JOURNAL OF CRUSTACEAN BIOLOGY, 30(2): 251-256, 2010
THE OLDEST BRACHYURA (DECAPODA: HOMOLODROMIOIDEA: GLAESSNEROPSOIDEA)
KNOWN TO DATE (JURASSIC)
Carrie E. Schweitzer and Rodney M. Feldmann
(CES, [email protected]) Department of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW,
North Canton, Ohio 44720;
(RMF, [email protected]) Department of Geology, Kent State University, Kent, Ohio 44242
ABSTRACT
The oldest confirmed brachyurans, referable to the Homolodromioidea and Glaessneropsoidea, are Early to Middle Jurassic in age.
Geographically, all of the occurrences are in Europe, with one exception, in Tanzania.
KEY WORDS: Brachyura, Decapoda, Homolodromioidea, Jurassic, origins
DOI: 10.1651/09-3231.1
tidae Beurlen, 1932; Prosopidae Von Meyer, 1860;
Tanidromitidae Schweitzer and Feldmann, 2008 [imprint
2007].
INTRODUCTION
The search for the oldest brachyuran has had many false starts.
As better and more complete diagnoses for the various
anomuran and brachyuran groups have become available, and
as more fossils and more complete fossils are known, it has
become possible to determine which organisms belong within
Brachyura and which are most likely anomurans or some other
type of decapod. Recent workers have removed several
candidates for the oldest brachyuran from the group (Schweitzer
and Feldmann, 2005; Feldmann and Schweitzer, 2010). Herein
we report on the oldest confirmed members of Brachyura, those
occurrences that fit the definition of Brachyura without doubt.
Krobicki and Zatoń (2008) provided an overview of the
Jurassic roots of crabs, but that work did not take into
account recent advances in the systematics and phylogeny
of these animals. An important implication for our findings
is that it permits the constraint of molecular studies of
Brachyura and other decapod taxa, with an earliest possible
divergence time based upon actual evidence for these
groups. These findings also permit paleontologists to focus
on an age horizon and therefore to target localities in the
search for ancestral Decapoda within the anomuran and
brachyuran lineages; these two clades are shown to be
sisters in most recent analyses (Ahyong and O’Meally,
2004; Ahyong et al., 2007). Recognition of an Early to
Middle Jurassic origin for Brachyura can allow workers to
begin investigating the dynamics of the origin and radiation
of this ecologically important group of animals.
Institutional abbreviations: BMNH, The Natural History
Museum, London, UK; BSP, Bayerische Staatsammlung
für Paläontologie, München, Germany; OUM, Geological
Collections, Oxford University Museum.
Homolodromioidea incertae sedis
Eoprosopon Förster, 1986
Type and Sole Species.—Eoprosopon klugi Förster, 1986,
by monotypy.
Eoprosopon klugi Förster, 1986
Fig. 1
Material Examined.—Cast of BSP 1986 I 19, holotype.
Discussion.—Förster (1986) described Eoprosopon from
the Pliensbachian of Germany. He referred it to Prosopidae,
and Guinot (1995) later removed it to Homolodromiidae
based upon its possession of a deep cervical groove, long
dactyls of pereiopods 2 and 3, reduced pereiopods 4 and 5,
and a flattened pleon with epimeres. All of these features
are present, but the nature of the pleon suggests that
placement into Homolodromiidae is not warranted. Examination of the pleonites indicates the presence of discrete,
obvious swellings in the area of the epimeres (Fig. 1),
which are not present on any extant homolodromiid. Each
individual pleonite 5-7 is rectangular and long, they fit
closely together, and they maintain their length laterally.
Pleonite 7 has clear articulating rings. The telson is slightly
longer than wide. In extant homolodromiids, both male and
female, the telson is very long and longer than wide, e.g.,
illustrations in Guinot (1995). Pleonites 5-7 are short in
both males and females, and they narrow laterally into
triangular tips. Weak articulating rings may be present if
present at all.
Thus, the pleon of Eoprosopon is unlike that of
Homolodromiidae. However, the features of Eoprosopon
seem to suggest clearly that it is a brachyuran. It possesses
one pair of chelae; the walking legs are long and achelate;
the last pair of pereiopods and possibly the fourth pair also
SYSTEMATICS
Infraorder Brachyura Linnaeus, 1758
Section Dromiacea De Haan, 1833
Superfamily Homolodromioidea Alcock, 1900
Included Families.—Homolodromiidae Alcock, 1900; Bucculentidae Schweitzer and Feldmann, 2009a; Goniodromi251
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010
Fig. 1. Eoprosopon klugi Förster, 1986, cast of BSP 1986 I 19, holotype. Scale bar 5 1 cm.
are reduced; the dorsal carapace has well-developed
regions and cervical and branchiocardiac grooves; and the
carapace is dorsoventrally compressed. The pleon extends
posteriorly from the carapace, and does have epimeres,
both of which are atypical of the brachyurans. However,
these features are seen in the extant homolodromiids. Thus,
Eoprosopon is the earliest known specimen that can be
referred to Brachyura with reasonable confidence, probably
best referred to the Homolodromioidea incertae sedis.
Examination of the type specimen, perhaps with some
additional preparation, or recovery of more material might
permit referral of the specimen to a family with confidence.
To date, the appendages and pleon of the Jurassic
homolodromioids are unknown. Garassino et al. (2005)
had referred a specimen with preserved appendages to
Pithonoton sp.; however, Feldmann et al. (2006) have
already suggested that it is probably a porcellanid.
Unfortunately, Eoprosopon lacks well-preserved front and
lateral views that could permit determination of the
presence and size of augenrests or orbital structures
necessary to place it within a family.
Homolodromiidae Alcock, 1900
Homolus Eudes-Deslongschamps, 1835
Protocarcinus Woodward and Salter, 1865, fig. 15.
Palaeinachus Woodward, 1866, p. 493, pl. 24, fig. 1.
Type and Sole Species.—Homolus auduini Eudes-Deslongchamps, 1835, by monotypy.
Diagnosis.—Carapace longer than wide, widest about 70%
the distance posteriorly in branchial region; rostrum with
long lateral rostral spines and downturned central rostral
spine; augenrests large, directed forward, protected by three
spines; dorsal carapace ornamented by long spines; cervical
groove deep, nearly straight; mesogastric region wellornamented by spines and muscle scars.
Discussion.—Eudes-Deslongschamps (1835) named the
genus and its type and sole species in 1835. In 1865,
Woodward and Salter referred a species to Protocarcinus,
P. longipes, and then later Woodward (1866) placed that
species within the new genus Palaeinachus. He did not use
the name Protocarcinus because: 1) the specimen in his
opinion bore no relation to Carcinus, and 2) ‘‘the prefix
‘Proto’ is objectionable in Palaeontology’’ (Woodward, 1866,
p. 493). Examination of specimens in the Natural History
Museum, London, suggests that Homolus auduini and
Palaeinachus longipes are synonymous as suggested by
Glaessner (1929) and Guinot (1995). The two species (H.
auduini and P. longipes) appear to be conspecific based upon
possession of three large nodes on the anterior half of
carapace (one on anterior extension of mesogastric, and one
each on protogastric), a pair of nodes on the posterior end of
the mesogastric region, a straight path of the cervical groove,
and possession of pseudorostral spines. All three generic
names are available according to Nomenclator Zoologicus.
However, Homolus is the earlier name, was fully described
and illustrated, and was designated with an associated
species; therefore, we use it here. Wehner (1988) referred
the species to Foersteria, but that name has been replaced by
Gabriella Collins et al., 2005, and the species is not referable
to that genus (Schweitzer and Feldmann, 2009b).
Homolus is referable to Homolodromiidae based upon
possession of large, forward-directed augenrests protected
SCHWEITZER AND FELDMANN: THE OLDEST CRABS
by spines; lateral rostral spines that are very long and a
downturned central rostral spine; a forward-directed outerorbital spine; a well-defined branchiocardiac and a
moderately defined postcervical groove; an inflated subhepatic region; a well-defined, straight cervical groove;
lateral margins that are subparallel and diverge posteriorly;
relatively short first pereiopods; a dorsal carapace that has
high flanks that become markedly narrower posteriorly,
possibly indicating a poorly calcified posterior portion; and
an abdomen that is partially visible in dorsal view. In fact,
Homolus is remarkably similar to extant homolodromiids.
Bouvier (1896) had already noted this remarkable similarity. No other decapod family exhibits this combination of
characters.
Homolus differs from extant homolodromiids in possessing large spine-like swellings on the dorsal carapace. In
addition, the augenrest is larger and the opening for the
eyestalk is smaller in Homolus than in extant homolodromiids. Given the approximately 165 million year difference
in age, these differences seem relatively minor. Homolus is
the oldest confirmed member of Homolodromiidae.
The history of the specimens referable to this species is
complicated. Eudes-Deslongchamps (1835) referred to two
specimens in his description. One specimen, which was
designated as a paralectotype (Morris, 1980), is deposited
in the Natural History Museum (BMNH In. 57979) and is
noted on the labels as having been purchased as part of the
Tesson Collection in 1857. The specimen is from Ranville,
Calvados, France, one of the original localities of EudesDeslongchamps. Morris or possibly another curator apparently believed it to be one of the original specimens of
Eudes-Deslongchamps as evidenced by notes on one of the
labels associated with this specimen, indicating it as a
syntype and as figured by Eudes-Deslongchamps (1835).
The other specimen described by Eudes-Deslongchamps
(1835) was collected at Langrune near Caen; the disposition of that specimen is not known. Apparently, Morris
considered the Langrune specimen to be the lectotype,
based upon his designation of BMNH In. 57979 as a
paralectotype. We suggest that BMNH In. 57979 be
considered as the lectotype for Homolus auduini, given
that the whereabouts of the Langrune specimens of EudesDeslongchamps are unknown.
The Woodward specimen apparently has been lost.
Withers (1932) reported that the specimen was originally
held by the Natural History Museum in London but was
returned to Prof. T. Bell in the mid-1800s. Only a plaster
cast remains there now as well as in the Museum National
d’Histoire Naturelle in Paris. The cast shows that the pleon
of Homolus auduini extended posteriorly from the posterior
margin of the carapace and shows details of the rostrum and
pereiopods.
Van Straelen (1924 [imprint 1925]) erected the genus
Avihomola to embrace several species, the earliest of which
was Bajocian. He provided a diagnosis for the genus, but
did not name a type species. The many species which he
referred to Avihomola were originally referred to either
Prosopon von Meyer, 1835, or Pithonoton von Meyer,
1842, but the variation among them is enormous. Glaessner
(1929) did not recognize the genus, although under the
253
International Code of Zoological Nomenclature (1999:
Articles 11 and 12), the name does appear to be available.
All of the species referred to it were originally referred to
other genera, and they have usually been treated as
belonging to the genus to which they were originally
referred in generic revisions, especially because Van
Straelen did not designate a type species. Thus, most of
the species referred to Avihomola have since been referred
to other genera. The remaining species that Van Straelen
referred to Avihomola are known from poor, incomplete
material. At this time, it appears, therefore, that Avihomola
is a nomen nudum.
Beurlen (1932) reported Early Jurassic specimens of
Goniodromites Reuss, 1858 [imprint 1857]. Schweitzer and
Feldmann (2008 [imprint 2007]) have already suggested
that, based upon the drawings of these specimens, they must
be eubrachyuran taxa and are most likely not Jurassic in age.
Unfortunately, the specimens may have been stored in what is
now Kaliningrad, Russia, which was heavily bombed during
World War II, making it unlikely that the specimens survive
(G. Schweigert, personal communication).
Homolus auduini Eudes-Deslongchamps, 1835
Fig. 2
Homolus auduini Eudes-Deslongchamps, 1835: 39, pl. 1, figs. 4-6.
Protocarcinus longipes Woodward and Salter, 1865: pl. 2, fig. 15; Bouvier,
1896, 80, fig. 36; Glaessner, 1929, 348; Beurlen and Glaessner, 1930,
66, fig. 15; Withers, 1932, 319, pl. 10, fig. 2-4.
Palaeinachus longipes Woodward, 1866: 493, pl. 24, fig. 1; A. MilneEdwards and Bouvier, 1902, pl. 10, fig. 4.
Pithonoton auduini (Deslongchamps): Hée, 1924, 148, pl. 6, fig. 4.
Avihomola auduini (Eudes-Deslongchamps): Van Straelen, 1924 [imprint,
1925], 339, pl. 10, fig. 9.
Prosopon auduini (Deslongchamps): Glaessner, 1933, 180; Morris, 1980, 15.
Foersteria auduini (Deslongchamps): Wehner, 1988, 30; Guinot, 1995,
265.
Gabriella audini [sic] (Deslongchamps): Collins et al., 2005, p. 125.
Homolodromiidae incertae sedis: Schweitzer and Feldmann, 2009a, table 2.
Protocarcinus auduini (Deslongchamps): Schweitzer and Feldmann,
2009b, 8.
Description (Translated from Eudes-Deslongchamps,
1835: 39).— I have found only once an almost entire
carapace of this crustacean; it has an elongate quadrilateral
form; it is strongly inflated above and rounded at its
anterior end, which is terminated by a bilobed rostrum
which is a little damaged on the specimen; the length is of
seven lines, and the width is of five [what is meant by
‘‘lines’’ is unknown]. A transverse groove is strongly
expressed, concave forward, dividing it into two approximately equal regions. The anterior [region], rounded in
front and on the sides, is ornamented with nine small
conical tubercles strongly projecting, disposed in a very
elegant manner. The rostrum is large, of two lobes separated
by a large longitudinal groove. I cannot distinguish whether
or not the front was ornamented with spines; I do not have
any expressed in the drawings; but it is possible that they
exist, and in another specimen they might be seen.
The posterior region is deprived of tubercles; it offers
many well-pronounced projections, separated by grooves;
one of these projections, situated immediately behind the
transverse groove, is only slightly stretched lengthwise, but
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010
Fig. 2. Homolus auduini Eudes-Deslongchamps, 1835. A, digital image of drawing of Bouvier (1896) of Woodward specimen showing lateral rostral
spines, pereiopods, and abdomen; B-D, BMNH In. 57979, paralectotype of Morris (1980) and suggested as the lectotype, dorsal carapace (B), anterior view
(C), and left lateral view (D); E, OUM J.71902, dorsal carapace. Scale bars 5 1 cm. B-C courtesy P. Crabbe, Natural History Museum, London, UK.
stretched strongly laterally. Behind this and the axis, one
sees another [region], very narrow, in the form of a heart,
with the point directed posteriorly. The shell of this little
carapace is very thin and very white; seen under a strong
lens, the entire surface appears covered in projected points,
regularly placed.
The elongate form of this carapace, and especially the
transverse groove which divides it in two, brings to mind
somewhat the appearance of the carapace of the macruran
crustaceans; it has, nevertheless, the most similarities with
the form of the crustaceans of the genus Homola which is
placed with the brachyurans and with which I place it. I
followed the opinion of Mr. Audouin and Mr. Milne
Edwards, who, looking with me last year at this little fossil
carapace, regarded it as being like the homolid genera. I
have found it at Langrune (three miles north of Caen) in a
bank resting on coral-bearing limestone, and it is like the
pisolite of England.
Mr. Tesson found a little later, in the Ranville quarry,
another carapace of the Audoin homolid; it is about half
again the size of the first described in this article. The
tubercles are disposed in the same manner; it offers, what is
more, two small [tubercles] close together, situated behind
and three [tubercles] close to the transverse groove; these
latter could have been easily destroyed in my small
specimen.
Description of Lectotype.—Carapace rectangular, longer
than wide, width about 80% length, maximum width about
70% the distance posteriorly in branchial regions; moderately vaulted transversely and longitudinally; flanks steep,
high.
Rostrum with two long lateral rostral spines and
downturned central rostral spine; rostral width about 43%
maximum carapace width measured at base of lateral
rostral spines. Augenrest concave, smooth, directed forward, with three spines arranged around outer margin:
upper, outer, and lower augenrest spines; fronto-orbital
width including augenrest about 85% maximum carapace
width. Lateral margins straight, diverging weakly posteriorly; weak swelling anterior to intersection of cervical
groove, margin incised at cervical groove; weak swelling
posterior to cervical groove; incised at intersection of
branchiocardiac groove; weakly convex posterior to
branchiocardiac groove. Posterior margin unknown.
Mesogastric region with long anterior process, widening
posteriorly; spine at about half-length of anterior process;
pair of spines at anterior end of widened area followed
posteriorly by pair of muscle scars. Metagastric region
wide, rectangular, with muscle scars and pair of spines on
top of muscle scars; urogastric region flattened, short;
cardiac region triangular, with two spines anteriorly,
posterior portion unknown, intestinal region unknown.
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SCHWEITZER AND FELDMANN: THE OLDEST CRABS
Table 1. Earliest confirmed Brachyura, ages, and geographic occurrences (data in references cited above).
Family
Genus
Age
Geographic occurrence
Homolodromiidae
Tanidromitidae
Tanidromitidae
Longodromitidae
Longodromitidae
Homolodromioidea
Homolus
Gabriella
Tanidromites
Planoprosopon
Abyssophthalmus
Eoprosopon
Bathonian
Bajocian-Bathonian
Latest Bajocian
Bajocian
Early Bajocian
Pliensbachian
UK, France
Tanzania (Förster, 1985)
UK
France
France, Germany (Schweigert, 2006)
Germany
Protogastric and hepatic regions confluent, spine on
anterior margin nearly on margin of augenrest.
Cervical groove deep, extending nearly straight across
carapace. Postcervical groove composed of two discontinuous, weak grooves bounding posterior margin of metagastric region, with deep, arcuate pit at each lateral end.
Branchiocardiac groove extending weakly convex forward
in oblique axial path to cardiac region. Epibranchial region
rectangular laterally, drawing into fingerlike projection
directed at cardiac region axially. Remainder of branchial
region undifferentiated. Subhepatic region inflated,
bounded posteriorly by ventral extension of cervical
groove; flank becoming less high posteriorly.
Remainder of carapace and appendages unknown.
Material Examined.—BMNH In. 57979, paralectotype of
Morris (1980) and suggested as the lectotype herein; cast of
Woodward specimen deposited in BMNH; OUM J.71902.
Occurrence.—BMNH In. 57979, suggested lectotype:
Bathonian, Ranville, Calvados, France, couches à polypiers, Tesson collection, purchased July, 1857, original of
Eudes-Deslongchamps; Woodward specimen: Bathonian
Forest Marble, Great Oolite, Malmesbury, Wiltshire; OUM
J.71902: Lower Cornbrash, Ornithella obovata zone, Upper
Bathonian, Stratton Audley, Oxfordshire, UK.
DISCUSSION
The oldest confirmed Brachyura are Early and Middle
Jurassic in age, and arrayed in at least three families and
two superfamilies (Table 1). Both Planoprosopon Schweitzer et al., 2007, and Coelopus Étallon, 1861, of Longodromitidae Schweitzer and Feldmann, 2009a, are reported
from the Middle Jurassic of France, with ages of Bajocian
and Bathonian respectively (Schweitzer and Feldmann,
2009b; 2010). Gabriella Collins et al., 2005 and Tanidromites Schweitzer and Feldmann, 2008 [imprint 2007] of
Tanidromitidae are known from the Bajocian/Bathonian
and the latest Bajocian respectively (Schweitzer and
Feldmann, 2009b). Thus, based upon available evidence,
the origin of Brachyura appears to lie in the Early Jurassic
within Homolodromioidea (Eoprosopon), with radiation
into two superfamilies and multiple families by the Middle
Jurassic. In the Late Jurassic, an evolutionary explosion
within Brachyura occurred, with dozens of species and
genera known, primarily from Europe (Feldmann et al.,
2006; Feldmann and Schweitzer, 2009; Schweitzer et al.,
2007; Schweitzer and Feldmann, 2008 [imprint 2007],
2009a-d).
ACKNOWLEDGEMENTS
Museum work in Europe was funded by NSF grant EF 0531670 to
Feldmann and Schweitzer. M. Nose provided access to collections at the
Bayerische Staatsammlung für Paläontologie, München, Germany; M.
Munt and C. Mellish, arranged our visit and photography at The Natural
History Museum, London, UK; P. Crabbe provided images of some of the
specimens illustrated herein from The Natural History Museum, London;
and S. De Grave arranged the examination and loan of material from the
Geological Collections, Oxford University Museum, UK. G. Schweigert,
Staatliches Museum für Naturkunde, Stuttgart, Germany, and H.
Karasawa, Mizunami Fossil Museum, Japan, provided constructive
reviews of the manuscript. Our thanks go to all of these individuals.
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RECEIVED: 30 September 2009.
ACCEPTED: 13 October 2009.