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Number 3432, 9 pp., 1 table February 27, 2004

Family-Group Names for Termites (Isoptera)

MICHAEL S. ENGEL1 AND KUMAR KRISHNA2

ABSTRACT
Thirty-nine available family-group names are identified within the insect order Isoptera
(termites). For all names the correct author, date, type genus, and combining stem are provided
for the first time. This nomenclatural compilation is done to stabilize the usage of family-
group names in the Isoptera in advance of a world catalog. Several problems of priority are
identified and discussed. The little understood subfamily Foraminitermitinae is diagnosed;
while generally believed by many authors to be a new, unnamed subfamily, it was in fact
established by Holmgren nearly a century ago. The subfamilies Syntermitinae and Sphaer-
otermitinae are newly proposed for the mandibulate genera of nasute termites and for Sphaer-
otermes, respectively. The classification of Isoptera is briefly outlined.

INTRODUCTION lowed, thereby jeopardizing the stability of


termite classification. In fact, the numerous
It has long been recognized that the cur-
rent usage of family-group names in the clas- errors that appear in Snyders (1949) catalog
sification of termites (Isoptera) is not fully in have been perpetuated by subsequent au-
accord with the rules of zoological nomen- thors, who, rather than checking the original
clature (ICZN, 1999). Not only has the au- literature and applying the rules of nomen-
thorship and date of many names been in- clature, have lifted from Snyder or from oth-
correctly stated (e.g., by Snyder, 1949) and er catalogs which were in turn derived from
the Principle of Coordination (ICZN, 1999: Snyder (e.g., Roonwal and Chhotani, 1989;
Art. 36) ignored, but in many instances more Chhotani, 1997).
critical issues of priority have not been fol- In our preparation of a revised catalog of

1 Department of Ecology and Evolutionary Biology, and Division of Entomology, Natural History Museum, Snow

Hall, 1460 Jayhawk Boulevard, University of Kansas, Lawrence, Kansas 660457523. e-mail: msengel@ku.edu
2 Division of Invertebrate Zoology, American Museum of Natural History; City College and the Graduate School,

City University of New York. e-mail: krishn@amnh.org

Copyright q American Museum of Natural History 2004 ISSN 0003-0082


2 AMERICAN MUSEUM NOVITATES NO. 3432

the world termite fauna (Krishna and Engel, stem: Termit. Note: Latreille (1805, 1810)
in prep.), we have gone to great lengths to subsequently changed the name of his fam-
attend to these nomenclatural difficulties ille to Termitinae but the name was made
(e.g., Engel and Krishna, 2001a, 2001b, in available in 1802 (ICZN, 1999: Art. 11.7).
press a & b; Engel et al., 2003). In order to 2. Calotermitinae Froggatt, 1897: 516.
bring the usage of family-group names into Type genus: Kalotermes Hagen, 1853 [Cal-
accord with ICZN (1999) rules, we provide otermes Hagen, 1858 is an unjustified emen-
herein a catalog of all such names that have dation: see Engel and Krishna, 2001a; ICZN,
been applied in the Isoptera. This has been 2002]. Name emended to Kalotermitinae
done to correct the various circulating errors, Froggatt, 1897, in accordance with ICZN
to highlight the most egregious problems, (1999: Art. 29.1). Combining stem: Kaloter-
and to lay a standardized foundation for the mit.
forthcoming catalog. In addition, we have 3. Glyptotermitinae Froggatt, 1897: 518.
provided systematic descriptions for three Type genus: Glyptotermes Froggatt, 1897.
subfamilies (two new), which are otherwise Combining stem: Glyptotermit.
not employed by isopterists today but must 4. Rhinotermitinae Froggatt, 1897: 518.
be validated prior to the publication of the Type genus: Rhinotermes Hagen, 1858.
catalog (Krishna and Engel, in prep.). We Combining stem: Rhinotermit.
have also included those published names 5. Heterotermitinae Froggatt, 1897: 550.
that are otherwise unavailable at the end of Type genus: Heterotermes Froggatt, 1897.
the listing of available names. It must be not- Combining stem: Heterotermit.
ed that additional unavailable names have 6. Mastotermitinae Desneux, 1904a: 284.
been used on internet sites and employed by Type genus: Mastotermes Froggatt, 1897.
GenBank, but none of these have met the Combining stem: Mastotermit.
criteria of valid publication. While being un- 7. Hodotermitini Desneux, 1904a: 284.
available, like the three Holmgren names
Type genus: Hodotermes Hagen, 1853. Com-
considered below, these names have not been
bining stem: Hodotermit.
published in the sense defined by the ICZN
8. Stolotermitinae Holmgren, 1910a: 285.
(1999) (unlike Holmgrens names which
Type genus: Stolotermes Hagen, 1858. Com-
were published in accordance with the
bining stem: Stolotermit. Note: This name
ICZN) and are therefore excluded from this
study. unfortunately has priority over Termopsidae
Holmgren, 1911, a fact discovered too late
TAXONOMIC CATALOG to be included in the petition to conserve
Termopsis and Termopsidae relative to Mas-
Below we list in order of priority all fam-
totermitidae (Engel et al., 2003). Therefore,
ily-group names proposed for termites. The
names are presented in their original forms, a second petition was prepared to the ICZN
regardless of present day rank or suffix. The to conserve Termopsidae relative to Stoloter-
type genus is provided along with the correct mitidae (Engel and Krishna, in press a), since
combining stem for composing a family- it would be destabilizing for nomenclature to
group name. Daggers () indicate names pro- use Stolotermitidae in place of what is today
posed for fossil taxa. Some names were be- referred to as Termopsidae.
lieved to have been nomina nuda by Snyder 9. Leucotermitinae Holmgren, 1910a: 285.
(1949) when they were first proposed; how- Type genus: Leucotermes Silvestri, 1901.
ever, prior to 1931 family-group names could Combining stem: Leucotermit.
be made available by simple formation from 10. Coptotermitinae Holmgren, 1910a:
an available genus-group name (ICZN, 1999: 285. Type genus: Coptotermes Wasmann,
Art. 12.2.4) and need not have had a formal 1896. Combining stem: Coptotermit. Note:
description. Proposed again as new in Holmgren (1910b).
11. Serritermitinae Holmgren, 1910a: 285.
AVAILABLE NAMES Type genus: Serritermes Wasmann, 1897.
1. Termitina Latreille, 1802: 293. Type ge- Combining stem: Serritermit.
nus: Termes Linnaeus, 1758. Combining 12. Termitogetoninae Holmgren, 1910a:
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 3

286. Type genus: Termitogeton Desneux, 22. Macrotermitinae Kemner, 1934: 69.
1904b. Combining stem: Termitogeton. Type genus: Macrotermes Holmgren, 1909.
13. Microcerotermitinae Holmgren, Combining stem: Macrotermit. Note: See
1910b: 145. Type genus: Microcerotermes comments for Acanthotermitinae (above).
Silvestri, 1901. Combining stem: Microcer- 23. Amitermitinae Kemner, 1934: 110.
otermit. Type genus: Amitermes Silvestri, 1901.
14. Eutermitinae Holmgren, 1910b: 146. Combining stem: Amitermit.
Type genus: Eutermes Heer, 1849. Combin- 24. Miro-capritermitinae Kemner, 1934:
ing stem: Eutermit. Note: This name has 166. Type genus: Mirocapritermes Holm-
priority over Nasutitermitinae, despite the gren, 1914. Combining stem: Mirocapriter-
synonymy of its type genus with Nasutiter- mit. Note: Although Kemner (1934) hy-
mes (ICZN, 1999: Art. 40.1). Eutermes phenated the name in its original spelling, the
should also take precedence over Nasutiter- ICZN (1999) does not allow hyphenation and
mes, compounding the confusion. Use of Eu- the family-group name must be considered a
termitinae in place of Nasutitermitinae, how- single word.
ever, would be destabilizing for termite no- 25. Nasutitermitinae Hare, 1937. Type ge-
menclature. A petition to the ICZN has been nus: Nasutitermes Dudley, 1890. Combining
prepared to conserve Nasutitermes and Na- stem: Nasutitermit. Note: See comments
sutitermitinae (Engel and Krishna, in press above on Eutermitinae.
b). Current usage should be maintained until 26. Electrotermitinae Emerson, 1942: 10.
an authoritative decision is rendered. Type genus: Electrotermes Rosen, 1913.
15. Termopsinae Holmgren, 1911: 35. Combining stem: Electrotermit.
Type genus: Termopsis Heer, 1849. Combin- 27. Porotermitinae Emerson, 1942: 10.
ing stem: Termops. Note: See comments on Type genus: Porotermes Hagen, 1858. Com-
Stolotermitinae (above). bining stem: Porotermit.
16. Psammotermitinae Holmgren, 1911: 28. Apicotermitinae Grasse and Noirot,
64. Type genus: Psammotermes Desneux, 1954 [1955]: 360. Type genus: Apicotermes
1902. Combining stem: Psammotermit. Holmgren, 1912. Combining stem: Apicoter-
17. Pseudomicrotermitinae Holmgren, mit.
1912: 5. Type genus: Pseudomicrotermes 29. Cubitermitini Weidner, 1956: 99. Type
Holmgren, 1912. Combining stem: Pseudom- genus: Cubitermes Wasmann, 1906. Combin-
icrotermit. ing stem: Cubitermit.
18. Foraminitermitinae Holmgren, 1912: 30. Mirotermitini Weidner, 1956: 99. Type
5. Type genus: Foraminitermes Holmgren, genus: Mirotermes Wasmann, 1897. Com-
1912. Combining stem: Foraminitermit. bining stem: Mirotermit.
19. Stylotermitinae Holmgren and Holm- 31. Capritermitini Weidner, 1956: 100.
gren, 1917: 141. Type genus: Stylotermes Type genus: Capritermes Wasmann, 1897.
Holmgren and Holmgren, 1917. Combining Combining stem: Capritermit.
stem: Stylotermit. 32. Indotermitidae Roonwal and Sen-Sar-
20. Arrhinotermitinae Sjostedt, 1926: 8. ma In Roonwal, 1958: 81. Type genus: In-
Type genus: Arrhinotermes Wasmann, 1902. dotermes Roonwal and Sen-Sarma In Roon-
Combining stem: Arrhinotermit. wal, 1958. Stem: Indotermit.
21. Acanthotermitinae Sjostedt, 1926: 8. 33. Cretatermitinae Emerson, 1968: 278.
Type genus: Acanthotermes Sjostedt, 1900. Type genus: Cretatermes Emerson, 1968.
Combining stem: Acanthotermit. Note: This Combining stem: Cretatermit.
name has priority over Macrotermitinae; 34. Prorhinotermitinae Quennedey and
however, a petition has been submitted to Deligne, 1975: 265. Type genus: Prorhino-
conserve the usage of Macrotermitinae (En- termes Silvestri, 1909. Combining stem:
gel and Krishna, 2001b), which has been ap- Prorhinotermit.
proved by the ICZN (2003). Macrotermitinae 35. Lutetiatermitinae Schluter, 1989: 61.
is to be used whenever Macrotermes and Type genus: Lutetiatermes Schluter, 1989.
Acanthotermes are placed into the same fam- Combining stem: Lutetiatermit.
ily-group taxon. 36. Carinatermitinae Krishna and Gri-
4 AMERICAN MUSEUM NOVITATES NO. 3432

maldi, 2000: 134. Type genus: Carinater- 11.7.1.1). Note: This name was proposed as
mes Krishna and Grimaldi, 2000. Combining new again in Holmgren (1910a) and Holm-
stem: Carinatermit. gren (1910b). [A genus-group name, Proter-
37. Archeorhinotermitinae Krishna and mes, was not proposed until 1910, at which
Grimaldi, 2003: 2. Type genus: Archeorhin- time Holmgren (1910a) did not include Pro-
otermes Krishna and Grimaldi, 2003. Com- termes in his Protermitidae, placing it instead
bining stem: Archeorhinotermit. in Metatermitidae (in fact, Protermes was
38. Syntermitinae Engel and Krishna, never included in Protermitidae). Protermi-
herein (see below). Type genus: Syntermes tidae was used exclusively for what Holm-
Holmgren, 1909. Combining stem: Synter- gren considered to be the basal termites.
mit. Thus, Protermitidae was not subsequently
39. Sphaerotermitinae Engel and Krishna, validated in the later publications of Holm-
herein (see below). Type genus: Sphaeroter- gren.]
mes Holmgren, 1912. Combining stem:
Sphaerotermit. SYSTEMATIC DESCRIPTIONS
Below we provide descriptions for three
UNAVAILABLE NAMES little understood subfamilies of Termitidae,
one hitherto little understood and two new.
Some authors (e.g., Banks, 1920; Roonwal
The Foraminitermitinae was originally pro-
and Chhotani, 1989) have erroneously equat-
posed by Holmgren (1912) but has not been
ed the following three names with some of
generally recognized. The genera Foramini-
the present families or subfamilies. All three
termes and Labritermes, however, have in-
are unavailable and therefore do not enter
creasingly been recognized as distinctive and
into zoological nomenclature and should not
worthy of exclusion from Termitinae (e.g.,
be used.
Krishna, 1963; Noirot, 2001; Bitsch and
1. Metatermitidae Holmgren, 1909: 99.
Noirot, 2002), where they had been placed.
Unavailable, as it was not based on an avail-
The subfamily is characterized here to make
able genus-group name (ICZN, 1999: Art.
its diagnostic characters more widely under-
11.7.1.1).
stood. Similarly, the mandibulate genera of
2. Mesotermitidae Holmgren, 1909: 100.
the nasute termites are excluded from Na-
Unavailable, as it did not include Mesoter-
sutitermitinae and a new subfamilial name is
mes Haase, 1890 (ICZN, 1999: Art. 11.7.1.1)
required to accommodate this group (pro-
and was therefore not based on a genus-
posed as Syntermitinae, below). Lastly,
group name. Note: This name was proposed
Sphaerotermes is excluded from the Macro-
as new again in Holmgren (1910a) and
termitinae owing to a unique combination of
Holmgren (1910b). [Mesotermes was an
plesiomorphic and apomorphic traits (see be-
available genus-group name at that time, but
low).
recognized by Handlirsch (1906) as a neu-
ropteran and not a termite. Holmgren (1909, Subfamily FORAMINITERMITINAE Holmgren
1910a) clearly did not intend this name to be
based on the fossil neuropteran Mesotermes Foraminitermitinae Holmgren, 1912: 5. Type ge-
but established it instead for a series of ex- nus: Foraminitermes Holmgren, 1912.
plicitly included, living termite genera (see Combining stem: Foraminitermit.
Holmgren, 1910a, 1910b), which he believed DIAGNOSIS [Derived from Krishna and Ad-
to occupy an intermediate phylogenetic po- ams (1982)]: Imago. Head densely pilose;
sition in his scheme of termite relationships; fontanelle punctiform, situated at tip of a
thus, Mesotermes was not included by infer- cornical projection. Forecoxa with longitu-
ence of the generic stem of Mesotermitidae dinal ridge, without protuberance; tibial
(ICZN, 1999: Art. 11.7.1.1) and the name spurs 322. Soldier. Left mandible with up-
was never made available.] per, inner cutting edge finely serrated, and
3. Protermitidae Holmgren, 1909: 100. with blunt teeth appearing as crenulations be-
Unavailable, as it was not based on an avail- low serrations; right mandible smooth, with-
able genus-group name (ICZN, 1999: Art. out teeth, base of molar region with one or
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 5

TABLE 1
Hierarchical Outline of Termite Classification
All family-group names indicated; synonymic names italicized.
6 AMERICAN MUSEUM NOVITATES NO. 3432

two very tiny, pointed, thornlike spines or the inner scaly membrane (except reduced in
projections. Labrum with hyaline tip. Work- Syntermes).
er. Gut with mixed segment absent; four INCLUDED GENERA: Syntermes Holmgren,
malpighian tubules separately attached at 1909; Cornitermes Wasmann, 1897; Procor-
transverse mesenteron-proctodeum junction, nitermes Emerson In Snyder, 1949; and La-
extending forward for short distance and biotermes Holmgren, 1912. Other mandibu-
then looping around toward hindgut; mal- late genera of Nasutitermitinae may eventu-
pighian tubules swollen basally; first proc- ally be included here as well.
todeal segment (P1) tubular and narrow prox-
imally, almost same diameter as midgut, and SPHAEROTERMITINAE, new subfamily
dilated and saclike distally, with malpighian
tubules forming jumbled mass (5 pseudom- TYPE GENUS: Sphaerotermes Holmgren,
alpighian knot of Noirot, 2001; Bitsch and 1912.
Noirot, 2002). DIAGNOSIS: Imago. Head with light, small,
INCLUDED GENERA: Foraminitermes Holm- punctiform fontanelle. Labrum uniformly
gren, 1912 and Labritermes Holmgren, 1914. sclerotized, transparent; postclypeus with
COMMENTS: This group was first recog- median line. Soldier. Head with sides round-
nized by Holmgren (1912), who had diffi- ed; inconspicuous fontanelle present in mid-
culty placing it with any other group of Ter- dle of head. Labrum with sides rounded, with
mitinae (referring to it also as the Foramin- hyaline point. Left mandible with a few ser-
itermes-reihe) (also recognized as distinc- rations and blunt tooth near base; right man-
tive by other authors; e.g., Krishna, 1963; dible with inner cutting-edge smooth. Pron-
Krishna and Adams, 1982; Noirot, 2001). otum much narrower than head; anterior and
Noirot (2001) suggested that Foraminitermes lateral margins even, without projecting
belonged to a new subfamily, while a family- spines. Worker. Right mandible with 67
group name had already been proposed for it ridges on molar plate. Labrum uniformly
nearly a century ago. sclerotized (Sands, 1998: lacking the trans-
verse apical band of sclerotization seen in
SYNTERMITINAE, new subfamily Macrotermitinae, e.g., Donovan et al., 2000);
postclypeus strongly convex, with distinct
TYPE GENUS: Syntermes Holmgren, 1909 median line. Gizzard with small pulvilli at-
(not 1910a as cited by Snyder, 1949; Con- tached posteriorly on columns, crenulated
stantino, 1995, 1998; and other authors). crests of columns I and II poorly developed;
DIAGNOSIS: Imago. Fontanelle generally setae of pulvilli poorly developed (Noirot,
large, round; slightly depressed or forming a 2001). Conspicuous, backward-oriented bris-
convex plate. Postclypeus moderately or tles at the P3-P4 (paunch-colon) junction
strongly convex, with a median line. Fore- (Noirot, 2001) (unique among the Isoptera).
and hindwing with short R1, sometimes INCLUDED GENERA: Includes only Sphaer-
R21R3 joining costal margin very near the otermes Holmgren, 1912.
suture. Pronotum with anterior margins pro- COMMENTS: This is a remarkable subfamily
jecting laterally into points in Syntermes; in sister to the Macrotermitinae (e.g., three
Procornitermes, Cornitermes and Labioter- nymphal instars, unique worker instar, male
mes anterolateral corners without points, workers larger than females) but unlike the
margins even. Arolium absent. Soldier. Na- macrotermitines, Sphaerotermes is not asso-
sus (frontal tube) well developed, tip flat; ciated with Termitomyces fungi and lacks the
frontal pore wide and conspicuous; well-de- transverse band of sclerotization across the
veloped, functioning biting-mandibles; la- labrum of the worker and the imago (Sands,
brum simple, narrow, with hyaline tip. 1998). While these are indeed plesiomor-
Worker. Gut with mixed segment doubled, phies relative to Macrotermitinae, Sphaero-
dual lobes overlapping the first proctodeal termitinae has some remarkable apomorphic
segment (P1) (except reduction to single in traits that support its recognition [e.g., unique
Procornitermes); enteric valve with spines in possession of conspicuous, backward-orient-
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 7

ed bristles at the P3-P4 (paunch-colon) junc- 25]: 152. Bruxelles [Brussels]: P. Wytsman.
tion: Noirot, 2001]. [publ. date: 15 November 1904]
Donovan, S.E., D.T. Jones, W.A. Sands, and P.
ACKNOWLEDGMENTS Eggleton. 2000. Morphological phylogenetics
of termites (Isoptera). Biological Journal of the
We are grateful to Valerie Krishna, David Linnean Society 70: 467513.
Dudley, P.H. 1890. The termites of the Isthmus of
Nickle, and Charles D. Michener for reviews
PanamaPart II. Transactions of the New York
of the manuscript. Support was provided by Academy of Sciences 9: 157180. [publication
a National Science Foundation grant to cat- date June 1890, not 28 April 1890 as cited by
alog the termites of the world (NSF DEB- Constantino (2002): results of 5 May 1890
9870097 to K. Krishna and D. Grimaldi). meeting are published on same page as Dud-
This is contribution number 3344 of the Di- leys article and date on top of facing page is
vision of Entomology, Natural History Mu- date of previous Academy meeting, not of pub-
seum and Biodiversity Research Center, Uni- lication; journal cover indicates an earliest pos-
versity of Kansas. sible publication date of June 1890]
Emerson, A.E. 1942. The relations of a relict
South African termite (Isoptera: Hodotermiti-
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