Zootaxa 4052 (3): 383–393
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ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.4052.3.8
http://zoobank.org/urn:lsid:zoobank.org:pub:12401B21-CB99-4786-BFF9-24FD2D3E65F0
A new species of Sunipea Griffin & Tranter, 1986 (Crustacea: Brachyura:
Inachidae) and a redescription of Sunipea indicus (Alcock, 1895)
WILLIAM SANTANA
Laboratório de Sistemática Zoológica, Pró-Reitoria de Pesquisa e Pós-Graduação, Universidade Sagrado Coração – USC, Rua Irmã
Arminda, 10-50, Jd. Brasil, 17011-160, Bauru, SP, Brasil. E-mail: william_santana@yahoo.com.br
Abstract
A new species of spider crab, Sunipea callistus, n. sp., from Cape Guardafui, Somali Republic, is described and illustrated.
Sunipea indicus (Alcock, 1895) is also redescribed and figured. Sunipea callistus n. sp. differs from S. indicus in the structures of the carapace, third maxillipeds, thoracic sternum, male abdomen, and chelipeds. Despite a number of characteristics that separate both species, these differences are more pronounced in males and are less evident in females. A
diagnosis for the genus is provided.
Key words: Apocremnus, Aepinus, Majoidea, Majidae, Indian Ocean
Introduction
Alcock (1895) described Apocremnus indicus based on material from two localities in the Indian Ocean: Andaman
Islands and Sri Lanka (Ceylon). He believed it was a member of Apocremnus A. Milne-Edwards, 1878, and close
to the American Apocremnus septemspinosus A. Milne-Edwards, 1878 (type species). Apocremnus A. MilneEdwards, 1878, however, was pre-occupied by Apocremnus Fieber, 1858 (Hemiptera); and Rathbun (1897)
proposed Aepinus as a replacement name. Griffin & Tranter (1986: 59) established a new genus, Sunipea, to
accommodate Aepinus indicus which they argued was quite different from the American type species. Although
Griffin & Tranter (1986) provided a description and discussion for the genus, they did not redescribe Sunipea
indicus, probably because of the detailed description and figures by Alcock (1895: 188, pl. IV figs. 2, 2a) as well as
discussion of this species over the years (see Griffin 1972, 1974; Griffin & Tranter 1974). Alcock’s description,
however, lacks several important morphological details by modern standards, especially when we compare it with
Sunipea callistus n. sp. A more detailed description is thus necessary to facilitate morphological comparisons and
future studies about Sunipea indicus.
I herein redescribe Sunipea indicus and describe a new species, Sunipea callistus n. sp. A detailed diagnosis
for the genus and both species is also provided. Abbreviations used are as follow: stn, station; det., determiner; cl,
carapace length (from the rostral notch to the posterior margin of the carapace); cw, carapace width; m, meters;
mm, millimeters. The material herein studied is deposited in the U.S. National Museum of Natural History,
Smithsonian Institution, Washington, D.C. (USNM).
Taxonomy
Superfamily Majoidea
Family Inachidae
Accepted by P. Castro: 2 Nov. 2015; published: 3 Dec. 2015
383
Sunipea Griffin & Tranter, 1986
Sunipea Griffin & Tranter, 1986: 59.
Sunipea—Davie 2002: 298; Števčić 2005: 97; Ng et al. 2008: 113; De Grave et al. 2009: 36.
Type species. Apocremnus indicus Alcock, 1895, by original designation; gender masculine.
Included species. Sunipea indicus (Alcock, 1895); Sunipea callistus n. sp.
Diagnosis. Hepatic region of the carapace with 2 long spines directed anterolaterally; metagastric, cardiac,
mesobranchial, intestinal regions with distinct spines, some of which are sometimes capitate; branchial region with
1 anteroventrolateral spine. Rostrum bifurcated. Supraorbital spines long, directed anteriorly; postorbital spines
reduced, present as a crenulated tubercle. Ocular peduncle constricted medially, armed with sharp tubercles or
distinct anterior lamella in distal half. Antennal article with long spine in anterolateral angle, slightly directed
anteriorly. Anterolateral margin of merus of third maxilliped medially expanded with distinct spine. Male gonopod
1 stout, with acute apex on lateral margin, a distinct mesial lobe.
Sunipea indicus (Alcock, 1895)
(Figures 1B, D; 2B, D; 3B, D; 4B, D, F; 5C, D)
Apocremnus indicus Alcock 1895: 188, pl. IV figs. 2, 2a.
Aepinus indicus—Rathbun 1911: 247; 1925: 92; Griffin 1972: 68, fig 3; Griffin 1974:6 [pro parte]; Griffin & Tranter 1974:
164; Takeda 1977: 122, fig. 4B.
Sunipea indicus—Griffin & Tranter 1986: 59, figs 15a, d [pro parte]; Davie 2002: 298; Komai et al. 2002: 20; Ng et al. 2008:
113; Poore et al. 2008: 60; Komatsu 2011: 252, fig 17E.
Material examined. Somali Republic, Cape Guardafui, Anton Bruun, stn 463, 11°24’N-51°35’E, 17.xii.1964, 75–
150 m, ovigerous female (USNM 135111). South of Baia Di Hafun, Anton Bruun, stn 445, 09°41’N-51°03’E,
16.xii.1964, D.J.G. Griffin det., 60–70 m, female (USNM 135109). South of Baia Di Hafun, Anton Bruun, stn 444,
09°36’N-51°01’E, 16.xii.1964, D.J.G. Griffin det., 80 m, 2 males (USNM 135108). Seychelles, Amirante Islands,
Sealark, stn E10 [approx. 6°00'S 53°10'E (Lowry & Stoddart 1992)], 11.x.1905, 40–155m, female (USNM 41388).
Mozambique, Mozambique Channel, Anton Bruun, stn 371D, 24°26’S-35°21’E, 18.viii.1964, D.J.G. Griffin det.,
165 m, ovigerous female (USNM 135106).
Diagnosis. Carapace covered with subequal tubercles, irregularly distributed, concentrated on hepatic,
branchial regions. Rostrum bifurcated, curved, divergent. Ocular peduncle with distinct anterior lamella in distal
half. Third, fourth antennal articles not reaching tip of rostrum; epistome with few distinct tubercles. Merus with
anterolateral margin strongly expanded, with distinct medial spine. Thoracic sternite III–VII with verrucose
tubercles forming carinas Sterno–abdominal cavity encircling telson, forming sharp carina, with 2 distinct tubercles
on each side of the telson. Female abdominal somites 2–6, telson covered with tubercles. Male gonopod 1 slightly
curved, divergent, with mesial lobe leaf-like, directed anteromesially, with a large aperture, distinct velvet on
lateral margin absent.
Redescription. Carapace piriform, almost two times longer than wide; surface covered with subequal, small
tubercles, irregularly distributed, mostly concentrated on hepatic, branchial regions; hooked setae sparsely
distributed, with dense clusters in rostrum, between orbits, hepatic, gastric, branchial regions; hepatic region with
long spine directed anterolaterally; metagastric, cardiac, mesobranchial, intestinal regions with one spine each,
branchial region with 1 anteroventrolateral spine. Carapace spines, including rostrum, sometimes resembling blunt
tubercles, spines longer, capitate in some specimens. Gastric, hepatic, branchial, cardiac, intestinal regions well
defined. Gastric region elongated forming constricted neck. Rostrum bifurcated, curved, divergent, longer in males,
anteriorly directed; rostrum, orbital dorsal margins with few tubercles. Supraorbital spines long, directed
anteriorly; postorbital spines very short, with crenulated tips. Ocular peduncle constricted medially, with distinct
anterior lamella in distal half. Antennular fossae longitudinally ovate, with smooth margins. Interantennular
septum elongated, compressed laterally, forming distinct ventrally directed lobe. Antennal flagella short; third,
fourth antennal articles long, slender, not reaching tip of rostrum. First, second antennal articles fused to epistome;
second article with long spine in anterolateral angle, slightly directed anteriorly, with few sub-equal tubercles near
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mesial margin. Epistome with few distinct tubercles, markedly longer than wide. Epistomial spine, interantennular
septum separated by small gap. Mouthfield subquadrate, protruded anteriorly, with crenulated anterolateral angles.
Pterygostomian region subtriangular, with row of strong tubercles forming carinae, smaller tubercles present;
separated from subhepatic region by prominent groove. Subhepatic region with long anteroventraly directed spine,
small tubercles evenly distributed.
FIGURE 1. A–D, Habitus, dorsal and ventral views. A, C, Sunipea callistus n. sp., male holotype cl 14.6 mm, cw 10.5 mm
(USNM 1191775). B, D, Sunipea indicus, male (USNM 135108). Cheliped digitally removed in C. Scale bars, 5.0 mm.
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FIGURE 2. A–D, Habitus, dorsal and ventral views. A, C, Sunipea callistus n. sp., ovigerous female paratype, cl 7.4 mm, cw
4.6 mm (USNM 135110). B, D, Sunipea indicus, ovigerous female (USNM 135106). Scale bars, 5.0 mm.
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FIGURE 3. A, B, Habitus, dorsal view. C, D, Male sternum and abdomen. A, C, Sunipea callistus n. sp., male holotype cl 14.6
mm, cw 10.5 mm (USNM 1191775). B, D, Sunipea indicus, male (USNM 135108). Dotted lines represent broken spines. Scale
bars, 5.0 mm.
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FIGURE 4. A, B, Female abdomen. C, D, third maxilliped. E, F, chelipeds. A, Sunipea callistus n. sp., ovigerous female
paratype, cl 7.4 mm, cw 4.6 mm (USNM 135110). B, Sunipea indicus, ovigerous female (USNM 135111). C, E, Sunipea
callistus n. sp., male holotype cl 14.6 mm, cw 10.5 mm (USNM 1191775). D, F, Sunipea indicus, male (USNM 135108). Scale
bars, A-D, 2.5 mm; E, F, 5.0 mm.
Third maxillipeds completely covering buccal frame. Exopod long, reaching distal margin of merus; dorsal
face with sparse tubercles parallel to lateral margin, otherwise irregularly distributed; mesial margin with strong
spine in distal third. Ischium longer than broad; mesial margin almost straight; crista dentata with row of short
setae, small acute teeth; dorsal face of ischium longitudinally concave, with distinct, small, well-spaced tubercles,
sharp tubercles on lateral margin. Merus longer than half of ischium, with smaller tubercles; anterolateral margin
strongly expanded, with distinct medial spine, few long setae present; anterolateral angle crenulated, mesial margin
with few small sharp tubercles, otherwise smooth. Palp shorter than merus; carpus compressed, longer, propodus,
dactylus cylindrical, similar in size, without tubercles, with long setae along mesial, lateral margins, propodus
slender; carpus with few long setae on lateral margin.
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FIGURE 5. Abdominal view of first male gonopod. A, B, Sunipea callistus n. sp., male holotype cl 14.6 mm, cw 10.5 mm
(USNM 1191775). C, D, Sunipea indicus, male (USNM 135108). Scale bars, 2.0 mm.
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Male thoracic sternite I to IV fused, broadly triangular, with 3 acute tubercles in anterior tip, few strong
verrucose tubercles irregularly distributed. Posterior half of fused sternites I–IV strongly sloping down in ventral
view, covered with strong verrucose tubercles, forming a carina along lateral margin. Sterno-abdominal cavity
encircling telson, forming sharp carina, with 2 distinct tubercles on each side of the telson. Male sternites V–VII
with strong, verrucose tubercles, mostly fused, forming line along sternite sutures, surface in females smooth.
Margin of male episternites IV–VII irregular, smooth on females.
Chelipeds equal, long, robust in males; females with shorter, more slender chelipeds. Palm, fingers strongly
compressed laterally, with carinate edges, palm with small, few granules on proximal margin. Dactylus, fixed
finger distinctly shorter than palm, cutting edges with sub-equal teeth; without gap when closed. Dactylus with a
line of tubercles on lateral face. Carpus, merus, ischium densely covered with tubercles. Merus with row of
stronger tubercles on mesial face, short hooked setae present.
Walking legs slender, cylindrical. First leg longest; other legs progressively decreasing in length. Dactylus
slightly curved, densely setose, with small granules on lower edge. Propodus, carpus, merus, ischium with
subequal tubercles sparsely distributed.
Male abdominal somites 1–6, telson free, slightly raised medially forming low longitudinal ridge. Somite 3
with several strong tubercles; somites 4, 5 with verrucose tubercles medially, somite 6 with sparse tubercles.
Female abdominal somites 1–3, telson free, 4–6 fused. In males, surfaces of abdominal somites 2, 4–6 with few
tubercles, somite 3 densely covered with strong tubercles; female abdominal somites 2–6 covered with distinct,
subequal, uniformly distributed tubercles. Female abdomen markedly arched, with row of setae on margin. Telson
subtriangular, terminating in rounded apex with sparse tubercles in males; female telson transversally oval, with
uniformly distributed tubercles. Gonopod 1 (= male pleopod 1) stout, slightly curved, divergent, with acute apex on
lateral margin; mesial lobe long, leaf-like, directed anteromesially, with large aperture surrounded with several
small setae. Lateral margin smooth. Gonopod 2 (= male pleopod 2) slender, straight, very short (one fifth of G1
length).
Distribution. Indo-West Pacific region: from East Africa and Red Sea to Andaman Sea, Indonesia (Sumbawa
and Kai Is.), Ogasawara Is., Philippines, and Australia (Griffin & Tranter 1986; Poore et al. 2008).
Sunipea callistus n. sp.
(Figures 1A, C; 2A, C; 3A, C; 4A, C, E; 5A, B)
Sunipea indicus—Griffin 1974: 7 [pro parte]; Griffin & Tranter 1986: 59 [pro parte].
Type material. Somali Republic, Cape Guardafui, Anton Bruun, stn 463, 11°24’N-51°35’E, 17.xii.1964, 75–150
m, male holotype, cl 14.6 mm, cw 10.5 mm (USNM 1191775). Somali Republic, Cape Guardafui, Anton Bruun,
stn 463, 11°24’N-51°35’E, 17.xii.1964, 75–150 m, male paratype, cl 13.7 mm, cw 9.4 mm (USNM 1191774).
Cape Guardafui, Anton Bruun, stn 453, 11°11’N-51°14’E, 17.xii.1964, 47–49 m, ovigerous female paratype, cl 7.4
mm, cw 4.6 mm (USNM 135110).
Diagnosis. Carapace uniformly, densely covered with subequal tubercles. Rostrum bifurcated, straight. Ocular
peduncle with sharp tubercles distally. Third, fourth antennal articles reaching the tip of rostrum; epistome without
tubercles. Merus of the third maxillipeds with distinct crenulated margin medially ending in a spine. Sparse small
tubercles irregularly distributed in thoracic sternite III; male thoracic sternites IV–VII with tubercles irregularly
distributed. Male sterno-abdominal cavity longer than telson, leaving a gap between telson, anterior crenulated
margin. Female abdominal somites 1–3 with tubercles; female telson without tubercles. Male gonopod 1 straight,
slightly converging, with mesial lobe acute, upturned, forming a distinct groove, lateral margin with distinct velvet
in distal third.
Description. Carapace piriform, 1.5 times longer than wide in males, females slender; surface densely covered
with subequal tubercles uniformly distributed; hooked setae sparsely, with dense clusters between orbits, rostrum,
hepatic, gastric, branchial regions; hepatic region with one long spine directed anterolaterally; metagastric, cardiac,
mesobranchial, intestinal regions with 1 long, sometimes capitate, spine each; branchial region with 1
anteroventrolateral spine. Gastric region delimited by groove without tubercles, more evident in males; hepatic,
branchial, cardiac, intestinal regions well defined. Rostrum bifurcated, straight, short, anteriorly directed; rostrum,
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orbital dorsal margins with tubercles. Supraorbital spines long, directed anteriorly; postorbital spines very short.
Ocular peduncle constricted medially, armed with sharp tubercles distally. Antennular fossae longitudinally ovate,
with smooth margins. Interantennular septum elongated, compressed laterally, forming distinct ventrally directed
lobe. Antennal flagella short; third, fourth antennal articles long, slender, reaching the tip of rostrum. First, second
antennal articles fused to epistome; second article with long spine in anterolateral angle, slightly directed
anteriorly, with sub-equal tubercles near lateral margin. Epistome without tubercles, distinctly longer than wide.
Epistomial spine, interantennular septum separated by small gap. Mouthfield subquadrate, protruded anteriorly,
with crenulated anterolateral angles. Pterygostomian region subtriangular, with row of strong tubercles forming
carinae, smaller tubercles present; separated from subhepatic region by shallow groove. Subhepatic region with
long spine, crenulated tip in some specimens; small tubercles evenly distributed.
Third maxillipeds completely covering buccal frame. Exopod long, nearly reaching distal margin of merus;
dorsal face with row of small tubercles parallel to lateral margin, otherwise irregularly distributed; mesial margin
with strong spine in distal half. Ischium longer than broad; mesial margin almost straight; crista dentata with row of
short setae, small acute teeth; dorsal face of ischium concave longitudinally, with distinct, well-spaced tubercles.
Merus longer than half of ischium, with smaller tubercles; anterolateral margin strongly expanded, with distinct
crenulated margin medially ending in a spine, few long setae present; mesial margin crenulated, otherwise smooth.
Palp shorter than the merus; carpus compressed, longer, ornamented with small tubercles dorsally, few long setae
on lateral margin; propodus, dactylus cylindrical, similar in size, without tubercles, with long setae along mesial,
lateral margins; propodus slender.
Male thoracic sternite III broadly triangular, with 3 acute tubercles in the anterior tip, sparse tubercles
irregularly distributed. In males, sternite IV strongly sloping down in ventral view, sparsely covered with tubercles,
sterno-abdominal cavity longer than telson, leaving gap between telson, anterior crenulated margin. Male sternites
V–VII with strong tubercles, surface in females smooth. Episternites IV–VII with crenulated margins.
Chelipeds equal, distinctly long, robust in males, females with shorter, more slender chelipeds. Palm, fingers
compressed laterally. Row of tubercles in lateral margin forming carina, smaller tubercles sparsely distributed;
mesial margin forming distinct carina. Dactylus, fixed finger distinctly shorter than palm, cutting edges with subequal teeth; in males, fingers leaving strong gap in proximal half when closed, dactylus with small, blunt, distinct
basal tooth, absent in females. Movable, fixed fingers smooth on mesial, lateral faces, inconspicuous carina on
lateral face of dactylus. Carpus, merus, ischium densely covered with tubercles. Merus with row of stronger
tubercles on mesial face, short hooked setae present.
Walking legs slender, cylindrical. First leg longest. Dactylus slightly curved, densely setose, with small
granules on lower edge. Propodus, carpus, merus, ischium with subequal tubercles evenly distributed.
Male abdominal somites 1–6, telson free, slightly raised medially forming low longitudinal ridge. Female
abdominal somites 1–3, telson free, 4–6 fused. In males, surface of abdominal segments 1–6, in females, 1–3
covered with distinct tubercles, female fused somites with tubercles concentrated in the proximal half. Female
abdomen markedly arched, with row of setae on margin. Telson subtriangular, terminating in rounded apex in
males, with few tubercles; female telson transversally oval, without tubercles. Gonopod 1 (= male pleopod 1) stout,
straight, slightly converging, with acute apex on lateral margin; mesial lobe long, acute, upturned, with aperture
forming distinct groove. Distinct velvet on lateral margin in distal third. Gonopod 2 (= male pleopod 2) slender,
straight, very short (one fifth of G1 length).
Type locality. Cape Guardafui, Somali Republic, 11°24’N-51°35’E, 75–150 m of depth.
Distribution. Known so far only from the type locality.
Etymology. The specific epithet is derived from the Greek adjective kallistos for “the most beautiful.”
Remarks. Sunipea callistus n. sp. is easily distinguishable from S. indicus by (i) a carapace that is uniformly
and densely covered with subequal tubercles (Figs. 1A, 2A, 3A) (carapace with subequal tubercles irregularly
distributed and concentrated in some regions in S. indicus; Figs. 1B, 2B, 3B); (ii) rostrum straight (Figs. 1A, 2A,
3A) (rostrum curved, divergent in S. indicus; Figs. 1B, 2B, 3B); (iii) ocular peduncle with sharp tubercles distally
(Figs. 1A, 3A) (ocular peduncle with a distinct anterior lamella in distal half in S. indicus; Figs. 1B, 3B); (iv) third
and fourth antennal articles that reach the rostrum length (Fig. 3A) (third and fourth antennal articles do not reach
the tip of the rostrum in S. indicus; Fig. 3B); (v) epistome without tubercles (Figs. 1C, 2C) (with a few distinct
tubercles in S. indicus; Figs. 1D, 2D); (vi) the merus of the third maxillipeds with a distinct crenulated anterior
margin that ends in a spine medially and a mesial crenulated margin (Fig. 4C) (the merus of the third maxillipeds
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are without an anterior crenulated margin and a medial spine, with a crenulated anterolateral angle and mesial
margin that contains a few small sharp tubercles in S. indicus; Fig. 4D); (vii) sparse small tubercles that are
irregularly distributed in thoracic sternite III (Figs. 1C, 2C, 3C) (verrucose tubercles in S. indicus; Figs. 1D, 2D,
3D); (viii) male thoracic sternites IV–VII with tubercles irregularly distributed (Figs. 1C, 3C) (the strong verrucose
tubercles form carinas in S. indicus; Figs. 1D, 3D); (ix) the male sterno-abdominal cavity is longer than the telson,
thus leaving a gap between the telson and the anterior crenulated margin (Figs. 1C, 3C) (the sterno-abdominal
cavity encircles the telson, forming a keen carina with two distinct tubercles on each side of the telson in S. indicus;
Figs. 1D, 3D); (x) the chelipeds of the males with a distinct gap between the fingers in the proximal half (Fig. 4E)
(chelipeds without gap in S. indicus; Fig. 4F); (xi) the female abdominal somites 1–3 only have tubercles (Figs. 2C,
4A) (female abdominal somites 2–6 are covered with tubercles in S. indicus; Figs. 2D, 4B); (xii) the female telson
is without tubercles (Figs. 2C, 4A) (the female telson is covered with tubercles in S. indicus; Figs. 2D, 4B); and
(xiii) the male gonopod 1 is straight and slightly converging, with mesial lobe acute, upturned, forming a distinct
groove, and the lateral margin has a distinct velvet in the distal third (Figs 5A, B) (gonopd 1 slightly curved,
divergent, with mesial lobe leaf-like, directed anteromesially, with a large aperture and the distinct velvet on lateral
margin absent in S. indicus; Figs 5C, D).
Although several characters can be diagnostic between the two species, these differences are much more
noticeable in males than females. For example, the difference in the width of the carapace, which is noticeably
wider in males of S. callistus n. sp. than in S. indicus, is much less evident in females (Fig. 2). Also, differences in
the rostrum, chelipeds and abdomen of the females of both species are not as noticeable as in males (Figs 2, 4A, B).
This may be why Griffin (1974: 7) identified the present material as S. indicus. Griffin (1974: 7) treated the
differences he observed in the rostrum as only morphological variation, commenting that “the rostral spines are
sometimes straight and sometimes outwardly curved” (see also Griffin & Tranter 1974: 165). There is no mention
about the other differences that are listed, although he observed that some specimens did not possess intact gastric
and cardiac spines (Griffin & Tranter 1974). The similarities of the females and the fact that carapace spines,
including the rostrum, are frequently broken and sometimes resemble blunt tubercles could also be misleading.
Griffin & Tranter (1984: 59) also described the interantennular partition as without a spine for the genus. In the
material studied, however, all specimens of both species had an interantennular septum that had a distinct ventrally
directed lobe. Other than that, the description presented here agrees with the description of the genus presented by
Griffin & Tranter (1984).
Alcock (1895: 189) described Apocremnus indicus on the basis of two specimens, a male of 9 mm cl and a
female 6 mm cl, without indicating which was the holotype. Davie (2002: 298) mentioned that the syntypes are
deposited in the Indian Museum, the present Zoological Survey of India (ZSI). This material was not available for
me to study, thus, although needed, no inference about lectotypes can be made.
Acknowledgments
I am grateful to Marcos Tavares (Museu de Zoologia da Universidade de São Paulo) for his guidance and
discussions about taxonomy. I am also indebted to Rafael Lemaitre (USNM) for making material from his
institution available for study and for providing working space. Thanks also to Karen Reed for her kind assistance
during my visits to the USNM and to Daniel Lima (Universidade Estadual Paulista – UNESP) and Jessica Colavite
(Universidade Regional do Cariri—URCA) for the help during figures preparation. I thank FAPESP (Fundação de
Amparo à Pesquisa do Estado de São Paulo) for the financial support through grants number 2008/11280-6, 2013/
01201-0 and 2014/15549-0. This work greatly benefited from the comments of P.K.L. Ng (Lee Kong Chian Natural
History Museum) and from Peter Castro (California State Polytechnic University, Pomona).
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