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Stud Neotrop Fauna & Environm (1999) Vol.

34: 114128

0165-0521/99/3402-0114$15.00
Swets & Zeitlinger

THE NEOTROPICAL GENUS NOTODIAPTOMUS KIEFER, 1936


(CALANOIDA: DIAPTOMIDAE): REDESCRIPTION OF THE TYPE
SPECIES NOTODIAPTOMUS DEITERSI (POPPE, 1891) AND
DESIGNATION OF A NEOTYPE
Edinaldo N. Santos-Silva1,2, Geoff A. Boxshall2 and Carlos E.F. Rocha3
1Department

of Aquatic Biology, INPA, Manaus, Brazil


of Zoology, The Natural History Museum, London, England
3Department of Zoology, USP, So Paulo, Brazil

2Department

ABSTRACT
Notodiaptomus Kiefer, 1936 is the most abundant and speciose genus of freshwater calanoids in the Neotropical
region. The type species, N. deitersi (Poppe, 1891) is in need of redescription but the type material is no longer
extant and the type locality no longer exists. In order to avoid possible confusion over the identity of the type species
of the genus, a neotype is designated. This is an adult from Baia Pedra Branca because this locality is near Cuiab,
in the State of Mato Grosso, Brazil, the original type locality. The neotype is described and this description forms
the basis for a complete diagnosis of the genus Notodiaptomus. Like most diaptomid genera Notodiaptomus is
characterized by prosome morphology, by the fifth legs in both sexes and by the geniculate antennule. Sexual
dimorphism in the rostrum is reported here and the homology of the modified setal elements on the geniculate right
antennule of the is analysed.
KEYWORDS: Notodiaptomus deitersi, neotype, taxonomy, Calanoida, freshwater, Mato Grosso, Brazil.

INTRODUCTION
The genus Notodiaptomus Kiefer, 1936 is the most
widely distributed and most species rich genus of
freshwater calanoids in the Neotropics. In 1983 Dussart and Defaye listed 28 species in this genus; the
number of nominal species is currently 39. As with
most diaptomids, species have been erected mainly
on morphological details of adult , with particular emphasis on the sexually dimorphic characters of
the leg 5, right antennule and the posterior part of the
prosome. Except for a few recently described spe-

cies, little attention has been given to . Descriptions are very variable regarding the level of detail
provided in the text as well as in the quality of illustrations. This makes identification very difficult. The
aim of this paper is to provide a thorough redescription to modern standards of accuracy, of the typespecies Notodiaptomus deitersi (Poppe, 1891), to
improve knowledge of the genus and to enhance understanding of the distribution of this poorly characterized species.

MATERIALS AND METHODS


Address correspondence to: E.N. Santos-Silva, Coordenao de Pesquisas em Biologia Aqutica, Instituto Nacional de Pesquisas da Amaznia, C.P. 478, 69011-970,
Manaus, AM, Brasil. E-mail: nelson@inpa.gov.br

The specimens were collected in the Baia Pedra Branca (15o5219S, 56o0835W), a locality in the vicinity of Cuiab, Mato Grosso, Brazil.

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI


Samples were taken using a plankton net (55 m
mesh), concentrated and fixed in the field with 6%
formalin. Specimens were examined whole or dissected as temporary preparations mounted in lactophenol. Pieces of broken coverslips were used to
prevent compression by the coverslip. Observations
were made using a differential interference contrast
microscope (Olympus BH-2) and all drawings were
made with the aid of camera lucida. All measurements were made with an ocular micrometer. Additional preparations were made for SEM studies (Phillips XL-30), following the protocol used by
Felgenhauer (1987) and Huys and Boxshall (1991).
The specimens were washed several times in distilled water, dehydrated through graded acetone and
critical-point dried. Samples were mounted on stubs
and coated with gold or gold-palladium.
Neotype: dissected on 20 slides and deposited
in the INPA (INPA CR 550a-t) was selected from
larger sample of and from Baia Pedra Branca. Additional material from the same locality and
date: 1 adult dissected on 20 slides (INPA CR
551a-t) and 231 (INPA CR 552) and 57
(INPA - CR 676) in alcohol; 20 and 20 in
the MZUSP (MZUSP 12.823); 20 and 20
in the NHM-London (NHM 1998.2389 and
1998.2398); 20 and 20 in the MNHM
Paris (MNHN-Cp1685); 20 and 20 in the
USNM Washington, DC (USNM 243686).
Abbreviations used in the text: INPA-CR-Instituto Nacional de Pesquisas da Amaznia, Seo Crustacea, Manaus; MZUSP-Museu de Zoologia da Universidade de So Paulo, So Paulo; NHM-The
Natural History Museum, London; MNHM-Musum
National dHistoire Naturelle, Paris; USNM-National Museum of Natural History, Washington, DC;
SEM-Scanning Electron Microscope; s-seta; ae-aesthetasc; cs-conical seta; vs-vestigial seta; ms-modified seta; p-process; N-number of actual segments;
A-ancestral segments.

RESULTS
Notodiaptomus deitersi (Poppe, 1891)
Neotype
An adult ; Baia Pedra Branca (15 o5219S,
56o0835W), Mato Grosso, Brazil, in the vicinity of
Cuiab, the original type locality. The description is

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supplemented by the examination of additional material, including , from the same locality.
Description
Length of neotype: excluding caudal setae: 1100 m,
(mean length 1098 m (N=20), standard deviation
(SD)27.6). Body (Fig.1A) smaller and more slender
than . Rostrum (Fig. 5D) with paired rostral filaments; asymmetrical with process on right side of
basal region; defined from frontal margin of dorsal
cephalic shield by complete suture; ornamented with
pair of sensillae adjacent to suture. Cephalosome
with incomplete suture dorsally. Fifth free pedigerous somite with small and more or less symmetrical
lateral wings, each with small sensilla at apex (Fig.
1A). In dorsal view, body widest at first free pedigerous somite (Fig. 1A). Sensillae distributed on prosomal somites as figured (Fig. 1A).
Urosome: (Fig. 1A) 5-segmented. Genital somite
symmetrical with one sensilla at right posterior corner and another on left side inserted more medially
than on right. Genital aperture at ventrolateral posterior corner of genital somite on left side. Anal somite
with weak operculum; two sensillae on each side
(Fig. 1A). Caudal rami symmetrical, longer than
wide, with 6 setae; setules along medial margin.
Right antennule: 22-segmented (Fig. 2AD); segments 13 to 18 (XV to XX) swollen and modified.
Segmentation pattern and setal armature presented in
Table 2. On segments 17 and 18 (XIX, XX), one of
those setal elements is probably a seta-like aesthetasc; on segment 20 (XXIVXXV), two setae are
inserted posteriorly; on 21 (XXVI), one is inserted
ventrally. Vestigial setae on segments 2 (III), 3 (V)
and 5 (VII) comprising circle of thin cuticle and
minute seta at centre of circle. Tips of large setae on
segments 3, 7, 9 and 14 blunt (Fig. 2B, see arrowheads). Seta on segment 19 very small (Fig. 2C).
Modified setae on segments 10 and 11 similar to each
other but different from those on segments 13, 17, 18
and 19 (Fig. 2C, D). Modified seta forming strong
process on segment 13 (Figs. 2D, 8A). Modified setae on segments 17, 18 and 19 similar (Fig. 2C, see
arrowheads). Segments 15 and 16 each with spinous
process on frontal margin (Figs. 2B, D).
Left antennule: 25-segmented (Fig. 3A, B); only
second and last segments compound, representing the
ancestral segments IIIV and XXVIIXXVIII respectively, armature of segments presented in Table
2. Insertion of one of those setae is posterior on seg-

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E.N. SANTOS-SILVA ET AL.

Fig. 1. Notodiaptomus deitersi neotype, adult : A, habitus, dorsal. Adult : B, habitus, dorsal.

ments 22 and 23 (XXIV, XXV), and ventral on segment 24. Tips of large setae on segments 3, 7, 9 and
14 blunt, as in right antennule (Fig. 3A). Segmentation and armature pattern different between right and
left antennules. Both with compound second (IIIV)

and last segments (XXVIIXXVIII). Right antennule


with additional fusions: segments 19 and 20 representing ancestral segments XXIXXIII and XXIV
XXV respectively. Armature pattern differing with
segments 13 and 15 of left antennule lacking aes-

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

117

Fig. 2. Notodiaptomus deitersi neotype, adult : A, right antennule, detail of last segment (ancestral XXVIIXXVIII); B, right antennule, ventral, arrowheads indicate blunt setae or, on proximal segments, vestigial setae; C, right antennule, detail of segments 17,
18 and 19 (ancestral segments XIX, XX and XXIXXIII respectively), anterior, arrowheads indicate modified elements around
geniculation; D, right antennule, detail of segments 10-16 (ancestral segments XIIXVII), ventral, arrowheads indicate spinuous
processes, and extra aesthetascs not present on left antennule.

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E.N. SANTOS-SILVA ET AL.

Fig. 3. Notodiaptomus deitersi neotype, adult : left antennule


drawn in two sections as indicated by asterisks, A, segments 117 (ancestral segments IXIX), anterior; B, segments 1825 (ancestral segments XXXXVII), anterior.
Blunt setae, conical setae and vestigial setae are indicated
by arrowheads.

thetascs and segments 13 to 18 each lacking proximal seta. Modified setae present on segments 10, 11,
13, 17, 18 and 19 unmodified in left antennule.
Smooth hyaline membrane present anteriorly on segment 20.
Antenna: biramous (Figs. 5E, 6A). Coxa bearing
inner seta. Basis with two setae at posterior inner

corner. Endopod 2-segmented (Fig. 5E): first segment with two setae on inner margin, ornamented
with oblique row of spinules on outer side and pore
between row of spinules and setae; compound second segment bilobed; inner lobe bearing 8 setae distally, outer lobe with 7 distal setae and patch of
spinules on outer margin. Exopod 7-segmented (Fig.
6A), segmentation and armature pattern: segment 1
(ancestral segment I), 1 seta (s); segment 2 (ancestral
segments IIIV), 3 s; segment 3 (V), 1 s; segment 4
(VI), 1 s; segment 5 (VII), 1 s; segment 6 (VIII), 1 s;
segment 7 (ancestral segments IX-X), 4 s (distal part
derived from segment X elongate, bearing 3 long apical setae).
Mandible: (Figs. 7C, E, F) with strongly sclerotized coxal gnathobase carrying prominent lobe on
caudal margin (Fig. 7E); cutting blade with acute
caudal and triangular subcaudal teeth, and group of 6
multicusped teeth, apicalmost bearing spinules, additional row of spinules present on dorsal side; single
dorsal seta located near apical margin (Fig. 7F). Mandibular palp (Fig. 7C) with basis bearing four setae
on inner margin, three inserted more distally. Endopod 2-segmented; first segment with lobe bearing
four setae; second with 9 distal setae (one reduced),
and two rows of spinules. Exopod 4-segmented with
1, 1, 1, 3 setal formula.
Maxillule: (Fig. 6B, C, D) with praecoxal arthrite
carrying 10 stout marginal spines, plus five sub-marginally (four of them more slender), ornamentation
comprising patch of spinules present sub-marginally
(Fig. 6C). Coxal epipodite with 9 setae. Coxal endite
with 4 distal setae. Basal exite represented by outer
seta. Proximal basal endite well defined, bearing 4
setae distally. Distal endite fused to basis, with 4
setae and row of marginal spinules. Endopod 1-segmented and bilobate: proximal lobe with 3 setae on
margin, distal lobe with 5 setae (Fig. 6D). Exopod
unsegmented, with 5 distal setae and row of spinules
on margin (Fig. 6D).
Maxilla: (Fig. 7A) with praecoxa and coxa medially fused, but separate laterally. Proximal praecoxal
endite with 5 setae and one small spine; distal endite
bearing 3 setae and row of spinules. Coxal endites
each carrying 3 setae plus row of spinules on distal
margin. Allobasis well developed, armed with 3 setae. Three free endopod segments bearing 5 setae in
total.
Maxilliped: (Fig. 5B) well developed, comprising
syncoxa, basis and six free endopod segments. Prae-

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

119

Fig. 4. Notodiaptomus deitersi neotype, adult : A, left first leg, posterior; B, left second leg, posterior; C, left third leg, posterior; D,
third exopod segment of third leg, anterior; E, third endopod segment of third leg, anterior; F, third exopod segment of second
leg, anterior; G, third endopod segment of second leg, anterior; H, first and second segments of second leg endopod showing
Schmeils organ, lateral.

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E.N. SANTOS-SILVA ET AL.

Fig. 5. Notodiaptomus deitersi neotype, adult : A, fourth leg, posterior, arrowhead indicates presence of seta on basis; B, left maxilliped, medial; C, adult rostrum, ventral; D, adult rostrum, ventral, arrowhead indicates process on basal region; E, Adult ,
right antenna endopod, insertion of exopod indicated by asterisk; F, third exopod segment of fourth leg, anterior; G, third
endopod segment of fourth leg, anterior. H, external genital area; AM, arthrodial membrane; AH, anterior hinge; OP, opercular
pad; LP, lateral process; GP, gonoporal slit; GS, gonoporal slit.

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

121

Fig. 6. Notodiaptomus deitersi neotype, adult : A, right antenna exopod; B, right maxillule; C, detail of right maxillule arthrite; D,
detail of right maxillule endopod and exopod. Adult : E, left fifth leg, posterior; F, fifth leg, detail of endopod.

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E.N. SANTOS-SILVA ET AL.

Fig. 7. Notodiaptomus deitersi neotype, adult : A, right maxilla; B, fifth leg, posterior. Arrowheads indicate: patch of tubercles on left
basis; patch of tubercles and outgrowth with a deep groove on right basis; acute outgrowth on first exopod segment. C, palp of
right mandible; D, detail of right fifth leg endopod and groove, posterior; E, right coxal gnathobase of mandible with cutting
blade; F, detail of right mandible cutting blade; G, detail of left fifth leg endopod and exopod segments, posterior.

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

coxal endite with one seta and row of spinules. Coxal


endites represented by 8 setae in 3 groups on medial
margin: setal formula 2, 3, 3 plus patches of spinules
each; distal inner angle of syncoxa produced into
rounded lobe with setules around margin. Basis with
3 setae on medial margin and subterminal rows of
spinules on one side and setules on other. Endopod
6-segmented; first segment reduced, setal formula: 2,
3, 2, 1 + 1, 4.
Swimming legs: (Figs. 4AH, 5A, F, G) biramous
with 3-segmented rami, except endopod 2-segmented in leg 1: legs scarcely ornamented except by setule rows along outer and inner margins of endopodal
segments; legs 24 with distal row of spines on anterior surface of third exopod and two rows on third
endopod (Figs. 4D-G, 5F, G). Schmeils organ
present on posterior surface of second endopodal segment of leg 2 (Fig. 4H). Outer seta present on posterior surface of basis of leg 4 (Fig. 5A). Spine and seta
formula as follows:
TABLE 1. Notodiaptomus deitersi, neotype . Spine and setaformula.

Leg 1
Leg 2
Leg 3
Leg 4

Coxa

Basis

Exopod

01
01
01
01

00
00
00
10

I1; 01; I,I,4


I1; I1; I,I,5
I1; I1; I,I,5
I1; I1; I,I,5

Endopod
0-1; 1,2,3
01, 02; 2,2,3
01; 02;2,2,3
01; 02; 2,2,3

Fifth legs: asymmetrical (Fig. 7B, D, G, 8C, D).


Right fifth leg (Fig. 7B, 8C, D) with rudimentary
praecoxa present; coxa with conical process directed
posteriorly and projecting over basis, process with
slender sensilla on tip. Basis with outgrowth on posterior surface and with deep oblique groove reaching
endopodal lobe, groove finely ornamented with
minute tubercles along its edges (Fig. 7D); inner margin with semicircular lamella covered with fine tubercles extending onto adjacent segment surface; setae on outer margin inserted anteriorly; distal inner
corner bearing endopodal lobe. Exopod 2-segmented; first exopod segment with acute sclerotized outgrowth posteriorly on distal margin projecting over
second exopod segment; distal outer corner produced
into triangular outgrowth; second segment with
curved ridge on posterior surface, lateral spine located in distal quarter of segment, row of spinules on
inner margin. Claw inserted distally, strong and
curved proximally, slightly curved distally with

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spinules along inner margin; endopodal lobe with


comb of spinules on inner anterior surface.
Left fifth leg: (Fig. 7B, G, 8D) not reaching right
first exopod segment; coxa with small conical process posteriorly, and triangular sensilla at outer distal
corner: basis with seta on outer margin, inner margin
slightly concave, ornamented with patch of tubercles. Exopod 2-segmented; first segment sub-triangular, outer margin curved, inner margin with semicircular outgrowth bearing long setules; second
segment ending in heavily chitinized digitiform process; inner margin with curved process carrying setules on margin and spinulose seta inserted distally on
anterior surface; anterior surface concave with patch
of spinules. Endopod 1-segmented, conical, with row
of spinules on inner distal margin.

Adult
Length: excluding caudal setae: 1125 m, (mean
length 1148 m (N=20), SD28.5). Body (Fig. 1B)
larger than . Rostrum (Fig. 5C) broader than in ;
symmetrical; with pair of sensillae at incomplete
proximal suture. Cephalosome with incomplete suture dorsally. Fifth free pedigerous somite symmetrical, with curved posterolateral wings, each with sensilla at tip (Fig. 1B). In dorsal view, body widest at
first free pedigerous somite (Fig. 1B). Sensillae distributed on prosomal somites as figured (Fig.1B).
Urosome: (Fig. 1B) 3-segmented. Dorsal: genital
double-somite nearly symmetrical, longer than other
somites together, swollen in anterior region with two
sensillae, one on each side; right posterior corner
expanded over next somite. Second urosomite small,
right and left posterior corners expanded laterally.
Anal somite with weakly developed operculum with
two sensillae on each side. Caudal rami symmetrical,
with setules along inner margin. External genital area
(Fig. 5H) ventral: delimited anteriorly by broad symmetrical opercular pad, and laterally by well developed, posteriorly-directed lateral processes. Paired
gonoporal plates and slits located adjacent to midline, between lateral processes. Extensive area of relatively flexible cuticle (arthrodial membrane?),
present anterior to opercular pad.
Antennules: symmetrical; similar to left antennule (Table 2). All other appendages as in except
leg 5. Leg 5 (Fig. 6E, F) symmetrical; coxa with
conical posterior process in distal left corner with
triangular sensilla at tip; basis sub-triangular, outer

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E.N. SANTOS-SILVA ET AL.

Fig. 8. Notodiaptomus deitersi neotype, adult , SEM pictures: A, detail of modified seta forming strong process on segment 13
(ancestral segment XV); B, detail of the segments around the geniculation of the right antennule, arrows indicate the modified
elements; C, fifth leg, detail of the right endopod; D, fifth leg, lateral, arrows indicate the tip of the left endopod, the outgrowth
in the posterior surface of the left basis and the deep oblique groove.

margin smaller than inner margin with long seta extending beyond distal margin of first exopod; first
exopod segment larger than second; second segment
with lateral spine; distinct third exopod segment with
two terminal setae, lateral smaller, not reaching beyond middle of medial. Terminal claw with medial
and lateral rows of denticles. Endopod 1-segmented
bearing two setae on oblique tip plus row of spines
subterminally on anterior surface.
Remarks
Notodiaptomus deitersi (Poppe, 1891), was briefly
described by Poppe. It is possible that there has been
some confusion with subsequent identification of

Notodiaptomus deitersi. This species was recorded


from Lake Parnagua, Piaui in the northeastern Brazil
by Spandl (1926), and from the lakes Recreio and S
Mariana, Mato Grosso, Brazil by Matsumura-Tundisi
(1986); from Paraguay by Lowndes (1934) and from
Argentina by Brehm (1959), Ringuelet and Martinez
de Ferrato (1967). The material reported by Lowndes,
Brehm and Matsumura-Tundisi differs in some characters from Notodiaptomus deitersi sensu Poppe,
1891 as described by Spandl (1926).
The only feature clearly figured by Poppe (1891)
in the original description, that we can not confirm in
the material examined, is the presence of 2-segmented endopod in leg 5 of the . We have observed a

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

discontinuity in the cuticle of the inner margin of the


endopod which presumably indicates the plane of fusion of the endopod segments. All other features described by Poppe (1891) are present in the specimens
observed.
Some authors, when reporting on material identified as N. deitersi (Poppe, 1891), have presented descriptions that match Poppes in most details (Spandl,
1926) but others report features that differ from
Poppes (1891) original description. Lowndes
(1934), studying material from Paraguay, drew attention to the pyramidal projection on the penultimate somite of the body which was covered with
minute spinules, to the presence of hyaline membrane
on the antepenultimate segment of the right antennule, and to a spine (treated here as conical setae) as
long as those on segments 10 and 11. He also found
that the had a 2-segmented endopod in the fifth
leg. Brehm (1959) noted several features that agree
with Lowndes (1934), but he didnt observe the pyramidal projection in the penultimate somite of the
body, only a patch of spinules. He reported a semicircular membrane on the basis of right fifth leg of
and also a strongly chitinized region in the distal
inner margin of the second exopod segment of the
same leg. It is remarkable that Brehm stated that the
specimens observed by him are closely related to N.
venezolanus. Ringuelet and Martinez de Ferrato
(1967) also found the pyramidal projection covered
by spinules on the fourth somite of the body and 2segmented endopod in the fifth leg. MatsumuraTundisi (1986) figured the pyramidal projection on
the fourth prosomal somite covered by spinules,
but found no hyaline lamella or process on the antepenultimate segment of the right antennule. The
endopod of the fifth leg was 1-segmented.
Given this degree of inconsistency in morphological accounts of N. deitersi we consider that it is premature to summarise existing distribution records.
Only the examination of material previously identified as N. deitersi will provide verification that these
identifications match Poppes description (1891) and
the neotype designation presented here.
Notodiaptomus Kiefer, 1936
Diagnosis
Diaptomidae, Diaptominae. and appendages
similar except right antennule and leg 5. Rostrum
sexually dimorphic. and left antennule 25-seg-

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mented, exhibiting similar segmentation pattern (all


ancestral segments expressed except IIIV and
XXVIIXXVIII) and setal armature; lacking aesthetascs on segments 13 and 15 (XV and XVII) and
setae on segments 13 to 18 (XVXX) present in .
and antennules with vestigial setae on ancestral
segments III, V and VII; large setae on segments 3,
7, 9 and 14 (V, IX, XI and XVI) each with blunt tip;
conical setae on segments 8 and 12 (X and XIV).
Antenna endopod 2-segmented; first segment with
oblique row of spinules and conspicuous pore: exopod 7-segmented; segments 2 and 7 representing ancestral segments IIV and IXX respectively. Swimming legs poorly ornamented, biramous with
3-segmented rami, except endopod 2-segmented in
leg 1; legs 14 with lateral spine on each exopod
segment, except leg 1 lacking spine on exopod segment 2; legs 24 ornamented with distal row of
spinules on anterior surface of third exopod and two
rows on third endopod segment. Schmeils organ
present on posterior surface of second endopod segment of leg 2. Leg 4 with seta on posterior surface of
basis. : cephalosome with incomplete suture dorsally; rostrum asymmetrical with process on right
side of basal region; nearly symmetrical lateral wings
on fifth free pedigerous somite; urosome 5-segmented; caudal rami symmetrical, longer than wide, with
setules along medial margin; right antennule modified, 22-segmented; segmentation pattern (ancestral
segments expressed except IIIV, XXIXXII,
XXIVXXV and XXVIIXXVIII) and setal armature of segments different from left and antennules (Table 2). On segments 17 and 18 (XIX, XX),
one of those setae is probably a seta-like aesthetasc;
on segment 20 (XXIVXXV), two setae are inserted
posteriorly; on segment 21 (XXVI), one seta inserted
ventrally; segments 13 to 18 (XV to XX) swollen and
modified; small spinous processes present on segments 15 and 16 (XVII and XVIII); conical setae
asE; modified setae on segments 10, 11, 13, 17, 18
and 19 (XII, XIII, XV, XIX, XX and XXI); very
small setae on segment 19 (XXII); modified setae on
segments 10, 11 and 13 forming processes, that on 13
stronger and more robust than those on segments 10
and 11; modified setae on segments 17, 18 and 19 are
similar among them, but differing from those on segments 10 and 11. One of those 2 setal elements on
segments 17 and 18 is probably a seta-like aesthetasc.
Left antennule 25-segmented; second and last segments compound; armature is presented in Table 2.

E.N. SANTOS-SILVA ET AL.

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TABLE 2. Segmentation pattern and armature of antennules in the genus Notodiaptomus Kiefer, 1936 based primarily on the type
species but including known interspecific variation within the genus. A, ancestral segments; N, number of actual segments;
s, seta; ae, aesthetasc; vs, vestigial seta; ms, modified seta; p, process. (*) shows the armature observed in the neotype.
A
I
II
III
IV
V
VI
VII
VIII
IX
X
XI
XII
XIII
XIV
XV
XVI
XVII
XVIII
XIX
XX
XXI
XXII
XXIII
XXIV
XXV
XXVI
XXVII
XXVIII

right antennule

1s+1ae

2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18

3s,1ae,1vs
1s,1vs
1s
1s,1ae,1vs
1s
1s,1ae
1s,1cs
2s,1ae
1s,1ms
1s,1ms
1s,1ae,1cs
1s,1ae,1ms
2s,1ae
2s,1ae,1p
2s,1ae,1p
1 or 2s(*),1ms
2s,1ms

19

2s,1ae,2ms

20
21

4s
2s

22

4s,1ae

One of those setae on segments 22 and 23 (XXV) is


inserted posteriorly, and ventrally on segment 24;
lacking aesthetasc on segments 13 and 15.
Fifth leg asymmetrical; rudimentary praecoxa
present; coxa with conical process projecting over
basis with slender sensilla on tip; outgrowth on posterior basal surface with deep oblique groove ornamented with minute tubercles along edge; semicircular lamella on inner margin of basis covered with fine
tubercles extending onto adjacent segment surface;
endopodal lobe on inner corner of basis. Exopod 2segmented; acute sclerotized outgrowth present on
distal margin of first exopod segment and distal corner produced into triangular outgrowth; second segment with curved ridge on posterior surface, lateral
spine in distal third of segment; claw strong and
curved proximally, with row of spinules along inner
margin. Endopodal lobe with comb of spinules on inner anterior surface. Left fifth leg; coxa with conical
process posteriorly with triangular sensilla on apex;
inner margin of basis with patch of tubercles.
larger than ; all appendages, except leg 5 and
right antennule, as in ; rostrum broad, symmetrical

left antennule

antennules

1s+1ae

1s+1ae

2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24

3s,1ae,1vs
1s,1vs
1s
1s,1ae,1vs
1s
1s,1ae
1s,1cs
2s,1ae
1s
1(*) or 2s
1s,1ae,1cs
1s
1s,1ae
1s
1s,1ae
1s
1s
1s,1ae
1s
1s
2s
2s
2s

3s,1ae,1vs
1s,1vs
1s
1s,1ae,1vs
1s
1s,1ae
1s,1cs
2s,1ae
1s
1 or 2s
1s,1ae,1cs
1s
1s,1ae
1s
1s,1ae
1s
1s
1s,1ae
1s
1s
2s
2s
2s

25

4s,1ae

4s,1ae

without process on basal region; symmetrical lateral


wings on fifth free pedigerous somite; urosome 3segmented; genital double-somite inflated anteriorly
and nearly symmetrical; anal somite with weakly developed operculum; caudal rami symmetrical with
setules along inner margin; external genital area with
prominent lateral processes on opercular pad; antennules symmetrical 25-segmented similar to left antennule of ; leg 5 symmetrical, coxa with conical
process with sensilla at tip, basis subtriangular with
long setae reaching beyond distal margin of first exopod, first exopod segment larger than second, second
with lateral spine, distinct third exopod segment
small, with two terminal setae, lateral smaller; terminal claw with denticles on medial and lateral margins, endopod 1-segmented with two setae and oblique comb of spines subterminally on anterior
surface.
Remarks
The genus Notodiaptomus was established by Kiefer
(1936) to accommodate eleven species originally
placed in Diaptomus Westwood, 1836 sensu lato.

REDESCRIPTION OF NOTODIAPTOMUS DEITERSI

Four of these species, Notodiaptomus nordestinus


(Wright, 1935), N. henseni (Dahl, 1894), N. iheringi
(Wright, 1935) and N. deitersi (Poppe, 1891) had
previously been considered part of nordestinus-group
created by Wright (1935); the other seven added by
Kiefer (1936) were N. amazonicus (Wright, 1935),
N. cearensis (Wright, 1936), N. santaremensis
(Wright, 1927), N. carteri (Lowndes, 1934), N. anisitsi (Daday, 1905), N. incompositus (Brian, 1925)
and N. inflatus (Kiefer, 1933). Kiefer did not provide
a formal diagnosis to the new genus but grouped
these species on the basis of the following features:
the nature of the final segment of the left exopod
of the fifth leg;
the geniculate antennule carried conspicuous
spines on segments 10, 11, 13, 14 and 16, which
were very similar in all species;
the antepenultimate segment of the geniculate
antennule is ornamented with, at most, one hyaline membrane.
Kiefer (1936) considered the nature of the left exopod on the fifth leg to be the most significant
character.
Kiefer did not designate a type species for the
genus Notodiaptomus. In the absence of an original
designation there has been some confusion about
the type of the genus. Ringuelet (1958) formally
designated Diaptomus deitersi Poppe, 1891 as the
genotype of the genus Notodiaptomus. Under the
International Code of Zoological Nomenclature this
subsequent designation is valid. Then, Dussart &
Defaye (1983) proposed that par souci de priorit,
cest N. gibber (Poppe, 1889) qui pourrait tre prise
comme espce-type. But Diaptomus gibber was
transferred to Notodiaptomus by Pallares in 1963,
several years after the creation of the genus, and was
not originally included in Notodiaptomus by Kiefer
(1936). Consequently, following Article 67(g) of the
Code it can not be accepted as the type of the genus.
So, the designation by Ringuelet (1958) of Diaptomus
deitersi as type species of the genus Notodiaptomus is
valid.

DISCUSSION
Sexual dimorphism in the rostrum of calanoid copepods has been observed and figured by many other
authors. In the family Aetideidae, for instance, there
are extreme examples of this sexual dimorphism; in

127

Batheuchaeta Brodsky, 1950 for example, has the


rostrum well developed in , but absent in
(Markhaseva, 1996). In the family Diaptomidae some
authors have noted or figured the dimorphic rostra.
Brandorff (1973), Dussart & Robertson (1984), Reid
(1985), Reddy (1994) and Santos-Silva et al. (1996),
for example, all figured differences in and rostra but didnt discuss the significance of this. It appears that this dimorphism may be widespread in the
family since it is known in Notodiaptomus and Mastigodiaptomus Light, 1939 (Santos-Silva et al., 1996)
and in Neodiaptomus Kiefer, 1932, Arctodiaptomus
Kiefer, 1932 and Sinodiaptomus Kiefer, 1932 (Reddy, 1994). The extent of expression of this sexual
dimorphism across the genera of the family Diaptomidae, remains to be established.
The geniculate right antennule of diaptomids
provides many characters of taxonomic importance
at both specific and generic levels. The modifications of setal elements on the segments either side of
the geniculation (Fig. 2C) are probably highly conserved and of little significance, since they correspond closely to modifications known in other calanoid families. These may be deep rooted calanoid
characters. Similarly, the 10 proximal segments appear highly conserved and all species of Notodiaptomus thus far studied exhibit identical setation, including the possession of the three vestigial setae
(Santos-Silva, unpublished). The most informative
section of the right antennule lies between segments 10 (XII) and 16 (XVIII) (Fig. 2D). Setal elements are modified on segments 10, 11 and 13, and
the precise apomorphic form of each element can be
important. Segments 15 (XVII) and 16 (XVIII) have
novel spinous processes on the frontal margin; their
presence and relative size can be important. The form
of the long setae on segments 3(V), 7(IX), 9(XI) and
14(XVI) may also be significant since the apex appears to be strongly rounded in some species of Notodiaptomus.
Preliminary studies of six Notodiaptomus species
have revealed that the basic setal formula of the
and left antennule is identical to that of N. deitersi,
with one exception. Segment 11 (XIII) commonly
carries one seta only, as in N. deitersi, but in some
congeners, such as N. amazonicus (Wright, 1935) and
N. cearensis (Wright, 1936), two setae are present.
The loss of the proximal seta on this segment may be
a useful apomorphy in phylogenetic analysis of Notodiaptomus, and even other genera of Diaptomidae.

128

E.N. SANTOS-SILVA ET AL.

ACKNOWLEDGEMENTS
This research has been supported by CAPES and PDEE
grant BEX0290/98-0. We are grateful to Rony Huys for his
guidance in drawing techniques and for numerous discussions; to Danielle Defaye (Musum National dHistoire
Naturelle, Paris) for her comments on genital structures
and on the manuscript; to Alex Ball and Chris Jones from
the Electron Microscope Unit/The Natural History Museum for their assistance and; to Vangil Pinto da Silva from
the University of Mato Grosso who kindly collected and
provided the material.

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Received: 12 October 1998


Accepted: 24 November 1998