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Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part I. Lithodoidea, Lomisoidea and Paguroidea

Authors:
5
ANNOTATED CHECKLIST OF ANOMURAN DECAPOD CRUSTACEANS OF
THE WORLD (EXCLUSIVE OF THE KIWAOIDEA AND
FAMILIES CHIROSTYLIDAE AND GALATHEIDAE OF THE GALATHEOIDEA)
PART I – LITHODOIDEA, LOMISOIDEA AND PAGUROIDEA
Patsy A. McLaughlin
Shannon Point Marine Center, Western Washington University,
1900 Shannon Point Road, Anacortes, WA 98221-4042, USA
Email: hermit@fi dalgo.net
Tomoyuki Komai
Natural History Museum and Institute, Chiba,
955-2 Aoba-cho, Chuo-ku, Chiba, 260-8682 Japan
Email: komai@chiba-muse.or.jp
Rafael Lemaitre
Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,
4210 Silver Hill Road, Suitland, Maryland 20746, USA
Email: lemaitrr@si.edu
Dwi Listyo Rahayu
UPT Mataram, Indonesian Institute of Sciences (LIPI),
Teluk Kodek, Pemenang, Lombok Barat 83352, NTB, Indonesia
Email: dwilistyo@yahoo.com
THE RAFFLES BULLETIN OF ZOOLOGY 2010 Supplement No. 23: 5–107
Date of Publication: 31 Oct.2010
© National University of Singapore
INTRODUCTION
The preamble to the Systema Brachyurorum (Ng et
al., 2008), the catalysis for this and similar annotated
checklists, began with the readers’ attentions directed to
previous catalogues that had contributed substantially to the
associations of scientifi c names with the brachyuran crabs
they represented. Among the paguroid Anomura MacLeay
1838, two publications similarly have been indispensible.
Alcock’s (1905b) catalog provided not only information
on the holdings in the Indian Museum, but a listing of all
species known at the time; Gordan’s (1956) bibliography
of paguroids since Alcock’s publication made available to
the scientifi c community a thorough literary review of fi fty
years of research. A similar cornerstone for lithodids was
Dawson’s (1989) comprehensive bibliography that included
both systematic and fi shery orientated references. However,
during the more than half a century since these monographic
summations, the total number of species has increased
several folds. Only in the Lomisoidea Bouvier, 1895 has
the species number remained only one. In contrast to the 16
genera and 95 species documented by Dawson, recognized
lithodoid species now total 129, in 15 genera (Acantholithus
Stimpson, 1858 formally having been placed in synonymy
with Paralomis White, 1847). Most of these additional taxa
have been assigned to the lithodid genera Lithodes Latreille,
1806 and Paralomis. Although a direct comparison with the
number of species and subspecies (varieties) listed by Gordan
(1956) is not realistic as hers was simply a compilation
of names in the existing literature of the time, there is no
doubt that increased genera and species recognition has been
greatest in the Paguroidea Latreille, 1802: Coenobitidae
Dana 1851, two genera and 17 species; Diogenidae Ortmann,
1892, 20 genera and 428 species; Paguridae Latreille 1802,
75 genera and 542 species; Parapaguridae Smith, 1882, 10
genera and 76 species; Pylocheles Bate, 1888, 10 genera
and 52 species; Pylojacquesidae McLaughlin & Lemaitre,
2001, two genera and two species. However, these numbers,
particularly in the species category, will undoubtedly change
as taxonomic revisions in progress and those contemplated
are completed and published. Perhaps it should be noted that
while the estimated number of paguroid genera 120 provided
by De Grave et al. (2009), agreed with the number of 120
documented in the present checklist, the number of species
differed somewhat, 1,069 in the De Grave et al. (2009)
checklist, and 1,106 in the present study, with 36 new species
added following publication of De Grave et al. (2009).
6
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
HISTORY OF CLASSIFICATION
Two anomuran species, Cancer bernhardus Linnaeus,
1758 and Cancer maja Linnaeus, 1758 were among the
rst decapods offi cially recognized in Linnaeus’ 1758 tenth
edition of Systema Naturae. Cancer bernhardus subsequently
was transferred to the genus Pagurus Fabricius, 1775, while
Cancer maja was assigned still later to Lithodes Latreille,
1806 (as L. arctica Latreille, 1806). As late as Stimpson’s
(1858) classifi cation, the Lithodidae Samouelle, 1819 and
Paguridae were considered separate major taxa; Lomis H.
Milne Edwards, 1837, when specifi cally mentioned, was
aligned with the Lithodidae [see McLaughlin et al. (2007);
Lemaitre & McLaughlin (2009) for specifi cs]. Although
disagreement on the placements of the anomuran families
and interpretations of their relationships continued, the
derivations by Boas (1880b, 1924) and Bouvier (1894 b,
1895a, 1897a) of the Lithodidae from the Paguridae became
firmly implanted in the scientific literature. But it was
Bouvier’s proposed transformation from pagurid to lithodid
pleon that gained general acceptance. In the taxonomic
section of his monumental treatise on the Decapoda,
Balss’ (1957) classifi cation of the Paguridea included the
families Pylochelidae, Paguridae, Coenobitidae, Lomisidae
and Lithodidae. At that point in classifi catory history, the
Paguridae was composed of the subfamilies Pagurinae
sensu Dana (= Diogeninae) and Eupagurinae sensu Brandt
(= Pagurinae).
However, the interpretation of phylogenetic relationships
within the Paguridea, quickly changed and with it the
classifi cation. Based on rather limited developmental data,
MacDonald et al. (1957) divided the Paguridea into two
superfamilies, the Coenobitoidea Dana, 1851, including the
Pylochelidae, Coenobitidae and Diogenidae, and Paguroidea
Latreille, 1802, including the Paguridae, Parapaguridae
and Lithodidae and suggested early divergence of two
lineages from an ancestral anomuran; the Lomisidae was
not considered. Polyphyly in the Paguroidea was first
challenged by McLaughlin (1983b), but it was not until
Martin & Davis (2001) accepted McLaughlin’s (1983b)
argument, that the Paguroidea was reconciled as a single
superfamily, retaining all six paguroid families. Martin &
Davis’ (2001) classifi cation also supported removal of the
Lomisidae to its own family and superfamily as had been
proposed by McLaughlin (1983a). Despite very contentious
disagreements between traditional morphologists and
molecular phylogenetists regarding the relationships among
the Paguroidea s.l. that continue today, only two substantive
changes have occurred to date in the formal classifi cation
of the superfamily. The fi rst was the addition of the family
Pylojacquesidae McLaughlin & Lemaitre, 2001, considered
unique among anomurans in having a chitinous, strongly
toothed mandible that could refl ect an unusual neotenous
or paedomorphic condition. The second was the recently
proposed classifi cation of McLaughlin et al. (2007) in which
the family Lithodidae was formally transferred to its own
superfamily with families Hapalogastridae Brandt, 1850
and Lithodidae. This superfamilial rank for the Lithodoidea
Samouelle, 1819, although in use by some (e.g. De Grave et
al., 2009; Ahyong et al., 2009, Ahyong, 2010b, Ahyong et
al., 2010), is not universally accepted. The positions of the
Lomisoidea as a distinct superfamily and the Porcellanidae
Haworth, 1825 as a family of the superfamily Galatheoidea
Samouelle, 1819 were unchanged in McLaughlin et al.’s
(2007) classifi cation.
Although the formal classifi cation within the Lithodoidea
(or Lithodidae) has not been a source of major contention,
the evolutionary relationship between pagurids and lithodids
most certainly has. The historical account of this presumed
relationship has been reviewed in detail by McLaughlin
& Lemaitre (1997), McLaughlin et al., (2004), Ahyong &
O’Meally (2004), McLaughlin et al. (2007) and need not
be repeated. Suffi ce it to say that if a close phylogenetic
relationship truly does exist, the unresolved question lies in
the determination of the direction of transformation — from
“hermit to king” (e.g., Cunningham et al., 1992; Richter &
Scholtz, 1994; Morrison et al., 2002) or “king to hermit”
(e.g., McLaughlin & Lemaitre, 1997; McLaughlin et al.,
2004; McLaughlin et al., 2007).
Despite the Lomisoidea being represented by the monotypic
genus, Lomis, it did not escape early uncertainty as to its
taxonomic placement and phylogenetic relationship, or the
lack of accuracy in its reported morphology. Because of its
superfi cial resemblance to the Porcellanidae, Lamarck initially
assigned his species to the genus Porcellana Lamarck, 1801.
Henri Milne Edwards (1837) recognized several characters
that set the taxon apart from the porcellanids and therefore
established the genus, Lomis, for the species. Unfortunately,
he was not careful in his morphological assessment and
aligned Lomis with the Lithodidae, a presumed relationship
that was perpetuated for more than a century. Both Bouvier
(1894a, 1895a) and Boas (1926) speculated on the evolution
of Lomis. Bouvier (1894a, 1895a) derived Lomis from
an ancestor intermediate between the paguroid genera
Mixtopagurus A. Milne-Edwards, 1880 and Paguristes Dana,
1851; whereas Boas was of the opinion that it probably had
originated from the oldest symmetrical hermit crab of the
Pylochelidae. Pilgrim (1965) thoroughly investigated the
work of earlier carcinologists, correcting many of the existing
errors and inadequacies. He found no justifi cation for the
inclusion of Lomis in the Porcellanidae or for Bouvier’s
(1894a, 1895a) derivation of the genus from an asymmetrical
ancestor. Nor could Pilgrim fi nd justifi cation for inclusion
of Lomis in the remaining Galatheoidae. He was left with
only the alternative of including Lomis in the Paguridea s.l.
In contrast, McLaughlin (1983a) found the inclusion Lomis
in the Coenobitoidea untenable, and removed the genus and
family to its own superfamily.
Classifi cation of the Paguroidea is by far the most convoluted.
When first established by Fabricius (1775), Pagurus
included a number of Linnaeus’ (1758) non crab-like
species of Cancer. It was thirty-fi ve years later that Pagurus
bernhardus was designated by Latreille (1810) as the type
species of the genus. In a monographic revision, H. Milne
Edwards (1836) removed, as distinct genera, “Coenobites”,
Cancellus H. Milne Edwards, 1836, and Birgus Leach,
7
THE RAFFLES BULLETIN OF ZOOLOGY 2010
1816, and subdivided the genus Pagurus into three
subgenera: Pagures Ordinaires, Pagures Appendiculés, and
Pagures Armés. Pagurus bernhardus was listed fi rst in his
Pagures Ordinaires. Subsequently, he (H. Milne Edwards,
1848) subdivided Pagures Ordinaires still further. Pagurus
bernhardus was included, with 13 additional species with
enlarged right chelipeds, in his section Dextres. The two
other sections, Senestres for species with enlarged left
chelipeds, and Æquimanus for species with generally equal
chelipeds, encompassed the majority of species recognized
at the time, although the subgenera Appendiculés, for species
with sexually modifi ed pleopods, and Armés for species
with spinose intercalary rostriform processes remained as
originally defi ned.
Brandt (1851) proposed the replacement name Eupagurus
for H. Milne Edwards’ (1848) non-Latin Dextres, but did not
address other aspects of H. Milne Edwards’ classifi cation
except to mention in a footnote that a syntopic review of
Pagurus would be forthcoming; it apparently was never
published. About the same time, Dana (1851) published
a revision of the Milne Edwards’ (1848) classification.
For Pagures Ordinaires, Dextres, Dana (1851) proposed
the generic name Bernhardus, with Pagurus bernhardus
as the type, but renamed by Dana, Bernhardus typicus.
Dana disagreed with H. Milne Edwards’ (1836, 1848)
interpretation of the sections Senestres and Æquimanus,
choosing to divide the taxa using the terminal structure
of the left chela rather than chela symmetry. Dana (1851)
proposed the genus Calcinus Dana, 1851 for species with
calcareous-tipped chelae. For the remaining species with
corneous-tipped chelae Dana believed they constituted the
true genus Pagurus. Stimpson (1857) called attention to the
fact that Eupagurus and Bernhardus were synonyms and
Brandt’s (1851) taxon had priority over Dana’s (1851) by a
few weeks; subsequently Stimpson (1858) designated Cancer
bernhardus Linnaeus, 1758 the type of Eupagurus Brandt,
1851. Consequently, Eupagurus became the generic name
for species characterized by enlarged right chelae, whereas
species with corneous-tipped left chelae continued to be
referred to Pagurus sensu Dana (1851).
Although Paul’son (1875) proposed the genus Dardanus
for one of the species assigned to Pagurus (sensu Dana),
Paul’son’s genus was placed in synonymy with Dana’s (1851)
Pagurus by Kossmann (1880). It was not until considerably
later that Benedict (1896) realized that Dana’s ( 1851)
Pagurus contained none of the species originally assigned to
the genus by Fabricius (1775) and pointed out that Latreille’s
(1810) designation of Pagurus bernhardus as the type species
of Pagurus (sensu Fabricius) had priority over its assignment
as the type species of either Bernhardus or Eupagurus.
Around the turn of the century, American authors such as
Rathbun (1899) and Holmes (1900) began using Pagurus
as it had been interpreted by Benedict (1896), and Benedict
(1901b) suggested the replacement name Pagurias Benedict,
1901 for Pagurus (sensu Dana). However, before Pagurias
could become established in the carcinological literature of
the time, Rathbun (1903) recognized that Paul’son’s (1875)
Dardanus was available for Dana’s (1851) taxon. Thus
American and Russian carcinologists replaced Eupagurus
with Pagurus (sensu Fabricius, 1775) and Pagurus (sensu
Dana, 1851) with Dardanus Paul’son, 1875. However, most
Asian and European carcinologists continued to refer species
to Eupagurus Brandt, 1851 and Pagurus (sensu Dana, 1851).
The ensuing nomenclatorial chaos prevailed until the matter
was brought before the International Commission (Forest &
Holthuis, 1955). In its subsequent ruling, the Commission
validated the generic names Pagurus (sensu Fabricius) with
family Paguridae Latreille, 1802, and Dardanus Paul’son,
1875 with family Diognidae Ortmann, 1892 (ICZN Opinion
472), and placed the names Eupagurus Brandt and Bernhardus
Dana on the Offi cial Index of Rejected and Invalid Names
in Zoology (Hemming, 1958).
INFRAORDER ANOMURA MACLEAY, 1838
Infraorder Anomura MacLeay, 1838
= Anomaux Latreille, 1816
(not in Latin, invalid name)
= Anomalia Latreille, 1817
= Anomala Schinz, 1823
= Anomoures H. Milne Edwards, 1832
(not in Latin, invalid name)
= Anomura MacLeay, 1838
Extant families, subfamilies and tribes of the superfamilies
Lithodoidea, Lomisoidea and Paguroidea. –
Superfamily Lithodoidea Samouelle, 1819
Hapalogastridae Brandt, 1850 {1}
= Hapalogastrina Brandt, 1850
Lithodidae Samouelle, 1819 {1}
= Lithodiadae Samouelle, 1819
(invalid original spelling ICZN Opinion 511)
= Lithodinae Samouelle, 1819
= Lithodeacea De Haan, 1833
= Lithodea Brandt, 1848
= Elithodea Brandt, 1848
= Holaspidura Brandt, 1848
= Cryptolithodea Brandt, 1848
= Lithodina Brandt, 1850
= Ostracogastrica Brandt 1850
= Lithodideorum Stimpson, 1858
= Lithodinès Bouvier, 1895
(not in Latin, invalid name)
= Lithodinés Bouvier, 1896
(not in Latin, invalid name)
Superfamily Lomisoidea Bouvier, 1895
= Lomisinés Bouvier, 1894
(not in Latin, invalid name)
Lomisidae Bouvier, 1895
= Lomoidea Glassner, 1969 (incorrect stem)
= Lomidae Glassner, 1969 (incorrect stem)
Superfamily Paguroidea Latreille, 1802
= Paguridea Latreille, 1802
= Coenobitoidea Dana, 1851
8
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Coenobitidae Dana, 1851
Diogenidae Ortmann, 1892
= Diogeninae Ortmann, 1892
= Dardaninae Schmitt, 1926
Paguridae (correction by Samouelle, 1819, of Pagurii
Latreille, 1802)
= Pagurinae Latreille, 1802
Parapaguridae Smith, 1882
Pylochelidae Bate, 1888
= Pomatochelidae Stebbing, 1914
Pylochelinae Bate, 1888
Pomatochelinae Stebbing, 1914
Trizochelinae Forest, 1987
Parapylochelini Forest, 1987
Cancellochelini Forest, 1987
Trizochelini Forest, 1987
Mixtopagurini Bouvier, 1895
Pylojacquesidae McLaughlin & Lemaitre, 2001
DESCRIPTIVE TERMS AND CURRENT STATUS
LITHODOIDEA
General morphology. The body shape is crab-like with
the carapace generally well calcifi ed and covering the entire
cephalothorax. Regions of the dorsal carapace are usually
well defined and the integument provided with spines,
tubercles or granules except in Cryptolithodes Brandt,
1848. The rostrum is variable in size and armature, but
always present; external orbital spines (cf. Macpherson,
1988) are well developed or not. Ocular peduncles are
generally short, with corneas pigmented; no ocular acicles
are developed. Antennular peduncles often are as long as
or longer than antennal peduncles. Antennal peduncles
exhibit supernumerary segmentation and antennal acicles
are well developed, reduced or absent. Third maxillipeds
are pediform and widely separated basally; the ischium has
a well-developed crista dentata and accessory tooth. Gills are
phylobranchiate, 11 on each side: 5 pairs of arthrobranchs
on arthroidal membranes of the third maxillipeds, chelipeds
and pereopods 2–4; 1 pleurobranch is developed on the body
wall above each fourth pereopod.
Chelipeds usually, but not always, are markedly unequal, the
right largest. Pereopods 2–4 are developed as walking legs,
each with terminal claw; pereopod 5 is reduced and most
frequently carried dorsally under the carapace. The pleon is
short, weakly to fi rmly bent under the cephalothorax; with
the tergite of the second pleomere provided with paired
marginal and lateral plates and single median plate, or
nodules in place of actual calcifi ed plates; tergites 3–5 are
primarily membranous in the Hapalogastridae, incompletely
to completely calcifi ed in the Lithodidae. Internally, male
and females reproductive organs and lobules of the midgut
cecum (hepatopancreas), together with other principal organ
systems, are located in the cephalothorax as they are in
brachyurans.
Adult males are completely lacking pleopods; females each
are provided with a pair of small pleopods developed on fi rst
pleonal segment, segments 2–5 each have an unpaired, left
uniramous pleopod. Uropods are entirely absent in adults of
both sexes. The telson is reduced to a small calcifi ed plate.
Development. – Lithodoid larval development is similar to
that of other anomurans in that the developing embryos hatch
from the parental eggs as swimming larvae (zoeae) provided
with carapace, rostrum and cephalothoracic appendages that
include eyes, antennules, antennae, mandibles, maxillules,
maxillae, and fi rst and second maxillipeds. The elongate
pleon is composed of fi ve pleomeres and a telson. Although
lithodoid zoeae often tend to hatch in a somewhat slightly
advanced stage than typical fi rst zoeae (eyes already at least
partially stalked), they generally pass through two or more
zoeal stages, during which the third maxilliped and buds
of the thoracic appendages develop in the cephalothoracic
region, while the pleon adds a sixth pleomere, pleopodal
buds, and in some taxa uniramous uropods (McLaughlin
& Lemaitre, 2001a; McLaughlin et al., 2004). The zoeal
phase is followed by a molt to the megalopa, and it is at this
stage that lithodoid and paguroid developmental differences
begin to become apparent although these differences were
initially observed only from adult morphology and regrettably
misunderstood. Specifi cally, the adult condition suggested
that the membranous pleon of the ancestral paguroid was
progressively invaded by calcifi ed nodules that ultimately
fused to form calcifi ed plates in lithodoids. Initial fusion
occurred primarily in the fi rst and second tergites with the
third through fi fth tergites invaded fi rst with granules and
successively with increasingly larger and more numerous
nodules. Marginal and lateral plates of the adult tergites
were formed by these latter invasions. Ultimately the median
nodules also fused resulting in solid plates (Bouvier, 1894
b, 1895a, 1897a).
However, in a series of developmental studies, Crain &
McLaughlin (2000a, b), McLaughlin & Lemaitre (2001a),
McLaughlin & Paul (2002) and McLaughlin et al. (2004)
were able to demonstrate conclusively that it was not the
calcification and fusion of pleonal tergites in paguroid
evolution that gave rise to lithodoid pleonal armament,
but rather decalcifi cation and sundering. The chitinous or
weakly calcifi ed, but entire, individual tergal plates seen in
the lithodoid megalopae underwent calcium loss and other
changes in fi rst or second juvenile crab stages in certain
genera, coupled with accessory marginal plate development
that resulted in the patterns of plates and/or nodules
recognized in the adults (see McLaughlin et al., 2004, for
detailed account).
In addition to the evidence provided by tergal plate
decalcification, there are other aspects of lithodoid
development that set this superfamily apart from its “crab-
like cousins” and from paguroids. The most signifi cant is seen
in the uropods. Uropods typically fi rst develop in the third
zoeal stage, and remain biramous appendages, albeit with
modifi cations, through adult life. However this is not the case
in lithodoids. From the data available, if uropods develop,
9
THE RAFFLES BULLETIN OF ZOOLOGY 2010
they appear in the third zoeal stage as would be expected.
However in genera such as Acantholithodes Holmes, 1900,
and Cryptolithodes, both of which have four zoeal stages,
uropods never develop. When zoeal uropods occur in
lithodoids, they are uniramous structures that are lost entirely
with the molt to fi rst crab, except in Placetron Schalfeew,
1892. When total loss occurs in that genus presently is not
known, but uropods are lacking in adults.
Current status. – The hypothesized evolutionary relationship
between hermit crabs and king crabs, as has been previously
noted, is fraught with inconsistencies and disagreements.
The sources of these disparities are multifold, but include
archaic and inaccurate morphological data, conflicting
spermatological information, ignored developmental
documentation, a paucity of comprehensive molecular data,
and the dangerous assumption that one molecular “shoe fi ts
all.” As emphasized by Boero (2010) not all species are
molecularly the same. Brief perusal of major phylogenetic
assessments published over the last decade (e.g. Schram,
2001; Morrison et al., 2002; Dixon et al., 2003; Ahyong &
O’Meally 2004; Porter et al., 2005; McLaughlin et al., 2007;
Tsang et al., 2008, in press; Ahyong et al., 2009; Bracken
et al, 2009) demonstrates the remarkable discrepancies
provided, particularly by the molecular data. Nonetheless,
these data confi rmed the “nesting” of the Lithodoidea with
the Paguroidea in the four most recent analyses, all of which
employed refi ned molecular methodologies (Tsang et al.,
2008, in press; Ahyong et al., 2009; Bracken et al., 2009). It
has been argued, and possibly justifi ably, that this apparent
relationship between lithodoids and paguroids is artifi cial
because of the very poor species representations for each.
Tsang et al.’s (2008) dataset included seven paguroids and
one lithodid; Bracken et al.’s (2009) included three paguroids
and one lithodid; Ahyong et al.’s (2009) included 13
paguroids and two lithodoids. However, in their presentation
at the Anomura III Symposium in Tokyo, 2009, Tsang et
al. (in press) increased their species representation to four
lithodids, eight pagurids, eight diogenids + coenobitids,
six parapagurids and four pylochelids. That none of
these investigations were able to recover a monophyletic
Paguroidea should not come as surprising. The monophyly
the authors refer to is that presented by McLaughlin (1983b)
as a challenge to the polyphyly proposed by MacDonald et
al. (1957) based on their limited larval data. Both claims are
archaic and should be relegated to their places in the history of
anomuran historical “side-steps.” Knowledge and understand
of morphological and developmental processes have been
advanced considerably in the ensuing quarter century. And
more recently our “toolbox” has been enhanced by molecular
methodologies. Unfortunately, the advancements on both
fronts have not been incorporated into the analytical processes
employed by either molecular or morphological practitioners.
For example, Ahyong, et al. (2009) found their resulting
polyphyly of asymmetrical paguroids “diffi cult to reconcile
with somatic morphology. ‘A priori’, the suite of associated
modifi cations required for gastropod shell habitation, present
in all asymmetrical hermit crabs is compelling evidence
for monophyly.” Perhaps recognition that not all paguroids
are asymmetrical, gastropod shell-inhabitants and that the
pagurid taxon apparently most closely aligned with their two
lithodid exemplars is a polychaete tube-dweller could provide
a slightly broader interpretation of their results.
It would be akin to burying one’s head in the sand to
completely disregard aspects of the molecular hypothesis
regarding the relationship between lithodoids and paguroids.
It would be equally irresponsible to accept the molecularly
implied relationships without current knowledge of the
morphological and developmental incongruities that exist
between the two superfamilies. In place of the “ancient
history” both morphologists and molecular biologists have
been relying on for explanations of complex issues, isn’t it
about time that the two schools share and work together to
nd the most realistic and parsimonious explanations to the
puzzles of this very multifaceted relationship?
LOMISOIDEA
General morphology. – The body shape is crab-like with the
carapace generally fl attened. Regions of the dorsal carapace
are not well defi ned but the integument is provided with
tufts of short setae. The rostrum is triangular. The ocular
peduncles are mesially fl attened, slightly compressed dorso-
ventrally and well calcifi ed; the corneas are reduced; ocular
orbits and ocular acicles are absent. The antennal peduncles
exhibit supernumerary segmentation; the antennal fl agella
are thick and provided by paired, long setae. An epistomial
spine is present. The third maxillipeds are pediform, widely
separated basally; the ischium has a crista dentata but no
accessory tooth. Gills are phylobranchiate, 14 on each
side: 5 pairs of arthrobranchs on the arthroidal membranes
of the third maxillipeds, chelipeds and pereopods 2–4; 4
pleurobranchs, one on the body wall above the fi rst through
fourth pereopods.
The chelipeds are equal, broad, depressed. Pereopods 2–4 are
developed as walking legs each with terminal claw; pereopod
5 is reduced and carried dorsally under the carapace. The
pleon is symmetrical, with well calcifi ed tergites, weakly
bent under the cephalothorax; the second tergite clearly
visible in dorsal view.
Adult males have paired fi rst and second pleopods modifi ed as
gonopods, pleopods 3–5 and uropods are vestigial or entirely
lacking; females have a pair of pleopods developed on each
of pleonal segments 2–5; the terminal pleomere has a pair of
sexually dimorphically uropods. The telson is reduced to a
small calcifi ed plate in both sexes. One apomorphy unique to
Lomis hirta (Lamarck, 1818) is this dimorphic development of
uropods — well developed and elongate in females, vestigial
in males (McLaughlin et al., 2007).
Development. – Cormbie (1993) and McLaughlin et al.
(2004) provided limited information about development in
the Lomisoidea. Lomis hirta larvae hatch in an advanced
state, with the eyes at least partially stalked and buds
of pereopods 1–4 or 1–5 already present. There is some
additional disagreement between Cormie’s (1993) account
10
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
and that of McLaughlin et al. (2004) with the former author
nding only a four-segmented pleon and poorly developed
pleopod buds. McLaughlin et al. found the pleon to consist
of fi ve pleomeres with fused telson but with the upcoming
separation already indicated in some specimens; pleopods
were moderately well developed and weakly biramous.
McLaughlin et al. were of the opinion that even if errors
in observation occurred between the two reports, interstage
morphological variation was likely to occur in L. hirta as
in other anomurans.
Current status. – The phylogenetic relationship of the
Lomisoidea with other major anomuran taxa is still
unresolved.
PAGUROIDEA
General morphology. – The dorsal surface of the
cephalothorax is covered by a chitinized to moderately
well calcifi ed carapace; the anterior portion of this carapace,
the shield, is partially to completely delineated laterally
and posterolaterally by the cervical groove and posteriorly
by the linea transversalis; near the anterolateral margin the
linea anomurica is usually apparent. The dorsal surfaces
of the shield and posterior carapace may be pitted, weakly
grooved, spinulose or tuberculate, but not divided into regions
corresponding to the internal organs of the cephalothorax.
A rostrum may or may not be developed, but rarely is it
prominently produced; an intercalary rostral process is
occasionally developed; the anterior margin of the shield
may or may not show development of lateral projections.
The ocular peduncles are three-segmented, with the basal
segments fused in the midline and occasionally produced into
spinose projections; no ocular orbits are developed; corneas
are well developed, reduced or rarely absent; ocular acicles
are present. The antennular peduncles are innermost and
three-segmented; the antennal peduncles each are provided
with a supernumerary segment. Gills are bi- or quadriserial
phylobranchiae, varying in number from 14 to eight pairs;
epipods occasionally are present.
Chelipeds are equal, subequal or grossly unequal. Pereopods 2
and 3 are developed as ambulatory legs; pereopod 4 is reduced
in size and modifi ed, usually with a propodal rasp. Pereopod
5 is also reduced, usually provided with a terminal rasp, and
often carried within the carcinoecium. The paguroid pleons
may be elongate, straight or fl exed, bulbous, or reduced;
entirely membranous, provided with calcifi ed tergites on all
six pleomeres, or with calcifi cation reduced but not entirely
absent. Internally, space in the paguroid pleon is occupied by
the male or female reproductive organs and the cecal lobes
of the midgut gland (hepatopancreas).
Adult males may be provided with a pair of gonopores on
the coxae of the fi fth pereopods or only a single gonopore
may be present; rarely, males also may exhibit, presumably
nonfunctional, female gonopores on the coxae of the third
pereopods. In addition to the male gonopores, a sexual
tube of variable length and complexity may be extruded
from either the right or the left gonopore or from both, or
one or the other gonopore may be masked by a tuft of stiff
setae. Adult females most commonly are provided with a
pair of gonopores on the coxae of the third pereopods, but
occasionally only a single left gonopore is present. Females
rarely, if ever, are also provided with male gonopores. Males
of certain genera develop paired fi rst and second pleopods
modifi ed as gonopods, or occasionally only second pleopods
so modifi ed. Males also may be provided with unpaired
pleopods on pleomeres 2–5 or reduced combinations thereof,
usually occurring on the left side of the pleon, but occasionally
on either side; in certain genera, male pleopods are totally
lacking. Females not infrequently are provided paired and
modifi ed fi rst pleopods, occasionally asymmetrically paired
second pleopods. Unpaired female pleopods usually occur
only on the left side of the pleon on pleomere 2–5, but in
certain genera may develop on either side. Unpaired pleopods
are never entirely lost, but their number may decrease to 2
or 3.
The pleon terminates in paired symmetrical or asymmetrical,
biramous uropods, almost always modifi ed by propodal rasps.
The sixth pleonal somite and its uropodal appendages are
most frequently contiguous with the telson; however, in a
few species of two pagurid genera, the telson is separated
from the sixth somite by a lobe of tissue. The reason for this
peculiarity has not yet been determined.
Development. – Larval development in the majority of
paguroids is similar to that of other anomurans in that
the developing embryos hatch from the parental eggs as
swimming larvae (zoeae) provided with carapace, rostrum
and cephalothoracic appendages that include eyes, antennules,
antennae, mandibles, maxillules, maxillae, and fi rst and
second maxillipeds. The elongate pleon typically is composed
of fi ve pleomeres and a telson to which a sixth is added in a
subsequent zoeal stage as is the third maxilliped, pereopodal
and pleopodal buds and uropods. Although four zoeal stages
are common before the molt to megalopa, this number may
be reduced to one or two or extended to several. Rarely has
direct development been documented.
Developmental “clocks” are variable and phase predictability
is challenging. Unlike brachyuran larvae where setal
variability is rare, paguroid setal variability is the norm.
Despite the general developmental patterns paguroids share,
familial and/or generic specializations make zoeal, megalopal,
and early juvenile stages potentially informative from
evolutionary standpoints but caution is required. Development
in one species is not necessarily going to be the same for
potentially allied species. For example, the significant
differences observed in the fi rst zoeal stages of four pylochelid
species (two species of Trizocheles Forest, 1987, one each
of Pylocheles A. Milne-Edwards, 1880, and Pomatocheles
Miers 1879) substantially strengthened Lemaitre et al.’s
(2009) proposition of three distinct evolutionary pathways
within the Pylochelidae. Clear differences also exist between
diogenid and pagurid larval developmental patterns and
even among species of the polyphyletic genus Pagurus.
Unfortunately, interest in the information available from
11
THE RAFFLES BULLETIN OF ZOOLOGY 2010
larval studies has waned, thus another phylogenetic resource
is being underutilized.
Current status. – Most evolutionary hypotheses regarding
the Paguroidea have focused on the presumed derivation of
lithodids from a Pagurus-like ancestor, or the more broadly
explored the role of carcinization in paguroid evolution.
Only relatively recently have molecular techniques been
applied and, as previously alluded to, with mixed results.
Relationships suggested by molecules often have been in
direct confl ict with those supported by morphological or
developmental data, or have surprised phylogenetists with
information known from morphology for decades. However,
one recent exception is striking. Albeit limited in scope,
developmental and morphological data for four species of the
paguroid family Pylochelidae are in accord with molecular
analyses of three of the four same species. All three sources
confi rm a broad separation of the subfamilies Pylochelinae
and Trizochelinae.
A different application of molecular methodology has
provided the fi rst thorough and in-depth review of speciation
in a genus of the Paguroidea. Malay & Paulay (2010)
constructed mitochondrial and nuclear gene phylogenies
of the majority of species of Calcinus using replicates
from multiple locations spanning the known ranges of the
species. Their study demonstrated the practical usefulness
of sequence data in examining speciation, diversifi cation
and even the evolution of coloration. It can only be hoped
that their technique will be applied to other elements of the
paguroid puzzle.
CHECKLIST
SUPERFAMILY LITHODOIDEA
SAMOUELLE, 1819 {1, 2}
Family Hapalogastridae Brandt, 1850 {1}
Acantholithodes Holmes, 1895 {2}
= Acantholithodes Holmes, 1895
(type species Dermaturus hispidus Stimpson, 1860,
by original designation; gender masculine)
Acantholithodes hispidus (Stimpson, 1860)
[Dermaturus]
Dermaturus Brandt, 1850 {3}
= Dermaturus Brandt, 1850 (type species Dermaturus
mandtii Brandt, 1850, by monotypy; gender
masculine)
Dermaturus mandtii Brandt, 1850
Hapalogaster Brandt, 1850
= Hapalogaster Brandt, 1850
(type species Hapalogaster mertensii Brandt, 1850,
by monotype; gender masculine)
Hapalogaster cavicauda Stimpson, 1859
Hapalogaster dentata (De Haan, 1849) [Lomis]
Hapalogaster grebnitzkii Schalfeew, 1892
Hapalogaster mertensii Brandt, 1850
Oedignathus Benedict, 1895
= Oedignathus Benedict, 1895
(type species Oedignathus gilli Benedict, 1895,
by monotypy; gender masculine) {4}
Oedignathus inermis (Stimpson, 1860) [Hapalogaster]
= Hapalogaster brandtii Schalfeew, 1892
= Oedignathus gilli Benedict, 1895
Placetron Schalfeew, 1892
= Placetron Schalfeew, 1892 (type species Placetron
wosnessenskii Schalfeew, 1892, by monotypy;
gender masculine)
= Lepeopus Benedict 1895 (type species Lepeopus
forcipatus Benedict, 1895, by monotypy; gender
masculine) {5}
Placetron wosnessenskii Schalfeew, 1892
= Lepeopus forcipatus Benedict, 1895
NOTES
{1} McLaughlin et al. (2007) removed the family
Lithodidae from the superfamily Paguroidea and
elevated the family to the rank of superfamily with
families Hapalogastridae and Lithodidae.
{2} The status of Acantholithodes is controversial. In his
original description of Dermaturus hispidus, Stimpson
(1860) commented that it might be necessary to
separate it [the species] generically from Dermaturus,
but he had not seen the typical species of Brandt’s
(1850) genus. Holmes (1895) established the genus
Acantholithodes for Stimpson’s (1860) D. hispidus,
differentiating it from Dermaturus by presence of
spines on the body and legs and by the large, spiny
prolongation of the outer side of the second segment of
the antenna, characters entirely lacking in Dermaturus.
Although A. hispidus rarely is encountered, the genus
has been recognized as a valid Northeastern Pacifi c
taxon since its original description (e.g., Rathbun,
1904; Schmitt, 1921; Hart, 1982; Williams et al.,
1989; McLaughlin et al., 2005).
{3} Dawson & Yaldwyn (1985) and Dawson (1989)
considered Acantholithodes a synonym of Dermaturus
and apparently that opinion was shared by Macpherson
(1988) who listed only Dermaturus among the four
genera of the subfamily Hapalogastrinae. Additionally,
the date of publication given by De Grave et al. (2009)
of 1894 is incorrect. Holmes’ paper was published in
1895.
{4} Makarov (1938) and Dawson (1989) both specifi cally
cited Oedignathus inermis as the type species.
However, in his original description of the genus,
Benedict (1895) included only his own new species
O. gilli. Thus O. gilli is the type species of the genus
Oedignathus by monotypy, despite it being a junior
synonym of O. inermis.
12
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Family Lithodidae Samouelle, 1819 {1, 2}
Cryptolithodes Brandt, 1848
= Cryptolithodes Brandt, 1848 (type species
Cryptolithodes typicus Brandt, 1848, by monotypy;
gender masculine)
Cryptolithodes expansus Miers, 1879
Cryptolithodes sitchensis Brandt, 1853
= Cryptolithodes alta-fi ssura Bate, 1864
Cryptolithodes typicus Brandt, 1848
= Cryptolithodes brevifrons Miers, 1879
Glyptolithodes Faxon, 1895
= Glyptolithodes Faxon, 1895
(type species Rhinolithodes cristatipes Faxon,
1893, by monotypy; gender masculine)
Glyptolithodes cristatipes (Faxon, 1893)
[Rhinolithodes]
Lithodes Latreille, 1806
= Lithodes Latreille, 1806,
(type species Cancer maja Linnaeus, 1758, by
monotypy; gender masculine)
= Pseudolithodes Birstein & Vinogradov, 1972
(type species Pseudolithodes zenkevitchi Birstein
& Vinogradov, 1972, by original designation;
gender masculine)
Lithodes aequispinus Benedict, 1895 {2, 3}
= Paralithodes longirostris Navozov-Lavrov,
1929
Lithodes aotearoa Ahyong, 2010
Lithodes australiensis Ahyong, 2010
Lithodes ceramensis Takeda & Nagai, 2004
Lithodes chaddertoni Ahyong, 2010
Lithodes confundens Macpherson, 1988
Lithodes couesi Benedict, 1895 {2}
Lithodes ferox Filhol, 1885 {4}
= Lithodes tropicalis A. Milne-Edwards, 1883
(nomen nudum)
= Lithodes tropicalis Bouvier, 1895
= Pseudolithodes pyriformis Birstein & Vinogradov,
1972
Lithodes formosae Ahyong & Chan, 2010
Lithodes galapagensis Hall & Thatje, 2009
Lithodes jessica Ahyong, 2010
Lithodes longispina Sakai, 1971
Lithodes macquariae Ahyong, 2010
Lithodes maja (Linnaeus, 1758) [Cancer]
=
Cancer horridus Pennant, 1777
= Cancer spinosus Ascanius, 1776
= Cancer spinosus Amboinensis Seba, 1759
= Cancer spinosus maximus, orientalis Seba, 1759
(non binominal, not valid name)
= Parthenope Maja Fabricius, 1798
= Inachus Maja Fabricius, 1798
= Maja eriocheles Lamarck, 1801 (invalid name
ICZN 511)
= Lithodes arctica Latreille, 1806 (invalid name
ICZN 511)
= ? Lithode douteuse H. Milne Edwards, 1837 (not
Latin, not valid name) {5}
= ? Lithodes dubius Brandt, 1848 {5}
= Lithodes Maia Samouelle, 1819
(misspelling of maja)
= Lithodes maia Smith, 1879
(misspelling of maja)
= Maia vulgaris Bosc, 1802
(suppressed generic name ICNZ Opinion 511)
Lithodes mamillifer Macpherson, 1988
Lithodes manningi Macpherson, 1988
Lithodes megacantha Macpherson, 1991
Lithodes murrayi Henderson, 1888
Lithodes nintokuae Sakai, 1978
Lithodes panamensis Faxon, 1893
Lithodes paulayi Macpherson & Chan, 2008
Lithodes rachelae Ahyong, 2010
Lithodes richeri Macpherson, 1990
Lithodes robertsoni Ahyong, 2010
Lithodes santolla (Molina, 1782) [Cancer] {6}
= Lithodes antarctica Hombron & Jacquinot,
1842 {6}
= Pseudolithodes zenkevitschi Birstein & Vinogradov,
1972
Lithodes turkayi Macpherson, 1988
Lithodes turritus Ortmann, 1892
Lithodes unicornis Macpherson, 1984 {7}
Lithodes wiracocha Haig, 1974
Lopholithodes Brandt, 1848
= Lopholithodes Brandt, 1848 (type species
Lopholithodes mandtii Brandt, 1848 by monotypy;
gender masculine)
= Ctenorhinus Gibbons, 1855 (type species
Ctenorhinus setimanus Gibbons, 1855, by
monotypy; gender masculine)
= Echinocerus White, 1848 (type species Echinocerus
cibarius White, 1848, by monotypy; gender
masculine)
Lopholithodes foraminatus (Stimpson, 1859)
[Echinocerus]
Lopholithodes mandtii Brandt, 1848
= Echinocerus cibarius White, 1848
= Ctenorhinus setimanus Gibbons, 1855
Neolithodes A. Milne-Edwards & Bouvier, 1894 {8}
= Neolithodes A. Milne-Edwards & Bouvier, 1894
(type species Lithodes Grimaldii A. Milne-
Edwards & Bouvier 1894, by original designation;
gender masculine)
Neolithodes agassizii (Smith, 1882) [Lithodes]
Neolithodes asperrimus Barnard, 1947
Neolithodes brodiei Dawson & Yaldwyn, 1970
Neolithodes bronwynae Ahyong, 2010
Neolithodes capensis Stebbing, 1905
Neolithodes diomedeae (Benedict, 1895)
[Lithodes] {2}
= Neolithodes martii Birstein & Vinogradov, 1972
Neolithodes duhameli Macpherson, 2004
Neolithodes fl indersi Ahyong, 2010
Neolithodes grimaldii (A. Milne-Edwards & Bouvier,
1894) [Lithodes]
= Lithodes goodei Benedict, 1895 {2}
Neolithodes nipponensis Sakai, 1971
Neolithodes vinogradovi Macpherson, 1988
13
THE RAFFLES BULLETIN OF ZOOLOGY 2010
Neolithodes yaldwyni Ahyong & Dawson, 2006
Paralithodes Brandt, 1848
= Paralithodes Brandt, 1848 (type species Lithodes
brevipes H. Milne Edwards & Lucas 1841, by
monotypy; gender masculine)
Paralithodes brevipes (H. Milne Edwards & Lucas, 1841)
[Lithodes]
Paralithodes californiensis (Benedict, 1895)
[Lithodes] {2, 9}
Paralithodes camtschaticus (Tilesius, 1815) [Maja]
= Lithodes spinosissimus Brandt, 1848
= Lithodes Japonicus De Haan 1849
= Paralithodes rostrofalcatus MacKay, 1932
Paralithodes platypus (Brandt, 1850) [Lithodes]
Paralithodes rathbuni (Benedict, 1895)
[Lithodes] {2, 9}
Paralomis White, 1856
= Paralomis White, 1856 (type species Lithodes
granulosa (Hombron & Jacquinot, 1846, by
original designation; gender feminine) {10}
= Acantholithus Stimpson, 1858 (type species
Lithodes hystrix De Haan, 1849, by original
designation; gender masculine)
= Leptolithodes Benedict, 1895 [type species
Paralomis aculeata Henderson, 1888 by
subsequent designation by Ahyong et al. (2010);
gender masculine] {2, 11}
= Pristopus Benedict, 1895 [type species Pristopus
verrilli Benedict, 1895, by subsequent designation
by Ahyong et al. (2010) designation; gender
masculine] {2, 12}
Paralomis aculeata Henderson, 1888
Paralomis africana Macpherson, 1982
Paralomis alcockiana Hall & Thatje, 2009
Paralomis anamerae Macpherson, 1988
Paralomis arae Macpherson, 2001
Paralomis arethusa Macpherson, 1994
Paralomis aspera Faxon, 1893
Paralomis birsteini Macpherson, 1988
Paralomis bouvieri Hansen, 1908
Paralomis ceres Macpherson, 1989
Paralomis chilensis Andrade, 1980
Paralomis cristata Takeda & Ohta, 1979
Paralomis cristulata Macpherson, 1988
Paralomis cubensis Chace, 1939
Paralomis danida Takeda & Bussarawit, 2007
Paralomis dawsoni Macpherson, 2001
Paralomis diomedeae (Faxon, 1893) [Echinocerus]
Paralomis dofl eini Balss, 1911
Paralomis echidna Ahyong, 2010
Paralomis elongata Spiridonov, Türkay, Arntz & Thatje,
2006
Paralomis erinacea Macpherson, 1988
Paralomis formosa Henderson, 1888
Paralomis gowlettholmes Ahyong, 2010 {13}
Paralomis granulosa (Hombron & Jacquinot, 1846)
[Lithodes] {10}
= Lithodes granulata Jacquinot & Lucas, 1854
{10}
= Lithodes verrucosa Dana, 1852
Paralomis grossmani Macpherson, 1988
Paralomis haigae Eldredge, 1976
Paralomis hirtella de Saint Laurent & Macpherson,
1997
Paralomis histrix (De Haan, 1849) [Lithodes]
Paralomis hystrixoides Sakai, 1980
Paralomis inca Haig, 1974
Paralomis indica Alcock & Anderson, 1899
Paralomis investigatoris Alcock & Anderson, 1899
Paralomis jamsteci Takeda & Hashimoto, 1990
Paralomis japonicus Balss, 1911
Paralomis kyushupalauensis Takeda, 1985
Paralomis longidactylus Birstein & Vinogradov, 1972
{14}
Paralomis longipes Faxon, 1893
Paralomis makarovi Hall & Thatje, 2009
Paralomis manningi Williams, Smith & Baco, 2000
Paralomis medipacifi ca Takeda, 1974
Paralomis mendagnai Macpherson, 2003
Paralomis microps Filhol, 1884
= Rhinolithodes biscayensis Bouvier, 1895 {15}
Paralomis multispina (Benedict, 1895) [Leptolithodes]
{2}
Paralomis nivosa Hall & Thatje, 2009
Paralomis ochthodes Macpherson, 1988
Paralomis odawarai (Sakai, 1980) [Lopholithodes]
Paralomis otsuae Wilson, 1990
Paralomis pacifi ca Sakai, 1978
Paralomis papillata (Benedict, 1895) [Leptolithodes]
{2}
Paralomis pectinata Macpherson, 1988
Paralomis phrixa Macpherson, 1992
Paralomis poorei Ahyong, 2010
Paralomis roeleveldae Kensley, 1981
Paralomis seagranti Eldredge, 1976
Paralomis serrata Macpherson, 1988
Paralomis spectabilis Hansen, 1908
Paralomis spinosissima Birstein & Vinogradov, 1972
Paralomis staplesi Ahyong, 2010
Paralomis stella Macpherson, 1988
Paralomis stevensi Ahyong & Dawson 2006
Paralomis taylorae Ahyong, 2010
Paralomis truncatispinosa Takeda & Miyake, 1980
= Paralomis heterotuberculata Tung, Yang & Li,
1984
Paralomis tuberipes Macpherson, 1988
Paralomis verrilli (Benedict, 1895) [Pristopus] {2}
Paralomis webberi Ahyong, 2010
Paralomis zealandica Dawson & Yaldwyn, 1971
= Paralomis shinkaimaruae Takeda & Hatanaka,
1984
Phyllolithodes Brandt, 1848
= Phyllolithodes Brandt, 1848 (type species
Phyllolithodes papillosus Brandt, 1848 by
monotypy; gender masculine)
= Petaloceras White, 1856 (type species Petaloceras
bellianus White, 1856, by monotypy; gender
masculine)
14
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Phyllolithodes papillosus Brandt, 1848
= Petaloceras bellianus White, 1856
= Phyllolithodes bicornis Bate, 1866
Rhinolithodes Brandt, 1848
= Rhinolithodes Brandt, 1848 (type species
Rhinolithodes wosnessenskii Brandt, 1848, by
monotypy; gender masculine)
Rhinolithodes wosnessenskii Brandt, 1848
Sculptolithodes Makarov, 1934
= Sculptolithodes Makarov, 1934 (type species
Sculptolithodes derjugini Makarov, 1934, by
monotypy; gender masculine)
Sculptolithodes derjugini Makarov, 1934
NOTES
{1} In his table of gill elements in lithodids, Macpherson
(1988b: 18) indicated that pleurobranchs were entirely
absent. This is incorrect. Only in a few genera of
the Paguridae are pleurobranchs lost completely;
lithodoids all have a pleurobranch on the thoracic
wall above each fourth pereopod (McLaughlin et al.,
1997).
{2} Although correctly cited by Dawson (1989), as noted
by Shirley (2002), the date of publication of Benedict’s
report on the lithodids collected by the U.S. Fish
Commission steamer Albatross has been incorrectly
cited by numerous authors including Hart (1982),
Macpherson (1988b) and Williams et al. (1989).
Benedict’s paper was issued as a “separate” (1016)
for volume 17 of the Proceedings of the U.S. National
Museum. The separate’s cover carried the date 1894;
however, according to the table of contents for the
volume, Benedict’s paper was actually published on
29 January 1895.
{3} Also correctly cited by Dawson (1989), and noted
by Shirley (2002), the spelling of the specifi c name
of this taxon has been incorrectly cited by numerous
authors including Macpherson (1988b) and Williams
et al. (1989). Benedict (1895) described the species
as Lithodes aequispinus, incorrectly assigning a
masculine ending to the feminine noun spina. It does
not appear that this is an inadvertent error on the
author’s part; endings of Latin nouns are not changed
to agree in gender with generic name. Nonetheless,
Bouvier (1896) corrected the spelling to the original
form of the noun, and for many years this species was
reported under the name Lithodes aequispina. The
current International Code of Zoological Nomenclature
(ICZN1999: Art. 32.2) is unambiguous in stating that
the original spelling is the correct spelling unless it is
shown to be an inadvertent error; thus L. aequispinus
as the original spelling is correct.
{4} Macpherson (1988b) corrected the error in authorship
of L. ferox made by both Dawson & Yaldwyn (1985)
and Dawson (1989). Those authors attributed the
species to A. Milne-Edwards (1883), although in
his “References” Macpherson incorrectly cited the
publication date as 1884. Authorship of L. ferox
correctly belongs to Filhol (1885). As pointed out
by Macpherson (1988b), A. Milne-Edwards (1883)
mentioned only that a distinct species from off West
Africa was designated Lithodes tropicalis. As Milne-
Edwards’ remark was not accompanied by either a
description or an illustration, the name was a nomen
nudum. The species was first formally described
by Bouvier (1895). However, in the interim, Filhol
(1885) described Lithodes ferox from the same area.
Although Filhol’s specimens proved to be juveniles
of L. tropicalis, Filhol’s name had priority and L.
tropicalis became a junior synonym.
{5} Among the plates of Seba’s (1734–1765; 1828–1831)
Locupletissimi Rerum Naturalium Thesauri &
Planches de Seba (Holthuis 1969), was a fi gure of
a lithodid considered by H. Milne Edwards (1837:
186) to depict the same species as an unidentifi ed
specimen in the collections of the Muséum national
d’Histoire naturelle. Because the specimen was in
poor condition, and he was unsure of its conspecifi city
with the unidentifi ed museum specimens H. Milne
Edwards gave it the name “Lithode douteuse” using
a question mark to indicate his doubt. Brandt (1848)
reported the species in its Latinized form as Lithodes
dubius M. Edw. but although he repeated H. Milne
Edwards’ (1837) diagnosis, noted that he had not seen
the species. Subsequently, Brandt (1851: 93, footnote)
suggested that “Lithode douteuse” was probably a
specimen of Lithodes arcticus, but because of his
uncertainty, retained H. Milne Edwards’ question
mark.
{6} Despite being fi rst described by Molina (1782) as
Cancer santolla, this species was reported under
the name “Lithodes antarctica Jacquinot” for many
years. As discussed by Macpherson (1988), the date
of publication of Jacquinot’s species was uncertain, as
the expedition results were published in several parts
and the illustrations published separately. Based on
the mention by H. Milne Edwards & Lucas (1842-
1844: vol. 6(1): 32, pl. 7) of Lithodes antarctica, the
correct date was taken to be 1844. However, Clark &
Crosnier (2000) painstakenly traced the accurate dates
of publication of the 28 livraisons that comprised the
Atlas of the “Voyage au pole …” and determined that
the correct date of publication was 1842. Authorship
was corrected by Holthuis (2002) from Jacquinot
in Hombron & Jacquinot to Hombron & Jacquinot,
October, 1842.
{7} Dawson & Yaldwyn (1985) and Dawson (1989)
considered L. unicornis to be junior synonym of L.
murrayi. While acknowledging the morphological
similarities between the two species, Macpherson
(1988) believed that certain characters appeared
to confirm their distinctness and we accept his
judgment.
15
THE RAFFLES BULLETIN OF ZOOLOGY 2010
{8} Davie (2002) cited monotypy as the type designation
for Neolithodes, and the source as A. Milne-Edwards
& Bouvier (1894). Actually, the type was fi xed by
A. Milne-Edwards & Bouvier (1894a) by original
designation. These authors included two species,
N. grimaldii and N. agassizi (Smith, 1882) in their
remarks accompanying the original description,
both species initially having been included in the
genus Lithodes. A. Milne-Edwards & Bouvier
(1894a) described L. grimaldii on page 62 with
accompanying plate 3. However, in an appendix to
that report (A. Milne-Edwards & Bouvier, 1894a:
91) the authors introduced the description of their
new genus, Neolithodes with the following statement.
“En comparant Lithodes Grimaldii (voir ci-dessus, p.
62) avec les autres représentants de la sous-famille
des Lithodinés nous avons pu nous convaincre qu’il
diffère de ceux-ci par la structure de son abdomen
et par la forme de son rostre; comme ces differences
sont d’ordre essentiellement générique dans la sous-
famille, il y a lieu de ranger l’espèce de l’Hirondelle
dans un nouveau genre pour lequel nous proposons le
nom de Neolithodes.” The description following was
identical to the one published by A. Milne-Edwards &
Bouvier (1894b), but the latter included two different
illustrations. Despite being titled “Neolithodes genre
nouveau de la sous-famillle des Lithodinés” the second
description was published on 10 July 1894, whereas
the fi rst appeared in print on 1 June 1894.
{9} The original descriptions of Paralithodes californiensis
and P. rathbuni were based on two female specimens
of the former and one male of the latter, respectively.
Benedict (1895) emphasized the appreciable similarities
between the two taxa, but indicated that he was
hesitant to consider them a single species. Although
three additional females were subsequently reported
from off San Diego, California (Schmitt, 1921), Dr.
Schmitt, curator of Crustacea the National Museum
of Natural History for many years, expressed serious
doubts about their distinctiveness and suggested that
perhaps they simply represented males and females
of a single taxon (personal communication to PMcL).
Both species are still reported as part of the eastern
Pacific fauna (e.g., Anderson & Cailliet, 1974;
Wicksten, 1987; Zaklan, 2002); however, their true
status is unclear.
{10} Both authorship and publication date for this species
have been sources of disagreement. Macpherson
(1988b) reemphasized Haig’s (1955) contention that
authorship was established in Hombron & Jacquinot,
1847, page 4, Section Crustacea, volume 3, where the
species was attributed to Jacquinot. Although the text
was not published until 1854, publication of the fi gures
preceded it, and while no actual date of publication
was specifi ed, White’s (1847) inclusion of the species
in his list of Crustacea in the British Museum was
accepted as an indication of the correct date. However,
in their detailed account of the published zoological
results of the expedition of the “Astrolabe” and
“Zélée”, Clark & Crosnier (2000) determined that
Paralomis granulosa (as Lithodes) was fi rst published
in July of 1846 as part of Atlas plate 8. The text of
volume 3 Crustacés was published by Jacquinot and
Lucas in 1854 where the species was cited as Lithodes
granulata. Holthuis (2002) subsequently confi rmed
the correct authorship as Lithodes granulosa Hombron
& Jacquinot, 1846.
{11} Benedict (1895) established the genus Leptolithodes
for certain species previously assigned to Paralomis
that he believed differed from the designated type
species, Lithodes granulosus. Although he included
three previously described taxa and added two new
species of his own, Benedict did not designate a
type species. In a review of the lithodids of Taiwan,
Ahyong et al. (2010) designated Paralomis aculeatus
as the type species of Leptolithodes.
{12} Benedict (1895) established the genus Pristopus,
which he considered similar to Leptolithodes. To
Pristopus, Benedict assigned Henderson’s Paralomis
formosa and his own new species Pristopus verrilli.
Again, Benedict did not designate a type species.
Ahyong et al. (2010) designated Paralomis verrilli
as the type species of Pristopus.
{13} The specifi c epithet, gowlettholmes, is treated as a
noun in apposition by the author, and is correct as
written (Ahyong, per. comm.).
{14} Birstein & Vinogradov (1972) described their new
species as Paralomis longidactylus, the “us” ending
being correct for a masculine noun. However, Takeda
et al. (1984) changed the specific ending to “a”,
presumably because the generic name Paralomis
is feminine. The original spelling has been retained
(ICZN Art. 32.2); dactylus is a noun and as such, need
not agree in gender with its generic name (ICZN Art.
31.2.1).
{15} Authorship of Rhinolithodes biscayensis, a synonym
of Paralomis microps, was credited to A. Milne-
Edwards & Bouvier (1894a) by Bouvier (1896), an
error repeated by Dawson (1989), and to A. Milne-
Edwards & Bouvier (1900) by Dawson & Yaldwyn
(1985). As pointed out by Macpherson (1988b) the
original description of the species by the two authors
was never published. The fi rst published description
and illustrations were presented by Bouvier (1895); no
mention was made of co-authorship, thus the species
is correctly attributed to Bouvier.
SUPERFAMILY LOMISOIDAE BOUVIER, 1895
Family Lomisidae Bouvier, 1895
= Lomisinés Bouvier 1894 (not Latin, not valid
name) {1}
16
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
= Lomisidae Bouvier 1895
= Lomidae Glassner, 1969 (incorrect stem)
Lomis H. Milne Edwards, 1837
= Porcellana hirta (type species Porcellana hirta
Lamarck, 1818, by monotypy; gender feminine)
Lomis hirta (Lamarck, 1818) [Porcellana]
NOTES
{1} Lemaitre & McLaughlin (2009) erred when they
corrected the date for the Lomisoidea from 1895 to
1894. They were citing the date that Bouvier published
the family group name in the French vernacular,
but that was not a valid name. Only when the name
was Latinized by Bouvier in 1895 did it become an
acceptable family-group name.
SUPERFAMILY PAGUROIDEA LATREILLE, 1802
Family Coenobitidae Dana, 1851
Birgus Leach, 1816 {1}
= Birgus Leach (type species Cancer latro Linnaeus,
1767, by monotypy; gender masculine)
Birgus latro (Linnaeus, 1767) [Cancer]
= Cancer crumenatus Rumphius, 1705 (pre-linnean
name; not valid)
= Cancer crumenatus orientalis Seba, 1759 {2}
= Cancer (Astacus) latro Linnaeus, 1767
= Birgus laticauda Latreille, 1829
Coenobita Latreille, 1829 {3}
= Coenobita Latreille, 1829 (type species Pagurus
clypeatus Fabricius, 1787, by monotypy, gender
masculine) {4}
= Eremita Osbeck, 1765 (type species Eremita
javanica Osbeck, 1765, by monotypy) {4}
= Carcinion Jarocki, 1825 (type species Pagurus
clypeatus Olivier = Coenobita olivieri Owen,
1839, by monotypy) {4}
= Coénbite Latreille, 1825 (not Latin, unavailable
name)
= Cenobites Berthold 1827 (type species Pagurus
clypeatus Fabricius, 1787, by monotypy) {4}
= Cenobita H. Milne Edwards 1837 (incorrect
spelling of Coenobita)
Coenobita brevimanus Dana, 1852
= Cenobita clypeata var. brevimanus Dana, 1852
(incorrect spelling of Coenobita)
= Coenobita hilgendorfi Terao 1913 (unnecessary
replacement name) {5}
= Coenobita clypeata var. puerto-galerae Yap-
Chiongco, 1938
Coenobita carnescens Dana, 1851
= Cenobita carnescens Dana, 1851 (incorrect
spelling of Coenobita)
Coenobita cavipes Stimpson, 1858
= Cenobita cavipes Stimpson, 1858 (incorrect
spelling of Coenobita)
= Coenobita baltzeri Neumann, 1878
Coenobita clypeatus (Fabricius, 1787) [Pagurus]
{6}{8}
= Cancellus Terrestris Bahamensis Catesby, 1743
(pre-Linnean name, not valid) {7}
Coenobita compressus H. Milne Edwards, 1836
= Cenobita compressa H. Milne Edwards, 1836
(incorrect spelling of Coenobita)
= Cenobita intermedia Streets, 1871 (incorrect
spelling of Coenobita)
= Cenobita panamensis Streets, 1871 (incorrect
spelling of Coenobita)
Coenobita compta White, 1847 (nomen nudum) {9}
Coenobita laeviuscula White, 1847 (nomen nudum)
{9}
Coenobita longitarsis De Man, 1902
Coenobita olivieri Owen, 1839
= Caenobita olivieri Owen, 1839 (incorrect spelling
of Coenobita)
Coenobita perlatus H. Milne Edwards, 1837
= Cenobita perlata H. Milne Edwards, 1837
(incorrect spelling of Coenobita)
= Coenobita perlatus var. affi nis Miers, 1880
Coenobita purpureus Stimpson, 1858
= Cenobita purpurea Stimpson, 1858 (incorrect
spelling of Coenobita)
Coenobita pseudorugosus Nakasone, 1988
Coenobita rubescens Greeff, 1884
Coenobita rugosus H. Milne Edwards, 1837
= Cenobita rugosa H. Milne Edwards, 1837
(incorrect spelling of Coenobita)
= Coenobita subrugosa Neumann, 1878
= Coenobita rugosa var. wagneri Dofl ein, 1900
Coenobita scaevola (Forskål, 1775) [Cancer]
= Coenobita rugosa var. Jousseaumi Bouvier,
1890
= Coenobita rugosa var. granulata Bouvier, 1890
Coenobita spinosus H. Milne Edwards, 1837
= Cenobita spinosa H. Milne Edwards, 1837
(incorrect spelling of Coenobita)
= Cenobita brunnea Dana, 1851 (incorrect spelling
of Coenobita)
= Birgus hirsutus Hess, 1865
Coenobita variabilis Mc Culloch, 1909
Coenobita violascens Heller, 1862
= Cenobita violascens Heller, 1862 (incorrect
spelling of Coenobita)
NOTES
{1} The date of publication usually given for Birgus is
1815. But as pointed out by Davie (2002) while the
volume of the Transactions of the Linnean Society,
London is dated 1815, the actual publication date is
24 January 1816.
17
THE RAFFLES BULLETIN OF ZOOLOGY 2010
{2} The publication date for volume 3 of Seba’
Locupletissimi rerum naturalium thesauri …
routinely was given as 1761 by carcinologists and
1758 by ichthyologists. However, Holthuis (1969)
demonstrated conclusively that the correct date for the
original publication was 1759. A title reprint edition
was published in 1761.
{3} Morgan & Holthuis (1989) pointed out that while some
authors routinely considered Coenobita masculine,
a similar number thought it to be feminine. Thus
the literature is “sprinkled” with reports of the same
specifi c names ending in “us” or “a” depending upon
the author(s). As explained by Morgan & Holthuis,
Coenobita is later ecclesiastical Latin, meaning “a
monk”, and while the majority of Latin names ending
in “a” are feminine in gender, Coenobita is masculine.
Consequently, the correct ending for species-group
names in Coenobita is the masculine suffi x “us.”
{4} Morgan & Holthuis (1988), concerned that generic
name Coenobita might be replaced by one of three
existing senior, but obscure, synonyms, applied to
the International Commission for conservation of the
name Coenobita. They argued: 1) Although Jarocki’s
(1825) Carcinion was a senior objective synonym, it
was an entirely forgotten name whose reintroduction
would cause considerable undesirable confusion; 2)
Berthold’s (1827) Cenobites was simply a translation
of Latreille’s (1825) vernacular Cénobite and its
reintroduction into carcinological literature would
result in great confusion. Osbeck’s (1765) Eremita
posed a somewhat different problem. Despite the
generalities of Osbeck’s description, which could
justify Eremita’s possible status as a nomen dubium,
the name is the oldest name ever applied to a
hermit crab thus posing a serious threat to paguroid
nomenclature. In Opinion 1575, March 1990, the
Commission placed Coenobita and its type species,
C. clypeatus, on the Offi cial List of Valid Names in
Zoology, and the names Carcinion, Cenobites and
Eremita with its type species E. javanick Osbeck,
1765, on the Offi cial Index of Rejected and Invalid
Names in Zoology.
{5} Terao (1913) proposed the replacement name, C.
hilgendorfi for Coenobita clypeatus sensu Latreille
(1826), stating: “on Hilgendorf’s authority we learn
that Cancer clypeatus Herbst (1791) is not identical
with Latreille’s Coenobita clypeata (1826) but is the
same as Coenobita diogenes M.–Edwards (1837).
… Since, now, this C. clypeatus (Herbst) is clearly
a species distinct from C. clypeata Latr., the latter
must receive a new specifi c name.” However, Terao’s
(1913) interpretation of C. clypeatus was based on a
misidentifi cation of C. clypeatus (= C. brevimanus).
{6} Until recently, considerable confusion has existed
regarding the correct authorship of Coenobita
clypeatus. For example, Latreille (1826) and H. Milne
Edwards (1837) credited Herbst (1791), whereas
Hilgendorf (1869) believed the author to be H. Milne
Edwards (1837) and Rathbun (1919) attributed the
species to Latreille, but incorrectly cited with date
and reference to Olivier (1811). The correct author
is Fabricius (1787).
{7} Catesby’s initial publication is usually cited with the
date 1754, which is the date of the second edition.
Catesby’s (1743, pl. 33, fi gs. 1, 2) illustrations were
reproduced by Latreille (1818) as Pagurus diogenes
Linnaeus.
{8} Not only has authorship of Coenobita clypeatus been a
problem, the actual identity of the species has been as
well, although the problem went unrecognized for two
hundred years. Fabricius (1787) based his description
of the species on Herbst’s (1791), at the time
unpublished, illustrations of two specimens reported
to be from the East Indies. Subsequent carcinologists
incorrectly assumed that both fi gures represented
the same specimen, one simply an enlargement. In
a review Herbst’s collection in the Naturhistorisches
Forschungsinstitut Museum für Naturkunde zu
Berlin, Hilgendorf (1869) was convinced that the two
specimens of Coenobita present in the collection did
not represent C. clypeatus. As a result of his inaccurate
interpretation of the taxon, Hilgendorf concluded that
Herbst (1791) had erred in citing the type locality. He
“corrected” the type locality to West Indies. Although
the West Indian Coenobita had been known for many
years as Coenobita diogenes (Linnaeus), Rathbun
(1919) pointed out that Linnaeus’ (1758) Pagurus
diogenes represented a species of the marine genus
Petrochirus, and she adopted ‘Coenobita clypeatus
Herbst’ for the terrestrial species. It was not until
Herbst’s (1791) specimens were critically reexamined
(McLaughlin & Holthuis, 2002) was it realized that
Herbst had confounded two species, C. rugosus
H. Milne Edwards, 1837, and C violascens Heller,
1862, both Indo-Pacifi c taxa, under the specifi c name
clypeatus. An application was then submitted to the
ICZN to replace the two existing East Indies syntypes
with a West Indies neotype. By so doing, Coenobita
clypeatus, as commonly used for many years would
be retained, as would the two common East Indies
species. In Opinion 2052 (2003), the ICZN set aside
all previous type fi xations for the nominal species
Pagurus clypeatus Fabricius, 1767, and designated the
Caribbean specimen USNM 126773 as the neotype.
Additionally, Pagurus clypeatus Fabricius, 1787,
was added to the Offi cial List of Specifi c Names
in Zoology, with an endorsement recording that the
species is defi ned by its designated neotype. Cenobita
rugosus H. Milne Edwards, 1837, and Cenobita
violascens were similarly added to the Offi cial List.
{9} In his list of Crustacea in the British Museum, Adam
White (1847: 59–62) listed as new species, Coenobita
compta from the collection of Mr. Broderip, and
18
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Coenobita laeviuscula from Port Essington. Neither
taxon was described nor illustrated and mention
of neither could be found in any of White’s other
publications.
Family Diogenidae Ortmann, 1892
Allodardanus Haig & Provenzano, 1965
= Allodardanus Haig & Provenzano, 1965
(type species Allodardanus rugosus Haig &
Provenzano, 1965, by original designation; gender
masculine)
Allodardanus bredini Haig & Provenzano, 1965
Allodardanus midas McLaughlin & Gore, 1985
Allodardanus rugosus Haig & Provenzano, 1965
Aniculus Dana, 1852
= Aniculus Dana, 1852 (type species Pagurus
aniculus Fabricius, 1787, by original designation;
gender masculine)
Aniculus aniculus (Fabricius, 1787)
= Aniculus typicus Dana, 1852
Aniculus elegans Stimpson, 1858
= Aniculus longitarsis Streets, 1871
Aniculus erythraeus Forest, 1984
Aniculus hopperae McLaughlin & Hoover, 1996
Aniculus maximus Edmondson, 1952
Aniculus miyakei Forest, 1984
Aniculus retipes Lewinsohn, 1982
Aniculus sibogae Forest, 1984
Aniculus ursus (Olivier, 1812) [Pagurus] {1}
Aeropaguristes Rahayu & McLaughlin, 2010
= Aeropaguristes Rahayu & McLaughlin, 2010 (type
species Pagurus setosus Filhol, 1885 by original
designation; gender masculine) (replacement
name) {2}
= Stratiotes Thomson, 1899 (junior homonym of
Stratiotes Putzeys, 1846, Coleoptera)
Areopaguristes abbreviatus (Dechancé, 1963)
[Paguristes]
Areopaguristes breviantennatus (Rahayu, 2005)
[Stratiotes]
Areopaguristes cyanops (Forest, 1978) [Paguristes]
Areopaguristes diffi cilis (Forest, 1952) [Paguristes]
Areopaguristes engyops (Barnard, 1947) [Paguristes]
Areopaguristes hewatti (Wass, 1963) [Paguristes]
Areopaguristes hispidus (A. Milne-Edwards & Bouvier,
1892) [Paguristes]
Areopaguristes hummi (Wass, 1955) [Paguristes]
Areopaguristes iris (Forest & de Saint Laurent, 1968)
[Paguristes]
Areopaguristes japonicus (Miyake, 1961) [Paguristes]
Areopaguristes mclaughlinae (Ayón Parente & Hendrickx,
2006) [Stratiotes]
Areopaguristes micheleae (Rahayu, 2005) [Stratiotes]
Areopaguristes ngochoae (Rahayu, 2005) [Stratiotes]
Areopaguristes nigroapiculus (Komai, 2009)
[Stratiotes]
Areopaguristes orbis (Komai, 2009) [Stratiotes]
Areopaguristes perspicax (Nobili, 1906) [Paguristes]
= Paguristes jousseaumii var. perspicax Nobili,
1906
Areopaguristes pilosus (H. Milne Edwards, 1836)
[Pagurus]
Areopaguristes rubrodiscus (Forest, 1952) [Paguristes]
Areopaguristes setosus (H. Milne Edwards, 1848)
[Pagurus]
= Pagurus setosus Filhol, 1885
Areopaguristes taenia (Komai, 1999) [Paguristes]
Areopaguristes tuberculatus (Whitelegge, 1900)
[Paguristes]
Areopaguristes virilis (Forest, 1952) [Paguristes]
Bathynarius Forest, 1989
= Bathynarius Forest, 1989 (type species Clibanarius
anomalus A. Milne-Edwards & Bouvier, 1893, by
original designation; gender masculine)
Bathynarius albicinctus (Alcock, 1905) [Clibanarius]
Bathynarius anomalus (A. Milne-Edwards & Bouvier,
1893) [Clibanarius]
Bathynarius izuensis Komai & Takeda, 2004
Bathynarius pacifi cus Forest, 1993
Bathynarius wolffi Forest, 1993
Calcinus Dana, 1851 {3}
= Calcinus Dana, 1851 (type species Cancer tibicen
Herbst, 1791, by original designation; gender
masculine)
Calcinus albengai Poupin & Lemaitre, 2003
Calcinus anani Poupin & McLaughlin, 1998
Calcinus argus Wooster, 1984
Calcinus californiensis Bouvier, 1898
Calcinus chilensis (H. Milne Edwards, 1836)
[Pagurus] {4}
Calcinus dapsiles Morgan, 1989
Calcinus elegans (H. Milne Edwards, 1836) [Pagurus]
= ? Pagurus fasciatus Bell, 1853 {5}
Calcinus explorator Boone, 1930
Calcinus gaimardii (H. Milne Edwards, 1848)
[Pagurus]
Calcinus gouti Poupin, 1997
Calcinus guamensis Wooster, 1984
Calcinus haigae Wooster, 1984
Calcinus hakahau Poupin & McLaughlin, 1998
Calcinus hazletti Haig & McLaughlin, 1984
Calcinus imperialis Whitelegge, 1901
Calcinus inconspicuus Morgan, 1991
Calcinus isabellae Poupin, 1997
Calcinus kurozumii Asakura & Tachikawa, 2000
Calcinus laevimanus (Randall, 1840) [Pagurus]
= Pagurus tibicen H. Milne Edwards, 1836
(preoccupied name)
= Pagurus lividus H. Milne Edwards, 1848
= Calcinus herbstii De Man, 1887
Calcinus latens (Randall, 1840) [Pagurus]
= Pagurus cristimanus H. Milne Edwards, 1848
= Calcinus intermedius De Man, 1881
= Calcinus terrae-reginae Haswell, 1882
= Calcinus abrolhensis Morgan, 1988 {6}
Calcinus laurentae Haig & McLaughlin, 1984
Calcinus lineapropodus Morgan & Forest, 1991
19
THE RAFFLES BULLETIN OF ZOOLOGY 2010
Calcinus mclaughlinae Poupin & Bouchard, 2006
Calcinus minutus Buitendijk, 1937
Calcinus morgani Rahayu & Forest, 1999
= Calcinus areolatus Rahayu & Forest, 1999
Calcinus nitidus Heller, 1865
Calcinus obscurus Stimpson, 1859 {7}
Calcinus orchidae Poupin, 1997
Calcinus paradoxus Bouvier, 1922 {8}
Calcinus pascuensis Haig, 1974
Calcinus pulcher Forest, 1958
Calcinus revi Poupin & McLaughlin, 1998
Calcinus rosaceus Heller, 1861
Calcinus seurati Forest, 1951
Calcinus sirius Morgan, 1991
Calcinus spicatus Forest, 1951
Calcinus talismani A. Milne-Edwards & Bouvier, 1892
Calcinus tibicen (Herbst, 1791) [Cancer]
= Calcinus sulcatus H. Milne Edwards, 1836 {9}
Calcinus tropidomanus Lewinsohn, 1981
Calcinus tubularis (Linnaeus, 1767) [Cancer]
= Pagurus ornatus Roux, 1830
= Clibanarius rouxi Heller, 1863
Calcinus urabaensis Campos & Lemaitre, 1994
Calcinus vachoni Forest, 1958
Calcinus vanninii Gherardi & McLaughlin, 1994
Calcinus verrillii (Rathbun, 1901) [Clibanarius]
Cancellus H. Milne Edwards, 1836
= Cancellus H. Milne Edwards, 1836 (type species
Cancellus typus H. Milne Edwards, 1836, by
original designation; gender masculine)
Cancellus canaliculatus (Herbst, 1804) [Cancer]
Cancellus frontalis Forest & McLaughlin, 2000
Cancellus investigatoris Alcock, 1905
Cancellus laticoxa Forest & McLaughlin, 2000
Cancellus macrothrix Stebbing, 1924
Cancellus mayoae Forest & McLaughlin, 1998
Cancellus
ornatus Benedict, 1901
Cancellus panglaoensis McLaughlin, 2008
Cancellus parfaiti A. Milne-Edwards & Bouvier, 1891
Cancellus quadraticoxa Morgan & Forest, 1991
Cancellus rhynchogonus Forest & McLaughlin, 2000
Cancellus sphaerogonus Forest & McLaughlin, 2000
Cancellus spongicola Benedict, 1901
Cancellus tanneri Faxon, 1893
Cancellus typus H. Milne Edwards, 1836
Cancellus viridis Mayo, 1973
Ciliopagurus Forest, 1995
= Ciliopagurus Forest, 1995 (type species Cancer
strigatus Herbst, 1804, by original designation;
gender masculine)
Ciliopagurus albatrossi Forest, 1995
Ciliopagurus alcocki Forest, 1995
Ciliopagurus babai Forest, 1995
Ciliopagurus caparti (Forest, 1952) [Trizopagurus]
Ciliopagurus galzini Poupin & Malay, 2009
Ciliopagurus haigae Forest, 1995
Ciliopagurus hawaiiensis (McLaughlin & Bailey-Brock,
1975) [Trizopagurus]
Ciliopagurus krempfi (Forest, 1952) [Trizopagurus]
Ciliopagurus liui Forest, 1995
Ciliopagurus macrolepis Forest, 1995
Ciliopagurus major Forest, 1995
Ciliopagurus pacifi cus Forest, 1995
Ciliopagurus plessisi Forest, 1955
Ciliopagurus shebae (Lewinsohn, 1969) [Trizopagurus]
Ciliopagurus strigatus (Herbst, 1804) [Cancer]
= Pagurus annulipes H. Milne Edwards, 1848
{10}
Ciliopagurus tenebrarum (Alcock, 1905) [Aniculus]
Ciliopagurus tricolor Forest, 1995
Ciliopagurus vakovako Poupin, 2001
Clibanarius Dana, 1852
= Clibanarius Dana, 1852 (type species Cancer
clibanarius Herbst, 1791, by original designation;
gender masculine)
Clibanarius aequabilis (Dana, 1851) [Pagurus]
Clibanarius
africanus Aurivillus, 1898
Clibanarius albidigitus Nobili, 1901
Clibanarius ambonensis Rahayu & Forest, 1993
Clibanarius antennatus Rahayu & Forest, 1993
Clibanarius antillensis Stimpson, 1859 {11}
= Clibanarius brasiliensis Dana, 1852
Clibanarius arethusa De Man, 1888
Clibanarius astathes (Stebbing, 1924) [Calcinus]
Clibanarius bimaculatus (De Haan, 1849) [Pagurus]
Clibanarius bistriatus Rahayu & Forest, 1993
Clibanarius boschmai Buitendijk, 1937
Clibanarius carnifex Heller, 1861
Clibanarius chapini Schmitt, 1926 {12}
Clibanarius clibanarius (Herbst, 1791) [Cancer]
= Clibanarius vulgaris Dana, 1852
Clibanarius cooki Rathbun, 1900
Clibanarius corallinus (H. Milne Edwards, 1848)
[Pagurus]
= Pagurus globosi-manus Dana, 1851
= Clibanarius corallinus var. spinatus Yap-
Chiongco, 1938
Clibanarius cruentatus (H. Milne Edwards, 1848)
[Pagurus]
Clibanarius danai Rahayu & Forest, 1993
Clibanarius demani Buitendijk, 1937
Clibanarius digueti Bouvier, 1898
Clibanarius englaucus Ball & Haig, 1972
Clibanarius elongatus (H. Milne Edwards, 1848)
[Pagurus] {13}
Clibanarius erythropus (Latreille, 1818) [Pagurus]
= Pagurus misanthropus Risso, 1827
Clibanarius eurysternus (Hilgendorf, 1879) [Pagurus]
Clibanarius fonticola McLaughlin & Murray, 1990
Clibanarius foresti Holthuis, 1959
Clibanarius harisi Rahayu, 2003
Clibanarius hirsutimanus Kobjakova, 1971{14}
Clibanarius humilis (Dana, 1851) [Pagurus]
Clibanarius inaequalis (De Haan, 1849) [Pagurus]
Clibanarius infraspinatus (Hilgendorf, 1869)
[Pagurus]
Clibanarius janethaigae Hendrickx & Esparza-Haro,
1997
20
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Clibanarius laevimanus Buitendijk, 1937
Clibanarius lineatus (H. Milne Edwards, 1848)
[Pagurus] {15}
= Clibanarius panamensis (Stimpson, 1859)
= Clibanarius lordi Miers, 1877
Clibanarius longitarsus (De Haan, 1849) [Pagurus]
= ? Pagurus asper H. Milne Edwards, 1848
Clibanarius merguiensis De Man, 1888 {16}
= Clibanarius aequabilis var. merguiensis De Man,
1888
= Clibanarius misanthropus var. merguiensis De
Man, 1888
Clibanarius nathi Chopra & Das, 1940
Clibanarius olivaceus Henderson, 1915
Clibanarius pacifi cus Stimpson, 1858 {17}
Clibanarius padavensis De Man, 1888
Clibanarius ransoni Forest, 1953
Clibanarius rhabdodactylus Forest, 1953
= Clibanarius zebra var. rhabdodactylus Forest,
1953
Clibanarius rosewateri Manning & Chace, 1990 {18}
Clibanarius rubroviria Rahayu, 1999
Clibanarius rutilus Rahayu, 1999
Clibanarius sachalinicus Kobjakova, 1955 {19}
Clibanarius sclopetarius (Herbst, 1796) [Cancer]
= Pagurus tuberculosus H. Milne Edwards, 1836
= Pagurus cubensis de Saussure, 1858
= Clibanarius carnescens Miers, 1877
= Clibanarius formosus Ives, 1892
Clibanarius senegalensis Chevreux & Bouvier, 1892
Clibanarius serenei Rahayu & Forest, 1993
Clibanarius signatus Heller, 1861
= Clibanarius semistriatus Heller, 1862
Clibanarius similis Rahayu & Forest, 1993
Clibanarius snelliusi Buitendijk, 1937
Clibanarius striolatus Dana, 1852
Clibanarius taeniatus (H. Milne Edwards, 1848)
[Pagurus]
Clibanarius tricolor (Gibbes, 1850) [Pagurus]
= Clibanarius hebes Verrill, 1908
=
Clibanarius brachyops Bouvier, 1918
Clibanarius virescens (Krauss, 1843) [Pagurus]
= Clibanarius philippinensis Yap-Chiongco, in
Estampador, 1937
Clibanarius vittatus (Bosc, 1802) [Pagurus]
= Pagurus symmetricus Randall, 1840
= Clibanarius cayennensis Miers, 1877
= Clibanarius speciosus Miers, 1877
Clibanarius willeyi Southwell, 1910 {20}
Clibanarius zebra Dana, 1852 [Pagurus]
Dardanus Paul’son, 1875 {21}
= Dardanus Paul’son, 1875 (type species Pagurus
depressus Heller, 1861, by original designation)
= Pagurias Benedict, 1901 (unnecessary replacement
name)
Dardanus arrosor (Herbst, 1796) [Cancer]
= Aniculus chiltoni Thompson, 1930
Dardanus aspersus (Berthold, 1846) [Pagurus]
= Pagurus diogenes De Haan, 1849
Dardanus australis Forest & Morgan, 1991
Dardanus brachyops Forest, 1962
Dardanus calidus (Risso, 1827) [Pagurus]
= ? Cancer hungarus Herbst, 1791{22}
Dardanus callichela Cook, 1989
Dardanus corrugatus Cook, 1989
Dardanus crassimanus (H. Milne Edwards, 1836)
[Pagurus]
= Pagurus sculptipes Stimpson, 1858
= Pagurus pavimentatus Hilgendorf, 1879
Dardanus dearmatus (Henderson, 1888) [Pagurus]
Dardanus deformis (H. Milne Edwards, 1836)
[Pagurus]
= Pagurus cavipes White, 1847 (nomen nudum)
= Pagurus cavipes White 1847
= Pagurus cultratus White, 1847 (nomen nudum)
= ? Glaucothoe carinata Henderson, 1888 {23}
Dardanus fucosus Biffar & Provenzano, 1972
Dardanus gemmatus (H. Milne Edwards, 1848)
[Pagurus]
Dardanus guttatus (Olivier, 1812) [Pagurus] {1}
= ? Pagurus catephractus Boone, 1935 {24}
Dardanus hessii (Miers, 1884) [Pagurus]
Dardanus imbricatus (H. Milne Edwards, 1848)
[Pagurus]
Dardanus imperator (Miers, 1881) [Pagurus]
Dardanus impressus (De Haan, 1849) [Pagurus]
Dardanus insignis (de Saussure, 1858) [Pagurus]
= Aniculus arrosor var. petersi A. Milne-Edwards,
1880
Dardanus jacquesi Asakura & Hirayama, 2002
Dardanus janethaigae Ayón Parente & Hendrickx,
2009
Dardanus lagopodes (Forskål, 1775) [Cancer]
= Pagurus sanguinolentus Quoy & Gaimard, 1824
= Pagurus affi nis H. Milne Edwards, 1836
= Pagurus euopsis Dana, 1852
= Pagurus depressus Heller, 1861
= Dardanus helleri Paul’son, 1875
Dardanus longior Asakura, 2006
Dardanus magdalenensis Ayón Parente & Hendrickx,
2009
Dardanus megistos (Herbst, 1804) [Cancer]
= Pagurus punctulatus Miers, 1879
= Dardanus jordani Schmitt, 1921{25}
Dardanus nudus Ayón Parente & Hendrickx, 2009
Dardanus pectinatus (Ortmann, 1892) [Pagurus]
= Pagurus arrosor var. pectinata Ortmann, 1892
Dardanus pedunculatus (Herbst, 1804) [Cancer]
= Pagurus asper De Haan, 1849
= Dardanus haani Rathbun, 1903
= Neopagurus horai Kamalaveni, 1950
Dardanus pilosus Ayón Parente & Hendrickx, 2009
Dardanus robustus Asakura, 2006
Dardanus rufus Buitendijk, 1937
Dardanus sanguinocarpus Degener, 1925
Dardanus scutellatus (H. Milne Edwards, 1848)
[Pagurus]
= Pagurus fabimanus Dana, 1852
= Pagurus watasei Terao, 1913
21
THE RAFFLES BULLETIN OF ZOOLOGY 2010
Dardanus setifer (H. Milne Edwards, 1836) [Pagurus]
Dardanus sinistripes (Stimpson, 1859) [Pagurus]
= Pagurus peruensis Balss, 1921
Dardanus squarrosus Cook, 1989
Dardanus stimpsoni Ayón Parente & Hendrickx, 2009
Dardanus sulcatus Edmondson, 1925
Dardanus tinctor (Forskål, 1775) [Cancer]
= Pagurus varipes Heller, 1861
Dardanus umbella Asakura, 2006
Dardanus undulatus (Balss, 1921) [Pagurus]
Dardanus venosus (H. Milne Edwards, 1848)
[Pagurus]
= Pagurus arrosor var. divergens Moreira, 1905
{26}
Dardanus vulnerans (Thallwitz, 1892) [Pagurus]
Dardanus woodmasoni (Alcock, 1905) [Pagurus]
Diogenes Dana, 1851
= Diogenes Dana, 1851 [type species Pagurus miles
Fabricius, 1787, by subsequent designation by
Dana (1852); gender masculine] {27}
Diogenes alias McLaughlin & Holthuis, 2001 {28}
Diogenes avarus Heller, 1865
Diogenes bicristimanus Alcock, 1905
Diogenes biramus Morgan, 1987
Diogenes brevirostris Stimpson, 1858
Diogenes capricorneus Grant & McCulloch, 1907
Diogenes costatus Henderson, 1893
Diogenes crosnieri Dechancé, 1964
Diogenes custos (Fabricius, 1798) [Pagurus]
= Diogenes affi nis Henderson, 1893
Diogenes defl ectomanus Wang & Tung, 1980
Diogenes denticulatus Chevreux & Bouvier, 1892
= Diogenes pugilator var. cristata Balss, 1921
= ? Diogenes pugilator var. subcristata Balss,
1921
Diogenes dorotheae Morgan & Forest, 1991
Diogenes dubius (Herbst, 1804) [Cancer]
Diogenes edwardsii (De Haan, 1849) [Pagurus]
Diogenes extricatus Stebbing, 1910
Diogenes fasciatus Rahayu & Forest, 1995
Diogenes foresti Rahayu & Hortle, 2002
Diogenes goniochirus Forest, 1956
Diogenes granulatus Miers, 1880
Diogenes guttatus Henderson, 1888
Diogenes inglei McLaughlin & Clark, 1997
Diogenes investigatoris Alcock, 1905
Diogenes izananiae Asakura, 2006
Diogenes jousseaumei (Bouvier, 1897) [Troglopagurus]
= Diogenes setocristatus Morgan & Forest, 1991
= Diogenes stenops Morgan & Forest, 1991
Diogenes jubatus (Nobili, 1903) [Troglopagurus]
= Diogenes platyops Rahayu & Forest, 1995
Diogenes karwarensis Nayak & Neelakantan, 1989
Diogenes klaasi Rahayu & Forest, 1995
Diogenes laevicarpus Rahayu, 1996
Diogenes lanaris Yap-Chiongco, in Estampador, 1937
Diogenes leptocerus Forest, 1956
Diogenes lophochir Morgan, 1989
Diogenes maclaughlinae Nayak & Neelakantan, 1985
Diogenes manaarensis (Henderson, 1893)
[Troglopagurus]
Diogenes merguiensis De Man, 1888
Diogenes mercatoris Forest, 1952
Diogenes miles (Fabricius, 1787) [Pagurus]
= Pagurus diaphanus Fabricius, 1798
Diogenes mixtus Lanchester, 1902
= Diogenes intermedius De Man, 1892 (preoccupied
name)
= Diogenes hainanica Wang & Dong, 1977
= Diogenes plavoeti McLaughlin & Clark, 1997
Diogenes moosai Rahayu & Forest, 1995
Diogenes nitidimanus Terao, 1913
Diogenes ovatus Miers, 1881
= Diogenes pugilator var. ovata Miers, 1881
Diogenes pallescens Whitelegge, 1897
= Diogenes gardineri Alcock, 1905
= Diogenes serenei Forest, 1956
Diogenes paracristimanus Wang & Dong, 1977
Diogenes patae Asakura & Godwin, 2006
Diogenes penicillatus Stimpson, 1858
Diogenes persicus (Nobili, 1905) [Troglopagurus]
Diogenes planimanus Henderson, 1893
= Diogenes custos var. planimanus Henderson,
1893
Diogenes pugilator Roux, 1828
= Pagurus algarbiensis Brito Capello, 1875
= Pagurus arenarius Lucas, 1846
= Pagurus Bocagii Brito Capello, 1875
= Pagurus curvimanus Clément, 1874
= Pagurus dillwynii Bate, 1851
= Diogenes pugilator var. gracillima Balss 1921
(misspelling of gracillimanuus)
= Diogenes varians var. gracillimanus Miers,
1881
= Diogenes intermedius Bouvier, 1891
= Pagurus Lafonti Fischer, 1872
= Pagurus ponticus Kessler, 1860
= Diogenes pugilator orientalis Codreanu &
Balcesco, 1968
Diogenes rectimanus Miers, 1884
Diogenes senex Heller, 1865
Diogenes serripes (Costa, 1838) {29}
Diogenes spinicarpus Rahayu & Forest, 1995
Diogenes spinifrons (De Haan, 1849)
Diogenes spinulimanus Miers, 1880
Diogenes tomentosus Wang & Tung, 1980
Diogenes tirmiziae Siddiqui & McLaughlin, 2003
Diogenes tumidus Rahayu & Forest, 1995
Diogenes viridis Haig & Ball, 1988
Diogenes violaceus Henderson, 1893
Diogenes waltairensis Kamalaveni, 1950
= Diogenes custos var. waltairensis Kamalaveni,
1950
Isocheles Stimpson, 1858
= Isocheles Stimpson, 1858 (type species Bernhardus
aequimanus Dana, 1852, by original designation;
gender masculine)
22
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
= Holopagurus Holmes, 1900 (type species
Holopagurus pilosus Holmes, 1900, by original
designation; gender masculine)
Isocheles aequimanus (Dana, 1852) [Bernhardus]
Isocheles pacifi cus Bouvier, 1907
= Isocheles Wurdemanni var. pacifi cus Bouvier,
1907
Isocheles pilosus (Holmes, 1900) [Holopagurus]
Isocheles sawayai Forest & de Saint Laurent, 1968
Isocheles wurdemanni Stimpson, 1859
Loxopagurus Forest, 1964
= Loxopagurus Forest, 1964 (type species Pagurus
loxochelis Moreira, 1901, by original designation;
gender masculine)
Loxopagurus loxochelis (Moreira, 1901) [Pagurus]
Paguristes Dana, 1851
= Paguristes Dana, 1851 (type species Paguristes
hirtus Dana, 1851, by subsequent designation by
Stimpson, 1858; gender masculine)
= Pagurites Lörenthey & Beurlen, 1929 (misspelling
of Paguristes)
Paguristes acanthomerus Ortmann, 1892
Paguristes aciculus Grant, 1905
Paguristes agulhasensis Forest, 1954
Paguristes albimaculatus Komai, 2001
Paguristes alegrias Morgan, 1987
Paguristes alcocki McLaughlin & Rahayu, 2005
Paguristes anahuacus Glassell, 1938
Paguristes anomalus Bouvier, 1918
= Paguristes anaryballus Provenzano, 1965
Paguristes angustithecus McLaughlin & Provenzano,
1974
Paguristes antennarius Rahayu, 2006
Paguristes arostratus Rahayu, 2006
Paguristes aulacis Rahayu & Forest 2009
Paguristes aztatlanensis Glassell, 1937
Paguristes bakeri Holmes, 1900
= ? Paguristes holmesi Glassell, 1937 {30}
Paguristes
balanophilus Alcock, 1905
Paguristes barbatus (Heller, 1862) [Clibanarius]
Paguristes barnardi Forest, 1954
Paguristes brachyrostris Rahayu, 2006
Paguristes brevicornis (Guérin, 1830) [Pagurus]
Paguristes brevirostris Baker, 1905
= Paguristes brevirostrus Mower & Shepard, 1988
(misspelling of brevirostris)
Paguristes cadenati Forest, 1954
Paguristes calvus Alcock, 1905
Paguristes ciliatus Heller, 1862
Paguristes crinitimanus McLaughlin, 2008
Paguristes dampierensis McLaughlin, 2008
Paguristes depressus Stimpson, 1859
= Paguristes depressus Stimpson, 1858 (nomen
nudum)
Paguristes digitalis Stimpson, 1858
= Paguristes kagoshimensis Ortmann, 1892
Paguristes digueti Bouvier, 1893
Paguristes doederleini Komai, 2001
Paguristes eremita (Linnaeus, 1767) [Cancer]
= Pagurus oculatus Fabricius, 1775
= Pagurus maculatus Risso, 1827
Paguristes erythrops Holthuis, 1959
Paguristes fagei Forest, 1952
Paguristes fecundus Faxon, 1893
Paguristes foresti Scelzo, 1971
Paguristes frontalis (H. Milne Edwards, 1836)
[Pagurus]
Paguristes gamianus (H. Milne Edwards, 1836)
[Pagurus]
Paguristes geminatus McLaughlin, 2008
Paguristes gonagrus (H. Milne Edwards, 1836)
[Pagurus]
Paguristes grayi Benedict, 1901
Paguristes hernancortezi McLaughlin & Provenzano,
1974
Paguristes holguinensis Manjón-Cabeza, García Raso &
Martínez Iglesias, 2002
Paguristes incomitatus Alcock, 1905
Paguristes inconstans McLaughlin & Provenzano,
1975
Paguristes insularis Forest, 1966
Paguristes jalur Morgan, 1992
Paguristes jousseaumei Bouvier, 1892
= Paguristes jousseaumei var. glabra Nobili, 1906
= Paguristes jousseaumei var. intermedia Nobili,
1906
Paguristes kimberleyensis Morgan & Forest, 1991
Paguristes lapillatus McLaughlin & Provenzano, 1975
Paguristes laticlavus McLaughlin & Provenzano, 1975
Paguristes lauriei McLaughlin & Hogarth, 1998
Paguristes lewinsohni McLaughlin & Rahayu, 2005
Paguristes limonensis McLaughlin & Provenzano,
1975
Paguristes longirostris Dana, 1851
Paguristes longisetosus Morgan, 1987
Paguristes lymani A. Milne-Edwards & Bouvier, 1893
Paguristes maclaughlinae Martinez-Iglesias & Gómez,
1989
Paguristes macrops Rahayu & Forest, 2009
Paguristes macrotrichus Forest, 1954
Paguristes markhami Sandberg, 1996
Paguristes marocanus A. Milne-Edwards & Bouvier,
1892
Paguristes mauritanicus Bouvier, 1906
Paguristes meloi Nucci & Hebling, 2004
Paguristes microphthalmus Forest, 1952
Paguristes miyakei Forest & McLaughlin, 1998
Paguristes moorei Benedict, 1901
Paguristes mundus Alcock, 1905
Paguristes ocellus Komai, 2010
Paguristes oculiviolaceus Glassell, 1937
Paguristes ortmanni Miyake, 1978
Paguristes oxyacanthus Forest, 1952
Paguristes oxyophthalmus Holthuis, 1959
Paguristes palythophilus Ortmann, 1892
Paguristes paraguanensis McLaughlin & Provenzano,
1975
Paguristes parvus Holmes, 1900
23
THE RAFFLES BULLETIN OF ZOOLOGY 2010
Paguristes pauciparus Forest & de Saint Laurent, 1968
Paguristes petalodactylus Rahayu, 2007
Paguristes perplexus McLaughlin & Provenzano, 1974
Paguristes perrieri Bouvier, 1895
Paguristes planatus A. Milne-Edwards & Bouvier,
1893
Paguristes praedator Glassell, 1937
Paguristes pugil McCulloch, 1913
Paguristes puncticeps Benedict, 1901
Paguristes puniceus Henderson, 1896
= Paguristes puniceus var. unispinosa Balss, 1912
Paguristes purpureantennatus Morgan, 1987
Paguristes pusillus Henderson, 1896
Paguristes robustus Forest & de Saint Laurent, 1968
Paguristes rostralis Forest & de Saint Laurent, 1968
Paguristes rubropictus A. Milne-Edwards & Bouvier,
1892
= Paguristes maculatus var. rubro-picta A. Milne-
Edwards & Bouvier, 1892
= Paguristes oculatus var. brunneo-pictus A. Milne-
Edwards & Bouvier, 1900
Paguristes runyanae Haig & Ball, 1988
Paguristes sanguinimanus Glassell, 1938
Paguristes sayi A. Milne-Edwards & Bouvier, 1893
Paguristes seminudus Stimpson, 1858
Paguristes sericeus A. Milne Edwards, 1880
= Paguristes rectifrons Benedict, 1901
= Paguristes tenuirostris Benedict, 1901
Paguristes simplex Rahayu & McLaughlin, 2006
Paguristes sinensis Tung & Wang, 1966 {31}
Paguristes skoogi Odhner, 1923
Paguristes spectabilis McLaughlin & Provenzano,
1975
Paguristes spinipes A. Milne-Edwards, 1880
= Paguristes armatus Hay, 1917
= Paguristes visor Henderson, 1888
Paguristes squamosus McCulloch, 1913
Paguristes starcki Provenzano, 1965
Paguristes streaensis Pastore, 1984
Paguristes
subpilosus Henderson, 1888
Paguristes sulcatus Baker, 1905
Paguristes syrtensis de Saint Laurent, 1971
Paguristes tomentosus (H. Milne Edwards, 1848)
[Pagurus]
Paguristes tortugae Schmitt, 1933
Paguristes tosaensis Komai, 2010
Paguristes triangulatus A. Milne-Edwards & Bouvier,
1893
Paguristes triangulopsis Forest & de Saint Laurent,
1968
Paguristes triton McLaughlin, 2008
Paguristes turgidus (Stimpson, 1857) [Clibanarius]
= Paguristes turgides Gordan, 1956 (misspelling of
turgidus)
Paguristes ulreyi Schmitt, 1921
= Paguristes occator Glassell, 1937
Paguristes versus Komai, 2001
Paguristes wassi Provenzano, 1961
Paguristes weddellii (H. Milne Edwards, 1848)
[Pagurus]
= Paguristes hirtus Dana, 1851
Paguristes werdingi Campos & Sanchez, 1995
Paguristes zebra Campos & Sanchez, 1995
Paguristes zhejiangensis Wang & Tung, 1982
= Paguristes pusillus zhejiangensis Wang & Tung,
1982
Paguropsis Henderson, 1888
= Paguropsis Henderson, 1888 (type species
Paguropsis typicus Henderson, 1888, by monotypy;
gender feminine)
= Chlaenopagurus Alcock, 1899 (type species
Chlaenopagurus andersoni Alcock 1899, by
monotypy; gender masculine)
Paguropsis typica Henderson, 1888 {32}
= Paguropsis andersoni (Alcock, 1899)
[Chlaenopagurus] {33}
Petrochirus Stimpson, 1858
= Petrochirus Stimpson, 1858 (type species Pagurus
granulatus Olivier, 1812, by original designation;
gender masculine)
Petrochirus californiensis Bouvier, 1895
Petrochirus diogenes (Linnaeus, 1758) [Cancer] {34}
= Petrochirus bahamensis (Herbst, 1791)
[Cancer]
= Petrochirus granulatus (Olivier, 1812)
[Pagurus] {1}
Petrochirus pustulatus (H. Milne Edwards, 1848)
[Pagurus]
= Isocheles gracilis Miers, 1881
= Petrochirus cavitarius Osorio, 1887
Pseudopaguristes McLaughlin, 2002
= Pseudopaguristes McLaughlin, 2002 (type species
Pseudopaguristes janetkae McLaughlin, 2002, by
original designation; gender masculine)
Pseudopaguristes araeos Rahayu, 2007
Pseudopaguristes asper Rahayu, 2005
Pseudopaguristes bicolor Asakura & Kosuge, 2004
Pseudopaguristes bollandi Asakura & McLaughlin,
2003
Pseudopaguristes calliopsis (Forest & de Saint Laurent,
1968) [Paguristes]
Pseudopaguristes hians (Henderson, 1888)
[Paguristes]
= Diogenes desipiens Lanchester, 1902
Pseudopaguristes invisisacculus (McLaughlin &
Provenzano, 1974) [Paguristes]
Pseudopaguristes janetkae McLaughlin, 2002
Pseudopaguristes kuekenthali (De Man, 1902)
[Paguristes]
Pseudopaguristes laurentae (Morgan & Forest, 1991)
[Paguristes]
= Paguristes brachytes Komai, 1999
= Pseudopaguristes gracilis Rahayu, 2005
Pseudopaguristes monoporus (Morgan, 1987)
[Paguristes]
Pseudopaguristes shidarai Asakura, 2004
24
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea
Pseudopagurus Forest, 1952
= Pseudopagurus Forest, 1952 (type species
Pagurus granulimanus Miers, 1881, by original
designation; masculine)
Pseudopagurus granulimanus (Miers, 1881)
Pseudopagurus biafrensis (Monod, 1927)
Strigopagurus Forest, 1995
= Strigopagurus Forest, 1995 (type species Pagurus
strigimanus White, 1847, by original designation;
gender masculine)
Strigopagurus bilineatus Forest, 1995
Strigopagurus boreonotus Forest, 1995
= Clibanarius multipunctatus Wang & Tung,
1986
Strigopagurus elongatus Forest, 1995
Strigopagurus poupini Forest, 1995
Strigopagurus strigimanus (White, 1847) [Pagurus]
= Pagurus strigimanus White, 1847 (nomen
nudum)
Tisea Forest & Morgan, 1991
= Tisea Forest & Morgan, 1991 (type speciesTisea
grandis Forest & Morgan, 1991, by original
designation; gender feminine)
Tisea grandis Forest & Morgan, 1991
Trizopagurus Forest, 1952
= Trizopagurus Forest, 1952 (type species Clibanarius
melitai Chevreux & Bouvier, 1892, by original
designation; gender masculine)
Trizopagurus melitai (Chevreux & Bouvier 1892)
[Clibanarius]
Trizopagurus magnifi cus (Bouvier, 1898) [Clibanarius]
= Clibanarius chetyrkini Boone, 1932
Trizopagurus rubrocinctus Forest & Garcia Raso, 1990
Incerta sedis
Pagurus lar Heller, 1862
NOTES
{1} Volume 8 of Olivier’s (1811–1812) Encyclopédie
Methodique (Dictionnaire Encyclopédique
Méthodique) was published in two parts. Although
his paguroid descriptions most frequently have been
cited with the 1811 date, as pointed out by Lewinsohn
(1982a), Sherborn & Woodward (1906) showed that
the second part (pages 361–722 and containing the
“Pagures”) was not published until 1812.
{2} Forest & McLaughlin (2000) described in detail the
confusion that had existed regarding the New Zealand
species Paguristes pilosus and P. setosus and the
errors made by Filhol (1885b, d) and Thomson (1899)
that led to the establishment, by Thomson, of the
genus Stratiotes with Pagurus setosus Filhol, 1885,
as its type. Although Alcock (1905b) questionably
considered Thomson’s (1899) Stratiotes setosus a
synonym of H. Milne Edwards’ Paguristes setosus, it
was Forest & McLaughlin (2000) that fi nally placed
the two taxa in synonymy. However, when Rahayu
(2005) found the gill number of Paguristes species
in the collection of the Muséum national d’Histoire
naturelle variable, she restricted Paguristes to species
having 13 pairs of gills and reinstated Stratiotes for
species with only 12 pairs. Unfortunately Stratiotes
Thomson was found to be a junior homonym of
Stratiotes Putzeys, 1846, and has been replaced by
Areopaguristes Rahayu & McLaughlin, 2010.
{3} A newly published molecular analysis of the genus
Calcinus (Malay & Paulay, 2010) has indicated
that several previously unrecognized species are
distinguishable, not only at the molecular level, but
also by coloration.
{4) Poupin & Bouchard (2006) considered the status of
C. chilensis uncertain.
{5} On the authority of L. B. Holthuis, Biffar &
Provenzano (1972) reported Pagurus fasciatus Bell,
1853 (as Dardanus) to be a synonym of Calcinus
elegans. The enlarged left cheliped, setose ventral
margins of the propodi of the ambulatory legs, and the
blue banding of these legs would support this opinion.
However, its reported collection site of Falmouth,
England may justify Ingle’s (1993) opinion that it
should be regarded incerta sedis. It is entered in this
checklist as a questionable synonym.
{6} Although Morgan’s (1988) Australian Calcinus
abrolhensis proved to be conspecifi c with C. latens,
the Hawaiian population thought to represent this
latter taxon apparently represents a distinct species.
{7} The date of publication for Calcinus obscurus has been
given in the older literature as 1862 (e.g., Gordan,
1956). However, Stimpson’s description appeared in
a separate published in 1859, and that is the correct
date of publication.
{8} Calcinus paradoxus, represented by a single specimen
was thought to represent a Calcinus species by Ingle
(1993) but its placement was questioned by both
the Bouvier (1922) and by Joseph Poupin (personal
communication).
{9} Calcinus sulcatus H. Milne Edwards, 1836, was put
into synonymy with Calcinus tibicen by Hilgendorf
(1869) because he thought that Herbst (1791) had erred
in his type locality. It was shown by McLaughlin &
Holthuis (2002) that it was Hilgendorf, not Herbst
who was in error regarding Herbst’s type locality for
another presumed western Atlantic species, Coenobita
clypeatus Herbst. Thus is possible that further
examination of Herbst’s material will show Calcinus
sulcatus is a species distinct from C. tibicen.
25
THE RAFFLES BULLETIN OF ZOOLOGY 2010
{10} Eupagurus annulipes Stimpson, 1860, became a
secondary junior homonym of Pagurus annulipes H.
Milne Edwards, 1836, when the ICZN (Opinion 472)
placed Eupagurus on the Offi cical Index of Rejected
and Invalid Generic Names in Zoology. However,
Milne Edwards’ species had been transferred from
Pagurus to Aniculus, then Trizopagurus, and fi nally
to Ciliopagurus (Forest, 1995), and was never
considered conspecifi c with Stimpson’s (1860) taxon.
Consequently, no replacement name was required.
{11} Although Clibanarius antillensis Stimpson, 1859, was
the acknowledged junior synonym of C. brasiliensis
Dana, 1852a, Forest & de Saint Laurent (1968: 99)
cited common usage as their reason for selecting
Stimpson’s name.
{12} Clibanarius chapini was considered a subspecies of
C. tricolor by Forest & de Saint Laurent (1968), a
ranking dismissed by Manning & Chace (1990).
{13} The status and validity of Pagurus elongatus H. Milne
Edwards, 1848, are uncertain. H. Milne Edwards
(1848) in his original description said that it was very
close to Pagurus tuberculosus H. Milne Edwards,
1836 [= Clibanarius sclopetarius (Herbst, 1791)];
the type locality was given as the Hogolu Islands, the
former name of the Truk Islands of the western Pacifi c.
However Fize & Serène (1955) list C. tuberculosus
(H. Milne Edwards, 1848) among their Atlanto-
Mediterranean species of Clibanarius. Clibanarius
sclopetarius is a western Atlantic species.
{14} The status and validity of Kobjakova’s (1971)
Clibanarius hirsutimanus are uncertain.
{15} Rahayu & Forest (1992) determined that Clibanarius
lineatus was not the Indo-Pacifi c species that it had
been thought to be, but rather H. Milne Edwards’
(1848) Pagurus lineatus collected by Gaudichaud
off the west coast of South America, and known
regionally as Clibanarius panamensis (Stimpson,
1859).
{16} Originally described as a subspecies of Clibanarius
aequabilis Dana, 1851, De Man’s Clibanarius
aequabilis merguiensis was elevated to full specifi c
rank by Forest (1953) when he determined that C.
aequabilis was an Atlantic species.
{17} Clibanarius pacifi cus is another enigmatic species.
According to McLaughlin et al. (2007) the species has
not been accurately reported from anywhere except the
type localities of Tanegashiima and Anami-Oshima,
Japan. All recent investigations by one of the authors
(TK) failed to locate any specimens fi tting Stimpson’s
(1858) description.
{18} In their discussion of Clibanarius rosewateri Manning
and Chace (1990: 37) used the manuscript name C.
acensionis; however, the etymology for this species
clearly indicated that C. acensionis was a laspus
calami on the parts of the authors.
{19} As with C. hirsutimanus, the status and validity of
Kobjakova’s (1955) Clibanarius sachalinicus are
uncertain.
{20} Clibanarius willeyi Southwell, 1910 appears to have
been overlooked. It was not mentioned by Fize and
Serène (1955) in their monographic treatment of
Indo-Pacific species or by Gordan (1956) in her
bibliographic account of pagurids not covered by
Alcock (1905b). Whether it will prove to be a forgotten
senior synonym has yet to be determined.
{21} As previously mentioned, although Paul’son (1875)
proposed the genus Dardanus for one of the species
assigned to Pagurus (sensu Dana), Paul’son’s
genus soon thereafter was placed in synonymy with
Dana’s Pagurus. It was not until considerably later
that Benedict (1896) realized that Dana’s Pagurus
contained none of the species originally assigned to the
genus by Fabricius (1775). As Bernhardus had been
shown to be a synonym of Eupagurus and both species
had the same type species originally designated for
Pagurus sensu Fabricius, Benedict (1896) urged
carcinologists to return to using the name Pagurus
for species then assigned to Eupagurus. However,
while Benedict’s recommendation was accepted by
most American and Russian researchers the same was
not true for most European and Asian workers. This
nomenclatorial confusion was resolved only after the
question of the true interpretation of Pagurus was
brought before the ICZN (Forest & Holthuis, 1955).
In its subsequent ruling (Opinion 472), Pagurus sensu
Fabricius and Dardanus Paul’son were placed on the
Offi cial Index of Valid Names in Zoology.
{22} Sakai (1999) suggested that Herbst’s (1791) Cancer
hungarus might be conspecific with Dardanus
calidus; however, the type apparently is no longer
extant, and Herbst’ brief description is inadequate
for confi rmation. It is questionably included as a
synonym.
{23} Initially described as a distinct species, Glaucothoe
carinata subsequently was shown to be the megalopal
stage of a Dardanus species. Its true specifi c identity
is not known with certainty, thus it is included
only questionably in the synonymy of Dardanus
deformis.
{24} McLaughlin et al. (2005: 303) pointed out that
Boone’s species had not been found since the original
description and suggested that D. cataphractus most
probably was actually a specimen of D. guttatus, a
species that had been reported occasionally in the
Hawaiian Islands.