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New Eocene Brachyura (Crustacea: Decapoda) from Egypt
Rodney M. Feldmann, Carrie E. Schweitzer, Olga Bennett, Ovidiu D. Franţescu, Nick Resar, and
Ashley Trudeau
With 15 figures and 5 tables
Feldmann, R.m., SchweitzeR, c.e., Bennett, O., FRanţeScu, O.d., ReSaR, n. & tRudeau, a. (2011):
New Eocene Brachyura (Crustacea: Decapoda) from Egypt. – N. Jb. Geol. Paläont. Abh., DOI:
10.1127/0077-7749/2011/0202; Stuttgart.
Abstract: Examination of E. lőRenthey’s decapod material in the Staatliches Museum für
Naturkunde, Stuttgart, as well as other museums, has resulted in several new genera and species
from the Eocene of Egypt. The Eocene Egyptian decapod fauna was robust, unsurprising given the
diverse Eocene faunas known from Italy, Spain, and North America. The revision of Palaeocarpilius
is included. New genera include Bryocarpilius, Hemsut, Laticarpilius, and Tumidomaia; new species
include Bryocarpilus aspidorsalis, Laticarpilius aegypticus, Lophoranina cinquecrista, Ocalina
delicata, Planobranchia egyptensis, and Tumidomaia mokattamensis. There are numerous new com-
binations for Hemsut, Liopsalis, Paraocalina, Harpactocarcinus, L aticarpilius, and Bryocarpilius.
Key words: Eocene, Decapoda, Brachyura, Raninoida, Eubrachyura, Egypt, Tethys.
1. Introduction
The decapod crustacean fauna of Egypt, in the
vicinity of Cairo, attracted a great deal of interest in
the latter half of the 19th and early 20th centuries.
Numerous authors studied and described new species
from Eocene localities collectively referred to as
Gebel Mokattam. During that period, 19 taxa were
recognized. lőRenthey (1909) summarized all these
works and provided excellent illustrations of decapod
fauna of the region. Subsequent to that work, only
two new species have been added to the list (Van
StRaelen 1929[1930]; andeRSOn & Feldmann 1995).
However, subsequent to the work of lőRenthey,
considerable work has been done on the systematics
and classication of decapod crustaceans that bears on
the identity of this fauna resulting in the recognition
of new taxonomic combinations, denition of new
genera, and identication of new species (Table 1).
Thus, it is the purpose of this work to reinvestigate
the crab fauna of the Mokattam Eocene in light of the
advances that have occurred since that early work.
This re-examination has also permitted more careful
comparison with congeneric taxa throughout the
Tethyan region which has resulted in reassignment of
some species lying outside Egyptian borders. Because
the original and subsequent locality data was sketchy,
little has been done to update the stratigraphic and
geographic occurrence data.
Where possible, material documenting the fauna
has been studied, primarily in museums in Europe;
however, a single specimen of interest was also
discovered in the collections of the United States
National Museum of Natural History. Species for
which the actual specimens are missing or could
not be located have been studied using original and
©2011 E. Schweizerba rt’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbar t.de
DOI: 10.1127/0077-7749/2011/0202 0077-7749/2011/0202 $ 7.75
N. Jb. Geol. Paläont. Abh. Fast Track Article
published online September 2011 October 2011
2 R.M. Feldmann et al.
subsequent descriptions and illustrations. By far, the
most extensive collection of type material is held in
the Staatliches Museum für Naturkunde, Stuttgart,
Germany. Study of this and other collections permitted
preparation of modern descriptions and photographic
illustrations of the brachyuran fauna based upon actual
specimens and casts of them.
2. Locality and horizon
The specimens described herein were collected from
Mokattam, Egypt, unless otherwise indicated. The
rocks from Gebel Mokattam, near Cairo, range from
Lutetian (middle Eocene) to late Eocene (Priabonian)
age (GinGeRich 1992). Sections of the Gebel Mokattam
area illustrated by GinGeRich (1992) show decapods
at two levels. Lobocarcinus ReuSS, 1857, a crab,
and Callianassa leach, 1814, a ghost shrimp, were
shown as occurring in the upper Mokattam Formation
(GinGeRich 1992: gs. 6-7, 10) of Lutetian age, and
Callianassa occurred in the Maadi Formation of
Priabonian age (GinGeRich 1992: 16). Thus, the
decapods discussed here can be constrained to the
Lutetian to Priabonian stages, because we have no
more specic location data.
Institutional abbreviations: BMNH, The Natural
History Museum, London, UK; MGSB, Museo Geológico
del Seminario de Barcelona, Spain; KSU D, Decapod
Comparative Collection, Department of Geology, Kent
State University, Kent, Ohio, USA; SMF, Senckenberg
Forschungsinstitut und NaturMuseum, Frankfurt am Main,
Germany; SMNS, Staatliches Museum für Naturkunde,
Stuttgart, Germany; USNM, United States National
Museum of Natural History, Smithsonian Institution,
Washington, D.C., USA.
Table 1. Decapod species reported from Egypt. List from Glaessner (1929) and subsequent literature. All are from the Eocene of
the Mokattam area unless otherwise noted. * indicates Miocene occurrences.
Original Designation Current Designation
Callianassa nilotica FRaaS, 1867 Callianassa nilotica
Callinanssa longa nOetlinG, 1885 Callinanssa longa
Callianassa fraasi nOetlinG, 1885 Callianassa fraasi
Callianassa mokattamensis nOetlinG, 1885 Callianassa mokattamensis
Callianassa transversoplicata nOetlinG, 1885 Callianassa transversoplicata
Pagurus (Clibanarius?) dubius nOetlinG, 1885 Pagurus dubius
Pagurus mezi lőRenthey, 1907b Petrochirus mezi
Lophoranina spp. (pereiopod fragments) Lophoranina spp. (pereiopod fragments)
Lophoranina cinquecristata n. sp. Lophoranina cinquecristata
Calappa sp. (subfossil) Calappa sp.
Xanthopsis cuvillieri Van StRaelen, 1929 [1930] Eriosachila cuvillieri
Hepatiscus schweinfurthi nOetlinG, 1885 Hemsut schweinfurthi
Typilobus trispinosus lőRenthey, 1907b Typilobus trispinosus
Neptunus granulatus A. milne-edwaRdS, 1860* Portunus alphonsei KaRaSawa et al., 2008
Neptunus monspeliensis A. milne-edwaRdS, 1860* Portunus monspeliensis
Neptunus sp. Neptunus sp.
Micromaja laevis lőRenthey, 1907b Planobranchia laevis
Micromaja tuberculata BittneR, 1875 Tumidomaia tuberculata
Lambropsis wanneri lőRenthey, 1907b Lambropsis wanneri
Lobocarcinus ? aegypticus lőRenthey, 1907b Lobocarcinus aegypticus
Lobocarcinus lumacopius andeRSOn & Feldmann, 1995 (Eocene of
Fayum, Egypt) Lobocarcinus lumacopius
Lobocarcinus paulinowuerttembergensis VOn meyeR, 1847 Lobocarcinus paulinowuerttembergensis
Palaeocarpilius macrocheilus deSmaReSt, 1822 Specimens may be referable to Bryocarpilius apsidorsalis n. g., n. sp.
Palaeocarpilius simplex StOliczKa, 1871 Laticarpilius aegypticus n. g., n. sp.
Plagiolophus markgra lőRenthey, 1907b Karasawaia markgra
Cancer fraasi lőRenthey, 1907b Montezumella fraasi
Goniocypoda transsilvanica BittneR, 1893 Goniocypoda transsilvanica
Bryocarpilius apsidorsalis n. g., n. sp. Bryocarpilius apsidorsalis n. g., n. sp.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 3
3. Systematic paleontology
Infraorder Brachyura linnaeuS, 1758
Section Raninoida ahyOnG et al., 2007
Superfamily Raninoidea de haan, 1839
Family Raninidae de haan, 1839
Genus Lophoranina FaBiani, 1910
(= Palaeonotopus BROcchi, 1877, p. 4, nomen
oblitum; see GlaeSSneR 1969: R498).
Type species: Ranina marestiana KöniG, 1825, by original
designation.
Included species: Lophoranina aculeata (A. milne-
edwaRdS, 1881), as Ranina; L. aldrovandii (Ranzani,
1820), as Ranina; L. bakerti A. milne-edwaRdS, 1872,
nomen nudum, based upon the holotype of L. reussi; L.
barroisi (BROcchi, 1877), as Palaeonotopus; L. bishopi
SquiReS & demetRiOn, 1992; L. bittneri (lőRenthey, 1902),
as Ranina; L. cristaspina VeGa, cOSma et al., 2001; L.
georgiana (RathBun, 1935), as Ranina; L. kemmelingi Van
StRaelen, 1924 [imprint 1925], as Ranina (Lophoranina);
L. laevifrons (BittneR, 1875), as Ranina; L. levantina lewy,
1977; L . marestiana (KöniG, 1825), as Ranina; L. maxima
BeSchin et al., 2004; L. persica witheRS, 1932; L. porifera
(wOOdwaRd, 1866), as Ranina; L. precocia Feldmann et al.,
1996; L. quinquespinosa (RathBun, 1945), as Ranina; L.
raynorae BlOw & manninG, 1996; L. reussi (wOOdwaRd,
1866), as Ranina; L. rossi BlOw & manninG, 1996; L.
soembaensis Van StRaelen, 1938; L. straeleni Vía, 1959;
L. tchihatchef (A. milne-edwaRdS, 1866), as Ranina; L.
toyosimae yaBe & SuGiyama, 1935.
Lophoranina cinquecrista n. sp.
Fig. 1
1885 Ranina cf. Marestiana KöniG, 1825. – nOetlinG, p.
488.
Material examined: Holotype, SMNS 67887.
Etymology: The trivial name is derived from the Old
French word creste, meaning crest, and the French word
cinque, meaning ve, in reference to the presence of ve
distinctive convex forward ridges on the anterior portion of
the carapace.
Diagnosis: Anteriormost of carapace with terraced ridges
more broadly spaced (ca. 1.5 mm) and with coarser and more
broadly spaced granules than remainder of carapace, with
ridges spaced at about 1.2 mm; anteriormost ridge extends
across mesial 20% of carapace width as forward-directed
chevron; next ve ridges convex forward, continuous
across carapace, with chevron-shaped axis diminishing in
height so that ridges four and ve exhibit very faint axial
inections; remainder of terraced ridges irregularly arrayed;
some continuous across carapace, others discontinuous, still
others bifurcate.
Description: Carapace elongate ovoid; length (39.7
mm) greater than width (28.5 mm); moderately vaulted
longitudinally and transversely. Surface with closely spaced
transverse terraced ridges with granular anterior margins
and smooth, sloping posterior margins.
Front incompletely preserved, fronto-orbital width 21.8
mm; outermost corner with broad projection bearing one
long triangular and one blunt spine; outermost projection
separated from prominent triangular spine by deep
V-shaped notch. Lateral margins entire, smoothly convex,
joining straight, narrow posterior margin in smooth arc.
Anteriormost part of carapace with terraced ridges
more broadly spaced (ca. 1.5 mm) and with coarser and
more broadly spaced granules than over remainder of
carapace with ridges spaced at about 1.2 mm. Anteriormost
ridge extending across mesial 20% of carapace width as
forward-directed chevron. Next ve ridges convex forward,
continuous across carapace, with chevron-shaped axis
diminishing in height so that ridges four an ve exhibit very
faint axial inections.
Remainder of terraced ridges irregularly arrayed; some
Fig. 1. Lophoranina cinquecristata n. sp., holotype, SMNS
67887, dorsal carapace. Scale bars = 1 cm.
4 R.M. Feldmann et al.
continuous across carapace, others discontinuous, and still
others bifurcate.
Carapace regions not dened except metagastric region
bounded by short, arcuate branchiocardiac grooves.
Venter, pereion, and appendages not preserved.
Discussion: Species of Lophoranina are distinguished
from one another on the basis of carapace outline, nature
of the spinulation on the front, breadth of the front, and
development of terraced ridges – the single feature that
most clearly distinguishes the genus from other raninids.
The terraced ridges vary in size, spacing, orientation,
and continuity. Lophoranina cinquecrista n. sp. exhibits
a combination of characters that distinguish it from all
other species within the genus. The new species is widest
at midlength and has a smoothly convex anterolateral and
posterolateral margin with no inection between the two.
The front is incomplete; however, the left anterolateral
corner is preserved, which conrms that the frontal region
is very short. The most distinctive feature is that the anterior
part of the carapace is traversed by one short and ve
complete terraced lines that are relatively widely spaced and
sharply convex forward axially. The three ridges between
the branchiocardiac grooves are complete and concave
forward. The remaining ridges are more closely spaced and
may be continuous, discontinuous, or bifurcating as they
cross the carapace in gently concave-forward arcs. This
combination of features is seen on no other species within
the genus.
The two species that appear to be most closely related
to Lophoranina quinquecrista also exhibit convex forward
ridges in the anterior part of the carapace. Lophoranina
maxima, from the Lutetian (middle Eocene) of Vicenza,
Italy, exhibits one medial and two complete terraced ridges
whose axial regions are convex forward; an additional
medial ridge may also have this form. The ridges between
the branchiocardiac grooves may be discontinuous or
straight and continuous. Specimens referred to this species
had previously been referred to as L . reussi by BuSulini
et al. (1983) or L. cf. [L.] reussi by BeSchin et al. (1988).
Subsequent examination by BeSchin et al. (2004) revealed
that the convex forward pattern was not seen on specimens
of L. reussi. They concluded that this pattern was sufciently
distinctive to warrant erection of Lophoranina maxima. We
agree. The other species that has convex forward ridges is
L. soembaensis, from the lower Eocene of the Lesser Sunda
Islands, north of Australia. It has ve complete terraced
ridges with convex forward axes. The convexity decreases
posteriorly. The ridges between the branchiocardiac grooves
are continuous and concave forward. To our knowledge,
no other species of Lophoranina exhibits terraces in the
anterior region with forward directed axes. In an attempt
to conrm the morphology of L. soembaensis, one of us
(RMF) contacted S. dOnOVan, Nationaal Natuurhistorisch
Museum, Leiden, The Netherlands to request a photograph
or loan of the holotype of the species. Van StRaelen (1938)
indicated that the type, and sole specimen, was deposited in
the Rijks Geologisch-Mineralogisch Museum at Leyden. To
date, the specimen has not been located.
Considering the characters diagnostic of Lophoranina
quinquecrista, the other species of Lophoranina possess
characteristics that readily distinguish them from the
new species. Lophoranina aculeata has relatively few,
concave forward ridges and large, simple spines on the
front. Lophoranina aldrovandii bears short terraced lines
forming a scalloped pattern throughout. The terraced lines
on Lophoranina barroisi are more or less continuous and
describe sinuous patterns. Very large anterolateral spines
on a carapace which is widest in the front ¼ and which
has uniformly spaced ridges characterizes L. bishopi.
Lophoranina bittneri has a fronto-orbital width of 59%
of maximum width and continuous terraced ridges in the
anterior 2/3 of the carapace that are concave forward. The
front on L. cristaspina is broad, granular, and lacks ridges.
The other species with a very broad, generally unadorned
front is L. precocia. This species, which also has very large
anterolateral spines, is the oldest member of the genus.
It was described (Feldmann et al. 1996) from the Late
Cretaceous of Chiapas, Mexico. Lophoranina cristaspina
was also described from Chiapas, in Eocene rocks.
Lophoranina georgiana has a hexagonal outline,
is widest in the anterior 1/3 of the carapace, and has
uniformly spaced terraced ridges. The sole specimen of
L. kemmelingi consists of the right posterolateral corner of
the carapace and, although it is sufcient to recognize the
genus, no reasonable comparison can be made with other
species. Lophoranina laevifrons has a very short front,
and most ridges are concave forward and uniformly spaced
throughout. The terraced ridges on L. levantina are sinuous
with concave forward axes, and the carapace is widest in
the anterior one quarter. The type species of the genus,
Lophoranina marestiana, has a broad, granular front and
uniform continuous and discontinuous ridges over the entire
carapace surface. Lophoranina persica consists only of a
fragment of the left posterolateral corner; it bears generally
straight, strong ridges with coarse spine bases. Although
the terraced ridges on L. porifera are numerous, generally
continuous, and evenly spaced, the species is poorly known
and comparison is difcult. That is even more true for L.
quinquespinosa which is known only from a fragment of
pereiopod 1. The generic assignment is probably sound, but
species comparison is not possible.
Lophoranina raynorae is readily recognized as having
a distinct front which marks the widest part of the carapace
and which bears an ornate axial crest and has numerous
short, discontinuous terraced lines. Lophoranina reussi
is characterized by a narrow front, a fronto-orbital width
which is 46% the maximum width, and equally spaced,
sinuous ridges. An ornate fronto-orbital eld on a carapace
that is widest in the anterior third characterizes L. rossi.
Lophoranina straeleni, which has a prominent inection
at the anterolateral corner, bears a strongly spinose
anterolateral margin. Although the original description of
L. tchihatchef is sparse and the illustration has not been
seen, BeSchin et al. (2004) noted that the anterior half of the
carapace bears regular ridges. Lophoranina toyosimae is
widest in the anterior ¼ and bears deeply concave forward,
generally complete, broadly spaced terraced ridges.
Lophoranina arose in the Late Cretaceous
(Maastrichtian) and is also known in rocks ranging from
Eocene to Miocene. The genus reached its acme in the
Eocene when it was represented by 18 of 24 species. None
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 5
is known from Paleocene rocks. Species are distributed
through Tethyan regions in North America, Europe, and
Asia. Thus, the Egyptian occurrence of Lophoranina
quinquecrista in Eocene rocks does not expand the range
of the genus.
Section Eubrachyura de Saint lauRent, 1980
Superfamily Leucosioidea SamOuelle, 1819
Family Leucosiidae SamOuelle, 1819
Genus Typilobus StOliczKa, 1871
Type species: Typilobus granulosus StOliczKa, 1871.
Included species: Typilobus alponensis BeSchin, de
anGeli & zORzin, 2009; T. ? belli quayle & cOllinS,
1981; T. ? birshteinyi FöRSteR & mundlOS, 1982; T. boscoi
Vía, 1959; T. ? corrodatus (nOetlinG, 1885), as Ilia; T.
granulosus StOliczKa, 1871; T. johnfei (hu & taO, 1996),
as Chumaoia; T. kishimotoi KaRaSawa, 1998; T. marginatus
mORRiS & cOllinS, 1991; T. ? modregoi Vía, 1959; T.
moralejai mülleR, 1993; T. ? obscurus quayle & cOllinS,
1981; T. ? prevostianus (deSmaReSt, 1822), as Leucosia; T.
sadeki witheRS, 1925; T. ? semseyanus lőRenthey, 1898;
T. trispinosus lőRenthey, 1907b; T. unispinatus maRtinS-
netO, 2001.
Diagnosis: Carapace circular to transversely oval, about
as wide as long or wider than long; front bilobed; lateral
margin smooth or granular, sometimes bearing tubercles;
posterior margin with blunt spine on each side; dorsal
surface strongly convex, granular, with more or less dened
regions; male abdomen with seven free somites.
Discussion: mülleR (1993) noted that the genus as then
known embraced a wide range of morphotypes. In fact,
the variation in development of lateral and posterior spines
and the development of grooves dening frontal, gastric,
hepatic, cardiac, and branchial regions is great enough
to warrant reconsideration and redenition of the genus.
Unfortunately, the present work is based almost entirely on
published descriptions and illustrations, most of which are
drawings, so that it is not appropriate to undertake such a
revision at this time.
Typilobus trispinosus lőRenthey, 1907b
Fig. 2
1904 Typilobus trispinosus nov. sp. – lőRenthey, p. 165
(in Hungarian).
1907a Typilobus trispinosus nov. sp. – lőRenthey, p. 34
1907b Typilobus trispinosus nov. sp. – lőRenthey, p. 205,
pl. 2, g. 1 (in Hungarian).
1909 Typilobus trispinosus nov. sp. – lőRenthey, p. 117,
pl. 2, g. 1.
1929 T. trispinosus löRenthey, 1909. – GlaeSSneR, p. 389.
2010 T. ? trispinosus lőRenthey, 1909. – SchweitzeR et
al., p. 91.
Material examined: Holotype and sole specimen, SMNS
67890.
Diagnosis: Carapace typical of genus and bearing ne
and moderately coarse granules overall, becoming coarser
posteriorly; a prominent spine near posterior end of inated
cardiac region; axial as well as lateral spines on posterior
margin.
Description: Carapace circular, 1.48 cm wide with
greatest width at midlength, 1.5 cm long excluding broken
rostral area and posterior spines; length/width ratio 0.99.
Anterolateral and posterolateral margins entire, smooth.
Posterolateral corners marked by broad, blunt spines about
0.2 cm long. Posterior axial spine broken, but appears to be
narrower and shorter than posterolateral spines.
Carapace surface strongly vaulted transversely
and longitudinally. Regions dened by narrow, deep grooves
dening frontal, hepatic, gastric, cardiac, and branchial
regions. Frontal region about 0.4 cm wide, frontal width/
width ratio 0.27; anterior margin broken; orbits not known;
posterior margin weakly concave forward at intersection
with gastric region. Hepatic regions appear triangular
in dorsal view and bearing two longitudinal rows of ne
granules. Gastric regions undifferentiated, forming an
elongate hexagonal outline about 0.4 cm wide at front, about
0.84 cm wide at junction with posterior corner of hepatic
region, and about 0.4 cm wide at junction with cardiac
region. Cardiac region more inated than other regions,
Fig. 2. Typilobus trispinosus lőRenthey, 1907b. SMNS
67890, dorsal carapace. Scale bars = 1 cm.
6 R.M. Feldmann et al.
broadens posteriorly and bears a single prominent node
approximately at level of greatest width. Intestinal region
not apparent. Branchial regions large, undifferentiated.
Surface of carapace with granules of two sizes; ne
granules uniformly distributed over carapace and coarser
granules increase in density from widest part of gastric
region to the posterior margin; coarse granules most evident
on axial regions.
Venter, pleon, and appendages not preserved.
Discussion: This species is known only from a single
specimen from Mokattam. It is distinguished from other
species within the genus by having a prominent axial spine on
the posterior margin which, coupled with the posterolateral
spines, accounts for the trivial name. Additionally, the
surface of the carapace bears coarse and ne granules,
unlike the ornamentation on other species. Typilobus
alponensis, from the Ypresian of Italy, is smoother, has a
more angular outline, and bears posterolateral swellings
rather than spines, thus distinguishing it from the Egyptian
species. Typilobus ? belli has a bilobed frontal region and
a nearly equidimensional hexagonal gastric area, whereas
T. trispinosus has a simple frontal region and an elongate,
hexagonal gastric region. FöRSteR & mundlOS (1982) named
T. birshteinyi based upon two illustrations in BiRShtein
(1956: 92, gs. 18-19). BiRShtein referred to the specimens
simply as Leucosiidae. The regions on T. birshteinyi are
more tumid than those on T. trispinosus; the outline is less
circular, and there seem to be no posterolateral spines.
Rather than having a circular outline, such as is seen in
T. trispinosus, T. boscoi has lateral margins bearing ve
short spines, excluding the posterolateral spines. Tipilobus
corrodatus exhibits three small nodes on the cardiac region
and two nodes on the hepatic region, whereas there is one
relatively large node on the cardiac region and no discrete
nodes on the hepatic region of T. trispinosus. The type
species of the genus, T. granulosus, is transversely ovate,
bears small spines on the lateral margins, has rounded
margins of the gastric region and a somewhat posteriorly
projected cardiac region, whereas the Egyptian species is
circular, has smooth margins, angular margins of the gastric
region, and a cardiac region that is not projected posteriorly.
Typilobus johnfei is wider than long, bears a deeply pitted
surface, and lacks nearly all the grooves reminiscent of
other species within the genus. Thus, its assignment to
Typilobus is questionable. The species was originally placed
in a monotypic genus, Chumaoia hu & taO, 1996, in the
Goneplacidae. KaRaSawa subsequently (1997) recognized
it as a synonym of Typilobus (p. 67, footnote in Japanese).
KaRaSawa & KatO (2003: 146) conrmed that it was a
leucosiid, not a goneplacid.
Typilobus kishimotoi has a somewhat more angular
outline than T. trispinosus, and the former also has nodes on
the gastric and branchial regions and has a more spinose
lateral margin and small posterolateral spines that the
latter does not exhibit. The lateral margin of T. marginatus
is sharp and strongly granular, the grooves bounding the
gastric and hepatic regions are apparently absent, and
the surface is coarsely granulose, features not seen on T.
trispinosus. The outline of T. modregoi is transversely
ovate, bears only two spines on the posterior margin,
and has a nearly equidimensional gastric area. Typilobus
trispinosus is circular in outline, has three posterior spines,
and exhibits an elongate gastric region. The groove pattern
of T. moralejai is very weak and the surface of the carapace
is uniformly nely granular, whereas the groove pattern
is distinct and the granules are of two different sizes on T.
trispinosus. The posterior margin of the former species is
not preserved, so that interpretation of the posterior spines is
not possible. Although only examined from the illustration
in quayle & cOllinS (1981: pl. 104, g. 13), the specimen
of T. ? obscurus appears to have been illustrated with the
posterior oriented toward the top of the plate. It differs
signicantly from the Egyptian species in that the grooves
dening the branchial, cardiac, and posterior part of the
gastric region are very ne and the anteriormost grooves are
not in evidence. The surface is uniformly, nely pustulose.
Typilobus ? prevostianus bears an axial and two oblique
ridges on the gastric region and is very nely punctate.
There seems to be a posterior axial node developed as an
extension of the cardiac region but no posterolateral spines.
These are features not seen on the Egyptian species. As
a note of clarication, deSmaReSt (1822: 114) mistakenly
referred to the illustration of T. ? prevostianus as pl. 9,
g. 13, whereas it should be pl. 9, g. 14. The outline of
T. sadeki is angular; there is a clear inection between the
anterolateral and posterolateral margins. The ornamentation
on the surface is uniformly ne and there appears to be
no posterolateral spines. The rounded outline, variable
granulation, and possession of three posterior spines of
T. trispinosus distinguish it from T. sadeki. Typilobus ?
semseyanus exhibits a uniformly granular carapace surface
and has small spines on the lateral margin in the branchial
region; neither of these features is seen on T. trispinosus.
Typilobus unispinatus was described (maRtinS-netO 2001:
243) as having an anterolateral spine and lacks spines on the
posterolateral corners. These features were not illustrated
by maRtinS-netO but rather were illustrated by a very
simple sketch by BeuRlen (1958: pl. 3, g. 4) wherein he
referred the specimen to Randallia sp. ? Placement of this
specimen in Typilobus is problematic.
Typilobus is known from Eocene to Miocene occurrences
in Europe, north Africa, the Middle East, Pakistan,
Turkmenistan, Japan, and Taiwan. One questionable
occurrence is reported from the Miocene of Brazil.
Family Matutidae de haan, 1841
Included fossil genera: Ashtoret Galil & claRK, 1994;
Eomatuta de anGeli & maRchiORi, 2009; Hemsut new
genus; Szaboa mülleR & Galil, 1998.
Diagnosis: Carapace not much wider then long, widest at
about midlength, regions undifferentiated, ornamented with
discrete tubercles not arranged in rows; lateral spine may
be present; anterolateral and posterolateral margins often
with discrete spines; front trilobate or with four spines;
posterior margin narrow; pereiopods 2-5 paddlelike; chelae
with few spines on upper margin and rows of tubercles on
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 7
outer surface; sternum obovate, narrow, widest anterior to
midlength; sterno-abdominal cavity extending onto sternite
4; abdominal somites 3-5 fused in males (after BellwOOd
1996; SchweitzeR & Feldmann 2000).
Discussion: The family does not have an extensive fossil
record. Recently, it has been summarized by SchweitzeR
et al. (2010) and SchweitzeR & Feldmann (2010b). Fossil
occurrences are limited to tropical and subtropical,
primarily carbonate environments. Extant taxa range from
the Red Sea, to the Indian Ocean, to the Indo-Pacic north
to Japan, and south to Australia, and also in West Africa
(Galil & claRK 1997).
Genus Hemsut nov.
Type species: Hepatiscus schweinfurthi nOetlinG, 1885.
Etymology: The genus is named for the Egyptian goddess of
fate and protection. This follows the lead of Galil & claRK
(1994) in naming matutid genera for female goddesses or
mythical beings and notes the occurrence of the genus in
Egypt. The gender is feminine.
Diagnosis: Carapace about as long as wide, maximally
wide about 40% the distance posteriorly; front sinuous;
fronto-orbital width about half maximum carapace width;
anterolateral margin initially straight, then followed by ve
blunt, small, but well developed spines, last spine nearly on
posterolateral margin; posterolateral margins long, centrally
with one spine that originates on the branchial regions
but projects laterally so that it could be considered as a
posterolateral spine; protogastric and mesogastric tubercles
small; cardiac and epibranchial tubercles large.
Discussion: The new genus is placed among the Matutidae
based upon its possession of a carapace that is about as
long as wide, poorly dened carapace regions with discrete
tubercles or swellings, and long, straight posterolateral
margins. Hemsut n. g. also bears supercial similarity
to many genera within Aethridae dana, 1851, such as
Eriosachila BlOw & manninG, 1996; Hepatiscus BittneR,
1875; and Matutites BlOw & manninG, 1996; however, in
Aethridae, the carapace is generally wider than long, the
carapace regions are better dened, and the posterolateral
margin is short and concave. In addition in most aethrids,
the suborbital margins do not extend beyond the upper
orbital margins (i.e. Hepatus latReille, 1802; Osachila
StimpSOn, 1871) as they do in matutids and Hemsut. The
placement of Hemsut in Matutidae could be made stronger
by recovery of pereiopods 2-5, which are paddle-like in
matutids and lanceolate in aethrids.
The recognition of Matutidae from the Eocene of Egypt
reinforces the known Tethyan distribution of the family
in both the fossil and Recent record. The other known
Eocene occurrence is Italy; the family is reported from
the Oligocene of France and the Miocene of Hungary and
Japan (SchweitzeR & Feldmann 2010b, for summary and
references).
Fig. 3. Hemsut schweinfurthi (nOetlinG, 1885) n. comb.,
SMNS 67861. 1 – dorsal carapace; 2 – anterior view; 3 –
right lateral view. Scale bars = 1 cm.
8 R.M. Feldmann et al.
Hemsut schweinfurthi (nOetlinG, 1885) n. comb.
Fig. 3
1885 Hepatiscus schweinfurthi NOetlinG, p. 488, pl. 4,
g. 1.
1929 H. Schweinfurthi nOetlinG, 1885. – GlaeSSneR, p.
210.
2010 H. schweinfurthi nOetlinG, 1885. – SchweitzeR et
al., p. 85.
Material examined: SMNS 67861.
Diagnosis: As for genus.
Description of material: Carapace ovate, slightly wider
than long (13.5 mm length; 14 mm width); length to posi-
tion of maximum width 6.2; dorsal carapace vaulted trans-
versely and longitudinally, regions weakly dened, cara-
pace surface ornamented by eight large tubercles.Rostrum
broken. Fronto-orbital width about half carapace width (7.4
mm). Orbits circular, directed anterolaterally and upward,
suborbital margin extending beyond upper-orbital margin,
with inner suborbital spine. Anterolateral margin convex,
initially straight, then followed by ve blunt, small, well de-
ned spines, last spine nearly on posterolateral margin. Pos-
terolateral margins straight, with long spine centrally that
originates on branchial region. Posterior margin narrow,
smooth, straight, 7.3 mm wide. Cervical groove shallow, U-
shaped, concave forward axially, becoming weakly convex
forward laterally. Protogastric regions weakly inated, with
a small tubercle centrally; mesogastric region with long an-
terior process, widened posteriorly, with one large tubercle
posteriorly. Hepatic region indistinct, depressed below level
of protogastric regions. Urogastric region depressed, poorly
dened; cardiac region pentagonal, bears largest tubercle
on carapace; intestinal region poorly dened, at. Epibran-
chial region poorly dened, with large tubercle, remain-
der of branchial region undifferentiated, with lateral spine
that extends as posterolateral spine. Pterygostomial region
smooth. Abdomen and appendages not preserved.
Discussion: The specimen described here bears close re-
semblance to the holotype of Hemsut schweinfurthi in size
and shape of the carapace, characteristic occurrence and
form of tubercles dening regions. The anterolateral mar-
gins are marked with ve equally spaced spines, being
widest at the third anterolateral spine, and being strongly
convergent posteriorly. The type of H. schweinfurthi and
the specimen described herein were collected from the
same locality and, according to nOetlinG (1885), the spe-
cies is not very rare. A total of four were found prior to
the specimen described here. There are, however, some dif-
ferences between the type specimen and the new material,
such as presence of spines at the posterolateral corners and
two small spines at the outer angle of orbital margin of the
holotype. nOetlinG (1885: 488) described the front of his
specimens as being wide, sinuous anteriorly, horizontally
projected and slightly sulcate. This is different from the il-
lustration (pl. 4, g. 1a), which shows a markedly anteriorly
projected, broadly arcuate front. Examination of the type
specimen will be necessary to reconcile these differences.
Absence of these features on the studied specimen may be
due to erosion of the margins.
Hemsut schweinf urthi bears supercial resemblance
to Eriosachila cuvillieri (Van StRaelen, 1929 [1930], also
from the Eocene of Mokattam, Egypt. Eriosachila cuvillieri
although not well illustrated, appears to t the diagnosis for
Eriosachila and the Hepatidae well in possessing a strongly
arcuate anterolateral margin with three spines and a concave
posterolateral margin. These features differentiate it from
Hemsut schweinfurthi.
Superfamily Majoidea SamOuelle, 1819
Family Majidae SamOuelle, 1819
Subfamily Majinae SamOuelle, 1819
Diagnosis: Carapace typically pyriform, widest in branchial
regions which are well developed; surface convex, often
highly ornamented. Front narrow, typically produced as a
bid rostrum, but may be truncate and downturned. Orbits
complete or nearly complete with supraorbital eave, often
an intercalated spine, and a postorbital lobe or spine; basal
antennal article does not form orbital oor. Anterolateral
margin may have strong spine. Epistome well developed
with quadrate buccal cavity. Legs spiny or granular;
chelipeds often shorter than walking legs. Pleon of males
and females with unfused somites. [Modied from daVie
(2002) and pOORe (2004)].
Discussion: SchweitzeR & Feldmann (2010b) discussed
the status of Micromaiinae BeuRlen, 1930, and concluded
that the three genera embraced within it by BeuRlen,
Micromaia BittneR, 1875; Mithracia Bell, 1858; and
Pisomaia lőRenthey in lőRenthey & BeuRlen, 1929, did
not exhibit a unique set of characters distinguishing them
from other subfamilies within Majidae. The latter two
genera have not been investigated in detail so that their
placement remains in question. Micromaia, as dened
by BittneR, can be assigned to Majinae with condence.
However, the concept of Micromaia has evolved since its
naming to permit inclusion of taxa that bear little or no
resemblance to the type species, M. tuberculata BittneR,
1875. As a result, SchweitzeR & Feldmann (2010b) erected
Planobranchia to embrace two species originally assigned
to Micromaia, M. laevis lőRenthey, 1907b, and M. simplex
Remy in GOROdiSKi & Remy, 1959.
Examination of type and illustrated material
representing the remaining species assigned to Micromaia
conrms that there are at least three species groups, two
of which do not conform to the original denition of the
genus. One of the disparate groupings is represented in
the Egyptian fossil decapod fauna and, as a result, a new
genus is proposed herein to embrace a specimen previously
referred to Micromaja tuberculata by lőRenthey (1898) as
well as two species from the Eocene of Spain.
Genus Tumidomaia nov.
Type species: Micromaia batalleri Vía, 1959.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 9
Included species: Tumidomaia batalleri (Vía, 1959),
as Micromaia; T. mokattamensis n. sp.; T. priabonensis
(Oppenheim, 1901), as Micromaia.
Diagnosis: Carapace pyriform; rostrum axially sulcate
with divergent rostral spines; orbits with prominent
supraorbital eaves; prominent spine on anterolateral margin
arising from hepatic region; carapace regions well dened
with metagastric and urogastric regions narrower than
mesogastric and cardiac regions; intestinal region relatively
long; epibranchial region bilobed; metabranchial region
with anteriorly directed extension separating mesobranchial
region from cardiac region.
Etymology: The generic name is derived from the Latin
tumidus meaning swollen, and Maia a common sufx
within Majidae.
Discussion: The new genus reects the signicant difference
in morphology exhibited by the three included species and
species of Micromaia as dened by BittneR (1875). Regions
on species of Tumidomaia are very clearly dened, the
mesogastric and urogastric regions are distinctly narrower
than the metagastric and cardiac regions, and the intestinal
region is relatively long. Micromaia is characterized by have
regions that are either indistinct or very weakly dened,
with axial regions of generally uniform width, and with an
intestinal region that is very short or not expressed. Further,
the bilobed epibranchial region and the anteriorly directed
extension of the metabranchial region are characters unique
to Tumidomaia. This combination of characters amply
justies creation of the new genus.
Tumidomaia batalleri, from the Eocene of Spain, is
clearly represented by the most complete, and best preserved
material (Fig. 4.1), exhibiting all the diagnostic characters of
the genus. Tumidomaia priabonensis, from the Priabonian
(Eocene) of Italy, is less completely preserved in the anterior
region, but it does exhibit a sulcate, bid rostrum, well
developed orbital eaves, and the conformation of regions is
diagnostic of the new genus (BeSchin et al. 2006: 106, pl. 2,
g. 7) (Fig. 4.3). Tumidomaia mokattamensis n. sp. is less
well preserved than specimens of the other two species, but
the regional development is as diagnosed for the genus. The
anterolateral spine arising from the hepatic region, the bid
rostrum, and the distinctive orbital eaves also are present.
Tumidomaia mokattamensis n. sp.
Fig. 4.2
1898 Micromaja tuberculata Bittn . – lőRenthey, p. 41.
1898 Micromaja tuberculata Bittn. – lőRenthey, p. 31,
pl. 2, g. 2 only.
Fig. 4. Tumidomaia spp. 1 – Tumidomaia batalleri (Vía,
1959) n. comb., cast of MGSB 23413; 2 –Tumidomaia
mokattamensis n. sp., holotype, SMNS 61862; 3 –
Tumidomaia priabonensis (Oppenheim, 1901) n. comb.,
photo courtesy A. de anGeli. Scale bars = 1 cm.
10 R.M. Feldmann et al.
1904 Micromaja tuberculata Bittn. – lőRenthey, p. 165
(in Hungarian).
1907a Micromaja tuberculata Bittn. – lőRenthey, p. 34.
1907b Micromaja tuberculata Bittn. – lőRenthey, p. 201
(in Hungarian).
1909 Micromaja tuberculata Bittn. – lőRenthey, p. 122.
Type: Holotype, SMNS 61862.
Etymology: The trivial name alludes to the site from which
the specimen was collected.
Diagnosis: As for genus.
Description: Carapace pyriform; moderately vaulted
transversely and longitudinally; regions tumid, moderately
well dened; surface uniformly pustulose.
Front narrow, 15% maximum width measured in bran-
chial region about 60% total length from front, axially
sulcate, with pair of short, stout spines curving adaxially.
Orbits with long, swollen eave lacking ssures; postorbital
spine may be present, poorly preserved. Anterolateral mar-
gin short, convex, bearing prominent, triangular spine aris-
ing from hepatic region. Posterolateral margin smoothly
convex with short, concave posterolateral corner where it
joins the broad, biconvex-forward posterior margin which is
56% maximum width.
Fig. 5. Planobranchia laevis (lőRenthey, 1907b), holotype, SMNS 67891. 1 – dorsal carapace; 2 –oblique anterior view of
right orbit and carapace; 3 – closeup of right orbit and rostrum. Scale bars = 1 cm.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 11
Mesogastric and protogastric regions conuent except
for pair of subtle post-rostral ridges dening margins of an-
terior process of mesogastric region. Margins of mesogas-
tric/protogastric regions dened by deep, V-shaped cervi-
cal groove extending to hepatic region and there curving in
concave-forward arc around hepatic region. Hepatic region
longer than wide, separated from axial regions by distinct
groove. Metagastric region wider than long, transversely
vaulted, with concave-forward anterior margin and nearly
straight lateral and posterior margins. Urogastric region
quadrate, lower and narrower than metagastric region. Car-
diac region circular, highest region on carapace, with two
prominent nodes at midlength. Intestinal region long, nar-
row, elevated slightly above metabranchial regions.
Epibranchial region dened by shallow grooves extend-
ing posterolaterally from cervical groove and anterolater-
ally to midpoint of cervical groove surrounding hepatic re-
gion. Mesobranchial region large, inated, elevated above
metabranchial region. Metabranchial region broad posteri-
orly with narrow anterior projections separating mesogas-
tric region from intestinal and cardiac regions.
Ventral surface, pleon, and appendages not preserved.
Measurements: Length 15.7 mm; maximum width 12.4
mm; frontal width 1.9 mm; fronto-orbital width 4.1 mm;
posterior width 7.0 mm.
Discussion: Tumidomaia mokattamensis is readily distin-
guishable from the other two species within the genus based
upon overall shape, development of tubercles, and details of
the rostral spines. Although all are united by the denitional
characters of the genus, T. batalleri has more slender rostral
spines than does the Egyptian species. In addition, the tu-
bercles are more densely spaced and more variable in size
than those on T. mokattamensis. The variation in size and
density of tubercles serves to distinguish the mesogastric
and protogastric regions from one another in the type spe-
cies, whereas they are difcult to distinguish on T. mokatta-
mensis. The tubercles on T. priabonensis are more coarsely
and densely spaced, the metagastric and urogastric regions
are shorter than those of T. mokattamensis.
Genus Planobranchia SchweitzeR & Feldmann,
2010b
Type species: Micromaja laevis lőRenthey, 1907b, by
original designation.
Included species: Planobranchia laevis (lőRenthey,
1907b), as Micromaja; P. simplex (Remy in GOROdiSKi &
Remy, 1959), as Micromaia.
Diagnosis: See SchweitzeR & Feldmann (2010b: 407).
Planobranchia laevis lőRenthey, 1907b
Fig. 5
1904 Micromaja laevis nov. sp. – lőR enthey, p. 165 (in
Hungarian).
1907a Micromaja laevis nov. sp. – lőRenthey, p. 35.
1907b Micromaja laevis nov. sp. – lőRenthey, p. 208, pl. 1,
g. 2 (in Hungarian).
1909 Micromaja laevis nov. sp. – LőRenthey p. 117, pl. 1,
g. 2 (in German).
1929 M. laevis löRenthey, 1909. – GlaeSSneR, p. 256.
2010 M. laevis lőRenthey, 1909. – SchweitzeR et al., p.
95.
2010b Planobranchia laevis (lőRenthey, 1909). –
SchweitzeR & Feldman n, p. 407, g. 2B.
Material examined: Holotype, SMNS 67891; cast of the
holotype, KSU D 403.
Emended description: Carapace longer (>22 mm) than
wide (16.4 mm), pyriform, regions moderately well dened.
Front narrow, poorly preserved; appears to be slightly down-
turned and axially sulcate. Upper orbital margin appears to
be a smooth eve; tiny postorbital spine visible on right orbit
(Fig. 5.3). Anterolateral margin short, weakly convex. Pos-
terolateral margin smoothly convex, rimmed, merging with
posterior margin which terminates in an axial obtuse angle.
Mesogastric and protogastric regions conuent, swol-
len, triangular with apex directed posteriorly, about 37%
maximum width, bearing two small spines at level of maxi-
mum width and a prominent transversely elongate axial
spine near posterior end of region. Gastric muscle forms
broadly concave forward arc around posterior termination
of mesogastric region. Metagastric region well dened as
concave forward, moderately swollen region with muscle
pits. Urogastric region narrow, equidimensional, bearing a
transversely elongated central node. Cardiac region promi-
nent, elevated to same level as mesogastric/protogastric re-
gions, bearing two nodes at widest point dened by a trans-
verse ridge extending to mesobranchial regions. Intestinal
region long, low, parallel-sided ridge bearing single medial
axial node. Hepatic regions prominently swollen, irregular-
ly nodose. Cervical groove deeply concave forward around
mesogastric region and gently convex forward posterior to
hepatic region. Epibranchial regions obliquely ovoid, bi-
lobed swellings bounded anteriorly by cervical groove and
posterolaterally by arcuate row of ne beads. Mesogastric
regions large, strongly inated, nely granular. Metabran-
chial region depressed below mesobranchial region, ir-
regularly ridged, punctate surface. Mesobranchial regions
separated from axial regions by smooth, axially convex
groove. Metabranchial region separated from axial regions
by smooth, arcuate groove.
Venter, pleon, and appendages not preserved.
Discussion: Planobranchia laevis differs from the only
other member of the genus, P. simplex, from Senegal, in
that the former exhibits relatively broader gastric and
cardiac regions, a more pronounced transverse ridge on the
cardiac region, a longer intestinal region, a granular surface
on the mesobranchial region, and a punctuate surface on
the metabranchial region, whereas that of P. simplex has
narrower axial regions, a more subtle transverse ridge
on the cardiac region, a nearly smooth surface on the
mesobranchial region, and an impunctate surface on the
12 R.M. Feldmann et al.
metabranchial region. The specimens of both species are
preserved as molds of the interior of the carapace so that the
comparison is direct.
Examination of the illustrations of the two species (Sch-
weitzeR & Feldmann 2010b: g. 2A, B) conrm that the
illustration of Planobranchia laevis was truncated along the
anterior side. Re-examination of the specimen upon which
the photo was based, reveals subtle, but distinct evidence of
a narrow, axially sulcate front with the termination missing
(Fig. 5.2) and some indication of the anatomy of the orbit
on the right side of the specimen. The presence of a supra-
orbital eave and a tiny postorbital spine are consistent with
placement in the Majinae. Other details of the front are not
preserved.
Fig. 6. Planobranchia? egyptensis n. sp., holotype, (BMNH) In. 59436. 1 – dorsal carapace; 2 – left lateral view; 3 – left
oblique anterior view; 4 – ventral view; 5 – abdominal somites. Scale bars = 1 cm.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 13
Planobranchia? egyptensis n. sp.
Fig. 6
Etymology: The trivial name egyptensis is in reference to
its discovery location in Egypt.
Type: Holotype, (BMNH) In. 59436.
Locality and horizon: The label indicates that the specimen
was collected from Egypt, limestone bed.
Diagnosis: Carapace sub-triangular, relatively unorna-
mented; branchial regions highly inated; cardiac region
well-developed, sub-hexagonal, with two sub-medial nodes;
intestinal region axially inated, depressed lateral to inat-
ed area, then a spherical inated area lateral to depressed
area.
Description: Carapace sub-triangular; maximum width
positioned at mid-branchial region, narrows toward anterior
end; moderately vaulted transversely and longitudinally;
branchial regions strongly inated. Carapace with scattered
tubercles, otherwise relatively smooth.
Rostrum and orbits not preserved; anterolateral margin
broken. Protogastric region moderately inated, with a
central, broad tubercle. Mesogastric region with anterior
narrow projection, widening posteriorly, with central
tubercle. Posterior to protogastric region and along lateral
margin hepatic or subhepatic region developed; otherwise,
posterior to protogastric and mesogastric regions, axial
area of carapace broadly depressed and laterally concave,
resulting in an arcuate, depressed metagastric region
and a narrow and depressed urogastric region bounded
laterally by deep grooves which extend posteriorly to bound
inated cardiac region and centrally inated intestinal
region. Cardiac region hexagonal with two tubercles;
intestinal region broad, axially inated. Branchial regions
undifferentiated; anteriorly with an arcuate depression
lateral to urogastric region; posteriorly with weak oblique
ridge extending from cardiac region posterolaterally to
posterolateral margin; ornamented with scattered tubercles.
Posterolateral margin strongly convex, with granular rim,
curving into nearly straight posterior margin, which then
curves rather abruptly into a posterior projection that meets
the pleon.
Very little of ventral surface preserved. Sternites 1 and 2
small, fused, triangular; fused to sternite 3, with transverse
ridge at suture. Sternite 4 longer than wide, with apparent
concavity anteriorly. Five pleonites visible, rst somite
apparently obscured by carapace; each pleonite raised in
center with transverse ridge at posterior margin; pleonites
increasing in length posteriorly so that sixth is longest.
Margins of pleurae straight, sub-parallel to medial plane.
Discussion: Almost all of the margins are broken on the
specimen, and the critical areas of the rostrum and orbits
are not preserved. The relatively smooth carapace, the
depressed area bounding the axial regions, and the well-
dened and undifferentiated branchial regions are similar
to species of Planobranchia, so we tentatively assign
the species to that genus. We acknowledge that there are
several differences between the new species and species
of Planobranchia. Planobranchia is characterized by
a more inated urogastric and metagastric region, and
the metagastric region is narrower in other species of
Planobranchia than in P.? egyptensis. Planobranchia?
egyptensis lacks the lateral ridges extending from the cardiac
region as in other species of the genus, and it has much more
laterally inated branchial regions and a straighter posterior
margin with a posterior extension, not seen in other species
of Planobranchia. However, because the new material is
incomplete, we hesitate to refer it to a new genus or any
other genus at this time until more complete material can be
collected. Planobranchia is well-known from Eocene rocks
of Africa, so the referral does not extend the range of the
genus.
Superfamily Cancroidea latReille, 1802
Family Cheiragonidae ORtmann, 1893
Genus Karasawaia VeGa et al., 2008
Type species: Plagiolophus markgra lőRenthey, 1907b,
by original designation.
Diagnosis: Carapace quadrate, about as wide as long;
anterolateral margin with at least three spines excluding
outer-orbital spine; posterior margin entire; posterior
margin wide, entire, rimmed; protogastric regions wide,
with arcuate ridge posteriorly; hepatic region bilobate,
composed of two triangular elements; mesogastric,
metagastric, urogastric, and cardiac regions well-dened,
cardiac region with transverse ridge; epibranchial region
arcuate, composed of at least two segments; mesobranchial
region inated; metabranchial region attened, continuous
with intestinal region.
Karasawaia markgra (lőRenthey, 1907b)
Fig. 7
1904 Plagiolophus markgra nov sp. – lőRenthey, p. 165
(in Hungarian).
1907a Plagiolophus markgra nov. sp. – lőRenthey, p. 35.
1907b Plagiolophus markgra nov. sp. – lőRenthey, p.
225, pl. 1, g. 5 (in Hungarian).
1909 Plagiolophus markgra nov. sp. – lőR enthey, p.
135, pl. 1, g. 5 (in German)
1929 P. Markgra löRenthey, 1909. – GlaeSSneR, p. 329.
2004 Stintonius markgra (lőRenthey). – KaRaSawa &
SchweitzeR, p. 146, g. 1.7.
2008 Karasawaia markgra (lőRenthey, 1907 [1909]). –
VeGa et al., p. 56.
2010 K. markgra (lőR enthy, 1907 [German version
1909]). – SchweitzeR et al., p. 103.
Material examined: SMNS 67892, holotype; cast of
holotype, KSU D 401.
Discussion: KaRaSawa & SchweitzeR (2004) revised
14 R.M. Feldmann et al.
Glyphithyreus (= Plagiolophus) and placed Plagiolophus
markgra within Stintonius cOllinS, 2002 with some
reservations. Later, VeGa et al. (2008) erected the genus
Karasawaia to accommodate both P. markgra as well
as new specimens they recovered from Eocene rocks
of Mexico, which they referred to P. markgra. There
appear to be sufcient differences between the Mexican
material and the type material from Egypt to suggest that
the Mexican specimens warrant at least a different species.
The anterolateral margins appear to be much longer in the
Mexican material, although this is difcult to determine
because it appears that the specimen is broken and displaced
or sheared, because the anterolateral margins are not
symmetrical (VeGa et al. 2008, pl. 2, g. 13). Examination
of the Mexican material will be necessary to determine
appropriate placement for it. In any case, the Egyptian
species is the type for Karasawaia, and we illustrate a cast
of the holotype of the type species here, the rst time that
we know of that the actual specimen has been illustrated
rather than a drawing (de lőRenthey, 1909, pl. 1, g. 5).
Genus Montezumella RathBun, 1930
Type species: Montezumella tubulata RathBun, 1930, by
subsequent designation of GlaeSSneR (1969).
Included species: See SchweitzeR et al. (2010).
Montezumella fraasi (lőRenthey, 1907b)
Fig. 8
Material examined: SMNS 67893/1, holotype; 67893/2,
gured specimen (lőRenthey 1909, pl. 2, g. 3a); cast of
holotype, KSU D 404.
Diagnosis: Carapace ovate, longer than wide; front
quadrilobed, lobes with tiny spines or serrations; anterolateral
margin with four spines; posterolateral margin entire;
carapace regions moderately well developed, attened,
ornamented with granules or scabrous rows of granules;
axial regions particularly well-marked; epibranchial region
composed of two or three segments arranged in an arc.
Discussion: Montezumella is a distinctive genus within
Cheiragonidae that was summarized by SchweitzeR &
SalVa (2000). Herein we illustrate a cast of the holotype of
Montezumella fraasi lőRenthey, 1907b, originally referred
to Cancer (Fig. 8; illustrated by lőRenthey 1909, pl. 2,
g. 3b). Note that the specimen is longitudinally sheared
somewhat; however, it exhibits the diagnostic ornamentation
and regional development of the genus.
Superfamily Xanthoidea macleay, 1838
Family Xanthidae macleay, 1838
Diagnosis: Carapace hexagonal or transversely ovate;
carapace length/width falling into two groups, length either
about two-thirds to three quarters or 90-100% maximum
carapace width, widest two-thirds to three quarters
the distance posteriorly except in some Xanthinae and
Polydectinae in which position of maximum width is before
Fig. 7. Karasawaia markgra (LőRenthey, 1907b), cast of
holotype (SMNS 67892), cast numbered KSU D 401.
Fig. 8. Montezumella fraasi (LőRenthey, 1907b), cast of
holotype (SMNS 67893/1), cast numbered KSU D 404.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 15
the midlength; regions usually well-dened but dorsal
carapace can be smooth; frontal margin bilobed, with axial
notch, ranging from one-quarter to about 40% maximum
carapace width; fronto-orbital width generally about half to
two-thirds maximum carapace width; anterolateral margins
with between two and six lobes or spines; anterolateral
margin well-differentiated from posterolateral margin,
anterolateral margin often strongly convex; posterolateral
margin often concave. Modied after daVie (2002). The
following diagnosis is quoted from pOORe (2004: 460)
and includes only those characters of the dorsal carapace.
“Carapace hexagonal, transversely oval or sometimes
subcircular; dorsal regions generally well-dened; surface
usually ridged or granular. Frontal margin bilobed.
Anterolateral margins generally with two to six teeth or
lobes, sometimes weakly differentiated.”
Subfamily Actaeinae alcOcK, 1898
Diagnosis: The following diagnosis is quoted from pOORe
(2004: 463) and includes only those characters of the dorsal
carapace. “Carapace with areas well separated, separate [sic]
by deep grooves, with mosaic-like tubercles or squamiform
or nodiform tubercles, without setae; posterolateral margins
nearly straight.”
Discussion: In addition to the diagnosis, the systematic
key (pOORe 2004: 462) includes the observations that the
carapace of Actaeinae has three spines on the anterolateral
margin and a weakly projecting front, both characters
reinforcing the denition of the group and supporting
placement of Lambropsis in the subfamily.
Genus Lambropsis lőRenthey, 1907b
Type species: Lambropsis wanneri lőRenthey, 1907b, by
monotypy.
Included species: Lambropsis wanneri lőRenthey, 1907b.
Diagnosis: Carapace wider than long, length 80% maximum
width. Front axially sulcate; frontal width 26% maximum
width; orbits transversely ovate with two small supraorbital
protuberances, a larger outer orbital spine, and a blunt
suborbital spine; fronto-orbital width 51% maximum width.
Anterolateral margin with three spines, last one largest.
Posterolateral margin convex anteriorly, bearing a single
spine, becoming concave where it meets straight posterior
margin. Regions well delimited; protogastric region very
large, intestinal region depressed; cervical groove deep,
continuous. Surface with blunt nodes or spines.
Discussion: Assignment of the genus to suprageneric cate-
gories has been difcult. lőRenthey (1909) considered it to
be referable to the Parthenopidae macleay, 1838; however,
the conformation of the front and orbits, the development
of the cervical groove, denition of regions, and form of
the posterior margin are inconsistent with denitions of that
family (RathBun 1925; GlaeSSneR 1969; daVie 2002; pOORe
2004). GlaeSSneR (1969: R532) considered it to be “of un-
certain systematic position or status.” Further, he indicated
that BeuR len (1930) had considered the genus to be a xan-
thid. SchweitzeR et al. (2010) placed it in incertae sedis.
The genus is herein referred to Actaeinae within Xanthi-
dae based upon examination of systematic keys and diagno-
ses in daVie (2002) and pOORe (2004). Lambropsis exhibits
all the characters of the dorsal carapace that are diagnostic
of that subfamily. Absent the ventral surface of the cara-
pace, this seems to be the best placement at this time.
Lambropsis wanneri lőRenthey, 1907b
Fig. 9
1904 Lambropsis wanneri nov. gen. Wanneri nov. sp. –
lőRenthey, p. 165 (in Hungarian).
1907a Lambropsis wanneri nov. gen. Wanneri nov. sp. –
LőRenthey, p. 35
1907b Lambropsis wanneri nov. gen. Wanneri nov. gen. –
LőRenthey, p. 213, pl. 1 , g. 1 (in Hungarian).
1909 Lambropsis Wanneri. – LőRenthey, p. 124, pl. 1 ,
g. 1 (German version).
Material examined: Holotype, SMNS 67894; cast of the
holotype, KSU D 402.
Diagnosis: As for genus.
Description: Carapace wider (17.1 mm) than long (13.6
mm), length 80% maximum width; greatest width at
spines on anterolateral corner, 75% total length from front;
strongly vaulted longitudinally, particularly in front half;
weakly vaulted transversely.
Front narrow (4.4 mm), projected beyond orbits; about
26% maximum width, axially sulcate, bounded by short,
rounded projections. Orbits transversely ovoid, directed
forward; fronto-orbital width (8.7 mm) 51% maximum
width. Upper orbital margin sinuous with inner and medial
projections dened by broad, shallow reentrants; outer
orbital spine directed anteriorly; suborbital spine broad,
short. Eye occupies entire orbit.
Anterolateral margin weakly convex, bearing three
spines excluding outer orbital spine. Anteriormost two spines
broad, short, triangular. Posteriormost spine at anterolateral
corner longer, triangular, prominent. Posterolateral margin
as long as anterolateral margin, convex anteriorly with
single, broad, triangular spine and becoming concave near
intersection with wide (6.0 mm), straight, posterior margin.
Frontal area as wide as frontal margin and extending
posteriorly to level of posteriormost extent of orbits; weakly
elevated and separated by axial sulcus. Protogastric regions
large, triangular, width (11.1 mm) 65% maximum width,
swollen, dening highest points on carapace; separated
from long, narrow pyriform mesogastric region by broad,
smoothly shallow grooves. Hepatic region small, bearing
two nodes including rst anterolateral spine. Cervical
groove deep, smooth, straight and inclined posteromesially
from lateral margin to corners of mesogastric region and
broadly concave forward medially. Metagastric region as
wide as mesogastric region, short, well dened laterally,
poorly dened at intersections with small, narrow urogastric
16 R.M. Feldmann et al.
region. Cardiac region prominent, pentagonal, with three
spines arrayed as triangle with apex directed posteriorly.
Intestinal region not dened as it crosses long, smooth,
sloping surface as wide as posterior margin. Epibranchial
region small, triangular, bearing three spines including last
two anterolateral spines. Mesobranchial region narrows
axially, inated. Metabranchial region generally circular,
set below level of mesobranchial region, bearing two blunt
protuberances at approximate level of anterior pair of
cardiac spines.
Carapace surface coarsely pitted which appears to be
hollow nodes with broken tips.
Venter, pleon, and appendages not preserved.
Discussion: The genus contains a single species; it bears
relatively strong resemblance to some species of Actaea de
haan, 1833; however, as dened Actaea has a broader front,
about 33% maximum width, whereas the front of Lambropsis
is about 26% maximum width. Because ornamentation on
species within Actaea and other genera within the subfamily
is quite variable, it is reasonable to conclude that the densely
nodose surface of L. wanneri would fall well within the
range of variation of ornamentation seen in Actaeinae.
Superfamily Carpilioidea ORtmann, 1893
Family Carpiliidae ORtmann, 1893
Included genera: Bryocarpilius n. g.; Carpilius
deSmaReSt, 1823; Eocarpilius BlOw & manninG, 1996;
Holcocarcinus witheRS, 1924; Laticarpilius n. g.; Liopsalis
VOn meyeR, 1862; Ocalina RathBun, 1929; Palaeocarpilius
A. milne-edwaRdS, 1862; Paraocalina BeSchin et al., 2007;
Proxicarpilius cOllinS & mORRiS, 1978.
Discussion: SchweitzeR (2003) revised the family as it
occurs in the fossil record, and a few genera and species have
been added since (SchweitzeR et al. 2010). Reexamination
of the Egyptian material, some of which was originally
described by lőRenthey (1909) and some of which was
labeled by him and deposited in the SMNS, suggests
that the species that have been historically assigned to
Palaeocarpilus A. milne-edwaRdS, 1862, might be better
reassigned to new genera (Table 2).
Carpiliidae is a largely extinct family but the sole living
relative, Carpilius, has a nearly cosmopolitan distribution
(Fig. 11, data from wetzeR et al. 2003), mainly in shallow
tropical and subtropical waters. Comparison of the
distributions of members of the family over geologic time
(Figs. 10-11) shows that ecological preferences appear to
have changed little over the past 50 million years.
Genus Palaeocarpilius A. milne-edwaRdS, 1862
sensu stricto
Type species: Cancer macrochelus deSmaReSt, 1817, by
subsequent designation of GlaeSSneR (1929).
Included species: Palaeocarpilius aquitanicus A. milne-
edwaRdS, 1862; P. ignotus A. milne-edwaRdS, 1862 (nger
only); P. intermedius StuBBleField, 1927; P. laevis imaizumi,
1939; P. macrochelus; P. mississippiensis (RathBun, 1935);
P. rugifer StOliczKa, 1871; P. valrovinensis (de GReGORiO,
1895).
Diagnosis: Carapace wider than long, length about 75%
maximum width; widest about 70% the distance posteriorly,
smooth; front downturned, triangular, may have a small
axial sulcus at tip, about 40% maximum carapace width;
orbits small, circular, entire, fronto-orbital width about
60% maximum carapace width; anterolateral margins
long, very tightly convex, with seven to nine spines or
blunt projections including outer-orbital spine, with well-
developed ridge extending onto dorsal carapace from last
Fig. 9. Lambropsis wanneri (LőRenthey, 1907b), cast of
holotype (SMNS 67894), cast numbered KSU D 402. 1 –
dorsal carapace; 2 – oblique anterior view. Scale bars = 1
cm.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 17
anterolateral spine; posterolateral margin initially at about
80º angle, then becoming more gentle, at about 60º angle to
axis; chelae generally massive, with spines on upper margin
(modied after SchweitzeR 2003).
Discussion: Examination of the species currently referred
to Paleocarpilius (SchweitzeR et al. 2010) indicated that the
range of variation within the genus has gotten very broad,
with the genus embracing species with anterolateral margins
ranging from entire to having nine spines; triangular and
entire fronts to those with four blunt spines; length/width
ratios of 65-78%, and nearly straight posterolateral margins
to those composed of two distinct segments. We herein
restrict Palaeocarpilius to those species with a triangular
Fig. 10. Eocene and Oligocene occurrences of Carpiliidae. Br = Bryocarpilius, C = Carpilius, E = Eocarpilius, H =
Holcocarcinus, La = Laticarpilius, Li = Liopsalis, O = Ocalina, Pa = Palaeocarpilius, Po = Paraocalina, Pr = Proxicarpilius.
Base maps from ScOteSe (2001) with data from SchweitzeR et al. (2010) and references therein.
18 R.M. Feldmann et al.
front, at least seven anterolateral spines, a length to width
ratio of about 75%, and a posterolateral margin that is
composed of two segments, initially at about 80º angle,
then becoming more gentle, at about 60º. Other species with
less ornate margins or different carapace or margin shapes
have been referred to other genera. This rearrangement
should permit easier identication and placement of new
specimens within the various genera of Carpiliidae; we
have provided a table of distinguishing characters to assist
with identication (Table 3).
Palaeocarpilius brodkorbi (lewiS & ROSS 1965) was
reported from the Eocene of Florida. Examination of
Fig. 11. Miocene and Recent occurrences of Carpiliidae. C = Carpilius, E = Eocarpilius, Pa = Palaeocarpilius. Base maps
from ScOteSe (2001, 2006) with data from SchweitzeR et al. (2010) and references therein.
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 19
the holotype (USNM 649417) suggests that it might be
better placed within Paraocalina BeSchin et al., 2007.
Palaeocarpilius brodkorbi possesses a clearly quadrilobed
front, not a triangular front as in Palaeocarpilius sensu
stricto. In addition, it is wider overall than Palaeocarpilius
and the position of maximum width is further anteriorly
than in Palaeocarpilius spp. Palaeocarpilius brodkorbi
possesses about six weak lobes on the anterolateral margins,
not seven to nine spines or lobes as in Palaeocarpilius
spp. Paraocalina has a carapace size and shape much
more similar to that of P. brodkorbi (Table 3) and it has
a quadrilobed front as in P. brodkorbi. In the type species
of Paraocalina, there are ve anterolateral lobes. Thus,
it seems that Paraocalina is a better t for this species,
resulting in the new combination Paraocalina brodkorbi
Vía (1941) named Menippe almerae for moderately well-
preserved material. Examination of the holotype (MGSB
8886) suggests that the coxa does not articulate directly
with the merus, although the specimen is not well enough
preserved to make this observation unequivocal. The front
is quadrilobed, the anterolateral margins are ornamented
with at least 6 spines, and the surface is uniformly punctate.
Thus, the best placement for this species is Harpactocarci-
nus A. milne-edwaRdS, 1862, within Zanthopsidae, which
can accommodate these features (SchweitzeR 2003), result-
ing in Harpactocarcinus almerae new combination.
Table 2. All species referred by SchweitzeR et al. (2010) to Palaeocarpilius and their current placement. Data from
GlaeSSneR (1929) and species references.
Species (under original generic designation) Current Placement Geographic Occurrence Age
Palaeocarpilius macrochelus Palaeocarpilius (type species) Italy, Hungary, Romania, Egypt? Eocene, Oligocene
P. aquitanicus Palaeocarpilius France Oligocene
P. ignotus Palaeocarpilius France Eocene (Bartonian)
P. intermedius Palaeocarpilius Tanzania (Zanzibar) early Miocene
P. laevis Palaeocarpilius Japan early Miocene
P. mississippiensis Palaeocarpilius Mississippi, USA late Eocene
P. rugifer Palaeocarpilius western India late Eocene
P. valrovinensis Palaeocarpilius Italy Eocene
Laticarpilius aegypticus n. sp. Laticarpilius Egypt Eocene
P. bispinosus Laticarpilius eastern India Eocene
Cancer klipsteini Liopsalis (type species) Germany Eocene
P. anodon Liopsalis Italy Eocene
P. simplex Liopsalis western India Eocene
P. aquilinus Bryocarpilius (type species) Libya Eocene
Bryocarpilius apsidorsalis n. sp. Bryocarpilius Egypt Eocene
P. brodkorbi Paraocalina Florida, USA Eocene
Menippe almerae Harpactocarcinus Spain Eocene
Table 3. Ratios of measurements and important differentiating features of genera among the Carpiliidae. L = carapace
length, W = carapace width, FW = frontal width, FOW = fronto-orbital width, L2 = length from anterior of carapace to
position of maximum carapace width, Q = quadrilobed, T = triangular, AM = anterolateral margins, PLM = posterolateral
margins. *denotes measurements taken from illustrations. **denotes measurements provided by original author. *** axially
two lobes may be fused so that front is trilobed.
Genus Front L/W L2/L FW/W FOW/W AM Other distinguishing features
Bryocarpilius n. g. Q, T 66 60 36 50 8-9 spines Strongly longitudinally vaulted
Carpilius Q*** 76 51 24 38 1 blunt Broadly convex PLM
Eocarpilius Q 77 60 38 55 Entire PLM nearly straight
Holcocarcinus* T 69 46 40 54 2 blunt Two transverse carapace ridges
Laticarpilius n. g. T 60-70 55-65 45-50 55-60 2 blunt AM composed of two arcs
Liopsalis* T 70-78 55-67 36-50 60-70 Entire
Ocalina Q 63-73 60 35-40 50-55 > 9 blunt Large spherical carapace ornamentation
Palaeocarpilius T 75 70 40 60 7-9 spines
Paraocalina Q 65** -- 45* 60** 5+ lobes PLM nearly straight
Proxicarpilius T 73 50 40 65 4 spines Axial regions moderately dened
20 R.M. Feldmann et al.
Genus Laticarpilius nov.
Type species: Laticarpilius aegypticus n. sp., by original
designation.
Included species: Laticarpilius aegypticus n. sp.;
L. bispinosus (SatSanGi and chanGKaKOti, 1989), as
Palaeocarpilius; probably specimens of Palaeocarpilus
simplex sensu Vía (1969).
Etymology: The generic name is a combination of the
Latin word latus, meaning broad, and the generic name
Carpilius, a common stem within Carpiliidae, referring to
the very wide carapace in this genus and the very low ratio
of carapace length to width.
Diagnosis: Carapace ovoid, length 60-70% maximum
width, widest 55-65% the distance posteriorly; rostrum
triangular, downturned, frontal width 45-50% carapace
width; orbits small, circular, rimmed, fronto-orbital width
55-65% maximum carapace width; anterolateral margin
composed of two arcs, one gentle, then becoming more
strongly convex in posterior one-third, with two broadly
separated blunt projections that mark the beginning of the
inection in convexity and the anterolateral angle; last blunt
projection extending into rim along posterolateral margin
and onto dorsal carapace; posterolateral margin initially at
80º to axis, then curving at 60º to axis; chelae stout, smooth.
Discussion: Palaeocarpilius bispinosus from the middle
Eocene of eastern India as well as a new species described
here differ from Palaeocarpilius sensu stricto in possessing
an anterolateral margin with two segments, only two
anterolateral spines instead of seven to nine, and a much
longer than wide carapace, with a length to width ratio of
under 70% as compared to 75% (Table 3). Thus, we erect
a new genus, Laticarpilius, to embrace these two species.
Examination of the illustrations of Eocene specimens from
Spain that Vía (1969) referred to Palaeocarpilius simplex
StOliczKa, 1871, indicates that these specimens are not
referable to that species and are probably better referred to
a species within Laticarpilius. Palaeocarpilius simplex has
entire anterolateral margins, whereas the specimens of Vía
(1969) appear to have two blunt anterolateral projections and
a margin with two segments. Examination of the specimens
will be needed to conrm this and formalize the placement.
Laticarpilius aegypticus n. sp.
Fig. 12
1904 Palaeocarpilius simplex StOl. – lőRenthey, p. 165
(in Hungarian).
1907a Palaeocarpilius simplex StOl. – lőRenthey, p. 35.
1907b Palaeocarpilius simplex StOl. – lőRenthey, p. 216,
pl. 1, gs. 3-4 (in Hungarian).
1909 Palaeocarpilius simplex StOliczKa. – lőRenthey, p.
127, pl. 1, gs. 3-4 (German version).
Types: The holotype is herein designated as SMNS
67895/1, a male, illustrated by lőRenthey (1907b, 1909: pl.
1, g. 3). Four paratypes include SMNS 67895/2, illustrated
by lőRenthey (1907b, 1909: pl. 1, g. 4), mature female;
SMNS 61863, a mature male; SMNS 61864, a male, and
SMNS 61863, an immature female. A fth paratype, USNM
Acc. 167630, is deposited in USNM, collected from the
Fayum Desert, Egypt, in 1907. Rocks exposed there include
marine and non-marine units (Van cOuVeRinG & haRRiS
1991; GinGeRich 1992); without a specic locality, it seems
reasonable to suggest a middle to late Eocene age.
Etymology: The trivial name is derived from the Latin
Aegyptus, referring to the type locality.
Diagnosis: Carapace ovoid, length 65% maximum width,
widest about 65% the distance posteriorly; rostrum
triangular, downturned, frontal width about 45% carapace
width; orbits small, circular, rimmed, fronto-orbital width
about 55% maximum carapace width; anterolateral margin
composed of two arcs, one gentle, then becoming more
strongly convex in posterior one-third, with two broadly
separated blunt projections that mark the beginning of the
inection in convexity and the anterolateral angle, last blunt
projection extending into rim along posterolateral margin
and onto dorsal carapace; posterolateral margin initially at
80º to axis, then curving at 60º to axis; chelae smooth.
Description: Carapace ovoid, wider than long, length 65%
maximum width, widest about 65% the distance posteriorly
on carapace; strongly vaulted longitudinally, moderately
convex transversally; carapace generally smooth, regions
and grooves generally not dened, cuticle preserved as
white, chalky material, variously punctate, granular, or
smooth.
Rostrum triangular, extending well beyond orbits, 45%
maximum width, strongly downturned, entire; orbits small,
sub-circular, deeper laterally than axially, directed weakly
anterolaterally, entire, fronto-orbital width about 55%
maximum carapace width. Anterolateral margin gently
convex, rounded, smooth for rst two-thirds of length,
then arcing more strongly convexly for posterior one third;
margin itself thin so that posterior one third is broken on
all available specimens; with two broadly separated blunt
projections that mark the beginning of the inection in
convexity and the anterolateral angle, last blunt projection
extending into rim along posterolateral margin and onto
dorsal carapace. Posterolateral margin initially concave,
at about 80º angle to axis of animal, thickly rimmed,
rim extending onto carapace; then arcing more strongly
posteriorly making about 60º angle to axis. Posterior margin
straight, 32% maximum width, rimmed.
Regional denition very weak; arcuate array of pits
in hepatic region parallel to anterolateral margin; cardiac
region dened as a subtle elevated area, urogastric area
anterior to cardiac region and anterior-most cardiac
region laterally bounded by two weakly dened grooves,
probably vestigial remnant from either postcervical or
branchiocardiac grooves.
Other regions and grooves not dened.
Male rst and second sternites fused; third sternite wid-
er than long, separated from sternite 2 by complete suture,
separated from sternite 4 by marginal notches; sternite 4
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 21
Fig. 12. Laticarpilius aegypticus n. g., n. sp. 1-2 – Holotype SMNS 67895/1, male, dorsal carapace (1) and ventral surface
showing abdomen (2); 3-4 – paratype SMNS 67895/2, female, dorsal carapace (3) and ventral surface showing abdomen
(4); 5-8 – paratype SMNS 61863, male, dorsal carapace (5), ventral surface showing abdomen (6), anterior view showing
triangular, downturned front (7), and right lateral view showing moderate longitudinal convexity of carapace (8). Scale bars
= 1 cm.
22 R.M. Feldmann et al.
long, sub-rectangular, with long episternal projections; ster-
nites ve and 6 about as long as wide; sternite 7 barely vis-
ible; sternite 8 not visible. First male pleonite very short,
much wider than long; second pleonite longer than rst,
wider than long; third, fourth, and fth pleonites fused, third
pleonite widest, extending beyond other three into rounded
lobes, weak demarcations visible between pleonites 3/4 and
4/5; sixth pleonite longest, longer than wide; telson about as
long as wide, extending to about one-third the distance ante-
riorly along the coxae of pereiopod one; male pleon entirely
lling the distance between coxae 5. Female pleon wide,
nearly covering sternum, all somites free, becoming longer
posteriorly so that sixth is longest.
Chelipeds moderately heterochelous; merus and coxa
articulating directly; merus longer than high; coxa short,
smooth; manus stout, longer than high, becoming higher
distally; xed nger with black tips, with some denticles on
occlusal surface; movable nger nearly entirely black, with
large blunt molariform tooth near proximal end on right
chela. Pereiopods 2-5 much more slender than pereiopod 1.
Carapace slightly deformed by dorso-ventral
compression.
Measurements: Measurements taken on specimens of
Laticarpilius aegypticus n. sp. are presented in Table 4.
Discussion: The presence of two very small, blunt
projections on the anterolateral margin is weak, but can
be conrmed. There is a swelling at the point of inection
of the convexity of the anterolateral margin, and there is a
swelling at the anterolateral corner, at the end of the ridge.
However, the broken nature of the anterolateral margins
makes it difcult to see the presence of these projections
without viewing the specimens under a microscope as they
are visible as swellings in the cuticle, often with breaks
around them. Interestingly, handedness in the species is
about evenly distributed among specimens. Two are left
handed and three are right handed. In general, most studies
of handedness in carpilioid brachyurans have indicated
tendencies toward right handedness (SchweitzeR &
Feldmann 2010a: 176).
Genus Liopsalis VOn meyeR, 1862
Type species: Cancer klipsteini VOn meyeR, 1842, p. 589,
by monotypy.
Included species: Liopsalis anodon (BittneR, 1875), as
Palaeocarpilius; L. klipsteini; L. simplex (StOliczKa, 1871),
as Palaeocarpilius.
Diagnosis: Carapace ovate, wider than long, length
70-78% maximum carapace width, widest 55-67% the
distance posteriorly at position of anterolateral corner; front
triangular, downturned, weakly sinuous, 36-50% maximum
carapace width; orbits circular, directed forward, 60-70%
maximum carapace width; anterolateral margins entire,
tightly convex; rim along posterolateral margin and onto
dorsal carapace, posterolateral margin initially at 80º to
axis, then curving at 60º to axis; upper margin of chela with
stout spines.
Discussion: VOn meyeR (1842) described Cancer klipsteini
from the Eocene of Germany and later referred that species
to a new genus, Liopsalis VOn meyeR, 1862 (p. 163). In that
same year, A. milne-edwaRdS (1862) referred the species to
Palaeocarpilius. The species has been referred to Palaeo-
carpilius ever since, and GlaeSSneR (1929: 291) included
Liopsalis as a synonym of Palaeocarpilius, although he
did not include it as a synonym in the Treatise (GlaeSSneR
1969). Vía (1969) used Liopsalis as a subgenus of Palaeo-
carpilius to embrace those species with entire anterolateral
margins. Examination of both VOn meyeR’s (1862) and A.
milne-edwaRdS’ descriptions and illustrations of Cancer
klipsteini suggests that whereas it is very similar to Palaeo-
carpilius spp. in terms of its overall carapace shape and its
triangular front (Table 3), C. klipsteini has an entire antero-
lateral margin, completely lacking spines or nodes, a major
differentiating character within Carpiliidae. Thus, we sug-
gest going a step further than Vía (1969), elevating Liopsalis
VOn meyeR, 1862, to generic level to accommodate this spe-
cies. Cancer klipsteini VOn meyeR, 1842, is the type species
of Liopsalis, as it was the sole species for which VOn meyeR
(1862) erected the genus.
Two other species appear to be referable to Liopsalis,
based upon the descriptions and illustrations in the original
publications. Palaeocarpilius simplex, described from the
Eocene of western India, is based upon incomplete material
that was described as having an entire anterolateral mar-
gin and triangular front. Palaeocarpilius anodon, from the
Eocene of Italy, was illustrated similarly, so much so that
it was eventually synonymized with P. simplex. Herein we
reinstate P. anodon as it has different carapace ratios from
P. simplex and is known from quite a different geographic
Table 4. Measurements and ratios taken on species of Laticarpilius aegypticus n. gen., n. sp. *denotes measurements
taken from photograph. L = carapace length, W = maximum carapace width (approximate, specimens are broken in this
region), FW = frontal width, FOW = fronto-orbital width, PW = posterior width, L2 = length to approximate position of
maximum width (specimens are broken in this region), H = handedness of rst pereiopod.
Specimen L W FW FOW PW L2 Gender H
SMNS 67895/1 (Holotype) 62.5 98.1 43.9 52.6 28.1 48.0 Male Left
SMNS 67895/2 53.9 87.0 40.8 49.8 27.4 44.1 Female Left
SMNS 61863 67.2 104.7 45.2 56.1 30.1 45 Male Right
SMNS 61864 53.3 79.3 36.8 46.2 26.9 37 Male Right
SMNS 61865 53 87.0 --- 49.8 30.8 33 Female Right
USNM Acc. 167630* 57 87.0 41 50 -- 37 Unknown Unknown
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 23
area. The smooth, arcuate, entire anterolateral margins of
both of these species suggests that they are better placed
within Liopsalis than Palaeocarpilius. The range of Liop-
salis with all three species included would be the Eocene of
Italy, eastern India, and Germany.
Genus Bryocarpilius nov.
Type species: Palaeocarpilius aquilinus cOllinS &
mORRiS, 1973.
Included species: Bryocarpilius apsidorsalis n. sp.; B.
aquilinus (cOllinS & mORRiS, 1973) n. comb.
Etymology: The genus name is derived from the Greek
word bryo, meaning swell, and the genus name Carpilius, a
common stem within Carpiliidae, referring to the extremely
vaulted carapace in this genus. The gender is masculine.
Diagnosis: Carapace ovate, length about two-thirds maxi-
mum width, widest about 60% the distance posteriorly; very
strongly vaulted longitudinally; front quadrilobed, weakly
projected into a sinuous triangular shape, with two closely
spaced axial lobes and blunt inner orbital lobes, about 36%
maximum width, fronto-orbital width about half maximum
carapace width; anterolateral margin with eight or nine even-
ly spaced spines including outer orbital spine; posterolateral
margin rimmed anteriorly, sometimes with one small spine,
at 80º angle to axis, then arcing posteriorly at about 60º angle
to carapace; mesogastric region weakly to moderately de-
ned, sometimes with weak arcuate ridge; chela with row of
spines on upper surface, distal half of ngers black.
Discussion: cOllinS & mORRiS (1973) described Palaeo-
carpilius aquilinus based upon well preserved material
from the Eocene of Libya. The highly vaulted carapace,
triangular but quadrilobed nature of the front, wide dorsal
carapace exhibiting lower length to width ratio, and posi-
tion of maximum width placed farther anteriorly (Table 3)
suggest that it is better placed within a new genus, result-
ing in Bryocarpilius aquilinus. Bryocarpilius differs from
Laticarpilius new genus and Liopsalis in its highly vaulted
carapace and possession of numerous anterolateral spines
and from Laticarpilius in its quadrilobed front. The genus
is known from the Eocene of North Africa, including Libya
and Egypt.
Fig. 13. Bryocarpilius apsidorsalis n. g., n. sp., holotype, SMNS 61866. 1 – dorsal carapace; 2 – posterior view showing
pleon; 3 – anterior view showing orbits, front, and chelae; 4 – right lateral view showing extreme longitudinal vaulting of
carapace. Scale bars = 1 cm.
24 R.M. Feldmann et al.
Bryocarpilius apsidorsalis n. sp.
Fig. 13
Etymology: The trivial name is derived from the Latin
words apsis, meaning vault, and dorsalis, meaning of the
back, alluding to the strong convexity of the dorsal carapace.
Type: Holotype, SMNS 61866.
Diagnosis: Carapace ovate, length about 68% maximum
width, widest about 60% the distance posteriorly; very
strongly vaulted longitudinally; front quadrilobed, weakly
projected into a sinuous triangular shape, with two closely
spaced axial lobes and inner orbital lobes, about 36%
maximum width, fronto-orbital width about half maximum
carapace width; anterolateral margin with eight, evenly
spaced spines including outer orbital spine; metagastric
region very weakly dened, broadly inated.
Description: Carapace transversely ovoid; 37.0 mm long
and 54.6 mm wide, length about 68% maximum width,
widest at position of seventh anterolateral spine, about 60%
the distance posteriorly on carapace; vaulted transversely
and much more so longitudinally; smooth, lacking nodes or
spines on dorsal surface, nely pitted. Front quadrilobed,
front weakly projected into a sinuous triangular shape, axial
two lobes closely spaced and projected beyond remainder
of front, other two lobes serve as inner-orbital projections,
narrows distally, extends beyond orbits; frontal width 19.5
mm, about 36% maximum carapace width. Orbits circular,
entire, directed forward, fronto-orbital width about half
maximum carapace width. Anterolateral margin with eight
spines including outer orbital spine; spines small triangu-
lar, separated by arcuate sulci, similar in size and evenly
spaced, last spine extending onto carapace as ridge that also
denes origin of posterolateral margin. Posterolateral mar-
gin initially oriented at about 80º angle to axis, then arcing
posteriorly at about 60º angle to carapace. Posterior margin
nearly straight. Metagastric region very weakly dened,
broadly inated; urogastric and anterior-most cardiac re-
gion bounded laterally by postcervical or branchiocardiac
groove.
Merus of pereiopod 1 short; carpus triangular. Chelae
weakly heterochelous, right larger; mani stout, longer than
high, becoming higher distally, upper surfaces with spines;
ngers with black tips. Merus of pereiopod 2 much longer
than high, much more slender than chelipeds.
Sternites 4-7 visible; sternite 8 covered by pleon. Male
pleonite 1 narrow, wider than long; pleonite 2 longer than
1, wider than long; pleonites 3-5 fused, 3 wider than 4 or 5,
sutures between 3/4 and 4/5 barely visible; pleonite 6 about
as wide as long telson triangular about as wide as long.
Discussion: Bryocarpilius apsidorsalis n. sp. differs from
the type species, B. aquilinus, in possessing 8 spines rather
than 9 on the anterolateral margin, lacking a posterolateral
spine, and in lacking an arcuate metagastric ridge.
Genus Ocalina RathBun, 1929
Type species: Ocalina oridana RathBun, 1929, by
monotypy.
Included species: Ocalina delicata n. sp.; O. oridana
RathBun, 1929; O. haldixoni cOllinS & dOnOVan,
2006; O. straeleni (Remy in Remy & teSSieR, 1954), as
Palaeocarpilius; O. sublevis cOllinS & dOnOVan, 2005.
Diagnosis: Carapace wider than long, length 63-73%
maximum carapace width, maximum width position about
60% the distance posteriorly; surface ornamented with large
tubercles, most prominently on anterior and anterolateral
surfaces; front with two prominent lobes anking axis
and two smaller lobes near inner orbital corner, 35-40%
maximum carapace width; orbits rimmed, granular,
fronto-orbital width 50-55% maximum carapace width;
anterolateral margin lobulate or with at least nine large,
blunt projections; posterior margin with one or a few blunt
projections anteriorly; carapace regions poorly dened;
chelae with rows of large granules.
Discussion: As with other members of the family, species of
Ocalina have an ovoid, wider than long carapace with little
or no denition of regions. Margins are variable, but many
bear nodes or small spines on the anterolateral margin.
Ocalina is distinguished from other genera within the
family by having a beaded front and beaded, rimmed orbits.
Additionally, the surface of the frontal and anterolateral
margins bear coarse granular or tubercular ornamentation,
rather than being smooth as in other genera.
Ocalina delicata n. sp.
Fig. 14
Etymology: The trivial name is derived from the Latin
delicatus = dainty, and alludes to the delicate nature of the
ornamentation on the frontal and anterolateral regions of
the carapace.
Material: Holotype, SMF X/m 69a1.
Locality and horizon: The locality accompanying the
specimen indicates that it is an Eocene fossil crab from
Egypt. Based upon its lithology and comparison with the
preservational style exhibited by other specimens from
Egypt, we speculate that it was collected from Gebel
Mokattam, just northeast from Cairo, Egypt.
Diagnosis: Carapace with delicate granules on frontal
and anterolateral surfaces; anterolateral margin nely
spinose or nodose. Lateral cardiac nodes with pair of small
protuberances.
Description: Carapace ovoid, wider (44.0 mm) than
long (32.2 mm), length about 73% maximum carapace
width; moderately vaulted transversely, strongly vaulted
longitudinally particularly in anterior; regions not well
dened on cuticle exterior, more so on mold of interior of
carapace. Margin of carapace along frontal and anterolateral
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 25
surfaces markedly granular. Remainder of carapace
apparently smooth.
Front broad, 41% maximum width measured at about
midlength, downturned, projected slightly in advance of
orbits, bearing two rounded projections anking midline,
two smaller nodes near orbits, and smaller inner orbital
spines. Orbits small, circular, directed anterolaterally,
complete, weakly rimmed. Fronto-orbital width 56%
Fig. 14. Ocalina delicata n. sp., holotype, SMF X/m 69a1. 1 – dorsal carapace; 2 – anterior oblique view; 3 – left oblique
lateral view; 4 – right oblique lateral view; 5 – right lateral view showing strong longitudinal convexity; 6 – right oblique
frontal view. Scale bars = 1 cm.
26 R.M. Feldmann et al.
maximum width. Anterolateral margin convex, becoming
more convex posterior to point of maximum width; bearing
numerous small spines of which the last three are rounded
nodes and others are sharper. Anterolateral corner forms
granular, transverse ridge extending a short distance onto
carapace surface. Posterolateral margin concave anteriorly
and weakly convex posteriorly, smooth. Posterior margin
weakly convex, about 33% maximum width.
Epigastric regions subtle transverse swellings. Proto-
gastric, mesogastric, and hepatic regions undifferentiated.
Posterior termination of mesogastric region and axial part
of cervical groove dened on mold of interior by prominent
triangular gastric muscle scars and axial node. Meta/uro-
gastric regions undifferentiated, bounded laterally by later-
ally concave muscle scars. Cardiac region prominent, pen-
tagonal, bounded on anterolateral edges by muscle scars;
three nodes form triangle with apex directed posteriorly on
surface of cardiac region; lateral cardiac nodes each bear
two small protuberances. Intestinal region and branchial re-
gions undifferentiated.
Venter, pleon, and appendages not preserved.
Measurements: Measurements, in mm, taken on Ocalina
delicata and the other species within the genus are given
in Table 5.
Discussion: Ocalina delicata n. sp. conforms closely to the
denition of the genus (RathBun 1929) in regard to the fea-
tures preserved on the specimen from Egypt. Certainly, the
ornamentation of the carapace and the general absence of
regional development are diagnostic of the genus. However,
this species differs from the others in the genus sufciently
to warrant erection of a new species.
Ocalina delicata n. sp. differs from the type species,
O. oridana in being much less strongly ornamented. Of
all the species of the genus, O. oridana is by far the most
coarsely tubercular. RathBun (1929: 1) described the orbital
margin as being complete, or nearly so. That feature is not
evident in either O. haldixoni or O. sublevis; that region is
not seen in the illustration of O. straeleni nor is it exposed
in O. delicata. The type species is also more than twice the
size of any of the other species, based upon the available
material. Variation in size of the various species is interest-
ing in that O. straeleni is the smallest, being about half the
length of O. delicata, O. haldixoni, and O. sublevis which
are, in turn, half the length of the type species (Table 1).
Ocalina haldixoni has a prominent sulcate axis extend-
ing from the frontal margin to the anterior projection of
the mesogastric region which is weakly outlined and bears
moderately broad, blunt tubercles. These features are not
present in O. delicata.
Ocalina straeleni was originally assigned to Palaeocar-
pilius, but GuinOt (1968: footnote, p. 325) noted that the
species bore resemblance to Ocalina. That reassignment
was formalized by SchweitzeR (2003). The smooth cara-
pace surface and very small size of O. straeleni distinguish
the species from O. delicata. Finally, the rugose nature of
the carapace of Ocalina sublevis coupled with the promi-
nent rectilinear anterolateral tubercles readily distinguish
that species from O. delicata.
Ocalina delicata and O. straeleni are known from mid-
dle Eocene (Lutetian) occurrences in tropical and subtropi-
cal Africa whereas the remaining species are from middle
to late Eocene tropical and subtropical regions in the Ca-
ribbean and Florida, U.S.A. Thus, the Tethyan distribution
is well documented. cOllinS & dOnOVan (2005) suggested
that the genus arose in the eastern part of its range, which
was at that time the locality of O. straeleni, in the middle
Eocene and dispersed westward where it is known from
rocks as young as late Eocene. Further, with the possible
exception of Ocalina straeleni for which the lithology of
the surrounding rocks is not known to us, all the specimens
are preserved in limestones containing a diverse fauna and
abundant shelly fragments.
Superfamily Goneplacoidea macleay, 1838
Family Hexapodidae mieRS, 1886
Genus Goniocypoda wOOdwaRd, 1867
Type species: Goniocypoda edwardsi wOOdwaRd, 1867, by
monotypy.
Diagnosis: Carapace subrectangular, corners rounded,
especially anterolateral corner; front narrow, ared; fronto-
orbital width occupying entire frontal margin of carapace;
orbital margins rimmed, outer edge of orbit intersecting
anterolateral corner; reentrants at posterolateral corner
small; axial regions sometimes with small nodes.
Goniocypoda transsilvanica BittneR, 1893
Fig. 15
Table 5. Measurements and ratios taken on species of Ocalina. *denotes measurements taken from illustrations and adjusted
for published scales. **denotes measurements provided by original author. L = carapace length, W = Carapace width, FW
= frontal width, FOW = fronto-orbital width PW = posterior width.
Species L W L/W FW FW/W FOW FOW/W PW
O. delicata 32.2 44.0 0.73 18.0 0.41 24.7 0.56 15.9
O. oridana** 74 108 0.69 40.6 0.38 52 0.48 ---
O. haldixoni* 24.8 36.2 0.69 --- --- 20.8 0.57 ~13.3
O. straeleni* 14.1 21.7 0.65 9.1 0.42 13.0 0.60 ~7.4
O. sublevis* 27.6 42.5 0.65 --- --- 20.8 0.49 ~12.0
New Eocene Brachyura (Crustacea: Decapoda) from Egypt 27
Material examined: SMNS 67889 and cast KSU D 405.
Discussion: lőRenthey (1904, 1907a, b, 1909) referred
a specimen from the Eocene of Egypt to Goniocypoda
transsilvanica BittneR, 1893, a species originally named
from the Eocene of what is now a quarry near Cluj-
Napoca, Romania. The specimen illustrated by lőRenthey
is incomplete, lacking the front and much of the orbital
margins (Fig. 15). However, the Egyptian specimen clearly
has orbits that occupy the entire frontal margin; is much
wider than long; has a wide, sinuous posterior margin; and
is relatively unornamented. This ts the diagnosis for the
genus and is similar to the drawing of G. transsylvanica
BittneR (1893: pl. 2, g. 4). Until more complete material
from Egypt is collected, this referral seems acceptable;
however, the Egyptian specimen differs from the Romanian
specimen in not narrowing posteriorly.
Acknowledgements
Examination of type and comparative material in museum
collections in the USA and Europe was supported by
NSF grant EF-0531670 to Feldmann and SchweitzeR.
G. SchweiGeRt, Staatliches Museum für Naturkunde,
Stuttgart, Germany; M. munt and C. melliSh, The Natural
History Museum, London, UK; P. aRtal and S. calzada,
Museo Geológico del Seminario de Barcelona, Spain; A.
lORd and C. FRanz, Senckenberg Museum, Frankfurt am
Main, Germany; R. lemaitRe and K. Reed, Division of
Crustacea and J. thOmpSOn and M. FlORence, Paleobiology,
United States National Museum, Smithsonian Institution,
Washington, D.C., USA; all facilitated access and loans
from the collections at their respective institutions. A.
de anGeli, Vicenza, Italy, provided a photograph of
Tumidomaia priabonensis that we have used herein. S.
dOnOVan, Nationaal Natuurhistorisch Museum, Leiden,
The Netherlands, conducted a search for the holotype of
Lophoranina soembaensis. Useful reviews and editorial
assistance were provided by G. SchweiGeRt, and H.
KaRaSawa, Mizunami Fossil Museum, Mizunami, Japan.
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Manuscript received: July 29th, 2011.
Revised version accepted by the Stuttgart editor: August
26th, 2011.
Addresses of the authors:
ROdney m. Feldmann, OlGa Bennett, OVidiu d. FRanţeScu,
nicK ReSaR, aShley tRudeau, Department of Geology, Kent
State University, Kent, Ohio 44242, USA;
e-mail: rfeldman@kent.edu
caRRie e. SchweitzeR, Department of Geology, Kent State
University at Stark, 6000 Frank Ave. NW, North Canton,
Ohio 44720, USA;
e-mail: cschweit@kent.edu
