Maja Lamarck, 1801

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 : 113-118

Maja Lamarck, 1801

Maja Lamarck, 1801: 154 .

Maia – Bouvier, 1940: 319, fig. 195 [incorrect spelling of Maja ].

Meria Griffith & Pidgeon, 1833: 165 [erroneous spelling of Maja ].

Mamaia Stebbing, 1905: 23 [substitute name for Maja ; type species Cancer squinado Herbst, 1788 ].

Diagnosis. Carapace rounded in adults; dorsal surface covered by tubercles, short spines; gastric, branchial regions not distinct, separated by shallow grooves ( Figs. 4 , 5 , 6A, B ). Intestinal region with small median spine, not always discernible ( Figs. 4 , 5 , 6A, B ). Pseudorostral spines short, divergent, forming a V ( Figs. 4 , 5 , 6A, B ). Supraorbital eave usually with anterior part longitudinally narrow, rectangular, not prominently expanded; antorbital spine sharp ( Figs. 4 , 5 , 6A, B ). Intercalated spine distinct, separated from supraorbital eave and postorbital spine by narrow gaps; postorbital spine strong; hepatic region with 1 strong spine, much shorter than postorbital spine; 1 smaller spine below ( Figs. 36A–E , 39A–D ). Lateral carapace margin with 3 large spines, numerous spinules, branchial region with 1 low spine or tubercle ( Figs. 4 , 5 , 6A, B ). Posterior carapace margin with 2 short median spines ( Figs. 4 , 5 , 6A, B ). Eyes relatively long, thin, slightly curved, with ovoid cornea ( Figs. 36A–E , 39A–D ). Antennal flagellum short, slender. Basal antennal article as broad as long (quadrate), with 1 long distal spine, 1 blunt median lateral spine; proximal outer angle rounded; inner, outer lateral margins entire; antero-external crested rim of antennular fossa prominently overlapping with subdistal part of basal antennal article as hook-shaped structure by about one-third of its width ( Fig. 39A–D ). Epistome much wider than long, anterior margin with 2 large round tubercles; posterior margin composed of 4 rectangular plates separated by deep fissures ( Figs. 39A–D , 41A–C ). Suborbital margin separated from basal antennal article and margin of postorbital tooth by 2 deep fissures ( Fig. 39A–D ). Outer surface of third maxilliped covered by short setae; ischium squarish to subrectangular; postero-external angle of merus relatively broad, ‘inserting’ into shallow concavity on outer margin of ischium; anterointernal part of ischium rounded, auriculiform ( Figs. 43A–F , 44 ). Male chelipeds long in adult males, surfaces of merus, carpus covered with distinct tubercles, granules; carpus elongate, with low granulated longitudinal ridge; propodus of palm elongated, curved, smooth, without lateral cristae, slightly enlarged, distinctly longer than fingers; fingers long, slender, gently curved, with narrow basal gape when closed ( Figs. 4 , 5 , 6A, B , 53A–F ). Ambulatory legs relatively short, thick; merus without dorsal subdistal spine; dactylus relatively short, curved, usually covered with dense short setae except for corneous distal quarter ( Figs. 4 , 5 , 6A, B, E , 55A–C ). Thoracic sternum wide; surfaces of somites 5–8 almost smooth or with scattered very small granules; sternites 3, 4 slightly depressed; margin between sternites

2, 3 demarcated by small notch; anterior margin of sternoabdominal cavity not forming complete rim ( Figs. 47A–G , 52A, B ). Male abdomen subrectangular, with 6 free somites and telson; somites 3, 4 distinctly wider than somites 5, 6 and telson; telson subtriangular with convex margins ( Fig. 47A–G ). Male press-button abdominal locking mechanism submedian in position on sterno-abdominal cavity ( Fig. 52A, B ). Female abdomen dome-shaped, covering most of thoracic sternum. G1 very long, slender, gently curved throughout length, distal part dilated, subspatuliform, with short but distinct subdistal dorsal process, with scattered very short setae ( Fig. 7A–Q ).

Type species. Cancer squinado Herbst, 1788 , by subsequent designation under the plenary powers of the International Commission on Zoological Nomenclature ( ICZN) in Opinion 511. Gender of genus feminine; name 1260 on Official List .

Remarks. The complex nomenclatural history of Maja Lamarck, 1801 , has been discussed at length by Holthuis (1958) and the ICZN has ruled on the validity of the name as well as its associated type species (ICZN Opinion 511). Mamaia Stebbing, 1905 , is now treated as an unnecessary replacement name. Balss (1957: 1627) spelled the name as “ Mamaja Stebbing” but this must be regarded as erroneous.

According to Ng et al. (2008), Maja Lamarck, 1801 , currently contains 19 species, of which M. brachydactyla Balss, 1922 , M. capensis Ortmann, 1894 (= Mamaia queketti Stebbing, 1908 ), M. crispata Risso, 1827 (= Maia verrucosa H. Milne Edwards, 1834 , Cancer majoides Nardo, 1847 ), M. erinacea Ninni, 1924 , M. goltziana d’Oliveira, 1889 , and M. squinado (Herbst, 1788) (=? Cancer cornutus Fabricius, 1787 ) are known only from the eastern Atlantic, Mediterranean and/ or South Africa. The other 13 species are only from the Indo-West Pacific. Of these, we are of the opinion Maja tuberculata De Haan, 1839 , is probably a junior synonym of M. squinado (Herbst, 1788) (see discussion for the latter species).

The excellent series of Maja specimens and species available for this study allows us to now reappraise the taxonomy of the genus. Maja has traditionally been defined for species with an orbital region that has a distinct supraorbital eave, a prominent intercalated spine and a well developed postorbital spine which has the inner surface excavated to form a depression in which the cornea fits; the base of the antenna is inserted inside the orbital hiatus, and the median row of the dorsal carapace surface has small or large spines (see Griffin & Tranter, 1986).

As to the homogeneity of Maja, Serène & Lohavanijaya (1973: 50) comment that “further observations could demonstrate that the Indo-Pacific species of Maja belong to a different new genus”. There is certainly a substantial degree of variety in the general form of the various Maja species recognised thus far, but to date, no one has found reliable characters to separate them. A recent molecular study focussing on European Maja species by Sotelo et al. (2009) also tested several South African and Asian species ( Maja “ South Africa ”, “ Maja japonica ” [from Singapore], Maja spinigera [from Taiwan] and “ Maja kominatoensis ” [from the Philippines]). The specimens they used from South Africa are here identified as Maja cornuta (ex Maja capensis ); “ Maja japonica ” from Singapore is referred to Holthuija miersii (new genus, see below); while “ Maja kominatoensis ” from Philippines is Alcomaja irrorata n. sp. (new genus, see below). Interestingly, in the Sotelo et al. (2009) study, the Atlantic species of Maja and M. cornuta formed one clade, with Holthuija miersii and Paramaya spinigera forming another, while Alcomaja irrorata aligned with another group further away. While this work is preliminary for the Indo- West Pacific species, it nevertheless supports the revised generic system proposed here.

Of the six Maja species now recognised from the Atlantic, Mediterranean and South Africa, M. erinacea Ninni, 1924 , is here regarded as a synonym of M. crispata (see discussion under the latter species). Of the five remaining species, three – M. squinado , M. brachydactyla and M. cornuta share the following diagnostic characters: the supraorbital eave has the anterior part longitudinally narrow and rectangular in shape ( Fig. 36A–D ); each branchial region only has one spine ( Figs. 4 , 5 ), the suborbital margin is separated from the basal antennal article and margin of postorbital tooth by distinct fissures ( Fig. 39A–C ); the antero-external crested rim of the antennular fossa prominently overlaps the surface of article ( Fig. 39A–C ); the epistome is broader than long ( Figs. 39A–C , 41A, B ), thoracic sternites 3 and 4 are separated by low broad cleft ( Fig. 47A–G ); the carpus of the chelipeds is relatively long, covered with distinct tubercles and granules, with the chelae slender, elongate, not prominently inflated ( Fig. 53A–E ); the merus of the ambulatory legs is unarmed ( Figs. 4 , 5 , 6E ); the dactylus of ambulatory legs is short, stout, with most of the surface covered with short, stiff setae except for the pectinated tip ( Fig. 55A, B ); and the G1 is very long, straight, gently curved, with the distal part slightly expanded and tapering without prominent projections or folds ( Fig. 7A–N ). They are clearly congeneric.

Maja crispata share the characters of the carapace, orbit, antennae, epistome, male thoracic sternum, male abdomen and gonopods as M. squinado , M. brachydactyla and M. cornuta but possesses several unusual characters that make its placement in Maja s. str. more difficult. Its carapace shape is distinctively more pyriform and dorsoventrally flattened ( Fig. 6A, B ) (versus carapaces that are more rounded and swollen, Figs. 4 , 5 ); the presence of a window-like structure on each side of the gastric region which has no or few granules and marks the area below the carapace where the gastric ossicles are anchored ( Fig. 6A, B ) (versus no such structure visible, Figs. 4 , 5 ); the ocular peduncle is proportionately longer and more slender ( Figs. 36E , 39D ) (versus relatively shorter and stouter, Figs. 36A–D , 39A–C ); the supraocular eave is relatively much wider and more foliaceous ( Fig. 39D ) (versus more narrow, Fig. 39A–C ); the ischium of the third maxilliped is squarish ( Fig. 43F ) (versus rectangular, Fig. 43A–E ); the carpus of the cheliped is rugose but not tuberculated or granulated ( Fig. 53F ) (versus prominently tuberculated, Fig. 53A–E ); and the dactylus of the ambulatory legs are covered with long coarse setae ( Fig. 55C ) (versus with short setae, Fig. 55A, B ). However, in view of the very similar male thoracic sternal, male abdominal and gonopodal characters shared by M. squinado , M. brachydactyla , M. cornuta and M. crispata , we prefer to place all four species in Maja s. str. for the time being. This is supported by the genetic study by Sotelo et al. (2009) that showed that M. crispata to be the sister species to M. squinado s. str. Interestingly, both species are from the Mediterranean Sea.

The fifth Atlantic species, M. goltziana , differs significantly from the other four taxa in that the antero-external crested rim of the antennular fossa overlaps further across the surface of the basal antennal article, reaching more than half its width ( Fig. 39E, F ); its epistome is as long as broad ( Figs. 39E, F , 41D ), each branchial region only has two spines ( Fig. 6C, D ), the merus of the ambulatory legs is armed with a strong subdistal dorsal spine ( Fig. 6C, D, F ), the male anterior thoracic sternum is relatively narrower transversely ( Fig. 47H, I ), and the distal part of the G1 has a different structure, with the distal part lacking a subdistal dorsal process ( Fig. 7R–T ). It is here referred to a new genus, Neomaja . The genetic study by Sotelo et al. (2009) also shows this species forming a separate clade from the other European Maja species.

In Indo-West Pacific species that have been referred to Maja , the basal antennal article is distinctly longer than broad, the male thoracic sternites 3 and 4 are separated by a deep V-shaped cleft; the dactylus of the adult ambulatory legs is slender, with scattered long or short setae; the carpus of the chelipeds is short or long with the chelae differently structured (may be inflated); and the G1 is shaped otherwise. With regards to whether the antero-external crested rim of the antennular fossa extends substantially over the basal antennal article, only in a small number of species (here referred to two new genera, Holthuijia and Ovimaja ) have structures similar to that seen in Maja s. str. In all other Indo-West Pacific species, the antero-external crested rim of the antennular fossa does not or just slightly overlaps the basal antennal article.

Even then, these Indo-West Pacific “ Maja ” species can still be easily separated into seven groups, which we here recognise as separate genera: Paramaya De Haan, 1837 , Paramaja Kubo, 1936 , and five new genera – Alcomaja , Holthuija , Sakaija , Planaja and Ovimaja .

One species, Maja bisarmata Rathbun, 1916 , is not a Maja as defined at present. Although Rathbun (1916) had defined Maja for species in which the antennal flagellum is included inside the orbit, for M. bisarmata it is actually just outside the orbit ( Fig. 57G, H ). While the antennal flagellum for M. bisarmata is not prominently excluded from the orbit as in normal Leptomithrax Miers, 1876 ( Fig. 57A–F ), it is also clearly not inside the orbit as in true Maja ( Fig. 2 ). In fact, it is just outside the margin of the orbit, but still different from typical Leptomithrax . This is in fact the same condition as in Leptomithrax kiiensis T. Sakai, 1969 . Tune Sakai (1969, 1976) clearly interprets his species as having the antennal flagellum excluded from the orbit, and hence placed it in Leptomithrax . Maja bisarmata Rathbun, 1916 , is very similar to Leptomithrax kiiensis T. Sakai, 1969 , and both are congeneric. The two species are here referred to a new genus, Rathbunaja , together with two new species from the Philippines and Papua New Guinea, respectively.

Much has been written about the position of the antennal flagellum, one of the key characters separating Maja s. lato from Leptomithrax Miers, 1876 . As noted above, all Maja s. lato species are supposed to have the antennal flagellum inside the orbit ( Fig. 2 ), while that of Leptomithrax species is outside ( Fig. 57 ). However, while this is generally true, it is not always easy to draw a clear line. As discussed, the antennal flagellum of Rathbunaja species is just outside the orbit, like the type species Leptomithrax , L. longimanus ( Miers, 1876) ( Fig. 57A ). In most other species of Leptomithrax , the antennal flagellum is positioned far from the orbit ( Fig. 57B–F ). On the basis of the structure of the antennal flagellum (as well as characters of the third maxillipeds, male abdomen and chelipeds), Leptomithrax , as recognised at present, should eventually be separated into three genera (excluding Rathbunaja ) (see later discussion for the latter genus). In Maja s. str., as well as Neomaja , Paramaja , Holthuija , Planaja and Ovimaja , the antennal flagellum is clearly positioned on the inner surface of the orbit, just before the distal margin of the basal antennal article ( Fig. 2A, B, E, G, H ). In Alcomaja , Paramaya and Sakaija , the antennal flagellum is positioned right at the margin of the basal antennal article ( Fig. 2C, D, F ), and it is not possible to state for certain whether it is just inside or outside the orbit.

The intestinal regions of the carapace of all genera are covered with small granules or tubercles, but only in Paramaya is there a prominent median spine in all species. In Maja , Neomaja , Sakaija and Planaja , the intestinal region has a small spine that may be eroded. In some species in these genera, e.g. Sakaija africana , the intestinal spine is always present and is very strong ( Fig. 29 ). Species of Paramaja , Alcomaja , Holthuija and Ovimaja do not have visible intestinal spines, but this may be because the whole surface is covered with granules of similar sizes. Species of Paramaja , Alcomaja and Planaja also usually have a pair or several pairs of short spines along the median part of the intestinal region.

The supraorbital eave of Holthuija is relatively large and prominently expanded, with the margin of the anterior part foliaceous and the margin convex ( Fig. 37D–J ). In Planaja , the supraorbital eave is expanded ( Fig. 38A ), but not to the degree observed in Holthuija . In all the other genera, the anterior part of the supraorbital eave is relatively narrow longitudinally and more rectangular in general shape ( Figs. 36 , 37A–C, K–O , 38B, C ). In Paramaya , the hepatic spine is slightly shorter, as long as or longer than the postorbital spine ( Fig. 37A–C ). In all other genera, the hepatic spine or tooth is always distinctly smaller than the postorbital spine or tooth ( Figs. 36 , 37D–O , 38 ).

The lateral margin of the carapace (excluding the outer branchial spine) always has three or four spines but they are not always easy to enumerate. In Holthuija and Paramaya , there are always three clearly defined spines since these are large with the granules around them small ( Figs. 21 , 22 , 24 , 25 ). In Maja s. str., the lateral margin has three large spines (e.g., Figs. 4B, C , 5C, D ), but those in between them are also relatively distinct such that in large specimens, it is not always easy to distinguish these from the other spines (e.g., see Fig. 4 D-F). In genera like Paramaja and Alcomaja , the lateral carapace spines are only clearly discernible in juveniles and smaller specimens (e.g., Figs. 10A, B , 11 A-C, 12A), becoming lower and more rounded in adults, so much so that they sometimes cannot be distinguished (e.g., Figs. 9A, B , 10C, D , 11 D-G, 12C-E). Planaja is interesting because it has about six or seven spines along the lateral margin but these are all relatively small and while distinct, these are only slightly larger than the surrounding spinules and sharp granules ( Fig. 34A, B ). Ovimaja is unusual because only the lateral spines are relatively short, and the number varies between 3 and 4, depending on whether you count sharp tubercles as spines ( Fig. 34C, F, G ). In general, the same situation applies for the branchial spines for Ovimaja .

The posterior carapace margin always has two spines, which vary in size. In some genera (see later), the spines can be short and much eroded to the point that they are indiscernible in larger specimens. In Maja , Neomaja , Alcomaja , Planaja and Ovimaja , the spines are short and usually visible ( Figs. 4 , 5 , 6A–D ), although in very large specimens of Paramaja , they are not visible from dorsal view because of their inflated branchial regions (e.g., Figs. 8 , 9 , 10C, D , 11D, F, G , 12D, E ). In some Sakaija species , the spines can become so reduced that they are barely visible in larger specimens ( Fig. 28 A-C). On the other hand, some species of Sakaija (e.g. S. africana and S. longispina n. sp.) always have very long intestinal spines (see Figs. 29 , 32A, B ). In Paramaya and Holthuija , the intestinal spines are consistently long and always distinct even in large specimens, with those of Paramaya extremely long ( Figs. 21 , 22 , 24 , 25 ).

Paramaya is the only genus that has the proximal outer angle of the basal antennal article as a lobe-like projection that has a sharp or acute tip that may be produced into a spine. This is accentuated by the fact that it is clearly separated from the suborbital margin by a broad cleft ( Fig. 40A–C ). In Holthuija , the proximal angle of the basal antennal article is also dilated and lobiform (never spiniform) but it is more closely appressed against the suborbital margin ( Fig. 40D–I ). The antero-external crested rim of the antennular fossa projects onto the basal antennal article to various degrees but there is a distinct pattern. In Maja s. str. and Neomaja , the projection is prominent and extends into and overlaps the article as a prominent hook-like structure ( Fig. 39A–F ). The same is true for Paramaya and Ovimaja although the shapes of their basal articles are different ( Fig. 40A–C, O ). In genera like Paramaja , Alcomaja and Sakaija , the antero-external crested rim of the antennular fossa slightly overlaps the basal antennal article as a simple projection ( Figs. 39G–O , 40J–M ). In Holthuija and Planaja , the antero-external crested rim of the antennular fossa extends deeper into the basal antennal article, but the structure is only gently curved and not hooked ( Fig. 40D–I, N ). The outer edge of the basal antennal article in Paramaya is separated from the suborbital margin by a broad U-shaped sinus ( Fig. 40 A-C). In other genera in which this structure is distinct, it is separated only by a fissure ( Figs. 39A–F , 40D–I, N, O ). At the same time, the suborbital margin is fused and confluent with the margin of postorbital tooth in Paramaja , Alcomaja and Sakaija ( Figs. 39G–O , 40J–M ), but in Maja s. str., Neomaja , Paramaya , Holthuija , Planaja and Ovimaja , they are separated by distinct clefts or fissures ( Figs. 39A–F , 40D–I, N, O ).

The structure of the epistome is distinctive for some genera. Although there is a tendency for the epistome to become slightly broader transversely with increased body size, it generally stays the same shape. In genera like Maja s. str. and Paramaja , the adult epistome is always rectangular and much broader than long ( Figs. 39A–D, G–K , 41A–C, E, F ). In Neomaja and Alcomaja , the epistome is somewhat more quadrangular, being only slightly wider than long ( Figs. 39E, F, L–O , 41D, G, H ). The same is true for Holthuija , Sakaija , Planaja and Ovimaja ( Figs. 40D–O , 42C–H ). Only in Paramaya does the epistome possess a different shape, being distinctly longer than broad ( Figs. 40A–C , 42A, B ). The anterior margin of the epistome is adjacent to the posterior margin of the antennular fossae and it is usually unarmed or smooth. In Maja s. str., Neomaja and Paramaya , there is a large lobiform structure at the outer margin of the anterior epistomal margin ( Figs. 41A–D , 42A, B ). In Ovimaja , this projection is large and tuberculiform ( Fig. 42H ). In Paramaja , Alcomaja , Holthuija , Sakaija and Planaja , the anterior epistomal margin may be cristate or even distinctly raised, but there are no prominent lobes or projections present ( Figs. 41E–H , 42C–G ). In Holthuija , however, there is usually one or a few small tubercles clustered just below the two corners of the anterior margin although the margin itself is not prominently raised ( Fig. 42C–E ). The posterior margin of the epistome in these genera is generally similar, with four plates separated by distinct fissures; the median ones being relatively broader ( Figs. 41 , 42C–H ). However, in Paramaya , the median plates are characteristically more strongly produced downward ( Fig. 42A, B ).

The form of the third maxilliped is distinct in several genera. In Paramaya , the postero-external angle of the merus (which joins the ischium) is very narrow and elongate, and is “inserted” into a deep concavity on the outer margin of the ischium ( Fig. 45I–L ). As a consequence of this, the antero-internal part of the ischium is acutely triangular ( Fig. 45I–L ). In all other genera, the postero-external angle of the merus is proportionately broader and the antero-internal part of the ischium is rounded and auriculiform ( Figs. 43 , 44 , 45A–H , 46 ).

The form of the male anterior thoracic sternum is diagnostic for some genera. In Paramaya , the thoracic sternites 1–4 are elongate, with the lateral junction between thoracic sternites 2 and 3 deep ( Fig. 50A–C ). In Ovimaja , while thoracic sternites 1–4 are not elongated, the lateral junction between sternites 2 and 3 is very deep, forming a distinct waist-like structure and the abdominal margins are concave accordingly ( Fig. 51I ). In Holthuija , the depression on thoracic sternite 4 is not evenly rounded like in the other genera but appears as an oblique depression parallel to the outer margin ( Fig. 50D–I ). The anterior margin of the male sterno-abdominal cavity on thoracic somite 4 of most of the genera is normally slightly raised, with the median part flat ( Figs. 47–49 , 50A–C , 51A–E, I ). Only in Holthuija and Planaja is the entire margin of the male sterno-abdominal cavity on thoracic somite 4 prominently raised and entire, forming a clear rim ( Figs. 50D–I , 51F, G ). In Holthuija , this rim is distinct but is relatively lower ( Fig. 50D–I ). In Planaja , the rim is very pronounced and distinct ( Fig. 51F, G ).

The merus and carpus of the cheliped in Maja s. str. and Neomaja are prominently tuberculate, being covered with prominent short sharp tubercles as well as small granules ( Fig. 53A–H ). In all the other genera, the merus and carpus are lined with small to very small granules, and while some raised areas appear uneven or crenulated, there are never prominent tubercles covering the surface ( Figs. 53I–P , 54 ). The carpus of Maja , Neomaja and Paramaya is proportionately much longer than those in the other genera ( Figs. 4 , 5 , 6A–D , 21 , 22 ), and these proportions do not appear to change much even after considering size and sex.

The structure of the G1 is important in separating groups of genera. In Maja s. str., Neomaja , Paramaja and Paramaya , the G1 is long, slender and gently curved to almost straight long most of its length ( Figs. 7 , 15 , 23 ). In Maja s. str. and Neomaja , the distal part lined only with scattered short spinules or very short setae ( Fig. 7 ); it is covered with scattered spinules and spines in Paramaja ( Fig. 15 ), and devoid of spines, spinules or setae in Paramaya , with a distinct dorsal projection ( Fig. 23 ). In Alcomaja , the G1 is generally gently curved with a distinct subdistal flap or projection on the lower margin, and with prominent spines and short setae or spinules present on the distal surface ( Figs. 19 , 20 ). In Ovimaja , the G1 is very elongated and strongly curved, the distal part has two prominent folds forming a petal-like structure without spines or long setae ( Fig. 35F–L ). In Holthuija , the G1 is relatively stout, curved or straight, but the distal part is slightly to distinctly wider, numerous spines and spinules ( Fig. 27 ). Planaja has an atypical curved G 1 in which the distal part is partially pectinated and lined with strong spines ( Fig. 35A–E ), and in this respect, resembles many species of Leptomithrax s. lato (see Griffin, 1966b). The G1s of Sakaija species are very characteristic, they are relatively slender, curved and the distal part is lined with numerous long setae that are usually weakly or distinctly plumose ( Fig. 33 ). The G1 structures, especially the distal parts, are often curved or gently twisted, and for comparisons between species to be useful, they have to be positioned correctly. In Maja s. str. and Neomaja , for example, the distal dilated part may appear slender from one view (e.g., with the dorsal or sternal surface horizontal with the flat surface ( Fig. 7K ) but appears higher and more dilated when it is tilted slightly ( Fig. 7M ). Interestingly, the G1 structure does not vary substantially even between male specimens substantially different in size (e.g., see Paramaja turgida n. sp. and Planaja plana n. sp., Figs. 15N–U , 35A–E ).

A comment about the names Paramaya De Haan, 1837 , and Paramaja Kubo, 1936 , is necessary. The spellings of the two names are nearly identical and were probably intended to be that way. Why De Haan spelled his name “ Paramaya ” (as a subgenus of Maja ) with a “y” and not a “j” is confusing, but it is clearly intentional. De Haan (1837) used the name Paramaya three times in his book, always as a subgenus (but with different generic arrangements) and with the same Japanese species, spinigera – “ MAJA (MAJA) SPINIGERA , n. sp. T. XXIV. f. 4. ♀. ( Paramaya )” (De Haan, 1837: 93), “ Maja (Paramaya) spinigera , n.” (De Haan, 1837: errata), and “PISA (Paramaya) spinigera n.” (De Haan, 1837: Pl. 24, fig. 4). Maja (Maja) spinigera De Haan, 1837 , is therefore its type species. Some authors (e.g., Manning & Holthuis, 1981: 307; Yamaguchi & Baba, 1993: 359) incorrectly spelled the name as “ Paramaja ” or “ Paramya ” but this must be regarded as an incorrect emendation of Paramaya . Rathbun (1905) commented that the name Paramaya De Haan, 1837 , was available for use. Kubo (1936) later established the name Paramaja for a very different species, Paramaja kominatoensis Kubo, 1936 , also from Japan. Balss (1957: 1628) noted that Paramaja Kubo, 1936 , was a homonym of De Haan’s name and proposed a replacement name, Paramajella . This is not the case. Despite the close similarity of the two names, the current zoological code ( ICZN, 1999) treats them as separate names, the use of “y” and “j” for the two genera deemed sufficient to differentiate them. This is most unfortunate and can certainly give rise to confusion.

The morphological characters used here agree with the molecular analysis by Sotelo et al. (2009) who showed that M. squinado , M. brachydactyla , M. crispata and M. cornuta (as Maja SA 1 and Maja sp. SA2) belonged to one distinct clade. The sister clade to this is Maja goltziana , which is here transferred to its own genus, Neomaja as it has a suite of distinct morphological characters (see later).

In summary, Maja Lamarck, 1801 , is now restricted to four primarily Atlantic and Mediterranean species. The other species previously placed in the genus are transferred to nine other genera, seven of which are new:

Maja Lamarck, 1801 – Maja squinado (Herbst, 1788) , Maja brachydactyla Balss, 1922 , Maja cornuta ( Linnaeus, 1758) , Maja crispata ( Risso, 1827)

Neomaja n. gen. – Neomaja goltziana (d’Oliviera, 1889)

Paramaja Kubo, 1936 – Paramaja kominatoensis Kubo, 1936 , Paramaja gibba ( Alcock, 1895) , Paramaja turgida n. sp.

Paramaya De Haan, 1837 – Paramaya spinigera (De Haan, 1837) , Paramaya ouch n. sp., Paramaya coccinea n. sp.

Alcomaja n. gen. – Alcomaja irrorata n. sp., Alcomaja confragosa ( Griffin & Tranter, 1986) , Alcomaja gracilipes ( Chen & Ng, 1999) , Alcomaja nagashimaensis ( Sakai, 1969) , Alcomaja latens n. sp., Alcomaja miriky n. sp., Alcomaja desmondi n. sp.

Holthuija n. gen. – Holthuija miersii ( Walker, 1887) , Holthuija suluensis ( Rathbun, 1916) , Holthuija cognata n. sp., Holthuija aussie n. sp., Holthuija pauli n. sp., Holthuija poorei n. sp.

Sakaija n. gen. – Sakaija japonica ( Rathbun, 1932) , Sakaija sakaii ( Takeda & Miyake, 1969) , Sakaija serenei n. sp., Sakaija santo n. sp., Sakaija africana ( Griffin & Tranter, 1986) ; Sakaija longispinosa n. sp.

Planaja n. gen. – Planaja plana n. sp.

Ovimaja n. gen. – Ovimaja compressipes ( Miers, 1879)

Rathbunaja n. gen. – Rathbunaja bisarmata ( Rathbun, 1916) , Rathbunaja kiiensis (T. Sakai, 1969) , Rathbunaja brevipes n. sp., Rathbunaja ursus n. sp.

Figures

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. 5. General habitus, Maja cornuta. A, dried female (96.8 × 89.5 mm) (ZMS) (lectotype of Maja squinado var. capensis Ortmann, 1894, and neotype of Cancer cornutus Linnaeus, 1758), South Africa; B, dried female (71.0 × 60.3 mm) (ZMS) (paralectotype of Maja squinado var. capensis Ortmann, 1894), South Africa; C, male (88.8 × 86.8 mm) (NHM 1928.12.1.177) (lectotype of Mamaia queketti Stebbing, 1908), South Africa; D, male (115.3 × 103.4 mm) (ZRC 2013.1184), South Africa.

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. 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. 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. 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. 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. 57. Basal antennal articles and position of flagellum. A, Leptomithrax longimanus, holotype male (38.0 × 36.2 mm) (NHM 1852.12), New Zealand; B, Leptomithrax tuberculatus Whitelegge, 1900, female (36.6 × 28.7 mm) (ZRC 1965.10.13.77), Australia; C, Leptomithrax sternocostulatus (H. Milne Edwards, 1851), male (35.1 × 27.0 mm) (ZRC 1965.10.13.74), Australia; D, Leptomithrax edwardsii, male (75.8 × 69.4 mm) (ZRC 2001.56), Taiwan; E, Leptomithrax bifidus, male (32.5 × 27.5 mm) (NHM 1961.11.13.28), Japan; F, Leptomithrax longipes (G. M. Thomson, 1902), male (55.4 × 39.3 mm) (AM P86814), Australia [by S.T. Ahyong]; G, Rathbunaja bisarmata, holotype male (20.3 × 15.6 mm) (USNM 48220), Philippines; H, Rathbunaja bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines.

Fig. 2. Basal antennal article and position of flagellum.A, Maja brachydactyla, male (98.4 × 89.0 mm) (ZRC 2009.1130), U.K.; B, Paramaja kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; C, Alcomaja irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; D, Paramaya ouch n. sp., paratype ovigerous female (68.3 × 54.2 mm, with rhizocephala) (ZRC 2013.1317); E, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497), Singapore; F, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; 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. 29. General habitus, Sakaija africana. A, as Maja africana (after Griffin & Tranter, 1986: pl. 14a); B, male (27.3 × 21.2 mm) (MNHN-IU-2010-504), Madagascar; C, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; D, ovigerous female (23.8 × 18.4 mm) (MNHN-IU-2010-505), Madagascar.

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. 38. Frontal regions of carapaces. A, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; B, Ovimaja compressipes, holotype female (51.2 × 40.5 mm) (NHM 1860.15), China; C, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

Fig. 21. General habitus, Paramaya species. A, P. spinigera (as Maja (Paramaya) spinigera, after De Haan, 1837: pl. 24, fig. 4); B, P. spinigera, dried female (70.1 × 58.8 mm) (KPM NH4195), Japan; C, P. spinigera, male (64.7 × 51.2 mm) (NHM 1961.6.5.117), Japan; D, P. spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; E, P. ouch n. sp., paratype male (39.7 × 29.6 mm) (ZRC 2011.0045), Philippines; F, P. ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines.

Fig. 22. General habitus. A, Paramaya coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; B, “Maja spinigera” (ca. 70 mm carapace length), Sri Lanka, not preserved (photograph: R. Pethiyagoda); C, D, “Maja spinigera” (after Alcock & Anderson, 1898: pl. 34, fig. 3).

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. 10. General habitus, Paramaja gibba. A, lectotype female (25.1 × 21.2 mm) (NHM 1896.5.14.9), India; B, as Maja gibba (after Alcock & Anderson, 1898: pl. 21 fig. 5); C, male (65.0 × 59.8 mm) (NSMT-Cr 16618), Andaman Sea; D, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal.

Fig. 11. General habitus, Paramaja gibba. A, juvenile male (10.6 × 8.1 mm) (MNHN-IU-2010-54), Madagascar; B, male juvenile (16.4 × 12.5 mm) (MNHN-IU-2010-55), Madagascar; C, male (24.6 × 20.9 mm) (MNHN-IU-2010-61), Madagascar; D, male (58.6 × 52.2 mm) (MNHN-IU-2010-63), Madagascar; E, juvenile female (33.9 × 29.4 mm) (NMV J54316), Australia; F, 1 ovigerous female (45.1 × 40.6 mm) (MNHN-IU-2010-467), Madagascar; G, female (46.7 × 41.6 mm) (NMV J61055), Australia.

Fig. 9. General habitus, Paramaja kominatoensis. A, dried male (60.4 × 52.5 mm) (KPM NH124171), Japan; B, dried female (61.0 × 54.6 mm) (KPM NH124171), Japan; C, male (73.0 × 66.0 mm) (PCM), Taiwan; D, male (53.1 × 46.8 mm) (SMF 47728), Japan; E, male (50.5 × 44.6 mm) (SMF 47738), Japan.

Fig. 34. General habitus. A, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; B, Planaja plana n. gen. n. sp., paratype male (38.3 × 30.7 mm) (ZRC 2013.1370), Philippines; C–E, Ovimaja compressipes, holotype female (51.2 × 40.5 mm) (NHM 1860.15), China; F, Ovimaja compressipes (type of Maja linapacanensis Rathbun, 1916, 27.0 × 20.0 mm, USNM 48225), Philippines; G, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan. A–C, F, G, overall dorsal views; D, antennae, antennules, orbit and epistome; E, thoracic sternum.

Fig. 8. General habitus, Paramaja kominatoensis. A, holotype male (63.0 × 57.0 mm) (after Kubo, 1936: fig. 2); B, neotype male (56.6 × 40.4 mm) (SMF 47808), Japan; C, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; D, male (76.9 × 69.6 mm) (SMF 47738), Japan; E, female (52.7 × 47.5 mm) (SMF 47736), Japan; F, female (62.0 × 58.0 mm) (SMF 47810), Japan.

Fig. 12. General habitus, Paramaja turgida n. sp. A, paratype male (26.2 × 21.3 mm) (ZRC 2001.0616b); B, paratype female (33.5 × 29.0 mm) (ZRC 2013.1230); C, paratype male (67.9 × 60.5 mm) (MNHN), Philippines; D, paratype female (62.6 × 57.2 mm) (ZRC 2013.1241), Philippines; E, holotype male (74.1 × 66.8 mm) (NMCR), Philippines; F, male (25.1 × 19.6 mm) (MNHN-IU-2013-4051), Solomon Islands.

Fig. 28. General habitus, Sakaija species. A, S. japonica, holotype male (16.2 × 13.0 mm) (USNM 48252), Japan; B, S. japonica male (22.6 × 17.9 mm) (ZRC 2013.1269), Tashi fish port, Ilan County, northern Taiwan; C, S. japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; D, S. sakaii (as Maja sakaii, after T. Sakai 1976: pl. 85 fig. 2); E, S. sakaii (as Maja sakaii, after T. Sakai, 1976: text-fig. 125a); F, S. sakaii (as Maja sp., after T. Sakai, 1932: text-fig. 5); G, S. sakaii, male (9.0 × 6.6 mm) (NSMT-Cr 8094), Japan.

Fig. 32. General habitus, Sakaija species. A, S. longispinosa n. sp., holotype ovigerous female (11.4 × 8.6 mm) (NMV J63792), Australia; B, S. longispinosa n. sp., paratype ovigerous female (14.2 × 10.7 mm) (NMV J63197), Australia; C, S. santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; D, S. santo n. sp., paratype female (13.1 × 10.4 mm) (ZRC 2013.1245), 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. 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.

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. 51. Anterior thoracic sternums and abdomens. A, Sakaija japonica, holotype male (16.2 × 13.0 mm) (USNM 48252), Japan; B, Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; C, Sakaija africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; D, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR), Philippines; E, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; F, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; G, Planaja plana n. gen. n. sp., paratype male (38.3 × 30.7 mm) (ZRC 2013.1370), Philippines; H, Ovimaja compressipes, holotype female (51.2 × 40.5 mm) (NHM 1860.15), China; I, 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. 15. Left G1s, Paramaja species. A, P. kominatoensis (after Kubo, 1936: fig. 1B); B-E, P. kominatoensis, male (76.9 × 69.6 mm) (SMF), Japan; F-J, P. kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; K-M, P. gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; N-R, P. turgida n. sp., paratype male (67.9 × 60.5 mm) (MNHN), Philippines; S-U, P. turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines. Scales: B, C, F, G, K, N, O, S = 5.0 mm; D, E, H-J, L, M, P-R, T, U = 1.0 mm.

Fig. 23. Left G1s, Paramaya species. A–C, P. spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; D–F, P. ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines; G–I, P. coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu. Scale bars = 5.0 mm [A, D, G]; 1.0 mm [B, C, E, F, H, I].

Fig. 19. Left G1s, Alcomaja species. A–C, A. irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; D–F, A. gracilipes, holotype male (45.4 × 43.5 mm) (IOCAS K33B-34), South China Sea; G, A. confragosa (as Maja confragosa, after Griffin & Tranter, 1986: fig. 73a). Scale bars = 1.0 mm [A–C].

Fig. 20. Left G1s, Alcomaja species. A, B, A. nagashimaensis (as Maja nagashimaensis, after T. Sakai, 1976: text-fig. 129); C, D, A. nagashimaensis, holotype male (34.0 × 28.5 mm) (USNM 125886), Japan; E–G, A. nagashimaensis, male (37.2 × 32.4 mm) (CBM- ZC9928), Japan; H–J, A. nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines; K–N, A. latens n. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon Islands; O–Q, A. miriky n. sp., holotype male (26.4 × 20.9 mm) (MNHN-IU-2010-929), Madagascar; R–T, A. desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines. Scale bars = 1.0 mm [H, K–R]; 2.0 mm [E]; 0.5 mm [C, F, G, I, J, S, T]; 0.25 mm [D].

Fig. 35. Left G1s. A–C, Planaja plana n. gen. n. sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; D, Planaja plana n. gen. n. sp., paratype male (38.3 × 30.7 mm) (ZRC 2013.1370), Philippines; E, Planaja plana n. gen. n. sp., paratype male (42.8 × 34.3 mm) (AM), Philippines; F–H, Ovimaja compressipes (as Maja brevispinosa, after Dai, 1981: pl. 1 figs. 8, 9); I–L, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan. Scale bars = 1.0 mm [A, I]; 2.0 mm [F]; 0.5 mm [B–E, J, K]; 0.2 mm [G, H].

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. 33. Left G1s, Sakaija species. A–C, S. japonica, holotype male (16.2 × 13.0 mm) (USNM 48252), Japan; D–F, S. japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; G, S. japonica (as Maja japonica, after T. Sakai, 1976: text-fig. 126a); H, S. sakaii (as Maja sakaii, after Sakai, 1976: text-fig. 125b); I, J, S. sakaii, male (9.0 × 6.6 mm) (NSMT-Cr 8094), Japan; K–M, S. serenei n. sp., paratype male (14.3 × 10.9 mm) (ZRC 2013.1255), Philippines; N, O, S. africana (as Maja africana, after Griffin & Tranter, 1986: fig. 73c, d); P–R, S. africana, male (32.2 × 25.4 mm) (MNHN-IU-2010-928), Madagascar; S–U, S. santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu. Scale bars = 1.0 mm [D, K, P–R]; 0.5 mm [A, E, F, I, J, L, M, S–U]; 0.25 mm [B, C].