Maja crispata Risso, 1827

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 : 126-128

Maja crispata Risso, 1827

( Figs. 3C, D , 6A, B , 7O–Q , 36E , 39D , 41C , 43F , 47G , 52B , 53F , 55C )

Maja crispata Risso, 1827: 23 . – Neumann, 1996b: 828; d’Udekem d’Acoz, 1999: 189.

Maja verrucosa H. Milne Edwards, 1834: 328 , pl. 3 figs. 1–14. – Bouvier, 1940: 322. – Monod, 1956: 477. – Zariquiey Alvarez, 1968: 447, figs. 149b, 150a–f, 156a. – Muraoka, 1998: 27.

Cancer majoides Nardo, 1847: 3 . Maja erinacea Ninni, 1924: 49 , 50, 2 un-numbered figures.

(see Neumann, 1996b: 828; d’Udekem d’Acoz, 1999: 189, for references)

Material examined. Tunisia – 1 female ( NHM), station RBM Tun-303, Sidi Bou Said, Yacht Club, among rocks of jetty, 1 m or less water, coll. R. B. Manning & R. Ingle , 21 February 1974 . — 1 male, 1 female ( USNM 265589 ), station R26 ST. 1, 34°29.5’N 11°14.5’E, 34–36 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 18 July 1973 . — 1 female ( USNM 265651 station), station RBM TUN 184 , algae covered boulders above Posidonia meadows, isolated cove about 2 km west of Korbous town , coll. Tunisian Marine Decapod Project, 13 July 1973 . — 1 male ( USNM 265654 ), station R24 ST. 3, off Kerkenah , 34°25.2’N 11°10.2’E, 37–42 m, coll. Jeddi, 8 July 1973 . — 1 male ( USNM 265655 ), station R12 ST. 10, off Zarat , Gabès Governorate, 33°55.2’N 10°31.8’E, 21 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 28 June 1973 . — 2 females ( USNM 265658 ), station R28 ST. 3, off Kerkenah , 34°36’N 11°31.5’E, 29–31 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 7 July 1973 . — 3 males, 1 female ( USNM 265659 ), station R25 ST. 3, off Kerkenah , 34°28.8’N 11°14.5’E, 40–43 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 9 July 1973 . — 1 female ( USNM 265660 ), station R19 ST. 9, off Djerba, 34°13.5’N 10°40.2’E, 36 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 20 July 1973 . — 1 male ( USNM 265661 ), station R14 ST. 2, off Gabes, 34°30’N 10°21’E, coll. Tunisian Marine Decapod Project, RV El Maghreb, 28 June 1973 . — 2 males, 1 juvenile ( USNM 265662 ), station R27 ST. 6, off Kerkenah , 34°31.5’N 11°30.5’E, 40–41 m, coll. Tunisian Marine Decapod Project, RV El Maghreb, 8 July 1973 . — 1 male ( USNM 265868 ), station RBM TUN 211 D, shallow rocky platform with grass and algae, La Marsa, Public Beach at Marsa Plage, coll. R. B. Manning, 21 August 1973 . Spain – 1 juvenile male ( NHM 1957.8.12.53), Barcelona Habour, coll. R. Zariquiey Alvarez . — 1 juvenile female ( NHM 1956.3.22.200), Cap l’Abeille, Banyuls , coll. I. Gordon. — 1 male ( NHM), Playa Garvet, by Port Bau , coll. I. Gordon. — 1 male ( NHM 1955.2.28.241), Cadaqués Bay , coll. I. Gordon. — 2 females ( NHM 1953.10.28.77–78), Cadaqués Bay, coll. R. Zariquiey Alvarez . — 1 male ( NHM 1965.12.9.164), Vedra Strait, Ibiza , coll. Imperial College, London, 1963 . Portugal – 1 female ( USNM 258341 ), Armacao De Pera, Algarve , south coast of Portugal, coll. 1 November 1974 . France – 1 male ( NHM 1952.3.18.13), Villefranche , inshore waters, coll. Captain Zotton. Italy – 1 female ( NHM 1959.5.26.126), Bay of Naples , coll. I. Gordon. — 1 male, 1 female ( NHM 1959.5.26.124–125), Bay of Naples , coll. Naples Zoological Station. — 3 males, 1 female ( NHM 98.5.7.265/6), Naples , coll. Norman. — 1 male, 2 females ( NHM 1954.11.4.175–177), L’Arenella, near Naples , from fishermen, coll. I. Gordon. — 1 male ( NHM 1978.209 ), Santa Marinella , 42°02’N 11°51’E, Roma, coll. M. Gianinotta. — 1 male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Station Porto Cesareo, Station di Biologia Marina dei Salento, Lecce, no other data . — 1 male (77.1 × 64.3 mm) (MNHN-IU-2013-4040), Messina , coll. M. Vialleton, no date . — 1 male (48.5 × 39.0 mm) (MNHN-IU-2013-4041), Station Porto Cesareo, Costra Neretina Provenance , Lecce, 10–15 m, no other data . — 1 female ( USNM 152285 ), station RBM ITAL12 , west side of Bay of Carini , 15 m, Sicily, coll. R. B. Manning, 16 June 1974 . — 1 carapace ( USNM 152286 ), station RBM ITAL28 , Trapani , rocky flat south of and outside Harbor, Sicily, coll. R. B. Manning, 20 June 1974 . — 1 male ( USNM 152287 ), station RBM ITAL36 , fish market, Syracuse , Sicily, coll. R. B. Manning, 21 June 1974 . — 3 males ( USNM 152288 ), station RBM ITAL36 , fish market, Syracuse , Sicily, coll. R. B. Manning, 21 June 1974 . — 1 ovigerous female ( USNM 205791 ), Mazara Del Vallo , Sicily, coll. R. B. Manning, 2 September 1985 . — Greece – 1 male, 1 female ( NHM 1968.638 ), Chios Islands , coll. G. Potts. — 1 male ( NHM 1968.637 ), Chios Islands , coll. G. Potts. — 1 female ( NHM 96.5.30.3), Smyrna , coll. Y. Holswood. Croatia – 1 male ( ZRC), Rovinj, Figarola , coll. Senckenberg Museum class, don. SMF, 23 August 1989 . — 1 female ( ZRC), Rovinj , coll. Senckenberg Museum class, don. SMF, 20 August 1989 . — 1 male ( ZRC), Rovinj , coll. Senckenberg Museum class, don. SMF, August 1993 . Slovenia – 1 male, 1 female ( NHM 98.5.7.301), Pirano Adriatic, don. Vienna Museum , coll. Norman. Macedonia – 1 female ( NHM 1919.19.4.281), coll. Waterston. Israel – 1 male ( USNM 152564 ), Mikhmoreth , near rocks, 3–4 m, coll. D. Popper, 12 February 1972 .

Diagnosis. Medium-sized species (adult carapace length 60–70 mm). Carapace distinctly pyriform, much longer than broad more rounded in large specimens; dorsal surface gently convex; uniformly covered with granules and short spines except for window-like patch on each side of gastric region which is weakly or not armed ( Fig. 6A, B ). Pseudorostral spines short, straight, strongly diverging ( Figs. 6A, B , 36E ). Supraorbital eave relatively wide, antorbital spine sharp, almost straight or gently curved upwards; intercalated spine short, touching postorbital spine basally; postorbital spine largest, long, sharp; hepatic spine slender, sharp with 2 or 3 small spines or tubercles basally ( Figs. 6A, B , 36E ). Lateral margin of carapace with 3 strong spines and several small sharp granules and short spines ( Fig. 6A, B ). On median row, no long spines discernible, with 4 sharp tubercles: 2 gastric, 1 cardiac, 1 intestinal ( Fig. 6A, B ). Branchial region with 2 spines; posterior carapace margin with 2 distinct spines and one small median one ( Fig. 6A, B ). Basal antennal article very broad with 2 long distal spines, proximal triangular spine passing behind postorbital spine; outer margin with proximal tooth, appressed on suborbital tooth ( Fig. 39D ). Ischium of third maxilliped squarish, slightly longer than broad ( Fig. 43F ). Carpus of cheliped with low granules or low, uneven ridges, otherwise almost smooth ( Fig. 53F ). Ambulatory dactyli covered with relatively long setae ( Fig. 55C ). G1 long, gently curved; distal part dilated, gently curved downwards ( Fig. 7O–Q ).

Remarks. For the classification of this species in Maja s. str., see Remarks under the genus. The identity of Maja erinacea Ninni, 1924 , described from the Adriatic, is dubious. Türkay (in Števčić, 1990) had suggested that it was synonymous with Maja goltziana d’Oliviera, 1889 , but the problem is that the latter species is not known from the Mediterranean Sea. d’Udekem d’Acoz (1999: 189) noted that it is probably not M. goltziana . Ninni’s (1924) un-numbered figures are rather schematic, but the armature of the carapace spines does bear a marked resemblance to M. goltziana . One of the other diagnostic features of M. goltziana is that the ambulatory meri are armed with a prominent distal spine on the dorsal margin. Ninni (1924: 49–50) writes that in M. erinacea , “La base di quasi tutte le spine è ricoperta da peli, così pure il carapace e tutti I paia di piedi ad eccezione del primo paio. Questo è molto più piccolo degli altri, manca della piccola spina rivolta all’ insù e che trovasi invece sul quarto articolo sulla parte superiore di ogni altro paio di piedi” but it is not very clear if he actually says the ambulatory merus has the long sharp spine. In any case, some specimens of M. crispata are known to have relatively longer carapace spines ( Neumann, 1996a: fig. 2G), and M. goltziana is not known from the Mediterranean Sea. As such, we follow Neumann (1996a: 828) in treating Maja erinacea Ninni, 1924 , as a junior synonym of M. crispata , at least for the time being.

Maja crispata is known from Portugal to Mauritania and in the Mediterranean Sea , occurring at depths not deeper than 120 m. The larvae were described by Neumann (1993), with an unusual case of heteromorphosis recorded by Carmona- Suarez (1990) .

Figures

Fig. 3. General habitus. A, B, Maja squinado (after Seba, 1759: pl. 18 figs. 2, 3); C, D, Maja crispata (after Herbst, 1788: pl. 14, figures 84, 85; as Cancer squinado); E, Maja brachydactyla (after Herbst, 1803: pl. 56; as Cancer squinado).

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. 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].

Fig. 36. Frontal regions of carapaces. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; C, Maja brachydactyla, male (161.2 × 140.1 mm) (ZRC 2008.0179), Spain; D, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; E, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; F, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; G, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; H, Paramaja kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; I, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; J, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; K, Paramaja turgida n. sp., paratype male (67.9 × 60.5 mm) (MNHN), Philippines; L, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; M, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines; N, Alcomaja latens n. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon Islands; O, Alcomaja miriky n. sp., holotype male (26.4 × 20.9 mm) (MNHN-IU-2010-929), Madagascar.

Fig. 39. Antennae, antennules and epistomes. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; C, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; D, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; E, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN- IU-2013-4046), Congo; F, Neomaja goltziana, male (90.2 × 83.1 mm) (MNHN-IU-2013-4044), Gabon; G, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; H, Paramaja kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; I, Paramaja gibba, lectotype female (25.1 × 21.2 mm) (NHM 1896.5.14.9), India; J, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; K, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; L, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; M, Alcomaja gracilipes, male (41.4 × 35.4 mm) (ZRC 2013.1225), Philippines; N, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; O, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.

Fig. 41. Epistomes. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; C, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; D, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; E, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; F, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; G, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; H, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.

Fig. 43. Third maxillipeds. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja squinado, female (144.5 × 129.0 mm) (ZRC 2013.1126), Spain; C, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; D, Maja brachydactyla, male (161.2 × 140.1 mm), (ZRC 2008.0179), Spain; E, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; F, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; G, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; H,Neomaja goltziana, male (90.2 × 83.1 mm) (MNHN-IU-2013-4044), Gabon; I, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; J, Paramaja kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; K, Paramaja kominatoensis, dried male (60.4 × 52.5 mm) (KPM NH124171), Japan; L, Paramaja gibba, lectotype female (25.1 × 21.2 mm) (NHM 1896.5.14.9), India; M, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; N, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; O, Paramaja turgida n. sp., paratype male (71.4 × 64.4 mm) (NSMT-Cr 22329), Philippines; P, Paramaja turgida n. sp., paratype male (67.9 × 60.5 mm) (MNHN), Philippines.

Fig. 47. Male anterior thoracic sternums and abdomens. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, male (138.0 × 121.0 mm) (ZRC), Croatia; C, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; D, Maja brachydactyla, male (161.2 × 140.1 mm), (ZRC 2008.0179), Spain; E, Maja cornuta, male (88.8 × 86.8 mm) (NHM 1928.12.1.177) (lectotype of Mamaia queketti Stebbing, 1908), South Africa; F, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; G, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; H, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; I, Neomaja goltziana, male (90.2 × 83.1 mm) (MNHN-IU-2013-4044), Gabon.

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. 53. Chelae. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; C, Maja brachydactyla, male (161.2 × 140.1 mm), (ZRC 2008.0179), Spain; D, Maja cornuta, male (88.8 × 86.8 mm) (NHM 1928.12.1.177–) (lectotype of Mamaia queketti Stebbing, 1908), South Africa; E, Maja cornuta, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa; F, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; G, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; H, Neomaja goltziana, male (90.2 × 83.1 mm) (MNHN-IU-2013-4044), Gabon; I, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; J, Paramaja kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; K, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; L, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; M, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; N, Alcomaja gracilipes, male (41.4 × 35.4 mm) (ZRC 2013.1225), Philippines; O, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; P, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.

Fig. 55. First ambulatory dactyli and propodi. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; C, Maja crispata, male (63.1 × 51.9 mm) (MNHN-IU-2013-4042), Italy; D, Neomaja goltziana, male (73.4 × 65.0 mm) (MNHN-IU-2013-4046), Congo; E, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; F, Paramaja kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; G, Paramaja kominatoensis, dried male (60.4 × 52.5 mm) (KPM NH124171), Japan; H, Paramaja gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; I, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; J, Paramaja turgida n. sp., paratype female (62.6 × 57.2 mm) (ZRC 2013.1241), Philippines; K, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; L, Alcomaja irrorata, male (43.0 × 37.2 mm) (ZRC 2013.1223), Philippines; M, Alcomaja desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines; N, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.