Holthuija suluensis ( Rathbun, 1916 )

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 : 162

Holthuija suluensis ( Rathbun, 1916)

( Figs. 1A , 24E , 26A , 27D, E , 37F , 40F , 42D , 46B , 50F , 54E , 56D )

Maja suluensis Rathbun, 1916: 552 . – Serène, 1968: 57. – Griffin, 1976: 200, Fig. 7a . – Griffin & Tranter, 1986: 210, 218 (part), fig. 71f. – Ng et al., 2008: 117 (list).

Material examined. Holotype: female (32.4 × 41.2 mm) ( USNM 48224 a), station 5165, Observation Island, Tawi Tawi Group, Sulu Islands , 4°58.33’N 119°50.5’E, Philippines, coll. RV Albatross , 24 February 1908 . Paratypes: 2 females (22.5 × 30.6 mm, 22.4 × 30.9 mm) ( USNM 48224 a), same data as lectotype. Others : Philippines – 1 male (11.5 × 8.2 mm), 1 young female (26.8 × 20.8 mm) ( USNM 48507 ), station 5557, Jolo Island and vicinity, 19 fathoms, coll. RV Albatross , 18 September 1909 . — 1 female (36.3 × 27.4 mm) ( USNM 49697 ), station 5163, Tawi Tawi Group, Sulu Archipelago, 28 fathoms, coll. RV Albatross , 24 February 1908 .

Diagnosis. Carapace covered with small granules ( Fig. 24E ). Pseudorostral spines straight, strongly diverging ( Fig. 37F ). Antorbital spine long, curved, directed anteriorly; intercalated spine short, leaving narrow gaps on each side; postorbital spine very long, slightly curved outwards; hepatic spine directed anteriorly, half length of postorbital spine ( Figs. 24E , 37F ). Lateral margin with 3 long spines; 1 branchial spine; median row with 3 long spines: 2 gastric, 1 cardiac; 2 small spines on posterior carapace margin ( Fig. 24E ). Basal antennal article broad with median row of granules, 2 sharp distal spines ( Fig. 40F ). Epistome with 2 granulated lobes on anterior margin ( Fig. 40F ). G1 almost straight, distal part not dilated or curved [juvenile] ( Fig. 27D, E ).

Remarks. Rathbun (1916: 553) provided measurements for only one female specimen (the holotype) but did not mention how many more specimens she had. In the bottle are two other female specimens with the same data and they should be treated as paratypes. Rathbun (1916: 552) stated that the specimen was collected from 9 fathoms depth but the label says 46 fathoms. Griffin (1976: 200) said 9 to 28 fathoms.

The record by Griffin & Tranter (1986: 218) from Sulu Islands and Kei Islands ( Moluccas) almost certainly contains two species. They comment that the specimen they had from Moluccas has only one gastric spine in contrast to the material from Sulu Islands which has two spines (including the types). They suggested that this character may be variable and used other characters to differentiate H. suluensis from H. miersi . While the characters they use are valid, we are more inclined to believe that his Moluccas specimen is actually another species, possibly allied to H. pauli n. sp. (see below). Holthuija suluensis and H. pauli are superficially very close, but the large series of H. pauli on hand tells us that the position and number of gastric spines (two versus one) is very reliable and not variable, even between sexes or specimens very different in size. Otherwise, they are similar in general carapace form.

Serène & Vadon (1981: 119, 124, 128) records “ Maja miersi ” from the Philippines but their material will need to be checked to see if they are really this species or perhaps the allied H. pauli or H. suluensis . Maramura & Kosaka’s (2003: 35) record of “ Maja aff. suluensis ” from Japan will also need to be examined before its identity can be confirmed.

Figures

Fig. 1. Nomenclature of spines and other structures. A, dorsal surface of carapace, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; B, frontal surface of carapace, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines.

Fig. 24. General habitus, Holthuija species. A, Holthuija miersi (as Maia miersii, after Walker, 1887: pl. 6, fig. 1); B, H. miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; C, H. miersii, female (16.5 × 12.2 mm) (ZRC 1995.311), Singapore; D, H. miersii, female (21.4 × 16.7 mm) (NHM 1900.10.22.13), Singapore; E, H. suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; F, H. pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines.

Fig. 26. Lateral views of carapaces, Holthuija species. A, H. suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; B, H. pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; C, H. pauli n. sp., paratype male (35.0 × 27.0 mm) (ZRC 2013.1378), Philippines; D, H. aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; E, H. poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea [laterally inverted to match earlier figures]; F, H. poorei n. sp., paratype female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea laterally inverted to match earlier figures].

Fig. 27. Left G1s, Holthuija species. A–C, H. miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; D, E, H. suluensis, male (11.5 × 8.2 mm) (USNM 48507), Philippines; F–H, H. pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; I–K, H. cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; L–O, H. poorei n. sp., paratype male (26.1 × 20.1 mm) (NMV J63169), Timor Sea. Scale bars = 1.0 mm [A–F, I, L]; 0.5 mm [D, G, H, J, K, M–O]; 0.25 mm [E].

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. 46. Third maxillipeds. A, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; B, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; C, Holthuija pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; D, Holthuija cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; E, Holthuija aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; F, Holthuija poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea; G, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; H, Sakaija sakaii, male (9.0 × 6.6 mm) (NSMT-Cr 8094), Japan; I, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; J, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines; K, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; L, Sakaija longispinosa n. sp., paratype ovigerous female (14.2 × 10.7 mm) (NMV J63197), Australia; M, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; N, Planaja plana n. gen. n. sp., paratype male (38.3 × 30.7 mm) (ZRC 2013.1370), 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. 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. 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. 7. Left G1s. A–C, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia [gonopods laterally inverted as left G1 missing, right G1 drawn]; D–F, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; G–I, Maja brachydactyla, male (161.2 × 140.1 mm) (ZRC 2008.0179), Spain; J–L, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; M, N, Maja cornuta (as Mamaia queketti, after Barnard, 1950: fig. 13a); O–Q, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; R–T, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo. Scale bars = 5.0 mm [A, D, G, J, O, R]; 1.0 mm [B, C, E, F, H, I, K, L, P, Q, S, T].