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Paleontological Research, vol. 8, no. 3, pp. 143–154, September 30, 2004
6by the Palaeontological Society of Japan
Revision of the genus Glyphithyreus Reuss, 1859
(Crustacea, Decapoda, Brachyura, Xanthoidea) and
recognition of a new genus
HIROAKI KARASAWA1AND CARRIE E. SCHWEITZER2
1Mizunami Fossil Museum, Yamanouchi, Akeyo, Mizunami, Gifu 509-6132, Japan (e-mail: GHA06103@nifty.com)
2Department of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, Canton, Ohio 44720 U.S.A.
(e-mail: cschweit@kent.edu)
Received January 27, 2004; Revised manuscript accepted May 7, 2004
Abstract. The definition of the brachyuran genus Glyphithyreus Reuss, 1859 (FPlagiolophus Bell, 1858
non Pomel, 1857) is herein restricted such that the genus now embraces four species ranging from Paleo-
cene to Oligocene in age. Other species previously referred to the genus have been placed in other genera,
resulting in one new genus, Chirinocarcinus, and four new combinations, Chirinocarcinus wichmanni
(Feldmann et al., 1995), Lobonotus sturgeoni (Feldmann et al., 1995), Stintonius markgrafi (Lo˝renthey,
1907 [1909]), and Titanocarcinus bituberculatus (Collins and Jakobsen, 2003). The referral of Glyph-
ithyreus to the Panopeidae Ortmann, 1893, extends the range of that family into the Paleocene. The geo-
graphic range of Stintonius Collins, 2002, is extended from England to include Egypt as well.
Key words: Brachyura, Decapoda, New taxon, Xanthoidea
Introduction
Glyphithyreus Reuss, 1859, and Plagiolophus Bell,
1858, have been problematic since their introduction
into the fossil brachyuran (crab) nomenclature. The
two genera were named independently for species of
fossil brachyurans that are clearly congeneric (see
illustrations in Bell, 1858 and Reuss, 1859). Alphonse
Milne-Edwards (1865) was the first to recognize this
and synonymized the two, indicating that Plagiolo-
phus was the senior objective synonym based upon
priority. However, Via (1959) subsequently showed
that the name Plagiolophus hadbeenusedbyPomel
(1857) for a genus of eutherian mammal, thus render-
ing Plagiolophus Bell a junior homonym of Plagiolo-
phus Pomel. Glyphithyreus is thus the name with pri-
ority for the crab taxon as suggested by Via (1959).
All brachyuran species referred to Plagiolophus were
therefore referred to Glyphithyreus as a result of Via’s
(1959) suggestion. However, some authors maintained
the usage of Plagiolophus in the literature (Orr and
Kooser, 1971; Berglund and Feldmann, 1989) while
others used Glyphithyreus (Glaessner, 1969; Collins
and Morris, 1978; Squires et al., 1992; Feldmann et al.,
1995, 1998), contributing to the confusion in usage of
the two names.
The situation is made more problematic by the fact
that many of the species referred to Glyphithyreus
(¼Plagiolophus Bell) over the years are not con-
generic with Glyphithyreus, at least when compared to
the type species, G. formosus. It is the purpose of this
paper to provide a restricted definition of Glyphithyr-
eus, to evaluate each of the species that have been
referred to it as well as to Plagiolophus Bell, and to
recommend generic placement for each (Table 1).
This work has resulted in one new genus and four new
combinations. Karasawa and Kato’s (2003) placement
of Glyphithyreus within the Eucratopsinae Stimpson,
1871, of the Panopeidae Ortmann, 1893, is supported
and extends the range of both the family and sub-
family into the Paleocene, as predicted by Casadı
´o
et al. (in review).
Institutional abbreviations
CM—Carnegie Museum of Natural History, Pitts-
burgh, Pennsylvania, USA
GMC—Geological Museum Copenhagen, Copenha-
gen, Denmark
In.—The Natural History Museum, London, United
Kingdom
KSU—Paleontological collections at Kent State Uni-
versity, Kent, Ohio, USA
USNM—National Museum of Natural History,
Smithsonian Institution, Washington, DC, USA.
Systematic paleontology
Infraorder Brachyura Latreille, 1802
Section Heterotremata Guinot, 1977
Superfamily Cancroidea Latreille, 1802
Family Cheiragonidae Ortmann, 1893
Included genera.—Erimacrus Benedict, 1892
(extant); Montezumella Rathbun, 1930; Stintonius
Collins, 2002; Telmessus White, 1846 (extant).
Diagnosis.—Carapace as long as wide or longer
than wide, front bilobed or four-lobed with axial
notch; basal antennal article with a triangular projec-
tion extending into orbital hiatus; orbits with inner-
and outer-orbital spines and median fissure or notch;
lateral margins with 4 to 7 spines; posterior quarter of
dorsal carapace typically rectangular; posterior margin
typically with broad central concavity; sternum with
interrupted sutures between sternites 1 and 2 and
sternites 2 and 3; complex female genital opening not
covered by abdomen.
First pereiopods isochelous; carpus of first pereio-
pod with spinose outer margin and convex lower
margin, usually with a spine; distal margin with two
spines; mani of first pereiopods with small spines on
outer margin, often in rows (diagnosis after S
ˇtevc
ˇic
´,
1988; Schweitzer and Salva, 2000).
Genus Stintonius Collins, 2002
Type species.—Portunites subovata Quayle and
Collins, 1981, by monotypy.
Other species.—Stintonius markgrafi (Lo˝ renthey,
1907 [1909]), as Plagiolophus.
Diagnosis.—Carapace longer than wide; antero-
lateralmarginwithfourspineswhichbecomelarger
posteriorly; carapace regions well defined by narrow
grooves; protogastric region long; axial regions long,
especially urogastric region; hepatic region reduced;
subhepatic region very small; epibranchial region arc-
uate, comprised of two segments, innermost segment
small and triangular; carapace surface appearing to be
densely granulate (after Collins, 2002).
Discussion.—Collins (2002) erected the genus Stin-
tonius to accommodate the Eocene Portunites sub-
ovata Quayle and Collins, 1981, subsequent to the
suggestion by Schweitzer and Feldmann (1999, 2000)
that it did not belong within the genus Portunites and
may be better placed within the Cheiragonidae.
Collins (2002) questionably placed Stintonius within
the Cheiragonidae, based upon the similarity of
Table 1. All species historically referred to Glyphithyreus or Plagiolophus and their current generic placement.
Original Placement Current Placement Relevant Reference
Glyphithyreus formosus Reuss, 1859 (type species) Glyphithyreus Reuss, 1859
G. bituberculatus Collins and Jakobsen, 2003 Titanocarcinus A. Milne Edwards, 1864 this paper
G. sturgeoni Feldmann et al.,1998 Lobonotus A. Milne Edwards, 1864 this paper
?G. wichmanni Feldmann et al., 1995 Chirinocarcinus new genus this paper
Plagiolophus ellipticus Bittner, 1875 Glyphithyreus due to synonymy (Via, 1959)
P. wetherelli Bell, 1858 (¼G. affinis Reuss, 1859) Glyphithyreus due to synonymy (Via, 1959)
P. sulcatus Beurlen, 1939 Glyphithyreus due to synonymy (Via, 1969)
P. markgrafi Lo
´´ renthey, 1907 [1909] Stintonius Collins, 2002 this paper
P. weaveri Rathbun, 1926 Orbitoplax Tucker and Feldmann, 1990 Schweitzer, 2000
P. vancouverensis Woodward, 1896 Archaeopus Rathbun, 1908 Glaessner, 1929
P. bakeri Rathbun, 1935 Lobonotus A. Milne Edwards, 1864 Via, 1969; Fo
¨rster, 1970
P. ezoensis Nagao, 1941 Archaeopus Rathbun, 1908 Collins, Kanie, and Karasawa, 1993
P. vitiensis Rathbun, 1945 unknown; poorly preserved examination of holotype, USNM
498430
VFigure 1. Taxa currently or previously referred to Glyphithyreus Reuss, 1859. 1.Lobonotus sturgeoni (Feldmann et al., 1998) new
combination, dorsal carapace of holotype, CM 36036. 2.Glyphithyreus wetherelli (Bell, 1858), cast of GMC297 currently in KSU collection,
KSU 7035. 3.Glyphithyreus wetherelli (Bell, 1858), KSU 4841, Eocene, Sheppey, UK. 4.Glyphithyreus wetherelli (Bell, 1858), KSU 4854,
Eocene, Sheppey, UK. 5.Glyphithyreus ellipticus (Bittner, 1875), digital image from Bittner, 1875, plate II, figures 8a and b, a, dorsal view;
b, frontal view. 6.Glyphithyreus formosus (Reuss, 1859), digital image from Reuss, 1859, plate II, figure 1. 7.Stintonius markgrafi
(Lo˝ renthey, 1907 [1909]) new combination, digital image from Lo˝renthey, 1907 [1909], plate I, fig. 5a. Scale bars ¼1cm.
Hiroaki Karasawa and Carrie E. Schweitzer144
Revision of Glyphithyreus (Crustacea, Decapoda, Brachyura) 145
Stintonius to Montezumella Rathbun, 1930, which
Schweitzer and Salva (2000) had already placed within
the Cheiragonidae. Unfortunately, the specimen of
Stintonius subovata lacked preserved orbits and fron-
tal margin, which could have facilitated its placement
with no reservations into the Cheiragonidae. With the
referral of Plagiolophus markgrafi to Stintonius,itis
clear that the genus should definitely be placed within
the Cheiragonidae, based upon its bilobed front with
axial notch; at least one orbital fissure; inner- and
outer-orbital spines; and carapace that is longer than
wide. All of these features are diagnostic for the
Cheiragonidae (Schweitzer and Salva, 2000). Plagio-
lophus markgrafi has five anterolateral spines, fewer
than the extant taxa but one more than Montezumella,
the only other extinct genus. The referral of Stintonius
to the Cheiragonidae brings the number of known ex-
tinct genera to two; both are Eocene in age.
Stintonius markgrafi (Lo˝ renthey, 1907 [1909])
new combination
Figure 1.7
Plagiolophus markgrafi Lo˝ renthey, 1907 [1909], p. 137, pl. 1, figs. 5a,
b; Glaessner, 1929, p. 329.
Diagnosis.—Carapace longer than wide, widest at
position of last anterolateral spine, about half the dis-
tance posteriorly on carapace; front bilobed with axial
notch; inner- and outer-orbital spines well developed,
orbits with at least one fissure; regions well developed;
hepatic and branchial regions with oblique, ridgelike
ornamentation; anterolateralmarginwith5spines
excluding outer-orbital spine.
Discussion.—Plagiolophus markgrafi, described
from the Eocene of Egypt (Lo˝ renthey, 1907 [1909]),
cannot be accommodated by Glyphithyreus (¼Pla-
giolophus) for several reasons. The carapace of P.
markgrafi is longer than wide, not wider than long as
in Glyphithyreus,andP. markgrafi lacks the trans-
verse ridges separated by deeply depressed areas on
the branchial regions that typify Glyphithyreus.The
shape and development of carapace regions; the
longer than wide carapace; the bilobed nature of the
front, also with an axial notch; the presence of orbits
with fissures and inner- and outer-orbital spines; the
rectangular posteriormost end of the carapace; and
the possession of between 4 and 7 (5 in P. markgrafi)
anterolateral spines indicate that P. markgrafi is a
member of the Cheiragonidae.
Of the two known Eocene genera, Stintonius can
best accommodate P. markgrafi due to its possession
of tumid, well defined regions, which Montezumella
lacks, and its lack of scabrous ornamentation, which is
typical of Montezumella. However, note that S. mark-
grafi differs from Stintonius subovata in possessing
oblique, ridgelike ornamentation on the hepatic and
branchial regions. Examination of type material of S.
markgrafi may suggest that it should be referred to a
cheiragonid genus distinct from both Montezumella
and Stintonius.
Stintonius is now known from Eocene rocks of
England and Egypt.
Superfamily Xanthoidea MacLeay, 1838
Family Panopeidae Ortmann, 1893
Subfamily Eucratopsinae Stimpson, 1871
Discussion.—Glyphithyreus had previously been
placed within the subfamily Carcinoplacinae H. Milne
Edwards, 1852, of the family Goneplacidae (Balss,
1957; Glaessner, 1969; and many subsequent workers).
Glyphithyreus has well defined dorsal carapace
regions, which is not typical of the Goneplacidae.
Glyphithyreus lacks a straight frontal margin without a
median notch, an entire upper orbital margin with an
indistinct supraorbital angle, and a wide male abdo-
men with all free somites; all of these features are di-
agnostic characters of the subfamily Goneplacinae
MacLeay, 1838 (¼Carcinoplacinae) sensu Karasawa
and Kato, 2003. Thus, Karasawa and Kato (2003)
removed Glyphithyreus from the Goneplacinae of
the Goneplacidae to the panopeid subfamily Eucra-
topsinae Stimpson, 1871, because the carapace has
well defined dorsal regions, the front consists of two
rounded lobes, and the narrow male abdomen has
fused somites 3–5.
Distinction between the panopeid eucratopsine
genera and members of the Pseudorhombilidae is dif-
ficult based upon characters of the carapace, thoracic
sternum, male abdomen, and pereiopods. Major dif-
ferences between extant forms are only in the mor-
phology of the male gonopods (Hendrickx, 1998).
Recently, Schweitzer and Karasawa (2004) indicated
that the fronto-orbital width to carapace width ratio
and the frontal width to carapace width ratio in the
Eucratopsinae are consistently higher than in the
Pseudorhombilidae Alcock, 1900, and redefined both
taxa. The fronto-orbital width to maximum carapace
width ratio in the Eucratopsinae is about 63–81 per-
cent while in the Pseudorhombilidae it is about 53–59
percent. The frontal width in the Eucratopsinae occu-
pies about 30 to 43 percent of the maximum carapace
width but in the Pseudorhombilidae it is about 26–32
percent of the maximum carapace width. In Glyph-
ithyreus the fronto-orbital width to maximum carapace
Hiroaki Karasawa and Carrie E. Schweitzer146
width ratio is about 63–70 percent and the frontal
width to maximum carapace width ratio is about 30–
33 percent. Thus, the placement of Glyphithyreus in
the Eucratopsinae is acceptable.
Because Glyphithyreus is known from the Paleo-
cene of Pakistan (Collins and Morris, 1978), it is thus
the earliest known occurrence of the family and sub-
family. The Panopeinae Ortmann, 1893, was pre-
viously known to have a well established Eocene rec-
ord (Casadı
´oet al., in review). The supposition by
Casadı
´oet al. (in review) that the Panopeinae and the
Eucratopsinae diverged sometime before the Eocene
is therefore supported.
Genus Glyphithyreus Reuss, 1859
(¼Plagiolophus Bell, 1858 non Pomel, 1857)
Figure 1.2–1.6
Glyphithyreus Reuss, 1859, p. 4, pl. 2, figs. 1 –3.
Plagiolophus Bell, 1858, p. 19, pl. II, figs. 7 –13 (nom.preoccup.by
Plagiolophus Pomel, 1857).
Type species.—Plagiolophus wetherelli Bell, 1858 ¼
Glyphithyreus affinis Reuss, 1859, by monotypy under
ICZN, 1999, Article 67.8.
Species included.—Glyphithyreus ellipticus (Bittner,
1875) as Plagiolophus,G. formosus Reuss, 1859; G.
sulcatus (Beurlen, 1939), as Plagiolophus;G.wether-
elli (Bell, 1858), as Plagiolophus (¼G. affinis Reuss,
1859).
Diagnosis.—Carapace subquadrilateral, wider than
long, L/W about 0.75–0.80, widest in anterior one-
third of carapace; fronto-orbital margin about 63–70
percent maximum carapace width; front comprised of
two slightly rounded lobes, about one-third maximum
carapace width, with median notch; supraorbital an-
gle weakly defined; upper orbital margin concave,
rimmed, weakly notched medially or with two fissures;
anterolateral margin strongly convex with four spines
including outer-orbital spine, third spine largest; post-
erolateral margin sinuous, converging posteriorly; re-
gions granular dorsally, well defined by deep, smooth
grooves; epigastric regions well defined; mesogastric
region separated from metagastric region by V-shaped
groove; each epibranchial region inflated with broad
ridge forming convex-forward arc from metagastric
region to last anterolateral spine; broad transverse
ridge across cardiac and metabranchial regions,
forming nearly continuous ridge across carapace; epi-
branchial and cardiac/metabranchial ridges separated
by deep cavity; posterior end of carapace depressed to
level of cavity separating two branchial ridges.
Thoracic sternum relatively wide; male abdomen
narrow with somites 3–5 fused. Chelipeds massive,
elongate.
Discussion.—Bell (1858) described a new genus and
species, Plagiolophus wetherelli, from the Eocene
London Clay of England. Reuss (1859) described a
new genus, Glyphithyreus,andtwonewspecies,G.
formosus, the type species, and G. affinis. Alphonse
Milne Edwards (1865) synonymised Glyphithyreus af-
finis Reuss, 1859, with P. wetherelli, and indicated that
Glyphithyreus was the junior subjective synonym of
Plagiolophus. Via (1959) showed that the generic
name Plagiolophus was preoccupied by Plagiolophus
Pomel, 1857, for a genus of Mammalia, and first used
the junior subjective synonym of the valid name,
Glyphithyreus Reuss, 1859, instead of Plagiolophus
Bell, 1858. We concur with Via’s (1959) decision.
Glaessner (1929) recognized four species of Plagio-
lophus (¼Glyphithyreus), P. ellipticus Bittner, 1875;
P. markgrafi Lo˝ renthey, 1907 [1909]; P. weaveri
Rathbun, 1926; and P. wetherelli, the latter of which
he considered to be synonymous with both G. affinis
Reuss, 1859, and G. formosus Reuss, 1859. Glaessner
(1929) also moved P. vancouverensis Woodward,
1896, to Archaeopus Rathbun, 1908, of the family
Retroplumidae Gill, 1894. Four additional species,
Plagiolophus bakeri Rathbun, 1935, from the Eocene
of U.S.A.; P. ezoensis Nagao, 1941, from the Creta-
ceous of Japan; P. sulcatus Beurlen, 1939, from the
Oligocene of Hungary; and P. vitiensis Rathbun, 1945,
from the Miocene of Fiji, were subsequently de-
scribed. In his review of the Eocene decapods of the
world, Via (1969) placed P. ellipticus,P. ezoensis,P.
markgrafi,P. sulcatus,P. weaveri,andP. vitiensis
within Glyphithyreus. Both Via (1969) and Fo
¨rster
(1970) moved Plagiolophus bakeri to Lobonotus
A. Milne Edwards, 1864, of the family Xanthidae
MacLeay, 1838; we concur. Lobonotus is now placed
within the Pilumnidae Samouelle, 1819 (Schweitzer
et al., 2004).
More recently, Glyphithyreus ezoensis was assigned
to the retroplumid genus Archaeopus Rathbun, 1908
(Collins, Kanie, and Karasawa, 1993). Glyphithyreus
weaveri was moved to Orbitoplax Tucker and Feld-
mann, 1990, of the family Goneplacidae MacLeay,
1838 (Schweitzer, 2000). Collins and Morris (1978)
described G. wetherelli from the Paleocene of Pakistan
and treated G. formosus as a valid species, with which
we concur. The specimen they assigned to G. wether-
elli conforms to the general diagnosis of Glyphithyreus
in possessing broad orbits and marked transverse
ridges on the dorsal carapace; thus, it is the earliest
known occurrence of the genus.
The only known Miocene species of Glyphithyreus,
Revision of Glyphithyreus (Crustacea, Decapoda, Brachyura) 147
Glyphithyreus vitiensis, is here excluded from the ge-
nus because it lacks branchial ridges (USNM 498430),
which are diagnostic characters of the genus. Glyphi-
thyreus vitiensis possesses carapace characters most
like those of Xanthodius kambaraensis Rathbun, 1945
(Xanthidae), from the Miocene of Fiji. In recent
works, Glyphithyreus sturgeoni Feldmann et al., 1998,
from the Eocene of U.S.A. and ?G. wichmanni
Feldmann et al., 1995, from the Danian of Argentina,
have been described. However, both species lack very
distinctive branchial ridges, which are typical of Gly-
phithyreus, and they are assigned to other genera de-
scribed below.
Consequently, we recognize only four species of
Glyphithyreus (Table 1). Of these, the placement of G.
sulcatus is somewhat tentative and is based upon our
translation of Beurlen’s (1939) original description in
German and the very poorly reproduced illustration in
our copy of the work. The description of G. sulcatus
clearly indicates two transverse ridges on the bran-
chial regions, separated by a very deep cavity, which
is certainly characteristic of Glyphithyreus. Thus we
place the species in the genus until type material can
be examined. Glyphithyreus wetherelli is recorded
from the Paleocene of Pakistan (Collins and Morris,
1978) and the Eocene of England, Belgium, Denmark,
France, Spain, and Senegal (Bell, 1858; Remy in
Remy and Tessier, 1954; Via, 1969; Plaziat and Secre-
tan, 1971). Glyphithyreus wetherelli recorded from
Denmark (Ravn, 1903; Via, 1969; Plaziat and Secre-
tan, 1971) was assigned to the new species, Gly-
phithyreus bituberculatus (as Titanocarcinus bitu-
berculatus in this paper), by Collins and Jakobsen
(2003). Glyphithyreus ellipticus is only known from the
Eocene of Italy (Bittner, 1875), and Glyphithyreus
formosus is from the Cretaceous? of Germany (Reuss,
1859). Thus, Glyphithyreus is generally known from
Paleocene and Eocene deposits in the western Tethys
realm.
Family Goneplacidae MacLeay, 1838
?Subfamily Euryplacinae Stimpson, 1871
Diagnosis.—Carapace usually with poorly defined
carapace regions; front straight with shallow median
notch; supraorbital angle distinct; orbit sometimes
deep, large, with two, one or no orbital fissures; ante-
rolateral margin with two to five spines (after Kara-
sawa and Kato, 2003, p. 138–139). For remainder of
diagnosis, see Karasawa and Kato (2003).
Discussion.—The general shape of the carapace and
carapace regions; the spined nature of the antero-
lateral margins; and the clear distinction between the
anterolateral and posterolateral margins all indicate
that ?Glyphithyreus wichmanni is a member of the
Xanthoidea. Paleocene xanthoids are relatively un-
common, although they have received directed atten-
tion in recent years (Schweitzer, 2003a, b, in press).
A major problem for family, subfamily, and generic
placement of ?Glyphithyreus wichmanni is that the
ventral aspects of the carapace are unknown. Features
of the sternum, abdomen, and articulation of the per-
eiopods are very important in xanthoid classification
of both extant and fossil members (Guinot, 1978;
Davie, 2002; Karasawa and Kato, 2003; Schweitzer,
2003a, b, in press). Thus, proxy characters of the
dorsal carapace (Schweitzer and Feldmann, 2000;
Schweitzer, 2003a) must be used. The short antero-
lateral margin relative to the posterolateral margin
clearly excludes ?G. wichmanni from genera within
the Palaeoxanthopsidae Schweitzer, 2003a, and the
Zanthopsidae Via, 1959, both xanthoid families with
Paleocene representatives. The Hexapodidae Miers,
1886, have a fossil record extending into the Creta-
ceous (Schweitzer, in press), but the distinctively rect-
angular carapace and carapace dimensions of the
embraced genera (Schweitzer and Feldmann, 2001)
cannot accommodate ?G. wichmanni.
?Glyphithyreus wichmanni cannot be accom-
modated by the Eriphiidae MacLeay, 1838, because
members of that family have either very broad fronto-
orbital widths, occupying most of the maximum cara-
pace width, or two orbital fissures, neither of which
?G. wichmanni possesses. The Trapeziidae Miers,
1886, possess very broadly spaced orbits and smooth
dorsal carapaces which cannot accommodate ?G.
wichmanni. Similarly, the smooth dorsal carapace and
very long, convex anterolateral margins of the Carpi-
liidae Ortmann, 1893, cannot embrace ?G. wichmanni.
Members of the Pseudorhombilidae Alcock, 1900,
have two orbital fissures and an intraorbital spine,
none of which ?G. wichmanni possesses. Taxa within
the Platyxanthidae Guinot, 1977, possess two orbital
fissures and a more narrow front and a wider carapace
relative to the length than does ?G. wichmanni.The
Pseudoziidae Alcock, 1898, are much wider than long
and have poorly defined regions, while ?G. wichmanni
is only slightly wider than long (L/W ¼0.83), and has
well defined regions. Members of the Xanthidae
MacLeay, 1838 sensu stricto have long anterolateral
margins and concave posterolateral margins, neither
of which ?G. wichmanni exhibits. ?Glyphithyreus
wichmanni has very much shorter anterolateral mar-
gins than is typical of members of the Panopeidae
Ortmann, 1893, and its regions are in general better
developed than in members of the Panopeidae. In
Hiroaki Karasawa and Carrie E. Schweitzer148
addition, at least some panopeids have two orbital
fissures, which ?G. wichmanni lacks.
The two families to which ?G. wichmanni is most
likely referable are the Pilumnidae Samouelle, 1819,
and the Goneplacidae MacLeay, 1838. Members of
the Goneplacidae usually have very short antero-
lateral margins, a prominent character displayed by
?Glyphithyreus wichmanni. In addition, the Gonepla-
cidae is one of the few xanthoid families with a fossil
record extending into the Cretaceous (Schweitzer
et al., 2002), and which can therefore accommodate
?G. wichmanni without a range extension. The gone-
placid subfamily Euryplacinae Stimpson, 1871, as de-
fined by Karasawa and Kato, 2003, is quite variable in
terms of dorsal carapace morphology. Some genera
possess two orbital fissures, for example, Viaplax
Karasawa and Kato, 2003; some possess one orbital
fissure, for example, Stoaplax Vega et al., 2001; while
others possess none, as in Orbitoplax Tucker and
Feldmann, 1990. The dorsal carapace regions of
species of Orbitoplax are very well defined, while
those of Stoaplax are not, and extant genera such as
Nancyplax Lemaitre et al.,2001,arenearlysmooth.
In spite of this variability, all euryplacines possess
short anterolateral margins and relatively broad
fronto-orbital widths, which can accommodate ?G.
wichmanni. In addition, euryplacines possess from two
to five anterolateral spines, a well defined supraorbital
angle, and relatively large orbits, all of which can ac-
commodate ?G. wichmanni. The front of ?G. wich-
manni appears to be notched medially, another fea-
ture typical of euryplacines. Thus, the Euryplacinae
can best accommodate ?G. wichmanni,forwhicha
new genus has been erected below, and we tentatively
place it within the Euryplacinae until aspects of the
sternum, abdomen, and pereiopods can be examined.
Other subfamilies of the Goneplacidae cannot
accommodate ?Glyphithyreus wichmanni. Members of
the Goneplacinae MacLeay, 1838, usually have broad
orbits and narrow fronts, not exhibited by ?G. wich-
manni. The Carinocarcinoidinae Karasawa and Kato,
2003, possess transverse keels on a relatively smooth
dorsal carapace; neither are seen in ?G. wichmanni.
The Chasmocarcininae Sere
`ne, 1964 and Troglo-
placinae, Guinot, 1986, are typified by rectangular,
relatively featureless dorsal carapaces, which cannot
accommodate ?G. wichmanni. Fossil taxa within the
Mathildellinae Karasawa and Kato, 2003, have flat-
tened carapaces, two orbital fissures, and moderate
fronto-orbital widths of about half the carapace width,
none of which ?G. wichmanni possesses; however, the
arrangement of carapace regions in both Tehuacana
Stenzel, 1944, and Branchioplax Rathbun, 1916, is
similar to that of ?G. wichmanni.
Members of most subfamilies of the Pilumnidae
have very unusual shapes (Halimedinae Alcock, 1898;
Calmaniinae S
ˇtevc
ˇic
´, 1991; Eumedoninae Dana, 1853;
Rhizopinae Stimpson, 1858; see Davie, 2002) which
exclude ?G. wichmanni. The general arrangement
of carapace regions and proportions of the carapace
in ?G. wichmanni are similar to that seen in the
Pilumninae Samouelle, 1819. The Galeninae Alcock,
1898, a monogeneric subfamily, have poorly defined
carapace regions and a very broad carapace as com-
pared to the length, not seen in ?G. wichmanni.
Members of both the Pilumninae and the Galeninae
have longer anterolateral margins than does ?G.
wichmanni. In addition, the orbits of most pilumnines
are directed anterolaterally, while those of ?G. wich-
manni are directed forward. Thus, it is most likely that
?G. wichmanni is not a member of the Pilumnidae.
Chirinocarcinus new genus
Glyphithyreus Reuss, 1859 (part). Feldmann, Casadı
´o, Chirino-
Galvez, and Aguirre-Urreta, 1995, p. 14, figs. 11, 12.
Type species.—?Glyphithyreus wichmanni Feld-
mann, Casadı
´o, Chirino-Galvez, and Aguirre-Urreta,
1995, by monotypy.
Diagnosis.—Carapace slightly wider than long, L/W
about 0.83, widest at position of last anterolateral
spine about one-third the distance posteriorly on car-
apace; front appearing to have axial notch, about 35
percent maximum carapace width, projecting beyond
orbits; orbits circular, entire, directed forward, outer-
orbital angle projecting slightly; fronto-orbital width
about 63 percent maximum carapace width; antero-
lateral margin very short, with three spines excluding
outer-orbital spine, last spine largest; posterolateral
margin long, sinuous, convex; posterolateral reen-
trants well developed; posterior margin short, con-
cave; carapace regions developed as broadly swollen
areas separated by relatively deep grooves; urogastric
and cardiac regions ornamented with tubercles.
Etymology.—The genus name honors Luis Chirino-
Ga
´lvez, Chile, formerly a graduate student at Kent
State University, Kent, Ohio, who has contributed
much to our understanding of fossil crabs from South
America, especially those of Chile.
Discussion.—Glyphithyreus wichmanni, question-
ably referred to Glyphithyreus by Feldmann et al.
(1995), cannot be retained within that genus. The type
species of Glyphithyreus,G. wetherelli (Bell, 1858),
exhibits very distinctive, transverse ridges on the
dorsal carapace. These ridges are composed of the
Revision of Glyphithyreus (Crustacea, Decapoda, Brachyura) 149
arcuate epibranchial regions, which are linearly and
transversely swollen, and transverse ridges on the
mesobranchial region, which are nearly continuous
with the transversely swollen cardiac region. ?Glyphi-
thyreus wichmanni lacks these transverse ridges.
Glyphithyreus wetherelli has two orbital fissures, ob-
served on In. 59575 and In. 48229, while G. wichmanni
lacks orbital fissures. Glyphithyreus wetherelli has five
anterolateral spines, while G. wichmanni has only
three. Glyphithyreus wichmanni is more equant
than G. wetherelli, which is markedly wider than
long. Thus, ?G. wichmanni must be removed from
Glyphithyreus.
We herein place ?Glyphithyreus wichmanni in a
new genus, tentatively within the Euryplacinae, re-
sulting in the new combination Chirinocarcinus wich-
manni. Of the three fossil genera previously referred
to that subfamily (Karasawa and Kato, 2003), ?G.
wichmanni lacks orbital fissures, which are present in
the extinct genera Stoaplax and Viaplax, and absent in
Orbitoplax.Chirinocarcinus has moderately large
orbits, as in Viaplax, instead of very large orbits, as
in Orbitoplax and Stoaplax. No other fossil genus
has this particular combination of orbital fissures and
orbit size. In addition, the dorsal carapace of Chirino-
carcinus is more equant than members of Orbitoplax,
and the orbits of Orbitoplax and Stoaplax are rectan-
gular and very deep, not seen in Chirinocarcinus.If
Chirinocarcinus is confirmed as a member of the sub-
family, it would be the oldest member known, Paleo-
cene in age, whereas the other fossil taxa are Eocene.
Family Pilumnidae Samouelle, 1819
Genus Lobonotus A. Milne Edwards, 1864
Type species.—Lobonotus sculptus A. Milne
Edwards, 1864, by monotypy.
Other species.—Lobonotus bakeri (Rathbun, 1935),
as Plagiolophus;L. brazoensis Stenzel, 1935 (known
only from claws); L. mexicanus Rathbun, 1930; L.
natchitochensis Stenzel, 1935; L. sandersi (Blow and
Manning, 1997), as Eohalimede Blow and Manning,
1997; L. sturgeoni (Feldmann et al., 1998) as
Glyphithyreus.
Diagnosis.—see Schweitzer et al. (2004).
Discussion.—Schweitzer et al. (2002) examined the
genus Lobonotus and removed some species from it,
and they also referred Eohalimede sandersi Blow and
Manning, 1997, to the genus. Schweitzer et al. (2004)
restricted the genus to those species listed above,
other than L. sturgeoni.TheyplacedLobonotus with-
in the Pilumnidae based upon features of the sternum,
male abdomen, and dorsal carapace. Lobonotus
lobulata Feldmann et al., 1995, and Lobonotus ori-
entalis Collins and Morris, 1978, were each removed
to new genera, Lobulata and Pakicarcinus respectively
(Schweitzer et al., 2004). These actions restricted
the genus to only Central and North American
forms; the referral herein of Glyphithyreus sturgeoni,
described from the Eocene of North Carolina, USA,
to the genus maintains this geographic pattern. All of
the known occurrences, except for the Miocene L.
sculptus, are from Eocene rocks (Schweitzer et al.,
2002).
Lobonotus sturgeoni (Feldmann, Bice, Schweitzer
Hopkins, Salva, and Pickford, 1998) new combination
Fig. 1.1
Glyphithyreus sturgeoni Feldmann, Bice, Schweitzer Hopkins, Salva,
and Pickford, 1998, p. 13, figs. 17, 18.
Diagnosis.—Carapace length nearly 90 percent
width, small for genus; front axially notched, with six
small protuberances including inner orbital protuber-
ance; orbits broadly rimmed, with two well developed
fissures; anterolateral margin with 4 spines excluding
outer orbital spine; carapace regions well marked by
deep grooves; surface of carapace coarsely granular,
especially near lateral margins; cardiac region weakly
three-lobed.
Material examined.—CM 36036, holotype.
Discussion.—Glyphithyreus sturgeoni is here re-
ferred to the genus Lobonotus due to its possession of
numerous dorsal carapace characters similar to those
of the type species. These features include an equant
carapace that is steeply vaulted anteriorly; a notched
front; orbits with two well developed fissures; deep
grooves and well developed regions ornamented with
granules; anterolateral margin with 4 spines; and a
weakly three-lobed cardiac region. All of these fea-
tures are diagnostic for the genus. Lobonotus stur-
geoni may be differentiated from other species of the
genus by its small size (17.3 mm wide versus nearly 40
mm in Lobonotus mexicanus in Schweitzer et al.,
2002). The cardiac region of L. sturgeoni is more
weakly trilobed than in other members of the genus,
and the ornamentation appears to be less dense than
in other species. However, as the specimen is a mold
of the interior, much surface detail has probably been
lost. The front of Lobonotus sturgeoni clearly exhibits
six weak protuberances, which have not been de-
scribed in other species; the front typically has been
described as nearly straight. All of these differences
are clearly within the range of generic variation; the
speciesisreferredtoLobonotus with no reservations.
Hiroaki Karasawa and Carrie E. Schweitzer150
Genus Titanocarcinus A. Milne Edwards, 1864
Type species.—Titanocarcinus serratifrons A. Milne
Edwards, 1864, by subsequent designation of Glaess-
ner (1929).
Discussion.—Glyphithyreus bituberculatus Collins
and Jakobsen, 2003, cannot be accommodated by
Glyphithyreus because it lacks the branchial ridges
separated by a deep cavity and the very depressed
posteriormost end of the carapace diagnostic for
Glyphithyreus.Itmostcloselyresemblesspeciesof
Titanocarcinus, based upon its notched front; rimmed
orbits with two fissures; well developed carapace re-
gions separated by broad, smooth grooves; epibran-
chial region well subdivided into two areolae; equant
carapace; and anterolateral margins with three spines.
Collins and Morris (1978) suggested that Titano-
carcinus might be synonymous with Lobonotus;
Schweitzer et al. (2002) concurred. Schweitzer et al.
(2004) reevaluated Lobonotus but did not make a
decision on the Lobonotus and Titanocarcinus issue.
Such a decision is beyond the scope of this paper but
is being considered by the authors. For now, we
place the Paleocene Glyphithyreus bituberculatus in
Titanocarcinus based upon its possession of only three
anterolateral spines, as in T. serratifrons, the type
species, and T. raulinensis A. Milne Edwards, 1864;
and a cardiac region that is very weakly trilobed.
Species referred to Lobonotus have four or five ante-
rolateral spines and distinctly trilobed cardiac regions.
However, species of Titanocarcinus usually have a
longitudinal groove in the protogastric region, which
G. bituberculatus,aswellasspeciesofLobonotus,
lack. Historically, European species have been re-
ferred to Titanocarcinus and American forms have
been referred to Lobonotus (Collins and Morris, 1978;
Schweitzer et al., 2002); we follow that precedent for
the time being. As currently defined, Titanocarcinus is
known from Cretaceous to Miocene rocks (Glaessner,
1969).
Titanocarcinus bituberculatus (Collins and Jakobsen,
2003) new combination
Figure 2.1, 2.2
Glyphithyreus bituberculatus Collins and Jakobsen, 2003, p. 74, fig.
6, pl. 5, figs. 1– 5.
Diagnosis.—Carapace not much wider than long,
widest just posterior to last anterolateral spine;
front notched, about one-third maximum carapace
width; orbits rimmed, with two fissures; fronto-orbital
width about two-thirds maximum carapace width;
anterolateral margins with three spines excluding
outer-orbital spines; carapace regions well developed,
separated by broad, smooth grooves.
Sternum ovate; sternites 1–2 fused, no evidence of
suture; strong, entire groove marking suture between
sternites 2 and 3; very deep groove between sternites
3 and 4, medially interrupted; sternite 4 with deep
grooves marking fusion of episternites of sternite 3
with sternite 4; deep groove extending anteriorly from
sterno-abdominal cavity onto sternites 4 and 3; male
abdomen possibly covering entire space between
coxae of pereiopods 5, but unable to determine for
certain; all male abdominal somites free.
Figure 2. Titanocarcinus bituberculatus (Collins and Jakobsen, 2003) new combination. 1. Reproduced figure of plate 5, figure 2a,
from the Bulletin of the Mizunami Fossil Museum. 2. Reproduced figure of plate 5, figure 5b, from the Bulletin of the Mizunami Fossil
Museum. Scale bars ¼1cm.
Revision of Glyphithyreus (Crustacea, Decapoda, Brachyura) 151
Discussion.—Schweitzer et al. (2004) described
the male abdomen of species of Lobonotus as not en-
tirely covering the space between the coxae of per-
eiopods 5, based upon examination of the type of L.
mexicanus Rathbun, 1930 (USNM 371096). A small
portion of sternite 8 may have been exposed, but that
area of the specimen is covered with sediment that
cannot be prepared away. In the specimens of T.
bituberculatus, it is difficult to determine what the re-
lationship between the male abdomen and the coxae
of the fifth pereiopods was, as those coxae are not
preserved; the male abdomen is broadened in that
region, suggesting that it may have filled the entire
space between those coxae. However, it is not possible
to know for certain and it is similarly not possible
to know if any portion of sternite 8 was visible in T.
bituberculatus. If the male abdomen of T. bitubercula-
tus did in fact fill the entire space between the coxae of
the fifth pereiopods, the apparent relationship be-
tween Lobonotus and Titanocarcinus becomes prob-
lematic because these features of the male abdomen
and sternum are considered to be extremely important
at the genus, subfamily, and family level. If Titano-
carcinus and Lobonotus were to be synonymized,
these possible differences would have to be taken into
account.
The deep sternal grooves of T. bituberculatus cer-
tainly suggest affinity with members of a new family
(Schweitzer, in press) or Zanthopsidae Via, 1959.
Schweitzer (in press) suggested that these deep
grooves may be a primitive feature among some Xan-
thoidea, as they appear in many Eocene taxa as well
as the Platyxanthidae Guinot, 1977, which was con-
sidered by Guinot (1978) to possess many plesiomor-
phic characters. Similar groove patterns are found in
some members of various subfamilies of the Pilumni-
dae; perhaps they are plesiomorphic characters re-
tained by some members of the family. Investigation
of these groove patterns is ongoing.
Acknowledgements
A. Kollar, Carnegie Museum of Natural History,
Pittsburgh, PA, USA, loaned the specimen of Glyph-
ithyreus sturgeoni; we thank him. W. Blow, National
Museum of Natural History, Smithsonian Institution,
Washington, DC, kindly facilitated access to the
collections at that institution. R.M. Feldmann, Kent
State University, assisted with assembling the illustra-
tions and read an earlier draft of the manuscript. C.
Trocchio, Kent State University Stark Campus, as-
sisted with German translation. Y. Okumura, Mizu-
nami Fossil Museum, granted permission to reproduce
illustrations from the Bulletin of the Mizunami Fossil
Museum. Special thanks are due to H. Kato, Natural
History Museum and Institute, Chiba, for his review
of the manuscript.
References
Alcock, A., 1898: The family Xanthidae: the Brachyura Cyclo-
metopa, Pt. I, Materials for a Carcinological Fauna of
India, No. 3. Journal of the Asiatic Society of Bengal, vol.
67, no. II:1, p. 67–233.
Alcock, A., 1900: Materials for a Carcinological Fauna of In-
dia, No. 6. The Brachyura Catametopa, Grapsoidea. Jour-
nal of the Asiatic Society of Bengal, vol. 69, no. II(3), p.
279–456.
Balss, H., 1957: Decapoda. In,Klassen und Ordnungen des
Tierreichs, Band 5, Arthropoda, Abteilung 1, Buch 7, Lie-
ferung 12, p. 1505–1672.
Bell, T., 1858: A monograph of the fossil malacostracous Crus-
tacea of Great Britain. Part I. Crustacea of the London
Clay. 44 p. Palaeontographical Society, London.
Benedict, J. E., 1892: Corystoid crabs of the genera Telmessus
and Erimacrus.Proceedings of the United States National
Museum, vol. 15, p. 223–230, pls. 25–27.
Berglund, R. E. and Feldmann, R. M., 1989: A new crab, Ro-
gueus orri n. gen. and sp. (Decapod: Brachyura) from the
Lookingglass Formation (Ulatisian Stage: Lower Middle
Eocene) of southwestern Oregon. Journal of Paleontology,
vol. 63, p. 69–73.
Beurlen, K., 1939: Neue Decapoden-Krebse aus dem un-
garischen Tertia
¨r. Pala
¨ontologische Zeitschrift,vol.21,
nos. 1–4, p. 135–161, pl. 7.
Bittner, A., 1875: Die Brachyuren des Vicentinischen Tertia
¨r-
gebirges. Denkschriften der kaiserlichen Akademie der
Wissenschaften in Wien, vol. 34, p. 63–103.
Blow, W. C. and Manning, R. B., 1997: A new genus, Martinetta,
and two new species of xanthoid crabs from the Middle
Eocene Santee Limestone of South Carolina. Tulane
Studies in Geology and Paleontology, vol. 30, p. 171–180.
Casadı
´o, S., Feldmann, R. M., Parras, A. and Schweitzer, C. E.
(in review): Miocene fossil Decapoda (Crustacea: Bra-
chyura) from Patagonia, Argentina and their paleoeco-
logic setting. Annals of Carnegie Museum.
Collins, J. S. H., 2002: A taxonomic review of British decapod
Crustacea. Bulletin of the Mizunami Fossil Museum, no.
29, p. 81– 92.
Collins, J. S. H. and Jakobsen, S. L., 2003: New crabs (Crus-
tacea, Decapoda) from the Eocene (Ypresian/Lutetian)
Lillebælt Clay Formation of Jutland, Denmark. Bulletin
of the Mizunami Fossil Museum,no.30,p.63–96,8pls.
Collins, J. S. H., Kanie, Y. and Karasawa, H., 1993: Late Cre-
taceous crabs from Japan. Transactions and Proceedings of
the Palaeontological Society of Japan, new series, no. 172,
p. 292– 310.
Collins, J. S. H. and Morris., S. F., 1978: New lower Tertiary
crabs from Pakistan. Palaeontology, vol. 21, p. 957–981.
Dana, J. D., 1853: A review of the classification of Crustacea,
with reference to certain principles of classification. In,
United States Exploring Expedition during the Years 1838,
1839, 1840, 1841, 1842 under the Command of Charles
Wilkes, U.S.N., vol. 13, Crustacea, Part II, p. 1395–1450. C.
Sherman, Philadelphia.
Hiroaki Karasawa and Carrie E. Schweitzer152
Davie, P. J. F., 2002: Crustacea: Malacostraca: Eucarida (Part
2): Decapoda—Anomura, Brachyura. In,Wells,A.and
Houston, W. W. L. eds., Zoological Catalogue of Australia,
vol. 19.3B, p. 1–641. CSIRO Publishing, Melbourne,
Australia.
Feldmann, R. M., Bice, K. L., Schweitzer Hopkins, C., Salva,
E. W. and Pic kford, K., 1998: Decapod crustace ans from
the Eocene Castle Hayne Limestone, North Carolina: Pa-
leoceanographic implications. The Paleontological Society
Memoir vol. 48 (Supplement to Journal of Paleontology,
vol. 72),28p.
Feldmann, R. M., Casadı
´o, S., Chirino-Galvez, L. and Aguirre-
Urreta, M., 1995: Fossil decapod crustaceans from the
Jaguel and Roca Formations (Maastrichtian-Danian) of
the Neuque
´n basin, Argentina. The Paleontological Society
Memoir vol. 43 (Supplement to Journal of Paleontology,
vol. 69), 22 p.
Fo
¨rster, R., 1970: Zwei neue brachyure Krebse aus dem Pal-
a
¨oza
¨n der Haunsberges no
¨rdlich von Salzburg. Mitteilun-
gen der Bayerischen Staatssammlung fu
¨rPala
¨ontologie und
historische Geologie, vol. 10, p. 241–252.
Gill, T., 1894: A new Bassalian type of crabs. American Natu-
ralist, vol. 28, p. 1043–1045.
Glaessner, M. F., 1929: Crustacea Decapoda. In, Pompeckji,
F. J., ed., Fossilium Catalogus 1. Animalia, 464 p. Berlin.
Glaessner, M. F., 1969: Decapoda. In, Moore, R. C. ed., Treatise
on Invertebrate Paleontology, Part R, Arthropoda 4,p.
R399– R533, R626 –R628. Geological Society of America
and University of Kansas Press, Lawrence.
Guinot, D., 1977: Propositions pour une nouvelle classification
des Crustace
´sDe
´capodes Brachyoures. Comptes Rendus
de l’Acade
´mie des Sciences,Paris,Se
´rie D,vol.285,p.
1049– 1052.
Guinot, D., 1978: Principes d’une classification e
´volutive des
Crustace
´sDe
´capodes Brachyoures. Bulletin Biologique de
le France et de le Belgique,vol.112,p.211–292.
Guinot, D., 1986: Description d’un crabe cavernicole aveugle
de Nouvelle-Bretagne (Papouasie Nouvelle-Guine
´e),
Trogloplax joliverti gen. nov. sp. nov., et e
´stablissement
d’une sous-famille nouvelle, Trogloplacinae subfam. nov.
Comptes Rendus de l’Acade
´mie des Sciences, Paris, se
´rie 3,
vol. 303, p. 307 –312.
Hendrickx, M. E., 1998: A new genus and species of
‘‘goneplacid-like’’ brachyuran crab (Crustacea: Decapoda)
from the Gulf of California, Mexico, and a proposal for the
use of the family Pseudorhombilidae Alcock, 1900. Pro-
ceedings of the Biological Society of Washington,vol.111,
p. 634–644.
International Commission on Zoological Nomenclature, 1999:
International Code of Zoological Nomenclature, Fourth
Edition. 306 p. The International Trust for Zoological
Nomenclature, London.
Karasawa, H. and Kato, H., 2003: The family Goneplacidae
MacLeay, 1838 (Crustacea: Decapoda: Brachyura): sys-
tematics, phylogeny, and fossil records. Paleontological
Research,vol.7,p.129–151.
Latreille, P. A., 1802– 1803: Histoire Naturelle, Ge
´ne
´rale et Par-
ticulie
`re, des Crustace
´s et des Insectes. Volume 3, 467 p.
F. Dufart, Paris.
Lemaitre, R., Garcı
´a-Go
´mez, J., Von Sternberg, R. and
Campos, N. H., 2001: A new genus and a new species of
crab of family Goneplacidae MacLeay, 1838 (Crustacea:
Decapoda: Brachyura) from the tropical western Pacific.
Proceedings of the Biological Society of Washington,vol.
114, p. 951–963.
Lo˝ renthey, E., 1907 [1909]: Beitra
¨ge zur Kenntnis der eoza
¨nen
Decapodenfauna Aegyptens. Mathematische und natur-
wissenschaftliche Berichte aus Ungarn, vol. 25, p. 106–152.
MacLeay, W. S., 1838: On the brachyurous Decapod Crustacea
brought from the Cape by Dr. Smith. In,Smith,A.ed., Il-
lustrations of the Annulosa of South Africa; consisting
chiefly of the Objects of Natural History Collected during
an Expedition into the Interior of South Africa, in the Years
1834, 1835, and 1836; Fitted out by ‘‘The Cape of Good
Hope Association for Exploring Central Africa’’,p.53–71.
Smith, Elder, and Company, London.
Miers, E. J., 1886: Report on the Brachyura collected by H. M.
S. Challenger during the years 1873–1876. Report on the
Scientific Results of the Voyage of H. M. S. Challenger
during the years 1873– 1876, Zoology, vol. 17, 362 p., pls.
1–29.
Milne Edwards, A., 1862–1865: Monographie des crustace
´s
de la famille cance
´riens. Annales des Sciences Naturelles,
Zoologie, Se
´rie 4, vol. 18 (1862), p. 31–85, pls. 1 –10; vol. 20
(1863),p.273–324,pls.5–12;Se
´rie 5, vol. 1 (1864), p. 31–
88, pls. 3–9; vol. 3 (1865), p. 297–351, pls. 5–10.
Milne Edwards, H., 1852: De la famille des ocypodides (Ocy-
podidae). Second Me
´moire. Observations sur les affinite
´s
zoologiques et la classification naturelle des crustace
´s.
Annales des Sciences Naturelles, Zoologie, Se
´rie 3,vol.18,
p. 109–166.
Nagao, T., 1941: On some fossil Crustacea from Japan. Journal
of the Faculty of Science, Hokkaido
ˆImperial University,
Series 4, Geology and Mineralogy, vol. 6, p. 85–97, pl. 26.
Orr, W . N. and Kooser, M. A., 1971: Oregon Eocene decap od
Crustacea. The Ore Bin, vol. 33, p. 119–129.
Ortmann, A., 1893: Die Dekapoden-Krebse des Strassburger
Museums VII. Die Abtheilungen Brachyura (Brachyura
genuina Boas). 2. Unterabtheilung: Cancroidea, 2. Section:
Cancridea, 1. Gruppe: Cyclometopa. Zoologische Jahr-
bu
¨cher, Abtheilung fu
¨r Systematik, Geographie und Biol-
ogie der Thiere,vol.7,p.411–495.
Pomel, A., 1857: Note critique dur les caracte
`res et les limites
du genre Palaeotherium. Archives des Sciences Physique et
Naturelle, Gene
`ve,vol.5,p.200–207.
Plaziat, J.-C. and Secretan, S., 1971: La faune de Crustace
´sde-
capodes des calcaires a
`alveolines Ypresiens des Corbie
`res
Septentrionales (Aude). Geobios,vol.4,p.117–142.
Quayle, W. J. and Coll ins, J. S. H., 1 981: New Eoce ne crabs from
the Hampshire Basin. Palaeontology,vol.24,p.733–758.
Rathbun, M. J., 1908: Descr iptions of the fossil crabs from
California. Proceedings of the United States National Mu-
seum, vol. 35, p. 341– 349.
Rathbun, M. J., 1916: Description of a new genus and species of
fossil crab from Port Townsend, Washington. American
Journal of Science, vol. 41, p. 344 –346.
Rathbun, M. J., 1926: The fossil stalk-eyed Crustacea of the
Pacific slope of North America. Bulletin of the United
States National Museum, no. 138, I-VIIþ155 p., 39 pls.
Rathbun, M. J., 1 930: Fo ssil dec apod cr ustaceans f rom Mexi co.
Proceedings of the United States National Museum,vol.78,
p. 1–10, pls. 1–6.
Rathbun, M. J., 1935: Fossil Crustacea of the Atlantic and Gulf
Coastal Plain. Geological Society of America, Special
Paper, no. 2, p. 1– 160.
Rathbun, M. J., 1 945: Dec apoda C rustacea. In, Ladd, H. S. and
Revision of Glyphithyreus (Crustacea, Decapoda, Brachyura) 153
Hoffmeister, J. E., eds., Geology of Lau, Fiji. Bernice P.
Bishop Museum Bulletin, no. 181, p. 373–391, pls. 54–62.
Ravn, J. P . J., 1 903: Om foss ile T erebellide-rør fra Danmark.
Meddelelser fra Dansk Geologisk Forening,vol.4,p.383–
390.
Remy, J.-M. and Tessier, F., 1954: De
´capodes nouveaux de la
partie ouest de Se
´ne
´gal. Bulletin de la Societe
´Ge
´ologique
de France, Se
´rie 6,vol.4,p.185–191.
Reuss, A., 1859: Zur Kenntnis fossiler Krabben. Denkschriften
der kaiserlichen Akademie der Wissenschaften Wien,vol.
17, p. 1–90, pls. 1–24.
Samouelle, G., 1819: The Entomologist’s Useful Compendium,
or an Introduction to the Knowledge of British Insects.486
p. London.
Schweitzer, C. E., 2000: Tertiary Xanthoidea (Crustacea: De-
capoda: Brachyura) from the west coast of North America.
Journal of Crustacean Biology, vol. 20, p. 715–742.
Schweitzer, C. E., 2003a: Utility of proxy characters for classi-
fication of fossils: an example from the fossil Xanthoidea
(Crustacea: Decapoda: Brachyura). Journal of Paleontol-
ogy, vol. 77, p. 1107–1128.
Schweitzer, C. E., 2003b: Progress on the fossil Xanthoidea
MacLeay, 1838 (Decapoda, Brachyura). Contributions to
Zoology, vol. 72, p. 181–186.
Schweitzer, C. E., in press: The genus Xanthilites Bell and a
new xanthoid family (Crustacea: Decapoda: Brachyura:
Xanthoidea): new hypotheses on the origin of the Xan-
thoidea MacLeay, 1838. Journal of Paleontology.
Schweitzer, C. E. and Feldmann, R. M., 1999: Fossil decapod
crustaceans of the late Oligocene to early Miocene Pysht
Formation and late Eocene Quimper Sandstone, Olympic
Peninsula, Washington. Annals of Carnegie Museum,vol.
68, p. 215– 273.
Schweitzer, C. E. and Feldmann, R. M., 2000: New fossil por-
tunids from Washington, USA, and Argentina, and a re-
evaluation of generic and family relationships within the
Portunoidea Rafinesque, 1815 (Decapoda: Brachyura).
Journal of Paleontology, vol. 74, p. 636–653.
Schweitzer, C. E. and Feldmann, R. M., 2001: Differentiation
of the fossil Hexapodidae Miers, 1886 (Decapoda: Bra-
chyura) from similar forms. Journal of Paleontology,vol.
75, p. 330– 345.
Schweitzer,C.E.,Feldmann,R.M.andGingerich,P.,2004:
New decapods (Crustacea) from the Eocene of Pakistan
and a revision of Lobonotus A. Milne Edwards, 1864.
Contributions to the Museum of Paleontology, University
of Michigan, vol. 31, p. 89–118.
Schweitzer, C. E., Feldmann, R. M., Gonza
´les-Barba, G. and
Vega, F. J., 2002: New crabs from the Eocene and Oligo-
cene of Baja California Sur, Mexico and an assessment of
the evolutionary and paleobiogeographic implications of
Mexican fossil decapods. The Paleontological Society
Memoir, vol. 59 (Supplement to Journal of Paleontology,
vol. 76), 43 p.
Schweitzer, C. E. and Karasawa, H., 2004: Revision of Amy-
drocarcinus and Palaeograpsus (Decapoda: Brachyura:
Xanthoidea) with definition of three new genera. Paleon-
tological Research,vol.8,p.71–86.
Schweitzer, C. E. and Salv a, E. W., 2000: First recognition of
the Cheiragonidae (Decapoda) in the fossil record and
comparison of the family with the Atelecyclidae. Journal
of Crustacean Biology,vol.20,p.285–298.
Sere
`ne, R., 1964: Redescription du genre Megaesthesius
Rathbun et de
´finition des Chasmocarcininae, nouvelle
sous-famille des Goneplacidae (Decapoda Brachyura).
Crustaceana,vol.7,p.175–187.
Squires, R. L., Goedert, J. L. and Kaler, K. L., 1992: Paleontol-
ogy and stratigraphy of Eocene rocks at Pulali Point, Jef-
ferson County, eastern Olympic Peninsula, Washington.
Washington Division of Geology and Earth Resources
Report of Investigations, vol. 31, p. 1–27.
Stenzel, H. B., 1935: Middle Eocene an d Oli gocene decapod
crustaceans from Texas, Louisiana, and Mississippi. The
American Midland Naturalist, vol. 16, p. 379 –400.
Stenzel, H. B., 1944: A new Paleocene catometope crab from
Texas, Tehuacana tehuacana.Journal of Paleontology, vol.
18, p. 550–552.
S
ˇtevc
ˇic
´, Z., 1988: The status of the family Cheiragonidae
Ortmann, 1893. International Journal of Marine Biology
and Oceanography (EBALIA), XIV, new series, vol. 1987/
1988, p. 1–14.
S
ˇtevc
ˇic
´, Z., 1991: Note on some rare and aberrant Australian
crabs. The Beagle, Records of the Northern Territory Mu-
seum of Arts and Sciences, no. 8, p. 121–134.
Stimpson, W., 1858: Prodromus descriptionis animalium ever-
tebratorum, quae in Expeditione ad Oceanum Pacificum
Septentrionalem, a Republica Federata missa, Cadwala-
daro Ringgold et Johanne Rodgers Ducibus, observavit et
descripsit W. Stimpson. Pars V. Crustacea Ocypodoidea.
Proceedings of the Academy of Natural Sciences of Phila-
delphia, vol. 10, p. 93–110.
Stimpson, W., 1871: Preliminary report on the Crustacea
dredged in the Gulf Stream in the Straits of Florida, by
L.F. de Pourtales, Assist. U.S. Coast Survey. Bulletin of
the Museum of Comparative Zoo
¨logy at Harvard College,
vol. 2, p. 109–160.
Tucker, A. B. and Feldmann, R. M., 1990: Fossil decapod crus-
taceans from the lower Tertiary of the Prince William
Sound region, Gulf of Alaska. Journal of Paleontology,
vol. 64, p. 409 –427.
Vega, F. J., Cosma, T., Coutin
˜o, M. A., Feldmann, R. M.,
Nyborg, T. G., Schweitzer, C. E. and Waugh, D. A., 2001:
New middle Eocene decapods (Crustacea) from Chiapas,
Me
´xico. Journal of Paleontology, vol. 75, p. 929–946.
Via, L., 1959: De
´capodos fo
´siles del Eoceno espan
˜ol. Boletin
del Instituto Geolo
´gico y Minero Espan˜ola,vol.70,p.331–
402.
Via, L., 1969: Crusta
´ceos Deca
´podos del Eoceno espan
˜ol. Piri-
neos, nos. 91–94, 479 p.
White, A., 1846: Meeting of the Entomological Society, April
7, 1845. Annals of Natural History,vol.17,497p.
Woodward, H., 1896: On some podophthalmous Crustacea
from the Cretaceous formation of Vancouver and Queen
Charlotte Islands. Quarterly Journal of the Geological
Society of London, vol. 52, p. 221 –228.
Hiroaki Karasawa and Carrie E. Schweitzer154
