatus (Hölldobler, 1980). There is no
detectable evidence that Nothomyrmecia uses scent trails.
In conclusion it is clear that the aculeate night belongs to the ants, and that the correlated
evolution of pedestrian foraging and nocturnality greatly advanced evolutionary success in the
formicid lineage.
Hypogaeic foraging
Hypogaeic foraging involving the search for prey in the soil or leaf litter, in or under rotting
wood lying on the ground, or under stones, is rare among winged vespoid or sphecid wasps. It is,
however, the usual predatory mode of numerous ant species, some of which are so deeply
committed bionomically to life underground that they are seldom observed abroad, even on the
soil surface. Such hypogaeic elements are typified by most Amblyoponinae and many (but not
all) species of the other poneromorph subfamilies, the dorylomorphs, leptanillomorphs, and
some myrmicines. Their prey includes an array of soil-dwelling animals (mainly arthropods),
often including eggs or larvae. The distinction between nocturnal and diurnal activities seems
not to be significant in the most hypogaeic of these species.
Alternatively, several major ant lineages predominately include epigaeically foraging
species. These seek prey largely on vegetation or the ground surface, without penetration or
excavation of soil, litter or rotting wood. Many of them are nocturnal They include the
myrmeciomorph subfamilies Myrmeciinae and Pseudomyrmecinae, all species of which are
epigaeic, and the formicomorph subfamilies Dolichoderinae and Formicinae, most of which are
epigaeic.
The prey of vespoid and sphecid wasps rarely includes specialist soil-dwelling animals
(Krombein, 1967). These insects hunt epigaeically, usually on vegetation, or on the ground or
other exposed surfaces. Soil-dwelling arthropods, such as subterranean insects and their larvae,
or subsoil-inhabiting spiders and centipedes are seldom encountered, if at all.
These distinctions between ants and wasps appear to be related to the presence of wings in
foraging apterous aculeates, and their absence in foraging ants. It may reasonably be surmised
that penetration of the soil, litter or rotting wood, and the obstructed retrieval of immobilized
prey encountered there, would be maladaptively risky for winged foragers (especially regarding
potential damage to their wings), and that this explains the near-absence of such behavior in
winged aculeates.
Some groups of parasitic (as opposed to predatory) Apocrita include species with
permanently apterous, somewhat worker-ant-like females. Among aculeates these include
members of the chrysidoid families Dryinidae, Bethylidae and Chrysididae, and the vespoid
families Mutillidae and Typhiidae. Many of these females enter the soil or litter in search of
hosts (usually insect larvae), prior to oviposition. The hosts are not usually later withdrawn from
cover. The aptery of these species seems likely to be adaptively correlated with this hypogaeic
lifestyle. It has clearly evolved convergently and frequently. These exceptions support the
assertion that the presence of wings is not conducive to hypogaeic foraging.
The opportunities furnished by the evolution of apterous pedestrianism for the subsequent
acquisition of nocturnality and/or hypogaeic foraging activity were clearly variously exploited in
later formicid evolution. This is evidenced by the diversity among modern ants with respect to
these behaviors. It seems clear that there must have been early separation of lineages, some of
which were primarily epigaeic (and frequently nocturnal) in foraging behavior, and others which
were primarily hypogaeic predators. Myrmecia and Amblyopone today exemplify eventual
eusocial products of two such alternative groups. Haskins and Haskins (1951) pointed out that
these genera represent “widely divergent groups of ants” despite resemblance in many aspects of
their colony foundation and general social organization. The differences between them can
readily be explained by the present hypothesis that their distant Mesozoic ancestors responded
differently to the opportunities associated with pedestrianism - one lineage retaining wasp-like
epigaeic behavior, leading to Myrmecia (in which many species are also nocturnal) - the other
adopting a hypogaeic lifestyle, leading to Amblyopone. The similarities between them are
primarily derived from their distant common ancestry.
The petiole and elbowed antennae
A posteriorly constricted petiolar node comprising the first metasomal segment (2nd true
abdominal) is a synapomorphy of family Formicidae. The large remaining section of the
metasoma is termed the gaster. The 3rd abdominal segment often has its tergite and sternite fused
or firmly joined laterally. Its anterior part forms a ball-joint enclosed by the petiole (a
configuration serially analogous to the propodeal/petiolar joint in Apocrita, and with a similar set
of composite structures). The petiole is very narrowly constricted posteriorly in almost all
Formicinae and Dolichoderinae, but broadly articulated in some other ants, notably in subfamily
Amblyoponinae. The extra articulation clearly increases gastral rotation. Stinging ants probe
prey with the mandibles, apparently seeking chinks in the exoskeletal armor. When the jaws find
purchase the sting is swung far forwards under the body and applied near the mandibular tips.
The eggs of queens, and trophic eggs in workers, are removed from the abdominal tip in similar
manner. Venom-spraying formicines like Formica spray forwards with the gaster held under the
mesosoma (hence the ancient name “pismires”). The grooming behavior termed “abdominal tip
cleaning” in which the gastral apex is swung ventrally forwards to the mouthparts might,
incidentally, be an ant synapomorphy, apparently absent in other aculeates.
The antennae are elbowed in female ants with the scape notably elongate, often nearly as
long as the other articles together. This increases antennal mobility and reach, especially in
confined nest chambers. Scape length is functionally correlated with mandibular length, and is
considered an adaptation supporting antennation (olfactory scanning) of brood and potential
prey, and other workers, as in trophallaxis (Dlussky, 1975).
The non-eusocial, semisocial formicoids
This argument has now reached a stage where the pivotal formicid stock hypothecated here
has been prescribed. Its species are resident in colonies each comprising a group of coeval
dealate, fertilized females, variously and seasonally with their eggs, larvae and pupae, and also
seasonally with second-generation alate females and males. The winged virgin adults leave the
nests to mate and males afterwards disperse. Newly mated females typically seek potential nest
sites by flying and pedestrian search and join with other coeval females jointly to found further
colonies. Alternatively, they might enter established colonies (perhaps without first mating) to
supplement their personnel, contributing to the development of larger colonies than would
otherwise be possible. There would likely be strong adaptive support for this behavior. Dealation
in each female occurs after the mating flight, or upon entry to a new nest. It seems probable that
the mode of colony assembly would limit the prospects for development of very large colonies,
even considering that alternative strategies might be evolved.
Polygynous family groups like the above are common in social bees (Michener, 1974).
Twenty nine sphecoid and vespoid wasp species in which non-reproductive workers are lacking
and adult females share nests were listed by West-Eberhard (1978), who estimated the existence
of at least 39 such species.
These insects would initially have had wasp-like characteristics, but would become
increasingly ant-like as they diversified. Ocelli would initially be present in all individuals, and
elbowed antennae and a petiolate waist would be present in some species, though perhaps not
fully ant-like. Selection would probably favor increased adult longevity, perhaps to levels like
those known in living ants. The larvae would become progressively more ant like, some
probably with the crook-neck-squash shape seen in Myrmeciinae and Amblyoponinae. Foraging
in many species would be nocturnal, with dealate pedestrian foragers and visual navigation.
Comparative research on social wasps and bees demonstrates that reproductive competition
between the parental females in such communal nests would result in the evolution of more-or-
less morphologically undifferentiated reproductive and non-reproductive female castes, even (in
the early stages) if the non-reproductives had mated. Also, some virgin females might dealate
without mating to become workers. The resulting social arrangements (with 1-generational
matrifilial colonies, a worker caste and division of labor) are defined as “semisocial”. These
would be “caste-containing groups” in the sense of West-Eberhard (1978).
The asymmetries of relationship between the parental and daughter generations, related to
haplodiploid sex determination (which will not be explored here), would selectively favor the
presence in each colony of a single mated reproductive individual, the queen.
It will be argued below that this pre-eusocial stage would likely have been temporary in ant
phylogeny, but its significance would have been profound because, among other things, it would
have facilitated the evolution of efficient behavioral relationships and communication between
cooperating workers and between larvae and workers. Most importantly for this model, these
would include the communication of hunger by larvae to workers, in order to stimulate foraging
and prey-seeking behavior.
Considering its unique past acquisition of several highly adaptive major characteristics, this
lineage of “semisocial formicids” would have experienced great evolutionary success. It would
almost certainly have undergone adaptive radiation and taxonomic diversification, possibly with
the evolution of many species, genera and even higher taxa. There would be a diversity of
general lifestyles and degrees of crypsis in nesting and foraging, preferred nest sites, prey
preferences, mating flight parameters, modalities of reproductive behavior etc. The structure of
the mandibles in particular would have diversified in adaptive response to requirements for
dealing with different prey organisms in different groups, and some taxa would doubtless have
specialized on particular types of prey. Adult size would likely vary greatly between taxa.
Interspecific competition would be profound, as usual among related, similar species. This
would generate biological and ecological displacement and consequent adaptive shifts, which
would contribute to further diversification and radiation.
It is likely that some of the hypothecated semisocial formicids would have evolved fully
apterous females like those seen in other apocritan taxa (notably in Mutillidae, Tiphiidae and
Rhopalomastidae). Thus species with mesosomally-reduced workers and morphologically
undifferentiated reproductive females could have occurred. They would in some cases have
resembled workers of modern eusocial ants. This evolution could follow if the alate females
abandoned flight in mating, and instead attracted males using sexual calling pheromones. The
wings would thus lose adaptive function and natural selection would eliminate them. This would
probably be an all-or-nothing situation, with females either all standard (alate/dealate) or all
permanently apterous, depending on the species.
The evolution of high-level female polymorphism, with congenerational standard
reproductives and fully apterous ant-like workers (and appropriate caste-determining
physiological or sociobiological processes) would possibly have evolved in some taxa. The main
lineage, leading to eusociality must have retained winged females and allectomy, for that is the
synapomorphous condition in modern ants.
Untransformed non-eusocial formicids seem no longer to exist, but in evolutionary terms
they could well once have constituted a diverse, vigorous group.
Feeding, foraging and larval communication
Adult ants, like those of other aculeates, are liquid feeders. This is reinforced by use of the
labiomaxillary complex and infrabuccal pouch to remove large particulate inclusions from the
food, and by removal of fine particles from the digestive tract by the proventriculus, which
functions as a valve-dam. The proventriculus is variously and elaborately specialized for this
purpose among ants (Eisner, 1957) 1.
Ant larvae, however, are primitively entomophagus and feed on the solid masticated flesh of
prey (the many exceptions, including graminivory and liquid feeding by larvae, are derived
transformations of this character).
Thus, foraging adult ants are motivated by two behavioral drives: (1) to collect insect or
other prey for their larvae; (2) to locate and utilize liquid food for personal nourishment.
Unpublished field studies on Nothomyrmecia demonstrate that these drives are separately
satiated. Nocturnally-foraging workers delayed experimentally at honey baits early in their
progress imbibe the liquid avidly. They do not return to the nests when the drive for self-
nourishment is satiated, but proceed to hunt for insect prey. Their prey-getting drive is satiated
1
Eisner emphasised those taxa with the most derived proventricular structures (mainly genera of
subfamilies Formicinae, Dolichoderinae and Aneuretinae). There is need for further research among the
poneromorph subfamilies and those not yet studied. Amblyopone was shown to have a uniquely 6-partite
proventriculus while all other species studied were 4-partite, including Odontomachus, the only other
poneromorph reviewed. The two myrmicines and sole doryline examined had “degenerate” proventricular
structure, while that of Cerapachys (Cerapachyinae) was more complex.
when predation has succeeded, stimulating return to the nest by foragers carrying booty.
Otherwise that drive subsides at dawn when numbers of unsuccessful foragers return home.
The self-feeding drive is doubtless activated by worker hunger, but the prey-getting drive
might need to be triggered by larvae communicating their need for food to attendant workers,
which would be thus stimulated to leave the nest and to forage.
Such arrangements might be indicated by the fact that colonies of various ants are known
periodically briefly to cease foraging, later to resume. This is pronounced in the nocturnal
Nothomyrmecia macrops, where workers from previously active foraging colonies have been
observed not to depart their nests on nights when those from adjacent colonies were foraging
(and climatic conditions therefore not inimical), and to resume full activity the following night.
It is reasonable to suggest that larvae in these resting colonies were then well fed, were not
communicating hunger, and thus were not stimulating worker foraging behavior.
I have frequently maintained and studied colonies of Australian bulldog ants (Myrmecia) in
observation nests. On several occasions workers which would normally attack introduced live
insect prey have completely ceased such activity and ignored potential prey. This occurs
following pupation of the last larvae present in relevant nests. The enclosure of larvae in pupal
cocoons seems effectively to have removed larval influences, so the workers were no longer
motivated for prey-collection.
Authors have suggested that head waving and other body movements might be used by
larvae to communicate with workers. The demonstration by Cassill and Tschinkel (1995) of
olfactory communication of hunger by larvae to tending adults suggests a much more precise
mechanism. It raises the question that this communication might generally be pheromonal. More
research is desirable.
The evolution of formicid eusociality
The term “eusociality” refers to the level of social development attained by modern ants. It
requires the presence of overlapping generations, extreme division of reproductive labor, and
cooperative brood-care by non-reproductive adults. The workers of eusocial ants are not coeval
with their queens, but members of the following generation.
The queen (there is usually one per colony) is normally the mother of all other individuals
resident in a nest – all eggs, larvae, pupae and adults are her offspring - though males, which are
genetically haploid, may sometimes be derived from unfertilized, worker-laid eggs (Crozier and
Pamilo, 1996). Reproduction is the queen’s prerogative. She is the only member of the colony to
have previously mated. The sperm bank on which her reproduction depends is carried in her
spermatheca.
The workers are specialized brood-carers, and have in most cases lost the physical capacity
for reproduction. The ability to develop wings has been lost in their evolutionary derivation, and
the pterothoracic mesosomal sclerites specialized for flight have been deleted or fused, resulting
in drastically simplified mesosomal structure.
Eusociality has evolved homoplasiously several times elsewhere in the Aculeata. Classic
examples are wasps of subfamilies Stenogastrinae, Polistinae and Vespinae, members of the bee
subfamilies Meliponinae and Apinae and the remarkable sphecid Microstigmus comes. No other
eusocial species have fully apterous workers like those of ants.
Evolutionary transformation of the semisocial formicids posited above to fully eusocial ants
would require the daughter females to became non-reproductive workers. These second-
generation workers would eventually supplant the phylogenetically antecedent dealate queen-
coeval workers. Natural selection would act to favor individuals with diminished reproductive
organs and reduced alary and mesosomal complexity, ultimately to produce fully apterous non-
reproductive workers with simplified mesosomal structure. Physiological or sociobiological
mechanisms effecting caste determination would evolve in concert.
The first step to eusociality would require little more than the presence of unmated dealate
daughter workers in nests. If their supplementation of the worker force was adaptive, natural
selection would work to formalize the process. The other components of eusociality are already
latent in the lineage. Details, such as loss of the dealate queen-coeval worker caste and reduction
of worker mesosomal complexity would later be effected by natural selection, to produce all the
defining characteristics of modern formicid eusociality.
The hypothecated semisocial formicids can be considered neatly predisposed for evolution
of the daughter worker caste. The necessary first step is elegantly simple. It requires premature
dealation. There are at least two models by which this could occur:
(1) The larval manipulation hypothesis:
All females in semisocial formicid societies, including alate daughters, would inevitably be
programmed by evolution with the instincts (1) eventually to perform dealation and (2) to
respond to stimulation from hungry larvae by foraging. All that is required is for some daughter
alates to respond prematurely to the imperatives of larval stimulation by foraging, and to lose
their wings by premature dealation without having mated.
This process can be envisaged as larval manipulation because it would be in the interests of
larvae to increase the force of workers dedicated to their care.
(2) The parental manipulation hypothesis of Alexander (1974):
In this case the reproductive female(s) would impose the worker role on alate daughters by
forcibly removing their wings, as happens (in effect) in contemporary Diacamma species (see
below).
In either case, because mating requires flight, dealation of the daughter workers would
preclude them from future reproduction. The asymmetries of relationship to their mothers and
sisters, occasioned by haplodiploid sex determination (Crozier and Pamilo, 1996), would favor
their assumption of the worker role, to assist in the production of reproductive siblings from
eggs laid by their fertilized mothers. In this way daughter workers would appear almost
instantaneously. Assuming that the arrangement was competitively successful natural selection
would rapidly improve the functionality of this newly acquired eusociality.
These scenarios are presented as models for the evolution of formicid eusociality. Dealation
is a systematic component, and because of its presence the evolution of eusociality in ants has
followed an entirely different course from those hypothecated to have produced eusociality
elsewhere among winged eusocial aculeates.
The initial step is so simple that it could be considered an almost inevitable prospect (though
certainly not in any way pre-ordained) for formicids with the hypothecated semisocial lifestyle
suggested above. Either scenario could easily have occurred more than once in the formicid
clade.
Semisocial formicids with standard females and mesosomally-reduced workers, as
formulated above, could similarly evolve eusociality. They would have the advantage of pre-
existing caste determination mechanisms. Those with monomorphic, fully apterous females
would be unable to initiate eusociality, at least not along the lines suggested here.
Did the ant worker caste evolve more than once?
One of the “givens” of myrmecology is that evolution of the worker caste and resultant
eusociality was unique in ant phylogeny. This proposition is rarely mentioned, far less
questioned (though see Wilson, 1971: 74, for a suggested separate origin of amblyoponine
sociality). Its pervasiveness is evident. Consider that every published formicid “family tree” is
rooted in assumptions that (1) the ancestor of all ants was itself eusocial, with an apterous
daughter worker caste, and (2) all of the diversity of modern ants is derived from that single
ancestor, and (3) that its appearance predated the complex evolutionary branching which
established the major taxa of ants.
The possibility that the worker caste is not a synapomorphy of family Formicidae, but that
workers and eusociality might have been homoplasiously derived more than once from a group
of non-eusocial ancestral formicids, seems never to have challenged the entrenched position.
I propose that the worker caste in ants might not be uniquely evolved, but homoplasious.
This hypothesis has the following components: (1) that the daughter worker caste and eusociality
arose more than once from an ultimately monophyletic but diverse group of pre-eusocial,
semisocial, formicids of the kind hypothecated above; (2) that multiple derivation of eusocial
workers among the semisocial formicids would have been unexceptional, given the simplicity of
such evolution along the lines suggested in this essay; (3) that some of the major features of
modern ant diversity were already present among the antecedent non-eusocial formicids and
survive today to define some of the major lineages of family Formicidae. Each relevant lineage,
according to this model, originated from a taxon within the ancestral pre-eusocial group which
had acquired a daughter worker caste and eusociality separately from other such taxa; (4) That
the hypothecated non-eusocial formicids, although argued here likely to have been a diverse and
successful group, have become extinct as failed competitors of eusocial ants; (5) that the process
of worker mesosomal simplification proceeded separately with each acquisition of eusociality
and that these separate origins have comprehensively produced deceptively similar worker
mesosomal structures across the diversity of modern ants.
It is arguable that myrmecologists have been consistently deceived by the convergences in
worker mesosomal structure. As long as the observed reductions were accepted as being
homologous the worker caste and eusociality were accepted to be monophyletic. In fact, of
course, there is much diversity of mesosomal structure among worker ants.
This hypothesis is posited as a challenge to others to consider, and perhaps to test its
propositions and implications. Some further comments are in order.
It implies that the latest common ancestor of all extant ants was a semisocial wasp-like
formicid. It does not challenge the family Formicidae as a natural clade. If accepted it would
have little impact on the taxonomic classification of ants, except that some current taxa might be
raised in rank if their eusociality is considered to be independently derived. Bolton’s elevation of
the former “ponerine” tribes Amblyoponini and Ectatommini to subfamilies (the Ectatommini to
four: Ectatomminae, Heteroponerinae, Paraponerinae and Proceratiinae) might already have
partly accomplished this. Bolton’s formicomorph, myrmeciomorph, and leptanillomorph
subfamily groups and the subfamily Ponerinae could be separately derived lineages, and the
myrmicomorphs might be derived severally from stock shared with the poneromorphs.
Many ants have a second waist node, termed the postpetiole, formed by posterior
constriction of the 3rd true abdominal segment, and the related formation of an articular
exoskeletal ball-joint anteriorly on the 4th segment. This arrangement seems likely to have been
derived more than once.
Greater understanding of the processes of mesosomal reduction and of evolution of the
postpetiole is needed to identify cladistically useful characters. Is tubulation or some similar
modification of the 4th true abdominal segment a necessary precursor to the evolution of a
postpetiolar node and its articular condyle? 2 If so, is this configuration uniquely evolved, and
2
The term tubulation (in the sense of Taylor, 1978) refers to the configuration of the 4th true
abdominal segment. It is “tubulate” when the lateral margins of its tergite and sternite are aligned and
more-or-less fused to assemble a strong cylindrical structure. This is partly inserted into the exoskeleton of
the preceding segment 3 to furnish a rotatable joint. The inserted section is set off from the remainder of the
segmental exoskeleton by a girdling constriction. When segment 3 constitutes a postpetiolar node the
anterior section of segment 4 can be very small, forming a ball-joint articulating the large terminal
abdominal tagma (termed the “gaster”, but here 1 segment shorter than the formicomorph gaster). This
arrangement is an analog of the constricted preceding propodeal/petiolar and petiolar/postpetiolar joints.
what does the answer to that question say about affinities among the various postpetiolate
subfamilies of ants? Brown (1954) recognized separate “poneroid” and “myrmeciioid” lineages.
If his treatment of the myrmeciomorph subfamilies (sensu Bolton), including the Myrmeciinae
and Pseudomyrmecinae, is set aside, Brown’s hypothesis basically separates the abdominally
tubulate (and frequently postpetiolate) ants from the remainder (i.e. the Aneuretinae,
Dolichoderinae and Formicinae). In effect it posits two clades within the Formicidae. Ward and
Brady (2003) rigorously estimated that the subfamily Myrmeciinae diverged from other ant
lineages about 100mya. That lineage might have been established as a third major clade before it
evolved eusociality. This would imply separate evolution of postpetiolar structures, as seems
likely on anatomical and phylogenetic grounds (P.S. Ward, pers. comm.).
Some modern taxa seemingly could have evolved only from eusocial stock. The
dorylomorph army ant subfamilies (Ecitoninae, Aenictinae, Dorylinae and Leptanilloidinae) are
pertinent. Models for their derivation, culminating with the work of Brady (2003), have
consistently implied descent from unexceptional eusocial non-migratory primitive poneromorphs
(see Wilson, 1958). Brady’s analysis has them sharing ancestry with the Cerapachyinae (which
were also placed as dorylomorphs by Bolton, 2003).
The Diacamma enigma
The ponerine genus Diacamma has social organization strikingly different from those of all
other ants. It could be an informative exception, and must be considered in any broad discussion
of ant eusociality.
Diacamma is Indo-Australian, with perhaps 30 species. The bionomics of an Okinawan
species (the sole Japanese taxon, identified then as D. rugosum) were reported by Fukumoto,
Abe & Taki (1989). A second species (D. australe, Queensland, Australia), with essentially
identical biology, was reviewed by Peeters & Higashi (1989). The following account is based on
these reports and other research detailed below. The genus typically forms small colonies in
surface soil and leaf litter. Its females are uniformly apterous, and there is no general
morphological differentiation between the sole parental reproductive in each nest and associated
workers.
All Diacamma females at eclosion carry a pair of tiny bilateral lobate appendages housed in
small dorsolateral mesonotal pits, which are considered to be vestigial wings (Tulloch, 1934).
Their presence determines reproductive status. Females with appendages are able to function as
reproductives, to mate and produce diploid offspring. Those without appendages function as
non-reproductive workers. Reproductive status (i.e. caste) is controlled by the parental female
removing these appendages from prospective workers as they emerge from their pupal cocoons.
In this way vigorous resident reproductives are able totally to prevent their daughters from
becoming reproductives.
Colonies reproduce by fission. A group including the parental reproductive and workers
departs to establish a new nest. Cocoons containing pupae remain with workers in the original
nest. The females eclosing from these compete among themselves to retain their vestigial wings
which others attempt to remove. Eventually one (usually the first to emerge) remains intact – she
becomes the prospective new reproductive. She leaves the nest to mate nearby after sexually
calling a male partner. Following return she begins to lay fertilized eggs. Reproductives alone
retain wing vestiges in the nests (i.e. there is no equivalent of dealation after mating). The cycle
continues.
Note that the new colony is initially semisocial 3, with the reproductive and workers all
daughters of the same female. These ants are clearly eusocial, though with a distinctive caste-
3
This is the case also in all but the first departure in a series of honey bee swarms and presumably in
other swarm-founding aculeates, including ants with apterous reproductives and colony budding.
determination system, and a major semisocial phase in their colony cycle. If the species was
univoltine the semisocial phase could last for almost a year until the first offspring of the new
queen emerged.
Both the Fukumoto and Peeters groups demonstrated that reproductives by their presence
are able to inhibit the production of haploid (male) eggs by workers (a not unusual trait in queen
ants), and both considered their subject species to be “queenless”. Peeters and Higashi identified
the reproductive females as gamergates (mated reproductive workers).
The mesonotal appendages were termed “gemmae” by Peeters & Billen (1991) and shown
to be highly glandular by Billen & Peeters (1991), who presumed this to be related to “the
peculiar mechanism of reproductive regulation found in this queenless ant”. Mesosomal anatomy
of D. australe was discussed and illustrated by Bitsch & Peeters (1992) who confirmed that the
mesonotal appendages were vestigial mesothoracic wings. Gronenberg & Peeters (1993),
concluded that workers lacking gemmae can never replace a gamergate. Peeters, Billen &
Hölldobler (1992) discovered that the gemmae were not removed in workers of the Indian D.
vagans, and that mated top-ranked individuals dominated the nests and functioned as
reproductives. Similar reproductive dominance occurs in orphaned groups of the Japanese
species, with an alpha dominant worker monopolizing haploid egg production (Peeters & Tsuji,
1993). Sommer, Hölldobler & Rembold (1993) described a Malaysian species with bionomics
similar to the Japanese model. Following comparison of four Indian species, Veuille et al.
(1998) demonstrated that retention of gemmae is the derived condition versus gemma removal in
Diacamma. Recent literature, too extensive for citation here, has shown that other Diacamma
species have worker castes determined by the removal of gemmae.
There are two possible origins for this remarkable behavior. Either it is primary, derived
directly from a non-eusocial ancestor, or it is secondary, derived from a eusocial ancestor. Two
possible secondary models are:
(1) That Diacamma was derived from a conventionally eusocial ancestor through
acquisition of gemmae and associated mesosomal structures by its workers, followed by
replacement of the queen caste with gamergates. This seems improbable because gemmae, as
wing vestiges, would need to be developed on an already structurally-reduced mesosoma, and it
is not easy to understand why this would happen in eusocial ants with a pre-existing caste-
determining mechanism.
A shift in gene expression could theoretically reassign “queen” characters to the workers,
producing partial resurrection of an unreduced mesosoma and wing vestiges in a lineage
formerly with mesosomally-reduced workers. Apart from the possibility that ocelli were restored
in this way to the workers of Harpegnathos (which are uniquely ocellate among ponerine
workers) there seem to be no other examples of this phenomenon in poneromorph ants.
(2) That a secondary round of worker-caste evolution occurred from conventional eusocial
ancestry, starting with groups of coeval alate/dealate females, the descendents of which
eventually enacted a scenario like that outlined below. Potential startup groups could perhaps
originate from assemblies of pleometrotic colony foundresses.
Neither alternative seems very plausible and both models require the extinction of
conventionally eusocial ancestral stock.
One hesitates to suggest the primary option, but, consider the following evolutionary
scenario: (1) A monomorphic alate/dealate semisocial formicid has small colonies comprising
congenerational dealate workers and a single dealate reproductive female. They rear alate
females and males from eggs laid by the reproductive female. (2) The reproductive female co-
opts some of her alate daughters as workers by forcibly removing their wings. Unmutilated
daughters are able to fly and mate. They are the future reproductives. (3) The daughter (eusocial)
workers adaptively supercede the queen-coeval (semisocial) workers, and the evolving lineage
attains eusociality. Worker determination continues to be controlled by removal of wings from
virgin alates by the queen. (4) A descendent lineage abandons the mating flight and reproductive
females use pedestrian sexual calling to attract males. Wings are no longer adaptive for flight
and the lineage moves towards monomorphic aptery, undergoing evolutionary reduction in wing
size and simplification of mesosomal structure. (5) The wings are eventually not entirely lost,
but tiny lobate vestiges of the forewings remain, housed in small dorsolateral mesonotal pits.
Caste is now determined by the parental reproductive female removing these vestigial wings
from prospective workers etc.
Of these alternatives, the first secondary model seems the least implausible. The second
might satisfy those who believe that Diacamma females are phylogenetically degenerate queens.
The primary model would gain some support if Diacamma stood taxonomically apart from other
ants, but all relevant cladistic and other analyses place the genus firmly in the subfamily
Ponerinae as presently constituted (though this remains a probably artificial group likely
destined for more dismemberment such as that by Bolton, 2003). The primary model would not
be implausible if the taxonomic difficulties could be resolved. If accepted it would firmly
demonstrate that modern ant eusociality evolved at least twice from non-eusocial stock,
falsifying the common assumption that it is uniquely derived.
Pedestrianism in eusocial ants
The evolution of worker aptery and pedestrianism expedited both nocturnality and
hypogaeic foraging among the semisocial formicids posited here, contributing to their adaptive
success. Pedestrianism remains influential among formicids. One of its major advantages is that
much larger, heavier, prey can be carried or dragged by individuals than is possible in flight, and
foragers can readily cooperate in the transport of large prey.
The evolution of worker subcastes, including large-headed majors or “soldiers” would
otherwise be nearly impossible. They are unknown among winged social aculeates, except for
Australian halictine bees of genus Lasioglossum, which have a caste of large-headed, small-
winged males which stay on the nests, fight for possession of females and repel parasites
(Houston, 1970). The aerodynamic constraints of macrocephaly seem likely to have constrained
its development in winged aculeates. Ants, being pedestrian, suffer no such constraint, and
macrocephaly clearly does not inadaptively inhibit progression by large-headed soldiers, which
have evolved repeatedly and are present in several subfamilies.
The absence of wings has allowed ants to nest in a diversity of situations barely exploited by
flying aculeates.
Permanent substrate contact has occasioned repeated evolution of scent trail-laying
techniques important in foraging, nest migration and worker recruitment to food. These utilize a
range of chemicals and specialized glands unknown among other aculeates (Hölldobler, 1978,
and subsequent papers). Trail pheromones are produced by a rich diversity of glands, and their
taxonomic distribution could provide characters useful for phylogenetic analysis.
What happened to the ant ancestors?
The adage “extinction is the common lot, survival the exception” applies to the history
hypothecated above. Extinction of taxa related to the crown group is a necessary component at
each step in the phylogenetic assembly of modern ants, establishing the status of characters now
accepted as formicid synapomorphies.
The pre-eusocial formicids seem unlikely to have been very effective competitors of
eusocial ants. Their lifestyle would likely have been less efficient than eusociality. Colony size is
probably inherently limited, and potentials for behavioral and biological diversification more
restricted than in true ants.
Various ants are known to be highly effective biological competitors of other ants and to
locally eliminate other ant species. Consider the success of “tramp” species introduced by human
commerce to new localities. They regularly eliminate or restrict resident endemic ants or other
introduced species from their habitats. Their overwhelming competitive abilities involve not
only ecological success, but direct, targeted violent aggression (Holway et al., 2002). The
vagaries of the Argentine ant, Linepithema humilis and the Madeira ant Pheidole megacephala
in Bermuda (Haskins & Haskins, 1965, 1988), and the neotropical Little fire ant Wasmannia
auropunctata in New Caledonia (Fabres & Brown, 1978), Cameroon and the Galapagos Islands
are prime examples.
It is accepted here that the hypothecated non-eusocial formicids could have been eradicated
by competitive assault from true ants. Similar apocalyptic extinction has not occurred in the
other eusocial aculeates. Wasps (with some exceptions like Vespa) seem less prone than ants to
assault their relatives, and flying adult wasps can more readily escape attack than pedestrian
formicids. Even if their brood was lost they could survive, possibly to reestablish their nests.
Bees have moved away from larval entomophagy. As pollen collecting herbivores they are not
given to attacking other bees. I suggest that for these reasons a diversity of social life-styles
ranging from solitary to eusocial has survived among the wasps and bees, but not among ants.
The timing of events
The earliest phylogenetic events posited above would probably have occurred in rapid
evolutionary succession following acquisition of acellular nesting and metapleural glands. It is
now accepted from fossil and other evidence that eusocial ants first appeared in the Mesozoic
perhaps 140-100 million years ago (mya). Their rise by the late Cretaceous is evidenced by
several recently described fossils, which demonstrate that eusocial ants of modern appearance,
with close modern taxonomic affinities, were present in those times.
The fossil Kryomyrma neffi Grimaldi & Agosti (2000a) from late Cretaceous (Turonian)
New Jersey amber (estimated 92 mya) is an indubitable formicine worker, with detailed
resemblance to modern taxa.
The genus Afropone Dlussky, Brothers & Rasnitsyn (2004) with two species represented by
compression fossils of late Cretaceous age (Turonian, 93.5-89.3 mya) from Orapa, Botswana
was assigned by its authors to subfamily “Ponerinae”. Its metasomal structure, with a
differentiated postpetiole, is derived when compared to the usual ponerine configuration, and
resembles that of the modern myrmeciomorph subfamilies Myrmeciinae and
Pseudomyrmecinae, to which Afropone could be annectant. Its large eyes support this
assignment. 4
Brownimecia clavata Grimaldi, Agosti & Carpenter (1997) (White Oaks New Jersey amber;
Turonian, est. 94-90 mya) was considered by its authors to be a “ponerine”. The term
“poneromorph” is appropriate, since the former subfamily Ponerinae now comprises 6 modern
poneromorph subfamilies. No assessment is given as to whether the 4th abdominal segment is
tubulate or not. Such indication is required if fossils are claimed to be poneromorph.
Constriction of the 4th abdominal segment alone is insufficient for confident identification – the
defining character is the presence of its tubular structure. That said, the published Brownimecia
illustrations do evidence tubulation. Bolton (2003) placed Brownimecia in a monotypic
subfamily Brownimeciinae, and considered that “It may belong, speculatively, in a basal group
that is sister to the poneromorphs, or perhaps to the poneromorphs and leptanillomorphs
4
In modern poneromorphs the postpetiole is not usually strongly constricted posteriorly to form a
distinct node, but poneromorphs are engineered for postpetiolar development because of their tubulate 4th
abdominal exoskeleton and its insertion into the 3rd segment. A few poneromorphs are quite strongly
postpetiolate, including Paraponera (Paraponerinae) and some species of Cerapachys (Cerapachyinae) and
Proceratium (Proceratiinae).
together”. Both of these subfamily groups have tubulate abdomens, and many species are
postpetiolate.
Canopone dentata Dlussky (1999) (Medicine Hat, Alberta; Campanian, late Cretaceous, est.
83.5 – 70.6 mya) has a clearly tubulate abdomen and somewhat constricted postpetiole. Dlussky
was unsure of its placement as a “ponerine” or myrmeciine. I consider it to be poneromorph,
with some similarities to Paraponera. The presence of worker ocelli is unusual in
poneromorphs, but that character is present in living Harpegnathos.
A posteriorly constricted postpetiolar node is present in Afromyrma Dlussky, Brothers &
Rasnitsyn (2004) (Orapa, Botswana; Turonian, est. 94-90 mya). Its compression fossil is poorly
preserved but seems certainly to be myrmicomorph. Its was originally assigned to subfamily
Myrmicinae.
Eotapinoma macalpinei Dlussky (1999) from Canadian amber (Medicine Hat, Alberta;
Campanian, late Cretaceous, est. 83.5 – 70.6 mya) was considered a likely dolichoderine by its
author and a “putative dolichoderine” by Grimaldi & Agosti (2000b).
Gerontoformica cretacica Nel, Perrault, Perrichot & Neraudeau (2004) (Archingeay-les-
Nouillers, Charente-Maritime, SW France; Uppermost Albian amber, est. 100 mya) is not well
preserved, but the published photographs of its petiole and gaster closely recall Nothomyrmecia.
Nothing in the original paper challenges this resemblance. Gerontoformica is described as
having peg-like teeth on its clypeus (reminiscent of Amblyoponinae and Anomalomyrmini
(Leptanillinae)), small eyes and long mandibles. Its authors remarked that its first gastral (i.e. 3rd
true abdominal) segment had a “pretergite”, though their fig 2c seems to illustrate this character
for the 4th abdominal segment. They did not indicate whether abdominal segment 4 was tubulate
or not, but their photographs imply the latter. The fossil was said to lack a sting, but Fig. 2a,
seems to depict an unexserted sting shaft. Fig. 1c does not represent the head in detail sufficient
to illustrate the textural description of cephalic, mandibular and antennal particulars. The
discovery of more specimens from Archingeay is eagerly awaited. The authors are urged to
publish line illustrations backing-up their observations and photographs.
Using Bayesian dating analysis Ward & Brady (2003) indicate that the most recent common
ancestor of a clade containing Nothomyrmecia and Myrmecia is 74 my old (95% confidence
limits 53-101my).
Derivation of the several living dorylomorph army ant subfamilies from ancestral stock
considered antecedent also to the Cerapachyinae has recently been rigorously estimated to have
occurred in mid Cretaceous times (i.e. around 100mya), possibly in Gondwana (Brady, 2003),
providing further evidence that true ants of modern appearance and taxonomic affinity, with
behavior like that of highly specialized modern ants, were likely present by the late Cretaceous
These examples all imply that evolution from solitary wasp ancestry to ant eusociality could
have been complete within the first few million years of formicid history, with tens of millions
of years available to play out the details, including the extinction of non-eusocial formicids,
which probably disappeared promptly anyway following the rise of eusocial ant competitors. It
also seems likely that the earliest events in formicid evolution would have occurred within a
limited geographical range, perhaps entirely in Gondwana or one of its drifting continental
fragments, so the loss of groups standing as sister taxa to those of the emerging main formicid
lineage would have been local geographical events.
It appears that mid-Cretaceous aculeates would have included a cluster of related lineages
among which the prospective formicid synapomorphies first evolved and began successively to
accumulate. At the point of each character acquisition two sister taxa would be generated, one
lacking the new character, but possessing those previously evolved. There would probably at
times have been several contemporaneous taxa carrying incomplete sets of the relevant
characters. Also, some of the characters could have arisen more than once, later to be secured as
potential formicid synapomorphies by the surviving lineage. One lineage alone, according to this
model, followed the course prescribed here and that lineage is represented today only by the
living ants – the remaining taxa significant to this history have in the meantime become extinct.
It also appears prima facie that the fossil record has not yet yielded possible pre-eusocial ant
ancestors with incomplete sets of the relevant characters reviewed above. The hypothecated first
solitary protoformicid would be a strong candidate for recognition as a fossil. It would probably
have the general facies of a modern solitary vespid wasp, with metapleural glands or antecedent
structures, but probably not a petiolar waist node or elbowed antennae (though many modern
solitary vespids have somewhat elongated scapes). A fossilized dealate, queen-coeval forager
from an ancestral semisocial formicid colony of the type posited here would probably have a
more generally (perhaps substantially) ant-like facies but with some wasp-like attributes. It
would very likely be identified as a somewhat wasp-like dealate female eusocial ant, unless its
fossilization preceded acquisition of the petiolar node and/or elbowed antennae, or it had those
characters incompletely developed compared to modern ants. The extinct subfamily Armaniinae
Dlussky (reviewed by Bolton, 2003: 73, 259) requires consideration in these regards (P.S. Ward,
pers. comm.).
What is Sphecomyrma?
The extinct genus Sphecomyrma (type species S. freyi Wilson, Carpenter and Brown, 1967)
was based on two worker specimens enclosed in a piece of Turonian New Jersey amber of
estimated age 92 mya. Its characteristics are summarized in the original paper, and by Grimaldi,
Agosti & Carpenter (1997). They will not be reiterated here.
Further specimens from New Jersey are now known (of Turonian age, mid Cretaceous, ca
92 mya. – Grimaldi, Agosti & Carpenter, 1997). A second closely similar worker-based species
S. canadensis Wilson has been described from Alberta, Canada (78-79 mya), and a worker-based
related genus Cretomyrma Dlussky from the Taymyr peninsula, Siberia (of Santonian age, 85.5-
83.5 mya). Two male-based genera Dluskyidris Bolton and Baikuidris Dlussky (both from
Siberian Santonian amber) are believed to be relatives of Sphecomyrma. All are currently
classified in family Formicidae, subfamily Sphecomyrminae. There have been no reports of
fossil alate or dealate females.
Sphecomyrma has also been reported from late Albian Burmese amber (ca 100-105 mya)
(Grimaldi, Engel & Nascimbene, 2002), along with an enigmatic poneroid fossil which
represents the oldest known definitive ant. A further new species has recently been discovered in
New Jersey amber (Engel & Grimaldi, in prep.). These authors will report multiple workers of S.
freyi from a single piece of New Jersey amber, implying that the species was social. Engel (pers
com.) “would argue that Sphecomyrma was a likely social (perhaps even eusocial) early group of
ants”, a view with which I concur.
There has been much discussion concerning the taxonomic and phylogenetic status of
Sphecomyrma. It has been considered a taxon sister to the Formicidae, with the two families
comprising superfamily Formicoidea (Wilson, Carpenter & Brown,1967), assigned with the
other taxa indicated above to subfamily Sphecomyrminae in family Formicidae (Grimaldi,
Agosti & Carpenter, 1977; Bolton, 2003), or alleged perhaps not to be an ant at all, and to
require a separate family (Dlussky, 1975, and his other references in Grimaldi, Agosti &
Carpenter 1997). Baroni Urbani in 1988 considered it to be a nonsocial wasp (reported in
Grimaldi & Agosti, 2000b). In the morphological phylogenetic analysis of Ward & Brady (2003)
Sphecomyrma is sister to the extant ants sampled, with strong bootstrap support.
The argument centers on whether or not the relevant fossils possess metapleural glands
(which were identified with reservation in the original Wilson, Carpenter & Brown paper).
Recent researchers using recently collected specimens agree that metapleural glands are present,
and the affirmative evidence of Grimaldi, Agosti & Carpenter (1997) is convincing. It has also
been suggested that the antennae, which have shorter scapes than in all living female ants, could
indicate that Sphecomyrma might be neither eusocial nor an ant (see Grimaldi, Agosti &
Carpenter (1997) for summary). The last hypothesis is probably unfalsifiable.
Wilson (1971: 31) considered the wasp-like mandibles of Sphecomyrma an anomaly in
comparison with the very different mandibles of Amblyopone, which was presented as an
archetypical primitive ant. This difficulty is resolved by the model for ant evolution presented
here. It envisages a range of mandibular specializations among the hypothecated semisocial
formicids, easily accommodating both Sphecomyrma and Amblyopone. Note that broadly “wasp-
like” mandibles are present in virtually all male ants, despite the great range of mandibular
specializations seen among ant workers.
At the risk of introducing a further probably unfalsifiable hypothesis, I represent that the
various Sphecomyma fossils could be females or workers of a semisocial formicid, like those
hypothecated here.
It was suggested above that these insects would likely have retained strong wasp-like
characteristics along with emerging resemblances to modern ants, that ocelli would be present,
and that the antennae and waist nodes in some species might have transitional rather than fully
ant-like structure. Also that some species might either (1) like mutillids, have evolved
monomorphic wingless females with mesosomal reductions homoplasiously resembling those of
modern worker ants, or (2) have evolved a eusocial-ant-like apterous worker caste while
retaining standard reproductives (the two castes being congenerational within nests). All these
possibilities are plausible, and all could apply to Sphecomyrma. It could represent a species
meeting either of the two options numbered above.
Those interested in this proposition should consider it while re-reading the classic Wilson,
Carpenter & Brown paper. Its authors believed Sphecomyrma to be “truly intermediate between
primitive ants and the aculeate wasps” and stated that “if we are mistaken about the presence of
the metapleural gland, it would be possible to consider Sphecomyrma a wasp rather than an ant”.
In other words they considered the metapleural glands to be essentially the only definitively
formicid attribute of S. freyi.
Such observations regarding wasp/ant intermediacy would also apply exactly to the
semisocial formicids hypothecated here.
The “true” nature of Sphecomyrma might in fact be intractable because either: (1) the
discovery of alate or dealate female Sphecomyrma fossils would not falsify the above
hypothesis, since they could be related semisocial formicids of another species which had not
lost flight capacity. Neither would they prove that the Sphecomyrma fossils were eusocial
workers; or (2) alate or dealate fossils with “complete” mesosomal structure, collected in
indubitable conspecific association with mesosomally-reduced specimens like the known
Sphecomyrma fossils, could represent either a semisocial species with congenerational
alate/dealate females and apterous workers, or a fully eusocial ant with apterous mesosomally-
reduced daughter workers. They would indicate at best only that Sphecomyrma was either a
semisocial or a eusocial formicid.
The mesosoma in Sphecomyrma is unusual compared to those of all modern ants, but the
suggestion by Wilson, Carpenter and Brown (1967) that the suture between the meso- and
metathorax is “possibly movable” seems implausible.
Concluding remarks
Past discussions on the origins and evolution of formicid eusociality have reiterated the
Wheeler/Malyshev hypothesis (Wheeler, 1928: 63; Malyshev, 1968). It was reviewed without
significant support by Wilson (1971) and Hölldobler & Wilson (1990), and will not be further
considered here. Wilson’s suggestion (1971) that amblyoponine ants might have evolved in the
Wheeler/Malyshev manner via Scleroderma-like ancestry is untenable (Hölldobler & Wilson,
1990).
Various authors, notably Wilson (1971: 72), Hölldobler & Wilson (1990: 27) and Bourke &
Franks (1995: 73) have espoused the “subsocial route” for evolution of ant eusociality, largely
inspired by the seminal analysis of wasp sociality by Evans (1958). This model derives formicid
eusociality ultimately from groups comprising a parental female caring for her nondispersing
daughters. The model implies that such ancestral females would have initiated their colonies
alone, not in pleometrotic foundress groups. It is argued that this is the usual mode of
development of modern ant colonies, which is held in effect to reflect phylogeny and to support
the model. It is also held that semisocial groups of the kind posited in this essay would lack
kinship bonds sufficient for the semisocial model to operate under natural selection. Bourke and
Franks considered the semisocial option “a poor model for the origin of altruism in early ants”.
The first assumption is simplistic. Pleometrotic colony foundation by partially claustral
groups of coeval mated females could just as well be homologous with the ancestral condition,
as proposed above. Pleometrosis is known largely among “primitive” ants (Wheeler, 1933,
Wilson, 1971, Hölldobler & Taylor, 1983), and its general absence in “higher” ants, versus
solitary colony foundation, could reflect a phylogenetic succession.
The ability of queens to found colonies alone without foraging is another feature of ants
based on the presence of allectomy, which provided the opportunity for metabolization of the
redundant post-dealation flight muscles in order to sustain claustral, solitary foundress females
and their initial brood. The larvae in such founding colonies are fed by the queen with surplus
eggs, or trophic egg material, or nutrient provided by trophallaxis (processes broadly analogous
with milk feeding in mammals, by which female biomass is converted to larval biomass).
Solitary, claustral colony foundation is in fact commonly practiced mainly among the
formicomorph and myrmicomorph subfamilies. Their queens are usually much larger than
conspecific workers. The myrmeciomorphs, many poneromorphs and several known
myrmicines, are partially claustral, frequently with pleometrosis (see Wilson, 1971: 72). The
dorylomorphs and leptanillomorphs (possibly excluding the barely studied Anomalomyrmini
and a few cerapachyines) and some other “army ants”, like Onychomyrmex (Amblyoponinae),
have apterous queens and found new colonies by budding, as do those Ectatomminae,
Heteroponerinae and Ponerinae with apterous reproductives or gamergates. Most of these,
except the dorylomorph and leptanillomorph army ants, have relatively small queens, little larger
than workers. Solitary foundation is thus broadly correlated with the secondary presence of
strong size dimorphism between flying queens and their workers, where the ratio of available
bulk of maternal mesosomal flight musculature to biomass of the foundation brood is high. It
cannot be represented as the likely situation in the earliest formicids.
The initial groups of winged communal wasps posited in the current model would draw
their personnel from local populations, so would likely have significant relationship ties. Natural
selection would work upon such groups whatever their composition, and if individuals joining
positively selected groups were more likely than non-joiners to reproduce, their group-joining
behavior would be positively selected. Note that, in the earliest stages of group formation
discussed above, all joiners would have the opportunity to contribute eggs to the collective
cache, and their average chances for survival and reproduction could be better than if they had
not joined. Even if some individuals lost out in the reproductive stakes the group could still be
relatively adaptive, and the tendency by mated females to join groups could become selectively
established. Furthermore, if those groups with the most closely related personnel tended to be the
best survivors, selection would favor individuals which grouped with close relatives such as
sisters. This process could furnish the allegedly deficient kinship bonds.
Wheeler (1928: 107) considered that “The Ponerinae represent the primitive stock from
which the other ant subfamilies…have radiated”. He asked “Are the various subfamilies of ants
really derived from a common stock or are they polyphyletic like the various social subfamilies
of wasps and bees?” Careful reading indicates that he was talking about the “various subfamilies
of ants” as possible separate social lineages. He answered (p. 108) “I deem it…probable...that
the family (i.e. the Formicidae as a social lineage) as a whole is monophyletic”.
In reaching that conclusion Wheeler specifically rejected a phylogenetic scheme
diagrammatically proposed by Emery (1920: 370; the diagram reproduced by Wheeler, 1928:
107). This posited several lines of evolution from an “unknown hypothetical formicid stock” out
of which the following lineages were derived: (1) a “Prodorylinae” (i.e. Cerapachyinae) –
Dorylinae line, (2) a line comprising a hypothetical ancestral group the “Proponerinae”, (3) a
lineage labeled “?Dolichoderinae”, and (4) a lineage labeled “?Formicinae”. The “Proponerinae”
gave rise to several lines labeled: (1) Euponerinae; (2) Amblyoponini, Ectatommini, Proceratiini;
(3) Myrmeciini, Platythyreini; and (4) Myrmicinae. Wheeler expressed the view that this model
“indicated a polyphyletic origin of the subfamilies”, and that was his main reason for rejecting it.
Emery’s proposal was ultimately monophyletic, in that he clearly considered the hypothetical
stock to have been monophyletic, but it is not clear whether he considered the lineages he
depicted to have represented separate origins of sociality as Wheeler concluded.
Emery’s phylogenetic concept was in fact structurally close to that proposed here. His
“unknown hypothetical formicid stock” is schematically equivalent to the semisocial formicids
of the current model (and his “Proponerinae” is close to the “poneroid lineage” of Brown; 1954).
Wheeler (p. 108) rejected Emery’s model because: “if we take Emery’s point of view…the
hypothetical ancestors must have been very primitive and in all probability solitary or subsocial
Aculeates”. Thus Emery and Wheeler between them, came close to the phylogenetic hypothesis
developed in this essay (even though the latter opted for the subsocial route). A shift of emphasis
by Wheeler almost 80 years ago might have generated ideas similar to those expressed here.
Ideas, like evolution, can be very chancy!
The early stages of the evolutionary path to formicid eusociality proposed above relate well
to the “polygynous family hypothesis” of West-Eberhard (1978). The presence of allectomy,
however, projected the ant lineage on a later course completely different from those of other
socially advanced aculeates. The fact that the simple act of dealation can determine the potential
for future worker versus reproductive status in female ants provided a mechanism for ready
establishment of the worker caste and eusociality. That is the most important component of this
model!
Several recent studies have investigated the internal phylogeny of family Formicidae
cladistically, or by DNA sequence analysis and comparison, most notably Baroni Urbani, Bolton
& Ward (1992); Shattuck (1992); Grimaldi, Agosti & Carpenter (1997); Grimaldi & Agosti
(2000a), Brady (2003); Ward & Brady (2003); and Saux et al. (2004). More work in this area is
desirable, especially with the possibility of multiple evolution of eusociality in mind. Possibly
significant worker mesosomal and postpetiolar characters need to be better understood, along
with features related to the many pheromone-producing glands of ants, especially those involved
in the laying of scent trails. Further work on proventricular structure, especially in the
poneromorph, dorylomorph, leptanillomorph and myrmicomorph subfamilies might yield
phylogenetically useful characters.
There is a degree of prospective adaptation at each phylogenetic level in the model proposed
here. It is almost as if each step was automatically destined to be followed by the next. This is
especially true of the semisocial formicids with colonies comprising dealate reproductive
females and workers of one generation, and alate, potentially reproductive, females of the next.
Evolution of a eusocial worker caste following premature dealation of daughter alates in the
manner described above is so simple a process that its occurrence could be considered almost
predictable. Perhaps this should not amaze hymenopterists, for the various major groups of
aculeates have produced many separate social lineages, with a number culminating in eusociality
(in several groups of bees, vespine wasps, a sphecoid wasp and ants). This must have been, in
effect, preordained by the unique set of relevant characteristics phylogenetically assembled in
the ancestor common to all of these taxa.
Whether or not this model is considered plausible, it is clear that ants as aculeates are
endowed with a number of extraordinary characteristics. They have metapleural glands,
allectomy in alate reproductive females, permanently apterous workers and strong worker
dimorphism, often with large-headed soldiers; they lay scent trails, have acellular nests, and may
practice nocturnal and/or hypogaeic foraging behavior. In each case these characteristics are
unique to ants or very unusual, even rare, among other aculeates.
In conclusion ants, as my title implies, are very strange wasps indeed!
ACKNOWLEDGEMENTS
Sincere thanks are due to colleagues who gave advice on this paper or helped to polish the
arguments and presentation. All are gratefully acknowledged: Andrew Beattie, Denis Brothers,
James M. Carpenter, Ross Crozier, Michael S. Engel, Charles D. Michener, Ian Naumann,
Michael Schwarz, Steve Shattuck, Roy Snelling, John Trueman and Philip S. Ward, and my wife
Wendy Taylor. Michelle Hearn and Margaret Thornton provided much appreciated library
support. Newly reported field studies on Nothomyrmecia macrops were conducted with the late
Roger Bartell. Many of the ideas expressed here were discussed in my plenary address “Strange
Wasps Indeed” delivered to the XIII International Congress of IUSSI, Adelaide, Australia, 3
January, 1999. I especially thank Roy Snelling and Phil Ward for providing this fine opportunity
to celebrate the achievements of a distinguished friend and colleague.
LITERATURE CITED
Alexander, R.D. 1974. The evolution of social behaviour. Annual Review of Ecology and
Systematics 5: 325-381.
Baroni Urbani, C.B., Bolton, B. & Ward, P.S. 1992. The internal phylogeny of ants
(Hymenoptera: Formicidae). Systematic Entomology 17: 301-329.
Beattie, A.J., Turnbull, C.L., Hough, T. & Knox, R.B. 1986. Antibiotic production: a possible
function for the metapleural glands of ants (Hymenoptera: Formicidae). Annals of the
Billen, J.P.J. & Peeters, C. 1991. Fine structure of the gemma gland in the ant Diacamma
australe (Hymenoptera, Formicidae). Belgian Journal of Zoology 121: 203-210.
Bitsch, J. & Peeters, C. 1992. Moignons alaires et morphologie thoracique chez l'ouvrière de la
fourmi Diacamma australe (Fabricius)(Hym. Formicidae Ponerinae). Bulletin de la Societe
Entomologique de France 96: 213-221.
Bourke, A.F.G. & Franks, N.R. 1990. Social evolution in ants. 529 pp. Princeton, N.J.
Princeton University Press.
Brady, S.G. 2003. Evolution of the army ant syndrome: the origin and long-term evolutionary
stasis of a complex of behavioural and reproductive adaptations. Proceedings of the
National Academy of Sciences of the United States of America 100: 6575-6579.
Brown, W.L., Jr. 1954. Remarks on the internal phylogeny and subfamily classification of the
family Formicidae. Insectes Sociaux 1: 21-31.
Carpenter, J.M. 1991. Phylogenetic relationships and the origin of social behavior in the
Vespidae. in Ross, K.G. & Matthews, R.W. (eds). The social biology of wasps. 673 pp.
Ithaca, N.Y. Cornell University Press.
Cassill, D.L. & Tschinkel, W.R. 1995. Allocation of food to larvae via trophallaxis in colonies
of the fire ant Solenopsis invicta. Animal Behaviour 50: 801-813.
Casteels, P. 1998. Immune response in Hymenoptera. Pp 92-110 in Brey, P.T. & Hultmark, D.
(eds) Molecular mechanisms of immune responses in insects. 325pp. London. Chapman and
Hall.
Clark, J. 1951. The Formicidae of Australia, Volume 1 Subfamily Myrmeciinae. 230pp.
Melbourne. CSIRO Australia.
Crozier, R.H. & Pamilo, P. 1996. Evolution of social insect colonies. 306pp. Oxford. OUP
(Oxford series in ecology and evolution).
Dlussky, G.M. 1975. Formicidae in. Rasnitsyn, J.P. Hymenoptera Apocrita of the Mesozoic.
Transactions of the Palaeontological Institute of the Academy of Sciences of the USSR 147:
115-121 (in Russian).
Dlussky, G.M. 1999. New ants (Hymenoptera: Formicidae) from Canadian amber.
Palaeontological Journal 33: 409-412.
Dlussky, G.M., Brothers D.J. & Rasnitsyn, A.P. 2004. The first Late Cretaceous ants
(Hymenoptera: Formicidae) from southern Africa, with comments on the origin of the
Myrmicinae. Insect Systematics and Evolution 35: 1-13.
Eisner, T. 1957. A comparative morphological study of the proventriculus of ants
(Hymenoptera: Formicidae). Bulletin of the Museum of Comparative Zoology 116: 439-490.
Emery, C. 1920. La distribuzione geografica attuale delle formiche. Tentativo di spiegarne la
genesi col soccorso di ipotesi filogenetiche e paleogeografiche. Atti della R. Accademica
Lincei Memoria Classe di Scienze Fische, Matematiche e Naturale 13: 357-450.
Evans, H.E. 1958. The evolution of social life in wasps. Proceedings of the Tenth International
Congress of Entomology (Montreal, 1966) 2: 449-457.
Evans, H.E., & West-Eberhard, M.J. 1973. The wasps. 265 pp. Newton Abbot. David and
Charles.
Fabres, G. & Brown, W.L., Jr. 1978. The recent introduction of the pest ant Wasmannia
auropunctata into New Caledonia. Journal of the Australian Entomological Society 17:
139-142.
Fukumoto, Y., Abe, T. & Taki, A. 1989. A novel form of colony organization in the
“queenless” ant Diacamma rugosum. Physiology and Ecology Japan 26: 55-61.
Grimaldi, D.A. & Agosti, D. 2000a. A formicine in New Jersey Cretaceous amber
(Hymenoptera: Formicidae) and early evolution of the ants. Proceedings of the National
Academy of Sciences of the United States of America 97: 13678-3683.
Grimaldi, D.A. & Agosti, D. 2000b. The oldest ants are Cretaceous, not Eocene: comment.
Canadian Entomologist 132: 691-693.
Grimaldi, D.A. Agosti, D. & Carpenter, J.M. 1997. New and rediscovered primitive ants
(Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic
relationships. American Museum Novitates 3208: 1-43.
Grimaldi, D.A., Engel, M.S. & Nascimbene, P. 2002. Fossiliferous Cretaceous amber from
Myanmar (Burma): its rediscovery, biotic diversity, and palaeontological significance.
American Museum Novitates 3361: 1-72.
Gronenberg, W. & Peeters, C. 1993. Central projections of the sensory hairs on the gemma of
the ant Diacamma: substrate for behavioural modulation? Cell and Tissue Research 273:
401-415.
Haskins, C.P. & Haskins, E.F. 1951 Note on the method of colony foundation of the ponerine
ant Amblyopone australis Erichson. American Midland Naturalist 45: 432-445.
Haskins, C.P. & Haskins E.F. 1965. Pheidole megacephala and Iridomyrmex humilis in
Bermuda –equilibrium or slow replacement? Ecology 46: 736-740.
Haskins, C.P. & Haskins E.F. 1988. Final observations on Pheidole megacephala and
Iridomyrmex humilis in Bermuda. Psyche 95: 177-184.
Hölldobler, B.1978. Ethological aspects of chemical communication in ants. Pp75-115. in

Rosenblatt J.S., Hinde R.A., Beer C. & Busnel. M.C. (eds.) Advances in the study of
behavior, volume 8. New York. xiv + 261 pp. Academic Press.
Hölldobler, B. 1980. Canopy orientation: a new kind of orientation in ants. Science 210: 86-88.
Hölldobler, B. & Engel-Siegel, H. 1985. On the metapleural gland of ants. Psyche 91: 201-224.
Hölldobler, B & Haskins, C.P. 1977. Sexual calling behavior in primitive ants. Science 195:
793-794.
Hölldobler, B. & Taylor, R.W. 1983. A behavioral study of the primitive ant Nothomyrmecia
macrops Clark. Insectes Sociaux 30: 384-401.
Hölldobler, B. & Wilson, E.O. 1990. The Ants. 732 pp. Cambridge, Mass. Harvard Belknap
Press.
Holway, D.A., Lach, L., Suarez, A.V., Tsutsui, N.D. & Case, T. 2002. The causes and
consequences of ant invasions. Annual Review of Ecology and Systematics 33: 181-233.
Houston, T.F. 1970. Discovery of an apparent male soldier caste in a nest of a halictine bee
(Hymenoptera: Halictidae), with notes on the nest. Australian Journal of Zoology 18: 345-
351.
Hunt, J.H., Jeanne, R.L. & Keeping, M.G. 1983. Observations on Apoica pallens, a nocturnal
Neotropical social wasp. Insectes Sociaux 42: 223-236.
Ito, Y. 1993. Behaviour and social evolution of wasps: the communal aggregation hypothesis.
159 pp. Oxford. Oxford Series in Ecology and Evolution.
Kaaya, G.P. 1993. Inducible humoral antibacterial immunity in insects. Pp 71-89 in Pathak,
J.P.N. (ed) Insect immunity. 192pp. Dordtrecht. Kluwer Academic Publishers. (W. Junk
Series Entomologica 48).
Krombein, K.V. 1967. Trap-nesting wasps and bees: Life histories, nests, and associates. 570
pp. Washington, Smithsonian Press.
Mackintosh, J., Trimble, J.E., Jones, M.K., Karuso, P.H., Beattie, A.J. & Veal, D.A. 1995.
Antimicrobial mode of action of secretions from the metapleural gland of Myrmecia gulosa
(Australian bull ant). Canadian Journal of Microbiology 41: 136-144.
Malyshev, S.I. 1968. Genesis of the Hymenoptera and the phases of their evolution. Translated
from Russian by O.W. Richards & B. Uvarov. 319 pp. London, Metheun.
Matsuura, M. 1991. Vespa and Provespa. Pp. 232-62 in: Ross K.G. and Matthews R.W. (eds.)
The social biology of wasps. 673 pp. Ithaca, N.Y. Comstock.
Matsuura, M. & Yamane, S. 1984. Biology of the vespine wasps. 323 pp. Berlin. Springer-
Verlag
Michener, C.D. 1974. The social behavior of the bees: a comparative study. 404 pp.
Cambridge, Mass. Harvard University Press.
Michener, C. D. 2000. The bees of the world. 950 pp. Baltimore. Md. John Hopkins University
Press.
Naumann, I.D. & Masner, L. 1985. A revision of the termitophilous Australian genus
Leaiopria Dodd (Hymenoptera: Diapriidae). Journal of the Australian Entomological
Society 19: 143-149.
Naumann, I.D., van Achterberg, C., Houston, T.F., Michener, C.D. & Taylor, R.W. 1991.
Hymenoptera. Pp 916-1000 in: The insects of Australia, a textbook for students and
scientific workers. 1137pp. Melbourne. Melbourne University Press.
Nel, A., Perrault, G., Perrichot, V. & Neraudeau, D. 2004. The oldest ant in the Lower
Cretaceous amber of Charente-Martime (SW France)(Insecta: Hymenoptera: Formicidae).
Geologica Acta 2: 23-29.
Peeters, C. & Billen, J.P.J. 1991. A novel exocrine gland inside the thoracic appendages
('gemmae') of the queenless ant Diacamma australe. Experientia (Basel) 47: 229-231.
Peeters, C., Billen, J.P.J. & Hölldobler B. 1992. Alternative dominance mechanisms
regulating monogyny in the queenless ant genus Diacamma. Naturwissenschaften 79: 572-
573.
Peeters, C. & Higashi, S. 1989. Reproductive dominance controlled by mutilation in the
queenless ant Diacamma australe. Naturwissenschaften 76: 177-180.
Peeters, C. & Ito, F. 2001. Colony dispersal and the evolution of queen morphology in social
Hymenoptera. Annual Review of Entomology 46: 601-630.
Peeters, C. & Tsuji, K. 1993. Reproductive conflict among ant workers in Diacamma sp. from
Japan: dominance and oviposition in the absence of the gamergate. Insectes Sociaux 40:
119-136.
Saux, C., Fisher, B.L. & Spicer, G.S. 2004. Dracula ant phylogeny as inferred by nuclear 28S
rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae:
Amblyoponinae). Molecular Phylogenetics and Evolution 33: 457-468.
Schremmer, F. 1972. Beobachtungen zur Biologie von Apoica pallida (Olivier, 1791), einer
neotropischen sozialen Faltenwespe (Hymenoptera, Vespidae). Insectes Sociaux 19: 343-
357.
Schulz, T.R. 2000. In search of ant ancestors. Proceedings of the National Academy of Sciences
of the United States of America 97: 14028-14029.
Shattuck, S.O. 1992. Higher classification of the ant subfamilies Aneuretinae, Dolichoderinae
and Formicinae (Hymenoptera: Formicidae). Systematic Entomology 17: 199-206.
Sommer, K., Hölldobler B. & Rembold, H. 1993. Behavioral and physiological aspects of
reproductive control in a Diacamma species from Malaysia (Formicidae, Ponerinae).
Ethology 94: 162-170.
Taylor, R.W. 1978. Nothomyrmecia macrops: a living-fossil ant rediscovered. Science 201:
979-985.
Taylor, R.W. 1985. The ants of the Papuasian genus Dacetinops (Hymenoptera: Formicidae:
Myrmicinae. Pp. 41-67 in: Ball, G.E. (ed.) Taxonomy, phylogeny and zoogeography of
beetles and ants. 514pp. Dordrecht. W. Junk Series Entomologica 33.
Taylor, R.W. 1992. The nomenclature and distribution of some Australian and New Guinean
ants of the subfamily Formicinae (Hymenoptera: Formicidae). Journal of the Australian
Tschinkel, W.R. 1987. Seasonal life history and nest architecture of a winter-active ant,
Prenolepis imparis. Insectes Sociaux 34: 143-164.
Tulloch, G.S. 1934. Vestigial wings in Diacamma (Hymenoptera: Formicidae). Annals of the
Veal, D.A., Trimble, J.E. & Beattie, A.J. 1992. Antimicrobial properties of secretions from the
metapleural glands of Myrmecia gulosa (the Australian bull ant). Journal of Applied
Bacteriology 72: 188-194.
Veuille, M., Brusadelli, A., Brazier, L. & Peeters, C. 1998. Phylogenetic study of a
behavioural trait regulating reproduction in the ponerine ant Diacamma. P. 492 in Schwarz
M.P. & Hogendoorn. K. (eds.) Social insects at the turn of the millennium. Proceedings of
the XIII International Congress of IUSSI, Adelaide Australia, 29 December 1998 - 3
January 1999. 535 pp., Adelaide, XIII Congress of IUSSI.
Ward, P.S. & Brady, S.G. 2003. Phylogeny and biogeography of the ant subfamily
Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361-368.
Wehner, R.D., Harkness, R.D. & Schmid-Hempel, P. 1983. Foraging strategies in
individually searching ants Cataglyphis bicolor (Hymenoptera: Formicidae). 79pp.
Stuttgart. Gustav Fischer Verlag.
West-Eberhard, M.J. 1978. Polygyny and the evolution of social behavior in wasps. Journal of
the Kansas Entomological Society 51: 832-856.
Wheeler, W.M. 1928. The social insects. Their origin and evolution. 377 pp. London: Kegan,
Paul, Trubner.
Wheeler, W.M. 1933. Colony-founding among ants with an account of some primitive
Australian species. 179 pp. Cambridge, Mass: Harvard University Press.
Wilson, E.O. 1958. The beginnings of nomadic and group-predatory behavior in the ponerine
ants. Evolution 12: 24-31.
Wilson, E.O. 1971. The insect societies. 548 pp. Cambridge, Mass. Harvard Belknap Press.
Wilson, E.O., Carpenter, F.M. & Brown, W.L., Jr. 1967. The first Mesozoic ants, with the
description of a new family. Psyche 74: 1-19.
Trager, J. C., MacGown, J. A., Trager, M. D. 2007. Revision of the Nearctic endemic Formica
pallidefulva group, pp. 610-636. In Snelling, R. R., B. L. Fisher, and P. S. Ward (eds). Advances
in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50 years of
REVISION OF THE NEARCTIC ENDEMIC FORMICA PALLIDEFULVA GROUP
James C. Trager
Shaw Nature Reserve
P. O. Box 38 / Interstate 44 and Highway 100
Gray Summit, MO 63039, USA
james.trager@mobot.org
Joe A. MacGown
Department of Entomology & Plant Pathology
Mississippi State University, Box 9775
Mississippi State, MS 39762, USA
jmacgown@entomology.msstate.edu
and
Matthew D. Trager
Department of Entomology and Nematology
P.O. Box 110620
University of Florida
Gainesville, FL 32611, USA
ABSTRACT
We revise the Nearctic endemic Formica pallidefulva group based on study of types and other
museum specimens and material in J. Trager’s collection. The latter material originates from 30
years of accumulated samples, both from free-living colonies of F. pallidefulva group species,
and from “slave” populations in colonies of Polyergus lucidus s. l., which have single-species
host populations. Among the currently available names for the group, the four valid taxa are F.
archboldi, F. dolosa, F. incerta and F. pallidefulva. There is a fifth common, but previously
unrecognized new species, described here as Formica biophilica Trager, n. sp. Earlier
taxonomies of this group were constrained by typological thinking and inadequate treatment of
metric characters. For this study, well preserved individuals, nest series and types of all but one
taxon were studied (no types seen for F. pallidefulva). Analysis focused on form, length,
abundance and distribution of macrochaetae (pilosity); length and density of microchaetae
(pubescence); standard measurements and indices; distinct habitat preferences of the various
species; and host selection by Polyergus lucidus, s. l. Our results leave little doubt that F. incerta
n. stat., rev. stat. and F. biophilica n. sp. deserve recognition as species, that F. nitidiventris is a
synonym of F. pallidefulva and that F. schaufussi is a synonym of F. pallidefulva. Thus, the
current concept of schaufussi (incorrect in reference to the lectotype) must give way to the next
available name for the same population, dolosa n. stat.
In this paper, we provide diagnoses, qualitative morphological characteristics, tabulated
quantitative characters, natural history notes for all species and a key to the workers. As occurs
in other groups of closely related ant species (e.g. Umphrey, 1996; Steiner et al., 2006),
morphology is variable and partially overlapping, and species determinations will be more
certain when based on nest series consisting of three or more workers in prime condition.
Trager et al.: Formica pallidefulva group 611
Key words: Hymenoptera, Formicidae, Formicinae, Formica pallidefulva group, taxonomy,

morphometric characterization, natural history, habitat preferences, Nearctic, new species,
lectotype designation.
INTRODUCTION
The Formica pallidefulva species group is endemic to the eastern and central United States, with
two species extending west into the Great Plains and lower elevations in the Rocky Mountains
and north into eastern Canada. The species of the pallidefulva group are distinguished from
others in Formica by gracile body form and elongate appendages; integument moderately to
strongly shining; propodeum in profile with the angle between dorsal and posterior faces
completely obscured by rounding or at most indistinct (Fig. 6a-e); external mesometasternum of
uniform simple structure throughout the group (Fig. 1), this structure unadorned or rarely with a
few erect setae; male genitalia with elongate parameres; and behavior and habitat choice
generally thermophilous. They mainly inhabit grassland and open woodland habitats; only F.
pallidefulva Latreille is normally also associated with closed canopy forest. There are no social
parasites in the group, and all have a clear size difference between the workers and queens.
These species are hosts of dulotic Polyergus lucidus (sensu lato) and some F. sanguinea group
species. They are also temporary hosts during colony-foundation for some species in the F. rufa
/ microgyna group (Wheeler, 1904; Creighton, 1950). Colonies typically contain 500 or fewer
workers (Wheeler, 1904; Talbot, 1948). Nests are inconspicuous except when opening onto bare
ground, but may bear a small mound nestled among herbaceous vegetation of grasslands or
savannas, or occasionally a somewhat larger mound. Nests are quite often under stones or other
cover objects in cooler parts of the range, but generally not so further south. Recruitment of
individuals by social transport occurs during nest relocation, which may occur one or more times
per year in some colonies. Robson & Traniello (1998) have given a thorough accounting of
foraging and recruitment in F. incerta and F. dolosa (vouchers seen). Wheeler (1904) first
described the “top-heavy” nests composed of broad chambers and narrow interconnecting
passages of species in this group, and Talbot (1948) also described and illustrated them.
Mikheyev & Tschinkel (2000) have provided a modern study of nest architecture and economics
of nest building of one species. Milford (1999) reports the presence of a species in riparian
forests along the Rio Grande. (Presumably, F. pallidefulva was the subject of these two recent
studies, but vouchers were not available for identification). Bale et al.,(2004) reported on the
role of “F. schaufussi” in dispersing seeds of trilliums, and how the ants hastened their retrieval
and transport of these seeds when disturbed by yellowjackets (no vouchers seen).
Taxonomic history of the Formica pallidefulva group
The taxonomic history of this group begins with the description of the species Formica
“pallide-fulva” by the French naturalist Latreille (1802). F. schaufussi was first described as a
separate species by Mayr (1866). Emery (1893) relegated F. schaufussi to a subspecies of F.
pallidefulva, and described several new varieties and subspecies of F. pallidefulva. The
subgenus Neoformica was published twice in the same year by W. M. Wheeler to encompass
these taxa, first while designating F. pallidefulva as its type species (1913a) and later in the
formal description of the new subgenus in his revision of Formica (1913b). In the latter,
Wheeler described the varieties dolosa (cursorily described as var. meridionalis, an unavailable
name, by Wheeler in 1904, then renamed by him in 1912) and succinea. Buren (1944) first used
the taxon dolosa at the subspecies rank. Two further subspecies, F. p. delicata (Cole 1938) and
F. p. archboldi (M.R. Smith 1944), were added to the described forms in this group that
Creighton attempted to organize in his classic Ants of North America (1950).
Creighton popularized the subgenus name Neoformica and attempted to organize the taxa in
the group by reducing them (among those species treated in the present work) to three species;
the Floridian F. archboldi, and two widely-distributed species, the relatively non-pilose F.
pallidefulva (with northern and southern subspecies nitidiventris and pallidefulva) and the very
pilose F. schaufussi (with northern and southern subspecies schaufussi and dolosa). Leaving
aside for later some nomenclatural problems with Creighton’s scheme, it has become clear that
his classification was an oversimplification, since it synonymized the ecologically and
morphologically distinct F. incerta, and left unrecognized the new species described below.
Both of these species have rather variable arrays of macrochaetae and if, as seems to be the case
in many of the “identified” specimens we examined, abundance of pilosity is the primary or only
trait used to identify them, rubbed specimens or individuals near the extremes of intraspecific
variation within F. incerta or the new species may easily be misidentified as belonging to
Creighton’s concepts of either pallidefulva or schaufussi.
Finally, Wheeler’s (and Creighton’s) concept of Neoformica also encompassed F. moki
Wheeler, its subspecies grundmanni (now a synonym of xerophila) and xerophila M. R. Smith
(now a full species). These latter are all now considered members of the F. fusca group.
Excluding these taxa, the monophyly of the F. pallidefulva group may be supposed on
morphological (this study) and genetic grounds (Riitta Savolainen, personal communication).
Nevertheless, Buren (1968) synonymized this and the other subgenera into Formica, and we will
let this stand until a proper phylogeny of Formica is completed. Neoformica does continue to
appear in print, even as a genus name, especially in the non-taxonomic ant literature (e.g.
Abouheif & Wray, 2002)
Suggested and possible relatives of the Formica pallidefulva group
Several Formica species from outside of eastern North America have been suggested as
members or relatives of the F. pallidefulva group.
Among these, F. rufolucida Collingwood is the most similar to members of the pallidefulva
group in outward appearance, but we believe it is not a member of, or even a sister group to, the
pallidefulva group. Formica rufolucida is a rather shiny species with long scapes and rounded
propodeum from 7000 ft. elevation in the mountains of Burma (Myanmar). It was suggested as a
geographically disjunct member of the F. pallidefulva group and placed in Neoformica by
Collingwood (1962). Bolton (1995) questioned even the generic placement based on the
extraordinary location, but a paratype specimen of this species sent by Mr. Collingwood
confirms it as Formica. The thoracic sternal region of the specimen could not be examined, and
males are unknown, but the examined specimen otherwise has the proportions and sculpture
(although subtle) of the fusca group, not of the pallidefulva group.
The Iberian F. subrufa Roger has a superficial resemblance in the worker’s proportions,
sculpture and pilosity to F. archboldi, but the concave mesonotum of the workers and unusual
proportions of the sexual forms of this Spanish species are quite different from those of any
species in the pallidefulva group. Similarities between F. subrufa and the pallidefulva group
could well be symplesiomorphies, as these groups are both near the base of the cladogram in a
tentative, DNA-sequence based phylogeny of Formica being elucidated by Riitta Savolainen
(personal communication).
F. moki and F. xerophila are western North American species that also have long scapes and
a rounded propodeum. Despite their long limbs, these ants fall short of the gracile form of the
pallidefulva group, especially the mesosoma, and are quite different in vestiture, and especially,
sculpture. Furthermore, F. moki at least, is parasitized by Polyergus breviceps (s. l.), perhaps an
indication of its closer relationship to other hosts of this species. The mesometasternum of F.
moki differs in conformation and pilosity from that of pallidefulva group species described
below. R. Snelling writes (personal communication) that the genitalia of F. moki are similar to
fusca group males, lacking the elongated parameres of pallidefulva group males.
METHODS AND TERMINOLOGY
J. Trager collected specimens of these ants from the 1970’s till the present, in locations from
Long Island to Florida and Iowa to Colorado. Some colonies were reared for one or more years
to obtain males of certain identity. The Polyergus lucidus complex was another focus of
collection efforts over the same period, and colonies of these Amazon ants yielded host
populations that seemed in the field to belong to a single pallidefulva group species per
slavemaker colony, a suspicion confirmed in this study. Species of the pallidefulva group also
turned up in sampled host populations of dulotic Formica, particularly F. pergandei Emery over
much of its range and F. creightoni Buren in northern Missouri. In addition to Trager's extensive
collections, MacGown also collected specimens of this species group in the southeastern United
States from 1988 to the present, an area that sorely lacked distributional data. The result was a
good sampling of all species from much of their natural range.
In SPECIMENS EXAMINED under each species, we list for each State only the counties
from which specimens were collected, not the specific localities.
Museum specimens borrowed for study or examined in situ augmented the Trager material
(JCT). Particularly important were types of Gustav Mayr, Carlo Emery, William M. Wheeler
and Arthur Cole from the following museums (and the people who facilitated the loans): Museo
Civico de Historia Natural “Giacomo Doria”, Genova (MCSN-Roberto Poggi),
Naturhistorisches Museum Wien (NMW-Stefan Schoedl), Harvard University Museum of
Comparative Zoology (MCZ-Stefan Cover) and the Los Angeles County Museum of Natural
History (LACM-Roy Snelling). Other important entomological collections consulted included
those of Archbold Biological Station (AABS-Mark Deyrup), the Florida State Collection of
Arthropods (FSCA-Jim Wiley), the entomology department collections of the University of
Arkansas – Fayetteville (UAAM-Jim Whitfield), the University of Missouri – Columbia
(UMRM-Robert Sites), Mississippi State University (MEM-Joe MacGown), and the University
of Wisconsin – Madison (IRCW-Andrew Williams, Scott Sauer). The last is where the
voluminous collections of Mr. Williams’s Insect Associates of Prairie Plants study and the
Prairie Insect Survey of the Wisconsin Department of Natural Resources are housed. Other
colleagues too numerous to mention sent one or a few samples each from interesting localities or
as vouchers for their own studies.
Drawings for Figures 1 and 4-6 were done on a Leica MZ 16 stereomicroscope, using a
drawing tube. Figure 1 was drawn at 63×, Figures 4-6 were drawn at 50×, except for the
pubescence maps (Figure 4, insets) that were drawn at 100×.
Metric characteristics included measurements of body parts (mm) and indices calculated
from them, as well as tooth and hair counts, as listed below. Measurements and meristic
characters were acquired at 50× or 100× magnification, and data were entered in spreadsheets
and converted to hundredths of millimeters for analysis. Summarized morphometric data for all
specimens examined are presented in Tables 1 and 2.
MdT - number of mandibular teeth including any conspicuous denticles added to the basic
number of 7. Specimens with MdT=6 appeared in every case to be older individuals in
which any denticles and one of the (usually) smaller teeth were worn away.
P5L, P6L - lengths of 5th and 6th labial palp segments.
PnML, PpML, TML - lengths of longest erect hair (M = macrochaeta) on dorsum of
pronotum, propodeum and center dorsum of first gastral tergite.
TPL - length of a longest microchaeta (shorter, appressed hairs collectively called
pubescence) among those visible on the disc of the first gastral tergite.
HL - head length from distal tip of clypeus to posterior margin of head in full face view.
HW - maximum head width (exclusive of eyes) in full face view.
EL - compound eye length, in oblique view of the head, showing full surface of eye.
SL - scape length, with viewing angle perpendicular to the scape.

WL - length of mesosoma (traditionally called Weber's Length of thorax), from anterior
extremity of pronotum (exclusive of pronotal collar) to posterolateral extremity of
mesopleuron.
PnW - width of pronotum.
PnM, MnM, PpM - number of erect or suberect macrochaetae on pronotum, mesonotum
and propodeum.
TM - number of macrochaetae on first gastral tergite, exclusive of those along the posterior
border.
CI - Cephalic index, HW / HL × 100
SI - Scape index, SL / HW × 100 (This is a good surrogate for relative length of the limbs,
generally.)
OI - Ocular index, EL / HL × 100
CTI - Cephalothoracic index, HL / WL × 100
TWI - Thoracic width index PnW / WL × 100 (This is a good surrogate for overall
gracility.)
We first analyzed the morphometric data with discriminant function analysis (DFA), a
statistical procedure used to classify individuals and then predict group membership (species, in
this case) based on a set of predictor variables (Tabachnick & Fidell, 1996). Because this
method can consider a large number of variables, DFA can be applied to difficult taxonomic
relationships when there are only small differences among taxa for any one characteristic (e.g.
Bell, 1996; Burbrink, 2001; Floate & Whitham, 1995; Green et al., 1996). In this analysis we
performed stepwise DFA (using Wilks’ Lambda as a measure of variable contribution with enter
F at P = 0.05; removal F at P = 0.10) to test whether the specimens used to describe species in
this revision actually belonged to statistically distinct groups that differed across a set of
morphological measurements, thus verifying the morphologically-based taxonomy proposed in
this paper. We also conducted univariate ANOVA on some of the morphometric data to
distinguish between frequently confused pairs of species. All tests were performed with SPSS
11.5. The statistical analysis is meant to provide a quantitative complement to the morphological
descriptions, natural history information and key to the species provided below. It follows the
species accounts and precedes the key to species.
Species of the Formica pallidefulva group
Based on earlier observations, this study began with a working hypothesis of seven species
in the group, namely the Floridian archboldi and three north/south species pairs:
schaufussi/dolosa of well-drained, acid soils; nitidiventris/pallidefulva of mesic, neutral soils,
incerta/new species of open grassland habitats. Measurements and other characteristics obtained
during this study confirmed the separate species identities of incerta and the new species
(described below as F. biophilica n. sp. Trager), while the first two pairs of taxa were found to
represent extremes of regional variation within two single-species populations. Thus, there is a
total of five distinct species, as below. The correct names for each population were determined
through study of type specimens.
Formica archboldi M. R. Smith, 1944

Figures 4d, 5d, 6d
Formica pallidefulva subsp. archboldi Smith, M.R., 1944: 16. [Examined. Syntype workers] four
workers on two pins, labeled, Florida: Archbold Biological Station, 10 miles south Lake Placid,
Fla. X-7-43 T. C. Schneirla. Paratype No. 56765 U.S.N.M. (MCZ).
Formica (Neoformica) archboldi Creighton, 1950: 549. Raised to species.
DIAGNOSIS
Worker – The most evidently sculptured and the smallest member of the group,
characteristic of Florida’s upland pine and scrub habitats. F. archboldi has the largest eyes
relative to head size of all species in this group (see OI, Table 1). Head and gaster very dark
reddish brown, appearing blackish or dark gray-brown in the field; mesosoma often a little
lighter than head, and sometimes both lighter than gaster. Gastral dorsum at most weakly
shining, sheen dulled by numerous, shallow impressions (foveolae) and appressed pubescence
composed of moderately dense grayish appressed microchaetae (Fig. 4d). Dorsal sclerites of
mesosoma and gaster usually with moderately abundant, short, erect, brownish-gray erect
macrochaetae, and pale grayish appressed microchaetae. Erect macrochaetae on mesosoma and
usually on gaster are relatively short, straight and flattened with rounded, blunt or abruptly
tapering tips; less often, at least some of those on gaster a bit longer and slightly curved, these
less flattened and tapering.
Queen – Color, gastral pubescence and sculpture like the workers’, with the usual
differences in size. There is tessellation on the upper portion of head, pronotum, sides of
mesothorax, propodeum and gastral dorsum; wings, when present, clear brownish to clear smoky
gray. Pilosity longer and more flexuous than that of worker.
Male – Pubescence and pilosity abundant; mesosomal dorsum dull-punctate; entire body
black, legs reddish brown or mesosoma lighter, dusky yellowish brown; wings clear brownish to
clear smoky gray; pilosity like that of queen; gastral pubescence pale brownish yellow and very
dense. Averages smaller than the otherwise difficult-to-distinguish males of F. dolosa and F.
biophilica.
DISTINGUISHING FEATURES
Although a sample of this species in the Mayr collection is placed among his “schaufussi”
(anything in the group which was not typical F. pallidefulva is so placed in that collection), this
dark brown to nearly black species has rarely been misidentified since its description in 1944
and is unlikely to be confused with any congener in the field. Mounted specimens that are
shinier and less pilose than normal, or mislabeled ones (like those from “Virginia” in the Mayr
collection), might be confused with darker color variants of F. incerta or F. pallidefulva without
careful inspection. Occasionally other species, especially F. biophilica, may stain black when
mounted on pins, which could lead to confusion with F. archboldi.
ETYMOLOGY
This species is named after Richard Archbold, founder of Archbold Biological Station, Lake
Placid, Florida, the type locality.
RANGE AND HABITAT
Originally described from Archbold Biological Station in Highlands Co., Florida, this
species in fact appears to be more abundant in northern Florida. It is a characteristic ant of the
uplands of peninsular Florida and the eastern panhandle, and also recorded by D. R. Smith
(1979) as occurring in Georgia and Alabama. One collection in the Mayr collection labeled
“Virginia” is doubtless mislabeled. This ant is characteristic in relatively undisturbed, long-leaf
pine sandhills and in scrub or sand pine woodland. Less often it may be found in the transition
between these more open sandy habitats and drier portions of flatwoods, or southern live oak
woodlands. In northern Florida, F. archboldi coexists with F. dolosa and F. pallidefulva in
sandhill vegetation, but to the south on the Lake Wales Ridge (including the type locality), this
ant becomes more typical of moist soil among pond-edge vegetation in swales within the white
sand scrub vegetation, and is largely replaced in sandhill woodland by F. pallidefulva. It is also
reported from sandhill locations along the South Florida coasts, but probably is lacking from the
Everglades.
SPECIMENS EXAMINED
FLORIDA: Alachua; Baker; Citrus; Collier; Duval; Hernando; Highlands; Hillsborough;

Jackson; Leon; Levy; Liberty; Marion; Okochobee; Pasco; Putnam; Sumter; Suwannee; Volusia;
Wakulla.
D.R. Smith (1979) reports F. archboldi from ALABAMA: no county listed. GEORGIA: no
county listed.
NATURAL HISTORY
Within the pallidefulva group, this species is the most sensitive to human development and
habitat alteration. During eight years of residence in Gainesville FL, J. Trager watched this
species slowly disappear from sandhill woodland habitats near new housing developments, even
when efforts were made to protect the native vegetation in green space around the houses. On
the other hand, at Devil’s Millhopper Geological State Park during the same years, ecological
restoration efforts consisting of selective species removal (cutting down mesic-adapted trees)
and introduction of prescribed fire to maintain the open vegetation structure, resulted in a
resurgence of F. archboldi (and incidentally, F. dolosa) in the upland habitats of the site. A
healthy population of F. archboldi could be considered an indicator of high natural area
integrity, or at least a low level of habitat degradation, in Florida’s pine and pine-oak woodlands.
Nests are usually located beneath wiregrass clumps (or beneath beard grass or sedges in
scrub habitats), and occur less frequently in bare soil, at the base of a shrub or under oak-leaf
litter. The entrance is often marked with a small accumulation of plant fragments and/or
grasshopper dung. Returning foragers watched for ½-hour intervals in the afternoon foraging
period typically brought in more of these plant fragments and dung pellets than they did prey
items (J. Trager, unpublished). Worker pupae are typically enclosed in a light tan cocoon, sexual
pupae in a darker, thicker cocoon. This species is the host of the small, dull variant of Polyergus
lucidus that uses F. archboldi as its host in Florida (Trager & Johnson, 1985). F. archboldi lives
outside the range of dulotic or other parasitic Formica species.
The cricket Myrmecophila pergandei Bruner commonly inhabits the nests of F. archboldi.
Outside the nest, F. archboldi gathers honeydew from living plant surfaces and from leaf litter,
and also actively tends and defends Cinara aphids and Toumeyella scales on “grass-stage” long-
leaf pine saplings. The defense by F. archboldi as they tended scales was used to induce workers
to “attack” a termite offered on the end of a pine needle, then following the light-bodied prey as
a marker to follow as the worker returned to its nest. In two cases, this resulted in the discovery
of a colony of the Polyergus lucidus variety mentioned above.
Foraging occurs mainly between 8 a.m. and noon and between 4 p.m. and dusk, from March
through October. However, hemipteran colonies are tended around the clock. Peak foraging
activity occurs from April through June. Returning foragers carry a variety of freshly killed
insects into the nest. Most of these prey items are herbivorous insects, but also among them are
occasional individuals of Odontomachus brunneus Patton. Just how it is that this smaller and
less ferociously built Formica captures and kills this well-armed ponerine has not been observed.
Trager & Johnson (1985) report on habits of F. archboldi.
Sexuals occur in the nests from late April through June. The alates are not attracted to lights.
They apparently fly in early morning, around sunrise. Females must quickly dealate and
sequester themselves, as it appears no one has ever found either a newly mated female or
incipient colony of this locally abundant ant.
Formica biophilica James C. Trager new species

Figures 4b, 5b, 6b
Formica pallidefulva: Creighton, 1950, in part [Misidentification]

Formica schaufussi subsp. dolosa: Creighton, 1950, in part [Misidentification]
TYPE DATA
HL 1.43; HW 1.15; EL 0.42; SL 1.70; WL 2.34; PnW 0.95; PnM 6 ; PpM 14 ; TM 30
Type specimen label: ALABAMA, Chilton Co., Interstate-65 rest area 3 mi. E of Thorsby.
1-X-1983. M.B., J.R., B.R. DuBois. (JCT). Holotype and three paratypes on two pins. The
holotype and one paratype will be deposited at MCZ and the other two specimens will be added
to the excellent material of this species collected in Alabama and elsewhere by W. S. Creighton,
now housed at LACM.
DIAGNOSIS
Worker – Gracile, shiny, and the brightest yellow member of the group. Head, mesosoma
and legs light reddish- to pale brownish yellow; head and mesosoma not at all or only a little
lighter than gaster. Dorsal sclerites of mesosoma and especially the gaster with long, usually
curved, erect macrochaetae. Number of macrochaetae on propodeum usually exceeds the
number on the pronotum (20 of 32 specimens examined). Sheen of gaster readily visible through
pubescence composed of pale, slender, grayish hairs of medium density (Fig. 4b). Erect
macrochaetae on gaster long, commonly 0.25-0.30 mm, tapering to a point and curved (Fig. 6b).
Queen – Color, gastral pubescence and shininess like the workers’, with the usual
differences in size; with faint tessellation of upper portion of head, pronotum, sides of
mesothorax, propodeum and gastral dorsum; wings, when present, clear to light brownish.
Male – Pubescence and pilosity abundant; mesosomal dorsum dull-punctate; head and
gaster very dark brown, appearing black; mesosoma dusky yellowish brown, legs reddish brown;
wings clear to light brownish. A little brighter in color, especially mesosoma, less pilose, pilosity
also finer, and less pubescent than dolosa.
F. biophilica and F. incerta are sympatric in the southern part of the latter’s range. In the
field, F. biophilica appears more brightly and uniformly reddish-yellow in color, and (if several
workers of a colony are present) weakly polymorphic, whereas F. incerta is more brownish
yellow, smaller overall, and usually (though not invariably) more monomorphic within colonies.
Metrically, F. biophilica is more slender, has a proportionally longer mesosoma and narrower
head than F. incerta (compare SI, TWI and CI values, Table 1). F. biophilica usually has more
macrochaetae on the propodeum than on the pronotum (20 of 32 specimens), whereas F. incerta
usually has more macrochaetae on the pronotum than on the propodeum (22 of 31 specimens).
F. biophilica is also somewhat more brightly and uniformly colored, shinier and overall less
hairy, and has a sharper petiolar crest in profile than F. dolosa. The F. biophilica specimens
examined in this study also had a significantly higher CTI and OI (were more slender and had
relatively larger eyes) than F. dolosa (Table 1). In the South, the less pilose minor workers and
nanitics of F. biophilica are difficult to differentiate from F. pallidefulva. Often a rather squarer
propodeal profile and more uniform bright reddish yellow color indicate F. biophilica.
Additionally, on average the TWI of F. pallidefulva is larger than that of F. biophilica (Table 1).
Ecologically, specimens from fens, bogs, swamps and fresh or salt marshes are most likely to be
F. biophilica. In the northern part of the range of F. biophilica, its bright color will always
distinguish it from the at least partially brown F. incerta and F. pallidefulva.
ETYMOLOGY
The name biophilica is given in allusion to E. O. Wilson’s popularly inspirational coining

“biophilia”, meaning the love of other species as a part of human nature. Specimens from
Alabama, Dr. Wilson’s home state, were chosen as the type series to further honor his
contributions to myrmecology, conservation and behavioral biology.
RANGE AND HABITAT
Found in mesic to hydric open habitats, including fields, prairies, lawns, fens, bogs, marshes
and open woodlands, from the Carolinas to Missouri, south to northern Florida and central
Texas. Northward, its occurrence is more sporadic, especially in formerly glaciated regions,
where F. biophilica shifts to drier (thus warmer) loess and sandy grassland locations. It reaches
central Illinois in the Illinois River outwash sand prairies and reaches southeastern New York in
the sandy plains along the East Coast and on Long Island. The habitat overlaps that of F. incerta
in unglaciated prairies and eastern meadows, and overlaps that of F. dolosa in southern pine
woodland and savanna. F. biophilica is absent from the most xeric and infertile sites occupied by
F. dolosa. In the Ozarks and other southern U.S. hills, F. biophilica occurs in groundwater fens,
bogs, marshes and flatwoods. This is the only southern Formica that occurs in these wetland
habitats, where it nests in the elevated hummocks of organic matter formed by grass or sedge
tussocks. It is less common than F. pallidefulva in human habitats, but occasionally shows up in
lawns, parks and campuses, especially in parts of the South where fire ants are less abundant.
SPECIMENS EXAMINED
ARKANSAS: Logan; ALABAMA: Butler; Chilton; DeKalb; Lawrence; Mobile; Morgan;

St. Clair; Tuscaloosa; DELAWARE: Sussex; DISTRICT OF COLUMBIA: Washington;
FLORIDA: Alachua; Columbia; Gadsden; Okaloosa; Polk; Walton; GEORGIA: Clarke;
Habersham; Lumpkin; Rabun; ILLINOIS: Mason; LOUISIANA: Natchitoches; Tammany;
Washington; MISSISSIPPI: Alcorn; Bolivar; Chickasaw; Lafayette; Lee; Monroe; Oktibbeha;
Panola; Pontotoc; Tippah; MISSOURI: Franklin; Lincoln; Reynolds; Washington; NEW YORK:
Rockland; SOUTH CAROLINA: Pickens; TENNESSEE: Davidson; Monroe; Sevier; TEXAS:
Cass; Collin; Potter.
NATURAL HISTORY
Nests of F. biophilica have simple, cryptic openings in wetlands, grasslands or less often, in
open woodlands. The entrance is usually hidden amongst grass or sedges. In springtime,
colonies of F. biophilica may build a 10-25 cm diameter mound of soil and plant fragments
nestled against a grass or sedge clump, this collapsing in disuse during the hot, dry weather of
summer. In fens, bogs and wet meadows, when ground at the base is permanently or seasonally
saturated, F. biophilica nests in the upper parts of graminoid tussocks. One colony under a strip
of bark in unmowed grass in eastern Missouri contained four larvae of myrmecophilous
staphylinid beetles, probably Xenodusa cava LeConte (but not collected for determination).
This species has been found as host to the slavemaker Polyergus lucidus s. l. in Washington,
D.C., northern Georgia and east-central Missouri. The variety of this slavemaker parasitizing F.
biophilica has longer scapes and is somewhat less shiny and slightly more pubescent than typical
P. lucidus lucidus Mayr, which parasitizes F. incerta. F. biophilica occurs among the many
hosts of F. pergandei in the prairies of Missouri, but has only been observed in combination
with other host species. At one site, a F. pergandei nest contained a mélange of six slave species
including (in order of decreasing relative abundance) F. pallidefulva, F subsericea Say, F.
biophilica, F. dolosa, F. incerta and F. obscuriventris Mayr, certainly the most species-rich,
naturally occurring ant colony on record!
Sexuals have been collected in nests in Missouri, Texas and Georgia in mid-June to early
July, but no flight or colony-founding activity has been recorded. There is one example of a
queen-male bilateral gynandromorph in a Missouri collection. The worker pupae are always
enclosed in a pale tan cocoon, and the sexuals in a larger, darker cocoon.
Formica dolosa Buren, 1944 stat. nov.

Figures 1, 4e, 5e, 6e
Formica pallidefulva subsp. schaufussi var. meridionalis Wheeler, W. M. 1904: 370 [Unavailable
name.]
Formica pallidefulva subsp. schaufussi var. dolosa Wheeler, W. M. 1912: 90 [Unnecessary
replacement name for meridionalis; also unavailable.]
Formica pallidefulva subsp. schaufussi: Wheeler, W. M. 1913b: 552 (in part) [Misidentification.]
Formica pallidefulva subsp. schaufussi var. dolosa: Wheeler, W. M. 1913b: 554
Formica (Neoformica) schaufussi subsp. dolosa Buren, 1944: 309. [First available use of dolosa.]
Syntype workers, Bull Creek, Travis Co., Texas (W. M. Wheeler) (MCZ) [Examined. Three
workers on one pin, labeled “true types of dolosa” by S. Cover, and two gynes on one pin labeled
syntypes by S. Cover ]
Formica pallidefulva subsp. schaufussi: Emery, 1893: 654 [Misidentification.]
Formica schaufussi: Creighton, 1950: 551 [Misidentification.]
Formica schaufussi subsp. dolosa: Creighton, 1950: 551
Formica schaufussi: Robson & Traniello, 1998: (in part) [Vouchers examined.]
NOTE: We have selected a specimen in the Mayr collection (NMW) labeled “Nord
Amerika / Schaufuss” as lectotype of Formica schaufussi Mayr, as this corresponds to the
locality and collector information in Mayr’s (1866) description. This sample clearly belongs to
the much less pilose Formica pallidefulva. Thus, the name Formica schaufussi Mayr falls to the
synonymy of Formica pallidefulva, below.
DIAGNOSIS
Worker – The largest, most pilose, most densely pubescent and least shiny of reddish-
yellow members of the pallidefulva group (F. archboldi is duller, but always much darker and
averages smaller). Weakly bicolored; head, mesosoma and legs light coppery red (south) to
yellowish or reddish brown (north); gaster a little darker than head and mesosoma. Dorsal
sclerites of mesosoma with abundant erect pilosity (Fig. 6e); erect macrochaetae on gaster
abundant and long (longest macrochaetae 0.16-0.30 mm), straight to slightly curved. Mesosoma,
especially propodeal dorsum, pubescent; gaster dulled by long, dense, pale grayish, appressed
microchaetae (Fig. 4e). Gaster with small shallow foveolae in some samples, these nearly
lacking in others. The propodeal crest is nearly always rounded in F. dolosa. The larger workers
of this species are the largest eastern US Formica, matched within the genus only by the
allopatric and otherwise quite different F. ravida Creighton.
Queen – Color, gastral pubescence, abundant pilosity and lack of shininess like the
workers’, with the usual differences in size. Sculpture a little more accented with notable fine
tessellation of entire head, mesosoma and gastral dorsum; wings, when present, clear brownish
to dark smoky gray. Three mesoscutal spots present as in F. incerta, but these pale and diffuse.
Male – Pubescence dense and pilosity abundant; surface sculpture punctate; head and gaster
dark brown, mesosoma reddish brown to dark reddish brown with legs the same color; wings
dark smoky gray. Larger than the nearly similar F. incerta, in which the mesosoma is normally
about the same color as the head and gaster.
The propodeal crest of F. dolosa is nearly always rounded in profile, and is typically sharp
or even carinulate in the other species. This large, hairy, densely pubescent and faintly bicolored
ant is most likely to be confused with F. biophilica. Compared to F. biophilica, F. dolosa has
conspicuous appressed pubescence on the mesosoma, has more abundant, but slightly shorter
gastral pilosity (longest macrochaetae up to 0.30 mm), has longer, denser pubescence on the
gaster (compare Fig. 4b and 4e), and averages larger and heavier-bodied. The number of
macrochaetae on the pronotum usually exceeds that on the propodeum of F. dolosa, (46 of 54
specimens) whereas the number on the propodeum more often exceeds that on the pronotum of
F. biophilica (20 of 32 specimens). F. dolosa usually has relatively smaller eyes compared to F.
biophilica (Table 1). In the field, F. dolosa occupies the drier end of the habitat spectrum, the
two overlapping mainly in pine-oak woodlands of the Southeastern U.S., and in dry-mesic
prairies further north. In the Northeastern U.S., larger, more pilose workers of F. incerta are
often misidentified as F. dolosa, but F. dolosa averages larger and more pilose, has mesosomal
pubescence and denser gastral pubescence, has longer scapes and legs; is generally lighter, more
yellowish or reddish in color, and is more strictly associated with highly drained soils.
ETYMOLOGY
This name comes from the Latin adjective dolosus, meaning cunning or sly. Perhaps
Wheeler was referring to the fleetness of its escape when alarmed, as this species is very shy and
an excellent “escape artist”.
RANGE AND HABITAT
Widely distributed from New England across the Great Lakes region, west to Wisconsin and
Iowa and south to northern Florida, the Gulf Coast states and Texas. Records of this ant in
Colorado by Gregg are all misidentified F. incerta (L. Rericha, personal communication). F.
dolosa is decidedly most abundant on acid-soil sites. These include a variety of droughty or
well-drained habitats such as barrens, glades, prairies or open oak or pine woodlands on
silicaceous or loessic soils. Though reported (as schaufussi) from plowed fields and pastures in
the Northeast, F. dolosa is not usually common in such communities. J. Trager found F. dolosa
in calcareous glades in Alabama and Missouri, but it is not abundant in these sites. In stark
contrast to F. incerta and F. biophilica, F. dolosa does not nest in mesic habitats or in moist,
fertile soils.
SPECIMENS EXAMINED
ALABAMA: Lawrence; ARKANSAS: Logan; FLORIDA: Alachua; Bay; Columbia;

Escambia; Gilchrist; Jackson; Jefferson; Lake; Leon; Liberty; Okaloosa; Santa Rosa; Suwannee;
Walton; GEORGIA: Clarke; Lumpkin; ILLINOIS: Mason; MARYLAND: Allegany;
Dorchester; MASSACHUSETTS: Plymouth; Worchester MISSISSIPPI: Chickasaw; Choctaw;

Lafayette; Lee; Lowndes; Noxubee; Oktibbeha; Pontotoc; Scott; Tishomingo; Winston;
MISSOURI: Franklin; Johnson; Lincoln; Washington; NEW JERSEY: Ocean; NEW YORK:
Nassau; Suffolk; NORTH CAROLINA: Nash OHIO: Adams; SOUTH CAROLINA: Aiken;
Barnwell; McCormick; Oconee; TEXAS: Travis; WISCONSIN: Adams; Crawford; Dane;
Grant; Iowa; Marshall; Sauk; Walworth; Waukesha.
NATURAL HISTORY
Nests may be hidden beneath a rock or piece of wood, but most nest entrances are at the
base of a grass clump or other herbaceous plant. Some open onto bare ground, the entrance
surrounded by a crater of excavated soil adorned with plant fragments, charcoal bits or fine
gravel. J. MacGown collected F. dolosa in nests at the bases of large trees on relatively drier and
more open ridges in mixed forests in northern Mississippi, and from an infrequently mowed area
under loblolly pines near his house in Oktibbeha Co. Mississippi. The nest at the latter site was
a low mound about 45 cm across and about 15 cm high at the midpoint. Part of the mound was
inhabited by Camponotus castaneus Latreille.
In the East and Gulf Coast United States, F. dolosa is host to the slavemaker Polyergus
lucidus longicornis M. R. Smith. J. Trager’s collection contains samples of this slavemaker with
F. dolosa slaves from Massachusetts, New York, New Jersey, South Carolina and Mississippi.
In Missouri, F. dolosa is occasionally among the many hosts of F. pergandei, but we have only
observed them in combination with other host species (see “Natural History” of F. biophilica for
a case in point). In Florida, J. Trager observed F. dolosa and F. archboldi competing for
domination of colonies of Toumeyella scales on long-leaf pine “grass-stage” seedlings.
Occasionally, fights would arise in which the larger F. dolosa threw or chased F. archboldi
workers to the ground.
Winged sexuals were collected in nests in mid-June in Florida and Georgia, and one male
was found in a nest in western Missouri in August. Both worker and sexual pupae are always
enclosed in a cocoon.
Formica incerta Buren stat. rev., stat. nov.

Figures. 4c, 5c, 6c
Formica pallidefulva subsp. schaufussi var. incerta Emery, 1893 [Unavailable name]
Formica (Neoformica) pallidefulva subsp. schaufussi var. incerta: Wheeler, 1913b
Formica (Neoformica) pallidefulva subsp. incerta: Buren, 1944 [First available use of incerta]
Syntype workers, District of Columbia, iv-13-1886 (MCSN) [Examined. Five workers on three
pins labeled paratypes by A. Francoeur]
Formica (Neoformica) pallidefulva: Creighton, 1950, in part
Formica (Neoformica) schaufussi: Creighton, 1950, in part
Formica schaufussi: Robson & Traniello, 1998, in part [Vouchers examined] (JCT)
DIAGNOSIS
Worker – A relatively shiny grassland Formica with a relatively broad head (mean CI =
86.77), sides of head more convex (Fig. 5c) and scapes relatively short (mean SI = 132.99).
Head and gaster rich, dark brown (northeast) to brownish-yellow with darker tip (prairie region).
Mesosoma and legs yellowish-brown to light yellowish-brown. Mesosoma often a little lighter
than head, and both lighter than gaster. Specimens in the Great Plains portion of the range are
nearly concolorous brownish yellow except for the darker gastral apex. Mesosomal
macrochaetae of F. incerta typically conspicuously shortest on propodeum. Erect pilosity on
gaster relatively short, straight or only slightly curved, if curved, usually below the mid-point of
the length of the macrochaetae. Gaster shiny, but its sheen dulled by faint tessellation and
medium density pubescence (Fig. 4c) composed of pale grayish appressed microchaetae
differences in size. Color pattern differing from workers’ and from that of queens of all other
species in that there are three distinct, dark spots on the mesoscutum, one anteromedian and two
lateral over the parapsidal sulci. These may cover most of the mesoscutal area or may be reduced
to longitudinal dark elliptical marks. Upper portion of head, pronotum, sides of mesothorax,
propodeum and gastral dorsum with faint tessellation. Wings, when present, clear brownish to
clear smoky gray.
Male – Pubescence and pilosity abundant; mesosomal dorsum dull-punctate; entire body
uniform black or dull blackish brown, legs reddish brown; wings clear brownish to clear smoky
gray. Averages smaller than the nearly similar F. dolosa and smaller and of more uniform
blackish color than males of F. biophilica.
Metrically, F. incerta is distinguished from the other species in the group by a relatively
broad head and short scapes (CI and SI, Table 1). In the northeast part of its range, more pilose
F. incerta individuals may be confused with F. dolosa, and F. incerta specimens with little
pilosity may be confused with F. pallidefulva. The geographic range of F. pallidefulva
completely overlaps that of F. incerta, and most places where they are found together in the
field, F. incerta appears lighter in color and less shiny than F. pallidefulva, due to some faint
tessellation on the mesosoma and somewhat longer, denser pubescence on the gastral dorsum of
F. incerta. Mesosomal and gastral pilosity is usually much less abundant than in F. dolosa and
averages slightly less abundant than in F. biophilica. Also, F. incerta is darker and shinier than
sympatric F. dolosa. See F. biophilica account for the differences between F. incerta and that
species.
ETYMOLOGY
This name was coined by Emery from the Latin adjective incertus meaning uncertain. This
seems appropriate to describe Emery’s own and subsequent authors’ doubts regarding the
validity of this species.
RANGE AND HABITAT
This species occurs from New England and the Great Lakes States west to Minnesota,
Nebraska and low elevation grasslands of Colorado (and New Mexico?). It extends south in
eastern US to the balds, meadows and old fields of the southern Appalachians. F. incerta is
especially abundant in native mesic and dry-mesic grasslands, but also occurs in parks,
campuses and lawns, fields and forest clear-cuts. In the Northeast, it occurs in heathland and
sand barrens, and in the Midwest it is characteristic and abundant in prairie remnants, botanically
diverse old fields and meadows, and native prairie reconstructions.
SPECIMENS EXAMINED
(CANADA) ONTARIO: Lambton. (UNITED STATES) CONNECTICUT: Litchfield;

DELAWARE: Kent; DISTRICT OF COLUMBIA: Washington; ILLINOIS: DuPage; Madison;
IOWA: Dubuque; Johnson; Winneshiek; KENTUCKY: Laurel; MAINE: Androscoggan;
Cumberland; Kennebeck; MARYLAND: Allegany; Baltimore; MASSACHUSETTS: Essex;
Worchester; MICHIGAN: Livingston; MINNESOTA: Crow Wing; MISSOURI: Audrain;
Boone; Callaway; Franklin; Harrison; Jasper; Madison; St. Louis; NEBRASKA: Hall; NEW
JERSEY: Burlington; Essex; Gloucester; Salem; NEW YORK: Rockland; Suffolk; NORTH
CAROLINA: Cocke; Haywood; OHIO: Adams; Butler; Champaign; Delaware; Hamilton;
Hocking; Jackson; Montgomery; PENNSYLVANIA: Chester; Delaware; VIRGINIA: Farifax;
Rapahannock; Washington; WISCONSIN: Crawford; Washburn; Waushara.
NATURAL HISTORY
Nests are in bare soil, or beneath a grass clump, in the latter case often with a small,
irregular, conical (5-15 cm wide, 10-20 cm tall) mound of soil and plant fragments. This is often
the first Formica species to become abundant on restored native grasslands, “Conservation
Reserve Program” grassland plantings on former farmland and cut-over forests. A healthy
population of F. incerta may facilitate colonization by its parasites F. difficilis and F. pergandei,
if these occur nearby. It is less abundant than F. pallidefulva in lawns, campuses and parks.
This is often the most abundant Formica species in mesic tallgrass prairies from central
Illinois, Nebraska and south to Oklahoma and northeast Arkansas, and also in balds, meadows
and old fields at higher elevations of the southern Appalachian Mountains. Sweepnet samples
from all these types of habitats rarely fail to include F. incerta, and thus insect collections
housed at institutions near them may be rife with samples of individuals so captured. In the
Great Lakes Region and New England, this species is more associated with sandy soils and
pastureland and often nests under rocks. In the northern glaciated prairie region, F. incerta can
be a dominant ant in sand prairies, but is largely displaced from sites with moister, finer-textured
soils, which are dominated by aggressive, mound-building Formica species.
F. incerta appears to be the only host of Polyergus lucidus lucidus collections examined
from New England states, New Jersey, southern Ontario, Wisconsin and Missouri. F. incerta is
also frequent among the many hosts of F. pergandei and in western Missouri prairies commonly
occurs as a slave of this species, either alone or in mixed populations with F. subsericea. F.
incerta appears to be the primary host of alloparasitic (dispersing) queens of F. difficilis Emery,
the queen of which bears a superficial resemblance to F. incerta workers. Indeed, Wheeler
(1904) first used the term “temporary social parasitism” to describe the relationship he
elucidated between F. difficilis (as var. consocians) and F. incerta in Connecticut.
This ant often visits extrafloral nectaries of sunflowers, partridge peas and other prairie
plants. It also tends aphids and membracids on a variety of plants. F. incerta workers defend
these sugar sources from non-nest mates of their own species, from other, smaller ant species
and from some parasitoids. However, in areas where there are greater numbers of aggressive
mound-building prairie Formica species (e.g. F. montana Wheeler, F. obscuripes Forel), F.
incerta becomes more furtive and opportunistic in its honeydew gathering, as described below
for F. pallidefulva. Foraging strategy and recruitment to food sources has been well studied in F.
incerta (and incidentally, in F. dolosa) by Robson & Traniello (1998, and included references to
their earlier work). These authors identified their study subject as F. schaufussi in the articles,
but vouchers sent by Robson were examined for this revision. These were mostly F. incerta, but
also included a sample of F. dolosa.
In grasslands, especially those recently burned, northern flickers (Colaptes auratus) prey
heavily on Formica species, including F. incerta. This is especially so on sunny, late winter days
when workers migrate intranidally toward the surface, seeking warmth.
Alates occur in the nests in July and August in New England and the northern prairies, and a
few weeks earlier in the unglaciated prairie region and southern Appalachians. It is worth noting
that the maturation of alates of F. pallidefulva may precede that of F. incerta in by two or three
weeks, suggesting a possible temporal mechanism for reproductive isolation. Flights have not
been observed, but several mated queens have been captured walking about in mid to late
morning in Missouri. In the lab, these recently mated queens are “nervous” in captivity and often
fail to rear their first workers, in contrast to the ready adaptability to captive conditions of F.
pallidefulva queens. Worker pupae are typically enclosed in a light tan cocoon and sexual pupae
have darker tan cocoons. This is in contrast to the frequently naked worker pupae of F.
pallidefulva, as was earlier noted by both Wheeler (1904) and Talbot (1948).
Formica pallidefulva Latreille

Figures 4a, 5a, 6a
Formica pallide-fulva Latreille, 1802 [Types not seen, supposedly type-compared material in MCZ
examined]
Formica schaufussi Mayr, 1886 [Lectotype designated, labelled “N. Amer. / Schauf." (NHWC)] Syn.
nov.
Formica pallidefulva subsp. nitidiventris Emery, 1893 [Type examined.] “310 B Wokland(?). D.C.
with Polyergus lucidus. Paratype. Formica nitidiventris A. F.1968] Syn. nov.
Formica pallidefulva subsp. fuscata Emery, 1893. [Lectotype examined.] “Beatty PA. No. 314.
LECTOTYPE Formica pallidefulva fuscata A. F. 1968. Synonymy, under nitidiventris, by
Creighton, 1950: 551.
Formica pallidefulva var. succinea Wheeler, W. M. 1904 [Syntypes examined] Four workers on one
pin, Bee Creek. Travis Co. TEX. XI.9.02 M.C.Z. Type 5-8 8844 var. succinea Wheeler.
Synonymy by Creighton, 1950: 550.
Formica (Neoformica) pallidefulva: Wheeler, W. M., 1913 b [Vouchers examined] (MCZ)
Formica (Neoformica) pallidefulva subsp. delicata Cole, 1938 [Syntypes examined] 24 workers on 8
pins. Ten Sleep WY 9/31 A.C. Cole. Synonymy, under nitidiventris, by Creighton, 1950: 551.
Formica (Neoformica) pallidefulva: Creighton, 1950, in part [Vouchers examined] (MCZ, LACM)
Formica (Neoformica) pallidefulva subsp. nitidiventris: Creighton, 1950, in part [Vouchers examined]
(MCZ, LACM)
DIAGNOSIS
Worker – Includes conventional reddish or brownish yellow F. pallidefulva, as well as

darker populations known as F. p. nitidiventris and its synonyms F. p. fuscata and F. p. delicata.
This is the shiniest Formica of this group (though smaller workers of F. biophilica and some
series of F. incerta are also quite shiny). The mesosoma often lacks either appressed pubescence
or erect pilosity, or has relatively few, short, erect macrochaetae (Fig. 6a). Pubescence, even on
gaster, short and sparse (Fig. 4a). Sculpture faint to nearly smooth, best developed (to the point
of slightly weakening the sheen) in the northeastern part of the range, where the form fuscata
occurs. The gaster appears more voluminous than in other members of the group, and is quite
shiny, as reflected in the name nitidiventris. Color is highly variable, generally uniform dark
brown in Canada and New England, the Black Hills and western mountain areas, and
concolorous coppery yellow or weakly bicolored (gaster a little browner) in the deep South.
Various intermediate conditions occur in a broad band of territory from southern Missouri and
northern Arkansas, across the upper South to the foothills of the southern Appalachians in
Georgia and the Carolinas, and occasionally elsewhere. The transition area between typical
pallidefulva and typical nitidiventris is a 300-mile wide band straddling the Mason-Dixon Line.
In it, one may occasionally find single-queen colonies containing nearly the full range of color
variation. The extreme color forms are weakly distinguished morphometrically, with far northern
populations having slightly shorter scapes, but the variation is clinal through the zone of
transition (mean SI = 143.08 in the South, 140.94 in transition zone, 139.68 in the North).
differences in size. Sculpture very faint; pubescence short and sparse; pilosity sparse; wings,
when present, clear to amber. Mesoscutum lacking the three dark spots characteristic of incerta
queens, or these weakly distinct.
Male – Pubescence sparse; surface more shining than other species; rarely concolorous dark
brown (in those colonies with the most uniformly dark workers), most commonly in north and
Rocky Mountains and clinal transition zone with head and gaster blackish and mesosoma a little
lighter to clear yellowish brown; in south and Great Plains, concolorous honey-red or with only
the head notably darker; wings clear to amber-colored.
Most, if not all records of pallidefulva within the northern part of its range, where the color
form “nitidiventris” occurs, are usually F. biophilica or occasionally are lighter color morphs of
F. incerta. The short, sparse, gastral pubescence, lack or sparseness of pilosity on the mesosomal
dorsum, and the short, straight and flattened gastral pilosity of F. pallidefulva is distinctive for
this species in any of its color variants. The difference in SI strongly discriminates this species
from F. incerta (Table 1). The setal characteristics, the shininess and “globulous” gaster of this
species were expressly mentioned by Latreille (1802) in his original Latin description. Among
the species in the group, F. pallidefulva is the only one to frequently lack detectable
macrochaetae on the pronotum (34 of 57 specimens) and propodeum (33 of 57 specimens).
When mesosomal dorsal pilosity is present in F. pallidefulva, the macrochaetae average shorter
than in other species and are usually most numerous on the mesonotum rather than on the
pronotum or on the propodeum (Table 1). Bright-colored southern F. pallidefulva may be
distinguished from less pilose nanitic and small workers of F. biophilica by the nearly perfectly
rounded propodeum and straight, flattened gastral macrochaetae of F. pallidefulva (versus often
faintly right-angular propodeum and narrowly curviconical gastral macrochaetae in F.
biophilica).
ETYMOLOGY
This name was coined by Latreille from the Latin adjectives “pallidus” plus “fulvus”
meaning pale reddish yellow. This neatly describes the southern, lighter colored variants of this
species. Northeastern, Midwestern and western mountain populations of this species are
predominantly of darker, black-coffee-brown coloration, but even in these locations many
individuals and colonies are bicolored and some may have coloring closer to that of southern
populations.
RANGE AND HABITAT
Abundant and certainly the most widely distributed species of the group, F. pallidefulva
occurs farther north, west and south than others in the group, except that F. archboldi perhaps
extends farther south in Florida. F. pallidefulva occurs throughout the eastern United States and
southeastern Canada, then west across the US Great Plains to the lower-elevation Rocky
Mountains from Wyoming to New Mexico. F. pallidefulva also has considerable habitat latitude.
This ant lives in a variety of native and anthropogenic plant communities and soil types,
including dry-mesic to mesic grasslands, woodlands and forests, thickets, lawns, campuses and
parks. It is most abundant in mesic, wooded or partially wooded areas, from city parks to closed-
canopy forests. In the lower rainfall areas of the Great Plains, it is uncommon and probably
restricted to riparian woodlands (Milford, 1999). In the Rocky Mountains, it occurs at lower
elevations in meadows, mixed mesophytic forests and in parks and suburbs. F. pallidefulva does
not occur in bogs, wet meadows or fens, where it is replaced, in the South by F. biophilica and
in the North by other Formica species outside the pallidefulva-group such as F. montana, F.
glacialis Wheeler, and others.
SPECIMENS EXAMINED
(CANADA) ONTARIO: Lambton; Lowick; QUEBEC: Chateauguay. (UNITED STATES)

ALABAMA: Baldwin; Bibb; DeKalb; Lawrence; Morgan; Tuscaloosa; ARKANSAS: Logan;
Washington; COLORADO: Boulder; DELAWARE: Kent; FLORIDA: Alachua; Brevard; Clay;
Highlands; Liberty; GEORGIA: Clarke; Floyd; ILLINOIS: Tazewell; IOWA: Emmet; Johnson;
Winneshiek; KANSAS: Douglas; Reno; Wallace; KENTUCKY: Nelson; MARYLAND:
Allegany; Anne Arundel; Kent; Prince Georges; MASSACHUSETTS: Barnstable; MICHIGAN:
Calhoun; Livingston; MISSISSIPPI: Alcorn; Chickasaw; Itawamba; Lafayette; Lee; Lowndes;
Oktibbeha; Panola; Pontotoc; Tishomingo; Webster; Winston; MISSOURI: Audrain; Franklin;
Johnson; Lincoln; Ste. Genevieve; St. Louis; NEBRASKA: Hall; Dawson; NEW MEXICO:
Colfax; Otero; Rio Arriba; Union; NEW YORK: Orange; Suffolk; NORTH CAROLINA:
Yancey; OHIO: Franklin; Sandusky; OKLAHOMA: Latimer; PENNSYLVANIA: Alegheny;
Indiana; SOUTH CAROLINA: Anderson; Charleston; SOUTH DAKOTA: Jones; Pennington;
TEXAS: Bastrop; Cass; Culberson; Hemphill; VIRGINIA: Montgomery; WYOMING: Crook;
Washakie.
NATURAL HISTORY
Nests of F. pallidefulva may be located in bare soil of grassland and forest footpaths,
beneath leaf litter, under small diameter (<10 cm) fallen tree limbs, or under bark of a
decomposing stump. Less often the nest occupies a larger, punky, rotten log, especially during
late spring when the sexual brood is being reared. F. pallidefulva is the only species in this group
which normally inhabits closed-canopy mesic forests and which commonly nests in rotting
wood. In non-wooded settings F. pallidefulva may build a small mound nestled in or beside a
grass clump.
In the northern and Rocky Mountain parts of its range, at least, the dark brown form of F.
pallidefulva is the host to the slavemaker Polyergus lucidus montivagus Wheeler. We have seen
F. pallidefulva with this slavemaker in colonies from Long Island, southern Ontario, central
Illinois, northern Missouri and Rocky Mountain foothill locales in Colorado and New Mexico.
We have seen specimens that look like this Polyergus in the South (northern Mississippi) and we
suspect it uses the reddish, southern form of F. pallidefulva there, but have not yet been able to
confirm this with a nest collection. Formica creightoni raids this ant in northern Missouri oak
woodlands, where its usual F. neogagates group hosts are lacking. F. pallidefulva is the most
frequent of the many hosts of F. pergandei in Midwest woodlands and savannas.
In the lawns and gardens of the St. Louis, Missouri, area, F. pallidefulva is among the native
ants most sensitive to subterranean invasion and extermination by the introduced and rapidly
spreading invasive ant Tetramorium tsushimae (Steiner et al., 2006). However, throughout much
of the Southeast, F. pallidefulva often manages to coexist with low-density Solenopsis invicta
Buren, Solenopsis richteri Forel, and Solenopsis richteri × invicta populations.
Occasionally, workers and sexuals become covered with a mite (Oplitis sp.?) that reduces
their energy level and may cause the demise of the colony. In eastern Missouri, F. pallidefulva
colonies are commonly raided by the slavemaker/ant-predator F. rubicunda Emery, which uses
F. pallidefulva as prey only (F. rubicunda “enslaves” only F. subsericea). When nesting in
rotting wood, F. pallidefulva colonies may sometimes be pillaged by pileated woodpeckers. F.
pallidefulva often gathers honeydew beneath hemipteran-infested plants or from their leaf
surfaces, but they have not often been observed to gather honeydew directly from the
hemipterans. Furthermore, F. pallidefulva makes little effort to defend hemipterans, in contrast
to F. incerta and F. archboldi, which often tend and defend them.
Sexuals are present in the nests as early as April in Florida, but not until July in New
England and Canada. Males were observed to gather around the nest entrance around sunset in
Florida, and males are often lured to lights (but winged females are not) throughout the range. In
Missouri, J. Trager observed males, followed by females, flying from the nest, one by one,
shortly after sunrise. Despite rather frequent capture of males at lights after dusk, the actual
mating flight period is in the morning. In eastern Missouri, J. Trager has over the years caught
numerous recently mated females walking about in late morning or early afternoon. Dates of
these captures occur from 26 June into early July.
Unlike other species of this group, worker pupae most often lack cocoons in F. pallidefulva.
Wheeler (1904) first noted this contrast between F. pallidefulva and F. incerta in Connecticut,
and it was also later noted by Talbot (1948) in Michigan. This difference holds true in numerous
colonies of the two species sampled in Missouri by J. Trager. In some colonies of F.
pallidefulva, male pupae may also lack cocoons, but queen pupae nearly always are enclosed in
cocoons.
Morphometric analysis of species
Discriminant function analysis classified 97.3 percent of the specimens analyzed (177 of
182) consistently with their morphological species designations. The low rate of error
demonstrates that the species do comprise distinctive groups of individuals that can be
successfully identified using the morphological characteristics measured in this study. The five
species in the F. pallidefulva group can be distinguished from one another by a number of
morphological indices (Table 1) and body and hair measurements, summarized in Table 2.
Several body measurements distinguish among the species in this group, including species
pairs that are visually quite similar. The species differ significantly in the shape of the head and
size of the eye, as indicated by the significant pairwise differences among species for CI and OI
(Table 1). In particular, on average F. incerta has a proportionally broader head and F.
archboldi has proportionally larger eyes than the other species. As mentioned in the species
description above, F. biophilica is morphologically quite similar to F. incerta but these two
species are often differentiated from one another by the longer mesosoma and proportionally
narrower head of the former often combined with subtle differences in the number and
distribution of macrochaetae (Fig. 2). F. biophilica and F. pallidefulva also have a significantly
higher scape index than F. incerta (Table 1), as indicated by the proportionally longer scapes of
these two species compared with head width (Fig. 3).
Key to species (based mainly on worker)
1a Mesosomal dorsum without erect setae or with a small cluster on the mesonotum and/or a
few erect hairs elsewhere; gaster shiny with sparse, short appressed pubescence and relative
short, sparse, blunt pilosity (Figures 4a, 6a); color highly variable, ranging from bright
tawny or coppery yellow (southern United States) to very dark brown (mountains in
northern and western United States) and including many intermediate colorations; mesic
habitats, from gardens, parks and prairies to closed-canopy forests, or more shaded parts of
drier sites............................................................................................................... pallidefulva
b Mesosomal dorsum with several to many erect setae on pronotum, mesonotum and
propodeum; gastral tergite I shiny to dull with appressed pubescence of medium to high
density (Figures 4b-e, 4b-e); the average distance between the individual appressed setae
approximately equaling the average length of setae to much less; various open grassland,
heath, or barrens and open dry woodland habitats (mesic forests or forest edge in the
extreme southern U.S.) .......................................................................................................... 2
2a Mesosomal and gastral integument dulled by fine, but notable sculpture; mesosomal dorsum
feebly or (at some viewing angles) not shining; gaster usually with dense pubescence
(Figures 4d-e, 6d-e), appressed setae on gastral tergite I separated by about 0.5× (or less) the
average setal length (if gastral setae less abundant, then mesosomal and gastral integument
dulled by fine sculpture); propodeum usually with visible pubescence................................. 3
b Mesosomal and gastral integument somewhat shining, with fainter sculpture; gaster less
densely pubescent (Figures 4b-c, 6b-c); gastral tergite I with appressed setae separated by
about 0.5-1.0× their average length; propodeum with sparse pubescence or none................ 4
3a Reddish, often weakly bicolored; mesosoma with visible pubescence and gastral integument
dulled or even velvety looking due to dense pubescence; larger workers’ integument
appearing matte in dorsal or oblique dorsal view; numerous erect macrochaetae on all dorsal
surfaces, those on first tergite usually long, tapering and typically curved (Figure 6e); size
generally larger, HL 1.31-2.06, HW 1.05-1.70, and WL 2.12-3.48; eyes proportionally
smaller to head size (OI 28.58); widely distributed in eastern United States (rare west of
100th meridian); reddish-brown (northeastern United States) to light reddish (southeastern
and midwestern United States) ..................................................................................... dolosa
b Dark brown to blackish brown; with fine coriaceous sculpture on mesosoma and foveolae
on gastral dorsal surface; erect setae shorter and fewer, at least on mesosoma, erect setae
often blunt-tipped and without notable curvature, even on gastral tergite I (Figure 6d); size
smaller, HL 1.31-1.58, HW 1.03-1.39, and WL 2.08-2.65; eyes proportionally larger (OI
31.26); found in long-leaf pine sandhills and scrub or sand pine woodlands in southeastern
United States (Alabama, Florida, and Georgia) ........................................................archboldi
4a Length of the longest macrochaetae of gastral tergite I, 0.15-0.22 mm (Figure 4c, 6c, Table
2); color dingy yellowish or reddish brown with gaster or head and gaster a little to
significantly darker (midwestern United States, Great Plains) or more uniformly brown with
only the mesosoma a bit more yellow (northeastern United States); overall size averages
smaller (HL 1.15-1.64, HW 1.01-1.43, and WL 1.86-2.67); head wider and scapes shorter
(See CI and OI, Table 1, also Figures 2 & 3); queen with three distinct dark spots on
mesoscutum, one anteromedian and two lateral over parapsidal sulci (spots may be reduced
to longitudinal dark elliptical marks)............................................................................ incerta
b Erect setae of first gastral tergite long and curved, longest macrochaetae 0.20 – 0.30 mm
(Figure 4b, 6b, Table 2); color bright reddish-yellow or with gaster only slightly darker
(Beware: Gaster of this species sometimes stains black in mounted specimens.); mesosomal
dorsal integument a little more shining to quite smooth except in largest workers; overall
size averages larger (HL 1.37-1.82, HW 1.05-1.56, and WL 2.22-2.93); head more narrow
in smaller specimens and scapes longer (See CI and OI, Table 1, also Figures 2 and 3) in
smaller specimens (larger workers tend to have wider heads and shorter scapes); queen
lacking dark spots on mesoscutum .......................................................................... biophilica
ACKNOWLEDGEMENTS
We would like to thank the collection curators mentioned in the methods section for their careful
attention to requests for loans of specimens. Caitlin Argyros entered the hand-written
measurement data into a spreadsheet, carefully double-checking her work to the point of
perfection. Laura Rericha of the Cook County (Illinois) Forest Preserve District checked the
taxonomic scheme and key against her specimens collected in the Chicago Region, and most
kindly checked the identity of every sample of the Formica pallidefulva group in the Robert E.
Gregg collection at the Chicago Field Museum, helping confirm the western boundaries of
species distributions. J. Trager has benefited greatly over the years from conversations and
correspondence with André Francoeur and Stefan Cover regarding these and other Formica
species. The USDA-ARS Areawide Management of Imported Fire Ant Project supports Joe
MacGown’s work on ants, especially those of Mississippi.
LITERATURE CITED
Abouheif, E. & Wray, G. A. 2002. Evolution of the gene network underlying wing
polyphenism in Ants. Science 297: 249-252.
Bale, M.T., Zettler, J.A., Robinson, B.A., Spira, T.P. & Allen, C.R. 2004 Yellow jackets may
be an underestimated component of an ant-seed mutualism. Southeastern Naturalist 2: 609-
614.
Bell, D.A. 1996. Genetic differentiation, geographic variation and hybridization in gulls of the
Larus glaucescens-occidentalis complex. Condor 98: 527-546.
Burbrink, F.T. 2001. Systematics of the eastern ratsnake complex (Elaphe obsoleta).
Herpetological Monographs 15: 1-53.
Buren, W.F. 1944. A list of Iowa ants. Iowa State College Journal of Science 18: 277–312.
Buren, W.F. 1968. Some fundamental taxonomic problems in Formica (Hymenoptera:
Formicidae). Journal of the Georgia Entomological Society 3: 25–40.
Cole, A.C. 1938. Descriptions of new ants from the western United States. American Midland
Collingwood, C.A. 1962. Some ants (Hym. Formicidae) from north-east Asia. Entomologisk
Tidskrift 83: 215-230.
Zoology at Harvard College 104: 1-585.
Emery, C. 1893. Beiträge zur Kenntnis der nordamerikanischen Ameisenfauna. Zoologische
Jahrbücher. Abteilung für Systematik, Geographie und Biologie der. Tiere. 7: 633-682.
Floate, K.D. & Whitham, T.G. 1995. Insects as traits in plant systematics—their use in
discriminating between hybrid cottonwoods. Canadian Journal of Botany 73: 1-13.
Green, D.M., Sharbel, T.F., Kearlsley, J. & Kaiser, H. 1996. Postglacial range fluctuation,
genetic subdivision and speciation in the western North American spotted frog complex,
Rana pretiosa. Evolution 50: 374-390.
Latreille, P.A. 1802. “Formicoidea” in: Histoire naturelle, generale et particuliere des
crustaces et des insectes. Vol. 3. F. Dufart, Paris. 467 pp.
Mayr, G. 1866. Myrmecologische Beiträge. Sitzungsberichte der K. Akademie der
Wissenshaften. Mathematisch.- Naturwissenschaftliche Classe 53: 484-517.
Milford, E.R. 1999. Ant communities in flooded and unflooded riparian forests of the middle
Rio Grande. Southwestern Naturalist 44: 278-286.
Mikheyev, A.S. & Tschinkel, W.R. 2000. Nest architecture of the ant Formica pallidefulva:
structure, costs and rules of excavation. Insectes Sociaux 50: 30-36.
Robson, S.K. & Traniello, J.F.A. 1998. Resource assessment, recruitment behavior, and
organization of cooperative prey retrieval in the ant Formica schaufussi (Hymenoptera:
Formicidae). Journal of Insect Behavior 11: 1-22.
Smith, D.R. 1979. Superfamily Formicoidea [pp. 1323-1467]. In K.V. Krombein, P.D. Hurd,
Jr., D.R. Smith, and B.D. Burks (Eds.). Catalog of Hymenoptera in America North of
Mexico, Vol. 2: Apocrita (Aculeata). Smithsonian Institution Press, Washington D.C.
xvi+1199-2209.
Smith, M.R. 1944. Additional ants recorded from Florida, with descriptions of two new
subspecies. Florida Entomologist 27: 14-17.
Steiner, F.M., Schlick-Steiner, B.C., Trager, J.C., Moder, K., Sanetra, M., Christian, E. &
Staufer, C. 2006. Tetramorium tsushimae, a new invasive ant in North America. Biological
Invasions 8: 117-123.
Tabachnick, B.G. & Fidell, L.S. 1996. Using multivariate statistics (3rd ed.). New York:
Harper Collins.
Talbot, M. 1948. A comparison of two ants of the genus Formica. Ecology 29: 316–325.
Trager, J.C. & Johnson, C. 1985. A slave-making ant in Florida—Polyergus lucidus with
observations on the natural history of its host Formica archboldi (Hymenoptera:
Formicidae). Florida Entomologist 68: 261-266.
Umphrey, G. 1996. Morphometric discrimination among sibling species in the fulva–rudis–
texana complex of the ant genus Aphaenogaster (Hymenoptera: Formicidae) Canadian
Journal of Zoology 74: 528–559.
Wheeler, W.M. 1904. A new type of social parasitism among ants. Bulletin of the American
Museum of Natural History. 20: 347-375.
Wheeler, W.M. 1912. New names for some ants of the genus Formica. Psyche 19: 90.
Wheeler, W.M. 1913a. Corrections and additions to "List of type species of the genera and
subgenera of Formicidae." Annals of the New York Academy of Sciences. 23: 77-83.
Wheeler, W.M. 1913b. A revision of the ants of the genus Formica (Linné) Mayr. Bulletin of
the Museum of Comparative Zoology at Harvard College 53: 379-565.
Table 1. Summary of morphological index values for species in the F. pallidefulva group.
Means (and ranges) are presented for each measurement. We conducted univariate ANOVA
with Tukey’s HSD for post-hoc pairwise comparisons on each index for each species, and
subscript letters denote significant differences among homogeneous subsets based on those
analyses.
Species
Index F. archboldi F. biophilica F. dolosa F. incerta F. pallidefulva
CI 81.69 a 83.64 b 82.54 ab 86.77 c 82.07 ab

(78.6-87.5) (76.8-89.0) (71.9-88.5) (81.8-91.5) (76.9-88.9)
SI 147.43 c 142.53 b 145.30 bc 132.99 a 141.0 b

(139.4-154.6) (126.6-162.3) (125.0-160.7) (126.1-146.7) (126.4-153.7)
OI 31.26 d 29.38 bc 28.58 a 30.02 c 29.14 ab

(29.5-35.0) (27.9-30.8) (24.2-31.2) (27.4-31.9) (27.4-31.9)
CTI 61.72 b 61.41 b 60.42 a 62.04 b 61.70 b

(59.7-64.3) (58.2-64.7) (56.8-64.2) (58.8-64.7) (58.8-64.7)
TWI 40.12 a 40.55 a 40.59 a 41.90 b 41.89 b

(38.2-42.9) (37.7-43.8) (36.9-44.1) (38.2-44.8) (37.9-45.3)
Table 2. Summary of morphometric data for the F. pallidefulva group. Means (and ranges) are
provided for each measurement. The total number of individuals examined follows the specific
epithet, although actual sample sizes for each metric varied because not every measure could be
obtained from all specimens.
Species
Morphometric
F. archboldi F. biophilica F. dolosa F. incerta F. pallidefulva
n = 25 n = 32 n = 56 n = 31 n = 57
MdT 7.68 (7-9) 7.75 (7-9) 7.55 (6-10) 7.35 (6-9) 7.65 (6-10)
P5L 0.18 (0.16-0.20) 0.21 (0.18-0.24) 0.21 (0.16-0.24) 0.20 (0.16-0.22) 0.21 (0.17-0.24)
P6L 0.18 (0.17-0.22) 0.22 (0.19-0.28) 0.20 (0.17-0.24) 0.20 (0.16-0.22) 0.21 (0.16-0.24)
PnML 0.09 (0.03-0.14) 0.15 (0.08-0.24) 0.18 (0.10-0.25) 0.12 (0.06-0.16) 0.08 (0.02-0.12)
PpML 0.07 (0.05-0.11) 0.11 (0.06-0.17) 0.13 (0.07-0.19) 0.09 (0-0.11) 0.05 (0.03-0.08)
TML 0.16 (0.14-0.21) 0.23 (0.14-0.30) 0.22 (0.16-0.30) 0.18 (0.12-0.22) 0.13 (0.10-0.20)
TPL 0.05 (0.04-0.06) 0.05 (0.04-0.06) 0.06 (0.04-0.09) 0.05 (0.04-0.06) 0.02 (0-0.04)
HL 1.45 (1.31-1.58) 1.55 (1.37-1.82) 1.69 (1.31-2.06) 1.45 (1.15-1.64) 1.48 (1.16-1.68)
HW 1.19 (1.03-1.39) 1.30 (1.05-1.56) 1.40 (1.05-1.70) 1.26 (1.01-1.43) 1.21 (0.93-1.43)
EL 0.45 (0.41-0.51) 0.45 (0.40-0.53) 0.48 (0.40-0.55) 0.44 (0.37-0.48) 0.43 (0.34-0.50)
SL 1.75 (1.58-1.98) 1.84 (1.68-2.06) 2.02 (1.64-2.38) 1.68 (1.35-1.92) 1.71 (1.21-1.90)
WL 2.35 (2.08-2.65) 2.52 (2.22-2.93) 2.80 (2.12-3.48) 2.34 (1.86-2.67) 2.40 (1.90-2.71)
PnW 0.94 (0.83-1.09) 1.02 (0.89-1.23) 1.14 (0.87-1.37) 0.98 (0.77-1.11) 1.00 (0.80-1.15)
PnM 11.36 (1-23) 10.06 (2-20) 37.00 (14-64) 10.90 (4-17) 0.86 (0-5)
MnM 9.96 (3-19) 7.13 (2-15) 14.29 (5-32) 6.03 (0-12) 3.18 (0-10)
PpM 13.92 (6-31) 12.44 (6-35) 26.42 (12-42) 8.55 (0-16) 0.86 (0-5)
Figure 1. Mesometasternum of Formica dolosa drawn at approximately 45° angle with legs
removed. The scale bar equals 1 mm.
Figure 2. Morphological differences between F. biophilica and F. incerta: a) F. biophilica

(closed circles) generally has a longer mesosoma (WL) relative to head proportions than F.
incerta (open diamonds). b) Whereas F. biophilica rarely has more macrochaetae on the
propodeum than the pronotum (12 of 32 specimens), this is more often the case for the F. incerta
(22 of 31). The line indicates the relationship PnM = Ppm.
Figure 3. Scape length plotted against head width (= SI) for F. biophilica (closed circles), F.
incerta (open diamonds) and F. pallidefulva (plus signs). Note that in addition to the separation
of F. incerta from the other species, F. biophilica, on average, is somewhat larger than F.
pallidefulva, with a minimum scape length of approximately 1.7 mm.
Figure 4. Dorsal views of first gastral tergites of Formica pallidefulva group workers: A: F.
pallidefulva; b: F. biophilica; c: F. incerta; d: F. archboldi; e: F. dolosa. Scale bar = 1 mm.
Figure 5. Full-face views of heads of Formica pallidefulva group workers: a: F. pallidefulva; b:

F. biophilica; c: F. incerta; d: F. archboldi; e: F. dolosa. Scale bar = 1 mm.
Figure 6. Lateral view of mesosoma, petiole, and first gastral tergite of Formica pallidefulva
group workers: a: F. pallidefulva; b: F. biophilica; c: F. incerta; d: F. archboldi; e: F. dolosa.
Scale bar = 3 mm.
Ward, P. S. 2007. The ant genus Leptanilloides: discovery of the male and evaluation of
phylogenetic relationships based on DNA sequence data, pp. 637-649. In Snelling, R. R., B. L.
Fisher, and P. S. Ward (eds). Advances in ant systematics (Hymenoptera: Formicidae): homage
to E. O. Wilson – 50 years of contributions. Memoirs of the American Entomological Institute,
80.
THE ANT GENUS LEPTANILLOIDES: DISCOVERY OF THE MALE AND

EVALUATION OF PHYLOGENETIC RELATIONSHIPS BASED ON DNA SEQUENCE
DATA
Philip S. Ward
Department of Entomology,
University of California, Davis
Davis, CA 95616, USA
psward@ucdavis.edu
ABSTRACT
The male of the ant genus Leptanilloides is described, based on two specimens collected as
isolated individuals in sweep samples from montane Costa Rica. Although not associated with
ant workers or queens, these males are identified as Leptanilloides mckennae Longino on the
basis of DNA sequence similarity to a reference worker of this species from Costa Rica. The
male is slightly smaller than the worker, and has most traits expected of the dorylomorph group,
including tergosternal fusion of abdominal segments 2 and 3 (but not 4); exposed spiracles on
abdominal segments 5 and 6; a simplified volsella, lacking a cuspis; and absence of cerci.
Additional attributes include: mandibles elongate-triangular and edentate; anterior clypeal
lamella well developed; pronotum triangular in profile and much narrowed anteromedially;
mesopleuron lacking posterior oblique sulcus; postpetiole absent; and wing venation reduced
(forewing with pterostigma and one submarginal cell, but no discal cell). A molecular
phylogenetic analysis, based on 5.1 kb of sequence data from five nuclear genes (18S rDNA,
28S rDNA, wingless, abdominal-A, and LW-rhodopsin), provides strong independent evidence
that Leptanilloides is a member of the dorylomorph clade (parsimony bootstrap support of
100%; Bayesian posterior probability of 1.00), although relationships among taxa within that
group remain poorly resolved. The dorylomorphs are nested within the formicoid clade, a very
well supported group that contains the majority of extant ant subfamilies. Contrary to some
previous suggestions Leptanilloides is not closely related to Leptanilla. The latter appears to
stand well outside the formicoid clade, as sister taxon to all other extant ants sampled in this
study (72% parsimony bootstrap, 1.00 posterior probability). The results highlight the repeated
convergent evolution of similar morphology among subterranean ants.
Key words: Hymenoptera, Formicidae, phylogeny, morphology, dorylomorphs, doryline

section, Leptanilla, subterranean ants, 18S rDNA, 28S rDNA, wingless, abdominal-A, LW-
rhodopsin
INTRODUCTION
The study of ant assemblages living in subterranean habitats is one of the more exciting frontiers
in myrmecology. Underground environments seem to harbor a disproportionate share of
morphologically peculiar and phylogenetically interesting formicids (Brown et al., 1971; Bolton,
1990b; López et al., 1994; Ward, 1994). The ant subfamily Leptanilloidinae is one such group.
It comprises two genera of small, blind ants, whose biology, distribution, and phylogenetic
relationships are poorly understood (Bolton, 1990a, 2003; Brandão et al., 1999; Brady & Ward,
2005). Confined to the New World, these ants are rarely collected and are assumed to be
hypogaeic predators, possibly with legionary habits.
For many years the group was known only from a single species, Leptanilloides biconstricta
Mann (1923), described from Bolivia. The genus came to be placed in the tribe Cerapachyini
(Brown, 1975; Bolton, 1990a), and later in its own subfamily (Bolton, 1992). Our knowledge of
the group was considerably expanded by Brandão et al. (1999) who revised the subfamily and
described six new South American species, three of which were assigned to a new genus,
Asphinctanilloides. Longino (2003) described another new species, Leptanilloides mckennae,
from Costa Rica. This not only extended the known range of the subfamily from South America
to Central America, but also blurred the morphological distinctions between the two genera.
The phylogenetic position of Leptanilloidinae has been a point of uncertainty. Mann (1923)
and Borgmeier (1955) noted similarities to the Old World subfamily Leptanillinae, and to New
World army ants (Ecitoninae). Brown (1975) considered Leptanilloides to be a ponerine ant in
the tribe Cerapachyini, while Bolton (1990a, 1990c, 2003) provided compelling morphological
evidence that it belongs with the dorylomorphs (previously known as the doryline section), a
well defined monophyletic group that includes both cerapachyines and army ants, but not
Leptanillinae or Ponerinae. A recent morphological phylogenetic analysis of the dorylomorphs
(Brady & Ward, 2005) also concluded that Leptanilloidinae is part of this group, although a
close association with Leptanillinae could not be confidently rejected in that study.
Until recently all collections of Leptanilloides and Asphinctanilloides have consisted of
workers only, with males and queens being unknown. This paper provides the first description
of leptanilloidine males (see also postscript at the end of this paper). Although the specimens
were taken in sweep samples and were not associated with any specific ant colonies, their
identity as Leptanilloides was confirmed through comparative analysis of DNA sequence data.
The males in question were brought to my attention by John T. (“Jack”) Longino, who
encountered them when processing ants from Costa Rican sweep samples collected by John
Noyes. The males (two in number) were recognized as being unusual and it was hypothesized
that they represented the hitherto unknown males of Leptanilloides. Since the samples were
preserved in ethanol, and since workers of L. mckennae were also available in ethanol, the
opportunity was presented to test this hypothesis through comparison of DNA sequences.
Once the identity of the males as Leptanilloides was resolved, a broader inquiry into the
phylogenetic position of the genus was undertaken, using sequence data from five nuclear genes.
This yielded a number of interesting results, not only concerning Leptanilloides but also
regarding the phylogenetic relationships of other ant groups, including the enigmatic subfamily
Leptanillinae.
Critical material for this study was supplied by Jack Longino: an ethanol-preserved worker of
Leptanilloides mckennae from Monteverde, Costa Rica (July 1995; leg. D. McKenna), and two
unidentified male ants from Costa Rica, suspected to be Leptanilloides, and also held in ethanol.
The males were collected at Estación Cacao, Prov. Guanacaste (22 February 2003; leg. J. S.
Noyes), and Sendero Pilón, P. N. Arenal, Prov. Alajuela (26 February 2003; leg. J. S. Noyes),
Ward: Leptanilloides - description of male and phylogenetic evaluation 639
respectively. A fresh, ethanol-preserved worker of a second species, Leptanilloides nomada

Donoso, Vieira & Wild (2006) from Cotopaxi, Ecuador was provided by Alex Wild.
One of the two males, from Estación Cacao, was sacrificed for DNA extraction and
sequencing, but the head, wings, and genitalia of this specimen were retained for morphological
study and for comparison with the second male. Twenty-seven additional ant species were
targeted for sequencing (Table 1). These were chosen to provide broad coverage of higher-level
ant diversity, including nine other representatives of the dorylomorph group (belonging to the
genera Acanthostichus, Aenictus, Cerapachys, Cylindromyrmex, Dorylus, Neivamyrmex,
Simopone, and Sphinctomyrmex) and one representative of Leptanillinae (genus Leptanilla).
Three non-formicid outgroups were also included: a vespid (Mischocyttarus flavitarsis), a
bradynobaenid (Chyphotes mellipes), and an apid (Apis mellifera). All trees were rooted with
Apis mellifera.
Molecular methods
DNA sequence data from five nuclear genes were utilized: the small subunit (18S) and large
subunit (28S) ribosomal RNA genes, and the protein-encoding genes abdominal-A (abd-A),
wingless (wg), and long wavelength rhodopsin (LW Rh). Seventy-eight new sequences were
generated for this investigation, and the remainder were taken from Ward & Downie (2005).
Methods of DNA extraction, amplification, purification, and sequencing followed Ward &
Downie (2005). Sequences were collated with Sequencher (version 4.0.5), aligned with Clustal
X version 1.8 (Thompson et al., 1997), and manually edited with MacClade 4 (Maddison &
Maddison, 2000). GenBank accession numbers for each sequence are given in Table 1.
Phylogenetic analysis
An intron in the LW Rh gene fragment and hypervariable regions of the 28S gene were
removed prior to phylogenetic analyses, due to alignment difficulties. After exclusion of these
sites the five-gene dataset consisted of 5135 characters (base pairs), of which 1336 were variable
and 950 were parsimony-informative. Parsimony searches were carried out with PAUP*
(version 4.0b10) (Swofford, 2003), using TBR branch swapping and 1000 random taxon
additions. Gaps were treated as missing data. Bootstrap values were calculated using 1000 TBR
replicates and 10 random taxon additions per replicate. Bayesian estimation of the phylogeny
was conducted with Mr. Bayes v3b4 (Huelsenbeck & Ronquist, 2001), using a partitioned model
as described in Ward & Downie (2005). The MCMC (Markov chain Monte Carlo) analysis was
run for 10,000,000 generations, with sampling every 1000 generations, and a burn-in value of
1000 samples. The aligned data matrix used in phylogenetic analyses, with excluded sites
identified, is available at TreeBase (M2144).
Metric measurements and indices
The following measurements (in mm) and indices were employed. Measurements were taken at
50× with a Wild M5A microscope and a Nikon dual-axis micrometer.
HL Head length, measured in full-face (frontal) view, from the posterior margin to the
midpoint of the anterior clypeal margin, excluding the clypeal apron.
HW Maximum width of the head, including the eyes.
SL Length of scape, excluding the basal condyle and neck.
EL Length of compound eye, as seen in full-face view.
LHT Length of the metatibia in dorsal view, excluding the medioproximal lobe.
Table 1. List of taxa included in the molecular phylogenetic analysis, with GenBank accession numbers.
Taxon Locality 18S 28S wg LW Rh Abd-A
Formicidae
Amblyoponinae
Amblyopone pallipes California, USA AY703487 AY703554 AY703621 AY703755 AY703688
Dolichoderinae
Leptomyrmex erythrocephalus New South Wales, AS AY703494 AY703561 AY703628 AY703762 AY703695
Liometopum occidentale California, USA AY867449 AY867465 AY867434 AY867496 AY867481
Dorylomorph group
Acanthostichus kirbyi Rondônia, BR AY867438 AY867454 AY867423 AY867485 AY867470
Aenictus eugenii Swaziland AY867439 AY867455 AY867424 AY867486 AY867471
Cerapachys augustae Arizona, USA AY867440 AY867456 AY867425 AY867487 AY867472
Cerapachys larvatus Aust. Cap. Terr., AS AY703491 AY703558 AY703625 AY703759 AY703692
Cylindromyrmex striatus Bolívar, VZ AY867441 AY867457 AY867426 AY867488 AY867473
Dorylus helvolus KwaZulu-Natal, SF AY867442 AY867458 AY867427 AY867489 AY867474
Leptanilloides mckennae Puntarenas, CR AY867451 AY867467 -- AY867498 --
Leptanilloides male Guanacaste, CR AY867444 AY867460 AY867429 AY867491 AY867476
Leptanilloides nomada Cotopaxi, EC AY867443 AY867459 AY867428 AY867490 AY867475
Neivamyrmex nigrescens Arizona, USA AY867445 AY867461 AY867430 AY867492 AY867477
Simopone marleyi KwaZulu-Natal, SF AY867446 AY867462 AY867431 AY867493 AY867478
Sphinctomyrmex steinheili Aust. Cap. Terr., AS AY867447 AY867463 AY867432 AY867494 AY867479
Ectatomminae
Ectatomma opaciventre São Paulo, BR AY703492 AY703559 AY703626 AY703760 AY703693
Typhlomyrmex rogenhoferi Loreto, PE AY703496 AY703563 AY703630 AY703764 AY703697
Formicinae
Camponotus sp. nr. vicinus California, USA AY867448 AY867464 AY867433 AY867495 AY867480
Formica moki California, USA AY703493 AY703560 AY703627 AY703761 AY703694
Leptanillinae
Leptanilla sp. Eastern Cape, SF AY867436 AY867452 AY867421 AY867483 AY867468
Myrmeciinae
Myrmecia pyriformis South Australia, AS AY703500 AY703567 AY703634 AY703768 AY703701
Nothomyrmecia macrops South Australia, AS AY703501 AY703568 AY703635 AY703769 AY703702
Myrmicinae
Aphaenogaster occidentalis California, USA AY867450 AY867466 AY867435 AY867497 AY867482
Myrmica tahoensis California, USA AY703495 AY703562 AY703629 AY703763 AY703696
Paraponerinae
Paraponera clavata Maranhão, BR AY703489 AY703556 AY703623 AY703757 AY703690
Ponerinae
Hypoponera opacior California, USA AY703488 AY703555 AY703622 AY703756 AY703689
Platythyrea mocquerysi Madagascar AY867437 AY867453 AY867422 AY867484 AY867469
Proceratiinae
Proceratium stictum Queensland, AS AY703490 AY703557 AY703624 AY703758 AY703691
Pseudomyrmecinae
Pseudomyrmex gracilis Guanacaste, CR AY703529 AY703596 AY703663 AY703797 AY703730
Tetraponera punctulata Northern Terr., AS AY703514 AY703581 AY703648 AY703782 AY703715
other Vespoidea
Chyphotes mellipes California, USA AY703485 AY703552 AY703619 AY703753 AY703686
Mischocyttarus flavitarsis California, USA AY703486 AY703553 AY703620 AY703754 AY703687
Apis mellifera California, USA AY703484 AY703551 AY703618 AY703752 AY703685
CI Cephalic index: HW/HL.

REL Relative eye length: EL/HL.
REL2 Relative eye length, using HW: EL/HW.
SI Scape index: SL/HW.
IDENTIFICATION OF LEPTANILLOIDES MALE FROM DNA SEQUENCE DATA
The Leptanilloides mckennae worker readily yielded 18S and 28S sequences but it proved
difficult to obtain single-copy nuclear gene sequences from this specimen, possibly because of
its age and preservation history. It is likely that the worker had been stored at an ethanol
concentration of less than 95% (Longino, pers. comm.). Using primers that amplified short
stretches of DNA, I was able to obtain sequence for the LW Rh gene, but not for wg or abd-A.
This dictated use of a three-gene data set (18S, 28S, and LW Rh) to compare the putative
Leptanilloides male with the L. mckennae worker and with other taxa.
The results were unambiguous: the male proved to be identical to Leptanilloides mckennae
at the 18S and 28S genes, and almost identical (0.18% sequence divergence) at LW-rhodopsin.
The only difference detected at the LW-rhodopsin gene was a single base substitution in an
intron, which was 90 bp long in both the male and the L. mckennae worker. (The intron was 84
bp long in Leptanilloides nomada from Ecuador.) Over the concatenated three-gene data set the
male exhibited 0.02% sequence divergence from Leptanilloides mckennae, 0.79% from
Leptanilloides nomada, and moderate to large differences with all other sampled ants, with
uncorrected sequence divergences ranging from 3.31% (with Acanthostichus kirbyi) to 7.76%
(with Leptanilla sp.) (Table 2). Parsimony analysis of the sequence data placed the male as
sister to Leptanilloides mckennae, to the exclusion of Leptanilloides nomada, with 100%
bootstrap support (results not shown).
Table 2. Percent sequence divergence (uncorrected) between the putative Leptanilloides male and selected
other taxa, individually and for three genes combined.
Gene
Taxon 18S 28S LW Rh LW Rh all 3 genes

exon intron (incl. intron)
Leptanilloides mckennae 0.00 0.00 0.00 1.11 0.02

Leptanilloides nomada 0.27 0.50 2.62 8.24 0.79
Acanthostichus kirbyi 0.54 1.50 16.38 33.00 3.31
Cylindormyrmex striatus 0.65 1.83 15.72 41.91 3.60
Cerapachys augustae 0.49 2.05 18.34 52.47 4.22
Cerapachys larvatus 0.43 2.94 18.12 50.57 4.34
Simopone marleyi 0.49 3.17 16.59 39.61 4.07
Sphinctomyrmex steinheili 0.70 1.74 18.78 52.73 4.21
Neivamyrmex nigrescens 2.32 4.05 19.00 38.03 5.54
Aenictus eugenii 1.62 5.74 17.47 41.79 5.91
Dorylus helvolus 1.14 4.84 19.65 44.20 5.60
Ectatomma opaciventre 0.97 4.41 16.16 53.24 5.10
Nothomyrmecia macrops 0.89 4.84 19.21 50.92 5.54
Amblyopone pallipes 1.40 4.27 17.90 55.13 5.54
Leptanilla sp. 1.78 8.06 22.05 56.38 7.76
The possibility exists that the Leptanilloides male, while genetically very close to L.
mckennnae, belongs to a different (unknown) species. Without more information about the
diversity of Leptanilloides in Costa Rica and the degree of interspecific sequence divergence in
the genus, this possibility cannot be entirely dismissed. The virtual identity of the male with L.
mckennae, contrasted with its substantial divergence from Leptanilloides nomada, argues against
this interpretation. L. nomada is clearly not conspecific with L. mckennae but it appears to be
more closely related to the latter than to most other described species of the genus. Specifically,
it has the combination of a reduced postpetiole (smaller than petiole) and enlarged abdominal
segments 4-6, separated by distinct constrictions, that sets L. mckennae apart from most other
described species of Leptanilloides.
Thus I conclude that the sequenced male (from Estación Cacao) and the morphologically
similar specimen (from P. N. Arenal) represent the male caste of Leptanilloides mckennae
Longino.
DESCRIPTION OF LEPTANILLOIDES MCKENNAE MALE
Head broader than long, with large eyes that occupy the anterior half of sides of head; mandibles
slender, elongate-triangular, and overlapping at closure (Figure 1), masticatory margin edentate
and rounding into unarmed basal margin; external margin of mandible weakly concave;
maxillary palps 2-segmented; labial palps difficult to discern in situ but apparently similarly
reduced; genal teeth and hypostomal teeth lacking; anterior margin of clypeus with a prominent,
subtriangular, translucent lamella, bluntly pointed medially and with an anterior-posterior
extension at this midpoint that is subequal to scape width; posterolateral margin of clypeus well
marked, but posteromedial boundary obscure; antennal sockets horizontal and exposed, and
located close to the anterior clypeal margin; antenna 13-segmented, each segment much longer
than wide; scape and ultimate antennal segment subequal in length, each 0.12× total length of
antenna and 2.2× length of the second antennal segment; front of head immediately posterior to
antennal sockets slightly depressed, and furnished medially with a low, blunt, longitudinal ridge,
probably homologous with the elevated frontal carinae of the worker; lateral ocelli separated
from median ocellus by about their diameters.
Mesosoma with usual complement of sclerites (Figure 2); pronotum U-shaped in dorsal
view and reduced anteromedially to a thin horizontal strip, set well below the level of the
mesonotum; pronotum triangular in profile, with pointed posterior apex; mesonotum lacking
notauli, but with a darkened, weakly impressed anteromedial suture; parapsidal sutures present;
parascutal carina poorly developed and terminating before the transcutal cleft; axillae not
meeting medially, connected by a narrow furrow; tegula very small and inconspicuous;
mesopleuron lacking oblique transverse sulcus and hence not divided into anepisternum and
katepisternum, but lower third of mesopleuron with a broad longitudinal furrow (possibly an
artifact of cuticular collapse); boundary between metapleuron and propodeum effaced
posteriorly; metapleural gland reduced and inconspicuous; propodeal spiracle small, circular,
positioned at about midheight of propodeum and as far back as the posterior extremity of the
metanotum. Legs slender (LHT/HL ~1.5); mesotibia and metatibia each with a single spur,
barbulate and pectinate, respectively; tarsal claws lacking preapical tooth.
Wings somewhat infuscated and with reduced venation (Figure 3); pterostigma present;
forewing with one submarginal cell; closed discal (medial) cell lacking, i.e., m-cu crossvein
absent; veins M and Cu diverging distal to crossvein cu-a by a distance greater than the length of
the crossvein; hindwing lacking closed cells; anterior margin of hindwing with 3-4 hamuli; jugal
lobe absent.
Metasoma long and slender; abdominal segment 2 (petiole) subquadrate in profile (Figure
2), longer than high or wide, and only weakly constricted posteriorly (helcium thus apparently
quite broad); spiracle on abdominal segment 2 located at anterodorsal extremity; abdominal
segment 3 larger than petiole, and not developed as postpetiole nor separated from abdominal
segment 4 by a marked constriction; abdominal spiracle 3 located on anterior third of tergite;
abdominal segments 2 and 3 with tergosternal fusion; abdominal segment 4 lacking tergosternal
fusion; segment 4 with short but distinctly differentiated presclerites; spiracle present on anterior
half of tergite 4; abdominal segments 5 and 6 apparently lacking differentiated presclerites, and
not separated from succeeding segments by constrictions; abdominal spiracles 5 and 6 small and
inconspicuous but exposed, i.e., visible at anterior margins of respective tergites, under normal
distension; abdominal tergite 8 (pygidium) small and simple but visible dorsally, not wholly
covered by abdominal tergite 7; cerci absent; subgenital plate (abdominal sternite 9) with
posterior margin broadly concave but not bifurcate; anterior margin of subgenital plate with
mesial apodeme better developed than lateral apodemes; basal ring not hypertrophied; paramere
long and slender with upturned apex (Figure 2), about 1.5× petiole length; volsella simple, lobe-
like, lacking differentiated cuspis.
Body size small; total length, excluding appendages, approximately 2.7 mm; integument
mostly smooth and shiny, with scattered piligerous punctures; pilosity common on most of body,
suberect to decumbent. Color: yellowish-brown, dorsum of head a contrasting dark brown;
abdominal segments 4-8 medium brown.
Comparison of the two males
The two males were closely comparable in terms of external morphology, the chief
difference between them being larger eye size in the Arenal male (see measurements below).
Head shape, antennal proportions, integument sculpture, color, and overall habitus were
otherwise very similar. The preceding description is a composite for the head, wings and
genitalia, and is based on the Arenal male alone for other details.
Measurements (in mm) and indices:
HW HL SL LHT CI SI REL REL2

P.N. Arenal 0.639 0.472 0.242 0.724 1.35 0.38 0.54 0.40
Estación Cacao 0.592 0.452 0.221 — 1.31 0.37 0.47 0.36
Comments
To the extent that the Leptanilloides mckennae male is representative it suggests the
following provisional diagnosis for males of the subfamily Leptanilloidinae: mandibles
elongate-triangular and edentate; palp formula 2,2 or less; antennae 13-segmented; anterior
clypeal lamella present; pronotum triangular in profile, U-shaped in dorsal view, and narrow and
strap-like anteromedially; notauli absent; mesopleuron lacking oblique transverse suture; petiole
nodiform; postpetiole absent; abdominal spiracles 5 and 6 visible under normal distension;
posterior margin of subgenital plate emarginate; cerci absent; mesotibia and metatibia each with
single apical spur; forewing with one submarginal cell and no discal (medial) cell.
Several features of the Leptanilloides mckennae male match those of the worker caste of the
same species, such as the well developed, medially pointed clypeal lamella; the form of the tibial
spurs; the location of the abdominal spiracles; and overall body size (2.7 mm long versus ~3.1
mm in the worker). General traits of the Leptanilloides male that provide support for placement
of the genus in the dorylomorph group include tergosternal fusion of abdominal segments 2 and
3 (but not 4); exposure of abdominal spiracles 5 and 6 under normal distension; absence of cerci;
and simplified volsella. Interestingly the Leptanilloides male also shares some specific attributes
with army ant males including a triangular pronotum (in profile); reduced parascutal carina;
absence of an oblique transverse sulcus on the mesopleuron; and relatively small tegula. In other
respects, however, Leptanilloides lacks some of the key characteristics of male army ants such as
large size (relative to workers), elongate and sickle-shaped mandibles, robust metasoma, and
modified genitalia.
RELATIONSHIP OF LEPTANILLOIDES TO OTHER ANTS: MOLECULAR

EVIDENCE
Following identification of the sequenced male as Leptanilloides mckennae, it was possible to

use this specimen as the representative for this species in a more detailed (five-gene) molecular
phylogenetic analysis. This overcame the problems associated with the single L. mckennae
worker whose DNA was less well preserved.
Under both parsimony and Bayesian approaches the closeness of the two Leptanilloides
species is confirmed, and there is unequivocal support (parsimony bootstrap 100%, Bayesian
posterior probability 1.00) for placement of Leptanilloides in the dorylomorph group (Figure 4).
The parsimony analysis indicates that Leptanilloides is sister to all other dorylomorphs
(bootstrap support of 72%), but Bayesian inference places Leptanilloides as sister to
(Acanthostichus + Cylindromyrmex) with a posterior probability of 0.89. In general,
relationships among the dorylomorph taxa are not well resolved, except for the pairing of
Acanthostichus and Cylindromyrmex. Both parsimony and Bayesian trees recover army ant
monophyly, but without strong support (cf. Brady, 2003). The dorylomorphs are placed firmly
in the formicoid clade (parsimony bootstrap 100%, Bayesian posterior probability 1.00), a group
that is increasingly well supported by molecular data (Ward & Brady, 2003; Saux et al., 2004;
Ward & Downie, 2005).
Within the formicoid clade, the remaining taxa (representatives from Ectatomminae,
Myrmicinae, Formicinae, Dolichoderinae, Myrmeciinae, and Pseudomyrmecinae) appear to
form a clade that is sister to the dorylomorphs, but low support levels (parsimony bootstrap 57%,
Bayesian posterior probability 0.68) undermine the reliability of this hypothesis. Among the
remaining formicoids, two groups are well supported, however, at least under Bayesian
inference (posterior probability of 1.00): (Ectatomminae + Myrmicinae + Formicinae) and
(Pseudomyrmecinae + Myrmeciinae).
The ant genus Leptanilla—and by implication other Leptanillinae such as Protanilla and
Anomalomyrma—is not closely related to Leptanilloides and other dorylomorphs. It lies well
outside the formicoid clade, as the sister group of all other ants sampled in this study (parsimony
bootstrap 71%, Bayesian posterior probability 1.00). An earlier study, based on 28S data only,
also suggested a basal position for Leptanilla among the Formicidae (Saux et al., 2004). This
result should be viewed with caution, however, since Leptanilla shows divergent sequence
evolution and its placement adjacent to the aculeate outgroups might be an artifact of long-
branch attraction (see Bergsten, 2005).
DISCUSSION AND CONCLUSIONS
Three subterranean ant taxa, Leptanillinae, Leptanilloidinae, and Apomyrma, have workers of
similar appearance that share a suite of characteristics, including small size, pale integument,
absence of eyes, absence of frontal lobes, very narrow clypeus, exposed antennal sockets, short
scapes, reduced abdominal segment 3 (compared with segment 4), flexible promesonotal suture,
and well developed sting. The last two features are probably ancestral for ants as a whole, but
the others appear to be traits associated with adaptation to underground habitats. Saux et al.
(2004) demonstrated with 28S sequence data that Apomyrma, formerly associated with the
Leptanillinae (Bolton, 1990b, 2003), is actually a member of the subfamily Amblyoponinae, in
accordance with its original suggested placement (Brown et al., 1971). The present study
confirms that the superficial similarity between Leptanilloides and Leptanilla is no more than
that, and that both groups arise far apart on the ant tree (Figure 4).
Recent molecular studies (Ward & Brady, 2003; Ohnishi et al., 2004, Saux et al., 2004;
Ward & Downie, 2005; present work) have produced results consistent with the hypothesis
that the evolutionary history of ants involved an early diversification of poneromorph-like taxa,
within which a second stronger radiation occurred that generated most of modern ant diversity
(Figure 5). Highly specialized subterranean ants occur in both the early (“basal”) lineages and in
various groups within the younger formicoid clade. This indicates that colonization of, and
restriction to, the underground environment, with attendant convergence in morphology and
behavior, has occurred on multiple occasions during ant evolution. Given the frequency of
specialized hypogaeic species in lineages at the base of the ant phylogeny, the possibility could
be entertained that these taxa reflect the ancestral condition for Formicidae, but this implies one
or more transitions from subterranean to above-ground habits, which seems improbable.
Moreover, epigaeic taxa are prevalent throughout the fossil record of ants, beginning in the
Cretaceous (Grimaldi & Agosti, 2000; Nel et al., 2004). Nevertheless semi-cryptic nesting
habits (e.g., living in rotten logs) could have provided the ancestral ants with opportunities for
specialization in both directions.
POSTSCRIPT
As this manuscript was being prepared I learned of the discovery of additional new
Leptanilloides species in Cotopaxi, Ecuador (J. M. Vieira and D. A. Donoso, pers. comm.), with
ergatogyne queens and males. Descriptions of these taxa and castes appear elsewhere (Donoso
et al., 2006).
ACKNOWLEDGEMENTS
I am grateful to Jack Longino and Alex Wild for providing critical material of Leptanilloides for
morphological analysis and DNA sequencing. Other useful ant specimens were furnished by
Seán Brady, Jürgen Gadau, Randy Morgan, Dave Olson, Hamish Robertson, Eli Sarnat, and
Riitta Savolainen. I thank Seán Brady, David Donoso, Ted Schultz, Roy Snelling, and Robert
W. Taylor for helpful comments on the manuscript. The financial support of the U. S. National
Science Foundation (grants DEB-9903650 and DEB-0431330) is gratefully acknowledged.
Finally I wish to express my appreciation to Ed Wilson for his collegiality and support. His
writings on social insect biology have provided much inspiration and insight over the years, and
induced in me (as in many others) a lifelong fascination with myrmecology.
REFERENCES
Bergsten, J. 2005. A review of long-branch attraction. Cladistics 21: 163-193.

Bolton, B. 1990a. Abdominal characters and status of the cerapachyine ants (Hymenoptera,
Formicidae). Journal of Natural History 24: 53-68.
Bolton, B. 1990b. The higher classification of the ant subfamily Leptanillinae (Hymenoptera:
Formicidae). Systematic Entomology 15: 267-282.
Bolton, B. 1990c. Army ants reassessed: the phylogeny and classification of the doryline section
(Hymenoptera, Formicidae). Journal of Natural History 24: 1339-1364.
Bolton, B. 1992. Subfamily Leptanilloidinae Bolton, new subfamily. P. 317 in: Baroni Urbani,
C., Bolton, B., & Ward, P.S. The internal phylogeny of ants (Hymenoptera: Formicidae).
Borgmeier, T. 1955. Die Wanderameisen der neotropischen Region. Studia Entomologica 3: 1-
720.
Brady, S.G. 2003. Evolution of the army ant syndrome: the origin and long-term evolutionary
stasis of a complex of behavioral and reproductive adaptations. Proceedings of the National
Academy of Sciences U. S. A. 100: 6575-6579.
Brady, S.G. & Ward, P.S. 2005. Morphological phylogeny of army ants and other
dorylomorphs (Hymenoptera: Formicidae). Systematic Entomology 30: 593-618.
Brandão, C.R.F., Diniz, J.L.M., Agosti, D. & Delabie, J.H. 1999. Revision of the Neotropical
ant subfamily Leptanilloidinae. Systematic Entomology 24: 17-36.
Brown, W.L., Jr. 1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae,
tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini.
Search. Agriculture (Ithaca, New York) 5 (1): 1-116.
Brown, W.L., Jr., Gotwald, W.H., Jr. & Lévieux, J. 1971. A new genus of ponerine ants from
West Africa (Hymenoptera: Formicidae) with ecological notes. Psyche (Cambridge) 77:
259-275.
Donoso, D.A., Vieira, J.M. & Wild, A.L. 2006. Three new species of Leptanilloides Mann
from Andean Ecuador (Formicidae: Leptanilloidinae). Zootaxa 1201: 47-62.
Grimaldi, D. & Agosti, D. 2000. A formicine in New Jersey Cretaceous amber (Hymenoptera:
Formicidae) and early evolution of the ants. Proceedings of the National Academy of
Sciences U. S. A. 97: 13678-13683.
Huelsenbeck, J.P. & Ronquist, F. 2001. MRBAYES: Bayesian inference of phylogenetic trees.
Bioinformatics 17: 754-755.
Longino, J.T. 2003. A new Costa Rican species of Leptanilloides (Hymenoptera: Formicidae:
doryline section: Leptanilloidinae). Zootaxa 264: 1-6.
López, F., Martínez, M.D. & Barandica, J.M. 1994. Four new species of the genus Leptanilla
(Hymenoptera: Formicidae) from Spain—relationships to other species and ecological
issues. Sociobiology 24: 179-212.
Maddison, D.R. & Maddison, W.P. 2000. MacClade 4. Analysis of phylogeny and character
evolution. Sunderland, Massachusetts: Sinauer Associates.
Mann, W.M. 1923. Two new ants from Bolivia. Psyche (Cambridge) 30: 13-18.
Nel, A., Perrault, G., Perrichot, V. & Néraudeau, D. 2004. The oldest ant in the Lower
Cretaceous amber of Charente-Maritime (SW France) (Insecta: Hymenoptera: Formicidae).
Geologica Acta 2: 23-29.
Ohnishi, H., Imai, H.T. & Yamamoto, M.-T. 2004 (“2003”). Molecular phylogenetic analysis
of ant subfamily relationship inferred from rDNA sequences. Genes and Genetic Systems
78: 419-425.
Saux, C., Fisher, B.L. & Spicer, G.S. 2004. Dracula ant phylogeny as inferred by nuclear 28S
rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae:
Amblyoponinae). Molecular Phylogenetics and Evolution 33: 457-468.
Swofford, D.L. 2003. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods).
Version 4. Sunderland, Massachusetts: Sinauer Associates.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. 1997. The
ClustalX windows interface: flexible strategies for multiple sequence alignment aided by
quality analysis tools. Nucleic Acids Research 24: 4876-4882.
Ward, P.S. 1994. Adetomyrma, an enigmatic new ant genus from Madagascar (Hymenoptera:
Formicidae), and its implications for ant phylogeny. Systematic Entomology 19: 159-175.
Ward, P.S. & Brady, S.G. 2003. Phylogeny and biogeography of the ant subfamily
Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361-386.
Ward, P.S. & Downie, D.A. 2005. The ant subfamily Pseudomyrmecinae (Hymenoptera:
Formicidae): phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology 30:
310-335.
Figures 1-3. Leptanilloides mckennae male. 1, Frontal (full-face) view of head; 2, lateral view
of body; and 3, wing venation. Figures 2 and 3 are drawn to the same scale.
Figure 4. Phylogenetic relationships of Leptanilloides and other ant taxa, based on analysis of
five nuclear genes (18S, 28S, wg, abd-A, and LW Rh; total: 5.1 kb). Numbers above each node
are parsimony bootstrap values; those below are Bayesian posterior probabilities. The tree
depicted is the 50% majority-rule consensus tree obtained from parsimony bootstrap analysis.
The Bayesian tree (majority-rule consensus of all sampled trees after burn-in) is almost identical
in topology except that Leptanilloides appears as sister to (Acanthostichus + Cylindromyrmex)
with posterior probability of 0.89.
Figure 5. Relationships among major groups of extant ants, as inferred from DNA sequence
data. Based on the findings of Ward & Brady (2003), Ohnishi et al. (2004), Saux et al. (2004),
Ward & Downie (2005), and present study. The subfamilies Aneuretinae and Heteroponerinae
(see Bolton, 2003) are here treated as part of Dolichoderinae and Ectatomminae, respectively.
Asterisks signify groups containing genera whose workers are entirely hypogaeic and blind.
Yamane, S. 2007. Pachycondyla nigrita and related species in Southeast Asia, pp. 650-663. In
PACHYCONDYLA NIGRITA AND RELATED SPECIES IN SOUTHEAST ASIA
Seiki Yamane
Department of Earth and Environmental Sciences, Faculty of Science
Kagoshima University
Korimoto, Kagoshima, 890-0065 Japan
sky@sci.kagoshima-u.ac.jp
ABSTRACT
Pachycondyla nigrita and related species in Southeast Asia are revised. Four new species are
described: P. batak from N. Sumatra, P. flavipes from Myanmar, P. pilidorsalis from Borneo and
Malay Peninsula, and P. wallacea from Sulawesi, Lombok and Bali. Keys to species are presented,
though males and queens are not available for all the species.
Key words: Hymenoptera, Formicidae, Ponerinae, Pachycondyla, Brachyponera, Southeast Asia,

new species, keys.
Yamane: Pachycondyla nigrita group 651
INTRODUCTION
The taxonomic status of the ponerine genus-group taxon Brachyponera has a complicated history
(Bolton, 1995, 2003). Although it is currently synonymized with the genus Pachycondyla, now a
very large genus (Bolton, 1995), 'Brachyponera' is a compact group comprising species from the
Australian, Oriental and Ethiopian regions. The Oriental fauna currently includes only a few
species mainly from tropical areas, and has received little taxonomic treatment. In this group the
male characters may be important for species discrimination, but males are rarely encountered
when digging nests. In this short review I treat larger species of this group (i.e., P. nigrita and
related species) for which better material and information are available compared with other
species such as those of the P. chinensis-complex. Although this selection of species has not been
made upon a phylogenetic basis, the present paper may resolve some problems in this group that
have long been neglected. A more comprehensive review is being planned to include all the forms
of 'Brachyponera' and using materials from many museums/institutions in the world.
CHARACTERIZATION OF 'BRACHYPONERA'
Although Brachyponera was synonymized with Pachycondyla (see Bolton, 1995), it seems a
well-defined compact group. Andersen (2000) still uses Brachyponera as genus name, and
Shattuck (1999) as subgenus name. However, in this paper I follow Bolton (1995), who did not
recognize any subgenera in Pachycondyla.
Emery (1911) and Ogata (1987) presented diagnoses for the genus Brachyponera. Although
Ogata's description was based upon the then single Japanese species, B. chinensis, more
information was included than in Emery (1911). According to these authors and my own
observations, 'Brachyponera' has the following recognition characteristics (W: worker, Q: queen,
M: male):
1) Eye relatively large compared with those in Ponera and Hypoponera, located near the
anterior margin of cranium (W/Q), 2) basal portion of mandible with shallow and elliptical fovea
(but the fovea obsolete or absent in some species from Borneo, Bali, Krakatau and Sumatra)
(W/Q), 3) mandible rather slender, boomerang shaped (M), 4) anterior margin of clypeus weakly
and roundly produced, and medially shallowly emarginate (W/Q), 5) first funicular segment
(pedicel) longer than second in W and Q, but much shorter than second in M, 6) palpal formula 3,
3 (W/Q), 7) mesonotum completely surrounded by suture/groove, i.e., promesonotal suture and
metanotal groove deep and complete (W/Q), 8) mesopleuron with a transverse furrow always in
Q and sometimes also in W, 9) propodeum compressed, seen from back narrowed dorsally (W/Q),
10) dorsum of propodeum distinctly lower than promesonotum (this condition also occurs in some
Hypoponera species of similar size) (W/Q), 11) propodeum in profile demarcated from metanotum
by a groove that passes near propodeal spiracle (rarely this groove is obsolete) (W/Q), 12)
subpetiolar process with a backward-directed projection (Ogata described this projection as a pair
of teeth for P. chinensis, but actually it is generally a single process, which is flat with a round
apical margin, and in profile acute apically) (W/Q/M), 13) pygidium ending as a sharp spine
posteriorly (M).
Species treated in this paper:
All the 'Brachyponera' species in Southeast Asia are very similar to one another, and at this
moment it is difficult to sort them into species groups. However, for the convenience I have
recognized three artificial groups based on body size and some other minor characters among
them, i.e., those comprising large-, medium- and small-sized species.
In the present paper I deal with the large-sized species, which can be separated from the others
by a combination of the following character states in the worker caste: 1) Head width over 0.9 mm
in workers from mature colonies, 2) the number of ommatidia along the long axis of eye more than
10 (usually around 12; Figs. 1-3), 3) antennal scape long, in full-face view surpassing the posterior
border of head by more than 1/4 of its length (Figs. 4-8; only in P. flavipes sp. nov. scape shorter),
4) all the flagellar segments of antenna longer than broad (this condition also applies to some
medium- and small-sized species such as P. chinensis-complex), and 5) petiole relatively thick,
with more or less flat dorsal surface in profile (transitional states between thick and thin petioles
are found in other species). Among these the character states 2 to 5 may be associated with body
size. I have primarily adopted the state 2 to separate the large-sized species from the others.
The Southeast Asian forms may be assigned to the three groups as follows:
Large-sized species: Pachycondyla nigrita (continental Asia and Taiwan), P. wallacea sp.
nov. (Sulawesi, Lombok and Bali), P. flavipes sp. nov. (Myanmar), P. pilidorsalis sp.
nov. (Malay Peninsula and Borneo), and P. batak sp. nov. (N. Sumatra).
Medium-sized species: Pachycondyla chinensis (widely distributed in S.E Asia and temperate
eastern Asia) and a few new forms.
Small-sized species: Pachycondyla luteipes (continental Asia), P. jerdoni (N. India), and
some new forms. Prof. P. S. Ward (personal communication) suggested that P. obscurans
may also belong to this group.
The present paper is principally based on the Sk. Yamane (SKY) Collection at Kagoshima
University, Kagoshima, Japan. Two syntypes with the same collection data were borrowed from
the Natural History Museum, Vienna through the courtesy of Dr. Stefan Schödl, and one of them
was used for redescription.
Measurements were made for some body parts for the holotype and paratypes. Abbreviations
are as follows: HW (head width), HL (head length excluding mandibles), SL (scape length
excluding condylar bulb), EL (maximum eye length), MSL (malar space length: distance between
anterior margin of eye and anterior margin of cranium).
KEY TO SPECIES (WORKERS)
1. Pronotal side, metapleuron above, propodeum extensively with rather strong rugae. Pronotal
dorsum finely striate (Sulawesi, Lombok and Bali).................................. P. wallacea sp. nov.
-- Mesosoma only weakly microsculptured................................................................................ 2
2. Antennal scape shorter (Fig. 5), surpassing the posterior margin of head by less than 1/4 of its
total length. Legs yellowish brown to orangish, contrasting with jet black mesosoma
(Myanmar) .................................................................................................. P. flavipes sp. nov.
-- Antennal scape longer, surpassing the posterior margin of head by more than 1/4 of its total
length (Figs, 4, 6-8). Legs brown to dark brown, contrast with body color weaker................ 3
3. Mesosomal dorsum without standing hairs; standing hairs if any very few and shorter than
width of funicular segment 1 of antenna (Sumatra) ....................................... P. batak sp. nov.
-- Mesosomal dorsum usually with more than ten standing hairs, some of which are as long as
width of funicular segment 1 .................................................................................................. 4
4. Mesopleuron often with a transverse groove; the groove sometimes incomplete but at least
scar visible. Posterior faces of propodeum and petiole more strongly punctate. Gastral tergite
1 usually with more than ten standing hairs (Borneo; Malay Peninsula) ..................................
............................................................................................................. P. pilidorsalis sp. nov.
-- Mesopleuron usually without such a groove; the groove if any vestigial. Posterior face of
propodeum medially and petiole entirely smooth or very weakly punctate. Gastral tergite 1
with fewer standing hairs, the number, excluding those on the posterior margin of the tergite,
being usually less than ten (Thailand, Vietnam, Myanmar) ...................................... P. nigrita
KEY TO SPECIES (QUEENS)
Queen of P. flavipes not available.
1. Pronotal side densely striate; metapleuron and propodeal side partly and irregularly rugose ..
.................................................................................................................. P. wallacea sp. nov.
-- Lateral face of mesosoma not striate, not rugose .................................................................... 2
2. Mesosomal dorsum virtually without standing hairs (Fig. 13). Standing hairs on gastral
tergites 1 and 2 completely confined to posterior margin (Fig. 14) ............... P. batak sp. nov.
-- Mesosomal dorsum with standing hairs (Fig. 10). Gastral tergites 1 and 2 with standing hairs
on the disc as well as at the margin (Fig. 11).......................................................................... 3
3. Smaller, with head width ranging from 0.83 to 0.91 mm. Mesosoma dorsally with many
standing hairs, some of which are as long as or longer than funicular segment 1. Gastral
tergites 1 and 2 each with more than 10 long standing hairs................. P. pilidorsalis sp. nov.
-- Larger, head width ranging from 1.04 to 1.06 mm. Mesosoma dorsally with sparser hairs, most
of which are much shorter than width of funicular segment 1. Gastral tergites 1 and 2 with
sparser standing hairs that are usually shorter than width of funicular segment 1 ..... P. nigrita
KEY TO SPECIES (MALES)
Males available only for the following two species.
1. Body light yellowish brown, with creamy white legs (Fig. 17). Ocelli large, their diameter
longer than scape width; distance between anterior and posterior ocelli only slightly longer
than ocellar diameter.................................................................................................. P. nigrita
-- Body and legs blackish brown (legs only slightly paler than the body; Fig. 15). Ocelli small,
their diameter shorter than scape length; distance between anterior and posterior ocelli much
longer than ocellar diameter........................................................................... P. batak sp. nov.
ENUMERATION OF SPECIES
Pachycondyla nigrita (Emery, 1895)

Figures 6, 17,18
Ponera nigrita Emery, 1895: 459 (w) examined.

Euponera (Brachyponera) nigrita: Emery, 1900: 668.
Brachyponera nigrita: Bingham, 1903: 102.
Pachycondyla nigrita: Brown in Bolton, 1995: 307.
Worker (redescription of a syntype of Ponera nigrita Emery). Measurements (mm): Body

length in fully extended posture ca. 6; HW 1.05; HL 1.14; SL 1.14; EL 0.21; MSL 0.10. SL/HL
1.00, Number of ommatidia along longest axis of eye 12.
Antennal scape long, surpassing the posterior margin of head by more than 1/4 of its total
length; segments 3-7, 11 distinctly longer than wide; segments 8-10 only slightly longer than wide
(relative length of funicular segments much longer than in those of P. luteipes). Mandibular fovea
distinct. Lateral face of mesosoma very weakly sculptured or shining; lateral face of propodeum
well demarcated from metapleuron by a suture, somewhat distinctly microsculptured and
relatively mat. In profile petiole thick; its lower margin longer than lower margin of metapleuron.
Queen. The following description is based on specimens from Doi Suthep, N. Thailand.
Structure. Body distinctly larger than the worker, with the head width ranging from 1.04 to 1.06
mm. Eye much larger than in the worker (ca. 0.28 mm long and with ca. 22 ommatidia along the
long axis). Distance between posterior ocelli longer than that between anterior and posterior ocelli.
Mesopleuron above always with a distinct transverse groove. Compared with the worker, petiole
relatively thin, seen from above almost lacking dorsal flat portion. Body sculpture much as in the
worker. Mesopleuron and metapleuron smooth and shining; in one specimen from Taiwan
posterior face of propodeum and petiole also smooth medially, and mesopleuron partly and
metanotum densely punctate. Pilosity. Body covered with decumbent pubescence except on
mesopleuron and metapleuron where pubescence is absent or very sparse. Eye with short fine
standing hairs. Mesosoma dorsally and gastral tergites with very sparse standing hairs; long hairs
on tergites almost confined to their posterior margins; sternites with more long standing hairs on
the disc. Outer margin of forewing with dense short fringe; outer and posterior margins of
hindwing fringed with longer hairs. Coloration much as in the worker.
Male. Description based on the specimens from the same series as the queen. Structure. Head
including eyes much wider than long. Eyes very large (maximum diameter ca. 0.40 mm) with outer
margin strongly convex and inner margin weakly emarginate. Clypeus transverse with anterior and
posterior margins truncate; labrum anteriorly produced as a narrow lobe. Mandible long and
fragile, not opposable, basally wide and suddenly narrowed toward bluntly pointed apex. Ocelli
relatively large; distance between posterior ocelli slightly longer than their diameter, and as long
as ocello-occipital distance; antenna long; scape (segment 1) twice as long as segment 2 and
slightly shorter than segment 3. Pronotum short; mesoscutum large, with weak notauli and sharp
parapsidal lines; mesopleuron large, with a wide transverse furrow which is striate on the bottom;
scutellum dorsally convex with an anterior transverse zone which is much lower than the main
disc; axilla extensively concave; metanotum short. Propodeum rounded; dorsal, lateral and
posterior faces not clearly differentiated. Petiole relatively low, in profile rather strongly narrowed
above, seen from back almost as wide as high, with dorsal margin almost straight; subpetiolar
process flat, with acute posterior tooth in profile. Sculpture. Head superficially micropunctate.
Sculpture on pronotum, mesoscutum, metanotum and metapleuron still weaker; mesopleuron
extensively smooth and strongly shining; axilla coarsely rugose. Propodeum more strongly and
densely sculptured, but on its posterior face sculpture somewhat weaker. Anterior face of petiole
mat; posterior face weakly shining. Gaster only superficially and sparsely punctate and somewhat
shining. Pilosity similar to that in the queen. Mesopleuron, metapleuron and posterior face of
petiole almost hairless. Petiole extensively with dense obliquely standing short hairs on its anterior
and lateral faces; subpetiolar process below with standing hairs. Fringe on wings as in the queen.
Coloration. Body brown to pale brown, with ivory white femora and tibiae; tarsi especially fore
tarsus darker. Scape and pedicel yellowish; other segments dark brown.
TYPE MATERIAL
Worker, Carin Ghecu, Myanmar, 1300-1400 m L. Fea II-III 88 [printed]/Ponera nigrita

Emery [handwritten]/Emery [handwritten]/Collect. G. Mayr [printed]/nigrita [handwritten] det
Emery [printed] (examined).
ADDITIONAL MATERIAL
THAILAND. 2w, Doi Inthanon N.P. (HQ, 1800 m alt.), Chiang Mai Prov., 20 viii 1998, Sk.
Yamane; 1w, Doi Ang Kang (HQ, 1300 m alt.), Chiang Mai Prov., 21 viii 1998, Sk. Yamane; 10w,
Doi Suthep N.P., Chian Mai Prov., 21-22 xii 1997, F. & Sk. Yamane; 12w, same loc., 18 viii 1998,
H. Okido & Sk. Yamane; 5w, 1m, same loc. & date, H. Okido (TH98-H006); 10w, 2q, 8m, same
data, Sk. Yamane (TH98-SKY-06 & -07). VIETNAM. 30w, Cuc Phuong N.P., Nho Quan Distr.,
10-12 viii 1998, Sk. Yamane; 3w, same loc., 9 xi 2001, Sk. Yamane; 10w, same loc. (320 m alt.),
10 xi 2001, K. Eguchi (Eg01-VN-189); 8w, Bavi N.P. (670 m alt.), Hatai Distr., 12 xi 2001, K.
Eguchi (Eg01-VN-227); 11w, same loc., Sk. Yamane (VN01-SKY-96); 3w, same loc. (1100 m
alt.), 19 iv 2002, K. Eguchi (Eg02-VN-037); 18w, Tam Dao (900-1240 m alt.), Vinh Phuc Prov.,
9 viii 1998, Sk. Yamane; 1q, same loc., 8 viii 1998, H. Okido; 1w, Na Xan (500 m alt.), Nghe An
Prov., 14 iv 1999, T.V. Bui. TAIWAN. 2w, 1q, Wulai, Taipei Hsien, 2 viii 1981, M. Terayama;
10w, Fushan, Taipei Hsien, 22-23 ix 1997, Sk. Yamane.
REMARKS
The worker specimens from Doi Suthep, Chiang Mai, N. Thailand agreed well with the type
material. This species is very similar to P. pilidorsalis that occurs in Borneo and the southern part
of the Malay Peninsula. The former is distinguished from the latter by 1) more weakly sculptured
posterior faces of propodeum and petiole (W), 2) fewer and shorter standing hairs on mesosoma
and gastral tergites 1 and 2 (W, Q), 3) frequent lack of transverse groove on mesopleuron (W), and
3) much larger body size (Q).
P. nigrita shows variation in sculpturation on the mandible and propodeum in the worker.
Striation and punctation in these parts tend to be stronger and more extensive in the specimens
from N. Vietnam; in some specimens the major portion of the mandible is densely striate, and the
lateral and posterior faces of the propodeum are extensively and rather coarsely striate or
irregularly sculptured.
The subspecies P. nigrita nigritella (Starcke, 1930) from Sumatra may not be a race of P.
nigrita, since according to the original description the body size and eye size are smaller for P.
nigrita, and head is relatively long (HL/HW 1.14 for nigritella vs. 1.10 for typical nigrita). Starcke
himself (1930) suggested that nigritella is closely related to Brachyponera nigrita chinensis,
which is now given specific status (Brown, 1958).
Pachycondyla pilidorsalis Yamane, sp. nov.

Figures 7, 9-11
Worker. Measurements (mm) holotype in parentheses: HW 0.96-0.98 (0.96); HL 1.00 -1.05

(1.00); SL 1.07-1.09 (1.08); EL 0.19-0.20 (0.19); MSL 0.11-0.13 (0.11). SL/HL 1.03-1.08 (1.08);
Number of ommatidia along long axis of eye ca.12.
Very similar to P. nigrita and P. batak. Mesopleuron usually with a transverse groove.
Posterior faces of propodeum and petiole constantly punctate. Dorsum of mesosoma usually with
more than ten standing hairs, some of which are longer than width of antennal segment 2. Gastral
tergites 1 and 2 each with more than ten long standing hairs.
Queen. Similar to P. batak in body size and distinctly punctate metapleuron (in P. nigrita
metapleuron almost smooth). Specimens from Malay Pen. tend to be larger than those from Borneo
(HW: 0.83-0.91 vs. 1.04-1.06 mm). Mesosoma dorsally and gastral tergites with numerous long
standing hairs.
TYPE MATERIAL
Holotype: worker, Ulu Gombak (ca. 250 m alt.), Selangor Prov., Malaysia., 5 vii 1999, F. Ito
(FI99-291). Paratypes: 10w, 2q, from the same colony as holotype. The holotype is deposited in
the Entomological Collection at Forest Research Institute, Kuala Lumpur, Malaysia, and paratypes
in SKY Collection, Kagoshima University, Natural History Museum (NHMK), London (BMNH),
Museum of Comparative Zoology, Harvard University (MCZC) and Natural History Museum of
Los Angeles County (LACM).
ADDITIONAL MATERIAL
MALAY PENINSULA. 2w, Ulu Gombak, 24-29 x 1996, F. Ito; 10w, Bunga Buah (ca. 1000
m alt.), nr Genting Highland, 6 vii 1999, Sk. Yamane. JAVA. 2w, Cidaon, Ujung Kulon, 10 iii
1997, F. Ito. BORNEO. 5w, Poring (550-700 m alt.), Sabah, 16 iii 1995, Sk. Yamane; 54w, Poring
(600-900 m alt.), Sabah, 1996-97, T. Kikuta; 3w, 1q, Sayap Kinabalu (ca. 1000 m alt.), 15 vii 1996,
K. Eguchi (Eg96-BOR-47); 6w, same loc., 14-15 vii 1996, K. Eguchi & Sk. Yamane; 10w, 3q, G.
Rara, Tawau, Sabah, 11 xii 1996, K. Eguchi (Eg96-BOR-375); 11w, 1q, Tasek Merimbun, Brunei,
17 ii 1999, K. Eguchi (Eg99-BOR-148).
ETYMOLOGY
The name refers to the presence of numerous standing hairs on the dorsum of the bodies of the
queen and worker.
REMARKS
Diagnostic characteristics separating this species from two related species are given in the
keys. Since the male has not been collected the status of this species is still inconclusive. This
species nests in rotting wood in good forests.
Pachycondyla batak Yamane, sp. nov.

Figures 3, 4, 12-16
Worker. Measurements (mm), holotype in parentheses : HW 0.95-1.00 (0.99); HL 1.04-1.11

(1.08); SL 1.11-1.13 (1.13); EL 0.19-0.21 (0.20); MSL 0.11-0.12 (0.11). SL/HL 1.03-1.08 (1.05).
Structure. Similar to P. nigrita and P. pilidorsalis, but the following characteristics are useful
to recognize P. batak. Mesal half of mandible rather extensively smooth with scattered punctures.
Posterior face of propodeum superficially but extensively punctate. Both anterior and posterior
faces of petiole distinctly punctate. Dorsal face of mesosoma without standing hairs or with at most
five very short standing hairs. Gastral tergites 1 and 2 each with less than five standing hairs
(excluding those along posterior margin of the tergites).
Queen. Small, head width ca. 0.85 mm. Similar in size and structure to the queen of P.
pilidorsalis, but mesosoma dorsally and gastral tergites 1 and 2 virtually without standing hairs.
Male. Structure. Head small, much narrower than pronotum, posteriorly roundly narrowed.
Clypeus apically almost straight; labrum produced anteriorly. Lower frons with a short
longitudinal keel. Mandible reduced, basally widened. Eye large (maximum length 0.30 mm),
strongly convex, with short standing hairs, mesally weakly emarginate. Ocelli small, diameter as
long as the width of antennal segment 3; distance between posterior ocelli much longer than that
between anterior and posterior ocelli. Scape as long as antennal segment 3; pedicel (segment 2)
slightly longer than half the length of scape. Pronotum posterolaterally concave; mesoscutum
large, much higher than pronotum; scutellum much narrower than mesoscutum, with a wide and
deep furrow just behind mesoscutum; mesopleuron large, with a deep transverse groove of which
the bottom has many striae. Propodeum lower than meso-metanotum, weakly rounded above;
posterior face shallowly concave. Petiole relatively thin, in profile much narrowed above;
subpetiolar process in profile with an apically acute posterior projection. Sculpture. Head only
superficially punctate and shining. Pronotum, mesoscutum, scutellum and metanotum very weakly
sculptured and somewhat shining; mesopleuron almost entirely smooth and strongly shining;
metapleuron anteromedially coarsely puncto-striate; other part of metapleuron weakly sculptured.
Propodeum very densely punctate except for concave posterior face that is only superficially
sculptured. Anterior and posterior faces of petiole densely and irregularly sculptured; posterior
face very weakly sculptured and somewhat shining. Gaster only superficially sculptured. Pilosity.
Body rather densely covered with decumbent pubescence and obliquely standing short hairs; on
mesopleuron the pilosity very sparse. Coloration. Body dark reddish brown, with slightly paler
gaster and legs.
TYPE MATERIAL
Holotype. Worker, Parapat (900 m alt.), nr Danau Toba, N. Sumatra, Indonesia, 19 viii 2002,
Sk. Yamane (SU02-SKY-91). Paratypes. 7w, 4q, 5m from the same colony. The holotype is
deposited in Museum Zoologicum Bogoriense, Indonesia, and paratypes in SKY Collection
(NHMK), BMNH, MCZC and LACM.
ADDITIONAL MATERIAL
SUMATRA: 26w 2q, same loc. (SU02-SKY-85); 11w2q, same loc. (SU02-SKY-90); 16w,
Bt Lawang (lowland), G. Leuser N.P., Sk. Yamane (SU02-SKY-87).
ETYMOLOGY
Named for the Batak tribe who live in the area of the type locality.
REMARKS
This species may be most closely related to P. pilidorsalis, from which it is easily
distinguished by the lack of standing hairs on mesosoma and gastral tergites 1 and 2 (excluding
those arising along posterior margins of the tergites) in the queen. It is also separable from the
related P. nigrita by the smaller body (queen), and smaller ocelli and dark body color (male). On
the other hand, in the worker caste, P. batak differs from P. pilidorsalis and P. nigrita only in the
fewer numbers of standing hairs on mesosoma and tergites 1 and 2. This species nests in rotting
wood/branches on the ground in secondary forests.
Pachycondyla flavipes Yamane, sp. nov.

Figure 5
Worker. Measurements (mm), holotype in parentheses: HW 0.90-0.94 (0.94); HL 1.00-1.04

(1.04); SL 0.94-0.98 (0.97); EL 0.19-0.20 (0.19); MSL 0.08-0.10 (0.10). SL/HL 0.93-0.96(0.93).
Number of ommatidia along the long axis of eye ca 10.
Structure. Head slightly longer than wide, almost parallel-sided behind eyes, with the
posterior margin almost straight in full-face view. Clypeus with the anterior margin widely and
very shallowly emarginate. All the funicular segments of antenna longer than wide. Antennal
scape surpassing the posterior margin of head by less than 1/4 of its total length in full-face view.
Anterodorsal corner of pronotum weakly and roundly produced; mesonotum demarcated
anteriorly from pronotum by a wide and deep furrow; mesopleuron above sometimes with a weak
transverse groove; metanotal depression distinct. Propodeum in profile separated by a distinct
furrow from metapleuron and propodeum; posterior face distinctly margined laterally. Sculpture.
Dorsum of head finely and densely punctate. Clypeus almost smooth except in median portion that
is longitudinally and finely striate. Mandible extensively smooth with large sparse punctures, and
only basal portion finely striatopunctate. Dorsum of pronotum finely punctate; its lateral face and
mesonotum with still finer punctation; mesopleuron and metapleuron smooth and shining, with
rugulae in lower posterior area. Propodeum dorsally irregularly sculptured but its lateral face
smooth; posterior face of propodeum smooth except for margins and upper portion. Petiole
anteriorly micropunctate, posteriorly smooth. Gastral segments finely punctate and somewhat
shining. Pilosity. Body covered with decumbent pubescence except for mesopleuron, metapleuron,
and posterior faces of propodeum and petiole that are almost bare. Dorsum of head, mesosoma,
petiole almost without standing hairs; disc of gastral tergites 1 and 2 with a few standing hairs in
addition to those at posterior margins; other tergites and sternites also with standing hairs at
margins. Coloration. Mesosoma jet black; head, petiole and gaster blackish brown. Antennal
funiculus and mandible brown to reddish brown. Legs yellowish brown to orangish, strikingly
contrasting with jet black mesosoma.
TYPE MATERIAL
Holotype: worker, 27 ii 2002, between Kampetlet and Natmataung N.P. (1700 m alt.), Chin
State, Myanmar, 27 ii 2002, Sk. Yamane leg (MM02-SKY-33). Paratypes: 16 workers from the
same colony. The holotype is deposited in SKY Collection (NHMK), and paratypes in NHMK,
BMNH, MCZC, and LACM.
ETYMOLOGY
Named for the yellow legs of this species.
REMARKS
This species has the shortest relative length of antennal scape among the species treated here.
The scape surpasses the posterior margin of head by less than 1/4 of its total length.
Pachycondyla wallacea Yamane, sp. nov.

Figure 8
Worker. Measurements (mm), holotype in parentheses: HW 1.05-1.13 (1.11); HL 1.10-1.17

(1.15); SL: 1.10-1.15 (1.15); EL: 0.21-0.23 (0.22); MSL: 0.08-0.09 (0.08). SL/HL 0.97-1.01 1.00).
Structure. Head slightly longer than wide, in full-face view with posterior margin weakly
emarginate, almost parallel-sided behind eye. Mandible with six larger and three smaller teeth on
the masticatory margin; basal fovea absent or obsolete. Antennal scape rather long, surpassing the
posterior margin of head by more than 1/4 (often almost 1/3) of its total length in full-face view.
Pronotal dorsum bluntly demarcated from lateral face; seen from above anterolateral corner round.
Mesonotum distinctly demarcated from pronotum and propodeum by deep furrows, posteriorly
weakly emarginate. In profile propodeum demarcated by a distinct obliquely running suture from
metapleuron; posterior face well demarcated from lateral face by a rather sharp ridge, but
separation from dorsal face indistinct. Petiole thick, in posterior view round and slightly wider than
high. Sculpture. Dorsum of head minutely and densely punctate. Clypeus with much more
superficial sculpture and shining except in median sculptured portion. Greater part of mandible
finely and densely striate, leaving small area around apex and along mesal margin smooth.
Pronotum very finely striatopunctate dorsally; its lateral face strongly rugose. Mesonotum finely
and superficially punctate; mesopleuron almost smooth and shining, but rugulose below;
metapleuron smooth, but punctate below and rugose above. Propodeum dorsally irregularly
sculptured and mat; lateral face rugose; posterior face densely and transversely striate. Petiole with
anterior and posterior faces densely and superficially punctate. gastral segments densely and
weakly punctate evenly over the surface. Pilosity. Body except for lateral face of mesosoma
extensively covered with dense decumbent pubescence. Standing hairs sparse; gastral tergites 1
and 2 each with more than ten standing hairs.
Coloration. Body dark brown to blackish; gaster slightly lighter than head and mesosoma.
Antennal funiculus, mandible, subpetiolar process and legs yellowish brown to orangish.
Queen. Only one winged teneral individual is available, which is not suitable for detailed
description. Striation on mesosoma less conspicuous than in the worker; pronotal side very finely
and densely striate; striation on metapleuron above and propodeal side irregular; dorsum of
pronotum and mesonotum very finely punctate and matte.
TYPE MATERIAL
Holotype: worker, Kopi house, nr Semaya, W. Lombok, Indonesia, 29 x 1998, K. Eguchi leg.
(Eg98-LMB-1077). Paratypes: one queen and 12 workers from the same colony. The holotype is
deposited in Museum Zoologicum Bogoriense, Indonesia, and paratypes in SKY Collection
(NHMK), BMNH, MCZC, and LACM.
ADDITIONAL MATERIAL
INDONESIA: 2w, S.E. Sulawesi, Indonesia, 25 xi 2000, IKT Ginarsa; 10w, West Bali, 27 vi
1998, IKT Ginarsa.
ETYMOLOGY
Named for the famous "Wallace's Line" dividing the Asian from the Melanesian faunas.
REMARKS
This species is similar to P. atrata (Karavaiev, 1925) in having rugose pronotal sides (also see
Wilson, 1958). It is, however, easily distinguished from the latter in the worker caste by having the
pronotal dorsum and propodeum extensively rugose/striate, and the antennal scape much longer.
In P. atrata the scape surpasses the posterior margin of head by only 1/5 of its length, while in P.
wallacea by more than 1/4 (often almost 1/3) of its length. P. atrata is known from Amboina and
Boeton, eastern Indonesia, and excluded here.
ACKNOWLEDGEMENTS
I thank Roy Snelling for his invitation to contribute to this honorable volume, and P. S. Ward for
critical reading of an earlier draft. Thanks are extended to Stefan Schödl for his kind loan of the
type material of Ponera nigrita, and Gary Alpert for kindly sending me the Stärcke paper.
Katsuyuki Eguchi, Hirofumi Okido, I.K.T Ginarsa, Decha Wiwatwitaya, and Mamoru Terayama
helped me in collecting the material examined in the present paper.
LITERATURE CITED
Andersen, A.N. 2000. The ants of Northern Australia. A guide to the monsoonal fauna. v+106 pp.
CSIRO Publishing, Collingwood.
Bolton, B. 1995. A new general catalogue of the ants of the world. 504 pp. Harvard University
Press, Cambridge, Massachusetts.
Brown, W.L. 1958. A review of ants of New Zealand. Acta Hymenopterologica 1: 1-50.
Emery, C. 1895. Viaggio de Leonardo Fea in Birmania e regioni vicine.63. Formiche di Birmania,
del Tenasserim e del Monti carin, raccolte da L. Fea. Annali del Museo Civico de Storia
Naturale di Genova (2) 14 [34]: 450-483.
Emery, C. 1911. Hymenoptera, Fam. Formicidae, Subfam. Ponerinae. In: Wytsman, P., Genera
Insectorum 118: 1-125, 3 pls.
Karavaiev, W. 1925. Ponerinae (Fam. Formicidae) aus dem Indo-Australischen Gebiet. Konowia
4: 69-81, 115-131, 276-296.
Ogata, K. 1987. A generic synopsis of the poneroid complex of the family Formicidae in Japan
(Hymenoptera). Part I. Subfamilies Ponerinae and Cerapachyinae. Esakia (Fukuoka) 25:
97-132.
Shattuck, S.O. 1999. Australian ants. Their biology and identification. Monographs on
Invertebrate Taxonomy 3: i-xi, 1-226.
Stärcke, A. 1930. Verzeichnis der bis jetzt von der Insel Pulau Berhala bekannt gewordenen
Ameizen. Treubia 12: 371-381.
Wilson, E.O. 1958. Studies on the ant fauna of Melanesia III. Rhytidoponera in western Melanesia
and the Moluccas. IV. The tribe Ponerini. Bulletin of the Museum of Comparative Zoology
119: 303-371.
Figures 1-8. Pachycondyla workers. 1-3, Eyes: 1, Pachycondyla luteipes; 2, P.

chinensis-complex; 3, P. batak sp. nov. 4-8, heads in full-face view: 4, Pachycondyla batak sp.
nov.; 5, P. flavipes sp. nov.; 6, P. nigrita; 7, P. pilidorsalis sp. nov.; 8, P. wallacea sp. nov.
Figures 9-14. Winged queens. 9-11, Pachycondyla pilidorsalis; 12-14, P. batak. 9 & 12, Body in
profile; 10 & 13, Promesonotum in profile; 11 & 14, 1st and 2nd gastral tergites in profile.
Figures 15-18. Males. 15 & 16, Pachycondyla batak sp. nov.; 17 & 18, P. nigrita.
Yoshimura, M., Onoyama, K. 2007. A new sibling species of the genus Strumigenys, with a
redefinition of S. lewisi Cameron, pp. 664-690. In Snelling, R. R., B. L. Fisher, and P. S. Ward
(eds). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson – 50
years of contributions. Memoirs of the American Entomological Institute, 80.
A NEW SIBLING SPECIES OF THE GENUS STRUMIGENYS,

WITH A REDEFINITION OF S. LEWISI CAMERON
Masashi Yoshimura
Institute of Tropical Agriculture, Kyushu University
6-10-1 Hakozaki, Higashi-ku, Fukuoka 812-8581, Japan
myoshimura@ant-database.org
and
Keiichi Onoyama
Nishi 21, Minami 4-11-9,
Obihiro, Hokkaido, 080-2471, Japan
ABSTRACT
Strumigenys lewisi and two undetermined species, S. sp. 4 and S. sp. 9 from Japan that closely
resemble S. lewisi, were examined to clarify the taxonomic status of them. Measurements of the
queens were analyzed. Our analysis indicated that S. sp. 9 is an intraspecific variation of S. lewisi,
and that S. sp. 4 should be regarded as a new species, S. kumadori sp. nov. Morphological
differences between S. kumadori and S. lewisi were found in a pair of hairs on the apicoscrobe and
mesonotum of the worker, the eye, lateral ocelli, mesonotum, fore and hind wings of the queen,
and the mandible, genitalia, and hind wing of the male. All three castes of S. kumadori and S.
lewisi are characterized, and illustrated with scanning electron micrographs and microscope
photographs. Species distinctions and a taxonomic discussion of S. kumadori and S. lewisi are
given. Ecological and morphological discussions are also provided.
Key words: Hymenoptera, Formicidae, Strumigenys, worker, queen, male, SEM, Japan.
Yoshimura & Onoyama: A new Strumigenys from Japan 665
INTRODUCTION
The genus Strumigenys is known for having narrow and linear to sublinear mandibles, developed
petiolar spongiform tissue, and specialized feeding habits (Baroni Urbani & De Andrade, 1994;
Bolton, 1999). The dacetine ants of the World were revised by Bolton (2000) who recognized 466
valid species of the genus Strumigenys, about 220 of which were recorded from the
Malesian-Oriental-East Palaearctic area. However, even his great work could not include all
species belonging to the genus, since many remained undescribed.
Two undetermined species which closely resemble Strumigenys lewisi Cameron, 1886 have
been recognized in Japan. Both of the two species which are tentatively named S. sp. 4 and S. sp.
9 have been indistinguishable from S. lewisi in workers. S. sp. 4 was first recorded by Sonobe
(1977) as Strumigenys sp. 1 [Japanese name: Kita-uroko-ari] in his faunal survey in Miyagi
Prefecture, the northeast region of mainland of Japan. He mentioned in his discussion that S. sp.
1 is very similar to S. lewisi but different from the latter only in the queen by the clearly raised
mesosoma and much larger eyes. On the other hand, S. sp. 9 was firstly recorded by Onoyama
(1976) as S. sp. A in his faunal survey in Okinawa Island, one of the southern islands of Japan, and
he suggested that this is a sibling species which is distinguishable from S. lewisi only in the queen.
Keys and descriptions distinguishing among these three species using morphologies of queens
have been provided by Onoyama et al. (1992), and S. sp. A in Onoyama (1976) was renamed as S.
sp. 9 [Japanese name: Minami-uroko-ari] in the description. However, the taxonomic status of the
two unnamed species and the relationship among the former two and S. lewisi remained unclear.
Bolton (2000) mentioned in his introduction that dacetine queens are usually very like their
workers in key characters, and actually did not provide description of the queens. Both of the two
S. lewisi-like sibling species in Japan might be regarded as quite simply S. lewisi, under the
taxonomic system based on only worker morphologies.
The present study aims to clarify the taxonomic status of the two sibling species of S. lewisi
and their taxonomic relationship among them.
MATERIAL AND METHODS
Terminology, measurements and indices
Terminology basically follows Bolton (2000), Gauld & Bolton (1988), and Huber & Sharkey
(1993). Measurements and indices are illustrated in Figs 1 and 2. Measurements and indices for
workers and their abbreviations follow Bolton (2000). DA3L and DA3W, and their indices, DA3I,
were additionally defined as below.
Dorsal smooth area of A3 length (DA3L): Maximum longitudinal length of the dorsal smooth area
on abdominal tergite III in dorsal view.
Dorsal smooth area of A3 width (DA3W): Maximum width of the dorsal smooth area on
abdominal tergite III in dorsal view.
Dorsal smooth area of A3 index (DA3I): DA3L / DA3W × 100.
Measurements and indices, and their abbreviations for the queen and male are defined as below.
Head length (HL): same as worker and including ocelli in male.
Head width (HW): same as worker but including eyes in male.
Cephalic index (CI): same as worker.
Mandible length (ML): same as worker (only queen).
Mandible index (MI): same as worker (only queen).
Scape length (SL): same as worker.
Scape index (SI): same as worker.
Eye length (EL): maximum length of eye in lateral view.

Eye index (EI): EL/HW × 100
Head depth (HD): maximum depth of head in lateral view measured perpendicular to the full-face
view plane.
Diameter of lateral ocelli (DlO): mean length of the diameters of right and left ocelli.
Lateral ocelli index (DlOI): DlO/ HW × 100.
Pronotum height (PrH): maximum height of the pronotum in lateral view.
Mesoscutum width (MsW): maximum width of the mesoscutum in dorsal view.
Mesoscutum width index (MsWI): MsW/ HW × 100
Mesoscutum height (MsH): mesoscutal height on a line which passes through a mesopleural gland
orifice and intersect at right angle with the line joining ventral-most point of pronotum and
propodeal spiracle.
Mesoscutum height index (MsHI): MsH/ HW × 100.
Dorsal smooth area of A3 length (DA3L): same as worker (only queen).
Dorsal smooth area of A3 width (DA3W): same as worker (only queen).
Dorsal smooth area of A3 index (DA3I): same as worker (only queen).
Methods for morphological examinations
Usual observations were carried out under a stereoscopic microscope (OLYMPUS SZX12).
Measurements were made by a stereoscopic microscope (Nikon SMZ-U ZOOM 1:10) with a direct
reading digital micrometer apparatus (Kogaku Co.; the reading unit is 0.001 mm), and the values
were rounded to nearest 0.01 mm and presented in mm.
Observation with scanning electron microscope (JEOL JSM-6301F) was also carried out.
Specimens were treated in 99.9% ethanol at least for 24 hours to dehydrate, and sputter-coated
with gold before the observation.
Wings and male genitalia were photographed with a stereoscopic microscope using an
automatic photograph system (PM-10ADS) after treatments as below.
Fore and hind wings were removed from dry specimens mounted. The wings put on a slide for
examination.
Male genitalia with several anterior segments were removed from specimens which were
mounted or in spirit. The abdominal segments including genitalia were treated in 5% KOH for 15
minutes at 90°. Genitalia and subgenital plates were removed from abdominal segments after the
treatment. The KOH treatment was repeated once for the genitalia and subgenital plates. The
treated genitalia and subgenital plates were put under Neo-Shigaral (Shiga Kontyu Fukyusha) to
be photographed.
Statistical analysis was carried out with StatView 5.0 (SAS Institute Inc.) software. HW, EL,
DlO, and MsH were measured on 34 queens collected from 34 colonies from several locations.
Four Okinawan colonies were tentatively regarded as S. sp. 9 based on previous distribution
records (Terayama, 1999) to examine the morphological difference found between S. sp. 9 and S.
lewisi.
Material examined
To avoid confusion, only colony samples that include at least a worker and a queen collected
from a colony were used as materials for the present examination. Data of the materials examined
are as follows: colony codes are shown in brackets: the caste names are given as aq/aqq (alate
queen/queens), dq/dqq (dealate queen/queens), w/ww (worker/workers), and m/mm (male/males).
S. lewisi. Yoshimura collection: [998] 8 aqq, 5 dqq, 58 ww, 3 mm, Kawasaki, Kanagawa
Pref., 2.viii.2002, M.Yoshimura leg; [994] 1dq, 34 ww, same locality, 2.vii.2002, M.Yoshimura
leg.; [996] 3 dqq, 25 ww, same locality, 2.vii.2002, M.Yoshimura leg.; [995] 2 dqq, 39 ww, same
locality, 2.vii.2002, M.Yoshimura leg.; [989] 4 dqq, 9 ww, same locality, 2.vii.2002,
M.Yoshimura leg.; [672] 3 dqq, 15 ww, same locality, 25.viii.2000, M.Yoshimura leg.; [671] 1 dq,
42 ww, same locality, 25.viii.2000, M.Yoshimura leg.
Onoyama collection: [j6718-19] 1 dq, 1 w, Kanazawa University, Ishikawa Pref.,
10.viii.1990, S. Amano leg.; [j1047-52] 3 dqq, 3 ww, Manazurumisaki, Kanagawa Pref., 60 m alt.,
7.i.1973, K. Onoyama leg.; [j1186a-c] 2 dqq, 1 w, north of Kudacchi, Izu-oshima Island, Tokyo
Pref., 90 m alt., 28.xi.1974, K. Onoyama leg.; [j1187a-b] 1 dq, 1 w, same locality, 90 m alt.,
28.xi.1974, K. Onoyama leg.; [j2945-48] 2 aqq, 2 mm, Mitsune, Hachijoujima Island, Tokyo Pref.,
26.viii.1979, K. Masuko leg.; [j3547-49] 1 dq, 2 ww, Mt. Yourouzan, Yourou-cho, Gifu Pref., 150
m alt., 3.v.1979, K. Yamauchi leg.; [j3550-52] 1 dq, 2 ww, same locality, 560 m alt., 3.v.1979, K.
Yamauchi leg.; [j3565-68] 2 dqq, 2 ww, Kinkazan, Gifu Pref., 29.iii.1981, K. Kinomura leg.;
[j1370a-b] 1 aq, 1 w, Shizunami, Shizuoka Pref., 0 m alt., 12.viii.1975, K. Onoyama leg.; [j314a-c]
1 dq, 2 ww, Yoshidayama, Kyoto, Kyoto Pref., 90 m alt., 17.xii.1974, K. Onoyama leg.; [j316a-c]
1 aq, 1 dq, 1 w, same locality, 80 m alt., 7.i.1975, K. Onoyama leg.; [j1429a-b] 1 dq, 1 w, same
locality, 110 m alt., 25.xii.1975, K. Onoyama leg.; [j7550-54] 1 dq, 4 ww, Kyoutanabe, Kyoto
Pref., 80m alt., 19.ix.1993, K. Onoyama leg.; [j7625-35] 6 dqq, 5 ww, same locality, 80m alt.,
18.iv.1994, K. Onoyama leg.; [j7695-99] 2 dqq, 3 ww, same locality, 80m alt., 23.v.1994, K.
Onoyama leg.; [j7710-13] 2 dqq, 2 ww, same locality, 80m alt., 24.v.1994, K. Onoyama leg.;
[j9891-95] 1 dq, 21 ww, Mt. Yuzuriha, Awajishima Island, Hyogo Pref., 110 m alt., 6-viii-2003,
K. Onoyama leg.; [j1637a-b] 1 dq, 1 w, Hatakejima, Tanabe Bay, Wakayama Pref., 0 m alt.,
31.x.1973, K. Onoyama leg.; [j1671a-b] 1 dq, 1 w, same locality, 0 m alt., 1.xi.1973, K. Onoyama
leg.; [j1704a-b] 1 dq, 1 w, same locality, 0 m alt., 2.xi.1973, K. Onoyama leg.; [j7746-50] 2 dqq,
3 ww, Kii-oshima Island, Wakayama Pref., 1.iv.1976, K. Onoyama leg.; [j7787-88] 1 dq, 1 w,
same locality, 1.iv.1976, K. Onoyama leg.; [j5136-38] 1 dq, 2 ww, Hiroshima, Hiroshima Pref.,
30.iii.1985, K. Onoyama leg.; [j1929-30] 1 dq, 1 w, Hashirano, Iwakuni, Yamaguchi Pref.,
26.iii.1978, K. Onoyama leg.; [j1932-33] 1 dq, 1 w, same locality, 26.iii.1978, K. Onoyama leg.;
[j1989-90] 1 dq, 1 w, Oura, Tsushima Island, Nagasaki Pref., 170 m alt., 29.iii.1978, K. Onoyama
leg.; [j2065-66] 1 dq, 1 w, Tendou, Mt. Tatsura, Tsushima Island, Nagasaki Pref., 130 m alt.,
1.iv.1978, K. Onoyama leg.; [j1944-45] 1 dq, 1 w, Yobuko, Saga Pref., 10 m alt., 27.iii.1978, K.
Onoyama leg.
S. sp. 4. Yoshimura collection: [1006] 1 aq, 1 dq, 65 ww, 13 mm, Namie-machi, Futaba-gun,
Fukushima Pref., 11.viii.2002, M.Yoshimura leg.; [1005] 4 aqq, 8 ww, 1 m, same locality,
10.viii.2002, M.Yoshimura leg.; [1003] 1 dq, 44 ww, same locality, 9.viii.2002, M.Yoshimura
leg.; [1001] 1 dq, 7 ww, same locality, 9.viii.2002, M.Yoshimura leg.; [1000] 1 dq, 24 ww, same
locality, 9.viii.2002, M.Yoshimura leg.; [931] 1 dq, 12 ww, 1 m, same locality, 21.viii.2001,
M.Yoshimura leg.; [649] 1 aq, 4 dqq, 38 ww, 2 mm, Gozenyama, Ibaraki Pref., Japan,
23.viii.2000, M.Yoshimura leg.; [665] 1 dq, 22 ww, 2 mm, same locality, 23.viii.2000,
M.Yoshimura leg.; [670] 1 aq, 3 dqq, 65 ww, Kawasaki, Kanagawa Pref., 25.viii.2000,
M.Yoshimura leg.; [1159] 1 aq, 1 w, 6 mm, Matsumae, Shimane Pref., 29.viii.1985; [727] 1 dq, 14
ww, Manazurumisaki, Odawara, Kanagawa Pref., 30.iv.2001, M.Yoshimura leg.
Onoyama collection: [j2703-08] 3 mm, 3 ww, Kiyosumiyama, Chiba Pref., 25.vii.1980, K.

Masuko leg.; [j3579-81] 1 dq, 2 ww, Kinkazan, Gifu Pref., 11.ii.1983, K. Kinomura leg.;
[j8145-48] 6 dqq, 57 ww, Nagoya University., Aichi Pref., 4.iv.1992, K. Onoyama leg.; [j7373-83]
1 dq, 10 ww, Nagatani-ike, Seika-cho, Kyoto Pref., 90 m alt., 31.iii.1992, K. Onoyama leg.
S. sp. 9. Yoshimura collection: [1092] 3 dqq, 18 ww, Oppadake, Motobu, Okinawa I.,
27.vi.2002, M.Yoshimura leg.
Onoyama collection: [j4086-89] 2 dqq, 2 ww, Oku, Okinawa Island, 100 m alt., 4.iii.1984, K.
Onoyama leg.; [j4224-25] 1 dq, 1 w, Mt. Nishime, Okinawa Island, 380 m alt., 5.iii.1984, K.
Onoyama leg.; [j3427-28, 3685-86] 1 dq, 3 ww, Sueyoshi, Naha, Okinawa Island, 2.xii.1975, T.
Abe leg.
Strumigenys kumadori sp. nov.

[Japanese name: Kita-uroko-ari]
Strumigenys sp. 1: Sonobe, 1977.

Strumigenys sp.: Masuko et al., 1985: 11; Masuko, 1984.
Strumigenys lewisi: Munakata, 1972 [we confirmed his misidentification]; Bolton, 2000 (in part)
Strumigenys sp. 4: Onoyama et al., 1992; JADG, 2003a, 2003b, 2003c.
Worker (Figs 3-9). HL 0.67, HW 0.45, CI 67.7, ML 0.32, MI 48.5, SL 0.36, SI 80.5, DSA3L
0.15, DSA3W 0.19, DSA3I 127.3 (Holotype worker).
HL 0.62-0.67, HW 0.44-0.47 CI 69.0-72.3, ML 0.30-0.32, MI 46.0-50.5, SL 0.36-0.39, SI
79.4-86.6, DSA3L 0.13-0.15, DSA3W 0.17-0.20, DSA3I 113.6-140.0 (6 measured).
Ventrolateral margin of head at level of eye not extended outward. Antenna consisting of 6
segments. Fully closed mandible in full-face view curvilinear. On the mandible, a distinct, long
and spiniform preapical tooth present close to apical teeth. Apical teeth consisting of two distinct
spiniform teeth and three small intercalary teeth between them: basal one of the two spiniform
teeth longer than the apical one: basal one of the three intercalary teeth distinctly smaller than
apical two. With mesosoma in lateral view, the diameter of the excavated area of mesopleural
gland moderate, much less than the maximum width of the first coxa. Mesosoma except for
propodeal declivity without spongiform tissue. Propodeal declivity equipped with a broad and
conspicuous lamella; propodeal tooth very feeble and not sclerotized; posterior margin of the
lamella convex, and immediately under the propodeal tooth of the margin sometimes slightly
concave. Ventral margin of petiole in lateral view with longitudinal spongiform tissue. With
petiole in lateral view, anteriormost point of lateral spongiform lobe nearly reaches level of
anterior face of node.
Dorsal and lateral surfaces of pronotum entirely reticulate-punctate, sometimes with a small
patch above the fore coxa. Metapleuron and side of propodeum entirely smooth. Limbus distinct.
Abdominal tergite IV longitudinally sculptured at the basal portion, but not entirely covered.
With head in full-face view, a pair of distinct long flagellate hairs present on apicoscrobe;
posterior to the apicoscrobal hairs with laterally projecting distally plumose filiform hairs; anterior
to the apicoscrobal hairs without laterally projecting hair. With head in lateral view, dorsal surface
from level of eye to preoccipital margin with erect to reclinate ground-pilosity; hairs on
preoccipital margin distinctly differentiated from those on level of eye; from highest point of
vertex to preoccipital margin with the anteriorly directed ground-pilosity, which is very feebly
curved basely so that each hair is elevated and inclined upward away from the cephalic outline.
A pair of long, flagelliform hairs present on the pronotal humeri and mesonotum. Dorsum of hind
femur without short erect hairs, but with two or three (usually two) long erect flagellate hairs.
Dorsal surface of hind basitarsus with one freely projecting flagellate hair. The whole of the dorsal
surface of abdominal tergite IV with long filiform hairs. Basal portion of abdominal sternite IV
covered with matted hair-like tissue.
Body almost unicolorous, reddish brown to yellowish brown.
Queen (Figs 17-22, 41). HL 0.65-0.67, HW 0.47-0.50 CI 72.4-76.1, ML 0.29-0.33, MI

45.3-48.8, SL 0.36-0.38, SI 74.0-78.0, DlO 0.04, DlOI 7.49-8.79, EL 0.12-0.14, EI 25.7-29.1, HD
0.32-0.34, PrH 0.21-0.25, MsW 0.33-0.36, MsWI 66.8-75.6, MsH 0.12-0.15, MsHI 25.1-29.8,
DSA3L 0.13-0.16, DSA3W 0.22-0.23, DSA3I 135.8-169.1 (9 measured).
Generally similar to the worker with the usual caste differences. Head thicker than that of
queen of S. lewisi in lateral view. With head in full-face view, the ocelli distinctly developed,
situated at posterior 1/4 of the head. Eye relatively large. A distinct, long and spiniform preapical
tooth present close to apical teeth. Apical teeth consisting of two spiniform teeth and three small
intercalary teeth between them: basal one of the two spiniform teeth longer than another apical
one: basal one of the three intercalary teeth distinctly smaller than the apical two. With mesosoma
in lateral view, the highest point of the mesoscutum nearly situated on extension line of the
mesopleural wing process; mesopleural gland orifice distinct, but its maximum width not reaching
maximum width of the procoxa; the pits on the mesepisternum invisible. Metanotum in lateral
view slightly convex posteriorly. Propodeal spine developed and sclerotized, and under which the
lobe of spongiform tissue distinctly developed. With the spongiform tissue on propodeal declivity
in lateral view, its posterior margin concave under the propodeal spine, and the remaining convex
posteriorly. With mesoscutum in dorsal view, its anterior margin rounded, both lateral margins
weakly constricted at posterior 1/3, lateral corners by the constriction not strongly angular.
Transverse furrow on the mesoscutum weakly curved posteriorly. Mesoscutum wide (MsWI more
than 63), its width exceeding 3/4 of the head width in frontal view. With petiole in lateral view, the
lobe of spongiform tissue strongly developed.
Most of veins on both of the fore and hind wings absent or vestigial. Only costal (C) and
radial (R1) veins and r-rs cross vein clearly present on fore wing. Vestiges of the radial sector
(Rs), M+Cu, and cubital (Cu) veins sometimes visible as pigmented lines but not sclerotized. On
the hind wing, radial (R) vein present, but not reaching to costal margin; jugal lobe absent.
Head and mesosomal dorsum entirely reticulopunctate. Central part of mesepisternum and
most part of propodeum ventral to propodeal spiracle not punctate and smooth. Dorsal margin of
petiole reticulate- punctate. Dorsal surface of postpetiole not punctate and smooth. Limbs present
on abdominal tergite IV. Abdominal tergite IV longitudinally sculptured at the basal portion, but
sculpture not extended to posterior half.
Pairs of hairs on the pronotal humeri long and flagellate. Mesonotal dorsum with erect, and
straight or flagellate hairs. Dorsum of hind femur without short erect hairs, but with two or three
(usually two) long erect flagellate hairs. Dorsal surface of abdominal tergite IV with long filiform
hairs. Hair-like tissue on the basal portion of abdominal sternite IV dense. Fore and hind wings
densely hairy.
Body almost unicolorous, reddish brown to yellowish brown. Outer margins of ocelli
distinctly bordered by brown to black pigment on inside portions among the three.
Male (Figs 29-34, 42, 45-48). HL 0.46-0.48, HW 0.42-0.44, CI 87.3-95.6, SL 0.09-0.11, SI

21.9-24.9, DlO 0.05-0.07, EL 0.19-0.20, HD 0.34-0.36, PrH 0.23-0.26, MsW 0.40-0.43, MsWI
90.8-103.2, MsH 0.13-0.15, MsHI 30.7-35.4 (6 measured).
With head in full-face view, portion posterior to the eyes subglobose; anterior to the eyes
distinctly narrowed anteriorly. Ocelli distinct; the median ocellus situated about posterior 1/4 of
the head length, the lateral ocelli not reaching to the posterior border of the head. Eyes distinctly
developed and prominent, occupying central 1/3 of lateral margin of the head in full-face view.
Eye in lateral view broadened ventrally, and its outer margin expanded anteroventrally and
flattened posteriorly. Anterior tentorial pits indistinct. Anterior margin of the clypeus in full-face
view slightly convex, but nearly straight. Frontal carinae undeveloped and antennal insertions
exposed. Antennae long and filiform, consisting 13 segments. Scape short and broad. Pedicel
short and broadened apically. With mandible in full-face view, its apical portion abruptly curved
and narrowed; the basal lamella recognizable but very weakly projected; apical to the lamella
edentate. Mandible in lateral view very narrowly subtriangular. With labrum in full-face view; its
apical portion distinctly extended laterally; the distal lobes entirely reduced, and apical margin of
the labrum concave toward the midpoint. Palp formula 1, 1 (1 observed on SEM). Mesosoma in
lateral view shorter and higher than that of the queen. Mesoscutum distinctly developed and
strongly raised dorsally in lateral view. Mesoscutellum developed and extended posteriorly. With
the mesonotum in dorsal view, the median notal suture weakly impressed but mostly invisible; the
notauli weakly impressed; the parapsidal furrows distinctly impressed and continued to the distinct
transscutal suture, so that the axillae distinctly divided; anterior margin of the scuto-scutellar
suture distinctly sculptured longitudinally. With mesopleuron in lateral view, its anteroventral
margin distinctly more expanded than that of queen. Metanotum in lateral view slightly extended
posteriorly. With the propodeum in lateral view, a distinct spiracle situated at the midheight; the
posterior margin with distinct corner, but the spine or dent reduced; the lamella absent ventral to
the propodeal corner, even if its ventral portion with a carina along the propodeal declivity. With
the petiole in lateral view, the node more gently raised than that of worker and queen; the lateral
spongiform lobe entirely reduced; the longitudinal spongiform tissue feebly present. Ventral
surface of abdominal sternite III in lateral view usually with a distinct process and a weak lamella,
but rarely the process reduced. Abdominal segment IV in lateral view thicker than that of worker
and queen, the ventral expansion more gentle.
With genitalia in ventral view, the basal ring broader than long; lateral margins of the
parameral plate weakly concave; the cuspis of volsella distinctly shorter than the digitus. With
genitalia in lateral view, an anteriorly-directed process, such as the barb, present at apical 1/4 of its
ventral margin; the digitus of volsella gradually curved ventrally and not broadened at the corner.
Fore and hind wings similar to those of queen.
Head, pronotum, mesonotum, and metanotum entirely reticulate-punctate. Central part of
mesepisternum and most part of propodeum ventral to propodeal spiracle not punctate and smooth.
Dorsal margin of petiole reticulate-punctate. Dorsal surface of postpetiole not punctate and
smooth. Limbus absent.
Two pairs of standing filiform hairs present on the vertex. With head in lateral view, long and
frontally projecting hairs absent anterior to median ocellus. Mesonotum with long, erect, and
filiform to flagellar hairs. Dorsal surface of the petiole, abdominal tergite III and IV with sparse
filiform hairs.
Body almost unicolorous, blackish brown to reddish brown, legs same or lighter.
Type material. Holotype worker, Japan: Gozenyama, Ibaraki Pref., 11.viii.2002,

M.Yoshimura leg., specimen code [649-1]. (Type No. [OMNH TI 196], Osaka Museum of Natural
History). Paratypes. colony code [649] in Yoshimura collection:1 alate queen, 4 dealate queens,
37 workers, 2 males, same data as holotype.
Distribution. Japan: southern Hokkaido, Honshu, Kyushu; China: Peking; Korea; Taiwan
[We could not examine specimens collected from Korea; specimens collected in Peking (workers,
ix.1980, P. Hammond) and Taiwan (worker, Chuchin, Taipai, Taiwan, 13.vi.1992, C. C. Lin) were
examined].
Etymology. The species is named from the Japanese “Kumadori”, a traditional make-up for
the Kabuki actor.
Strumigenys lewisi Cameron

[Japanese name: Uroko-ari (=Minami-uroko-ari)]
Strumigenys lewisi Cameron, 1886: 229. Syntype workers and queen. Type locality: Japan, Nagasaki,
3.iii, George Lewis leg [syntype queen was examined].
Strumigenys lewisi: Ogata, 1991 [first description of male]; Bolton, 2000 (in part); JADG, 2003a,
2003b, 2003c.
Strumigenys sp. A: Onoyama, 1976.
Strumigenys sp. 9: Onoyama et al., 1992: JADG, 2003a, 2003b, 2003c.
Strumigenys sp. D: Terayama, 1999.
Worker (Figs 10-16). HL 0.65-0.71, HW 0.48-0.50 CI 69.8-76.0, ML 0.31-0.35, MI

46.9-50.0, SL 0.37-0.40, SI 76.4-82.0, DSA3L 0.12-0.13, DSA3W 0.19-0.20, DSA3I 140.0-155.6
(6 measured).
Ventrolateral margin of head at level of eye not extended outward. Antenna consisting of 6
segments. Fully closed mandible in full-face view curvilinear. On the mandible, a distinct, long
and spiniform preapical tooth present close to apical teeth. Apical teeth consisting of two distinct
spiniform teeth and three small intercalary teeth between them: basal of the two spiniform teeth
longer than the apical one: basal one of the three intercalary teeth distinctly smaller than apical
two. With mesosoma in lateral view, the diameter of the excavated area of mesopleural gland
moderate, much less than the maximum width of the first coxa. Mesosoma except for propodeal
declivity without spongiform tissue. Propodeal declivity equipped with a broad and conspicuous
lamella; propodeal tooth very feeble and not sclerotized; posterior margin of the lamella convex,
and immediately under the propodeal tooth of the margin sometimes slightly concave. Ventral
margin of petiole in lateral view with longitudinal spongiform tissue. With petiole in lateral view,
anteriormost point of lateral spongiform lobe nearly reaching level of anterior face of node.
Dorsal and lateral surfaces of pronotum entirely reticulate-punctate, sometimes with a small
patch above the fore coxa. Metapleuron and side of propodeum entirely smooth. Limbus distinct.
Abdominal tergite IV longitudinally sculptured at the basal portion, but not entirely covered.
With head in full-face view, a pair of long, curved distally plumose filiform hairs, rarely not
serrate, present on apicoscrobe; hairs posterior to the apicoscrobal hairs with shorter barbs,
laterally projecting filiform hairs; anterior to the apicoscrobal hairs without laterally projecting
hair. With head in lateral view, dorsal surface from level of eye to preoccipital margin with erect
to reclinate ground-pilosity; hair on vertex margin distinctly differentiated from that on level of
eye; from highest point of vertex to preoccipital margin with the anteriorly directed
ground-pilosity, which is very feebly curved basally so that each hair is elevated and inclined
upward away from the cephalic outline. A pair of hairs present on the pronotal humeri and
mesonotum; those on the humeri usually flagellate; those on the mesonotum usually short filiform
hairs but sometimes long curved. Dorsum of hind femur without short erect hairs, but with two or
three (usually two) long erect flagellate hairs. Dorsal surface of hind basitarsus with one freely
projecting flagellate hair. The whole of the dorsal surface of abdominal tergite IV with flagellate
hairs. Basal portion of abdominal sternite IV covered with matted hair-like tissue.
Queen (Figs 23-28, 43). HL 0.63-0.70, HW 0.45-0.52 CI 71.5-80.0, ML 0.29-0.33, MI

44.8-49.2, SL 0.33-0.38, SI 67.9-76.2, DlO 0-0.02, DlOI 0-4.94, EL 0.08-0.11, EI 16.6-21.6, HD
0.30-0.32, PrH 0.18-0.22, MsW 0.26-0.31, MsWI 50.4-60.6, MsH 0.07-0.13, MsHI 14.2-27.0,
DSA3L 0.12-0.15, DSA3W 0.19-0.24, DSA3I 146.9-188.7 (10 measured).
Generally similar to the worker with the usual caste differences. Head thinner than that of
queen of S. kumadori in lateral view. With head in full-face view, the ocelli weakly developed
situated at posterior 1/4 of the head with brown pigment around them, but ocelli often vestigial and
visible only with the pigments. Eye relatively small. A distinct, long spiniform preapical tooth
present close to apical teeth. Apical teeth consisting of two spiniform teeth and three, rarely two,
small intercalary teeth: basal one of the two spiniform teeth longer than another apical one: basal
one of the three (or two) intercalary teeth distinctly smaller than the apical two (or the apical one).
With mesosoma in lateral view, the highest point of the mesoscutum situated anterior to extension
line of the mesopleural wing process in most cases; mesopleural gland orifice distinct but its
maximum width not reaching maximum width of the procoxa; the pits on the mesepisternum
invisible. Metanotum in lateral view slightly convex posteriorly. Propodeal spine developed and
weakly sclerotized, and under which the lobe of spongiform tissue distinctly developed. With the
spongiform tissue on propodeal declivity in lateral view, its posterior margin weakly concave
under the propodeal spine. With mesoscutum in dorsal view, its anterior margin relatively sharp,
both lateral margins strongly constricted at posterior 1/3, lateral corners by the constriction
strongly angular. Transverse furrow on the mesoscutum nearly straight. Mesoscutum narrow
(MsWI less than 63), its width not reaching 3/4 of the head width in frontal view. With petiole in
lateral view, the lobe of spongiform tissue strongly developed.
Both of the fore and hind wings distinctly reduced in width at distal 1/2. Only costal (C) and
radial (R1) veins and r-rs cross vein clearly present on fore wing. Vestiges of the radial sector
(Rs), M+Cu, and cubital (Cu) veins sometimes visible as pigmented lines but not sclerotized. On
the hind wing, radial (R) vein present, reaching to costal margin and extended distally; jugal lobe
absent.
smooth. Limbus present on abdominal tergite IV. Abdominal tergite IV longitudinally sculptured
at the basal portion, but sculptures not extended to posterior half of the tergite.
Hairs on the pronotal humeri long and flagellate. Mesonotal dorsum with erect, straight or
flagellate hairs. Dorsum of hind femur without short erect hairs, but with 2 or 3 (usually 2) long
erect flagellate hairs. Dorsal surface of abdominal tergite IV with long filiform hairs. Hair-like
tissue on the basal portion of abdominal sternite IV developed. Fore and hind wings densely hairy.
Male (Figs 35-40, 44, 49-52). HL 0.44-0.46, HW 0.43-0.46, CI 95.9-101.3, SL 0.09-0.11, SI

20.8-23.0, DlO 0.05-0.06, EL 0.18-0.19, HD 0.36-0.37, PrH 0.21-0.22, MsW 0.40, MsWI
91.5-94.0, MsH 0.16-0.17, MsHI 35.1-38.7 (3 measured).
With head in full-face view, portion of posterior to the eyes subglobose; anterior to the eyes
distinctly narrowed anteriorly. Ocelli distinct; the median ocellus situated about posterior 1/4 of
the head length, the lateral ocelli not reaching to the posterior border of the head. Eyes distinctly
developed and prominent, occupying central 1/3 of lateral margin of the head in full-face view.
Eye in lateral view broadened ventrally, and its outer margin expanded anteroventrally and
flattened posteriorly. Anterior tentorial pits unclear. Anterior margin of the clypeus in full-face
view slightly convex, but nearly straight. Frontal carinae undeveloped and antennal insertions
exposed. Antennae long and filiform, consisting 13 segments. Scape short and broad. Pedicel
short and broadened apically. With mandible in full-face view, its apical portion gradually curved
and narrowed; the basal lamella distinctly recognized and strongly projected; apical to the lamella
edentate. Mandible in lateral view subtriangular, but broader than that of S. kumadori. With
labrum in full-face view; its apical portion distinctly extended laterally; the distal lobes entirely
reduced, and apical margin of the labrum concave toward the midpoint. Palp formula 1, 1 (1
observed on SEM). Mesosoma in lateral view shorter and higher than that of the queen.
Mesoscutum distinctly developed and strongly raised dorsally in lateral view. Mesoscutellum
developed and slightly extended posteriorly. With the mesonotum in dorsal view, the median notal
suture weakly impressed but mostly invisible; the notauli weakly impressed; the parapsidal
furrows impressed and continued to the distinct transscutal suture; anterior margin of the
scuto-scutellar suture distinctly sculptured longitudinally, but weak on the lateral portion, so that
the division of the axillae often indistinct. Metanotum in lateral view slightly extended posteriorly.
With the propodeum in lateral view, a distinct spiracle situated at the midheight; the posterior
margin with distinct corner, but the spine or dent reduced; the lamella absent ventral to the
propodeal corner, even if its ventral portion with a carina along the propodeal declivity. With the
petiole in lateral view, the node more gently raised than that of worker and queen; the lateral
spongiform lobe entirely reduced; the longitudinal spongiform tissue feebly present. Ventral
margin of abdominal sternite III without a distinct process except for its extreme anterior part.
Abdominal segment IV in lateral view thicker than that of worker and queen, the ventral expansion
more gentle.
With genitalia in ventral view, the basal ring broader than long; lateral margins of the
parameral plate weakly concave; the cuspis of volsella distinctly shorter than the digitus. With
genitalia in lateral view, an anteriorly directing process, such as the barb, present at apical 1/4 of
its ventral margin; the digitus of volsella abruptly curved ventrally and broadened at the corner.
Only costal (C) and radial (R1) veins and r-rs cross vain clearly present on fore wing.
Vestiges of the radial sector (Rs), M+Cu, and cubital (Cu) veins sometimes visible as pigmented
lines but not sclerotized. On the hind wing, radial (R) vein present, reaching costal margin and
extended distally; jugal lobe absent.
smooth. Limbus absent.
Two pairs of standing filiform hairs present on vertex. With head in lateral view, long and
frontally projecting hairs absent anterior to median ocellus. Mesonotum with long, erect, and
filiform to flagellar hairs present. Dorsal surface of the petiole, abdominal tergite III and IV with
sparse filiform hairs.
Body almost unicolorous, blackish brown to reddish brown, legs same or lighter.
Distribution. Japan: Honshu, Shikoku, Kyushu, Tsushima I., Yaku I., Amami Is, Okinawa
I., Ishigaki I., Iriomote I.; Korea; China; Taiwan.
DISCUSSION
Taxonomic status of Strumigenys kumadori sp. nov., S. lewisi, and S. sp. 9
In the present study, we confirmed that Strumigenys sp. 9 is within S. lewisi which has
polymorphic queens, and that S. kumadori has morphological differences distinct from S. lewisi.
Strumigenys sp. 9 is an intraspecific variation of S. lewisi, the queens of the former having
larger lateral ocelli and higher mesoscutum. In the previous studies, S. sp. 9 was distinguished
from S. lewisi by the thicker mesoscutum in the queen (Onoyama, 1976; Terayama, 1999; JADG,
2003a, 2003b, 2003c), and from S. kumadori by the smaller and indistinct lateral ocelli in the
queen. In our examination, however, S. sp. 9 was inseparable from S. lewisi by height (Fig. 53) and
width (Fig. 54) of the mesoscutum, and diameters of the ocelli (Fig. 55) and eyes (Fig. 56) in the
queens, though clearly separable from S. kumadori by three of those characters (Figs 54-56).
Large variation in S. lewisi (see below) probably had made the extension of this taxon confusing.
Strumigenys kumadori is distinguished from S. lewisi by the following characters: 1) in the
workers, all of two paired hairs on the apicoscrobe and anterior portion of the mesonotum are long
and distinctly flagellate in the former (Figs 5, 6), but both of the hairs are plumose-filiform or
filiform in the latter (Figs 12,13), 2) the eyes of the queens are relatively large in the former, but
relatively small in the latter (Fig. 56), 3) the lateral ocelli of queens are distinctly large (Figs 20,
55) with pigmented outer margins in the former, but relatively small (Figs 26, 55) or vestigial and
not distinctly bordered with pigment around them in the latter, 4) the mesoscutum of queen in
dorsal view is wide (MsWI>63) and weakly constricted at posterior 1/3, and lateral corners by the
constriction are not angular in the former, but narrow (MsWI<63) and strongly constricted, and the
lateral corners are angular in the latter, 5) the transverse furrow on mesoscutum of queen is curved
posteriorly in the former (Fig. 19), but nearly straight in the latter (Fig. 25), 6) fore wings of queen
are relatively broad in the former (Fig. 41), but are reduced and narrow in the latter (Fig. 43), 7) the
radial vein on the hind wing does not reach the costal margin in the former (Figs 41, 42), but does
so in the latter (Figs 43, 44), 8) the mandible of male is relatively narrow in the former (Figs 30,
32), but is relatively wide in the latter (Figs 36, 38), 9) on the male genitalia in lateral view, volsella
is gently curved and the corner is not broadened in the former (Fig. 46), but is abruptly curved and
the corner is distinctly broadened in the latter (Fig. 50).
In addition, there are not only morphological, but also geographical and ecological bases for
separating S. kumadori and S. lewisi. The distribution areas and sites of the two species overlap in
Honshu, Kyushu, and Korea (JADG, 2003a, 2003b, 2003c; Terayama, 1999), though we could not
examine specimens collected in Korea. We additionally confirm the presence of both species in
Taiwan. Nest habitats of the two species are the same or very similar (Masuko et al., 1985), and
we actually confirmed that both species were collected from a single leaf litter sample taken from
a quadrat of 0.5m x 0.5m. Therefore, these two have completely sympatric distribution.
Our collection data of the colonies suggest that polygyny is common in all sizes of colonies in
S. lewisi, while polygyny occurs in S. kumadori only in large colonies with more than 37 workers.
Masuko et al. (1985) reported that S. kumadori is predominantly monogynous, while S. lewisi is
predominantly polygynous and further reported (1999) that 17% of 96 colonies of S. kumadori
contained two or more queens regardless of the number of workers, and demonstrated that even in
polygynous colonies a single individual in each colony was fertile (i.e., functional monogyny).
Strumigenys lewisi and S. kumadori are very similar in worker characters so that they were
regarded as sibling species. Characters used by Bolton (2000) in his notes under the description
of S. godeffroyi and S. geminata also separate those two species from the other twelve species of
godeffroyi-complex (in Bolton, 2000). Both S. lewisi and S. kumadori can be separated from the
most similar species, S. geminata, by having a more feeble propodeal spine and the lateral surface
of pronotum completely reticulate-punctate or at most with a small smooth patch above the fore
coxa.
Latitudinal variations of S. lewisi
Two morphological indices, DlOI and MsHI, showed negative tendencies with latitudinal
gradient (Fig. 57). The relationship between DlOI and latitude (r=-0.614, n=34, P<0.0001) was
significant, although that between MsHI and latitude was not significant (r=-0.314, n=34,
P=0.071).
Reduction of the mesoscutum, ocelli, and wings suggest that the ability to fly is degenerated.
The degeneration in the queens gives us two hypotheses for S. lewisi; 1) a range of the dispersion
in northern population is smaller than that in southern population, 2) the northern population has
dispersed their genes relying on the flying ability of the males.
Morphological notes
We found that both S. lewisi and S. kumadori have three intercalary teeth which consist of
apical two small teeth and a basal, smaller denticle. Description of that state was not found in
previous studies.
In Bolton’s (1999, 2000) diagnosis for the genus Strumigenys, the numbers of intercalary teeth
is one or two. He also mentioned in his notes for comparing between the godeffroyi-group and
mayri-group that two is the usual number in the godeffroyi-group (Bolton, 2000). Some species
having three intercalary teeth are recorded in other species group in Bolton (2000), i.e.
horvathi-group and koningsbergeri-group, but neither diagnosis of the group agreed with
characters of S. kumadori and S. lewisi. The number of the intercalary teeth is an important
character to separate species group, therefore further study will be needed.
In addition, the usual state of hairs on apicoscrobe and mesonotum in S. lewisi did not agree
with those of godeffroyi-complex in subgroup A of the godeffroyi-group, although the complexes
were practically subdivided by Bolton (2000) to assist identification. Workers in the
godeffroyi-complex have flagellate hairs on both the apicoscrobe and mesonotum, but this state
was found in only S. kumadori. Both of the hairs are filiform in S. lewisi. Other character states
of S. lewisi agree with godeffroyi-complex, and hence none of the species-complexes agree with
the combination of the characters in S. lewisi.
In the present study, the characters in the queen have most distinctly separated between the
two species, S. kumadori and S. lewisi. The eyes, ocelli, mesonotum, and wings are useful
characters to distinguish these two species. Morphologies of queens may provide useful
taxonomic characters even in other species. While much developed, the male's mesosoma did not
provide useful characters to separate the two species, while queen's one did. The reduced male
mandible, however, provided useful distinguishing characters.
Fauna of the genus Strumigenys in Japan
Before the present study, 8 species and 2 undetermined species and a key to the 10 species
were provided for Japanese Strumigenys (JADG 2003c). Yoshimura & Onoyama (2003) added a
new record of S. godeffroyi Mayr, 1866 from Ogasawara Is. to the faunal list.
In the present study, the taxonomic status of the two undetermined species was clarified and
now the Japanese Strumigenys fauna includes 10 valid species as below. We also provide a new
key to species of Japanese Strumigenys which is a revised version of the key in JADG (2003c).
Species list of Japanese Strumigenys
S. emmae (Emery)
S. exilirhina Bolton
S. godeffroyi Mayr
S. kumadori sp. nov. (= S. sp. 4)
S. lacunosa Lin & Wu
S. lewisi Cameron (= S. sp. 9)
S. minutula Terayama & Kubota
S. solifontis Brown
S. stenorhina Bolton
S. strigatella Bolton
Key to species of the genus Strumigenys in Japan
1 Antenna with 4 segments; head in full-face view abruptly narrowed anteriorly at anterior 2/3
.................................................................................................................................. S. emmae
-- Antennae with 6 segments; head in full-face view gently narrowed anteriorly ...................... 2
2 Preapical tooth on the mandible reduced and not spiniform .................................. S. lacunosa
-- Preapical tooth on the mandible distinctly developed and spiniform...................................... 3
3 Lamella on propodeal declivity very narrow in lateral view; its maximum height below the
spiracle less than half the length of a propodeal spine ............................................................ 4
-- Lamella on propodeal declivity very broadly developed in lateral view; its maximum height
below the spiracle more than half the length of a propodeal spine ......................................... 5
4 Mandibles nearly straight and relatively long; ventral outline of head weakly arched in lateral
view .................................................................................................................... S. stenorhina
-- Mandibles curved and relatively short; ventral outline of head more strongly arched in lateral
view ......................................................................................................................S. exilirhina
5 Mandibles shorter than apical antennal segment, relatively strongly curved; spongiform
lamellae below propodeal spines well-developed; a relatively small species ........ S. minutula
-- Mandibles as long as or longer than apical antennal segment, relatively less-strongly curved;
spongiform lamellae below propodeal spines well-developed or not unusually developed;
larger species .......................................................................................................................... 6
6 Mesopleuron and sides of propodeum wholly finely punctate and opaque...........S. strigatella
-- Mesopleuron and sides of propodeum largely impunctate and smooth................................... 7
7 Mesosomal dorsum with dense decumbent hairs; lamella on propodeal declivity extended
above propodeal spine ......................................................................................... S. godeffroyi
-- Mesosomal dorsum with sparse or without decumbent hairs; lamella on propodeal declivity
not extended above propodeal spine ....................................................................................... 8
8 Outer margins of infradental propodeal lamellae each abruptly narrowed immediately below
propodeal spine; mandibles longer; a pair of hairs on pronotal humeri short and not flagellate;
mesosoma in dorsal view narrower........................................................................ S. solifontis
-- Outer margins of infradental propodeal lamellae not abruptly narrowed below the spines;
mandibles shorter; a pair of hairs on pronotal humeri long and flagellate; mesosoma in dorsal
view wider .............................................................................................................................. 9
9 Pairs of hairs on apicoscrobe and anterior portion of the mesonotum long and flagellate; lateral
ocelli of queen relatively large, bordered with black; eyes of queen relatively large, their
maximum length distinctly exceeding the length of propodeal spine; mesoscutum of queen in
dorsal view relatively wider (Fig. 19) .................................................... S. kumadori sp. nov.
-- Pairs of hairs on apicoscrobe and anterior portion of the mesonotum short and not flagellate;
lateral ocelli of queen relatively small, often indistinct; eyes of queen relatively small, their
maximum length close to that of propodeal spine; mesoscutum of queen in dorsal view
relatively narrower (Fig. 26) ........................................................................................S. lewisi
ACKNOWLEDGEMENTS
We would like to acknowledge Dr. Kazuo Ogata (Institute of Tropical Agriculture, Kyushu
University) for his survey and help to loan a type and reference specimens in The Natural History
Museum London and for his useful suggestions, Dr. Suzanne Ryder and Dr. George R. Else (The
Natural History Museum, London) for the loan of a type and reference specimens, Dr. Keiichi
Masuko (Senshu University), Dr. Katsusuke Yamauchi (Gifu University), and Mr. Yuzuru Kuboki
(Kyushu University) for offering material, Dr. Nobuo Kitamura (Obihiro University for
Agriculture and Veterinary Medicine) for his technical support for taking SEM photographs, Mr.
Keiki Tominaga, Ms. Makiko Tominaga (Fukushima Prefecture) and Ms. Mariko Yoshimura
(Fukuoka Prefecture) for their assistance in field surveys, and Ms. Takako Ikeda (Hokkaido
University) for her kind help.
This study was partially supported by a Grant-in-Aid for Scientific Research (C) (No.
15510190 to K. Onoyama) from the Japan Society for the Promotion of Science.
LITERATURE CITED
Baroni Urbani, C. & De Andrade, M.L. 1994. First description of fossil Dacetini ants with a
critical analysis of the current classification of the tribe. (Amber Collection Stuttgart:
Hymenoptera, Formicidae. VI: Dacetini.) Stuttgarter Beiträge zur Naturkunde Serie B
(Geologie und Paläontologie) 198: 1-65.
Bolton, B. 1994. Identification guide to the ant genera of the world. 222pp. Harvard University
Bolton, B. 1995. A new general catalogue of the ants of the world. 504 pp. Harvard University
Bolton, B. 1999. Ant genera of the tribe Dacetonini. Journal of Natural History 33: 1639-1689.
Bolton, B. 2000. The ant tribe Dacetini. Memoirs of the American Entomological Institute 65:
1-1028.
Cameron, P. 1886. On a new species of Strumigenys (S. lewisi) from Japan. Proceedings of the
Manchester Literary and Philosophical Society 25: 229-232.
Gauld, I. & Bolton, B. 1988. The Hymenoptera. xii + 322 pp. Oxford.
Huber, J.T. & Sharkey, M.J. 1993. Structure. Pp. 13-59 in: Goulet, H. & Huber, T.J. (eds).
Hymenoptera of the world: An identification guide to families. 668pp. Research Branch
Agriculture Canada Publication 1894/E, Ottawa.
Japanese Ant Database Group (JADG). 2003a. Ants of Japan. 224 pp. Gakken, Tokyo.
Japanese Ant Database Group (JADG). 2003b. Super Visual Encyclopedia. Ants of Japan.
196pp. Gakken, Tokyo (in Japanese).
Japanese Ant Database Group (JADG) 2003c. Ant image database 2003. CD-ROM.
Idengaku-Fukyukai (The Association for Propagation of the Knowledge of Genetics),
Mishima.
Masuko, K. 1984. Studies on the predatory biology of Oriental dacetine ants (Hymenoptera:
Formicidae): I. Some Japanese species of Strumigenys, Pentastruma, and Epitritus, and a
Malaysian Labidogenys, with special reference to hunting tactics in short-mandibulate forms.
Insect Sociaux 31: 429-451.
Masuko, K. 1999. The number of queens in colonies of Strumigenys sp. (kita-uroko-ari): the result
of collection and dissection. Ari 23: 4-7 (in Japanese).
Masuko, K., Yamaoka, H., Kannari, T. & Usuba, S. 1985. Ants of Mt. Kiyosumi (4). Kiyosumi
11: 9-12 (in Japanese).
Munakata, M. 1972. Records of a cicada, Melampsalta radiator, and an ant, Strumigenys lewisi
from Hokkaido. Kontyu , 40: 314 (in Japanese).
Ogata, K. 1991. A generic synopsis of the poneroid complex of the family Formicidae
(Hymenoptera). Part II. Subfamily Myrmicinae. Bulletin of the Institute of Tropical
Agriculture, Kyushu University 14: 61-149.
Onoyama, K. 1976. A preliminary study on the ant fauna of Okinawa-Ken, with taxonomic notes
(Japan; Hymenoptera: Formicidae). Pp. 121-141 in: Ikehara, S. (ed). Ecological studies of
nature conservation of the Ryukyu Islands - (II). 141pp. Naha, Okinawa.
Onoyama, K., Ogata, K. & Terayama, M. 1992. Genus Strumigenys. Pp. 63-66 in: The
Myrmecological Society of Japan Editorial Committee (ed.). A guide for the identification of
Japanese ants. III. Myrmicinae and supplement to Leptanillinae. (Hymenoptera:
Formicidae). 94pp. The Myrmecological Society of Japan, Tokyo (in Japanese).
Sonobe, R. 1977. Ant fauna of Miyagi Prefecture, Japan. Japanese Journal of Ecology 27:
111-116.
Terayama, M. 1999. Family Formicidae. Pp. 133-317 in: Yamane, S., Ikudome, S. & Terayama,
M. (eds). Identification guide to the Aculeata of the Nansei Islands, Japan. 831pp. Hokkaido
University Press, Sapporo.
Yoshimura, M. & Onoyama, K. 2003. A new record of a dacetine ant, Strumigenys godeffroyi
Mayr, 1866 (Hymenoptera: Formicidae) from Japan. Edaphologia 71: 9-10.
Figures 1-2 Illustration of measurements for Strumigenys, (1) head of worker in full-face view, (2)
mesosoma of queen in lateral view.
Figures 3-9 Worker of Strumigenys kumadori sp. nov., (3) head and body in lateral view, (4), head
in full-face view, (5) mesosoma in lateral view, (6) apicoscrobal hair, (7) abdominal segments I to
IV in lateral view, (8) mandible in oblique ventral view, (9) pronotum in dorsal view.
Figures 10-16 Worker of Strumigenys lewisi, (10) head and body in lateral view, (11) head in full
face view, (12) mesosoma in lateral view, (13) apicoscobal hair, (14) abdominal segments I to IV
in lateral view, (15), pronotum in oblique dorsal view, (16) mandible in oblique ventral view.
Figures 17-22 Queen of Strumigenys kumadori sp. nov., (17) head and body in lateral view, (18),
head in full-face view, (19) mesosoma in dorsal view, (20) ocellar area in frontal view, (21)
mandible in oblique ventral view, (22) abdominal segments I to IV in lateral view.
Figures 23-28 Queen of Strumigenys lewisi, (23) head and body in lateral view, (24) head in
full-face view, (25) mesosoma in dorsal view, (26) ocellar area in frontal view, (27) mandible in
oblique ventral view, (28) abdominal segments I to IV in lateral view.
Figures 29-34 Male of Strumigenys kumadori sp. nov., (29) head and body in lateral view, (30)
head in full-face view, (31) mesosoma in lateral view, (32) head in lateral view, (33) mesosoma in
dorsal view, (34) abdominal segments I to IV in lateral view.
Figures 35-40 Male of Strumigenys lewisi, (35) head and body in lateral view, (36) head in
full-face view, 37) mesosoma in lateral view, (38) head in lateral view, (39) mesosoma in dorsal
view, (40) abdominal segments I to IV in lateral view.
Figures 41-44 Wings of S. kumadori sp. nov. and S. lewisi, (41) queen, (42) male of S. kumadori
sp. nov., (43) queen, (44) male of S. lewisi.
Figures 45-52 Male genitalia of S. kumadori sp. nov. and S. lewisi, genitalia of S. kumadori sp.
nov. in (45) ventral, (46) lateral, (47) dorsal view, (48) subgenital plate of S. kumadori sp. nov.,
genitalia of S. lewisi in (49) ventral, (50) lateral, (51) dorsal view, (52) subgenital plate of S. lewisi.
Figure 53 Relationship between head width (HW) and mesoscutum height (MsH) among
Strumigenys kumadori sp. nov., S. lewisi and S. sp. 9 in queens. MsH of S. kumadori is relatively
higher than those of the latter two, but the index does not separate the species.
Figure 54 Relationship between head width (HW) and mesoscutum width (MsW) among
Strumigenys kumadori sp. nov., S. lewisi and S. sp. 9 in queens. MsW of S. kumadori is distinctly
wider than those of the latter two. The index does not distinguish between S. lewisi and S. sp. 9.
Figure 55 Relationship between head width (HW) and diameter of lateral ocelli (DlO) among
Strumigenys kumadori sp. nov., S. lewisi and S. sp. 9 in queens. The lateral ocelli of S. kumadori
is distinctly larger than those of the latter two. The index is inseparable between S. lewisi and S.
sp. 9.
Figure 56 Relationship between head width (HW) and eye length (EL) among Strumigenys
kumadori sp. nov., S. lewisi and S. sp. 9 in queens. The eye of S. kumadori is distinctly larger than
those of the latter two. The index is inseparable between S. lewisi and S. sp. 9.
Figure 57 Latitudinal variation of two indices, diameter of lateral ocelli (DlOI) and Mesoscutum
height (MsHI), in queen of Strumigenys lewisi. Both of the indices are standardized to 0-1, and are
shown as DlOI (S) and MsHI (S). Both of the indices showed negative correlation with latitude,
and that of DlOI is significant (r=-0.614, n=34, P<0.0001).

Advances in Ant System A Tics (Hymenoptera Formicidae) - 2007

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Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O.

EDWARD O. WILSON AND HIS CONTRIBUTIONS TO ANT SYSTEMATICS

A REVIEW OF THE ANT GENUS METAPONE FOREL

Key words: Hymenoptera, Formicidae, Metapone, new species, Madagascar, taxonomy,

KEY TO WORKERS OF METAPONE FROM MADAGASCAR

Metapone emersoni Gregg

Metapone madagascarica Gregg

Metapone vincimus, new species

Diagnosis - Similar to M. madagascarica but readily distinguished by the following

Type Material. – Holotype worker from MADAGASCAR, 30km N of Antalaha, 3km W

ANT DIVERSITY IN ARID AUSTRALIA: A SYSTEMATIC OVERVIEW

Australia North America Southern

DIVERSITY OF THE AUSTRALIAN ANT FAUNA

TOTAL 1119 3606 5920 5.3

DIVERSITY WITHIN MAJOR ARID GENERA

Table 3. Classification, distribution and diversity of major species-groups and complexes of

a subspecies of I. rufoniger, which belongs to an unrelated group; see below) is extremely

Table 4. Classification, distribution and diversity of species-groups of Melophorus. Figures are

Table 5. Classification, distribution and diversity of major species-groups and complexes of

consobrinus complex (14, S)

Table 6. Classification, distribution and diversity of major species-groups of Monomorium that

Table 7. Classification, distribution and diversity of major species-groups and complexes of

Appendix 1. Key to species-groups of Melophorus, based on minor workers unless otherwise

1. Hind tibia without a conspicuous apical spur ....................................................................... 2

3. Mesosoma glabrous .................................................................................................. Group G

8. Mesosoma entirely glabrous .................................................................................................. 9

21. Mesosoma barrel-shaped, without even a propodeal suture ............................hirsutus group

22. Head conspicuously sculptured, dull or feebly shiny........................................................... 23

HOW TO CONDUCT LARGE-SCALE TAXONOMIC REVISIONS IN FORMICIDAE

Key words: Hymenoptera, Formicidae, taxonomy, revisions, methodology.

The aims of this essay

Good taxonomy ultimately relies upon:

i the detection of unique characters that diagnose a taxon,

In short: the function of taxonomy is to establish identity.

INITIATION PHASE OF THE PROJECT

Select a group to study

Choice of a study group may depend upon:

i museums and other state or local institutions,

i unique characters that discriminate individual species,

First draft key

1 Antenna with 11 segments............................................................................................... 2

A few guidelines for key construction:

Culmination of initial phase

By this stage the following results have been achieved:

INTERMEDIATE PHASE OF THE PROJECT

i to build up a complete list of previously established names,

Xus aus Green

Xus bus Green

Xus cus (Schwarz)

Xus dus Blanc

Schwarz, E. 1888. Beschreibungen nicht-existierender Insekten Deutschlands: 1000 pp. Berlin.

1 Antenna with 11 segments............................................................................................... 2

Also the file covering the taxon is changed from

Xus cus (Schwarz)

1 Antenna with 11 segments............................................................................................... 2

1 Head four times longer than broad................................................................................ fus

2 Antenna with 11 segments............................................................................................... 3

1 Antenna with 11 segments............................................................................................... 2

2 Head four times longer than broad................................................................................ fus

iv The result of running the type-specimen through the key is ambivalent.

Finalizing the key

1 Antenna with 11 segments............................................................................................... 2