259 

A REVIEW OF THE AUSTRALIAN MAJID SPIDER CRABS 
(CRUSTACEA, BRACHYURA) 

By D. J. G. Griffin 

(Zooloey Department, University of Tasmania, Hobart) 
(Plates XV-XVII & Text-figures 1-3) 

SUMMARY 

An historical account is given of taxonomic studies of spider or masking 
crabs in Australia and overseas. The basis of the review is a key to all but 
two of the species known to occur in Australia. The key includes information 
on synonymies, geographic and bathymetric distribution and references to 
descriptions and illustrations of the species. Characters which are important in 
the classification are briefly reviewed. Zoogeographical relationships of the 
fauna are discussed. 

The family Majidae is currently divided into seven subfamilies mainly 
on the basis of orbital configuration, form of the rostrum, abdomen and first 
pleopod of the male. All but one of the subfamilies are represented in the 
Australian fauna which is considered to comprise 95 species in 45 genera. 
This is about twice the number of species and genera listed by Haswell (1882c) 
in his "Catalogue." About one-third of the species have been recorded from 
Australia on only a single occasion. Numerous genera and species are in need 
of detailed reinvestigation. 

The fauna is rather clearly partitioned into a tropical group with widespread 
Indo-West Pacific relationships and a temperate group related to tropical 
Australia and/or the Indo-West Pacific rather than to temperate regions outside 
Australia. There are no clear boundaries between these two faunas but rather 
quite broad transition areas. Thirty-seven species and five genera, most of 
which are temperate, are restricted to Australia. 

1. INTRODUCTION 

One of the most characteristic features of spider crabs of the family Majidae 
is the presence on the carapace and legs of special curled or "hooked" hairs 
which aid in the attachment of various kinds of epifauna and flora, especially 
seaweeds (for example see McNeill 1923). These organisms are placed in 
position by the crab with the aid of the chelipeds which are able to reach up on 
to the dorsal surface of the carapace. For this reason majid crabs are sometimes 
called "masking crabs" or "seaweed crabs." Of special interest in these crabs 
are the orbits which may be expanded in various ways and surrounded by 
a seemingly complex array of spines. The legs are often long and slender and 
the carapace usually triangular or pyriform. Spider crabs range in size from a 
few millimetres to more than a metre in carapace length and are found in 
almost all seas and oceans. They form a relatively important part of the benthos 
and may be locally very abundant on the shelf although some species extend 
considerable distances down the continental slope to depths as great as 1000 
fathoms. 

Substantial revisions of the family Majidae have been undertaken on a 
world-wide scale by Dana (1851), Miers (1879c), Alcock (1895) and Balss 
(1929). Sakai (1938), Stephensen (1945) and Garth (1958) have adapted 
Balss's scheme to the faunas of Japan, the Iranian Gulf and America respectively. 
In the course of these revisions the Majidae have been arranged in several very 
different fashions (see historical reviews by Miers 1879c, and Garth 1958). 
Early workers such as Dana and Miers grouped the species in several families 
and a large number of subfamilies 260 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

Many morphological characters have been used in these different classi- 
fications and some have been shown to be unreliable guides to phylogenetic 
relationships (see Section III). Characters concerned with the form of the orbit 
still dominate the classification but Garth has found, in the Pacific American 
fauna, groups of genera showing vast agreements in the form of the male first 
pleopod. This appendage is now realised to be of great value in the taxonomy 
of nearly all families of crabs. 

The first Australian spider crabs to be described were made known by the 
European naturalists Herbst, Latreille and H. Milne Edwards in the early 
nineteenth century. In the second half of that century the work of Dana, 
Stimpson, Miers and Haswell resulted in substantial additions to knowledge 
of the Australian spider crab fauna. Dana in 1852 dealt with the Crustacea 
of the U.S. Exploring Expedition of 1838-1842, Stimpson with the Crustacea 
of the North Pacific Exploring Expedition led by Ringgold and Rodgers 
(Stimpson 1857 GRIFFIN 261 

given by Rathbun (1925), Garth (1958) and Griffin (in press, b). Terminology, 
unless otherwise indicated, follows that used by Rathbun and Garth and by 
Griffin in previous papers. Although most information used in the key is taken 
from the literature, a considerable amount of material comprising most of the 
species represented in Australia, has been examined at the Australian Museum 
over the past three years. 

II. THE FAMILY 

The Majidae belong, with the Hymenosomidae (fiat-back crabs) and the 
Parthenopidae (caltrop crabs), to the superfamily Oxyrhyncha. All three 
families possess an anteriorly narrowed and produced carapace with prominently 
expanded branchial regions, large epistome, quadrate buccal cavity and 
longitudinally folded antennules. The characters which chiefly distinguish majid 
crabs from other families of the Brachyura are: (1) the possession of specially 
mobile chelipeds; (2) fusion of the well-developed second article of the antenna 
(usually called the basal antennal article) to the epistome and often also to 
the front; (3) the comparatively incomplete orbits; (4) articulation of the palp 
of the external maxillipeds at the anteromedial angle of the merus; and 
(5) the much greater length of the first pleopods of the male relative to the 
second pair. The family has usually been considered in the past to share with 
other Oxyrhyncha nine pairs of gills on each side but Hartnoll (1964) has 
recently shown that this number is sometimes reduced in the Majidae to seven 
or eight. 

It should be mentioned that the term Maioidea used by Dana and Miers is 
partially synonymous with Oxyrhyncha (as now understood) and not with 
the term Majidae (until recently frequently spelt Maiidae). In Dana's and 
Miers's time the Maioidea included only the Majidae and the Parthenopidae; 
de Haan's (1839) addition of the Hymenosomidae did not gain general 
acceptance until after Rathbun (1925). 

As to the evolution and relationships of the Oxyrhyncha, Glaessner (1960: 
46) states, "The Oxyrhyncha, or spider crabs, are . . . unrepresented before 
the Tertiary except by unidentifiable fragments. The pointed rostrum . . . and 
the prominent mesogastral-cardiac ridge, together with the elongate cephalothorax, 
place this group much closer to the Oxystomata than to the Brachyrhyncha, 
but it is more advanced in the organisation of its mouthparts." 

III. THE SUBFAMILIES 

The subfamilial arrangement adopted here is that proposed by Garth (1958) 
which is a modification of the schemes of Alcock (1895) and Balss (1929). In 
the delimitation of subfamilies and in the arrangement of genera within them, 
particular attention has been paid to the structure of the orbit, i.e., the degree 
of expansion of the anterolateral part of the carapace above the origin of the 
eyestalk (the "supraorbital eave"), the presence or absence, and form of, 
a spine behind the eave (the "postorbital spine") and the degree of expansion 
of the basal antennal article. These are characters used before Alcock by Miers 
(1879c) and to some extent by Dana (1851). H. Milne Edwards (1834) 
considered the length of the ambulatory legs to be important and the classification 
proposed by de Haan (1839) was based on the shape of the merus of the 
third maxillipeds. Use of these latter characters was criticised by Miers. In 
1861 Claus proposed a classification based on the form of the basal antennal 
article. Use of this character was criticised by Stimpson in 1871. Balss laid 
particular stress on a further feature associated with the structure of the 
orbit: the presence or absence of a spine (the "intercalated spine") between the 
eave and the postorbital lobe. However, later workers on the group 
(Sakai 1938 and Garth 1958) have considered that Balss laid too much stress 
on this feature (see below). The relative size of this spine is sometimes 
variable with either growth or geographical locality (Garth 1958; Griffin, 
unpublished). Balss also introduced three other characters: the number of 
free segments of the abdomen (basically seven but sometimes reduced by 

262 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

coalescence or fusion involving segments four to six inclusive), the degree 
of development of the interantennular spine (the true rostrum of the zoea) 
and the degree of fusion of the (pseudo) rostral spines. 

In the key presented here genera within each subfamily are arranged 
in a series which begins with forms with a double rostrum and seven free 
abdominal segments in both sexes and ends with those in which the rostral 
spines are partly or wholly fused and the number of free segments in the 
abdomen is reduced. In this respect the key resembles those provided by 
Garth (1958). Implicit in such a presentation is the concept that parallel 
evolution has proceeded independently within each subfamily. The genera at the 
beginning of the series are considered to be primitive and those at the end 
advanced. This is in accordance with the ideas of Balss who considered 
that a well developed interantennular spine and the presence of an intercalated 
spine were also primitive features. 

Some aspects of orbital structure require further clarification. The term 
"commencing orbit," used in respect of some of the Ophthalmiinae, Acanthony- 
chinae and Pisinae, is intended to contrast with the unformed orbits of the 
Inachinae and some members of the Acanthonychinae on the one hand and 
with the complete or almost complete orbits of the Majinae and Mithracinae 
on the other. The commencing orbit is thus intermediate between two extremes. 
It involves partial enlargement of the supraorbital eave, most commonly by 
the development of a prominent spine either anteriorly or, less often, posteriorly; 
this spine scarely conceals the eyestalk from dorsal view. 

In some inachines there is above the origin of the eyestalk a variously 
developed spine, considered by Balss to represent the intercalated spine. 
However, in the Inachinae, this spine is very seldom separated by distinct 
fissures from the surrounding parts of the orbit as it is in the Majinae and 
some Mithracinae. Further, in some inachine genera which appear to be 
natural groups, some species possess a spine above the eyestalk and some do 
not. It would seem better to call this spine merely a supraorbital spine and 
to disregard, at least for the present, any possible homology between it in 
the Inachinae and the intercalated opine of other majids. 

A similar difficulty exists in many of the Pisinae. In some the postorbital 
lobe is provided on the upper anterior edge close to the base with a prominent 
accessory lobe and in a few species this lobe is quite distinct from the 
postorbital lobe. In other pisines where the eave is virtually unexpanded a small 
denticle is present on the supraorbital margin. There this lobe or denticle 
appears to be the intercalated spine and is generally so treated here. 

To sum up, emphasis on the intercalated spine led Balss to divide the 
Inachinae and the Pisinae each into two further subfamilies. In at least the 
Inachinae it is difficult to work out the homologies of a spine above the 
orbit in those species in which it is present and in the Pisinae Garth has 
found that such a division, in the Pacific American forms, is not supported 
by the male first pleopods. The term supraorbital spine is used, especially 
in the Inachinae, to denote a spine above the orbit where homologies are 
not clear. In other groups the term preorbital spine is restricted to mean an 
anterior outgrowth of the supraorbital eave and the term antorbital spine is used 
to denote a posterior outgrowth (see figure in Griffin in press, a). 

Figure 1. Generalised morphology of the subfamilies Inachinae, Ophthal- 
miinae and Acanthonychinae. Front of the carapace from above 
at left and from below at right; whole carapace from above in 
the middle. Abbreviations: a,, antennule; a 2 , antenna; a.o., 
antorbital spine; b.a., basal antennal article; e, eye; i, intercalated 
spine; m, mouthfield; p.o., postorbital lobe; pr, preorbital spine: 
r, rostrum; s.o., supraorbital eave. 

GRIFFIN 

NACHINAE 

OPHTHALMIINAE 

ACANTHONYCHINAE 

264 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

The characters of the six subfamilies represented in Australia are now 
given (the number of the couplet in the key at which genera of each subfamily 
begin is indicated in brackets after the name): 

1NACHINAE (4): Orbits undeveloped, or eave weakly expanded only; a 
supraorbital spine sometimes present. A postorbital spine sometimes present but 
affording no concealment to the cornea of the retracted eyestalk. Basal antennal 
article extremely slender and usually long. Eyestalk very long and visible 
almost to its base in both dorsal and ventral view. Carapace subtriangular or 
sometimes subpyriform or occasionally circular. Rostrum sometimes of a 
single spine, and frequently short. Ambulatory legs frequently extremely 
long and slender. 

OPHTHALMIINAE (26): Orbits consisting of a well developed and laterally 
expanded eave or of a greatly elongated spine. Postorbital spine short. An 
intercalated spine never present. Basal antennal article moderately expanded only. 
Eyestalk very long, often concealed in dorsal view but always largely visible 
in ventral view. Carapace elongate and often truncate in front and provided 
with a medial expansion or spine posteriorly. Rostral spines distinct and usually 
short. Ambulatory legs seldom very long. 

ACANTHONYCHINAE (29): Orbits undeveloped, eave rounded or forwardly 
produced as a preorbital spine. Postorbital spine, if present, simple and 
affording no concealment to eyestalk. Basal antennal article not very wide and 
characteristically truncate triangular. Eyestalks extremely short and often sunk 
in sides of rostrum. Carapace basically pyriform but characteristically twice 
expanded laterally at hepatic and branchial margins. Rostrum consisting either 
of two distinct spines or sometimes of a huge "beak." Ambulatory legs short 
or at most of moderate length. 

PISINAE (36): Orbits with a weekly expanded eave either produced anteriorly 
as a preorbital spine or acute or sometimes posteriorly acute, or else weakly 
expanded midway along and almost confluent with the postorbital lobe. Inter- 
calated spine present or absent, sometimes a small denticle or lobe close to 
the postorbital lobe. Postorbital lobe always well developed and cupped but 
not completely concealing the eyestalk from dorsal view. Basal antennal article 
slightly to broadly expanded and usually produced into a spine anterolateral^. 
Eyestalk typically short and bulbous with a large cornea. Carapace always 
pyriform although sometimes very wide. Rostrum frequently bifid for distal half 
only. Ambulatory legs often very long and slender. 

MAJINAE (62): Orbit well developed, comprising above a laterally expanded 
eave which is always acute posteriorly and sometimes produced into a spine, 
an intercalated spine and a postorbital spine which is sometimes simple and 
sometimes cupped and occasionally armed with an accessory spine on the upper 
anterior edge near the base. Basal antennal article moderately broad and 
rectangular and frequently armed with spines at both anteromedial and 
anterolateral angles. Eyestalks of moderate length and generally slender: only 
the distal half is visible in dorsal view. Carapace pyriform and sometimes 
broad. Rostrum always of two distinct spines. Ambulatory legs generally of 
moderate length. 

MITHRACINAE (81): Orbits extremely well developed with the eave and 
basal antennal article laterally expanded and usually tubular, completed behind 
by the prominent postorbital lobe which is often cupped. An intercalated 
spine is sometimes present. Eave sometimes forwardly produced into a preorbital 
spine or lobe, and sometimes prolonged posteriorly into an antorbital spine. 
Eyestalk moderately long but only the distal portion generally visible in either 
dorsal or ventral view. Carapace basically pyriform but anteriorly broadened 
due to expansion of orbits. Rostrum of two distinct spines which are often 
contiguous and sometimes fused into a broad lamella. Ambulatory legs seldom 
of great length. 

Figure 2. Generalised morphology of the subfamilies Pisinae, Majinae and 
Mithracinae. Arrangement and abbreviations as in text-fig. 1. 

GRIFFIN 

265 

P I S I N A E 

M A J I N A E 

MITHRAC IN AE 

266 REVIEW OF AUSTRALIAN MAUD SPIDER CRABS 

The seventh subfamily, the OREGONIINAE contains three genera which 
are boreal in distribution and not represented in Australia. They resemble 
the Inachinae in most features but are characterised by the terminally broadened 
male abdomen, the seventh segment being quadrate and deeply inserted into 
the sixth segment, and the longitudinally grooved male first pleopod, the groove 
being margined by rows of filamentous setae. A detailed treatment is given 
by Garth (1958). 

IV. THE GENERA 

The characters which are of value at the generic level are essentially 
the same as those used at the subfamilial level and already discussed. It need 
only be emphasised that the application of these characters at the generic level 
is invariably more rigorous than at the higher levels. Thus congeneric species 
are usually considered to share very similar orbital structure, form of the rostrum 
and of the male first pleopods and to agree in the number of free segments 
in the abdomen. They also generally resemble each other closely in carapace 
shape and ornamentation, form of the third maxillipeds, shape of the male 
abdomen (a character introduced by Shen in 1932) and often shape of the 
male chela but seldom in actual size and in relative length of the ambulatory 
legs. 

The remaining part of this section and most of the succeeding one will be 
devoted to a discussion of the several lower taxa of the Australian Majidae 
which are considered to pose important problems at the present time. 

Of the 24 majid genera listed by Haswell (1882c), ten now either have 
quite different meanings or are known under different names. These are as 
follows : 

1. Stenorhynchus Lamarck. Of the three species included here by Haswell, two 
have been shifted to Achaeus and one, S. curvirostris, has not been collected 
since its discovery (see next section). Lamarck's genus is currently recognised 
for two species confined to the Americas and north-west Africa (Garth 1958, 
Garth & Holthuis 1963); since Haswell mentions the rostrum being formed of two 
spines, not one, it is reasonable to assume that he intended to refer to 
Stenorhynchus Latreille (now consideied a synonym of Macropodia Leach, 1814). 

2. Halimus Latreille. The species included here by Haswell are now known 
as species of Naxia Latreille, Halimus being a junior synonym of that genus. 
The composition of the abdomen (of both sexes) and several other 
characters in these species require investigation to determine their relationships 
to the New Zealand representative of the genus (Griffin, in press, b). The 
comparatively well developed orbits and distally expanded ambulatory propodi 
set this genus apart from other inachines. Haswell's subgenus Microhalimus, 
with its single species, M. deflexifrons, was included as a subgenus of Naxia 
by McCulloch (1908). For the sake of convenience, Microhalimus is here 
not given subgeneric status though it is true that M. deflexifrons appears to 
differ in some important features of the orbit from other species of Naxia, 
particularly in the number and arrangement of the spines above and behind 
the orbit. Such disagreement is especially obvious when it is compared with 
those species with which it agrees in the slight expansion of the ambulatory 
propodi. 

3. Naxia H. Milne Edwards. Haswell listed one species under this genus. 
Pisa serpulifer Guerin. This was transferred to Naxioides A. Milne Edwards 
(of which Naxia Miers is a synonym) by Rathbun (1914) and later (Rathbun 
1924) set apart in a genus of its own, Paranaxia; H. Milne Edwards's genus 
is a synonym of the latter. 

4-6. Gonatorhynchus Haswell, Paramithrax H. Milne Edwards and Chlorinoides 
Haswell. Gonatorhynchus has recently (Griffin 1963b) been sunk in Paramithrax 
which has been recognised as monotypic. There appear to be some important 
similarities in orbital structure between P. harhicornis and Anacinetops stimpsoni 
although the species are at present placed in different subfamilies. The meaning 
attributed Paramithrax in Haswell's time has also been changed through the 
removal of several species to Chlorinoides (see Griffin in Dress, a). These latter 
species are the ones often placed in Acanthophrys A. Milne Edwards, a genus 
synonymous with Hyastenus White. 

GRIFFIN 267 

7-9. Egeria Latreille, Cyclomaia Stimpson and Parathoe Miers. These have been 
replaced respectively by Phalangipus Latreille, Cyclax H. Milne Edwards and 
Perinia Dana without change in meaning. In addition Eucinetops Stimpson no 
longer appears in the Australian lists, Anacinetops Miers having been accepted 
for E. stimpsoni Miers (see Balss 1935). One genus listed by Haswell among 
the Oxyrhyncha, the monotypic Plcuruphricus A. Milne Edwards, has recently 
been included among the Cymopolidae (now known as the Palicidae 268 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

Antilibinia and Taliepus have in the past been considered subgenera of 
Epialtus H. Milne Edwards (see Garth 1958:207). A new genus may be 
required for the Australian and Philippine species. 

5. Hyastenus White. As the basis for Balss's (1929) Hyasteniinae, this extremely 
large genus (Balss in 1935 listed 38 species) is usually considered to be 
characterised especially by an absence of an intercalated spine from the orbit 
and by the distinctness of all seven abdominal segments. However, Laurie 
(1906) has noted that in his H. irami there is a distinct lobe between the 
eave and postorbital lobe. H. convexus Miers also appears to possess such a 
structure and many species of Hyastenus have on the upper anterior edge of the 
postorbital lobe a well developed accessory lobe. Sakai (1938:280) has pointed 
out that the abdomen of the female of H. diacanthus (de Haan) consists of 
only five segments. Such features are usually considered sufficient to warrant 
generic separation. 

6. Micippa Leach and Paramicippa H. Milne Edwards. When Paramicippa 
was first set up in 1834, two species, Micippa platipes Riippell and P. tuberculosa 
Milne Edwards were included. The first was designated type cpecies of 
Paramicippa by Miers (1879c: 662) and a definition of that genus g.i.'en at the 
same time. The second species was alone included in Paramicippa in Miers's 
later revision of the two genera (Miers 1885). Such a procedure was quite 
invalid. Since M. platipes is now generally accepted as belonging tc Micippa 
(Sakai 1938, Buitendijk 1939), Paramicippa should fall to that geius. If a 
separate genus is to be established for P. tuberculosa because of the pronounced 
bifid nature of its rostrum or its comparatively long eyestalk or for any other 
reason, a new name must be used. The situation awaits further investigation 
and P. tuberculosa is here included amongst species of Micippa. 

Finally, among the genera introduced to the Australian lists in this paper, 
two require further mention; in one way they may be regarded as merely 
replacing names already in use. 

1. Scyramathia A. Milne Edwards. In 1918 Rathbun placed Hyastenus fultoni 
Grant, 1905 in this genus which has recently been recognised as a synonym 
of Rochinia A. Milne Edwards (Garth 1958: 282). Examination of material 
of Grant's species (11 specimens including relatively large males and females 
from the Australian Museum's collections: reg. no. P.4515, 20 miles east of 
Babel I., Tasmania, 65-70 fms., "Endeavour" Expedition) reveals the male first 
pleopod as being of the "pisoidiform" type (Garth 1958:249; see also his 
pi. Q) with a truncate but poorly expanded apex and similar to that of Rochinia 
occidentalis (Faxon) (see Garth 1958: pi. Q fig. 7). H. fultoni also resembles 
this species in shape and type of ornamentation of the carapace, relative length 
of the rostrum, form of the orbit and in several other features (see Rathbun 
1925: pis. 228, 229 fig. 5). It seems to fit very satisfactorily into Rochinia 
which genus is now added to the Australian fauna. In the Indo-West-Pacific, 
species of Rochinia are known also from Japan, India and South Africa. 

2. Chorilibinia Lockington. Garth (1958: 282) has recently orphaned the 
Australian and New Guinean C. gracilipes Miers, 1879 and the Indian C. 
andamanica Alcock, 1895 by the transference, to the mithracine Stenocionops 
Desmarest, 1823, of Chorilibinia angusta Lockington, 1877, type species (by 
monotypy) of Chorilibinia Lockington, 1877. The Indian and Australian species 
are completely different from C. angusta and resemble species of Libinia Leach 
and Libidoclea H. Milne Edwards & Lucas among the American Pisinae. 
However, examination of C. gracilipes (numerous specimens including relatively 
large males and females from the Australian Museum's collections; reg. no. P. 
14931, Albany Passage, N. Queensland, Melbourne Ward, August 1928) 
shows that it differs from such American species in two important characters. 
First, the female abdomen comprises only five free segments (fourth to sixth 
inclusive fused) in contrast to the seven of Libinia and Libidoclea. Secondly, 
the male first pleopod is of the pisoidiform type with a tapered, acute apex 
and subterminal aperture in contrast to the scyriform type possessed by the 
American species. C. andamanica and C. gracilipes resemble each other in 
structure of the orbit, form of the rostrum, shape of the carapace and 
comparative length of the ambulatories; C. andamanica also has a five-segmented 
abdomen in the female (Alcock 1895). The form of the male first pleopod 

GRIFFIN 269 

is unfortunately unknown for the Indian species. A new genus thus seems 
to be required for these two Indo-West-Pacific species. The name Chlorolibinia 
published by Haswell (1882:17) is surely a mistake for Lockington's genus 
as the latter's name follows the name of the genus. Being an incorrect 
subsequent spelling. Chlorolibinia is unavailable for use under article 33 (b) 
of the International Code of Zoological Nomenclature. The name Austrolibinia 
is therefore proposed and the genus is diagnosed below. 

AUSTROLIBINIA n. gen. 
Chorilibinia Lockington; Miers 1879b: 7; (part: C. gracilipes Miers, 1879). 

Alcock 1895: 221 (part: C. andamanica Alcock, 1895). 
Chlorolibinia Haswell, 1882c: 17; incorrect subsequent spelling of Chorilibinia 
Lockington, 1877. 

Carapace pyriform, armed with a few slender spines and bearing posteriorly 
a broad, medially acute lobe. Rostrum united in basal half, consisting distally 
of two acute, divergent spines. Supraorbital eave well expanded, anterolaterally 
and posterolaterally acute, separated from the large, cupped postorbital lobe 
by a very narrow fissure; intercalated spine absent. Basal antennal article of 
moderate width, provided with a prominent lobe laterally at its base. Ambulatory 
legs long and slender, the first about twice carapace length. Chelipeds shorter 
than the first ambulatory leg in both sexes, chelae moderately inflated in male. 
Abdomen of seven segments in the male, of five in the female, fourth to 
sixth fused. First pleopod of male slender, tapering, apically acute, aperture 
subterminal (based on C. gracilipes only). 

Type species: Chorilibinia gracilipes Miers, 1879. 
V. THE SPECIES 

The characters which are of value at the specific level are for the most 
part different in each genus. For instance, in the genus Notomithrax, the form 
of the crests on the carpus of the cheliped is of importance; in nearly all majines 
the number and arrangement of the spines on the carapace are a reliable 
guide but appear to be of little use in many pisines. Several characters which 
are widely used at the specific level unfortunately differ with age and/or sex. 
These include the shape of the chela and abdomen which change with growth, 
often in a single moult, and also differ according to sex; relative proportions 
of the carapace and length of the spines which change with growth, the 
carapace becoming wider (particularly in pisines and majines) and the spines 
shorter and blunter; and the number of free segments in the abdomen which 
is often different in males and females of the one species. In Huenia species 
the shape of the carapace is strikingly different in males and females. Cases 
in which sexual dimorphism has resulted in the original description of two 
species, one based on the male and one on the female, are, as in other groups 
of animals, not infrequent. The shape of the merus of the third maxilliped and 
of the basal antennal article are often used diagnostically but in some species 
the degree of spinulation or tuberculation of these two structures may change 
during growth. 

Among the 45 species listed by Haswell (1882c), 16 have since suffered 
specific name changes. Some of these have been mentioned in the preceding 
section. Some species, particularly among the Acanthonychinae, are now 
recognised as highly polymorphic so that numerous names are reduced to a 
single valid one. This is true, for example, in the genera Oncinopus de Haan, 
Menaethius Latreille and Huenia de Haan, although the number of Australian 
species currently recognised is the same as that listed by Haswell. Specific name 
changes have taken place, since Haswell's time, particularly in the genera Naxia 
Latreille, Chlorinoides Haswell, Micippa Leach and Tiarinia Dana (see references 
in key). Special mention should be made of the relatively recent recognition 
that Platymaia wyvillethomsoni Miers is a western Pacific species distinct 
from the Indian Ocean P. alcocki Rathbun {P. wyvillethomsoni of Alcock and 
later authors) (Rathbun 1918); Caiman's (1900) Torres Strait material of 
Xenocarcinus tuberculatus is correctly referable to X. depressus Miers (Gordon 
1934, Sakai 1965), whilst Xenocarcinus tuberculatus is a western Pacific species 
distinct from the Indian Ocean X. alcocki Laurie (Sakai 1965); Paramicippa 
hispida Baker (Eruma hispida of McCulloch) is synonymous with Anacinetops 

270 REVIEW OF AUSTRALIAN MAUD SPIDER CRABS 

stimpsoni Miers (Balss 1935); Schizophyrs dama (Herbst) is a western species 
in Australia distinct from the widespread S. aspera (H. Milne Edwards) (Balss 
1935, Yaldwyn 1964); Cyclax spinicinctus Heller (? = C. perryi Dana) is 
distinct from C. suborbicularis (Stimpson) (Forest & Guinot 1961); Micippa 
platipes Riippell and M. philyra (Herbst), often considered synonymous, are 
in fact distinct (Buitendijk 1939); Notomithrax ursus (Herbst) includes Para- 
mithrax latreillei Miers (Bennett 1964, Griffin 1963a) and definitely occurs 
in Australia (McNeill 1953); the Australian material of Paramithrax peronii 
mentioned by Haswell (1882c) is correctly referable to Notomithrax minor 
(Filhol) (Bennett 1964); Leptomithrax australiensis Miers and L. spinulosis 
Haswell are both synonyms of L. gaimardii (H. Milne Edwards) (Griffin 
1963b); Achaeus jissifrons (Haswell) includes A. tenuicollis Miers (Griffin and 
Yaldwyn 1965); and that Sargassocarcinus foliatus Ward also occurs in Japan 
where it has been known under the older but generically inaccurate name of 
Mimulus cristatus Balss (Sakai 1965). In this paper advantage is taken of the 
remarks of previous carcinologists to consider Cancer aragnoides Rumphius 
(specific name misspelt arachnoides by later workers), Cancer longipes Linnaeus, 
Egeria indica Leach and E. herbstii H. Milne Edwards a single species, 
Phalangipus longipes (Miers 1884: 182, Alcock 1895: 224 and Grant & 
McCulloch 1906: 27). Similarly, Naxia cerastes Ortmann is considered synony- 
mous with Naxia taurus Pocock and included as Naxioides taurus (Alcock 
1895: 220, Caiman 1900: 37). Both these species need investigation. 

It is brought out later in this paper that a large number of species 
considered to be restricted to Australia have been recorded only once from 
little material. Nothing is to be gained by listing these GRIFFIN 271 

1. Entomonyx spinosus Miers. This quite widespread Indo-West Pacific species, 
of which Macrocoeloma nummifer Alcock is a synonym, has been recorded 
only once from Australia (as Acanthophrys spinosus by Balss 1929). Entomonyx 
is resurrected as a monotypic genus to accommodate it (Griffin, in press, a). 

2. Criocarcinus superciliosus (Herbst). This widespread Indo-West Pacific 
ophthalmine is included in the key on the basis of material from the Great 
Barrier Reef identified by F. A. McNeill (in prep.). 

3. Achaeus sp. A single specimen in the Australian Museum's collections 
appears to constitute an unnamed species of Achaeus. Additional characters 
of this species have been given elsewhere (Griffin & Yaldwyn 1965). 

4. Chlorinoides goldsboroughi Rathbun. This species, previously known only 
from Hawaii, is included in the key on the basis of two specimens in the 
Australian Museum's collections taken off New South Wales. Additional 
characters for separating this species from the closely related Japanese 
C. brevispinosa Yokoya have recently been given by Sakai (1965:88). 

VI. THE KEY 

The following key to genera and species includes all species recorded 
in the literature from Australian localities, with the exception of Stenorhynchus 
curvirostris and Inachus australis (see above). 

Apart from the characters given for each taxon, the following information is 
also presented. In the case of genera, the name is followed by the world 
geographic range; references to important accounts of the genus as a whole, 
or merely of non-Australian species; and the approximate number of known 
valid species. In the case of species the name is followed by specific synonyms 
(as recorded in the Australian literature); size; distribution within Australia; 
distribution outside Australia; bathymetric range (for details of last four see 
below); further distinguishing characters or other information which may be of 
assistance in more positively identifying the species; and finally (in brackets), 
one or two references to descriptions and illustrations of the species. (Note: Six 
species are figured in this paper 272 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

Bathymetric Distribution: The depth at which species occur is divided 
into five categories: littoral: intertidal; sublittoral: down to 10 fms; shallow 
offshore: 10-50 fms; lower shelf: 50-100 fms; and slope: over 100 fms. In the 
case of species which extend down the slope, the deepest recorded occurrence 
is given. 

Finally, an attempt has been made in the key, at least at the generic level 
and above, to use as far as possible characters which are considered important 
from the phylogenetic point of view; in most cases ready separation is 
achieved. The key is set out with contrasting couplets adjacent so as to permit 
easier comparison of the divisions. 

GRIFFIN 273 

KEY TO SUBFAMILIES OF MAJID BRACHYURA AND AUSTRALIAN 
GENERA AND SPECIES 

1 Eyes either without orbits, or with incomplete or commencing 

orbits. Basal antennal article rather slender- 2 

274 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

8(7) Eyestalks small, retractile beneath edge of carapace. Supraorbital 
eave unexpanded, without lobes or spines. Ambulatory legs 
slender and moderately long, propodi 1 & 2 dilated and com- 
pressed, dactyli of all ambulatories subchelate 

Oncinopus de Haan, 1839. 

Widespread Indo-West Pacific; monotypic: O. aranea de Haan, 
1839 (= O. subpcllucidus Stimpson, 1857; O. angulatus Haswell, 
1880); small; S, SE, NE, N; littoral to shallow offshore; 
carapace weakly calcified (Hale 1927: 125, fig. 122; Sakai 
1965: 66, pi. 27, fig. 1). 

GRIFFIN 275 

13(12) Rostral spines stout, almost straight. Carpus of cheliped with 
lateral ridge rounded . . . . N. aurita (Latreille, 1825) 
(= Halimus laevis Haswell, 1880); large; SW, S, SE; littoral 
(Hale 1927: 129, fig. 127). 

276 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

19(18) Carapace smooth, lacking prominent tubercles. Dactyli of ambula- 
tories 3 and 4 almost semicircular, ventrally spinulated for 

distal 2/3 A. lacertosus Stimpson, 1857 

(= A. breviceps Haswell, 1880); small; SE, NE, NW; Indo-West 
Pacific; sublittoral (Stimpson 1907: 20, pi. Ill fig. 7). 

GRIFFIN 277 

26(25) Supraorbital eave hardly expanded and bearing only a small 
antorbital spine. Rostrum of two weakly curved spines less 
than 1/3 postrostral carapace length. Branchial margin with 
small tubercles or unarmed . . . Zewa McCulloch, 1913 .... 27 
Australia, Japan; see McCulloch (1913), Sakai (1938); 4; 
Australian species, both of which are not known overseas, 
lack prominent marginal branchial tubercles. 

278 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

31(29) Supraorbital eave with a strong, forwardly directed preorbital spine. 
Branchial regions, at least, with prominent lateral expansions. 
Rostrum slender, abdomen in male of 7 free segments 32 

GRIFFIN 279 

37(36) Abdomen of seven segments in both sexes. Carapace pyriform. 
Supraorbital eave laterally expanded. Ambulatory legs generally 
short, seldom more than twice carapace length, usually less .... 38 

280 

REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

42(37) Median suborbital tooth much defiexed and bounded by a broad 
U sinus on both sides. Lobe on first abdominal segment in both 
sexes arcuate and occupying nearly whole width of segment . . . 
P. australiensis Rathbun, 1918 
Small; restricted to Australia, known only from Platypus B. 
(NE) in 7-9 fms: further distinguished by absence of accessory 
spinule on rostral spines (Rathbun 1918: 15, pi. VI). 

GRIFFIN 

281 

48(47) Rostral spines as long as postrostral portion of carapace. Surface 
of carapace eroded but indistinctly tuberculate . . . 

H. sebac White, 1847 
Small; N, NW; Indian O., Indo-Malaya; sublittoral (Alcock 
1895: 213). Figured here pi. XV, figs, b, c. 

282 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

55(52) Carapace strongly tuberculate. Rostral spines almost as long as 
postrostral carapace length ....//. brockii de Man, 1888 
Medium; in Australia known only from Torres Strait (N); 
Indian O., Indo-Malaya; sublittoral (de Man 1888: 221, pi. 7 
fig. 1). 

GRIFFIN 

60(43) Supraorbital eave well expanded posterolaterally, separated from 
postorbital lobe by an extremely narrow fissure. Rostrum bifid 
for distal half only. Carapace armed with a few slender spines 
Ambulatory legs long, the first at least twice carapace length, 

slender, smooth Austrolibinia, n. gen. 

India, New Guinea, Australia, 2: A. gracilipes (Miers, 1879) 
n. comb.; small; NE, N, NW; New Guinea; shallow offshore; 
further distinguished by 2 medial gastric and 1 medial cardiac 
spine, a broad, acute intestinal lobe and 2 dorsal branchial 
spines (Miers 1879b: 7, pi. IV figs. 4, 4a). 

284 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

GRIFFIN 285 

Sternum or abdomen, or both, excavated as rimmed pits in adult 
males and juveniles. Carapace strongly tuberculate. Three or four 

marginal branchial spines 70 

70(69) Postorbital lobe acuminate. Four marginal branchial spines . . . 

L. tuberculatus (Whitelegge, 1900) 
Medium to large; SE; New Zealand, Kermadecs; shallow off- 
shore to lower shelf; Australian forms with short marginal and 
long dorsal spines (Whitelegge 1900: 146, pi. XXXIV figs. 1, 2). 

286 

REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

75(74) Antorbital lobe a narrow, flattened, distally rounded lamella, 
preorbital lobe much wider and somewhat longer. Rostral 
spines weakly curved 76 

GRIFFIN 

81(61) Intercalated spine present 

288 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

87(86) Carapace minutely granular dorsally with a few tubercles, antero- 
lateral margins with about 3 small spinules. Merus of cheliped 

dorsally carinate; ambulatory legs tuberculate 

M. curtispina Haswell, 1880 
Small; NE, N; Singapore; sublittoral; further distinguished by 
basally vertically deflexed and apically inflexed rostrum (Haswell 
1880a: 446, pi. 25 figs. 1, la). 

GRIFFIN 289 

92(91) Carapace with distinct erect tubercles and granules dorsally, 
branchial region with six obtuse tubercles laterally . . . 

T. cornigera (Latreille, 1825) 
(= T. mammillata Haswell, 1880); medium to large; NW; 
Indian O., Indo-Malaya, lapan; littoral; carapace very wide 290 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

GRIFFIN 291 

out most strikingly by the fact that a larger proportion of it is shared with 
other Indo-West Pacific areas than with temperate Australian provinces. 
Thus, of the north-eastern species only 18% extend southward compared 
with 50% which are shared with the Indian Ocean, 47% with Japan and 
36% with Indo-Malaya. Similarly, for the north-western fauna, 28% extend 
southward whereas 72% are shared with the Indian Ocean, 48% with Japan 
and 60% with Indo-Malaya. However, the proportion of species which 
are distributed throughout the Australian tropical area is not very high. For 
example 42% of the species found in north-eastern Australia are shared with 
north-western Australia and the proportion falls to 29% if Torres Strait is 
excluded. Indian Ocean species are represented to approximately equal extents 
(about 60%) in both the north east and north west but the Japanese species 
are definitely best represented in the north east (also about 60%). Widespread 
Indo-West Pacific species which are also widely distributed in the Australian 
tropics include Oncinopus aranea, Menacthius monoceros, several species of 
Hyastenus, Schizophrys aspera, two species of Chlorinoides, three of Micippa 
and two of Tiarinia. 

The low degree of restriction in the tropical fauna is evidenced by fewer 
than 30% of the species in either of the tropical provinces which are not 
found outside Australia and about 30% which are restricted to any one 
province. Restricted Australian species found in the tropics include species of 
Zewa, Hyastenus minimus, Phalangipus australiensis and two species of 
Chlorinoides. 

The temperate provinces contain fewer species and overall there is 
a much greater restriction of these species both to Australia and to particular 
provinces. Thus, only nine species are found in both south-western and south- 
eastern Australia out of a total of 34 south-eastern and 20 south-western species; 
40% of the south-eastern and 50% of the south-western species are not known 
outside Australia. Eight species appear to be widespread temperate forms (two 
species of Naxia, Ephippias endeavouri, Paratymolus latipes, two species of 
Leptomithrax and Chlorinoides spatulifer). The relationships of the temperate 
species are either with tropical Australia (e.g., species of Paratymolus, Zewa, 
Huenia, Xenocarcinus, Hyastenus, Chlorinoides and Micippa) or with the Indo- 
West Pacific (e.g. species of Achaeopsis, Platymaia, Cyrtomaia, Pugettia, Doclea, 
Eurynome and Leptomithrax). The relationships with temperate regions outside 
Australia are slight. Only one species which does not have a tropical distribution, 
Achaeopsis thomsoni, is shared with South Africa. Five species, all of which 
are found in south-eastern Australia, are shared with New Zealand. Of the 
nine genera shared with New Zealand, seven are more or less widespread 
in the Indo-West Pacific. The strong restricted element in the Australian 
temperate fauna is exemplified by species of Naxia, Ephippias endeavouri, 
Paramithrax barhicornis and Tumulosternum longimanus. Of the 37 species 
restricted to Australia, 69% are temperate. 

If only species which penetrate the transition zones between tropical and 
temperate regions are considered, there is indeed only a very slight partitioning 
of the fauna into eastern and western elements. Such eastern species probably 
number no more than six (e.g. Naxia tumida and Notomithrax minor) and 
western ones no more than five (e.g., Paranaxia serpulifera and Schizophrys 
dama). If more stenothermal species are considered a division between eastern 
and western regions is quite clear. 

Distant relationships of the fauna are shown, at the specific level, by nine 
species shared with South Africa, 10 with the Red Sea and eight with Hawaii; 
one species, Achaeopsis thomsoni, is also found in the Atlantic. The vast 
majority of these widespread species are represented in the tropical fauna 
of Australia. 

Figure 3. Known Australian distribution of eight species of majid spider crab. 
(Each circle represents a single recorded locality). 

292 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

The zoogeographical features shown by the species are strongly emphasised 
at the generic level. Thus 58% of the 43 genera have widespread tropical 
Indo-West PaciSc representation. A further 16%, also represented in the 
Indo-West Pacific, have wider relationships, four genera (Achaeus, Achaeopsis, 
Eurynome and Maja) being found in the Atlantic and three (Pugettia, Herbstia 
and Rochima) in the eastern Pacific. An additional 7% (three genera GRIFFIN 293 

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298 REVIEW OF AUSTRALIAN MAJID SPIDER CRABS 

EXPLANATION OF PLATES. 

Plate xv (a) Hyastenus auctus Rathbun. HOLOTYPE, male, carapace length 
from photo (including rostral spines in all cases) about 30 mm. Sulu 
Sea, Philippine Islands, Albatross Exped., U.S. National Museum 
no. 48214 (Photo: U.S. Nat. Mus.). 

(b) Hyastenus sebae White. LECTOTYPE (selected on the advice of 
Dr. I. Gordon), dorsal view, female, carapace length from photo 
about 11.5 mm. Corregidor, Philippine Islands, British Museum 
(N.H.) no. 43.6 (Photo: British Museum). 

(c) Hyastenus sebae White. LECTOTYPE, ventral view (Photo: British 
Museum). 

(d) Huenia bifurcata Streets. Male, carapace length 22 mm., setae 
cleaned from shaft of rostrum only. New South Wales, Australian 
Museum no. P. 14961 (Photo: Anthony Healy). 

(e) Huenia bifurcata Streets. Female, carapace length 21 mm, uncleaned. 
Port Jackson, N.S.W., Aust. Mus. no. G. 5102 (Photo: Anthonv 
Healy). 

Plate xvi (a) Schizophrys aspera (H. Milne Edwards). Dorsal view, male, 
carapace length 60.5 mm. Lord Howe Island, W.R.B. Oliver 
collection, Dominion Museum, Wellington (Photo: Athol Beswick). 

(b) Schizophrys aspera (H. Milne Edwards). Ventral view of sam£ 
specimen (Photo: Athol Beswick). 

Plate xvii (a) Paranaxia serpulifera (Guerin). Male, carapace length 102 mm. 
Darnley Island, Torres Strait, Aust. Mus. no. G. 2469 (Photo: 
Anthony Healy). 

(b) Tiarinia elegans Hasweli. HOLOTYPE, male, carapace length 14.5 
mm. Off Broughton Island, near Port Stephens, N.S.W., 25 fms. 
Aust. Mus. no. G. 5140 (Photo: Anthony Healy). 

AUSTRALIAN ZOOLOGIST. Vol. XIII 

PLATE XV 

M 

"AUSTRALIAN ZOOLOGIST. Vol. XIII 

PI ATE XVI 

SPIDER CRABS (for explanation see page 298). 

AUSTRALIAN ZOOLOGIST. Vol. XIII 

PLATE XVII 

SPIDER CRABS (for explanation see page 298).