Holthuija pauli, Ng & Forges, 2015

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 : 162-163

Holthuija pauli n. sp.

( Figs. 24F , 26B, C , 27F–H , 37G , 40G , 46C , 50G , 54F , 69E, F )

Material examined. Holotype: male (37.3 × 28.0 mm) ( NMCR), Balicasag Island , Panglao , Bohol, Philippines, 200– 300 m, coll. fishermen in tangle nets, July 2003 . Paratypes: Philippines – 10 males (27.0 × 19.7 mm, 27.6 × 20.2 mm, 27.9 × 20.3 mm, 30.6 × 22.8 mm 32.4 × 24.0 mm, 33.2 × 24.1 mm, 34.2 × 26.1 mm, 34.4 × 25.3 mm, 34.5 × 25.9 mm, 41.0 × 30.8 mm,) , 3 females (41.9 × 31.4 mm, 41.9 × 32.4 mm, 34.8 × 26.2 mm), 1 ovigerous female (34.7 × 26.4 mm), 1 damaged female (28.0 × 20.8 mm) ( ZRC 2013.1371 ), same data as holotype . — 1 female (27.8 × 21.5 mm) ( ZRC 2001.411 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, December 2000 . — 1 male ( ZRC 2013.1266 , ex part ZRC 2001.0577 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, 28 November 2001 . — 4 males (27.5 × 19.9 mm, 28.2 × 20.7 mm, 31.5 × 23.4 mm, 31.9 × 23.8 mm), 1 ovigerous female (35.0 × 26.0 mm) ( ZRC 2001.0590 ), Balicasag Island , Panglao, Bohol, coll. fishermen with tangle nets, 28 November 2001 . — 1 female (23.8 × 17.6 mm) ( ZRC 2013.1376 ), Balicasag Island, Panglao , Bohol, coll. fishermen in tangle nets, June 2002 . — 1 juvenile female with bopyrid ( ZRC 2013.1314 ), Balicasag Island, Panglao , Bohol, 200–300 m, coll. fishermen with tangle nets, June 2002 . — 1 male (35.0 × 27.0 mm) ( ZRC 2013.1378 ), Balicasag Island, Panglao , Bohol, coll. fishermen in tangle nets, June 2002 . — 1 male, 2 females (1 ovigerous) (NSMT-Cr 15392), Balicasag Island, Panglao , Bohol, coll. local fisherman via M. Takeda & H. Komatsu, February 2003 . — 2 females (33.8 × 25.3 mm, 40.7 × 30.7 [carapace badly damaged]) ( ZRC 2013.1375 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, July 2003 . — 1 damaged male (19.5 × 15.6 mm) ( ZRC 2013.1377 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, July 2003 . — 1 female (38.8 × 28.9 mm) ( ZRC 2013.1379 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, 25–30 July 2003 . — 1 female (35.5 × 27.2 mm) ( ZRC 2013.1380 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, November 2003 . — 2 males ( AM), Balicasag Island, Panglao , Bohol, 200–300 m, coll. fishermen with tangle nets, January 2004 . — 1 male ( ZRC 2013.1324 ), Balicasag Island, Panglao , Bohol, 200–300 m, coll. fishermen with tangle nets, February 2004 . — 1 female (19.9 × 14.7 mm) ( ZRC 2013.1384 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, March 2004 . — 1 male, 2 females ( ZRC 2013.1222 ), Balicasag Island, Panglao , Bohol, 200–300 m, coll. fishermen with tangle nets, April 2004 . — 1 male (29.0 × 21.9 mm) ( ZRC 2013.1373 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, May 2004 . — 4 males (smallest 23.4 × 17.4 mm, largest 42.0 × 31.8 mm), 1 female with rhizocephalan (28.2 × 21.4 mm), 2 females (33.4 × 25.7 mm, 45.9 × 35.0 mm) ( ZRC 2013.1372 ), Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, 29 May 2004 . — 1 female ( ZRC 2013.1374 ), Balicasag Island, Panglao , Bohol, 200–300 m, coll. fishermen with tangle nets, 29 February 2005 . Others: 1 badly damaged male (21.5 × 17.8 mm) ( ZRC 2013.1323 , ex part ZRC 2000.585 ) Balicasag Island, Panglao , Bohol, coll. fishermen with tangle nets, 28 November 2001 .

Diagnosis. Carapace covered with small granules ( Fig. 24F ). Pseudorostral spines long, strongly diverging and usually curved upwards ( Figs. 37G ). Antorbital spine curved; intercalated spine triangular with large gaps; postorbital spine very long; hepatic spine relatively small ( Figs. 24F , 37G ). Lateral margin with 3 long spines, posterior longest; 1 branchial long spine; median row with 2 spines: 1 gastric, 1 cardiac; 2 short spines on posterior carapace margin ( Fig. 24F ). Basal antennal article broad, granulated, with longitudinal furrow, 2 strong distal spines ( Fig. 40G ). Epistome with 2 low tubercles on anterior margin ( Fig. 40G ). G1 almost straight with distal part slightly dilated ( Fig. 27F–H ).

Etymology. The name honors the memory of Paul Beaudemoulin, father-in-law of the second author, who passed away in October 2012.

Remarks. Of the Holthuija species , H. pauli n. sp. is distinct in possessing very long carapace spines which are prominently curved obliquely upwards. The large number of specimens we have examined show that the length and orientation of these spines are not variable, even for small specimens and females. It also has proportionately, the longest ambulatory legs ( Fig. 24F ). In the other species, the ambulatory legs, particularly the merus, are shorter ( Figs. 24A–D , 25 ).

The G1 structure of H. pauli most closely resembles that of H. cognata n. sp., with the distal part laterally flattened, even though their carapace and legs are quite different (carapace relatively broader with longer and more curved spines, and with proportionately longer legs in H. pauli ). The G1 of H. pauli , however, is relatively more slender than that of H. cognata ( Fig. 27F–H versus Fig. 27I–K ).

Holthuija pauli might be confused with small specimens of Paramaya spinigera and Paramaya ouch , especially with regard to the relatively long carapace spines which are curved and the very divergent pseudorostral spines. Paramaya species , however, have long distal spines on the meri of ambulatory legs (absent in H. pauli ) as well as many other characters at the generic level (see discussion for Paramaya ).

In life, H. pauli is beige and light reddish-brown overall, with the third maxillipeds and abdomen brighter in colour ( Fig. 69E, F ).

Figures

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. 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. 69. Colours in life. A, Alcomaja nagashimaensis, male (specimen not examined), Philippines; B, Alcomaja miriky n. sp., paratype male (17.3 × 12.7 mm) (MNHN-IU-2010-60), Madagascar; C, D, Holthuija miersii, ovigerous female (50.2 × 40.4 mm) (ZRC 2013.1272), Singapore; E, Holthuija pauli n. sp., paratype male (31.5 × 23.4 mm) (ZRC 2001.0590), Philippines; F, Holthuija pauli n. sp., female (38.8 × 28.9 mm) (ZRC 2013.1379), Philippines.

Fig. 25. General habitus, Holthuija species. A, H. cognata n. sp., holotype male (29.5 × 23.7 mm) (CBM-ZC3662), Japan; B, H. aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; C, H. aussie n. sp., paratype juvenile female (11.9 × 9.1 mm) (NMV J53958), Australia; D, H. aussie n. sp., juvenile female (14.3 × 11.9 mm) (NMV J61058), Australia; E, H. poorei n. sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea; F, H. poorei n. sp., paratype female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea.