Maja squinado (Herbst, 1788)

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 : 119-122

Maja squinado (Herbst, 1788)

( Figs. 3A, B , 4A, B , 6E , 7A–C , 36A , 39A , 41A , 43A, B , 44 , 47A, B , 53A , 55A )

Cancer squinado Herbst, 1788: 214 (in part, Mediterranean) (not Cancer squinado Herbst, 1788 : pl. 14, figures 84, 85 = Maja crispata Risso, 1827 ).

Maja tuberculata De Haan, 1839 : Pl. F.

Maja squinado – d’Udekem d’Acoz, 1999: 189 – Ng et al., 2008: 117 (list).

[For complete synonymy, see Neumann, 1998: 1669; d’Udekem d’Acoz, 1999: 189]

Material examined. Neotype (here designated): male (147.1 × 126.3 mm) (SMF-4548), Rovinj, Croatia, 20–30 m, coll. 21 August 1989. Croatia – 1 male ( SMF 11088), Rovinj, Croatia, Sorelle, 3–4 m, coll. University of Frankfurt class, August 1981. — 1 male ( SMF 40613), west of Figarola , Rovinj, Croatia, 45°5.532’N 13°36.852’E – 45°5.532’N 13°36.852’E, 28.5-30.5 mm, coll. F.B. Burin, 18 September 2009. — 1 male (138.0 × 121.0 mm) ( ZRC), 50 m, on mud and sand, Rijeka Bay , near Island of Krk , coll. trawler, 2 April 2014. Spain – 1 female ( NHM 1954.11.4.178), Rosas. — 1 female (144.5 × 129.0 mm) ( ZRC 2013.1126 ), ca. 70 km north of Barcelona , 41°40’N 2°47’E, northwestern Mediterranean , ca. 5 m, coll. 1960. Italy – 1 female ( USNM 205743 ), Mazara del Vallo , Mediterranean , 14–85 m, coll. R. B. Manning, 4 September 1985. Gilbraltar – 1 male, 2 females ( USNM 57436 ), Straits of Gilbraltar, coll. March 1923. Tunisia – 2 males ( USNM 265648 ), station R28 ST. 10, off Kerkenah, 34°29.5’N 11°40.2’E, coll. Tunisian Marine Decapod Project, RV Dauphin, 7 July 1973. Israel – 1 male (84.8 × 70.3 mm) ( USNM 152563 ), 31°46.2’N 35°13.8’E, coll. fishermen.

Diagnosis. Large species (adult carapace length in excess of 80–90 mm). Carapace pyriform; dorsal surface strongly convex; uniformly covered with granules and short spines ( Fig. 4A, B ). Pseudorostral spines long, strong, gently diverging in smaller specimens, subparallel in larger adults ( Figs. 4A, B , 36A ). Supraorbital eave relatively narrow, antorbital spine sharp, almost straight to gently curved upwards; intercalated spine stout, short, touching postorbital spine; postorbital spine biggest, long, sharp; hepatic spine sharp with 2 accessory spines at base ( Figs. 4A, B , 36A ). Lateral margin of carapace with 3 spines and several small sharp granules and short spines ( Fig. 4A, B ). On median row 7 strong spines: 3 gastric, 2 subgastric, 1 cardiac, 1 intestinal, with 1 sharp granule between cardiac and intestinal spines; spines relatively lower in large specimens ( Fig. 4A, B ). Branchial region with 2 spines; posterior carapace margin with 2 spines ( Fig. 4A, B ). Basal antennal article very broad with 2 long distal spines; outer margin with proximal spine, appressed on suborbital tooth ( Fig. 39A ). Ischium of third maxilliped rectangular, much longer than broad ( Fig. 43A, B ). Carpus of cheliped with distinct tubercles and granules ( Fig. 53A ). Ambulatory dactyli covered with short setae ( Fig. 55A ). G1 long, almost straight to slightly curved; distal part straight ( Fig. 7A–C ).

Remarks. Maja squinado (Herbst, 1788) is the type species of the genus (ICZN Opinion 511, see Holthuis, 1958). In a detailed morphological and biometric study using extensive material from the Mediterranean and the Atlantic coast of Europe, Neumann (1998) was able to recognise three species in what had been generally confused under Maja squinado : M. squinado (Herbst, 1788) s. str. which is restricted to the Mediterranean, M. brachydactyla Balss, 1922 , which occurs from Ireland to Mauritania (see also Sotelo et al., 2007, 2008, 2009); and M. capensis Ortmann, 1894 (present M. cornuta ), which is found from the Gulf of Guinea to South Africa and just entering southwestern Indian Ocean. He also noted that there is probably overlapping of the distributions of M. brachydactyla and M. cornuta (as M. capensis ) along the Senegal coast but more work will have to be done on this.

The excellent study by Neumann (1998), and the series of molecular studies by Sotelo et al. (2007, 2008, 2009) have helped resolve the identity of M. brachydactyla Balss, 1922 , a large species that has long been regarded as synonymous or merely a subspecies of M. squinado (see Manning & Holthuis, 1981: 307). It is clearly a distinct species and the examination of the present material confirms the observations of Neumann (1998). As it turns out, the commercially more valuable species that occurs from Norway to the Atlantic coasts of Spain that had previously been known as “ Maja squinado ” should now be called M. brachydactyla . Maja squinado s. str. is restricted to the Mediterranean Sea. Even locations immediately outside the Mediterranean Sea such as in Morocco are only known to have M. brachydactyla . Interestingly, it seems that M. brachydactyla s. str. can also be found in the Mediterranean itself. Recently, Abelló et al. (2014) conclusively recorded this species from the Alboran Sea in western Mediterranean; so it appears at least in this part of the ocean, both species may be sympatric.

Maja squinado and M. brachydactyla are close and small specimens are difficult to separate. Large adults, however, can be separated by the shape of their carapaces, with those of M. squinado generally more pyriform and elongate ( Fig. 4A, B ) whereas M. brachydactyla is more rounded ( Fig. 4C–F ). Neumann (1998) uses the shape of the pseudorostral spines as a character – subparallel in M. squinado ( Fig. 4A ) but clearly diverging in M. brachydactyla ( Fig. 4C–F ) and this holds in large specimens, especially males. Smaller specimens of M. squinado , however, have pseudorostral spines that are somewhat intermediate in condition, and certainly cannot be described as subparallel ( Fig. 4B ). As such, both the carapace shape and pseudorostral spine form should be used as characters, with the G1 structure valuable when adult males are available.

There are, however, problems with two names that Neumann (1998) did not treat that have a major bearing on the current nomenclature of Maja s. str.: Cancer cornutus Linnaeus, 1758 , and Maja tuberculata De Haan, 1839 .

The name Cancer cornutus was first used by Linnaeus (1758: 629) who described the species very briefly: “29. C. brachyurus, thorace aculeato: rostro spinis corniformibus barbatis, manibus teretibus. M. L. U. Habitat in Oceano Indico.” Several years later, Linnaeus (1764: 445) elaborated on this short diagnosis: “ Cancer brachyurus, thorace aculeato: rostro spinis corniformibus barbatis, manibus teretibus. Syst. Nat. p. 629· n. 29. Habitat in Mari Indico. Thorax ovatus, adspersus punctis acuto spinosis. Latera utrinque ciliata spinis 6, validis. Disci spinae 3 transversae & una anterior; ommes erectae, reliquis aequales. Spinae 2 minores, erectiusculae, ad basin thoracis. Rostri loco Spinae 2 inter oculos, reliquis majores, erectiusculae, distantes subtus barbatae. Rostrum subtus inter oculos, acutum, reflexum; & duae spinae coadunatae juxta oculos. Cauda inflexa, ovata, articulis 7. linearibus. Brachia teretia, laevia, glabra, nuda. Manus pedibus reliquis minores, basi extrorsum bidentatae.” [crab-like, carapace spinate: rostrum spinate, horn-shaped with whiskers, chelipeds smooth. Inhabits Indian Seas. Carapace ovate, covered with sharp spines. Lateral margins each with six large spines. Three transverse and one anterior spine on carapace, all straight, remainder are same length. Two small straight spines at base of carapace. Rostrum with two intraocular spines, other spines short, arranged closely like whiskers. The rostrum bends below eyes, spines concentrated near eyes. Abdomen rigid, ovate, with seven straight segments. Hands smooth, glabrous, without setae. Chelipeds smaller than other articles, base bidentate. Feet fully hirsute. Digits hirsute.]

The relatively detailed description by Linnaeus (1764) matches that of Maja squinado , M. brachydactyla or M. cornuta . The only unusual character seems to be the comment that rostrum bends below the eyes. In Maja species , the two most obvious frontal structures are the prominent “rostral horns”. As has been discussed earlier, these may not be the true rostral structures (they are referred to here as pseudorostral spines). The actual rostrum is the spine that is between the base of the pseudorostral spines and that is directed sharply downwards. This is almost certainly why Linnaeus (1764) described the rostrum twice. The first time, he described the pseudorostral spines as “rostral intraocular spines”, and the second time, he describes the actual rostrum, noting that it “bends below the eyes”. Some authors have speculated that Linnaeus (1764) was actually saying that the pseudorostrum bends below the eyes, a character that defines Indo-West Pacific genera like Micippa Leach, 1817 . Henri Milne Edwards (1834: 331) lists Linnaeus’ taxon (as “ Cancer cornatus ”) as a species of Micippa and Miers (1885: 11, footnote) suggests that Cancer cornutus Linnaeus, 1758 , may be Micippa thalia (Herbst, 1803) . The provenance given by Linnaeus (1758: 629) – Indian Seas – is surprising as most Maja species are Atlantic. One Maja species , however, does enter the western Indian Ocean in South Africa – M. cornuta . While it is possible the specimen or specimens of Cancer cornutus Linnaeus had on hand may have been mislabelled (and they were actually Atlantic in origin), it is also likely that he had Maja material from South Africa as it was part of the then trade routes.

Ng et al. (2008: 117) listed Cancer cornutus Fabricius, 1787 , as a doubtful synonym of Maja squinado (Herbst, 1788) . Fabricius (1775: 407) actually first used the name “ Cancer cornutus ” with the following comments: “C. thorace aculeato, rostro spinis corniformibus, barbatis, manibus rotundatis. Linn. Syst. Nat. 11. 1047. 46. Mus . Lud. Ulr. 445.” It is clear from this account that he was citing Linnaeus’ (1758) species. In his next publication when he used the name “ Cancer cornutus ”, Fabricius (1787: 325) writes: “C. thorace aculeato, rostro spinis corniformibus barbatis, manibus rotundatis. Habitat et in mari Norwagico.” This time, he makes no mention of Linnaeus. He also reports the species from Norwegian Seas for the first time. In his next mention of Cancer cornutus , however, Fabricius (1798: 356) again refers to Linnaeus, writing: “6. I. thorace aculeato, rostro spinis corniformibus, barbatis, manibus rotundatis. Cancer cornutus Ent. Syst. 2. 462. 80. * Linn. Syst. Nat. 2. 1047. 46. Mus . Lud. Ulr. 445. Habitat in mari mediterraneo, Oceano.” It is thus very clear from these publications that Fabricius (1787) never intended to use the name “ Cancer cornutus ” as a novel name even though he did not specifically mention Linnaeus in that paper.

Herbst (1788: 214–217, pl. 14 figs. 84, 85) chose to describe a new species, Cancer squinado , but also recognised Linnaeus’ taxon, C. cornutus , as separate. He treated Cancer squinado again a few years later with a more detailed description and another figure (Herbst, 1803: 23–27, pl. 56). His detailed descriptions and figures of Cancer squinado suggest he had three different taxa. One form, which was relatively smaller was characterised by the presence of two window-like patches on the carapace. This was the form he first figured (Herbst, 1788: pl. 14 figs. 84, 85) (reproduced here as Fig. 3C, D ) and is clearly referrable to Maja crispata (see also Neumann, 1996a). Herbst’s next figure (1803: pl. 56) (reproduced here as Fig. 3E ) shows a more rounded animal that can easily be identified with Maja brachydactyla . The larger form Herbst (1788, 1803) discusses (depicting M. brachydactyla ) also certainly includes M. squinado as he comments that the species is fished in Napoli (=Naples) in the Mediterranean.

Maja crispata is not only relatively smaller at adult size, it also has two distinct patches on the sides of the gastric region that are not only darker in colour than the other parts of the carapace but are also unarmed. These are evident even in long-preserved specimens. Maja squinado (and M. brachydactyla as well as Neomaja goltziana ) does not have these patches. Charpentier (1829: 160–162) discussed at length what he believed were two forms of Cancer squinado and opted to recognise them as separate species, treating the smaller form (figured in Herbst’s pl. 14 figs. 84 and 85) as “ Inachus cornutus ( Linnaeus, 1758) ”. Since Herbst (1788) also cited the plates in the non-binomial work (and hence not available for nomenclature purposes) of Seba (1759: pl. 18 figs. 2, 3), the latter’s specimen can also be part of his original type series.

The types of Cancer squinado Herbst, 1788 , are no longer extant ( Neumann, 1998). Katushi Sakai (1999: 40) also indicated there are no extant types in the Herbst cabinets in the Museum fur Naturkunde in Berlin; and a recent check of the collection by its curator, Oliver Coleman, and Tan Heok Hui (ZRC) confirmed this. Neumann (1998: 1669) argues that Seba’s figures “clearly show an adult male specimen with squinado -characters (i.e. comparatively long lateral carapace spines and long, parallel rostral horns)”, and as such selected this specimen as the lectotype of Cancer squinado Herbst, 1788 . These figures (reproduced here in Fig. 3A, B ) show a specimen resembling M. squinado in general carapace form, although its rostral spines appear to be more divergent than parallel. In any case, there is no extant specimen as Seba’s material is lost (see Koh & Ng, 2008: 341). A neotype for Cancer squinado Herbst, 1788 , is designated here to stabilise the taxonomy of this species (see later).

So what is Cancer cornutus Linnaeus, 1758 ? From all the available evidence, and assuming the original provenance (Indian Ocean) is correct, it can only be what is today known as Maja capensis Ortmann, 1894 (= Mamaia queketti Stebbing, 1908 ) (type localities of both species Port Elizabeth in South Africa). Recognising these names as synonyms means Linnaeus’ name has priority, and the South African spider crab must hence be known as Maja cornuta ( Linnaeus, 1758) . While the name Cancer cornutus Linnaeus, 1758 , has actually not been used as an available name since 1900, invoking Article 23.9.1 of the Code ( ICZN, 1999: 27) to suppress this name in favour of the younger name, M. capensis is not necessary. This is because the name Maja capensis Ortmann, 1894 , itself is rarely cited, and has been in general use only in recent years since Neumann’s (1998) revision.

Linnaeus’ specimens of this species are no longer extant and studies of his extant brachyuran material in Uppsala University failed to uncover any specimen of Maja (S. H. Tan, pers. comm.) (see also Holm, 1957). Ortmann (1894: 40–41) established the name “ Maja squinado var. capensis ” on the basis of three female specimens from Port Elizabeth in South Africa. No measurements were provided for any specimen and all are syntypes. Ortmann compared this variety with M. squinado and M. crispata (as M. verrucosa ) and indicated that while he believed it to be intermediate between the two species, he was more inclined to think it was closer to the former species. Neumann (1998) did not indicate if he examined the types. Ortmann’s (1894) material is deposited in the Museum of Zoology at Strasbourg (MZS) and two dried specimens were located. To stabilise the taxonomy of the species in question here, we here designate the lectotype female (96.8 × 89.5 mm) of Maja squinado var. capensis Ortmann, 1894 , in the Strasbourg Museum as the neotype of Cancer cornutus Linnaeus, 1758 . As a result of this action, all three names are synonyms, with the oldest name being Maja cornuta ( Linnaeus, 1758) .

The second outstanding taxonomic problem associated with M. squinado s. str. and M. brachydactyl a pertains to the identity of Maja tuberculata De Haan, 1839 . The species was named when De Haan (1839: pl. F) figured two mouthparts of a species he named Maja tuberculata but without any description, indication of provenance or diagnostic characters. The name is nevertheless available under the current zoological code ( ICZN, 1999). Yamaguchi & Baba (1993: 695, fig. 24) figured some dried and mounted mouthparts still extant in the Leiden Museum (RMNH D 43524, Fig. 44A, B ) and suggested they may be the type of M. tuberculata . Herklots (1861: 19) in his study of the material of De Haan in the Leiden Museum had indicated that he had a new species of Maja (as Maia nov. sp.) from “Iles moluques” but he did not formally name it, attribute it to De Haan’s “ M. tuberculata ” or refer to the any material or specimens. Whether these dried mouthparts in RMNH formed the basis of Herklots’ “ Maia nov. sp. ” is not known. There is no other evidence for Yamaguchi & Baba’s (1993) suspicion.

Through the courtesy of Charles Fransen, the first author examined the box that supposedly contains the mouthparts of Maja tuberculata . The mouthparts remain attached and arranged in a row, and on the original background label is written “ Maja tuberculata ”. While this label is old, it is not known if this was written by De Haan or later on by someone else. De Haan (1839: pl. F) figured the various mouthparts of Maja squinado (right first to third maxillipeds, right first and second maxillas and right mandible), and only the last two figures (which depicted a right third maxilliped and right first maxilliped) belonged to Maja tuberculata . The third maxilliped of M. tuberculata figured is smaller than that of M. squinado figured on the same plate (both were drawn life-size), and there are no obvious morphological differences between them. In the extant material supposedly of M. tuberculata , the various mouthparts are present, but there are two left third maxillipeds in the box ( Fig. 44A, B ). One is still attached to the box and labelled as Maja tuberculata , while the other was loose. No right third maxillipeds were present. The two extant left third maxillipeds, however, are of different sizes; the attached one is 11.1 mm (measured from the anterior margin of the merus to the posterior margin of the basis) ( Fig. 44E ), the other 17.3 mm ( Fig. 44F ). They clearly belong to two separate specimens. On the basis of the structure and size, the larger left third maxilliped ( Fig. 44F ) corresponds well with the size of the right third maxilliped figured as Maja squinado by De Haan (1839) ( Fig. 44C ). The smaller left third maxilliped ( Fig. 44E ) agrees in size with the right third maxilliped figured as Maja tuberculata by De Haan (1839) ( Fig. 44C ). On this evidence, it would appear that the smaller left third maxilliped, as well as the associated series of mouthparts, are the types of Maja tuberculata De Haan, 1839 . What happened to the right third maxilliped of Maja tuberculata figured by De Haan (1839) is not known – it is probably lost.

The figure of the now missing right third maxilliped of M. tuberculata by De Haan (1839) is quite accurate ( Fig. 4D ) and agrees with the general structure of the extant left third maxilliped. The third maxilliped of M. tuberculata measures about 11.1 mm long and on the basis of the Maja specimens examined in this study, it would have to belong to a large species measuring about 70–90 mm in carapace length.

Comparing the third maxilliped structure of M. tuberculata ( Fig. 44 ) with species of Maja sensu lato, it is clear that it almost identical in shape, proportions and setation to Maja squinado , M. brachydactyla and M. cornuta (ex M. capensis ) from the eastern Atlantic and South Africa. The third maxillipeds of the other two Atlantic species, Neomaja goltziana (which can grow to 80 mm in carapace length) ( Fig. 43G, H ) and Maja crispata (which can exceed 60 mm in carapace length) ( Fig. 43F ), are very different from that of M. tuberculata ( Fig. 44E ). In the Indo-West Pacific, only Paramaja gibba , Paramaja kominatoensis , Paramaja turgida , Paramaya spinigera and Paramaya ouch are known to grow to 80 mm in carapace length, but their third maxillipeds ( Figs. 43L–P , 45I–K ) are very different in structure compared to that of M. tuberculata . As such, the suggestion by Yamaguchi & Baba (1993) that Maja tuberculata may have originated from the “ Moluccas ” and it is Herklots’ (1861) unnamed species seems unfounded.

The available data strongly suggests that De Haan’s Maja tuberculata was merely based on a small specimen of M. squinado , which also tends to be more prominently granulated on the carapace when younger (see Neumann, 1998). Since the distributions of M. squinado and M. brachydactyla partially overlap ( Neumann, 1998), we cannot be sure which of the Atlantic species M. tuberculata is actually conspecific with. It poses some potential nomenclatural problems if Maja tuberculata De Haan, 1839 , is the same as M. brachydactyla Balss, 1922 , since it is a senior name. Maja cornuta ( Linnaeus, 1758) and M. squinado (Herbst, 1788) are older names. For practical reasons, we regard Maja tuberculata De Haan, 1839 , as a subjective junior synonym of Cancer squinado Herbst, 1788 , at least for the time being.

As the types of Cancer squinado Herbst, 1788 , are lost (K. Sakai, 1999), and even the lectotype selected by Neumann (1998) is no longer extant (see earlier discussion); we here designate a male specimen measuring 147.1 by 126.3 mm (SMF-4548) from Croatia as the neotype of Cancer squinado Herbst, 1788 ( Fig. 4A ). This will stabilise the taxonomy of the genus, the identities of the European species and objectively separate it from the allied M. brachydactyla Balss, 1922 . As discussed earlier, the types of Maja tuberculata De Haan, 1839 , may also have to be suppressed in the future by an application to the International Commission for Zoological Nomenclature.

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. 4. General habitus, Maja species. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja squinado, male (84.8 × 70.3 mm) (USNM 152563), Israel; C, Maja brachydactyla, male (61.5 × 53.8 mm) (MNHN-IU-2013-4039), Morocco; D, Maja brachydactyla, male (161.2 × 140.1 mm), (ZRC 2008.0179), Spain; E, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; F, Maja brachydactyla, ovigerous female (119.0 × 103.0 mm) (ZRC 2009.1130), U.K.

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. 44. Third maxillipeds. Maja squinado. A, B, mouthparts of specimen in Leiden labelled as “Maja tuberculata De Haan, 1839” (RMNH D 43524); C, mouthparts of Maja squinado and Maja tuberculata (after De Haan, 1839: Pl. F); D, right third maxilliped of Maja tuberculata (after De Haan, 1839: Pl. F); E, left third maxilliped ostensibly of Maja tuberculata mounted on card (RMNH D 43524, part); F, free left third maxilliped ostensibly of Maja tuberculata pinned in box (RMNH D 43524, part).

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. 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.

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.