On the Indian species of Eurycarcinus A. Milne-Edwards, 1867, Heteropanope Stimpson, 1858, and Pilumnopeus A. Milne-Edwards, 1867 (Decapoda: Brachyura: Pilumnidae)

Trivedi, Jigneshkumar; Mitra, Santanu; Patel, Pooja; Maheta, Niketa; Patel, Krupal; Ng, Peter K. L.

doi:10.1590/2358-2936e2021004

Abstract

Five species of pilumnid crabs assigned to Eurycarcinus A. Milne-Edwards, 1867, Heteropanope Stimpson, 1858, and Pilumnopeus A. Milne-Edwards, 1867, have been reported from India: E. orientalis A. Milne-Edwards, 1867, E. bengalensis Deb, 1999, H. glabra Stimpson, 1858, H. neolaevis Deb, 1995, and P. convexus (Maccagno, 1936). The identity of E. bengalensis is confused and the species had been provisionally transferred to Heteropanope. Examination of the types however, confirms the affinities of the species with Eurycarcinus and consequently extends the range of the genus to the eastern Indian Ocean. A re-examination of the types of H. neolaevis show that it is a junior subjective synonym of Aniptumnus quadridentus (De Man, 1895), and is the first record of the genus from India. Eurycarcinus integrifrons De Man, 1879, is also formally recorded for the first time from India.

Keywords
Aniptumnus ; Eurycarcinus ; Gujarat; Heteropanope ; India; new records; Pilumnopeus ; synonymy; taxonomy; West Bengal

INTRODUCTION

Davie (1989) revised the taxonomy of the Indo-West Pacific pilumnid genera HeteropanopeStimpson, 1858, and Pilumnopeus A. Milne-Edwards, 1867, and transferred several species to his new genus, Benthopanope Davie, 1989. The three genera were distinguished by carapace morphology, structure of the frontal margin, basal antennal article, male thoracic sternum, and male pleon (Davie, 1989). Until then, the taxonomy of these similar-looking genera was confused. Although Davie (1989: 130) noted that the Indian Ocean Eurycarcinus A. Milne-Edwards, 1867, was close to Heteropanope, but, as he did not have access to specimens, was unable to discuss the characters distinguishing these two genera.

In India, two species of Eurycarcinus have previously been recorded: E. natalensis (Krauss, 1843) Andhra Pradesh (Dev Roy and Nandi, 2007; Dev Roy, 2008); Tamil Nadu (Thomas, 1969; Dev Roy, 2008; Kathirvel and Gokul, 2010); West Bengal (Deb, 1995; Dev Roy and Nandi, 2012); Andaman and Nicobar Islands (Thomas, 1969; Bakus, 1994; Venkataraman et al., 2004), and E. orientalis A. Milne-Edwards, 1867 Gujarat (Chhapgar, 1957; Trivedi et al., 2012; Pandya and Vachhrajani, 2013); Maharashtra (Chhapga 1957; Pati et al., 2012); Goa (Dev Roy and Bhadra, 2008; Dev Roy, 2013); Karnataka (Chhapgar, 1957; Haragi et al., 2010); Kerala (Dev Roy, 2013); Andhra Pradesh (Dev Roy and Nandi, 2007; Dev Roy, 2008; Rath and Dev Roy, 2009); Orissa (Dev Roy and Rath, 2017); West Bengal (Dev Roy and Nandi, 2012); Andaman and Nicobar Islands (Dev Roy and Nandi, 2012). Two species in these genera were later described from specimens collected in the Bay of Bengal, India: Heteropanope neolaevis Deb, 1995, and Eurycarcinus bengalensis Deb, 1999. Subsequently, Trivedi et al. (2015) and Gosavi et al. (2017) recorded Heteropanope glabraStimpson, 1858, and Pilumnopeus convexus (Maccagno, 1936), respectively, from India.

Ng et al. (2018) revised Eurycarcinus, and distinguished it from Heteropanope by the following characters: thoracic sternite 8 remains clearly exposed when the male pleon is closed, male pleonal somite 3 is proportionately wider with the lateral margin distinctly projecting on lateral side, and male thoracic sternites 3 and 4 are transversely wider with the sterno-pleonal cavity close to the margin between sternites 3 and 4. On the basis of the published figures of Deb (1995, 1999), Ng et al. (2018) suggested that E. bengalensis can be assigned to Heteropanope on the basis of carapace shape but was uncertain about H. neolaevis because its gonopod structure was atypical for the genus. Trivedi et al. (2018: 59) noted that the records of E. natalensis from different parts of India were suspect and more likely belong to species of Heteropanope instead. Eurycarcinus natalensis is known for certain only from southern and eastern Africa (Ng et al., 2018). Trivedi et al. (2018: 59) also expressed doubts about the records of E. orientalis from eastern India, a species otherwise only known for certain from the western Indian Ocean (Ng et al., 2018).

The present paper examines and describes new material of the above genera including the types of H. bengalensis and H. neolaevis. Eurycarcinus integrifrons De Man, 1879 is formally recorded here for the first time from India. Eurycarcinus bengalensis is shown to be a valid taxon but the type material contains two species, with the paratype females belonging to E. integrifrons instead. In addition, H. neolaevis is shown to be a junior subjective synonym of Aniptumnus quadridentatus (De Man, 1895).

MATERIAL AND METHODS

The specimens examined are deposited in the Zoological Reference Collection, Department of Life Sciences, Hemchandracharya North Gujarat University, Patan, Gujarat, India (LFSc.ZRC); Zoological Survey of India, Kolkata, India (ZSI); and the Zoological Reference Collection of the Lee Kong Chian Natural History Museum, National University of Singapore (ZRC). The terminology used in the description follows Davie et al. (2015) and Ng et al. (2018). The following abbreviations are used: CL = carapace length; CW = carapace width; G1 = male first gonopod; G2 = male second gonopod; coll. = collector. All the measurements are in millimeters (mm).

TAXONOMY

Family Pilumnidae Samouelle, 1819

Genus Eurycarcinus A. Milne-Edwards, 1867

Eurycarcinus orientalis A. Milne-Edwards, 1867

(Figs. 1, 9A-C)

Figure 1.
Eurycarcinus orientalis A. Milne-Edwards, 1867, male (CW 41.5 mm, CL 26.9 mm) (LFSc.ZRC-64), Kamboi, India. A, habitus, dorsal view; B, frontal view; C, anterior thoracic sternum (sternites 1-4) and pleon; D, chelae, outer view; E, posterior thoracic sternum and pleon.

Eurycarcinus orientalis A. Milne-Edwards, 1867: 277; Jones, 1986: 162, pl. 47; Vousden, 1987: 36, tabs. 4, 7; Apel and Türkay, 1992: 194, 204, 205; Ismail and Ahmed, 1993: 158; Apel 1994a: 43, 44; Apel, 2001: 97, 98; Al-Khayat and Jones, 1996: 806, fig. 5; Al-Khayat and Jones, 1999: 58, 61; Tirmizi and Ghani, 1996: 30-32, fig 10; Cooper, 1997: 168-170, figs. 4, 5, 15; Hornby, 1997: 16 (? part); Ng et al., 2008: 140 (list); Naderloo and Türkay, 2012: 37; Al-Khafaji et al., 2017: 363, fig. 3; Naderloo, 2017: 304, figs. 26.10c, 26.11, 26.13; Ng et al., 2018: 484, figs. 1C, 3B, 4B, 5B, 6B, E, 7B, E, H, 8B, F, G; Trivedi et al., 2018: 59 (list).

Eurycarcinus grandidieri - Alcock, 1898: 211, 212 [not Eurycarcinus grandidierii A. Milne-Edwards, 1867 = E. natalensis (Krauss, 1843)].

Eurycarcinus sp. - Basson et al., 1977: 58, 228 (in list), fig. 38; Titgen, 1982: 131.

Type locality. Mumbai, India.

Material examined. 5 males (CW 27.6-41.5 mm, CL 17.5-26.9 mm), 4 females (CW 28.5-6.4 mm, CL 17.5-23.0 mm), LFSc.ZRC-64, Kamboi (22°12’59"N 72°36’59”E), Gujarat State, India, 11 April 2014, coll. J. Trivedi.

Remarks. The specimens examined in the present study agree with the description and figures of the species provided by Ng et al. (2018). Eurycarcinus orientalis resembles E. natalensis in carapace shape and dentition of the anterolateral border but can be separated from the latter in the following characters: the carapace is high (Fig. 1A) (not prominently raised in E. natalensis; cf. Ng et al., 2018: figs. 1A; 6A, D); suborbital and pterygostomial regions are prominently granulated (Fig. 1B) (less granulated in E. natalensis; cf. Ng et al., 2018: fig. 6D); G1 distal part gently recurved, tip gently curved (Fig. 9A, B) (distal part prominently recurved, tip hook shaped in E. natalensis; cf. Ng et al., 2018: fig. 8A, D, E).

Eurycarcinus orientalis is so far reported from the Gulf of Aden, Persian Gulf, Gulf of Oman, Pakistan, India, and Thailand (Naderloo, 2017; Ng et al., 2018). In India, the species is recorded from Gujarat, Maharashtra, Karnataka (Chhapgar, 1957) and Kerala (Dev Roy, 2013). Trivedi et al. (2018) commented that all the records of this species from the east coast including Andhra Pradesh (Dev Roy and Nandi, 2007), Orissa (Dev Roy and Rath, 2017), West Bengal (Dev Roy and Nandi, 2012), and Andaman and Nicobar Islands (Dev Roy and Nandi, 2012) require re-examination.

Trivedi et al. (2018) commented that the records of E. natalensis from India: Andhra Pradesh (Dev Roy and Nandi, 2007), Tamil Nadu (Thomas, 1969), West Bengal (Dev Roy and Nandi, 2012), and Andaman and Nicobar Islands (Thomas, 1969) are doubtful. Some of these may have been based on earlier incorrect identifications of Indian material by Alcock (1898) as Eurycarcinus grandidieri, the latter species being a junior synonym of E. natalensis. On the basis of the available specimens examined here, these records are probably a species of Heteropanope or Eurycarcinus and require re-examination, but, based on biogeography, they are likely to be E. bengalensis or E. integrifrons (see discussion for these species).

Eurycarcinus integrifrons De Man, 1879

(Figs. 2, 9D-F)

Figure 2.
Eurycarcinus integrifrons De Man, 1879, male (CW 28.2 mm, CL 19.9 mm) (LFSc.ZRC-63), Lakhpat, India. A, habitus, dorsal view; B, frontal view; C, anterior thoracic sternum (sternites 1-4) and pleon; D, chelae, outer view; E, posterior thoracic sternum and pleon.

Eurycarcinus integrifrons De Man,1879: 55, 56; Apel, 1994b: 415, 433, 434; Al-Ghais and Cooper, 1996: 425, 426; Apel, 2001: 97; Naderloo and Sari, 2007: 344, tab. 1; Özcan et al., 2010: 507, fig. 2; Naderloo and Türkay, 2012: 36; Naderloo et al., 2013: 449, 456, tab 2; Naderloo, 2017: 303, figs. 26.10b, 26.11, 26.12; Ng et al., 2018: 484, figs. 1C,3B, 4B, 5B, 6B, E, 7B, E, H, 8B, F, G; Trivedi et al., 2018: 59 (list).

Eurycarcinus orientalis - Alcock, 1898: 210, 211 (part); Chhapgar, 1957: 436, 437, pl. 11d-f; Tirmizi et al., 1986: 8-10, fig. 3a-d; Tirmizi and Ghani, 1996: 30-32, fig. 10; ? Hornby, 1997: 16 (part) [not Eurycarcinus orientalis A. Milne-Edwards, 1867].

Litocheira (amoyensis Gordon?) - Stephensen, 1946: 169-171, fig. 46; Titgen, 1982: 252 (list) [not Litocheira amoyensis Gordon, 1931 = Heteropilumnus amoyensis (Gordon, 1931)].

Eurycarcinus sp. (? integrifrons) - Apel and Türkay, 1992: 194-195, 204-205.

Eurycarcinus bengalensisDeb, 1999: 376 (part).

Type locality. Unknown, probably India (cf. Naderloo 2017).

Material examined. 5 males (CW 13.2-28.2 mm, CL 9.66- 9.9 mm), 5 females (CW 11.3-25.2 mm, CL 18.6-8.14 mm), LFSc.ZRC-63, Lakhpat (23˚50’01"N 68˚46’26”E), Gujarat State, India, 27 March, 2015, coll. J. Trivedi; 2 ovigerous females (CW 13.9 mm, CL 10.4 mm; CW 21.3 mm, CL 15.3 mm), ZSI-C3349/2 (part), Chamta Block, Sunderbans Tiger Reserve, West Bengal State, India (paratypes of Eurycarcinus bengalensisDeb, 1999).

Remarks. The specimens examined in the present study agree with the description and figures of the species in Ng et al. (2018). Eurycarcinus integrifrons differs from E. natalensis and E. orientalis in the following characters: carapace subquadrate (Fig. 2A) (transversely ovate in E. natalensis and E. orientalis; cf. Ng et al., 2018: figs. 1A, B, C; 6A, B); frontal margin weakly bilobed (Fig. 2B) (distinctly bilobed in E. natalensis and E. orientalis; cf. Ng et al., 2018: figs. 1A-C; 6A, B).

The species is so far reliably reported from Gulf of Aden, Persian Gulf, Gulf of Oman, and Pakistan (Naderloo, 2017; Ng et al., 2018). The present study is actually the first confirmed record of the species from India, even though Naderloo (2017) suggested that the type may have originally been from there. The specimens from Gujarat (Kolak and Umarsadi) and Maharastra (Mumbai) (recorded as E. orientalis) in Chhapgar (1957) should be referred to E. integrifrons; his figures leave no doubt. These records of Chhapgar (1957) thus extend the known range of E. integrifrons nearly 900 kilometers further south along the coast of western India. The two ovigerous female paratypes of E. bengalensisDeb, 1999, are here also re-identified as E. integrifrons, extending the range of this species to West Bengal state (see remarks for next species).

Eurycarcinus bengalensis Deb, 1999

(Figs. 3, 4, 9G-I)

Figure 3.
Eurycarcinus bengalensisDeb, 1999, holotype male (CW 25.1 mm, CL 16.9 mm) (ZSI-C3349/2), Chamta Block, India. A, habitus, dorsal view; B, frontal view; C, anterior thoracic sternum (sternites 1-4) and pleon; D, chelae, outer view; E, posterior thoracic sternum and pleon.

Figure 4.
Eurycarcinus integrifrons De Man, 1879, paratype ovigerous female of Eurycarcinus bengalensisDeb, 1999 (CW 21.3 mm; CL 15.3 mm) (ZSI-C3349/2), Chamta Block, India. A, habitus, dorsal view; B, frontal view; C, chelae, outer view; D, anterior thoracic sternum (sternites 1-4) and pleon.

Eurycarcinus bengalensisDeb, 1999: 376 (part), fig. 2.

Heteropanope bengalensis - Ng et al., 2018: 474; Trivedi et al., 2018: 59.

Material examined. Holotype, male (CW 25.1 mm; CL 16.9 mm), ZSI-C3349/2, Chamta Block, Sunderbans Tiger Reserve, West Bengal State, India, coll. S. Bhuinya. Paratypes, 2 males (CW 11.5 mm; CL 7.8 mm, CW 11.7 mm; CL 8.6 mm), data as per holotype.

Remarks.Deb (1999) described E. bengalensis on the basis of a holotype and an unspecified number of specimens collected from Sundarbans Tiger Reserve, West Bengal state, India. Ng et al. (2018) provisionally transferred this species to Heteropanope on the basis of the figure and description of the type specimen given in Deb (1999). In the present study, the type specimens of the species were examined and identified as Eurycarcinus because the thoracic sternite 8 of the specimen remains exposed when the male pleon is closed (Fig. 3E) and pleonal somite 3 is relatively wider, with the lateral margin projecting (Fig. 3C), which are characteristics of Eurycarcinus.

The type series of E. bengalensis is mixed. Of the five specimens, the two paratype males agree well with the holotype male in most aspects and are clearly conspecific. The two ovigerous females (CW 13.9 mm, CL 10.4 mm; CW 21.3 mm, CL 15.3 mm), however, belong to E. integrifrons instead. The carapace shape and features (Fig. 4) are clearly of this species.

Eurycarcinus bengalensis s. str. is similar to E. integrifrons in the following characters: the carapace is not prominently swollen and high in frontal view (Figs. 2A, 3A, 4A), the frontal margin is weakly bilobed (Figs. 2B, 3B, 4B), there are distinct clumps of short setae between the first to third anterolateral teeth (Figs. 2A, 3A, 4A), the supraorbital margin is smooth without granules, the sub-orbital and pterygostomial regions are smooth and glabrous with dense setae only along the sutures (Figs. 2B, 3B, 4B), and the posterior margin of the epistome has the median lobe truncate, protruding anteriorly and separated from lateral margin by distinct rounded angle (Figs. 2B, 3B, 4B). Eurycarcinus bengalensis, however, differs from E. integrifrons in several characters that cannot be explained by variation: the carapace of E. bengalensis is proportionately wider (Figs. 3A, 4A) (more quadrate in E. integrifrons; Fig. 2A); the anterolateral teeth are lobiform and wide, separated from each other by a fissure (Figs. 3A, 4A) (more narrow dentiform anterolateral margin with the teeth separated by a U-shaped hiatus in E. integrifrons; Fig 2A); the male chela is proportionately shorter (Fig. 3D) (more elongate in E. integrifrons; Fig 2D); male pleonal somite 6 is distinctly trapezoidal in shape (Fig. 3C) (less so in E. integrifrons; Fig. 2C); male pleonal somites 1-3 are proportionately less broad (Fig. 3E) (proportionately wider in E. integrifrons; Fig. 2E); and the distal part of the G1 is relatively shorter with the tip gently upcurved (Fig. 9G, H) (G1 distal part relatively longer with the tip almost straight or gently curved downwards in E. integrifrons; Fig. 9D, E).

The relative width of the carapace and proportions of male pleonal somites 1-3 of E. bengalensis (Figs. 3A, 4A, 3E) are actually closer to the condition in E. orientalis (Figs. 1A, E) but the other characters (notably carapace shape, anterolateral armature, posterior margin of the epistome, male pleon shape and G1) do not match.

The characters possessed by E. bengalensis are interesting. While its carapace closely resembles species of Heteropanope as indicated by Ng et al. (2018), the male sternal and pleonal characters are those of Eurycarcinus. It is possible that some, if not all the records of “Eurycarcinus natalensis” and “Eurycarcinus orientalis” from the eastern Indian Ocean (e.g., Dev Roy, 2008; Rath and Dev Roy, 2009; see under E. orientalis) actually belong to E. bengalensis instead. The material will need to be examined.

Genus Pilumnopeus A. Milne-Edwards, 1867

Pilumnopeus convexus ( Maccagno, 1936 )

(Figs. 5, 10A-C)

Figure 5.
Pilumnopeus convexus (Maccagno, 1936), male (CW 15.3 mm, CL 11.3 mm) (LFSc.ZRC-70), Kuda, India. A, habitus, dorsal view; B, chelae, outer view; C, anterior thoracic sternum (sternites 1-4) and pleon; D, posterior thoracic sternum and pleon.

Heteropanope convexaMaccagno, 1936: 176, 177.

Pilumnopeus salomonensis Ward, 1942: 96, pl. 6 fig. 11; Davie, 1989: 143; Ng et al., 2008: 141 (list).

Pilumnopeus vauquelini - Stephensen, 1946: 141, fig 35a, b; Guinot, 1967: 275; Basson et al., 1977: 228, 231; Titgen, 1982: 252 (list); Hornby, 1997: 15; Naderloo and Türkay, 2012: 37 [not Pilumnus vauquelini Audouin, 1826 = Pilumnopeus vauquelini (Audouin, 1826)].

Pilumnopeus indica - Barnard, 1955: 30, fig. 12 [not Heteropanope indica De Man, 1887 = Benthopanope indica (De Man, 1887)].

Pilumnopeus convexa - Davie, 1989: 142, 143, fig. 7A-C.

Pilumnopeus convexus - Cooper, 1997: 171-173, figs. 6, 16; Apel, 2001: 98; Ng et al., 2008: 141 (list); Naderloo and Türkay, 2012: 37; Ghory et al., 2013: 301-312, figs. 1-5; Naderloo et al., 2013: 449, tab. 1; Gosavi et al., 2017: 429-433, figs. 2-7; Naderloo, 2017: 311, figs. 26.21, 26.22a, 26.23.

Figure 6.
Heteropanope glabraStimpson, 1858, male (CW 14.4 mm, CL 10.1 mm) (LFSc.ZRC-66), Ghogha mangroves, India. A, habitus, dorsal view; B, chelae, outer view; C, anterior thoracic sternum (sternites 1-4) and pleon; D, posterior thoracic sternum and pleon.

Type locality. Ethiopia.

Material examined. 1 male (CW 15.3 mm; CL 11.3 mm), 2 females (CW 11.9 mm; CL 8.4 mm; CW 12.8 mm; CL 8.6 mm), LFSc.ZRC-70, Kuda (21˚37’33"N 72˚18’17”E), Gujarat State, India, 12 October, 2016, coll. J. Trivedi: 1 male (CW 9.6 mm; CL 6.4 mm), 1 female (CW 5.9 mm; CL 4.3 mm), ZRC 2012.0422, Somar Goth, Pakistan, 26 January 2009, coll. F. Ghory; 1 female (CW 8.7 mm; CL 5.9 mm) ZRC 2018.0067, Buleii, Pakistan, 4 March 1986, coll. Q. Kazmi; 1 female (CW 14.4 mm; CL9.8 mm) (paralectotype of Pilumnopeus salomonensis Ward, 1942), ZRC 2012.0794, Salomón Islands, Chagos Archipelago, 1936, coll. R. Viader and G. Antelme; 2 males (CW 9.0 mm; CL 6.3 mm; CW 8.0 mm; CL 5.6 mm) ZRC 2018.1364, Persian Gulf, Iran, May 2010, coll. M. Safaei.

Remarks. The specimens examined (Figs. 5, 10A-C) in the present study compared well with the description and figures in Ghory et al. (2013). Pilumnopeus convexus is close to P. serratifrons but varies from the latter in having a CW/CL ratio of 1.4 to 1.5 and a less projecting front with a shallow central notch (Davie, 1989). The specimens were collected (Figs. 6, 11D-F) from Ekkakula mangroves located in Odisha State.

The species is so far reported from South Africa, Red Sea, Gulf of Aden, Persian Gulf, and Gulf of Oman (Naderloo, 2017). In India, the species was recorded from Gujarat (Gosavi et al., 2017) located on the west coast of India.

Genus Heteropanope Stimpson, 1858

Heteropanope glabra Stimpson, 1858

(Figs. 6, 10D-F)

Heteropanope glabraStimpson, 1858: 35.

Eurycarcinus maculatusAlcock, 1898: 212; Deb, 1999: 375.

Heteropanope glabra - Davie, 1989: 130-134, figs.1a-j, 2; Tan and Ng, 1994: 84 (list); Ng et al., 2008: 140 (list); Naderloo and Türkay, 2012: 37; Naderloo et al., 2013: 449, tab. 1; Trivedi et al., 2015: 1-5, figs. 1, 2; Naderloo, 2017: 307, figs. 26.10d, 26.15, 26.16; Trivedi et al., 2018: 59 (list). [For complete synonymy see Davie, 1989: 130; Sakai, 1976: 503; Dai and Yang, 1991: 377].

Type locality. Hong Kong.

Material examined. 4 males (CW 14.4-6.0 mm: CL 10.1-4.0 mm), 6 females (CW 12.3-7.2 mm: CL 8.3-5.4 mm), LFSc.ZRC-66, Ghogha (21°40’41"N 72°17’06”E), Gujarat State, India, 14 April, 2014, coll. J. Trivedi.

Remarks. The specimens examined in the present study (Fig. 6) agree with the description and figures in Davie (1989). The G1 tip (Fig. 10D, E) is slightly longer compared to that figured in Davie (1989). This difference can easily be explained by the size of the present specimens.

This species is so far reported from East Africa, Persian Gulf, Gulf of Oman, Pakistan, India, Andaman Sea, Thailand, Hong Kong, Taiwan, Mergui Archipelago, Singapore, Australia, and New Caledonia (Sakai, 1976;Davie, 1989; Dai and Yang, 1991; Trivedi et al., 2015; Naderloo, 2017).

Genus Aniptumnus Ng, 2002

Aniptumnus quadridentatus (De Man, 1895 )

(Figs. 7, 8, 10G-I)

Figure 7.
Aniptumnus quadridentatus (De Man, 1895), male (CW 13.9 mm; CL 9.6 mm) (ZSI-C1503/2) Matla River, India (lectotype of Heteropanope neolaevisDeb, 1995); A, habitus, dorsal view; B, chelae, outer view.

Figure 8.
Aniptumnus quadridentatus (De Man, 1895), male (CW 13.9 mm; CL 9.6 mm) (ZSI-C1503/2) Matla River, India (lectotype of Heteropanope neolaevisDeb, 1995); A, fourth ambulatory leg merus and basis-ischium lateral view; B, pleon; C, frontal margin; D, posterior thoracic sternum and pleon.

Figure 9.
A- C, Eurycarcinus orientalis A. Milne-Edwards, 1867, male (CW 41.5 mm, CL 26.9 mm) (LFSc.ZRC-64), Kamboi, India; D- F, Eurycarcinus integrifrons De Man, 1879, male (CW 28.2 mm, CL 19.9 mm) (LFSc.ZRC-63), Lakhpat, India; G-I, Eurycarcinus bengalensisDeb, 1999, holotype male (CW 25.1 mm, CL 16.9 mm) (ZSI-C3349/2), Chamta Block, India. A, D, G, ventral view of left G1; B, E, H, ventral view of distal part of left G1; C, F, I, left G2.

Figure 10.
A-C, Pilumnopeus convexus (Maccagno, 1936), male (CW 15.3 mm, CL 11.3 mm) (LFSc.ZRC-70), Kuda, India; D-F, Heteropanope glabraStimpson, 1858, male (CW 14.4 mm, CL 10.1 mm) (LFSc.ZRC-66), Ghogha mangroves, India; G-I, Aniptumnus quadridentatus (De Man, 1895), male (CW 13.9 mm; CL 9.6 mm) (ZSI-C1503/2) Matla River, India (lectotype of Heteropanope neolaevisDeb, 1995). A, D, G, ventral view of left G1; B, E, H, ventral view of distal part of left G1; C, F, I, left G2.

Pilumnus seminudus - De Man, 1887: 65 [not Pilumnus seminudus Miers, 1884 = Glabropilumnus seminudus (Miers, 1884)].

Pilumnus quadridentatus De Man, 1895: 537, fig. 6; Nobili, 1906: 278.

Parapilumnus quadridentatusBalss, 1933: 39 (list); Tan and Ng, 1994: 84 (list).

Heteropanope neolaevisDeb, 1995: 220; 1999: 374, figs. 3, 4; Ng et al., 2018: 475, 481-482; Trivedi et al., 2018: 59 (list).

Aniptumnus quadridentatus - Ng, 2002: 213, figs. 1, 2; Ng and Clark, 2008: figs. 13-18; Ng et al., 2008: 140 (list).

Type locality. Pontianak, West Kalimantan, Indonesia.

Material examined. Lectotype, male (CW 13.9 mm; CL 9.6 mm), ZSI-C1503/2, Matla River, Gangetic Delta, West Bengal State, India, December 1916, coll. S.W. Kemp. Paralectotypes, 4 males (CW 8.0-9.3 mm; CL 5.8-6.5 mm), same data as holotype.

Remarks.Ng et al. (2018: 481-482) clarified the date of publication for Heteropanope neolaevis, noting that the correct spelling of the name and citation should be Heteropanope neolaevisDeb, 1995

Deb (1995: 220) listed “50 examples” of this species (as H. neolaevis) from an estuarine area in the Matla River, Gangetic Delta, West Bengal State, India, and no types were designated. Deb (1999: 374) noted that she had “Several (about 50) specimens including holotype and paratypes from Gangetic delta, collected by S. W. Kemp, Z.S.I. Regd. No. C1503/2”. She did not specify the sex or size of the holotype. In the ZSI, there is one male specimen labeled as holotype which has the same data as indicated by Deb (1999) and is here recognized as the lectotype since no holotype was noted in the original paper (Deb, 1995).

Ng et al. (2018: 475) commented that the species is neither Heteropanope nor Pilumnopeus or Benthopanope, and the G1 figured (Deb, 1999: fig. 4) was unusual, being short and stout with the tip rounded. The lectotype and paralectotypes of H. neolaevis were examined and the Heteropanope neolaevis of Deb is here identified as Aniptumnus quadridentatus (De Man 1895). De Man (1895) described Pilumnus quadridentatus from a good series of specimens from the port of Pontianak, West Borneo, Indonesia. He also referred a specimen earlier obtained from Mergui (De Man, 1887) to this species. Nobili (1906) reported the species from Djibouti in the Red Sea, but his specimen should be re-examined in order to confirm its identification. Balss (1933) first referred it to ParapilumnusDe Man, 1895, but Ng (2002) reviewed the status of Parapilumnus and showed that this genus was actually not a pilumnid but an acidopsid. Ng (2002) selected a lectotype for P. quadridentatus and made it the type for a new genus, Aniptumnus, characterized by its subtruncate G1 tip, presence of sharp granules on the ventral margin of the basis-ischium and merus of the fourth ambulatory leg.

The types of H. neolaevis agree with the descriptions and figures of A. quadridentatus by De Man (1895), Ng (2002), and Ng and Clark (2008) and are considered conspecific. The carapace, cheliped and ambulatory leg characters all agree (Figs. 7a, b, 8a). The G1 of H. neolaevis figured by Deb (1999: Fig. 4) is inaccurate, being much shorter and stouter in her illustration. The actual G1 closely resembles that of A. quadridentatus, except that the tip is more rounded and less produced (Fig. 10G, H), but this can easily be explained by variation. In addition, the male sternite 8 of the types of H. neolaevis are exposed when the male pleon is closed (Fig. 8D; see also Ng and Clark, 2008; Hsueh et al., 2009).

Aniptumnus quadridentatus is a mangrove species, occurring in brackish waters. The type locality, Pontianak, is a port at the opening of the Kapuas River in Borneo and is surrounded by mangroves. In Malaysia and Singapore, the species is often found among fouling communities in mangrove and estuarine habitats. When present, it often occurs in large numbers.

ACKNOWLEDGEMENTS

The second author (SM) is most thankful to the Director, Zoological Survey of India, Kolkata, for encouragement during this study and providing research facilities. The authors are also thankful to Ms. Dhruva Trivedi for illustrations of specimens. We are most grateful to Peter Davie and Paul Clark for their many important suggestions which have significantly improved this paper.

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Publication Dates

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08 Feb 2021
Date of issue
2021

History

Received
22 May 2020
Accepted
29 Sept 2020

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[2] Al-Ghais, S. and Cooper, R. 1996. Brachyura (Grapsidae, Ocypodidae, Portunidae, Xanthidae and Leucosiidae) of Umm Al Quwain mangal, United Arab Emirates. Tropical Zoology, 9: 409-430.

[3] Al-Khafaji, K.K.; Khary, D.S. and Al-Malik, T.H.Y. 2017. First records of two pilumnid crabs: Pilumnopeus convexus Maccagno, 1936 and Eurycarcinus orientalis Edwards, 1867 (Crustacea, Decapoda, Pilumnidae) from Khor Al-Zubair Canal, South of Iraq. International Journal of Marine Science, 7(36): 361-365.

[4] Al-Khayat, J.A. and Jones, D.A. 1996. Two new genera, Manningis and Leptochryseus (Decapoda: Camptandriinae), and descriptions of the first zoea of six brachyurans from the Arabian Gulf. Journal of Crustacean Biology, 16: 797-813.

[5] Al-Khayat, J.A. and Jones, D.A. 1999. A comparison of the macrofauna of natural and replanted mangroves in Qatar. Estuarine, Coastal and Shelf Science, 49: 55-63.

[6] Apel, M. 1994a. Effects of the 1991 Gulf War oil spill on the crab fauna of intertidal mudflats in the western Arabian Gulf. Courier Forschungsinstitute Senckenberg, 166: 40-46.

[7] Apel, M. 1994b. Biology, ecology and taxonomy of brachyuran and paguridean Crustacea. p. 406-437. In: F. Krupp; A.H. Abuzinada and I.A. Nader (eds), Establishment of a marine habitat and Wildlife Sanctuary for the Gulf region final report for phase II. Jubail & Frankfurt, Frankfurt a. Main, CFC/WCWCD.

[8] Apel, M. 2001. Taxonomie und Zoogeographie der Brachyura, Paguridea und Porcellanidae (Crustacea: Decapoda) des Persisch-Arabischen Golfes. Frankfurt am Main, Germany, Johann Wolfgang Goethe Universität, Biologie und Informatik, Doctoral dissertation, 260p. [Unpublished]

[9] Apel, M. and Türkay, M. 1992. The intertidal crabs and hermit crabs (Crustacea: Decapoda: Brachyura and Paguridea) in the study area and their condition after the oil spill. Final report for phase I CEC-Project: establishment of a Marine Habitat and Wildlife Sanctuary for the Gulf Region, p 187-205.

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[13] Chhapgar, B.F. 1957. On the marine crabs (Decapoda, Brachyura) of Bombay State, Part I. Journal of Bombay Natural History Society, 54: 399-439.

[14] Cooper, R.T. 1997. Mangal-associated Brachyura (Ocypodidae, Grapsidae, Portunidae, Majidae, Xanthidae and Leucosiidae) from the north-eastern coastal Islands of Abu Dhabi, United Arab Emirates. Crustaceana, 70: 155-179.

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