THE OLDEST BRACHYURA (DECAPODA: HOMOLODROMIOIDEA: GLAESSNEROPSOIDEA) KNOWN TO DATE (JURASSIC) Carrie E. Schweitzer and Rodney M.

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1 JOURNAL OF CRUSTACEAN BIOLOGY, 30(2): , 2010 THE OLDEST BRACHYURA (DECAPODA: HOMOLODROMIOIDEA: GLAESSNEROPSOIDEA) KNOWN TO DATE (JURASSIC) Carrie E. Schweitzer and Rodney M. Feldmann (CES, Department of Geology, Kent State University Stark Campus, 6000 Frank Ave. NW, North Canton, Ohio 44720; (RMF, Department of Geology, Kent State University, Kent, Ohio 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: / 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. SYSTEMATICS Infraorder Brachyura Linnaeus, 1758 Section Dromiacea De Haan, 1833 Superfamily Homolodromioidea Alcock, 1900 Included Families. Homolodromiidae Alcock, 1900; Bucculentidae Schweitzer and Feldmann, 2009a; Goniodromitidae Beurlen, 1932; Prosopidae Von Meyer, 1860; Tanidromitidae Schweitzer and Feldmann, 2008 [imprint 2007]. 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 251

2 252 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 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

3 SCHWEITZER AND FELDMANN: THE OLDEST CRABS 253 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 ) and is noted on the labels as having been purchased as part of the Tesson Collection in The specimen is from Ranville, Calvados, France, one of the original localities of Eudes- Deslongchamps. 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 as a paralectotype. We suggest that BMNH In be considered as the lectotype for Homolus auduini, given that the whereabouts of the Langrune specimens of Eudes- Deslongchamps 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 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 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 Palaeinachus longipes Woodward, 1866: 493, pl. 24, fig. 1; A. Milne- Edwards 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 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

4 254 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010 Fig. 2. Homolus auduini Eudes-Deslongchamps, A, digital image of drawing of Bouvier (1896) of Woodward specimen showing lateral rostral spines, pereiopods, and abdomen; B-D, BMNH In , 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.

5 SCHWEITZER AND FELDMANN: THE OLDEST CRABS 255 Table 1. Earliest confirmed Brachyura, ages, and geographic occurrences (data in references cited above). Family Genus Age Geographic occurrence Homolodromiidae Homolus Bathonian UK, France Tanidromitidae Gabriella Bajocian-Bathonian Tanzania (Förster, 1985) Tanidromitidae Tanidromites Latest Bajocian UK Longodromitidae Planoprosopon Bajocian France Longodromitidae Abyssophthalmus Early Bajocian France, Germany (Schweigert, 2006) Homolodromioidea Eoprosopon Pliensbachian 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 , paralectotype of Morris (1980) and suggested as the lectotype herein; cast of Woodward specimen deposited in BMNH; OUM J Occurrence. BMNH In , 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 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. 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