Rare deep-water crabs (Crustacea: Decapoda) from Indian waters, with description of one new species

Abstract

Deep-water brachyuran crabs collected from the Indian Exclusive Economic Zone by the Fishery Oceanographic Research Vessel “Sagar Sampada” in the southeastern Arabian Sea (50 m depth), southwestern Bay of Bengal (307 m depth), and in the vicinity of the Andaman and Nicobar Archipelagos (271-535 m depth) were studied. They are referred to Sphaerodromia kendalli (Alcock and Anderson, 1894) and Sphaerodromia nux Alcock, 1900 (family Dromiidae), Intesius brevipes sp. nov. (family Mathildellidae), Tunepugettia corbariae B.Y. Lee, Richer de Forges and P.K.L. Ng, 2019 (family Epialtidae), Cyrtomaia suhmii Miers, 1885 (family Inachidae), Chaceon alcocki Ghosh and Manning, 1993 (family Geryonidae), and Sphenomerides trapezioides (Wood-Mason, 1891) (family Trapeziidae). Sphaerodromia nux and T. corbariae are new records from Indian waters. The first male specimen of C. alcocki is reported, with a description of the male first gonopod. Intesius brevipes sp. nov. is the fifth species of the genus, differing from the four known congeners in the form and setation of the carapace, distinctly curved and granular anterolateral margins, and shorter pereopods with much slender merus and carpus articles.

Keywords
Brachyura; deep-sea; India; new species; taxonomy

INTRODUCTION

The deep-water brachyuran fauna of India is moderately diverse and was substantially documented during the colonial era (Wood-Mason, 1885; 1888; Wood-Mason and Alcock, 1891; Alcock, 1894; 1895; 1896; 1898; 1899a; 1899b; 1900a; 1900b; 1901; 1905; Alcock and Anderson, 1894; 1899; Anderson, 1896; McArdle, 1900; 1901; MacGilchrist, 1905; Lloyd, 1907; Kemp and Sewell, 1912). In comparison, the recently published literature revealed few reports on the deep-water Brachyura based on the collections of the deep-sea by-catch species along the peninsular coasts of India (Kumar et al., 2013; Ng and Kumar, 2015; 2016; Mendoza and Devi, 2017; Ng et al., 2017a; 2018a; 2018b; Ng and Mitra, 2019; Mendoza et al., 2020).

The Centre for Marine Living Resources & Ecology, Ministry of Earth Sciences, Government of India, Kochi has been conducting dedicated deep-water faunal surveys in the Indian Exclusive Economic Zone (EEZ) since the 10^th Five-year plan period (2002-2007). These surveys have resulted in a collection of deep-water crustaceans which includes new species and new geographical records (see Ng et al., 2019; Padate et al., 2020a-c). The present study reports seven deep-water species from the Indian EEZ, including one new species; all from specimens collected between 2010-2016.

MATERIAL AND METHODS

The study area comprises the continental shelf off the southwest coast of India in the southeastern Arabian Sea, the upper continental slope region in the southwestern Bay of Bengal, and the upper continental slopes of the eastern and western sides of the Andaman and Nicobar Islands (Fig. 1).

Figure 1.
Map of the study area indicating the sites of sample collection in the Indian EEZ.

This study is based on the archived samples collected between 2010-2016 using an Expo model trawl net and an High-Speed Demersal Trawl II net (Crustacean Version with 30 mm cod end mesh size) on-board the FORVSS during the following cruises: no. 279 in the western Bay of Bengal, no. 280 in the Andaman Sea and southeastern Bay of Bengal, no. 321 in the southeastern Arabian Sea, and no. 334 (leg I) and no. 349 (leg II) in the southeastern Bay of Bengal.

The specimens were cleaned to remove debris, examined, and photographed with an Olympus TG-5 underwater camera. Microscopic diagnostic characters were photographed using a Leica M80 stereo-zoom microscope equipped with Leica MC170 HD microscope camera and Leica Application Suite imaging software. Photographic plates were made in Adobe^® Photoshop CS5. Measurements were done with vernier callipers to an accuracy of 0.01 mm.

The morphological terminology used is from Guinot and Richer de Forges (1982), Serène (1984), Ghosh and Manning (1993), McLay (2001), Crosnier and Ng (2004), Ng et al. (2017b), and Lee et al. (2019). The measurements (in millimetres) are of the maximum carapace length (excluding posterior margin ornamentation), post-rostral carapace length and maximum carapace width (at base of lateral spine).

Material examined in this study are deposited in the Referral Centre collection of the Centre for Marine Living Resources and Ecology (CMLRE), Kochi, Kerala, India, and preserved in 70 % ethanol.

Abbreviations used in the following text are: CL = carapace length; coll. = collected by; CW = carapace width; FORVSS = Fishery Oceanographic Research Vessel “Sagar Sampada”; G1 = male first gonopod; G2 = male second gonopod; IO/SS/BRC = Indian Ocean/Sagar Sampada/Brachyura; P2 to P5 = pereopods 2 to 5 (first to fourth ambulatory legs), respectively; PCL = post-rostral carapace length; stn = station.

TAXONOMY

Infraorder Brachyura Latreille, 1802

Superfamily Dromioidea De Haan, 1833

Family Dromiidae De Haan, 1833

Genus Sphaerodromia Alcock, 1899

Sphaerodromia kendalli ( Alcock and Anderson, 1894 )

(Figs. 2A-C, 3)

Dromidia kendalliAlcock and Anderson, 1894: 175 (type locality: off Nellore coast, Bay of Bengal, 14°5’55”N 80°25’20”E). - Alcock and Anderson, 1896: pl. 24, fig. 1, 1a. - Guinot and Tavares, 2003: 102, 108. - Ng et al., 2008: 36 (list).

Sphaerodromia kendalli - Alcock, 1900a: 153. - Alcock, 1901: 39, 78, pl. 4, fig, 18, 18a. - Ihle, 1913: 92 (list). - Balss, 1922: 106. - Sakai, 1936: 15. - Sakai, 1976: 28, fig. 16. - McLay, 1991: 465 (discussion), table 1 (comparison). - McLay and Crosnier, 1991: 185-186 (discussion). - McLay, 1993: 125 (list), 126, 127 (key), 127, figs. 2a-i, 15a. - McLay, 2001: 821 (key), tab. 1. - Guinot and Tavares, 2003: 105. - Ng et al., 2008: 36 (list). - Trivedi et al., 2018: 36 (tab. 1).

Dromia (Sphaerodromia) kendalli - Alcock, 1899a: 5 (list), 16.

Material examined. 3 males (CL 37.8 mm, CW 43.4 mm; CL 38.6 mm, CW 45.4 mm; CL 43.1 mm, CW 48.6 mm) (IO/SS/BRC/00294), southwest Bay of Bengal, FORVSS stn. 27911, 13.22°N 80.51°E, 307 m, coll. Dr. R. Raghu Prakash, HSDT (CV), 29 August 2010.

Remarks. Sphaerodromia kendalli was originally described from a female (18 × 19 mm) from the western Bay of Bengal at 205 m depth (Alcock and Anderson, 1896, pl. 24 fig. 1, 1a). The original published account (Alcock and Anderson, 1894) and the subsequent publications (Alcock, 1900a; 1901; Sakai, 1936; 1976) provided short descriptions of the Indian and Japanese specimens. McLay’s review (1993) of the family Dromiidae included a detailed re-description of the material collected from the Philippines (MUSORSTOM 3 expedition) and Indonesia (KARUBAR expedition) supplemented with illustrations of the Philippine material (McLay, 1993: figs. 2a-i, 15a).

The present specimens collected from approximately 100 km south of the type locality agree largely with the description of the type specimens as well as subsequent reports, with the exception of a slightly wider carapace, CW/CL 1.13-1.18 (Fig. 2A, B) (vs. 0.97 in the Indonesian male (McLay, 1993); female specimens - 1.06 (holotype) (Alcock and Anderson, 1894), 1.13 (Japan) (Balss, 1922), and 1.02 (Philippines) (McLay, 1993)), the absence of granules on the subhepatic region (Fig. 2C) (vs. granules present in the holotype, and the Philippine material (McLay, 1993)), the relatively longer chelipeds, 1.74 times CL (vs. 1.33 times CL in the holotype female (Alcock, 1901)). The above differences in the present specimens as compared to the type material could be attributed to the comparatively larger size of the present material.

Distribution. Bay of Bengal at 205-307 m depth (Alcock and Anderson, 1894; Alcock, 1900a; 1901; present study), Japan (Balss, 1922; Sakai, 1936; 1976), Philippines, Southeast Molucca Islands, Indonesia at 205-214 m depth (McLay, 1993).

Figure 2.
A-C, Sphaerodromia kendalli (Alcock and Anderson, 1894), male (CL 38.6 mm, CW 45.4 mm) (IO/SS/BRC/00294), southwest Bay of Bengal, India. D, E, Sphaerodromia nux Alcock, 1899, male (CL 29.3 mm, CW 33.1 mm) (IO/SS/BRC/00295), Bay of Bengal west of South Andaman Island, India. A, D, dorsal habitus; B, E, denuded dorsal carapace; C, denuded lateral carapace. Scale bars: 10 mm.

Figure 3.
Sphaerodromia kendalli (Alcock and Anderson, 1894). A-N, P, male (CL 38.6 mm, CW 45.4 mm) (IO/SS/BRC/00294), southwest Bay of Bengal, India. A, Antennule, ventral view; B, antenna, ventral view; C, third maxilliped, ventral view; D, cheliped epipod and arthrobranch, lateral view; E, right chela and carpus, outer view; F, G, P2-P3 dactyli, dorsal view; H, I, P4-P5 subchela, dorsal view; J, thoracic sternum in ventral view indicating the position of the gonopods; K, L, G1 dorsal and ventral views; M, N, G2 dorsal and ventral views; P, male telson, ventral view. O, male (CL 43.1 mm, CW 48.6 mm) (IO/SS/BRC/00294), southwest Bay of Bengal, India, posterior pleonal somites 3-5 and telson in dorsal view indicating the form and position of vestigial pleopods. Scale bars: E-G, J = 10 mm, A-C, K-N, P = 5 mm, D, O = 2 mm, H-I = 1 mm.

Sphaerodromia nux Alcock, 1900

(Figs. 2D, E, 4)

Sphaerodromia nuxAlcock, 1900a: 154 (type locality: Gulf of Martaban). - Alcock, 1901: 40, 78, pl. 4, fig. 19. - Ihle, 1913: 92 (list). - Lewinsohn, 1984: 115, pl. 3, fig. B, pl. 4, figs. C-D. - McLay, 1991: 463, 465 (discussion), tab. 1 (comparison). - McLay and Crosnier, 1991: 186 (discussion). - McLay, 1993: 126 (list), 127 (key). - Crosnier, 1994: 347 (remarks). - McLay, 2001: 821 (key, description). - Guinot and Tavares, 2003: 107, 108, figs. 22B, 23A-B. - Ng et al., 2008: 36 (list). - Trivedi et al., 2018: 36 (tab. 1).

Material examined. 1 male (CL 29.3 mm, CW 33.1 mm) (IO/SS/BRC/00295), west of South Andaman Island, Bay of Bengal, FORVSS stn. 349 (leg II) 09, 11.93°N 92.28°E, 290 m, coll. Dr. Aneesh Kumar K.V., HSDT (CV), 12 April 2016.

Remarks. Sphaerodromia nux was originally described from supposedly juvenile specimens (1 male of undetermined size, 1 female 10 × 11 mm) from the Gulf of Martaban off Myanmar at 129 m depth (Alcock, 1900a). The original description by Alcock (1900a) described only the carapace, and a subsequent monograph by Alcock (1901) provided a short description of the third maxilliped, chelipeds, and the vestigial male pleopods on pleonal somites 3-5 supplemented with a line illustration of the dorsal habitus (Alcock, 1901: fig. 19). Lewinsohn (1984) redescribed this species from adult specimens from Madagascar, and reported differences from the type specimen in the absence of a gap between the anterior margin of the third maxillipeds and the buccal cavern. McLay (1991) provided a detailed comparison between S. nux and Sphaerodromia ducoussoiMcLay, 1991, and supplemented with a table comparing the diagnostic morphological characters of all Sphaerodromia species. Crosnier (1994) commented that S. nux possessed well-developed podobranchs on the cheliped and pereopod 2, and vestigial bud-like podobranchs on pereopod 3. McLay (2001) amended Lewinsohn’s redescription and added morphological details of female specimens from the Philippines. A systematic revision of the family Dromiidae by Guinot and Tavares (2003) included illustrations of the vestigial male pleopods, P5 coxa and G2.

The present specimen agrees with the updated description by McLay (2001), except for the presence of lines of compressed granules on the P2-P3 meri (Fig. 4F-G) and a relatively higher dactylus length/propodus length of 0.90 for P2-P3. The present observation is a new record in Indian waters.

Distribution. Gulf of Martaban at 129 m depth (Alcock, 1900a; 1901), Madagascar (Guinot and Tavares, 2003), and Andaman Sea, India at 290 m depth (present study).

Superfamily Goneplacoidea MacLeay, 1838

Family Mathildellidae Karasawa and Kato, 2003

Genus Intesius Guinot and Richer de Forges, 1981

Intesius brevipes sp. nov.

(Fig. 5)

Zoobank: urn:lsid:zoobank.org:act:8F73C88D-767D-4FFD-92E8-7BA05BD80F39

Material examined. Holotype: female (CL 34.5 mm, CW 42.2 mm) (IO/SS/BRC/00296), west of Rutland Island, Andamans, Bay of Bengal, FORVSS stn. 334 (leg 1) 11, 11.35°N 92.39°E, 535 m, coll. Dr. S. Venu, HSDT (CV), 11 January 2015.

Description of holotype (female). Carapace sub-hexagonal with edges rounded off, 1.22 times wider than long; dorsal surface moderately convex anteriorly, gently convex transversely, microscopically granular, bearing short moderately dense tomentum. Regions demarcated by broad shallow grooves; gastric regions slightly prominent, separated from cardiac region by gastro-cardiac groove, metagastric region with pair of indistinct pits; branchial regions faintly demarcated, subdivided into anterior and posterior parts by broad lateral extension of gastro-cardiac groove; cardiac region as prominent as gastrics, separated from intestinal region by shallow submedian depression (Fig. 5A).

Figure 4.
Sphaerodromia nuxAlcock, 1899, male (CL 29.3 mm, CW 33.1 mm) (IO/SS/BRC/00295), Bay of Bengal west of South Andaman Island, India. A, Antennule and antenna; B, third maxilliped; C, right chela and carpus, outer view; D, P2 merus, dorsal view; E, P2 dactylus, dorsal view; F, P2-P3 meri, ventral view; G, P3 dactylus, dorsal view; H, I, P4-P5 subchela, dorsal view; J, posterior pleonal somites and telson, ventral view; K, pleonal somites 3-5 and telson in dorsal view indicating the form and position of vestigial pleopods; L, M, G1 dorsal and ventral views; N, O, G2 dorsal and ventral views. Scale bars: B-C = 5 mm, A, D-O = 2 mm.

Frontal margin narrow (0.28 times CW), gently deflexed, distinctly projecting, granular, medially notched giving it a bilobed appearance, separated from orbital margin by narrow sulcus (Fig. 5B, C). Supraorbital margin concave, distinctly granular, interrupted by 2 distinct fissures into slightly elevated inner lobe and slightly depressed outer lobe; external orbital tooth short, minutely granular, separated from distinctly granular infraorbital margin by lateral fissure, inner infraorbital angle not extending to level of frontal margin (Fig. 5B, C).

Anterolateral margins distinctly curved, divided into 4 teeth; first tooth shortest, bearing small granules, separated from the external orbital tooth by long granule; remaining teeth spinose - second tooth largest, bearing long granules, separated from first tooth by 2 long granules, and from third tooth by 2 short granules; third tooth subequal to second, bearing smaller granules, separated from fourth tooth by short granules; fourth tooth spinose, bearing 1 posterior granule (Fig. 5B, C). Posterolateral margins gently convex, longer than anterolateral margins, converging posteriorly, anterior halves bearing small scattered granules (Fig. 5A, B).

Figure 5.
Intesius brevipes sp. nov., holotype, female (CL 34.5 mm, CW 42.2 mm) (IO/SS/BRC/00296), Bay of Bengal west of Rutland Island, Andamans, India. A, Dorsal habitus; B, dorsal carapace, anterior portion; C, anterior frontal view indicating the position of antennules, antennae, and third maxilliped; D, right chela and carpus, outer view; E, left chela and carpus, outer view; F, thoracic sternum in ventral view indicating position of gonopores; G, pleon and telson, ventral view; H, gonopores, enlarged view. Scale bars: A-G = 10 mm, H = 5 mm.

Antennules folding slightly obliquely. Basal antennal segment elongated rectangular, movable, located in orbital hiatus, flagellum long, extending to second anterolateral tooth (Fig. 5C).

Epistome small, wide, posterior margin elevated, interrupted by 3 notches (Fig. 5C). Endostomial ridges well developed posteriorly. Buccal frame subquadrate. Third maxillipeds sparsely setose, microscopically granular when denuded; ischium sub-rectangular, 1.57 times longer than wide, bearing distinct submedian longitudinal groove, cutting edge thickly setose; merus subquadrate, 0.92 times longer than wide, length 0.60 times ischial length, bearing strong oblique groove, antero-external angle slightly produced, rounded; palp articulating at antero-internally on merus; exopod slender, flagellum well developed (Fig. 5C).

Female chelipeds unequal, heterochelous, sparsely setose. Right cheliped 1.69 times CL. Merus triangular in cross-section, anterior, posterior margins and ventral surface microscopically granular, posterior margin with 4 proximal elongate granules and subdistal granular spine; carpus with distinctly granular inner margin, large granular spine at antero-internal angle, dorsal and outer surfaces microscopically granular; palm massive, microscopically granular, dorsal margin distinctly granular proximally; fingers microscopically granular, subequal to upper palm length, pollex bearing one indistinct longitudinal groove each on inner and outer margins, dactylus bearing two indistinct longitudinal grooves each on both margins, occlusal margin of dactylus with large proximal molar tooth fitting into socket on pollex, remaining margins with short dentition, distal tips curved, blunt, crossing in closed position (Fig. 5D). Left cheliped 1.32 times CL, ornamentation on merus and carpus similar to that of right cheliped, palm slender, with distinct elongate patch of granules on dorsal margin, outer surface bearing several rows of slightly smaller granules, inner surface with minute granules, ventral margin granular; fingers distinctly granular, longer than upper palm length, grooves similar to those on large cheliped, occlusal margins with short, sharp dentition, dactylus with patch of large granules on proximal one-fourth, distal tip fitting into depression on pollex in closed position (Fig. 5E).

P2-P5 slender, microscopically granular, bearing longer setae (compared to carapace), their lengths 1.55, 1.76, 1.83, and 1.49 times CL, respectively. Length/width ratios for merus, carpus, propodus and dactylus are as follows: P2 = 4.67, 2.64, 3.41, and 7.86; P3 = 5.09, 2.29, 3.86, and 8.78; P4 = 5.02, 2.10, 3.90, and 9.26; P5 = 4.73, 1.96, 2.91, and 6.60. Pereopod meri bearing large granules on anterior margins, those on P5 largest, posterior margins with dense setation, surfaces sparsely setose; carpi with short dense setation on margins, surfaces sparsely setose; propodi with long dense setation on margins, surfaces sparsely setose; dactyli lanceolate, with long dense setation on margins, distal tips corneous (Fig. 5A).

Thoracic sternum wide (width 0.56 times CW), sternites 1-3 and lateral half of sternite 4 microscopically granular, other portions smooth, covered with long silky setae; first 2 sternites completely fused, sternite 2 and 3 separated by sinuous suture, sternites 3-7 separated by medially interrupted sutures, sternite 7 and 8 separated by complete suture, midline dividing posterior half of sternite 7 and entire 8 (Fig. 5F).

Female pleon ovate, widest at somite 5 (width 0.35 times CW), with 6 free somites and telson, margins bearing dense long setation; somite 6 three times longer than wide; telson bluntly triangular, 1.45 times wider than long (Fig. 5G). Gonopores large, ovate, extending across much of sternite 6, margins elevated by sternal prominence with oblique suture anteriorly (Fig. 5H).

Color (Coloration preserved in formalin for ca. 5 years; recently transferred to 70 % ethanol). Light brown with slightly darker setae; large cheliped fingers dark brown on distal eight-tenths, small cheliped fingers dark brown on distal three-quarters.

Remarks. Specimens from the genus IntesiusGuinot and Richer de Forges, 1981 are extremely rare, with only 18 specimens (including the present study). The genus is characterized by a squarish carapace with less prominent anterolateral teeth, male pleon with strongly ankylosed somites 3-5 and a mobile sixth somite (Guinot and Richer de Forges, 1981a; Davie, 1998; Crosnier and Ng, 2004). Hitherto known only from the Western Pacific region (eastern Australia, New Caledonia, Norfolk Ridge, Philippines, French Polynesia, and Mariana Islands), the present observation is the first record of this genus from the Indian Ocean region. Intesius brevipes sp. nov. differs from the existing congeners in:

1. (1) sub-hexagonal carapace, CW/CL ratio 1.22 (Fig. 5A) (vs. quadrangular in I. crosnieri, CW/CL ratio 1.17 (Davie, 1998: figs. 1A, 2A) and I. lucius, CW/CL ratio 1.15-1.17 (Crosnier and Ng, 2004: figs. 3B, 7); subcircular in I. pilosus, CW/CL ratio 1.07-1.20 (Davie, 1998: figs. 1B, 2B); rectangular in I. richeri, CW/CL ratio 1.26 (Crosnier and Ng, 2004: figs. 2, 3A));

2. (2) relatively less dense short setation on carapace that does not conceal the edges (Fig. 5A, B), similar to I. lucius (Crosnier and Ng, 2004: figs. 3B, 7) (vs. dense setation in I. crosnieri (Davie, 1998: figs. 1A, 2A), I. pilosus (Davie, 1998: figs. 1B, 2B), and I. richeri (Crosnier and Ng, 2004: figs. 2, 3A));

3. (3) distinctly projecting, bilobed frontal margin separated from the supraorbital margin by a distinct notch (Fig. 5A-C) (vs. very distinctly projecting, strongly bilobed frontal margin separated from the supraorbital margin by a wider notch in I. pilosus (Guinot and Richer de Forges, 1981a: pl. 7, fig. 1, 1a; Davie, 1998: figs. 1B, 2B); less distinctly projecting, weakly bilobed frontal margin separated from the supraorbital margin by a narrow notch in I. crosnieri (Davie, 1998: figs. 1A, 2A), I. lucius (Crosnier and Ng, 2004: figs. 3B), and I. richeri (Crosnier and Ng, 2004: fig. 3A));

4. (4) slightly elevated inner supraorbital lobe (Fig. 5A-C) similar to I. pilosus (Davie, 1998: fig. 2B) (vs. indiscernible inner supraorbital lobe in the remaining species (Davie, 1998: fig. 2A; Crosnier and Ng, 2004: fig. 3A-B));

5. (5) distinctly curved anterolateral margins (Fig. 5A-C) (vs. moderately curved in its first part and then runs sub-parallel to longitudinal axis in I. pilosus (Guinot and Richer de Forges, 1981a: pl. 7, fig. 1, 1a; Davie, 1998: figs. 1B, 2B); gently convex in I. lucius (Crosnier and Ng, 2004: figs. 3B, 7); gently convex in its first part and then runs sub-parallel to longitudinal axis in I. crosnieri (Davie, 1998: figs. 1A, 2A) and I. richeri (Crosnier and Ng, 2004: figs. 2, 3A));

6. (6) more salient anterolateral teeth of carapace, particularly second and third teeth, bearing distinct granules (Fig. 5A-C), similar to I. pilosus (Davie, 1998: figs. 1B, 2B) (vs. moderately salient teeth bearing moderately large granules in I. crosnieri (Davie, 1998: figs. 1A, 2A) and I. richeri (Crosnier and Ng, 2004: figs. 2, 3A); less developed teeth bearing short granules in I. lucius (Crosnier and Ng, 2004: figs. 2, 3A));

7. (7) palm of large chela distinctly granular proximally on dorsal margin (Fig. 5D), similar to I. pilosus (Davie, 1998: fig. 1B) (vs. finely granular dorsal margin and outer surface in I. richeri (Crosnier and Ng, 2004: fig. 2); sharply granular dorsal and proximal portions of outer surface in I. crosnieri (Davie, 1998: fig. 1A); granular dorsal and ventral margins in I. lucius (Crosnier and Ng, 2004: fig. 7));

8. (8) pereopods 2-5 bearing relatively short setae (Fig. 5A), similar to I. lucius (Crosnier and Ng, 2004: fig. 7) and I. richeri (Crosnier and Ng, 2004: fig. 2) (vs. long dense setae in I. crosnieri (Davie, 1998: fig. 1A) and I. pilosus (Davie, 1998: fig. 1B));

9. (9) relatively shorter P4-5 (pereopod length/ CL ratio 1.83 for P4, 1.49 for P5) with much more slender pereopod meri and carpi (length/width ratio 5.02 and 2.10 for P4, 4.73 and 1.96 for P5) (vs. longer P4-5 (pereopod length/ CL ratio 1.99-2.20 for P4, 1.78-1.97 for P5) with less slender meri (length/width ratio 3.88-4.48 for P4, 3.55-4.35 for P5) and carpi (length/width ratio 1.87-2.06 for P4, 1.48-1.83 for P5) in the remaining congeners).

Etymology. The species name is derived from the combination of Latin terms “brevis” and “pede” alluding to the relatively shorter P4-P5. Gender is masculine.

Distribution. Known only from the type locality in the southeastern Bay of Bengal, India at 535 m depth (present study).

Superfamily Majoidea Samouelle, 1819

Family Epialtidae MacLeay, 1838

Genus Tunepugettia Ng, Komai and Sato, 2017

Tunepugettia corbariae B.Y. Lee, Richer de Forges and P.K.L. Ng, 2019

(Fig. 6)

Tunepugettia corbariae B.Y. Lee, Richer de Forges and P.K.L. Ng, 2019: 21, figs. 10A, 11D-F, 12A-D (type locality: Ainto Bay, southeast New Britain, Solomon Sea, Papua New Guinea, MADEEP Expedition, 06°08’S 149°10’E, 430-620 m depth).

Material examined. 1 female (PCL 26.9 mm, CW 20.8 mm) (IO/SS/BRC/00297), off Middle Andaman Island, Andaman Sea, FORVSS stn. 28016, 12.83°N 93.21°E, 441 m, coll. Dr. Vinu Jacob, Expo model trawl, 17 September 2010.

Remarks. Tunepugettia corbariae was originally described from a male (31.4 × 21.2 mm) collected from Ainto Bay, Papua New Guinea at 430-620 m during the MADEEP Expedition (Lee et al., 2019).

The present female specimen largely agrees with the original description, with exception of a relatively narrow carapace, PCL/CW ratio 1.29 (vs. Papuan female specimens, PCL/CW ratio 1.45-1.67 (Lee et al., 2019)) and slightly less prominent protuberances on the carapace (Fig. 6A-C). The above differences in the present specimen as compared to the Papuan material could be attributed to the comparatively smaller size of the present material. The present observation is a new record in Indian waters.

Distribution. Papua New Guinea at 280-1085 m depth (Lee et al., 2019) and Andaman Sea, India at 441 m depth (present study).

Family Inachidae MacLeay, 1838

Genus Cyrtomaia Miers, 1886

Cyrtomaia suhmii Miers, 1886

(Fig. 7)

Cyrtomaia suhmiiMiers, 1886: 16, pl. 3, fig. 2, 2a-c (type locality: off Tulur Islands, South China Sea, 4°33’0”N 127°6’0”E). - Rathbun, 1918: 6 (remarks). - Richer de Forges and Guinot, 1990: 525, 527 (discussion). - Richer de Forges and Ng, 2007: 56 (list). - Ng et al., 2008: 111 (list). - Promdam, 2011: 7, figs. 1, 2A-F.

Cyrtomaia suhmi - Rathbun, 1893: 230 (remarks). - Griffin, 1974: 9. - Guinot and Richer de Forges, 1981b: 1096 (remarks). - Guinot and Richer de Forges, 1982: 16 (key), 21, figs. 10, 11A-B, 23B. - Guinot and Richer de Forges, 1985: 116, figs. 11B, 12A, B, 14 A-C, pl. V, fig. A-D, F-I. - Griffin and Tranter, 1986a: 24 (key), 30, fig. 9e-g. - Griffin and Tranter, 1986b: 352, figs. 1, 2. - Richer de Forges and Guinot, 1988: 42-44 fig. 2B, C, pl. 2F, G. - Poore, 2004: 360 (key), 361, fig. 108g. - Richer de Forges and Ng, 2007: 62 (remarks).

Cyrtomaia Suhmi var. curvicerosBouvier, 1915: 9-15, pl. 1.

Cyrtomaja suhmi typica - Serène and Lohavanijya, 1973: 45, 46 (key).

Cyrtomaja suhmi curvicornis - Serène and Lohavanijya, 1973: 45, 46 (key).

Cyrtomaia curviceros - Sakai, 1976: 176 (key), 181. - Guinot and Richer de Forges, 1981b: 1096 (remarks). - Guinot and Richer de Forges, 1982: 24, fig. 12A-D. - Richer de Forges and Guinot, 1988: 42, 43 (remarks), fig. 2A. - Ng et al., 2001: 13, 54, 75, 81, fig. 3h. - Richer de Forges and Ng, 2007: 56 (list), 62, 63 (remarks).

Material examined. 1 male (PCL 67.1 mm, CW 78.5 mm) (IO/SS/BRC/00298), off Great Nicobar Island, Bay of Bengal, FORVSS stn. 28037, 6.64°N 93.68°E, 271 m, coll. Dr. Vinu Jacob, Expo model trawl, 24 September 2010.

Remarks. Cyrtomaia suhmii was originally described from a damaged male specimen (25 × 28 mm) collected from the South China Sea, Indonesia at 915 m depth during the HMS Challenger Expedition (Miers, 1886). Doflein (1904) recognized two subspecies from the “Valdivia” collections in the Nicobar Islands, Indonesia (C. suhmi typica) and the Western Indian Ocean (C. suhmi platyceros) based on the differences in the form of carapace, rostrum, female pleon as well as the carapace ornamentation and texture.

Figure 6.
Tunepugettia corbariae B.Y. Lee, Richer de Forges and P.K.L. Ng, 2019, female (PCL 26.9 mm, CW 20.8 mm) (IO/SS/BRC/00297), Andaman Sea off Middle Andaman Island, India. A, Dorsal habitus; B, C, carapace dorsal and lateral views, respectively; D, ventral view indicating the position of antennules, antennae, and third maxilliped; E, gonopores and pleonal locking mechanism, enlarged; F, pleon and telson, ventral view. Scale bars: A-C = 10 mm; D-F = 2 mm.

Figure 7.
Cyrtomaia suhmii Miers, 1885, male (PCL 67.1 mm, CW 78.5 mm) (IO/SS/BRC/00298), Bay of Bengal off Great Nicobar Island, India. A, Dorsal habitus; B, ventral view indicating the position of antennules, antennae and third maxilliped; C, thoracic sternum in ventral view indicating G1 position; D, pleon and telson, ventral view; E, right G1, sub-ventral view. Scale bars: A-B = 50 mm, C-D = 10 mm, E = 1 mm.

Bouvier (1915) recognized a separate subspecies from Japan, C. suhmi curviceros based on arched pseudorostral and gastric spines, and poorly developed branchial spines. Serène and Lohavanijaya (1973) provided a key to the species and subspecies of the genus Cyrtomaia including these subspecies. Griffin (1974) reported this species from the Andaman Sea off Thailand, along with morphological differences from the holotype with respect to the armature of the carapace and the basal antennal article. He also elaborated upon its synonymy and morphological relationships with congeners. Griffin and Brown (1976) reported a large female from Australian waters having long protogastric spines, one supraorbital spine and one mesogastric spine. Sakai (1976) elevated C. suhmi curviceros to a distinct species. Guinot and Richer de Forges (1982) redescribed the holotype, attempted to resolve the taxonomic status of several of its subspecies, and compared other published descriptions with the holotype. They also elevated C. suhmi typica to a distinct species, C. gaillardi. Guinot and Richer de Forges (1985) reported morphological similarities of specimens collected during the MUSORSTOM I and II expeditions in the Philippines to both C. suhmi and C. curviceros and emphasized upon reviewing the identity of C. curviceros. Griffin and Tranter (1986a) suggested that C. suhmi curviceros is an adult stage of C. suhmi and synonymized it with the latter. Griffin and Tranter (1986b) reported specimens from Australian waters and the Straits of Malacca having a large adult size, smooth carapace, and a short eyestalk. Richer de Forges and Guinot (1990) established a new species, Cyrtomaia griffini, based on the material collected off the eastern coast of Australia, also including previously published reports (Griffin and Brown, 1976; Griffin and Tranter, 1986a; 1986b). Promdam (2011) reported this species in the Andaman Sea off Thailand and corroborated C. curviceros to be a synonym of C. suhmii on the basis of both adult and juveniles resembling these taxa, respectively.

The present specimen (Fig. 7A, B, E) largely agrees with the original description by Miers (1886), the holotype specimen (see Guinot and Richer de Forges, 1982: figs. 11A-B, 23B; Richer de Forges and Guinot, 1988: fig. 2B, C), the Philippine specimens (see Guinot and Richer de Forges, 1985: figs. 11B, 12, 14A), line diagram of G1 of male specimen from Tasman Sea (see Griffin and Tranter, 1986a: fig. 9e-g), the Western Australia specimen (Griffin and Tranter, 1986b: figs. 1, 2), and the Thailand specimen (see Promdam, 2011: figs. 1, 2D). However, it differs from the southeastern Arabian Sea specimens in Alcock’s report, in the presence of spine on the “last abdominal tergum” (Fig. 7D) (vs. smooth tergum in the latter specimens in Alcock’s report).

Distribution. Indonesia at 915 m depth (Miers, 1886), Japan, Taiwan, Andaman Sea off Thailand at 502-512 m (Griffin, 1974) and southeastern Bay of Bengal at 271 m (present study).

Superfamily Portunoidea Rafinesque, 1815

Family Geryonidae Colosi, 1923

Genus Chaceon Manning and Holthuis, 1989

Chaceon alcockiGhosh and Manning, 1993 (Fig. 8)

Chaceon alcockiGhosh and Manning, 1993: 714, figs. 1-3 (type locality: off Travancore coast, 8°37’N 75°37’30”E). - Ng et al., 2008: 147 (list). - Jose et al., 2019: 749, figs. 1-3.

Geryon affinis - Alcock, 1899a: 85 [not Geryon affinisMilne Edwards and Bouvier, 1894].

Material examined. 1 male (CL 146.4 mm, CW 161.1 mm) (IO/SS/BRC/00300), off Alleppey (Kerala), Arabian Sea, FORVSS stn. 32121, 9.25°N 76.17°E, 55 m, coll. Dr. Vinu Jacob, HSDT (CV), 13 December 2013.

Remarks.Alcock (1899a) erroneously reported a female geryonid (110.5 × 126.5 mm) from off Travancore (Kerala) coast at 410-520 m as Geryon affinis. Ghosh and Manning’s re-examination of this specimen (1993) resulted in the establishment of a new species, Chaceon alcocki, characterized by short frontal and anterolateral spines, the absence of distal spines dorsally on the P2-P5 meri, and laterally compressed pereopod dactyli. Jose et al. (2019) reported another female specimen (146 × 156 mm) collected from a commercial trawl landing along Kerala coast, and was reportedly caught at 260-300 m depth.

Figure 8.
Chaceon alcockiGhosh and Manning, 1993, male (CL 146.4 mm, CW 161.1 mm) (IO/SS/BRC/00300), southeast Arabian Sea off Alleppey (Kerala), India. A, Dorsal habitus; B, ventral view indicating the position of antennules, antennae and third maxilliped; C, right chela and carpus, outer view; D, left chela and carpus, outer view; E, P5 merus, dorsal view; F, P5 carpus and propodus, dorsal view; G, H, P5 dactylus, dorsal and lateral views; I, pleon and telson, ventral view; J, right G1and G2 in situ, ventral view; K, L, right G1 distal portion, ventral and dorsal views. Scale bars: A = 50 mm, B-F, J = 10 mm, G-I = 20 mm, K-L = 2 mm.

The present specimen is the first male and the largest specimen reported for this species. The G1 is stout, curved, tapering distally; inner margin is covered with a row of short setae along proximal half, distal quarter with a short spinulose patch terminating sub-distally; outer margin with spinulose patch on distal one-third terminating sub-distally, followed by a small patch of long setae appearing like a spinose projection (Fig. 8J). G2 shorter than G1 in length, slender, tapering distally, with spinose process subdistally on inner margin (Fig. 8J). Among the closely related congeners, published descriptions of only two species, Chaceon macphersoniManning and Holthuis, 1988 and Chaceon goreniGalil and Manning, 2001 were supplemented with illustrations of the gonopods. Chaceon alcocki differs from C. macphersoni and C. goreni in the presence of setae on both inner and outer margins (vs. row of setae on inner margin in C. goreni (see Galil and Manning, 2001: fig. 3C-E); absence of setae on either margin in C. macphersoni (see Manning and Holthuis, 1988: fig. 4G, H)).

Distribution. Known only from the southeastern Arabian Sea off Kerala at 50-520 m depths (Ghosh and Manning, 1993; Jose et al., 2019; present study).

Superfamily Trapezioidea Miers, 1886

Family Trapeziidae Miers, 1886

Genus Sphenomerides Rathbun, 1897

Sphenomerides trapezioides ( Wood-Mason, 1891 )

(Fig. 9)

Sphenomerus trapezioidesWood-Mason in Wood-Mason and Alcock, 1891: 263 (type locality: between North and South Sentinel Islands, Bay of Bengal. - Wood-Mason, 1892: pl. 5, fig. 2. - Alcock, 1898: 228. - Poupin et al., 2018: 108 (basionym).

Sphenomerides trapezoides [sic] - Serène, 1968, 89. - Serène, 1973: 207, figs. 6, 27, 28, pl. 5A-D. - Serène, 1984: 289, fig. 196, pl. 42D. - Castro et al., 2004: 34 (in key), 60, pl. 4, fig. E. - Ng et al., 2008: 185 (list). - Castro, 2013: 461, fig. 2F-H.

Figure 9.
Sphenomerides trapezioides (Wood-Mason, 1891), female (CL 6.9 mm, CW 9.4 mm) (IO/SS/BRC/00299), Bay of Bengal off Great Nicobar Island, India. A, Dorsal habitus; B, ventral view indicating the position of antennules, antennae and third maxilliped; C, pleon and telson, ventral view; D, gonopores, enlarged view. Scale bars: A = 5 mm, B-D = 1 mm.

Material examined. 1 female (CL 6.9 mm, CW 9.4 mm) (IO/SS/BRC/00299), Bay of Bengal off Great Nicobar Island, FORVSS stn. 28037, 6.64°N 93.68°E, depth 271 m, coll. Dr. Vinu Jacob, Expo model trawl, 24 September 2010.

Remarks. Sphenomerides trapezioides was described from a male specimen (8.5 × 11 mm) collected off the west coast of the Andamans at 403-439 m depths (Wood-Mason and Alcock, 1891), and subsequently figured by Wood-Mason (1892: pl. 5 fig. 2). Alcock (1898) provided a brief description based on additional specimens from the Andaman Sea. Serène (1973) reported one male and two female specimens from the Indonesian reefs, supplemented with line illustrations of the P4 dactylus and G1, and photographs of carapace and chelipeds. Serène (1984) studied a large collection from Madagascar supported with a line illustration of the G1 and a photograph of the dorsal habitus. Castro et al. (2004) provided details on the synonymy, morphological diagnosis, and notes on the geographical distribution. Castro (2013: fig. 2G) reported the first evidence of S. trapezioides living inside a siliceous sponge, as well as the presence of mucus-gathering setae typical of the family Trapeziidae (Castro, 2013: fig. 2H).

Distribution. Bay of Bengal, off Andamans at 403-439 m depth (Wood-Mason and Alcock, 1891) and Nicobars at 271 m (present study), Andaman Sea at 238-531 m (Alcock, 1898), Haruku Island, Indonesia at 138-152 m and Madagascar at 220-362 m (Serène, 1984; Castro, 2013).

ACKNOWLEDGEMENTS

The authors are grateful to the Director, Centre for Marine Living Resources and Ecology, Kochi, for providing an opportunity to work on deep-water crustacean resources of the Indian EEZ. The authors are thankful to Shri. N. Saravanane, Scientist F, CMLRE and Project Co-ordinator of the ‘Resource Exploration and Inventorization System’ project. We thank the chief scientists, fishing hands and participants of the FORV “Sagar Sampada” cruise numbers 279, 280, 321, 334 (leg I) and 349 (leg II) for meticulously collecting the specimens. The authors are grateful to the VLIZ Library, Flanders Marine Institute, Ostend, Belgium and Dr. Josileen Jose, Principal Scientist, Central Marine Fisheries Research Institute, Kochi, India for sharing valuable taxonomic literature. The authors wish to express their gratitude to the Associate Editor and the two anonymous reviewers for the critical review that improved the quality of the manuscript.

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Publication Dates

• Publication in this collection
  21 July 2021
• Date of issue
  2021

History

• Received
  18 Nov 2020
• Accepted
  28 Apr 2021