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New species of Cletocamptus and a new and fully
illustrated record of C. sinaloensis (Copepoda:
Harpacticoida) from Brazil
SAMUEL GO
´MEZ
Instituto de Ciencias del Mar y Limnologı´a, Mazatla´n, Sinaloa, Mexico
(Accepted 14 July 2005)
Abstract
Two new species of harpacticoid copepods, Cletocamptus levis sp. nov. and Cletocamptus nudus sp. nov.
are described from Brazil. Cletocamptus nudus sp. nov. seems to be closely related to C. schmidti based
on the armature formula of mouth parts, P3 and P4, but can be separated based on the
ornamentation of the anal operculum, relative length of exopodal and baseoendopodal setae of the
female P5, and length ratio of the female P1 EXP:ENP. Cletocamptus levis sp. nov. seems to be more
closely related to C.axi by the combination of a P1 ENP shorter than the P1 EXP, the armature
formula of the mandibular palp, length: width ratio of caudal rami, and relative length of the female
P3 ENP 2, male P2–P4 EXP 1, and number of segments of the male P3 ENP. It can be separated by
the spinule ornamentation on the anal operculum, shape of female rostrum, relative length of outer
spinules of female P2–P4 EXP, relative length of the setae on the male P5 EXP, armature of male P6,
and by the dimorphic male rostrum of C. levis sp. nov. which is similar to that observed for C.
retrogressus and C.albuquerquensis. Additionally, a fully illustrated record of the Mexican C. sinaloensis
is reported for the first time from Brazilian waters, and C. affinis mongolicus Ste
ˇrba, 1968 is given full
species status.
Keywords: Cletocamptus, Copepoda, Harpacticoida, taxonomy, new species
Introduction
Mesochra (5Cletocamptus)deitersi was originally described by Richard (1897) from the
Naposta Grande River (Argentina). Since Richard’s description, C.deitersi has been
reported from inland brines as well as coastal estuaries and mangroves from several sites in
North, Central, and South America as well as in India, China, Ethiopia, Hawaii, Australia,
Iran, and Malaysia (Go´ mez et al. 2004). Nevertheless, the cosmopolitanism of C. deitersi is
best explained by the intraspecific variability of the species, and above all, by the fact that
virtually all specimens attributed to C. deitersi around the globe are morphologically similar
and cannot be differentiated on the basis of Richard’s (1897) original description (Go´mez
et al. 2004).
Correspondence: Samuel Go´mez, Instituto de Ciencias del Mar y Limnologı´a, Joel Montes Camarena s/n, Mazatla´n 82040,
Sinaloa, Mexico. Email: samuelgomez@ola.icmyl.unam.mx
Published 28 October 2005.
Journal of Natural History, 2005; 39(34): 3101–3135
ISSN 0022-2933 print/ISSN 1464-5262 online #2005 Taylor & Francis
DOI: 10.1080/17415970500264335
Several authors (Dexter 1995; Sua´ rez-Morales et al. 1996; Gee 1999; Mielke 2000a,
2001) have suggested that C. deitersi consists of a number of morphologically
indistinguishable sibling species. This was partially confirmed by Rocha-Olivares et al.
(2001) and Castro-Longoria et al. (2003). Their findings instigated the morphological
analysis of the known populations of Cletocamptus previously identified with the
Argentinian C. deitersi. After thorough examination of the available literature, Go´mez
et al. (2004) decided to consider as species inquirendae C. deitersi (Richard, 1897), C. dadayi
(Delachaux, 1917), C. brehmi Kiefer, 1933, C. bermudae Willey, 1930, C. cfr. bicolor,
Herbst, 1960, C. gabrieli Lo¨ ffler, 1961, C. ecuadorianus Lo¨ ffler, 1963, C. kummleri
(Delachaux, 1917), and the C. deitersi material of Daday (1902), Kiefer (1936), Herbst
(1960), Hamond (1973), Dussart (1974), Tai and Song (1979), and Sua´ rez-Morales et al.
(1996). Go´mez et al. (2004) considered Ranga Reddy and Radhakrishna’s (1979), Dussart
and Frutos’ (1986), Sitjar’s (1988), and Zamudio-Valde´z’s (1991) records of the species as
doubtful. They also suggested a more detailed description of dorsal and ventral
ornamentation of body somites of C. axi Mielke, 2000, which following Go´mez et al.
(2004) is a valid species. They also considered C. schmidti Mielke, 2000 as a valid species
different from C. deitersi based on the fact that this species resembles Cletocamptus
specimens (previously identified with C. deitersi) from Sa˜ o Luis Island (Maranhao, Brazil)
(USNM 250011) in the armature formula of female P1–P5, but suggested a detailed
redescription of the dorsal and ventral ornamentation of body somites of C. schmidti.
Additionally, Go´ mez et al. (2004) noticed that the small seta on the male P6 of some
specimens of C. axi and C. schmidti was found also in specimens from Sa˜o Luis Island
(Jansen lagoon, Maranhao, Brazil) (USNM 242172), suggesting that this character may
provide useful phylogenetic information.
The present contribution deals with the description of two new species of Cletocamptus
and a new and fully illustrated record of C. sinaloensis Go´mez, Fleeger, Rocha-Olivares and
Foltz, 2004, from Canane´ ia (Sa˜o Paulo, Brazil) and Sa˜o Luis Island (Maranhao, Brazil).
Material and methods
Specimens of C.nudus sp. nov. and C.sinaloensis analysed in this study were borrowed from
the USNM collection. The vials were labelled and contained a number of specimens as
follows:
Cletocamptus deitersi, USNM 250011; Brazil; Maranhao; Ilha de Sa˜ o Luis, near Raposo,
in mangroves. This vial contained 13 Cletocamptus specimens and three specimens of an
unidentified Laophontidae. The Cletocamptus specimens were shown to belong to two
different species: 11 specimens are herein attributed to C. sinaloensis Go´ mez, Fleeger,
Rocha-Olivares and Foltz, 2004, and two specimens were attributed to C. nudus sp. nov.
described below.
Cletocamptus deitersi, USNM 242172; Brazil; Maranhao; Ilha de Sa˜o Luis; Lagoa Jansen.
This vial contained five Cletocamptus specimens that were attributed to C. sinaloensis.
Cletocamptus deitersi, USNM 278124; Brazil; Rio de Janeiro, Lagoa de Freitas; P. S.
Young, June 1996; J. W. Reid, 1 July 1996; 2f; Acc. #41. This vial contained only one CIII
(third copepodid) and was not included in the present study.
Specimens of C. levis sp. nov. from Canane´ ia, Sa˜o Paulo (Brazil) were provided by Dr
Guilherme R. Lotufo.
Morphological observations and drawings were made from whole and dissected
specimens. Intraspecific variability in armature formulae of P1–P6 was assessed only from
3102 S. Go´mez
dissected specimens. Aberrations, deformed setae/spines and/or segments were not
considered as intraspecific variability. Only the presence or lack of well-developed and/or
reduced setae/spines, and different patterns of spinular ornamentation of body somites
were considered as intraspecific variability. Observations and drawings were made using a
Leica compound microscope equipped with drawing tube at a magnification of 10006.
Additional observations were made also at 12506. The type material has been deposited in
the National Museum of Natural History (Smithsonian Institution) (USNM) and in the
Copepoda collection of the Museu de Zoologia, Universidade de Sa˜o Paulo (MZUSP).
The terminology proposed by Huys and Boxshall (1991) for morphological descriptions
was adopted. The following abbreviations are used in the text and tables: P1–P6, first to
sixth swimming legs; EXP, exopod; ENP, endopod; P1(P2–P4) EXP(ENP) 1(2, 3)
denotes the proximal (middle, distal) exopodal (endopodal) segment of P1, P2, P3, or P4.
Taxonomic account
Family CANTHOCAMPTIDAE Sars, 1906 (incertae sedis)sensu Por, 1986
Genus Cletocamptus Schmankewitsch, 1875
Cletocamptus nudus sp. nov.
(Figures 1–5)
Type material
One female holotype (USNM 250011) preserved in alcohol and one dissected female
paratype (USNM 1010094).
Type locality
Sa˜o Luis Island, Maranhao, Brazil (2u319S, 44u169W).
Etymology
The specific epithet refers to the bare anal operculum of the species.
Description
Female. Habitus (Figure 1A, B) tapering posteriorly; total body length measured from tip
of rostrum to posterior margin of caudal rami ranging from 675 to 740 mm (mean
707.5 mm, n52; holotype 740 mm). Rostrum defined at base, triangular, with pair of setules
subapically and ornamented with small spinules distally. Cephalic shield (Figure 1A, B)
with small, fine spinules along margin dorsally and with comparatively longer spinules
laterally. Dorsal and lateral surface of free thoracic somites (P2–P4-bearing somites) with
transverse rows of minute spinules, with longitudinal row of small spinules close to
posterior margin and with long spinules along posterior margin. Dorsal and lateral surface
of first urosomite (P5-bearing somite) with transverse rows of minute spinules, with row of
small spinules close to posterior margin and with long spinules along posterior margin.
Genital somite with subcuticular rib dorsally and laterally indicating former division
between first and second genital somites (Figure 1A, B), but completely fused ventrally
(Figure 2A); dorsal and lateral surface of first and second genital somite with transverse
New species of Cletocamptus from Brazil 3103
rows of minute spinules, with row of long spinules along posterior margin of both somites,
and with relatively longer spinules laterally, ventrally with spinules. Fourth and fifth uroso-
mite as in second genital somite dorsally, ventral surface with short transverse spinule rows.
Dorsal surface of anal somite (Figure 1A, D) with transverse rows of minute spinules and
with dorsolateral strong spinules close to joint with caudal rami; rounded anal operculum
Figure 1. Cletocamptus nudus sp. nov., female. (A) Habitus, dorsal; (B) habitus, lateral; (C) anal somite and right
caudal ramus, lateral; (D) anal somite and caudal rami, dorsal. Scale bars: 100 mm.
3104 S. Go´mez
without ornamentation. Caudal rami (Figures 1A–D, 2A) about 1.5 times longer than
wide; dorsal and ventral surface smooth, except for inner set of spinules close to insertion
site of seta VII and close to posterior margin dorsally and ventrally; with seven elements.
Antennule (Figure 3A) six-segmented; surface of segments smooth except for two rows
of spinules on first segment. Armature formula, 1-(1), 2-(8), 3-(6), 4-(1 +[1 +ae]), 5-(1),
6-(9 +[1 +ae]).
Antenna (Figure 3B) with small coxa. Allobasis armed with two abexopodal setae. Free
endopodal segment ornamented with inner strong spinules proximally and subdistally; with
Figure 2. Cletocamptus nudus sp. nov., female. (A) Urosome, ventral (P5-bearing somite omitted); (B) P5. Scale
bar: 100 mm.
New species of Cletocamptus from Brazil 3105
two lateral inner spines and a slender seta (the latter indicated in Figure 3B), and five distal
elements. Exopod one-segmented; about seven times longer than wide; with few spinules,
and with one lateral and one apical smooth seta.
Mandible (Figure 3C) robust; chewing edge with bicuspidate teeth, four (or five?)
multicuspidate teeth, one pyriform element and one lateral seta. Palp one-segmented, with
two setae unequal in length and one small seta arising nearby.
Figure 3. Cletocamptus nudus sp. nov., female. (A) Antennule; (B) antenna; (C) mandible; (D) maxillule; (E)
maxilla; (F) maxilliped. Scale bar: 10 mm.
3106 S. Go´mez
Figure 4. Cletocamptus nudus sp. nov., female. (A) P1; (B) P2. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3107
Maxillule (Figure 3D) robust; arthrite of praecoxa with few spinules, with one surface
seta, six (or seven?) distal spines and one lateral strong seta, the latter spinulose. Coxa with
some spinules and with two slender setae. Basis with some median spinules and with two
apical setae; exopod and endopod represented by three setae each.
Figure 5. Cletocamptus nudus sp. nov., female. (A) P3; (B) P4. Scale bar: 100 mm.
3108 S. Go´mez
Maxilla (Figure 3E): syncoxa with minute spinules along inner margin and close to joint
with allobasis; with two endites, each bearing three setae. Allobasis drawn into strong claw
bearing one accompanying seta. Endopod represented by three setae.
Maxilliped (Figure 3F) subchelate. Syncoxa with row of spinules and with a small seta on
inner distal corner. Basis without armature and with one anterior and one posterior longi-
tudinal row of spinules. Endopod drawn into long and slender claw with one accompanying
small seta. The shape of the maxilliped shown in Figure 3F is an artefact of mounting.
P1 (Figure 4A): praecoxa with spinules close to joint with coxa. The latter with four
transverse spinule rows on anterior face, and with spinule row near outer distal corner on
posterior face. Basis with median spinule row, and with stronger spinules at base of exopod,
between rami and at base of inner basal spine. Exopod three-segmented. Endopod two-
segmented, of about the same length as exopod.
P2 (Figure 4B): praecoxa as in P1. Coxa with median row of small spinules and with
strong spinules close to outer distal corner anteriorly, and with some spinules close to outer
distal corner posteriorly. Basis with spinules between rami and at base of endopod, and
with stronger spinules at base of exopod; outer element spine-like. Exopod three-
segmented and ornamented as figured. Endopod two-segmented, reaching proximal
quarter of EXP 2; ENP 1 small, slightly wider than long and with outer spinules; ENP 2
with long spinules as shown, and with one outer spine, one apical and one inner seta.
P3 (Figure 5A): praecoxa and coxa as in P2. Basis as in P2 except for seta-like outer
element. Exopod as in P2 except for two inner setae on P3 EXP 3. Endopod as in P2 except
for comparatively shorter ENP 2.
P4 (Figure5B): coxa and basis as in P2. Exopod as in P2, except for location of inner seta on
P4 EXP 3. Endopod two-segmented, barely reaching the middle of P4 EXP 1; ENP 1 small,
slightly wider than long; ENP 2 with inner and outer slender spinules and with two apical setae.
P5 (Figure 2B): exopod and baseoendopod fused. Baseoendopodal lobe about two times
longer than exopod, with sets of spinules along inner and outer margin, with spinules at
base of apical seta; with one outer, one apical, and four inner setae; relative length of setae
as figured. Exopod with spinules as figured, with five setae plus outer seta of basis.
Armature formula of female P1–P5 as in Table I.
P6 (Figure 2A) represented by median plate in anterior half of first genital somite, each
vestigial leg represented by an outer long and slender seta, and an inner minute element
(indicated in Figure 2A). Copulatory pore in the middle of genital double somite.
Male. Unknown.
Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004
(Figures 6–13)
Material examined
Four females and one male preserved in alcohol, one dissected male and one dissected
female, and two males and two females prepared for SEM (USNM 1010091), and one
Table I. Armature formula of female P1–P5 of Cletocamptus nudus sp. nov.
P1 P2 P3 P4 P5
EXP I-0; I-1; I,I1,1 I-0; I-1; II,I1,1 I-0; I-1; II,I1,2 I-0; I-1; II,I1,1 5
ENP 0-1; 0,I1,1 0-0; I,1,1 0-0; I,1,1 0-0; 0,2,0 6
New species of Cletocamptus from Brazil 3109
male preserved in alcohol, one dissected female and three dissected males (USNM
242172).
Distribution
Mexico: Ensenada del Pabello´ n lagoon, Sinaloa (24u199–24u359N, 107u289–107u459W), El
Yugo estuary, Sinaloa (23u189140N, 106u299W), Urı´as system, Sinaloa (23u119060N,
106u259060W); Brazil: Jansen lagoon, Sa˜ o Luis Island, Maranhao (2u319S, 44u169W).
Figure 6. Cletocamptus sinaloensis Go´mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) Habitus, dorsal; (B)
habitus, lateral; (C) anal somite and right caudal ramus, dorsal; (D) left caudal ramus, lateral. Scale bars: 100mm.
3110 S. Go´mez
Description
The material presented herein matches the description of C. sinaloensis given by Go´mez
et al. (2004). The body length of the Brazilian material (from 450 to 640 mm; mean
569 mm, n57) falls within the range reported earlier by Go´ mez et al. (2004) (from 510 to
750 mm; mean 658 mm, n511). The detailed description of Cletocamptus specimens is
necessary for a correct identification and species separation. Lack of detail and careless
descriptions of Cletocamptus species attributed to C. deitersi worldwide have obliged several
authors to establish synonymies based on poor descriptions. In order to strongly support
the assignation of the Brazilian material to C. sinaloensis, the female (Figure 6A, B) and
male habitus (Figure 11A, B), female (Figure 6C, D) and male caudal rami (Figure 11A, B,
Figure 7. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) Urosome, ventral
(P5-bearing somite omitted); (B) P5. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3111
D), ventral view of the female (Figure 7A) and male urosome (Figure 11D); female mouth
parts (Figure 8A–F), male A1 (Figure 12A), female P1–P5 (Figures 7B, 9A, B, 10A, B),
and male P1–P6 (Figures 11C, D, 12B, C, 13A–C) of Cletocamptus sinaloensis from Jansen
Lagoon are illustrated in detail. The written description has been given elsewhere (Go´mez
et al. 2004) and it is not presented here.
Variability
Male. Two males (USNM 242172) possess a minute seta on each P6 (see Figure 11D).
Female. USNM 242172 possesses two inner setae on the second exopodal segment of left
P2; the setae of P6 of another paratype are somewhat longer (not shown).
Figure 8. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) Antennule; (B)
antenna; (C) mandible; (D) maxillule; (E) maxilla; (F) maxilliped; (G) rostrum. Scale bar: 100 mm.
3112 S. Go´mez
Figure 9. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) P1; (B) P2. Scale
bar: 100 mm.
New species of Cletocamptus from Brazil 3113
Figure 10. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) P3; (B) P4. Scale
bar: 100 mm.
3114 S. Go´mez
Cletocamptus levis sp. nov.
(Figures 14–25)
Type material
One female holotype (MZUSP 16669) preserved in alcohol, one male allotype dissected
and mounted on to four slides (MZUSP 16670), four female dissected paratypes (MZUSP
16671, MZUSP 16675, MZUSP 16673, MZUSP 16674), and seven female paratypes
preserved in alcohol (MZUSP 16672), 3 October, 1998, coll. Guilherme Lotufo.
Type locality
Canane´ia, Sa˜ o Paulo, Brazil (25u019S, 47u579W).
Figure 11. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, male. (A) Habitus, dorsal;
(B) habitus, lateral; (C) P6 and third urosomite, ventral; (D) urosome (P5-bearing somite omitted) showing P6
with small seta. Scale bars: 100 mm.
New species of Cletocamptus from Brazil 3115
Figure 12. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, male. (A) Antennule; (B) P1;
(C) P2. Scale bar: 100 mm.
3116 S. Go´mez
Etymology
The specific epithet refers to the variation in spinular ornamentation of the body somites
and anal operculum.
Description
Female. Habitus (Figures 14A, B, 15A, 16A) tapering posteriorly; total body length
measured from tip of rostrum to posterior margin of caudal rami ranging from 500 to
680 mm (mean 588 mm, n510; holotype 585 mm). Rostrum (Figure 18G) defined at base,
triangular, with pair of setules subapically and with small spinules distally. Cephalic shield
with long, fine spinules along lateral margin (Figures 14B, 16A), and without (Figure 14A)
or with smaller spinules along posterior dorsal and dorsolateral margin (Figures 15A, 16A).
Dorsal and lateral spinular ornamentation of free thoracic somites (P2–P4-bearing somites)
as in Figures 14A, B, 15A or 16A, B. Dorsal and lateral surface of first urosomite (P5-
bearing somite) ornamented as in Figures 14A, B, 15A, B, 16A, B. Genital somite with
Figure 13. Cletocamptus sinaloensis Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, female. (A) P3; (B) P4; (C)
P5. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3117
Figure 14. Cletocamptus levis sp. nov., female. (A) Habitus, dorsal; (B) habitus, lateral; (C) anal somite and right
caudal ramus, lateral; (D) anal somite and caudal rami, dorsal. Scale bar: 167 mm (A, B); 100 mm (C, D).
3118 S. Go´mez
Figure 15. Cletocamptus levis sp. nov., female. (A) Habitus, dorsal; (B) urosome, lateral. Scale bar: 167 mm (A);
100 mm (B).
New species of Cletocamptus from Brazil 3119
subcuticular rib dorsally and laterally indicating former division between first and second
genital somites (Figures 14A, B, 15A, B, 16A, B), but completely fused ventrally
(Figure 17A). Dorsal and lateral surface ornamentation of first and second genital
somites, and fourth and fifth urosomites variable (Figures 14A, B15A, B, 16A, B).
Dorsal surface of anal somite with transverse spinule rows (Figures 14A, D, 15A, 16B);
Figure 16. Cletocamptus levis sp. nov., female. (A) Habitus, lateral; (B) urosome, dorsal. Scale bar: 167 mm (A);
100 mm (B).
3120 S. Go´mez
rounded anal operculum with variable number of spinules (see Figures 14A, D, 15A, 16A).
Caudal rami (Figures 14A–D, 15A, B, 16A, B, 17A) about 1.5 times longer than wide;
dorsal surface smooth, except for some small spinules close to insertion of seta VII
(Figures 14A, D, 15A, 16B); with (Figure 16B) or without (Figures 14A, B, 15A) spinules
close to posterior margin dorsally; with spinules close to posterior margin ventrally; without
Figure 17. Cletocamptus levis sp. nov., female. (A) Urosome, ventral; (B) P5; (C) baseoendopodal lobe of P5,
another specimen. Scale bar: 100 mm (A, B); 85 mm (C).
New species of Cletocamptus from Brazil 3121
Figure 18. Cletocamptus levis sp. nov., female. (A) Antennule; (B) antenna; (C) mandible; (D) maxillule; (E)
maxilla; (F) maxilliped; (G) rostrum; (H) chewing edge of mandible, another view. Scale bar: 100 mm.
3122 S. Go´mez
Figure 19. Cletocamptus levis sp. nov., female. (A) P1; (B) P2. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3123
Figure 20. Cletocamptus levis sp. nov., female. (A) P3; (B) P4. Scale bar: 100 mm.
3124 S. Go´mez
Figure 21. Cletocamptus levis sp. nov., male. (A) Habitus, dorsal; (B) urosome, lateral. Scale bar: 200 mm (A);
86 mm (B).
New species of Cletocamptus from Brazil 3125
Figure 22. Cletocamptus levis sp. nov., male. (A) Antennule; (B) urosome, ventral (P5-bearing somite omitted);
(C) P5. Scale bar: 143 mm (A); 167 mm (B); 100 mm (C).
3126 S. Go´mez
(Figure 14A, D) or with (Figures 15A, 16B, 17A) long spinules medially on inner margin of
caudal rami; with seven setae.
Antennule (Figure 18A) six-segmented; surface of segments smooth except for two rows
of spinules on first segment. Armature formula, 1-(1), 2-(9), 3-(6), 4-(1 +[1 +ae]), 5-(1),
6-(9 +[1 +ae]).
Antenna (Figure 18B): allobasis with two abexopodal setae. Free endopodal segment
with inner strong spinules proximally and subdistally; with two lateral inner spines and a
slender seta (the latter indicated in Figure 18B), and five distal elements. Exopod one-
segmented; about seven times longer than wide; with few spinules, and with one inner, one
outer (arrowed in Figure 18B) and one apical seta.
Mandible (Figure 18C, H) robust; chewing edge with bicuspidate teeth, four (or five?)
multicuspidate teeth, one pyriform element and one lateral seta. Palp (Figure 18C) one-
segmented, with two long setae unequal in length and one small seta arising nearby.
Maxillule (Figure 18D) robust; arthrite of praecoxa with few spinules, one surface seta,
seven distal spines and one lateral strong seta, the latter spinulose. Coxa with some spinules
and with two slender setae. Basis with some median spinules and three apical setae; exopod
and endopod represented by three setae each.
Maxilla (Figure 18E): syncoxa with minute spinules along inner and outer margin and
close to joint with allobasis; with two endites, each bearing three setae. Allobasis drawn into
strong claw bearing one accompanying seta. Endopod represented by three setae.
Maxilliped (Figure 18F) subchelate. Syncoxa with row of spinules and a small seta on
inner distal corner. Basis without armature and with one anterior and one posterior
longitudinal spinule row. Endopod drawn into long and slender claw with one
accompanying small seta.
Figure 23. Cletocamptus levis sp. nov., male. Anal somite and caudal rami, dorsal. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3127
P1 (Figure 19A): praecoxa with spinules close to joint with coxa. The latter with anterior
transverse spinule rows and with spinules in outer distal corner posteriorly. Basis with
median spinule row and with stronger spinules at base of exopod, between rami and at base
of inner basal spine. Exopod three-segmented. Endopod two-segmented, ENP 2 reaching
the middle of EXP 3.
P2 (Figure 19B): praecoxa as in P1. Coxa with median row of small spinules and with
strong spinules close to outer distal corner anteriorly, and with some spinules close to outer
Figure 24. Cletocamptus levis sp. nov., male. (A) P1; (B) P2. Scale bar: 100 mm.
3128 S. Go´mez
distal corner posteriorly. Basis with spinules between rami and at base of endopod and with
stronger spinules at base of exopod; outer element spine-like. Exopod three-segmented and
ornamented as figured. Endopod two-segmented, reaching tip of EXP 1; ENP 1 small,
slightly wider than long; ENP 2 with long spinules as figured and with one outer spine, one
apical and one inner seta.
Figure 25. Cletocamptus levis sp. nov., male. (A) P3; (B) P4. Scale bar: 100 mm.
New species of Cletocamptus from Brazil 3129
P3 (Figure 20A): praecoxa and coxa as in P2. Basis as in P2 except for seta-like outer
element. Exopod as in P2. Endopod as in P2 except for comparatively shorter ENP 2.
P4 (Figure 20B): basis as in P2. Exopod as in P2, except for lack of inner seta on EXP 3.
Endopod two-segmented, barely reaching the middle of EXP 1; ENP 1 small, slightly wider
than long; ENP 2 with inner and outer slender spinules and two apical setae.
P5 (Figure 17B, C): exopod and baseoendopod fused. Exopodal lobe reaching insertion
site of outermost baseoendopodal seta; baseoendopodal lobe with spinule patches along
inner and outer margin, with spinules at base of apical and subapical setae; with one outer,
one apical, and four inner setae; relative length of setae variable (compare relative length of
arrowed setae on Figure 17B, C). Exopod with spinules as figured and with five setae plus
outer seta of basis.
Armature formula of female P1–P5 as in Table II.
P6 (Figure 17A) represented by median plate in anterior half of first genital somite, each
vestigial leg represented by a long and slender outer seta and a minute inner element.
Copulatory pore in the middle of genital double somite.
Male. General body shape (Figure 21A) as in female except for genital-double somite.
Lateral margin of cephalic shield with long, slender spinules; with few small spinules along
posterior margin dorsally. Rostrum (see Figure 22A) sexually dimorphic, more slender than
in female. Prosomites (P2–P4-bearing somites) smooth dorsally (as in the female shown in
Figure 15A), except for long, slender spinules along posterior margin and for transverse row
of minute spinules close to posterior margin of P4-bearing somite. First to fifth urosomites
(Figure 21A, B) with transverse spinule rows as in the female (see Figure 16A, B). Third
and fourth urosomites with strong spinules ventrally; fifth urosomite with one transverse
row of strong spinules ventrally (Figure 22B). Anal somite (Figures 21A, B, 22B, 23) as in
female; anal operculum with some dorsal spinules and comparatively smaller spinules along
posterior margin (as in the female shown in Figure 15A). Caudal rami (Figures 21A, B,
22B, 23) as in female.
Antennule (Figure 22A) six-segmented; subchirocer. Last segment with two acute teeth.
Antenna, mandible, maxillule, maxilla and maxilliped (not shown) as in female.
P1 (Figure 24A) as in female, except for dimorphic inner projection and slender inner
spine of basis, ornamentation of the outer seta of ENP 2 (with spinules in the female, but
with setules in the male) and relative length of apical seta of ENP 2 (nearly three times as
long as supporting ramus in the female, but less than two times as long as supporting ramus
in the male).
P2 (Figure 24B) as in female except for dimorphic (slightly stronger) inner spine on ENP
2 and comparatively stronger outer spines of exopod.
P3 (Figure 25A): exopod as in female except for comparatively stronger outer spines.
Endopod three-segmented; ENP 1 as in female; ENP 2 with strong spinules, with inner
apophysis reaching far beyond ENP 3, the latter with two setae.
P4 (Figure 25B) as in female except for comparatively stronger outer exopodal spines.
Table II. Armature formula of female P1–P5 of Cletocamptus levis sp. nov.
P1 P2 P3 P4 P5
EXP I-0; I-1; I,I1,1 I-0; I-1; II,I1,1 I-0; I-1; II,I1,1 I-0; I-1; II,I1,0 5
ENP 0-1; 0,I1,1 0-0; I,1,1 0-0; I,1,1 0-0; 0,2,0 6
3130 S. Go´mez
P5 (Figure 22C): both P5 fused; exopodal and baseoendopodal lobes fused; the former
with transverse row of strong spinules at its base and along inner and outer margin, with
four setae plus outer seta of basis. Baseoendopodal lobe with strong spinules subdistally
and apically and with three elements.
P6 (Figure 22B) represented by a plate, without armature.
Discussion
At present the valid species within Cletocamptus are C. retrogressus Schmankewitsch, 1875,
C. confluens (Schmeil, 1894), C. albuquerquensis (Herrick, 1894), C. trichotus Kiefer, 1929,
C. feei (Shen, 1956), C. affinis Kiefer, 1957, C. gravihiatus (Shen and Sung, 1963), C.
helobius Fleeger, 1980, C. merbokensis Gee, 1999, C. axi Mielke, 2000, C. schmidti Mielke,
2000, C. deborahdexterae Go´ mez, Fleeger, Rocha-Olivares and Foltz, 2004, C. stimpsoni
Go´mez, Fleeger, Rocha-Olivares and Foltz, 2004, C. sinaloensis Go´mez, Fleeger, Rocha-
Olivares and Foltz, 2004, and C. fourchensis Go´ mez, Fleeger, Rocha-Olivares and Foltz,
2004. Go´mez et al. (2004) considered as species inquirendae some species within the genus
and they also considered as doubtful some records of C. deitersi. Additional doubtful
records of C. deitersi are those of Brehm (1936, 1965), Chappuis (1936), Ringuelet (1958a,
1958b, 1960, 1962), Ringuelet et al. (1965), Oliveira and Miranda (1971), Apostolov
(1984), Ruber et al. (1994), and Loftus and Reid (2000).
To the best of my knowledge, only C. trichotus, C. feei, C. affinis, C. gravihiatus, C.
helobius, C. axi, C. schmidti, C. merbokensis, C. deborahdexterae, C. stimpsoni, C. sinaloensis,
and C. fourchensis have been reported only from or near their type localities. Cletocamptus
confluens,C. retrogressus, and C. albuquerquensis have been reported repeatedly from distant
localities (for a list of references see Mielke 2000b, p 138–139 for C. confluens, Mielke 2001,
p 6–8 for C. retrogressus, and Herrick 1894, Pallares 1962, Chappuis 1933, Lang 1948,
p 1278, and Brehm 1954 for C. albuquerquensis). Cletocamptus affinis was described from
Turkey in 1957, and later a new subspecies, C. affinis mongolicus Ste
ˇrba, 1968, was
described from Mongolia. Ste
ˇrba (1968, p 67) noticed a number of differences between his
C. affinis mongolicus and Kiefer’s C. affinis: ventral spinular ornamentation of female
urosome, number of setae on the caudal rami, and number and relative size of the setae on
P1 ENP 1 (similar to C. feei,C. gravihiatus,andC. helobius in that P1 ENP 1 lacks the inner
seta) and P3 ENP 2 (similar to C. stimpsoni in the number of setae). On the basis of current
research it is suggested to elevate C. affinis mongolicus Ste
ˇrba, 1968 to full species rank as
Cletocamptus mongolicus Ste
ˇrba, 1968.
Based on erroneous identifications of Cletocamptus species worldwide, C. deitersi was
supposed to be a cosmopolitan species distributed in North, Central, and South America as
well as in India, China, Ethiopia, Hawaii, Australia, Iran, and Malaysia (see Go´ mez et al.
2004 for a list of references). However, recent studies (Rocha-Olivares et al. 2001; Castro-
Longoria et al. 2003; Go´ mez et al. 2004) have partially proved that its alleged cosmopolitan
distribution is the result of incompleteness of Richard’s (1897) description of the species
(all specimens attributed to this species around the globe are morphologically similar and
cannot be differentiated on the basis of Richard’s 1897 original description), and the high
intraspecific variability of the genus (Go´ mez et al. 2004; see also Dexter 1995; Sua´rez-
Morales et al. 1996; Gee 1999; Mielke 2000a, 2001). Similarly, C. confluens seems to be
distributed in Eastern and Western Europe, in Northern and Southern Africa, close to the
Persian Gulf, India and Australia (Mielke 2000b, p. 139, Figure 7). It is possible that this
species ‘‘may in reality represent an assemblage of barely discernible species’’ (Mielke
New species of Cletocamptus from Brazil 3131
2000b). The same could be the case for C. albuquerquensis and C. retrogressus and a
thorough revision of these species is still pending.
Upon preliminary examination of the Brazilian material of C. sinaloensis, only two
differences were observed when compared to the Mexican material of the species: the
armature formula of the antennal exopod and the presence of spinules on the male fifth
urosomite ventrally. The antennal exopod was described as having two setae only and the
male fifth urosomite was described as lacking ventral spinules. Upon re-examination of the
material of C. sinaloensis deposited in the Copepoda collection of the Institute of Marine
Sciences and Limnology (Mazatlan Marine Station), three setae were observed on the
antennal exopod (one lateral, and one apical, small seta masked by a bigger apical element)
(Figure 8B), and the presence of spinular ornamentation on the male fifth urosomite
(Figure 11D) was confirmed. The mouth parts, female and male P1–P6 are identical. The
occurrence of the same species in the Atlantic coast of Brazil and in the Mexican Tropical
Pacific seems to be a rather common phenomenon among benthic copepods as evidenced
by the presence of Cyclopina caissara Lotufo, 1994 as reported recently by Go´mez and
Martı´nez-Arbizu (2004).
Cletocamptus nudus sp. nov. is similar to C. fourchensis in the bare anal operculum.
Nevertheless, C. nudus sp. nov. differs from C. fourchensis in the relative size of the ventral
spinules of the second genital segment, and fourth and fifth urosomites. The most obvious
difference between these two species is the armature formula of the antennal exopod (with
two setae in C. nudus sp. nov., and three setae in C. fourchensis) and armature formula of
P3–P4 EXP 3 (P3 and P4 with two and one inner seta in C. nudus sp. nov., and with one
and without inner seta in C. fourchensis). The female P5 of these two species differ in the
relative length of the baseoendopodal setae, the distalmost inner apical seta being about
three-quarters and half length of the apical element in C. nudus sp. nov. and C. fourchensis,
respectively. Cletocamptus nudus sp. nov. seems to possess a very small element close to the
single seta representing P6. The same was observed in some specimens of the Brazilian
material of C. sinaloensis sp. nov. (not shown) but this character still has to be verified.
Cletocamptus nudus sp. nov. seems to be related to C. schmidti,C. stimpsoni, and C.
retrogressus in the armature formula of P3 EXP 3 and P4 EXP 3 (with two and one inner
seta) but differs from the two latter species in the armature formula of P3 ENP 2 (with
three setae in C. nudus sp. nov. and five elements in C. retrogressus and C. stimpsoni). In this
regard, C. nudus sp. nov. seems to be more closely related to C. schmidti by the combination
of the armature formula of the antennal exopod, mandible, P3 and P4, as well as in the
shape of the lateral element on the maxillulary arthrite. Cletocamptus schmidti and C. nudus
sp. nov. can be separated based on the ornamentation of the anal operculum (without
spinules in C. nudus, but with small spinules in C. schmidti), relative length of exopodal and
baseoendopodal setae of the female P5, and above all on the length ratio of the female P1
EXP:ENP (P1 ENP of C. nudus sp. nov. being slightly longer than the exopod, but barely
reaching beyond the tip of P1 EXP 2 in C. schmidti). Unfortunately, the male of C. nudus
sp. nov. remains unknown and cannot be compared with the male of C. schmidti.
Mielke (2000a) described C. axi and C. schmidti from three sampling stations in the
Galapagos Archipelago (Mielke 2000a, p. 282, Figure 7). In one sampling station (locality
IX.4 of Santa Cruz; Mielke 2000a, Figure 7) he found these two species living together.
Despite the fact that C. axi and C. schmidti revealed some intraspecific variability, Mielke
(2000a) was able to define and separate these two species since ‘‘no intermediate animal
was found’’ (Mielke 2000a). Even though the specimens of C. levis sp. nov. did not show
any variability in the armature formula of mouth parts and swimming legs (except for some
3132 S. Go´mez
slight variability in the relative length of the setae of P5 of one female paratype, see
Figure 17B, C), three different types of dorsal spinular ornamentation of body somites and
caudal rami were observed (compare Figures 14A, B, 15A, B, 16A, B), with the specimen
shown in Figure 15 being intermediate. It is noteworthy that the ventral spinular
ornamentation of the female urosome was found to be constant (i.e. no variability was
observed in the female ventral spinular ornamentation of urosome). On the other hand,
more or less the same number of specimens of each type was observed, therefore making it
difficult to decide which type of somitic spinular ornamentation should define this species.
However, because the dorsal surface spinular ornamentation of body somites of the only
male observed (see Figure 21A, B) is similar to that of the female shown in Figure 16A, B, it
is suggested that this type could define C. levis sp. nov. in terms of dorsal spinular
ornamentation of body somites. Nevertheless, it has to be noted that the male lacks the
inner long spinules on the caudal rami, which are well developed in the females shown in
Figures 15A and 16B, but very small in the female shown in Figure 14A, D.
Previous to this paper, eight Cletocamptus species have been described from America (C.
helobius,C.sinaloensis,C.fourchensis,C.deborahdexterae,C.stimpsoni,C.axi,C.schmidti,and
C.albuquerquensis). Cletocamptus levis sp. nov. is similar to C.sinaloensis,C.fourchensis,C.
deborahdexterae,andC.axi in the armature formula of the mandibular palp (with two setae
arising from the one-segmented palp, plus a small seta arising nearby), shape of the lateral
spinulose element of the maxillulary arthrite, and armature formula of P1–P4. However, C.
levis sp. nov. can be separated from C.sinaloensis,C.fourchensis,andC.deborahdexterae by the
P1 EXP:ENP length ratio (P1 ENP longer or as long as exopod in C.sinaloensis,C.fourchensis,
and C.deborahdexterae, but P1 ENP 2 hardly reaching the middle of P1 EXP 3 in C.levis
sp. nov.) and by the P5 BENP:EXP length ratio (exopod reaching the middle of the
baseoendopodal lobe, far below the insertion site of the outermost seta in C.sinaloensis,
C.fourchensis,andC.deborahdexterae, but reaching the insertion site of the outermost
baseoendopodal seta, far above the middle of the baseoendopodal lobe in C.levis sp. nov.).
Cletocamptus levis sp. nov., C.helobius,C.axi,C. schmidti,andC.albuquerquensis are the only
American representatives of the genus in which the P1 ENP is shorter that the P1 EXP (P1
ENP 2 barely reaching the middle of P1 EXP 3 in Cletocamptus levis sp. nov., C.helobius,C.axi,
and C.albuquerquensis, and P1 ENP barely reaching beyond the tip of P1 EXP2 in C. schmidti).
Cletocamptus levis sp. nov. seems to be more closely related to C.axi than to C.albuquerquensis
sensu Pallares (1962) and C.helobius by the armature formula of the mandibular palp, length:
width ratio of caudal rami, and relative length of female P3 ENP 2, P2–P4 EXP 1, and
number of segments of the male P3 ENP. Cletocamptus axi and C. levis sp. nov. can be
separated by the spinule ornamentation of the anal operculum (larger in C. axi), shape of
female rostrum (oval-shaped in C. axi,triangularinC. levis sp. nov.) , relative length of outer
spinules of female P2–P4 EXP (longer in C. axi), and relative length of the setae on the male
P5 EXP. Cletocamptus levis sp. nov. showed the typical sexual dimorphism for the genus in A1,
basis of P1, outer spines of P2–P4, P2 ENP, P3 ENP, P5 and P6. It is noteworthy that C.levis
showed sexual dimorphism in the rostrum, being more slender in the male than in the female
(compare Figures 18G and 23A). The same has been observed only for C.retrogressus (Mielke
2001, p 4, Figure 2B) and C.albuquerquensis (Pallares 1962, p. 242, La´m. I, Figure8).
Acknowledgements
I am indebted to Dr Chad Walter and Dr Frank Ferrari (Smithsonian Institution) for the
loan of the examined material of Cletocamptus nudus sp. nov. and C. sinaloensis, and to Dr
Guilherme Lotufo for providing me with some specimens of C. levis sp. nov. from
New species of Cletocamptus from Brazil 3133
Canane´ia, Sa˜ o Paulo, Brazil. I am also grateful to Dr Janet W. Reid for pointing out
relevant records of Cletocamptus deitersi.
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New species of Cletocamptus from Brazil 3135
