The Oldest Brachyura (Decapoda: Homolodromioidea: Glaessneropsoidea) Known to Date (Jurassic)

Home , Decapoda, Dromioidea, Glaessneropsoidea, Goniodromites, Homolodromiidae

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 Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021 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

INTRODUCTION tidae Beurlen, 1932; Prosopidae Von Meyer, 1860; The search for the oldest brachyuran has had many false starts. Tanidromitidae Schweitzer and Feldmann, 2008 [imprint As better and more complete diagnoses for the various 2007]. anomuran and brachyuran groups have become available, and as more fossils and more complete fossils are known, it has Homolodromioidea incertae sedis become possible to determine which organisms belong within Eoprosopon Fo¨rster, 1986 Brachyura and which are most likely anomurans or some other type of decapod. Recent workers have removed several Type and Sole Species.—Eoprosopon klugi Fo¨rster, 1986, candidates for the oldest brachyuran from the group (Schweitzer by monotypy. 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. Eoprosopon klugi Fo¨rster, 1986 Krobicki and Zaton´ (2008) provided an overview of the Fig. 1 Jurassic roots of crabs, but that work did not take into account recent advances in the systematics and phylogeny Material Examined.—Cast of BSP 1986 I 19, holotype. of these animals. An important implication for our findings Discussion.—Fo¨rster (1986) described Eoprosopon from is that it permits the constraint of molecular studies of the Pliensbachian of Germany. He referred it to Prosopidae, Brachyura and other decapod taxa, with an earliest possible and Guinot (1995) later removed it to Homolodromiidae divergence time based upon actual evidence for these based upon its possession of a deep cervical groove, long groups. These findings also permit paleontologists to focus dactyls of pereiopods 2 and 3, reduced pereiopods 4 and 5, on an age horizon and therefore to target localities in the and a flattened pleon with epimeres. All of these features search for ancestral Decapoda within the anomuran and are present, but the nature of the pleon suggests that brachyuran lineages; these two clades are shown to be placement into Homolodromiidae is not warranted. Exam- sisters in most recent analyses (Ahyong and O’Meally, ination of the pleonites indicates the presence of discrete, 2004; Ahyong et al., 2007). Recognition of an Early to obvious swellings in the area of the epimeres (Fig. 1), Middle Jurassic origin for Brachyura can allow workers to which are not present on any extant homolodromiid. Each begin investigating the dynamics of the origin and radiation of this ecologically important group of animals. individual pleonite 5-7 is rectangular and long, they fit Institutional abbreviations: BMNH, The Natural History closely together, and they maintain their length laterally. Museum, London, UK; BSP, Bayerische Staatsammlung Pleonite 7 has clear articulating rings. The telson is slightly fu¨r Pala¨ontologie, Mu¨nchen, Germany; OUM, Geological longer than wide. In extant homolodromiids, both male and Collections, Oxford University Museum. 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 SYSTEMATICS triangular tips. Weak articulating rings may be present if Infraorder Brachyura Linnaeus, 1758 present at all. Section Dromiacea De Haan, 1833 Thus, the pleon of Eoprosopon is unlike that of Superfamily Homolodromioidea Alcock, 1900 Homolodromiidae. However, the features of Eoprosopon seem to suggest clearly that it is a brachyuran. It possesses Included Families.—Homolodromiidae Alcock, 1900; Buc- one pair of chelae; the walking legs are long and achelate; culentidae Schweitzer and Feldmann, 2009a; Goniodromi- the last pair of pereiopods and possibly the fourth pair also

251 252 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010 Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021

Fig. 1. Eoprosopon klugi Fo¨rster, 1986, cast of BSP 1986 I 19, holotype. Scale bar 5 1 cm. are reduced; the dorsal carapace has well-developed long lateral rostral spines and downturned central rostral regions and cervical and branchiocardiac grooves; and the spine; augenrests large, directed forward, protected by three carapace is dorsoventrally compressed. The pleon extends spines; dorsal carapace ornamented by long spines; cervical posteriorly from the carapace, and does have epimeres, groove deep, nearly straight; mesogastric region well- both of which are atypical of the brachyurans. However, ornamented by spines and muscle scars. these features are seen in the extant homolodromiids. Thus, Eoprosopon is the earliest known specimen that can be Discussion.—Eudes-Deslongschamps (1835) named the referred to Brachyura with reasonable confidence, probably genus and its type and sole species in 1835. In 1865, best referred to the Homolodromioidea incertae sedis. Woodward and Salter referred a species to Protocarcinus, Examination of the type specimen, perhaps with some P. longipes, and then later Woodward (1866) placed that additional preparation, or recovery of more material might species within the new genus Palaeinachus. He did not use permit referral of the specimen to a family with confidence. the name Protocarcinus because: 1) the specimen in his To date, the appendages and pleon of the Jurassic opinion bore no relation to Carcinus,and2)‘‘the prefix homolodromioids are unknown. Garassino et al. (2005) ‘Proto’ is objectionable in Palaeontology’’ (Woodward, 1866, had referred a specimen with preserved appendages to p. 493). Examination of specimens in the Natural History Pithonoton sp.; however, Feldmann et al. (2006) have Museum, London, suggests that Homolus auduini and already suggested that it is probably a porcellanid. Palaeinachus longipes are synonymous as suggested by Unfortunately, Eoprosopon lacks well-preserved front and Glaessner (1929) and Guinot (1995). The two species (H. lateral views that could permit determination of the auduini and P. longipes) appear to be conspecific based upon possession of three large nodes on the anterior half of presence and size of augenrests or orbital structures carapace (one on anterior extension of mesogastric, and one necessary to place it within a family. each on protogastric), a pair of nodes on the posterior end of the mesogastric region, a straight path of the cervical groove, Homolodromiidae Alcock, 1900 and possession of pseudorostral spines. All three generic Homolus Eudes-Deslongschamps, 1835 names are available according to Nomenclator Zoologicus. However, Homolus is the earlier name, was fully described Protocarcinus Woodward and Salter, 1865, fig. 15. and illustrated, and was designated with an associated Palaeinachus Woodward, 1866, p. 493, pl. 24, fig. 1. species; therefore, we use it here. Wehner (1988) referred Type and Sole Species.—Homolus auduini Eudes-Deslong- the species to Foersteria, but that name has been replaced by champs, 1835, by monotypy. Gabriella Collins et al., 2005, and the species is not referable to that genus (Schweitzer and Feldmann, 2009b). Diagnosis.—Carapace longer than wide, widest about 70% Homolus is referable to Homolodromiidae based upon the distance posteriorly in branchial region; rostrum with possession of large, forward-directed augenrests protected SCHWEITZER AND FELDMANN: THE OLDEST CRABS 253 by spines; lateral rostral spines that are very long and a International Code of Zoological Nomenclature (1999: downturned central rostral spine; a forward-directed outer- Articles 11 and 12), the name does appear to be available. orbital spine; a well-defined branchiocardiac and a All of the species referred to it were originally referred to moderately defined postcervical groove; an inflated sub- other genera, and they have usually been treated as hepatic region; a well-defined, straight cervical groove; belonging to the genus to which they were originally lateral margins that are subparallel and diverge posteriorly; referred in generic revisions, especially because Van relatively short first pereiopods; a dorsal carapace that has Straelen did not designate a type species. Thus, most of high flanks that become markedly narrower posteriorly, the species referred to Avihomola have since been referred possibly indicating a poorly calcified posterior portion; and to other genera. The remaining species that Van Straelen an abdomen that is partially visible in dorsal view. In fact, referred to Avihomola are known from poor, incomplete Homolus is remarkably similar to extant homolodromiids. material. At this time, it appears, therefore, that Avihomola Bouvier (1896) had already noted this remarkable similar- is a nomen nudum. ity. No other decapod family exhibits this combination of Beurlen (1932) reported Early Jurassic specimens of characters. Goniodromites Reuss, 1858 [imprint 1857]. Schweitzer and Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021 Homolus differs from extant homolodromiids in posses- Feldmann (2008 [imprint 2007]) have already suggested sing large spine-like swellings on the dorsal carapace. In that, based upon the drawings of these specimens, they must addition, the augenrest is larger and the opening for the be eubrachyuran taxa and are most likely not Jurassic in age. eyestalk is smaller in Homolus than in extant homolodro- Unfortunately, the specimens may have been stored in what is miids. Given the approximately 165 million year difference now Kaliningrad, Russia, which was heavily bombed during in age, these differences seem relatively minor. Homolus is World War II, making it unlikely that the specimens survive the oldest confirmed member of Homolodromiidae. (G. Schweigert, personal communication). The history of the specimens referable to this species is complicated. Eudes-Deslongchamps (1835) referred to two Homolus auduini Eudes-Deslongchamps, 1835 specimens in his description. One specimen, which was designated as a paralectotype (Morris, 1980), is deposited Fig. 2 in the Natural History Museum (BMNH In. 57979) and is Homolus auduini Eudes-Deslongchamps, 1835: 39, pl. 1, figs. 4-6. noted on the labels as having been purchased as part of the Protocarcinus longipes Woodward and Salter, 1865: pl. 2, fig. 15; Bouvier, Tesson Collection in 1857. The specimen is from Ranville, 1896, 80, fig. 36; Glaessner, 1929, 348; Beurlen and Glaessner, 1930, Calvados, France, one of the original localities of Eudes- 66, fig. 15; Withers, 1932, 319, pl. 10, fig. 2-4. Palaeinachus longipes Woodward, 1866: 493, pl. 24, fig. 1; A. Milne- Deslongchamps. Morris or possibly another curator appar- Edwards and Bouvier, 1902, pl. 10, fig. 4. ently believed it to be one of the original specimens of Pithonoton auduini (Deslongchamps): He´e, 1924, 148, pl. 6, fig. 4. Eudes-Deslongchamps as evidenced by notes on one of the Avihomola auduini (Eudes-Deslongchamps): Van Straelen, 1924 [imprint, labels associated with this specimen, indicating it as a 1925], 339, pl. 10, fig. 9. syntype and as figured by Eudes-Deslongchamps (1835). Prosopon auduini (Deslongchamps): Glaessner, 1933, 180; Morris, 1980, 15. Foersteria auduini (Deslongchamps): Wehner, 1988, 30; Guinot, 1995, The other specimen described by Eudes-Deslongchamps 265. (1835) was collected at Langrune near Caen; the disposi- Gabriella audini [sic] (Deslongchamps): Collins et al., 2005, p. 125. tion of that specimen is not known. Apparently, Morris Homolodromiidae incertae sedis: Schweitzer and Feldmann, 2009a, table 2. considered the Langrune specimen to be the lectotype, Protocarcinus auduini (Deslongchamps): Schweitzer and Feldmann, based upon his designation of BMNH In. 57979 as a 2009b, 8. paralectotype. We suggest that BMNH In. 57979 be Description (Translated from Eudes-Deslongchamps, considered as the lectotype for Homolus auduini, given 1835: 39).— I have found only once an almost entire that the whereabouts of the Langrune specimens of Eudes- carapace of this crustacean; it has an elongate quadrilateral Deslongchamps are unknown. form; it is strongly inflated above and rounded at its The Woodward specimen apparently has been lost. anterior end, which is terminated by a bilobed rostrum Withers (1932) reported that the specimen was originally which is a little damaged on the specimen; the length is of held by the Natural History Museum in London but was seven lines, and the width is of five [what is meant by returned to Prof. T. Bell in the mid-1800s. Only a plaster ‘‘lines’’ is unknown]. A transverse groove is strongly cast remains there now as well as in the Museum National expressed, concave forward, dividing it into two approxi- d’Histoire Naturelle in Paris. The cast shows that the pleon mately equal regions. The anterior [region], rounded in of Homolus auduini extended posteriorly from the posterior front and on the sides, is ornamented with nine small margin of the carapace and shows details of the rostrum and conical tubercles strongly projecting, disposed in a very pereiopods. elegant manner. The rostrum is large, of two lobes separated Van Straelen (1924 [imprint 1925]) erected the genus by a large longitudinal groove. I cannot distinguish whether Avihomola to embrace several species, the earliest of which or not the front was ornamented with spines; I do not have was Bajocian. He provided a diagnosis for the genus, but any expressed in the drawings; but it is possible that they did not name a type species. The many species which he exist, and in another specimen they might be seen. referred to Avihomola were originally referred to either The posterior region is deprived of tubercles; it offers Prosopon von Meyer, 1835, or Pithonoton von Meyer, many well-pronounced projections, separated by grooves; 1842, but the variation among them is enormous. Glaessner one of these projections, situated immediately behind the (1929) did not recognize the genus, although under the transverse groove, is only slightly stretched lengthwise, but 254 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010 Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021

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 Description of Lectotype.—Carapace rectangular, longer sees another [region], very narrow, in the form of a heart, than wide, width about 80% length, maximum width about with the point directed posteriorly. The shell of this little 70% the distance posteriorly in branchial regions; moder- carapace is very thin and very white; seen under a strong ately vaulted transversely and longitudinally; flanks steep, lens, the entire surface appears covered in projected points, high. regularly placed. Rostrum with two long lateral rostral spines and The elongate form of this carapace, and especially the downturned central rostral spine; rostral width about 43% transverse groove which divides it in two, brings to mind maximum carapace width measured at base of lateral somewhat the appearance of the carapace of the macruran rostral spines. Augenrest concave, smooth, directed for- crustaceans; it has, nevertheless, the most similarities with ward, with three spines arranged around outer margin: the form of the crustaceans of the genus Homola which is upper, outer, and lower augenrest spines; fronto-orbital placed with the brachyurans and with which I place it. I width including augenrest about 85% maximum carapace followed the opinion of Mr. Audouin and Mr. Milne width. Lateral margins straight, diverging weakly poster- Edwards, who, looking with me last year at this little fossil iorly; weak swelling anterior to intersection of cervical carapace, regarded it as being like the homolid genera. I groove, margin incised at cervical groove; weak swelling have found it at Langrune (three miles north of Caen) in a posterior to cervical groove; incised at intersection of bank resting on coral-bearing limestone, and it is like the branchiocardiac groove; weakly convex posterior to pisolite of England. branchiocardiac groove. Posterior margin unknown. Mr. Tesson found a little later, in the Ranville quarry, Mesogastric region with long anterior process, widening another carapace of the Audoin homolid; it is about half posteriorly; spine at about half-length of anterior process; again the size of the first described in this article. The pair of spines at anterior end of widened area followed tubercles are disposed in the same manner; it offers, what is posteriorly by pair of muscle scars. Metagastric region more, two small [tubercles] close together, situated behind wide, rectangular, with muscle scars and pair of spines on and three [tubercles] close to the transverse groove; these top of muscle scars; urogastric region flattened, short; latter could have been easily destroyed in my small cardiac region triangular, with two spines anteriorly, specimen. posterior portion unknown, intestinal region unknown. 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 (Fo¨rster, 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 ACKNOWLEDGEMENTS anterior margin nearly on margin of augenrest. Museum work in Europe was funded by NSF grant EF 0531670 to Cervical groove deep, extending nearly straight across Feldmann and Schweitzer. M. Nose provided access to collections at the Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021 carapace. Postcervical groove composed of two discontin- Bayerische Staatsammlung fu¨r Palaöntologie, Mu¨nchen, Germany; M. uous, weak grooves bounding posterior margin of meta- Munt and C. Mellish, arranged our visit and photography at The Natural History Museum, London, UK; P. Crabbe provided images of some of the gastric region, with deep, arcuate pit at each lateral end. specimens illustrated herein from The Natural History Museum, London; Branchiocardiac groove extending weakly convex forward and S. De Grave arranged the examination and loan of material from the in oblique axial path to cardiac region. Epibranchial region Geological Collections, Oxford University Museum, UK. G. Schweigert, rectangular laterally, drawing into fingerlike projection Staatliches Museum fu¨r Naturkunde, Stuttgart, Germany, and H. Karasawa, Mizunami Fossil Museum, Japan, provided constructive directed at cardiac region axially. Remainder of branchial reviews of the manuscript. Our thanks go to all of these individuals. region undifferentiated. Subhepatic region inflated, bounded posteriorly by ventral extension of cervical groove; flank becoming less high posteriorly. Remainder of carapace and appendages unknown. REFERENCES Ahyong, S. T. and D. O’Meally 2004. Phylogeny of the Decapoda Material Examined.—BMNH In. 57979, paralectotype of Reptantia: Resolution using three molecular loci. The Raffles Bulletin of Morris (1980) and suggested as the lectotype herein; cast of Zoology 52: 673-693. Woodward specimen deposited in BMNH; OUM J.71902. ———, J. C. Y Lai., D. Sharkey, D. J. Colgan, and P. K. L. Ng. 2007. Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status Occurrence.—BMNH In. 57979, suggested lectotype: of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45: 576-586. Bathonian, Ranville, Calvados, France, couches a` poly- Alcock, A. 1900. Materials for a carcinological fauna of India 5: The piers, Tesson collection, purchased July, 1857, original of Brachyura Primigenia or Dromiacea. Journal of the Asiatic Society of Eudes-Deslongchamps; Woodward specimen: Bathonian Bengal 68(II:3): 123-169. Forest Marble, Great Oolite, Malmesbury, Wiltshire; OUM Beurlen, K. 1932. Brachyurenreste aus dem Lias von Bornholm mit J.71902: Lower Cornbrash, Ornithella obovata zone, Upper Beitra¨gen zur Phylogenie und Systematik der Brachyuren Dekapoden. Palaöntologische Zeitschrift 14: 52-66. Bathonian, Stratton Audley, Oxfordshire, UK. Beurlen, K., and M. F. Glaessner. 1930. Systematik der Crustacea Decapoda auf stammesgeschichtlicher Grundlage. Zoologisches Jahr- buch 60: 49-84, 22 pls. DISCUSSION Bouvier, E. L. 1896. Sur l’origine homarienne des crabes: e´tude comparative des dromiace´s vivants et fossils. Bulletin de la Socie´te´ The oldest confirmed Brachyura are Early and Middle philomathique de Paris 8: 34-110. Jurassic in age, and arrayed in at least three families and Collins, J. S. H., A. J. Ross, G. Genzano, and H. Manzian. 2006. Earleria two superfamilies (Table 1). Both Planoprosopon Schweit- gen. nov. and Gabriella gen. nov., replacement names for Foersteria zer et al., 2007, and Coelopus E´ tallon, 1861, of Long- Arai and Brinkmann-Voss, 1980 (Cnidaria, Hydrozoa, Mitrocomidae) and Foersteria Wehner, 1988 (Crustacea, Decapoda, Prosopidae), junior odromitidae Schweitzer and Feldmann, 2009a, are reported homonyms of Foersteria Sze´pligeti, 1896 (Insecta, Hymenoptera, from the Middle Jurassic of France, with ages of Bajocian Braconidae). Bulletin of the Mizunami Fossil Museum 33: 125-126. and Bathonian respectively (Schweitzer and Feldmann, De Haan, W. 1833-1850. Crustacea, pp. i-xvii, i-xxxi, ix-xvi, 1-243, pls. A-J, 2009b; 2010). Gabriella Collins et al., 2005 and Tanidro- L-Q, 1-55, circ. tab. 2. In, P. F. von Siebold (ed.), Fauna Japonica sive mites Schweitzer and Feldmann, 2008 [imprint 2007] of Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui summum in India Batava Imperium Tenent, Suscepto, Tanidromitidae are known from the Bajocian/Bathonian Annis 1823-1830 Collegit, Notis, Observationibus et Adumbrationibus and the latest Bajocian respectively (Schweitzer and Illustravit:. J. Mu¨ller et Co., Lugduni Batavorum [5 Leyden]. Feldmann, 2009b). Thus, based upon available evidence, E´ tallon, A. 1861. Notes sur les Crustace´s Jurassiques du bassin du Jura. the origin of Brachyura appears to lie in the Early Jurassic Me´moires de la Societe´ de l’Agriculture, des Sciences et Lettres de la within Homolodromioidea (Eoprosopon), with radiation Haute Saoˆne 9: 129-171, pl. 2. Eudes-Deslongchamps, J. A. 1835. Me´moire pour servir a` l’histoire into two superfamilies and multiple families by the Middle naturelle des Crustace´s fossiles. Me´moire de la Societe´ Linneénne de Jurassic. In the Late Jurassic, an evolutionary explosion Normandie 5: 37-46, pl. 1. within Brachyura occurred, with dozens of species and Feldmann, R. M., and C. E. Schweitzer. 2009. Revision of Jurassic genera known, primarily from Europe (Feldmann et al., Homoloidea De Haan, 1839, from the Ernstbrunn and Sˇtramberk 2006; Feldmann and Schweitzer, 2009; Schweitzer et al., limestones, Austria and the Czech Republic. Annalen des Naturhistor- ischen Museums in Wien (A) 111: 183-206. 2007; Schweitzer and Feldmann, 2008 [imprint 2007], ———, and ———. 2010. Is Eocarcinus the earliest brachyuran? Journal 2009a-d). of Crustacean Biology 30: 241-250. 256 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 30, NO. 2, 2010

———, I. Laza˘r, and C. E. Schweitzer. 2006. New crabs (Decapoda: ———, and ———. 2008 [imprint 2007]. A new classification for some Brachyura: Prosopidae) from Jurassic (Oxfordian) sponge bioherms of Jurassic Brachyura (Crustacea: Decapoda: Brachyura: Homolodromioi- Dobrogea, Romania. Bulletin of the Mizunami Fossil Museum 33: 1-20. dea): Families Goniodromitidae Beurlen, 1932 and Tanidromitidae new Fo¨rster, R. 1985. Fru¨he Anomuren und Brachyuren (Decapoda, Crustacea) family. Senckenbergiana lethaea 87: 119-156. aus dem mittleren Dogger. Mitteilungen der Bayerischen Staatssamm- ———, and ———. 2009a. Revision of the Prosopinae sensu Glaessner, lung fu¨r Palaöntologie und Historische Geologie 25: 45-60. 1969 (Crustacea: Decapoda: Brachyura) including 4 new families and 4 ———. 1986. Der erste Nachweis eines brachyuren Krebses aus dem Lias new genera. Annalen des Naturhistorischen Museums in Wien (A) 110: (Oberes Pliensbach) Mitteleuropas. Mitteilungen der Bayerischen 55-121. Staatssammlung fu¨r Palaöntologie und Historische Geologie 26: 25-31. ———, and ———. 2009b. Revision of Gabriella with new species. Garassino, A., A. De Angeli, and G. Schweigert. 2005. Brachyurans from Neues Jahrbuch fu¨r Geologie und Palaöntologie (Abhandlungen) 252 the Upper Jurassic (Kimmeridgian-Tithonian) of Pfalzpaint and Breiten- (1): 1-16. hill (Bavaria, S. Germany. Atti della Societa` Italiana di Scienze ———, and ———. 2009c. New species of Longodromitidae Schweitzer Naturale e del Museo Civico di Storia naturale di Milano 146(I): 69-78. and Feldmann, 2009, from the Ernstbrunn Formation, Late Jurassic Glaessner, M. F. 1929. Crustacea Decapoda, pp. 1-464. In, J. F. Pompeckj (Tithonian), Austria. Annalen des Naturhistorischen Museums in Wien (ed.), Fossilium Catalogus: Animalia, pars 41. W. Junk, Berlin. (A) 111: 207-224. ———. 1933. Die Krabben der Juraformation. Zentralblatt fu¨r Miner- ———, and ———. 2009d. Revision of the genus Cyclothyreus Remesˇ,

alogie, Geologie und Palaöntologie, Abteilung B: 178-191. 1895 (Decapoda: Brachyura: Dromioidea). Neues Jahrbuch fu¨r Geologie Downloaded from https://academic.oup.com/jcb/article/30/2/251/2419233 by guest on 02 October 2021 Guinot, D. 1995. Crustacea Decapoda Brachyura: Re´vision des Homo- und Palaöntologie (Abhandlungen) 253: 357-372. lodromiidae Alcock, 1900. In, A. Crosnier (ed.), Re´sultats des ———, and ———. 2010. The genus Coelopus E´ tallon, 1861 (Brachyura: Campagnes Musorstom, 13. Me´moires du Museum National d’Histoire Glaessneropsoidea: Longodromitidae) with new species. Neues Jahr- Naturelle, Paris 163: 155-282. buch fu¨r Geologie und Palaöntologie, (Abhandlungen) 253: 357-372. Heé, A., 1924. Catalogue critique des Crustace´s jurassiques du Calvados et ———, ———, and I. Lazar. 2007. Decapods from Jurassic (Oxfordian) de l’Orne. Bulletin de la Societe´ Linneénne de Normandie (7)6: 126- sponge megafacies of Dobrogea, Romania and reconsideration of 157, pls. 3-6. Nodoprosopon Beurlen, 1928. Neues Jahrbuch fu¨r Geologie und International Commission on Zoological Nomenclature. 1999. Interna- Palaöntologie, (Abhandlungen) 244: 99-113. tional Code of Zoological Nomenclature. International Trust for Van Straelen, V. 1924 [imprint 1925]. Contribution a` l’e´tude des Crustace´s Zoological Nomenclature, London. Dećapodes de la pe´riode jurassique. Me´moires d’Acade´mie Royale de Krobicki, M., and M. Zaton´. 2008. Middle and Late Jurassic roots of Belgique, (Science), [collected in number 4, series 2, 7: 1-462, pls. 1-10. brachyuran crabs: palaeoenvironmental distribution during their early von Meyer, H. 1835. Briefliche Mitteilungen. In, Leonhardt und Bronn’s evolution. Palaeogeography, Palaeoclimatology, Palaeoecology 263: 30-43. Neues Jahrbuch fu¨r Mineralogie, Geologie, und Palaöntologie, 1834: Linnaeus, C. [von] 1758. Systema Naturae per Regna tria Naturae, secundum 329. C. F. Winter, Stuttgart. classes, ordines, genera, species, cum characteribus, differentiis, synony- ———. 1842. U¨ ber die in dem dichten Jurakalk von Aalen in Wu¨rtemburg mis, locis (ed. 10), 1: 1-824. Laurentii Salvii, Holmiae [5 Stockholm]. Milne-Edwards, A., and E. L. Bouvier. 1902. Report on the results of vorkommenden Spezies des Crustaceengenus Prosopon. Beitra¨ge zur dredging by the U.S. Coast Survey Streamer ‘Blake.’ 39. Les Dromiace´s Petrefaktenkunde, 5: 70-75, pl. 15. et Oxystomes. Memoirs of the Museum of Comparative Zoology at ———. 1860. Die Prosoponiden oder die Familie der Maskenkrebse. Harvard College, Cambridge. Palaeontographica 7: 183-222, pl. 23. ¨ Morris, S. F. 1980. Catalogue of the type and figured specimens of fossil Wehner, G., 1988. Uber die Prosoponiden (Crustacea, Decapoda) des Jura: Crustacea (excl. Ostracoda), Chelicerata, Myriapoda and Pycnogonida 1-154, 8 pls., 1 insert. Dissertation zur Erlangung des Doktorgrades der in the British Museum (Natural History). Trustees of the British Fakulta¨t fu¨r Geowissenschaften der Ludwig-Maximilians-Universita¨t zu Museum (Natural History), 53 pp., 3 pls. Mu¨nchen. Reuss, A. E. 1858 [imprint 1857]. U¨ ber kurzschwa¨nzige Krebse im Jurakalke Withers, T. H. 1932. A liassic crab, and the origin of the Brachyura. Ma¨hrens. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaf- Annals and Magazine of Natural History, series 10, 9: 313-323, pls. ten, (Mathematisch-Naturwissenschaftliche Classe) 31: 5-13. 9, 10. Schweigert, G. 2006. A specimen of Prosopon hebes v.Meyer, 1840 Woodward, H. 1866. On the oldest known British crab (Palaeinachus (Decapoda: Brachyura: Prosopidae) from the Middle Jurassic of SW longipes) from the forest marble of Malmesbury, Wilts. Quarterly Germany. Neues Jahrbuch fu¨r Geologie und Palaöntologie (Mitt.) 2006 Journal of the Geological Society, 22: 493-494, pl. 14. (6): 361-370. ———, and J. Salter. 1865. Catalogue and chart of fossil Crustacea. Schweitzer, C. E., and R. M. Feldmann. 2005. Decapods, the Cretaceous- London. Palaeogene Boundary, and Recovery, pp. 17-53. In, S. Koenemann, and R. A. Jenner (eds.), Crustacea and Arthropod Relationships,Crustacean RECEIVED: 30 September 2009. Issues Vol. 16. Taylor and Francis Group, Boca Raton, FL. ACCEPTED: 13 October 2009.