Paramaya spinigera (De Haan, 1837)

Ng, Peter K. L. & Forges, Bertrand Richer De, 2015, Revision of the spider crab genus Maja Lamarck, 1801 (Crustacea: Brachyura: Majoidea: Majidae), with descriptions of seven new genera and 17 new species from the Atlantic and Indo-West Pacific, Raffles Bulletin of Zoology 63, pp. 110-225 : 155-156

Paramaya spinigera (De Haan, 1837)

( Figs. 6G , 21A–D , 23A–C , 37A , 40A , 42A , 45I , 50A , 52H , 54A , 56A )

Maja (Maja) spinigera De Haan, 1837: 93 .

Maja (Paramaya) spinigera – De Haan, 1837: errata.

Maja (Paramaya) spinigera – De Haan, 1837: pl. 24, fig. 4.

Maia spinigera – Adams & White, 1848: 15. – Parisi, 1915: 289.

Maja spinigera – Ortmann, 1893: 51. – Doflein, 1902: 656. – Ninni, 1924: 47, un- numbered figure on p. 48. – Balss, 1924: 34. – T. Sakai, 1934: 297. – T. Sakai, 1936: 98, pl. 25, fig. 3. – T. Sakai, 1938: 397, pl. 30, fig. 1. – T. Sakai, 1965: 84, pl. 38. – Serène, 1968: 57. – Utinomi, 1969: 77, pl. 39-3. – Holthuis & T. Sakai, 1970: 121, pl. 14. – Nishimura & Suzuki, 1971: 109, pl. 38-2. – Serène & Lohavanijaya, 1973: 50 (key). – Takeda, 1975: 127. – T. Sakai, 1976: 237, pl. 83. – Matsuzawa, 1977: pl. 96 fig. 4. – Takeda, 1982: 128. – Miyake, 1983: 43, pl. 14 fig. 4. – Dai et al., 1986: 136, pl. 18(3). – Matsuda et al., 1986: 130. – Dai & Yang, 1991: 151, pl. 18(3). – Yamaguchi & Baba, 1993: 359, Fig. 116. – Takeda, 1993: 43. – Huang, 1994: 582. – Muraoka & Odawara, 1995: 34. – Muraoka, 1998: 27. – Minemizu, 2000: 216. – Takeda et al., 2000: 139. – Ng et al., 2001: 12. – Maramura & Kosaka, 2003: 34. – Takeda & Ueshima, 2006: 74. – Yang et al., 2008: 780. – Ng et al., 2008: 117 (list).

Material examined. Japan – 1 female ( NSMT 5999 ), Kii- Nagashima, Kaino , Mie Prefecture, coll. M. Saba, 1970s . — 1 carapace only ( NSMT 7614 ), Kii-Nagashima, Kaino , Mie Prefecture, coll. M. Saba, 16 December 1976 . — 1 male, 1 female ( NSMT 5967 ), Kii-Nagashima, Kainoura , Mie Prefecture, coll. M. Saba, 1970s . — 1 male ( NSMT 6134 ), Kii-Nagashima, Kainoura , Mie Prefecture, in lobster gill-nets, coll. M. Saba, 25 February 1979 . — 1 male ( NSMT 10178 ), Kii-Nagashima, Kaino , Mie Prefecture, coll. M. Saba, 18 December 1978 . — 1 female ( SMF 47734), Toba , Mie Prefecture, coll. N. Yamashita, 1983 . — 1 male, 1 female ( SMF 9289 ), Toba , Mie Prefecture, coll. Nakamura, May 1980 . — 1 male ( NSMT 6032 ), Kii-Nagashima , Mie Prefecture, coll. M. Saba, 1970s . — 1 male ( NSMT 6752 ), Kii-Nagashima , Mie Prefecture, coll. M. Saba, 1970s . — 1 female ( NHM 1961.11.13.7/9, part), Seto, Shirahama, Japan, coll. I. Gordon. — 1 male, 1 female ( NHM 1961.11.13.7/9, part), Seto, Shirahama, Japan, coll. I. Gordon. — 1 male (64.7 × 51.2 mm) ( NHM 1961.6.5.117), Tosa Bay , coll. K. Sakai, April 1961 . — 1 male ( USNM 18854 ), Miura, Atami Province, Yokaito coast, Japan, from Mr Sakamoto, from Garrett Droppers . — 1 male ( SMF 30766), Katahara , Aichi, 80–150 m, coll. N. Akashi, 1978-1983 . — 1 male, 1 female ( SMF 7627 ), Kii Peninsula , coll. K. Sakai. — 2 males, 4 females, 1 ovigerous female ( SMF 46532), Kii Minabe , 33°44.944’N 135°19.65’E, coll. K. Sakai, 15 February 1983 . — 1 male ( SMF 47732), Shikoku, Usa, Ikenoura , Kochi Prefecture, coll. K. Sakai, 18 October 1984 . — 1 female ( SMF 47747), None, Cape Muroto , Kochi Prefecture, ca. 33°26.617’N 134°16.75’E, coll. K. Matsuzawa, 20 January 1993 . — 1 ovigerous female ( SMF 47746), Mitsu , Kochi Prefecture, 33°16.6’N 134°10.6’E, coll. 9 February 1974 . — 1 ovigerous female ( SMF 47748), Mitsu , Kochi Prefecture, 33°16.6’N 134°10.6’E, coll. 13 December 1994 . — 1 male ( SMF 47757), Mitsu, Muroto City , Kochi Prefecture, 33°16.6’N 134°10.6’E, no other data . — 1 female ( SMF 47741), Tokushima, Shikoku, Tsubakidomari, Anan , Kochi Prefecture, 33°51.678’N 134°43.603’E, coll. T. Nakano, 18 April 1983 . — 1 female ( SMF), Minabe, Wakayama Prefecture, coll. K. Sakai, 18 February 1984 . — 1 male ( SMF), Minabe, Wakayama Prefecture, T. Sakai Collection . — 1 female ( SMF 47742), Honshu, Wakayama Prefecture, Kii Mimase , coll. K. Sakai, 1 March 1988 . — 1 male ( NSMT 15905 ), no other data . — 1 dried female (70.1 × 58.8 mm) ( KPM NH4195 ), T. Sakai Collection . — 1 juvenile female ( USNM 48486 ), station 4880, eastern channel of Korean Straits, vicinity of Ohi-Shima , 59 fathoms, coll. RV Albatross, 2 August 1906 . Taiwan – 7 males (85.0 × 66.4 mm, 78.2 × 62.1 mm, 73.6 × 55.3 mm, 68.3 × 53.4 mm, 73.8 × 58.4 mm, 62.8 × 49.0 mm, 72.8 × 57.3 mm), 1 ovigerous female (63.0 × 48.6 mm) ( ZRC 1999.738 ), Longtong, near Keelung, northern Taiwan , in tangle nets for lobsters, coll. S.-H. Wu, May 1999 .

Diagnosis. Pseudorostral horns relatively short ( Fig. 21A–D ). Hepatic, lateral and branchial spines long; median row with 5 spines: 3 gastric, 1 cardiac, 1 intestinal; 2 strong spines on posterior carapace margin ( Fig. 21A–D ). Surface of thoracic sternum relatively smooth, with very low rounded granules ( Fig. 50A ). Chela of adult male with distinct carina on dorsal and ventral margins ( Fig. 54A ). Ambulatory meri in adults relatively stout, short ( Fig. 21A–D ). G1 gently curved ( Fig. 23A–C ).

Remarks. Yamaguchi & Baba (1993: 359) recorded two syntype female specimens (87.4 × 72.9 mm, 76.9 × 66.4 mm) in RMNH and selected the larger one as lectotype. The excellent figures in De Haan (1837: pl. 24, fig. 4) and Yamaguchi & Baba (1993: fig. 116) leave no doubt that the present specimens from Taiwan are conspecific with Paramaya spinigera s. str.

Alcock (1895: 239) and Alcock & Anderson (1898: pl. 34, fig. 3) recorded “ Maia spinigera ” from a specimen from off Beluchistan ( Pakistan) in the Indian Ocean (sex and size not mentioned) ( Fig. 22C, D ). We also have a photograph of a large specimen (not preserved) collected from Sri Lanka in deeper waters off a reef by aquarium collectors ( Fig. 22B ). The specimen was collected with a number of other relatively deeper reef animals at night (see Ng, 1994). Alcock’s (1895) and Alcock & Anderson’s (1898) specimen has relatively short ambulatory legs (notably the merus), suggesting it is unlikely to be real Paramaya spinigera (or either of the new species) as presently defined. However, this may be a consequence of its apparently small size (judging by the figure). The specimen is not in the NHM and is probably still in the Indian Museum (present day Zoological Survey of India). The larger specimen photographed from Sri Lanka has proportionately longer legs and a spiny carapace, not unlike P. coccinea n. sp. and P. ouch n. sp., although on the basis of geography, they are probably different species. For the moment, we do not know their precise identities.

Figures

Fig. 6. General habitus and right first ambulatory legs. A, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; B, Maja crispata, male (48.5 × 39.0 mm) (MNHN-IU-2013-4041), Italy; C, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; D, Neomaja goltziana, male (90.2 × 83.1 mm) (MNHN-IU-2013-4044), Gabon; E, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; F, Neomaja goltziana, male (74.7 × 66.7 mm) (MNHN-IU-2013-4043), Senegal; G, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; H, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines.

Fig. 21. General habitus, Paramaya species. A, P. spinigera (as Maja (Paramaya) spinigera, after De Haan, 1837: pl. 24, fig. 4); B, P. spinigera, dried female (70.1 × 58.8 mm) (KPM NH4195), Japan; C, P. spinigera, male (64.7 × 51.2 mm) (NHM 1961.6.5.117), Japan; D, P. spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; E, P. ouch n. sp., paratype male (39.7 × 29.6 mm) (ZRC 2011.0045), Philippines; F, P. ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines.

Fig. 23. Left G1s, Paramaya species. A–C, P. spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; D–F, P. ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; G–I, P. coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu. Scale bars = 5.0 mm [A, D, G]; 1.0 mm [B, C, E, F, H, I].

Fig. 37. Frontal regions of carapaces. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; C, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; D, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; E, Holthuija miersii, male (22.3 × 16.5 mm) (CBM ZC4001), Singapore; F, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; G, Holthuija pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; H, Holthuija cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; I, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; J, Holthuija poorei n. sp., paratype female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea; K, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; L, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; M, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines; N, Sakaija longispinosa n. sp., holotype ovigerous female (11.4 × 8.6 mm) (NMV J63792), Australia; O, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu.

Fig. 40. Antennae, antennules and epistomes. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; C, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; D, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; E, Holthuija miersii, male (22.3 × 16.5 mm) (CBM ZC4001), Singapore; F, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; G, Holthuija pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; H, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; I, Holthuija poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea; J, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; K, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; L, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines; M, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; N, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; O, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 42. Epistomes. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; C, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; D, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; E, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; F, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; G, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; H, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 45. Third maxillipeds. A, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; B, Alcomaja gracilipes, holotype male (45.4 × 43.5 mm) (IOCAS K33B-34), South China Sea [third max laterally inverted to match]; C, Alcomaja gracilipes, male (41.4 × 35.4 mm) (ZRC 2013.1225), Philippines; D, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; E, Alcomaja nagashimaensis, holotype male (34.0 × 28.5 mm) (USNM 125886), Japan; F, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines; G, Alcomaja latens n. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon Islands; H, Alcomaja miriky n. sp., holotype male (26.4 × 20.9 mm) (MNHN-IU-2010-929), Madagascar; I, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; J, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; K, Paramaya ouch n. sp., paratype male (39.7 × 29.6 mm) (ZRC 2011.0045), Philippines; L, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu.

Fig. 50. Male anterior thoracic sternums and abdomens. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; C, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; D, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; E, Holthuija miersii, male (22.3 × 16.5 mm) (CBM ZC4001), Singapore; F, Holthuija suluensis, male (11.5 × 8.2 mm) (USNM 48507), Philippines; G, Holthuija pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; H, Holthuija cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; I, Holthuija poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea.

Fig. 52. Male sterno-abdominal cavities. A, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; B, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; C, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; D, Paramaja gibba n. sp., male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; E, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; F, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; G, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines; H, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; I, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; J, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; K, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; L, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 54. Chelae. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya ouch n. sp., paratype male (76.5 × 61.3 mm) (ZRC 2001.0577), Philippines; C, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; D, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; E, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; F, Holthuija pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; G, Holthuija cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; H, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; I, Holthuija poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea; J, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; K, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines; L, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; M, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; N, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; O, Ovimaja compressipes, holotype female (51.2 × 40.5 mm) (NHM 1860.15), China; P, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 56. First ambulatory dactyli and propodi. A, Paramaya spinigera, dried female (70.1 × 58.8 mm) (KPM NH4195), Japan; B, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; C, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; D, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; E, Holthuija cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan [figure laterally inverted]; F, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; G, Holthuija poorei, n. sp. paratype female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea; H, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; I, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines [figure laterally inverted]; J, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; K, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; L, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 22. General habitus. A, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; B, “Maja spinigera” (ca. 70 mm carapace length), Sri Lanka, not preserved (photograph: R. Pethiyagoda); C, D, “Maja spinigera” (after Alcock & Anderson, 1898: pl. 34, fig. 3).