Holthuija, 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 : 160

Holthuija n. gen.

Diagnosis. Carapace ovate; dorsal surface inflated, regions distinct, covered by granules or spines; gastric and branchial regions clearly delimited by grooves ( Figs. 24–26 ). Intestinal region without median spine ( Figs. 24–26 ). Pseudorostral spines relatively long, diverging, forming a V ( Figs. 24 , 25 , 37D–J ). Supraorbital eave large, anterior part strongly expanded, foliaceous, margin convex; spine; antorbital spine long, sharp ( Fig. 37D–J ). Intercalated spine distinct, separated from supraorbital eave and postorbital spine by distinct gaps; postorbital spine strong, lobiform; hepatic region with 1 strong spine, much shorter than postorbital spine; 1 tubercle below ( Fig. 37D–J ). Lateral carapace margin with 3 large spines, 1 branchial spine ( Figs. 24 , 25 ). Posterior carapace margin with 2 distinct median spines ( Figs. 24 , 25 ). Eyes relatively long, slender, with ovoid cornea ( Fig. 37D–J ). Antennal flagellum short, slender. Basal antennal article longer than broad, rectangular; surface with several tubercles, with 2 spines distally; inner and outer lateral margins with low granules; proximal outer angle rounded; antero-external crested rim of antennular fossa overlaps distal part of basal antennal article by about a third of its width ( Fig. 40D–I ). Epistome slightly wider than long, anterior margin with 2 low lobes and granular extension below each; posterior margin composed of 4 rectangular plates separated by shallow fissures ( Figs. 40D–I , 42C–E ). Suborbital margin separated from basal antennal article and margin of postorbital tooth by fissures ( Fig. 40D–I ). Outer surface of third maxilliped covered by short setae; ischium subrectangular, slightly longer than broad; postero-external angle of merus relatively broad, “inserted” into shallower concavity on outer margin of ischium; antero-internal part of ischium rounded, auriculiform ( Fig. 46A–F ). Male chelipeds relatively short in adult males, surfaces of merus and carpus almost smooth; carpus elongate; propodus of palm elongated, not inflated, curved, smooth, palm longer than fingers; fingers long, slender, gently curved, with small basal gape when closed ( Figs. 24 , 25 , 54D–I ). Ambulatory legs relatively short, slender; merus without dorsal subdistal spine; dactylus elongate, curved, covered with long setae except for corneous tip ( Figs. 24 , 25 , 56C–G ). Thoracic sternum wide; surfaces of somites 5–8 with numerous rounded tubercles and granules; sternites 3 and 4 with obliquely longitudinal depressions; margin between sternites 2 and 3 demarcated by deep notch; anterior margin of sterno-abdominal cavity forming complete rim ( Figs. 50D–I , 52I ). Male abdomen subrectangular, with 6 free somites and telson; somites 3 and 4 subequal to or wider than somites 6 and telson ( Fig. 50D–I ). Male press-button abdominal locking mechanism submedian in position on sterno-abdominal cavity ( Fig. 52I ). Female abdomen dome-shaped, covering most of thoracic sternum. G1 long, relatively stout, almost straight or gently curved, distal part with scattered short setae ( Fig. 27 ).

Type species. Maia miersii Walker, 1887 , by present designation.

Etymology. The name is derived from an arbitrary combination of the name Holthuis, with the genus name Maja . This honours the major contribution by the late Lipke B. Holthuis in resolving the convoluted nomenclature of the genus. Gender feminine.

Remarks. Six species are included in Holthuija n. gen. H. miersii ( Walker, 1887) , H. suluensis ( Rathbun, 1916) , H. pauli n. sp., H. cognata n. sp., H. aussie n. sp. and H. poorei n. sp. These species are all easily distinguished by their prominently foliate supraorbital eave, which is the most expanded in all the genera treated here. All species also have a strongly granulated carapace surface, epistome, suborbital and pterygostomial regions, third maxilliped, thoracic sternum and abdomen. The suborbital tooth is also sharply separated from the postorbital tooth by a distinct fissure; and the G1 is generally simple, with a simple distal part and only scattered short setae.

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. 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.

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. 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. 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. 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. 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].