New Fossil Decapod Crustaceans from the Remy Collection, Muséum National D’Histoire Naturelle, Paris

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New fossil decapod crustaceans from the Remy Collection, Muséum national d’Histoire naturelle, Paris

Carrie E. SCHWEITZER Kent State University Stark Campus, Department of Geology, 6000 Frank Ave. NW, North Canton, 44720 Ohio (USA) [email protected]

Rodney M. FELDMANN Kent State University, Department of Geology, Kent, 44242 Ohio (USA) [email protected]

Schweitzer C. E. & Feldmann R. M. 2010. — New fossil decapod crustaceans from the Remy Collection, Muséum national d’Histoire naturelle, Paris. Geodiversitas 32 (3): 399-415.

ABSTRACT Unpublished material originally identifi ed by J.-M. Remy, as well as evaluation of his type material deposited in the Muséum national d’Histoire naturelle, Paris, has KEY WORDS resulted in two new genera, Remyranina n. gen. and Planobranchia n. gen.; two new Crustacea, species, Liocarcinus heintzi n. sp. and Szaboa lamarei n. sp., and fi ve new combina- Decapoda, Brachyura, tions, Remyranina ornata n. comb., Calappilia? gorodiskii n. comb., Mursia simplex n. Cretaceous, comb., Planobranchia simplex n. comb., and Planobranchia laevis n. comb. ese taxa Eocene, Oligocene, include the fi rst Oligocene record for the Matutidae, and the extinct genus Szaboa France, is re-diagnosed. e Remy collection, comprised primarily of fossils from France Africa, and equatorial Africa, consists of numerous Calappidae and Carpilioidea, especially new species, new genera, abundant today in tropical environments, suggesting that at least for some taxa in new combinations. these groups, environmental tolerances have changed little since the Eocene.

RÉSUMÉ Nouveaux crustacés décapodes fossiles de la Collection Remy, Muséum national d’Histoire naturelle, Paris. L’étude du matériel non publié et la révision du matériel type de J.-M. Remy déposé au Muséum national d’Histoire naturelle, Paris, a permis de proposer deux genres nouveaux, Remyranina n. gen. et Planobranchia n. gen., deux espèces nouvelles, MOTS CLÉS Liocarcinus heintzi n. sp. et Szaboa lamarei n. sp., et cinq combinaisons nouvelles, Crustacea, Remyranina ornata n. comb., Calappilia? gorodiskii n. comb., Mursia simplex n. Decapoda, Brachyura, comb., Planobranchia simplex n. comb., et Planobranchia laevis n. comb. Un Matu- Crétacé, tidae de l’Oligocène est identiﬁ é pour la première fois et une nouvelle diagnose du Éocène, Oligocène, genre éteint Szaboa est proposée. La collection Remy, principalement constituée de France, fossiles de France et d’Afrique équatoriale, regroupe de nombreux Calappidae et Afrique, Carpilioidea. Ces crustacés, particulièrement abondants dans les environnements espèces nouvelles, genres nouveaux, tropicaux actuels, suggèrent que les tolérances environnementales de certaines combinaisons nouvelles. espèces de ces groupes ont peu changé depuis l’Éocène.

GEODIVERSITAS • 2010 • 32 (3) © Publications Scientiﬁ ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 399 Schweitzer C. E. & Feldmann R. M.

INTRODUCTION the matrix of the specimen of Liocarcinus heintzi n. sp. contains what appears to be miliolids. e e Muséum national d’Histoire naturelle, Paris calcilutite and calcirudite both have been reported houses a collection of decapod specimens de- to contain abundant terrigenous input (Puech- scribed or preliminarily identifi ed by Jean-Marcel maille & Vigneaux 1966), seen in the matrix of Remy during the mid-twentieth century. Our the specimen of Szaboa lamarei n. sp. us, the work and that of others over the past decade label information corresponds well with the actual has already revised and illustrated some of the lithology of the rock surrounding the specimens, taxa described in Remy’s works, bringing them so we can only assume it to be correct. up-to-date with the most recent classifi cations (see Collins & Morris 1975; Schweitzer 2003, DAX (LANDES, FRANCE) 2005; Feldmann & Portell 2007; Ahyong 2008; Remy reported that the specimen collected from Karasawa et al. 2008). Herein we evaluate the this locality, simply noted as Dax (Landes) was of remainder of the taxa described or preliminarily Eocene age, and an old label noted it as having been identifi ed by Remy, resulting in several new taxa part of the Nummulitique de Dax, collection of and combinations. A. Milne-Edwards. e Eocene age was probably Remy’s work has been particularly important derived from the association with a nummulitic for decapod studies because it includes some of rock unit. Indeed, Eocene rocks are well known the few decapod crustaceans known from the from that region of France, the southwestern re- continent of Africa (Remy 1960; Remy & Tessier gion of Aquitaine (Sztràkos et al. 1997, 1998). 1954; Gorodiski & Remy 1959). Some other works Fossil crabs have previously been reported from have described taxa from Egypt (Noetling 1885; the region (A. Milne-Edwards 1862-1865; Burger Lőrenthey 1909; Anderson & Feldmann 1995) et al. 1945; Daguin 1948; Boulanger et al. 1967; and other areas, but relatively few decapods are Sztràkos et al. 1998; Odin 2005). We herein sim- known from the continent. ply refer to the age of the specimen as Paleogene until more material with more precise locality data is collected. LOCALITIES AND GEOLOGIC SETTING

Discussed here are the localities for the previously ABBREVIATIONS unpublished specimens described herein. Remy’s BSP B ayerische Staatsammlung für Paläontolo- published work contains localities for the specimens gie und historische Geologie München, from Africa and Israel. Munich; IMDASlf.Nr. Staatliches Museum für Naturkunde, MONSÉGUR (GIRONDE, FRANCE) Stuttgart; Two decapod specimens were collected from Mon- KSU decapod collections, Kent State Univer- sity, Kent, Ohio; ségur, Liocarcinus heintzi n. sp. and Szaboa lamarei MR Természettudományi Múzeum, Föld-és n. sp. is locality belongs to the Calcaire à Astéries Ősléntár, Stéfania ut 14, Budapest (Hun- Formation (Rupelian). is formation is composed garian National Museum, Paleontological of carbonates in the northern region of Aquitaine and Geological Collection); (France), including calcilutites, calcarenites, and MNHN collections de Paléontologie du Muséum calcirudites, in that stratigraphic order (Puech- national d’Histoire naturelle, Paris. maille & Vigneaux 1966; Maillet & Puechmaille 1967). Magne (1947) conﬁ rmed the Oligocene SYSTEMATICS age based upon various fossils. e calcarenite has been noted to contain abundant miliolid foramini- Order DECAPODA Latreille, 1802 ferans (Maillet & Puechmaille 1967), and indeed, Infraorder BRACHYURA Linnaeus, 1758

400 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

Section EUBRACHYURA Remyranina ornata (Remy, 1960) n. comb. de Saint Laurent, 1980 (Fig. 1A) Subsection RANINOIDA De Haan, 1839 Superfamily RANINOIDEA De Haan, 1839 Raninella ornata Remy, 1960: 57, fi g. 2; fi gs 8, 9 of the unnumbered plate. Family RANINIDAE De Haan, 1839 Subfamily RANININAE De Haan, 1839 TYPE MATERIAL. — Holotype (dorsal carapace) by monotypy (MNHN R03847, coll. Tessier). Genus Remyranina n. gen. TYPE LOCALITY. — From probable Eocene rocks on the seacliff below Kraïebouen hill, near Fresco, 5.03°N, TYPE SPECIES. — Raninella ornata Remy, 1960. 5.31°W, Ivory Coast, Africa (Remy 1960). ETYMOLOGY. — e generic name combines the name MEASUREMENTS. — Measurements (in mm) taken on of the author of the type species and prominent French the holotype of Remyranina ornata n. comb.: maximum paleontologist, Jean-Marcel Remy, and Ranina, the type carapace length, > 18.7; maximum carapace width, 14.3; genus of the family. frontal width, 21.0; fronto-orbital width, 10.0. OCCURRENCE. — e sole specimen was collected from probable Eocene rocks of Ivory Coast, Africa. EMENDED DESCRIPTION Carapace small for family, longer than wide, mod- DIAGNOSIS. — Small raninid with expanded, moderately erately vaulted transversely, weakly arched longitu- broad fronto-orbital margin; ovoid, slightly longer than wide, with carapace ornamentation granular in anterior dinally; regions not well defi ned; surface granular in half and strongly terraced in posterior half. anterior half and strongly terraced posteriorly. Front narrow, about 17% carapace width; fronto-orbital DESCRIPTION. — As for emended description of spe- margin moderately broad, about 68% carapace cies. width; maximum carapace width, measured at mi- DISCUSSION dlength, less than 80% maximum carapace length; Although the specimen referred to the new genus posterior margin broken, estimated to be about 54% clearly allies it with the Ranininae, the combina- maximum width. Front projected slightly in advance tion of characters is unique and readily justifi es of postorbital spines; rostrum sulcate, trifi d, with axial creation of the new taxon. e genus to which spine projected in advance of lateral spines; axis of it is most closely allied is Lophoranina Fabiani, rostrum with subtle granular ridge. Orbital margin 1910. ese two genera are the only ones in the not well preserved; apparently sinuous with at least Raninidae that are characterized by possession one orbital fi ssure at about midpoint; terminating of a trifi d rostrum and development of terraced laterally in prominent outer orbital spine. Lateral ornamentation. However, Lophoranina tends to margin of outer orbital spine convex, widest near be widest near the front or at the level of the tip and narrowing to tip and to intersection with last anterolateral spine, whereas Remyranina anterolateral margin. Anterolateral margin widen- n. gen. is broadest at the midlength, well pos- ing posteriorly; with two spines (broken) dividing terior to the anterolateral spines. Furthermore, margin into thirds; general outline of carapace widest the antero lateral spines and the postorbital spines posterior to spines. Posterolateral margins nearly on Lopho ranina are generally small, whereas straight, converging posteriorly, rimmed by beaded those on Remy ranina n. gen. are large. Finally, marginal rim. Posterior margin broken. the terraced lines of Lophoranina extend nearly Surface of carapace divided into granular anterior to the front of the carapace, well in advance of half and terraced posterior half. Mesogastric region the branchiocardiac grooves, whereas those on discernable as extremely subtle pyriform swelling Remyranina n. gen. are confi ned to the posterior widening posteriorly from rostral ridge to anterior half of the carapace. ere seems to be no other termination of prominent, deep, arcuate branchio- genus of raninids with which Remyranina n. gen. cardiac pits. Regions on remainder of front half of could be confused. carapace indistinguishable; surface with fi ne gran-

GEODIVERSITAS • 2010 • 32 (3) 401 Schweitzer C. E. & Feldmann R. M.

ules, somewhat coarser axially and fi ner laterally. Calappilia? gorodiskii Posterior half of carapace crossed by prominent, (Remy in Gorodiski & Remy, 1959) n. comb. continuous to discontinuous, digitate, sinuous ter- (Fig. 1B) race structures; terrace structures generally concave forward; tips of digitations broken, fi ne, about 2.6 Atele cyclus gorodiskii Remy in Gorodiski & Remy, 1959: digitations per mm, forward-directed. Regions not 318, pl. 19, fi g. 2. — Salva & Feldmann 2001: 49. defi ned in posterior half. TYPE MATERIAL. — Holotype (dorsal carapace; MNHN Remainder of carapace, ventral surface, and ap- R03840, coll. Gorodiski), paratype of M’Bathié (Senegal) pendages not observed. not present in the MNHN Collections. TYPE LOCALITY. — Pougnar (Senegal), Upper Lutetian DISCUSSION (Gorodiski & Remy 1959). e description of this species given by Remy (1960: 57) is quite complete and little can be added to it. DISCUSSION e emended description has been written to use e type specimen originally referred to Atele cyclus consistent, modern terminology to facilitate com- gorodiskii is quite incomplete. All of the margins parison with other raninids. are broken, and nothing remains of the front and e nature of the fi ne structure developed on orbits. Dorsal carapace regions are reasonably well the terraced ornamentation on Remyranina ornata preserved, however, and indicate narrow, elongate n. comb. is very much like that seen on Lophoranina axial regions bordered by deep axial grooves; small, georgiana (Rathbun, 1935) (Feldmann et al. 1996: narrow protogastric regions; and large, granular fi g. 5), L. precocia Feldmann, Vega, Tucker, García- ornamentation on all of the carapace regions. e Barrera & Avendaño, 1996 and L. cristaspina Vega, carapace appears to have been ovate, but it is not Cosma, Coutiño, Feldmann, Nyborg, Schweitzer & possible to know for certain because of the broken Waugh, 2001. In those three species and, in all like- margins. Comparison of the nature of the carapace lihood, other species within the genus, the digitate regions with members of the Atelecyclidae (e.g., il- margins of terraced structures are broken ends of fi ne lustrations in Salva & Feldmann 2001) indicates that spines that are directed anteriorly (Vega et al. 2001: the species is not a member of that family. Species fi g. 4.7). ese small spines were interpreted by Feld- of Atele cyclus Leach, 1814, have arcuate swellings mann et al. (1996) to function as hooks to anchor parallel to the lateral margins of the cardiac region, the organism in very fi ne sediment or to capture bits which are absent in Atele cyclus gorodiskii. Other genera or organic debris as camoufl age (Savazzi 1981). It is within the Atelecyclidae and the related Cancridae possible that Remyranina ornata n. comb. also pos- Latreille, 1802, have narrow axial regions, but they sessed small spines that had a similar function. are not as long as those of Atele cyclus gorodiskii, and the cardiac and intestinal regions are never as long in those families as in Atele cyclus gorodiskii. Subsection HETEROTREMATA Guinot, 1977 e Calappidae can better accommodate Atele- Superfamily CALAPPOIDEA De Haan, 1833 cyclus gorodiskii. Members of that family possess Family CALAPPIDAE De Haan, 1833 narrow, elongate axial regions bordered by deep axial grooves; small, narrow protogastric regions; and Genus Calappilia A. Milne-Edwards, 1873 often possess large, granular ornamentation on all of the carapace regions, all of the observable features TYPE SPECIES. — Calappilia verrucosa A. Milne-Edwards, on Atele cyclus gorodiskii. Herein we tentatively refer 1873, by subsequent designation of Glaessner (1929). the species to Calappilia, a common Eocene genus characterized by all of these features. In addition, SPECIES INCLUDED. — See Feldmann et al. (2005). Calappilia possesses spines on the posterolateral DIAGNOSIS AND DISCUSSION. — See Feldmann et al. margin as does A. gorodiskii, and lacks a posterola- (2005) for a recent discussion. teral fl ange of spines, as is seen in many species of

402 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

A B

C D

FIG. 1. — Fossil Brachyura from Monségur (Gironde, France) : A, Remyranina ornata (Remy, 1960) n. comb., cast of holotype, MNHN R03847, dorsal carapace; B, Calappilia? gorodiskii (Remy in Gorodiski & Remy, 1959) n. comb., cast of holotype, MNHN R03840, dorsal carapace; C, D, Calappidae gen. et sp. indet., dorsal (C) and ventral (D) surfaces, MNHN R03836; E, Mursia simplex (Remy, 1960) n. comb., cast of holotype, MNHN R03849, dorsal carapace; F, Szaboa lamarei n. sp., holotype, MNHN R03779, dorsal carapace. Scale bars: 1 cm.

GEODIVERSITAS • 2010 • 32 (3) 403 Schweitzer C. E. & Feldmann R. M.

Calappa Weber, 1795. More complete specimens along the entire axial region. Cenomanocarcinus spp. will be necessary to confi rm generic placement for have a transverse ridge on the cardiac and branchial this species; however, placement within the Calap- regions which C. simplex lacks. Cenomanocarcinus pidae seems reasonable. spp. have an arcuate epibranchial ridge extending from the last anterolateral spine and terminating along the axis, which C. simplex lacks. Genus Mursia Leach, 1823 Cenomanocarcinus simplex is better accommodated within the Calappidae and Mursia. Cenomanocarcinus TYPE SPECIES. — Mursia cristiata H. Milne Edwards, simplex possesses large, circular, rimmed orbits; a 1837, by subsequent monotypy (see Ng et al. 2008). granular anterolateral margin with a small lateral FOSSIL SPECIES INCLUDED. — Mursia armata De Haan, spine; and fi ve dorsal carapace ridges with large 1837 (also extant); M. aspina Schweitzer & Feldmann, transverse granules. ese features are typical of 2000; M. australiensis Campbell, 1971 (also extant); Mursia spp.; thus, we refer the species to Mursia, M. bekenuensis Collins, Lee & Noad, 2003; M. circularis resulting in n. comb. (Karasawa, 1989); M. creutzbergi Collins & Donovan, Mursia simplex Mursia simplex 2004; M. granulosa Collins & Donovan, 2002; M. lien- n. comb. diff ers from other species of the genus in harti (Bachmayer, 1962); M. marcusana Rathbun, 1926; having less granular ornamentation overall. M. macdonaldi Rathbun, 1918; M. minuta Karasawa, 1993; M. obscura Rathbun, 1918; M. simplex (Remy, 1960) n. comb.; M. takahashii Imaizumi, 1952; M. yaquinensis (Remy, 1960) n. comb. Rathbun, 1926. Mursia simplex (Fig. 1E) DIAGNOSIS. — Carapace about as wide as long or wider than long, ovate; orbits circular, directed forward, front Cenomanocarcinus simplex Remy, 1960: 56, text-fi g. 1, narrow; anterolateral margins with small spines and fi g. 3 of the plate. long lateral spine or entire; carapace ornamented with fi ve or seven ridges, one axial and two or three on each TYPE MATERIAL. — Holotype (dorsal carapace) by side; ridges can be granular; carapace may be granular monotypy (MNHN R03849, coll. Tessier). between ridges; merus of cheliped may have long distal TYPE LOCALITY spine; manus with spines on upper margin; sternum . — Probable Eocene rocks on the seacliff ovate, narrow. below Kraïebouen hill, near Fresco, 5.03°N, 5.31°W, Ivory Coast, Africa (Remy 1960).

DISCUSSION DISCUSSION Mursia is easily diagnosed by the presence of well- One other Eocene species of Mursia is known, developed carapace ridges and generally granular from Pacifi c coastal North America (Schweitzer & ornamentation. It has been a widespread genus from Feldmann 2000). e records for the genus span Eocene to recent time, recorded from tropical to tem- tropical and temperate localities, so the occurrence perate regions (Schweitzer & Feldmann 2000). of an Eocene species in West Africa extends the geo- Remy (1960) referred a small, ovate specimen to graphic range of the genus but not the geologic or Cenomanocarcinus simplex. is species cannot be ecological range. e North American occurrence is referred to Cenomanocarcinus for several reasons (see late Eocene in age; it is not possible at this time to illustrations in Schweitzer et al. 2003 and Guinot et al. determine whether the African occurrence is older 2008). Species of Cenomanocarcinus are characterized as it is simply reported as probably Eocene. by ridges on the carapace, but they are arrayed both transversely and longitudinally. Cenomanocarcinus simplex has carapace ridges that are axially and obliquely Calappidae gen. et sp. indet. arrayed on the lateral regions of the carapace. Species (Fig. 1C, D) of Cenomanocarcinus have well-defi ned axial, protogastric, and hepatic regions, whereas C. simplex has axial MATERIAL EXAMINED. — Carapace (MNHN R03836, regions that are defi ned only by a deep lateral groove coll. Milne-Edwards).

404 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

MEASUREMENTS. — Measurements (in mm) taken on the DISCUSSION dorsal carapace of MNHN R03836: maximum carapace e specimen described here appears to be ref- width = 35.8; maximum carapace length > 33.0; fronto- erable to the family based upon its ovate shape; orbital width = 19.7; frontal width = 7.5. closely spaced, circular orbits; well-defi ned axial OCCURRENCE. — Paleogene rocks of Dax, Landes, in regions that are bounded by deep grooves; narrow Aquitaine, France. sternum with deep sterno-abdominal cavity; and sternal sutures 4/5 and 5/6 defi nitely interrupted. DESCRIPTION e specimen described here seems to lack poste- Carapace ovate, about as wide as long, widest at about rolateral spines, eliminating it from the speciose midlength; moderately vaulted both longitudinally Eocene genus Calappilia and also from Calappa, and transversely. Front narrow, about 20% maximum but those margins are heavily damaged. Species of carapace width, broken. Orbits directed forward; rims Mursia generally have granular ornamentation on unknown; fronto-orbital width 55% maximum cara- the dorsal carapace, which the specimen described pace width. Anterolateral and posterolateral margins here lacks. However, the specimen is damaged on confl uent; anterolateral portion appearing to have had all of the margins, so it is not possible to place it small spines or protuberances; posterolateral portion within a genus. Although the dorsal carapace is possibly rimmed; posterior margin broken. poorly preserved, the sternum is quite well preserved. Protogastric regions weakly infl ated, merged with Recovery of more specimens will help to place this mesogastric region. Metagastric and urogastric regions specimen within a taxon. confl uent, about same height as mesogastric region, lateral margins concave. Cardiac region rounded- triangular, elongate, two swellings anteriorly. Lateral Superfamily LEUCOSIOIDEA Samouelle, 1819 margins bounded by deep, lateral groove. Intestinal Family MATUTIDAE De Haan, 1841 region poorly defi ned. Hepatic region weakly infl ated centrally. Epibranchial region arcuate, weakly infl ated. Genus Szaboa Müller & Galil, 1998 Remainder of branchial regions undiff erentiated.

ird maxillipeds elongate. TYPE SPECIES. — Matuta inermis Brocchi, 1883 by Male sternum long, narrow. Sternites 1-2 fused, monotypy. no evidence of suture, triangular, separated from OTHER SPECIES INCLUDED. — Szaboa lamarei n. sp. sternite 3 by complete suture. Sternite 3 about as MATERIAL EXAMINED. — Cast of Szaboa inermis, MR45- long as wide, deeply depressed axially, separated 2-1. from sternite 4 by deep, oblique grooves. Sternite 4 much longer than wide, deeply grooved axially, EMENDED DIAGNOSIS. — Carapace wider than long, sterno-abdominal cavity extending about one-quarter length about 95% maximum width, widest 35-45% the the distance onto sternite, with short episternal distance posteriorly on carapace; front with axial spine, about 20% maximum carapace width; orbits deepest projections, directed anterolaterally. Sternite 5 axially, outer-orbital spine triangular, directed forward; wider than long, suture 4/5 incomplete, with short fronto-orbital width about 60% maximum carapace episternal projections, directed laterally; sternite 6 width; lower orbital rim extending well beyond upper- wider than long, suture 5/6 incomplete, with short orbital rim, with two blunt spines, pterygostomial regions episternal projections, directed posterolaterally; with sub-orbital swelling; anterolateral margin with several small spines; posterolateral and posterior margins sternite 7 wider than long, suture 6/7 incomplete, rimmed; protogastric, mesogastric, cardiac, epibranchial, directed strongly posterolaterally; sternite 8 short, and branchial regions with swellings; stridulating ridges wide; directed strongly posterolaterally, suture 7/8 on inner surface of manus. may or may not have been incomplete: preservation insuﬃ cient to determine. DISCUSSION Remainder of carapace and appendages un- We herein follow Schweitzer & Feldmann (2000) known. and Martin & Davis (2001) in placing the Ma-

GEODIVERSITAS • 2010 • 32 (3) 405 Schweitzer C. E. & Feldmann R. M.

tutidae within the Leucosioidea. Galil & Clark spines, at least one with granular ornament. Poste- (1994) extensively revised the matutid genera, rolateral margin weakly concave, probably weakly and Martin & Davis (2001) recognized the family rimmed. Posterior margin narrow, about 30% status of the Matutidae. Schweitzer & Feldmann maximum carapace width, weakly rimmed. (2000) followed Bellwood (1996) in recognizing Epigastric region weakly infl ated, spherical. Proto- the Matutidae and applied the family to the fossil gastric regions triangular, apex directed posteriorly, record as had Müller & Galil (1998). Despite this with transverse swelling centrally. Mesogastric region work, there is a limited fossil record for the family. with weak anterior extension, highly infl ated pos- Only four fossil species are known, the two spe- teriorly. Urogastric region depressed below level of cies of Szaboa mentioned here, Eomatuta granosa mesogastric and cardiac regions. Cardiac region most De Angeli & Marchiori, 2009, and one unnamed infl ated of all regions, produced into conical elevation. species attributed to the genus Ashtoret by Kara- Intestinal region very long, not well diff erentiated. sawa (2002). Hepatic region small, fl attened. Epibranchial region with large spherical swelling centrally. Remainder of branchial regions undiff erentiated, with small swell- Szaboa lamarei n. sp. (Fig. 1F) ing just adjacent to posterolateral margin. Pterygostomial region with suborbital swelling. Re- TYPE MATERIAL. — Holotype (carapace; MNHN R03779, mainder of carapace and appendages unknown. coll. Remy).

MEASUREMENTS. — Measurements (in mm) taken on DISCUSSION the holotype and sole specimen of Szaboa lamarei n. sp.: Szaboa lamarei n. sp. is easily distinguished from the maximum carapace width = 20.3; maximum carapace only other species in the genus, S. inermis, in possess- length = 19.3; fronto-orbital width = 11.3; posterior width = 6.3; length to position of maximum width = 6.7. ing much larger carapace swellings and apparently larger anterolateral swellings, although the latter is ETYMOLOGY. — e trivial name is that used by Jean- diffi cult to determine because the sole specimen of Marcel Remy on the label but that was never published, S. lamarei is abraded. e features of the two species in honour of Pierre Lamare (French geologist). We honor are essentially identical except that the ornamentation the work of both geologists by using Remy’s intended name. of S. lamarei is much more robust. Because the species of S. lamarei is an abraded mold of the interior, TYPE LOCALITY. — Monségur, Gironde, Rupelian, Cal- collection of additional material could provide more caire à Astéries Formation. details of the morphology of this species. DIAGNOSIS. — Outer-orbital spines triangular, directed e new species of Szaboa is accommodated within forward; carapace swellings large; lower orbital margin the Matutidae based upon its lower orbital margins with large swelling. extending well beyond the upper orbital margins, seen in extant species (Galil & Clark 1994) and in DESCRIPTION Szaboa inermis (Müller & Galil 1998: fi g. 1); its Carapace obovate, about as wide as long, length obovate carapace that is about as wide as long; and about 95% maximum carapace width, widest about its wide orbits that occupy most of the frontal mar- 35% the distance posteriorly on carapace at about gin of the carapace. e new species is superfi cially level of posterior edge of protogastric regions; similar to species of Eriosachila Blow & Manning, fl attened transversely and moderately vaulted lon- 1996, of the Aethridae Dana, 1851, but Eriosachila gitudinally. petiti Blow & Manning, 1996, the type species of Front abraded, broken. Orbits wide, outer orbital the genus, has small orbits with a lower orbital mar- spine triangular, directed forward; upper orbital gin that does not extend beyond the upper orbital margin sinuous; lower orbital margin extending margin and has well-defi ned carapace regions not well anterior of upper orbital margin, with central typical of the Matutidae. us, the new species is swelling. Anterolateral margins with several rounded best accommodated within the Matutidae.

406 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

Of the Matutidae, the new species is the only one not examined all available representative specimens of early Oligocene age. Eomatuta granosa is early of all species of Micromaia, Mithracia, and Pisomaia, Eocene in age, known from Italy (De Angeli & it is premature to comment on their relationships Marchiori 2009). e other two fossil species are in detail. at work is in progress. Miocene in age (Müller & Galil 1998; Karasawa 2002). e occurrence extends the geographic range somewhat, although Szaboa inermis was already Subfamily MAJINAE Samouelle, 1819 known from Hungary; thus, the probable range and distribution pathway to the current Indo-Paciﬁ c Genus Planobranchia n. gen. occurrences remains Tethyan. TYPE SPECIES. — Micromaja laevis Lőrenthey, 1909: 119, pl. 1, ﬁ g. 2a, b, by present designation.

Superfamily MAJOIDEA Samouelle, 1819 INCLUDED SPECIES. — Planobranchia laevis (Lőrenthey, Family MAJIDAE Samouelle, 1819 1909); P. simplex (Remy in Gorodiski & Remy, 1959).

ETYMOLOGY. — e generic name alludes to the depressed Subfamily MICROMAIINAE Beurlen, 1930 and fl attened metabranchial region which serves to readily distinguish the genus from closely related forms. e DISCUSSION gender is feminine. Beurlen (1930) provisionally united the genera DIAGNOSIS. — Carapace pyriform, widest at midlength Micromaia Bittner, 1875, Mithracia Bell, 1858 and of branchial region; moderately vaulted transversely and Pisomaia Lőrenthey in Lőrenthey & Beurlen, 1929, longitudinally. Gastric regions only weakly diff erenti- within the subfamily Micromaiinae. e basis for this ated; defi ned laterally by prominent V-shaped groove placement is not clear and the comparison of referred converging from anterior margin of orbits to urogastric taxa suggests that they bear little resemblance to one region, the narrowest part of axial regions. Cardiac region nearly as wide as widest part of gastric regions, hexago- another. e conformation of axial regions on the nal to ovoid in outline; bearing two nodes on medial type species of the three genera are totally diff erent transverse ridge. Intestinal region well defi ned, long, from one another; the outlines, particularly with approximately as wide as urogastric region. Epibranchial reference to the posterior margin are dissimilar; and and mesobranchial regions strongly infl ated, separated the orbital architecture of the two taxa on which it from one another by subtle arcuate attachment scar expressed on mold of the interior of the carapace; wid- is present on the type species, Micromaia tuberculata est part of these regions converge as angular projections Bittner, 1875, and on Pisomaia tuberculata Lőrenthey toward urogastric region. Metabranchial region extends in Lőrenthey & Beurlen, 1929, are diff erent. us, from widest part of cardiac region posterolaterally around there seems to be no unifying set of morphological posterior margin of metabranchial region and clearly criteria to justify the Micromaiinae. Further, Glaess- defi ned axially by posterior margin of cardiac region and intestinal region; depressed below other regions. ner (1969) characterized the orbital structure of Surface of carapace weakly ornamented by fi ne granules the subfamily as being like the Inachinae which he or pits; lacking strong tubercles. described as lacking orbits. at is certainly not the case in the several examples of Micromaia examined DISCUSSION by us and well illustrated by Beschin et al. (1985). Planobranchia laevis n. comb. and P. simplex n. comb. Investigation into the composition and validity of were both originally assigned to Micromaia based the subfamily is ongoing. upon overall similarities in outline and, in the absence Discussion of the Micromaiinae is warranted be- of other majid genera that the species more closely cause the type species of the new genus, Planobranchia resembled, the assignments seemed reasonable. How- n. gen., was originally placed within Micromaia. As ever, the two species exhibit several morphological discussed below, the species referred to the new genus features that readily distinguish them from other spe- bear close resemblance to members of the Majinae, cies assigned to Micromaia. Bittner (1875) originally and that placement is adopted here. Because we have defi ned Micromaia tuberculata, both the genus and

GEODIVERSITAS • 2010 • 32 (3) 407 Schweitzer C. E. & Feldmann R. M.

species, simultaneously. us, species referable to DISCUSSION Micromaia must conform to the concept of the genus e description by Remy in Gorodiski & Remy as defi ned by Bittner (1875: 76) within his species- (1959: 317) adequately describes the species and level description. e elements of his description of will not be repeated here. e genus Planobranchia Micromaia tuberculata that we take as characterizing n. gen. embraces two species that are quite similar the genus Micromaia include a pyriform outline; very to one another and confi rm their affi nities and their poorly defi ned regions, sometimes defi ned only by distinction from species of Micromaia as discussed the pattern of tubercles; axial regions that are more above. Planobranchia simplex n. comb. has a broader, or less uniform in width, and with the intestinal re- more infl ated epibranchial and mesobranchial region gion being very short or not diff erentiated; branchial than does the type species. e cardiac region of the regions that are undiff erentiated and uniformly in- Senegal specimen is narrower, about 21% maximum fl ated; and a uniform, dense array of coarse, often width, whereas the cardiac region of P. laevis n. comb. truncated-conical tubercles over the entire carapace. (Fig. 2B) is about 27% maximum width. e trans- e orbits are within the range of typical majids: verse ridge on the cardiac region of P. simplex n. comb. directed anterolaterally and arising posterior to the is more subtle than that of P. laevis n. comb., and the bifi d, weakly downturned rostrum. ridge on the latter species extends laterally to intersect e two species herein removed from Micromaia the margin of the mesobranchial region. Finally, the referred to Planobranchia n. gen. have regions that are metabranchial region on P. simplex n. comb. is ir- weakly defi ned by grooves, rather than tubercle pat- regularly elevated but smooth, whereas that surface terns; axial regions that are quite variable in width, and on P. laevis n. comb. is also irregularly elevated but with the cardiac region being nearly the widest region; is covered with tiny pits or perforations. e frontal a distinct, long, narrow intestinal region; a depressed, regions on both type specimens are broken, so that fl attened metagastric region that is clearly diff erentiated comparison is not possible. from the epibranchial and mesobranchial regions; and Both species of Planobranchia n. gen. were collected an overall absence of strong ornamentation. Although from Eocene rocks in North and West Africa, on the the fronts of both type specimens of P. laevis n. comb. south margin of the Tethyan seaway. e type species and P. simplex n. comb. are absent, the orbital margin was collected near Cairo, Egypt (Lőrenthey 1909). appears to be quite similar to other majids and, at least Species of Micromaia are known primarily from on P. simplex n. comb., there appears to be a slight Italy, but also from other localities on the northern indication of a bifi d rostrum. us, representatives margin of the Tethys seaway. of the two genera are readily distinguishable, based upon characters that are taxonomically important at the generic level in majids. Superfamily PORTUNOIDEA Rafi nesque, 1815 Family MATHILDELLIDAE Karasawa & Kato, 2003 Planobranchia simplex n. comb. Genus Branchioplax Rathbun, 1916 (Remy in Gorodiski & Remy, 1959) (Fig. 2A) Branchioplax? ballingi Remy in Remy & Tessier, 1954 Micromaia simplex Remy in Gorodiski & Remy, 1959: 317, (Fig. 2C) fi g. 2. Branchioplax ballingi Remy in Remy & Tessier, 1954: 189, TYPE MATERIAL . — Holotype (carapace) by monotypy text-fi g. 1, pl. 11, fi g. 6. — Karasawa et al. 2008: 125. (MNHN R03839, coll. Gorodiski). Casts of the type (KSU D1097) are deposited in the Department of Geo- TYPE MATERIAL. — Holotype (fragmental dorsal carapace) logy, Kent State University. by monotypy (MNHN R03793, coll.Tessier).

TYPE LOCALITY. — N’Gueyène, western Senegal, Upper TYPE LOCALITY. — Marigot de Balling, Senegal, Paleocene Lutetian (Gorodiski & Remy 1959). (Remy & Tessier 1954).

408 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

DISCUSSION nant, 1777), also Recent; L. depurator (Linnaeus, 1758), e holotype is only a partial specimen, retaining also Recent; L. holsatus (Fabricius, 1798), also Recent; the urogastric, hepatic, branchial, and parts of the L. kuehni (Bachmayer, 1953); L. lancetidactylus (Smirnov, 1929) (see Garassino & Novati 2001); L. marmoreus mesogastric and cardiac regions. e anterolateral (Leach, 1814), also Recent; L. oroszyi (Bachmayer, 1953); margin is broken, but shows the bases of four rela- L. praearcuatus Müller, 1996; L. pusillus (Leach, 1815), tively robust spines. e arcuate epibranchial region also Recent; L. rakosensis (Lőrenthey, 1929). and the marked muscle scars on the mesogastric DIAGNOSIS. — Carapace wider than long, length about region and along the urogastric region are what prob- 80% maximum width measured about half the distance ably led Remy to refer the species to Branchioplax, posteriorly on carapace; front variably ornamented; orbits which is characterized by both features. However, with two fi ssures on upper orbital margin; anterolateral the apparently large anterolateral spines are larger margins with fi ve fl attened spines including outer-orbital than those seen in species of Branchioplax. In ad- spine, fi rst four spines directed forward, last spine may be directed laterally or anterolaterally; regions generally dition, the holotype of B. ballingi, while broken, well defi ned, epibranchial region markedly arcuate and has a length to width ratio of about 67%. Other infl ated; regions may be ornamented with transverse species of Branchioplax, including B. washingto- ridges or granules; posterolateral reentrant large; posterior niana, the type species, are more equant, having margin nearly straight; merus of cheliped without distal ratios of 80-85%. It is possible that B. ballingi is spine on inner margin, carpus with large spine on inner angle, dactylus with three ridges; dactyl of pereiopod 5 more closely related to members of the Portunidae lanceolate (after Poore 2004). or Macropipidae, but more complete material will be necessary to fully assess its placement. For now, DISCUSSION we questionably refer it to Branchioplax. Species of the genus are quite variable in their dorsal carapace ornamentation, both fossil and extant. e extant Liocarcinus corrugatus is characterized by Family CARCINIDAE MacLeay, 1838 transverse ridges on the dorsal carapace, and such ornamentation is seen on the fossil species L. kuehni Subfamily POLYBIINAE Paul’son, 1875 and possibly on L. rakosensis. e illustration of L. rakosensis in Lőrenthey & Beurlen (1929: pl. 13, DISCUSSION fi g. 1a) shows six anterolateral spines; however, it Karasawa et al. (2008) provided diagnoses for the has been reported that some of the illustrations in Carcinidae and Polybiinae. e new species de- that volume are inaccurate (P. Müller pers. comm. scribed below is well accommodated within the 2004). Liocarcinus oroszyi is less well preserved than Polybiinae based upon its length to width ratio some other species and appears to have had some of about 80%; front occupying about 30% maxi- granular ornamentation on the dorsal carapace mum carapace width; fronto-orbital width of about (Bachmayer 1953). e other fossil specimens are 75% maximum carapace width; well-developed compression specimens, making details of the cara- posterolateral reentrants; fi ve anterolateral spines pace and ornamentation diffi cult to interpret. including the outer-orbital spine; and hexagonal e new species is referred to Liocarcinus based shape. No other portunoid family accommodates upon its possession of a wider than long carapace; this combination of features. fi ve anterolateral spines, the fi rst four of which are directed forward; marked posterolateral reentrants; granular ornamentation; and well-defi ned carapace Genus Liocarcinus Stimpson, 1871 regions. ese are typical of Liocarcinus. e referral of the new material to Liocarcinus marks the second TYPE SPECIES. — Portunus holsatus Fabricius, 1798, by Oligocene occurrence of the genus. Liocarcinus original designation. lancetidactylus had already been noted from the INCLUDED FOSSIL SPECIES. — Liocarcinus atropatanus Oligocene of the Caucasus (Smirnov 1929; Kara- (Aslanova & Dschafarova, 1975); L. corrugatus (Pen- sawa et al. 2008). Species of this genus are known

GEODIVERSITAS • 2010 • 32 (3) 409 Schweitzer C. E. & Feldmann R. M.

from the fossil record in Europe and what is now concave; posterolateral reentrant large, rimmed; southern Russia, and extant species are Atlantic posterior margin nearly straight, rimmed. and Indo-Pacifi c in distribution (Ingle 1980; Poore Epigastric region infl ated, equant. Mesogastric 2004). us, the genus may have had a Tethyan region with long anterior process terminating at pos- distribution and dispersal route in the past. terior end of epigastric regions, widened posteriorly, posterior portion pentagonal, posterior margin nearly straight. Protogastric regions hexagonal, weakly in- Liocarcinus heintzi n. sp. fl ated. Urogastric region short, wide. Cardiac region (Fig. 2D) wide, triangular, apex directed posteriorly, with two large, circular swellings anteriorly. Hepatic region de- TYPE MATERIAL. — Holotype (dorsal carapace; MNHN pressed, small, fl attened. Epibranchial region arcuate, R03778, coll. Remy). extending from last anterolateral spine to near cardiac ETYMOLOGY. — e trivial name is that used by Jean- region, ornamented with granules. Mesobranchial Marcel Remy on the museum label in honour of Émile region with longitudinal, granular swelling parallel Heintz of the MNHN, Paris, a specialist of fossil mam- and adjacent to axis, smooth laterally. Metabranchial mals, but that was never published. We honour both scientists by using Remy’s intended name. region infl ated parallel to posterior margin axially, disappearing laterally, granular. TYPE LOCALITY. — Monségur, Gironde, Aquitaine, Remainder of carapace and appendages un- SW France, Rupelian (Stampian), Calcaire à Astéries known. Formation.

DIAGNOSIS. — Carapace with granular ornamentation DISCUSSION on infl ated areas of regions; lateral margins crispate; Liocarcinus heintzi n. sp. diff ers from all other species anterolateral spines curving anteriorly. in the genus in possessing granular ornamentation on MEASUREMENTS. — Measurements (in mm) taken on the most infl ated portions of the carapace regions and the dorsal carapace: maximum carapace width = 8.4; in having crispate anterolateral margins terminating maximum carapace length = 6.9; length to position in upturned, forward directed anterolateral spines. of maximum width = 3.3; fronto-orbital width = 6.4; e specimen is rather incomplete, with a broken frontal width = 2.5. front and damaged orbits so a complete description is not possible. However, it seems that Liocarcinus is DESCRIPTION the best generic placement for it at this time. Carapace hexagonal, wider than long, length about 80% maximum width, widest at position of last anterolateral spine, about half the distance poste- Family MACROPIPIDAE riorly on carapace; carapace weakly vaulted longi- Stephenson & Campbell, 1960 tudinally and transversely; infl ated areas granular; Genus Pleolobites Remy, 1960 lateral margins crispate, upturned. Front damaged, about 30% maximum carapace Pleolobites erinaceous Remy, 1960 width. Orbits wide, apparently with two fi ssures, (Fig. 2E, F) outer-orbital spine triangular, directed forward; fronto-orbital width 75% maximum carapace Pleolobites erinaceous Remy, 1960: 59, text-fi gs 4, 5; fi gs 10, width. 11 of the plate. — Glaessner 1969: R513, fi g. 321.3. — Karasawa et al. 2008: 101. Anterolateral margins with fi ve spines including outer-orbital spine; second spine broad, triangular, TYPE MATERIAL. — Holotype (MNHN R03782, coll. directed forward; third spine smaller, triangular, Tessier); paratypes, 12 specimens (MNHN R03781, directed forward; fourth spine about same size as coll. Tessier). third; fi fth spine apparently directed anterolater- OCCURRENCE. — Eocene of Fresco, Ivory Coast (Remy ally. Posterolateral margin appearing to be weakly 1960).

410 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

ACB

FIG. 2. — Fossil Brachyura from Monségur (Gironde, France): A, Planobranchia simplex (Remy, 1960) n. comb., cast of holotype, MNHN R03839, dorsal carapace; B, Planobranchia laevis (LCrenthey, 1909) n. comb., cast of holotype, IMDASlf.Nr.25520/2005 (dorsal carapace); C, Branchioplax? ballingi Remy in Remy & Tessier, 1954, cast of holotype, MNHN R03793, fragmental dorsal carapace; D, Liocarcinus heintzi n. sp., holotype, R03778, dorsal carapace; E, F, Pleolobites erinaceous Remy, 1960, casts of holotype, MNHN R03782 (E), and MNHN R03781 (F). Scale bars: 1 cm.

GEODIVERSITAS • 2010 • 32 (3) 411 Schweitzer C. E. & Feldmann R. M.

Ursquilla yehoachi (Remy & Avnimelech, 1955) (Fig. 3)

Eryon yehoachi Remy & Avnimelech, 1955: 311, pl. 14a. Ursquilla yehoachi – Hof 1998: 258, ﬁ gs 2-4.

TYPE MATERIAL. — Holotype by monotypy (MNHN R62691, coll. Yehoach). Hof (1998) listed additional specimens.

TYPE LOCALITY. — Road from Beersheba to Mount Se- dom, valley of Wadi Seiyal, Israel, Campanian (Upper Cretaceous) (Remy & Avnimelech 1955: 312).

DISCUSSION At the time of preparation of the Treatise on Inverte- brate Paleontology, classifi cation of the stomatopods was considered to be quite simple (Holthuis & Manning 1969: R543). Holthuis & Manning (1969) referred the 24 genera of fossil and extant stomatopods to only two families. Since that time, FIG. 3. — Fossil Stomatopoda from Monségur (Gironde, France), extensive work on the group has resulted in a pro- Ursquilla yehoachi (Remy & Avnimelech, 1955), cast of holotype, liferation of suprageneric taxa so that Martin & MNHN R62691. Scale bar: 1 cm. Davis (2001) recognized seven superfamilies and 17 families of living forms. Far less proliferation TYPE LOCALITY. — Probable Eocene rocks on the sea- of fossil taxa has resulted owing to the paucity of cliff below Laga-Ghirobo hill, near Fresco, Ivory Coast, Africa (Remy 1960). work on fossil stomatopods since 1969. Hof (1998) restudied the type material of Eryon DISCUSSION yehoachi as well as additional unidentifi ed specimens Karasawa et al. (2008) distinguished Pleolobites and placed them into a new family and genus of from the similar Rhachiosoma Woodward, 1871. stomatopod. Holthuis & Manning (1969: R541) ey also noted the undocumented specimen in had already tentatively assigned the holotype of BSP indicating a Paleocene age for the genus and Eryon yehoachi to Squilla; thus, recognition that species. However, because the only well-documented it belonged within the Stomatopoda and not the material for Pleolobites erinaceous, the sole species Decapoda had been recognized for quite some time. of the genus, is indicated as Eocene, we can only Later, Ahyong (2008) synonymized Ursquillidae Hof, regard it as Eocene at this time. 1998, with Squillidae based upon comprehensive phylogenetic analysis including most fossils.

Subclass HOPLOCARIDA Calman, 1904 Order STOMATOPODA Latreille, 1817 DISCUSSION Suborder UNIPELTATA Latreille, 1825 Superfamily SQUILLOIDEA Latreille, 1802 Remy’s work has provided some of the few fossil Family SQUILLIDAE Latreille, 1802 decapods known from Africa. e collections he Genus Ursquilla Hof, 1998 described from Africa, as well as the specimens he identiﬁ ed but never published from France, docu- TYPE SPECIES. — Eryon yehoachi Remy & Avnimelech, ment a fauna with strong tropical inﬂ uence as might 1955, by original designation. be expected. e entire fauna contains three species

412 GEODIVERSITAS • 2010 • 32 (3) New fossil decapod crustaceans from the Remy Collection

of the Calappidae, a matutid, and three members Brachyura) with a reappraisal of the status of the of the Carpilioidea, each from diﬀ erent families family. Geological Journal of the Linnean Society 113: (Schweitzer 2003). Modern members of these 165-193. BESCHIN C., BUSULINI A., DE ANGELI A. & TESSIER G. groups are predominantly found in tropical areas 1985. — Il genere Micromaia Bittner (Crustacea, De- (Rathbun 1930; Galil & Clark 1994; Schweitzer & capoda) nel Terziario dell’area dei Berici e dei Lessini, Feldmann 2000). e region of western Africa and con descrizione di tre nuove specie. Lavori – Società France has thus embraced a fauna tending toward Veneziana di Scienze Naturali 10: 97-119. the modern, at least in terms of family and ecologi- BEURLEN K. 1930. — Vergleichende Stammesgeschichte Grundlagen, Methoden, Probleme unter besonderer cal composition, since the Eocene. Berücksichtigung der höheren Krebse. Fortschrifte in der Geologie und Paläontologie 8: 317-586. BITTNER A. 1875. — Die Brachyuren des vicentinischen Acknowledgements Tertiärgebirges. Denkschriften der kaiserlichen Akademie A. Lauriat-Rage facilitated our work at the MNHN der Wissenschaften Mathematisch-Naturwissenschaftliche Klasse 34: 63-105, pls 1-5. and the original loan of the material and photographs BOULANGER D., MASSIEUX M., PLAZIAT J. C., & TOU- of specimens in 2001. NSF grant EF-0531670 MARKINE M. 1967. — Essai de corrélation stratigra- funded comparative work in museums in Europe phique entre les marnes à Xanthopsis dufourii de la and the United States. Important comparative casts Chalosse, les couches à Nummulites atacicus des Cor- were made in the Természettudományi Múzeum, bières septentrionales et du Mont-Cayla et le Lutétien du bassin de Paris. Bulletin de la Société géologique de Föld-és Ősléntár, Budapest, Hungary; Museo Civico France (7) 9: 367-370 “G. Zannato”, Montecchio Maggiore, Italy; and BURGER J. J., CUVILLIER J. & SCHOEFFLER J. 1945. — the Department of Paleontology, Geological Sur- Stratigraphie du Nummulitique de la Chalosse de vey of Austria, Vienna. Our thanks to P. Müller Montfort. Bulletin de la Société géologique de France (Természettudományi Múzeum); C. Beschin and 5 (4-6): 207-220. COLLINS J. S. H. & MORRIS S. F. 1975. — A new crab A. De Angeli (Museo Civico “G. Zannato”, Mon- Costacopluma concava from the Upper Cretaceous of tecchio Maggiore, Italy); A. Garassino (Museo Nigeria. Palaeontology 18: 823-829. Civico de Storia Naturale di Milano, Italy); and I. DAGUIN F. 1948. — L’Aquitaine occidentale. Géologie ré- Zorn (Geological Survey of Austria) for facilitating gionale de la France 5. Hermann & Cie, Paris, 232 p. access to these respective collections. Construc- DE ANGELI A. & MARCHIORI L. 2009. — Eomatuta granosa n. gen., n. sp. (Decapoda, Brachyura, Matu- tive reviews of the manuscript were provided by tidae), nuovo crostaceo dell’Eocene dei Monti Berici H. Karasawa (Mizunami Fossil Museum) and an (Vicenza, Italia settentrionale). Lavori – Società Vene- anonymous reviewer. ziana di Scienze Naturali 34: 105-110. FELDMANN R. M. & PORTELL R. W. 2007. — First report of Costacopluma Collins and Morris, 1975 (Decapoda: Brachyura: Retroplumidae) from the REFERENCES Eocene of Alabama, U.S.A. Journal of Crustacean Biology 27: 90-96. AHYONG S. T. 2008. — Phylogenetic analysis of the FELDMANN R. M., VEGA F., TUCKER A. B., GARCIA- Squilloidea (Crustacea: Stomatopoda). Invertebrate BARRERA P. & A VENDAÑO J. 1996. — e oldest Systematics 19: 189-208. record of Lophoranina (Decapoda: Raninidae) from ANDERSON J. L. & FELDMANN R. M. 1995. — Lobo- the late Cretaceous of Chiapas, southeastern Mexico. carcinus lumacopius (Decapoda: Cancridae), a new Journal of Paleontology 70: 296-303. species of cancrid crab from the Eocene of Fayum, FELDMANN R. M., SCHWEITZER C. E. & ENCINAS A. Egypt. Journal of Paleontology 69: 922-932. 2005. — New decapods from the Navidad Forma- BACHMAYER F. 1953. — Die Decapodenfauna des tor- tion (Miocene) of Chile. Journal of Crustacean Biology tonischen Leithakalkes von Deutsch-Alternburg 25: 427-449. (Niederösterreich). Mitteilungen der Geologischen GALIL B. S. & CLARK P. F. 1994. — A revision of the Gesellschaft in Wien 44 (dated 1951, published 1953): genus Matuta Weber, 1795 (Crustacea: Brachyura: 237-262. Calappidae). Zoologische Verhandelingen 294: 3-55, BELLWOOD O. 1996. — A phylogenetic study of the pls 1-14. Calappidae H. Milne Edwards 1837 (Crustacea: GARASSINO A. & NOVATI M. 2001. — Liocarcinus lan-

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Submitted on 19 January 2009; accepted on 20 October 2009.

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