Hephthopelta Alcock, 1899

Ng, Peter K. L. & Castro, Peter, 2016, Revision of the family Chasmocarcinidae Serène, 1964 (Crustacea, Brachyura, Goneplacoidea), Zootaxa 4209 (1), pp. 1-182 : 48-52

Genus Hephthopelta Alcock, 1899

Hephthopelta Alcock, 1899: 76 ; 1900: 327.— Tesch 1918: 202 [in key], 233.— Serène 1964a: 239.—Ng et al. 2008: 76 [in list].— De Grave et al. 2009: 32 [in list].

Type species. Hephthopelta lugubris Alcock, 1899 (gender feminine) Other species included:

Hephthopelta occidentalis n. sp.

Diagnosis. Carapace subtrapezoidal, strongly globose, only slightly wider than long; front bilobed, with shallow to well-defined median cleft; anterolateral margins arcuate, minutely granular, entire or with small tubercle. Epistome slightly depressed; median lobe of posterior margin distinctly broad, with deep median fissure, semicircular lateral margins. Eye peduncle filling orbit, distinctly short, gently tapering to cornea, immobile; cornea reduced, pigmented. Third maxillipeds fill buccal cavern; merus subquadrate to elongated, outer margin nearly straight, anteriorly flattened, anteroexternal angle rounded; ischium ovate to elongated, slightly longer than merus. Chelipeds subequal in length, slightly dissimilar in female, heterochelous in males; fingers of minor chela distinctly flattened laterally, broad throughout most of length, shear-like, cutting margins with low, broad teeth. Ventral surface of cheliped merus with 3 large teeth (sometimes additional small tooth or large tubercle) on outer margin in both sexes, inner margin unarmed. Inner margin of carpus with long distal tooth, short proximal part rounded to dentiform. Inner margin of basis-ischium of adult major cheliped gently granuliform. Inner margin of basis-ischium of adult major cheliped gently granuliform ( Fig. 42 G). Meri of P2, P3 with short teeth, meri of other ambulatory legs with microscopic granules, unarmed. P5 dactylus straight to gently curved. Fused thoracic sternites 1, 2 transversely narrow, high, triangular; fused thoracic sternites 3, 4 transversely narrow. Male pleon with lateral margins of somite 6, fused somites 3‒5 nearly straight; postero-lateral regions prominently swollen; telson proportionally short. Sterno-pleonal cavity of male deep, press-button for pleonal holding as small, short tubercle posterior to thoracic sternal suture 4/5 near edge of sterno-pleonal cavity. Male thoracic sternite 8 distinctly short, quadrate; “supplementary plate” short, rectangular; structures tightly appressed with most of penis concealed. G1 with stout basal part, distal part tapering, with short spinules. G2 longer than G1; distal segment long, evenly broad throughout length to tip. Somites of female pleon with convex lateral margins; telson proportionally short, narrow (unknown in H occidentalis ). Sterno-pleonal cavity of female deep, vulvae widely separated, on outer margins of cavity close to suture 5/6.

Remarks. Hephthopelta was established by Alcock (1899) for one species, H. lugubris , from the Indian Ocean. At least ten more species have since been added over the years to the genus (Ng et al. 2008). Of these, all are from the Indo-West Pacific region except for Hephthopelta superba Boone, 1927 , described from the western Atlantic and now placed in Deltopelta n. gen. (see above).

Examination of the good series of “ Hephthopelta ” specimens on hand shows that the 10 Indo-West Pacific species can be separated into six distinct groups on the basis of the carapace form, eyestalks, structures of the third maxillipeds, chelipeds, ambulatory legs, male thoracic sternum, gonopods, and vulvae. We here recognise six genera: Hephthopelta s. str. (for H. lugubris Alcock, 1899 , and H. occidentalis n. sp.), Angustopelta n. gen. (for H. aurita Rathbun, 1932 , H. cribrorum Rathbun, 1932 , A. modesta n. sp. and A. robusta n. sp.), Chinommatia n. gen. (for H. cavimanus Rathbun, 1914 , C. bicuspida n. sp., H. littoralis Tesch, 1918 , H. bruuni Serène, 1964 , and C. turpis n. sp.), Notopelta n. gen. (for H. mortenseni Serène, 1964 ), Statommatia n. gen. (for H. apta Rathbun, 1914 , S. granulosa n. sp., H. knudseni Serène, 1964 , and H. pubescens Chen, 1987 ) and Tenagopelta n. gen. (for T. pacifica n. sp., T. potens (Davie & Richer de Forges, 2013) , and T. brachyphallus n. sp.).

The six genera are all very distinct, easily distinguished by a suite of characters associated with the morphology of the carapace, chelipeds, ambulatory legs, thoracic sternum, male pleon, gonopods, and vulvae ( Table 2 ). Hephthopelta s. str. is closest to Tenagopelta n. gen. because of the rounded form of its carapace and prominently inflated globose appearance when viewed frontally (e.g., Fig. 24 A). The G2 of members of both genera also have well-developed distal segments and are longer than the G1. Members of both genera are also found in deep waters with the former occurring in much greater depths (exceeding 500 m). Compared to Hephthopelta s. str., Tenagopelta n. gen. has mobile eyestalks (immobile in Hephthopelta s. str.); a relatively wider thoracic male sternum (notably sternites 1‒4) but also evident in females ( Figs. 54 F; 88C) (distinctly narrower in Hephthopelta s. str., e.g., Figs. 52 A; 91A); the epistome is relatively narrow with a triangular median lobe and sinuous lateral margins with marked fissures) (e.g., Fig. 26 F) (relatively broad, truncated and nearly rectangular with the lateral margins semicircular without fissures in Hephthopelta s. str., e.g., Fig. 24 A); the merus of the third maxilliped is more quadrate (e. g., Fig. 33 G) (subovate in Hephthopelta s. str., e.g., Fig. 31 A); the base of the basal segment of the G2 is distinctly elongated (e.g., Fig. 79 O) (short and rounded in Hephthopelta s. str., e.g., Fig. 71 M); and the distal segment of the G2 is distinctly narrowed medially with the proximal part wider than the distal part (e.g., Fig. 79 E) (G2 distal segment is of uniform width in Hephthopelta s. str., e.g., Fig. 71 E). There are also several other characters that separate the two genera but they may prove to be more species-specific when more species are discovered in the future: the teeth on the inner margin of the cheliped merus are typically shorter in Tenagopelta n. gen. ( Fig. 42 I) than in Hephthopelta ( Fig. 42 G) and male pleonite 6 relatively narrower (e.g., Fig. 54 F) than in Hephthopelta (e.g., 52A) with the swollen area of postero-lateral regions of somite 3‒5 less marked in Tenagopelta n. gen. (e.g., Fig. 61 E) than in Hephthopelta (e.g., Fig. 59 A).

The remaining species previously placed in Hephthopelta may be placed in two major groups, both with G2s distinctly shorter than the G1 and the distal segment of the G2 poorly developed. One group has immobile eyestalks that are completely immobile in their orbits, the ischium and merus of the third maxilliped are short and quadrate and the G2 is short, half or less than half the length of the G1 ( Angustopelta n. gen. and Statommatia n. gen.). The other group has mobile eyestalks, the ischium and merus of the third maxilliped are more elongated, and the G2 is about three-quarters the length of the G1 ( Chinommatia n. gen. and Notopelta n. gen.).

The two genera with immobile eyestalks are easily distinguished by the form of their thoracic sterna and gonopods. One genus ( Angustopelta n. gen.) has a relatively narrow anterior male thoracic sternum (sternites 1‒4) that is also evident in adult females (e.g., Fig. 52 D); the female sterno-pleonal cavity is relatively deep, with the vulvae on the steep outer margin of the sterno-pleonal cavity (e.g., Fig. 91 B); and the G1 is stout and medially bent (e.g., Fig. 73 A). The second genus ( Statommatia n. gen.) has a relatively narrower anterior male thoracic sternum (sternites 1‒4) also evident in adult females (e.g., Fig. 54 B); the female sterno-pleonal cavity is relatively shallow, with the vulvae mesial in the flat portion of thoracic sternite 6 (e.g., Fig. 91 E); and the G1 is more slender, with the distal half almost lanceiform and straight (e.g., Fig. 77 A).

Inner margin of major Gently granuliform Gently granuliform (e.g., Gently granuliform (e.g., With long prolongation Gently granuliform (e.g., Gently granuliform (e.g., cheliped basis-ischium in (e.g., Fig. 42 G) Fig. 42 H) Fig. 42 J, K) visible from dorsal view Fig. 42 M, N) Fig. 42 I) adults ( Figs. 15 G; 42L)

……continued on the next page Character Hephthopelta s. str. Angustopelta n. gen. Chinommatia n. gen. Notopelta n. gen. Statommatia n. gen. Tenagopelta n. gen. Fingers of minor chela Distinctly laterally Subcircular in Subcircular in Subcircular in cross-section, Subcircular in Subcircular in

flatteneđ, broađ cross-section, not cross-section, not not đistinctly laterally cross-section, not cross-section, not throughout most of đistinctly laterally đistinctly laterally flatteneđ, gently tapering to đistinctly laterally đistinctly laterally length; shear-like, cutting flatteneđ, gently tapering flatteneđ, gently tapering tip; scissor-like, cutting flatteneđ, gently flatteneđ, gently tapering margins with low, broađ to tip; scissor-like, cutting to tip; scissor-like, cutting margins with đistinct sharp tapering to tip; to tip; scissor-like, cutting teeth (e.g., Fig. 40 E) margins with đistinct margins with đistinct teeth ( Fig. 45 B) scissor-like, cutting margins with đistinct

sharp teeth (e.g., Fig. sharp teeth (e.g., Fig. margins with đistinct sharp teeth (e.g., Fig.

41D) 43B) sharp teeth (e.g., Fig. 47 B)

45D)

The group with mobile eyestalks also contains two genera ( Chinommatia n. gen. and Notopelta n. gen.), separated by the form of their eyestalks, thoracic sterna, and chelipeds. Chinommatia n. gen. has an eye peduncle that is evenly subcylindrical in cross-section (e.g., Fig. 25 B); a proportionately wider anterior male thoracic sternum (sternites 1‒4) that is also evident in adult females (e.g., Fig. 53 A); and the basis-ischium of the male major cheliped does not have an elongated process (e.g., Fig. 14 B). In contrast, Notopelta n. gen. has the median part of the eye peduncle distinctly constricted and dorso-ventrally flattened, appearing reniform ( Fig. 25 G); a proportionately broader anterior male thoracic sternum (sternites 1‒4) ( Figs. 53 F); and the male major cheliped basis-ischium has a markedly long hook-like process (e.g., Fig. 42 L).

Distribution. Indo-West Pacific region: Indian and western Pacific oceans.

Figures

FIGURE 14. Overall dorsal view of species of Chinommatia n. gen. A, C. cavimanus, Philippines: holotype male 11.4 × 13.9 mm (USNM 46413); B, C. cavimanus, Philippines: male 10.1 × 12.2 mm (ZRC 2015.228); C, C. cavimanus, Philippines: female 7.6 × 9.0 mm (ZRC 2015.225); D, C. cavimanus, Philippines: male 8.1 × 10.2 mm (ZRC 2015.228); E, C. cavimanus, Philippines: female 7.4 × 8.6 mm (ZRC 2015.227); F, South China Sea: male 6.1 × 7.3 mm (IOCAS-MBM 166773); G, C. bruuni, Vietnam: holotype female 4.0 × 5.0 mm (after Serène 1964: pl. 22 B); H, C. bruuni, Indonesia: male 3.4 × 4.0 mm (ZMUC CRU- 1524); I, C. bruuni, Malaysia: male 4.3 × 5.5 mm (ZRC 2013.0020); J, C. bruuni, Malaysia: female 5.5 × 7.2 mm (ZRC 2013.0020).

FIGURE 15. Overall dorsal view of species of Chinommatia n. gen. and Notopelta n. gen. A, Chinommatia littoralis, Indonesia: holotype female 3.5 × 4.9 mm (NNM-ZMA); B, Chinommatia bicuspida n. sp., Papua New Guinea: holotype male 10.7 × 12.9 mm (MNHN-IU- 2011 - 0954); C, Chinommatia bicuspida n. sp., Fiji: male 12.7 × 10.6 mm (MNHN-IU- 2013 - 9056); D, Chinommatia bicuspida n. sp., Papua New Guinea: paratype female 16.7 × 20.4 mm (MNHN-IU- 2011 - 3461); E, F, Chinommatia turpis n. sp., Vanuatu: holotype male 3.1 × 4.3 mm (MNHU-IU- 2013 - 9044); G, H, Notopelta mortenseni, Indonesia: holotype male 5.0 × 6.9 mm (ZMUC CRU- 7397); I, Notopelta mortenseni, Indonesia: female 5.9 × 4.1 mm (ZMUC CRU- 007768).

FIGURE 24. Frontal view of species of Hephthopelta and Angustopelta n. gen. A, Hephthopelta lugubris, Australia: male 7.9 × 8.3 mm (NMV J 61081); B, Hephthopelta occidentalis n. sp., Madagascar: holotype male 6.1 × 6.6 mm with sacculinid (MNHN-IU- 2010 - 1031); C, Angustopelta aurita, Philippines: holotype female 18.9 × 19.1 mm (USNM 46405); D, Angustopelta aurita, Philippines: male 9.2 × 10.6 mm (ZRC 2015.251); E, Angustopelta cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); F, Angustopelta robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616); G, Angustopelta modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038); H, Angustopelta aff. modesta n. sp., Papua New Guinea: female 9.0 × 10.2 mm (ZRC 2015.259).

FIGURE 25. Frontal view of species of Chinommatia n. gen. and Notopelta n. gen. A, Chinommatia cavimanus, Philippines: male 10.1 × 12.2 mm (ZRC 2015.228); B, Chinommatia bicuspida n. sp., Papua New Guinea: paratype female 16.7 × 20.4 mm (MNHN-IU- 2011 - 3461); C, Chinommatia bruuni, Java Sea: male 3.4 × 4.0 mm (ZMUC CRU- 1524); D, Chinommatia bruuni, Malaysia: female 5.5 × 7.2 mm (ZRC 2013.0020), Malaysia; E, Chinommatia littoralis, Indonesia: holotype female 3.5 × 4.9 mm (NNM-ZMA); F, Chinommatia turpis n. sp., Vanuatu: holotype male 3.1 × 4.3 mm (MNHU-IU- 2013 - 9044); G, Notopelta mortenseni, Indonesia: holotype male 5.0 × 6.9 mm (ZMUC CRU- 7397).

FIGURE 26. Frontal view of species of Statommatia n. gen. and Tenagopelta n. gen. A, Statommatia apta, Philippines: holotype 7.8 × 9.4 mm (USNM 46389); B, Statommatia apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); C, Statommatia malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a); D, Statommatia knudseni, New Caledonia: holotype male 4.7 × 6.9 mm (ZMUC CRU- 6974); E, Statommatia granulosa n. sp., New Caledonia: holotype male 4.1 × 4.7 mm (MNHN-IU- 2013 - 9454); F, Tenagopelta pacifica n. sp., Philippines: holotype male 10.3 × 11.9 mm (NMCR); G, Tenagopelta brachyphallus n. sp., Australia: holotype male 6.1 × 6.9 mm (NMV J 61680); H, Tenagopelta potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504).

FIGURE 31. Third maxilliped of species of Hephthopelta and Angustopelta n. gen. A, B, Hephthopelta lugubris, Australia: male 7.9 × 8.3 mm (NMV J 61081); C, Hephthopelta occidentalis n. sp., Madagascar: holotype male 6.1 × 6.6 mm with sacculinid (MNHN-IU- 2010 - 1031); D, Angustopelta aurita, Philippines: holotype female 18.9 × 19.1 mm (USNM 46405); E, Angustopelta aurita, Philippines: male 9.2 × 10.6 mm (ZRC 2015.250); F, Angustopelta cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); G, Angustopelta robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616); H, Angustopelta modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038); I, Angustopelta modesta n. sp., Solomon Islands: paratype male 11.5 × 13.2 mm (MNHU-IU- 2013 - 9039).

FIGURE 33. Third maxilliped of species of Statommatia n. gen. and Tenagopelta n. gen. A, Statommatia apta, Philippines: holotype male 7.8 × 9.4 mm (USNM 46389); B, Statommatia apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); C, Statommatia knudseni, New Caledonia: holotype male 4.7 × 6.9 mm (ZMUC CRU- 6974); D, Statommatia pubescens, South China Sea: holotype male 5.0 × 7.9 mm (after Chen 1998: fig. 23 - 5); E, Statommatia malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a); F, Statommatia granulosa n. sp., New Caledonia: holotype male 4.1 × 4.7 mm (MNHN-IU- 2013 - 9454); G, Tenagopelta pacifica n. sp., Philippines: holotype male 10.3 × 11.9 mm (NMCR); H, Tenagopelta brachyphallus n. sp., Australia: holotype male 6.1 × 6.9 mm (NMV J 61680); I, Tenagopelta potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504).

FIGURE 40. Chelae of species of Chasmocarcinops and Hephthopelta. A, B, Chasmocarcinops gelasimoides, Andaman Sea: male 8.8 × 9.5 mm (PMBC 16849); C, D, Chasmocarcinops gelasimoides, Singapore: male 9.5 × 10.3 mm (ZRC 2000.1341); E, F, Hephthopelta lugubris, Australia: male 7.9 × 8.3 mm (NMV J 61081); G, H, Hephthopelta occidentalis n. sp., Madagascar: holotype male 6.1 × 6.6 mm, with sacculinid (MNHN-IU- 2010 - 1031).

FIGURE 41. Chelae of species of Angustopelta n. gen. A, B, A. aurita, Philippines: male 9.2 × 10.6 mm (ZRC 2015.250); C, D, A. cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); E, F, A. cribrorum, Taiwan: male 9.9 × 11.6 mm (ZRC 2015.214); G, H, A. robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616); I, J, A. modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038).

FIGURE 42. P 4, P 5, and merus of cheliped of species of Angustopelta n. gen. A, A. robusta n. sp., Vanuatu: paratype male 12.3 × 13.3 mm (ZRC 2015.255); B, A. robusta n. sp., Vanuatu: paratype female 13.4 × 15.1 mm (MNHN-IU- 2010 - 5616); C, A. robusta n. sp., Vanuatu: paratype male 12.5 × 14.8 mm (MNHN-IU- 2013 - 9041); D, A. modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038); E, A. modesta n. sp., Solomon Islands: paratype female 13.1 × 15.3 mm (ZRC 2015.258); F, A. aff. modesta n. sp., Papua New Guinea: female 9.3 × 9.9 mm (MNHN-IU- 2011 - 3307); G, Hephthopelta lugubris, Papua New Guinea: female 9.8 × 10.9 mm (MNHN-IU- 2011 - 2657), major cheliped; H, Angustopelta aurita, Philippines: female 10.9 × 12.7 mm (ZRC 2015.248), minor cheliped; I, Tenagopelta pacifica n. sp., Indonesia: male 11.0 × 12.4 mm (MNHU-IU- 2013 - 9053), minor cheliped; J, Chinommatia cavimanus, Philippines: male 9.7 × 11.5 mm (ZRC 2015.234), minor cheliped; K, Chinommatia cavimanus, Philippines: male 7.6 × 9.3 mm (ZRC 2015.231), minor cheliped; L, Notopelta mortenseni, Indonesia: holotype male 5.0 × 6.9 mm (ZMUC CRU- 7397), major cheliped; M, Statommatia apta, Philippines: males 8.5 × 10.3 mm (ZRC 2015.237), minor cheliped; N, Statommatia malagasy n. sp., Madagascar: male 8.8 × 11.0 mm (MNHN-IU- 2010 - 4370 b), minor cheliped.

FIGURE 45. Chelae of species of Statommatia n. gen. and Notopelta n. gen. A, B, Notopelta mortenseni, Indonesia: holotype male 5.0 × 6.9 mm (ZMUC CRU- 7397); C, D, Statommatia apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); E, F, S. knudseni, New Caledonia: holotype male 4.7 × 6.9 mm (ZMUC CRU- 6974); G, H, Statommatia malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a); I, J, Statommatia granulosa n. sp., New Caledonia: holotype male 4.1 × 4.7 mm (MNHN-IU- 2013 - 9454).

FIGURE 47. Chelae of species of Tenagopelta n. gen. A, B, T. pacifica n. sp., Philippines: holotype male 10.3 × 11.9 mm (NMCR); C, D, T. brachyphallus n. sp., Australia: paratype male 6.9 × 7.6 mm (NMV J 54594); E ‒ H, T. potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504); I, J, T. potens, Papua New Guinea: male 8.8 × 10.0 mm (ZRC 2015.242).

FIGURE 52. Anterior thoracic sternum and male pleon of species of Hephthopelta and Angustopelta n. gen. A, Hephthopelta lugubris, Australia: male 7.9 × 8.3 mm (NMV J 61081); B, C, Hephthopelta occidentalis, Madagascar: holotype male 6.1 × 6.6 mm with sacculinid (MNHN-IU- 2010 - 1031); D, Angustopelta aurita, Philippines: male 9.2 × 10.6 mm (ZRC 2015.250); E, Angustopelta cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); F, A. cribrorum, Taiwan: male 9.9 × 11.6 mm (ZRC 2015.214); G, Angustopelta robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616); H, A. modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038).

FIGURE 53. Anterior thoracic sternum and male pleon of species of Chinommatia n. gen. and Notopelta n. gen. A, Chinommatia cavimanus, holotype male 11.4 × 13.9 mm (USNM 46413); B, Chinommatia cavimanus, Philippines: male 10.1 × 12.2 mm (ZRC 2015.228); C, Chinommatia bruuni, Indonesia: male 3.4 × 4.0 mm (ZMUC CRU- 1524); D, Chinommatia bicuspida n. sp., Papua New Guinea: holotype male 10.7 × 12.9 mm (MNHN-IU- 2011 - 0954); E, Chinommatia turpis n. sp., Vanuatu: holotype male 3.1 × 4.3 mm (MNHU-IU- 2013 - 9044); F, Notopelta mortenseni, Indonesia: holotype male 5.0 × 6.9 mm (ZMUC CRU- 7397).

FIGURE 54. Anterior thoracic sternum and male pleon of species of Statommatia n. gen. and Tenagopelta n. gen. A, Statommatia apta, Philippines: holotype 7.8 × 9.4 mm (USNM 46389); B, Statommatia apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); C, Statommatia malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a); D, E, Statommatia knudseni, New Caledonia: holotype male 4.7 × 6.9 mm (ZMUC CRU- 6974); F, Tenagopelta pacifica n. sp., Philippines: holotype male 10.3 × 11.9 mm (NMCR); G, Tenagopelta brachyphallus n. sp., Australia: holotype male 6.1 × 6.9 mm (NMV J 61680); H, Tenagopelta potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504).

FIGURE 59. Thoracic sternites 7 and 8, male pleon and sterno-pleonal cavity of species of Hephthopelta and Angustopelta n. gen. A, Hephthopelta lugubris, Australia: male 7.9 × 8.3 mm (NMV J 61081); B, Hephthopelta lugubris, Solomon Is: male 8.3 × 9.1 mm (MNHN-IU- 2013 - 9040); C, Hephthopelta occidentalis n. sp., Madagascar: holotype male 6.1 × 6.6 mm with sacculinid (MNHN-IU- 2010 - 1031); D, Angustopelta aurita, Philippines: male 9.2 × 10.6 mm (ZRC 2015.250); E, Angustopelta cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); F, Angustopelta robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616); G, Angustopelta modesta n. sp., Solomon Islands: holotype male 13.3 × 15.1 mm (MNHU-IU- 2013 - 9038); H, Angustopelta modesta n. sp., Solomon Islands: paratype male 11.5 × 13.2 mm (MNHU- IU- 2013 - 9039); I, Angustopelta cribrorum, Taiwan: male 9.9 × 11.6 mm (ZRC 2015.214); J, Angustopelta robusta n. sp., Vanuatu: holotype male 9.3 × 10.7 mm (MNHN-IU- 2010 - 5616). Abbreviations: pb = press buttom male pleonal locking mechanism.

FIGURE 61. Thoracic sternites 7 and 8 and male pleon of species of Statommatia n. gen. and Tenagopelta n. gen. A, Statommatia apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); B, Statommatia knudseni, New Caledonia: holotype 4.7 × 6.9 mm (ZMUC CRU- 6974); C, Statommatia malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a); D, Statommatia granulosa n. sp., New Caledonia: holotype male 4.1 × 4.7 mm (MNHN-IU- 2013 - 9454); E, Tenagopelta pacifica n. sp., Philippines: holotype male 10.3 × 11.9 mm (NMCR); F, Tenagopelta brachyphallus n. sp., Australia: holotype male 6.1 × 6.9 mm (NMV J 61680); G, Tenagopelta potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504).

FIGURE 71. Gonopods of Hephthopelta lugubris. A ‒ E, Australia: male 7.9 × 8.3 mm (NMV J 61081); F ‒ I, Solomon Is: male 8.3 × 9.1 mm (MNHN-IU- 2013 - 9040); J, Solomon Is: male 7.3 × 7.8 mm (MNHN-IU- 2013 - 9037); K ‒ M, Fiji: male 6.3 × 6.5 mm (MNHN-IU- 2013 - 9028). A, B, F, J, K, ventral view of left G 1; C, H, ventral view of distal part of left G 1; D, I, dorsal view of distal part of left G 1; E, M, left G 2; G, L, dorsal view of left G 1. Scales: A, B, E, F, G, J ‒ M = 0.5 mm; C, D, H, I = 0.1 mm.

FIGURE 73. Gonopods of species of Angustopelta n. gen. A, A. aurita, Philippines: male 7.7 × 9.5 mm (ZRC 2015.248); B ‒ E, A. aurita, Philippines: male 9.6 × 11.2 mm (ZRC 2015.250); F ‒ H, A. cribrorum, Japan: holotype male 13.6 × 14.9 mm (USNM 46388); I, A. cribrorum, Taiwan: male 9.9 × 11.6 mm (ZRC 2015.214). A, B, F, ventral view of left G 1; C, D, G, H, distal part of left G 1; E, I, left G 2. Scales: A, B, E, F, I = 0.5 mm; C, D, G, H = 0.1 mm.

FIGURE 77. Gonopods of species of Statommatia n. gen. A ‒ D, S. apta, Philippines: male 5.9 × 7.1 mm (ZRC 2015.239); E ‒ H, S. knudseni, New Caledonia: holotype male 4.7 × 6.9 mm (ZMUC CRU- 6974); I ‒ L, S. malagasy n. sp., Madagascar: holotype male 9.9 × 12.4 mm (MNHN-IU- 2010 - 4370 a). A ‒ C, ventral view of right G 1 [left one damaged]; E, I, ventral view of left G 1; B, dorsal view of distal part of right G 1; C, ventral view of distal part of right G 1; F, J, ventral view of distal part of left G 1; G, K, dorsal view of distal part of left G 1; D, H, L, left G 2. Scales: A, D, E, H, I, L = 0.5 mm; B, C, F, G, J, K = 0.1 mm.

FIGURE 79. Gonopods of species of Tenagopelta n. gen. A ‒ E, T. pacifica n. sp., Philippines: paratype male 10.2 × 11.4 mm (ZRC 2015.212); F ‒ J, T. brachyphallus n. sp., Australia: holotype male 6.1 × 6.9 mm (NMV J 61680); K ‒ N, T. potens, Papua New Guinea: male 11.8 × 13.8 mm (MNHN-IU- 2011 - 1504). A, F, K, ventral view of left G 1; B, G, L, dorsal view of left G 1; C, H, M, ventral view of distal part of left G 1; D, I, N, dorsal view of distal part of left G 1; E, J, O, left G 2. Scales: A, B, E, F, G, J, K, L, O = 0.5 mm; C, D, H, I, M, N = 0.1 mm.

FIGURE 91. Vulvae of species of Hephthopelta, Angustopelta n. gen., Chinommatia n. gen., Statommatia n. gen. and Tenagopelta n. gen. A, Hephthopelta lugubris, Australia: ovigerous female 6.3 × 6.6 mm (NMV J 61083); B, Angustopelta aurita, Philippines: female 8.7 × 10.0 mm (ZRC 2015.253); C, Angustopelta robusta n. sp., Vanuatu: paratype female 13.4 × 15.1 mm (MNHN-IU- 2010 - 5616); D, Chinommatia cavimanus, Philippines: female 7.6 × 9.0 mm (ZRC 2015.228); E, Statommatia apta, Philippines: female 8.5 × 10.6 mm (ZRC 2015.238); F, Statommatia malagasy n. sp., Madagascar: paratype female 9.4 × 8.1 mm (ZRC 2015.215); G, Tenagopelta pacifica n. sp., Philippines: female 9.2 × 10.4 mm (ZRC 2015.187); H, Tenagopelta brachyphallus n. sp., Australia: female 6.9 × 7.6 mm (NMV J 61680).

Tables

TABLE 2. Comparison of characters of Hephthopelta s. str. and five new genera described for species previously included in Hephthopelta s. lato.

Character Carapace shape	Hephthopelta s. str. Subtrapezoidal; slightly wider than long; strongly globose (e.g., Figs. 10A; 24A)	Angustopelta n. gen. Subtrapezoidal; wider than long; dorsal surface gently convex (e.g., Figs. 12A; 24E)	Chinommatia n. gen. Subtrapezoidal to subovate; wider than long; dorsal surfaces gently convex (e.g., Figs. 14B; 25B)	Notopelta n. gen. Subtrapezoidal; distinctly wider than long; dorsal surfaces gently convex (Figs. 15G; 25G)	Statommatia n. gen. Subtrapezoidal to subovate; distinctly wider than long; dorsal surfaces gently convex (e.g., Figs. 16A; 26A)	Tenagopelta n. gen. Subtrapezoidal; slightly wider than long; globose (e.g., Figs. 17A; 26H)
Anterolateral margins	With small tubercle or entire (e.g., Fig. 10A, C)	With sharp tubercle or tooth, or entire (e.g., Fig. 11A, E)	Usually entire, occasionally with low tooth or granule (e.g., Figs. 14E, F; 15F)	With straight, granular, submarginal crest (Fig. 15G, H)	Entire, never armed, sometimes granular (e.g., Fig. 16A)	With low tubercle or entire (e.g., Fig.17A, D)
Eye peduncles	Distinctly short, gently tapering to cornea; immobile (e.g., Fig. 24A)	Short, gently tapering to cornea; immobile (e.g., Fig. 24E)	Short to long, gently tapering to cornea; mobile (e.g., Fig. 25B, F)	Short, median part of peduncle distinctly constricted, dorso-ventrally flattened, appearing reniform; mobile (Fig. 25G)	Short, gently tapering to cornea; immobile (e.g., Fig. 26B)	Distinctly short, gently tapering to cornea; mobile (e.g., Fig. 26H)
Posterior margin of epistome	Median lobe distinctly broad (e.g., Fig. 24B)	Median lobe broad (e.g., Fig. 24D)	Median lobe broad to narrow (e.g., Fig. 25A, D)	Median lobe narrow (Fig. 25G)	Median lobe broad to narrow (e.g., Fig. 26B, D)	Median lobe broad (e.g., Fig. 26G)
Third maxillipeds	Ischium ovate, elongated; merus subquadrate to elongated (e.g., Fig. 31A–C)	Ischium quadrate to rectangular; merus rounded to quadrate (e.g., Fig. 31D–I)	Ischium rectangular; merus ovate (e.g., Fig. 32A–E)	Ischium rectangular; merus ovate (Fig. 32F)	Ischium rectangular to quadrate; merus ovate to quadrate (e.g., Fig. 33A–F)	Ischium rectangular; merus quadrate (e.g., Fig. 33G–I)
Ventral surface of cheliped merus	Outer margin with 3 or 4 long sharp teeth; inner margin unarmed (Fig. 42G)	Outer margin with 3 or 4 sharp teeth; inner margin with strong distal tooth (Fig. 42H)	Outer margin with 4 or more sharp tubercles or dentiform granules, sometimes low; inner margin unarmed (Fig. 42J, K)	Outer margin with row of sharp tubercles; inner margin unarmed (Fig. 42L)	Outer margin granuliform to dentiform with more than 4 sharp tubercles; inner margin unarmed (Fig. 42M, N)	Outer margin with 3 or 4 sharp teeth; inner margin unarmed (Fig. 42I)
Inner margin of cheliped carpus	Distal angle with long distal tooth (e.g., Fig. 10D, I); proximal part rounded (e.g., Fig. 10F) to dentiform (e.g., Fig. 10I)	Distal angle with long tooth (e.g., Fig. 11E); proximal part rounded (e.g., Fig.11E) to slightly angular (e.g., Fig. 11C)	Distal angle with short tooth (e.g., Fig. 14A, B, E); proximal part rounded (e.g., Fig. 14A, E)	Distal angle with short tooth (Fig. 15G); proximal part rounded (Fig. 15G)	Distal angle with short tooth (e.g., Fig. 16A, B); proximal part rounded (e.g., Fig. 16A, E)	Distal angle with long tooth (e.g., Fig. 17A, B); short proximal part angular (e.g., Fig. 17B) or prominently dentiform (e.g., Fig. 17A, C)