Ng & Richer de Forges: Revision of the spider crab genus Maja
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 63: 110–225

Date of publication: 29 May 2015
http://zoobank.org/urn:lsid:zoobank.org:pub:40BCDD62-D35E-46D1-95A3-2CC0DF219DEE

Revision of the spider crab genus Maja Lamarck, 1801 (Crustacea:
Brachyura: Majoidea: Majidae), with descriptions of seven new generaand 17 new species from the Atlantic and Indo-West PacificPeter K. L. Ng1* and Bertrand Richer De Forges2
Abstract. The taxonomy of spider crabs of the genus Maja Lamarck, 1801, is revised, and a total of 36 species
in 10 genera are now recognised from the eastern Atlantic, Mediterranean and Indo-West Pacific. The present
revision describes seven genera and 17 species as new. Two genera previously synonymised under Maja: Paramaya
De Haan, 1837, and Paramaja Kubo, 1936, are here treated as valid taxa. The confused nomenclature of Cancer
cornutus Linnaeus, 1758, is resolved, and the name replaces Maja capensis Ortmann, 1894, and Mamaia queketti
Stebbing, 1908. All genera and species are diagnosed and figured, and keys are provided for their identification.
Key words. Crustacea, Brachyura, Maja, taxonomy, revision, new genera, new species, Indo-West Pacific, Atlantic

INTRODUCTION

problematic because they were only briefly described without
any figures. Griffin (1976) also treated them in passing but
without clear figures or photographs of the types. Overall,
the state of taxonomy of Maja remains unsatisfactory.

Spider crabs of the genus Maja Lamarck, 1801 (Majoidea:
Majidae Samouelle, 1819) are common in the eastern Atlantic,
Mediterranean and Indo-West Pacific, occurring in shallow
as well as deep waters. Some large species from the Atlantic,
e.g., Maja squinado (Herbst, 1788) and M. brachydactyla
Balss, 1922, are valuable as fishery resources and have long
been exploited. However, the majority of species occur in
the Indo-Pacific and most are small to medium-sized, noneof which have commercial value. Ng et al. (2008) listed 19
species of which the identities of two were doubtful. The
Atlantic species have been reappraised and revised in recent
years (d’Udekem d’Acoz, 1999; Neumann, 1998) and theiridentities have been clarified. With regard to the Indo-West
Pacific taxa, Serène & Lohavanijaya (1973), T. Sakai (1976)
and Griffin & Tranter (1986) treated various groups of species
and reviewed their taxonomy, providing keys to most or all
the species. However, these keys are inaccurate in parts,
incomplete and/or difficult to use. Several species have been
described from juvenile specimens (e.g., Maja gibba Alcock,1895), and because it was generally believed that carapace
spinulation varies with size and geography (e.g., see Griffin,
1976; Griffin & Tranter, 1986), the identities of several taxa
remain unclear. The Philippine species named by Rathbun(1916) — M. suluensis Rathbun, 1916, M. linapacanensis
Rathbun, 1916, and M. bisarmata Rathbun, 1916, areOver the last decade, the authors have obtained numerous
Maja specimens from many parts of the Indo-West Pacific.
This includes a major collection of crabs collected from tangle
nets around the island of Balicasag in the Philippines (seeNg et al., 2009; Mendoza et al., 2010). Study of this material
in particular has allowed us to resolve many outstanding
taxonomic problems with Maja.
We can now separate the recognised species of Maja into
10 distinct genera, seven of which are new. In addition,
we describe 17 new species from the South China Sea,
Philippines, Taiwan, Japan, Australia, Papua New Guinea,
New Caledonia, Vanuatu, Solomon Islands and Madagascar.
MATERIAL AND METHODSThe terminology used here essentially follows that used by
Griffin (1966a) (see also Griffin & Tranter, 1986) (Fig. 1).
Some additional comments, however, are necessary. The
postorbital spine, as defined by Griffin (1966a) in Maja isactually one of two major spines on the hepatic region andthe anterior one is here regarded as the actual postorbital
spine. Otherwise we follow Griffin’s (1966a) system here sothat appropriate comparisons can be made when other generain the Majidae are considered. There is always a marked
constriction between the hepatic region and the rest of thecarapace, which also marks the start of the convex lateral
margin. The lateral margin is always armed with three orfour distinct spines, which may be reduced; and there aresometimes smaller ones interspersed between them. The
lateralmost spine is usually the largest. There is usually one

Lee Kong Chian Natural History Museum, National University of Singapore, Lower
Kent Ridge Road, Singapore 119260; Email: dbsngkl@nus.edu.sg (*corresponding
author)1

5, rue Félix Franchette, 98800 Nouméa, New Caledonia; Email: b.richerdeforges@
gmail.com
2© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Fig. 1. Nomenclature of spines and other structures. A, dorsal surface of carapace, Holthuija suluensis, holotype female (32.4 × 41.2 mm)(USNM 48224a), Philippines; B, frontal surface of carapace, Paramaya ouch n. sp., holotype male (76.8 × 60.0 mm) (NMCR), Philippines.

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Ng & Richer de Forges: Revision of the spider crab genus Maja
more following spine that is clearly part of the lateral series,
but is positioned higher on the branchial region. Herein werefer to this as the branchial spine. The gastric and cardiac
regions are usually armed with at least one spine of varying
size, although in some species, it is only apparent as a tubercle
or large granule in adult specimens. The intestinal region
may or may not be armed. The posterior carapace marginis usually lined with granules but in many species, there
is a pair of relatively longer and more prominent medianspines. The presence of a spine or tubercle on the various
regions sometimes poses problems because it is not always
discernible. In some species, because the spine is variablein strength (especially with size), it may be reduced and/or
eroded to be just a granule, and cannot be separated fromthe rest of the granules on the surface.

median part of the thoracic sternum. The ventral surfaceof the G1s figured is the view of the in situ structure, i.e.
the abdominal view. The dorsal surface of the G1 refers to
the thoracic sternal view. The structure of the male second
gonopod (G2) has not been useful in the taxonomy of thevarious taxa treated here. All of them have a relatively
short G2 that is between 0.15 and 0.20 times the length of
the G1, with the distal part subspatuliform to cup-like, andnever with a distinct elongated distal segment.
The measurements provided, in millimetres, are of the
post-pseudorostral carapace length (from the base of spines
to the posterior carapace margin, not including medianposterior spines) against the maximum caparace width(at base of spines), respectively. Precise measurements ofthese dimensions can be problematic in many species as theposterior carapace margin and widest points at the base ofspines are often unevenly granular, with the bases of large
spines gently sloping, not clearly demarcated and covered
with different sized granules. In addition, the base of thepseudorostral spines is also not always distinct as the cleft
between them can be deep. Detailed diagnoses are presented
for all genera so comparisons can be made. For species,
only diagnostic characters useful (or potentially useful) to
separate congeners are presented.

There is some subjectivity with regard to what are heretermed spines or teeth, and tubercles or granules. Griffin(1966a) and his subsequent papers generally use these terms
relatively interchangeably; with spines being any major
projection from the carapace margin surface; and granules
for any low small structures present. In brachyurans, theterm spine is normally used when the cross-section of anelongated structure is oval or round, and the term tooth is often
applied when the structure is distinctly laterally flattened.
In majoids, this is not always easy to distinguish. As such,
we will generally use the term spine in the descriptions
and characterize it with additional terms as necessary. The
distinction between an elongated tubercle and low spine is
not always clear; and a dorsolaterally flattened spine thatis lobiform can just as well be called a tooth. We therefore
use the term “spine” whenever possible for any elongatedstructure that is longer than broad.

Material examined is deposited in the Kanagawa Prefectural
Museum of Natural History (KPM), Iriuda, Japan; National
Museum of Nature and Science (NSMT), Tsukuba, Tokyo,
Japan; Muséum national d’Histoire naturelle (MNHN), Paris,
France; The Natural History Museum (NHM), London, U.K.;
The Naturalis (ex Rijksmuseum van Natuurlijke Historie,
RMNH), Leiden, The Netherlands; Senckenberg Museum
Forschunginstitut (SMF), Frankfurt, Germany; Museumof Zoology, Strasbourg (MZS), France; Shirshov Instituteof Oceanography (SIO), Russian Academy of Sciences,
Moscow, Russia; Crustacean Collection of the Philippine
National Museum (NMCR), Manila, Philippines; National
Institute of Water & Atmospheric Research (NIWA),
Wellington, New Zealand; U.S. National Museum of Natural
History (USNM), Smithsonian Institution, Washington D.C.,
U.S.A.; and Australian Museum (AM), Sydney, Australia;
Museum Victoria (NMV), Melbourne, Australia; Instituteof Oceanology, Chinese Academy of Science (IOCAS),
Qingdao, China; Peikuan Crab Museum (PCM), Tashi,
Taiwan; and the Zoological Reference Collection (ZRC),
Lee Kong Chian Natural History Museum (formerly Raffles
Museum of Biodiversity Research), National University ofSingapore.

The term “basal antennal article” is used for the largest
antennal article observed. As discussed by Ng et al. (2008:11), the “basal antennal article” is actually formed by fusionof the true articles 2 and 3, and is not to be confused withthe first article which has the osmoregulatory green gland.
Although it can be somewhat misleading, it has been so
extensively used that to change terminology now would
serve no useful purpose. As such, this term is used here
for convenience.
The male anterior thoracic sternum refers to sternites 1–4.
Sternites 1 and 2 are always fused and separated fromsternite 3 by a groove. The relative width of sternites 3 and
4 are useful characters to separate some genera and species.
The male sterno-abdominal cavity refers to the medianlongitudinal depression in which the abdomen sits. The
surfaces surrounding the sterno-abdominal cavity (notably
the anterior part on sternite 4) can be armed or structured indifferent ways and can be used to separate genera and species.

Much of the examined material is derived from the expeditions
to the western Pacific organised by MNHN with colleagues
from the area. The names of these expeditions are written
in capitals, sometimes with a year after it if there were aseries of collections from one area (e.g., PANGLAO 2004).
When cruises are named after the research ships used, thename of the vessel is italicised (e.g., Albatross).

The dorsal margin of the articles of the ambulatory leg refers
to the extensor or anterior margin. The male first gonopod(G1) in the species revised here are all more or less straightor curve outwards, i.e. they are directed away from the112

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TAXONOMY

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 mostof thoracic sternum. G1 very long, slender, gently curved
throughout length, distal part dilated, subspatuliform, withshort but distinct subdistal dorsal process, with scatteredvery short setae (Fig. 7A–Q).

Family Majidae Samouelle, 1819
Subfamily Majinae Samouelle, 1819
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].

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.

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 fromsupraorbital 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 3large 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 1long 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 composedof 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; propodusof 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 shortsetae 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

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 nameas 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 synonymof 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, aprominent intercalated spine and a well developed postorbital
spine which has the inner surface excavated to form adepression in which the cornea fits; the base of the antenna
is inserted inside the orbital hiatus, and the median row ofthe dorsal carapace surface has small or large spines (seeGriffin & 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 foundreliable 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 (Maja113

Ng & Richer de Forges: Revision of the spider crab genus Maja

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.

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“South Africa”, “Maja japonica” [from Singapore], Maja
spinigera [from Taiwan] and “Maja kominatoensis” [from thePhilippines]). The specimens they used from South Africa arehere identified as Maja cornuta (ex Maja capensis); “Majajaponica” 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, theAtlantic species of Maja and M. cornuta formed one clade,with Holthuija miersii and Paramaya spinigera forming
another, while Alcomaja irrorata aligned with another groupfurther away. While this work is preliminary for the IndoWest Pacific species, it nevertheless supports the revised
generic system proposed here.

with long coarse setae (Fig. 55C) (versus with short setae,
Fig. 55A, B). However, in view of the very similar malethoracic sternal, male abdominal and gonopodal charactersshared by M. squinado, M. brachydactyla, M. cornuta andM. 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 thesurface of the basal antennal article, reaching more thanhalf 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 isarmed with a strong subdistal dorsal spine (Fig. 6C, D, F),
the male anterior thoracic sternum is relatively narrowertransversely (Fig. 47H, I), and the distal part of the G1 hasa 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 theother European Maja species.

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 thefollowing diagnostic characters: the supraorbital eave has
the anterior part longitudinally narrow and rectangular inshape (Fig. 36A–D); each branchial region only has one
spine (Figs. 4, 5), the suborbital margin is separated fromthe basal antennal article and margin of postorbital tooth by
distinct fissures (Fig. 39A–C); the antero-external crested rimof the antennular fossa prominently overlaps the surface ofarticle (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 theG1 is very long, straight, gently curved, with the distal partslightly expanded and tapering without prominent projections
or folds (Fig. 7A–N). They are clearly congeneric.

In Indo-West Pacific species that have been referred to
Maja, the basal antennal article is distinctly longer thanbroad, the male thoracic sternites 3 and 4 are separated bya deep V-shaped cleft; the dactylus of the adult ambulatory
legs is slender, with scattered long or short setae; the carpusof 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 (herereferred to two new genera, Holthuijia and Ovimaja) havestructures similar to that seen in Maja s. str. In all otherIndo-West Pacific species, the antero-external crested rimof the antennular fossa does not or just slightly overlaps thebasal antennal article.

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 isdistinctively 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 eachside of the gastric region which has no or few granules andmarks 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 shorterand stouter, Figs. 36A–D, 39A–C); the supraocular eave isrelatively 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

Even then, these Indo-West Pacific “Maja” species canstill 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 forM. bisarmata is not prominently excluded from the orbit
as in normal Leptomithrax Miers, 1876 (Fig. 57A–F), it isalso 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 having115

Ng & Richer de Forges: Revision of the spider crab genus Maja
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, andboth are congeneric. The two species are here referred toa new genus, Rathbunaja, together with two new species
from the Philippines and Papua New Guinea, respectively.

The lateral margin of the carapace (excluding the outerbranchial 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 arelarge 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 arealso relatively distinct such that in large specimens, it is notalways easy to distinguish these from the other spines (e.g.,
see Fig. 4D-F). In genera like Paramaja and Alcomaja, thelateral carapace spines are only clearly discernible in juveniles
and smaller specimens (e.g., Figs. 10A, B, 11A-C, 12A),
becoming lower and more rounded in adults, so much sothat they sometimes cannot be distinguished (e.g., Figs. 9A,
B, 10C, D, 11D-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, theseare only slightly larger than the surrounding spinules andsharp 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 countsharp tubercles as spines (Fig. 34C, F, G). In general, thesame situation applies for the branchial spines for Ovimaja.

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 Majas. 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 justoutside the orbit, like the type species Leptomithrax, L.
longimanus (Miers, 1876) (Fig. 57A). In most other speciesof Leptomithrax, the antennal flagellum is positioned far
from the orbit (Fig. 57B–F). On the basis of the structureof 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 theorbit, just before the distal margin of the basal antennal
article (Fig. 2A, B, E, G, H). In Alcomaja, Paramaya andSakaija, the antennal flagellum is positioned right at themargin of the basal antennal article (Fig. 2C, D, F), and it
is not possible to state for certain whether it is just insideor outside the orbit.

The posterior carapace margin always has two spines, whichvary in size. In some genera (see later), the spines can beshort 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 theirinflated branchial regions (e.g., Figs. 8, 9, 10C, D, 11D, F,
G, 12D, E). In some Sakaija species, the spines can becomeso reduced that they are barely visible in larger specimens(Fig. 28A-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).

The intestinal regions of the carapace of all genera are covered
with small granules or tubercles, but only in Paramaya is therea prominent median spine in all species. In Maja, Neomaja,
Sakaija and Planaja, the intestinal region has a small spinethat may be eroded. In some species in these genera, e.g.
Sakaija africana, the intestinal spine is always present andis 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 ofshort spines along the median part of the intestinal region.

Paramaya is the only genus that has the proximal outer angleof the basal antennal article as a lobe-like projection thathas 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 moreclosely 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 theshapes of their basal articles are different (Fig. 40A–C,
O). In genera like Paramaja, Alcomaja and Sakaija, theantero-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, theThe supraorbital eave of Holthuija is relatively large andprominently 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 thedegree observed in Holthuija. In all the other genera, theanterior 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 spineor tooth (Figs. 36, 37D–O, 38).

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RAFFLES BULLETIN OF ZOOLOGY 2015
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 isseparated from the suborbital margin by a broad U-shapedsinus (Fig. 40A-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 inParamaja, 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).

sternites 1–4 are not elongated, the lateral junction between
sternites 2 and 3 is very deep, forming a distinct waist-likestructure 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 normallyslightly 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 structure of the epistome is distinctive for some genera.
Although there is a tendency for the epistome to becomeslightly 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 andmuch 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). Onlyin 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 orsmooth. In Maja s. str., Neomaja and Paramaya, there is alarge 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, theanterior 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 medianones being relatively broader (Figs. 41, 42C–H). However,in Paramaya, the median plates are characteristically morestrongly produced downward (Fig. 42A, B).

The merus and carpus of the cheliped in Maja s. str. and
Neomaja are prominently tuberculate, being covered withprominent 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 mucheven after considering size and sex.
The structure of the G1 is important in separating groups ofgenera. In Maja s. str., Neomaja, Paramaja and Paramaya,
the G1 is long, slender and gently curved to almost straightlong most of its length (Figs. 7, 15, 23). In Maja s. str.
and Neomaja, the distal part lined only with scattered shortspinules or very short setae (Fig. 7); it is covered with
scattered spinules and spines in Paramaja (Fig. 15), anddevoid of spines, spinules or setae in Paramaya, with adistinct 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 apetal-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 G1 in
which the distal part is partially pectinated and lined withstrong 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 withnumerous 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, thedistal 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

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, andis “inserted” into a deep concavity on the outer margin of
the ischium (Fig. 45I–L). As a consequence of this, theantero-internal part of the ischium is acutely triangular (Fig.
45I–L). In all other genera, the postero-external angle ofthe merus is proportionately broader and the antero-internalpart 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 areelongate, with the lateral junction between thoracic sternites
2 and 3 deep (Fig. 50A–C). In Ovimaja, while thoracic117

Ng & Richer de Forges: Revision of the spider crab genus Maja
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).

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.

A comment about the names Paramaya De Haan, 1837, andParamaja Kubo, 1936, is necessary. The spellings of the two
names are nearly identical and were probably intended to bethat way. Why De Haan spelled his name “Paramaya” (asa subgenus of Maja) with a “y” and not a “j” is confusing,but it is clearly intentional. De Haan (1837) used the nameParamaya 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 establishedthe name Paramaja for a very different species, Paramaja
kominatoensis Kubo, 1936, also from Japan. Balss (1957:
1628) noted that Paramaja Kubo, 1936, was a homonymof De Haan’s name and proposed a replacement name,Paramajella. This is not the case. Despite the close similarityof the two names, the current zoological code (ICZN, 1999)
treats them as separate names, the use of “y” and “j” for thetwo genera deemed sufficient to differentiate them. This ismost unfortunate and can certainly give rise to confusion.

Key to genera studied
1. Antennal flagellum positioned just outside of orbit, near rim(Fig. 57G, H); West Pacific........................Rathbunaja n. gen.
– Antennal flagellum clearly inside orbit (Fig. 2)......................2
2. Ambulatory merus with sharp subdistal spine on dorsal margin(Figs. 6F, G).............................................................................3– Ambulatory merus unarmed on dorsal margin (Fig. 6E, H)....
..................................................................................................4
3. Basal antennal article as long as broad; epistome as long as
broad (Fig. 39E, F); Eastern Atlantic............. Neomaja n. gen.
– Basal antennal article longer than broad; epistome longer thanbroad (Fig. 40A–C); Indo-West Pacific....................................
......................................................... Paramaya De Haan, 1837
4. Suborbital margin confluent with margin of postorbital tooth,not demarcated by fissure or cleft (Fig. 39G–O)....................5– Suborbital margin clearly separated from margin of postorbital
tooth by distinct deep fissures or cleft (Fig. 39A–F)..............7
5. Dorsal surface of carapace covered with small uniformly sized
small rounded granules (Fig. 28); G1 sinuous with numerous
long setae on distal surface (Fig. 33); Indo-West Pacific.........
........................................................................... Sakaija n. gen.
– Dorsal surface of carapace covered with numerous small andlarge granules of various types (Figs. 8–12); G1 straight or
curved with scattered setae on distal surface (Figs. 15, 19, 20);
Indo-West Pacific.....................................................................6

The morphological characters used here agree with themolecular analysis by Sotelo et al. (2009) who showed thatM. squinado, M. brachydactyla, M. crispata and M. cornuta(as Maja SA1 and Maja sp. SA2) belonged to one distinct
clade. The sister clade to this is Maja goltziana, which ishere transferred to its own genus, Neomaja as it has a suiteof distinct morphological characters (see later).

6. Pseudorostral spines relatively short, stout (Fig. 36G-K);
branchial regions of adults very swollen laterally and dorsally,
cardiac region prominently swollen, regions poorly demarcated;
lateral carapace spines undiscernible in adults (Figs. 8–12); G1
very long, mostly straight, distal part conical (Fig. 15); IndoWest Pacific........................................... Paramaja Kubo, 1936
– Pseudorostral spines long, slender (Fig. 36L–O); branchialregions distinctly less swollen, cardiac region gently convex
but not swollen, regions clearly demarcated; lateral carapace
spines, even if short, always visible in adults (Figs. 16, 17);
G1 proportionately shorter, more curved, part of dorsal foldbends downwards to form a distinct ventral projection (Figs.
19, 20); Indo-West Pacific............................. Alcomaja n. gen.

In summary, Maja Lamarck, 1801, is now restricted to four
primarily Atlantic and Mediterranean species. The otherspecies previously placed in the genus are transferred tonine 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

7. Basal antennal article as broad as long (Fig. 39A–D); G1
very long, distal part with pad-like folds (Fig. 7A–Q); Eastern
Atlantic to Western Indian Ocean...........Maja Lamarck, 1801
– Basal antennal article longer than broad (Fig. 40); G1
otherwise...................................................................................8
8. Carapace ovoid, distinctly longer than broad (Fig. 34C, F, G);basal antennal article with several large rounded granules andtubercles (Fig. 40O); male thoracic sternites 1–4 not elongated,
lateral junction between sternites 2 and 3 very deep, forming
waist-like structure (Fig. 51I); margin of male sterno-abdominal
cavity almost flat, without rim (Fig. 52L); G1 strongly bent,
distal part dilated with 2 folds (Fig. 35F–L); West Pacific......
..........................................................................Ovimaja n. gen.

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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 mudand sand, Rijeka Bay, near Island of Krk, coll. trawler, 2April 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, 2females (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. TunisianMarine 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.

Carapace pyriform, not distinctly ovoid (Figs. 24, 25, 34A,B); basal antennal article with small granules, never large
tubercles (Fig. 40N); male thoracic sternites 1–4 elongated,
lateral junction between sternites 2 and 3 relatively shallow(Fig. 51F, G); margin of male sterno-abdominal cavity on
thoracic somite 4 raised, entire, forming clear rim (Fig. 52K);
G1 otherwise............................................................................99. Lateral carapace margin with 3 spines (Figs. 24, 25); supraoculareave very prominent, large, anterior part prominently foliaceous(Fig. 37D–J); G1 relatively stout, straight or gently bent, distal,part simple, with scattered spinules and short setae (Fig. 27);
Indo-West Pacific........................................... Holthuija n. gen.
– Lateral carapace margin with 6 spines (Fig. 34A, B); supraoculareave otherwise (Fig. 38A); G1 slender, curved, distal part sharply
tapering, margins distinctly chitinised, line with numerous shortspinules (Fig. 35A–E); West Pacific................Planaja n. gen.

Diagnosis. Large species (adult carapace length in excessof 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 broadwith 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 partstraight (Fig. 7A–C).

Key to species of Maja Lamarck, 1801, s. str.
1. Supraorbital eave relatively broad, as wide as long; dorsoanterior surface of carapace with 2 unarmed, relatively smoothdark oval window-like patches, one on each side of gastricregion (Figs. 6A, B, 36E); ischium of third maxilliped ca. 1.5
times as long as wide (Fig. 43F); Mediterranean and adjacentAtlantic waters................................ Maja crispata Risso, 1827
– Supraorbital eave relatively narrow, longer than wide; dorsoanterior surface of carapace evenly granulated, without any
unarmed or ovate patches (Figs. 4, 36A–D); ischium of third
maxilliped twice as long as wide (Fig. 43A–E); eastern Atlantic
to western Indian Ocean..........................................................2
2. Carapace of adults longer than wide; adult pseudorostral spines
subparallel; restricted to Mediterranean (Fig. 36A)..................
.................................................. Maja squinado (Herbst, 1788)– Carapace of adults rounded, as long as large; adult pseudorostralspines diverging, forming a V (Fig. 36B–D); Atlantic to South
Africa........................................................................................3
3. Spines of lateral carapace margin relatively short; antorbital
spine almost straight (Figs. 4C–F, 36B, C); northern Atlantic
to mid-Atlantic....................... M. brachydactyla (Balss, 1922)– Spines of lateral carapace margin relatively long; antorbital spine
curved (Figs. 5, 36D); southern Atlantic to South Africa........
.....................................................M. cornuta (Linnaeus, 1758)

Remarks. Maja squinado (Herbst, 1788) is the type speciesof the genus (ICZN Opinion 511, see Holthuis, 1958). In adetailed morphological and biometric study using extensive
material from the Mediterranean and the Atlantic coast ofEurope, 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 theMediterranean, 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 notedthat 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.

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) (notCancer 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

The excellent study by Neumann (1998), and the series ofmolecular studies by Sotelo et al. (2007, 2008, 2009) havehelped resolve the identity of M. brachydactyla Balss, 1922,
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Ng & Richer de Forges: Revision of the spider crab genus Majaa large species that has long been regarded as synonymousor merely a subspecies of M. squinado (see Manning &Holthuis, 1981: 307). It is clearly a distinct species and theexamination of the present material confirms the observationsof Neumann (1998). As it turns out, the commercially morevaluable species that occurs from Norway to the Atlantic
coasts of Spain that had previously been known as “Majasquinado” should now be called M. brachydactyla. Maja
squinado s. str. is restricted to the Mediterranean Sea. Even
locations immediately outside the Mediterranean Sea suchas 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 thispart of the ocean, both species may be sympatric.

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 sevenstraight segments. Hands smooth, glabrous, without setae.
Chelipeds smaller than other articles, base bidentate. Feetfully hirsute. Digits hirsute.]

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 relatively detailed description by Linnaeus (1764)
matches that of Maja squinado, M. brachydactyla or M.
cornuta. The only unusual character seems to be the commentthat rostrum bends below the eyes. In Maja species, the twomost 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 thatis 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 intraocularspines”, and the second time, he describes the actual rostrum,noting that it “bends below the eyes”. Some authors havespeculated that Linnaeus (1764) was actually saying thatthe pseudorostrum bends below the eyes, a character thatdefines 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 specimensof 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.

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 utrinqueciliata spinis 6, validis. Disci spinae 3 transversae & una
anterior; ommes erectae, reliquis aequales. Spinae 2 minores,
erectiusculae, ad basin thoracis. Rostri loco Spinae 2 interoculos, 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-shapedwith 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.

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 isclear 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, hemakes no mention of Linnaeus. He also reports the species
from Norwegian Seas for the first time. In his next mentionof 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.

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 thoseof 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 pseudorostralspines as a character – subparallel in M. squinado (Fig. 4A)
but clearly diverging in M. brachydactyla (Fig. 4C–F) andthis holds in large specimens, especially males. Smaller
specimens of M. squinado, however, have pseudorostralspines that are somewhat intermediate in condition, andcertainly cannot be described as subparallel (Fig. 4B). As
such, both the carapace shape and pseudorostral spine formshould be used as characters, with the G1 structure valuable
when adult males are available.

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has actually not been used as an available name since 1900,
invoking Article 23.9.1 of the Code (ICZN, 1999: 27) tosuppress 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 generaluse only in recent years since Neumann’s (1998) revision.

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 andanother 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 patcheson 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.

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) andindicated 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 depositedin 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 ofCancer cornutus Linnaeus, 1758. As a result of this action,
all three names are synonyms, with the oldest name being
Maja cornuta (Linnaeus, 1758).

Maja crispata is not only relatively smaller at adult size,
it also has two distinct patches on the sides of the gastricregion that are not only darker in colour than the other partsof 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 havethese 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 thesmaller 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 second outstanding taxonomic problem associatedwith M. squinado s. str. and M. brachydactyla 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 ordiagnostic 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 typeof M. tuberculata. Herklots (1861: 19) in his study of thematerial of De Haan in the Leiden Museum had indicatedthat 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 materialor 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.

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 withsquinado-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 thanparallel. In any case, there is no extant specimen as Seba’s
material is lost (see Koh & Ng, 2008: 341). A neotype forCancer squinado Herbst, 1788, is designated here to stabilise
the taxonomy of this species (see later).

Through the courtesy of Charles Fransen, the first author
examined the box that supposedly contains the mouthpartsof Maja tuberculata. The mouthparts remain attached andarranged in a row, and on the original background label iswritten “Maja tuberculata”. While this label is old, it is notknown if this was written by De Haan or later on by someone
else. De Haan (1839: pl. F) figured the various mouthpartsof Maja squinado (right first to third maxillipeds, right firstand second maxillas and right mandible), and only the last

So what is Cancer cornutus Linnaeus, 1758? From all theavailable 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,
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which of the Atlantic species M. tuberculata is actuallyconspecific with. It poses some potential nomenclaturalproblems if Maja tuberculata De Haan, 1839, is the sameas 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 synonymof Cancer squinado Herbst, 1788, at least for the time being.

two figures (which depicted a right third maxilliped and rightfirst maxilliped) belonged to Maja tuberculata. The third
maxilliped of M. tuberculata figured is smaller than that ofM. 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, areof different sizes; the attached one is 11.1 mm (measured
from the anterior margin of the merus to the posterior marginof the basis) (Fig. 44E), the other 17.3 mm (Fig. 44F). They
clearly belong to two separate specimens. On the basis of thestructure 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 sizewith 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.

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 thegenus, 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 anapplication to the International Commission for Zoological
Nomenclature.
Maja brachydactyla Balss, 1922
(Figs. 2A, 3E, 4C–F, 7D–I, 36B, C, 39B, 41B, 43C, D,
47C, D, 52A, 53B, C, 55B)
Cancer cornutus – Fabricius, 1787: 325 (not Cancer cornutus
Linnaeus, 1758).
Maja squinado var. brachydactyla Balss, 1922: 74.
Maia gigantea Baudouin, 1931: 256.
Maja squinado – Takeda, 1982: 129. – Takeda, 1993: 43. – Muraoka,
1998: 27, pl. 14 fig. 3 (not Cancer squinado Herbst, 1788).
Maja brachydactyla – Neumann, 1998: 1672. – d’Udekem d’Acoz,
1999: 188. – Abelló et al., 2014:e77. – Moro et al., 2014: 52.
[For complete synonymy, see Neumann, 1998: 1672; d’Udekem
d’Acoz, 1999: 188–189].

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 measuresabout 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 thatit 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 thatof M. tuberculata (Fig. 44E). In the Indo-West Pacific, only
Paramaja gibba, Paramaja kominatoensis, Paramaja turgida,
Paramaya spinigera and Paramaya ouch are known to growto 80 mm in carapace length, but their third maxillipeds (Figs.
43L–P, 45I–K) are very different in structure compared tothat of M. tuberculata. As such, the suggestion by Yamaguchi& Baba (1993) that Maja tuberculata may have originatedfrom the “Moluccas” and it is Herklots’ (1861) unnamed
species seems unfounded.

Material examined. Spain – 1 male (161.2 × 140.1 mm),
1 ovigerous female (143.2 × 127.6 mm) (ZRC 2008.0179),
northwest of Baiona, from fishermen, 25 January 2008.
— 1 female (NHM 1956.5.2.49), Lanzarote. — 1 male
(USNM 122017), northwestern Spain. Portugal – 1 male
(NHM 98.1.28.1), Madeira. England – 2 males (98.4 ×
89.0 mm, 92.6 × 75.5 mm), 1 female (127.8 × 111.9 mm),
1 ovigerous female (119.0 × 103.0 mm) (ZRC 2009.1130),
Worbarrow Bay, south Dorset, 5 m, 50°37.06’N 2°11.23’W,
U.K., coll. S. De Grave, 5 June 2005. — 1 female (NHM
1983.46), Thorn Island, Pembrokeshire, 10 m, coll. 10
August 1981. — 1 female (NHM 1983.46), Knoll Pins,
Lundy, 22 m, coll. 3 September 1982. — 1 female (NHM
1985.11), Eastbourne, coll. 8 August 1984. — 1 female(NHM 1983.44), Littlehampton, 50°39.11’N 00°36.9’W, coll.
29 July 1983. — 1 female (NHM 1959.10.15.1), Langston
Harbour, Portsmouth. — 2 males, 2 females (NHM 1965.12),
Durdle Door, Dorset, coll. 6 June 1965. — 2 females (NHM
1983.448), Start Bay, Devon. — 2 females (NHM 1983.44),
Devon, Start Bay, off Hall Sands, 12 m, 10 May 1981. — 1
female (NHM 1981.54), Devon, Brixham Harbour, 5 m, coll.
9 July 1981. — 1 female (NHM 1902.3.10.2), Plymouth.
— 1 female (NHM 1981.237), Plymouth. — 1 male (NHM
98.5.7.264), Falmouth. — 1 male (NHM 1984.454), mouth

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 granulatedon the carapace when younger (see Neumann, 1998). Since
the distributions of M. squinado and M. brachydactyla
partially overlap (Neumann, 1998), we cannot be sure122

RAFFLES BULLETIN OF ZOOLOGY 2015
of Fal river, Trefusis Point, 50°9.8’N 5°3.1’W, coll. 4
July 1984. — 2 females (NHM), Cornwall. — 1 male, 1female (NHM 1981.37), Belle Creve Bay, Guernsey, coll.
11 June 1981. — 1 male, 3 females (NHM), Guernsey,coll. September 1982. — 1 male (NHM), Rocquaire Bay,
Guernsey, coll. 18 August 1982. — 1 male, 4 females(NHM 1951.1.25.134–136), English Channel coast. — 3
males, 4 females (NHM), southern England. — 1 male
(USNM 6773), England, from A. M. Norman. — 1 male,
1 female (USNM), Channel Islands, coll. E. Lovett. — 1
male (NSMT 3805), Portsmouth, Devon, English coasts,
no other data. Ireland – 1 male (NHM 1982.13), Galway
Bay, 10 m, coll. 27 August 1981. Morocco – 1 male (61.5
× 53.8 mm) (MNHN-IU-2013-4039), coll. Lt. Dyé, 1905.

110.0 × 100.0 mm with spines) (ZMS), Port Elizabeth, South
Africa, obtained from natural history dealer Rolle in Berlin,
1891. South Africa – 1 dried female (71.0 × 60.3 mm, 80.0 ×
70.0 mm with spines) (ZMS) (paralectotype of Maja squinado
var. capensis Ortmann, 1894), same data as lectotype. — 1
male (88.8 × 86.8 mm) (NHM 1928.12.1.177) (lectotype,
here designated, of Mamaia queketti Stebbing, 1908), Port
Elizabeth, Stebbing Collection, coll. Fitzsimons, 5 November
1907. — 1 female (NHM 1913.3.29.4), Durban, coll. H. C.
Chubb. — 2 males (NHM 1928.12.1.178–179), Bird Island,
45 fathoms, coll. H. W. Bek Marley. — 1 ovigerous female(114.9 × 103.8 mm) (ZRC 2005.0014), Kenton on Sea, Bay
World, Port Elizabeth, eastern part of Cape of Good Hope,coll. S. Warren, 1 December 2003. — 1 male (115.3 ×
103.4 mm) (ZRC 2013.1184), station ACEP 1.4, Transvaal,
off Durban, 29°58.56’S 31°04.98’E – 29°57’S 31°06.3’E,
65 m, trawl, coll. S. Fennessy, 18 March 2010. — 2 males(79.8 × 68.9 mm, 53.8 × 45.1 mm) (ZRC 2013.1183), station
ACEP 2.4, Transvaal, 29°29.9’S 31°52.8’E – 29°26.56’S
31°53.5’E, 184 m, trawl, coll. S. Fennessy, 19 March 2010.

Diagnosis. Large species (adult carapace length in excessof 90–100 mm). Carapace pyriform, more rounded in large
specimens; dorsal surface strongly convex; uniformly covered
with granules and short spines (Fig. 4C–F). Pseudorostralspines long, gently curved to almost straight, gently diverging(Figs. 4C–F, 36B, C). Supraorbital eave relatively narrow,antorbital spine sharp, almost straight to gently curved
upwards; intercalated spine stout, short, touching postorbital
spine basally; postorbital spine largest, long, sharp; hepatic
spine sharp with 2 accessory spines basally (Figs. 4C–F,
36B, C). Lateral margin of carapace with 3 spines andseveral small sharp granules and short spines (Fig. 4C–F).
On median row 7 strong spines: 3 gastric, 2 subgastric, 1
cardiac, 1 intestinal, with 1 sharp granule between cardiac andintestinal spines; spines relatively lower in large specimens(Fig. 4C–F). Branchial region with 2 spines; posterior
carapace margin with 2 short spines (Fig. 4C–F). Basalantennal article very broad with 2 long distal spines; outermargin with proximal spine, appressed on suborbital tooth(Fig. 39B). Ischium of third maxilliped rectangular, much
longer than broad (Fig. 43C, D). Carpus of cheliped with
distinct tubercles and granules (Fig. 53B, C). Ambulatory
dactyli covered with short setae (Fig. 55B). G1 long, gently
curved; distal part gently curved downwards (Fig. 7D–I).

Diagnosis. Large species (adult carapace length in excess of
80 mm). Carapace pyriform; dorsal surface strongly convex;
uniformly covered with granules and short spines (Fig. 5).
Pseudorostral spines long, straight, strongly diverging (Figs.
5, 36D). Supraorbital eave relatively narrow, antorbital
spine sharp, curved upwards; intercalated spine stout, short,
touching postorbital spine basally; postorbital spine largest,long, sharp; hepatic spine sharp with 2 accessory spines
basally (Figs. 5, 36D). Lateral margin of carapace with 3strong spines and several small sharp granules and shortspines (Fig. 5). On median row 7 strong spines: 3 gastric,
2 subgastric, 1 cardiac, 1 intestinal, with 1 sharp granulebetween cardiac and intestinal spines (Fig. 5). Branchialregion with 2 spines; posterior carapace margin with 2
spines (Fig. 5). Basal antennal article very broad with 2 long
distal spines; outer margin with proximal spine, appressedon suborbital tooth (Fig. 39C). Ischium of third maxilliped
rectangular, much longer than broad (Fig. 43E). Carpus ofcheliped with distinct tubercles and granules (Fig. 53D, E).
Ambulatory dactyli covered with short setae (Fig. 5). G1
long, gently curved; distal part gently curved downwards(Fig. J–N).

Remarks. Small specimens of M. brachydactyla tend to be
more spiny overall, with the lateral spines relatively stronger(Fig. 4C) compared to larger ones (Fig. 4D-F). See also
discussion for M. squinado.

Remarks. Maja cornuta (previously M. capensis) occurs
in the sublittoral zone on the Indian Ocean side of South
Africa. Compared with M. brachydactyla, the carapace ofM. cornuta is not as inflated, the pseudorostral spines are
relatively longer and more slender, the antorbital spine ismore distinctly curved anteriorly, there is a relatively wider
gap between the intercalated, supra- and postorbital spines(the three spines are closely appressed in M. brachydactyla),
and all the spines on carapace margin, including the hepaticone, are proportionately longer and sharper. The G1 structures
are also slightly different between the two species: in M.
brachydactyla it is straight with a large terminal opening(Fig. 7D–I), while in M. cornuta, it is slender and curved
with a more slender tip (Fig. 7J–L). See also discussion forM. squinado.

Maja cornuta (Linnaeus, 1758)(Figs. 5, 7J–N, 36D, 39C, 43E, 47E, F, 53D, E)
Cancer cornutus Linnaeus, 1758: 629. – Linnaeus, 1764: 445.
Maja squinado var. capensis Ortmann, 1894: 40.
Mamaia queketti Stebbing, 1908: 3, pl. 27.
Mamaia capensis – Barnard, 1950: 59, fig. 13a.
Maja capensis – Neumann, 1998: 1675.
? Maja capensis – Ng et al., 2008: 117 (list).
[For complete synonymy, see Neumann, 1998: 1675, as Maja
capensis Ortmann, 1894]

Material examined. Lectotype of Maja squinado var.
capensis (here designated) and neotype (here designated)of Cancer cornutus Linnaeus, 1758: male (96.8 × 89.5 mm,123

Ng & Richer de Forges: Revision of the spider crab genus Maja

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).

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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.

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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.

Maja crispata Risso, 1827
(Figs. 3C, D, 6A, B, 7O–Q, 36E, 39D, 41C, 43F, 47G,
52B, 53F, 55C)

Maja erinacea Ninni, 1924: 49, 50, 2 un-numbered figures.
(see Neumann, 1996b: 828; d’Udekem d’Acoz, 1999: 189, forreferences)

Maja crispata Risso, 1827: 23. – Neumann, 1996b: 828; d’Udekem
d’Acoz, 1999: 189.
Maja verrucosa H. Milne Edwards, 1834: 328, pl. 3 figs. 1–14. –Bouvier, 1940: 322. – Monod, 1956: 477. – Zariquiey Alvarez,
1968: 447, figs. 149b, 150a–f, 156a. – Muraoka, 1998: 27.
Cancer majoides Nardo, 1847: 3.

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

R.B. Manning, 21 June 1974. — 1 ovigerous female (USNM
205791), Mazara Del Vallo, Sicily, coll. R.B. Manning,
2 September 1985. — Greece – 1 male, 1 female (NHM
1968.638), Chios Islands, coll. G. Potts. — 1 male (NHM
1968.637), Chios Islands, coll. G. Potts. — 1 female (NHM
96.5.30.3), Smyrna, coll. Y. Holswood. Croatia – 1 male
(ZRC), Rovinj, Figarola, coll. Senckenberg Museum class,don. SMF, 23 August 1989. — 1 female (ZRC), Rovinj, coll.
Senckenberg Museum class, don. SMF, 20 August 1989. — 1
male (ZRC), Rovinj, coll. Senckenberg Museum class, don.
SMF, August 1993. Slovenia – 1 male, 1 female (NHM
98.5.7.301), Pirano Adriatic, don. Vienna Museum, coll.
Norman. Macedonia – 1 female (NHM 1919.19.4.281), coll.
Waterston. Israel – 1 male (USNM 152564), Mikhmoreth,near rocks, 3–4 m, coll. D. Popper, 12 February 1972.
Diagnosis. Medium-sized species (adult carapace length
60–70 mm). Carapace distinctly pyriform, much longer thanbroad more rounded in large specimens; dorsal surface gently
convex; uniformly covered with granules and short spines
except for window-like patch on each side of gastric region
which is weakly or not armed (Fig. 6A, B). Pseudorostralspines short, straight, strongly diverging (Figs. 6A, B, 36E).
Supraorbital eave relatively wide, antorbital spine sharp,
almost straight or gently curved upwards; intercalated spine
short, touching postorbital spine basally; postorbital spine
largest, long, sharp; hepatic spine slender, sharp with 2 or 3
small spines or tubercles basally (Figs. 6A, B, 36E). Lateral
margin of carapace with 3 strong spines and several small
sharp granules and short spines (Fig. 6A, B). On median row,
no long spines discernible, with 4 sharp tubercles: 2 gastric,
1 cardiac, 1 intestinal (Fig. 6A, B). Branchial region with 2
spines; posterior carapace margin with 2 distinct spines andone small median one (Fig. 6A, B). Basal antennal article
very broad with 2 long distal spines, proximal triangularspine passing behind postorbital spine; outer margin withproximal tooth, appressed on suborbital tooth (Fig. 39D).
Ischium of third maxilliped squarish, slightly longer thanbroad (Fig. 43F). Carpus of cheliped with low granules or
low, uneven ridges, otherwise almost smooth (Fig. 53F).
Ambulatory dactyli covered with relatively long setae (Fig.
55C). G1 long, gently curved; distal part dilated, gently
curved downwards (Fig. 7O–Q).
Remarks. For the classification of this species in Maja s. str.,
see Remarks under the genus. The identity of Maja erinacea
Ninni, 1924, described from the Adriatic, is dubious. Türkay
(in Števčić, 1990) had suggested that it was synonymouswith Maja goltziana d’Oliviera, 1889, but the problem is thatthe latter species is not known from the Mediterranean Sea.
d’Udekem d’Acoz (1999: 189) noted that it is probably not
M. goltziana. Ninni’s (1924) un-numbered figures are rather
schematic, but the armature of the carapace spines does beara marked resemblance to M. goltziana. One of the otherdiagnostic features of M. goltziana is that the ambulatory
meri are armed with a prominent distal spine on the dorsal
margin. Ninni (1924: 49–50) writes that in M. erinacea, “Labase di quasi tutte le spine è ricoperta da peli, così pure il
carapace e tutti I paia di piedi ad eccezione del primo paio.
Questo è molto più piccolo degli altri, manca della piccola127

Ng & Richer de Forges: Revision of the spider crab genus Maja
spina rivolta all’ insù e che trovasi invece sul quarto articolo
sulla parte superiore di ogni altro paio di piedi” but it is
not very clear if he actually says the ambulatory merus has
the long sharp spine. In any case, some specimens of M.
crispata are known to have relatively longer carapace spines(Neumann, 1996a: fig. 2G), and M. goltziana is not known
from the Mediterranean Sea. As such, we follow Neumann(1996a: 828) in treating Maja erinacea Ninni, 1924, as a
junior synonym of M. crispata, at least for the time being.

spine; dactylus relatively short, curved, covered with dense
short setae except for corneous distal quarter (Figs. 6C, D,
F, 55D). Thoracic sternum wide; surfaces of somites 5–8
almost smooth; sternites 3 and 4 slightly depressed; margin
between sternites 2 and 3 demarcated by notch; anterior
margin of sterno-abdominal cavity not forming complete rim(Figs. 47H, I, 52C). Male abdomen subrectangular, with 6
free somites and telson; somites 3 and 4 distinctly wider thansomites 5, 6 and telson; telson semicircular (Fig. 47H, I).
Male press-button abdominal locking mechanism submedianin position on sterno-abdominal cavity (Fig. 52C). Female
abdomen dome-shaped, covering most of thoracic sternum.
G1 very long, slender, gently curved, distal part dilated,
sub-spatuliform, without subdistal process, with scatteredvery short setae (Fig. 7R–T).

Maja crispata is known from Portugal to Mauritania and in
the Mediterranean Sea, occurring at depths not deeper than
120 m. The larvae were described by Neumann (1993), withan unusual case of heteromorphosis recorded by CarmonaSuarez (1990).
Neomaja n. gen.

Type species. Maja goltziana d’Oliviera, 1889, by present
designation.

Diagnosis. Carapace pyriform; dorsal surface covered by
granules and tubercles, spines large; gastric and branchialregions not distinctly demarcated, separated by shallow
grooves; gastric and cardiac regions each with 1 distinct spine(Fig. 6C, D). Intestinal region with distinct median spine(Fig. 6C, D). Pseudorostral spines long, strongly divergent,forming a V (Fig. 6C, D). Supraorbital eave with anterior
part longitudinally narrow, rectangular, not prominently
expanded; antorbital spine sharp (Fig. 36F). Intercalated
spine relatively short, stout, separated from supraorbital eaveand postorbital spine by narrow gaps; postorbital spine verystrong, long. Hepatic region with 1 strong spine, shorter thanpostorbital spine; 1 smaller spine below (Fig. 36F). Lateralcarapace margin with 3 large spines, branchial region withstrong spine (Fig. 6C, D). Posterior carapace margin with
2 median spines (Fig. 6C, D). Eyes relatively long, thin,slightly curved, with ovoid cornea (Fig. 36F). Antennal
flagellum short, slender. Basal antennal article as broad as
long, quadrate, with 1 long distal spine, 1 low, lobiform
proximal lateral tooth; proximal outer angle rounded; innerand outer lateral margins entire; antero-external crested
rim of antennular fossa prominently overlaps subdistal partof basal antennal article as hook-shaped structure by morethan half its width (Fig. 39E, F). Epistome as long as wide,
anterior margin with 2 large rounded tubercles; posterior
margin composed of 4 rectangular plates separated by deep
fissures (Figs. 39E, F, 41D). Suborbital margin separated frombasal antennal article and margin of postorbital tooth by 2
distinct gaps, basally wide but with anterior parts adjoined
to form fissures (Figs. 39E, F, 41D). Outer surface of third
maxilliped covered by short setae; ischium subrectangular,
distinctly longer than broad; postero-external angle of merus
relatively broad, “inserted” into shallow concavity on outermargin of ischium; antero-internal part of ischium rounded,auriculiform (Fig. 43G, H). Male chelipeds long in adultmales, surfaces of merus and carpus covered with distinct
tubercles and granules; carpus elongate, with granulated
longitudinal ridge; propodus of palm elongated, curved,
smooth, without lateral cristae, slightly enlarged, longer thanpalm; fingers long, slender, gently curved, with narrow basal
gape when closed (Figs. 6C, D, 53G, H). Ambulatory legs
relatively short, thick; merus with strong, stout subdistal

Etymology. The genus name alludes to the new taxon being
different from Maja s. str. gender feminine.
Remarks. Maia goltziana was compared to Maja spinigera
De Haan, 1837 (presently in Paramaya) by Bouvier (1940:
323, 324) who commented that in the former species, “…uneforte épine au bout distal du mérus des pattes ambulatoires”.
The species is also characterised by possessing relatively
shorter antennae (Fig. 39E, F) compared to other Maja s.
str. species (Fig. 39B, C). The differences in the form of theepistome (relatively longer) (Figs. 39E, F, 41D), presenceof distal spines on dorsal margins of the ambulatory meri
(Fig. 6C, D, F), and proportionately broader male abdomen(Fig. 47H, I) suggest it should be referred to its own genus,
Neomaja. Its affinities, however, are clearly with Maja s.
str., notably in the relatively broad and short basal antennal
article (Fig. 39E, F), form of the chelipeds (including theprominent granulation on the carpus) (Fig. 53G, H), andgeneral structure of the G1 (Fig. 7R–T). This taxonomy isconsistent with the molecular study of Sotelo et al. (2009)
which shows that M. squinado, M. brachydactyla, M. crispataand M. cornuta (as Maja sp. “SA”) form a distinct clade,separate but sister to M. goltziana.
Neomaja goltziana (d’Oliveira, 1889)(Figs. 6C, D, F, 7R–T, 36F, 39E, F, 41D, 43G, H, 47H, I,
52C, 53G, H, 55D)
Maja goltziana d’Oliviera, 1889: 3. – Capart, 1951: 100. – Monod,
1956: 478, Fig. 644–645. – Rossignol, 1957: 116. – Crosnier,
1967: 339, figs. 31, 32. – Longhurst, 1958: 88. – Zariquiey
Alvarez, 1968: 447. – Massuti, 1970: 127. – d’Udekem d’Acoz,
1999: 189. –Moro et al., 2014: 52.
Maïa goltziana – Le Loeuff & Intès, 1968.
Maia goltziana – Maurin, 1968a: 30. – Maurin, 1968b: 484.
[see d’Udekem d’Acoz, 1999: 189 for references]

Material examined. Morocco – 1 female (48.0 × 40.5 mm)(MNHN-IU-2013-4037), station 80, 30°21’N 09°56’W,
11–40 m, coll. Vanneau, 31 August 1925. Portugal – 2 males,
1 female (NHM1966.4.6.62–64), off Malha, Portuguese
coast, 3–4 miles offshore, 40 fathoms, coll. G. Palmer,
128

RAFFLES BULLETIN OF ZOOLOGY 2015

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 × 126.3mm) (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.

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Ng & Richer de Forges: Revision of the spider crab genus Maja

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].

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Paramaja Kubo, 1936

26–27 August 1965. Congo – 1 male (73.4 × 65.0 mm), 1
female (70.9 × 64.6 mm) (MNHN-IU-2013-4046), west of
Pointe Noire, 80 m, coll. July 1958. Senegal – 1 male (74.7
× 66.7 mm) (MNHN-IU-2013-4043), station 9, 14°27’N
17°35’W, Dakar, 180–220 m, coll. trawl, L. Amaro, 21
February 1981. Gabon – 1 male (90.2 × 83.1 mm) (MNHNIU-2013-4044), 40 m, coll. July 1959. Cameroon – 1 male
(67.3 × 57.7 mm) (MNHN-IU-2013-4045), coasts, 3°54’N
8°50’E, 65–70 m, coll. 25 August 1963. Nigeria – 1 male,
2 females (1 badly damaged) (NHM 1957.5.26.115–117),
Afons Deep, coll. A.R. Longhurst. Azores – 1 male (SMF9832), Canary Islands, La Palma, 160 m, coll. E. Santaella,
11 June 1974. — 10 juveniles (SMF 24519), Nilmundung,
32°00.54’N 31°53.70’E – 32°00.55’N 31°52.61’E, 196-199
m, coll. FS Meteor, 1 June 1993.

Paramaja Kubo, 1936: 361.
Paramajella Balss, 1957: 1628 [unnecessary replacement namefor Paramaja Kubo, 1936].
Maja – T. Sakai, 1938: 296 (part). – T. Sakai, 1965: 83 (part).
(not Maja Lamarck, 1801).

Diagnosis. Carapace rounded in adults; dorsal surface
evenly inflated, covered by granules or tubercles; gastricand branchial regions clearly delimited by grooves (Figs.
8–12). Intestinal region without distinct median spine (Figs.
8–14). Pseudorostral spines relatively short, diverging,forming a V (Figs. 8–13). Supraorbital eave with anterior
part longitudinally narrow, rectangular, not prominently
expanded; antorbital spine sharp (Fig. 36G–K). Intercalated
spine distinct, separated from supraorbital eave andpostorbital spine by distinct gaps; postorbital spine strong,
lobiform; hepatic region with 1 strong spine, much shorter
than postorbital spine; 1 or more smaller spines below (Fig.
36G–K). Lateral carapace margin with 3 or 4 spines andnumerous tubercles, branchial region with larger tubercle,
not discernible from other surrounding tubercles in large
specimens (Figs. 8–13). Posterior carapace margin with 2
short median spines, not visible from dorsal view when adultdue to swollen carapace (Figs. 8–12). Eyes relatively short,
slender, with large ovoid cornea (Fig. 36G–K). Antennal
flagellum short, slender. Basal antennal article longer thanbroad, rectangular; surface with several tubercles, with 2 blunt
spines distally; inner and outer lateral margins may have low
granules or lobes, or entire; proximal outer angle rounded;
antero-external crested rim of antennular fossa touches but
does not significantly overlap distal part of basal antennal
article (Fig. 39G–K). Epistome much wider than long, anterior
margin unarmed; posterior margin composed of 4 rectangularplates separated by shallow fissures (Figs. 39G–K, 41E,
F). Suborbital margin separated from basal antennal article
by short fissure, confluent with margin of postorbital tooth(Fig. 39G–K). Outer surface of third maxilliped coveredby short setae in adults; ischium subrectangular, distinctly
longer than broad; postero-external angle of merus relatively
broad, “inserted” into shallower concavity on outer margin ofischium; antero-internal part of ischium rounded, auriculiform(Fig. 43I–P). Male chelipeds relatively short in adult males,surfaces of merus and carpus almost smooth or with verysmall granules; carpus short, with low, uneven longitudinalridge; propodus of palm elongated, curved, smooth, without
lateral cristae in young, short, inflated in adults, palm longer
than fingers; fingers long, slender, gently curved, with distinct
basal gape when closed (Figs. 8–12, 53I–L). Ambulatory
legs relatively long, slender; merus without dorsal subdistal
spine; dactylus elongate, curved, covered with long setae
when young except for corneous tip, almost completely
smooth in adults (Figs. 6H, 8–12, 55E–J). Thoracic sternum
wide; surfaces of somites 5–8 with numerous prominent
rounded tubercles and granules; sternites 3 and 4 slightly
depressed; margin between sternites 2 and 3 demarcatedby deep notch; anterior margin of sterno-abdominal cavity
not forming complete rim (Figs. 48, 52D). Male abdomen
subrectangular, with 6 free somites and telson; somites 3–6
and telson subequal in width (Fig. 48). Male press-button

Diagnosis. Medium-sized species (60–70 mm carapace
length). Carapace pyriform; dorsal surface gently convex;
uniformly covered with granules and short spines, those on
posterior part of branchial and intestinal regions spiniform(Fig. 6C, D). Pseudorostral spines long, straight, strongly
diverging (Figs. 6C, D, 36F). Supraorbital eave relatively
narrow, antorbital spine sharp, gently curved upwards;
intercalated spine stout, short, touching postorbital spine
basally; postorbital spine largest, long, sharp; hepatic spine
sharp with 1 short accessory spines basally (Figs. 6C, D,
36F). Lateral margin of carapace with 3 strong spines,
second longest, and several small sharp granules and shortspines (Fig. 6C, D). On median row 5 spines: 3 gastric, 1
cardiac, 1 intestinal (Fig. 6C, D). Branchial region with 1strong spine; posterior carapace margin with 2 long spines(Fig. 6C, D). Basal antennal article very broad with 2 long
distal spines; outer margin with proximal lobiform tooth,appressed on suborbital tooth (Fig. 39E, F). Ischium of third
maxilliped rectangular, much longer than broad (Fig. 43G,
H). Carpus of cheliped with distinct tubercles and granules(Fig. 53G, H). Ambulatory merus with strong subverticalsubdistal spine on dorsal margin (Fig. 6F); dactylus covered
with short setae (Fig. 55D). G1 long, gently curved; distal
part gently curved upwards (Fig. 7R–T).
Remarks. Neomaja goltziana was originally described from
Portuguese waters (d’Oliviera, 1889). Although the year of
publication is usually cited as 1888, volume 36 (where thepaper was published) is dated July 1888 to June 1889, andthe year “1889” was printed at the bottom of title page ofthe paper.
Neomaja goltziana is superficially similar to Paramaya
spinigera, especially with regards to its spiny carapace and theambulatory meri possessing a strong distal spine on the dorsal
margin. Bouvier (1940: 323) comments that “une forte épine
au bout distal du mérus des pattes ambulatoires”. However,as discussed under the genus, there are many substantial
differences between the two species at the generic level.
Neomaja goltziana was described from Portuguese waters.
Manning & Holthuis (1981: 307) reported a distribution from
Mediterranean to Congo, between 15 to 200 m.

131

Ng & Richer de Forges: Revision of the spider crab genus Majaabdominal locking mechanism submedian in position onsterno-abdominal cavity (Fig. 52D). Female abdomen
dome-shaped, covering most of thoracic sternum. G1 very
long, slender, almost straight for most of length, distal partgently curved, subconical with rounded tip, with small, low
subdistal dorsal flap, with scattered setae (Fig. 15).

–Type species. Paramaja kominatoensis Kubo, 1936, byoriginal designation and monotypy. Gender feminine.

may merge basally (Figs. 8, 9, 13A–C, 14A–E); distal partof G1 relatively long, tip gently bent laterally (Fig. 15A–J);
Japan, South Korea and Taiwan................................................
....................................... Paramaja kominatoensis Kubo, 1936
In adults, pseudorostral horns dorso-ventrally flattened (Figs.
13D–F, 36I–K); carapace dorsal surface relatively less swollen,with numerous small and large distinct tubercles and granules
which are never swollen (Figs. 10–12, 13D–F, 14F–H); distal
part of G1 not as above...........................................................2

2. Adult carapace rounded (Fig. 12); distal part of G1 relatively
shorter, tip in line with rest of G1 (Fig. 15N–U); Philippines,
Lesser Sunda Islands and Solomon Islands..............................
.............................................................Paramaja turgida n. sp.
– Adult carapace ovate (Figs. 10, 11); distal part of G1 relatively
longer, tip gently bent laterally (Fig. 15K–M); Indian Ocean.
................................................Paramaja gibba (Alcock, 1895)

Remarks. Kubo (1936: 361) established a new genus fora new species from Kominato, Japan, with the following
comments: “The present genus stands somewhat intermediate
between the two genera Maja and Majopsis (sic for Maiopsis
Faxon, 1893), and it is easily distinguished from the genus
Maja in lacking any spines on carapace, and from thegenus Majopsis in having no accessory spine on rostralspines, but two spines on distal margin of basal segmentof outer antennae, and fingers of chela not canaliculate
within.” Although none of these characters work at present
to distinguish Paramaja from Maja s. str. or allied genera,
it can still be separated by other features.

Paramaja kominatoensis Kubo, 1936
(Figs. 2B, 8, 9, 13A–C, 14A–E, 15A–J, 36G, H, 39G, H,
41E, 43I–K, 48A–E, 53I, J, 55E, F, 68A)
Paramaja kominatoensis Kubo, 1936: 361, figs. 1, 2.
Maja kominatoensis – Miyake, 1936: 418, pl. 28 figs. 1, 2. – T.
Sakai, 1938: 300, pl. 38 fig. 3. – Serène, 1968: 57. – Serène &Lohavanijaya, 1973: 50 (key). – T. Sakai, 1976: 240, text fig.
128, pl. 84 fig. 2. – Matsuzawa, 1977: pl. 96 fig. 1. – Miyake,
1983: 47, pl. 16 fig. 2. – Ikeda, 1998: 117, pl. 41. – Muraoka,
1998: 27, pl. 5 fig. 1. – Ng et al., 2001: 12. – Maramura &Kosaka, 2003: 34. – Yang et al., 2008: 780. – Ng et al., 2008:
117 (list).
Maja gibba – T. Sakai, 1976: 239 (part), text fig. 127a, b, pl. 84
fig. 1. – Dai et al., 1986: 137, pl. 18(4). – Dai & Yang, 1991:
152, pl. 18(4). – Huang, 1994: 583. – Muraoka, 1998: 27. –Maramura & Kosaka, 2003: 34. – Yang et al., 2008: 780. (notMaja gibba Alcock, 1895).

Paramaja Kubo, 1936, is here resurrected to accommodate
Indo-Pacific species which have a uniformly inflated carapace
which has the dorsal surface covered with granules andtubercles: P. gibba (Alcock, 1895), P. kominatoensis Kubo,
1936, and P. turgida n. sp. The genus is also characterisedby features of its suborbital margin, epistome, male abdomen
and G1 structure.
Paramaja is closest to Alcomaja n. gen., with which it sharesa suite of carapace, antennal, epistomal, third maxilliped,
pereiopodal, thoracic sternal and male abdominal characters.
The single most important character separating Paramaja
from Alcomaja is the form of the G1. In Paramaja, the G1
is very long and mostly straight, the distal part being conical(Fig. 15). In Alcomaja, the G1 is proportionately shorter,
more curved, and part of the dorsal fold bends downwards
to form a distinct ventral projection (Figs. 19, 20). Speciesof Alcomaja also have relatively longer and more slenderpseudorostral spines (Fig. 36L–O) than in Paramaja (Fig.
36G–K). Adults of the two genera are easy to separate,with Paramaja growing to much larger sizes (over 60–70
mm carapace length) while Alcomaja species are smaller.
In Paramaja specimens, even when small (20–30 mm
carapace length), the branchial and cardiac regions areproportionately more inflated and the lateral and branchialspines are relatively shorter (Figs. 10A, B, 11A–C, 12A,
B). Specimens of Alcomaja of comparable size on the other
hand have less inflated branchial and cardiac regions, andthe lateral and branchial spines are stronger and longer(Figs. 16, 17). That said, juvenile female specimens of thetwo species can be difficult to separate.

Material examined. Neotype (here designated): male (56.6
× 40.4 mm) (SMF 47808), None, west of Cape Muroto,
Kochi Prefecture, 33°26.617’N 134°16.75’E, Japan, coll.
K. Matsuzawa, 13 December 1999. Japan – 1 dried male
(62.6 × 56.1 mm) (KPM NH0104298), probably Mimase,Tosa Bay, T. Sakai Collection. — 1 dried male (60.4 × 52.5mm), 1 dried female (61.0 × 54.6 mm) (KPM NH124171),
Minabe town, Wakayama Prefecture, Japan, T. Sakai
Collection. — 2 males (41.0 × 34.6 mm, 53.1 × 46.8 mm),
1 female (63.5 × 58.5 mm) (SMF 47728), Ashizuri, 200
fathoms, coll. RV Taisho Maru, 24 November 1958. — 2
males (50.5 × 44.6 mm, 76.9 × 69.6 mm) (SMF 47738),
Tosa Bay, Kochi Prefecture, T. Sakai collection (TS00320),
coll. 1970s. — 1 female (62.0 × 58.0 mm) (SMF 47810),
None, Kochi Prefecture, 33°26.617’N 134°16.75’E, coll. K.
Matsuzawa, 12 November 1988. — 1 female (52.7 × 47.5mm) (SMF 47736), Tosa Bay, Kochi Prefecture, 190 m,
coll. RV Toyohata-Maru, 7 April 1989. — 1 female (25.7
× 21.0 mm) (SMF 47744), Tosa Bay, Kochi Prefecture,
250–300 m, coll. K. Sakai, 3–14 November 1963. South
Korea – 1 female (53.1 × 40.0 mm) (SMF 47605), Cheju
Island, 33°32.415’N 126°30.783’E, East China Sea, bottom,
coll. M. Matsuo, 7 May 1991. Taiwan – 1 dried male (ca.
60 mm carapace length) (PCM), Tashi, northern Taiwan,coll. K.-X. Li, 2000s. — 1 dried male (73.0 × 66.0 mm,
79.0 mm including pseudorostrum) (PCM, one G1 in ZRC

Key to species of Paramaja Kubo, 1936
1. In adults, pseudorostral horns with cross-section rounded (Figs.
8, 9, 13A–C, 36G, H); carapace dorsal surface usually strongly
swollen, covered with large rounded or sharp tubercles which132

RAFFLES BULLETIN OF ZOOLOGY 2015
Females of P. kominatoensis are similar to males in nonsexual characters except that their pereipods are relatively
shorter and more slender (Figs. 8E, F, 9B versus Figs.
8A–C, 9A, C).

2013.1407), Tashi Port, northern Taiwan, coll. K.-X. Li,from deep-water trawlers, 2000s.
Diagnosis. Adult carapace with branchial and gastricregions rounded; dorsal surface inflated to strongly inflated,covered by large rounded tubercles and/or wart-like or
sharp granules, some which may merge basally; those on
median row especially large, forming low rounded or sharp
crest (Figs. 8, 9, 13A–C, 14A–E). Adult pseudorostralspines gently curved outwards to almost straight, strongly
diverging; cross-section circular (Figs. 8, 9, 13A–C, 36G,
H). Supraorbital eave relatively narrow, granulated; antorbital
spine triangular, sharp; intercalated spine irregular, slightly
shorter than supraorbital spine; postorbital spine relatively
wide, gently curved anteriorly, triangular with some granules
at base of anterior margin; hepatic area prominently inflatedwith several large sharp tubercles (Figs. 8, 9, 36G, H).
Adult male anterior thoracic sternum relatively broad (Fig.
48A–E); lateral margins of anterior edge of male sternoabdominal cavity almost straight or gently curved (Fig.
48A–E). Distal part of G1 relatively long, tip gently bent
laterally (Fig. 15A–J).

Paramaja kominatoensis is close to P. gibba s. str. (and P.
turgida n. sp.), and some authors (e.g., Griffin & Tranter,1986) have suggested that they may be synonymous. The
most distinctive feature of P. kominatoensis is the presenceof large wart-like granules on the median part of thecarapace, especially on the gastric, cardiac, intestinal andinner branchial regions (Ikeda, 1998: pl. 41, fig. 1a–f; present
Figs. 8B, C, 13A–C, 14A–C); but as discussed above, thesecharacters vary to some degree and cannot be relied on.
In P. gibba s. str. and P. turgida n. sp., these granules arealways relatively smaller, sharper and well separated fromeach other. Paramaja kominatoensis also almost always has
a proportionately more strongly inflated carapace, especiallywhen viewed frontally or laterally (Ikeda, 1998: pl. 41,
figs. 1e, 1f; present Figs. 13A–C, 14A–D), compared to P.
gibba s. str. or P. turgida n. sp. (Figs. 13D–F, 14F–H). This
“inflatedness” is attenuated by the large wart-like granulesthat may be present on the carapace in P. kominatoensis(Figs. 13A–C, 14A–C). In addition, in adult Paramaja
kominatoensis, the pseudorostral spines are circular incross-section (Figs. 13A–C, 36G, H) (versus dorso-ventrally
flattened in P. gibba and P. turgida, Figs. 13D–F, 36I–K).

Remarks. Kubo (1936) described the species on the basis ofa single large male 63.0 by 57.0 mm collected from a depthof 370 m from off Kominato in Chiba, Japan. The extant
material of Kubo is in the Tokyo University of Fisheries,
and a list of taxa present in this collection was publishedby Takeda & Ueshima (2006). A recent search there again
failed to find his type specimen (T. Komai, pers. comm.).
In view of the variation in carapace form observed in theJapanese material, and its close similarity with P. gibba(Alcock, 1895) and P. turgida n. sp., we have decided to
select a neotype for Paramaja kominatoensis Kubo, 1936.
The neotype selected is a male 56.6 × 40.4 mm in the SMF
collected from None in Kochi Prefecture, Japan. Although
it not the largest specimen we have examined, it is the mostcomplete and is an adult male.

The G1 structure of P. kominatoensis closely resembles
that of P. gibba, with the tips of the distal part gently bent
laterally (Fig. 15A–J). However, the distal part of the G1
in P. kominatoensis is proportionately longer (Fig. 15A,
D, H) compared to that of P. gibba (Fig. 15L). Comparedto the G1 of P. kominatoensis, that of P. turgida n. sp. is
proportionately shorter with the tip in line with the rest of
the structure (Fig. 15N–U) (versus longer with the tip gently
bent laterally, Fig. 15A–J).
The male specimen from Japan (KPM NH0104298) is one of
the two males listed by T. Sakai (1976). Although the attachedlabel does not carry any locality data, T. Sakai (1976: 240)
noted he had only two males, both from Tosa Bay. Althoughgenerally regarded as a Japanese endemic, the two dried
specimens in the Peikuan Crab Museum were supposedlycollected from Tashi Fish Port just off northeastern Taiwan.
There is also a record of this species from Taiwan (Chou etal., 1999) but the specimen cannot be located so we cannot
confirm its identity. The species has also been listed from
East China Seas by Yang et al. (2008). It is recorded here
for the first time from South Korea.

Kubo’s (1936: fig. 1) (Fig. 8A) figure of P. kominatoensis
shows a specimen with a strongly inflated carapace but the
granules on it do not appear to be as swollen, large or wartlike as those illustrated by T. Sakai (1976) or Ikeda (1998)(see present Figs. 8B, C, 68A). The convexity and size ofthe granules on the carapace varies, as the excellent seriesof specimens in SMF demonstrate. Smaller specimens all
have relatively less swollen carapaces, and the granules areproportionately smaller and sharper (e.g., Fig. 9D, E). Larger
specimens vary, with some having the sharper granules (e.g.,
Figs. 9A–C), some with wart-like granules and having a moreswollen carapace (e.g., Figs. 8B, C, E, F), and a few with
granules and tubercles intermediate in condition (e.g., Figs.
8D) (like in Kubo’s type, Fig. 8A). The lateral margins ofanterior edge of the male sterno-abdominal cavity are usuallystraight or almost so in P. kominatoensis (Fig. 48A, B) but
can also be gently curved (Fig. 48C–E). This variation in
the sterno-abdominal cavity does not seem to be associatedwith size. The cross-section of the pseudorostral spines inall thes specimens, however, is circular (Fig. 13A–C), notdorsoventrally flattened like in P. turgida n. sp. (Fig. 13D–F).

The color in life is cream with a band of longitudinal big
granules bright red (Fig. 68A) (see also Ikeda, 1998: pl. 41).
Paramaja gibba (Alcock, 1895)(Figs. 10, 11, 13D, 14F, G, 15K–M, 36I, 39I, J, 41F, 43L,
M, 48F, G, 52D, 53K, 55H, 68B, C)
Maia gibba Alcock, 1895: 239, pl. 4, fig. 5, 5a. – Alcock &Anderson, 1898: pl. 21 fig. 5. – Alcock, 1899: 56.

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Indian Ocean, 12°17.7’N 53°08.9’E 12°21.8’N 53°05.6’E,
depth 375–380 m, coll. RV Vitiaz, trawl, 27 October 1988.
— 1 female (67.7 × 63.2 mm) (SIO), cruise 17, station 2804,
Saya de Malha, Indian Ocean, 11°06’S 62°14’E–11°09’S
62°13’E, 235-230 m, coll. RV Vitiaz, 7 January 1989.

Maja gibba – Serène, 1968: 57. – Serène & Lohavanijaya, 1973:
50 (not specimen figured Pl. IX B = Maja compressipes). – T.
Sakai, 1976: 239 (part). – Griffin & Tranter, 1986: 216 (part).
– Poore et al., 2008: 62. – Ng et al., 2008: 117 (list).

Material examined. Lectotype (here designated): juvenile
female (25.1 × 21.2 mm) (NHM 1896.5.14.9), code zoo20121157T, Andaman Sea, 250 fms, coll. Investigator. Others:
India – 1 male (79.5 × 77.9 mm) (ZRC 2013.1232), Gulf
of Bengal, V. Kakati, 2011. Thailand – 1 male (65.0 × 59.8mm) (NSMT-Cr 16618), Andaman Sea, 400 m, coll. Y.
Tamura, 5 June 1987. Australia – 1 juvenile female (33.9
× 29.4 mm) (NMV J54316), off Point Cloates, 22°37.07’S
113°29.03’E – 22°37.2’S et 113°28.57’E, Western Australia,
355–382 m, coll. G. Poore, CSIRO RV Southern Surveyor,
9 December 2005. — 1 female (46.7 × 41.6 mm) (NMV
J61055), Kulurnburu L29 Transect, 13°15.9’S 123°22.45’E –13°16.35’S 123°21.4’E, northwestern Australia, 390–394 m,
beam trawl, coll. D. Bray, CSIRO RV Southern Surveyor, 6July 2007. Madagascar – 1 male (24.6 × 20.9 mm) (MNHNIU-2010-61), station CP 3189, 12°30’S 48°18’E, 346–376 m,
coll. MIRIKY, 27 June 2009. — 1 ovigerous female (47.6 ×
44.1 mm) (ZRC 2013.1405, ex MNHN-IU-2010-508), station
CP 3192, 578–580 m, 12°26’S-48°13’E, coll. MIRIKY,
27 June 2009. — 1 ovigerous female (47.1 × 42.6 mm)(MNHN-IU-2010-507), station DW 3198, 440–447 m,
12°05’S-48°59’E, coll. MIRIKY, 28 June 2009. — 1 male(58.6 × 52.2 mm) (ZRC 2013.1406, ex MNHN-IU-2010-63),
station CP 3183, 420-436 m, 12°38’S 48°14’E, coll. MIRIKY,
26 June 2009. — 1 male (61.3 × 54.8 mm), 3 ovigerous
females (45.1 × 40.6 mm, 46.3 × 43.3 mm, 45.0 × 40.7mm) (MNHN-IU-2010-467), station CP 3183, 420–436 m,
12°38’S-48°14’E, coll. MIRIKY, 26 June 2009. – 1 male(37.8 × 33.1 mm), 1 ovigerous female (47.1 × 44.3 mm), 1
female (24.4 × 20.3 mm) (MNHN-IU-2010-510), CP 3223,
430–488 m, 12°46’S-48°11’E, coll. MIRIKY, 2 July 2009. —1 ovigerous female (48.7 × 44.5 mm) (MNHN-IU-2010-506),
station DW 3212, 367–369 m, coll. MIRIKY, 30 June 2009.
— 2 males (60.4 × 53.4 mm, 62.8 × 57.8 mm), 1 ovigerous
female (47.7 × 42.2 mm) (MNHN-IU-2010-926), station DW
3217, 391–438 m, 12°33’S-47°56’E, coll. MIRIKY, 30 June
2009. — 1 ovigerous female (44.6 × 42.2 mm), 1 juvenile
male (22.4 × 18.8 mm) (MNHN-IU-2010-927), station CP
3190, 415–416 m, 12°31’S-48°15’E, coll. MIRIKY, 27 June
2009. — 1 male (50.8 × 45.7 mm) (MNHN-IU-2010-509)
MIRIKY, CP3289, 332–379 m, 14°29’S-47°26’E, 14 July
2009. — 1 ovigerous female (43.5 × 40.1 mm) (MNHNIU-2010-58) (specimen barcoded), station CP 3183, 420–436
m, 12°38’S-48°14’E, coll. MIRIKY, 26 June 2009. — 1
female (20.9 × 16.9 mm) (MNHN-IU-2010-931), station
CP 3293, 268–408 m, 14°30’S-47°26’E, coll. MIRIKY, 14
July 2009. — 1 juvenile male (16.4 × 12.5 mm) (MNHNIU-2010-55), station CP 3223, 430–488 m, 12°46’S-48°11’E,
coll. MIRIKY, 2 July 2009. — 1 juvenile male (10.6 × 8.1mm) (MNHN-IU-2010-54) (specimen barcoded), station
CP 3223, 430–488 m, 12°46’S 48°11’E, coll. MIRIKY,
2 July 2009. — 1 juvenile male (9.9 × 8.0 mm) (MNHNIU-2010-56), station CP 3184, 492–524 m, 12°40’S 48°12’E,
coll. MIRIKY, 26 June 2009. Indian Ocean – 1 male (64.7 ×60.7 mm) (SIO), cruise 17, station 2560, near Socotra Island,

Diagnosis. Adult carapace with branchial and gastric regions
rounded; dorsal surface convex, covered by numerous
rounded and sharp granules, those on median row larger,
tuberculate, forming low raised crest (Figs. 10, 11, 13D,
14F, G). Adult pseudorostral spines usually almost straight,
gently diverging, appears dorso-ventrally flattened (Figs. 10,
11, 13D, 36I, J). Supraorbital eave large, curved with very
long, sharp antorbital spine; 1 strong postorbital tooth; 1
intercalated spine large nearly as long as supraorbital spine;
short hepatic spine; 3 spines on lateral margin (Figs. 10,
11, 36I, J). Adult male anterior thoracic sternum relatively
broad (Fig. 48F, G); lateral margins of anterior edge ofmale sterno-abdominal cavity gently curved (Fig. 48F, G).
Distal part of G1 relatively shorter, tip gently bent laterally(Fig. 15K-M).
Remarks. Maja gibba was described by Alcock (1895:
239, pl. 4 fig. 5) from the Andaman Sea (463 m) on thebasis of two specimens, supposedly one male 32.0 mm(carapace length, with spines) and one female 41.0 mm, bothwith relatively short ambulatory legs with all the articles
prominently setose. The small figure of Alcock (see also
Alcock & Anderson, 1898: pl. 21 fig. 5; present Fig. 10B)
shows a posterior carapace margin lined with short spines,
a relatively long ocular peduncle; four spines on the lateralcarapace margin and the branchial regions separated bydeep grooves. The NHM specimen (Fig. 10A) was labelled
as a syntype and agrees with the measurements of Alcock(1895) of the male (without the pseudorostrum). Tune Sakai(1976: 240) also noted that he examined this “cotype” malespecimen in NHM. However, this NHM specimen is actuallya juvenile female, but since the abdomen is subrectangularand resembles that of a male, he (and T. Sakai) probably
accidentally misidentified the sex of the specimen. It is here
designated as the lectotype of Maja gibba Alcock, 1895
(Fig. 10A).
Interestingly, the species has not been reported from the
Bay of Bengal area since its original description. Specimensthat have been referred to “Maja gibba” from the Pacific byvarious authors after Alcock (1895) should be referred toother taxa. Serène & Lohavanijaya (1973: 50) provided a key
to the genus and noted that their Maja gibba has spines on
the median row of the carapace. The specimen they figured
from the South China Sea, however, has a longitudinally
ovoid carapace that is covered with flattened tubercles andshort, densely setose ambulatory legs which have a large
flattened carpus; features diagnostic of Maja compressipes(Miers, 1879) (here referred to Ovimaja n. gen.).
Tune Sakai (1976: 239) reported the species from Japan andhis colour plate depicted a male 54 mm in carapace lengthwith short setose ambulatory legs (T. Sakai, 1976: pl. 84, fig.
1) with his text figure showing a specimen with three lateral
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spines and two posterior carapace margin spines (T. Sakai,
1976: 240, text-fig. 127). Tune Sakai (1976) commented that
he had examined a type specimen in the NHM of Maja gibba
as well. Griffin (1976: 200) recorded a male as “Maja gibba”
from between Negros and Siquijor Islands in Philippines;
while small specimens were recorded from off Bali and theSouth China Sea by Griffin & Tranter (1986: 216). In theirkeys to the genus, T. Sakai (1976: 236) stated that for his“M. gibba” and M. kominatoensis, the “carapace without
median spine”. This was also cited by Griffin & Tranter(1986: 210) in their key to Maja. Their specimens are here
provisionally referred to P. turgida.

spines and granules, they never reach the degree seen in
P. kominatoensis, in which the granules can become large
and swollen, with those along the median row fused basally(Fig. 8B, C). Nor does the degree of inflation of the carapace
vary to the degree seen in P. kominatoensis. As discussed
earlier, the carapace of P. kominatoensis is almost always
more swollen and inflated (Figs. 13A–C, 14A–D) than P.
gibba or P. turgida (Fig. 13D–F, 14F–H), and this is obvious
from lateral and frontal views. The material of P. turgida
also demonstrates that the form of the pseudorostrum, maleabdomen and G1 structure are relatively constant characters.
Understanding the degree of variation in P. turgida alsomeans that we are confident that the material from the Indian
Ocean, Japan, Taiwan and Philippines belong to three species.
Paramaja kominatoensis is clearly a distinct species. The
Indian Ocean appears to have only one species, P. gibba
s. str. The Philippine material is referred to a new species,
Paramaja turgida. However, most of the characters discussed
and regarded as of species-identification value are useful
only for adults. Juvenile specimens of these species about
30–40 mm in carapace length cannot be easily separated.

Tune Sakai (1976: 239–240) had small and large specimensof “Maja gibba” from Japan, East China Sea and Philippinesand commented that the spines on the carapace median row
and lateral margins are relatively sharper and longer in smaller
specimens and become very short in large adults, with the
granules on the surface also becoming more rounded. Griffin& Tranter (1986) discussed the possible conspecificity of what
they regarded as Maja gibba Alcock, 1895, with Paramaja
kominatoensis Kubo, 1936, noting that the large wart-likegranules on the carapace supposedly diagnostic of the latter
species may be variable once a good series of specimens
become available. As discussed under P. kominatoensis, thisis indeed the case, but the species can still be separated byother characters.

See also remarks for P. turgida for discussion of differenceswith P. gibba.
Paramaja turgida n. sp.
(Figs. 6H, 12, 13E, F, 14H, 15N–U, 36J, K, 39K, 43N–P,
48H, I, 53L, 55I, J, 68D–F)

The present study has an excellent series of specimensof various sizes and sexes of P. turgida n. sp. from thePhilippines and confirms T. Sakai’s (1976) observationsof changes in the strength and sharpness of the carapace
spines and granules – large male and female specimens
have almost indiscernible median and lateral spines, and the
granules are relatively lower and eroded. The specimens of
P. turgida from the Philippines are consistent in how theyvary in carapace and ambulatory leg form and armature; andspecimens of similar sizes do not vary much. Comparing thegood series of specimens of P. turgida from the Philippinesof similar sex and sizes, several other variation patterns areapparent. The eyes are proportionately less elongated inlarge specimens (also true for P. gibba). The proportions of
the ambulatory legs, especially the first two, become very
long and elongated in large adult males, with the propodus
becoming very elongate, almost devoid of setae; and thechelipeds also become enlarged with the palm inflated(Figs. 12, 53L). This is also true for P. gibba (Figs. 10, 11,53K). The rostrums in smaller specimens of P. gibba andP. turgida form a more distinct V, with the pseudorostralspines gently curved outwards (Figs. 10A, 11A–C, E,
12A–C, 36K). In large specimens, the rostrums becomestraighter and less curved (Figs. 9C, D, 11D, 12E, 36, I, J).
While the lateral margins of male abdominal somites 4–6
are parallel in adult males of P. gibba and P. turgida, withthe abdomen rectangular (Fig. 48G, I), the lateral margins
become gently convex in very large males and the abdomen
becomes slightly oval (Fig. 48F, H).

Maja gibba – Griffin, 1976: 200. – T. Sakai, 1976: 239 (part). –Griffin & Tranter, 1986: 216 (part). (not Maja gibba Alcock,1895).
Maja kominatoensis – Ng et al., 2008: 123, Fig. 90. (not Paramaja
kominatoensis Kubo, 1936).

Material examined. Holotype: male (74.1 × 66.8 mm)(NMCR), Balicasag Island, Panglao, Bohol, Philippines,
200–300 m, coll. fishermen with tangle nets, 28 November
2001. Paratypes: Philippines – 1 male (71.4 × 64.4 mm)(NSMT-Cr 22329), 1 male (55.4 × 48.4 mm), 1 ovigerous
female (62.9 × 57.2 mm) (AM, ex ZRC 2001.0615), samedata as holotype. — 1 young male (26.2 × 21.3 mm)(ZRC 2001.0616b), Balicasag Island, Panglao, Bohol,
coll. fishermen with tangle nets, 28 November 2001. — 1
male (69.2 × 64.0 mm), 1 ovigerous female (47.1 × 41.1mm) (ZRC 2013.1185), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, June 2002.
— 1 ovigerous female (dried) (58.4 × 53.4 mm) (ZRC
2009.0972), Banacon Island, Bohol, coll. local fisherman,
2000. — 3 females (NSMT-Cr 15388), Balicasag Island,
Panglao, Bohol, coll. local fisherman via M. Takeda & H.
Komatsu, February 2003. — 2 males (34.3 × 29.2 mm, 56.4
× 49.7 mm), 1 female (60.1 × 55.6 mm, with anemone)(ZRC 2013.1239), Balicasag, Panglao, Bohol, 200–300 m,
coll. local fishermen with tangle nets, January 2004. — 1
female (61.3 × 56.6 mm) (ZRC 2013.1240), Balicasag,
Panglao, Bohol, 200–300 m, coll. local fishermen with tangle
nets, February 2004. — 2 males (70.6 × 62.2 mm, 57.8 ×
51.5 mm), 2 ovigerous females (60.2 × 53.1 mm, 62.2 ×
55.2 mm) (ZRC 2012.1203), Balicasag, Panglao, Bohol,

The series of specimens of P. turgida show that while there
is variation in the sharpness and length of the carapace135

Ng & Richer de Forges: Revision of the spider crab genus Maja

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.

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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.

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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.

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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 × 40.6mm) (MNHN-IU-2010-467), Madagascar; G, female (46.7 × 41.6 mm) (NMV J61055), Australia.

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Fig. 12. General habitus, Paramaja turgida n. sp. A, paratype male (26.2 × 21.3 mm) (ZRC 2001.0616b); B, paratype female (33.5 × 29.0mm) (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.

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Fig. 13. Frontal views of carapaces, Paramaja species. A, P. kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan;
B, P. kominatoensis, dried male (60.4 × 52.5 mm) (KPM NH124171), Japan; C, P. kominatoensis, dried male (73.0 × 66.0 mm) (PCM),
Taiwan; D, P. gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; E, P. turgida n. sp., holotype male (74.1 × 66.8 mm)(NMCR), Philippines; F, P. turgida n. sp., paratype male (67.9 × 60.5 mm) (MNHN), Philippines.

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Fig. 14. Lateral views of carapaces, Paramaja species. A, P. kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298), Japan; B,
P. kominatoensis, female (62.0 × 58.0 mm) (SMF 47810), Japan; C, P. kominatoensis, female (63.5 × 58.5 mm) (SMF 47728), Japan;
D, P. kominatoensis, male (50.5 × 44.6 mm) (SMF 47738), Japan; E, P. kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan[laterally transposed to match other figures]; F, P. gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; G, P. gibba, female(67.7 × 63.2 mm) (SIO), Indian Ocean; H, P. turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines.

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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.

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200–300 m, coll. local fishermen with tangle nets, March
2004. — 1 female (61.9 × 58.4 mm) (ZRC 2013.1235),
northwest Panglao, Bohol, 120–160 m, tangle nets, coll. J.
Arbasto, July 2004–May 2005. — 1 male (70.4 × 63.9 mm)(ZRC 2012.1201), station CP 2332, off Pamilacan Island,
Bohol Sea, 9°38.8’N 123°45.9’E, coll. PANGLAO 2005, 22
May 2005. — 1 male (70.9 × 62.2 mm) (ZRC 2012.1202),
station CP 2372, Bohol/Sulu Seas, 9°31.4’N 124°00.6’E,
PANGLAO 2005, 24 May 2005. — 1 male (70.5 × 63.1mm) (ZRC 2012.1204), station CP 2397, off Aligbay Island,
Maribojoc Bay, Bohol Sea, 9°34.9’N 123°41.7’E, coll.
PANGLAO 2005, 31 May 2005. — 1 male (36.4 × 31.2mm) (MNHN), station CP 2406, rock-sand-coral substrate,
9°38.8’N 123°46.5’E – 9°39.7’N 123°46.6’E, 331–334
m, Bohol Sea, coll. PANGLAO 2005, 1 June 2005. — 1
female (59.6 × 54.5 mm) (MNHN), station CP 2383, sandy
substrate, 8°43.1’N 123°19.2’E – 8°44.0’N 123°18.7’E,
253–338 m, Bohol Sea, coll. PANGLAO 2005, 29 May
2005. — 1 male (28.0 × 22.5 mm) (ZRC 2013.1237), station
CP 2331, muddy substrate, 9°37.4’N 123°45.9’E – 9°38.9’N
123°46.9’E, 268.2–273 m, Bohol Sea, coll. PANGLAO
2005, 22 May 2005. — 1 female (31.0 × 26.5 mm) (ZRC
2013.1231), station CP 2341, sandy-muddy substrate,
9°26.1’N 123°48.4’E – 9°24.9’N 123°49.3’E, 381–544
m, Bohol Sea, coll. PANGLAO 2005, 23 May 2005. — 1
female (62.6 × 57.2 mm) (ZRC 2013.1241), station CP 2363,
sandy substrate, 9°0.1’N 123°26.5’E – 9°0.3’N 123°25.8’E,
439–452 m, Bohol Sea, coll. PANGLAO 2005, 26 May
2005. — 1 female (51.0 × 45.1 mm) (ZRC 2013.1236),
station CP 2392, off Balicasag Island, sandy-muddy substrate,
9°30.1’N 123°43.4’E – 9°29.5’N 123°42.6’E, 242–313 m,
Bohol Sea, coll. PANGLAO 2005, 30 May 2005. — 1 female(50.5 × 44.8 mm, with rhizocephalan, photographed) (ZRC
2013.1236), station CP 2331, muddy substrate, 9°37.4’N
123°45.9’E – 9°38.9’N 123°46.9’E, 268–273 m, Bohol Sea,
coll. PANGLAO 2005, 22 May 2005. — 1 female (33.5 ×
29.0 mm) (ZRC 2013.1230), station CP 2396; sandy substrate,
9°37.1N 123°445’E – 9°36.7N 123°42.9’E, 467–609 m,
Bohol Sea, coll. PANGLAO 2005, 31 May 2005. — 1 female(ZRC 2013.1234), station CP2405, Maribojoc Bay, Panglao,
9°39.0’N 123°46.1’E, 387–453 m, coll. PANGLAO 2005,
1 June 2005. — 1 male (67.9 × 60.5 mm) (MNHN), station
CC 2743, eastern coast of Luzon Island, coll. AURORA
2007, 2 June 2007. — 1 female (42.2 × 37.6 mm) (MNHNIU-2013-4055), station CP 53, 13°59’N 120°18’E, 215–216
m, coll. RV Coriolis, MUSORSTOM 2, 27 November 1980.
— 1 male (33.7 × 28.4 mm) (USNM 48508), station 5403,
between Leyte and Cebu, 182 fathoms, coll. RV Albatross,
16 March 1909. Non-type: Philippines –fragments (USNM
48509), station 5536, 279 fathoms, coll. RV Albatross, 19
August 1909. Solomon Islands – 1 male (25.1 × 19.6 mm)(MNHN-IU-2013-4051), station CP 1851, 10°28’S 162°00’E,
297–350 m, coll. RV Alis, SALOMON 1, 6 October 2001.

Supraorbital eave narrow, granulated, with antorbital spine
triangular, sharp; intercalated spine irregular, slightly shorter
than supraorbital spine; postorbital spine wide, pointing
anteriorly, triangular with some granules at base of anterior
margin; hepatic area inflated with several granules, largest
spiniform, pointing outwards (Figs. 12, 13E, F, 36J, K). Adult
male anterior thoracic sternum relatively narrow (Fig. 48H,I); lateral margins of anterior edge of male sterno-abdominal
cavity straight (Fig. 48H, I). Distal part of G1 relatively
shorter, tip not bent, in line with rest of G1 (Fig. 15N–U).
Etymology. From the Latin “turgidus” for swollen, alluding
to the carapace appearance of the species.
Remarks. Paramaja turgida n. sp. is closest to P. gibba,
but can be separated by its proportionately narrower maleanterior thoracic sternum (Fig. 48H, I) (versus relatively
broader in P. gibba, Fig. 48F, G) and the distal part of theG1 is relatively shorter with the tip in line with the rest of
the structure (Fig. 15N–U) (versus relatively longer with the
tip gently bent laterally in P. gibba, Fig. 15K–M).
The chelipeds of adult females of P. turgida are slender andnot inflated, while the ambulatory legs are distinctly shorterand more setose (Fig. 12B, D). The smallest ovigerous female
measured 47.1 by 41.1 mm (ZRC 2013.1185).
A young male specimen from the Solomon Islands (25.1 ×
19.6 mm, MNHN-IU-2013-4051) (Fig. 12F) is tentatively
referred to P. turgida. The G1 resembles that of P. turgida, but
at this size, it is not easy to separate from P. kominatoensis.
The less inflated carapace shape of the specimen is closest
to P. turgida and as such, we provisionally refer it here.
Adult males will need to be collected to be certain about
their identity as the Solomon Islands is some distance fromthe Philippines.
See also discussion for P. gibba and P. kominatoensis.
Alcomaja n. gen.
Diagnosis. Carapace ovate; dorsal surface convex, coveredby granules or tubercles; gastric and branchial regionsdelimited by distinct grooves (Figs. 16, 17). Intestinal region
without distinct median spine (Figs. 16, 17). Pseudorostralspines long, diverging, forming a V, (Figs. 18, 36L–O).
Supraorbital eave with anterior part longitudinally narrow,rectangular, not prominently expanded; antorbital spine sharp(Figs. 18, 36L–O). Intercalated spine distinct, separated
from supraorbital eave and postorbital spine by distinct
gaps; postorbital spine strong, lobiform; hepatic region
with 1 strong spine, much shorter than postorbital spine;
1 or more smaller spines below (Figs. 18, 36L–O). Lateralcarapace margin with 3 spines and numerous tubercles or
granules, 1 branchial spine or tubercle usually discernible,
may be low in large specimens (Figs. 16, 17). Posterior
carapace margin with 2 short median spines (Figs. 16, 17).
Eyes relatively long, slender, with ovoid cornea (Figs. 18,
36L–O). Antennal flagellum short, slender. Basal antennal
article longer than broad, rectangular; surface with several

Diagnosis. Adult carapace with branchial and gastricregions ovate; dorsal surface convex, covered by numerous
rounded and sharp granules, those on median row larger,
tuberculate, forming low crest (Figs. 12, 13E, F, 14H).
Adult pseudorostral spines almost straight, gently diverging,appears dorso-ventrally flattened (Figs. 12, 13E, F, 36J, K).
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Key to species of Alcomaja n. gen.

tubercles or granules, with 2 spines distally; inner and outer
lateral margins may have low granules or lobes, or entire;
proximal outer angle rounded; antero-external crested rim ofantennular fossa touches does not substantially overlap basal
antennal article (Fig. 39L–O). Epistome as wide as long,anterior margin with low lobes or low swellings; posterior
margin composed of 4 rectangular plates separated by shallow
fissures (Figs. 39L–O, 41G, H). Suborbital margin separated
from basal antennal article by short fissure, confluent with
margin of postorbital tooth (Fig. 39L–O). Outer surfaceof third maxilliped covered by very short setae in adults;
ischium subrectangular, distinctly longer than broad; posteroexternal angle of merus relatively broad, “inserted” into
shallower concavity on outer margin of ischium; anterointernal part of ischium rounded, auriculiform (Fig. 45A–H).
Male chelipeds relatively short in adult males, surfaces ofmerus and carpus almost smooth to covered with very small
granules, low longitudinal ridges with low tubercles; carpus
short, with low, uneven longitudinal ridge; propodus of palm
elongated, curved, smooth, without lateral cristae in young,short, inflated in adults, palm longer than fingers; fingers
long, slender, gently curved, with distinct basal gape whenclosed (Figs. 16, 17, 53M–P). Ambulatory legs relatively
long, slender; merus without dorsal subdistal spine; dactylus
elongate, curved, covered with long setae when young
except for corneous tip, almost completely smooth in adults(Figs. 16, 17, 55K–N). Thoracic sternum wide; surfaces ofsomites 5–8 with numerous prominent rounded tubercles
and granules; sternites 3 and 4 distinctly depressed; margin
between sternites 2 and 3 demarcated by deep notch; anterior
margin of sterno-abdominal cavity not forming completerim (Figs. 49, 52E–G). Male abdomen subrectangular, with
6 free somites and telson; somites 3 and 4 subequal to orslightly wider than somites 5, 6 and telson (Fig. 49). Male
press-button abdominal locking mechanism submedian inposition on sterno-abdominal cavity (Fig. 52E–G). Female
abdomen dome-shaped, covering most of thoracic sternum.
G1 long, slender, gently curved, distal part subconicalwith rounded tip, dorsal part folding downwards to form
ventral projection, with scattered setae on ventral surface
and numerous short setae on dorsal surface (Figs. 19, 20).

1. Dorsal surface of carapace and ambulatory legs densely covered
with long, stiff setae (Fig. 17F); G1 almost straight with distinct
ventral fold (Fig. 20R–T); Philippines......................................
..........................................................Alcomaja desmondi n. sp.
– Dorsal surface of carapace and ambulatory legs covered with
scattered short, hooked setae, sometimes dense; G1 curved....
..................................................................................................2
2. Dorsal surface of carapace covered with small rounded granules,
lateral and branchial spines strong even as adults, with 2 hepatic
spines; adult ambulatory legs relatively short, with first leg ofadult males not substantially elongated (Figs. 17A–E)..........3– Dorsal surface of carapace covered with small and large granules,
sometimes sharp, lateral and branchial spines distinct but low
to absent in adults, with 2 hepatic spines; adult ambulatory
legs long, with first leg of adult males prominently elongated(Figs. 16A–E)...........................................................................5
3. Carapace relatively more elongate; pseudorostral spines long(Figs. 17E, 36O); G1 gently curved with distal part gently
curved downwards (Fig. 20O–Q); Madagascar........................
...............................................................Alcomaja miriky n. sp.
– Carapace relatively broad; pseudorostral spines relatively shorter(Figs. 17A–D, 36M, N); G1 similar to above or otherwise...44. Intercalated spine acutely triangular (Fig. 36M); G1 strongly
curved, with distal part gently curved downwards (Fig. 20A–J);
Japan and Taiwan.......................................................................
..............................Alcomaja nagashimaensis (T. Sakai, 1969)– Intercalated spine basally subquadrate (Fig. 36N); G1 gently
curved, with distal part gently curved upwards (Fig. 20K–M);
Solomon Islands.................................... Alcomaja latens n. sp.
5. Dorsal surface of carapace with distinct small and large sharp
and rounded granules; intercalated tooth triangular (Figs. 16D,
E,18A, B); G1 gently curved, distal part relatively short (Fig.
19G); Moluccas and Western Australia....................................
.......................Alcomaja confragosa (Griffin & Tranter, 1986)– Dorsal surface of carapace as above or otherwise (Figs. 16A–C,
18C–F); intercalated tooth subtruncate basally (Fig. 18C–F):
G1 gently or strongly curved...................................................6
6. Dorsal surface of carapace with distinct small and large
sharp, rounded granules (Figs. 16B, C, 18C, D); pseudorostralspines with rounded cross-section (Fig. 18C, D); ischium ofthird maxilliped distinctly granular (Fig. 45B, C); G1 gently
curved, distal part straight (Fig. 19D–F); South China Sea andPhilippines................Alcomaja gracilipes (Chen & Ng, 1999)– Dorsal surface of carapace with small and large rounded
granules, some of which merge basally (Figs. 16A, 18E, F);
pseudorostral spines dorsoventrally flattened (Fig. 18E, F);
ischium of third maxilliped smooth (Fig. 45A); G1 strongly
curved, distal part curved upwards (Fig. 19A–C); Philippines............................................................ Alcomaja irrorata n. sp.

Type species. Alcomaja irrorata n. sp. by present designation.
Etymology. The genus is named after Alfred William Alcock.
The name is an arbitrary combination of his family namewith Maja. Gender feminine.
Remarks. Alcomaja n. gen. is established to accomodate
A. irrorata n. sp., Maja gracilipes Chen & Ng, 1999, M.
confragosa Griffin & Tranter, 1986, M. nagashimaensis T.
Sakai, 1969, A. desmondi n. sp., A. miriky n. sp., and A.
latens n. sp.

Alcomaja irrorata n. sp.
(Figs. 2C, 16A, 18E, F, 19A–C, 39L, 41G, 45A, 49A,
52E, 53M, 55K, L)

In the form of the suborbital margin, basal antennal article,
epistome, third maxilliped, pereiopods, thoracic sternum
and male abdomen, Alcomaja is most similar to Paramaja
Kubo, 1936. The differences have been discussed under theremarks for that genus.

Material examined. Holotype: male (52.6 × 45.7 mm)(NMCR), Balicasag, Panglao, Bohol, Philippines, 200–300 m,
coll. local fishermen with tangle nets, June 2002. Paratypes:
Philippines – 1 male (45.5 × 39.5 mm, with rhizocephala),
6 males (47.7 × 43.1 mm, 48.4 × 42.2 mm, 46.7 × 41.5 mm,
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Remarks. In general appearance, A. irrorata n. sp. is closest
to A. gracilipes (Chen & Ng, 1999), described from theSouth China but is now also recorded from the Philippines.
However, there are several key differences between thetwo species. Most significantly, while the G1 structures aresuperficially similar, it is more strongly curved in A. irrorata(Fig. 19A–C) with that of A. gracilipes relatively straighter
along the median part (Fig. 19D–F). This difference isconsistent even for smaller specimens. In addition, the outersurface of the ischium of the third maxilliped is smooth oralmost so in A. irrorata (Fig. 45A) (distinctly granulous in
A. gracilipes, Fig. 45B, C); the pseudorostral spines in A.
irrorata is relatively stouter with the cross-section more
dorso-ventrally flattened (Fig. 18E, F) (relatively more
slender with a rounded cross-section in A. gracilipes, Fig.
18C, D); the postorbital spine is usually relatively widerin A. irrorata (Fig. 8E, F) (more slender in A. gracilipes,
Fig. 18C, D); and the ambulatory legs are relatively longer
in adult A. irrorata than in A. gracilipes (Fig. 16A versus
Fig. 16B, C).

44.7 × 39.1 mm, 44.2 × 38.0 mm, 44.7 × 38.4 mm), 2 young
males (34.4 × 29.8 mm, 28.7 × 23.9 mm), 2 young females(31.5 × 26.9 mm, 29.7 × 26.1 mm), 10 ovigerous females(43.7 × 38.2 mm, 44.1 × 38.7 mm, 41.1 × 36.9 mm, 44.1
× 36.9 mm, 44.1 × 38.2 mm, 43.7 × 39.2 mm, 45.1 × 40.8mm, 44.0 × 37.6 mm, 44.1 × 39.2 mm, 41.6 × 36.6 mm),
1 male (28.1 × 23.6 mm, with bopyrid) (ZRC 2013.1229),
Balicasag, Panglao, Bohol, Philippines, 200–300 m, coll.
local fishermen with tangle nets, June 2002. — 2 ovigerous
females (42.6 × 37.4 mm, 43.0 × 38.0 mm) (USNM, ex
ZRC 2001.0434), Balicasag, Panglao, Bohol, 200–300 m,
coll. local fishermen with tangle nets, Balicasag, Panglao,
Bohol, 200–300 m, coll. local fishermen with tangle nets,
December 2000. — 12 males (44.6 × 39.7 mm, 44.4 ×
39.7 mm, 45.3 × 39.9 mm, 47.5 × 43.4 mm, 46.4 × 40.2mm, 40.5 × 35.0 mm, 44.4 × 39.0 mm, 43.9 × 38.3 mm,
44.8 × 39.2 mm, 47.0 × 40.4 mm, 46.5 × 40 .1 mm, 44.1
× 39.1 mm); 10 ovigerous females (44.3 × 38.9 mm, 42.5
× 38.1 mm, 39.3 × 34.1 mm, 37.9 × 35.2 mm, 43.7 × 37.7mm, 45.3 × 39.6 mm, 45.3 × 40.7 mm, 42.9 × 37.9 mm,
40.5 × 35.6 mm, 37.7 × 32.7 mm), 1 young female (32.2
× 26.8 mm) (ZRC 2001.0616), Balicasag, Panglao, Bohol,
200–300 m, coll. local fishermen with tangle nets, 28
November 2001. Others: 1 young female (29.7 × 25.4 mm),
1 ovigerous female (26.2 × 22.5 mm) (ZRC 2013.1222, partex ZRC 2001.0616), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, 28 November 2001. — 1 male(45.6 × 41.1 mm) (AM), Balicasag Island, Panglao, Bohol,
coll. fishermen with tangle nets, July 2003. — 1 male withrhizocephalan (38.9 × 33.5 mm) (ZRC 2013.1221), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets,
November 2003. — 4 males, 2 females (NSMT-Cr 15387),
Balicasag Island, Panglao, Bohol, coll. local fisherman via
M. Takeda & H. Komatsu, February 2003. — 1 male (43.0 ×
37.2 mm) (ZRC 2013.1223), station CP 2343, off Pamilacan
Island, Bohol Sea, 9°27.4’N 123°49.4’E, 273–356 m, coll.
PANGLAO 2005, 23 May 2005.

Alcomaja irrorata is also similar to A. confragosa (Griffin& Tranter, 1986). However, the intercalated spine is moreacutely triangular in A. confragosa (Fig. 18A, B) (relatively
stouter in A. irrorata, Fig. 18E, F); the postorbital spine
is relatively more narrow in A. confragosa (Fig. 18A, B)(relatively stouter in A. irrorata, Fig. 18E, F); the basal
antennal article is lined with granules on the outer margin in
A. confragosa (see Griffin & Tranter, 1986: fig. 72d) (withonly several granules on the inner margin in A. irrorata, Fig.
39L); the ischium of the third maxilliped is relatively shorter
with the outer surface is weakly granular in A. confragosa(see Griffin & Tranter, 1986: fig. 72c) (ischium relatively
longer with the outer surface smooth or almost so in A.
irrorata, Fig. 45A); and the G1 is relatively shorter with the
distal half more prominently curved and the distal part after
the subdistal flap proportionately shorter in A. confragosa(Fig. 19G, Griffin & Tranter, 1986: 215, fig. 73a, b) (G1relatively longer with the distal half gently curved and the
distal part after the subdistal flap proportionately longer in
A. irrorata, Fig. 19A–C).

Diagnosis. Carapace pyriform; dorsal surface of carapace andambulatory legs covered with short, hooked setae; with small
and large rounded granules, some of which merge basally(Fig. 16A). Pseudorostral spines dorsoventrally flattened in
adults (Fig. 18E, F). Intercalated spine subtruncate basally;
antorbital spine gently curved. Lateral and branchial spines
strong in juveniles, weak in adults (Fig. 18E, F). Subhepatic
region with rugose and unevenly granulated surface (Fig.
39L). Basal antennal article weakly granulated, with 6 small
granules, 2 stout spines distally (Figs. 39L, 41G). Ischium ofthird maxilliped smooth or rugose, without distinct granules(Fig. 45A). Adult male ambulatory legs long, especiallymerus and propodus first leg which can be twice length ofcarapace; surfaces almost smooth or with setae when young(Figs. 16A, 55K, L). G1 strongly curved, distal part curved
upwards (Fig. 19A–C).

For A. irrorata, we observed that male specimens under
35 mm length are still juveniles, with adults possessing
inflated chelae only at lengths of 44 mm or longer. In small
specimens, the armature is relatively sharper, in particular
the three lateral and two branchial spines and the seriesof large granules along the median row. The smallest
ovigerous female, however, is only 26.2 by 22.5 mm (ZRC
2013.1222a). Another slightly larger female specimen fromthe same site (29.7 × 25.4 mm, ZRC 2013.1222b), however,was still immature.
The spines on the posterior carapace margin and on the lateral
margins are always present but get relatively smaller andblunter as the specimens get larger. The ambulatory legs are
relatively longer in adult males, with the surfaces generallysmooth (Fig. 55K). Smaller male and female specimens,
however, have the articles distinctly more setose, particularly
the dactylus (Fig. 55L).

Etymology. The name is derived from the Latin “irroratus”
which means covered with granules, alluding to theappearance of the species.

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Although juveniles of A. irrorata (cf. 1 juvenile female 29.7
× 26.1 mm, 1 male 28.7 × 23.9 mm, ZRC 2013.1229) aresuperficially similar in general form to young Paramaja(of P. gibba and P. turgida), the pseudorostral spines areinvariably relatively longer in A. irrorata, the carapace is
far more granulous but less swollen, the spines are relatively
stronger, and the ambulatory legs are proportionately longer.

(MNHN-IU-2013-4048), station CP 61, 184–202 m, 14°02’N
120°18’E, RV Vauban, MUSORSTOM 1, 27 March 1976. –1 female (32.7 × 27.1 mm) (MNHN-IU-2013-4058), station
CP 88, 14°00’N 120°17’E, 183–187 m, coll. RV Coriolis,
MUSORSTOM 3, 31 May 1985. — 1 female (35.7 ×
29.8 mm) (MNHN-IU-2013-4056), station CP96, 14°00’N
120°17’E, 190–194 m, RV Coriolis, MUSORSTOM 3, 1 June
1985. — 1 male (31.7 × 26.6 mm) (MNHN-IU-2013-4057),
station CP 98, 14°00’N 120°18’E, 194–205 m, coll. RV
Coriolis, MUSORSTOM 3, 1 June 1985. — 1 male (31.7
× 26.6 mm) (ZRC 2001.432), Balicasag Island, Panglao,
Bohol, coll. fishermen with tangle nets, December 2000.
— 1 male (41.4 × 35.4 mm) (ZRC 2013.1225), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets,
June 2002. — 1 young female (24.1 × 20.5 mm) (ZRC
2013.1224), Balicasag Island, Panglao, Bohol, coll. fishermenwith tangle nets, June 2002. — 1 female (32.1 × 27.5mm) (ZRC 2013.1227), Balicasag Island, Panglao, Bohol,
coll. fishermen with tangle nets, 2003. — 1 female (ZRC
2013.1228), Balicasag Island, Panglao, Bohol, 200–300
m, coll. fishermen with tangle nets, May 2004. — 1 male
(NSMT-Cr 22327), Balicasag Island, Panglao, Bohol, coll.
local fisherman via M. Takeda & H. Komatsu, February
2003. — 1 female (NSMT-Cr 22326), Balicasag Island,
Panglao, Bohol, coll. local fisherman via M. Takeda & H.
Komatsu, February 2003. — 1 male (41.9 × 35.2 mm) (ZRC
2013.1226), northwest Panglao, Bohol, coll. J. Arbasto withtangle nets, 2006.

Alcomaja confragosa (Griffin & Tranter, 1986)(Figs. 16D, E, 18A, B, 19G)
Maja confragosa Griffin & Tranter, 1986: 213, figs. 72, 73a, b,
pl. 16. – Poore et al., 2008: 61. – Ng et al., 2008: 117 (list).

Material examined. Australia – 1 young female (21.5 ×
18.2 mm) (NMV J53978), off Point Cloates, 22°37.06’S
113°29.03’E – 22°37.02’S 113°28.56’E, Western Australia,
355–382 m, coll. G. Poore, CSIRO RV Southern Surveyor,
9 December 2005.
Diagnosis. Carapace pyriform; dorsal surface of carapace andambulatory legs covered with short, hooked setae; with small
and large rounded granules, some of which merge basally(Fig. 16D, E). Pseudorostral spines with rounded cross-section(Fig. 18A, B). Intercalated spine acutely triangular; antorbital
spine gently curved (Fig. 18A, B). Lateral and branchialspines strong in juveniles, weak in adults (Fig. 16D, E).
Subhepatic region with distinctly granulated surface. Basalantennal article with numerous distinct granules. Ischiumof third maxilliped distinctly granulated, especially onouter margin, with 2 large blunt distal spines. Adult maleambulatory leg condition not known (Fig. 16E). G1 gently
curved, distal part relatively short (Fig. 19G).

Diagnosis. Carapace pyriform; dorsal surface of carapace
and ambulatory legs covered with short, hooked setae;
with distinct small and large sharp and rounded granules(Figs. 16B, C, 18C, D). Pseudorostral spines with roundedcross-section (Fig. 18C, D). Intercalated spine subtruncate
basally; antorbital spine gently curved (Fig. 18C, D).
Lateral and branchial spines strong in juveniles, weak inadults. Subhepatic region with strongly granulated surface(Fig. 39M). Basal antennal article with numerous distinct
granules (Fig. 39M). Ischium of third maxilliped distinctly
granulated on surface, with 2 distinct distal spines (Fig. 45B,
C). Adult male ambulatory legs long, especially merus andpropodus first leg which can be twice length of carapace;surfaces almost smooth or with sparse setae (Figs. 16B, C).
G1 gently curved, distal part straight (Fig. 19D–F).

Remarks. A small female specimen collected from thenorthern part of Western Australia agrees very well withthe type male described in Griffin & Tranter (1986). The
holotype of Maja confragosa was described from Indonesia,
in the Eastern Mollucas. The new material comes from theTropic of Capricorn near 22°S in the same depth range,around 300 m.
Differences with the allied A. irrorata n. sp. and A. gracilipes(Chen & Ng, 1999) are discussed under these two species.
Alcomaja gracilipes (Chen & Ng, 1999)(Figs. 16B, C, 18C, D, 19D–F, 39M, 45B, C, 49B, C,
53N)

Remarks. The specimens from the Philippines agree wellwith the holotype of A. gracilipes (Chen & Ng, 1999)
described from the South China Sea. The characters outlined
in Chen & Ng (1999) serve to easily distinguish A. gracilipes
from A. confragosa and A. irrorata n. sp. (see also discussionof A. irrorata).

Maja gracilipes Chen & Ng, 1999: 754, figs. 1, 2. – Yang et al.,
2008: 780. – Ng et al., 2008: 117 (list).

Material examined. Holotype: male (45.4 × 43.5 mm)(IOCAS K33B-34), station 6080, 180 m, on gravel, South
China Sea, coll. 21 April 1959. Philippines – 1 female (39.8
× 29.2 mm) (MNHN-IU-2013-4049), station CP 27, 14°00’N
120°19’E, 188–192 m, coll. RV. Vauban, MUSORSTOM
1, 22 March 1976. — 1 male (31.9 × 25.5 mm) (MNHNIU-2013-4054), station CP 35, 13°59’N 120°18’E, 186–187
m, coll. RV Vauban, MUSORSTOM 1, 23 March 1976.
— 1 male (24.8 × 19.4 mm), 1 female (16.3 × 13.2 mm)That both A. irrorata and A. gracilipes occur in the Philippines(with some specimens sympatric) does pose problems but
we know too little about the ecology of these species to
speculate further. In the steep slopes along Balicasag,
Philippines, the dominant species is clearly A. irrorata,
with only a few specimens of A. gracilipes obtained. On theother hand, only A. gracilipes was observed in the trawledmaterial from the French expeditions to the Philippines,147

Ng & Richer de Forges: Revision of the spider crab genus Maja
albeit there were only a few specimens. The type male wasalso collected by trawling in the South China Sea. Only one
specimen of A. irrorata from off Pamilacan Island (male 43.0
× 37.2 mm, ZRC 2013.1223,) was collected by trawling in
the Philippines. Whether this suggests they have different
habitat preferences is not known.

Island, Panglao, Bohol, 200–300 m, coll. fishermen withtangle nets, November 2003. — 1 female (AM), Balicasag
Island, Panglao, Bohol, 200–300 m, coll. fishermen withtangle nets, December 2003. — 3 males (22.6 × 18.7 mm,
26.4 × 21.6 mm, 28.3 × 24.1 mm) (ZRC 2013.1381), 19
May 2004. — 1 male (28.9 × 24.8 mm) (ZRC 2013.1388),
28 May 2004. — 1 ovigerous female (ZRC 2013.1386),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, May 2004. — 1 male (ZRC 2013.1382),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, 29 May 2004. — 1 male (AM), Balicasag
Island, Panglao, Bohol, 200–300 m, coll. fishermen withtangle nets, 4 June 2004. — 1 male (26.1 × 22.0 mm) (ZRC
2013.1390), northwest Panglao, Bohol, 120–160 m, tangle
nets, coll. J. Arbasto, 1 December 2007. — 1 male (27.7 ×
23.8 mm), 1 ovigerous female (20.9 × 17.4 mm), 1 female(22.2 × 19.2 mm) (ZRC 2013.1399), northwest Panglao,
Bohol, 120–160 m, tangle nets, coll. J. Arbasto, 2007. — 1
female (15.0 × 11.2 mm) (MNHN-IU-2013-4053), station
CP 31, 14°00’N 120°16’E, 187–195 m, coll. RV Vauban,
MUSORSTOM 1, 22 March 1976. — 1 ovigerous female(15.8 × 12.7 mm), 1 female (12.1 × 9.1 mm) (MNHNIU-2013-4052) CP57, 13°53’N-120°13’E, 96–107 m, coll.
RV Vauban, MUSORSTOM 1, 26 March 1976. Taiwan –1 male (29.5 × 24.2 mm) (ZRC 2013.1385), Tashi fishingport, Ilan Province, sandy mud, 20 m, coll. Hwang 28 April
1989. — 1 male (23.2 × 18.4 mm) (ZRC 2013.1383), Tashi
fishing port, Ilan Province, sandy mud, 20 m, coll. Hwang,
4 July 2011.

Alcomaja nagashimaensis (T. Sakai, 1969)(Figs. 17A–C, 20A–J, 36M, 39O, 41H, 45E, F, 49D, E,
52G, 53P, 55N, 69A)
Maja nagashimaensis T. Sakai, 1969: 256, pl. 1, fig. 2, text-fig.
5b, b’. – Serène & Lohavanijaya, 1973: 50 (key). – T. Sakai,
1976: 241, pl. 85, fig. 1, text-fig. 129. – Serène & Vadon,
1981: 129. – Griffin & Tranter, 1986: 217. – Muraoka, 1998:
27. – Ng et al., 2008: 117 (list).
Maja nagasimaensis [sic] – Maramura & Kosaka, 2003: 34.

Material examined. Holotype: male (34.0 × 28.5 mm)(USNM 125886), on rocky shore, Kii Nagashima, Mie
Prefecture, Honshu Island, Japan, coll. Y. Yamamoto.
Others: Japan – 1 male (37.2 × 32.4 mm) (CBM-ZC9928),
Ohse-Zaki, Suruga Bay, 35°3.53’N 138°48.26’E, 135–142
m, in 1 m ORI dredge, cruise KT04-6, station OS-1, coll. T.
Akiyama, RV Tansei-Maru, 4 May 2004. Philippines – 2
males (27.3 × 23.3 mm, 28.7 × 23.9 mm), 1 ovigerous female(23.9 × 20.6 mm) (ZRC 2013.1393, ex part ZRC 2001.0435),
Balicasag Island, Panglao, Bohol, coll. fishermen with tangle
nets, December 2000. — 2 ovigerous females (24.7 × 20.6mm, 24.6 × 22.3 mm), 3 males (23.4 × 19.5 mm, 21.9 × 18.4mm, 15.7 × 7.3 mm) (ZRC 2001.0434), Balicasag Island,
Panglao, Bohol, coll. fishermen with tangle nets, December
2000. — 1 male (30.5 × 26 mm) (ZRC 2013.1394, ex part
ZRC 2001.430), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, 28 November 2000. — 4 males(27.7 × 22.8 mm, 26.8 × 22.1 mm, 22.3 × 17.7 mm, 22.9 ×
17.6 mm [with bopyrid], 7 ovigerous females (28.2 × 24.0mm, 25.1 × 20.8 mm, 22.9 × 18.7 mm, 25.1 × 21.1 mm, 23.1
× 20.0 mm, 22.6 × 19.1 mm) (ZRC 2001.0585a), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets, 28
November 2001. — 1 female (ZRC 2013.1389, ex part ZRC
2001.0601), Balicasag Island, Panglao, Bohol, 200–300 m,
coll. fishermen with tangle nets, 28 November 2001. — 1
juvenile male (ZRC 2013.1315), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, June
2002. — 2 males, 2 females (ZRC 2013.1402), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets,
June 2002. — 2 males (NSMT-Cr 15390a), Balicasag Island,
Panglao, Bohol, coll. local fisherman via M. Takeda & H.
Komatsu, February 2003. — 3 males (NSMT-Cr 13032),
Balicasag Island, Panglao, Bohol, coll. local fisherman via
M. Takeda & H. Komatsu, February 2003. — 1 male (20.8
× 16.8 mm), 3 females (23.7 × 19.4 mm, 23.2 × 19.3 mm,
23.1 × 18.2 mm), 1 ovigerous female (23.9 × 19.9 mm)(ZRC 2013.1400), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, July 2003. — 1 male (19.9 ×
16.6 mm) (ZRC 2013.1391), Balicasag Island, Panglao,
Bohol, coll. fishermen with tangle nets, July 2003. — 1
male (ZRC 2013.1387), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, 25–30 July
2003. — 1 male, 1 female (ZRC 2013.1392), Balicasag

Diagnosis. Carapace pyriform; dorsal surface of carapace
and ambulatory legs covered with short, hooked setae; withsmall rounded granules (Fig. 17A–C). Pseudorostral spines
with rounded cross-section (Fig. 36M). Antorbital spine
almost straight; intercalated spine acutely triangular, shorter
than supraorbital and postorbital spines, with large open gaps
between them; postorbital spine large, longer than others;
1 distinct hepatic spine with small basal spine (Fig. 36M).
Lateral and branchial spines strong in juveniles and adults(Fig. 17A–C). Subhepatic region with rugose and weakly
granulated surface (Fig. 39O). Basal antennal article withlow granules and 2 tubercles, with 2 distal spines (Fig. 39O,
41H). Ischium of third maxilliped almost smooth to slightly
rugose (Fig. 45E, F). Ambulatory legs relatively short, firstleg never twice length of carapace; surfaces distinctly setose(Figs. 17A–C, 55N). G1 strongly curved, with distal partgently curved downwards (Fig. 20A–J).
Remarks. The type male (Fig. 17A) is very delicate and inpoor condition, and appears to have dried up at some stage.
The gonopods are still present, although not in a good state.
The lateral and branchial spines of the type are relatively
stouter (Fig. 17A) compared to the other specimens we haveexamined from Japan, Taiwan and Philippines (Fig. 17B,
C), but this is probably due to variation. The G1 drawn by
T. Sakai (1969: text-fig. 5b) (present Fig. 20A, B) appears
to have been drawn in situ and the curvature is therefore
less strong compared to that when drawn ex-situ (Fig. 20C,
D). The subdistal process observed seems to have dried up
previously and looks folded, unlike the original condition
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Fig. 16. General habitus, Alcomaja species. A, A. irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR), Philippines; B, A. gracilipes,
holotype male (45.4 × 43.5 mm) (IOCAS K33B-34), South China Sea; C, A. gracilipes, male (41.4 × 35.4 mm) (ZRC 2013.1225),
Philippines; D, A. confragosa, female (21.5 × 18.2 mm) (NMV J53978), Australia; E, A. confragosa, holotype male (as Maja confragosa,
after Griffin & Tranter 1986: pl. 16a).

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Fig. 17. General habitus, Alcomaja species. A, A. nagashimaensis, holotype male (34.0 × 28.5 mm) (USNM 125886), Japan; B, A.
nagashimaensis, male (37.2 × 32.4 mm) (CBM-ZC9928), Japan; C, A. nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Japan;
D, A. latens n. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon Islands; E, A. miriky n. sp., holotype male (26.4
× 20.9 mm) (MNHN-IU-2010-929), Madagascar; F, A. desmondi n. sp., holotype male (35.4 × 28.3 mm) (NMCR), Philippines.

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Fig. 18. Frontal regions of carapace showing different structures of intercalated teeth, Alcomaja species. A, A. confragosa, holotype male(as Maja confragosa, after Griffin & Tranter, 1986: pl. 16a); B, A. confragosa, female (21.5 × 18.2 mm) (NMV J53978), Australia; C,
A. gracilipes, holotype male (45.4 × 43.5 mm) (IOCAS K33B-34), South China Sea; D, A. gracilipes, male (41.4 × 35.4 mm) (ZRC
2013.1225), Philippines; E, A. irrorata n. sp., paratype male (44.7 × 39.1 mm) (ZRC 2013.1229), Philippines; F, A. irrorata n. sp., holotype
male (52.6 × 45.7 mm) (NMCR), Philippines.

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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].

figured (T. Sakai, 1969: text-fig. 5b’) which is more similar
to that in the fresh material on hand. In any case, the shapeand structure (Fig. 20C, D) agrees well with the G1s from
Philippines and Taiwan and we have no doubt they areconspecific.

first leg never twice length of carapace; surfaces distinctly
setose (Fig. 17D). G1 gently curved, with distal part gently
curved upwards (Fig. 20K–N).
Etymology. The name is derived from the Latin for “hidden”,
alluding to a general absence of external characters to separate
the new species from A. nagashimaensis.

The chelae are only enlarged in larger males. Smaller males
about 20 mm or less in carapace length (e.g., ZRC 2013.1391)
have slender chelae.

Remarks. Alcomaja latens n. sp. is very close to A.
nagashimaensis (T. Sakai, 1969) and the two species can only
be effectively separated by the form of the G1. In Alcomaja
latens, the dorsal part is gently bent upwards (Fig. 20K–N)
while in A. nagashimaensis, it is gently curved downwards(Fig. 20A–J). At around the same size as the holotype of A.
latens, a male specimen of A. nagashimaensis already has
distinctly inflated chelipeds.

Maja nagashimaensis was described from Japan, with
Serène & Vadon (1981) reporting this species from thePhilippines, and Griffin & Tranter (1986) recording it fromthe Sulu Islands in the Philippines. It is now also recorded
from Taiwan.
Alcomaja latens n. sp.
(Figs. 17D, 20K–N, 36N, 45G, 49F)

Alcomaja miriky n. sp.
(Figs. 17E, 20O–Q, 36O, 45H, 49G, 69B)

Material examined. Holotype: male (25.3 × 21.4 mm)(MNHN-IU-2013-4050), station DW 2190, 8°24’S 159°27’E,
140–263 m, Solomon Islands, coll. RV Alis, SALOMON
2, 24 October 2004.

Material examined. Holotype male (26.4 × 20.9 mm)(MNHN-IU-2010-929), station DW 3215, 12°32’S 47°54’E,
314–433 m, Madagascar, coll. MIRIKY, 30 June 2009.
Paratype: Madagascar – 1 male (17.3 × 12.7 mm) (ZRC
2013.1409, ex MNHN-IU-2010-60), station CP 3284,
14°51’S 46°59’E, 236–297 m, coll. MIRIKY, 13 July 2009.

Diagnosis. Carapace pyriform; dorsal surface of carapace
and ambulatory legs covered with short, hooked setae; withsmall rounded granules (Fig. 17D). Pseudorostral spines withrounded cross-section (Fig. 36N). Antorbital spine almost
straight; intercalated spine subtruncate basally; postocularspine strong, pointing anteriorly and outwards; 1 distinct
hepatic spine with small basal spine (Fig. 36N). Lateral andbranchial spines strong in juveniles and adults (Fig. 17D).
Subhepatic region with rugose and weakly granulated surface.
Basal antennal article with low granules and 2 tubercles, with
2 distal spines. Ischium of third maxilliped almost smooth toslightly rugose (Fig. 45G). Ambulatory legs relatively short,

Diagnosis. Carapace elongated, pyriform; dorsal surface of
carapace and ambulatory legs covered with short, hooked
setae; with small rounded granules (Fig. 17E). Pseudorostralspines relatively long, curved, with rounded cross-section(Fig. 36O). Antorbital spine almost straight; intercalated
spine acutely triangular; postocular spine very long; 1 distinct
hepatic spine (Fig. 36O). Lateral and branchial spines strong
in juveniles and adults (Fig. 17E). Subhepatic region withrugose, weakly granulated surface. Basal antennal article
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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) (CBMZC9928), 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].

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Etymology. The name is formed to honor Desmond Griffin,
a carcinologist who devoted a large part of his life to thetaxonomy of Indo-West Pacific majoid crabs.

with low granules and 2 tubercles, with 2 relatively long
distal spines. Ischium of third maxilliped almost smooth toslightly rugose (Fig. 45H). Ambulatory legs relatively short,first leg never twice length of carapace; surfaces distinctly
setose (Fig. 17E). G1 gently curved with distal part gently
curved downwards (Fig. 20O–Q).

Remarks. Alcomaja desmondi n. sp. superficially resembles
Holthuija suluensis (Rathbun, 1916), in general shape,
especially since both species have strong lateral and branchialspines as well as two gastric spines on the carapace.
Excluding the generic level differences, A. desmondi hasa more rounded carapace, especially on the branchialregions; the pseudorostral spines are more diverging inHolthuija suluensis; the first gastric spine is placed moreanteriorly in A. desmondi; the antorbital spine is straightand directed outwards in A. desmondi (versus curved anddirected anteriorly in H. suluensis), and the setae covering
the ambulatory legs in A. desmondi are longer and stiffer
compared to H. suluensis.

Etymology. The species is named after the cruise thatcollected it, MIRIKY. The name is used as a noun inapposition.
Remarks. Alcomaja miriky n. sp. is very close to A.
nagashimaensis (T. Sakai, 1969). The main differences are
as follow: the G1 of A. miriky is prominently less curved(Fig. 20O–Q versus Fig. 20A–J); the distal spines of thebasal antennal article in A. miriky are relatively longer andsharper; the overall carapace is proportionately longer (Fig.
17E versus Fig. 17A–C); there is only one hepatic spine in
A. miriky versus two in A. nagashimaensis (Figs. 17E, 36O
versus Figs. 17A–C, 36M); there is one accessory spineon the postorbital spine in A. miriky which is absent in A.
nagashimaensis (Figs. 17E, 36O versus Figs. 17A–C, 36M);
and the pseudorostral spines of A. miriky are relatively longer
and more curved (Figs. 17E, 36O versus Figs. 17A–C, 36M).

Within Alcomaja, A. desmondi stands out in several features.
It is unique among members of the genus in possessing
long stiff setae covering its carapace and legs; and the G1
structure is atypical, being almost straight, much stouter withan unusually broad ventral flap (Fig. 20R–T). Since the restof its characters agree with Alcomaja as presently defined,
it seems best to leave it in this genus for the time being

Alcomaja desmondi n. sp.
(Figs. 17F, 20R–T, 36L, 39N, 45D, 49H, 52F, 53O, 55M)

The dense stiff setae on the specimens of A. desmondi from
Balicasag trap a thick layer of silicate sponge spicules, whichmakes their handling very difficult. It appears to be a species
living in the deep sea on the deep steep rocky slopes ofBalicasag Island at depths exceeding 300 m.

Material examined. Holotype: male (35.4 × 28.3 mm(NMCR), Balicasag Island, Panglao, Bohol, Philippines,
200–300 m, coll. local fishermen with tangle nets, June
2002. Paratypes: Philippines – 2 males (36.6 × 31.7 mm,
27.1 × 23.5 mm) (ZRC 2013.1273), same data as holotype.
— 2 males (27.7 × 21.3 mm, 36.6 × 29.1 mm), 1 ovigerous
female (38.0 × 32.6 mm) (ZRC 2001.0588), Balicasag Island,
Panglao, Bohol, 50–500 m, coll. local fishermen with tangle
nets, 28 November 2001. — 1 ovigerous female (37.6 ×
32.0 mm) (NSMT-Cr 22328), Balicasag Island, Panglao,
Bohol, coll. local fisherman via M. Takeda & H. Komatsu,
February 2003.

Paramaya De Haan, 1837
Maja (Paramaya) De Haan, 1837: pl. 24, fig. 4.
Maja (Paramaya) – De Haan, 1837: errata.
Paramaya – Rathbun, 1905: 73.
Maja – T. Sakai, 1938: 296 (part). – T. Sakai, 1965: 83 (part).
(not Maja Lamarck, 1801).

Diagnosis. Carapace ovate; dorsal surface convex, coveredby granules or tubercles; gastric and branchial regions
distinct, clearly delimited by distinct grooves (Figs. 21,
22A–C). Intestinal region with long, distinct medianspine (Figs. 21, 22A–C). Pseudorostral spines very long,
diverging (Figs. 21, 22A–C, 37A–C). Supraorbital eavewith anterior part longitudinally narrow, rectangular, notprominently expanded; antorbital spine sharp (Fig. 37A–C).
Intercalated spine distinct, separated from supraorbital eaveand postorbital spine by wide gaps; postorbital spine strong;
hepatic region with 1 strong spine, slightly shorter or longer
than postorbital spine; 1 smaller spine below (Fig. 37A–C).
Lateral carapace margin with 3 long spines and numerous
granules and spinules, branchial region with 3, sometimes
4 spines (Figs. 21, 22A–C). Posterior carapace margin with
2 long median spines (Figs. 21, 22A–C). Eyes relatively
long, slender, with ovoid cornea (Fig. 37A–C). Antennal
flagellum short, slender. Basal antennal article longer thanbroad, rectangular; surface with several low tubercles, with
2 spines distally; inner and outer lateral margins smoothto uneven; proximal outer angle acute, may be produced

Diagnosis. Carapace pyriform; dorsal surface of carapace
densely covered with long, stiff setae; with small rounded
granules (Fig. 17F). Pseudorostral spines with rounded crosssection (Fig. 36L). Antorbital spine strong, curved upwards,
appears straight from dorsal view; intercalated spine acutely
triangular; postorbital spine very long (Fig. 36L). Lateraland branchial spines strong in juveniles and adults; 3 spines
on median row: 2 gastric, 1 cardiac; 2 distinct spines onposterior carapace margin (Fig. 17F). Subhepatic region withnumerous very small rounded granules on surface (Fig. 39N).
Basal antennal article with numerous granules (Fig. 39N).
Ischium of third maxilliped almost smooth to slightly rugose(Fig. 45D). Ambulatory legs relatively short, first leg never
twice length of carapace; surfaces strongly setose, covered
with long coarse setae (Figs. 17F, 55M). G1 almost straightwith distinct ventral fold (Fig. 20R–T).

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into spine; antero-external crested rim of antennular fossa
overlaps distal part of basal antennal article by about a thirdof its width, forming hook-like structure (Fig. 40A–C).
Epistome rectangular, longer than broad, anterior margin
with 2 tubercles; posterior margin composed of 4 rectangularplates separated by shallow fissures (Figs. 40A–C, 42A,
B). Suborbital margin separated from basal antennal article
by U-shaped cleft, separated from margin of postorbital
tooth by deep fissure (Fig. 40A–C). Outer surface of third
maxilliped covered by setae; ischium subrectangular, longer
than broad; postero-external angle of merus very narrow,elongate, “inserted” into deep concavity on outer margin ofischium; antero-internal part of ischium acutely triangular(Fig. 45I–L). Male chelipeds relatively long in adult males,surfaces of merus and carpus almost smooth; carpus elongate;
propodus of palm elongated, may be inflated, curved, smooth,may have lateral cristae, palm longer than fingers (Fig. 54A,C); fingers long, slender, almost straight to gently curved,
with or without distinct basal gape when closed (Figs. 21,
22A–C, 54A–C). Ambulatory legs relatively long, slender;
merus with long dorsal subdistal spine; dactylus elongate,
curved, covered with long setae except for corneous tip (Figs.
6G, 21, 22A–C, 56A, B). Thoracic sternum longer thanwide; surfaces of somites 5–8 with small granules; sternites
3 and 4 distinctly depressed; margin between sternites 2
and 3 demarcated by deep cleft; anterior margin of sternoabdominal cavity not forming complete rim (Figs. 50A–C,
52H). Male abdomen subtriangular, with 6 free somites
and telson; somites 3 and 4 wider than somites 5, 6 andtelson (Fig. 50A–C). Male press-button abdominal locking
mechanism submedian in position on sterno-abdominal cavity(Fig. 52H). Female abdomen dome-shaped, covering mostof thoracic sternum. G1 very long, slender, gently curved;
distal part with rounded tip, dorsal part folding upwards toform distinct dorsal projection, with scattered very shortsetae (Fig. 23).

His name is a nomen nudum and is not possible to identify
at the moment.
Key to species of Paramaya De Haan, 1837
1. Pseudorostral horns relatively short; ambulatory meri in adults
relatively stout, short (Fig. 21A–D); G1 gently curved (Fig.
23A–C); Japan and Taiwan.......................................................
...................................... Paramaya spinigera (De Haan, 1837)– Pseudorostral horns very long; ambulatory meri in adults
relatively slender, long (Figs. 21E, F, 22A); G1 otherwise...2
2. Surface of thoracic sternum relatively smooth, with very low
rounded granules (Fig. 50C); G1 gently curved (Fig. 23G–I);
Vanuatu............................................ Paramaya coccinea n. sp.
– Surface of thoracic sternum distinctly granulated (Fig. 50B);
G1 more strongly curved especially distally (Fig. 23D–F);
Philippines..............................................Paramaya ouch n. sp.

Paramaya spinigera (De Haan, 1837)(Figs. 6G, 21A–D, 23A–C, 37A, 40A, 42A, 45I, 50A,
52H, 54A, 56A)
Maja (Maja) spinigera De Haan, 1837: 93.
Maja (Paramaya) spinigera – De Haan, 1837: errata.
Maja (Paramaya) spinigera – De Haan, 1837: pl. 24, fig. 4.
Maia spinigera – Adams & White, 1848: 15. – Parisi, 1915: 289.
Maja spinigera – Ortmann, 1893: 51. – Doflein, 1902: 656. – Ninni,
1924: 47, un-numbered figure on p. 48. – Balss, 1924: 34. –T. Sakai, 1934: 297. – T. Sakai, 1936: 98, pl. 25, fig. 3. – T.
Sakai, 1938: 397, pl. 30, fig. 1. – T. Sakai, 1965: 84, pl. 38. –Serène, 1968: 57. – Utinomi, 1969: 77, pl. 39-3. – Holthuis &T. Sakai, 1970: 121, pl. 14. – Nishimura & Suzuki, 1971: 109,
pl. 38-2. – Serène & Lohavanijaya, 1973: 50 (key). – Takeda,
1975: 127. – T. Sakai, 1976: 237, pl. 83. – Matsuzawa, 1977:
pl. 96 fig. 4. – Takeda, 1982: 128. – Miyake, 1983: 43, pl.
14 fig. 4. – Dai et al., 1986: 136, pl. 18(3). – Matsuda et al.,1986: 130. – Dai & Yang, 1991: 151, pl. 18(3). – Yamaguchi& Baba, 1993: 359, Fig. 116. – Takeda, 1993: 43. – Huang,
1994: 582. – Muraoka & Odawara, 1995: 34. – Muraoka, 1998:
27. – Minemizu, 2000: 216. – Takeda et al., 2000: 139. – Ng
et al., 2001: 12. – Maramura & Kosaka, 2003: 34. – Takeda& Ueshima, 2006: 74. – Yang et al., 2008: 780. – Ng et al.,
2008: 117 (list).

Type species. Maja (Maja) spinigera De Haan, 1837, bymonotypy.
Remarks. The genus Paramaya was established by De
Haan (1837: 24) to accommodate Maja (Maja) spinigera.
As discussed earlier (see remarks for Maja), the nameis available even though it is very similar in spelling toParamaja Kubo, 1936. In the present study, we add two new
species to Paramaya, P. ouch n. sp. from the Philippines
and P. coccinea n. sp. from Vanuatu.

Material examined. Japan – 1 female (NSMT 5999), KiiNagashima, Kaino, Mie Prefecture, coll. M. Saba, 1970s.
— 1 carapace only (NSMT 7614), Kii-Nagashima, Kaino,
Mie Prefecture, coll. M. Saba, 16 December 1976. — 1
male, 1 female (NSMT 5967), Kii-Nagashima, Kainoura,
Mie Prefecture, coll. M. Saba, 1970s. — 1 male (NSMT6134), Kii-Nagashima, Kainoura, Mie Prefecture, in lobster
gill-nets, coll. M. Saba, 25 February 1979. — 1 male (NSMT
10178), Kii-Nagashima, Kaino, Mie Prefecture, coll. M.
Saba, 18 December 1978. — 1 female (SMF 47734), Toba,
Mie Prefecture, coll. N. Yamashita, 1983. — 1 male, 1female (SMF 9289), Toba, Mie Prefecture, coll. Nakamura,
May 1980. — 1 male (NSMT 6032), Kii-Nagashima, Mie
Prefecture, coll. M. Saba, 1970s. — 1 male (NSMT 6752),
Kii-Nagashima, Mie Prefecture, coll. M. Saba, 1970s. — 1
female (NHM 1961.11.13.7/9, part), Seto, Shirahama, Japan,
coll. I. Gordon. — 1 male, 1 female (NHM 1961.11.13.7/9,
part), Seto, Shirahama, Japan, coll. I. Gordon. — 1 male

Members of Paramaya are distinctive with their very longcarapace spines, elongate epistome and spinose ambulatory
meri, and cannot be confused with any other genus. The
carapaces of most of the Paramaya specimens examined
were not extensively covered with animals or debris, andspecies generally only have sparse stiff setae on their bodies.
Their ambulatory legs, however, are densely covered withlong stiff setae and these often trap debris.
White (1847: 7) recognised a new species: “PARAMAYA
Dehaanii, n. s.” but did not provide any data or description.

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Ng & Richer de Forges: Revision of the spider crab genus Maja
(64.7 × 51.2 mm) (NHM 1961.6.5.117), Tosa Bay, coll. K.
Sakai, April 1961. — 1 male (USNM 18854), Miura, Atami
Province, Yokaito coast, Japan, from Mr Sakamoto, fromGarrett Droppers. — 1 male (SMF 30766), Katahara, Aichi,
80–150 m, coll. N. Akashi, 1978-1983. — 1 male, 1 female(SMF 7627), Kii Peninsula, coll. K. Sakai. — 2 males, 4
females, 1 ovigerous female (SMF 46532), Kii Minabe,
33°44.944’N 135°19.65’E, coll. K. Sakai, 15 February 1983.
— 1 male (SMF 47732), Shikoku, Usa, Ikenoura, Kochi
Prefecture, coll. K. Sakai, 18 October 1984. — 1 female(SMF 47747), None, Cape Muroto, Kochi Prefecture, ca.
33°26.617’N 134°16.75’E, coll. K. Matsuzawa, 20 January
1993. — 1 ovigerous female (SMF 47746), Mitsu, Kochi
Prefecture, 33°16.6’N 134°10.6’E, coll. 9 February 1974. —1 ovigerous female (SMF 47748), Mitsu, Kochi Prefecture,
33°16.6’N 134°10.6’E, coll. 13 December 1994. — 1
male (SMF 47757), Mitsu, Muroto City, Kochi Prefecture,
33°16.6’N 134°10.6’E, no other data. — 1 female (SMF
47741), Tokushima, Shikoku, Tsubakidomari, Anan, Kochi
Prefecture, 33°51.678’N 134°43.603’E, coll. T. Nakano,
18 April 1983. — 1 female (SMF), Minabe, Wakayama
Prefecture, coll. K. Sakai, 18 February 1984. — 1 male(SMF), Minabe, Wakayama Prefecture, T. Sakai Collection.
— 1 female (SMF 47742), Honshu, Wakayama Prefecture,
Kii Mimase, coll. K. Sakai, 1 March 1988. — 1 male (NSMT
15905), no other data. — 1 dried female (70.1 × 58.8 mm)(KPM NH4195), T. Sakai Collection. — 1 juvenile female(USNM 48486), station 4880, eastern channel of Korean
Straits, vicinity of Ohi-Shima, 59 fathoms, coll. RV Albatross,
2 August 1906. Taiwan – 7 males (85.0 × 66.4 mm, 78.2
× 62.1 mm, 73.6 × 55.3 mm, 68.3 × 53.4 mm, 73.8 ×
58.4 mm, 62.8 × 49.0 mm, 72.8 × 57.3 mm), 1 ovigerous
female (63.0 × 48.6 mm) (ZRC 1999.738), Longtong, near
Keelung, northern Taiwan, in tangle nets for lobsters, coll.
S.-H. Wu, May 1999.

The specimen was collected with a number of other relatively
deeper reef animals at night (see Ng, 1994). Alcock’s (1895)
and Alcock & Anderson’s (1898) specimen has relatively
short ambulatory legs (notably the merus), suggesting it isunlikely to be real Paramaya spinigera (or either of the
new species) as presently defined. However, this may be
a consequence of its apparently small size (judging by the
figure). The specimen is not in the NHM and is probably
still in the Indian Museum (present day Zoological Surveyof India). The larger specimen photographed from Sri Lanka
has proportionately longer legs and a spiny carapace, notunlike P. coccinea n. sp. and P. ouch n. sp., although on
the basis of geography, they are probably different species.
For the moment, we do not know their precise identities.
Paramaya ouch n. sp.
(Figs. 1B, 2D, 21E, F, 23D–F, 37B, 40B, 45J, K, 50B,
54B, 70A)
Material examined. Holotype: male (76.8 × 60.0 mm)(NMCR), Balicasag Island, Panglao, Bohol, Philippines,
coll. local fishermen in tangle nets, July 2003. Paratypes:
Philippines – 2 ovigerous females (75.1 × 58.0 mm, 63.2
× 49.3 mm) (ZRC 2013.1312), same data as holotype. — 1
male (NSMT-Cr 16646), Balicasag Island, Panglao, Bohol,
coll. local fisherman via T. Kase, 6–10 March 1999. —2 males, 1 female (NSMT-Cr 15391), Balicasag Island,
Panglao, Bohol, coll. local fisherman via T. Kase, 6–10
March 1999. — 2 young females (19.0 × 13.7 mm, 28.0
× 20.6 mm) (ZRC 2013.1282, ex part ZRC 2001.0411),
Balicasag Island, Panglao, Bohol, coll. fishermen in tangle
nets, December 2000. — 3 males (largest 76.5 × 61.3 mm,
one smaller male with rhizocephalan), 2 ovigerous females(larger 60.6 × 74.1 mm), 3 females (ZRC 2001.0577),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenin tangle nets, 28 November 2001. — 4 males (65.0 × 50.4
mm, 53.3 × 40.1 mm, 69.0 × 52.9 mm, 70.7 × 51.7 mm[with rhizocephala], 1 ovigerous female (64.9 × 51.8 mm),
3 ovigerous females (with rhizocephalan) (56.9 × 40.9 mm,
68.9 × 53.5 mm, 58.7 × 42.5 mm), 1 young female (30.0
× 22.5 mm) (ZRC 2013.1187), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen in tangle nets, June 2002.
— 1 juvenile male (2013.1313), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, June
2002. — 1 male (56.1 × 40.5 mm), 1 female (47.7 × 36.1mm) (AM, ex ZRC 2001.0374), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen in tangle nets, December
2000. — 1 female (46.5 × 36.9 mm) (ZRC 2013.1299),
Balicasag Island, Panglao, Bohol, coll. fisherman with tangle
nets, 25 October 2003. — 4 males (54.5 × 42.1 mm, 71.4 ×
56.5 mm, 75.8 × 62.0 mm, 48.7 × 36.8 mm), 4 females (55.0
× 43.0 mm, 66.5 × 52.5 mm, 57.6 × 44.5 mm, 65.5 × 50.8mm) (ZRC 2012.1207), Balicasag, Panglao, Bohol, 200–300
m, coll. local fishermen with tangle nets, November 2003 –May 2004. — 1 male (32.7 × 25.2 mm) (ZRC 2013.1283),
Balicasag Island, Panglao, Bohol, coll. fisherman with tangle
nets, February 2004. — 1 female (42.6 × 34.5 mm) (ZRC
2013.1300), Balicasag Island, Panglao, Bohol, coll. fishermanwith tangle nets, May 2004. — 1 male (33.0 × 26.1 mm)(ZRC 2013.1298), station PN 1, Balicasag Island, Panglao,

Diagnosis. Pseudorostral horns relatively short (Fig. 21A–D).
Hepatic, lateral and branchial spines long; median row with5 spines: 3 gastric, 1 cardiac, 1 intestinal; 2 strong spines
on posterior carapace margin (Fig. 21A–D). Surface ofthoracic sternum relatively smooth, with very low rounded
granules (Fig. 50A). Chela of adult male with distinct carina
on dorsal and ventral margins (Fig. 54A). Ambulatory meriin adults relatively stout, short (Fig. 21A–D). G1 gently
curved (Fig. 23A–C).
Remarks. Yamaguchi & Baba (1993: 359) recorded two
syntype female specimens (87.4 × 72.9 mm, 76.9 × 66.4mm) in RMNH and selected the larger one as lectotype.
The excellent figures in De Haan (1837: pl. 24, fig. 4) andYamaguchi & Baba (1993: fig. 116) leave no doubt thatthe present specimens from Taiwan are conspecific withParamaya spinigera s. str.
Alcock (1895: 239) and Alcock & Anderson (1898: pl. 34,
fig. 3) recorded “Maia spinigera” from a specimen from off
Beluchistan (Pakistan) in the Indian Ocean (sex and size notmentioned) (Fig. 22C, D). We also have a photograph of alarge specimen (not preserved) collected from Sri Lanka indeeper waters off a reef by aquarium collectors (Fig. 22B).
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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.

52.0 mm) (ZRC 2013.1306), station P4, Balicasag Island,
Panglao, Bohol, 200–300 m, coll. fishermen with tangle
nets, 4 June 2004. — 1 male (65.0 × 50.6 mm), 1 female(70.0 × 54.6 mm), 2 ovigerous females (69.4 × 53.6 mm,
71.2 × 56.5 mm) (ZRC 2012.1209), Balicasag, ca. 100 m,
9°31.1’N 123°41.5’E, Panglao, Bohol, 200–300 m, coll. local
fishermen with tangle nets, coll. PANGLAO 2004, June 2004.
— 1 male (69.5 × 53.7 mm), 1 female (41.5 × 30.6 mm)(ZRC 2012.1208), Maribojoc Bay, Panglao, coll. J. Arbasto,
November 2003 – April 2004. — 1 ovigerous female (68.3

Bohol, 200–300 m, coll. fishermen with tangle nets, January
2004. — 1 young female (ZRC 2013.1281), Balicasag Island,
Panglao, Bohol, 200–300 m, coll. fishermen with tangle
nets, February 2004. — 1 female (53.3 × 43.4 mm) (ZRC
2013.1305), station PN1, Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, 28 May 2004.
— 1 female (53.3 × 43.4 mm) (ZRC 2013.1305), station
PN 1, Balicasag Island, Panglao, Bohol, 200–300 m, coll.
fishermen with tangle nets, 28 May 2004. — 2 males (37.2
× 29.3 mm, 50.3 × 38.5 mm [damaged]), 1 female (62.7 ×
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Ng & Richer de Forges: Revision of the spider crab genus Maja

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).

Remarks. The character used in their keys by Serène& Lohavanijaya (1973), T. Sakai (1976) and Griffin &Tranter (1986) concerning a proximal tooth on the outermargin of antenna looks very variable. On some specimens
there are only two branchial spines. Even on the juvenile
specimens (female 28 mm carapace length, ZRC 2013.1282)
the characteristics of P. ouch n. sp. are well marked: thepseudorostral spines are long and very divergent but lesscurved upward than in the adults; and the spines on thelateral margin of the carapace are very long.

× 54.2 mm, with rhizocephala) (ZRC 2013.1317), northwest
coast of Panglao, Bohol, 120–160 m, coll. J. Arbasto, tangle
nets, 1 December 2007. — 1 ovigerous female (63.7 × 50.8mm, with rhizocephala), 1 young female (56.4 × 44.1 mm)(USNM), northwest coast of Panglao, Bohol, coll. J. Arbasto,
tangle nets, 2006. — 1 male (39.7 × 29.6 mm, photographed)(ZRC 2011.0045), northwest coast of Panglao, Bohol, 80–300
m, coll. J. Arbasto, January-March 2011.
Diagnosis. Pseudorostral horns very long (Fig. 21E, F).
Hepatic, lateral and branchial spines very long; median row
with 4 or 5 spines: 2 or 3 gastric, 1 cardiac, 1 intestinal; 2strong spines on posterior carapace margin (Fig. 21E, F).
Surface of thoracic sternum distinctly granulated (Fig. 50B).
Chela of adult male slender, without distinct carina (Fig.
54B). Ambulatory meri in adults relatively slender, long
(Fig. 21E, F). G1 more strongly curved especially distally(Fig. 23D–F).

Paramaya ouch is superficially similar to P. spinigera but
can be distinguished easily by the following characters: thespines are proportionately much longer in P. ouch than in
P. spinigera (Fig. 21E, F versus Fig. 21A–D); the antorbital
spine is relatively shorter and not as curved as in P. ouch(Fig. 37B) than in P. spinigera (Fig. 37A); and the G1 of P.
ouch is more distinctly curved distally (Fig. 23D–F) (versus
less curved in P. spinigera, Fig. 23A–C).

Etymology. The name is derived from the old English word
for jewel and ornament, alluding to the beautiful arrangementof red spines and tubercles of the species. It also indirectly
alludes to the exclamation for pain, associated with carelesshandling of the present spiny species. Used as a noun inapposition.

The structure of the chela is interesting because in P.
spinigera, the adult male chela is distinctly inflated andthe margins are clearly carinated (Fig. 54A). In adult P.
ouch, the male chela is not inflated and there is no distinct
carina, although in some of the largest males, the proximal
part appears to be weakly carinated (Fig. 54B). Whether the158

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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].

Diagnosis. Pseudorostral horns very long (Fig. 22A). Hepatic,lateral and branchial spines very long; median row with 5
spines: 3 gastric, 1 cardiac, 1 intestinal; 2 strong spines onposterior carapace margin (Fig. 22A). Surface of thoracic
sternum relatively smooth, with very low rounded granules(Fig. 50C). Chela of adult male slender, without carina (Fig.
54C). Ambulatory meri in adults relatively slender, long
(Fig. 22A). G1 gently curved (Fig. 23G–I).

carina is associated with size or dominance, or is a usefulspecies character cannot be ascertained.
Paramaya ouch is very common on the reef slopes ofBalicasag Island. The large series of males, females andjuveniles were useful to observe the variation of spines
with size.
Paramaya coccinea n. sp.
(Figs. 22A, 23G–I, 37C, 40C, 42B, 45L, 50C, 54C, 56B,
70B)

Etymology. The name is derived from the Latin for scarlet,
alluding to the bright red colour of the species when alive.

Material examined. Holotype: male (69.0 × 55.6 mm)(MNHN), northeast of Malo Island, near Santo, 201–212
m, 15°39.0’S 167°15.8’E, Vanuatu, coll. RV Alis, SANTO
2006, 5 October 2006.

Remarks. Paramaya coccinea n. sp. is close to P. ouch n.
sp., with both species having long pseudorostral spines andvery long spines on the margins of the carapace. There areseveral differences. The surfaces of the thoracic sternum
are granular in P. ouch (Fig. 50B) but smooth to almost
smooth in P. coccinea (Fig. 50C). The ambulatory legs159

Ng & Richer de Forges: Revision of the spider crab genus Maja
are also longer and relatively more slender in P. coccinea(Fig. 22A) than in P. spinigera (Fig. 21A–D). The G1 of
P. coccinea resembles that of P. spinigera (Figs. 23A–D,
G–I), and both differ from that of P. ouch which is morestrongly curved distally (Fig. 23D–F). The live colorationis also different: totally red in P. coccinea (Fig. 70B) but
red and cream in P. ouch (Fig. 70A).

Female abdomen dome-shaped, covering most of thoracic
sternum. G1 long, relatively stout, almost straight or gently
curved, distal part with scattered short setae (Fig. 27).
Type species. Maia miersii Walker, 1887, by present
designation.
Etymology. The name is derived from an arbitrary
combination of the name Holthuis, with the genus nameMaja. This honours the major contribution by the late Lipke
B. Holthuis in resolving the convoluted nomenclature of thegenus. Gender feminine.

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

Remarks. Six species are included in Holthuija n. gen.
H. miersii (Walker, 1887), H. suluensis (Rathbun, 1916),
H. pauli n. sp., H. cognata n. sp., H. aussie n. sp. and H.
poorei n. sp. These species are all easily distinguished bytheir prominently foliate supraorbital eave, which is themost expanded in all the genera treated here. All species
also have a strongly granulated carapace surface, epistome,
suborbital and pterygostomial regions, third maxilliped,
thoracic sternum and abdomen. The suborbital tooth is also
sharply separated from the postorbital tooth by a distinct
fissure; and the G1 is generally simple, with a simple distal
part and only scattered short setae.
Key to species of Holthuija n. gen.
1. Carapace with 2 median gastric spines (e.g., Fig. 26A).........2– Carapace with single median gastric spine (e.g., Fig. 26C)...3
2. Carapace with relatively small lateral spines, 1 cardiac spine(Fig. 24E); pseudorostral spines directed horizontally (Fig.
26A); Philippines.............Holthuija suluensis (Rathbun, 1916)– Carapace with relatively long lateral spines, 2 or 3 cardiac spines(Fig. 25B–D); pseudorostral spines gently curved downwards(Fig. 26D); Eastern Indian Ocean......... Holthuija aussie n. sp.
3. Carapace with relatively short lateral spines, pseudorostralspines almost straight or gently curved outward, appears levelfrom lateral view and not prominently curved upward (Figs.
24A–D, 25A)............................................................................4– Carapace with relatively long lateral spines, pseudorostralspines strongly curved outward and usually distinctly bent or
curved upward (Figs. 24F, 25E, F, 26B, E, F).......................5
4. Pseudorostral spines gently curved outward (Fig. 24A–D);
G1 strongly curved with small flap distally (Fig. 27A–C);
Singapore.............................. Holthuija miersii (Walker, 1887)– Pseudorostral spines almost straight (Fig. 25A); G1 almost
straight with distal part dilated (Fig. 27I–K); Japan.................
............................................................ Holthuija cognata n. sp.
5. Dorsal surface of carapace with low granules (Fig. 24F); G1
almost straight with distal part slightly dilated (Fig. 27F–H);
Philippines, Taiwan.................................Holthuija pauli n. sp.
– Dorsal surface of carapace with distinct rounded granules (Fig.
25E, F); G1 curved, distal part simple (Fig. 27L–O); Eastern
Indian Ocean.........................................Holthuija poorei n. sp.

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Holthuija miersii (Walker, 1887)(Figs. 2E, 24A–D, 27A–C, 37D, E, 40D, E, 42C, 46A,
50D, E, 52I, 54D, 56C, 69C, D)

Comparisons of material from Japan and Singapore show thatwhile they are superficially resemble each other in having
a relatively elongate carapace and similar arrangement ofspines, their G1 structures are very different. The G1 of H.
miersii s. str. is medially curved while that of the Japanesematerial is straight. In addition the two median spines on theposterior carapace margin are always short in the Japanese
specimens while it is proportionately longer and stronger in
H. miersii s. str. In addition, the subhepatic, pterygostomianand thoracic sternal surfaces are relatively more strongly
granulated in H. miersii s. str. (Fig. 50D, E) compared to
the less granulated condition in the Japanese material (Fig.
50H). The Japanese material previously identified as “Majamiersi” is here referred to Holthuija cognata n. sp. The G1
structure of H. cognata actually resembles that of H. paulin. sp., but the two species have different carapace features.

Maia miersii Walker, 1887: 109, pl. 6, figs 1–3.
Maja miersi – Serène, 1968: 57. – Serène & Lohavanijaya, 1973:
50 (key).
Maja miersii – Rathbun, 1924: 6. – Griffin, 1966a: 284 (key). –Griffin & Tranter, 1986: 217. – Ng et al., 2008: 117.

Material examined. Singapore – 1 female (21.4 × 16.7 mm)(NHM 1900.10.22.13), Pasir Panjang, near Sungei Pandan,
sandy mud, 6 fathoms, coll. F.P. Bedford & W.F. Lanchester,
1900. — 1 male (32.6 × 25.6 mm) (ZRC 2000.1497), Southern
Islands, dredge, coll. D. Lane, 1991. — 2 females (14.7 ×
10.7 mm, 16.5 × 12.2 mm) (ZRC 1995.311), dredged off
Pulau Semakau, coll. P.K.L. Ng, 1991. — 1 male (20.1
× 14.7 mm) (ZRC 1985.325), Damar Laut, station B54,
5–8 fms, mud with gorgonians, coll. Singapore Fisheries
Research Station, 1955–56. — 1 male (22.3 × 16.5 mm)(CBM-ZC4001), southern Singapore, coll. T. Komai, dredge,
9 May 1997. — 1 ovigerous female (50.2 × 40.4 mm) (ZRC
2013.1272), station 4714-TB2, Semakau Island, dredge,
coll. Comprehensive Marine Biodiversity Survey team, RV
Galaxea, 15 May 2013.

Johnson (1958: 115) had checked on the type specimens of
the four decapod species Walker described from Singapore,including Maia miersii, and noted that “none of them ispresent in there at the moment”. With Paul Clark, the second
author searched the collections of NHM for Walker’s typemale but could not locate the specimen. It is probably nolonger extant, but a neotype is not necessary in light of the
distinctive features of the species described here.

Diagnosis. Carapace covered with small granules (Fig.
24A–D). Pseudorostral spines relatively short, diverging,curved outwards (Fig. 37D, E). Antorbital spine long, sharp;
intercalated spine triangular, sharp; postorbital spine very
long, sharp, slightly curved upwards; hepatic area with
2 sharp spines pointed outwards, first one longest (Figs.
24A–D, 37D, E). Lateral margin with 3 spines; 1 sharp
branchial spine; 2 median spines: 1 gastric, 1 cardiac; 2spines on posterior carapace margin (Fig. 24A–D). Basalantennal article granulated, with 2 long blunt spines distally(Fig. 40D, E). Epistome with 2 large tubercles on anterior
margin (Fig. 40D, E). G1 strongly curved with small flap
distally (Fig. 27A–C).

The recent specimen of H. miersii (ZRC 2013.1272) (Fig.
69C, D) was collected inside a large rock from around 20
m of water, in an area that is very rocky with very strong
currents. This may be the preferred habitat of the species.
In life, the species is dull brown with patches of purple.
The ambulatory dactyli are reddish brown; with parts of theventral surfaces and abdomen purple (Fig. 69C, D).
Holthuija cognata n. sp.
(Figs. 25A, 27I–K, 37H, 46D, 50H, 54G, 56E)
Maja miersii – Muraoka, 1998: 27.
Maja miersi – T. Sakai, 1938: 298–299, pl. 38 fig. 2. – T. Sakai,
1976: 237, pl. 82, fig. 3. – Miyake, 1983: 47, pl. 16 fig. 1.
– Huang, 1994: 582. – Muraoka, 1998: 27. – Maramura &Kosaka, 2003: 34.

Remarks. Walker (1887) described Maia miersii fromSingapore on the basis of a male 33 mm in carapace length(excluding pseudorostrum) (Fig. 24A). The largest male fromSingapore examined here (ZRC 2000.1497) agrees well withhis description and figures. The largest known specimen isa recently collected female specimen and differs from thesmaller specimens in several features: the antorbital spine
is directed somewhat more upwards, the posterior carapace
spines are pointed more upwards, the ocular peduncle isrelatively longer and more prominently curved upwards,
it has two rows of large rounded granules starting fromthe basal antennal article to the epistome which are more
prominent, the suborbital tooth is more strongly granulated,
and the ambulatory legs are more setose. These differences
are not significant and probably associated with its large size.

Material examined. Holotype: male (29.5 × 23.7 mm)(CBM-ZC3662a), off Minabe, Kii Peninsula, 10–20 m, Japan,
gill net, coll. T. Komai, 28 March 1997. Paratypes: Japan –1 male (32.2 × 24.4 mm), 2 females (36.5 × 27.8 mm, 36.6
× 28.2 mm) (CBM-ZC3662b), same data as holotype. — 1
female (30.2 × 23.6 mm) (CBM-ZC4119), Arita, Kushimoto,Kii Peninsula, in lobster net, coll. S. Yamaguchi, 19 February
1978. — 1 female (26.3 × 19.1 mm) (NSMT-Cr 15470), northof Kuchinoshima Island, 30°5.77’N 130°4.54’E, 96m, coll.
H. Saito, 28 May 1999. — 2 males (29.7 × 25.4 mm, 31.0
× 24.7 mm with broken rostrum) (NHM 1961.11.13.5–6),
Seto, Shirahama, Japan, coll. gill nets, I. Gordon & Harada,
1950s. — 1 male (ZRC), Honshu, Wakayama, Kii Minabe,
33°45.712’N 135°18.927’E, coll. K. Sakai, 26 October 1988.
— 1 male (31.9 × 26.0 mm), 1 female (38.4 × 30.0 mm)(SMF 47733), Honshu, Kii-Minabe, Wakayama, 33°45.712’N
135°18.927’E, coll. K. Sakai, 26 October 1988. — 1

Tune Sakai (1938) first recorded this species as “Maja miersi”
from Japan, and because material from Singapore (andSoutheast Asia) has not been reported since, the Japanese
accounts by T. Sakai and his colleagues (see synonymy for
next species) are generally the ones used to define this taxon.
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female (42.7 × 33.0 mm) (SMF 47591), Odawara Harbour,
35°14.265’N 139°9.058’E, coll. H. Watabe, 3 August 1970.
— 1 male (39.4 × 30.2 mm) (SMF 47737), Shikoku, Tosa
Bay, Ashizuri, 32°43.55’N 133°5.028’E, coll. K. Sakai,
13–14 November 1965. — 1 dried male (36.8 × 29.0 mm),
1 dried female (38.0 × 29.6 mm) (SMF 47730), Honshu,
Wakayama, Kii Minabe, 33°45.712’N 135°18.927’E, coll.
K. Sakai, 15 February 1983.

spine; median row with 3 long spines: 2 gastric, 1 cardiac; 2small spines on posterior carapace margin (Fig. 24E). Basalantennal article broad with median row of granules, 2 sharp
distal spines (Fig. 40F). Epistome with 2 granulated lobes
on anterior margin (Fig. 40F). G1 almost straight, distal partnot dilated or curved [juvenile] (Fig. 27D, E).
Remarks. Rathbun (1916: 553) provided measurements for
only one female specimen (the holotype) but did not mention
how many more specimens she had. In the bottle are twoother female specimens with the same data and they shouldbe treated as paratypes. Rathbun (1916: 552) stated that the
specimen was collected from 9 fathoms depth but the labelsays 46 fathoms. Griffin (1976: 200) said 9 to 28 fathoms.

Diagnosis. Carapace covered with small granules (Fig. 25A).
Pseudorostral spines almost straight, diverging (Fig. 37H).
Antorbital spine long, curved forwards; intercalated spine
short, triangular leaving gaps on each side; postorbital spine
strong, straight, enlarged proximally; hepatic spine shorter
than postorbital spine, pointed outwards (Figs. 25A, 37H).
Lateral margin with 3 low spines, last one longest; 1 branchialspine; median row with 2 spines: 1 gastric, 1 cardiac; 2small spines on posterior carapace margin (Fig. 25A). Basalantennal article granulated, with 2 long blunt spines distally.
Epistome with 2 distinct tubercles on anterior margin. G1
almost straight with distal part dilated (Fig. 27I–K).

The record by Griffin & Tranter (1986: 218) from Sulu
Islands and Kei Islands (Moluccas) almost certainly contains
two species. They comment that the specimen they had
from Moluccas has only one gastric spine in contrast to thematerial from Sulu Islands which has two spines (including
the types). They suggested that this character may be variable
and used other characters to differentiate H. suluensis from
H. miersi. While the characters they use are valid, we are
more inclined to believe that his Moluccas specimen is
actually another species, possibly allied to H. pauli n. sp.
(see below). Holthuija suluensis and H. pauli are superficiallyvery close, but the large series of H. pauli on hand tells usthat the position and number of gastric spines (two versusone) is very reliable and not variable, even between sexes or
specimens very different in size. Otherwise, they are similarin general carapace form.

Etymology. The species name alludes to the close relationshipof the new species with H. pauli n. sp. and H. miersii.
Remarks. Although the carapace of H. cognata n. sp. isclose to H. miersii, the G1 structure of H. cognata (Fig.
27I–K) is closest to H. pauli n. sp. (Fig. 27F–H), differingslightly only in proportions. Old records of “Maja miersi”
from Japan and the East China Sea (Huang, 1994) are almost
certainly H. cognata instead. See discussion for H. miersii.
Holthuija suluensis (Rathbun, 1916)(Figs. 1A, 24E, 26A, 27D, E, 37F, 40F, 42D, 46B, 50F,
54E, 56D)

Serène & Vadon (1981: 119, 124, 128) records “Majamiersi” from the Philippines but their material will need to
be checked to see if they are really this species or perhaps
the allied H. pauli or H. suluensis. Maramura & Kosaka’s(2003: 35) record of “Maja aff. suluensis” from Japan will
also need to be examined before its identity can be confirmed.

Maja suluensis Rathbun, 1916: 552. – Serène, 1968: 57. – Griffin,
1976: 200, Fig. 7a. – Griffin & Tranter, 1986: 210, 218 (part),
fig. 71f. – Ng et al., 2008: 117 (list).

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

Material examined. Holotype: female (32.4 × 41.2mm) (USNM 48224a), station 5165, Observation Island,
Tawi Tawi Group, Sulu Islands, 4°58.33’N 119°50.5’E,
Philippines, coll. RV Albatross, 24 February 1908. Paratypes:
2 females (22.5 × 30.6 mm, 22.4 × 30.9 mm) (USNM
48224a), same data as lectotype. Others: Philippines – 1
male (11.5 × 8.2 mm), 1 young female (26.8 × 20.8 mm)(USNM 48507), station 5557, Jolo Island and vicinity, 19
fathoms, coll. RV Albatross, 18 September 1909. — 1 female(36.3 × 27.4 mm) (USNM 49697), station 5163, Tawi Tawi
Group, Sulu Archipelago, 28 fathoms, coll. RV Albatross,
24 February 1908.

Material examined. Holotype: male (37.3 × 28.0 mm)(NMCR), Balicasag Island, Panglao, Bohol, Philippines, 200–
300 m, coll. fishermen in tangle nets, July 2003. Paratypes:
Philippines – 10 males (27.0 × 19.7 mm, 27.6 × 20.2 mm,
27.9 × 20.3 mm, 30.6 × 22.8 mm 32.4 × 24.0 mm, 33.2 ×
24.1 mm, 34.2 × 26.1 mm, 34.4 × 25.3 mm, 34.5 × 25.9mm, 41.0 × 30.8 mm,), 3 females (41.9 × 31.4 mm, 41.9 ×
32.4 mm, 34.8 × 26.2 mm), 1 ovigerous female (34.7 × 26.4mm), 1 damaged female (28.0 × 20.8 mm) (ZRC 2013.1371),
same data as holotype. — 1 female (27.8 × 21.5 mm) (ZRC
2001.411), Balicasag Island, Panglao, Bohol, coll. fishermenwith tangle nets, December 2000. — 1 male (ZRC 2013.1266,
ex part ZRC 2001.0577), Balicasag Island, Panglao, Bohol,
coll. fishermen with tangle nets, 28 November 2001. — 4
males (27.5 × 19.9 mm, 28.2 × 20.7 mm, 31.5 × 23.4
mm, 31.9 × 23.8 mm), 1 ovigerous female (35.0 × 26.0mm) (ZRC 2001.0590), Balicasag Island, Panglao, Bohol,

Diagnosis. Carapace covered with small granules (Fig. 24E).
Pseudorostral spines straight, strongly diverging (Fig. 37F).
Antorbital spine long, curved, directed anteriorly; intercalated
spine short, leaving narrow gaps on each side; postorbital
spine very long, slightly curved outwards; hepatic spine
directed anteriorly, half length of postorbital spine (Figs.
24E, 37F). Lateral margin with 3 long spines; 1 branchial
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Remarks. Of the Holthuija species, H. pauli n. sp. is
distinct in possessing very long carapace spines which areprominently curved obliquely upwards. The large numberof specimens we have examined show that the length andorientation of these spines are not variable, even for small
specimens and females. It also has proportionately, thelongest ambulatory legs (Fig. 24F). In the other species, theambulatory legs, particularly the merus, are shorter (Figs.
24A–D, 25).

coll. fishermen with tangle nets, 28 November 2001. — 1
female (23.8 × 17.6 mm) (ZRC 2013.1376), Balicasag
Island, Panglao, Bohol, coll. fishermen in tangle nets, June
2002. — 1 juvenile female with bopyrid (ZRC 2013.1314),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, June 2002. — 1 male (35.0 × 27.0 mm)(ZRC 2013.1378), Balicasag Island, Panglao, Bohol, coll.
fishermen in tangle nets, June 2002. — 1 male, 2 females(1 ovigerous) (NSMT-Cr 15392), Balicasag Island, Panglao,
Bohol, coll. local fisherman via M. Takeda & H. Komatsu,
February 2003. — 2 females (33.8 × 25.3 mm, 40.7 × 30.7
[carapace badly damaged]) (ZRC 2013.1375), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets,
July 2003. — 1 damaged male (19.5 × 15.6 mm) (ZRC
2013.1377), Balicasag Island, Panglao, Bohol, coll. fishermenwith tangle nets, July 2003. — 1 female (38.8 × 28.9 mm)(ZRC 2013.1379), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, 25–30 July 2003. — 1 female(35.5 × 27.2 mm) (ZRC 2013.1380), Balicasag Island,
Panglao, Bohol, coll. fishermen with tangle nets, November
2003. — 2 males (AM), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, January 2004.
— 1 male (ZRC 2013.1324), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, February
2004. — 1 female (19.9 × 14.7 mm) (ZRC 2013.1384),
Balicasag Island, Panglao, Bohol, coll. fishermen with tangle
nets, March 2004. — 1 male, 2 females (ZRC 2013.1222),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, April 2004. — 1 male (29.0 × 21.9 mm)(ZRC 2013.1373), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, May 2004. — 4 males (smallest
23.4 × 17.4 mm, largest 42.0 × 31.8 mm), 1 female withrhizocephalan (28.2 × 21.4 mm), 2 females (33.4 × 25.7mm, 45.9 × 35.0 mm) (ZRC 2013.1372), Balicasag Island,
Panglao, Bohol, coll. fishermen with tangle nets, 29 May
2004. — 1 female (ZRC 2013.1374), Balicasag Island,
Panglao, Bohol, 200–300 m, coll. fishermen with tangle
nets, 29 February 2005. Others: 1 badly damaged male(21.5 × 17.8 mm) (ZRC 2013.1323, ex part ZRC 2000.585)
Balicasag Island, Panglao, Bohol, coll. fishermen with tangle
nets, 28 November 2001.

The G1 structure of H. pauli most closely resembles that of
H. cognata n. sp., with the distal part laterally flattened, eventhough their carapace and legs are quite different (carapace
relatively broader with longer and more curved spines, andwith proportionately longer legs in H. pauli). The G1 of H.
pauli, however, is relatively more slender than that of H.
cognata (Fig. 27F–H versus Fig. 27I–K).
Holthuija pauli might be confused with small specimens of
Paramaya spinigera and Paramaya ouch, especially withregard to the relatively long carapace spines which are curved
and the very divergent pseudorostral spines. Paramaya
species, however, have long distal spines on the meri of
ambulatory legs (absent in H. pauli) as well as many othercharacters at the generic level (see discussion for Paramaya).
In life, H. pauli is beige and light reddish-brown overall,with the third maxillipeds and abdomen brighter in colour(Fig. 69E, F).
Holthuija aussie n. sp.
(Figs. 25B–D, 26D, 37I, 40H, 42E, 46E, 54H, 56F)
Maja suluensis – Poore et al., 2008: 62. (not Maja suluensis Rathbun,1916). (not Maja suluensis Rathbun, 1916).

Material examined. Holotype: ovigerous female (42.1 × 34.4mm) (NMV J63752), Arafura Sea, 10°58.03’S 136°47.82’E,
107–108 m, coll. B. A. Glasby, CSIRO RV Southern
Surveyor, 17 October 2012. Others: 1 juvenile female (14.3 ×
11.9 mm [right branchial region with bopyrid parasite) (NMV
J61058), Imperieuse L23 Transect, northwestern Australia,
18°27.62’S 120°08.68’E – 18°27.72’S 120°08.68’E, 80–81
m, coll. M. Gomon, CSIRO RV Southern Surveyor, 19 June
2007. — 1 juvenile female (11.9 × 9.1 mm) (NMV J63958),
off Red Bluff, Western Australia, 24°02.62’S 113°01.62’E– 24°02.83’S 113°01.73’E, 100 m, coll. G. Poore, CSIRORV Southern Surveyor, 8 December 2005.

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

Diagnosis. Carapace covered with small granules (Fig.
25B–D). Pseudorostral spines long, diverging (Fig. 37I).
Antorbital spine long, sharp, strongly curved upwards andanteriorly; intercalated spine triangular, with distinct gaps oneach side; postorbital spine long, strong; hepatic spine short;median row with 5 long spines: 2 gastric, 3 cardiac (2 may
be small in young specimens) (Figs. 25B–D, 37I). Lateral
margin with 3 long spines increasing towards posterior; 1very long branchial spine; 2 long upwardly directed spines
on posterior carapace margin (Fig. 25B–D). Basal antennal
article with 2 big blunt distal teeth; on inner margin 2 lobate

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

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teeth; external margin serrulated (Fig. 40H). Epistome with
2 tubercles on anterior margin (Fig. 40H). G1 not known.

the species comes from the northwestern Australian coast,in the Arafura Sea, very close to Indonesian waters.

Etymology. The species is named after a common nickname
for Australians – “Aussie”. The name is used as a noun inapposition.

The identities of two juvenile specimens (J61058 and J53958)
collected at 18°S and 24°S, is uncertain as they differ fromthe type in several aspects. The following differences areobserved: the antorbital spine is not as curved in the juveniles
as found in the adults; there are wide gaps on each side of
the intercalated spine in the juveniles versus narrow gaps in

Remarks. Holthuija aussie n. sp. is the only species of theMaja group to possess three cardiac spines. The holotype of

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.

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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.

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Fig. 26. Lateral views of carapaces, Holthuija species. A, H. suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines;
B, H. pauli n. sp., holotype male (37.3 × 28.0 mm) (NMCR), Philippines; C, H. pauli n. sp., paratype male (35.0 × 27.0 mm) (ZRC
2013.1378), Philippines; D, H. aussie n. sp., holotype ovigerous female (42.1 × 34.4 mm) (NMV J63752), Arafura Sea; E, H. poorei n.
sp., holotype male (24.3 × 18.7 mm) (NMV J63749), Timor Sea [laterally inverted to match earlier figures]; F, H. poorei n. sp., paratype
female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea laterally inverted to match earlier figures].

m, coll. B.A. Glasby, CSIRO RV Southern Surveyor, 17
October 2012.

the adults; all the lateral spines are shorter in the juvenile
specimens; only one cardiac spine versus 3 in the adult;
the distal spines on the basal antennal article are shorterand not so sharp; the cornea shape is different, being moreelongate in the juveniles. They could belong to a new species
but as neither specimens are adult, it was decided that thiswas best left to a later date when better material becomes
available. They are here provisionally referred to Holthuija
aussie for convenience.

Diagnosis. Carapace covered with relatively larger rounded
granules (Fig. 25E, F). Pseudorostral spines long, strongly
diverging (Fig. 37J). Antorbital spine long, straight;
intercalated spine strong, with convex margins, with large
gaps on each side; postorbital spine long; hepatic spine short,pointed outwards; median row with 2 spines, 1 gastric, 1
cardiac; 2 small spines on posterior carapace margin (Figs.
25E, F, 37J). Lateral margin with 3 large spines, increasing insize towards posterior; 1 long branchial spine, curved upwards(Fig. 25E, F). Basal antennal article with 2 big blunt distalteeth, external margin smooth, carinated, internal margin
with one proximal lobate tooth (Fig. 40I). Epistome with
2 low tubercles on anterior margin (Fig. 40I). G1 curved,
distal part simple (Fig. 27L–O).

Holthuija poorei n. sp.
(Figs. 25E, F, 26E, F, 27L–O, 37J, 40I, 46F, 50I, 54I,
56G)
Material examined. Holotype: male (24.3 × 18.7 mm) (NMV
J63749), Timor Sea, Ocean Shoals Commonwealth Marine
Reserve, 12°04.5’S 127°26.62’E – 12°04.53’S 127°26.62’E,
59–66 m, coll. B.A. Glasby, RV Solander, 3 October 2012.
Paratypes: 1 male (26.1 × 20.1 mm) (NMV J63196), samedata as holotype. — 1 female (27.0 × 22.5 mm) (NMV
J63751), Arafura Sea, 10°58.03’S 136°47.82’E, 107–108

Etymology. The species is named after our “Aussie” friend,
Gary Poore, former curator of Crustacea at Museum Victoria.

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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].

Remarks. Holthuija poorei n. sp. resembles H. pauli n.
sp. in many aspects, but is easily separated by possessing
a distinctly more granular carapace (Fig. 26E, F versus
Fig. 25E, F). Holthuija poorei also has relatively shorter
ambulatory legs (Fig. 25E, F versus Fig. 24F), and the G1 isrelatively more elongated and curved with the tip distinctly
dilated (Fig. 27L–D versus Fig. 27F–H).

The material of H. poorei n. sp. was collected off northwestern
Australian coast, in the Timor Sea and Arafura Sea, both at
the boundary of Indonesian waters.
Sakaija n. gen.
Diagnosis. Carapace pyriform to rounded in adults; dorsal
surface inflated, covered by rounded granules; gastric andbranchial regions delimited by grooves (Figs. 28–30, 31A,
167

Ng & Richer de Forges: Revision of the spider crab genus Maja
Remarks. Sakaija gen. nov. is established to accommodate
the following species: Sakaija japonica (Rathbun, 1932) fromJapan, S. sakaii (Takeda & Miyake, 1969) from Japan, S.
serenei n. sp. from Philippines, S. santo n. sp. from Vanuatu,
S. africana (Griffin & Tranter, 1986) from South Africa, andS. longispinosa n. sp. from the western coast of Australia.

B, D, E, 32). Intestinal region with small median spine,often reduced to blunt tubercle (Figs. 28–30, 31A, B, D, E,
32). Pseudorostral spines relatively short, diverging,(Figs.
28–30, 32, 37K–O). Supraorbital eave with anterior part
longitudinally narrow, rectangular, not prominently expanded;
antorbital spine sharp (Fig. 37K–O). Intercalated spinedistinct, separated from supraorbital eave and postorbital
spine by distinct gaps; postorbital spine strong, lobiform;
hepatic region with 1 spine, shorter than postorbital spine; 1or more smaller spines below (Fig. 37K–O). Lateral carapace
margin with 2 short spines and numerous granules, somesharp; branchial region with 1 spine (Figs. 28–30, 31A,
B, D, E, 32). Posterior carapace margin with 2 very shortmedian spines, sometimes almost undiscernible in adults(Figs. 28–30, 32). Eyes relatively short, slender, with ovoid
cornea (Fig. 37K–O). Antennal flagellum short, slender.
Basal antennal article longer than broad, rectangular; surface
granulated, with 2 spines distally; inner and outer lateral
margins with low granules; proximal outer angle rounded;
antero-external crested rim of antennular fossa touches but
does not significantly overlap distal part of basal antennal
article (Figs. 28F, 40J–M). Epistome slightly wider thanlong, anterior margin unarmed; posterior margin composedof 4 rectangular plates separated by shallow fissures (Figs.
28F, 40J–M, 42F). Suborbital margin separated from basal
antennal article by short fissure, confluent with marginof postorbital tooth (Fig. 40J–M). Outer surface of third
maxilliped covered by scattered setae in adults; ischium
subrectangular, just longer than broad; postero-externalangle of merus relatively broad, “inserted” into shallower
concavity on outer margin of ischium; antero-internal part ofischium rounded, auriculiform (Fig. 46G–L). Male chelipeds
relatively short in adult males, surfaces of merus and carpus
smooth; carpus short, with low longitudinal ridge; propodusof palm smooth, short, inflated in adults, palm longer thanfingers; fingers slender, gently curved, with distinct basal
gape when closed (Figs. 28–30, 32, 54J–M). Ambulatory legs
slender or stout, slender; merus without dorsal subdistal spine;
dactylus elongate, curved, covered with long setae except forcorneous tip (Figs. 28–30, 31C, F, 56H–J). Thoracic sternum
wide; surfaces of somites 5–8 with numerous prominent
rounded tubercles and granules; sternites 3 and 4 distinctly
depressed; margin between sternites 2 and 3 demarcated bydeep notch; anterior margin of sterno-abdominal cavity notforming complete rim (Figs. 51A–E, 52J). Male abdomen
subrectangular, with 6 free somites and telson; somites 3–6
and telson subequal in width (Fig. 51A–E). Male pressbutton abdominal locking mechanism submedian in position
on sterno-abdominal cavity (Fig. 52J). Female abdomen
dome-shaped, covering most of thoracic sternum. G1 short,
slender, gently curved, distal part slightly spatuliform, tip
rounded, margins lined with numerous long setae (Fig. 33).

The primary diagnostic feature for members of this genus is
the characteristic slender and sinuous or curved G1, the distal
part of which is lined with numerous long setae. Membersof Sakaija, however, can also be recognised by a suite ofcharacters: a distinctly pyriform carapace that has relatively
short pseudorostral spines, the suborbital margin confluent
with the margin of the postorbital tooth, and shape of themale thoracic sternum and abdomen. Noteworthy is also that
all the species are relatively small, the largest being about
30 mm carapace length (S. africana), and small species
maturing at carapace lengths of only 10 mm (e.g., S. serenei).
Key to species of Sakaija n. gen.
1. Pseudorostral spines very long; posterior carapace margin with
2 spines and long intestinal spine (Fig. 29); G1 gently curved,
distal part with long setae, with low subdistal swelling (Fig.
33N–R); Western Indian Ocean.................................................
.............................. Sakaija africana (Griffin & Tranter, 1986)– Pseudorostral spines not elongate; posterior carapace margin
with 2 short spines, sometimes not discernible, intestinal spine
short or not visible (Figs. 28, 30, 32); G1 otherwise.............2
2. Carapace with very long lateral, branchial and intestinal spines(Fig. 32A, B); Eastern Indian Ocean.........................................
........................................................... Sakaija longispina n. sp.
– Carapace with relatively short lateral, branchial and intestinal
spines (Figs. 28, 30, 32C, D)...................................................3
3. Carapace with 2 subhepatic spines, one of which is small (Fig.
32C, D); G1 sinuous, distal part turned backwards towards
median line of sternum, with very long setae (Fig. 33S–U);
Vanuatu and New Caledonia.....................Sakaija santo n. sp.
– Carapace with single visible hepatic spine (Figs. 28, 30); G1
otherwise...................................................................................4
4. Carapace with posterior part rounded, prominently swollen;
lateral spines low; ambulatory legs relatively short, stout (Figs.
28A–C, 31A–C); G1 distinctly curved, distal part long, taperingwith very long setae (Fig. 33A–G); Japan and Taiwan............
.............................................Sakaija japonica (Rathbun, 1932)– Carapace more evenly ovate in appearance; lateral spines short
but distinct; ambulatory legs slender, more elongate (Figs.
28D-G, 30); G1 otherwise.......................................................5
5. Cardiac spine long, intestinal spine distinct (Fig. 28D, E, G);
G1 gently curved, distal part relatively short, subspatuliform,
lined with long setae (Fig. 33H–J); Japan.................................
................................. Sakaija sakaii (Takeda & Miyake, 1969)– Cardiac and intestinal spines very low, almost undiscernible(Figs. 30, 31D, E); G1 gently curved, distal part long, taperingwith very long setae (Fig. 33K–M); Philippines, New Britain
and Papua New Guinea...........................Sakaija serenei n. sp.

Type species. Maja japonica Rathbun, 1932, by present
designation.
Etymology. The genus name is derived from an arbitrary
combination of the family name Sakai with Maja. It honours
the significant work done by the late Tune Sakai in advancing
our knowledge of Asian Majoidea. Gender feminine.
168

RAFFLES BULLETIN OF ZOOLOGY 2015
Sakaija japonica (Rathbun, 1932)(Figs. 28A–C, 31A–C, 33A–G, 37K, 40J, 42F, 46G, 51A,
B, 52J, 54J, 56H)

28A–C, 31A, B). Basal antennal article long with several
granules, 2 strong distal spines (Fig. 40J). Ischium of third
maxilliped weakly granulated (Fig. 46G). Ambulatory legs
relatively short (Figs. 31C, 56H). G1 distinctly curved, distal
part long, tapering with very long setae (Fig. 33A–G).

Maja sp. – T. Sakai, 1932: 50, pl. 2, fig. 5, text-fig. 5.
Maja japonica Rathbun, 1932: 33. – T. Sakai, 1965: 83, pl. 37, fig.
1. – Serène, 1968: 57. – Takeda & Miyake, 1969: 512, pl. 17,
fig. A, B, pl. 18, fig. B. – Takeda, 1973: 42. –Takeda, 1975:
127. – T. Sakai, 1976: 238, pl. 82, fig. 1; text-figs 126a, b. –Miyake, 1983: 44. – Dai et al., 1986: 150, pl. 18(1), text-fig.
76. – Griffin & Tranter, 1986: 216. – Huang, 1989: 339. – Dai& Yang, 1991: 135, pl. 18(1), text-fig. 76. – Takeda, 1993:
43. – Huang, 1994: 582. – Muraoka, 1998: 27. – Ng et al.,2001: 12. – Takeda, 2001: 235. – Maramura & Kosaka, 2003:
34. – Yang et al., 2008: 780. – Ng et al., 2008: 117 (list).
Maja japonica Yokoya, 1933: 157, text-fig. 56.
Maja nipponensis T. Sakai, 1934: 297, text-fig. 11. – T. Sakai,
1936: 100, text-fig. 46. – T. Sakai, 1938: 299, pl. 38, fig. 1,
text-fig. 41.
Maja sakaii – Miyake, 1983: 44, pl. 15 fig. 5.

Remarks. Despite its age, the type specimen is in a relatively
good condition although the G1 appears rather frail and noteasy to study. However, it agrees well with those from Japanand Taiwan, and we are confident of their conspecificity.
For a detailed discussion of its taxonomy, see remarks for
S. sakaii.
In Japan, S. japonica has been reported from depths ofbetween 85 and 120 m. Outside Japan, it has been recorded
from Hong Kong (Griffin & Tranter, 1986), although their
specimens should be re-examined.
Sakaija sakaii (Takeda & Miyake, 1969)(Figs. 28D–G, 33H–J, 46H)

Material examined. Holotype: male (16.2 × 13.0 mm)(USNM 48252), off Honshu, Japan, coll. RV Albatross, 7
May 1900. Others: Japan – 1 male (18.7 × 14.6 mm) (NHM
1961.6.5.120), Sagami Bay, Japan, coll. T. Sakai. — 1 male(with bopyrid) (15.6 × 13.4 mm), 1 female (20.3 × 15.4 mm)(NSMT-Cr 13177), Tosa Bay, coll. RV Kotoka-Maru, 21January 1998. — 1 male (with bopyrid) (NSMT-Cr 1132),
off Tsushima Islands, station 10, 33°57.7’N 129°11.6’E,
sand and shell substrate, coll. Tsushima Expedition, 26 July
1968. — 1 male, 1 female (USNM 120722), off Ashizuri
Peninsula, Shikoku Island, 300 m, coll. T. Sakai & K. Sakai,
27–31 March 1966. — 1 juvenile female (carapace only)(USNM 48483), station 4893, 10–20 miles southwest of Goto
Island, Eastern Seas, 106 fathoms, coll. RV Albatross, 9August 1906. — 1 male, 2 females (SMF), Tosa Bay, Kochi,Usa, coll. RV Toyohata-Maru, K. Sakai. Taiwan – 1 male
(22.3 × 17.8 mm) (ZRC 2013.1267), Tashi fishing port, Ilan
County, northern Taiwan, coll. 11 July 1989. — 1 broken
male (20.8 × 16.7 mm) (ZRC 2013.2371), Tashi fishingport, Ilan County, northern Taiwan, coll. S.-H. Wu, 1997.
— 1 ovigerous female (20.4 × 16.4 mm) (ZRC 2013.1270),
Tashi fishing port, Ilan County, northern Taiwan, coll. S.-H.
Wu, 23 January 1999. — 1 male (22.6 × 17.9 mm) (ZRC
2013.1269), Tashi fishing port, Ilan County, northern Taiwan,coll. S.-H. Wu, 8 March 1999. — 1 male (18.5 × 13.9 mm)(ZRC 2013.1268), Tashi fishing port, Ilan County, northern
Taiwan, coll. Hwang, 18 July 2011.

Maja sp. – T. Sakai, 1932: 50, pl. 2 fig. 5, text fig. 5.
Maja japonica – T. Sakai, 1934: 297, text-figs. 10. – T. Sakai, 1936:99, pl. 25, fig. 2, text-fig. 45. – T. Sakai, 1938: 299, pl. 30, fig.
2. – Takeda, 1982: 128. (not Maja japonica Rathbun, 1932).
Maja sakaii Takeda & Miyake, 1969: 512, pl. 17, fig. C. – Serène& Lohavanijaya, 1973: 50 (key). – T. Sakai, 1976: 237, fig.
125a, b, pl. 85 fig. 2. – Huang, 1989: 339. – Takeda, 1993:
43. – Huang, 1994: 583. – Yang et al., 2008: 780. – Ng et al.,
2008: 117 (list).

Material examined. Japan – 1 male (9.0 × 6.6 mm) (NSMTCr 8094), station 12, Shimoda, Shizuoka Prefecture, coll.
M. Takeda, 9 November 1981.
Diagnosis. Carapace branchial area gently inflated, ovate;with relatively short lateral and branchial spines (Fig. 28D,
E, G). Pseudorostral spines short, diverging (Fig. 28D–G).
Supraorbital eave gently curved; antorbital spine short,triangular, sharp; intercalated spine triangular, separated
from adjacent spines by large gaps; postorbital spine longest,
triangular, pointed anteriorly; hepatic spine short, pointed
outwards (Figs. 28A–C). Lateral margin with 2 short spines,
posterior one larger; branchial region with 1 spine; medianrow with 4 low spines or tubercles: 2 gastric (very low), 1
cardiac (long), 1 intestinal (low); 2 short spines on posterior
carapace margin (Fig. 28D, E, G). Basal antennal article
long with several granules, 2 strong distal spines (Fig. 28F).
Ischium of third maxilliped weakly granulated (Fig. 46H).
Ambulatory legs relatively long, slender (Fig. 28D, G). G1
gently curved, distal part relatively short, subspatuliform,
lined with long setae (Fig. 33H–J).

Diagnosis. Carapace branchial area strongly inflated,rounded; with relatively short lateral and branchial spines(Figs. 28A–C, 31A, B). Pseudorostral spines short, diverging(Fig. 37K). Supraorbital eave strongly curved; antorbital
spine short, triangular, sharp; intercalated spine triangular,
separated from adjacent spines by large gaps; postorbital
spine longest, triangular, pointed anteriorly; hepatic spine
short pointed outwards (Figs. 28A–C, 37K). Lateral margin
with 2 short spines, posterior one larger (often becoming
tubercles in larger specimens); branchial region with 1 low
spine or tubercle; median row with 4 low spines or tubercles:
2 gastric (posterior one larger), 1 cardiac, 1 intestinal; 2short spiniform granules on posterior carapace margin (Figs.

Remarks. The identities of S. japonica and S. sakaii havebeen confusing. Tune Sakai (1932) first reported a 14.5 by
13.0 mm male specimen as “Maja sp.” from Sagami Bay
that he thought was close to Maja japonica Rathbun, 1932,
but believed it to be a juvenile, and hence was uncertainof its identity. Tune Sakai (1934) later realised that there
were two forms in Japan and both Rathbun (1932) and169

Ng & Richer de Forges: Revision of the spider crab genus Maja
Yokoya (1933) had used the same name “Maja japonica”
independently of each other. Tune Sakai (1934), however,identified one male specimen from the East China Sea as“M. japonica Rathbun, 1932”, and another female specimen
from Nagasaki as “M. japonica Yokoya, 1933”. He notedthat their carapaces, ambulatory leg and G1 structures
were starkly different. Believing that his “M. japonica”
was conspecific with Rathbun’s (1932) species, and since
the name Maja japonica Yokoya, 1933, was preoccupied
by Maja japonica Rathbun, 1932, T. Sakai (1934) gave areplacement name, Maja nipponensis for Yokoya’s species(see also T. Sakai, 1936, 1938). Tune Sakai (1965) examined
the type of Maja japonica Rathbun, 1932, and realised it was
actually conspecific with Maja japonica Yokoya, 1933, as
was M. nipponensis T. Sakai, 1934. He also noted that the
species he had identified as “Maja japonica” in 1934 wasan undescribed species, but he did not name it.

has the same kind of G1 although the distal part is slightly
less expanded and has less setae (Fig. 33I, J), differences
easily attributed to their size. The kind of G1 in S. sakaii,
however, is quite different from that of S. japonica. While
the G1 of S. japonica is dorsoventrally flattened, like mostof the G1s of this genus (with the exception of S. santo n.
p.), that of S. sakaii is actually more laterally flattened, suchthat the longitudinal groove is not on the inner margin (asin S. japonica, Fig. 33A–F) but is fully exposed from theabdominal (ventral) view (Fig. 33H–J). In the form of the
G1, S. sakaii is closer to S. santo from Vanuatu and New
Caledonia (Fig. 33S–U) (see discussion for that species).
Sakaija sakaii (like S. japonica) is superficially similar
to Alcomaja nagashimaensis in being a relatively smaller
species in which males have inflated chelipeds. Alcomaja
nagashimaensis can easily be separated not only by thepreviously discussed generic characters, it can also be
distinguished by the carapace possessing proportionately
longer and sharper lateral and median spines; the fingersof the adult chelae are less curved and the basal antennal
article is smoother (Figs. 17A–C, 39O, 41H).

Takeda & Miyake (1969: 512) agreed that what T. Sakai(1934, 1936, 1938, 1965) had originally identified as “Majajaponica” was a separate but unnamed species for which
they chose a new name, Maja sakaii. However, since theyalso designated an ovigerous female measuring 12.9 by 10.0mm from the East China Sea as the holotype of M. sakaii,
they were in effect establishing a new species and not justproviding a replacement name.

The figure of S. sakaii in Miyake (1983) more closely
resembles S. japonica with its rounded carapace and isreferred there. Huang’s (1989, 1994) records from the East
China Sea are probably true S. sakaii. Serène & Vadon’s(1981) record from the Philippines is here referred to S.
serenei. Griffin & Tranter’s (1986) records from the Lesser
Sunda Islands, Moluccas, Ambon and Java Sea are uncertain.
Whether their material is really S. sakaii, or a separate Sakaija
species will have to be determined through a re-examinationof their material.

Sakaija japonica is certainly similar to S. sakaii, and mostof the distinguishing characters originally highlighted by T.
Sakai (1934) still apply (see also Takeda & Miyake, 1969).
The shape of the carapace is diagnostic, with M. japonica
more rounded (Fig. 28A–C) while S. sakaii is distinctly
pyriform (Fig. 28D, E, G). The difference in carapace shape
does not appear to be completely size-dependent in membersof this genus. In S. serenei n. sp., for which a good series of
specimens is available for study, the carapace shape changes
only slightly with larger specimens more rounded (Fig. 30)
but never reaching the condition of S. japonica (Fig. 28A–C).
Takeda & Miyake’s (1969) adult holotype female of S. sakaii
measures 12.9 by 10.0 mm but its carapace is still distinctly
pyriform. Another key character is the proportions of theambulatory legs. In S. japonica, the merus is notably shorterand stouter (Figs. 28A–C, 31C) while those of S. sakaii are
relatively more slender and longer (Fig. 28D, G).

Sakaija africana (Griffin & Tranter, 1986)(Figs. 2F, 29, 33N–R, 37L, 40K, 46I, 51C, 54M, 70C)
Maja africana Griffin & Tranter, 1986: 211, 71a–e, 73c, d, Pl.
14. – Ng et al., 2008: 117 (list).

Material examined. Madagascar – 1 male (28.7 × 22.3mm) (MNHN-IU-2010-57) (specimen barcoded), station 27,
between Nosy Bé and Cap Du Leven, MIRIKY, 12°31’S
48°22’E, 298–301 m, coll. MIRIKY, June 2009. — 1 male(32.2 × 25.4 mm) (MNHN-IU-2010-928), station DW 3228,
260–319 m, 12°55’S 48°11’E, 2 July 2009. — 1 male (26.3
× 19.9 mm), 2 ovigerous females (25.7 × 20.2 mm, 23.8 ×
18.4 mm) (MNHN-IU-2010-505), station DW 3211, 12°32’
47°52’E, 244–300 m, coll. MIRIKY, 30 June 2009. — 1
male (27.3 × 21.2 mm) (MNHN-IU-2010-932), no otherdata, coll. MIRIKY, 25 June–14 July 2009. — 1 male (27.3
× 21.2 mm), 1 ovigerous female (25.7 × 19.9 mm) (ZRC
2013.1408, ex MNHN-IU-2010-504), station CP 3188,
12°31’S 48°22’E, 298–301 m, coll. MIRIKY, 27 June 2009.

The G1 structures are quite different. The G1 structures
figured for the two species in T. Sakai (1934: text fig.
11) show two different structures and have been used inall subsequent papers by T. Sakai (1934, 1938, 1976) toseparate the two species. The G1 of S. japonica provided(as Maja nipponensis; T. Sakai, 1934: text fig. 11b) is thatof a small male from Sagami Bay, measuring 14.5 by 13.0mm (cf. T. Sakai, 1932: 50, 51) (Fig. 33G). In larger males,
the distal part of the G1 is more elongate and has evenmore long setae (Fig. 33A–F). The G1 of S. sakaii figured(as Maja japonica; Sakai, 1934: text fig. 11a), apparently
from a small male, is drawn at a marginal view and froma different angle from that of S. japonica s. str., and shows
an ovate distal part densely lined with setae (Fig. 33H). The
young male specimen of S. sakaii examined in this study

Diagnosis. Carapace branchial area gently inflated, ovate;with relatively short lateral and branchial spines (Fig.
29). Pseudorostral spines very long, diverging (Fig. 37L).
Supraocular eave narrow, strongly curved; antorbital toothstrong, triangular pointed outwards; intercalated spine basally170

RAFFLES BULLETIN OF ZOOLOGY 2015
subtruncate, separated from other spines by narrow gaps;
postorbital spine large, pointed anteriorly; hepatic spine low,
tuberculiform (Figs. 29, 37L). Lateral margin with 2 shortspines, subequal or anterior one slightly smaller; 1 large
branchial spine; median row with 3 low spines: 1 gastric,
1 cardiac, 1 long, posteriorly directed intestinal; posterior
carapace margin with 2 long spines (Fig. 29). Basal antennal
article with 2 large distal spines, external margin with 4
blunt teeth (Fig. 40K). Ischium of third maxilliped almost
smooth (Fig. 46I). Adult ambulatory legs relatively long,
slender (Fig. 29). G1 gently curved, distal part with longsetae, with low subdistal swelling (Fig. 33N–R).

(smallest 13.9 × 11.3 mm, 14.6 × 11.1 mm; largest 17.1 × 13.8mm), 1 female (ZRC 2013.1256), 2 males (larger 18.8 × 15.1mm), 1 female (NSMT-Cr 22330), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, March
2004. — 1 ovigerous female (ZRC 2013.1244), Balicasag
Island, Panglao, Bohol, 200–300 m, coll. fishermen withtangle nets, Balicasag Island, Panglao, Bohol, 200–300 m,
coll. fishermen with tangle nets, May 2004. — 11 males, 14
females (ZRC 2013.1255), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, May 2004. — 2
males (ZRC 2013.1258), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, 29 May 2004.
— 2 males (ZRC 2013.1259), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, 29 May
2004. — 3 males, 2 females (ZRC 2013.1243), station P4,
9°31.1’N 123°41.5’E, Panglao, Bohol, coll. tangle nets fromlocal fishermen, 31 May 2004. — 1 male, 1 female (AM),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, 4 June 2004. — 1 male (19.3 × 15.4 mm),
1 female (15.3 × 12.4 mm) (ZRC 2013.1261), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets, June
2004. — 1 male (17.7 × 14.6 mm) (ZRC 2013.1262), station
1, Maribojoc Bay, Bohol, 9°36.1’N 123°45.0’E, 90–200 m,
coll. tangle nets from local fishermen, 30 May 2004. — 1
ovigerous female (14.4 × 10.7 mm) (ZRC 2013.1242),
northwest Panglao, Bohol, coll. J. Arbasto with tangle nets,
2006. Papua New Guinea – 1 ovigerous female (18.2 × 14.8mm) (MNHN-IU-2011-3898), Jacquinot Bay, New Britain,
station DW 3770, 05°34’S 151°32’E, 220–294 m, coll. RVAlis, BIOPAPUA, 16 October 2010.

Remarks. Sakaija africana was described on the basis of
one male specimen, 24 mm in carapace length, collected off
Durban, Indian Ocean coast of South Africa. It is the largest
known species of Sakaija. This species seems common in
the Mozambique Channel at a depth of around 300 m.
Male and female specimens smaller than the holotype (Fig.
29B, D) generally have relatively shorter ambulatory legs
compared to large males (Fig. 29C); with females also
having proportionately shorter pseudorostral spines. The G1
figured by Griffin & Tranter (1986: fig. 73c, d) appears tohave been drawn in situ and at an angle and not with thestructure lying flat on its dorsal or ventral surfaces. The
two lobes depicted on their fig. 73d (present Fig. 33N, O)
is where the folds overlap (Fig. 33Q, R). They otherwise
agree very well in form and structure.
Sakaija serenei n. sp.
(Figs. 30, 31D–F, 33K–M, 37M, 40L, 46J, 51D, 54K,
56I, 70D)

Diagnosis. Carapace branchial area gently inflated, ovate;with relatively short lateral and branchial spines (Figs. 30,
31D, E). Pseudorostral spines short, diverging (Fig. 37M).
Supraorbital eave distinctly curved with triangular antorbital
spine; intercalated spine shorter, triangular; postorbital spine
longest, slightly curved outwards and upwards; hepatic spine
short (Fig. 37M). Lateral margin with 2 large tubercles or
short spines and several smaller granules; 1 low but strong
branchial spine; median row with 3 tubercles: 2 gastric, 1
cardiac, intestinal not easily discernible; posterior carapace
margin with 2 low tubercles, sometimes not visible (Figs. 30,
31D, E). Basal antennal article covered with granules, with
2 distal blunt spines (Fig. 40L). Ischium of third maxilliped
covered with low granules (Fig. 46J). Adult ambulatory legs
relatively long (Figs. 30, 56I). G1 gently curved, distal partlong, tapering with very long setae (Fig. 33K–M).

Maja japonica – Serène & Lohavanijaya, 1973: 50, figs. 93–97,
pl. 9 fig. A.
Maja sakaii – Serène & Vadon, 1981: 124.

Material examined. Holotype: male (17.4 × 14.7 mm)(NMCR), Balicasag Island, Panglao, Bohol, Philippines,
coll. fishermen with tangle nets, July 2003. Paratypes:
Philippines – 5 males (18.3 × 14.4 mm, 16.0 × 13.5 mm,
15.3 × 11.8 mm, 16.9 × 13.2 mm, 16.5 × 13.1 mm), 3
ovigerous females (15.2 × 11.6 mm, 14.9 × 11.5 mm, 14.4
× 11.3 mm) (ZRC 2013.1263), same data as holotype. — 3
males, 1 female (ZRC 2001.0413) Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, Balicasag
Island, Panglao, Bohol, 200–300 m, coll. fishermen withtangle nets, December 2000. — 2 males, 2 females (ZRC
2001.0601) Balicasag Island, Panglao, Bohol, 200–300 m,
coll. fishermen with tangle nets, 28 November 2001. — 1
female (NSMT-Cr 15389), Balicasag Island, Panglao, Bohol,
coll. local fisherman via M. Takeda & H. Komatsu, February
2003. — 8 males, 5 females (1 ovigerous) (ZRC 2013.1265),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, November 2003. — 1 male, 1 female (ZRC
2013.1260), Balicasag Island, Panglao, Bohol, 200–300 m,
coll. fishermen with tangle nets, December 2003. — 1 male
(ZRC 2013.1257), Balicasag Island, Panglao, Bohol, 200–300
m, coll. fishermen with tangle nets, January 2004. — 6 males

Etymology. The species is named after the renowned Frenchcarcinologist Raoul Serène, who first recorded this species
as Maja japonica from the Philippines.
Remarks. The good series of S. serenei on hand shows thevariation in carapace shape as the specimens increase insize. The carapace is relatively more pyriform (Fig. 30A, B)
in smaller specimens of S. serenei (ca. carapace length 12mm) but as they get larger (ca. carapace length 16 mm), thecarapace becomes somewhat more rounded in appearance(Fig. 30C). Even so, it never gets as rounded or as broad as
similarly sized specimens of S. japonica (Fig. 28A–C). The
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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.

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Sakai (1934: text fig. 11b) (as M. nipponensis) is that of asmall male measuring 14.5 by 13.0 mm, but in larger adultmales, the distal part of the G1 is slightly more elongate
and more setose (Fig. 33G). However, the G1 structure ofadult S. japonica clearly differs from that of S. serenei bybeing distinctly more strongly curved outwards (Fig. 33A–F
versus Fig. 33K–M).

carapaces are also different in the degree of their inflation andthe depth of grooves separating the regions. Sakaija japonica
differs in having the branchial region more evenly swollen,with the grooves separating it from the gastric, cardiac andintestinal regions visible but shallow and narrow, so muchso that the surfaces of all these regions look evenly convex(Figs. 28A–C, 31A, B). In S. serenei on the other hand, thegrooves separating the branchial region from the gastric,
cardiac and intestinal regions are distinct, relatively broader,
and the regions are clearly separated (Figs. 30, 31D, E). Since
the branchial region is more inflated, the dorsal branchialspine in S. japonica is directed obliquely vertically (Figs.
28A–C, 31A, B) while that of S. serenei is directly obliquely
laterally (Figs. 30, 31D, E). Significantly, the ambulatory legs,
notably the meri, are proportionately shorter and stouter in S.
japonica (Figs. 28A–C, 31C) compared to S. serenei where
they are relatively more slender and longer (Figs. 30, 31F).

Serène & Lohavanijaya (1973: 50, figs. 93–97, pl. 9 fig. A)
recorded “Maja japonica” from off Nhatrang, Vietnam, on
the basis of a 16.0 by 11.5 mm male. Their excellent figuresleave no doubt that their specimen is S. serenei. Serène &Vadon (1981) reported “Maja sakaii” from the Philippines,
but in view of the present study, their records are almost
certainly referrable to S. serenei n. sp. instead.
An ovigerous female from Papua New Guinea (MNHNIU-2011-3898) is tentatively referred to S. serenei. Although
it was collected some distance from the Philippines, it agrees
best with S. serenei with regards to the carapace shape andambulatory leg proportions.

The G1 structures of S. japonica and S. serenei aresuperficially similar, both are curved and have the distal part
lined with long setae. The G1 of S. japonica figured by T.

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 × 18.4mm) (MNHN-IU-2010-505), Madagascar.

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Sakaija longispinosa n. sp.
(Figs. 32A, B, 37N, 46L)

(Fig. 46L). Ambulatory legs relatively short (Fig. 32A, B).
G1 not known.

Material examined. Holotype: ovigerous female (11.4 × 8.6mm) (NMV J63792), Northwest Shelf, between Port Hedlandand Dampier, 18°45.00’S 118°24.00’E, 142 m, Australia,
coll. RV Soela, 5 June 2007. Paratype: 1 ovigerous female(14.2 × 10.7 mm) (NMV J63197), same data as holotype.
Others: Australia – 1 ovigerous female (13.4 × 9.7 mm)(NMV J60779), northwestern coast, Adele L26 transect
14°33.41’S 122°54.22’E, 135–165 m, coll. RV Southern
Surveyor, 4 July 2007.

Etymology. The name alludes to the long carapace spinesof the species.
Remarks. This species has proportionately, the longest lateraland branchial spines in any congener, which easily distinguish
it. As such, we are confident in referring it to a new species
even in the absence of males. One ovigerous specimen(NMV J60779) is tentatively referred to S. longispinosa. It
appears to be somewhat more setose compared to the others
and several of its major spines are broken.

Diagnosis. Carapace branchial area gently inflated, ovate;with very long lateral and branchial spines (Fig. 32A, B).
Pseudorostral spines long, diverging (Fig. 37N). Supraoculareave distinctly curved; antorbital spine strong, curved distally
upwards; intercalated spine short, with very wide gaps
between spines; postorbital spine long, pointed anteriorly;
hepatic spine small, pointed outwards (Fig. 32A, B). Lateral
margin with 2 distinct spines, anterior one small; branchialspine very long; median row with 4 spines: 2 gastric (theanterior one reduced), 1 long cardiac, 1 long intestinal;
posterior carapace margin with 1 strong median spine and 2
adjacent small spinules or sharp spines (Fig. 32A, B). Basalantennal article with 2 blunt distal teeth, with 3 rows of 4
granules. Ischium of third maxilliped distinctly granulated

Sakaija santo n. sp.
(Figs. 32C, D, 33S–U, 37O, 40M, 46K, 51E, 54L, 56J,
70E)
Material examined. Holotype: male (9.4 × 6.6 mm)(MNHN), station AT 61, west of Malo Island, Santo,
15°39.2’S 167°01.4’E, 266–281 m, Vanuatu, coll. RVAlis, SANTO 2006, 4 October 2006. Paratypes: Vanuatu– 2 females (13.2 × 10.1 mm, 13.7 × 10.6 mm) (ZRC
2013.1253), station AT 24, 23 September 2006. — 3 females(13.1 × 10.4 mm, 12.8 × 10.0 mm, 12.6 × 9.0 mm) (ZRC
2013.1245), station AT 64, west of Malo Island, Santo,
15°39.6’S 167°01.9’E, 249–252 m, coll. RV Alis, SANTO

Fig. 30. General habitus, Sakaija serenei n. sp. A, paratype male (14.3 × 10.9 mm) (ZRC 2013.1255), Philippines; B, paratype male (17.7
× 14.6 mm) (ZRC 2013.1262), Philippines; C, holotype male (17.4 × 14.7 mm) (NMCR), Philippines; D, paratype female (14.2 × 10.9mm) (ZRC 2013.1255), Philippines.

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2006, 4 October 2006. — 1 female (11.2 × 8.4 mm) (ZRC
2013.1253), Santo, coll. September–October 2006. — 1 male(9.7 × 7.5 mm), 2 females (13.4 × 9.9 mm, 11.5 × 8.7 mm)(ZRC 2013.1254), Santo, coll. September–October 2006.
— 1 female (13.4 × 10.4 mm) (ZRC 2013.1251), station
AT 61, west of Malo Island, Santo, 15°39.6’S 167°01.9’E,
249–252 m, coll. RV Alis, SANTO 2006, 4 October 2006.
— 1 female (12.8 × 9.5 mm) (ZRC 2013.1250), station
AT 64, west of Malo Island, Santo, 15°39.6’S 167°01.9’E,
249–252 m, coll. RV Alis, SANTO 2006, 4 October 2006.
— 1 female (8.5 × 6.3 mm) (ZRC 2013.1247), station ZB 6,
southwest of Urélapa Island, Santo, 15°36.8’S 167°01.3’E,
on patches of sand, 30 m, coll. RV Alis, SANTO 2006, 28
September 2006. — 2 females (11.9 × 9.4 mm, 12.5 × 9.9mm) (ZRC 2013.1246), station AT 86, Scorff Passage, Santo,
15°31.9’S 167°16.2’E, 176–246 m, coll. RV Alis, SANTO
2006, 12 October 2006. — 1 male (damaged, 11.1 × 8.5mm) (ZRC 2013.1248), station AT 117, Scorff Passage,
Santo, 15°32.6’S 167°15.5’E, 123–196 m, coll. RV Alis,
SANTO 2006, 18 October 2006. — 1 female (12.5 × 9.7mm) (ZRC 2013.1249), station AT 117, Scorff Passage,
Santo, 15°32.6’S 167°15.5’E, 123–196 m, coll. RV Alis,
SANTO 2006, 18 October 2006. Others: New Caledonia– 1 male (22.2 × 17.8 mm) (MNHN-IU-2013-1762), station
CP 3884, 521–567 m, 22°22’S 171°38’E, coll. EXBODI, 18
September 2011. — 1 female (20.1 × 15.8 mm) (MNHNIU-2014-4010), station DW 2491, 24°44’S 159°40’E, coll.
EBISCO, 18 6 October 2010.

Fig. 31. Frontal and lateral views of carapaces, and right first ambulatory legs, Sakaija species. A–C, S. japonica male (22.6 × 17.9 mm)(ZRC 2013.1269), Taiwan; D–F, S. serenei n. sp., paratype male (17.7 × 14.6 mm) (ZRC 2013.1262), Philippines.

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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.

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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., paratypemale (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].

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Diagnosis. Carapace branchial area gently inflated, ovate;with short lateral and branchial spines (Fig. 32C, D).
Pseudorostral spines long, diverging (Fig. 37O). Supraorbital
eave distinctly curved; antorbital spine strong, triangular;intercalated spine short, with large gaps between spines;
postorbital spine strong, triangular, directed anteriorly;
hepatic spine small, pointed outwards (Fig. 37O). Lateral
margin with 2 spines, anterior one small, with granulebetween them; 1 long branchial spine; median row with 3
spines: 1 gastric (with granule anterior to it), 1 low cardiac;
1 intestinal (sometimes very low); posterior carapace margin
with 3 short spines, median one longest (Fig. 32C, D).
Basal antennal article with 2 low distal spines, with 2 rowsof 3 or 4 granules (Fig. 40M). Ischium of third maxilliped
distinctly granulated (Fig. 46K). Ambulatory legs long,
slender (Figs. 32C, D, 56J). G1 sinuous, distal part turned
backwards towards median line of sternum, with very longsetae (Fig. 33S–U).

region with 1 short spine and 1 submedian tubercle (Fig.
34A, B). Posterior carapace margin with 2 very short medianspines (Fig. 34A, B). Eyes relatively short, slender, withlarge ovoid cornea (Fig. 38A). Antennal flagellum short,
slender. Basal antennal article longer than broad, rectangular;
surface with numerous granules, with 2 spines distally; innerand outer lateral margins granulated; proximal outer angle
rounded; antero-external crested rim of antennular fossa
touches overlap distal part of basal antennal article by abouta third of its width (Fig. 40N). Epistome wider than long,anterior margin with 2 low granular lobes; posterior margincomposed of 4 rectangular plates separated by shallow fissures(Figs. 40N, 42G). Suborbital margin separated from basal
antennal article and margin of postorbital tooth by deep
fissures fissures (Fig. 40N). Outer surface of third maxilliped
covered by short setae in adults; ischium subrectangular,
just longer than broad; postero-external angle of merus
relatively broad, “inserted” into shallower concavity on outermargin of ischium; antero-internal part of ischium rounded,auriculiform (Fig. 46M, N). Male chelipeds relatively shortin adult males, surface of merus and carpus finely granulated;
carpus long; propodus of palm elongated, curved, smooth,not inflated, palm as long as fingers; fingers long, slender,
gently curved, with distinct basal gape when closed (Figs.
34A, B, 54N). Ambulatory legs relatively short, slender;
merus without dorsal subdistal spine; dactylus elongate,
curved, covered with long setae except for corneous tip(Figs. 34A, B, 56K). Thoracic sternum wide; surfaces ofsomites 5–8 with numerous prominent rounded tubercles
and granules; sternites 3 and 4 depressed; margin between
sternites 2 and 3 demarcated by deep notch; anterior marginof sterno-abdominal cavity forming prominent complete rim(Figs. 51F, G, 52K). Male abdomen subtriangular, with 6 freesomites and telson; somites 3 and 4 wider than somite 6 andtelson (Fig. 51F, G). Male press-button abdominal locking
mechanism submedian in position on sterno-abdominal
cavity (Fig. 52K). Female abdomen dome-shaped, covering
most of thoracic sternum. G1 long, slender, distal part bent
approximately 90°, distal part chitinised with rows of short
stiff setae, tapering to sharp tip (Fig. 35A–E).

Etymology. The species is name is derived from the nameof the island from which the species was collected, Espiritu
Santo in Vanuatu. The name is used as a noun in apposition.
Remarks. In the general carapace morphology andproportions of the ambulatory legs, S. santo n. sp. is closest
to S. serenei n. sp. They differ markedly in the form of theG1, which is gently curved and dorsoventrally flattened in
S. serenei with the distal part elongate (Fig. 33K–M), but
is less curved, more laterally flattened with the distal part
ovoid in S. santo (Fig. 33S–U). In addition, the posterior
carapace margin of S. santo is always conspiciously marked
by a small but distinct median spine (with 2 smaller lateralones) (Fig. 32C, D), absent or undiscernible in S. sakaii(Fig. 28D, E, G). This median spine may be low in somespecimens of S. santo but is always visible. In S. sakaii, theposterior carapace margin has two median spines that may
be very low (Fig. 28D, E, G). In S. santo, there are threespines with the median one longest, and while the lateralones may become reduced, the median one is always distinct(Fig. 32C, D). The G1 structure of S. santo (Fig. 33S–U) is
most similar to that of S. sakaii s. str. but the setae on the
ovoid distal part are relatively shorter in the latter species(Fig. 33H–J).

Type species. Planaja plana n. sp., by present designation.

Planaja n. gen.

Etymology. The name is an arbitrary combination of theLatin “planus” for flat and Maja. Gender feminine.

Diagnosis. Carapace subovoid; dorsal surface gently inflated,covered by sharp granules or low tubercles; gastric andbranchial regions clearly delimited by grooves (Fig. 34A,
B). Intestinal region with small median spine (Fig. 34A, B).
Pseudorostral spines relatively long, slender, diverging (Figs.
34A, B). Supraorbital eave large, anterior part expanded,
foliaceous, margin convex, antorbital spine long, sharp (Fig.
38A). Intercalated spine distinct, separated from supraorbital
eave and postorbital spine by distinct gaps; postorbital spine
strong, long, gently recurved; hepatic region with 1 low but
distinct spine, much shorter than postorbital spine; 1 or more
tubercles below (Fig. 38A). Lateral carapace margin with 6
or 7 short sharp spines (posterior ones smaller), branchial

Remarks. The main differences between Planaja gen.
nov. and the other genera treated here have been discussed
under Holthuija and Sakaija. The most diagnostic featuresof the genus are the relatively flatter carapace in which thedorsal surface is only gently convex, the supraorbital eave
is expanded, possessing more spines (six) on the lateralcarapace margin, as well as the short and thick ambulatory
legs (see key to genera). Another unique character of thetype species is that the distal quarter of the G1 is distinctly
chitinised and lined with numerous short setae (Fig. 35A–E),
a feature not present in other genera in this study. In thisrespect, the G1 of Planaja resembles species of Leptomithrax
like L. longimanus and L. edwardsii (unpublished data).

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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 × 40.5mm) (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.

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Planaja plana n. sp.
(Figs. 2G, 34A, B, 35A–E, 38A, 40N, 42G, 46M, N, 51F,
G, 52K, 54N, 56K)

Panglao, Bohol, coll. fishermen with tangle nets, December
2000. — 8 males (38.2 × 31.7 mm, 44.4 × 36.8 mm, 38.5
× 31.7 mm, 38.7 × 31.0 mm, 38.2 × 31.0 mm, 30.4 × 24.3mm, 26.3 × 21.2 mm, 34.4 × 27.3 mm), 4 ovigerous females(44.7 × 37.2 mm, 42.4 × 34.5 mm, 36.1 × 29.7 mm, 44.5
× 38.6 mm) (ZRC 2001.0430), Balicasag Island, Panglao,
Bohol, coll. fishermen with tangle nets, December 2000.
— 1 male (39.3 × 30.3 mm) (ZRC 2001.0591), Balicasag
Island, Panglao, Bohol, coll. fishermen with tangle nets,
28 November 2001. — 2 males (40.2 × 31.8 mm, 35.8 ×
27.9 mm), 1 female (44.4 × 34.8 mm) (USNM), Balicasag

Material examined. Holotype: male (43.7 × 37.1 mm)(NMCR), Balicasag Island, Panglao, Bohol, Philippines,
coll. fishermen with tangle nets, July 2003. Paratypes:
Philippines – 3 males (42.9 × 35.0 mm, 38.3 × 30.1 mm,
33.3 × 26.9 mm), 2 ovigerous females (47.9 × 40.9 mm, 45.1
× 37.8 mm) (ZRC 2013.1319), same data as holotype. — 1
male (40.2 × 33.2 mm) (ZRC 2001.0435), Balicasag Island,

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].

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Remarks. Planaja plana is superficially similar to Sakaija
japonica in general form, especially with regard the relatively
short ambulatory legs. Planaja plana, however, has six orseven lateral carapace spines behind the hepatic one andits dorsal surface is conspicuously less convex. Smaller
specimens of P. plana about 20 mm in carapace width (e.g.,
22.2 × 17.5 mm, 22.7 × 18.3 mm; ZRC 2013.1325) have aslightly more inflated carapace, and the G1 is less strongly
bent outwards with the distal part relatively shorter (Fig. 35D).

Island, Panglao, Bohol, coll. fishermen with tangle nets, June
2002. — 1 juvenile male (ZRC 2013.1316), Balicasag Island,
Panglao, Bohol, 200–300 m, coll. fishermen with tangle nets,
June 2002. — 2 males (42.5 × 34.9 mm, 38.1 × 29.8 mm),
1 ovigerous female (48.8 × 41.4 mm), 1 female (45.3 × 37.8mm) (ZRC 2013.1369), Balicasag Island, Panglao, Bohol,
coll. fishermen with tangle nets, July 2003. — 2 males, 1
female (NSMT-Cr 22331), Balicasag Island, Panglao, Bohol,
200–300 m, coll. fishermen with tangle nets, February
2004. — 2 males (22.2 × 17.5 mm, 22.7 × 18.3 mm) (ZRC
2013.1325), Balicasag Island, Panglao, Bohol, 200–300 m,
coll. fishermen with tangle nets, May 2004. — 1 male (42.1
× 35.1 mm), 2 ovigerous females (44.8 × 37.2 mm, 44.9
× 37.3 mm) (ZRC 2013.1368), Balicasag Island, Panglao,
Bohol, station PN1, coll. P.K.L. Ng from fishermen withtangle nets, 29 May 2004. — 1 male (42.8 × 34.3 mm),
1 female (AM), station PN1, Balicasag Island, Panglao,
Bohol, coll. fishermen with tangle nets, January 2004. —1 female (ZRC 2013.1320), Stn PN 1, Balicasag Island,
Panglao, Bohol, coll. fishermen with tangle nets, 3 January
2004. — 1 female (37.1 × 30.2 mm) (ZRC 2013.1321),
Balicasag Island, Panglao, Bohol, coll. fishermen with tangle
nets, February 2004. — 2 males (35.3 × 28.8 mm, 33.6 ×
26.7 mm), 2 females (41.0 × 32.0 mm, 28.9 × 23.0 mm)(ZRC 2013.1326), Balicasag Island, Panglao, Bohol, coll.
fishermen with tangle nets, March 2004. — 1 male (38.3
× 30.7 mm) (ZRC 2013.1370), Balicasag Island, Panglao,
Bohol, coll. fishermen with tangle nets, 28 May 2004. — 2
males (41.0 × 34.0 mm, 45.4 × 36.5 mm) (ZRC 2012.1206),
Balicasag, Panglao, Bohol, 200–300 m, coll. local fishermenwith tangle nets, 29 May 2004. — 1 ovigerous female (45.2
× 36.9 mm) (ZRC 2012.1205), station DW 2402, Dipolog
Bay, off Balicasag Island, Bohol Sea, 9°30.8’N 123°41.5’E,
coll. PANGLAO 2005, 31 May 2005. Others: Taiwan – 1
female (46.6 × 40.9 mm) (ZRC 2001.0057), Nangfangao
port, Su-Ao, Ilan County, northeastern Taiwan, coll. P.K.L.
Ng, 5 November 2000.

Surprisingly, this new genus and new species was obtained inlarge numbers in Balicasag, all collected by tangle nets. None
were collected from trawls or dredges from the Philippines.
The one specimen from Taiwan, a female (46.6 × 40.9 mm,
ZRC 2001.0057), was obtained by a commercial trawl.
Ovimaja n. gen.
Diagnosis. Carapace pyriform, much longer than broad;dorsal surface evenly inflated, covered by low granules
or tubercles; gastric and branchial regions delimited bygrooves (Fig. 34C, F, G). Intestinal region without medianspine (Fig. 34C, F, G). Pseudorostral spines relatively short,diverging (Figs. 34C, D, F, G, 38B, C). Supraorbital eavewith anterior part longitudinally narrow, rectangular, notprominently expanded; antorbital spine sharp (Fig. 38B,
C). Intercalated spine distinct, separated from supraorbital
eave and postorbital spine by distinct gaps; postorbital
spine strong, lobiform; hepatic region with 1 distinct spine,much shorter than postorbital spine; 2 or small spines
below (Fig. 38B, C). Lateral carapace margin with about
3 short spines and some sharp tubercles, branchial region
with 1 low spine, obscure in large specimens (Fig. 34C,
F, G). Posterior carapace margin with 2 very short medianspines (Fig. 34C, F, G). Eyes long, slender, with long ovoid
cornea bent at 30° from horrizontal (Fig. 38B, C). Antennal
flagellum short, slender. Basal antennal article longer thanbroad, rectangular; surface with several rounded granules,
with 3 low, blunt spines distally, outer lateral margin entire
or with a few small rounded granules; inner lateral margin
with 3 large rounded projections with overlaps antennularfossa; proximal outer angle rounded; antero-external crested
rim of antennular fossa overlaps halfway into distal part ofbasal antennal article, forming a hook-like structure (Fig.
40O). Epistome longer than wide, anterior margin with 2large rounded tubercles; posterior margin composed of 4
rectangular plates separated by deep fissures (Figs. 40O,
42H). Suborbital margin separated from basal antennal
article and margin of postorbital tooth by deep fissures (Fig.
40O). Outer surface of third maxilliped covered by setae inadults; ischium subrectangular, slightly longer than broad;postero-external angle of merus relatively broad, “inserted”
into shallower concavity on outer margin of ischium; anterointernal part of ischium rounded, auriculiform (Fig. 46O, P).
Male chelipeds relatively short in adult males, surfaces ofmerus and carpus smooth; carpus long; propodus of palm
short, smooth, fingers longer than palm; fingers long, slender,
gently curved, with basal gape when closed (Figs. 34C, F,
G, 54O, P). Ambulatory legs relatively short, slender; merus
without dorsal subdistal spine; carpus very short, cordiform

Diagnosis. Carapace pyriform; dorsal surface gently convex,
covered by numerous rounded granules (Fig. 34A, B).
Pseudorostral spines long, diverging (Fig. 38A). Supraorbital
eave expanded, with long sharp antorbital tooth; intercalated
spine triangular, sharp, shorter than others; postorbital tooth
triangular, very long, touching base of long, curved hepatic
spine; median row with 5, usually 6 short spines (Fig. 34A,
B, 38A). Lateral margin with 6 or 7 short spines, regularly
arranged; 1 short branchial spine and 1 submedian tubercle;
3 small spines on posterior carapace margin (Fig. 34A, B).
Ambulatory legs short, setose (Figs. 34A, B, 56K). Basalantennal article distinctly granulated with 2 strong spines
distally (Fig. 40N). Subhepatic area strongly granulated (Fig.
40N). Ischium of third maxilliped almost smooth, gently
rugose or with scattered low granules (Fig. 46M, N). G1
long, slender, distal part bent approximately 90°, distal part
chitinised with rows of short stiff setae, tapering to sharptip (Fig. 35A–E).
Etymology. From the Latin “planum” for flattened, alluding
to the relatively flatter carapace of the species.

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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 × 56.1mm) (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 latensn. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon Islands; O, Alcomaja miriky n. sp., holotype male (26.4 × 20.9mm) (MNHN-IU-2010-929), Madagascar.

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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 × 14.7mm) (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.

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with deep median groove; dactylus elongate, slightly curved,
covered with long setae except for corneous tip (Figs. 34C,
F, G, 56L). Thoracic sternum wide; surfaces of somites 5–8
almost smooth; sternites 3 and 4 slightly depressed; margin
between sternites 2 and 3 demarcated by very deep notch,
forming waist-like structure; anterior margin of sternoabdominal cavity not forming complete rim (Figs. 51I,
52L). Male abdomen subrectangular, with 6 free somites
and telson; somites 6 wider than all other somites and telson(Fig. 51I). Male press-button abdominal locking mechanism
laterally positioned on sterno-abdominal cavity (Fig. 52L).
Female abdomen dome-shaped, covering most of thoracic
sternum. G1 very long, slender, distal part elongated, curved
outwards, with 2 distal folds and sharp tip, distal part with
scattered very short setae (Fig. 35F–L).

holotype 31.0 × 24.0 mm) from the mid to late Early
Miocene (ca. 16–23 million years ago). The carapace of M.
robinsoni (cf. Jenkins, 1985: pl. 2 figs. 2a, 3) is similar to
O. compressipes, but the granules on the posterior half ofthe carapace appear to be more closely packed (less packed
in O. compressipes, Fig. 34C, F, G) and the lobulation on
the epistome and margins of the antenna is less distinct(Jenkins, 1985: pl. 2 fig. 2d, e) (cf. Fig. 40O).
Ovimaja compressipes (Miers, 1879) comb. nov.
(Figs. 2H, 34C–G, 35F–L, 38B, C, 40O, 42H, 46O, P,
51H, I, 52L, 54O, P, 56L, 70F)
Paramithrax (Leptomithrax) compressipes Miers, 1879: 8.
Maja compressipes – Griffin & Tranter, 1986: 211, 212, pl. 15a,b. – Chen & Ng, 1995: 758. – Rahayu & Ng, 2000: 888. –Maramura & Kosaka, 2003: 34. – Yang et al., 2008: 780. – Ng
et al., 2008: 117 (list).
Leptomithrax compressipes – Serène, 1968: 57. – Huang, 1994: 583.
Maja linapacanensis Rathbun, 1916: 553. – Serène, 1968: 57. –Griffin, 1976: 200. – Griffin & Tranter, 1986: 213 (discussion).
Maja gibba – Serène & Lohavanijaya, 1973: 163, pl. 9B. (notMaja gibba Alcock, 1895).
Maja brevispinosa Dai, 1981: 37, 38, pl. I: 8–10.
Maja brevispinosis [sic] – Dai et al., 1986: 136, pl. 18(2), figs.
77(1–4). – Dai & Yang, 1991: 151, pl. 18(2), figs. 77(1–4).
– Huang, 1994: 582.

Type species. Paramithrax (Leptomithrax) compressipes
Miers, 1879, by present designation.
Etymology. The genus name is derived from an arbitrary
combination of the Latin “ovum” for egg and Maja, alluding
to the egg-like appearance of the carapace of the type species.
Gender feminine.
Remarks. Ovimaja n. gen. is very different from Maja s.
str. and the various genera recognised here. Aside fromthe distinctively elongated carapace (Fig. 34C, F, G), thestructure of the basal antennal article is unique, with theinner margin lined with three prominent projections (Figs.
40O, 42H); the anterior margin of the epistome has twolarge projections (Figs. 40O, 42H); the male anterior thoracic
sternum is waist-like, with the region between sternites 2
and 3 deeply constricted (Figs. 51I, 52L); the carpus of theambulatory leg is very short, enlarged and flattened (Fig.
56L); and the G1 is very long and curved with the distal
part very slender (Fig. 35I–L). Ovimaja has a very different
male abdominal locking mechanism from the other genera.
In these genera, including Maja s. str., the press-button issubmedian in position (Fig. 52A–K). In Ovimaja, the pressbutton is positioned laterally on thoracic sternite 5, near themargin of the sterno-abdominal cavity (Fig. 52L).

Material examined. Holotype male (51.2 × 40.5 mm) (NHM
1860.15), Canton (= Guangzhou), China, coll. Horsfield,
East India Company. Others: Philippines – 1 dried carapace(27.0 × 20.0 mm) (with spines, length 31.8, width (rightside broken, 19.6 mm) (USNM 48225) (holotype of Majalinapacanensis Rathbun, 1916) [photographs examined],
Observatory Island, Linapacan Strait, north of Palawan, 46
fathoms, 11°37.25’N 119°48.75’E, coll. RV Albatross, 18
December 1908. Taiwan – 1 male (49.6 × 39.6 mm) (ZRC
2008.1318), Penghu Islands, coll. S.-H. Wu, 5 September
1974. — 1 male (55.7 × 42.7 mm) (ZRC 2013.1264), coll.
1990s. South China Sea – 1 young female (45.6 × 35.4 mm)(ZRC 1999.1260), southern part of South China Sea, coll.
commercial trawlers, 26 November 1982. Thailand – 1 male
(41.1 × 30.3 mm) (ZRC 2003.0162), Pattani Province, Saiburi
Crab Landing, Pattani Province, southeastern Thailand, coll.
Z. Jaafar et al., 7 June 2003. — 2 males (45.5 × 36.8 mm,
40.3 × 30.4 mm) (ZRC 2003.0196), Pattani Province, Saiburi

One fossil species described from South Australia, Majarobinsoni Jenkins, 1985, is almost certainly a species ofOvimaja. Described on the basis of nine carapaces (theFig. 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.

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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) (MNHNIU-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 × 26.0mm) (ZRC 2001.430), Philippines.

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Fig. 40. Antennae, antennules and epistomes. A, Paramaya spinigera, male (85.0 × 66.4 mm) (ZRC 1999.738), Taiwan; B, Paramaya ouchn. 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.

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Crab Landing, Pattani Province, southeastern Thailand, coll.
Z. Jaafar et al., 8 June 2003.

gastric, 1 cardiac; 1 spine on branchial region; 2 very small
spines on posterior carapace margin (Fig. 34C, F, G). Basalantennal article very broad, with 4 large blunt spines, 3 distal
and 1 on inner margin, 1 large proximal granule on inner
margin (Figs. 40O, 42H). Pterygostomian area prominently
granulated (Figs. 34D, 40O, 42H). Ambulatory legs short,with stout, short segments, covered with long hooked setae;
merus short (Fig. 34C, G, 40O, 56L). G1 long, slender,strongly curved, with 2 distal folds and sharp tip (Fig. 35F–L).

Diagnosis. Carapace dorsal surface covered with large
rounded granules, with groups of long hooked setae oneach granule; branchial area rounded, deliminated by deep
narrow grooves (Fig. 34C, F, G). Pseudorostral spines short,diverging outwards (Fig. 38B, C). Supraorbital eave large,
antorbital spine sharp, covered by hooked setae, with 2 rowsof large granules parallel to spines; intercalated spine short,triangular; postorbital spine largest, directed anteriorly; small
hepatic spine oriented outwards (Fig. 38B, C). Lateral margin
with 5–7 sharp granules; median row with 2 small spines: 1

Remarks. Maja compressipes was originally described from
Canton (present day Guangzhou) in China. It has since beenreported from other parts of China and Japan (e.g., Yang etFig. 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 × 45.7mm) (NMCR), Philippines; H, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.

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Ng & Richer de Forges: Revision of the spider crab genus Majaal., 2008; Maramura & Kosaka, 2003), Philippines (Rathbun,
1916, as Maja linapacanensis) and the Straits of Malacca(Rahayu & Ng, 2000), and we have material from the Gulfof Thailand, South China Sea and Taiwan.

short and cordiform. The photograph of a specimen attributed
to “Maja gibba” from Indonesia in Serène & Lohavanijaya(1973: 163, pl. IXB) is almost certainly Ovimaja compressipes
considering its setose body and very wide ambulatory carpus.

The species is very characteristic in having an elongated andnot inflated carapace; the carapace and ambulatory legs arecovered by a thick tomentum of long setae which gathers
a substantial amount of mud and debris; the basal antennal
article is very broad with large tubercles on its distal part;the ambulatory legs are very short and the carpus is very

Griffin & Tranter (1986: 213) suggested that Majalinapacanensis Rathbun, 1916, described only from a broken
carapace, is conspecific with M. compressipes, but they did
not formally synonymise them. Examination of the types ofboth species leaves no doubt the two taxa are synonymous.

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.

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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 × 56.1mm) (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 × 66.8mm) (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.

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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).

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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 × 26.0mm) (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 ouchn. 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.

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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., paratypeovigerous 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.

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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.

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Fig. 48. Male anterior thoracic sternums and abdomens, Paramaja. A, P. kominatoensis, dried male (62.6 × 56.1 mm) (KPM NH0104298),
Japan; B, P. kominatoensis, male (76.9 × 69.6 mm) (SMF 47738), Japan; C, P. kominatoensis, male (50.5 × 44.6 mm) (SMF 47738),
Japan; D, P. kominatoensis, dried male (73.0 × 66.0 mm) (PCM), Taiwan; E, P. kominatoensis, dried male (60.4 × 52.5 mm) (KPM
NH124171), Japan; F, P. gibba, male (79.5 × 77.9 mm) (ZRC 2013.1232), Bay of Bengal; G, P. gibba, male male (64.7 × 60.7 mm)(SIO), Indian Ocean; H, P. turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; I, P. turgida n. sp., paratype male (67.9
× 60.5 mm) (MNHN), Philippines.

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Fig. 49. Male anterior thoracic sternums and abdomens, Alcomaja species. A, A. irrorata n. sp., holotype male (52.6 × 45.7 mm) (NMCR),
Philippines; B, A.gracilipes, holotype male (45.4 × 43.5 mm) (IOCAS K33B-34), South China Sea; C, A. gracilipes, male (41.4 × 35.4mm) (ZRC 2013.1225), Philippines; D, A. nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines; E, A. nagashimaensis,
male (37.2 × 32.4 mm) (CBM-ZC9928), Japan; F, A. latens n. sp., holotype male (25.3 × 21.4 mm) (MNHN-IU-2013-4050), Solomon
Islands; G, A. miriky n. sp., holotype male (26.4 × 20.9 mm) (MNHN-IU-2010-929), Madagascar; H, A. desmondi n. sp., holotype male
(35.4 × 28.3 mm) (NMCR), Philippines.

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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 × 55.6mm) (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.

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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 × 40.5mm) (NHM 1860.15), China; I, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

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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.

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Fig. 53. Chelae. A, Maja squinado, neotype male (147.1 × 126.3 mm) (SMF-4548), Croatia; B, Maja brachydactyla, male (98.4 × 89.0mm) (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 × 103.4mm) (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 × 66.0mm) (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 × 28.3mm) (NMCR), Philippines; P, Alcomaja nagashimaensis, male (30.5 × 26.0 mm) (ZRC 2001.430), Philippines.

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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.

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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 × 56.1mm) (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.

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Fig. 56. First ambulatory dactyli and propodi. A, Paramaya spinigera, dried female (70.1 × 58.8 mm) (KPM NH4195), Japan; B, Paramaya
coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; C, Holthuija miersi, male (32.6 × 25.6 mm) (ZRC 2000.1497),
Singapore; D, Holthuija suluensis, holotype female (32.4 × 41.2 mm) (USNM 48224a), Philippines; E, Holthuija cognata n. sp., holotype
male (29.5 × 23.7 mm) (CBM-ZC3662), Japan [figure laterally inverted]; F, Holthuija aussie n. sp., holotype ovigerous female (42.1 ×
34.4 mm) (NMV J63752), Arafura Sea; G, Holthuija poorei, n. sp. paratype female (27.0 × 22.5 mm) (NMV J63751), Arafura Sea; H,
Sakaija japonica, male (22.3 × 17.8 mm) (ZRC 2013.1267), Taiwan; I, Sakaija serenei n. sp., holotype male (17.4 × 14.7 mm) (NMCR),
Philippines [figure laterally inverted]; J, Sakaija santo n. sp., holotype male (9.4 × 6.6 mm) (MNHN), Vanuatu; K, Planaja plana n. gen. n.
sp., holotype male (43.7 × 37.1 mm) (NMCR), Philippines; L, Ovimaja compressipes, male (49.6 × 39.6 mm) (ZRC 2008.1318), Taiwan.

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Rathbunaja n. gen.

abdomen dome-shaped, covering most of thoracic sternum.
G1 very long, slender, curved distally, with subdistal dorsal
flap (sometimes low), tip tapering, sharp; distal and subdistal
surface with numerous short setae (Figs. 66, 67A–G).

Diagnosis. Carapace distinctly pyriform; dorsal surface
covered by sharp granules, tubercles and short spines;
gastric and branchial regions distinct, separated by distinct
grooves; carapace and pereiopods covered with dense shortsetae, forming velvet-like pubescence (Figs. 59, 60). Cardiac
region with 2 short spines arranged transversely; intestinal
region with 2 or 3 short spines (Figs. 59, 60). Pseudorostrallong or short, divergent (Figs. 59, 60). Supraorbital eavewith anterior part longitudinally narrow, rectangular, notprominently expanded; antorbital spine short, directed
laterally (Figs. 59, 60, 62A). Intercalated spine distinct,
separated from supraorbital eave by wide keyhole shaped
gap, separated from postorbital spine by narrow slit-like
gap; postorbital spine strong; hepatic region with 1 strong
spine, subequal or shorter than postorbital spine, surroundedby short spines and tubercles (Figs. 59, 60, 62A). Lateralcarapace margin with 3 large spines and numerous spinules
and tubercles around them; branchial region with 2 spines,
outer one relatively larger (Figs. 59, 60). Posterior carapace
margin always with 2 distinct median spines (Figs. 59, 60).
Eyes relatively short, stout, with rounded cornea (Figs. 61A,
C, E, G, 62B). Antennal flagellum long, slender (Figs. 61A,
C, E, G, 62B). Basal antennal article longer than broad, with
2 long distal spines; proximal outer angle with small low
tooth; inner and outer lateral margins entire; antero-external
crested rim of antennular fossa slightly overlaps subdistal
part of basal antennal article (61A, C, E, G, 62B). Epistome
much wider than long, anterior margin with 2 low, flattened
tubercles; posterior margin composed of 4 rectangularplates, lateral plates separated from median plates by deep
fissures, median plate separated by shallow cleft (61A, C,
E, G, 62B). Suborbital margin separated from basal antennal
article by wide gap; suborbital tooth completely fused withpostorbital tooth (61A, C, E, G, 62B). Outer surface of third
maxilliped covered by short setae; ischium rectangular,smooth; postero-external angle of merus relatively broad,“inserted” into concavity on outer margin of ischium;antero-internal part of ischium rounded, relatively low, subor not auriculiform (Figs. 62C, 63). Male chelipeds very
long in adult males, surfaces of merus and carpus covered
with distinct tubercles and granules; carpus elongate, withlow granulated longitudinal ridge; palm elongated, curved,
smooth, without lateral cristae, slightly enlarged, withswellings on dorsal and ventral margins in very large males;
fingers long, slender, gently curved, with narrow basal gapewhen closed (Figs. 59, 60, 65C, G, H). Ambulatory leg
merus with distinct distal spine on dorsal margin; dactylus
relatively long, curved, covered with dense short and longsetae except for corneous distal quarter (Figs. 62D–G, 65B,
D, F, I–K). Thoracic sternum relatively wide; surfaces ofsomites 5–8 almost smooth; sternites 3 and 4 mediallydepressed; margin between sternites 2 and 3 demarcatedby small notch; anterior margin of sterno-abdominal cavity
forming incomplete rim with median part low (Figs. 64).
Male abdomen subrectangular, with 6 free somites and telson;
telson subtriangular to semicircular, with convex margins(Fig. 64). Male press-button abdominal locking mechanism
submedian in position on sterno-abdominal cavity. Female

Comparative material. Leptomithrax longimanus (Miers,
1876) – holotype male (38.0 × 36.2 mm) (NHM 1852.12),
New Zealand, coll. R. Gunn. — 1 male (54.3 × 42.4mm) (NIWA 6170), station KAH9704/127, 44°01.98’S
172°12.52’E, 31–34 m, New Zealand, coll. 5 January 1998.
— 1 male (50.7 × 48.9 mm) (ZRC 2013.1617), Godley
Head, east coast of South Island, New Zealand, in crab
trap, coll. C. McLay, May 2003. Leptomithrax australis(Jacquinot, in Jacquinot & Lucas, 1853) – 1 male (77.9
× 68.9 mm), 1 female (47.0 × 39.2 mm) (ZRC 2013.1618),
Banks Peninsula, east coast of South Island, New Zealand,
600 m, coll. C. L. McLay, May 2004. Leptomithrax longipes(G. M. Thomson, 1902) – 1 male (55.4 × 39.3 mm) (AMP86814), off Dunedin, New Zealand. — 1 male (39.8 × 39.3mm) (ZRC 2013.1719, ex NIWA 42151), 1 female (45.7 ×
36.7 mm) (ZRC 2013.1720 ex NIWA 44739), station KAH
0705/48, 44.4883°S 171.796°E – 44.5018°S 171.796°E,
63–65 m, New Zealand, coll. 18 May 2007. Leptomithrax
waitei (Whitelegge, 1900) – 1 male (29.9 × 23.7 mm), 1
female (23.5 × 17. 3 mm) (ZRC 2013.076, ex AM P64812),
station K96-04-14, 32°52’S 152°02’E, Australia, coll. FRV
Kapala, 13 March 1996. Leptomithrax edwardsii (De Haan,
1835) – 1 female (55.9 × 49.7 mm) (ZRC 2013.1186),
Amakusa, Shikizuki, Kyushu, Japan, from fishermen, coll.
J. Lai, September 2002. — 8 males (largest 69.8 × 62.8mm), 7 females (ZRC 2013.1401), Amakusa, Tomioka Port,
Kyushu, Japan, coll. J. Lai & S. Arakaki, 7–9 September
2002. — 1 male, 1 female (SMF 47751), None, Kochi
Prefecture, ca. 33°26.617’N 134°16.75’E, Japan, coll. K.
Matsuzawa, no other data. — 1 male (SMF 47754), None,
Kochi Prefecture, ca. 33°26.617’N 134°16.75’E, Japan,
coll. K. Matsuzawa, 11 December 1990. — 1 ovigerous
female (SMF 47753), Toyocho, None, Muroto Cape, Kochi
Prefecture, 33°26.617’N 134°16.75’E, Japan, coll. 12 June
1998. — 2 females (SMF 47756), None, Kochi Prefecture,ca. 33°26.617’N 134°16.75’E, Japan, coll. K. Matsuzawa,
24 March 1991. — 1 female (SMF 47755), None, Kochi
Prefecture, ca. 33°26.617’N 134°16.75’E, Japan, coll. K.
Matsuzawa, 18 May 1986. — 1 male (SMF 47740), Shikoku,
Tosa Bay, Kochi Prefecture, 33°29.049’N 133°35.707’E,
Japan, coll. 8 April 1988. — 1 male (SMF 47752), None, Cape
Muroto, Kochi Prefecture, 33°26.617’N 134°16.75’E, Japan,
coll. K. Matsuzawa, 18 November 1984. — 2 males, 1 female(SMF 47739), Tokushima, Shikoku, Ana, Tsubakidomari,
Kochi Prefecture, 33°51.678’N 134°43.603’E, Japan, coll.
K. Sakai, 19 April 1984. — 1 female (SMF 47809), None,west of cape Muroto, Kochi Prefecture, 33°26.617’N
134°16.75’E, Japan, coll. K. Matsuzawa, 12 June 1998.
— 1 female (SMF 47727), Tosa Bay, near Mimase, Kochi
Prefecture, 33°29.049’N 133°35.707’E, 150–200 m, Japan,
coll. K. Sakai. — 2 males, 1 crushed juvenile (SMF 47735),
Honshu, Wakayama Prefecture, Kii Minabe, 33°45.712’N
135°18.927’E, Japan, coll. K. Sakai, 1 March 1988. — 3
males (SMF 47731), Honshu, Wakayama Prefecture, Kii
Minabe, 33°45.712’N 135°18.927’E, Japan, coll. K. Sakai,203

Ng & Richer de Forges: Revision of the spider crab genus Maja

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.

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26 October 1988. — 2 males (SMF 46528), Tsubaki Donarii,
Anan City, Tokushina Prefecture, 33°50.62’N 134°42.75’E,
Japan, coll. K. Sakai. — 1 female (SMF 47749), Japan,
coll. K. Matsuzawa, 2005. — 1 dried male (SMF 47604),
Japan, no other data, T. Sakai Collection. — 1 male (75.8
× 69.4 mm) (ZRC 2001.56), Tashi, Taiwan. Leptomithrax
sinensis Rathbun, 1916 – holotype carapace (dried) (32.0
× 25.3 mm) (including spines, 38.3 × 28.8 mm) (USNM
48219), station 5311, South China Sea, near southern
Luzon, 21°33’N 116°15’E, 88 fathoms, Philippines, coll.
RV Albatross, 4 November 1908. Leptomithrax bifidus(Ortmann, 1893) – 1 dried male (36.5 × 29.4 mm) (KPM
NH4024), Kii-Nagashima, Japan, T. Sakai Collection, coll.
March 1969. — 2 males (32.5 × 27.5 mm, 27.5 × 20.6mm) (NHM 1961.11.13.28–29), Seto, Shirahama, Japan,
coll. gill nets, I. Gordon & Harada, 1950s. — 2 males(ZRC), Mitsu, off Muroto Cape, Kochi Prefecture, Japan,
33°16.6’N 134°10.6’E, in coral nets, coll. 8 July 1988. — 1
male (with rhizocephala), 2 females (SMF 47758), Mitsu,
off Muroto Cape, Kochi Prefecture, 33°16.6’N 134°10.6’E,
in coral nets, Japan, coll. 21 June 1988. — 1 male, 1 female(SMF 47758), Mitsu, off Muroto Cape, Kochi Prefecture,
33°16.6’N 134°10.6’E, in coral nets, Japan, coll. 21 June
1988. — 1 male (SMF 47743), Tosa, Usa, Kochi Prefecture,
33°26.316’N 133°28.003’E, 160 m, Japan, coll. K. Sakai, 10
May 1990. — 5 males (SMF 47729), Honshu, Wakayama

Prefecture, Kii Mimase, 33°45.712’N 135°18.927’E, Japan,
coll. K. Sakai, 1 March 1988. Leptomithrax gaimardii (H.
Milne Edwards, 1834) – 1 partially cracked female (ZRC
1965.10.13.38), trawled off Disaster Bay, New South Wales,
southern Australia, coll. M. Ward, 1930s. Leptomithrax
sternocostulatus (H. Milne Edwards, 1851) – 1 male (35.1
× 27.0 mm), 1 ovigerous female (34.0 × 25.7 mm) (ZRC
1965.10.13.74–75), Pittwater, New South Wales, southern
Australia, coll. M. Ward, December 1932. Leptomithrax
tuberculatus Whitelegge, 1900 – 1 female (36.6 × 28.7mm) (ZRC 1965.10.13.77), trawled off Newcastle, NewSouth Wales, southeastern Australia, coll. M. Ward, 1930s.
Leptomithrax eldredgei Richer de Forges & Ng, 2015
– holotype male (13.3 × 10.1 mm) (ZRC 1970.2.17.1), 2
ovigerous paratype females (21.1 × 17.4 mm, 1 broken
laterally), 1 paratype female (14.9 × 11.3 mm), 1 crushedparatype specimen (ZRC 1970.2.17.2–5), Cr 2/63, station 24,
T./10, Hong Kong, coll. & don. Fisheries Research Stationof Hong Kong, 1967.
Type species. Maja bisarmata Rathbun, 1916, by present
designation.
Etymology. The genus is named after Mary Jane Rathbun.
The name is an arbitrary combination of her family namewith Maja. Gender feminine.

Fig. 58. Third maxillipeds, Leptomithrax species. A, L. longimanus, holotype male (38.0 × 36.2 mm) (NHM 1852.12), New Zealand; B,
L. tuberculatus Whitelegge, 1900, female (36.6 × 28.7 mm) (ZRC 1965.10.13.77), Australia; C, L. sternocostulatus (H. Milne Edwards,
1851), male (35.1 × 27.0 mm) (ZRC 1965.10.13.74), Australia; D, L. edwardsii, male (75.8 × 69.4 mm) (ZRC 2001.56), Taiwan; E, L.
waitei, male (29.9 × 23.7 mm) (ZRC 2013.076, ex AM P64812), Australia; F, L. bifidus, male (32.5 × 27.5 mm) (NHM 1961.11.13.28),
Japan; G, L. longipes (G. M. Thomson, 1902), male (55.4 × 39.3 mm) (AM P86814), Australia [by S.T. Ahyong]; H, L. longipes (G. M.
Thomson, 1902) (after Griffin 1966b: fig. 15).

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Remarks. The position of the antennal flagellum, just outside
the orbit, is diagnostic of Rathbunaja n. gen. (Fig. 57G, H).
All other Leptomithrax species have the antennae positioned
well outside the orbit (Fig. 57A–F) (see Griffin, 1966b;
Griffin & Tranter, 1986). Leptomithrax Miers, 1876, itself
is a genus with 14 temperate to cold-water water species,
mostly from the southern seas (see Griffin, 1966b; Griffin& Tranter, 1986; Richer de Forges, 1993; Ng et al., 2008).
Northwestern Asia has four species, L. edwardsii (De Haan,
1835), L. bifidus (Ortmann, 1893) and L. sinensis Rathbun,
1916, and L. kiiensis T. Sakai, 1969 (Griffin, 1976; T. Sakai,
1976; Griffin & Tranter, 1986).

spines which may be short; the distal part of the basal antennal
article is relatively short with the flagellum positioned
just outside the orbit (Fig. 57A); the posterior edge of thesupraorbital eave is well separated from the postorbital
spine; the ischium of the third maxilliped is elongate andsubrectangular with the antero-lateral angle unarmed (Fig.
58A); the pollex of the chela in large adult males forms a
sharp angle with the rest of the palm; the surfaces of themale thoracic sternum and abdomen are level not distinctly
depressed; and the male abdomen is subrectangular with thelateral margins subparallel.
A second group corresponding to Austromithrax contains L.
mortenseni Bennett, 1964, L. depressus Richer de Forges,
1993, L. gaimardii (H. Milne Edwards, 1834) (= L. spinulosus
Haswell, 1880; L. australiensis Miers, 1876), L. globifer
Rathbun, 1918, L. sternocostulatus (H. Milne Edwards,
1851), L. tuberculatus Whitelegge, 1900, and L. waitei(Whitelegge, 1900). They share the following characters:
the posterior carapace margin has two spines; the distal
part of the basal antennal article is relatively wide, with theflagellum positioned on the outer edge and far from the orbit(Fig. 57B, C); the posterior edge of the supraorbital eave
is well separated from the postorbital spine; the ischium of
the third maxilliped is relatively shorter, subrectangular to
subtriangular, and the antero-lateral angle has a distinct spuror tooth (Fig. 58B, C, E); the pollex of the chela in large adultmales is smoothly contiguous with the rest of the palm; themale thoracic sternum and abdomen may or may not havedepressions; and the male abdomen is subtriangular with thelateral margins sloping towards the broader posterior part.
On the basis of these characters, three of the northeastern
Asian species (L. edwardsii, L. bifidus, L. sinensis) can bereferred to Austromithrax (cf. Figs. 57D, E, 58D, F). Thatsaid, this genus remains heterogeneous and the contained
species should be revised to see if other natural groupings
are apparent. Of these species, L. sinensis Rathbun, 1916,
may be a junior subjective synonym of L. bifidus (Ortmann,
1893). Richer de Forges & Ng (2015) discussed this matterat length when describing a new species, L. eldredgei, from
Hong Kong and commented that the type of L. sinensis isjust a carapace and only has short spines.

Bennett (1964) argued that three subgenera can be recognised
in Leptomithrax – Leptomithrax (Leptomithrax) s. str. (typespecies Paramithrax (Leptomithrax) longimanus Miers,
1876), Leptomithrax (Austromithrax) Bennett, 1964 (typespecies Leptomithrax (Austromithrax) mortenseni Bennett,
1964) and Leptomithrax (Zemithrax) Bennett, 1964 (typespecies Paramithrax longipes G. M. Thomson, 1902). As
discussed at length by Griffin (1966b: 61), Bennett (1964)
distinguished these subgenera on the basis of the positionand structure of the basal antennal article and flagellum, formof the supraorbital margin, whether the region between themerus and ischium of the third maxilliped is swollen, andthe presence or absence of depressions on the male thoracic
sternum and/or abdomen. Griffin (1966b) argued that whilesome of the characters were useful, others appeared to beless reliable and varied between species. In addition, he
was not certain about the placement of some species of
Leptomithrax and was concerned that some of the groupings
ended up with only a few species. As such, he preferred torecognise just one genus without subgenera for Leptomithrax.
He maintained this stand in his revision of the Indo-West
Pacific Majoidea (Griffin & Tranter, 1986: 208), although
he established a new monotypic genus, Teratomaia Griffin& Tranter, 1986, for Leptomithrax richardsoni Dell, 1960;
and transferred L. parvispinosus (Ward, 1933) to a redefined
Tumulosternum McCulloch, 1913.
Looking at the published descriptions of the various
Australian and New Zealand species, it is clear that there areat least three groups among the 11 species from this region.
They correspond to Bennett’s (1964) three groupings welland these should eventually be recognised as distinct genera.
Interestingly, McLay et al. (1995) conducted a cladistic
analysis of the fossil and extant species of Leptomithrax,
and recognised three groups of extant species which roughly
correspond to Bennett’s (1964) subgenera. However, wedefer from formally adopting Bennet’s subgenera as separate
genera as this should only be done as part of a revision of
the species of Leptomithrax. This is now being planned. It
is nevertheless useful to provide a synopsis of the genus.

The third group corresponding to Zemithrax contains
only L. longipes (G. M. Thomson, 1902) (= Leptomithrax(Zemithrax) molloch Bennett, 1964). Its defining suite ofcharacters are as follow: the posterior carapace margin has
only one spine or tooth; the posterior edge of the supraorbital
eave and postorbital spine almost touch each other; thedistal part of the basal antennal article is relatively narrow,with the flagellum positioned submedially and distant fromthe orbit (Fig. 57F); the ischium of the third maxilliped
is relatively short, subtriangular with the distal part muchwider than the proximal part, the anterolateral angle has adistinct tooth and the setal arrangement is very distinctive(Fig. 58G, H); the pollex of the chela in large adult males
is smoothly contiguous with the rest of the palm; the malethoracic sternum and abdomen have distinct depressions; andthe male abdomen is subtriangular with the lateral margins
sloping towards the broader posterior part.

Leptomithrax s. str. contains L. longimanus (Miers, 1876)(= Paramithrax (Leptomithrax) affinis Borradaile, 1916),
L. australis (Jacquinot, in Jacquinot & Lucas, 1853) (=Paramithrax (Leptomithrax) brevirostris Miers, 1879), andL. garricki Griffin, 1966. These species share the following
suite of characters: the posterior carapace margin has two
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Fig. 59. General habitus, Rathbunaja species. A, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM 48220), Philippines; B, R.
bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; C, R. kiiensis, holotype male (53.7 × 45.0 mm) (USNM 125885), Japan;
D, R. kiiensis, female (43.8 × 37.3 mm) (NSMT-Cr 6419), Kyushu-Palau Ridge; E, R. kiiensis, male (49.9 × 39.1 mm) (NSMT-Cr 5094),
Japan; F, R. brevipes n. sp., holotype male (24.0 × 18.4 mm) (MNHN-IU-2011-2408), Papua New Guinea.

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One of the characters used by Bennett (1964: 46, 53) toseparate Zemithrax from the others was what he describes
as a large boss (swelling) being present on the merus andischium at the junction where they articulate. However, he
admitted that while it was very obvious in the type species,
L. longipes (Fig. 58G, H), it was also present but less distinct
in other species, with L. globifer having the merus but notischium swollen, while in L. gaimardi (as L. australiensis),the ischium and merus are slightly swollen (Bennett, 1964:
51). In the specimens on hand, the third maxilliped of L.
tuberculatus has a small but distinct swelling only on themerus (Fig. 58B), whereas in L. sternocostulatus, the swellingis even smaller and less distinct (Fig. 58C). In L. bifidus, theswelling in the merus and ischium of the third maxilliped

is very prominent (Fig. 58F), not unlike that described andfigured for L. longipes (cf. Fig. 58G, H; Bennett, 1964: figs.
50, 53). In addition, the shape of the ischium also appears tovary from being short with the proximal part much narrower
than the distal part (e.g., L. tuberculatus) (Fig. 58B) to being
relatively longer with the difference in proportions lessobvious (e.g., L. sternocostulatus, Fig. 58C).
The northwestern Asian Leptomithrax kiiensis is very
distinct from all other Leptomithrax and belongs to its
own group, here named Rathbunaja n. gen. Rathbunaja isclosest to Leptomithrax s. str. in the male thoracic sternum
and abdomen not having any trace of depressions; the distal
part of the basal antennal article is relatively short with

Fig. 60. General habitus. A, Rathbunaja ursus n. sp., holotype male (50.1 × 41.5 mm) (NMCR, ex ZRC 2013.1275), Philippines; B,
Rathbunaja ursus n. sp., paratype male (41.6 × 32.8 mm) (ZRC 2013.1276), Philippines; C, Rathbunaja ursus n. sp., paratype female (49.4
× 38.4 mm) (NSMT-Cr 15386), Philippines; D, Rathbunaja ursus n. sp., paratype ovigerous female (46.8 × 36.6 mm) (ZRC 2001.0594),
Philippines

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the flagellum positioned just outside the orbit; the formof the third maxillipeds (ischium subrectangular without
tooth or projection on antero-lateral angle of the ischium);the pollex of the chela in large adult males forms a sharp
angle with the palm; and the subrectangular male abdomenwith the lateral margins subparallel. However, Rathbunaja

can easily be separated by the different carapace shape,
being less inflated (notably in the branchial regions) withthe pseudorostral and lateral spines proportionately longer(Figs. 59, 60); and having a prominent distal spine on thedorsal margin of the merus of the ambulatory leg (Fig. 65B,
D, F, I–K) (absent in Leptomithrax s. str.).

Fig. 61. Antennae, antennules, orbits and epistomes, Rathbunaja species. A, B, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM
48220), Philippines; C, D, R. bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; E, F, R. kiiensis, male (49.9 × 39.1 mm)(NSMT-Cr 5094), Japan; G, H, R. ursus n. sp., holotype male (50.1 × 41.5 mm) (NMCR, ex ZRC 2013.1275), Philippines.

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Key to species of Rathbunaja n. gen.

–

1. Pseudorostral spines short, weakly diverging, almost subparallel(Figs. 59F, 62A); ambulatory legs (notably merus and propodus)
short (Figs. 59F, 62D–G); Papua New Guinea.........................
........................................................ Rathbunaja brevipes n. sp.
– Pseudorostral spines long, diverging; ambulatory legs (notably
merus and propodus) relatively longer....................................2

Carapace relatively narrower (Fig. 59A–E); distal spines onbasal antennal article relatively shorter, less diverging (Fig.
61A, C, E); ischium of third maxilliped proportionately shorter(Figs. 62C, 63A–D); male telson more triangular in shape (Fig.
64A–C); G1 gently curved, not elongate, tip straight (Figs.
66).............................................................................................3

3. Ambulatory merus relatively longer, more slender (Figs. 59C, E,65F); G1 relatively more strongly curved, distal part relatively
longer (Fig. 66H–M); Japan......................................................
........................................Rathbunaja kiiensis (T. Sakai, 1969)– Ambulatory merus relatively shorter, stouter (Figs. 59A, B,
65B, D); G1 gently curved, distal part relatively shorter (Fig.
66A–G); Philippines...Rathbunaja bisarmata (Rathbun, 1916)

2. Carapace relatively broad (Fig. 60); distal spines on basal
antennal article relatively longer, more strongly diverging(Fig. 61G); ischium of third maxilliped proportionately longer(Fig. 63E–H); male telson proportionately broader, appears
semicircular (Fig. 64E); G1 very long, strongly curved, more
slender, tip bent inwards (Fig. 67A–D); Philippines................
............................................................. Rathbunaja ursus n. sp.

Fig. 62. Rathbunaja brevipes n. sp., holotype male (24.0 × 18.4 mm) (MNHN-IU-2011-2408), Papua New Guinea. A, frontal region, dorsal
view; B, antennae, antennules and epistome; C, left third maxilliped; D–G, first to fourth right ambulatory legs; H, left chela.

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Rathbunaja bisarmata (Rathbun, 1916)(Figs. 59A, B, 61A–D, 63A, B, 64A, B, 65A–D, 66A–G)

59A, B, 65B, D). Male telson subtriangular (Fig. 64A, B).
G1 gently curved, distal part relatively shorter (Fig. 66A–G).

Maja bisarmata Rathbun, 1916: 553. – Serène, 1968: 57. – Griffin,
1976: 199, Fig. 7b. – Griffin & Tranter, 1986: 211 (key).

Remarks. This is a very poorly known species and although
it has been treated as a valid Maja species by Griffin (1976)(who examined the type) and Griffin & Tranter (1986), are-examination of the holotype shows otherwise. Although
Rathbun (1916) placed this species in Maja, its antennal
flagellum is actually just outside the orbit. In fact, the species
is very close to Leptomithrax kiiensis, described by T. Sakai(1969) on the basis of a large male 55.0 by 44.5 mm fromKii Minable in Japan. Tune Sakai (1969, 1976) placed thisspecies in Leptomithrax because its antennal flagellum isjust outside the orbit, even if it is not to the same degree
as other congeners.

Material examined. Holotype: male (20.3 × 15.6 mm)(USNM 48220), station 5519, Point Tagolo Light, offnorthern Mindanao, 80°47’N 123°31.25’E, Philippines,
coll. RV Albatross, 9 August 1909. Others: Philippines – 1
male (37.1 × 29.6 mm) (ZRC 2013.1274), Balicasag Island,
Panglao, Bohol, 200–300 m, coll. fishermen with tangle
nets, 2 March 2004.
Diagnosis. Carapace relatively narrow (Fig. 59A, B).
Pseudorostral spines long, diverging (Fig. 59A, B). Medianrow with 2 median gastric spines; 2 cardiac spines; 2 intestinal
spines (Fig. 59A, B). Basal antennal article with 2 relatively
short distal spines not strongly diverging (Fig. 61A–D).
Ischium of third maxilliped proportionately shorter (Fig.
63A, B). Ambulatory merus relatively shorter, stouter (Figs.

The holotype male of Maja bisarmata is relatively small(Fig. 59A) and although its G1 is not fully developed (Fig.
61A–D), it shows enough features. A recent larger specimen
from the Philippines agrees well with the type male in its
carapace form, antennal basal article, epistome, structure

Fig. 63. Third maxillipeds, Rathbunaja species. A, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM 48220), Philippines; B, R.
bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; C, R. kiiensis, male (49.9 × 39.1 mm) (NSMT-Cr 5094), Japan; D,
R. kiiensis, ovigerous female (43.8 × 37.3 mm) (NSMT-Cr 6419), Kyushu-Palau Ridge; E, R. ursus n. sp., holotype male (50.1 × 41.5mm) (NMCR, ex ZRC 2013.1275), Philippines; F, R. ursus n. sp., paratype male (41.6 × 32.8 mm) (ZRC 2013.1276), Philippines; G,
R. ursus n. sp., paratype female (45.0 × 36.0 mm) (ZRC 2001.0594), Philippines; H, R. ursus n. sp., paratype ovigerous female (46.8 ×
36.6 mm) (ZRC 2001.0594), Philippines.

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Rathbunaja kiiensis (T. Sakai, 1969)(Figs. 59C–E, 61E, F, 63C, D, 64C, 65E–G, 66H–M)

of the third maxilliped, relatively short ambulatory legs,
thoracic sternum, male abdomen and structure of the G1,
and we are confident they are conspecific. These features
make it unlikely that Maja bisarmata is the juvenile formof the more common new species described here from thePhilippines, Rathbunaja ursus.

Leptomithrax kiiensis T. Sakai, 1969: 255, pl. 1 fig. 4. – Miyake,
1983: 48, pl. 16 fig. 4. – T. Sakai, 1976: 243, text fig. 130. –Matsuzawa, 1977: pl. 96 fig. 3. – Baba et al., 1986: 223, fig.
167. – Takeda, 1980: 280. – Takeda, 1993: 37. – Ikeda, 1998:
37, 118, 119, pl. 42.

Maja bisarmata Rathbun, 1916, is very close to Leptomithrax
kiiensis T. Sakai, 1969. Comparisons of Rathbun’s species
with material of L. kiiensis from Japan as well as thedescriptions and figures of the species by T. Sakai (1969,1976), Miyake (1983) and Ikeda (1998) show minimal
differences. The main difference appears to be the relative
lengths of the merus of the ambulatory leg, which in R.
kiiensis, is proportionately longer and more slender (Figs.
59C, E, 65F) than that of R. bisarmata (Figs. 59A, B, 65B,
D). The distal part of the G1 of R. kiiensis is more curved
and slightly longer than that of R. bisarmata (Fig. 66H–M
versus Fig. 66A–G). As such, both species are recognised forthe time being. As discussed under the genus, the distinctive
features of these two species (as well as two other new ones
here described) warrant the establishment of a new genus,
Rathbunaja, for them.

Material examined. Holotype: male (53.7 × 45.0 mm)(USNM 125885), Kii Minabe, Kii Peninsula, Wakayama
Prefecture, Honshu, Japan, coll. M. Ozaki. Others: Japan– 1 male (49.9 × 39.1 mm) (NSMT-Cr 5094), Kushimoto,Kii Peninsula, coll. 15 July 1977. — 1 ovigerous female(43.8 × 37.3 mm) (NSMT-Cr 6419), 26°43.8’N 135°20’E,
Kita-Koho Channel, Kyushu-Palau Ridge, 320 m, coll. 14
February 1978. — 1 female (44.0 × 34.8 mm) (NSMT-Cr
12211), station T13, 29°25.8’N 127°18.0’E – 29°24.4’N
127°17.9’E, East China Sea, 169 m, coll. 27 September 1997.
Diagnosis. Carapace relatively narrow (Fig. 59C–E).
Pseudorostral spines long, distinctly diverging (Fig. 59C–E).
Median row with 2 median gastric spines; 2 cardiac spines;
2 or 3 intestinal spines (Fig. 59C–E). Basal antennal article
with 2 relatively short distal spines not strongly diverging(Fig. 61E). Ischium of third maxilliped proportionately

Fig. 64. Male anterior thoracic sternums and abdomens, Rathbunaja species. A, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM
48220), Philippines; B, R. bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; C, R. kiiensis, male (49.9 × 39.1 mm)(NSMT-Cr 5094), Japan; D, R. brevipes n. sp., holotype male (24.0 × 18.4 mm) (MNHN-IU-2011-2408), Papua New Guinea; E, R. ursus
n. sp., holotype male (50.1 × 41.5 mm) (NMCR, ex ZRC 2013.1275), Philippines; F, R. ursus n. sp., paratype male (41.6 × 32.8 mm)(ZRC 2013.1276), Philippines.

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Fig. 65. Chelipeds and ambulatory legs, Rathbunaja species. A, B, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM 48220),
Philippines; C, D, R. bisarmata, male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; E, F, R. kiiensis, male (49.9 × 39.1 mm) (NSMTCr 5094), Japan; G, R. kiiensis, holotype male (53.7 × 45.0 mm) (USNM 125885), Japan; H, I, R. ursus n. sp., holotype male (50.1 ×
41.5 mm) (NMCR, ex ZRC 2013.1275), Philippines; J, R. ursus n. sp., paratype male (41.6 × 32.8 mm) (ZRC 2013.1276), Philippines;
K, R. ursus n. sp., paratype female (45.0 × 36.0 mm) (ZRC 2001.0594), Philippines.

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Fig. 66. Left G1s, Rathbunaja species. A–D, R. bisarmata, holotype male (20.3 × 15.6 mm) (USNM 48220), Philippines; E–G, R. bisarmata,male (37.1 × 29.6 mm) (ZRC 2013.1274), Philippines; H–J, R. kiiensis, holotype male (53.7 × 45.0 mm) (USNM 125885), Japan; K–M,
R. kiiensis, male (49.9 × 39.1 mm) (NSMT-Cr 5094), Japan. Scales: A, H, K = 1.0 mm; E = 5.0 mm; B–D, F, G, I, J, L, M= 0.5 mm.

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Fig. 67. Left G1s, Rathbunaja species. A–D, R. ursus n. sp., holotype male (50.1 × 41.5 mm) (NMCR, ex ZRC 2013.1275), Philippines;
E–G, R. brevipes n. sp., holotype male (24.0 × 18.4 mm) (MNHN-IU-2011-2408), Papua New Guinea. A, E = 5.0 mm; B-G = 1.0 mm;
F, G = 0.5 mm.

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nets, 2 March 2004. Paratypes: Philippines – 2 ovigerous
females (46.8 × 36.6 mm, 45.0 × 36.0 mm) (ZRC 2001.0594),
Balicasag Island, Panglao, Bohol, 200–300 m, coll. fishermenwith tangle nets, 28 November 2001. — 1 male (41.6 × 32.8mm), 1 ovigerous female (45.6 × 35.7 mm), 1 female (47.2
× 37.6 mm) (ZRC 2013.1276), Balicasag Island, Panglao,
Bohol, 200–300 m, coll. fishermen with tangle nets, June
2002. — 1 female (49.4 × 38.4 mm) (NSMT-Cr 15386),
Balicasag Island, Panglao, Bohol, coll. fishermen with tangle
nets, via H. Komatsu & M. Takeda, February 2003. — 1
male (47.5 × 39.5 mm) (ZRC 2013.1277), Maribohoc Bay,
Panglao, Bohol, coll. J. Arbasto, tangle nets, 30 May 2004.

shorter (Fig. 63C, D). Ambulatory merus relatively long,
slender (Figs. 59C, E, 65F). Male telson subtriangular (Fig.
64C). G1 relatively more curved, distal part relatively longer(Fig. 66H–M).
Remarks. The type of Rathbunaja kiiensis is the largest
known specimen. Although it is only slightly larger thananother male specimen (male 49.9 × 39.1 mm, NSMT-Cr5094), its chela is more prominently enlarged, with a distinct
swelling on each margin of the palm (Fig. 65G versus Fig.
65E). The G1 has not been figured before. The G1 of theholotype male is slightly less strongly curved than the othermale and the dorsal fold on the distal part of the G1 of theholotype male is also more pronounced (Fig. 66H–J versus
Fig. 66K–M). However, we do not regard these differences
as significant at the species-level.

Diagnosis. Carapace relatively broad (Fig. 60). Pseudorostralspines long, diverging (Fig. 60). Median row with 2 mediangastric spines; 2 cardiac spines; 2 intestinal spines (Fig. 60).
Basal antennal article with 2 relatively long distal spines,
strongly diverging (Fig. 61G). Ischium of third maxilliped
long (Fig. 63E–H). Ambulatory legs (notably merus andpropodus) relatively long (Figs. 60, 65I–K). Male telson
appears semicircular (Fig. 64E). G1 very long, strongly
curved, slender, tip bent inwards (Fig. 67A–D).

Rathbunaja brevipes n. sp.
(Figs. 59F, 62, 64D, 67E–G)
Material examined. Holotype: male (24.0 × 18.4 mm)(MNHN-IU-2011-2408), station DW 3719, 6°03’S 14°36’E,
410 m, Papua New Guinea, coll. RV Alis, BIOPAPUA, 7
October 2010.

Etymology. The name “ursus” alludes to the thick tomentum
covering the animal like the fur of a bear, «ursus». The nameis used as a noun in apposition.

Diagnosis. Carapace relatively narrow (Figs. 59F, 62A).
Pseudorostral spines short, weakly diverging, almost
subparallel (Fig. 59F). Median row with 2 median gastricspines; 2 cardiac spines; 3 intestinal spines (Fig. 59F). Basalantennal article with 2 relatively short distal spines notstrongly diverging (Fig. 62B). Ischium of third maxilliped
proportionately shorter (Fig. 62C). Ambulatory legs (notably
merus and propodus) short (Figs. 59F, 62D–G). Male telson
subtriangular (Fig. 64D). G1 with distal part distinctly curved,
dorsal fold low (Fig. 67E–G).

Remarks. The discovery of Rathbunaja ursus n. sp. from
Balicasag, Philippines, is rather surprising especially since
there was also a specimen of R. bisarmata (Rathbun, 1916)
collected from the same area. Most specimens of Rathbunaja
from the Philippines actually belong to this new species.
The two species are similar and are not easy to separate;
In R. ursus, however, the carapace is relatively broader(Fig. 60 versus Fig. 59A, B), the two distal spines on thebasal antennal article are more strongly diverging (Fig. 61G
versus Fig. 61A, C), the ischium of the third maxilliped is
proportionately longer (Fig. 63E–H versus Fig. 63A, B),
the anterior thoracic sternum (sternites 1–4) is relatively
longer (Fig. 64E, F versus Fig. 64A, B), the male telson is
proportionately broader (Fig. 64E, F versus Fig. 64A, B),
and the lateral margins of male abdominal somites 5 and 6
are more concave (Fig. 64E, F versus Fig. 64A, B). Most
significantly, their G1 structures are completely different,with that of R. ursus much longer, with the distal part morestrongly curved and slender (Fig. 67A–D versus Fig. 66A–G).
These differences cannot be explained by size or maturity.

Etymology. The species is named after its relatively shortambulatory legs. Used as a noun in apposition.
Remarks. Rathbunaja brevipes n. sp. is perhaps closest to
R. kiiensis but can easily be distinguished by possessing
relatively much shorter pseudorostral spines (Figs. 59F,
62A versus Fig. 59C–E); a relatively larger and broader
intercalated spine (Figs. 59F, 62A) (more triangular andsharper in R. kiiensis, Fig. 59C–E); proportionately shorter
lateral carapace spines (Fig. 59F versus Fig. 59C–E); andshorter ambulatory legs, especially the merus, which also
has a relatively weaker distal dorsal spine (Figs. 59F, 62D–G
versus Figs. 59C–E, 65F). Interestingly, the G1 structuresof the two species (Figs. 67E–G, 66H–M) are very similar.
The ambulatory legs of R. brevipes are also shorter thanthose on R. bisarmata (Figs. 59F, 62D–G versus Figs. 59A,
B, 65B, D).

GENERAL DISCUSSION
In the present revision, we separate Maja into 10 genera,
seven of which are new, and recognise 17 new species fromthe Indo-West Pacific. Considering that Maja s. lato has
never previously been revised, the number of new taxa is not
surprising. The most remarkable aspect of the present study
is the record of 11 species from one site, Balicasag Island,in the Philippines. Of these, nine are new to science. If theareas around Balicasag are considered, a total of 13 species
are known. The proportion of new species is very high. Thistiny island in the Bohol Sea has been previously regarded

Rathbunaja ursus n. sp.
(Figs. 60, 61G, H, 63E–H, 64E, F, 65H–K, 67A–D)
Material examined. Holotype: male (50.1 × 41.5 mm)(NMCR, ex ZRC 2013.1275), Balicasag Island, Panglao,
Bohol, Philippines, 200–300 m, coll. fishermen with tangle
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Fig. 68. Colours in life. A, Paramaja kominatoensis, female (ca. 57.8 mm carapace length) (after Ikeda, 1998: pl. 41-1b), Taiwan; B,
Paramaja gibba, male (58.6 × 52.2 mm) (MNHN-IU-2010-63), Madagascar; C, Paramaja gibba, small female (45.1 × 40.6 mm) (MNHNIU-2010-467), Madagascar; D, Paramaja turgida n. sp., holotype male (74.1 × 66.8 mm) (NMCR), Philippines; E, Paramaja turgida n.
sp., paratype male (70.4 × 63.9 mm) (ZRC 2012.1201), Philippines; F, Paramaja turgida n. sp., paratype female (50.5 × 44.8 mm) (ZRC
2013.1238), Philippines.

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

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Fig. 70. Colours in life. A, Paramaya ouch n. sp., paratype female (42.6 × 34.5 mm) (ZRC 2013.1300), Philippines; B, Paramaya
coccinea n. sp., holotype male (69.0 × 55.6 mm) (MNHN), Vanuatu; C, Sakaija africana, male (27.3 × 21.2 mm) (MNHN-IU-2010-504),
Madagascar; D, Sakaija serenei n. sp., paratype male (14.3 × 10.9 mm) (ZRC 2013.1255), Philippines; E, Sakaija santo n. sp., holotype
male (9.4 × 6.6 mm) (MNHN), Vanuatu; F, Ovimaja compressipes, male (55.7 × 42.7 mm) (ZRC 2013.1264), Taiwan.

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Ng & Richer de Forges: Revision of the spider crab genus Majaas perhaps the richest site for many groups of molluscs anddecapod crustaceans (see Ng et al., 2009; Mendoza et al.,2010). Almost all the new species were collected by tangle
nets, a technique that has proved especially effective insampling steep rocky slopes, and numerous new taxa havebeen discovered from this island using these nets over the last
decade (see Bouchet et al., 2009; Ng et al., 2009; Mendoza etal., 2010). The zone between 200 and 600 m deep is clearly
very rich in species, but cannot effectively be sampled withother methods. Of the new taxa found, Paramaya ouch n.
sp., Alcomaja irrorata n. sp., and Holthuija pauli n. sp.,were very common in Balicasag, but were not obtained bytrawling, even when done around the island (Bouchet et al.,2009; Richer de Forges et al., 2009). They have also notbeen collected by the Albatross and MUSORSTOM cruises
in the Philippines, which used trawls and dredges.

Croatia). Colin McLay and Shane Ahyong also provided
important reviews that improved the paper substantially,
while Michael Türkay was very helpful with many typematters and kindly passed many interesting specimens to thefirst author during his visit to SMF. Martyn Low was very
helpful as always in finding all the key literature to help solve
some messy nomenclatural problems. Tohru Naruse helped
us with photographs of some Taiwanese taxa, Zeehan Jaafar
helped with photographs of some USNM specimens, while
Rohan Pethiyagoda kindly passed us a photograph of the Sri
Lanka specimen. Chan Tin-Yam, Lin Chia-Wei, Jose C.E.
Mendoza and Tan Heok Hui kindly helped with photographsof Philippines, Vanuatu and Taiwan specimens. We thankCaptain Jean-François Barazer who helped us with the precise
coordinates for many localities. The expeditions we have been
involved in also obtained many important specimens. We
thank the crew and scientists on board the research vessels
DA-BFAR, Alis, Vizconde De Eza and Miriky who collectedmany important samples from the Indian Ocean and Pacific.
The various Philippine expeditions were jointly organised
by MNHN (through Philippe Bouchet); Bureau of Fisheries
and Aquatic Resources (BFAR), Manila (through Ludivina
Labe); National Museum of the Philippines (NMCR) (throughMarivene Manuel-Santos); University of San Carlos (USC,
Cebu) (through Danilo Largo); National Ocean University,
Keelung, Taiwan (through Chan Tin-Yam); RMBR; ASEAN
Research Centre for Biodiversity and Conservation (Manila);and the National University of Singapore, with key funding
from the Total Foundation and French Ministry of Foreign
Affairs. The SANTO 2006 expedition was operated by
MNHN, Pro Natura International (PNI) and Institut deRecherche pour le Développement (IRD) under a permit

The question is simple: why is this one small area so rich in
species? Ng et al. (2009) and Mendoza et al. (2010) argued
that it was probably a consequence of sampling bias because
many other such habitats have never been previously sampledwith such an efficient method. These results have serious
implications for our ongoing efforts to estimate global marine
biodiversity. Today, scientists are trying to accomplish thisusing different approaches and models (see Tittensor et al.,2010; O’Hara & Tittensor, 2010) or direct extrapolation
based on expert knowledge (see Appeltans et al., 2012).
These methods, have arrived at figures of between 8 and 10
million marine species. However, the case with Balicasag
suggests we may well be underestimating the actual numberof species known. The fact that these species-rich habitats –the steep slopes of islands from 80 m and below – are noteasily sampled by traditional methods suggests we may needto relook at our methods for marine species estimation (seealso Kitchingman & Lai, 2004; Ausubel, 2013).

issued to Philippe Bouchet (MNHN), and was part of the“Census of Marine Life’s Reefs programme funded by the
Total Foundation and Sloan Foundation. The Mozambiquematerial came from the MAINBAZA cruise under MNHN
(Philippe Bouchet chief scientist) and Instituto Espanolde Oceanografia, as part of the Mozambique-Madagascar
expeditions with Pro-Natura International funded by Total
Foundation, Prince Albert II of Monaco Foundation andStavros Niarchos Foundation. The Singapore material was
obtained and part of the second author’s time in ZRC was
funded, by the Comprehensive Marine Biodiversity Survey,a three year project of the National Parks Board (NParks),
Tropical Marine Science Institute (TMSI) and Lee Kong
Chian Natural History Museum; with financial support from
Shell Singapore Pte Ltd., Asia- Pacific Breweries (Singapore),and Hong Kong & Shanghai Banking Corporation (HSBC
Singapore).

ACKNOWLEDGEMENTS
This study has been helped by many friends and colleagues
who have helped us in one form or another (especially withmaterial), for which we are most grateful: Paul Clark (NHM),
Charles Fransen and Karen Van Dorp (RMNH), Karen Reed
and Rafael Lemaitre (USNM), Marivene Manuel-Santos(NMCR), Takehiru Sato (KPM), Hironori Komatsu (NSMT),
Tomoyuki Komai (CBM), Anouchka Sato, Paula Lefevre andLaure Corbari (MNHN), Chan Tin-Yam (National Taiwan
Ocean University, Keelung, Taiwan), N. Kakati (India),Tan Siong Kiat, Martyn Low, Zeehan Jaafar and Ng Ngan
Kee (ZRC), Shane Ahyong (AM), Jiang Wei (IOCAS),
Chang Hsien Chung (PCM), Wang Chia Hsiang (ex-Taiwan
Museum, Taipei), Oliver Coleman (Berlin Museum), Michael
Türkay and Kristin Arnold (SMF), Enrique Macpherson(Barcelona), Sean Fennessy (Oceanographic ResearchInstitute, Durban, South Africa), Gary Poore and Joanne
Taylor (NMV), Colin McLay (University of Canterbury,
New Zealand), Kareen Schnabel (NIWA), Marie Meisterand Dominique Wandhammer (MZS), Vasily Spiridonov(SIO), Marin Kirinčić (Natural History Museum Rijeka
Lorenzov, Rijeka, Croatia) and Zdravko Števčić (Rovinj,

LITERATURE CITED
Abelló P, Guerao G, Salmerón F & Rago JEC (2014) Maja
brachydactyla (Brachyura: Majidae) in the western
Mediterranean. Marine Biodiversity Records, 7: e77. doi:
10.1017/S17.
Adams A & White A (1848–1849) Crustacea. In: Adams A (ed.)
The zoology of the voyage of H.M.S. Samarang, under thecommand of Captain Sir Edward Belcher, C.B., F.R.A.S.,
F.G.S. during the years 1843–1846. Reeve, Benham, andReeve, London, i–viii [= Front Matter and Preface by Adams,

220

RAFFLES BULLETIN OF ZOOLOGY 2015
A.] + i–xv [= Introduction by Adams, A. and References toPlates] + 1–66 pp., pls. I–VI.
Alcock A (1895) Materials for a carcinological fauna of India, 1.
The Brachyura Oxyrhyncha. Journal of the Asiatic Society ofBengal, Calcutta, 64: 157–291, pls. 3–5.
Alcock A (1899) An account of the deep-sea Brachyura collectedby the Indian Marine Survey Ship Investigator. Trustees of the
Indian Museum, Calcutta. i–iv + 1–85 pp., pls. 1–4.
Alcock A & Anderson ARS (1898) Illustrations of the Zoologyof the Royal Indian Marine Survey Ship Investigator, under
the command of Commander T. H. Heming, R.N. Fishes.-Part
V, Plates XVIII-XXIV. Crustacea.-Part VI, Plates XXXIIIXXXV. Mollusca.-Part H, Plates VII-VIII. Published underthe Authority of Captain W. S. Goodridge, R.N., Director of
the Royal Indian Marine, Calcutta.
Appeltans W, Ahyong ST, Anderson G, Angel MV, Artois T,
Bailly N, Bamber R, Barber A, Bartsch I, Berta A, BłażewiczPaszkowycz M, Bock P, Boxshall G, Boyko CB, Brandão SN,
Bray RA, Bruce NL, Cairns SD, Chan T-Y, Cheng L, Collins
AG, Cribb T, Curini-Galletti M, Dahdouh-Guebas F, DaviePJF, Dawson MN, De Clerck O, Decock W, DeGrave S, de
Voogd NJ, Domning DP, Emig CC, Erséus C, Eschmeyer W,
Fauchald K, Fautin DG, Feist SW, Fransen CHJM, Furuya
H, Garcia-Alvarez O, Gerken S, Gibson D, Gittenberger A,
Gofas S, Gómez-Daglio L, Gordon DP, Guiry MD, HernandezF, Hoeksema BW, Hopcroft R, Jaume D, Kirk P, Koedam N,
Koenemann S, Kolb JB, Kristensen RM, Kroh A, Lambert G,
Lazarus DB, Lemaitre R, Longshaw M, Lowry J, Macpherson
E, Madin LP, Mah C, Mapstone G, McLaughlin PA, Mees J,
Meland K, Messing CG, Mills CE, Molodtsova TN, Mooi R,
Neuhaus B, Ng PKL, Nielsen C, Norenburg J, Opresko DM,
Osawa M, Paulay G, Perrin W, Pilger JF, Poore GCB, Pugh
P, Read GB, Reimer JD, Rius M, Rocha RM, Saiz-Salinas JI,
Scarabino V, Schierwater B, Schmidt-Rhaesa A, Schnabel KE,
Schotte M, Schuchert P, Schwab E, Segers H, Self-Sullivan
C, Shenkar N, Siegel V, Sterrer W, Stöhr S, Swalla B, Tasker
ML, Thuesen EV, Timm T, Todaro A, Turon X, Tyler S, Uetz
P, van der Land J, Vanhoorne B, van Ofwegen LP, van Soest
RWM, Vanaverbeke J, Walker-Smith G, Walter TC, Warren
A, Williams G, Wilson SP Costello MJ (2012) The magnitudeof global marine species diversity. Current Biology, 22(23):2189–2202.
Ausubel JH (2013) Wealth from the Oceans: Use, Stewardship,
and Security. Urban Coast Institute, Monmouth University andRockefeller Institute, 23 pp.
Baba K, Hayashi K-I & Toriyama M (1986) Decapod Crustaceansfrom continental shelf and slope around Japan. JapaneseFisheries Resource Conservation Association, Tosho Printing
Co., Ltd., Tokyo, 336 pp.
Balss H (1922) Crustacea VI: Decapoda Anomura (Paguridea) und
Brachyura (Dromiacea bis Brachygnatha). In: Michaelsen W(ed.) Beiträge zur Kenntnis der Meeresfauna Westafrikas, L.
Friederichsen & Co., Hamburg, Band 3, Lieferung 2: 37–67,
Figs. 1–7.
Balss H (1924) Ostasiatische Decapoden. V. Die Oxyrhynchen
und Schlussteil. Archiv für Naturgeschichte, 90(A5): 20–84.
Balss H (1957) Decapoda. VIII. Systematik. In: Dr. H. G.
Bronns, Klassen und Ordnungen des Tierreichs. Leipzig andHeidelberg, Winter. Fünfter Band, I. Abteilung, 7. Buch, 12.
Lief.: 1505–1672, figs. 1131–1199.
Barnard KH (1950) Descriptive catalogue of South African decapod
Crustacea (crabs and shrimps). Annals of the South African
Museum, 38: 1–837, figs. 11–154.
Baudouin M (1931) L’araignée de mer dite ‘Abraham’ en
Vendée. Espèce nouvelle: Maia gigantea. Comptes Rendus
de l’Association Française pour l’Avencement des Sciences,55: 256–258.

Bennett EW (1964) The marine fauna of New Zealand: Crustacea
Brachyura. New Zealand Department of Scientific and Industrial
Research, Bulletin, 153 [= New Zealand Oceanographic InstituteMemoir, 22]: 1–120, figs. 1–141, frontispiece.
Borradaile LA (1916) Crustacea. Part I.—Decapoda. In: British
Antartic (“Terra Nova”) Expedition, 1910. Natural History
Report. Zoology, Trustees of the British Museum, British
Museum (Natural History), London, 3(2): 75–110, figs. 1–16
Bouchet P, Ng PKL, Largo D & Tan SH (2009) PANGLAO
2004 – Investigations of the marine species richness in thePhilippines. Raffles Bulletin of Zoology, Supplement 20: 1–19.
Bouvier E-L (1940) Décapodes marcheurs. In: Faune de France,
Volume 37. Paul Lechevalier et Fils, Paris, 404 pp.
Capart A (1951) Crustacés Décapodes Brachyures. In: Expedition
océanographique Belge dans les eaux cötières Africaines del’Atlantique Sud (1948–1949), Résultats Scientifiques, 3(1):11–205.
Carmona-Suarez CA (1990) An unusual type of heteromorphosis
in a brachyuran crab Maja crispata Risso, 1827 (Decapoda,
Majidae). Crustaceana, 59: 220–223.
Charpentier T von (1829) Bemerkungen über einige Crustaceen
des mitteländischen Meeres. Verhandlungen der Gesellschaft
Naturforschender Freunde zu Berlin, 1: 157–173.
Chen HL & Ng PKL (1999) Description of a new spider crab,
Maja gracilipes, from the South China Sea, with notes on thetaxonomic validity of M. brevispinosis Dai, 1981 (Crustacea:
Decapoda: Brachyura: Majidae). Proceedings of the BiologicalSociety of Washington, 112(4): 754–758, Figs. 1, 2.
Chou W-R, Lai S-H & Fang L-S (1999) Benthic crustacean
communities in waters of southwestern Taiwan and theirrelationships to environmental characteristics. Acta Zoologica
Taiwanica, 10: 25–35.
Crosnier A (1967) Remarques sur quelques crustacés décapodes
benthiques Ouest-Africaines. Description de Heteropanope
acanthocarpus et Medaeus rectifrons spp. nov. Bulletin duMuséum national d’Histoire naturelle, sér. 2, 39(2): 320–344,
figs. 1–32.
Dai A-Y (1981) Two new species of Chinese Majidae (Crustacea:
Brachyura). Acta Zootaxonomica Sinica, 6(1): 36–38, figs. 1–10.
Dai A-Y & Yang S-L (1991) Crabs of the China Seas. China
Ocean Press, Beijing and Springer-Verlag, Berlin, 21 + 608
pp., figs. 1–295, pls. 1–74
Dai A-Y, Yang S-L, Song Y-Z & Chen G-X (1986) Crabs of the
China Seas. China Ocean Press, Beijing. 11 + 642 pp.
Doflein F (1902) Ostasiatische Dekapoden. Abhandlungen derBayerischen Akademie der Wissenschaften, München, 21(3):613–670, pls. 1–6.
Fabricius JC (1775) Systema Entomologiae, sistens InsectorumClasses, Ordines, Genera, Species, adjectis Synonymis, Locis,
Descriptionibus, Observationibus. Officina Libraria Kortii,
Flensburgi & Lipsiae. [1–30] + 1–832 pp.
Fabricius JC (1787) Mantissa insectorum sistens eorum species
nuper detectas adjectis characteribus genericis, differentiis
specificis, emendationibus, observationibus. Impensis Christ.
Gottl. Proft., Hafniae [= Copenhagen], 1: I–XX + 1–348 pp.
Fabricius JC (1798) Supplementum Entomologiae Systematicae.
Proft & Storch, Hafniae [= Copenhagen]. I–II + 1–572 pp.
Griffin DJG (1966a) A review of the Australian majid spider crabs(Crustacea, Brachyura). Australian Zoologist, 13(3): 259–298,
6 text-figs, pls. 15–17.
Griffin DJG (1966b) The marine fauna of New Zealand: spider
crabs, family Majidae (Crustacea, Brachyura). New ZealandDepartment of Scientific and Industrial Research Bulletin,
172(35): 1–111.
Griffin DJG (1976) Spider crabs of the family Majidae (Crustacea:
Brachyura) from the Philippine Islands. Journal of Natural
History, 10(2): 179–222, figs. 1–11, tables 1, 2.

221

Ng & Richer de Forges: Revision of the spider crab genus Maja
Griffin DJG & Tranter HA (1986) The Decapoda Brachyura of
the Siboga Expedition. Part VIII. Majidae. Siboga-Expeditie,
39C4: 1–335, 22 pls.
Griffith C & Pidgeon E (1833) The Classes Annelida, Crustacea, andArachnida, arranged by the Baron Cuvier, with supplementary
additions to each order by Edward Griffith and Edward Pidgeon.
In: The Animal Kingdom arranged in conformity with its
organization, by the Baron Cuvier, with supplementary additions
to each order, by Edward Griffith and others. London, Volume
13: i–viii, 1–540 (Crustacea: pls. 1–23).
Haan HM De (1833–1849) Crustacea. In: P. F. von Siebold,
Fauna Japonica, sive Descriptio animalium, quae in itinere per
Japoniam, jussu et auspiciis superiorum, qui summum in India
Batavia imperium tenent, suscepto, annis 1823–1830 collegit,
notis, observationibus a adumbrationibus illustravit. Lugduni
Batavorum, fasc. 1–8: I–xxi+vii–xvii+ix–xvi+1–243, pls. 1–55,
A–Q, circ., pl. 2. (For publication dates see Sherborn & Jentink,
1895; Holthuis, 1953; Holthuis & T. Sakai 1970).
Haswell WA (1880) On the Australian Brachyura Oxyrhyncha.
Proceedings of the Linnean Society of New South Wales, 4(4):431–458, pls. 25–27.
Herbst JFW (1782–1804) Versuch einer Naturgeschichte der
Krabben und Krebse nebst einer systematischen Beschreibung
ihrer verschieden Arten. Gottlieb August Lange, Berlin &Stralsund. Volumes 1–3: 1–274 + 1–226 +1–66 + 1–46, 1–54
+ 1–49, Pls. 1–62. [Vol. 1, Part 1 (Pp. 1–86, Pl. 1): 1782;
Vol. 1, Parts 2–5 (Pp. 87–182, Pls. 2–9): 1783; Vol. 1, Part 6(Pp. 183–206, Pls. 10–13): 1785; Vol. 1, Part 7 (Pp. 207–238,
Pls. 14–17): 1788; Vol. 1, Part 8 (Pp. 239–274, Pls. 18–21):
1790; Vol. 2, Part 1 (Pp. 1–48, Pls. 22–25): 1791; Vol. 2, Part
2 (Pp. 49–78, Pls. 26–29): 1792; Vol. 2, Part 3 (Pp. 79–98,
Pls. 30–33): 1793; Vol. 2, Part 4 (Pp. 99–146, Pls. 34–36):
1793; Vol. 2, Part 5 (Pp. 147–162, Pls. 37–40): 1794; Vol. 2,
Parts 6–end (Pp. 163–226, Pls. 41–46): 1796; Vol. 3, Part 1(1–66, Pls. 47–50): 1799; Vol. 3, Part 2 (Pp. 1–46, Pls. 51–54):
1801; Vol. 3, Part 3 (Pp. 1–54, Pls. 55–58): 1803; Vol. 3, Part
4 (Pp. 1–49, Pls. 59–62): 1804.] [For dates of publication, see
Rathbun, 1897: 157].
Herklots JA (1861) Catalogue de Crustacés qui ont servi de baseau système carcinologique de M. W. de Haan, rédigé d’après lacollection du Musée des Pays-Bas et les Crustacés de la faune
du Japon. Tijdschrift voor Entomologie, 4: 116–156. [Issued
as a separate: Herklots, J. A., 1861. Symbolae carcinologicae.
Études sur la classe des Crustacés. E. J. Brill, Leiden. 1–43 pp.]
Holm Å (1957) Specimina Linnaeana I Uppsala Bevarade
Zoologiska Samlingar Från Linnés Tid. Uppsala UniversitetsÅrsskrift, 6: 1–68.
Holthuis LB (1953) On the dates of publication of W. De Haan’s
volume on the Crustacea of P. F. von Siebold’s “Fauna
Japonica”. The Journal for the Society for the Bibliographyof Natural History, 3: 36–47, pl. 1.
Holthuis LB (1958) Proposed use of the Plenary Powers to validate
the generic name “Maja” Lamarck, 1801 (Class Crustacea,
Order Decapoda), and to designate a type species for thisgenus in harmony with current usage. Opinions rendered andDeclarations by The International Commission on Zoological
Nomenclature, 18(15): 257–272.
Holthuis LB & Sakai T (1975) Ph.F. von Siebold and Fauna
Japonica. A history of early Japanese Zoology. Academic
Press of Japan, Tokyo, i–xviii+part1: 1–132 (English), part 2:
207–323 (Japanese), pls. 1–32.
Huang L (1989) [Brachyura (Dromiacea, Gymnopleura, Oxystomata,Oxyrhyncha, Corystoidea)]. In: Dong Y (ed.) Fauna of Zhejiang.
Zhejiang Science and Technology, Zhejiang. Pp. 280–348.
Huang Z-G (ed.) (1994) Brachyura. In: Marine species and theirdistributions in China’s Seas. Department of Ocean Management

and Monitoring State Oceanic Administration. China Ocean
Press, Beijing. Pp. 576–600.
ICZN, International Commission on Zoological Nomenclature(1999) International Code of Zoological Nomenclature.
International Commission of Zoological Nomenclature.
Fourth Edition. Adopted by the XXI General Assembly of theInternational Union of Biological Sciences. International Trustfor Zoological Nomenclature, in association with the British
Museum (Natural History), London, 338 pp.
Ikeda H (1998) The Deep-Sea Crabs of Sagami Bay. Hayama
Shiosai Museum, Kanagawa, Japan, 180 pp.
Jacquinot H & Lucas H (1853) Crustacés. In: Voyage au Pôle
Sud et dans l’Océanie sur les corvettes l’Astrolabe et la Zélée;exécuté par ordre du roi pendant les années 1837-1838-18391840 sous le commandement de M. J. Dumont d’Urville. Gideet Baudry, Paris. Atlas, 1852, 9 pls. (en majeure partie par H.
Jacquinot) Zoologie, 3(3): 1–107 (par H. Lucas).
Jenkins RJF (1985) Fossil spider crabs from Australia. Special
Publication of the South Australian Department of Mines andEnergy, 5: 145–165.
Johnson DS (1958) The IndoWest Pacific species of the genus
Polyonyx (Crustacea, Decapoda, Porcellanidae). Annals of
Zoology, AGRA, 2(8): 95–118.
Kitchingman A & Lai S (2004) Inferences on potential seamountlocations from mid-resolution bathymetric data. In: Morato T &Pauly D (eds.) Seamounts: Biodiversity and Fisheries. Fisheries
Centre Research Reports. University of British Columbia,
Vancouver, Canada. Pp. 7–12.
Koh SK & Ng PKL (2008) A revision of the shore crabs of thegenus Eriphia (Crustacea: Brachyura: Eriphiidae). Raffles
Bulletin of Zoology, 56(2): 327–355.
Kubo I (1936) A new decapod crustacean from Japan, Paramaja
kominatoensis, gen. et sp. nov. Bulletin of the Japanese Societyof Scientific Fisheries, 4(6): 361–364.
Lamarck JBPA de (1801) Systême des animaux vertèbres, ou
Tableau général des classes, des ordres et des genres deces animaux; Présentant leurs caractères essentiels et leurdistribution, d’après la considération de leurs rapports naturels
et leur organisation, et suivant l’arrangement établis dans lesgaleries du Muséum d’Histoire Naturelle, parmi leur dépouilles
conservées; Précédé du discours d’ouverture du cours dezoologie, donné dans le Muséum National d’Histoire Naturellel’an 8 de la République. Déterville, Paris. Vol. 6: i–viii + 1–432.
Le Loeuff P & Intès A (1968) La faune benthique du plateau
continental de côte d’lvoire, récoltes au chalut, abondance–
repartition–variations saisonnières (Mars 1966–Février 1967).
Centre de Recherches Océanographiques, Abidjan, Document
Scientifique Provisoire, 25: 1–78, figs. 1–63.
Linnaeus C (1758) Systema naturae per regna tria naturae, secundumclasses, ordines, genera, species, cum characteribus, differentiis,
synonymis, locis. Tomus I. Editio Decima, Reformata. Laurentii
Salvii, Holmiae [= Stockholm], 823 pp.
Linnaeus C (1764) Museum S:ae R:ae M:tis Ludovicae Ulricae
Reginae Svecorum, Gothorum, Vandalorumque &c. In quo
animalia rariora, exotica, imprimis. Insecta & Conchilia
describuntur & determinantur, 720 pp.
Longhurst AR (1958) An ecological survey of the West African
marine benthos. Colonial Office, Fishery Publications, 11:
1–102, figs. 1–11.
Manning RB & Holthuis LB (1981) West African Brachyuran
Crabs (Crustacea: Decapoda). Smithsonian Contribution of
Zoology, 306: 1–379.
Marumura M & Kosaka A (2003) Catalogue of the Brachyuran
and Anomuran Crabs donated by the late Mr. Seiji Nagai to the
Wakayama Prefectural Museum of Natural History. Wakayama
Prefectural Museum of Natural History, 12–73 pp.

222

RAFFLES BULLETIN OF ZOOLOGY 2015
Masuda H, Hayashi K, Nakamura H & Kobayashi AM (1970) Les
Crustacés d’interêt commercial entre le Golfe de Cadix et leGolfe de Guinée. In: R. Letaconnoux & A. E. J. Went (eds.),
Symposium sur les resources vivantes du plateau continental
Atlantique africain du Détroit de Gibraltar au Cap Vert. Rapports
et Procès-Verbaux des Réunions, Conseil International pourl’Exploration de la Mer, 159: 126–127.
Matsuda H, Hayashi K, Nakamura H & Kobayashi AM (1986)
Marine Invertebrates. Tokai University Press, Tokyo, 256 pp.
Mat su z a wa K ( 1 9 7 7 ) S ea S h o re A n imals o f Muroto.
KansaiInsatsuKogyi, Japan, Un-numbered pages, 126 pls.
Maurin C (1968a) Écologie ichthyologique des fonds chalutables
atlantiques (de la Baie Ibero-Marocaine à la Mauritanie) et dela Mediterranée Occidentale. Revue des Travaux de l’InstitutPêches Maritimes, 32(1): 30–48.
Maurin C (1968b) Les crustacés captures par la “Thalassa” au large
des côtes nord-ouest africaines. Revue Roumaine de Biologie,
Série de Zoologie, 13(6): 479–493, figs. 1–9.
McLay CL, Feldmann RM & McKinnon DI (1995) New species ofMiocene spider crabs from New Zealand, and a partial cladistic
analysis of the genus Leptomithrax Miers, 1876 (Brachyura:
Majidae). New Zealand Journal of Geology and Geophysics,
38: 299–313.
Mendoza JCE, Naruse T, Tan S-H, Chan T-Y, Richer de Forges
B & Ng PKL (2010) Case studies on decapod crustaceansfrom the Philippines reveal deep, steep underwater slopes asprime habitats for ‘rare’ species. Biodiversity & Conservation,
19(2): 575–586.
Miers EJ (1876) Descriptions of some new species of Crustacea,
chiefly from New Zealand. Annals and Magazine of Natural
History, Series 4, 17(49): 218–229.
Miers EJ (1879) Descriptions of new or little-known species ofmaioid Crustacea (Oxyrhyncha) in the collection of the British
Museum. Annals and Magazine of Natural History, Series 5,4(19): 1–28, pls. 4, 5.
Miers EJ (1885) On the species of Micippa, Leach, and Paramicippa,
Milne Edwards. Annals and Magazine of Natural History,
Series 5, 15: 1–13, pl. 1.
Milne Edwards H (1834) Histoire naturelle des crustacés,comprenant l’anatomie, la physiologie et la classification de ces
animaux, Librairie Encyclopédique de Roret, Paris, 1: 1–468.
Milne Edwards H (1851) Observations sur le tégumentaire descrustacés décapodes, et sur la morphologie de ces animaux.
Annales des Sciences Naturelles, Series 3, Zoology, 16:
221–291, pls. 8–11.
Minemizu R (2000) Marine decapod and stomatopod crustaceans
mainly from Japan. Bun-Itsu Sougou Shuppan Ltd., Tokyo,
344 pp [In Japanese].
Miyake S (1936) On three crabs from off Tori-shima, Danjo-Group,Japan. Annotationes Zoologicae Japonenses, 15: 416–420, figs.
1, 2, pl. 28.
Miyake S (1983) Japanese Crustacean Decapods and Stomatopods
in color. Vol. II, Brachyura (Crabs). Hoikusha Publishing Co
Ltd., 277 pp.
Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires
de l’Institut Français d’Afrique Noire (IFAN), 45: 1–674, figs.
1–884, tables 1–10.
Moro L, Herrera R, Ortea J, Riera R, Bacallado JJ & Martín J(2014) Aportaciones al conocimiento y distribución de losDecápodos y Estomatópodos (Crustacea: Malacostraca) de las
Islas Canarias. Revista de la Academia Canaria de Ciencias,
26: 33–82.
Muraoka K (1998) Catalogue of the Brachyuran and Anomuran
Crabs donated by Prof. Dr. Tune Sakai to the Kanagawa
Prefectural Museum. Catalogue of the Collection in the
Kanagawa Prefectural Museum of Natural History, Odawara,
Kanagawa, Japan, No. 11: 1–67.

Muraoka K & Odawara T (1995) The Visual Guide to Crabs.
Narumi-Dou Shuppan Ltd., Tokyo, 159 pp. [in Japanese].
Nardo GD (1847) Sinonimia moderna delle specie registratenell’opera intitolata: Descrizione de’Crostacei, de’Testacei ede’Pesci che abitano le lagune e golfo Veneto, rappresentati infigure, a chiaro-scuro ed a colori dall’ Abate Stefano Chiereghini
Ven. Clodiense. Venezia [= Venice]. i–xi + 1–128 pp.
Neumann V (1993) The morphology of the first zoea stage of the
spider crab Maja crispata (Risso, 1827) (Crustacea: Decapoda:
Majidae). Senekenbergiana maritima, 23(4/6): 143–148.
Neumann V (1996a) Comparative investigations on the systematics
and taxonomy of European Maja species (Decapoda, Brachyura,
Majidae). Crustaceana, 69(7): 821–852.
Neumann V (1996b) Comparative gonopod morphology of theEuropean spider crabs of the genus Maja (Decapoda: Brachyura:
Majidae). Senckenbergiana Biologica, 75(1/2): 143–157.
Neumann V (1998) A review of the Maja squinado (Crustacea:
Decapoda: Brachyura) species-complex with a key to the eastern
Atlantic and Mediterranean species of the genus. Journal of
Natural History, 32: 1667–1684.
Nishimura S & Suzuki K (1971) Common seashore animals ofJapan in color. Hoikusha, Osaka, pp. 1–196, pls. 1–64.
Ng PKL (1994) First record of the synaxid lobster, Palinurellus
wieneckii (de Man, 1881) (Crustacea, Decapoda, Palinuridea)from Sri Lanka. Journal of South Asian Natural History,
Colombo, 1(1): 117–118.
Ng PKL, Guinot D & Davie PJF (2008) Systema Brachyurorum:
Part I. An annotated checklist of extant brachyuran crabs of
the world. Raffles Bulletin of Zoology, Supplement 17: 1–286.
Ng PKL, Mendoza JCE & Manuel-Santos M (2009) Tangle net
fishing, an indigenous method used in Balicasag Island, central
Philippines. Raffles Bulletin of Zoology, Supplement 20: 39–46.
Ng PKL, Wang C-H, Ho P-H & Shih H-T (2001) An annotatedchecklist of brachyuran crabs from Taiwan (Crustacea:
Decapoda). National Taiwan Museum. Special Publication
Series, Number 11: 1–86.
Ninni E (1924) Considerazioni sul genre Maja (Lam.) e sopra una
forma di Maja nuova per l’Adriatico. Bollettino bimestrale delR. Comitato Talassografico Italiano, 13: 39–50, 9 text-figs.
O’Hara T & Tittensor DP (2010) Environmental drivers of ophiuroid
species richness on seamounts. Marine Ecology, 31: 26–38.
D’Oliveira MP (1889) Nouveau Oxyrhynque du Portugal. O
Instituto, Coimbra, Series 2, Sciencias Physico-Mathematicas,
36: 78–79.
Ortmann AE (1893) Abtheilung: Brachyura (Brachyura genuinaBoas), I. Unterabtheilung: Majoidea und Cancroidea, 1: Section
Portuninea. Die Decapoden-Krebse des Strassburger Museums,
mit besonderer Berücksichtigung der von Herrn Dr. Döderlein
bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit
im Strassburger Museum aufbewahrten Formen. Theil VI [=Part 6]. Zoologische Jahrbücher, Abtheilung für Systematik,
Geographie, und Biologie der Thiere, 7(I): 23–88, pl. 3.
Ortmann AE (1894) Crustaceen. In: Semon R (ed.), Zoologische
Forschungreisen in Australien und dem Malayischen Archipel
in den Jahren 1891–93. V. Denkschriften der medizinischNaturwissenschaftlichen Geselschaft, Jena, 8: 1–80, pls. 1–3.
Parisi B (1915) I decapodi giapponesi del Museo di Milano. III.
Oxyrhyncha. Atti della Società Italiana di Scienze Naturali edel Museo Civico di Storia Naturale, Milano, 54(2–4): 281–296,
figs. 1–4, pl. 7.
Poore GCB, McCallum AW & Taylor J (2008) Decapod Crustaceaof the continental margin of southwestern and central Western
Australia: preliminary identifications of 524 species fromFRV Southern Surveyor voyage SS10-2005. Museum Victoria
Science Reports, 11: 1–106.

223

Ng & Richer de Forges: Revision of the spider crab genus Maja
Rahayu DL & Ng PKL (2000) Some new and interesting records of
spider crabs (Decapoda, Brachyura, Majidae) from Indonesia.
Crustaceana, 73(7): 887–890.
Rathbun MJ (1897) A revision of the nomenclature of the Brachyura.
Proceedings of the Biological Society of Washington, 11:
153–167.
Rathbun MJ (1905) Why not Paramaya? Proceedings of the
Biological Society of Washington, 18: 73.
Rathbun MJ (1916) New species of crabs of the families Inachidae
and Parthenopidae. Scientific results of the Philippine Cruiseof the Fisheries Steamer “Albatross,” 1907–1910.—No. 34.
Proceedings of the United States National Museum, 50(2135):
527–559.
Rathbun MJ (1918) Report on the spider crabs obtained by the F.I.S.
“Endeavour” on the coasts of Queensland, New South Wales,
Victoria, South Australia and Tasmania. Biological Results of
the Fishing Experiments Carried On by the F.I.S. “Endeavour,”
1909–14, 5(1): 3–29, figs. 1–3, pls. 1–15.
Rathbun MJ (1924) New species and subspecies of spider crabs.
Proceedings of the United States National Museum, 64: 1–5.
Rathbun MJ (1932) Preliminary descriptions of new species ofJapanese crabs. Proceedings of the Biological Society ofWashinginton, 45: 29–38.
Richer de Forges B (1993) Deep sea crabs of the Tasman seamounts(Crustacea: Decapoda: Brachyura). Records of the AustralianMuseum, 45: 11–24, figs. 1–11.
Richer de Forges B & Ng PKL (2015) The identity of Leptomithrax
sinensis Rathbun, 1916, and the description of of L. eldredgei,
sp. nov. from Hong Kong (Crustacea: Decapoda: Brachyura:
Majidae). Bishop Museum Bulletin of Zoology, Lucius G.
Eldredge Memorial Volume, Evenhuis NL & Carlton JT (eds.)
Honolulu, 9: 123–128.
Richer de Forges R, Tan SH, Bouchet P, Ng PKL, Chan T-Y &Saguil N (2009) PANGLAO 2005 — Survey of the deep-water
benthic fauna of Bohol Sea and adjacent waters. Raffles Bulletinof Zoology, Supplement 20: 21–38.
Risso A (1827) Histoire naturelle des principales productions del’Europe Méridionale et particulièrement de celles des environs
de Nice et des Alpes Maritimes. F.-G. Levrault, Paris. Volume
5 [= Animaux Articulés, Annelides, Crustacés, Arachnides,
Myriapodes et Insectes]: i–vii + 1–403, figs. 1–62, pls. 1–10.
Rossignol M (1957) Crustacés Décapodes marins de la région
de Pointe-Noire. In: Collignon J, Rossignol M & Roux Ch,
Mollusques, Crustacés, Poissons Marins des côtes d’A.E.F.
en collection au Centre d’Océanographie de l’Institut d’Études
Centrafricaines de Pointe-Noire, Paris. Pp. 71–136, figs. 1–20,
pls. 1–3.
Sakai K (1999) J.F.W. Herbst-collection of decapod Crustacea of
the Berlin Zoological Museum, with remarks on certain species.
Naturalists, Publications of Tokushima Biological Laboratory,
Shikoku University, 4: 1–45, pls. 1–21.
Sakai T (1932) Notes on some rare materials of Japanese
Oxyrhyncha. Science Reports of the Tokyo Bunrika Daigaku,
Section B, 1(4): 41–59, figs. 1–8, pls. 2, 3.
Sakai T (1934) Brachyura from the coast of Kyusyu, Japan. Science
Reports of the Tokyo Bunrika Daigaku, Section B, 1(25):281–330, figs. 1–26, pls. 17, 18.
Sakai T (1936) Crabs of Japan. 66 plates in life colours withdescriptions. Sanseido, Tokyo, pp. 1–239, 1–12 [literaturecited], 1–27 [bibliography and index], pls. 1–66. [dated 1935
but published in 1936]
Sakai T (1938) Studies on the Crabs of Japan. III. Brachygnatha,
Oxyrhyncha. Yokendo, Tokyo, pp. 193–364, figs. 1–55, pls.
20–41, table I.
Sakai T (1965) The Crabs of the Sagami Bay collected by His
Majesty the Emperor of Japan. Maruzen Co., Tokyo. i–xvi +1–206 pp. [English section], figs. 1–27, pls. 1–100, 1–92 [inJapanese] + 1–26 [Bibliography and Index in English] + 27–32[Index in Japanese] pp., 1 map.
Sakai T (1969) Two new genera and twenty-two new species ofcrabs from Japan. Proceedings of the Biological Society ofWashington, 82: 243–280, figs. 1–19, pls. 1, 2.
Sakai T (1976) Crabs of Japan and the Adjacent Seas. Kodansha
Ltd., Tokyo. In three volumes; English Text, pp. xxix+773 pp.,
Japanese Text, pp. 1–461, Plates volume, pp. 1–16, pls. 1–251.
Samouelle G (1819) The Entomologist’s Useful Compendium,or an Introduction to the Knowledge of the British Insects.
Thomas Boys, London. 496 pp.
Serène R (1968) The Brachyura of the Indo Pacific Region. In:
Prodromus for a Check List of the Non-planctonic Marine
Fauna of South East Asia. Special Publication of the Singapore
National Academy of Science, No. 1: 33–120.
Serène R & Lohavanijaya P (1973) The Brachyura (Crustacea:
Decapoda) collected by the Naga Expedition, including a reviewof the Homolidae. In: Brinton E, Newman W & Wooster WS(eds.) Scientific Results of Marine Investigations of the South
China Sea and the Gulf of Thailand, 1959–1961. Naga Report,
4(4): 1–187.
Serène R & Vadon C (1981) Crustacés décapodes: brachyoures.
Liste préliminaire, description de formes nouvelles et remarques
taxonomiques. In: Résultats des Campagnes MUSORSTOM,
1. Philippines (18–29 mars 1976). Mémoires ORSTOM, Paris,
91: 117–140.
Sherborn CD & Jentink FA (1895) On the dates of the partsof Siebold’s ‘Fauna Japonica’ and Giebel’s ‘Allegemeine
Zoologie’ (first edition). Proceedings of the Zoological Societyof London, 1895: 149–150.
Sotelo G, Morán P & Posada D (2007) Identification andcharacterization of microsatellite loci in the spiny spider crab
Maja brachydactyla. Conservation Genetics, 8: 245–247.
Sotelo G, Morán P & Posada D (2008) Genetic identification of thenortheastern Atlantic spiny spider crab as Maja brachydactyla
Balss, 1922. Journal of Crustacean Biology, 28(1): 76–81.
Sotelo G, Morán P & Posada D (2009) Molecular phylogeny
and biogeographic history of the European Maja spider crabs(Decapoda, Majidae). Molecular Phylogenetics and Evolution,
53: 314–319.
Stebbing TRR (1905) South African Crustacea. Part III. Marine
Investigations in South Africa, Cape of Good Hope, Departmentof Agriculture, Cape Town, 4: 21–123, pls. [Crustacea] 17–21,
21 bis, 22–26.
Stebbing TRR (1908) South African Crustacea (Part IV.). Annalsof the South African Museum, 6(1): 1–96, pls. 1–14.
Števčić Z (1990) Check-list of the Adriatic decapod Crustacea.
Acta Adriatica, 31(112): 183–274.
Takeda M (1973) Report on the crabs from the sea around theTsushima Islands collected by the research vessel “Genkai” forthe trustees of the National Science Museum, Tokyo. Bulletinof the Liberal Arts & Science Course, Nihon University Schoolof Medicine, 1: 17–68.
Takeda M (1975) Brachyura. In: Utinomi H (ed.) Aquatic
Invertebrates. Gakken Chukosei zukan, 9: 120–149 (fig. with
Latin names), 188–331 (descriptions). First Edition. Gakken,
Tokyo. [Second Edition in 1983 as The Aquatic Lower Animalsof Japan without changes].
Takeda M (1980) A small collection of crabs from the KyushuPalau
Submarine Ridge, with description of a new species of the
Homolidae. Micronesica, 16(2): 279–287.
Takeda M (1982) Keys to the Japanese and foreign crustaceans
fully illustrated in colors, Hokuryukan, Tokyo. Pp. vi+285,
1–58 (keys).
Takeda M (1993) Catalogue of crab specimens (Crustacea:
Decapoda: Brachyura) (Part 1). National Science Museum,
Tokyo, 81 pp.

224

RAFFLES BULLETIN OF ZOOLOGY 2015
Takeda M (2001) Annotated list of crabs from Tosa Bay, Southwest
Japan, collected by the R/V Kotaka Maru during the years
1997–2000. In: Fujita T, Saito H & Takeda M (eds) Deep-sea
fauna and pollutants in Tosa Bay. National Science Museum
Monographs, 20: 217–262.
Takeda M & Miyake S (1969) Crabs from the East China Sea.
III. Brachygnatha Oxyrhyncha. Journal of the Faculty ofAgriculture, Kyushu University, 15(4): 469–521, figs. 1–12,
pls. 17, 18.
Takeda M, Sakai S, Shinomiya S & Suga H (2000) Crabs fromthe Seto Inland Sea, Southern Japan. Memoirs of the National
Science Museum, Tokyo, 33: 135–144.
Takeda M & Ueshima R (2006) Crabs preserved in the Department
of Zoology, the University Museum, the University ofTokyo. Catalogue of the Invertebrate Collection depositedin the Department of Zoology, The University Museum, The
University of Tokyo. Ueshima R (ed.), The University Museum,The University of Tokyo, Material Reports No. 62: 61–105.
Thomson GM (1902) On a new species of Paramithrax from New
Zealand. Annals of the Magazine of Natural History, Series 7,10(61): 361–364, pls. 7, 8.
Tittensor DP, Mora C, Jetz W, Lotze HK, Ricard D, Vanden Berghe
E & Worm E (2010) Global patterns and predictors of marine
biodiversity across taxa. Nature, 466: 1098–1011.
d’Udekem d’Acoz C (1999) Inventaire des crustacés décapodesde l'Atlantique nord-oriental, de la Méditerranée et des eauxcontinentales adjacentes au nord de 25°N. Muséum Nationald’Histoire Naturelle/Service du Patrimoine Naturel, Collection
Patrimoines Naturels, Paris, 40, 383 pp.
Utinomi H (1969) Coloured illustrations of sea shore animals ofJapan. First Edition [1956], pp. xvii+168, pls. 1–64; Second
Edition [1969], pp. xix+166, pls. 1–64. Hoikusha, Osaka.

Walker AO (1887) Notes on a collection of Crustacea fromSingapore. Journal of the Linnean Society of London, 20(118):
107–117, pls. 6–9.
Ward M (1933) New genera and species of marine Decapoda
Brachyura from the coasts of New South Wales and Queensland.
Australian Zoologist, 7(5): 377–394, pls. 21–23.
White A (1847) List of specimens of Crustacea in the collectionof the British Museum. British Museum, London, viii+143 pp.
Whitelegge T (1900) Crustacea I. In: Etheridge R (ed.), Scientificresults of the trawling expedition of H.M.C.S. “Thetis”, off
the coast of New South Wales, in February and March, 1898.
Memoirs — Australian Museum, Sydney, 4(2): 133–199, pls.
32–35.
Yamaguchi T & Baba K (1993) Crustacean specimens collected
in Japan by Ph. F. Von Siebold and H. Bürger and held by
the Nationaal Natuurhistorisch Museum in Leiden and otherMuseums. In: Yamaguchi T (ed.) Ph.F. von Siebold and Natural
History of Japan Crustacea. The Carcinological Society ofJapan. pp. 145–570.
Yang CM (1979) A list of Brachyura in the Zoological ReferenceCollection of the Department of Zoology. Department of
Zoology, University of Singapore, 60 pp. (mimeographed).
Yang S, Chen H & Jiang W (2008) Crustacea: Decapoda Brachyura.
In: Liu R (ed.), Checklist of Marine Biota of China Seas. Instituteof Oceanology, Chinese Academy of Sciences. Pp. 761–810.
Yokoya Y (1933) On the distribution of decapod crustaceans
inhabiting the continental shelf around Japan, chiefly based
upon the materials collected by the S.S. Sôyô-Maru, during theyear 1923–1930. Journal of the College of Agriculture, Imperial
University of Tokyo, 12(1): 1–226, figs. 1–71, tabs. 1–4.
Zariquiey Álvarez R(1968) Crustáceos decápodos Ibéricos.
Investigación Pesquera, Barcelona, 32: i–xv + 1–510 pp.,figs. 1–164.

225