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Among Erymidae Van Straelen, 1925 (Van Straelen V. 1925. Contribution à l'étude des crustacés décapodes de la période jurassique. Mémoires de la Classe des Sciences de l'Académie royale de Belgique 7: 1–462), typical Mesozoic crustaceans, the genus Eryma Meyer, 1840 (Meyer H. von. 1840a. Briefliche Mittheilungen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefactenkunde 576–587) includes the largest number of species, mainly from Jurassic deposits. However, the lack of clear diagnoses for erymid genera has led to mistakes in generic assignments and to the establishment of redundant genera. The review of the concept of Eryma herein presents an attempt to clarify its diagnosis, mainly supported by the carapace groove pattern and the morphology of chelae of the first pair of pereiopods, and to emphasize its systematic implications. Thus, we maintain the synonymy of Klytia Meyer, 1840, Bolina Münster, 1839 (Münster G. 1839. Decapoda Macrura. Abbildung und Beschreibung der Fossilen Langschwänzigen Krebse in den Kalkschiefern von Bayern. Beiträge zur Petrefaktenkunde 2: 1–88) (sensu Étallon [Étallon A. 1859. Description des crustacés fossiles de la Haute-Saône et du Haut-Jura. Bulletin de la Société géologique de France 16: 169–205]), and Erymastacus Beurlen, 1928 (Beurlen K. 1928. Die Decapoden des Schwäbischen Jura mit Ausnahme der aus den oberjurassischen Plattenkalken stammenden. Palaeontographica 70: 115–278) with Eryma. Moreover, a review of the genera Protoclytiopsis Birshtein, 1958 (Birshtein JA. 1958. Ein Vertreter der ältesten Ordo der Crustacea Decapoda Protoclitiopsis antiqua gen. nov. sp. nov. aus dem Permo West-Sibiriens. Doklady Akademii Nauk, SSSR 122: 477–480), and Galicia Garassino and Krobicki, 2002 (Garassino A, Krobicki M. 2002. Galicia marianae n. gen., n. sp. (Crustacea, Decapoda, Astacidea) from the Oxfordian (Upper Jurassic) of the Southern Polish Uplands. Bulletin of the Mizunami Fossil Museum 29: 51–59), reveals the presence of a junction between the postcervical and branchiocardiac grooves. This feature is diagnostic of Eryma and supports the integration of these genera into the synonymy of Eryma. The addition of Protoclytiopsis to the synonymy of Eryma makes Eryma antiquum (Birshtein, 1958) nov. comb. the oldest representative of the genus and of the family, extending its stratigraphic range to the Late Permian (Changhsingian). Thus, this work also emphasizes that Erymidae crossed the Permian-Triassic boundary.

Parmi les Erymidae Van Straelen, 1925 (Van Straelen V. 1925. Contribution à l'étude des crustacés décapodes de la période jurassique. Mémoires de la Classe des Sciences de l'Académie royale de Belgique 7: 1–462), crustacés typiques du Mésozoïque, Eryma Meyer, 1840 (Meyer H. von. 1840a. Briefliche Mittheilungen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefactenkunde 576–587) est le genre regroupant le plus grand nombre d'espèces principalement dans les dépôts jurassiques. Cependant, le manque de clarté des diagnoses des genres d'Erymidae a mené à des erreurs dans les identifications génériques ainsi qu'à l'instauration de genres redondants. La révision du concept du genre Eryma présentée ici est donc l'occasion de proposer une diagnose clarifiée, reposant essentiellement sur l'architecture des sillons de la carapace et la morphologie des pinces de la première paire de péréiopodes, et d'en souligner les implications systématiques. Ainsi, la mise en synonymie avec Eryma des genres Klytia Meyer, 1840, Bolina Münster, 1839 (Münster G. 1839. Decapoda Macrura. Abbildung und Beschreibung der Fossilen Langschwänzigen Krebse in den Kalkschiefern von Bayern. Beiträge zur Petrefaktenkunde 2: 1–88) (sensu Étallon [Étallon A. 1859. Description des crustacés fossiles de la Haute-Saône et du Haut-Jura. Bulletin de la Société géologique de France 16: 169–205]) et Erymastacus Beurlen, 1928 (Beurlen K. 1928. Die Decapoden des Schwäbischen Jura mit Ausnahme der aus den oberjurassischen Plattenkalken stammenden. Palaeontographica 70: 115–278) est ici confirmée. De plus, la révision des genres Protoclytiopsis Birshtein, 1958 et Galicia Garassino and Krobicki, 2002 (Garassino A, Krobicki M. 2002. Galicia marianae n. gen., n. sp. (Crustacea, Decapoda, Astacidea) from the Oxfordian (Upper Jurassic) of the Southern Polish Uplands. Bulletin of the Mizunami Fossil Museum 29: 51–59) révèle la présence d'une jonction entre les sillons postcervical et branchiocardiaque. Ce caractère diagnostique d'Eryma justifie l'intégration de ces deux genres dans la synonymie d'Eryma. L'ajout de Protoclytiopsis à la synonymie d'Eryma fait d'Eryma antiquum (Birshtein, 1958) nov. comb. le plus ancien représentant du genre et de la famille, repoussant son extension stratigraphique à la fin du Permien (Changhsingien). Ainsi, ce travail met en lumière le fait que les Erymidae ont franchi la limite Permien-Trias.

Introduction

ErymaMeyer, 1840 is a genus of typical Mesozoic crustaceans. It is known by many fossils supporting the greatest number of described species among Erymidae Van Straelen, 1925 (Schweitzer et al., 2010; Hyžný et al., 2015). Eryma is well known in European Jurassic and Early Cretaceous deposits (Carter, 1886; Étallon, 1859, 1861; Oppel, 1861, 1862; Ferry, 1865; Krause, 1891; Lahusen, 1894; Harbort, 1905; Van Straelen, 1920, 1925; Hée, 1924; Beurlen, 1928; Woods, 1925–1931; Birshtein, 1956; Förster, 1965, 1966; Martill, 1991; Garassino, 1996; Schweigert et al., 2000; Fischer, 2003; Etter, 2004; Carpentier et al., 2006; Garassino and Schweigert, 2006; Charbonnier, 2009; Schweitzer et al., 2009; Charbonnier et al., 2010, 2012, 2014, in press; Bravi et al., 2014; Devillez et al., 2016), and some occurrences are also reported throughout the world: in Middle East (Roger, 1946; Förster and Seyed-Emani, 1982; Garassino, 1994), in Africa (Beurlen, 1933; Secrétan, 1984), in North America (Rathbun, 1923, 1926; Feldmann and Titus, 2006) in Latin America (Aguirre-Urreta and Ramos, 1981; Aguirre-Urreta, 1989; J. Luque, pers. com.), in Antarctic (Taylor, 1979) and in Japan (Karasawa et al., 2008; Kato et al., 2010; Devillez et al., 2016).

Currently, the carapace groove pattern of erymid lobsters is the main feature used for genera and species distinction (Van Straelen, 1925; Förster, 1966; Glaessner, 1969). However, the lack of clear generic diagnoses in the works of the nineteenth century and the different states of preservations of crustaceans has led to confusions about the concept of Eryma, and descriptions of redundant genera as consequences (BolinaMünster, 1839, Eryma and Klytia Meyer, 1840 sensu Étallon [1859]). Such confusions persist nowadays but some recent works have attempted to bring clear elements for erymid lobsters discrimination (Hyžný et al., 2015; Devillez et al., 2016).

This work aims to clarify the concept of Eryma on the basis of the carapace groove pattern and the morphology of the first pereiopod chelae. It also emphasizes systematic and stratigraphic implications of such clarification.

Material and methods

The studied material includes 42 specimens from the palaeontological collections of European institutions (Tab. 1). Except fossils from the Solnhofen Lithographic Limestones, most of the studied specimens are fragmentary with only isolated carapaces or chelae preserved. They were mostly studied using a CETI Belgium MEDO binocular microscope. Line drawings were prepared by the first author.

Systematic palaeontology

MALACOSTRACA Latreille, 1802

DECAPODA Latreille, 1802

ERYMIDA sensu Schram and Dixon, 2004

Superfamily ERYMOIDEA Van Straelen, 1925

Family ERYMIDAE Van Straelen, 1925

Preliminary remark. – Throughout the literature Erymidae has been included within Astacidea Latreille, 1802 (Van Straelen, 1925; Glaessner, 1969; Aguirre-Urreta, 1989; Schweigert et al., 2000; Garassino and Krobicki, 2002; Crônier and Courville, 2004; Feldmann and Titus, 2006; Garassino and Schweigert, 2006; Schweigert, 2013; Charbonnier et al., 2013), or within Glypheidea Zittel, 1885 (De Grave et al., 2009; Schweitzer et al., 2010; Wahle et al., 2012; Karasawa et al., 2013; Feldmann et al., 2015). Recent phylogenetic analysis of Charbonnier et al. (2015) questioned the assignment of Erymidae to Glypheidea. Then, following Hyžný et al. (2015), we consider the systematic position of the Erymidae as uncertain and do not list the taxonomic rank beyond the superfamily. Hence, following Schram and Dixon (2004), we only include the Erymidae in a separate clade, Erymida.

Genus Eryma Meyer, 1840

(Fig. 1A-C)

Eryma Meyer, 1840: 587.

Klytia Meyer, 1840: 19.

BolinaMünster, 1839,sensuÉtallon (1859: 192) (non Mertens, 1833).

ProtoclytiopsisBirshtein, 1958: 477. nov. syn.

GaliciaGarassino and Krobicki, 2002: 55. nov. syn.

ErymaOppel (1862: 20). − Zittel (1885: 693). − Méchin (1901: 74). − Van Straelen (1925: 233). − Rathbun (1926: 127). − Secrétan (1964: 61). − Förster (1966: 88). − Glaessner (1969: R455). − Aguirre-Urreta and Ramos (1981: 609). − Secrétan (1984: 516). − Aguirre-Urreta (1989: 513). − Crônier and Courville (2004: 1004). − Feldmann and Titus (2006: 63). − Feldmann and Haggart (2007: 1792). − Hyžný et al. (2015: 375). − Feldmann et al. (2015: 1).

ClytiaBeurlen (1928: 165).

ProtoclytiopsisFörster (1966: 86). − Feldmann et al. (2015: 10).

KlytiaGlaessner (1969: R456).

GaliciaFeldmann et al. (2015: 3).

Type species.–Macrourites modestiformisSchlotheim, 1822, by subsequent designation of Glaessner (1929).

Included species. – A list of fossil species of Eryma, including 60 species, has been done by Schweitzer et al. (2010). However, this work is a compilation, and the species of Eryma need a careful review as suggested by Bravi et al. (2014) and Hyžný et al. (2015). Such review requires the examination of the type material of all included species, so the proposition of a new list is beyond the aims of the present contribution.

Emended diagnosis.–Fusiform intercalated plate; deep cervical groove, joined to dorsal margin and to antennal groove; short gastro-orbital groove originating as a slight median inflexion of cervical groove; postcervical groove joined medially to branchiocardiac groove; branchiocardiac groove strongly inclined, joined to hepatic groove; concavo-convex hepatic groove, joined to cervical groove; inferior groove convex posteriorly, joined to hepatic groove and to ventral margin; inflated ω area; cephalic region with two divergent rows of tubercles: orbital row with strong distal spine and antennal row with strong distal antennal spine; chelate P1-P3; P1 chelipeds without prominent spines and with homogeneous ornamentation; P1 propodus dorso-ventrally compressed with narrow inner and outer margins; P1 with narrow dactylar bulge; fingers longer than P1 propodus, equal in length, narrowing gradually to distal extremity; index wider than dactylus (modified after Devillez et al., 2016).

Comments. – According to Hyžný et al. (2015), followed by Devillez et al. (2016), we distinguish two forms of P1 chelae: form I (Fig. 1B) with short rectangular propodus and straight fingers, slightly longer than the propodus; form II (Fig. 1C) with an elongate sub-rectangular or trapezoidal propodus bearing elongated fingers, usually curved inward.

The most recent diagnoses of Eryma proposed by Feldmann and Titus (2006), Hyžný et al. (2015) and Feldmann et al. (2015) are relatively broad, particularly about the characteristics of the groove pattern. So, among the erymid lobsters, Eryma is the only genus exhibiting a junction between postcervical and branchiocardiac grooves. Other erymids such as EnoploclytiaM'Coy, 1849 (Fig. 1F) and Pustulina Quenstedt, 1857 (Fig. 1G) show a reduced branchiocardiac groove only joined to dorsal margin, while it is well-developed and joined to the posterior extremity of the hepatic groove in PalaeastacusBell, 1850 (Fig. 1D) and StenodactylinaBeurlen, 1928 (Fig. 1E) (Devillez et al., 2016).

Discussion. – Meyer (1840) established two genera: (1) Eryma for small crustaceans from the Solnhofen Lithographic Limestones, previously assigned to GlypheaMeyer, 1835 by Münster (1839) (Fig. 2A), and (2) Klytia with Klytia ventrosa (Meyer, 1835) (Oxfordian, Haute-Saône, France) (Fig. 2C-D), and Klytia mandelslohi Meyer, 1840 (Oxfordian, Baden-Württemberg, Germany). According to Étallon (1859), the different states of preservation between the strongly compressed fossils from Solnhofen and the three-dimensionally preserved ones from Haute-Saône and Baden-Württemberg probably prevented Meyer (1840b) to assign them to the same genus. However, Klytia was clearly established based on the junction between postcervical and branchiocardiac grooves (see Meyer (1840b): 20]), which supports the synonymy between Eryma and Klytia. This synonymy was followed by numerous subsequent authors (e.g., Oppel, 1862; Van Straelen, 1925; Förster, 1966) even if some authors maintained the distinction between the two genera (e.g., Beurlen, 1928; Glaessner, 1929). After some hesitation, Glaessner (1969: R626) re-established the synonymy, which has been commonly accepted since then.

Münster (1839) erected Bolina with two new species from the Solnhofen Lithographic Limestones: Bolina pustulosa and Bolina angusta. Later, Étallon (1859) proposed an emended description for Bolina. He described the groove pattern as follows: there are three oblique, wide and deep transverse grooves; the first (cervical groove) is inflected forward and the two others (postcervical and branchiocardiac grooves) are dorsally close, joining each other at carapace mid-height. The carapace groove pattern of Bolina sensu Étallon (1859) clearly fits the diagnosis of Eryma and does not fit the descriptions of Bolina pustulosa and Bolina angusta, which are clearly not erymid lobsters. Moreover, Oppel (1861) proposed the new genera Stenochirus (type species: Bolina angusta) and Pseudastacus (type species: Bolina pustulosa) and remarked that BolinaMünster, 1839 was preoccupied by BolinaMertens, 1833 (Cnidaria).

Beurlen (1928) established the genus Erymastacus to accommodate erymid lobsters only known by P1 chelae bearing very long fingers, and previously assigned to Eryma. Later, Glaessner (1929) subsequently designated Glyphea ornati Quenstedt, 1857 as type species for Erymastacus. Throughout the literature this genus was regarded as a junior synonym of Eryma (Förster, 1966; Glaessner, 1969; Schweitzer et al., 2010; Feldmann et al., 2015) or as a distinct genus (Secrétan, 1964; Schweigert et al., 2000; Schweigert and Garassino, 2003; Hyžný et al., 2015). Careful examination of the lectotype of Glyphea ornati (Fig. 2B) leads us to regard it as P1 chelae of Eryma, belonging to form II because of the trapezoidal shaped propodus, the inward curvature of fingers and the index width greater than that of dactylus. In conclusion, we agree with Förster (1966), Glaessner (1969), Schweitzer et al. (2010), Feldmann et al. (2015) and Devillez et al. (2016), by maintaining the synonymy between Erymastacus and Eryma.

Birshtein (1958) erected the new genus Protoclytiopsis with Protoclytiopsis antiqua (Changhsingian, Siberia, Russia) as the type species. This species is known by a single laterally crushed carapace. Förster (1966) pointed out the close similarity between the carapace groove pattern of this species and that of Eryma spp. Finally, he included Protoclytiopsis within the subfamily Eryminae Beurlen, 1928. Förster's opinion was followed by Glaessner (1969) but not by Karasawa et al. (2013), Feldmann et al. (2012) and Feldmann et al. (2015), who assigned Protoclytiopsis within the family Clytiopsidae Beurlen, 1927 (pro Clytiopsinae Beurlen, 1927, which was elevated to family rank by Amati et al. [2004]).

Our examination of the photograph of the holotype of Protoclytiopsis antiqua reveals the presence of the typical grooves of the Erymidae: deep cervical groove, gastro-orbital groove present, postcervical and branchiocardiac grooves almost parallel. Moreover, the postcervical and branchiocardiac grooves are joined medially, as in Eryma (Fig. 2E-F). In conclusion, we consider Protoclytiopsis as a junior synonym of Eryma and the new combination Eryma antiquum (Birshtein, 1958) is herein proposed.

Garassino and Krobicki (2002) erected the new genus Galicia with Galicia marianae (Oxfordian; southern Poland) as the type species. Most of the authors assigned this genus to the family Erymidae (Garassino and Krobicki, 2002; Schweitzer et al., 2010; Karasawa et al., 2013; Feldmann et al., 2015) while others included it in the family Clytiopsidae because of the absence of intercalated plate (Feldmann et al., 2012). Our review of the holotype indicates a carapace shape and a groove pattern typical of an Erymidae (subcylindrical carapace, deep cervical groove, gastro-orbital groove present, postcervical and branchiocardiac grooves almost parallel). Moreover, the carapace groove pattern exhibits the junction between the postcervical and the branchiocardiac grooves which is diagnostic of the genus Eryma (Fig. 2G-H). Hence, we consider Galicia to be a junior synonym of Eryma.

In addition, Galicia marianae exhibits a carapace with the following morphological characters: a wide branchial region, a well-marked gastro-orbital groove, subparallel postcervical and branchiocardiac grooves, slightly concave forward, a short branch extends the postcervical groove under its junction with branchiocardiac groove, a slightly inflated ω area and a flat χ area, an ornamentation made of fine tubercles. The carapace groove pattern and ornamentation are very similar to those of the well-known Eryma ventrosum (Meyer, 1835) from the Terrain à Chailles Formation (Oxfordian, Haute-Saône, France) (Meyer, 1840b; Bronn, 1851–1852; Étallon, 1859; Van Straelen, 1925; Charbonnier et al., 2012) (Fig. 2C-D). Hence, we consider G. marianae to be a junior synonym of E. ventrosum.

Stratigraphic implications

Until now, Eryma extended from Sinemurian (Early Jurassic), with E. meyeriGarassino, 1996, to Albian (Early Cretaceous), with E. vocontiiDevillez et al., 2016.

However, our examination of the holotype MSNM i7606 of E. meyeri (Osteno, Italy) reveals morphological characters diagnostic of PalaeastacusBell, 1850: postcervical groove not joined medially to branchiocardiac groove but joined ventrally to hepatic groove, short and stout P1 chelae covered by spines (Fig. 3A-B). Thus, we propose the new combination: Palaeastacus meyeri (Garassino, 1996). Our review of the holotype MSNM il3517 of Pustulina sinemurianaGarassino, 1996, from the same locality (Osteno, Italy) reveals morphological characters diagnostic of Eryma: postcervical and branchiocardiac grooves joined medially, elongate P1 chelae (Fig. 3C-D). Thus, we propose the new combination: Eryma sinemuriana (Garassino, 1996).

Most of the described species of Eryma occurs in Jurassic (Fig. 2A-D, 2G-H, 3C-D) while only four species occurs in Early Cretaceous (Devillez et al., 2016): Eryma glaessneri (Van Straelen, 1936) and ErymasulcatumHarbort, 1905 in Hauterivian (Fig. 3I-L), ErymanipponKarasawa et al., 2008 in Barremian and E. vocontiiDevillez et al., 2016, in Albian (Fig. 3E-H). Other Eryma species found in Late Cretaceous deposits were described from North America (E. americanumRathbun, 1923, E. flectumRathbun, 1926, E. stantoniRathbun, 1935) and Lebanon (E. cretaceumRoger, 1946). After careful examination of the figures presented by Rathbun (1923, 1926, 1935) and according to Förster (1966: 125), we consider that the American species, only represented by fragments of specimens, are not representatives of Erymidae. As for the Lebanese species, we follow Charbonnier et al. (in press) with the placement in PustulinaQuenstedt, 1857.

Addition of ProtoclytiopsisBirshtein, 1958 to the synonymy of Eryma extends the stratigraphic range of the genus Eryma to the Late Permian (Changhsingian). Previously, Bachmayer and Malzahn (1983) had doubtfully assigned to Erymastacus a poorly preserved P1 chela from the Late Permian of Germany: Erymastacus (?) hoerstgenensis. As Schweitzer et al. (2010) regarded Erymastacus as synonymous with Eryma, they listed the species within representatives of Eryma. Later, Karasawa et al. (2013) removed the species from Erymidae because of its angular propodus and the shape of its fingers. Examination of the figure of the holotype of E. hoerstgenensis leads us to support the opinion of Karasawa et al. (2013).

Thus, Eryma antiquum (Birshtein, 1958) nov. comb., is the only erymid lobster reported in the Paleozoic and the oldest known representative of Eryma and Erymidae.

Finally, it is worth pointing out that the gap of almost 50 million years without reported Eryma between its two oldest representatives (E. antiquum nov. comb. from the Changhsingian and E. sinemuriana nov. comb.from the Sinemurian) shows the lack of fossils in the record of Eryma during the Trias.

Conclusions

A review of erymid genera leads us to consider ProtoclytiopsisBirshtein, 1958, and GaliciaGarassino and Krobicki, 2002, as junior synonyms of Eryma. Moreover, Galicia marianaeGarassino and Krobicki, 2002, is herein considered to be a junior synonym of Eryma ventrosum (Meyer, 1835) on the basis of its carapace groove pattern and ornamentation. These additions into the synonymy of Eryma result in the extension of its stratigraphic range to Changhsingian (Late Permian). Thus, Eryma survived the Permian-Triassic mass extinction, commonly considered as the biggest of all major extinctions (Raup, 1979; Erwin, 1994; Stanley and Yang, 1994), with the occurrence of Enoploclytia gardnerae (Rathbun, 1935) in Mexican Paleocene deposits (Vega et al., 2007), it is worth noting that Erymidae also survived the Cretaceous-Paleogene mass extinction. However, E. gardnerae is the last occurrence of erymid lobsters in the fossil record so they are considered to be extinct. The presence of an intercalated plate, an exclusive character of Erymidae, was recognized by Schram and Dixon (2004) in members of the extant family Enoplometopidae (Saint Laurent, 1988). This observation leads us to question the link between Erymidae and Enoplometopidae.

    Abbreviations
  • GPANM

    Naturhistorisches Museum, Wien (Austria)

  • KSGR

    Department of Stratigraphy and regional geology of the University of mining and metallurgy, Kraków (Poland)

  • MNHN.F

    Muséum national d'Histoire naturelle, Paris (France)

  • MSNM

    Museo di Storia Naturale di Milano (Italy)

  • NHMUK

    Natural History Museum, London (United Kingdom)

  • OSUG

    Observatoire des Sciences de l'Univers, Grenoble (France)

  • PIN

    Palaeontological Institute, Moscow (Russia)

  • SM

    Sedgwyck Museum, Cambridge (United Kingdom)

  • ω

    Attachment site of mandibular muscle

  • χ

    Attachment site of adductor testis muscle

  • P1-P3

    Pereiopods 1 to 3

Acknowledgments

We wish to thank Claire Mellish (Natural History Museum, London, United Kingdom), Fabienne Giraud-Guillot (Université Joseph Fournier − Institut Dolomieu, Grenoble, France), Liz Harper and Matt Riley (Sedgwyck Museum, Cambridge, United Kingdom) for the access to the collections of their respective institutions.

We are also grateful to A. Garassino (Museo di Storia Naturale di Milano, Italy), G. Schweigert (Staatliches Museum für Naturkunde, Stuttgart, Germany), F. Schram (University of Washington Seattle, Seattle, USA) and R. Feldmann (Kent State University, Kent, USA) who provided the pictures the type material of the erymid lobsters from Italy, Germany and Russia.

We also really thank Jean-Michel Pacaud (MNHN, Paris, France), for helping to solve some nomenclatural complications and Lilian Cazes (MNHN, Paris, France) for the photographs of some specimens.

Finally, we reiterate our thanks to Professor Rodney Feldmann for his constructive comments which greatly improved our original manuscript.

This paper is a contribution to the UMR 7207 CR2P, CNRS-MNHN-UPMC and to the “Département Histoire de la Terre” (Muséum national d'Histoire naturelle, Paris).

Cite this article as: Devillez J, Charbonnier S. 2017. The genus Eryma Meyer, 1840 (Crustacea: Decapoda: Erymidae): new synonyms, systematic and stratigraphic implications, Bull. Soc. géol. Fr. 188: 15.