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THE RAFFLES BULLETIN OF ZOOLOGY 2010 Supplement No.
23: 5107
Date of Publication: 31 Oct.2010
National University of Singapore
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@fidalgo.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
INTRODUCTION
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).
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 scientific names with the brachyuran crabs
they represented. Among the paguroid Anomura MacLeay
1838, two publications similarly have been indispensible.
Alcocks (1905b) catalog provided not only information
on the holdings in the Indian Museum, but a listing of all
species known at the time; Gordans (1956) bibliography
of paguroids since Alcocks publication made available to
the scientific community a thorough literary review of fifty
years of research. A similar cornerstone for lithodids was
Dawsons (1989) comprehensive bibliography that included
both systematic and fishery 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
McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

HISTORY OF CLASSIFICATION
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) classification.
Two anomuran species, Cancer bernhardus Linnaeus,

1758 and Cancer maja Linnaeus, 1758 were among the
first decapods officially 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 Stimpsons
(1858) classification, the Lithodidae Samouelle, 1819 and
Paguridae were considered separate major taxa; Lomis H.
Milne Edwards, 1837, when specifically mentioned, was
aligned with the Lithodidae [see McLaughlin et al. (2007);
Lemaitre & McLaughlin (2009) for specifics]. 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
Bouviers proposed transformation from pagurid to lithodid
pleon that gained general acceptance. In the taxonomic
section of his monumental treatise on the Decapoda,
Balss (1957) classification of the Paguridea included the
families Pylochelidae, Paguridae, Coenobitidae, Lomisidae
and Lithodidae. At that point in classificatory history, the
Paguridae was composed of the subfamilies Pagurinae
sensu Dana (= Diogeninae) and Eupagurinae sensu Brandt
(= Pagurinae).
Although the formal classification 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 &
OMeally (2004), McLaughlin et al. (2007) and need not
be repeated. Suffice 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
superficial 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 justification for the
inclusion of Lomis in the Porcellanidae or for Bouviers
(1894a, 1895a) derivation of the genus from an asymmetrical
ancestor. Nor could Pilgrim find justification 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.
However, the interpretation of phylogenetic relationships

within the Paguridea, quickly changed and with it the
classification. 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 McLaughlins (1983b)
argument, that the Paguroidea was reconciled as a single
superfamily, retaining all six paguroid families. Martin &
Davis (2001) classification 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 classification
of the superfamily. The first was the addition of the family
Pylojacquesidae McLaughlin & Lemaitre, 2001, considered
unique among anomurans in having a chitinous, strongly
toothed mandible that could reflect an unusual neotenous
or paedomorphic condition. The second was the recently
proposed classification 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
Classification 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-five 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,
THE RAFFLES BULLETIN OF ZOOLOGY 2010

1816, and subdivided the genus Pagurus into three
subgenera: Pagures Ordinaires, Pagures Appendiculs, and
Pagures Arms. Pagurus bernhardus was listed first 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 quimanus for species with generally equal
chelipeds, encompassed the majority of species recognized
at the time, although the subgenera Appendiculs, for species
with sexually modified pleopods, and Arms for species
with spinose intercalary rostriform processes remained as
originally defined.
American and Russian carcinologists replaced Eupagurus

with Pagurus (sensu Fabricius, 1775) and Pagurus (sensu
Dana, 1851) with Dardanus Paulson, 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 Paulson,
1875 with family Diognidae Ortmann, 1892 (ICZN Opinion
472), and placed the names Eupagurus Brandt and Bernhardus
Dana on the Official Index of Rejected and Invalid Names
in Zoology (Hemming, 1958).
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 classification
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 quimanus,
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
Brandts (1851) taxon had priority over Danas (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).
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
= 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
= Lithodins Bouvier, 1895
= Lithodins Bouvier, 1896
Although Paulson (1875) proposed the genus Dardanus

for one of the species assigned to Pagurus (sensu Dana),
Paulsons genus was placed in synonymy with Danas (1851)
Pagurus by Kossmann (1880). It was not until considerably
later that Benedict (1896) realized that Danas ( 1851)
Pagurus contained none of the species originally assigned to
the genus by Fabricius (1775) and pointed out that Latreilles
(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 Paulsons (1875)
Dardanus was available for Danas (1851) taxon. Thus
Superfamily Lomisoidea Bouvier, 1895

= Lomisins Bouvier, 1894
Lomisidae Bouvier, 1895
= Lomoidea Glassner, 1969 (incorrect stem)
= Lomidae Glassner, 1969 (incorrect stem)
Superfamily Paguroidea Latreille, 1802
= Paguridea Latreille, 1802
= Coenobitoidea Dana, 1851

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
Adult males are completely lacking pleopods; females each

are provided with a pair of small pleopods developed on first
pleonal segment, segments 25 each have an unpaired, left
uniramous pleopod. Uropods are entirely absent in adults of
both sexes. The telson is reduced to a small calcified 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 first and second maxillipeds. The elongate
pleon is composed of five pleomeres and a telson. Although
lithodoid zoeae often tend to hatch in a somewhat slightly
advanced stage than typical first 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. Specifically, the adult condition suggested
that the membranous pleon of the ancestral paguroid was
progressively invaded by calcified nodules that ultimately
fused to form calcified plates in lithodoids. Initial fusion
occurred primarily in the first and second tergites with the
third through fifth tergites invaded first 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).
DESCRIPTIVE TERMS AND CURRENT STATUS

LITHODOIDEA
General morphology. The body shape is crab-like with
the carapace generally well calcified 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 24; 1 pleurobranch is developed on the body
wall above each fourth pereopod.
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 decalcification and sundering. The chitinous or
weakly calcified, but entire, individual tergal plates seen in
the lithodoid megalopae underwent calcium loss and other
changes in first 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).
Chelipeds usually, but not always, are markedly unequal, the

right largest. Pereopods 24 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 firmly 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 calcified plates; tergites 35 are
primarily membranous in the Hapalogastridae, incompletely
to completely calcified 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.
In addition to the evidence provided by tergal plate

decalcification, there are other aspects of lithodoid
development that set this superfamily apart from its crablike cousins and from paguroids. The most significant is seen
in the uropods. Uropods typically first develop in the third
zoeal stage, and remain biramous appendages, albeit with
modifications, through adult life. However this is not the case
in lithodoids. From the data available, if uropods develop,

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 first crab, except in Placetron Schalfeew,
1892. When total loss occurs in that genus presently is not
known, but uropods are lacking in adults.
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 ones 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, isnt it
about time that the two schools share and work together to
find the most realistic and parsimonious explanations to the
puzzles of this very multifaceted relationship?
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 fits
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 &
OMeally 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 confirmed the nesting of the Lithodoidea with
the Paguroidea in the four most recent analyses, all of which
employed refined molecular methodologies (Tsang et al.,
2008, in press; Ahyong et al., 2009; Bracken et al., 2009). It
has been argued, and possibly justifiably, that this apparent
relationship between lithodoids and paguroids is artificial
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 difficult to reconcile
with somatic morphology. A priori, the suite of associated
modifications 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
LOMISOIDEA
General morphology. The body shape is crab-like with the
carapace generally flattened. Regions of the dorsal carapace
are not well defined but the integument is provided with
tufts of short setae. The rostrum is triangular. The ocular
peduncles are mesially flattened, slightly compressed dorsoventrally and well calcified; the corneas are reduced; ocular
orbits and ocular acicles are absent. The antennal peduncles
exhibit supernumerary segmentation; the antennal flagella
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 24; 4
pleurobranchs, one on the body wall above the first through
fourth pereopods.
The chelipeds are equal, broad, depressed. Pereopods 24 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 calcified tergites, weakly
bent under the cephalothorax; the second tergite clearly
visible in dorsal view.
Adult males have paired first and second pleopods modified as
gonopods, pleopods 35 and uropods are vestigial or entirely
lacking; females have a pair of pleopods developed on each
of pleonal segments 25; the terminal pleomere has a pair of
sexually dimorphically uropods. The telson is reduced to a
small calcified 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 14 or 15 already present. There is some
additional disagreement between Cormies (1993) account

and that of McLaughlin et al. (2004) with the former author
finding only a four-segmented pleon and poorly developed
pleopod buds. McLaughlin et al. found the pleon to consist
of five 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.
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 first and second pleopods
modified as gonopods, or occasionally only second pleopods
so modified. Males also may be provided with unpaired
pleopods on pleomeres 25 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
modified first pleopods, occasionally asymmetrically paired
second pleopods. Unpaired female pleopods usually occur
only on the left side of the pleon on pleomere 25, but in
certain genera may develop on either side. Unpaired pleopods
are never entirely lost, but their number may decrease to 2
or 3.
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 calcified 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.
The pleon terminates in paired symmetrical or asymmetrical,

biramous uropods, almost always modified 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 first and
second maxillipeds. The elongate pleon typically is composed
of five 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.
Chelipeds are equal, subequal or grossly unequal. Pereopods 2

and 3 are developed as ambulatory legs; pereopod 4 is reduced
in size and modified, 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 flexed, bulbous, or reduced;
entirely membranous, provided with calcified tergites on all
six pleomeres, or with calcification 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).
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 first 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
Adult males may be provided with a pair of gonopores on

the coxae of the fifth 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
10

larval studies has waned, thus another phylogenetic resource
is being underutilized.
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
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 conflict 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
confirm a broad separation of the subfamilies Pylochelinae
and Trizochelinae.
NOTES
A different application of molecular methodology has

provided the first 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, diversification
and even the evolution of coloration. It can only be hoped
that their technique will be applied to other elements of the
paguroid puzzle.
{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 Brandts
(1850) genus. Holmes (1895) established the genus
Acantholithodes for Stimpsons (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 Pacific
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 specifically

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.
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
11

Family Lithodidae Samouelle, 1819
{1, 2}
= 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,
{6}
1842
= 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. MilneEdwards & 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 flindersi Ahyong, 2010
Neolithodes grimaldii (A. Milne-Edwards & Bouvier,
1894) [Lithodes]
{2}
= Lithodes goodei Benedict, 1895
Neolithodes nipponensis Sakai, 1971
Neolithodes vinogradovi Macpherson, 1988
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-fissura 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
= 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}
12

= 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 medipacifica 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 pacifica 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
{2}
Paralomis verrilli (Benedict, 1895) [Pristopus]
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
= Petaloceras White, 1856 (type species Petaloceras
bellianus White, 1856, by monotypy; gender
masculine)
Neolithodes yaldwyni Ahyong & Dawson, 2006

Paralithodes Brandt, 1848
= Paralithodes Brandt, 1848 (type species Lithodes
brevipes H. Milne Edwards & Lucas 1841, by
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 dofleini Balss, 1911
Paralomis echidna Ahyong, 2010
Paralomis elongata Spiridonov, Trkay, 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}
13

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 MilneEdwards 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 Filhols specimens proved to be juveniles
of L. tropicalis, Filhols name had priority and L.
tropicalis became a junior synonym.
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
Rhinolithodes wosnessenskii Brandt, 1848
Sculptolithodes Makarov, 1934
= Sculptolithodes Makarov, 1934 (type species
Sculptolithodes derjugini Makarov, 1934, by
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 Benedicts
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).
Benedicts paper was issued as a separate (1016)
for volume 17 of the Proceedings of the U.S. National
Museum. The separates cover carried the date 1894;
however, according to the table of contents for the
volume, Benedicts paper was actually published on
29 January 1895.
{3}
{4}
Also correctly cited by Dawson (1989), and noted

by Shirley (2002), the spelling of the specific 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
authors 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.
Macpherson (1988b) corrected the error in authorship
of L. ferox made by both Dawson & Yaldwyn (1985)
and Dawson (1989). Those authors attributed the
14
{5}
Among the plates of Sebas (17341765; 18281831)

Locupletissimi Rerum Naturalium Thesauri &
Planches de Seba (Holthuis 1969), was a figure of
a lithodid considered by H. Milne Edwards (1837:
186) to depict the same species as an unidentified
specimen in the collections of the Musum national
dHistoire naturelle. Because the specimen was in
poor condition, and he was unsure of its conspecificity
with the unidentified 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 first 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 Jacquinots 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 (18421844: 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.

{8}
{9}
Davie (2002) cited monotypy as the type designation

for Neolithodes, and the source as A. Milne-Edwards
& Bouvier (1894). Actually, the type was fixed 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 reprsentants de la sous-famille
des Lithodins nous avons pu nous convaincre quil
diffre de ceux-ci par la structure de son abdomen
et par la forme de son rostre; comme ces differences
sont dordre essentiellement gnrique dans la sousfamille, il y a lieu de ranger lespce de lHirondelle
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 Lithodins the second
description was published on 10 July 1894, whereas
the first appeared in print on 1 June 1894.
results of the expedition of the Astrolabe and

Zle, Clark & Crosnier (2000) determined that
Paralomis granulosa (as Lithodes) was first published
in July of 1846 as part of Atlas plate 8. The text of
volume 3 Crustacs was published by Jacquinot and
Lucas in 1854 where the species was cited as Lithodes
granulata. Holthuis (2002) subsequently confirmed
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 Hendersons 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 specific epithet, gowlettholmes, is treated as a
noun in apposition by the author, and is correct as
written (Ahyong, per. comm.).
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.
{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. MilneEdwards & Bouvier (1894a) by Bouvier (1896), an
error repeated by Dawson (1989), and to A. MilneEdwards & 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 first published description
and illustrations were presented by Bouvier (1895); no
mention was made of co-authorship, thus the species
is correctly attributed to Bouvier.
{10} Both authorship and publication date for this species

have been sources of disagreement. Macpherson
(1988b) reemphasized Haigs (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 figures
preceded it, and while no actual date of publication
was specified, Whites (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
SUPERFAMILY LOMISOIDAE BOUVIER, 1895

Family Lomisidae Bouvier, 1895
= Lomisins Bouvier 1894 (not Latin, not valid
name) {1}
15

= Lomisidae Bouvier 1895
= Lomidae Glassner, 1969 (incorrect stem)
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
= Cenobita panamensis Streets, 1871 (incorrect
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
= Coenobita perlatus var. affinis Miers, 1880
Coenobita purpureus Stimpson, 1858
= Cenobita purpurea Stimpson, 1858 (incorrect
Coenobita pseudorugosus Nakasone, 1988
Coenobita rubescens Greeff, 1884
Coenobita rugosus H. Milne Edwards, 1837
= Cenobita rugosa H. Milne Edwards, 1837
= Coenobita subrugosa Neumann, 1878
= Coenobita rugosa var. wagneri Doflein, 1900
Coenobita scaevola (Forskl, 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
= 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
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,
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}
= Conbite Latreille, 1825 (not Latin, unavailable
name)
= Cenobites Berthold 1827 (type species Pagurus
clypeatus Fabricius, 1787, by monotypy)
{4}
= Cenobita H. Milne Edwards 1837 (incorrect
Coenobita brevimanus Dana, 1852
= Cenobita clypeata var. brevimanus Dana, 1852
= Coenobita hilgendorfi Terao 1913 (unnecessary
replacement name)
{5}
= Coenobita clypeata var. puerto-galerae YapChiongco, 1938
Coenobita carnescens Dana, 1851
= Cenobita carnescens Dana, 1851 (incorrect
NOTES
{1}
16
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.

{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
specific 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 suffix 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 Jarockis
(1825) Carcinion was a senior objective synonym, it
was an entirely forgotten name whose reintroduction
would cause considerable undesirable confusion; 2)
Bertholds (1827) Cenobites was simply a translation
of Latreilles (1825) vernacular Cnobite and its
reintroduction into carcinological literature would
result in great confusion. Osbecks (1765) Eremita
posed a somewhat different problem. Despite the
generalities of Osbecks description, which could
justify Eremitas 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 Official List of Valid Names in
Zoology, and the names Carcinion, Cenobites and
Eremita with its type species E. javanick Osbeck,
1765, on the Official 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 Hilgendorfs authority we learn
that Cancer clypeatus Herbst (1791) is not identical
with Latreilles 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 specific name. However, Teraos
(1913) interpretation of C. clypeatus was based on a
misidentification 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).
17
{7}
Catesbys initial publication is usually cited with the

date 1754, which is the date of the second edition.
Catesbys (1743, pl. 33, figs. 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 Herbsts (1791), at the time
unpublished, illustrations of two specimens reported
to be from the East Indies. Subsequent carcinologists
incorrectly assumed that both figures represented
the same specimen, one simply an enlargement. In
a review Herbsts collection in the Naturhistorisches
Forschungsinstitut Museum fr 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
Herbsts (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-Pacific taxa, under the specific 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 fixations 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 Official List of Specific Names
in Zoology, with an endorsement recording that the
species is defined by its designated neotype. Cenobita
rugosus H. Milne Edwards, 1837, and Cenobita
violascens were similarly added to the Official List.
{9}
In his list of Crustacea in the British Museum, Adam

White (1847: 5962) listed as new species, Coenobita
compta from the collection of Mr. Broderip, and

Coenobita laeviuscula from Port Essington. Neither
taxon was described nor illustrated and mention
of neither could be found in any of Whites other
publications.
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 pacificus Forest, 1993
Bathynarius wolffi Forest, 1993
{3}
Calcinus Dana, 1851
= 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
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 (Dechanc, 1963)
[Paguristes]
Areopaguristes breviantennatus (Rahayu, 2005)
[Stratiotes]
Areopaguristes cyanops (Forest, 1978) [Paguristes]
Areopaguristes difficilis (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 (Ayn Parente & Hendrickx,
2006) [Stratiotes]
Areopaguristes micheleae (Rahayu, 2005) [Stratiotes]
Areopaguristes ngochoae (Rahayu, 2005) [Stratiotes]
Areopaguristes nigroapiculus (Komai, 2009)
[Stratiotes]
Areopaguristes orbis (Komai, 2009) [Stratiotes]
18

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
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 pacificus 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 YapChiongco, 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
19

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 pacificus 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 Paulson, 1875
{21}
= Dardanus Paulson, 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 Ayn Parente & Hendrickx,
2009
Dardanus lagopodes (Forskl, 1775) [Cancer]
= Pagurus sanguinolentus Quoy & Gaimard, 1824
= Pagurus affinis H. Milne Edwards, 1836
= Pagurus euopsis Dana, 1852
= Pagurus depressus Heller, 1861
= Dardanus helleri Paulson, 1875
Dardanus longior Asakura, 2006
Dardanus magdalenensis Ayn Parente & Hendrickx,
2009
Dardanus megistos (Herbst, 1804) [Cancer]
= Pagurus punctulatus Miers, 1879
= Dardanus jordani Schmitt, 1921{25}
Dardanus nudus Ayn 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 Ayn 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
20

Dardanus setifer (H. Milne Edwards, 1836) [Pagurus]
Dardanus sinistripes (Stimpson, 1859) [Pagurus]
= Pagurus peruensis Balss, 1921
Dardanus squarrosus Cook, 1989
Dardanus stimpsoni Ayn Parente & Hendrickx, 2009
Dardanus sulcatus Edmondson, 1925
Dardanus tinctor (Forskl, 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 Dechanc, 1964
Diogenes custos (Fabricius, 1798) [Pagurus]
= Diogenes affinis Henderson, 1893
Diogenes deflectomanus 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
D i o g e n e s m a n a a re n s i s ( H e n d e r s o n , 1 8 9 3 )
[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 Clment, 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)
21

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 Manjn-Cabeza, Garca Raso &
Martnez 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 & Gmez,
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
= Holopagurus Holmes, 1900 (type species

Holopagurus pilosus Holmes, 1900, by original
Isocheles aequimanus (Dana, 1852) [Bernhardus]
Isocheles pacificus Bouvier, 1907
= Isocheles Wurdemanni var. pacificus 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 Lrenthey & 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 (Gurin, 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
22

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
Paguropsis typica Henderson, 1888
{32}
= P a g u ro p s i s a n d e r s o n i ( A l c o c k , 1 8 9 9 )
[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
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
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. MilneEdwards & Bouvier, 1892
= Paguristes oculatus var. brunneo-pictus A. MilneEdwards & 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
23

was Forest & McLaughlin (2000) that finally placed
the two taxa in synonymy. However, when Rahayu
(2005) found the gill number of Paguristes species
in the collection of the Musum national dHistoire
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.
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
Trizopagurus melitai (Chevreux & Bouvier 1892)
[Clibanarius]
Trizopagurus magnificus (Bouvier, 1898) [Clibanarius]
= Clibanarius chetyrkini Boone, 1932
Trizopagurus rubrocinctus Forest & Garcia Raso, 1990
{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 Ingles (1993) opinion that it
should be regarded incerta sedis. It is entered in this
checklist as a questionable synonym.
{6}
Although Morgans (1988) Australian Calcinus

abrolhensis proved to be conspecific 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, Stimpsons 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 Herbsts type locality for
another presumed western Atlantic species, Coenobita
clypeatus Herbst. Thus is possible that further
examination of Herbsts material will show Calcinus
sulcatus is a species distinct from C. tibicen.
Incerta sedis
Pagurus lar Heller, 1862
NOTES
{1}
{2}
Volume 8 of Oliviers (18111812) Encyclopdie

Methodique (Dictionnaire Encyclopdique
Mthodique) 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 361722 and containing the
Pagures) was not published until 1812.
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 Thomsons (1899) Stratiotes setosus a
synonym of H. Milne Edwards Paguristes setosus, it
24

{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 Officical Index of Rejected
and Invalid Generic Names in Zoology. However,
Milne Edwards species had been transferred from
Pagurus to Aniculus, then Trizopagurus, and finally
to Ciliopagurus (Forest, 1995), and was never
considered conspecific with Stimpsons (1860) taxon.
Consequently, no replacement name was required.
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
Kobjakovas (1955) Clibanarius sachalinicus are
uncertain.
{20} Clibanarius willeyi Southwell, 1910 appears to have
been overlooked. It was not mentioned by Fize and
Serne (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.
{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
Stimpsons name.
{21} As previously mentioned, although Paulson (1875)

proposed the genus Dardanus for one of the species
assigned to Pagurus (sensu Dana), Paulsons
genus soon thereafter was placed in synonymy with
Danas Pagurus. It was not until considerably later
that Benedict (1896) realized that Danas 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 Benedicts 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 Paulson were placed on the
Official Index of Valid Names in Zoology.
{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 Pacific.
However Fize & Serne (1955) list C. tuberculosus
(H. Milne Edwards, 1848) among their AtlantoMediterranean species of Clibanarius. Clibanarius
sclopetarius is a western Atlantic species.
{14} The status and validity of Kobjakovas (1971)
Clibanarius hirsutimanus are uncertain.
{15} Rahayu & Forest (1992) determined that Clibanarius
lineatus was not the Indo-Pacific 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).
{22} Sakai (1999) suggested that Herbsts (1791) Cancer

hungarus might be conspecific with Dardanus
calidus; however, the type apparently is no longer
extant, and Herbst brief description is inadequate
for confirmation. It is questionably included as a
synonym.
{16} Originally described as a subspecies of Clibanarius

aequabilis Dana, 1851, De Mans Clibanarius
aequabilis merguiensis was elevated to full specific
rank by Forest (1953) when he determined that C.
aequabilis was an Atlantic species.
{23} Initially described as a distinct species, Glaucothoe

carinata subsequently was shown to be the megalopal
stage of a Dardanus species. Its true specific identity
is not known with certainty, thus it is included
only questionably in the synonymy of Dardanus
deformis.
{17} Clibanarius pacificus 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 fitting Stimpsons
(1858) description.
{24} McLaughlin et al. (2005: 303) pointed out that

Boones 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.
{18} In their discussion of Clibanarius rosewateri Manning

and Chace (1990: 37) used the manuscript name C.
25

{25} In a study in progress, Manuel Ayon Parente has found
evidence to indicate that Schmitts (1921) Dardanus
jordani is conspecific with Dardanus megistos.
{31} Studies in progress by one of the current authors (TK)

indicate that Paguristes sinensis Tung & Wang, 1966,
may be transferred to the genus Areopaguristes.
{26} Gordan (1956) gave the date for Pagurus arrosor var.
divergens as 1905 and the pagination as 123145. De
Melo (1999) gave the date as 1905 and pagination as
125; Forest & de Saint Laurent (1968) and Biffar
& Provenzano (1972) gave the same pagination,
but a date of 1906. The correct date is 1905 and the
pagination 123145.
{32} Henderson (1888) incorrectly used the masculine

ending us in his original description of Paguropsis
typica.
{33} One of the authors (TK) has examined specimens from
Kerara, India and found that Paguropsis andersoni
(Alcock, 1905b) is not a synonym of P. typica and
should be reinstated.
{27} As pointed out by McLaughlin & Holthuis (2001),

although Dana (1851) specified Pagurus miles as the
type of the genus Diogenes, Danas species was not
P. miles of Fabricius (1787). The species had been
misidentified by both H. Milne Edwards (1836) and
Dana (1851, 1852a, c). Their P. miles was actually
Diogenes merguiensis De Man, 1888. While the
third edition of the ICZN Code required that such a
misidentification be referred to the Commission, the
fourth edition (1999) gave the revisers the option to fix
the status of the type species. McLaughlin & Holthuis
(2001) were of the opinion that the best interests of
stability in nomenclature were served by retaining
Diogenes miles (Fabricius, 1787) [not Diogenes miles
sensu H. Milne Edwards, 1836; Dana, 1851, 1852a,
c] as the type species of the genus Diogenes.
{34} Holthuis (1959) demonstrated that while this species

was best known as Petrochirus granulatus or P.
bahamensis, the correct name was P. diogenes,
described by Linnaeus (1758) as Cancer diogenes.
Linnaeus (1758) Cancer diogenes actually was
a composite taxon, based on the mirror image
reproductions of Rumphius (1705) [1745] illustrations
of an Indo-Pacific species of Coenobita and Catesbys
(1743) description and figure of Cancellus maximus
Bahamensis. Holthuis (1959) selected Catesbys
figured specimen as the lectotype of Cancer diogenes.
Herbsts (1791) Cancer bahamensis was based on
Catesbys same figured specimen, thus the two species
were objective synonyms. Holthuis (1959) noted that
the specific name diogenes Linnaeus had not been
in use for the past 40 years, so its reintroduction for
the American Petrochirus species would not result
in confusion. Herbsts (1791) species, bahamensis,
although senior, at the time was better known by
the specific name granulatus. Therefore Holthuis
(1959) found no reason not to strictly follow in Rule
of Priority.
{28} Holthuis (1959) determined that C. diogenes had

been based on two distinct species, one the western
Atlantic Petrochirus and the other an Indo-Pacific
species of Coenobita. Holthuis designated the
lectotype of Linnaeus (1758) composite taxon the
specimen figured by Catesby (1743) of his Cancellus
maximus Bahamensis, an invalid name for the taxon
known as Petrochirus bahamensis (Herbst, 1791).
The species reported in the literature as Pagurus
diogenes Fabricius, 1787, Cancer diogenes Herbst,
1791 and Diogenes diogenes of subsequent authors
was based on misidentifications of Linnaeus (1758)
Cancer diogenes and was replaced by Diogenes alias
McLaughlin & Holthuis, 2001.
Family Paguridae Latreille, 1802

Acanthopagurus de Saint Laurent, 1968
= Acanthopagurus de Saint Laurent, 1968 (type
species Anapagurus? dubius A. Milne-Edwards
& Bouvier, 1900, by original designation; gender
masculine)
{1}
Acanthopagurus dubius (A. Milne-Edwards & Bouvier,
1900) [Anapagurus]
= Catapaguroides macrophthalmus Bouvier, 1922
Agaricochirus McLaughlin, 1981
= Agaricochirus McLaughlin, 1981 (type species
Pylopagurus boletifer A. Milne-Edwards &
Bouvier, 1893, by original designation; gender
masculine)
Agaricochirus acanthinus McLaughlin, 1982
Agaricochirus alexandri (A. Milne-Edwards & Bouvier,
1893) [Pylopagurus]
Agaricochirus boletifer (A. Milne-Edwards & Bouvier,
1893) [Pylopagurus]
Agaricochirus cavimanus (Chace, 1939) [Pylopagurus]
Agaricochirus echinatus McLaughlin, 1982
{29} This species was credited to Costa (1838) by Hope

(1851) who provided a description. No description
could be found in the portion of Costas paper
available. Diogenes serripes (Costa, 1838) is the name
and description also given by Carus (1885). The type
locality is reported as Naples. Although D. serripes
has not been cited in any recent synonymy, it most
probably is another name for D. pugilator.
{30} Haig et al. (1970) considered P. holmesi a synonym
of P. bakeri. However, Glassells (1937) specimens
can no longer be found, and Hendrickx & Harvey
(1999) believe some question exists as to the accuracy
of the synonymy. It is included only questionably in
the synonymy of P. bakeri.
26

Agaricochirus erosus (A. Milne-Edwards, 1880)
[Eupagurus]
Agaricochirus gibbosimanus (A. Milne-Edwards, 1880)
[Eupagurus]
Agaricochirus hispidus (Benedict, 1892) [Eupagurus]
Alainopaguroides McLaughlin, 1997
= Alainopaguroides McLaughlin, 1997 (type species
Alainopaguroides lemaitrei McLaughlin, 1997, by
Alainopaguroides andamanensis McLaughlin, 2002
Alainopaguroides lemaitrei McLaughlin, 1997
Alainopaguroides megalophthalmus McLaughlin, 2006
Alainopagurus Lemaitre & McLaughlin, 1995
= Alainopagurus Lemaitre & McLaughlin, 1995
(type species Alainopagurus crosnieri Lemaitre &
McLaughlin, 1995 by original designation; gender
masculine)
Alainopagurus crosnieri Lemaitre & McLaughlin,
1995
Alloeopagurodes Komai, 1998
= Alloeopagurodes Komai, 1998 (type species
Alloeopagurodes spiniacicula Komai, 1998, by
Alloeopagurodes spiniacicula Komai, 1998
Anapagrides de Saint Laurent-Dechanc, 1966
= Anapagrides de Saint Laurent-Dechanc, 1966
[type species Eupagurus (Spiropagurus) facetus
Melin, 1939, by original designation; gender
masculine]
= Nanopagurus McLaughlin, 1986 (type species
Nanopagurus reesei McLaughlin, 1986, by
Anapagrides aequalis Komai, 1999
Anapagrides facetus (Melin, 1939) [Eupagurus
(Spiropagurus)]
A n a p a g r i d e s re e s e i ( M c L a u g h l i n , 1 9 8 6 )
[Nanopagurus]
Anapagurus Henderson, 1886
= Anapagurus Henderson, 1886 [type species Pagurus
laevis Bell, 1846, by subsequent designation by
Holthuis (1962); gender masculine]
Anapagurus adriaticus Garca-Gmez, 1994
Anapagurus alboranensis Garca-Gmez, 1994
Anapagurus atlantidii Garca-Gmez, 1994
Anapagurus bicorniger A. Milne-Edwards & Bouvier,
1892
Anapagurus bonnieri Nobili, 1905
Anapagurus breviaculeatus Fenizia, 1937
Anapagurus chiroacanthus (Lilljeborg, 1856)
[Pagurus]
= Anapagurus brevicarpus A. Milne-Edwards &
Bouvier, 1892
= Pagurus ferrugineus Norman, 1861
= Anapagurus chiroacanthus var. cristatus Fenizia,
1937
= Anapagurus chiroacanthus var. gracilis Fenizia,
1937
Anapagurus congolensis Garca-Gmez, 1994
Anapagurus curvidactylus Chevreux & Bouvier, 1892
Anapagurus hendersoni Barnard, 1947
Anapagurus hyndmanni (Bell, 1846) [Pagurus]

{2}
= Pagurus Hyndmanni Thompson, 1844 (nomen
nudum)
Anapagurus japonicus Ortmann, 1892
= Anapagurus pusillus var. japonicus Ortmann,
1892
Anapagurus laevis (Bell, 1846) [Pagurus]
{2}
= Pagurus laevis Thompson, 1844 (nomen
nudum)
Anapagurus longispina A. Milne-Edwards & Bouvier,
1900
= Anapagurus laevis var. longispina A. MilneEdwards & Bouvier, 1900
Anapagurus petiti Dechanc & Forest, 1962
Anapagurus smythi Ingle, 1993
Anapagurus vossi Garca-Gmez, 1994
Anapagurus wolffi Forest, 1961
Anisopagurus McLaughlin, 1981
= Anisopagurus McLaughlin, 1981 (type species
Eupagurus bartletti A. Milne-Edwards, 1880, by
Anisopagurus actinophorus Lemaitre & McLaughlin,
1996
Anisopagurus bartletti (A. Milne-Edwards, 1880)
[Eupagurus]
Anisopagurus hopkinsi Lemaitre & McLaughlin, 1996
Anisopagurus pygmaeus (Bouvier, 1918) [Eupagurus]
Anisopagurus vossi Lemaitre & McLaughlin, 1996
Bathypaguropsis McLaughlin, 1994
= Bathypaguropsis McLaughlin, 1994 (type species
Bathypaguropsis yaldwyni McLaughlin, 1994, by
Bathypaguropsis carinatus Komai & Takeda, 2004
Bathypaguropsis cruentus de Saint Laurent & McLaughlin,
2000
Bathypaguropsis foresti Komai & Lemaitre, 2002
Bathypaguropsis kuroshioensis (Miyake, 1978)
[Pagurus]
= Bathypaguropsis rahayuae McLaughlin, 1997
Bathypaguropsis marionensis McLaughlin, 1994
Bathypaguropsis microps (Balss, 1911) [Eupagurus]
Bathypaguropsis yaldwyni McLaughlin, 1994
Boninpagurus Asakura & Tachikawa, 2004
= Boninpagurus Asakura & Tachikawa, 2004 (type
species Boninpagurus acanthocheles Asakura &
Tachikawa, 2004, by original designation; gender
masculine)
Boninpagurus acanthocheles Asakura & Tachikawa,
2004
{3}
Bythiopagurus McLaughlin, 2003
= Bythiopagurus McLaughlin, 2003 (type species
Bythiopagurus macrocolus McLaughlin, 2003 by
Bythiopagurus macrocolus McLaughlin, 2003
Catapaguroides A. Milne-Edwards & Bouvier, 1892
= Catapaguroides A. Milne-Edwards & Bouvier,
1892 [type species Catapaguroides microps A.
Milne-Edwards & Bouvier, 1892, by subsequent
selection by Holthuis (1962); gender masculine]
27

Catapaguroides cristimanus de Saint Laurent, 1968
Catapaguroides declivis McLaughlin, 1997
Catapaguroides foresti McLaughlin, 2002
Catapaguroides fragilis (Melin, 1939) [Eupagurus
(Catapagurus)]
Catapaguroides hooveri McLaughlin & Pittman, 2002
Catapaguroides iejimensis Osawa & Takeda, 2004
Catapaguroides inermis de Saint Laurent, 1968
Catapaguroides iris Bouvier, 1922
Catapaguroides japonicus de Saint Laurent, 1968
Catapaguroides karubar McLaughlin, 1997
Catapaguroides kasei Osawa & Takeda, 2004
{4}
Catapaguroides megalops A. Milne-Edwards & Bouvier,
1892
Catapaguroides melini de Saint Laurent, 1968
Catapaguroides microps A. Milne-Edwards & Bouvier,
1892
{5}
Catapaguroides mortenseni de Saint Laurent, 1968
C a t a p a g u ro i d e s o l f a c i e n s ( A l c o c k , 1 9 0 5 )
[Cestopagurus]
Catapaguroides pectinipes (Lewinsohn, 1969)
[Cestopagurus]
Catapaguroides setosus Edmondson, 1951
{6}
Catapaguroides spinulimanus de Saint Laurent, 1968
Catapaguroides umbra Komai, 2009
Catapaguropsis Lemaitre & McLaughlin, 2006
= Catapaguropsis Lemaitre & McLaughlin, 2006
(type species Catapaguropsis queenslandica
Lemaitre & McLaughlin, 2006, by original
Catapaguropsis queenslandica Lemaitre & McLaughlin,
2006
Catapaguropsis brucei McLaughlin & Lemaitre, 2007
Catapagurus A. Milne-Edwards, 1880
{7}
= Catapagurus A. Milne-Edwards, 1880 (type
species Catapagurus sharreri A. Milne-Edwards,
= Hemipagurus Smith1881 [type species
Hemipagurus gracilis Smith, 1881, by subsequent
designation by Asakura (2001); gender
masculine]
{8}
C a t a p a g u r u s a l b a t ro s s a e ( A s a k u r a , 2 0 0 1 )
[Hemipagurus]
Catapagurus alcocki McLaughlin, in Hogarth et al.,
1998
{9}
Catapagurus danida McLaughlin, 2002
Catapagurus ensifer Henderson, 1893
Catapagurus franklinae McLaughlin, 2004
Catapagurus gracilis (Smith, 1881) [Hemipagurus]
Catapagurus gracilis var. intermedius A. Milne-Edwards
& Bouvier, 1893
{10}
Catapagurus granulatus Edmondson, 1951
Catapagurus haigae (Asakura, 2001) [Hemipagurus]
Catapagurus hirayamai (Asakura, 2001)
[Hemipagurus]
Catapagurus holthuisi McLaughlin, 1997
Catapagurus imperialis (Asakura, 2001)
[Hemipagurus]
{11}
Catapagurus insolitus Komai & Osawa, 2009
Catapagurus kosugei (Asakura, 2001) [Hemipagurus]
Catapagurus lewinsohni (Asakura, 2001)

[Hemipagurus]
Catapagurus maclaughlinae (Asakura, 2001)
[Hemipagurus]
{9}
Catapagurus misakiensis Terao, 1914
= Catapagurus japonicus Yokoya, 1933
Catapagurus oculocrassus McLaughlin, 1997
{11}
Catapagurus sharreri A. Milne-Edwards, 1880 {12}
= Hemipagurus socialis Smith, 1881
= Catapagurus socialis (Smith, 1882)
= Catapagurus Sparreri Verrill, 1883 (misspelling
of sharreri)
= Catapagurus sharrei Abele & Kim, 1986
(misspelling of sharreri)
Catapagurus spinicarpus de Saint Laurent & McLaughlin,
2000
Catapagurus tanimbarensis McLaughlin, 1997
Catapagurus toyoshioae (Asakura, 2001)
[Hemipagurus]
C a t a p a g u r u s t u b e rc u l o s u s ( A s a k u r a , 1 9 9 9 )
[Icelopagurus]
{7}
Catapagurus sp. 1
Catapagurus sp. 2
{7}
Catapagurus sp. 3
{7}
Catapagurus sp. 4
{7}
Catapagurus sp. 5
{7}
Ceratopagurus Yokoya, 1933
{13}
= Ceratopagurus Yokoya, 1933 (type species
Ceratopagurus pilosimanus Yokoya, 1933, by
Ceratopagurus pilosimanus Yokoya, 1933
Cestopagurus Bouvier, 1897
= Cestopagurus Bouvier, 1897 (type species
Cestopagurus coutieri Bouvier, 1897, by
Cestopagurus coutieri Bouvier, 1897
Cestopagurus puniceus Komai & Takeda, 2005
Cestopagurus timidus (Roux, 1830) [Pagurus]
= Catapaguroides acanthodes Fenizia, 1935
= Catapaguroides acutifrons A. Milne-Edwards &
Bouvier, 1892
= Catapaguroides crassipes Fenizia, 1935
= Catapaguroides crassipes var. tuberculatus
Fenizia, 1937
= Catapaguroides macrochirus Fenizia, 1937
= Catapaguroides macrochirus var. platichelus
Fenizia, 1937
= Catapaguroides timidus var. neapolitanus Fenizia,
1937
Chanopagurus Lemaitre, 2003
= Chanopagurus Lemaitre, 2003 (type species
Chanopagurus atopos Lemaitre, 2003, by original
Chanopagurus atopos Lemaitre, 2003
Cycetopagurus McLaughlin, 2004
= Cycetopagurus McLaughlin, 2004 (type species
Cycetopagurus morgani McLaughlin, 2004, by
Cycetopagurus morgani McLaughlin, 2004
28

Decaphyllus de Saint Laurent, 1968
= Decaphyllus de Saint Laurent, 1968 (type species
Decaphyllus spinicornis de Saint Laurent, 1968,
by original designation; gender feminine)
Decaphyllus barunajaya McLaughlin, 1997
Decaphyllus junquai de Saint Laurent, 1968
Decaphyllus maci McLaughlin, 1997
Decaphyllus similis de Saint Laurent, 1968
Decaphyllus spinicornis de Saint Laurent, 1968
Dentalopagurus McLaughlin, 2007
= Dentalopagurus McLaughlin, 2007 (type species
Dentalopagurus levii McLaughlin, 2007, by
Dentalopagurus levii McLaughlin, 2007
Dentalopagurus rutilocinctus Osawa & Chan, 2008
Diacanthurus McLaughlin & Forest, 1997
= Diacanthurus McLaughlin & Forest, 1997 (type
species Eupagurus spinulimanus Miers, 1876, by
Diacanthurus ecphyma McLaughlin & Forest, 1997
Diacanthurus ophthalmicus (Ortmann, 1892)
[Eupagurus]
Diacanthurus richeri McLaughlin & Forest, 1997
Diacanthurus rubricatus (Henderson, 1888)
[Eupagurus]
Diacanthurus spinulimanus (Miers, 1876) [Eupagurus]
= Eupagurus edwardsii Filhol, 1883
= Eupagurus intermedius Lenz, 1901
= Eupagurus norae Chilton, 1911
= Eupagurus chiltoni E. F. Thompson, 1930
Discorsopagurus McLaughlin, 1974
= Discorsopagurus McLaughlin, 1974 (type species
Pylopagurus schmitti Stevens, 1925, by original
Discorsopagurus cavicola Komai & Takeda, 1996
Discorsopagurus maclaughlinae Komai, 1995
Discorsopagurus schmitti (Stevens, 1925)
[Pylopagurus]
= Phylopagurus schmitti Hart 1930 (misspelling of
Pylopagurus)
Discorsopagurus tubicola Komai, 2003
Elassochirus Benedict, 1892
= Eupagurus (Elassochirus) Benedict, 1892 (type
species Bernhardus tenuimanus Dana, 1851, by
Elassochirus cavimanus (Miers, 1879) [Eupagurus]
= Eupagurus gotoi Terao, 1913
= Eupagurus (Elassochirus) munitus Benedict,
1892
Elassochirus gilli (Benedict, 1892) [Eupagurus]
= Eupagurus porcellanus Molander, 1914
Elassochirus tenuimanus (Dana, 1851) [Bernhardus]
Enallopaguropsis McLaughlin, 1981
= Enallopaguropsis McLaughlin, 1981 (type species
Pylopagurus guatemoci Glassell, 1937, by original
Enallopaguropsis guatemoci (Glassell, 1937)
[Pylopagurus]
= Pylopagurus hancocki Walton, 1954
Enallopaguropsis janetae McLaughlin, 1982

Enallopaguropsis williamsi Lemaitre & McLaughlin,
2003
Enallopagurus McLaughlin, 1981
= Enallopagurus McLaughlin, 1981(type species
Pylopagurus spinicarpus Glassell, 1938, by
Enallopagurus affinis (Faxon, 1893) [Pylopagurus]
E n a l l o p a g u r u s c o ro n a t u s ( B e n e d i c t , 1 8 9 2 )
[Eupagurus]
Enallopagurus provenzanoi Lemaitre & McLaughlin,
2003
Enallopagurus spinicarpus (Glassell, 1938)
[Pylopagurus]
Enneobranchus Garca-Gmez, 1988
= Enneobranchus Garca-Gmez, 1988 (type species
Enneobranchus flavioculatus Garca-Gmez, 1988,
Enneobranchus bermudensis Garca-Gmez, 1988
Enneobranchus flavioculatus Garca-Gmez, 1988
Enneobranchus markhami Garca-Gmez, 1988
Enneopagurus McLaughlin, 1997
= Enneopagurus McLaughlin, 1997 (type species
Enneopagurus garciagomezi McLaughlin, 1997,
Enneopagurus garciagomezi McLaughlin, 1997
Enneophyllus McLaughlin, 1997
= Enneophyllus McLaughlin, 1997 (type species
Enneophyllus spinirostris McLaughlin, 1997 by
Enneophyllus spinirostris McLaughlin, 1997
Forestopagurus Garca-Gmez, 1995
= Forestopagurus Garca-Gmez, 1995 (type species
Anapagurus drachi Forest, 1966, by original
Forestopagurus drachi (Forest, 1966) [Anapagurus]
Goreopagurus McLaughlin, 1988
= Goreopagurus McLaughlin, 1988 (type species
Pagurus piercei Wass, 1963 by original designation;
gender masculine)
Goreopagurus garthi McLaughlin & Haig, 1995
Goreopagurus piercei (Wass, 1963) [Pagurus]
Goreopagurus poorei Lemaitre & McLaughlin, 2003
Goreopagurus lemaitrei Nucci & de Melo, 2007
Hachijopagurus Osawa & Okuno, 2003
= Hachijopagurus Osawa & Okuno, 2003 (type
species Hachijopagurus rubrimaculatus Osawa
& Okuno, 2003, by original designation; gender
masculine)
Hachijopagurus rubrimaculatus Osawa & Okuno,
2003
Haigiopagurus McLaughlin, 2005
= Haigia McLaughlin, 1981(type species Pylopagurus
diegensis Scanland & Hopkins, 1969 by original
designation; gender masculine) (name preoccupied
by Haigia Steyskal, 1961, Insecta)
= Haigiopagurus McLaughlin, 2005 (replacement
name)
Haigiopagurus diegensis (Scanland & Hopkins, 1969)
[Pylopagurus]
29

Icelopagurus McLaughlin, 1997
= Icelopagurus McLaughlin, 1997 (type species
Icelopagurus crosnieri McLaughlin, 1997, by
Icelopagurus crosnieri McLaughlin, 1997
Icelopagurus undulatus McLaughlin, 2006
Iridopagurus de Saint Laurent-Dechanc, 1966
= Iridopagurus de Saint Laurent-Dechanc, 1966
(type species Spiropagurus iris A. MilneEdwards, 1880, by original designation; gender
masculine)
Iridopagurus caribbensis (A. Milne-Edwards & Bouvier,
1893) [Spiropagurus]
Iridopagurus dispar (Stimpson, 1859) [Spiropagurus]
= Spiropagurus dispar Stimpson, 1858 (nomen
nudum)
Iridopagurus globulus de Saint Laurent-Dechanc,
1966
Iridopagurus haigae Garca-Gmez, 1983
Iridopagurus iris (A. Milne-Edwards, 1880)
[Spiropagurus]
Iridopagurus margaritensis Garca-Gmez, 1983
Iridopagurus occidentalis (Faxon, 1893)
[Spiropagurus]
Iridopagurus reticulatus Garca-Gmez, 1983
Iridopagurus violaceus de Saint Laurent-Dechanc,
1966
Labidochirus Benedict, 1892
= Eupagurus Labidochirus Benedict, 1892 (type
species Pagurus splendescens Owen, 1839, by
= Libidochirus Benedict, 1892 (misspelling of
Labidochirus)
Labidochirus anomalus (Balss, 1913) [Eupagurus]
Labidochirus splendescens (Owen, 1839) [Pagurus]
Lithopagurus Provenzano, 1968
= Lithopagurus Provenzano, 1968 (type species
Lithopagurus yucatanicus Provenzano, 1968, by
Lithopagurus boucheti McLaughlin & Lemaitre, 2004
Lithopagurus tribulomanus McLaughlin & Lemaitre,
2004
Lithopagurus yucatanicus Provenzano, 1968
Lophopagurus (Australeremus) McLaughlin, 1981
= Australeremus McLaughlin, 1981 (type species
Eupagurus cookii Filhol, 1883, by original
Lophopagurus (Australeremus) cookii (Filhol, 1883)
[Eupagurus]
Lophopagurus (Australeremus) cristatus (H. Milne
Edwards, 1836) [Pagurus]
Lophopagurus (Australeremus) eltaninae (McLaughlin
& Gunn, 1992) [Australeremus]
Lophopagurus (Australeremus) indonesiensis
(McLaughlin, 1997) [Australeremus]
{14}
Lophopagurus (Australeremus) kirkii (Filhol, 1883)
[Eupagurus]
Lophopagurus (Australeremus) laurentae (McLaughlin
& Gunn, 1992) [Australeremus]
Lophopagurus (Australeremus) stewarti (Filhol, 1883)

[Eupagurus]
Lophopagurus (Australeremus) triserratus (Ortmann,
1892) [Eupagurus]
= Pylopagurus serpulophilus Miyake, 1978
Lophopagurus (Lophopagurus) McLaughlin, 1981
= Lophopagurus McLaughlin, 1981 (type species
Eupagurus thompsoni Filhol, 1885, by original
Lophopagurus (Lophopagurus) foresti McLaughlin &
Gunn, 1992
Lophopagurus (Lophopagurus) lacertosus (Henderson,
1888) [Eupagurus]
{15}
= Eupagurus crenatus Borradaile, 1916
Lophopagurus (Lophopagurus) nanus (Henderson, 1888)
[Eupagurus]
Lophopagurus (Lophopagurus) nodulosus McLaughlin
& Gunn, 1992
Lophopagurus (Lophopagurus) pumilus de Saint Laurent
& McLaughlin, 2000
Lophopagurus (Lophopagurus) thompsoni (Filhol, 1885)
[Eupagurus]
= Eupagurus thomsoni Thomson, 1898 (misspelling
of thompsoni)
Manucomplanus McLaughlin, 1981
= Manucomplanus McLaughlin, 1981 (type species
Eupagurus ungulatus Studer, 1883, by original
Manucomplanus cervicornis (Benedict, 1892)
[Eupagurus]
Manucomplanus longimanus (Faxon, 1893)
[Pylopagurus]
Manucomplanus spinulosus (Holthuis, 1959)
[Pylopagurus]
= Pagurus impressus zilchi Trkay, 1968
Manucomplanus ungulatus (Studer, 1883) [Eupagurus]
= Eupagurus corallinus Benedict, 1892
Manucomplanus varians (Benedict, 1892) [Eupagurus]
Michelopagurus McLaughlin, 1997
= Michelopagurus McLaughlin, 1997 (type species
Pagurodes limatulus Henderson, 1888, by original
Michelopagurus atlanticus (Bouvier, 1922)
[Pagurodes]
Michelopagurus chacei McLaughlin, 1997
Michelopagurus limatulus (Henderson, 1888)
[Pagurodes]
Michelopagurus richardi (Bouvier, 1922) [Pagurodes]
Micropagurus McLaughlin, 1986
= Micropagurus McLaughlin, 1986 (type species
Micropagurus devaneyi McLaughlin, 1986, by
Micropagurus n. sp. Lemaitre, 2010 (in press)
Micropagurus acantholepis (Stimpson, 1858)
[Eupagurus]
= Anapagurus australiensis Henderson, 1888
Micropagurus devaneyi McLaughlin, 1986
M i c ro p a g u r u s p o l y n e s i e n s i s ( N o b i l i , 1 9 0 6 )
[Anapagurus]
= Micropagurus vexatus Haig & Ball, 1988
30

Micropagurus propinquus Asakura, 2005
Micropagurus spinimanus Asakura, 2005
Munidopagurus A. Milne-Edwards & Bouvier, 1893
= Munidopagurus A. Milne Edwards & Bouvier,
1893 (type species Eupagurus macrocheles A.
Milne-Edwards, 1880, by monotypy; genus
masculine)
Munidopagurus macrocheles (A. Milne-Edwards, 1880)
[Eupagurus]
Nematopaguroides Forest & de Saint Laurent, 1968
= Nematopaguroides Forest & de Saint Laurent, 1968
(type species Nematopaguroides fagei Forest &
de Saint Laurent, 1968, by original designation;
gender masculine)
Nematopaguroides fagei Forest & de Saint Laurent,
1968
Nematopaguroides pusillus Forest & de Saint Laurent,
1968
Nematopagurus A. Milne-Edwards & Bouvier, 1892
= Nematopagurus A. Milne-Edwards & Bouvier,
1892 (type species Nematopagurus longicornis A.
Milne-Edwards & Bouvier, 1892, by monotypy;
gender masculine)
Nematopagurus alcocki McLaughlin, 1997
Nematopagurus australis (Henderson, 1888)
[Catapagurus]
Nematopagurus chanani McLaughlin, 2004
Nematopagurus chauseyensis McLaughlin, 1998
Nematopagurus crosnieri McLaughlin, 1998
Nematopagurus diadema Lewinsohn, 1969
Nematopagurus gardineri Alcock, 1905
= Nematopagurus holthuisi McLaughlin & Hogarth,
1998
= Nematopagurus pilosus Komai, 1999
Nematopagurus helleri (Balss, 1916) [Cestopagurus]
Nematopagurus indicus Alcock, 1905
Nematopagurus jacquesi McLaughlin, 2004
Nematopagurus kosiensis McLaughlin, 1998
= Nematopagurus shinnyoae Komai, 1999
Nematopagurus lepidochirus (Doflein, 1902)
[Eupagurus]
Nematopagurus lewinsohni Trkay, 1986
Nematopagurus sp. McLaughlin & Okuno, 2010 (in
press)
Nematopagurus longicornis A. Milne-Edwards &
Bouvier, 1892
Nematopagurus meiringae McLaughlin, 1998
Nematopagurus muricatus (Henderson, 1896)
[Catapagurus]
Nematopagurus ostlingochirus McLaughlin, 1997
Nematopagurus ricei McLaughlin, 2004
Nematopagurus richeri McLaughlin, 2004
Nematopagurus scutellichelis Alcock, 1905
Nematopagurus scutelliformis McLaughlin, 1997
Nematopagurus spinulosensoris McLaughlin & Brock,
1974
Nematopagurus spongioparticeps McLaughlin, 2004
Nematopagurus squamichelis Alcock, 1905
Nematopagurus tricarinatus (Stimpson, 1858)
[Eupagurus]
Nematopagurus vallatus (Melin, 1939) [Eupagurus

(Catapagurus)]
Orthopagurus Stevens, 1927
= Orthopagurus Stevens, 1927 [type species Pagurus
minimus Holmes, 1900; by subsequent designation
by Makarov (1938); gender masculine]
= Orthopaguropsis Serne, 1957 (type species
Orthopagurus harmsi Gordon 1935, by original
Orthopagurus minimus (Holmes, 1900) [Pagurus]
Ostraconotus A. Milne-Edwards, 1880
= Ostraconotus A. Milne-Edwards, 1880 (type
species Ostraconotus spatulipes A. MilneEdwards, 1880 by monotypy; gender masculine)
Ostraconotus spatulipes A. Milne-Edwards, 1880
Paguridium Forest, 1961
= Paguridium Forest, 1961 (type species Eupagurus
? minimum Chevreux & Bouvier, 1892, by original
designation; gender neuter)
{16}
Paguridium minimum (Chevreux & Bouvier, 1892)
[Eupagurus]
Paguritta Melin, 1939
= Paguritta Melin, 1939 (type species Eupagurus
(Paguritta) gracilipes Melin, 1939, by monotypy;
gender feminine)
Paguritta corallicola Lewinsohn, 1978
Paguritta gracilipes (Melin, 1939) [Eupagurus
(Paguritta)]
Paguritta harmsi (Gordon, 1935) [Orthopagurus]
Paguritta kroppi McLaughlin & Lemaitre, 1993
Paguritta morgani McLaughlin & Lemaitre, 1993
Paguritta scottae McLaughlin & Lemaitre, 1993
Paguritta vittata Komai & Nishi, 1996
Pagurixus Melin, 1939
= Pagurixus Melin, 1939 (type species Eupagurus
(Pagurixus) boninensis Melin, 1939, by monotypy;
gender masculine)
Pagurixus acanthocarpus Komai & Okuno, 2009
Pagurixus amsa Morgan, 1993
Pagurixus anceps (Forest, 1957) [Eupagurus]
Pagurixus aurantiaca Komai, 2010
Pagurixus boninensis (Melin, 1939) [Eupagurus
(Pagurixus)]
Pagurixus brachydactylus Komai & Osawa, 2006
Pagurixus carinimanus Komai & Osawa, 2006
Pagurixus cavicarpus Komai, 2010
Pagurixus concolor Komai & Osawa, 2006
Pagurixus crosnieri Komai, 2010
Pagurixus dissimilis Osawa & Komai, 2007
Pagurixus fasciatus Komai & Myorin, 2005
Pagurixus festinus McLaughlin & Haig, 1984
Pagurixus formosus Komai, 2010
Pagurixus granulimanus Morgan, 1993
Pagurixus haigae Komai & Osawa, 2007
Pagurixus handrecki Gunn & Morgan, 1992
Pagurixus hectori (Filhol, 1883) [Eupagurus]
= Eupagurus campbelli Filhol, 1885
Pagurixus icelus Komai, 2010
Pagurixus jerviensis McLaughlin & Haig, 1984
31

Pagurus alaini Komai, 1998
Pagurus alatus Fabricius, 1775
{18}
= Pagurus tricarinatus Norman, 1869
= Eupagurus variabilis A. Milne-Edwards &
Bouvier, 1892
Pagurus albidianthus de Saint Laurent & McLaughlin,
2000
Pagurus albus (Benedict, 1892) [Eupagurus]
= Pagurus fuscomaculatus Bouvier, 1895
Pagurus alcocki (Balss, 1911) [Eupagurus]
Pagurus aleuticus (Benedict, 1892) [Eupagurus]
Pagurus anachoretoides Forest, 1966
Pagurus anachoretus Risso, 1827
= Pagurus pictus H. Milne Edwards, 1836
= Clibanarius mediterraneus Kossmann, 1878
Pagurus angustus (Stimpson, 1858) [Eupagurus]
Pagurus annexus McLaughlin & Haig, 1993
Pagurus annulipes (Stimpson, 1860) [Eupagurus]
Pagurus arcuatus Squires, 1964
= Pagurus bankensis Nesis, 1964
Pagurus arenisaxatilis Harvey & McLaughlin, 1991
Pagurus armatus (Dana, 1851) [Bernhardus]
Pagurus benedicti (Bouvier, 1898) [Eupagurus]
= Eupagurus minutus Benedict, 1892 (preoccupied
name)
= Nympagurus galapagensis Boone, 1932
Pagurus beringanus (Benedict, 1892) [Eupagurus]
= Eupagurus newcombei Benedict, 1892
Pagurus bernhardus (Linnaeus, 1758) [Cancer]
= Bernhardus typicus Dana, 1851
= Pagurus ulidiae Thompson, 1844 (nomen
nudum)
= Pagurus ulidanus Bell, 1846
{19}
= Pagurus eblaniensis Kinahan, 1860
Pagurus boriaustraliensis Morgan, 1990
Pagurus bouvieri (Faxon, 1895) [Eupagurus]
= Eupagurus smithi A. Milne-Edwards & Bouvier,
1893 (preoccupied name)
Pagurus brachiomastus (Thallwitz, 1892) [Eupagurus]
Pagurus brandti (Benedict, 1892) [Eupagurus]
Pagurus brevidactylus (Stimpson, 1859) [Eupagurus]
= Pagurus miamensis Provenzano, 1959
= Pagurus miamensis uncifer Forest & de Saint
Laurent, 1968
Pagurus bullisi Wass, 1963
Pagurus capillatus (Benedict, 1892) [Eupagurus]
Pagurus capsularis McLaughlin, 1997
Pagurus carneus (Pocock, 1889) [Eupagurus]
Pagurus carolinensis McLaughlin, 1975
Pagurus carpoforaminatus (Alcock, 1905) [Eupagurus]
Pagurus caurinus Hart, 1971
Pagurus cavicarpus (Paulson, 1875) [Eupagurus]
= Eupagurus carpoforaminatus var. nephromma
Alcock, 1905
Pagurus chevreuxi (Bouvier, 1896) [Eupagurus]
Pagurus chinensis White, 1847 (nomen nudum) {20}
Pagurus compressipes (Miers, 1884) [Eupagurus]
Pagurus comptus White, 1847
{21}
= Pagurus gayi Nicolet, 1849
Pagurixus kermadecensis de Saint Laurent & McLaughlin,

2000
Pagurixus laevimanus (Ortmann, 1892) [Eupagurus]
Pagurixus longipes Osawa, Fujita & Okuno, 2006
Pagurixus maorus (Nobili, 1906) [Eupagurus]
Pagurixus nomurai Komai & Asakura, 1995
Pagurixus patiae Komai, 2006
Pagurixus paulayi Komai & Osawa, 2006
Pagurixus pilosus Komai, 2010
Pagurixus pseliophorus Komai & Osawa, 2006
Pagurixus pulcher Osawa, Fujita & Okuno, 2006
Pagurixus purpureus Komai & Okuno, 2009
Pagurixus ruber Komai & Osawa, 2006
Pagurixus rubrovittatus Komai, 2010
Pagurixus sculptus Komai, 2010
Pagurixus stenops Morgan, 1993
Pagurixus tweediei (Forest, 1956) [Eupagurus]
Pagurodes Henderson, 1888
= Pagurodes Henderson, 1888 [type species
Pagurodes inarmatus Henderson, 1888, by
subsequent designation by de Saint Laurent (1969);
gender masculine]
Pagurodes inarmatus Henderson, 1888
Pagurodofleinia Asakura 2005
= Dofleinia McLaughlin & Asakura, 2004 (type
species Catapagurus doederleini Doflein 1902,
by original designation; gender feminine) (name
preoccupied by Dofleinia Wassifieff, 1908,
Cnidaria)
= Pagurodofleinia Asakura, 2005 (replacement
name)
P a g u ro d o f l e i n i a d o e d e r l e i n i ( D o f l e i n , 1 9 0 2 )
[Catapagurus]
Pagurojacquesia de Saint Laurent & McLaughlin, 2000
= Jaquesia de Saint Laurent & McLaughlin,
1999 (type species Jacquesia polymorpha de
Saint Laurent & McLaughlin, 1999, by original
designation; gender feminine) (name preoccupied
by Jacquesia Mendes, 1944, Mollusca)
= Pagurojacquesia de Saint Laurent & McLaughlin,
2000 (replacement name)
Pagurojacquesia polymorpha (de Saint Laurent &
McLaughlin, 1999) [Jacquesia]
Pagurus Fabricius, 1775
{17}
= Pagurus Fabricius, 1775 [type species by
subsequent selection by Latreille (1810) Cancer
bernhardus Linnaeus, 1758 as defined by lectotype
selection by Forest & Holthuis (1955): specimen
figured by Swammerdam, 1737, ICZN Opinion
472]
= Pagures Ordinaires H. Milne Edwards, 1836 (not
in Latin, invalid name)
= Pagures Ordinaires, Dextres H. Milne Edwards,
1848 (not in Latin, invalid name)
= Eupagurus Brandt, 1851 [type species by
subsequent selection by Stimpson (1858) Cancer
bernhardus Linnaeus, 1758]
= Bernhardus Dana, 1851 (type species Bernhardus
typicus Dana, 1851 by original designation)
Pagurus acadianus Benedict, 1901
32

Pagurus ikedai Lemaitre & Watabe, 2005
Pagurus imafukui McLaughlin & Konishi, 1994
Pagurus imaii (Yokoya, 1939) [Eupagurus]
Pagurus imarpe Haig, 1974
Pagurus impressus (Benedict, 1892) [Eupagurus]
Pagurus indicus Sarojini & Nagabhushanam, 1972
Pagurus inermis (Chevreux & Bouvier, 1892)
[Eupagurus]
Pagurus insulae Asakura, 1991
Pagurus investigatoris (Alcock, 1905) [Eupagurus]
Pagurus iridocarpus de Saint Laurent & McLaughlin,
2000
Pagurus irregularis (A. Milne-Edwards & Bouvier,
1892) [Eupagurus]
Pagurus isochirus White, 1847 (nomen nudum) {20}
Pagurus japonicus (Stimpson, 1858) [Eupagurus]
= Eupagurus barbatus Ortmann, 1892
Pagurus kaiensis McLaughlin, 1997
Pagurus kennerlyi (Stimpson, 1864) [Eupagurus]
Pagurus kulkarnii Sankolli, 1962
Pagurus lanuginosus De Haan, 1849
Pagurus laurentae Forest, 1978
Pagurus lepidus (Bouvier, 1898) [Eupagurus]
Pagurus leptonyx Forest & de Saint Laurent, 1968
= Pagurus leptonix Fausto-Filho, 1970 (misspelling
of leptonyx)
Pagurus limatulus Fausto Filho, 1970
Pagurus liochele (Barnard, 1947) [Pylopagurus]
= Pagurus barnardi Forest, 1966
Pagurus longicarpus Say, 1817
= Pagurus truncatulus Rafinesque, 1817
Pagurus longimanus Wass, 1963
Pagurus lophochela Komai, 1999
Pagurus luticola Komai & Chan, 2006
Pagurus macardlei (Alcock, 1905) [Eupagurus]
Pagurus maclaughlinae Garca-Gmez, 1982
Pagurus maculosus Komai & Imafuku, 1996
Pagurus marshi Benedict, 1901
= Pagurus marschi Forest & de Saint Laurent, 1968
(misspelling of marshi)
Pagurus mbizi (Forest, 1955) [Eupagurus]
Pagurus meloi Lemaitre & Cruz Castao, 2004
Pagurus mertensii Brandt, 1851
Pagurus middendorffii Brandt, 1851
Pagurus minutus Hess, 1865
= Eupagurus dubius Ortmann, 1892
Pagurus moluccensis Haig & Ball, 1988
Pagurus nanodes Haig & Harvey, 1991
Pagurus nesiotes Haig & McLaughlin, 1991
Pagurus nigrivittatus Komai, 2003
Pagurus nigrofascia Komai, 1996
Pagurus nipponensis (Yokoya 1933) [Eupagurus]
Pagurus nisari Siddiqui & Komai, 2008
Pagurus novizealandiae (Dana, 1852) [Bernhardus]
Pagurus ochotensis Brandt, 1851
= Pagurus (Eupagurus) bernhardus var. B.
granulato-denticulata Brandt, 1851
= Eupagurus alaskensis Benedict, 1892
= Eupagurus ortmanni Balss, 1911
Pagurus parvispina Komai, 1997
Pagurus conformis De Haan, 1849

= Eupagurus megalops Stimpson, 1858
Pagurus confragosus (Benedict, 1892) [Eupagurus]
Pagurus confusus Komai & Yu, 1999
Pagurus constans (Stimpson, 1858) [Eupagurus]
= Pagurus sagamiensis Miyake, 1978
Pagurus cornutus (Benedict, 1892) [Eupagurus]
Pagurus criniticornis (Dana, 1852) [Bernhardus]
Pagurus cuanensis Bell, 1846
{19}
= Pagurus Cuanensis Thompson, 1844 (nomen
nudum)
= Pagurus spinimanus Lucas, 1846
= Pagurus lucasi Heller, 1863
= Eupagurus chiereghini Nardo, 1868
= Eupagurus placens Stebbing, 1924
Pagurus curacaoensis (Benedict, 1892) [Eupagurus]
Pagurus dalli (Benedict, 1892) [Eupagurus]
Pagurus dartevellei (Forest, 1958) [Pylopagurus]
Pagurus decimbranchiae Komai & Osawa, 2001
Pagurus defensus (Benedict, 1892) [Eupagurus]
Pagurus delsolari Haig, 1974
Pagurus dissimilis (A. Milne-Edwards & Bouvier, 1893)
[Eupagurus]
Pagurus edwardsii (Dana, 1852) [Bernhardus]
Pagurus emmersoni McLaughlin & Forest, 1999
Pagurus erythrogrammus Komai, 2003
Pagurus excavatus (Herbst, 1791) [Cancer]
{18}
= Pagurus angulatus Risso, 1816
= Eupagurus meticulosus Roux, 1830
Pagurus exiguus (Melin, 1939) [Eupagurus
(Pagurillus)]
Pagurus exilis (Benedict, 1892) [Eupagurus]
Pagurus filholi (De Man, 1887) [Eupagurus]
= Pagurus geminus McLaughlin, 1976
Pagurus fimbriatus Forest, 1966
Pagurus forbesii Bell, 1846
{19}
Pagurus forceps H. Milne Edwards, 1836
{21}
= Eupagurus comptus var. jugosa Henderson,
1888
= Eupagurus comptus var. latimanus Miers, 1875
Pagurus fungiformis Komai & Rahayu, 2004
Pagurus fuscomaculatus (Bouvier, 1895) [Eupagurus]
Pagurus gladius (Benedict, 1892) [Eupagurus]
Pagurus gordonae (Forest, 1956) [Eupagurus]
Pagurus gracilipes (Stimpson, 1858) [Eupagurus]
Pagurus granosimanus (Stimpson, 1859) [Eupagurus]
Pagurus gymnodactylus Lemaitre, 1982
Pagurus hartae (McLaughlin & Jensen, 1996)
[Parapagurodes]
Pagurus heblingi Nucci & de Melo, 2003
Pagurus hedleyi (Grant & McCulloch, 1906) [Eupagurus]
(replacement name for Eupagurus kirkii Miers, 1884,
preoccupied by Eupagurus kirkii Filhol 1883)
Pagurus hemphilli (Benedict, 1892) [Eupagurus]
Pagurus hirsutiusculus (Dana, 1851) [Bernhardus]
Pagurus hirtimanus (Miers, 1880) [Eupagurus]
= Pagurus hirtimanus White, 1847 (nomen
nudum)
= Eupagurus janitor Alcock, 1905
Pagurus holmi Ng & McLaughlin, 2009
33

Pagurus parvus (Benedict, 1892) [Eupagurus]
Pagurus pectinatus (Stimpson, 1858) [Eupagurus]
= Eupagurus seriespinosus Thallwitz, 1891
= Clibanarius japonicus Rathbun, 1903
Pagurus pergranulatus (Henderson, 1896) [Eupagurus]
Pagurus perlatus H. Milne Edwards, 1848
= Bernhardus obesocarpus Dana, 1852
Pagurus pilosimanus White, 1847 (nomen nudum)
{20}
Pagurus pilosipes (Stimpson, 1858) [Eupagurus]
Pagurus pilosiusculus White, 1847 (nomen nudum)
{20}
Pagurus pitagsaleei McLaughlin, 2002
Pagurus politus (Smith, 1882) [Eupagurus]
Pagurus pollexcavus Glassell, 1937 uncertain status
[22]
Pagurus pollicaris Say, 1817
= Eupagurus floridanus Benedict, 1892
Pagurus prideaux Leach, 1815
Pagurus protuberocarpus McLaughlin, 1982
Pagurus provenzanoi Forest & de Saint Laurent, 1968
Pagurus proximus Komai, 2000
Pagurus pubescens Kryer, 1838
= Pagurus thompsoni Bell, 1853
{15}
= Eupagurus kroyeri Stimpson, 1859
Pagurus pubescentulus (A. Milne-Edwards & Bouvier
1892) [Eupagurus]
= Eupagurus variabilis var. charcoti Bouvier,
1914
Pagurus pulchellus (A. Milne-Edwards & Bouvier, 1892)
[Eupagurus]
Pagurus quaylei Hart, 1971
Pagurus quinquelineatus Komai, 2003
Pagurus rathbuni (Benedict, 1892) [Eupagurus]
= Eupagurus (Trigonocheirus) polaris Sivertsen,
1932
Pagurus redondoensis Wicksten, 1982
Pagurus retrorsimanus Wicksten & McLaughlin, 1998
Pagurus rhabdotus Haig & Harvey, 1991
Pagurus rotundimanus Wass, 1963
Pagurus ruber (A. Milne-Edwards & Bouvier, 1892)
[Eupagurus]
Pagurus rubrior Komai, 2003
Pagurus samoensis (Ortmann, 1892) [Eupagurus]
Pagurus samuelis (Stimpson, 1857) [Eupagurus]
Pagurus setosus (Benedict, 1892) [Eupagurus]
Pagurus similimanus (Balss, 1921) [Eupagurus
(?Anapagurus)]
Pagurus similis (Ortmann, 1892) [Eupagurus]
Pagurus simulans Komai, 2000
Pagurus sinuatus (Stimpson, 1858) [Eupagurus]
Pagurus smithi (Benedict, 1892) [Eupagurus]
Pagurus souriei (Forest, 1952) [Eupagurus]
Pagurus spighti McLaughlin & Haig, 1993
Pagurus spilocarpus Haig, 1977
Pagurus spina Komai, 1994
Pagurus spinulentus (Henderson, 1888) [Eupagurus]
Pagurus stevensae Hart, 1971
Pagurus sticticus McLaughlin, 2008
Pagurus stimpsoni (A. Milne-Edwards & Bouvier, 1893)

[Eupagurus]
= Pagurus bonairensis Schmitt, 1936
= Pagurus hendersoni Wass, 1963
Pagurus tanneri (Benedict, 1892) [Eupagurus]
Pagurus townsendi (Benedict, 1892) [Eupagurus]
Pagurus traversi (Filhol, 1885) [Eupagurus]
Pagurus triangularis (Chevreux & Bouvier, 1892)
[Eupagurus]
Pagurus trichocerus Forest & de Saint Laurent, 1968
Pagurus trigonocheirus (Stimpson, 1858) [Eupagurus]
Pagurus tristanensis (Henderson, 1888) [Eupagurus]
Pagurus undosus (Benedict, 1892) [Eupagurus]
= Eupagurus trigonocheirus var. paulensis Balss,
1913
Pagurus venturensis Coffin, 1957
= Pagurus hirsutiusculus venturensis Coffin 1957
Pagurus vetaultae Harvey & McLaughlin, 1991
Pagurus villosus Nicolet, 1849
Pagurus virgulatus Haig & Harvey, 1991
Parapagurodes McLaughlin & Haig, 1973
= Parapagurodes McLaughlin & Haig, 1973 (type
species Parapagurodes makarovi McLaughlin
& Haig, 1973, by original designation; gender
masculine)
Parapagurodes laurentae McLaughlin & Haig, 1973
Parapagurodes makarovi McLaughlin & Haig, 1973
Phimochirus McLaughlin, 1981
= Phimochirus McLaughlin, 1981 (type species
Eupagurus operculatus Stimpson, 1859, by
Phimochirus californiensis (Benedict, 1892)
[Eupagurus]
= Eupagurus mexicanus Benedict, 1892
Phimochirus holthuisi (Provenzano, 1961)
[Pylopagurus]
Phimochirus leurocarpus McLaughlin, 1981
Phimochirus occlusus (Henderson, 1888) [Eupagurus]
P h i m o c h i r u s o p e rc u l a t u s ( S t i m p s o n , 1 8 5 9 )
[Eupagurus]
Pylopagurus samariensis Sanchez, 1978
Phimochirus randalli (Provenzano, 1961)
[Pylopagurus]
Phimochirus roseus (Benedict, 1892) [Eupagurus]
Phimochirus venustus (Bouvier, 1898) [Eupagurus]
Porcellanopagurus Filhol, 1885
= Porcellanopagurus Filhol, 1885 (type species
Porcellanopagurus edwardsi Filhol, 1885 by
Porcellanopagurus adelocercus McLaughlin & Hogarth,
1998
Porcellanopagurus belauensis Suzuki & Takeda, 1987
Porcellanopagurus chiltoni de Saint Laurent &
McLaughlin, 2000
Porcellanopagurus edwardsi Filhol, 1885
Porcellanopagurus filholi de Saint Laurent & McLaughlin,
2000
Porcellanopagurus foresti Zarenkov, 1990
Porcellanopagurus haptodactylus McLaughlin, 2000
34

Porcellanopagurus jacquesi McLaughlin, 1997
Porcellanopagurus japonicus Balss, 1913
Porcellanopagurus nihonkaiensis Takeda, 1985
Porcellanopagurus platei Lenz, 1902
Porcellanopagurus tridentatus Whitelegge, 1900
Porcellanopagurus truncatifrons Takeda, 1981
Propagurus McLaughlin & de Saint Laurent, 1998
= Propagurus McLaughlin & de Saint Laurent, 1998
(type species Pagurus gaudichaudii H. Milne
Edwards, 1836, by original designation; gender
masculine)
Propagurus deprofundis (Stebbing, 1924) [Eupagurus]
Propagurus gaudichaudii (H. Milne Edwards, 1836)
[Pagurus]
= Bernhardus barbiger A. Milne-Edwards, 1891
Propagurus haigae (McLaughlin, 1997) [Pagurus]
Propagurus miyakei (Baba, 1986) [Pagurus]
Propagurus obtusifrons (Ortmann, 1892) [Eupagurus]
= Pagurus yokoyai Makarov, 1938
Protoniopagurus Lemaitre & McLaughlin, 1996
= Protoniopagurus Lemaitre & McLaughlin, 1996
(type species Protoniopagurus bioperculatus
Lemaitre & McLaughlin, 1996, by original
Protoniopagurus bioperculatus Lemaitre & McLaughlin,
1996
Pseudopagurodes McLaughlin, 1997
= Pseudopagurodes McLaughlin, 1997 (type species
Pagurodes piliferus Henderson, 1888, by original
Pseudopagurodes piliferus (Henderson, 1888)
[Pagurodes]
Pseudopagurodes reconditus (Wang & McLaughlin,
2000) [Nematopaguroides]
Pteropagurus McLaughlin & Rahayu, 2006
= Pteropagurus McLaughlin & Rahayu, 2006 (type
species Pteropagurus inermis McLaughlin &
Rahayu, 2006 by original designation; gender
masculine)
Pteropagurus inermis McLaughlin & Rahayu, 2006
Pteropagurus spina McLaughlin & Rahayu, 2006
Pteropagurus spinulocarpus McLaughlin, 2007
Pumilopagurus McLaughlin & Rahayu, 2008
= Pumilopagurus McLaughlin & Rahayu, 2008
(type species Pumilopagurus tuberculomanus
McLaughlin & Rahayu, 2008, by original
Pumilopagurus tuberculomanus McLaughlin & Rahayu,
2008
Pygmaeopagurus McLaughlin, 1986
= Pygmaeopagurus McLaughlin, 1986 (type species
Pygmaeopagurus hadrochirus McLaughlin, 1986
Pygmaeopagurus hadrochirus McLaughlin, 1986
Pylopaguridium McLaughlin & Lemaitre, 2001
= Pylopaguridium McLaughlin & Lemaitre,
2001 (type species Pylopaguridium markhami
McLaughlin & Lemaitre, 2001 by original
designation; gender neuter)
Pylopaguridium markhami McLaughlin & Lemaitre,

2001
Pylopaguropsis Alcock, 1905
= Pylopaguropsis Alcock, 1905 (type species
Pylopagurus magnimanus Henderson, 1896, by
monotypy; gender feminine)
= Galapagurus Boone, 1932 (type species
Galapagurus teevana Boone, 1932, by monotypy;
gender masculine)
Pylopaguropsis atlantica Wass, 1963
Pylopaguropsis bellula Osawa & Okuno, 2006
Pylopaguropsis fimbriata McLaughlin & Haig, 1989
Pylopaguropsis furusei Asakura, 2000
Pylopaguropsis garciai McLaughlin & Haig, 1989
Pylopaguropsis granulata Asakura, 2000
Pylopaguropsis keijii McLaughlin & Haig, 1989
Pylopaguropsis laevispinosa McLaughlin & Haig,
1989
Pylopaguropsis lemaitrei Asakura & Paulay, 2003
Pylopaguropsis lewinsohni McLaughlin & Haig, 1989
Pylopaguropsis magnimanus (Henderson, 1896)
[Pylopagurus]
Pylopaguropsis pustulosa McLaughlin & Haig, 1989
Pylopaguropsis speciosa McLaughlin & Haig, 1989
Pylopaguropsis teevana (Boone, 1932) [Galapagurus]
Pylopaguropsis vicina Komai & Osawa, 2004
Pylopaguropsis zebra (Henderson, 1893) [Eupagurus]
Pylopagurus A. Milne-Edwards & Bouvier, 1893
= Pylopagurus A. Milne-Edwards & Bouvier, 1893
[type species Eupagurus discoidalis A. MilneEdwards, 1880, by subsequent designation by
Miyake (1978); gender masculine]
Pylopagurus discoidalis (A. Milne-Edwards, 1880)
[Eupagurus]
Pylopagurus gorei McLaughlin & Lemaitre, 2001
Pylopagurus holmesi Schmitt, 1921
= Pylopagurus longicarpus Walton, 1954
Pylopagurus macgeorgei McLaughlin & Lemaitre,
2001
Pylopagurus pattiae Lemaitre & Campos, 1993
Rhodochirus McLaughlin, 1981
= Rhodochirus McLaughlin, 1981 (type species
Pylopagurus rosaceus A. Milne-Edwards &
Bouvier, 1893, by original designation; gender
masculine)
Rhodochirus hirtimanus (Faxon, 1893) [Pylopagurus]
Rhodochirus rosaceus (A. Milne-Edwards & Bouvier,
1893) [Pylopagurus]
= Pylopagurus acutus Forest & de Saint Laurent,
1968
Scopaeopagurus McLaughlin & Hogarth, 1998
= Scopaeopagurus McLaughlin & Hogarth, 1998
(type species Scopaeopagurus megalochirus
McLaughlin & Hogarth, 1998, by original
Scopaeopagurus megalochirus McLaughlin & Hogarth,
1998
35

Solenopagurus de Saint Laurent, 1968
= Solenopagurus de Saint Laurent, 1968 (type
species Cestopagurus lineatus Wass, 1963, by
Solenopagurus diomedeae (Faxon, 1893)
[Catapagurus]
Solenopagurus lineatus (Wass, 1963) [Cestopagurus]
Solitariopagurus Trkay, 1986
= Solitariopagurus Trkay, 1986 (type species
Solitariopagurus profundus Trkay, 1986 by
Solitariopagurus profundus Trkay, 1986
Solitariopagurus triprobolus Poupin & McLaughlin,
1996
Solitariopagurus trullirostris McLaughlin, 2000
Solitariopagurus tuerkayi McLaughlin, 1997
Spiropagurus Stimpson, 1858
= Spiropagurus Stimpson, 1858 (type species
Pagurus spiriger De Haan, 1849 by monotypy;
gender masculine)
Spiropagurus elegans Miers, 1881
Spiropagurus fimbriatus Lewinsohn, 1982
Spiropagurus lophomeris Alcock, 1905
{23}
= Spiropagurus spiriger var. lophomeris Alcock,
1905
Spiropagurus profundorum Alcock, 1905
{23}
= Spiropagurus spiriger var. profundorum Alcock,
1905
Spiropagurus spinosicarpis Alcock, 1905
{23}
= Spiropagurus spiriger var. spinosicarpis Alcock,
1905
Spiropagurus spiriger (De Haan, 1849) [Pagurus]
Tarrasopagurus McLaughlin, 1997
= Tarrasopagurus McLaughlin, 1997 (type
species Tarrasopagurus rostrodenticulatus
McLaughlin, 1997, by original designation; gender
masculine)
Tarrasopagurus rostrodenticulatus McLaughlin, 1997
Tomopaguroides Balss, 1912
= Tomopaguroides Balss, 1912 (type species
Parapagurus valdividae Balss, 1911, by original
To m o p a g u r o i d e s v a l d i v i d a e ( B a l s s , 1 9 1 1 )
[Parapagurus]
Tomopaguropsis Alcock, 1905
= Tomopaguropsis Alcock, 1905 [type species
Tomopaguropsis lanata Alcock, 1905, by
subsequent selection by McLaughlin (1997);
gender feminine]
Tomopaguropsis lanata Alcock, 1905
Tomopaguropsis miyakei McLaughlin, 1997
Tomopaguropsis problematica (A. Milne-Edwards &
Bouvier, 1893) [Eupagurus]
Tomopagurus A. Milne-Edwards & Bouvier, 1893
= Tomopagurus A. Milne-Edwards & Bouvier, 1893
(type species Tomopagurus rubropunctatus A.
Milne- Edwards & Bouvier, 1893, by monotypy;
gender masculine)
= Benthopagurus Wass, 1963 (type species

Benthopagurus schmitti Wass, 1963, by original
Tomopagurus chacei (Wass, 1963) [Pylopagurus]
Tomopagurus cokeri (Hay, 1917) [Pagurus]
= Benthopagurus schmitti Wass, 1963
Tomopagurus cubensis (Wass, 1963) [Benthopagurus]
Tomopagurus maclaughlinae Haig, 1976
Tomopagurus merimaculosus (Glassell, 1937)
[Pagurus]
Tomopagurus purpuratus (Benedict, 1892) [Eupagurus]
= Pagurus bunomanus Glassell, 1937 (nomen
nudum)
Tomopagurus rubropunctatus A. Milne-Edwards &
Bouvier, 1893
= Pagurus rubrolineatus Wass, 1963
Tomopagurus wassi McLaughlin, 1981
Trichopagurus de Saint Laurent, 1968
= Trichopagurus de Saint Laurent, 1968 (type species
Catapaguroides ? trichophthalmus Forest, 1954,
by original designation; gender masculine) {24}
Trichopagurus trichophthalmus (Forest, 1954)
[Catapaguroides ?]
Trichopagurus macrochela Komai & Osawa, 2005
Turleania McLaughlin, 1997
= Laurentia McLaughlin & Haig, 1996 (type species
Laurentia albatrossae McLaughlin & Haig, 1996,
by original designation) (name preoccupied by
Laurentia Ragonot, 1888, Insecta)
= Turleania McLaughlin, 1997 (replacement
name)
Turleania albatrossae (McLaughlin & Haig, 1996)
[Laurentia]
Turleania boucheti McLaughlin, 2007
Tu r l e a n i a b a l l i ( M c L a u g h l i n & H a i g , 1 9 9 6 )
[Laurentia]
Turleania multispina McLaughlin, 1997
Turleania saliens Osawa & Fujita, 2008
Turleania senticosa (McLaughlin & Haig, 1996)
[Laurentia]
= Turleania similis Komai, 1999
Turleania sibogae (McLaughlin & Haig, 1996)
[Laurentia]
Turleania spinimanus Komai, 1999
Turleania tenebrosa Osawa & Fujita, 2008
Xylopagurus A. Milne-Edwards, 1880
= Xylopagurus A. Milne-Edwards, 1880 (type
species Xylopagurus rectus A. Milne-Edwards,
Xylopagurus anthonii Lemaitre, 1995
Xylopagurus caledonicus Forest, 1997
Xylopagurus cancellarius Walton, 1950
Xylopagurus philippinensis Forest, 1997
Xylopagurus rectus A. Milne-Edwards, 1880
Xylopagurus tayrona Lemaitre & Campos, 1993
Xylopagurus tenuis Lemaitre, 1995
Incertae sedis
Anapagurus acutus A. Milne-Edwards& Bouvier, 1893
{25}
36

Anapagurus marginatus A. Milne-Edwards & Bouvier,
1893
{25}
Catapagurus; still no mention was made of a type

species for Hemipagurus. After examining one of
the syntypes of Catapagurus sharreri A. MilneEdwards, 1880, Smith (1883) concluded that his own
C. socialis was conspecific with C. sharreri. It was
only when Asakura (2001) resurrected Hemipagurus
was H. gracilis cited as the type species of Smiths
(1881a) taxon. Although at the time of Asakuras
(2001) reinstatement, Hemipagurus contained only H.
gracilis, it is not the type species by monotypy (ICZN
Art. 69.4) but rather by designation by Asakura.
NOTES
{1}
To indicate their uncertainty regarding the generic

assignment of the new species to Anapagurus, A.
Milne-Edwards & Bouvier, 1900, separated the
generic name from the specific name, dubius, with a
question mark.
{2}
Thomas Bells A History of British Crustacea was

published in parts between 1844 and 1853. Part III,
which contained the pagurid species Anapagurus
hyndmanni and A. laevis was published in January
1846 (Gordon, 1959).
{3}
Pagurus pilosipes (Stimpson, 1858) will be transferred

to Boninpagurus by Komai & Okuno (in preparation)
as the senior synonym of B. acanthocheles.
{4}
Catapaguroides kasei Osawa & Takeda, 2004 appears

to be synonymous with C. foresti, but confirmation
is still needed.
{5}
De Saint Laurent (1968a) felt certain that the western

Pacific species she identified as Catapaguroides
microps was conspecific with the eastern Atlantic
population; however recent collections suggest that
the former may actually represent a distinct taxon.
{6}
Catapaguroides setosus Edmondson, 1951, was

questionably synonymized with C. fragilis by de
Saint Laurent (1968a) but reinstated by McLaughlin
& Pittman (2002) after comparing representatives of
the two species.
{7}
{8}
{9}
Catapagurus maclaughlinae Asakura, 2001, was

established on two females from the Seycheles and
distinguished from other species of Catapagurus
by weak development of carpal and meral spines
on the ambulatory legs. Subsequent discovery of a
male of C. maclaughlinae from a French Seycheles
expedition and comparison with an expanded sample
of C. alcocki has shown that the two taxa are
conspecific. Catapagurus maclaughlinae will be put
into synonymy with C. alcocki in an upcoming review
of the genus where justification for the synonymy will
be presented.
{10} Although C. gracilis var. intermedius was described by

A. Milne-Edwards & Bouvier (1893), this subspecies
was overlooked by Asakura (2001) in his generic
revision. In a reassessment currently in progress
McLaughlin has compared the two syntypes of this
subspecies with the nominal subspecies and found
no justification for two distinct taxa. Catapagurus
gracilis intermedius will be placed in synonymy in
the upcoming reassessment.
{11} In her ongoing study of species of Catapagurus,
McLaughlin has found C. imperalis to be conspecific
with C. oculocrassus and will be placed in synonymy
with this species.
Catapagurus A. Milne-Edwards, 1880, had long

been considered the senior synonym of Hemipagurus
Smith, 1881 until Asakura (2001) proposed the
resurrection of the latter. That reinstatement was
rejected by McLaughlin (2004) who showed that
the characters used by Asakura (2001) to distinguish
Catapagurus from Hemipagurus reflected only
intrageneric variability and growth related differences.
McLaughlin returned Hemipagurus to synonymy. Five
more species of Catapagurus are now being described
by the first author.
{12} Asakura (2001) inappropriately designated a lectotype

for C. sharreri; the same male had previously been so
designated by A. Milne-Edwards & Bouvier (1893:
130).
{13} It is possible that the type series of this genus was
a mixture of more than one species of different
sexes, because such a misidentification is not
uncommon in Yokoyas (1933) material. Despite
the collecting efforts of one of the present authors
(TK), no specimens that agree with Yokoyas original
description have been encountered in Japan.
In Smiths (1881a) initial description of Hemipagurus,

no species were mentioned. Subsequently, Smith
(1881b) repeated his description of the genus and
described the two species he assigned to it, H.
socialis Smith, 1881 and H. gracilis Smith, 1881;
he did not designate either species as the type for the
genus. The following year Smith (1882) determined
that Hemipagurus was identical with A. MilneEdwards (1880) Catapagurus, placed Hemipagurus
in synonymy and transferred its two species to
{14} The possibility exists that this taxon may be a junior

synonym of L. (A.) triserratus.
{15} Eupagurus thompsoni Filhol, 1885, became a
secondary junior homonym of Pagurus thompsoni
Bell, 1853 when the ICZN (Opinion 472) placed
Eupagurus on the Officical Index of Rejected and
37

Invalid Generic Names in Zoology. As discussed
by McLaughlin & Gunn (1992), Forest (in Pike,
1961) suggested that Filhols (1885b) P. thompsoni
was synonymous with P. lacertosus (Henderson,
1888). Forest (unpublished) subsequently concluded
that those two taxa were distinct, and both species
were transferred to Pylopagurus by Forest & de
Saint Laurent (1968) thus eliminating the need for a
replacement name for Filhols taxon (ICZN, 2000:
Art 59.2). Both species were ultimately transferred to
Lophopagurus McLaughlin, 1981, whereas Bells P.
thompsoni was found to be synonymous with Pagurus
pubescens Kryer, 1838.
isochirus and Pagurus pilosimanus without localities,

and Pagurus pilosiusculus from Mr. Drings collection,
Swan River. None of these taxa were provided with
even brief descriptions or illustrations, and no further
mention of any of them could be found in Whites
subsequent publications.
{21} The conspecificity of Pagurus comptus and P. forceps
has been the subject of debate since Lagerberg (1905)
first placed P. comptus in synonymy with P. forceps.
However, a recent study by Mantelatto et al. (2009),
using both morphological and genetic data, has
demonstrated conclusively that the two taxa represent
distinct species.
{16} Apparently uncertain about the generic placement of

their solitary juvenile specimen, Chevreux & Bouvier
(1892) described the taxon as Eupagurus ? minimum.
Forest (1961), having obtained additional, and mature
specimens, recognized the species as belonging to a
genus distinct from other known pagurid genera, and
extablished the genus Paguridium Forest, 1961 for
it.
{22} Glassells (1937) Pagurus pollexcavus was not

reported as occurring in the Gulf of California by
Hendrickx & Harvey (1999), nor was it included
the Lemaitre & Cruz Castaos (2004) checklist of
eastern Pacific species of Pagurus. But to the best
of our knowledge, this species has not been put
into synonymy with any other regional species or
considered incertae sedis for any reason. Therefore
we have listed its status simply as uncertain.
{17} As previously noted, when first established by

Fabricius (1775) Pagurus included a number of
Linnaeus (1758) non crab-like species of Cancer. It
was 35 years later that Pagurus bernhardus (Linnaeus,
1758) was selected by Latreille (1810) as the type
species of the genus and another quarter century
before the first detailed classification of Pagurus was
published (H. Milne Edwards 1836). As can be clearly
seen in the brief accounts of subsequent carcinologists
from Brandt (1851) to Rathbun (1903) and beyond,
inaccurate interpretations and misidentifications
resulted in nomenclatorial chaos that prevailed until
the matter was brought before the International
Commission (Forest & Holthuis, 1955). In its ruling,
the Commission validated the generic names Pagurus
(sensu Fabricius) and Dardanus Paulson (Opinion
472).
{23} McLaughlin (2002) elevated all of Alcocks (1905)

subspecies of Spiropagurus spiriger to full generic
rank.
{24} When Forest (1954) described Catapaguroides
? trichophthalmus, the genus Catapaguroides
appeared to have considerable overlap with a
similar genus, Cestopagurus. Not being certain of
its generic placement, Forest questionably assigned
trichophthalmus to Catapaguroides.
{25} Garca-Gmez (1994) examined the two female
specimens upon which A. acutus and A. marginatus
were based and concluded that they were inaccurately
assigned to Anapagurus. As they could not be
assigned with confidence to any taxon, Garca-Gmez
considered them incertae sedis.
{18} Forest (1955) placed Pagurus excavatus (Herbst,

1791) in synonymy with Pagurus alatus Fabricius,
1775. However, Ingle (1985) presented evidence
supporting Selbies (1921) opinion that the two
taxa were distinct, with P. excavatus restricted in its
northern distribution to the Bay of Biscay. Ingle (1985)
reinstated P excavatus and placed the morphologically
similar P. variabilis (A. Milne-Edwards & Bouvier,
1892) in synonymy with the northern P. alatus.
Family Parapaguridae Smith, 1882

Bivalvopagurus Lemaitre 1993
= Bivalvopagurus Lemaitre 1993 (type species
Parapagurus sinensis de Saint Laurent, 1972, by
Bivalvopagurus sinensis (de Saint Laurent, 1972)
[Parapagurus]
Oncopagurus Lemaitre, 1996
= Oncopagurus Lemaitre, 1996 (type species
Eupagurus bicristatus A. Milne-Edwards, 1880,
Oncopagurus africanus (de Saint Laurent, 1972)
[Parapagurus]
Oncopagurus bicristatus (A. Milne-Edwards, 1880)
[Eupagurus]
{1}
{19} Thomas Bells A History of British Crustacea was

published in parts between 1844 and 1853. Part
III, which included the pagurid species Pagurus
cuanensis, ulidianus and P. Forbesii, was published
in January 1846 (Gordon 1959).
{20} In his list of the Crustacea in the British Museum,
Adam White (1847: 5962) listed four new species
of Pagurus: Pagurus chinensis from China, Pagurus
38

Oncopagurus cidaris Lemaitre, 1996
Oncopagurus conicus Lemaitre, 2006
Oncopagurus curvispina (de Saint Laurent, 1974)
[Parapagurus]
{2}
Oncopagurus glebosus Lemaitre, 1997
Oncopagurus gracilis (Henderson, 1888)
[Parapagurus]
= Parapagurus bicristatus gracilis Henderson,
1888
{1}
= Sympagurus arcuatus A. Milne-Edwards &
Bouvier, 1893
= Pylopagurus exquisitus Boone, 1927
Oncopagurus haigae (de Saint Laurent, 1972)
[Parapagurus]
Oncopagurus indicus (Alcock, 1905) [Sympagurus]
= Sympagurus bicristatus var. indicus Alcock,
1905
{1}
Oncopagurus minutus (Henderson, 1896)
[Parapagurus]
Oncopagurus mironovi Zhadan, 1997
O n c o p a g u r u s m o n s t ro s u s ( A l c o c k , 1 8 9 4 )
[Parapagurus]
= Sympagurus arcuatus var. monstrosus Alcock,
1894
= ? Eupagurus brevimanus Yokoya, 1933
{3}
Oncopagurus oimos Lemaitre, 1998
Oncopagurus orientalis (de Saint Laurent, 1972)
[Parapagurus]
Oncopagurus stockmani Zhadan, 1997
Oncopagurus tuamotu Lemaitre, 1994
Oncopagurus sp. A Zhadan, 1997
{4}
Oncopagurus n. sp. 1
{5}
{5}
{5}
{5}
Paragiopagurus Lemaitre, 1996
= Paragiopagurus Lemaitre, 1996 (type species
Sympagurus diogenes Whitelegge, 1900, by
original designation; gender: masculine)
Paragiopagurus acutus (de Saint Laurent, 1972)
[Parapagurus]
= Parapagurus acutus acutus de Saint Laurent,
1972
{6}
Paragiopagurus bicarinatus (de Saint Laurent, 1972)
[Parapagurus]
= Parapagurus acutus bicarinatus de Saint Laurent,
1972
{6}
Paragiopagurus boletifer (de Saint Laurent, 1972)
[Parapagurus]
= Parapagurus sculptochela Zarenkov, 1990
Paragiopagurus bougainvillei (Lemaitre, 1994)
[Sympagurus]
Paragiopagurus diogenes (Whitelegge, 1900)
[Sympagurus]
Paragiopagurus fasciatus Lemaitre & Poupin, 2003
Paragiopagurus hirsutus (de Saint Laurent, 1972)
[Parapagurus]
= Parapagurus acutus hirsutus de Saint Laurent,
1972
{6}
Paragiopagurus hobbiti (Macpherson, 1983)

[Parapagurus]
Paragiopagurus insolitus Lemaitre, 1997
P a r a g i o p a g u r u s m a c ro c e r u s ( F o r e s t , 1 9 5 5 )
[Parapagurus]
Paragiopagurus pacificus (Edmondson, 1925)
[Sympagurus]
Paragiopagurus pilimanus (A. Milne-Edwards, 1880)
[Eupagurus]
Paragiopagurus rugosus (de Saint Laurent, 1972)
[Parapagurus]
Paragiopagurus ruticheles (A. Milne-Edwards, 1891)
[Eupagurus]
Paragiopagurus schnauzer Lemaitre, 2006
Paragiopagurus tuberculosus (de Saint Laurent, 1972)
[Parapagurus]
Paragiopagurus ventilatus Lemaitre, 2004
Paragiopagurus wallisi (Lemaitre, 1994) [Sympagurus]
{5}
Paragiopagurus n. sp. 1
{5}
{5}
{5}
Parapagurus Smith, 1879
= Parapagurus Smith, 1879 (type species
Parapagurus pilosimanus Smith, 1879, by
Parapagurus abyssorum (Filhol, 1885) [Pagurus] {7}
= Parapagurus abyssorum var. scabra Henderson,
1888
Parapagurus alaminos Lemaitre, 1986
Parapagurus andreui Macpherson, 1984
Parapagurus benedicti de Saint Laurent, 1972
= Parapagurus pilosimanus benedicti de Saint
Laurent, 1972
Parapagurus bouvieri Stebbing, 1910
Parapagurus foraminosus Lemaitre, 1999
Parapagurus furici Lemaitre, 1999
Parapagurus holthuisi Lemaitre, 1989
{8}
= Parapagurus abyssorum Henderson, 1888
Parapagurus janetae Lemaitre, 1999
Parapagurus latimanus Henderson, 1888
Parapagurus microps de Saint Laurent, 1972
Parapagurus nudus (A. Milne-Edwards, 1891)
[Sympagurus]
Parapagurus pilosimanus Smith, 1879
{9}
= Eupagurus jacobii A. Milne-Edwards, 1880
= Sympagurus grimaldii A. Milne-Edwards &
Bouvier, 1897
Parapagurus richeri Lemaitre, 1999
Parapagurus saintlaurentae Lemaitre, 1999
Parapagurus stenorhinus Lemaitre, 1999
Parapagurus wolffi Lemaitre, 1999
Probeebei Boone, 1926
= Probeebei Boone, 1926 (type species Probeebei
mirabilis Boone, 1926, by monotypy; gender
masculine)
= Planopagurus Wolff, 1960 (type species
Planopagurus galathea Wolff, 1960, by monotypy;
gender masculine)
39

Probeebei mirabilis Boone, 1926
= Planopagurus galathea Wolff, 1960
Strobopagurus Lemaitre, 1989
= Strobopagurus Lemaitre, 1989 (type species
Sympagurus gracilipes A. Milne-Edwards, 1891,
Strobopagurus breviacus Lemaitre, 2004
Strobopagurus gracilipes (A. Milne-Edwards, 1891)
[Sympagurus]
Strobopagurus sibogae (de Saint Laurent, 1972)
[Parapagurus]
= Parapagurus kilburni Kensley, 1973
Sympagurus Smith, 1883
= Sympagurus Smith, 1883 (type species Sympagurus
pictus Smith, 1883, by monotypy; gender
masculine)
Sympagurus acinops Lemaitre, 1989
Sympagurus affinis (Henderson, 1888) [Parapagurus]
Sympagurus andersoni (Henderson, 1896)
[Parapagurus]
= Parapagurus andersoni var. brevimanus Alcock,
1901
Sympagurus aurantium Lemaitre, 2004
Sympagurus brevipes (de Saint Laurent, 1972)
[Parapagurus]
Sympagurus burkenroadi Thompson, 1943
= Sympagurus papposus Lemaitre, 1996
Sympagurus chani Lemaitre, 2004
Sympagurus dimorphus (Studer, 1883) [Eupagurus]
= Parapagurus brevimanus Balss, 1911
= ? Eupagurus modicellus Stebbing, 1914
= Sympagurus arcuatus var. johnstoni Hale, 1941
= Sympagurus arcuatus var. mawsoni Hale, 1941
Sympagurus dofleini (Balss, 1912) [Parapagurus]
= Parapagurus ijimai Terao, 1913
= Parapagurus rectichela Zarenkov, 1990
Sympagurus pictus Smith, 1883
Sympagurus planimanus (de Saint Laurent, 1972)
[Parapagurus]
Sympagurus poupini Lemaitre, 1994
Sympagurus soela Lemaitre, 1996
Sympagurus spinimanus (Balss, 1911) [Parapagurus]
Sympagurus symmetricus Lemaitre, 2004
Sympagurus trispinosus (Balss, 1911) [Parapagurus]
= Parapagurus arcuatus var. trispinosa Balss,
1911
Sympagurus villosus Lemaitre, 1996
Tsunogaipagurus Osawa, 1995
= Tsunogaipagurus Osawa, 1995 (type species
Parapagurus chuni Balss, 1911, by original
Tsunogaipagurus chuni (Balss, 1911) [Parapagurus]
Tylaspis Henderson, 1885
{10}
= Tylaspis Henderson, 1885 (type species Tylaspis
anomala Henderson, 1885, by monotypy; gender
feminine)
Tylaspis anomala Henderson, 1885
Typhlopagurus de Saint Laurent, 1972
= Typhlopagurus de Saint Laurent, 1972 (type species
Typhlopagurus foresti de Saint Laurent, 1972, by
Typhlopagurus foresti de Saint Laurent, 1972
NOTES
{1}
In her revision of Parapagurus, de Saint Laurent

(1972) recognized three subspecies of A. MilneEdwardss bicristatus, Parapagurus bicristatus
bicristatus (A. Milne-Edwards, 1880), P. b. gracilis
Henderson, 1888, and P. b. indicus (Alcock, 1905b).
All were elevated to specific status by Lemaitre
(1989).
{2}
This species was described by de Saint Laurent

(1974) as Parapagurus curvispina and later placed
in Paragiopagurus by Lemaitre (1996). Examination
of type material, however, has revealed that it needs
to be formally assigned to Oncopagurus as a new
combination (Lemaitre, in prep.).
{3}
According to Lemaitre (1996), de Saint Laurent (1972)

questionably synonymized Eupagurus brevimanus
with Alcocks (1894) Parapagurus monstrosus.
Lemaitre (1996) found insufficient information to
confirm or deny the synonymy, and presumed that
Yokoyas (1933) specimens, like most of his other
material, were no longer extant.
{4}
Zhadan (1997: 67, fig. 9) did not provide a specific

name, and only noted that his material was intermediate
between O. tuamotu and O. cidaris
{5}
Formal descriptions are in preparation by one of the

authors (RL).
{6}
De Saint Laurents (1972) three subspecies,

Parapagurus acutus acutus de Saint Laurent, 1972,
P. a. bicarinatus de Saint Laurent, 1972, and P.
a. hirsutus de Saint Laurent, 1972, were assigned
to Sympagurus Smith, 1883 by Lemaitre (1989).
Subsequently Lemaitre (1996) elevated these three
subspecies to specific status and assigned them to
Paragiopagurus Lemaitre, 1996.
{7}
This is a senior homonym of Parapagurus abyssorum

Henderson, 1888.
{8}
Replacement name for Parapagurus abyssorum

Henderson, 1888, a junior homonym of Parapagurus
abyssorum (Filhol, 1885)
{9}
All the subspecies of Parapagurus pilosimanus Smith,

1879 proposed by de Saint Laurent (1972) were
elevated to specific status by Lemaitre (1989).
{10} De Grave et al. (2009) incorrectly attributed authorship

of the genus Tylaspis to Tizard, Mosely, Buchanan
& Murray (1885), presumably because they were the
editors of the narrative of the Challenger cruise.
However, the report on the Anomura (pp. 897901)
40

actually was written by Henderson, as Tizard et al.
acknowledged. The illustration and brief diagnosis of
Tylaspis n. gen. et sp. by Henderson (1885: 900901,
fig. 329) meet ICZN requirements for publication. The
correct author of this genus is therefore Henderson
(1885).
= Pylodietes miersi McGinitie & McGinitie, 1949

(misspelling of Pylocheles)
Subfamily Pomatochelinae Stebbing, 1914
Pomatocheles Miers, 1879
= Pomatocheles Miers, 1879 (type species
Pomatocheles jeffreysii Miers, 1879, by monotypy;
gender masculine)
Pomatocheles gaillardi Forest, 1987
Pomatocheles jeffreysii Miers, 1879
Pomatocheles stridulans Forest, 1987
Family Pylochelidae Bate, 1888

Subfamily Pylochelinae Bate, 1888
Bathycheles Forest, 1987
{1}
= Pylocheles (Bathycheles) Forest, 1987 (type species
Pylocheles (Bathycheles) incisus Forest 1987, by
Bathycheles crosnieri (Forest, 1987) [Pylocheles
(Bathycheles)]
Bathycheles cubensis (Ortiz & Gmez, 1986)
[Pylocheles]
= Pylocheles (Bathycheles) chacei Forest, 1987
Bathycheles incisus (Forest, 1987) [Pylocheles
(Bathycheles)]
Bathycheles integer (Forest, 1987) [Pylocheles
(Bathycheles)]
= Pylocheles (Bathycheles) profundus Forest,
1987
Bathycheles macgilchristi (Alcock, 1905) [Chiroplatea]
(misspelling of Cheiroplatea)
Bathycheles phenax McLaughlin & Lemaitre, 2009
Cheiroplatea Bate, 1888
= Cheiroplatea Bate, 1888 (type species Cheiroplatea
cenobita Bate, 1888, by monotypy; gender
masculine)
= Chiroplatea Ortmann, 1892 (misspelling of
Cheiroplatea)
Cheiroplatea cenobita Bate, 1888
Cheiroplatea laticauda Boas, 1926
Cheiroplatea mitoi Miyake, 1978
Cheiroplatea pumicicola Forest, 1987
Cheiroplatea scutata Ortmann, 1892
= Chiroplatea scutata Ortmann, 1892 (misspelling
of Cheiroplatea)
Cheiroplatea stenurus Forest, 1987
Pylocheles A. Milne-Edwards, 1880
= Pylocheles A. Milne-Edwards, 1880 (type species
Pylocheles agassizi A. Milne-Edwards, 1880, by
Pylocheles agassizi A. Milne-Edwards, 1880
= Pylocheles partitus Benedict, 1901
Pylocheles mortensenii Boas, 1926
= Pylocheles rigidus Yokoya 1933
Xylocheles Forest 1987
{2}
= Xylocheles Forest, 1987 (type species Pylocheles
(Xylocheles) macrops Forest, 1987, by original
Xylocheles macrops (Forest, 1987) [Pylocheles
(Xylocheles)]
Xylocheles miersi (Alcock & Anderson, 1899)
[Pylocheles]
Subfamily Trizochelinae Forest 1987

Tribe Cancellochelini Forest, 1987
Cancellocheles Forest, 1987
= Cancellocheles Forest, 1987 (type species
Pomatocheles sculptipes Miyake, 1978, by
Cancellocheles sculptipes (Miyake, 1978)
[Pomatocheles]
Tribe Mixtopagurini Bouvier, 1895
{3}
Mixtopagurus A. Milne-Edwards, 1880
= Mixtopagurus A. Milne-Edwards, 1880 (type species
Mixtopagurus paradoxus A. Milne-Edwards,
Mixtopagurus paradoxus A. Milne-Edwards, 1880
Tribe Parapylochelini
Parapylocheles Alcock, 1901
= Parapylocheles Alcock, 1901 (type species
Pylocheles scorpio Alcock, 1894, by monotypy;
gender masculine)
Parapylocheles scorpio (Alcock, 1894)
Tribe Trizochelini Forest, 1987
Forestocheles McLaughlin & Lemaitre, 2009
= Forestocheles McLaughlin & Lemaitre, 2009
(type species Trizocheles perplexus Forest, 1987,
Forestocheles perplexus (Forest, 1987) [Trizocheles]
Trizocheles Forest, 1987
= Trizocheles Forest, 1987 (type species Pylocheles
spinosus Henderson, 1888, by original designation;
gender masculine)
Trizocheles albatrossi Forest, 1987
Trizocheles balssi (Stebbing, 1914) [Pomatocheles]
Trizocheles boasi Forest, 1987
= Trizocheles gracilis Forest, 1987
Trizocheles brachyops Forest & de Saint Laurent, 1987
Trizocheles brevicaulis (Boas, 1926) [Mixtopagurus]
Trizocheles caledonicus Forest, 1987
Trizocheles hoensonae McLaughlin & Lemaitre, 2009
Trizocheles laurentae Forest, 1987
Trizocheles longicaulis (Boas, 1926) [Mixtopagurus]
Trizocheles loquax Forest, 1987
Trizocheles manningi Forest, 1987
Trizocheles mendanai McLaughlin & Lemaitre, 2009
41

Ahyong, S. T. & T.-Y. Chan, 2010. Lithodes formosae, a new species
of king crab from Taiwan (Crustacea: Decapoda: Lithodidae).
Zootaxa, 2332: 6168.
Trizocheles moosai Forest, 1987

Trizocheles mutus Forest, 1987,
Trizocheles pilgrimi Forest & McLaughlin, 2000
Trizocheles pulcher Forest, 1987
Trizocheles sakaii Forest, 1987
Trizocheles spinosus (Henderson, 1888)
= Trizocheless spinosus bathamae Forest & de Saint
Laurent, 1987
Ahyong, S. T. & E. W. Dawson, 2006. Lithodidae from the Ross

Sea, Antarctica, with descriptions of two new species (Crustacea:
Decapoda: Anomura). Zootaxa, 1303: 4568.
Ahyong, S. T. & D. OMeally, 2004. Phylogeny of the Decapoda
Reptantia: Resolution using three molecular loci and morphology.
Raffles Bulletin of Zoology, 52: 673693.
Ahyong, S. T., E. Macpherson & T.Y. Chan, 2010. Lithodoidea.
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Anomurans (Hippodea, Lithodoidea and Porcellanidae).
National Taiwan Ocean University, Keelung. Pp. 4266.
NOTES
{1}
{2}
{3}
Authorship of Bathycheles was incorrectly credited

to McLaughlin & Lemaitre, 2009, by De Grave et al.
(2009). Bathycheles was proposed by Forest (1987)
as a subgenus of Pylocheles. In their reassessment
of pylochelid classification, McLaughlin & Lemaitre
(2009) elevated Bathycheles to full generic rank, but in
accordance with ICZN Article 43.1 (1999) the author
and date for the taxon remain as Forest (1987).
Ahyong, S. T., K. E, Schnabel & E. W. Maas, 2009. Anomuran

Phylogeny: New insights from molecular data. In: Martin, J.
W., K. A. Crandall & D. L. Felder (eds.), Decapod Crustacean
Phylogenetics. Crustacean Issues 18. CRC Press, Boca Raton,
FL, New York, London. Pp. 399414.
Alcock, A., 1894. Natural history notes from H.M. Indian Marine
Survey Steamer Investigator, Commander R.F. Hoskyn, R.N.,
commanding.Series II., No. 1. On the results of deep-sea
dredging during the season 1890-91. Annals and Magazine of
Natural History, (6)13: 225245; 321334; 400411.
Authorship of Xylocheles was incorrectly credited

to McLaughlin & Lemaitre, 2009 by De Grave et al.
(2009). Xylocheles was proposed by Forest (1987)
as a subgenus of Pylocheles. In their reassessment
of pylochelid classification, McLaughlin & Lemaitre
(2009) elevated Xylocheles to full generic rank, but in
accordance with ICZN Article 43.1 (1999) the author
and date for the taxon remain as Forest ( 1987).
Alcock, A., 1901. Catalogue of the Indian decapod Crustacea

in the collection of the Indian Museum. Part 1. Brachyura.
Fascicle I. Introduction and dromides or Dromiacea (Brachyura
Primigenia). Indian Museum, Calcutta. 80 pp.
Alcock, A., 1905a. Marine Crustaceans. XIV. Paguridae. In:
Gardiner, J. S. (ed.). The fauna and geography of the Maldive
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the years 1899 and 1900, 2: 827835. University Press,
Cambridge.
Although the tribe name Mixtopaguriens was

Latinized by A. Milne-Edwards & Bouvier (1899),
Forest (1987) correctly assigned the authorship and
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11f (iii) of the ICZN Code (1985).
Alcock, A., 1905b. Anomura. Fasc. I. Pagurides. Catalogue of

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Alcock, A. & A. R. Anderson, 1899. Natural history notes from
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127.
Family Pylojacquesidae McLaughlin & Lemaitre, 2001

Pylojacquesia McLaughlin & Lemaitre, 2001
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species Pylojacquesia colemani McLaughlin &
Lemaitre, 2001, by original designation; gender
feminine)
Pylojacquesia colemani McLaughlin & Lemaitre, 2001
Lemaitreopsis McLaughlin, 2007
= Lemaitreopsis McLaughlin, 2007 (type species
Lemaitreopsis holmi McLaughlin, 2007, by
Lemaitreopsis holmi McLaughlin, 2007
Anderson, E. M. & G. M. Cailliet, 1974. Crab and snailfish

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Asakura, A., 1999. Icelopagurus tuberculosus, a distinct new
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of the Lomidae (Crustacea: Decapoda: Anomala). Journal of
Crustacean Biology, 3(3): 431437.
McLaughlin, P. A., 2007a. A new genus and new species in the

hermit crab family Pylojacquesidae (Crustacea: Anomura:
Paguroidea: Paguridae). Proceedings of the Biological Society
McLaughlin, P. A., 1983b. Hermit crabsAre they really

polyphyletic? Journal of Crustacean Biology, 3(4): 608621.
McLaughlin, P. A., 1986. Three new genera and species of hermit
crabs (Crustacea, Anomura, Paguridae) from Hawaii. Journal
of Crustacean Biology, 6(4): 789803.
McLaughlin, P. A., 2007b. A new species of Pteropagurus

McLaughlin & Rahayu and a new genus and species of
scaphopod dweller (Decapoda: Anomura: Paguroidea:
Paguridae). Zoosystema, 29(3): 503513.
McLaughlin, P. A., 1988. The rediscovery of Ceratopagurus Yokoya

and a new genus for Pagurus piercei Wass (Crustacea: Anomura:
Paguridae). Crustaceana, 55(3): 257267.
McLaughlin, P. A., 2007c. New records and a new species in the

genus Turleania McLaughlin, 1997 (Decapoda, Anomura,
Paguridae) from MUSORSTOM cruises, with a key to species.
Zoosystema, 29(3): 583593.
McLaughlin, P. A., 1994. A new genus and two new species of

deep-water hermit crabs (Decapoda: Anomura: Paguridae) from
the Southern Ocean. Proceedings of the Biological Society of
Washington, 107(3): 469481.
McLaughlin, P. A., 2008a. A new species of the hermit crab genus

Cancellus (Decapoda: Anomura: Paguroidea: Diogenidae) from
the PANGLAO Expeditions to the Philippine Islands. Raffles
Bulletin of Zoology, Supplement 19: 7380.
McLaughlin, P. A., 1997. Crustacea Decapoda: Hermit crabs of

the family Paguridae from the KARUBAR cruise in Indonesia.
In: Crosnier, A. & P. Bouchet (eds.), Rsultats des Campagnes
MUSORSTOM, 16. Mmoires du Musum national dHistoire
naturelle, 172: 433572.
McLaughlin, P. A., 2008b. Australian hermit crabs of the genera

Paguristes Dana, Stratiotes Thomson, and Pseudopaguristes
McLaughlin (Crustacea: Anomura: Paguroidea: Diogenidae).
In: Jones, D.S. (ed.), Crustaceans collected by the Western
Australian Museum/Woodside Energy Ltd. Partnership to
explore the marine biodiversity of the Dampier Archipelago,
Western Australia, 1998-2002. Records of the Western
Australian Museum, Supplement 73: 185273.
McLaughlin, P. A., 1998. Hermit crabs of the genus Nematopagurus

(Crustacea: Decapoda: Paguridae) from southeastern South
Africa and Madagascar: New records and new species.
Zoosystema, 20(2): 335358.
McLaughlin, P. A., 2000. Crustacea: Decapoda: Species of
Porcellanopagurus Filhol and Solitariopagurus Trkay
(Paguridae), from the New Caledonia area, Vanautu, and the
Marquesas: new records, new species. In: Crosnier, A. (ed.),
Rsultats des Campagnes MUSORSTOM, 21. Mmoires du
Musum national dHistoire naturelle, 184: 389414.
McLaughlin, P. A., 2008c. Hermit crabs (Crustacea: Anomura:

Paguroidea) of the Dampier Archipelago, Western Australia.
In: Jones, D.S. (ed.), Crustaceans collected by the Western
Australian Museum/Woodside Energy Ltd. Partnership to
explore the marine biodiversity of the Dampier Archipelago,
Western Australia, 1998-2002. Records of the Western
Australian Museum, Supplement 73: 273288.
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McLaughlin, P. A. & A. Asakura, 2004. Reevaluation of the hermit
crab genus Parapagurodes McLaughlin & Haig (Decapoda:
Anomura: Paguridae) and a new genus for Parapagurodes
doederleini (Doflein). Proceedings of the Biological Society
McLaughlin, P. A. & P. J. Hogarth, 1998. Hermit crabs (Decapoda:

Anomura: Paguridea) from the Seychelles. Zoologische
Verhandlingen, 318: 148.
McLaughlin, P. A. & L. B. Holthuis, 2001. In pursuit of J. F.
W. Herbsts species of Diogenes (Anomura: Paguridea:
Diogenidae). Journal of Crustacean Biology, 21(1): 257273.
McLaughlin, P. A. & J. Baily-Brock, 1975. A new Hawaiian

hermit crab of the genus Trizopagurus (Crustacea: Decapoda:
Diogenidae), with notes on its behavior. Pacific Science, 29(3):
259266.
McLaughlin, P. A. & L. B. Holthuis, 2002. Case 3183. Pagurus

clypeatus Fabricius, 1787 (currently Coenobita clypeatus;
Crustacea, Decapoda): proposed replacement of syntypes by a
neotype. Bulletin of Zoological Nomenclature, 59(1): 1723.
McLaughlin, P. A. & J. H. Brock, 1974. A new species of hermit

crab of the genus Nematopagurus (Crustacea: Decapoda:
Paguridae) from Hawaii. Proceedings of the Biological Society
McLaughlin, P. A. & J. P. Hoover, 1996. A new species of

Aniculus Dana (Decapoda: Anomura: Diogenidae) from Hawaii.
299305.
McLaughlin, P. A. & P. F. Clark, 1997. A review of the Diogenes

(Crustacea, Paguridea) hermit crabs collected by Bedford and
Lanchester from Singapore, and from the Skeat Expedition to
the Malay Peninsula, with a description of a new species and
notes on Diogenes intermedius De Man, 1892. Bulletin Natural
History Museum, London (Zoology), 63(1): 3349.
McLaughlin, P. A. & G. C. Jensen, 1996. A new hermit crab

species of the genus Parapagurodes from the eastern Pacific,
with a description of its first zoeal stage. Journal of Natural
History, 30: 841854.
McLaughlin, P. A. & K. Konishi, 1994. Pagurus imafukui, a
new species of deep-water hermit crab (Crustacea: Anomura:
Paguridea), with notes on its larvae. Publications of the Seto
Marine Biological Laboratory, 36(4): 211222.
McLaughlin, P. A. & J. Forest, 1997. Crustacea Decapoda:

Diacanthurus gen. nov., a new genus of hermit crabs (Paguridae)
with both Recent and fossil representation, and the descriptions
of two new species. In: Crosnier, A. (ed.), Rsultats des
McLaughlin, P. A. & R. Lemaitre, 1993. A review of the hermit

crab genus Paguritta (Decapoda: Anomura: Paguridae) with
descriptions of three new species. Raffles Bulletin of Zoology,
41(1): 129.
McLaughlin, P. A. & J. Forest, 1999. Hermit crabs of the genus

Pagurus Fabricius (Crustacea: Decapoda: Paguridae) from
southeastern South Africa. Annals of the South African Museum,
105(7): 297344.
McLaughlin, P. A. & R. Lemaitre, 1997. Carcinization fact or

fiction? I. Evidence from adult morphology. Contributions to
Zoology, Amsterdam, 67(2): 79123.
McLaughlin, P. A. & R. H. Gore, 1985. A new species of

Allodardanus (Crustacea: Decapoda: Diogenidae) from the
western Atlantic. Crustaceana, 49(1): 3641.
McLaughlin, P. A. & R. Lemaitre, 2001a. Revision of Pylopagurus

and Tomopagurus (Crustacea: Decapoda: Paguridae), with
descriptions of new genera and species. Part VI. Pylopagurus
Milne-Edwards and Bouvier, Haigia McLaughlin, and
Pylopaguridium new genus. Proceedings of the Biological
McLaughlin, P. A. & S. W. Gunn, 1992. Revision of Pylopagurus

and Tomopagurus (Crustacea: Decapoda: Paguridae), with the
descriptions of new genera and species: Part IV. Lophopagurus
McLaughlin and Australeremus McLaughlin. Memoirs of the
Museum of Victoria, 53(1): 4399.
McLaughlin, P. A. & R. Lemaitre, 2001b. Aspects of evolution in

the anomuran superfamily Paguroidea: one larval prospective.
Invertebrate Reproduction and Development, 38(3): 159169.
McLaughlin, P. A. & J. Haig, 1973. On the status of Pagurus

mertensii Brandt, with descriptions of a new genus and two
new species from California (Crustacea: Decapoda: Paguridae).
Bulletin of the Southern California Academy of Sciences, 72:
113136.
McLaughlin, P. A. & R. Lemaitre, 2001c. A new family for a

new genus and new species of hermit crab of the superfamily
Paguroidea (Decapoda: Anomura) and its phylogenetic
implications. Journal of Crustacean Biology, 21(4): 1062
1076.
McLaughlin, P.A. & J. Haig, 1984. A review of Pagurixus

(Decapoda, Anomoura, Paguridae) and descriptions of new
species. Crustaceana, 47(2): 121-148.
McLaughlin, P. A. & R. Lemaitre, 2004. The discovery of two

new species of Lithopagurus Provenzano, 1968 and the first
records of the genus in the western Pacific. Zoosystema, 26(3):
483494.
McLaughlin, P. A. & J. Haig, 1989. On the status of Pylopaguropsis

zebra Henderson, P. magnimanus (Henderson), and Galapagurus
teevanus Boone, with descriptions of seven new species of
Pylopaguropsis (Crustacea: Anomura: Paguridae). Micronesica,
22: 123171.
McLaughlin, P. A. & R. Lemaitre, 2007. A new and distinctive

species of the hermit crab genus Catapaguropsis (Crustacea:
Decapoda: Anomura: Paguroidea: Paguridae) from the South
China Sea. Zootaxa, 1560: 3141.
McLaughlin, P. A. & J. Haig, 1993. Two new species of the Pacific

component of the provenzanoi of Pagurus (Decapoda, Anomura,
Paguridae) and a key to the regional species. Bulletin of Marine
Science, 52(2): 642668.
McLaughlin, P. A. & R. Lemaitre, 2009. A new classification

of the Pylochelidae (Decapoda: Anomura: Paguroidea) and
descriptions of new taxa. Raffles Bulletin of Zoology. Supplement
20: 159231.
McLaughlin, P. A. & J. Haig, 1995. A new species of Gorepagurus

McLaughlin (Decapoda: Anomura: Paguridae) from the Pacific,
and a comparison with its Atlantic counterpart. Proceedings of
the Biological Society of Washington, 108(1): 6875.
McLaughlin, P. A. & T. Murray, 1990. Clibanarius fonticola n.

sp. (Anomura: Paguridea: Diogenidae), from a freshwater pool
on Espiritu Santo, Vanuatu. Journal of Crustacean Biology,
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McLaughlin, P. A. & J. Haig, 1996. A new genus for Anapagrides

sensu De Saint Laurent-Dechanc, 1966 (Anomura: Paguridae)
and descriptions of four new species. Proceedings of the
Biological Society of Washington, 109(1): 7590.
McLaughlin, P. A. & J. Okuno, 2010. A new and distinctive Japanese

species of Nematopagurus (Crustacea, Decapoda, Paguridae),
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McLaughlin, P. A. & C. Pittman, 2002. Reinstatement and

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Fig. 1. Lithodoidea. Representatives of Hapalogastridae Brandt, 1850 (A-G, in situ; H, preserved specimen): A, Acantholithodes hispidus
(Stimpson, 1860), Hood Canal, Washington (G. Jensen); B, Dermaturus mandtii Brandt, 1850, Pribilof Islands, Alaska (G. Jensen); C,
Hapalogaster cavicauda Stimpson, 1859, Cayucos, California (G. Jensen); D, Hapalogaster grebnitzkii Schalfeew, 1892, Prince William
Sound, Alaska (G. Jensen); E, Hapalogaster mertensii Brandt, 1850, Whidbey Island, Washington (G. Jensen); F, Oedignathus inermis
(Stimpson, 1860), Neah Bay,Washington (G. Jensen); G, H, Placetron wosnessenskii Schalfeew, 1892, Neah Bay, Washington (G) (G.
Jensen); Alaska, USNM 276166 (H) (R. Lemaitre). USNM = National Museum of Natural History, Smithsonian Institution, Washington,
D.C.
66
67
Fig. 2. Lithodoidea and Lomisoidea. Representatives of Lithodidae Samouelle, 1819 (A-G) and Lomisidae Bouvier, 1895 (H): A, Lithodes
aequispinus Benedict, 1895, Bering Sea, USNM 259215 (preserved specimen, R. Lemaitre); B, Neolithodes duhameli Macpherson, 2004,
Crozet Islands, Southern Ocean (J.-F. Dejouannet); C, Paralomis arae Macpherson, 2001, Fiji Islands (J.-F. Dejouannet); D, Paralomis
dofleini Balss, 1911, Sagami Sea, off Boso Pensinsula, Japan, CBM-ZC 8498 (T. Komai); E, Paralomis japonicus Balss, 1911, Sagami Sea,
off Boso Pensinsula, Japan, CBM-ZC 8499 (T. Komai); F, Paralomis mendagnai Macpherson, 2003, Salomon Islands (J.-F. Dejouannet);
G, Paralomis odawarai (Sakai, 1980), Sagami Sea, Boso Pensinsula, Japan, CBM-ZC 9843 (T. Komai); H, Lomis hirta (Lamarck, 1818),
Southern Australia ( Museum Victoria, Melbourne, M. Marmach). CBM = Natural History Museum and Institute, Chiba; USNM =
National Museum of Natural History, Smithsonian Institution, Washington, D.C.
68
69
Fig. 3. Paguroidea. Representatives of Coenobitidae Dana, 1851 (in situ except A, C, D): A, B, Birgus latro (Linnaeus, 1767), Tuamotu,
French Polynesia (A, J. Poupin; B, O. Gargominy); C, Coenobita clypeatus (Fabricius, 1787), Belize, ULLZ 11972 (D. L. Felder); D,
Coenobita cavipes Stimpson, 1858, Mayotte Island (J. Poupin & R. Clva); E, Coenobita perlatus H. Milne Edwards, 1837, Tuamotu,
French Polynesia (J. Poupin); F, Coenobita rugosus H. Milne Edwards, 1837, Moorea, French Polynesia (P. Bacchet); G, Coenobita
spinosus H. Milne Edwards, 1837, Wallis and Futuna (J. Poupin); H, Coenobita violascens Heller, 1862, Ryukyu Islands, Japan, specimen
not collected (T. Komai). ULLZ = University of Louisiana at Lafayette Zoological Collections.
70
71
Fig. 4. Paguroidea. Representatives of Diogenidae Ortmann, 1892 (F, H, in situ): A, Aniculus aniculus (Fabricius, 1787), Guam (G. Paulay);
B, Aniculus erythraeus Forest, 1984, Panama, eastern Pacific, LACM (D. L. Felder); C, Aniculus maximus Edmondson, 1952, Guam (G.
Paulay); D, Aniculus retipes Lewinsohn, 1982, Guam (G. Paulay); E, Aniculus ursus (Olivier, 1812), Guam (G. Paulay); F, Calcinus elegans
(H. Milne Edwards, 1836), Wallis and Futuna (J. Poupin); G, Calcinus gaimardii (H. Milne Edwards, 1848), Philippines (T.-Y. Chan); H,
Calcinus gouti Poupin, 1997, Tuamotu, French Polynesia (G. Paulay). LACM = Natural History Museum of Los Angeles County.
72
73
Fig. 5. Paguroidea. Representatives of Diogenidae Ortmann, 1892 (in situ except B, D, G, H): A, Calcinus guamensis Wooster, 1984,
Moorea, French Polynesia (J. Poupin); B, Calcinus laevimanus (Randall, 1840), Philippines (T.-Y. Chan); C, Calcinus latens (Randall,
1840), Hawaii, CBM-ZC 10013 (E. Myorin); D, Calcinus lineapropodus Morgan & Forest, 1991, Philippines (T.-Y. Chan), E, Calcinus
mclaughlinae Poupin & Bouchard, 2006, Clipperton Island (J. Poupin); F, Calcinus nitidus Heller, 1865, Moorea, French Polynesia (J.
Poupin); G, Calcinus pulcher Forest, 1958, Philippines (T.-Y. Chan); H, Calcinus seurati Forest, 1951, Kikai Island, Amami Islands,
Japan, CBM-ZC 9556 (T. Komai). CBM = Natural History Museum and Institute, Chiba.
74
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Fig. 6. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Cancellus panglaoensis McLaughlin, 2008, Philippines (T.-Y. Chan);
B, Ciliopagurus krempfi (Forest, 1952), Japan (T. Komai); C, Ciliopagurus tricolor Forest, 1995, Moorea, French Polynesia (A. Anker);
D, Clibanarius ambonensis Rahayu & Forest, 1993, Ryukyu Islands, Japan, CBM-ZC (T. Komai); E, Clibanarius corallinus (H. Milne
Edwards, 1848), Tuamotu, French Polynesia (G. Paulay); F, Clibanarius demani Buitendijk, 1937, Oura Bay, Ryukyu Islands, Japan (T.
Komai); G, Clibanarius englaucus Ball & Haig, 1972, Ryukyu Islands, Japan, CBM-ZC (T. Komai); H, Clibanarius rutilus Rahayu, 1999,
Bali, Indonesia, CBM-ZC 9990 (T. Komai). CBM = Natural History Museum and Institute, Chiba.
76
77
Fig. 7. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Clibanarius snelliusi Buitendijk, 1937, Ryukyu Islands, Japan,
CBM-ZC (T. Komai); B, Clibanarius striolatus Dana, 1852, Ryukyu Islands, Japan, CBM-ZC 9997 (T. Komai); C, Clibanarius virescens
(Krauss, 1843), Ryukyu Islands, Japan (T. Komai); D, Dardanus australis Forest & Morgan, 1991, Austral Islands, French Polynesia (J.
Poupin); E, Dardanus brachyops Forest, 1962, Ryukyu Islands, Japan, Okinawa Churaumi Aquarium (T. Komai); F, Dardanus deformis (H.
Milne Edwards, 1836), Ryukyu Islands, Japan, CBM-ZC 8000 (T. Komai); G, Dardanus gemmatus (H. Milne Edwards, 1848), Tuamotu,
French Polynesia (G. Paulay); H, Dardanus guttatus (Olivier, 1812), Ryukyu Islands, Japan, CBM-ZC 7999 (T. Komai). CBM = Natural
History Museum and Institute, Chiba.
78
79
Fig. 8. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Dardanus lagopodes (Forskl, 1775), Kagoshima Prefecture, Japan,
CBM-ZC 8687 (T. Komai); B, Dardanus robustus Asakura, 2006, Ogasawara Islands, Japan (H. Tachikawa); C, Dardanus scutellatus
(H. Milne Edwards, 1848), Ryukyu Islands, Japan, CBM-ZC 9995 (T. Komai); D, Dardanus woodmasoni (Alcock, 1905), Ryukyu
Islands, Japan, CBM-ZC 9989 (T. Komai); E, Diogenes alias McLaughlin & Holthuis, 2001, Kerara, India, CBM-ZC 10007 (T. Komai);
F, Diogenes cf. avarus Heller, 1865, Western Australia, CBM-ZC 9996 (T. Komai); G, Diogenes cf. tirmiziae Siddiqui & McLaughlin,
2003, Osumi Islands, Japan, CBM-ZC (T. Komai); H, Diogenes cf. manaarensis (Henderson, 1893), Kerara, India, CBM-ZC 10011 (T.
Komai). CBM = Natural History Museum and Institute, Chiba.
80
81
Fig. 9. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Diogenes miles (Fabricius, 1787), Kerara, India, CBM-ZC 10008
(T. Komai); B, Diogenes nitidimanus Terao, 1913, Japan, CBM-ZC (T. Komai); C, Diogenes sp. 1, Ryukyu Islands, Japan, CBM-ZC
(T. Komai ); D, Diogenes spinifrons (De Haan, 1849), Tateyama, Boso Peninsula, Japan, CBM-ZC 8423 (T. Komai); E, Diogenes sp. 2,
Tatsunokuchi, Nagasaki, Japan, CBM-ZC (T. Komai); F, Paguristes albimaculatus Komai, 2001, Ogasawara Islands, Japan, CBM-ZC
(T. Komai); G, Paguristes cadenati Forest, 1954, Gulf of Mexico ULLZ 7043 (D. L. Felder); H, Paguristes digitalis Stimpson, 1858,
Uraga Strait, Tokyo Bay, Japan, CBM-ZC 9999 (T. Komai). CBM = Natural History Museum and Institute, Chiba; ULLZ = University
of Louisiana at Lafayette Zoological Collections.
82
83
Fig. 10. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Paguristes gonagrus (H. Milne Edwards, 1836), Taiwan (T.-Y.
Chan); B, Paguristes jalur Morgan, 1992, Ryukyu Islands, Japan, CBM-ZC 9981 (T. Komai); C, Paguristes miyakei Forest & McLaughlin,
1998, Uraga Strait, Tokyo Bay, Japan (T. Komai); D, Paguristes palythophilus Ortmann, 1892, Uraga Strait, Tokyo Bay, Japan, CBMZC 4698 (T. Komai); E, Paguristes cf. puniceus Henderson, 1896, Uraga Strait, Japan, CBM-ZC (T. Komai); F, Paguristes cf. pusillus
Henderson, 1896, Taiwan, NTOU (T.-Y. Chan); G, Paguristes sericeus A. Milne Edwards, 1880, Gulf of Mexico, ULLZ 7118 (D. L.
Felder); H, Paguristes versus Komai, 2001, Izu Islands, Japan, CBM-ZC (T. Komai). CBM = Natural History Museum and Institute,
Chiba; NTOU = National Taiwan Ocean University; ULLZ = University of Louisiana at Lafayette Zoological Collections.
84
85
Fig. 11. Paguroidea. Representatives of Diogenidae Ortmann, 1892: A, Paguropsis typica Henderson, 1888, Philippines (T.-Y. Chan); B,
Pseudopaguristes bicolor Asakura & Kosuge, 2004, Satsunan Islands, Japan, CBM-ZC 9986 (T. Komai); C, Pseudopaguristes laurentae
(Morgan & Forest, 1991), Izu Islands, Japan, CBM-ZC (T. Komai); D, Pseudopaguristes monoporus (Morgan, 1987), Kochi Prefecture,
Japan, CBM-ZC 9993 (T. Komai); E, Areopaguristes japonicus (Miyake, 1961), Boso Peninsula, Japan, CMNH-ZC 672 (J. Okuno); F,
Areopaguristes nigroapiculus (Komai, 2009), Uraga Strait, Tokyo Bay, Japan, paratype, NSMT-Cr S45 (T. Komai); G, Areopaguristes
orbis (Komai, 2009), Izu Islands, Japan, paratype, CBM-ZC 9532 (T. Komai); H, Areopaguristes taenia (Komai, 1999), Ogasawara Islands,
Japan, CBM-ZC 9521 (T. Komai). CBM = Natural History Museum and Institute, Chiba; CMNH = Coastal Branch of the Natural History
Museum and Institute, Chiba; NSMT = National Museum of Nature and Science, Tokyo.
86
87
Fig. 12. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Anapagrides aequalis Komai, 1999, Sagami Sea, Japan, NSMT-Cr
S164 (T. Komai); B, Anapagurus cf. bonnieri Nobili, 1905, Ryukyu Islands, Japan (T. Komai); C, Boninpagurus sp., with rhizocephalan,
Sagami Bay, Japan, CBM-ZC 10000 (T. Komai); D, Catapaguroides japonicus de Saint Laurent, 1968, with rhizocephalan, Sagami Bay,
Japan, NSMT-Cr S165 (T. Komai); E, Catapaguroides sp., Ohsumi Islands, Japan (T. Komai); F, Catapagurus misakiensis Terao, 1914,
Izu Islands, Japan, Sagami Sea, Japan, NSMT-Cr S168 (T. Komai); G, Catapagurus sp., Ryukyu Islands, Japan, CBM-ZC (T. Komai); H,
Chanopagurus atopos Lemaitre, 2003, Taiwan (T.-Y. Chan). CBM = Natural History Museum and Institute, Chiba; NSMT = National
Museum of Nature and Science, Tokyo.
88
89
Fig. 13. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Decaphyllus spinicornis de Saint Laurent, 1968, Sagami Sea, Japan,
Boso Peninsula, Japan, CBM-ZC 8435 (T. Komai); B, Discorsopagurus tubicola Komai, 2003, Uraga Strait, Tokyo Bay, Japan, CBMZC 6813 (T. Komai); C, Elassochirus cavimanus (Miers, 1879), Iwate Prefecture, Japan, CBM-ZC 10005 (T. Komai); D, Iridopagurus
reticulatus Garca-Gmez, 1983, Carrie Bow Cay, Belize, ULLZ 10032 (D. L. Felder); E, Lophopagurus (Australeremus) triserratus
(Ortmann, 1892), Izu Islands, Japan, CBM-ZC (T. Komai); F, Nematopagurus gardineri Alcock, 1905, Ogasawara Islands, Japan, CBMZC (T. Komai); G, Nematopagurus kosiensis McLaughlin, 1998, Ogasawara Islands, Japan, CBM-ZC (T. Komai); H, Nematopagurus
cf. spinulosensoris McLaughlin & Brock, 1974, Sagami Sea, Japan, Sagami Sea, Japan, NSMT-Cr S178 (T. Komai). CBM = Natural
History Museum and Institute, Chiba; NSMT = National Museum of Nature and Science, Tokyo; ULLZ = University of Louisiana at
Lafayette Zoological Collections.
90
91
Fig. 14. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Nematopagurus tricarinatus (Stimpson, 1858), Ogasawara Islands,
Japan, CBM-ZC (T. Komai); B, Pagurixus anceps (Forest, 1957), Ryukyu Islands, Japan, CBM-ZC 9571 (T. Komai); C, Pagurixus
aurantiaca Komai, 2010, Ogasawara Islands, Japan, holotype, CBM-ZC 9644 (T. Komai); D, Pagurixus boninensis (Melin, 1939),
Ogasawara Islands, Japan, CBM-ZC (T. Komai); E, Pagurixus dissimilis Osawa & Komai, 2007, Kochi Prefecture, Japan, paratype,
CBM-ZC 9071 (T. Komai); F, Pagurixus haigae Komai & Osawa, 2007, Ryukyu Islands, Japan, holotype, CBM-ZC 8406 (T. Komai);
G, Pagurixus nomurai Komai & Asakura, 1995, Guam (G. Paulay); H, Pagurixus pseliophorus Komai & Osawa, 2006, Kochi Prefecture,
Japan, paratype, CBM-ZC 8533 (T. Komai). CBM = Natural History Museum and Institute, Chiba.
92
93
Fig. 15. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Pagurus alaini Komai, 1998, Ohsumi Islands, Japan, CBM-ZC 9982
(T. Komai); B, Pagurus angustus (Stimpson, 1858), Ryukyu Islands, Japan, CBM-ZC (T. Komai); C, Pagurus bernhardus (Linnaeus, 1758),
Belgium (C. dUdekem dAcoz); D, Pagurus brevidactylus (Stimpson, 1859), Gulf of Mexico, ULLZ 6753 (D. L. Felder); E, Pagurus
conformis De Haan, 1849, Uraga Strait, Japan, CBM-ZC 9998 (T. Komai); F, Pagurus erythrogrammus Komai, 2003, Kochi Prefecture,
Japan, CBM-ZC 9992 (T. Komai); G, Pagurus hirtimanus (Miers, 1880), Ryukyu Islands, CBM-ZC 8962 (T. Komai); H, Pagurus imafukui
McLaughlin & Konishi, 1994, Sagami Bay, Japan, NSMT-Cr S185 (T. Komai). CBM = Natural History Museum and Institute, Chiba;
NSMT = National Museum of Nature and Science, Tokyo; ULLZ = University of Louisiana at Lafayette Zoological Collections.
94
95
Fig. 16. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Pagurus japonicus (Stimpson, 1858), Izu Peninsula, Japan, CBM-ZC
6795 (T. Komai); B, Pagurus luticola Komai & Chan, 2006, Taiwan, holotype, NTOU A00805 (T.-Y. Chan); C, Pagurus nigrofascia
Komai, 1996, Chiba Port, Tokyo Bay, Japan, CBM-ZC 6772 (T. Komai); D, Pagurus quinquelineatus Komai, 2003, Boso Peninsula,
Japan, paratype, CBM-ZC 6622 (T. Komai); E, Pagurus rathbuni (Benedict, 1892), Sea of Japan, off Hokkaido, CBM-ZC 10002 (T.
Komai); F, Pagurus simulans Komai, 2000, Uraga Strait, Japan, paratype, CBM-ZC 5196 (T. Komai); G, Pagurus townsendi (Benedict,
1892), Hokkaido, Japan, HUMZ-C 1060 (T. Komai); H, Pagurus trigonocheirus (Stimpson, 1858), Hokkaido, Japan, HUMZ-C 1050 (T.
Komai). CBM = Natural History Museum and Institute, Chiba; HUMZ, Faculty of Fisheries, Hokkaido University; NTOU = National
Taiwan Ocean University.
96
97
Fig. 17. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Porcellanopagurus filholi de Saint Laurent & McLaughlin, 2000,
Uraga Strait, Tokyo Bay, Japan, CBM-ZC 10001 (T. Komai); B, Porcellanopagurus japonicus Balss, 1913, Izu Islands, Japan (T. Komai);
C, Porcellanopagurus nihonkaiensis Takeda, 1985, Sagami Bay, Japan, NSMT-Cr S194 (T. Komai); D, Porcellanopagurus platei Lenz,
1902, Hawaiian Islands, LACM CR 2006-013.1 (J. W. Martin); E, Propagurus obtusifrons (Ortmann, 1892), Uraga Strait, Tokyo Bay,
Japan, CBM-ZC 1668 (T. Komai); F, Pseudopagurodes piliferus (Henderson, 1888), Philippines (T.-Y. Chan); G, Pylopaguropsis granulata
Asakura, 2000, Ryukyu Islands, Japan, CBM-ZC 9994 (T. Komai); H, Pylopaguropsis lemaitrei Asakura & Paulay, 2003, Moorea, French
Polynesia (A. Anker). CBM = Natural History Museum and Institute, Chiba; LACM = Natural History Museum of Los Angeles County;
NSMT = National Museum of Nature and Science, Tokyo.
98
99
Fig. 18. Paguroidea. Representatives of Paguridae Latreille, 1802: A, Pylopaguropsis vicina Komai & Osawa, 2004, Ryukyu Islands,
Japan (T. Komai); B, Pylopaguropsis zebra (Henderson, 1893), Osumi Islands, Japan, CBM-ZC 3137 (T. Komai); C, D, Solitariopagurus
tuerkayi McLaughlin, 1997: C, Hawaiian Islands, LACM CR 2003-046.1 (R. B. Moffitt), D, infested with bopyrid, Ryukyu Islands,
Japan, CBM-ZC 8598 (T. Komai); E, Spiropagurus profundorum Alcock, 1905, Ohsumi Islands, Japan, CBM-ZC 9984 (T. Komai); F,
Trichopagurus trichophthalmus (Forest, 1954), Ryukyu Islands, Japan, CBM-ZC 9949 (T. Komai); G, Turleania saliens Osawa & Fujita,
2008, Kashiwa-jima, Ohtsuki, Kochi Prefecture, CBM-ZC 9991 (T. Komai); H, Turleania senticosa (McLaughlin & Haig, 1996), Ohsumi
Islands, Japan, CBM-ZC 9985 (T. Komai); I, Xylopagurus philippinensis Forest, 1997, Philippines (T.-Y. Chan). CBM = Natural History
Museum and Institute, Chiba; LACM = Natural History Museum of Los Angeles County.
100
101
Fig. 19. Paguroidea. Representatives of Parapaguridae Smith, 1882: A, Oncopagurus monstrosus (Alcock, 1894), Taiwan (T.-Y. Chan);
B, Paragiopagurus acutus (de Saint Laurent, 1972), Uraga Strait, Tokyo Bay, Japan, CBM-ZC 9898 (T. Komai); C, Paragiopagurus
bougainvillei (Lemaitre, 1994), Tuamotu, French Polynesia (J. Poupin); D, Paragiopagurus cf. diogenes (Whitelegge, 1900), French
Frigate Shoals, Hawaiian Islands (G. Paulay); E, Paragiopagurus fasciatus Lemaitre & Poupin, 2003, Austral Islands, French Polynesia
(J. Poupin); F, Paragiopagurus ventilatus Lemaitre, 2004, Taiwan (T.-Y. Chan); G, Paragiopagurus wallisi (Lemaitre, 1994), Austral,
Thiers Bank, French Polynesia (J. Poupin); H, I (in zoanthid carcinoecia), Parapagurus furici Lemaitre, 1999, Taiwan (T.-Y. Chan). CBM
= Natural History Museum and Institute, Chiba
102
103
Fig. 20. Paguroidea. Representatives of Parapaguridae Smith, 1882: A, Parapagurus latimanus Henderson, 1888, off Boso Peninsula,
Japan, CBM-ZC 2038 (T. Komai); B, Parapagurus richeri Lemaitre, 1999, Taiwan (T.-Y. Chan); C, Probeebei mirabilis Boone, 1926,
cephalothorax, pleon and proximal portions of chelipeds and ambulatory legs, off Peru, South Pacific, USNM 267810 (preserved specimen,
R. Lemaitre); D, Strobopagurus gracilipes (A. Milne-Edwards, 1891), Taiwan (T.-Y. Chan); E, Sympagurus brevipes (de Saint Laurent,
1972), Philippines (T.-Y. Chan); F, Tsunogaipagurus chuni (Balss, 1911), Izu Islands, Japan, CBM-ZC (T. Komai); G, Tylaspis anomala
Henderson, 1885, Japan (S. Ohta). CBM = Natural History Museum and Institute, Chiba; USNM = National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
104
105
Fig. 21. Paguroidea. Representatives of Pylochelidae Bate, 1888: A, Bathycheles incisus (Forest, 1897), Philippines (T.-Y. Chan); B,
Bathycheles integer (Forest, 1987), Philippines, MNHN Pg 7907 (T.-Y. Chan); C, Xylocheles macrops (Forest, 1987), Philippines (T.-Y.
Chan); D, Xylocheles miersi (Alcock & Anderson, 1899), Philippines (T.-Y. Chan); E, Parapylocheles scorpio (Alcock, 1894), Philippines
(T.-Y. Chan); F, Trizocheles boasi Forest, 1987, Philippines, in sponge habitat partially removed, NMCR (T.-Y. Chan). MNHN = Musum
national dHistoire naturelle, Paris; NMCR = National Museum of the Philippines, Manila.
106
107

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THE RAFFLES BULLETIN OF ZOOLOGY 2010 Supplement No.

ANNOTATED CHECKLIST OF ANOMURAN DECAPOD CRUSTACEANS OF

Dwi Listyo Rahayu

with Paralomis White, 1847). Most of these additional taxa

The preamble to the Systema Brachyurorum (Ng et

McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

al., 2009; Ahyong et al., 2009, Ahyong, 2010b, Ahyong et

Two anomuran species, Cancer bernhardus Linnaeus,

Although the formal classification within the Lithodoidea

However, the interpretation of phylogenetic relationships

Classification of the Paguroidea is by far the most convoluted.

THE RAFFLES BULLETIN OF ZOOLOGY 2010

American and Russian carcinologists replaced Eupagurus

Brandt (1851) proposed the replacement name Eupagurus

INFRAORDER ANOMURA MACLEAY, 1838

Although Paulson (1875) proposed the genus Dardanus

Superfamily Lomisoidea Bouvier, 1895

McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

Adult males are completely lacking pleopods; females each

DESCRIPTIVE TERMS AND CURRENT STATUS

However, in a series of developmental studies, Crain &

Chelipeds usually, but not always, are markedly unequal, the

In addition to the evidence provided by tergal plate

THE RAFFLES BULLETIN OF ZOOLOGY 2010

pagurid taxon apparently most closely aligned with their two

Current status. The hypothesized evolutionary relationship

McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

from either the right or the left gonopore or from both, or

Current status. The phylogenetic relationship of the

The pleon terminates in paired symmetrical or asymmetrical,

Chelipeds are equal, subequal or grossly unequal. Pereopods 2

Developmental clocks are variable and phase predictability

Adult males may be provided with a pair of gonopores on

THE RAFFLES BULLETIN OF ZOOLOGY 2010

Oedignathus Benedict, 1895

Current status. Most evolutionary hypotheses regarding

A different application of molecular methodology has

McLaughlin et al. (2007) removed the family

The status of Acantholithodes is controversial. In his

Dawson & Yaldwyn (1985) and Dawson (1989)

Makarov (1938) and Dawson (1989) both specifically

Acantholithodes Holmes, 1895

McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

= Lithodes Maia Samouelle, 1819

Cryptolithodes Brandt, 1848

THE RAFFLES BULLETIN OF ZOOLOGY 2010

Neolithodes yaldwyni Ahyong & Dawson, 2006

McLaughlin et al.: Checklist of world Lithodoidea, Lomisoidea and Paguroidea

Phyllolithodes papillosus Brandt, 1848

In his table of gill elements in lithodids, Macpherson

Although correctly cited by Dawson (1989), as noted

Also correctly cited by Dawson (1989), and noted

Among the plates of Sebas (17341765; 18281831)

Despite being first described by Molina (1782) as

Dawson & Yaldwyn (1985) and Dawson (1989)

THE RAFFLES BULLETIN OF ZOOLOGY 2010

Davie (2002) cited monotypy as the type designation

results of the expedition of the Astrolabe and

The original descriptions of Paralithodes californiensis

{14} Birstein & Vinogradov (1972) described their new

{10} Both authorship and publication date for this species

SUPERFAMILY LOMISOIDAE BOUVIER, 1895