# Review of the Late Cretaceous erymid lobsters (Crustacea: Decapoda) from the Western Tethys

Julien Devillez, Sylvain Charbonnier, Martina Kocová Veselská, Jean-Philippe Pezy

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## To cite this version:

Julien Devillez, Sylvain Charbonnier, Martina Kocová Veselská, Jean-Philippe Pezy. Review of the Late Cretaceous erymid lobsters (Crustacea: Decapoda) from the Western Tethys. Proceedings of the Geologists' Association, 2017, 128 (5-6), pp.779 - 797. 10.1016/j.pgeola.2017.08.006. hal-01689615

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# Review of the Late Cretaceous erymid lobsters (Crustacea: Decapoda) from the Western Tethys

Julien Devillez (corresponding author)

Muséum national d'Histoire naturelle, Paris
Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements (CR2P, UMR 7207), Sorbonne Universités, MNHN, UPMC, CNRS, 57 rue Cuvier F-75005 Paris (France)
julien.devillez@edu.mnhn.fr

Sylvain Charbonnier

Muséum national d'Histoire naturelle, Paris
Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements (CR2P, UMR 7207), Sorbonne Universités, MNHN, UPMC, CNRS, 57 rue Cuvier F-75005 Paris (France)
sylvain.charbonnier@mnhn.fr

Martina Kočová Veselská

Institute of Geology and Palaeontology
Charles University, Albertov 6, 128 43
Prague 2, Czech Republic
Veselskamartina@gmail.com

Jean-Philippe Pezy

Normandie Univ, UNICAEN, CNRS, M2C, 14000 CAEN, France
jean-philippe.pezy@unicaen.fr

## ABSTRACT

The Erymidae Van Straelen, 1925 had a worldwide distribution during the Jurassic and the Cretaceous. Twenty-seven species among four genera were reported in the Late Cretaceous, but confusions in diagnoses have led to wrong generic identifications of many erymid species. In the light of recent clarifications of the diagnoses of erymid genera, the Late Cretaceous species are here reviewed. We recognize thirteen of them including three European species which benefit of new descriptions: *Palaeastacus sussexiensis* (Mantell, 1824), *Enoploclytia leachi* (Mantell, 1822) and *Enoploclytia seitzi* Glaessner, 1932. Furthermore, a comparison of the Late Cretaceous erymid fauna with that from the Early Cretaceous emphasizes some changes. A decrease in diversity on genus and species level is noted with the absence of *Eryma* Meyer, 1840 in the Late Cretaceous and the predominance of *Enoploclytia* M'Coy, 1849 (including almost half of the species). Despite the scarcity of the fossil record, our palaeobiogeographic interpretations led us to propose hypotheses about the provenance of North American populations in the end of Early Cretaceous and in Late Cretaceous: (1) their supposed absence since the Callovian (Jurassic) is due to collecting and/or reporting bias or (2) they were the result of migrations of European populations. Moreover, the report of a chela assigned to *Stenodactylina cf. armata* (Secrétan, 1964) in France confirms the existence of migration routes between European and Malagasy faunas during the Late Cretaceous.

## KEY WORDS

Cretaceous, Erymidae, lobster, Mesozoic, paleobiogeography, Western Europe.
**1. Introduction**

Erymid lobsters are typical Mesozoic decapod crustaceans reported from Europe (e.g., Mantell 1833; Bell 1850, 1863; Oppel 1861, 1862; Lahusen 1894; Van Straelen 1925; Beurlen 1928; Glaessner 1931; Reuss 1854; Bachmayer 1959; Förster & Rieber 1982; Garassino 1996; Jagt & Fraaije 2002; Garassino & Krobicki 2002; Bravi *et al.* 2014), in the Middle East (Roger 1946; Förster & Seyed-Emani 1982; Garassino 1994; Charbonnier *et al.* 2017), in Africa (Beurlen 1933; Joleaud & Hsu 1935; Secrétan 1964, 1984; Charbonnier *et al.* 2012), in North America (Rathbun 1923, 1926; Stenzel 1945; Feldmann & McPherson 1980; Aguirre-Urreta & Ramos 1981; Aguirre-Urreta 1982, 1989; Schweitzer & Feldmann 2001; Feldmann & Titus 2006; Feldmann & Haggart 2007; Vega *et al.* 2013; J. Luque, pers. com.), in Japan (Karasawa *et al.* 2008; Kato *et al.* 2010), in Australia (Woodward 1877; Etheridge Jr. 1914; Woods 1957), and in Antarctica (Taylor 1979; Aguirre-Urreta 1989). Despite this worldwide distribution, the fossil record of erymid lobsters remains sparse and fragmentary partially due to the low fossilization potential of decapod crustaceans (scavenging, decortication, disarticulation, carapace fragmentation), and bias related to collection and reporting (Plotnick 1986; Bishop 1986; Plotnick *et al.* 1988; Tshudy *et al.* 1989; Müller *et al.* 2000; Stempien 2005; Mutel *et al.* 2008; Krause *et al.* 2011; Hyžný & Klompmaker 2015; Klompmaker *et al.* 2017). Currently, twenty-seven species are known in the Late Cretaceous. However, until recently, the diagnoses of erymid genera, mainly based on carapace groove pattern and morphology of the chelae of the first chelipeds (Van Straelen 1925; Förster 1966; Glaessner 1969), remained confused. Some recent studies tried to clarify the concepts of erymid genera and proposed new diagnoses (Hyžný *et al.* 2015; Devillez *et al.* 2016). Thus, considering these revisions, the present contribution aims to give a new look on Late Cretaceous erymid fauna with a focus on European taxa, which benefit of new descriptions, and to compare with Early Cretaceous erymid fauna.

**2. Material and methods**

The studied material includes 272 specimens from the Cretaceous (Table 1). They were mostly studied using a CETI Belgium MEDO binocular microscope. Most of them consist of isolated carapaces and first chelipeds. Almost all European specimens examined in this work are assigned to *Enoploclytia leachi* (Mantell, 1822) or *Palaeastacus sussexiensis* (Mantell, 1824). These species are represented by numerous fossils in the Late Cretaceous chalk formations from southeast United Kingdom and only *E. leachi* is abundantly present in the Bohemian Cretaceous Basin (Czech Republic). In both areas, the fossils are relatively undeformed and few specimens are preserved with some anatomical parts which are uncommonly preserved (pleon, antennulae, antennae, third maxillipeds).
## 2.1 Institutional abbreviations

<table><tr><td>BAS</td><td>British Antarctic Survey, Cambridge (United Kingdom).</td></tr><tr><td>BGR</td><td>Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin (Germany).</td></tr><tr><td>BM</td><td>Booth Museum of Natural History, Brighton (United Kingdom).</td></tr><tr><td>MNHN</td><td>Muséum national d'Histoire naturelle, Paris (France).</td></tr><tr><td>MUZ</td><td>Museo de Múzquiz, Coahuila (Mexico).</td></tr><tr><td>NHMUK</td><td>Natural History Museum, London (United Kingdom).</td></tr><tr><td>NM</td><td>Národní Muzeum, Prague (Czech Republic).</td></tr><tr><td>STI</td><td>Steinmann Institut für Geologie, Mineralogie und Paläontologie, Bonn (Germany).</td></tr><tr><td>SM</td><td>Sedgwick Museum of Earth Sciences, Cambridge (United Kingdom).</td></tr><tr><td>NPL</td><td>Non-Vertebrate Paleontology Laboratory, Jackson School of Geosciences, University of Texas, Austin (United States).</td></tr><tr><td>USNM</td><td>United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. (United States).</td></tr></table>

## 2.2 Anatomical abbreviations

a1 Antennulae.

a2 Antennae.

CL Carapace length (excluding rostrum).

CH Carapace height.

Mxp3 Third maxilliped.

ω Attachment site of mandibular muscle.

χ Attachment site of adductor testis muscle.

P1-P5 Pereiopods 1 to 5.

s1-s6 Pleonal somites 1 to 6.

# 3. Systematic Palaeontology

MALACOSTRACA Latreille, 1802

DECAPODA Latreille, 1802

ERYMIDA sensu Schram & Dixon, 2004

Superfamily ERYMOIDEA Van Straelen, 1925

Family ERYMIDAE Van Straelen, 1925

INCLUDED GENERA. — *Enoploclytia* M'Coy, 1849; *Eryma* Meyer, 1840; *Palaeastacus* Bell, 1850; *Pustulina* Quenstedt, 1857; *Stenodactylina* Beurlen, 1928; *Tethysastacus* Devillez, Charbonnier, Hyžný & Leroy, 2016.

PRELIMINARY REMARKS. — Currently, the phylogenetic position of Erymidae is debated. Indeed, many authors have included this family within the Astacidea Latreille, 1802 (Van Straelen 1925; Glaessner 1969; Aguirre-Urreta 1989; Schweigert et al. 2000; Garassino & Krobicki 2002; Crônier & Courville 2004; Garassino & Schweigert 2006; Feldmann & Titus 2006; Schweigert 2013; Charbonnier et al. 2013) while recent contributions recognized this family to belong to Glypheidea Zittel, 1885 (De Grave et al. 2009; Schweitzer et al. 2010; Wahle et al. 2012; Karasawa et al. 2013; Feldmann et
al. 2015). However, results of phylogenetic analysis by Charbonnier et al. (2015) suggest to remove Erymidae from Glypheidea. So, following Hyžný et al. (2015), we consider the systematic position of the Erymidae as uncertain. Then, according to Schram & Dixon (2004), we include the Erymidae in the separate clade Erymida, without listing taxonomic rank beyond superfamily.

## Genus *Eryma* Meyer, 1840

**TYPE SPECIES.** — *Macrourites modestiformis* Schlotheim, 1822, by subsequent designation of Glaessner (1929).

**DIAGNOSIS.** — An emended diagnosis was proposed by Devillez et al. (2016: 518) and does not need to be repeated here.

**REMARKS.** — Among the numerous species of *Eryma* listed by Schweitzer et al. (2010), four are known in the Late Cretaceous. Three of them are from the United States: *Eryma americanum* Rathbun, 1923, *Eryma flectum* Rathbun, 1926, and *Eryma stantoni* Rathbun, 1935. These species are based on fragments of P1 chelae and carapaces and, after careful examination of figures presented by Rathbun (1923, 1926, 1935), we follow Förster (1966: 125) and Devillez et al. (2016) who consider that *E. americanum*, *E. flectum* and *E. stantoni* do not belong to Erymidae. The last listed species is *Eryma cretaceum* Roger, 1946 from the Cenomanian of Lebanon. Based on the carapace groove pattern and the morphology of the P1 chelae, Charbonnier et al. (2017) transfered this species to *Pustulina* Quenstedt, 1857. We concur with this transfer.

In conclusion, to our current knowledge, there is no representative of *Eryma* in the Late Cretaceous and the last occurrence of this genus is *Eryma vocontii* Devillez, Charbonnier, Hyžný & Leroy, 2016 from the Albian (Early Cretaceous) of southeast France.

## Genus *Stenodactylina* Beurlen, 1928

**TYPE SPECIES.** — *Stenodactylina liasina* Beurlen, 1928, by monotypy.

**DIAGNOSIS.** — An emended diagnosis was given by Devillez et al. (2016: 522) and does not need to be repeated here.

**REMARKS.** — *Stenodactylina* includes thirteen species mostly from the Jurassic (Devillez et al. 2016). Only two species are reported from the Late Cretaceous: *Stenodactylina triglypta* (Stenzel, 1945) (Coniacian, Texas, United States) and *S. armata* (Secrétan, 1964) (Campanian, Madagascar). These species are known from only a few fossils. *Stenodactylina armata* is known from four incomplete P1 chelae initially assigned to *Enoploclytia* M'Coy, 1849 (Secrétan 1964; Förster 1966; Schweitzer et al. 2010; Charbonnier et al. 2012). These P1 chelae exhibit a strong, rectangular propodus, slightly compressed dorso-ventrally, with a slight concavity of
the outer margin at the basis of the index and with a strongly inflated dactylar bulge (Fig. 1A-B). As pointed out by Devillez *et al.* (2016), these characteristics fit the general morphology of the P1 chelae of the representatives of *Stenodactylina* and justify the assignation of the species to this genus. A fragment of P1 cheliped exhibiting a morphology very similar (propodus shape, dactylar bulge, finger bases, surface ornamentation) to that of *S. armata* (Fig. 1C) was found in Santonian deposits of Cognac (France). So, we identified this specimen as *S. cf. armata*, which supposes possible palaeogeographic link between the Western and the Southern Tethys.

*Stenodactylina triglypta* is known from two fragments of carapaces, designated by Stenzel (1945) as “syntype 1” (Fig. 1H) and “syntype 2” (Fig. 1D-F), and a fragment of a P1 chela, designated as “syntype 3” (Fig. 1G). They were initially assigned to *Enoploclytia* but the review of Devillez *et al.* (2016) demonstrated that the carapace groove pattern is characteristic of *Stenodactylina*: it does not show a junction between postcervical and branchiocardiac grooves, the postcervical groove is interrupted in hepatic region and the branchiocardiac groove is joined to the posterior extremity of the hepatic groove (Fig. 1F).

Since the publication of Stenzel (1945), no lectotype has been designated. Considering the quality of preservation of the carapace which allows the observation of almost all the carapace grooves, we select herein the specimen TMM-BED 21127, stored in the NPL collections, as lectotype for *S. triglypta* (“syntype 2”; Fig. 1C-E). The two paralectotypes are NPL TMM-BEG 21124 (“syntype 1”; Fig. 1H) and 21125 (“syntype 3”; Fig. 1G).

## Genus *Palaeastacus* Bell, 1850

**TYPE SPECIES.** — *Astacus sussexiensis* Mantell, 1824, by subsequent designation of Glaessner (1929).

**DIAGNOSIS.** — An emended diagnosis was proposed by Devillez *et al.* (2016: 525) and does not need to be repeated here.

**REMARKS.** — Among the numerous species of *Palaeastacus* listed by Schweitzer *et al.* (2010), eight are known from the Late Cretaceous. The type species *Palaeastacus sussexiensis* (Mantell, 1824) (Fig. 2A-F) is redescribed and discussed below. One listed species, *Palaeastacus plauensis* Geinitz, 1875 (Cenomanian, Germany), was previously included into the synonymy of *P. sussexiensis* based upon the shape and ornamentation of the P1 propodus and fingers (Förster 1966; Devillez *et al.* 2016). Two other species are from Europe: (1) *Palaeastacus macrodactylus* Bell, 1850 (Late Cretaceous chalk, United Kingdom) and (2) *P. cenomaniensis* Hée, 1924 (Cenomanian, France). Carefull examination of the figure presented by Bell (1850: pl. 38, fig. 6) shows that *Palaeastacus macrodactylus* exhibits P1 chelae with long slender fingers typical of *Enoploclytia* (see discussion under *Enoploclytia leachi* for more details). *Palaeastacus cenomaniensis* is known from isolated fragments of P1 chelae lost since World War II (Förster 1966). After the description and figures presented by Hée (1924),
the propodus has a square shape with a strongly tuberculate upper side, an almost smooth lower side and thin fingers. P1 chelae of *Palaeastacus* representatives have usually a rectangular or trapezoidal propodus, strongly ornamented on both sides, and wide fingers. Therefore, we do not consider *P. cenomaniensis* to be a representative of *Palaeastacus*. Moreover, the particular shape and ornamentation of the propodus leads us to follow Förster’s opinion by removing this species from the Erymidae.

The four remaining species were found in North America. *Palaeastacus kimzeyi* Rathbun, 1935 (Campanian, Texas, United States; Fig. 3A-C) is known from incomplete P1 chelae and by a fragment of carapace exhibiting the main characteristics of the genus: postcervical groove joined to median inflexion of the hepatic groove, sub-rectangular P1 propodus, slightly compressed dorso-ventrally and wide fingers. Careful examination of the holotype of *Palaeastacus trisulcatus* Schweitzer & Feldmann, 2001 (Cenomanian, California, United States; Fig. 3D-E) reveals an interruption of the branchiocardiac groove in the branchial region, and the presence of a cardiac groove rising from the postcervical groove and joined to the dorsal margin. This groove pattern is characteristic of *Pustulina* Quenstedt, 1857. Hence, we proposed the new combination *Pustulina trisulcata* (Schweitzer & Feldmann, 2001). As pointed out by Förster (1966), *Palaeastacus selmaensis* Rathbun, 1935 (Maastrichtian, Alabama, United States) is known from unrecognizable fragments, so we consider these fragments to be Incertae sedis. The last species, *Palaeastacus ornatus* Whiteaves, 1886 (Campanian, Canada) exhibits a groove pattern and longitudinal, tuberculate carinae along branchial and gastric regions, and dorsal margin, and there is no intercalated plate (Feldmann & McPherson 1980: pl. 2, fig. 1). These characteristics are typical of the Nephropidae Dana, 1852. So, we concur with Mertin (1941) and Feldmann & McPherson (1980) who considered this species to be a representative of *Palaeonephrops* Mertin, 1941.

In conclusion, only two species of *Palaeastacus* are herein recognized in the Late Cretaceous: *P. sussexiensis* and *P. kimzeyi*.

*Palaeastacus sussexiensis* (Mantell, 1824)
(Fig. 2A-F)

*Astacus sussexiensis* Mantell, 1824: pl. 29, fig. 15; 1833: 373, 379, fig. 2; 1844: 238, fig. 4. — Quenstedt 1852: 269; 1885: 411, fig. 129.

*Enoploclytia imagei* M'Coy, 1849: 331; 1854: 136. — Reuss 1854: 3. — Woodward 1877: 9. — Glaessner 1929: 146. — Schweitzer *et al.* 2010: 22.

*Enoploclytia brevimana* M'Coy, 1849: 332; 1854: 137. — Reuss 1854: 3. — Bronn 1851-1852: 352. — Woodward 1877: 9. — Schweitzer *et al.* 2010: 22.

*Palaeastacus dixoni* Bell, 1850: 344, 345, pl. 38, fig. 1-5. — Geinitz 1875: 292. — Schlüter 1879: 602. — Schweitzer *et al.* 2010: 25.

*Hoploparia scabra* Bell, 1863: 28, pl. 7, fig. 3-7. — Glaessner 1929: 221.
*Phlyctisoma granulatum* Bell, 1863: 36, pl. 11, fig. 9-10. — Glaessner 1929: 314. — Monaco & Garassino 2000: 297.

*Palaeastacus plauensis* Geinitz, 1875: 291, pl. 64, fig. 9.

*Astacus leachii* (pars.) — Mantell 1822: 223, pl. 30, fig. 3.

*Glyphea sussexiensis* — Roemer 1841: 105.

*Enoploclytia sussexiensis* — Morris 1854: 108. — Willet 1871: 42-43. — Woodward 1877: 10. — Woodward 1878: 377, pl. 38, fig. 1-4. — Schlüter 1879: 602. — Glaessner 1932: 578 — Van Straelen 1936: 12.

*Palaeastacus sussexiensis* — Glaessner 1929: 200. — Rathbun 1935: 23. — Förster 1966: 133, fig. 24, pl. 17, fig. 6. — Morris 1980: 13. — Aguirre-Urreta & Ramos 1981: 609, fig. 4c. — Morris 1987: 196, pl. 42, fig. 4-5. — Aguirre-Uretta 1989: 510, fig. 8-10. — Wittler 1998: 18, fig. 5. — Garassino & Schweigert 2006: 11. — Schweitzer *et al.* 2010: 26. — Karasawa *et al.* 2013: 79, 102. — Devillez *et al.* 2016: 526, fig. 6, fig. 7A-E.

*Palaeastacus ? plauensis* — Glaessner 1929: 290.

*Enoploclytia dixoni* — Woods 1930: 83, pl. 23, fig. 9-12, pl. 24, fig. 1-3. — Van Straelen 1936: 11.

*Enoploclytia (Palaeastacus) sussexiensis* — Mertin 1941: 161, fig. 4a.

*Enoploclytia (Palaeastacus) imagei* — Roberts 1962: 164.

*Palaeastacus scaber* — Förster 1966: 132, fig. 26, pl. 17, fig. 7-9. — Morris 1980: 13. — Schweitzer & Feldmann 2001: 174. — Garassino & Schweigert 2006: 11. — Schweitzer *et al.* 2010: 25. — Karasawa *et al.* 2013: 102.

*Palaeastacus cf. sussexiensis* — Taylor 1979: 30, fig. 10e, pl. 4 fig. d-f.

*Palaeastacus ? plavensis* — Schweitzer *et al.* 2010: 25.

*Pustulina granulata* — Schweitzer *et al.* 2010: 26.

*Pustulina scabra* — Schweitzer *et al.* 2010: 26.

**TYPE MATERIAL.** — Lectotype NHMUK 5601 (Fig. 2A-B), designated by Devillez *et al.* (2016); 13 paralectotypes NHMUK 5024, 5584, 5586, 5589, 5600, 5602, 5608, 5613, 5618, 5624, 5626, 5629, 10760.

**TYPE LOCALITY.** — Sussex, United Kingdom (precise type locality not indicated on the original labels).

**TYPE AGE.** — Cenomanian, Late Cretaceous.

**DESCRIPTION.** — *Carapace.* Subcylindrical carapace (lectotype: CL= 82 mm, CH= 41 mm); long spiny rostrum; fusiform and tuberculate intercalated plate; ridge on dorsal
margin of branchial region; deep cervical groove, joined to dorsal margin and to
antennal groove; deep antennal groove; short gastro-orbital groove originating as a
slight median inflexion of cervical groove; postcervical and branchiocardiac grooves
subparallel; dorsally deep, sinuous postcervical groove, shallowing ventrally, not joined
to dorsal margin, joined to hepatic groove; narrow branchiocardiac groove, strongly
inclined, shallower than postcervical groove, not joined to dorsal margin, joined to
hepatic groove; concavo-convex, narrow hepatic groove, joined to cervical groove;
inflated ω area; flat χ area; deep inferior groove, joined to hepatic groove.

*Ornamentation of carapace.* Carapace densely covered by rounded tubercles in branchial region and widely spaced coarse tubercles in cardiac, hepatic and cephalic regions; row of coarse tubercles parallel to dorsal margin in cardiac and branchial regions; two oblique rows of coarse tubercles in cardiac region; row of tubercles parallel to the intercalated plate in gastric region.

*Pleon and uropods.* Pleonal somites with terga ornamented by four pairs of strong spines and by rounded, coarse tubercles on the remaining surface (Fig. 2C-D); subtriangular pleura, with pointed ventral margin, ornamented with two strong spines; telson with rounded extremity; telson with longitudinal median ridge flanked by two wide lateral ridges interrupted at telson mid-length; telson covered by coarse rounded tubercles; uropods as long as telson, covered by tubercles; uropodal endopod with longitudinal carina; uropodal exopod with diaeresis.

*Thoracic appendages.* Mxp3 with the inferior side of the first two segments adorned with numerous small spines, the inferior distal extremities of the third segment is adorned by a strong spine; chelate P1; short, trapezoidal, slightly globose P1 propodus, with two divergent rows of spines on ventral and dorsal surfaces; inner margin with strong spines; wide fingers, slightly longer than propodus; occlusal margins with short, spaced teeth; index wider than dactylus; dactylus adorned with spines; spiny P1 carpus and merus (after Devillez *et al.* 2016: 526).

DISTRIBUTION. — *Palaeastacus sussexiensis* is a well-known species represented by numerous fossils, mainly from the Late Cretaceous chalk formations of southeast United Kingdom. It has also a wide stratigraphic range associated to a wide geographic distribution. Indeed, *P. sussexiensis* occurs in (1) the Aptian of Alexander Island (Antarctica; Fig. 2E; Taylor 1979) and Patagonia (Argentina; Aguirre-Urreta 1989), (2) the Albian of England (Bell, 1863; Woods, 1930; Förster 1966), (3) the Cenomanian of England (Fig. 2A, C-D; Bell 1850; Förster 1966; Morris 1987), Germany (Mertin 1941) and France (Van Straelen 1936; Förster 1966), and (4) the Turonian of England (Mantell 1822, 1833, 1844; Morris 1987), Germany (Wittler 1998) and France (Fig. 2F).

*Palaeastacus kimzeyi* Rathbun, 1935
(Fig. 3A-C)
*Palaeastacus kimzeyi* Rathbun, 1935: 23, pl. 2, figs. 15-21. — Förster 1966: 135. — Förster & Rieber 1982: 777. — Schweitzer & Feldmann 2001: 174, 176. — Garasino & Schweigert 2006: 11. — Schweitzer *et al.* 2010: 25.

*Enoploclytia kimzeyi* — Stenzel 1945: 414. — Secrétan 1964: 94, 95. — Taylor 1979: 25. — Feldmann 1981: tab. 1.

**TYPE MATERIAL.** — Holotype USNM PAL 73797 (Fig. 3A-B); 1 paratype USNM PAL 73841, and 9 paratypes USNM PAL 335146.

**TYPE LOCALITY.** — Farmersville, Texas, United States.

**TYPE AGE.** — Campanian, Late Cretaceous.

**DESCRIPTION.** — *Carapace*. Deep cervical groove, joined to antennal groove; deep antennal groove; short gastro-orbital groove; postcervical and branchiocardiac grooves subparallel; slightly curved postcervical groove, joined to hepatic groove; slightly curved branchiocardiac groove, joined to hepatic groove; concavo-convex hepatic groove, joined to cervical groove; inflated ω area; flat χ area; inferior groove joined to hepatic groove; carapace covered by small rounded tubercles, presence of coarse tubercles in cephalic and hepatic regions.

*Thoracic appendages*. Chelate P1; subrectangular, slightly globose P1 propodus, covered by rounded tubercles; both sides of P1 propodus with longitudinal depression parallel to the inner margin; strongly compressed inner margin; inflated, narrow dactylar bulge; wide fingers, covered by tubercles.

**DISCUSSION.** — Comparisons between *P. kimzeyi* and *P. sussexiensis* are only possible at level of the P1 propodus because the carapace of the first species is too fragmentary. The P1 propodus of *P. sussexiensis* shows no longitudinal depression along the inner margin on both sides. Besides, there are rows of tubercles on the dorsal and ventral surfaces of the P1 propodus (no row in *P. kimzeyi*).

## Genus *Enoploclytia* M'Coy, 1849
(Fig. 4A-B)

**TYPE SPECIES.** — *Astacus leachii* Mantell, 1822, by original designation.

**EMENDED DIAGNOSIS.** — An emended diagnosis was proposed by Devillez *et al.* (2016: 530). We add that the P1 merus exhibits strong, prominent distal process at outer side of the ventral extremity (Figs 5A-B, 6C, 7C).

**REMARKS.** — Among the numerous species of *Enoploclytia* listed by Schweitzer *et al.* (2010), twelve are known from the Late Cretaceous. Half of these species were found in Western Europe: *Enoploclytia brevimana* M'Coy, 1849 (Cenomanian, United
Kingdom, *Enoploclytia granulicauda* Schlüter, 1879 (Campanian, Germany), *Enoploclytia imagei* M'Coy, 1849 (Cenomanian, United Kingdom), *Enoploclytia leachi* (Mantell, 1822) (Cenomanian to Campanian, Western and Central Europe), *Enoploclytia paucispina* Schlüter, 1868 (Santonian, Germany) and *Enoploclytia seitzi* Glaessner, 1932 (Albian, Germany; Cenomanian, United Kingdom). *Enoploclytia leachi* and *E. seitzi* are described and discussed below and *E. granulicauda* is herein considered as a synonym of *E. leachi* based on its carapace groove pattern and ornamentation (see discussion about *E. leachi*). *Enoploclytia brevimana* and *E. imagei* exhibit morphological characters (carapace groove pattern, ornamentation, P1 chelae) typical of *Palaeastacus* and, according to Förster (1966) and Devillez *et al.* (2016), these species are considered to be synonyms with *P. sussexiensis*. *Enoploclytia paucispina* is known from a finely tuberculate fragment of carapace and from a P1 chela exhibiting a compressed propodus and wide, curved fingers armed with small, flattened teeth. Given that *Enoploclytia* has a globose P1 propodus bearing slender, straight fingers armed with long sharp teeth, we follow Mertin (1941) and Förster (1966) and do not consider *E. paucispina* to be an erymid lobster but a representative of *Oncopareia* Bosquet, 1854.

Two species are reported from the Campanian of Madagascar. *Enoploclytia collignoni* Secrétan, 1964 exhibits the typical carapace groove pattern and P1 chelae of the genus: postcervical and branchiocardiac grooves not joined; sinuous postcervical groove, joined to hepatic groove; reduced branchiocardiac groove; globose P1 propodus; slender and straight P1 fingers. *Enoploclytia armata* Secrétan, 1964 is known from some P1 chelae exhibiting the typical characteristics of *Stenodactylina* (see remarks about *Stenodactylina*). Thus, we follow Devillez *et al.* (2016), and we consider this species to be a representative of *Stenodactylina*.

The four remaining species were found in North America. *Enoploclytia sculpta* Rathbun, 1926 (Maastrichtian, Tennessee, United States) is known from a specimen showing a part of the pleon and a poorly preserved P1. The terga lack of relief and are densely covered by small pits (Rathbun 1926: pl. 76 fig. 3). Representatives of *Enoploclytia* usually have a pleon with longitudinal bulges on the pleural basis and a coarse, wide-spaced ornamentation. Thus, following Stenzel (1945) and Förster (1966), we conclude that this species is not a representative of *Enoploclytia* and probably not an erymid lobster. *Enoploclytia triglypta* Stenzel, 1945 (Coniacian, Texas, United States) exhibits a groove pattern diagnostic for *Stenodactylina* as discussed by Devillez *et al.* (2016) (see remarks about *Stenodactylina*). *Enoploclytia tumimana* Rathbun, 1935 (Maastrichtian, Alabama, United States) is known from several fragments of carapace and P1 (including fragments of merus, propodus and fingers). Careful examination of high resolution pictures of the holotype USNM PAL 73799 (Rathbun 1935: pl. 1, figs 10-11) leads us to confirm the typical characteristics of *Enoploclytia*: well-marked gastro-orbital groove; postcervical and branchiocardiac grooves not joined; reduced branchiocardiac groove; subrectangular, globose P1 propodus; straight, slender fingers. *Enoploclytia minor* (Woodward, 1900) (Maastrichtian, Canada) is known from a poorly preserved specimen showing the carapace and at least the P1-P3 (Feldmann & McPherson 1980: pl. 1, figs. 1-2). The carapace groove pattern is not recognizable but
the globose P1 propodus, bearing long slender fingers, allows us to maintain this species
into the genus.

Since the publication of the systematic list of Schweitzer *et al.* (2010), *Enoploclytia tepeyacensis* Vega, Garassino & Jaime, 2013 was described from Mexican Campanian deposits. This species is known from two carapaces exhibiting a groove pattern typical of *Enoploclytia* (well-marked gastro-orbital groove; postcervical and branchiocardiac grooves not joined; sinuous postcervical groove; reduced branchiocardiac groove). Some unidentified fragments attributed to *Enoploclytia* have also been reported from Campanian rocks of Seymour Island (Antarctica; Feldmann 1984).

In conclusion, with six species (*E. collignoni*, *E. leachi*, *E. minor*, *E. seitzi*, *E. tepeyacensis* and *E. tumimanus*), the genus *Enoploclytia*, has reached higher diversity during the Late Cretaceous (supposedly absent before the Cretaceous and four species known in the Early Cretaceous: *E. augustobonae* Devillez, Charbonnier, Hyžný & Leroy, 2016, *E. gigantea* Devillez, Charbonnier, Hyžný & Leroy, 2016, *E. seitzi*, *E. wintoni* Stenzel, 1945).

*Enoploclytia leachi* (Mantell, 1822)

(Figs 4C-J, 5, 6)

*Astacus leachi* Mantell, 1822: 221, pl. 29, fig. 1, 2, 5, pl. 30, figs. 1-2 (non 3), pl. 31, figs. 1-4; 1824: 11, pl. 29, fig. 15; 1833: 122, fig. 1. — Geinitz 1839: 14, pl. 7, fig. 4, 39, pl. 9, fig. 1. — Mantell 1844: 536, pl. 116, figs. 1-3. — Quenstedt 1852: 269, pl. 20, fig. 11; 1867: 321, pl. 25, fig. 11; 1885: 411, pl. 32, fig. 11. — Etheridge Jr. 1914: 272, pl. 24, fig. 2.

*Palaeastacus dixoni* Bell, 1850 (pars.): 344, pl. 38, fig. 5.

*Palaeastacus macrodactylus* Bell, 1850: 345, pl. 38, fig. 6.

*Enoploclytia heterodon* Schlüter, 1862: 724, pl. 11, figs. 2-4; 1879: 600. — Glaessner 1929: 146. — Secrétan 1964: 87.

*Enoploclytia granulicauda* Schlüter, 1879: 599, pl. 14, fig. 1-4. nov. syn. — Glaessner 1929: 146. — Van Straelen 1936: 11. — Förster 1966: 150, pl. 19, fig. 6. — Taylor 1979: 25, 29. — Jagt & Fraaije 2002: 92. — Schweitzer *et al.* 2010: 22.

*Glyphea leachi* — Roemer 1841: 105. — Schlüter 1862: 703.

*Klytia leachi* — Reuss 1845: 14, 103, pl. 6, figs. 1-6, pl. 42, fig. 3. — Geinitz 1849: 96. — Bronn 1852: 352. — Gümbel 1868: 752, 762. — Gosselet 1881: 267.

*Clytia leachi* — Bronn 1849: 578. — Reuss 1854: 1, pl. 1-5.

*Enoploclytia leachi* — M'Coy 1849: 331; 1854: 136. — Morris 1854: 108. — Schlüter 1862: 725. — Schlüter & von der Marck 1868: 295. — Willet 1871: 43. — Geinitz 1875: 292. — Woodward 1877: 10; 1878: 378, pl. 38, figs. 5-6. — Schlüter 1879: 602. — Zittel 1885: 694. — Fritsch & Kafka 1887: 27, figs. 48-52. — Fritsch 1877: 145; 1889: 91; 1893: 107. — Leonhard 1897: 63. — Wanderer 1908: 23. — Rathbun 1926: 188. — Glaessner 1929: 146. — Woods 1930: 85, pl. 24, fig. 4, pl. 25, fig. 1. — Glaessner 1932: 577. — Joleaud & Hsu 1935: 100. — Rathbun 1935: 19. — Van Straelen 1936: 11. — Secrétan
1964: 87. — Förster 1966: 148, pl. 19, figs. 4-5, 7-8 (non 3). — Taylor 1979: 24. — Morris 1980: 6; 1987: 196, pl. 42, fig. 3. — Aguirre-Urreta 1989: 516. — Wittler 1998: 17, figs. 3-4, 6. — Jagt & Fraaije 2002: 91, pl. 1, figs. a-c. — Vega *et al.* 2007: 1434. — Veselská 2010: 174, fig. 3. — Schweitzer *et al.* 2010: 22. — Karasawa *et al.* 2013: tab. 1.

*Hoploparia* sp. — Hallez 1869: 297, figs. 1-4.

*Xanthopsis leachi* — Gosselet 1883: pl. 22, fig. 3.

*Enoploclytia (Enoploclytia) leachi* var. *granulicauda* — Mertin 1941: 20, 242, figs. 5f-h, pl. 1, fig. 8.

*Enoploclytia (Enoploclytia) leachi* — Mertin 1941: 162, 242, figs. 4c, 5a-e, pl. 1, figs. 1, 5-7 (non 2-4, 8).

**TYPE MATERIAL.** — Lectotype NHMUK 5636 (Mantell coll.) (Fig. 4C), designated by Förster (1966); two paralectotypes NHMUK 5623, In.61020 (Fig. 4D).

**TYPE LOCALITY.** — Sussex, United Kingdom (precise type locality not indicated on the original labels).

**TYPE AGE.** — Turonian, Late Cretaceous.

**DESCRIPTION.** — *Carapace.* Subcylindrical carapace; spiny rostrum; fusiform and tuberculate intercalated plate; deep, wide cervical groove, joined to dorsal margin and to antennal groove; deep antennal groove; long gastro-orbital groove originating as a median inflexion of cervical groove, with two short and divergent branches, delimiting two gastro-orbital lobes (flat upper lobe, slightly inflated lower lobe); postcervical and branchiocardiac grooves subparallel; deep, sinuous postcervical groove, joined to dorsal margin and to hepatic groove, with ventral extension at carapace mid-height, deepening towards its junction to hepatic groove; narrow, deep branchiocardiac groove, joined to dorsal margin and not to postcervical groove, interrupted at almost carapace mid-height; concavo-convex hepatic groove, joined to cervical groove; inflated ω and χ areas; wide, deep inferior groove, joined to hepatic groove.

*Ornamentation of carapace.* Heterogeneous ornamentation; carapace entirely and densely covered by small tubercles; two oblique rows of coarse tubercles in cardiac region; row of coarse tubercles parallel to intercalated plate in gastric region; longitudinal row of coarse tubercles in gastric region; antennal row of coarse tubercles in antennal region preceded by an antennal spine.

*Cephalic appendages.* Strong a1 peduncle, with three segments, last segment (basipodite) articulated with two flagella, flagella composed of numerous short cylindrical articles; a2 with one long flagellum composed of numerous short cylindrical articles; stout scaphocerite.

*Pleon and uropods.* Somites with terga homogeneously ornamented by spaced tubercles; longitudinal bulge at the basis of the pleura; s2 with an elongated pleura, anteriorly subrectangular and posteriorly subtriangular; s3-s6 exhibiting subtriangular pleura, with sharply pointed ventral margins; telson with rounded extremity; telson with four longitudinal, sub-parallel carina; telson with a row of tubercles on each carina, a
strong tubercles adorn the proximal part of the two central carina; uropods as long as telson; uropodal endopod with longitudinal carina and irregularly ornamented by fine tubercles; uropodal exopod with longitudinal carina and diaeresis.

*Thoracic appendages.* Mxp3 partially known (Fig. 4E), the inferior distal extremities of the segments are adorned by a strong spine; P1-P3 chelate; massive globose P1 propodus, densely covered by small tubercles and with spaced coarse tubercles organized in longitudinal rows on dorsal and ventral surfaces; inner margin with short conical spines; narrow, inflated dactylar bulge; long slender P1 fingers, straight in dorsal view, slightly curved downward, equal in length; occlusal margins armed with slender sharp teeth; subtriangular P1 carpus, densely covered by small tubercles and some coarse tubercles irregularly arranged; upper margin with short conical spines; P1 merus covered by small tubercles; strong, prominent distal process at extern side of the ventral extremity of P1 merus (Figs 5A, 6C); short conical spines on the upper margin of P1 merus; P2-P3 with small chelae; P2-P5 surface almost smooth.

**COMPARISONS.** — The Late Cretaceous deposits of United Kingdom and Germany yielded several species of erymid lobsters which are here considered as junior synonyms of *Enoploclytia leachi*. An almost complete specimen of *E. leachi* was assigned to *Palaeastacus dixoni* by Bell (1850: pl. 38, fig. 5) although it exhibits main characteristics of the previous species: (1) wide cervical groove; (2) postcervical and branchiocardiac grooves not joined; (3) branchiocardiac groove interrupted near the carapace mid-height; (4) dense ornamentation made of small tubercles in branchial region; (5) two oblique rows of coarse tubercles in cardiac region; (6) s3-s6 with subtriangular pleura, exhibiting sharply pointed ventral margins and (7) P1 chelae with long, slender fingers, armed with teeth on their occlusal margins. Bell (1850: pl. 38, fig. 6) also described *Palaeastacus macrodactylus* based on a well-preserved pair of P1 chelae showing globose propodi, adorned with longitudinal rows of spiny tubercles on inner margins and on dorsal and ventral surfaces, the fingers are long and slender and have numerous teeth on their occlusal margins. Identical characteristics are known on the P1 chelae of *E. leachi*. Thus, we follow Förster (1966) and the synonymy between *P. macrodactylus* and *E. leachi* is maintained here.

Later, Schlüter (1862) described two species from Westphalia (Germany). The material of *Enoploclytia heterodon* Schlüter, 1862 (Coniacian), a carapace and a P1 chela, is lost since World War II (Förster 1966) but the carapace has been figured by Mertin (1941: pl. 1, fig. 5; Fig. 4G-H). It exhibits the typical groove pattern and the same ornamentation as in *E. leachi* (postcervical groove not joined to branchiocardiac groove; branchiocardiac groove interrupted near the carapace mid-height; carapace densely covered by tubercles; coarse tubercles arranged in rows in cephalic and cardiac regions; P1 chela with subrectangular propodus bearing long, slender fingers, numerous teeth on occlusal margins). Hence, according to Mertin (1941) and Förster (1966), these similarities justify the placement of *E. heterodon* into the synonymy of *E. leachi*. *Enoploclytia granulicauda* Schlüter, 1879 (Campanian) is based on a specimen showing an almost complete carapace, P1 chelae partially preserved and a part of the pleon (s1 and s2; Fig. 4I-J). This fossil clearly exhibits the typical groove pattern and
ornamentation of *E. leachi*: postcervical groove not joined to branchiocardiac groove; branchiocardiac groove interrupted near the carapace mid-height; carapace densely covered by small tubercles; coarse tubercles arranged in rows in cephalic and cardiac regions; P1 chela with subrectangular propodus; propodus covered with numerous small tubercles and coarse tubercles arranged in longitudinal rows; slender fingers. In conclusion, we consider *E. granulicauda* as a junior synomym of *E. leachi*.

*Enoploclytia leachi* is distinct from the five other Late Cretaceous species by its particular ornamentation: numerous fine tubercles and rows of coarse ones in cephalic and cardiac regions. Indeed, *Enoploclytia collignoni* and *E. seitzi* have a dense, coarse ornamentation made of tubercles and depressions, which gives a scabrous aspect to their respective carapaces (Charbonnier *et al.* 2012: fig. 7A-B; Fig. 7). *Enoploclytia tumimana* exhibits a heterogeneous carapace ornamentation made of coarse tubercles and numerous small pits; *Enoploclytia tepeyacensis* has coarser tubercles than *E. leachi* in branchial region; *Enoploclytia minor* has a P1 propodus covered by coarse tubercles (small tubercles and longitudinal rows of coarse ones in *E. leachi*). Furthermore, some other characteristics support the distinction between *E. leachi* and the other Late Cretaceous species: *E. collignoni* has a strongly prominent χ area (less inflated in *E. leachi*); the dorsal part of the postcervical groove of *E. tepeyacensis* is strongly inclined and almost horizontal at the level of the gastro-orbital groove (postcervical groove far less inclined in *E. leachi*); *E. seitzi* has a shallow and narrow branchiocardiac groove, a strongly elongated P1 propodus in comparison to its height (P1 propodus not so long in comparison to its height in *E. leachi*).

DISTRIBUTION. — *Enoploclytia leachi* is known from numerous fossils found in Western and Central Europe from the Cenomanian to the Campanian, especially in the Bohemian Cretaceous Basin of the Czech Republic. Indeed, *E. leachi* occurs (1) in the Cenomanian — Santonian of France (Fig. 5A-D; Hallez 1869; Van Straelen 1936; Förster 1966), (2) in the Turonian — Campanian of United Kingdom (Figs 4C-D, 5E; Mantell 1822, 1824, 1833, 1844; M'Coy 1849, 1854; Woodward 1878; Woods 1930; Förster 1966; Morris 1987), (3) in the Campanian of Belgium (Jagt & Fraaije 2002), (4) in the Cenomanian — Campanian of Germany (Fig. 4G, I; Schlüter 1862, 1879; Wanderer 1908; Förster 1966; Wittler 1998) and (5) in the Turonian — Coniacian of the Czech Republic (Figs 4E, 6; Geinitz 1839; Reuss 1845, 1854; Fritsch & Kafka 1887; Förster 1966, Veselská 2010). *E. leachi* belongs to the most common fossil lobsters found within the whole Bohemian Cretaceous Basin. Carefull examination of specimens from the Prague palaeontological collections led us to identify some fossils figured in Reuss's monography (1854): NM O7125 (Fig. 4E) corresponds to his pl.1, fig.2, NM O4062 (Fig. 6A) corresponds to his pl.2, fig.4 and NM O4063 corresponds to his pl.4, fig.10. Moreover, the specimen NM O4042 (Fig. 6B), one of the most complete and best-preserved from the Czech Republic, corresponds to the specimen 61 listed by Fritsch and Kafka (1887). This specimen was used as a model for their reconstruction of *E. leachi* on their figs 48-49. All the specimens figured here come from the lower-middle Turonian of Bílá Hora (Weissenberg) in Prague. This locality was Fritsch's
standard section from where many fossils were described, like inoceramids, ammonites, echinoids, decapod crustaceans and vertebrates (Fritsch 1872, 1878; Fritsch & Kafka 1887).

*Enoploclytia seitzi* Glaessner, 1932

(Fig. 7A-C)

*Enoploclytia scabrosa* Woodward, 1873: 524. — Woodward 1877: 10 — Glaessner 1929: 148. (nomina nuda).

*Enoploclytia seitzi* Glaessner, 1932: 578, fig. 1. — Secrétan 1964: 94. — Förster 1966: 147, pl. 19, fig. 1-2. — Taylor 1979: 25. — Morris 1980: 6. — Schweitzer *et al.* 2010: 22.

TYPE MATERIAL. — Lectotype BGR X 00266 designated by Förster (1966) (not recovered; Fig. 7A-B); 1 paralectotype BGR X 00267.

TYPE LOCALITY. — Wettensen, Lower Saxony, Germany.

TYPE AGE. — Albian, Early Cretaceous.

DESCRIPTION. — *Carapace*. Subcylindrical carapace (holotype: CL= 37 mm, CH= 16 mm); rostrum and intercalated plate not preserved; deep, sinuous cervical groove, joined to dorsal margin and to antennal groove; deep antennal groove; deep gastro-orbital groove originating as median inflexion of cervical groove; deep postcervical groove, strongly inclined, weakly sinuous, joined to dorsal margin and to hepatic groove, with a short ventral extension at carapace mid-height; narrow branchiocardiac groove, joined to dorsal margin and not to postcervical groove, extending near the carapace mi-height; concavo-convex hepatic groove, joined to cervical groove; inflated ω and χ areas; wide, deep inferior groove, joined to hepatic groove; carapace with scabrous surface, densely covered by coarse tubercles separated by strong depressions.

*Pleon and uropods*. Somites poorly preserved; terga ornamented by rounded tubercles and deep, irregular depressions; subtriangular pleura, with pointed ventral margin, directed backward, ornamented by deep depressions; telson and uropods not preserved

*Thoracic appendages*. Chelate P1; elongated, rectangular, globose P1 propodus, densely covered by coarse tubercles; inflated dactylar bulge; index and dactylus not preserved; elongated P1 carpus, densely covered by coarse tubercles; P1 densely covered by coarse tubercles; strong, prominent distal process at extern side of the ventral extremity of P1 merus (Fig. 7C).

DISCUSSION. — Woodward (1873) named *Enoploclytia scabrosa* based on a specimen found in the Cenomanian chalk of Dover (United Kingdom). However, he did not described nor figured this specimen, so this is a *nomen nudum* clearly identified by Glaessner (1929). Later, Glaessner (1932) described *Enoploclytia seitzi* based on specimens from the Albian of Westphalia (Germany). After carefull examination and
following Förster (1966), we identify the specimen NHMUK 59818, mentioned by
Woodward (1873) (see Fig. 7C), as a specimen of *E. seitzi* based on (1) the strong
inclination of the postcervical groove, (2) the shallow depth and narrow width of the
branchiocardiac groove and (3) the dense ornamentation made of tubercles separated by
depressions which give a scabrous aspect to the carapace. *Enoploclytia seitzi* is distinct
from *E. collignoni* by its slightly prominent χ bulge (strongly prominent in
*E. collignoni*), its homogeneous ornamentation made of tubercles separated by
depressions all over the carapace (heterogeneous in *E. collignoni*: wide depressions in
branchial regions and coarse tubercles and depressions in cephalic, cardiac and hepatic
regions) and its strongly elongated P1 propodus (shorter in *E. collignoni*).

*Enoploclytia seitzi* exhibits a groove pattern and an ornamentation different from *E. tepeyacensis*: (1) the cervical groove is more sinuous, (2) the postcervical groove is dorsally less inclined and less sinuous, (3) the branchiocardiac groove is narrower and shallower, (4) the ornamentation is homogeneous and made of tubercles separated by depressions.

*Enoploclytia seitzi* differs from *E. tumimana* by a higher sinuosity of the cervical groove, a homogeneous ornamentation with wider depressions and a more elongated P1 propodus. The poorly preserved *Enoploclytia minor* reduces the comparison to the P1 propodus: elongated and covered by tubercles and depressions in *E. seitzi* while it is shorter and covered by coarse tubercles in *E. minor*.

Genus *Pustulina* Quenstedt, 1857

TYPE SPECIES. — *Pustulina suevica* Quenstedt, 1857, by monotypy.

DIAGNOSIS. — An emended diagnosis was proposed by Devillez *et al.* (2016: 531) and does not need to be repeated here.

REMARKS. — Among of the species of *Pustulina* listed by Schweitzer *et al.* (2010), only *Pustulina dawsoni* (Woodward, 1900) is known from the Late Campanian – Early Maastrichtian formations of Hornby Island (Canada).

Two other species have been recently assigned to *Pustulina*: Charbonnier *et al.* (2017) transferred *Eryma cretaceum* to *Pustulina* (see remarks about *Eryma*) and we also add *Palaeastacus trisulcatus* to the genus based on its typical groove pattern showing a cardiac groove (see remarks under *Palaeastacus*; Fig. 3D-E).

**4. Comparison between Early and Late Cretaceous erymid faunas**

4.1. Palaeobiodiversity

The modifications in erymid lobsters diversity during the Cretaceous are difficult to
analyze mainly due to the scarcity of the fossil record and sampling bias. After the
review of the Early Cretaceous erymids (Devillez *et al.* 2016) and the present work on the Late Cretaceous taxa, 32 erymid species are now known from the Cretaceous deposits around the world (Table 2, Fig. 8A).

The Early Cretaceous erymid fauna is composed of 21 species distributed in six genera (*Eryma*, *Enoploclytia*, *Palaeastacus*, *Pustulina*, *Stenodactylina*, *Tethysastacus*) and appears to be more diversified than that of the Late Cretaceous (13 species distributed in *Enoploclytia*, *Palaeastacus*, *Pustulina*, and *Stenodactylina*). *Eryma* and *Tethysastacus* are not present in the Late Cretaceous. This decrease of the number of species confirm the Cretaceous decline of erymid lobsters suggested by Schweitzer & Feldmann (2014).

With six species of *Palaeastacus*, five of *Enoploclytia* and *Pustulina*, four of *Eryma*, and one of *Stenodactylina* and *Tethysastacus*, there is no domination of one erymid genus during the Early Cretaceous. However, with six species, *Enoploclytia* clearly dominates the erymid diversity in the Late Cretaceous.

## 4.2. Palaeobiogeography

Erymids were widespread in Early Cretaceous as the reports of their fossil occurrences from Europe, America, Australia, Antarctica, Madagascar and Japan document (Fig. 8B). Most of Early Cretaceous species are present in Western Europe (13 species; Devillez *et al.* 2016). For the Late Cretaceous, the erymid occurrences are almost exclusively concentrated in the Northern Hemisphere (Fig. 8C). The only known Late Cretaceous erymid fauna of the Southern Hemisphere was found in the Campanian deposits of Madagascar (Secrétan 1964; Charbonnier *et al.* 2012). Secrétan (1964) recognized the affinities of the Malagasy fauna with those of Europe, North America and Australia. This led her to suggest the presence of migration routes between Madagascar and other continents. The presence in France of a P1 chela attributed to *Stenodactylina cf. armata* (see remarks about *Stenodactylina*; Fig. 1C) confirms the existence of faunal exchanges between Europe and Madagascar.

During the Mesozoic, the presence of erymid lobsters seems to have been discontinuous in North America. Three occurrences are reported in the Middle and Late Jurassic: *Eryma foersteri* Feldmann 1979 (Callovian), *Eryma jungostrix* Feldmann & Titus, 2006 (Oxfordian) and *Stenodactylina walkerae* (Feldmann & Haggart, 2007) (middle Bajocian – early Callovian) and then no erymid is known between the Oxfordian and the Albian. At the end of the Early Cretaceous and in the Late Cretaceous, most of North American species (*Enoploclytia gigantea* Devillez, Charbonnier, Hyžný & Leroy, 2016, *E. tumimana*, *E. wintoni*, *Palaeastacus kimzeyi*, *P. walkeri* (Whitfield, 1883), *Stenodactylina triglypta*) occurred in the north of the Gulf of Mexico, which included the flooded part of the south of the United States (Texas, Louisiana, Mississippi, Alabama, Georgia, Florida). The gap in the erymid record between the Oxfordian and the Albian could be explained by (1) a collecting and/or reporting bias or by (2) the real absence of erymid lobsters in North America. In the second case, the settlement of a new erymid fauna in the north part of the Gulf of Mexico could be the result of migration of populations from the other area. Eurasia and
South America were the closest continents at the end of the Early Cretaceous, but only the Eurasiatic fauna included species of *Enoploclytia*, *Palaeastacus* and *Stenodactylina* during the Early Cretaceous (there was only some species of *Palaeastacus* at the extreme south of South America). Moreover, there were no connection between North America and South America at the end of the Early Cretaceous, while the northern part of the Atlantic Ocean was not completely opened (Scotese 2014a). Thus, a migration of Eurasiatic erymid populations to the west, through the northern part of the Atlantic, and then to the south, along the east coast of North America until they reached the Gulf of Mexico, was certainly possible.

We notice that no fossils of erymid lobsters are reported from the deposits of the Western Interior Seaway (WIS). This epicontinental sea has started to flood the North America in the Aptian and linked the Boreal Ocean with the Atlantic Ocean during almost all the Late Cretaceous (Gill & Cobban 1966, 1973; Molenaar & Rice 1988; Eicher & Diner 1989; Scotese 2014a & b). The absence of erymid lobsters in the WIS, while five species are present at its south-eastern extremity (Texas), is probably due to collecting and/or reporting biases. Indeed, the connection between the Gulf of Mexico and the WIS remained relatively stable during the Late Cretaceous (Sageman & Arthur 1994; Robinson Roberts & Kirschbaum 1995; Scotese 2014b), so no geographic barrier could have prevent the settlement of an erymid fauna in the WIS. The last three North American species (*Enoploclytia minor*, *Pustulina dawsoni*, *P. trisulcata*) occurred along the west coast of United States and Canada, in the Pacific Ocean (Fig. 8C).

**5. Conclusions**

The present review of the Late Cretaceous erymid lobsters includes 13 species with a worldwide distribution. Four genera are represented: *Enoploclytia*, with six species (*E. collignoni*, *E. leachi*, *E. minor*, *E. seitzi*, *E. tepeyacensis*, *E. tumimana*), *Palaeastacus*, with two species (*P. kimzeyi*, *P. sussexiensis*), *Pustulina* with three species (*P. cretacea*, *P. dawsoni*, *P. trisulcata* nov. comb.), *Stenodactylina* with two species (*S. armata*, *S. triglypta*).

We emphasize a change in the composition of erymid fauna between Early and Late Cretaceous. To our knowledge, there is a fewer number of species in the Late Cretaceous. Almost half of them are assigned to *Enoploclytia* while no genus clearly included more species than others in the Early Cretaceous. Moreover, *Eryma*, well-represented during the Jurassic and the Early Cretaceous, is absent in the Late Cretaceous. This decrease in erymid diversity suggests a progressive decline of these lobsters, previously suggested by Schweitzer & Feldmann (2014).

The scarcity and the discontinuity of the record of the Erymidae is a limit for the palaeobiogeographic interpretations. This difficulty prevent us to know if the presence of an erymid fauna in North America was continuous between the Late Jurassic and the Early Cretaceous or if the Cretaceous populations result of migrations from the other continent. In this case, the area of origin could have been Europe because of its connections, the geographic proximity and affinities of its erymid fauna (all the
Cretaceous North American genera are represented in Western Europe in the first half of the Early Cretaceous). Moreover, we have noted that almost all the erymids known during the Late Cretaceous are reported from the northern hemisphere. The only erymids of the southern hemisphere were found in Madagascar and the report of a P1 chela of *S. cf. armata* in France confirms the existence of migration between Europe and Madagascar as suggested by Secrétan (1964). This asymmetric repartition between the two hemispheres is probably due to collecting and/or report biases. So, only new reports, especially in the southern hemisphere, could allow us to identify the migration channels of the erymid lobsters during the Cretaceous and the geographic origins of the different faunas.

**Acknowledgements**

The authors are gratefull to Claire Mellish (Natural History Museum, London, United Kingdom), Liz Harper and Matt Riley (Sedgwyck Museum, Cambridge, United Kingdom), Lee Ismail and John Cooper (Booth Museum, Brighton, United Kingdom) and Jan Sklenář (Národní Muzeum, Prague, Czech Republic) for the access to the fossils housed in the collections of their respective institutions.

We also greatly thank Hilary Blagbrough (British Antarctic Survey, Cambridge, United Kingdom), Angela Ehling and Tanja Wodtke (Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin, Germany), Christian Neumann (Museum für Naturkunde, Berlin, Germany), Georg Heumann, Georg Oleschinski and Jes Rust (Steinmann Institut für Geologie, Mineralogie und Paläontologie, Bonn), Rodney Feldmann (Kent State University, Kent, USA), Thomas Jorstad, Matthew Miller and Laurence O'Reilly (Department of Paleobiology, Smithsonian Institution, Washington, D.C), Ann Molineux and Angella Thompson (Non-vertebrate Paleontology Laboratory, Jackson School of Geosciences, University of Texas, Austin, USA) and Francisco Vega (Instituto de Geologia Mexico, Mexico) for searching and supplying photographs of the specimens stored in the collections of their institutions.

We also really thank Jean-Michel Pacaud (MNHN, Paris, France), for helping to solve some nomenclatural complications, Colas Bouillet and Yohan Despres for the preparation of some studied specimens (MNHN, Paris, France) and Lilian Cazes and Philippe Loubry (MNHN, Paris, France) for the photographs of some figured specimens.

Finally, we also thank Günter Schweigert (Stattliches Museum für Naturkunde, Stuttgart, Germany) and Matúš Hyžný (Comenius University, Bratislava, Slovakia) for their valuable reviews of this work and for their useful advices.

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SCHWEIGERT G. 2013. — A new record of the enigmatic lobster genus *Stenodactylina* Beurlen, 1928 (Crustacea: Decapoda: Erymidae) from the Middle Jurassic of southwestern Germany. *Paläontologische Zeitschrift* 87: 409-413.

SCHWEIGERT G., DIETL G. & RÖPER M. 2000. — Die Panzerkrebse des Familie Erymidae VAN STRAELEN (Crustacea, Decapoda) aus dem Nusplinger Plattenkalk (Ober-Kimmeridgium, Schwäbische Alb) im Vergleich mit fränkischen Vorkommen. *Stuttgarter Beiträge zur Naturkunde* B (285): 1-25.

SCHWEITZER C. E. & FELDMANN R. M. 2001. — New Cretaceous and Tertiary decapod crustaceans from western North America. *Bulletin of Mizunami Fossil Museum* 28: 173-210.

SCHWEITZER C. E. & FELDMANN R. M. 2014. — Lobster (Decapoda) diversity and evolutionary patterns through time. *Journal of Crustacean Biology* 34 (6): 173-210.

SCHWEITZER C. E., FELDMANN R. M., GARASSINO A., KARASAWA H. & SCHWEIGERT G. 2010. — Systematic list of fossil decapod crustacean species. *Crustaceana Monographs* 10: 1-222.
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## Captions

TABLE 1. — List of the examined material of Late Cretaceous erymid lobsters.

TABLE 2. — List and occurrences of the Early and Late Cretaceous erymid species.

FIG. 1. — Late Cretaceous specimens of *Stenodactylina* Beurlen, 1928. **A-B**, holotype MNHN.F.R03913 of *Stenodactylina armata* (Secrétan, 1964) from the Campanian of Belo-sur-Tsiribihina, Madagascar: right P1 chela in dorsal view (A), and inner view (B); **C**, specimen MNHN.F.A66891 of *Stenodactylina* cf. *armata* from the Santonian of Cognac, France, dorsal view of right P1; **D-H**, *Stenodactylina triglypta* (Stenzel, 1945) from the Coniacian of Savoy, USA: lectotype TMM-BEG 21127 (D-F) showing part and counter part of carapace, and line drawing; paralectotype TMM-BEG 21124 (G) showing incomplete P1 chela; paralectotype TMM-BEG 21125 (H) showing fragment of right side of carapace with superimposed line drawing of grooves. Scale bars: 20 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; $b_1$: hepatic groove; c: postcervical groove; d: gastro-orbital groove; $e_1$: cervical groove; i: inferior groove; ω: attachment site of mandibular muscle. Photographs: C. Lemzaouda (**A**, **B**), P. Loubry (**C**), Non-vertebrate Paleontology Laboratory, Jackson School of Geosciences, The University of Texas at Austin (**D**, **E**, **G**, **H**). Line drawing: J. Devillez.

FIG. 2. — *Palaeastacus sussexiensis* (Mantell, 1824). **A-B**, lectotype NHMUK 5601 (Mantell coll.) from the Cenomanian of Sussex, United Kingdom: right side of carapace with parts of pereiopods (A) and line drawing of carapace (B); **C-D**, lectotype BM 007750 (Willet coll.) of *Palaeastacus dixoni* Bell, 1850 from the Cenomanian of Glynde, United Kingdom: subcomplete specimen in right lateral view (C) and dorsal view (D); **E**, specimen BAS KG.103.134 from the Aptian of Alexander Island, Antarctica, left lateral view; **F**, specimen MNHN.F.A58710 from the Turonian of Saint Florentin, France, dorsal view of left P1 chela. Scale bars: 10 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; $b_1$: hepatic groove; c: postcervical groove; d: gastro-orbital groove; $e_1$: cervical groove; i: inferior groove. Preparation: Y. Despres (**F**). Photographs: J. Devillez (**A**, **C**, **D**), H. Blagbrough (**E**), L. Cazes (**F**). Line drawing: J. Devillez.

FIG. 3. — *Palaeastacus kimzeyi* Rathbun, 1935 from the Campanian of Farmersville, United States (**A-C**) and *Pustulina trisulcata* (Schweitzer & Feldmann, 2001) n. comb. from the Cenomanian of Sites, United States (**D-E**). **A-B**, holotype USNM PAL 73797: P1 propodus, dorsal surface (A) and ventral surface (B); **C**, paratype USNM PAL 73841: fragment of carapace, left lateral view with superimposed line drawing of
grooves; **D-E**, holotype USNM PAL 512150, subcomplete specimen, left side of carapace and pleon with superimposed line drawing of grooves (D) and dorsal view (E). Scale bars: 10 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; b₁: hepatic groove; c: postcervical groove; cd: cardiac groove; d: gastro-orbital groove; e₁e: cervical groove; i: inferior groove. Photographs: L. O'Reilly. Line drawings: J. Devillez.

FIG. 4. — Historical specimens of *Enoploclytia leachi* (Mantell, 1822). **A**, line drawing of carapace groove pattern characteristic of *Enoploclytia* M'Coy, 1849; **B**, line drawing of P1 chela characteristic of *Enoploclytia*; **C**, lectotype NHMUK 5636 (Mantell coll.) from the Turonian of Sussex, United Kingdom, left P1 chela; **D**, paralectotype NHMUK In.61020 from the Late Cretaceous of Sussex, United Kingdom, fragment of pleon; **E-F**, specimen NM O7125 (figured by Reuss 1854: pl. 1, fig. 2) from the Turonian of Prague, Czech Republic: general view showing the third maxilliped (E) and line drawing of carapace (F); **G-H**, syntype of *Enoploclytia heterodon* Schlüter, 1862 from the Santonian of Dülmen, Germany: original figure of Mertin (1941: pl. 5, fig. 1) (G) and line drawing of carapace (H); **I-J**, holotype STIPB-Schlüter-124 of *Enoploclytia granulicauda* Schlüter, 1879 from the Campanian of Coesfeld, Germany: general view (I) and line drawing of the carapace (J). Scale bars: 20 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; b₁: hepatic groove; c: postcervical groove; d: gastro-orbital groove; e₁e: cervical groove; i: inferior groove; ip: intercalated plate; Mxp3: third maxilliped; ω: attachment site of mandibular muscle; χ: attachment site of adductor testis muscle. Photographs: J. Devillez (**C**, **D**), M. Kočová Veselská (**E**), G. Oleschinski (**I**). Line drawings: J. Devillez.

FIG. 5. — New specimens of *Enoploclytia leachi* (Mantell, 1822). **A-B**, specimen MNHN.F.B16580 from the Cenomanian of La Ferté-Bernard, France: right lateral view showing the merus with prominent distal process (A) and left lateral view (B); **C-D**, specimen MNHN.F.A66890 (Pezy coll.) from the Cenomanian of Villers-sur-Mer, France: general view (C) and line drawing of carapace (D); **E**, specimen BM 007766 from the Late Cretaceous of Brighton, United Kingdom. Scale bars: 20 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; b₁: hepatic groove; c: postcervical groove; d: gastro-orbital groove; e₁e: cervical groove; i: inferior groove; ip: intercalated plate; ω: attachment site of mandibular muscle; χ: attachment site of adductor testis muscle. Preparation: Y. Despres (**A-B**). Photographs: P. Loubry (**A-C**), J. Devillez (**E**). Line drawing: J. Devillez.

FIG. 6. — *Enoploclytia leachi* (Mantell, 1822) from the Turonian of Bílá Hora, Czech Republic. **A**, specimen NM O4062 (figured by Reuss 1854: pl. 2, fig. 4), carapace and pleon, left lateral view; **B**, specimen NM O4042 (listed by Fritsch & Kafka 1887),
subcomplete specimen, dorsal view; C, specimen NM O7123, subcomplete specimen,
left lateral view. Scale bars: 20 mm. Photographs: M. Kočová Veselská.

FIG. 7. — *Enoploclytia seitzi* Glaessner, 1932. **A-B**, holotype BGR X 00266 (not found) from the Albian of Wettensen, Germany: original figure of Förster (1966: pl. 19, fig. 1) (A) and line drawing of carapace (B); **C**, specimen NHMUK 5918 (initially named as *Enoploclytia scabrosa* by Woodward 1873 – nomen nudum) from the Cenomanian of Dover, United Kingdom: right lateral view showing the prominent distal process of the P1 merus. Scale bars: 10 mm. Abbreviations: a: branchiocardiac groove; b: antennal groove; b₁: hepatic groove; c: postcervical groove; d: gastro-orbital groove; e₁e: cervical groove; i: inferior groove; ω: attachment site of mandibular muscle; χ: attachment site of adductor testis muscle. Photographs and line drawing: J. Devillez.

FIG. 8. — Stratigraphic (A) and geographic distribution (B, C) of the erymid lobsters
during the Cretaceous. Palaeogeographic maps of the Middle Albian (B) and Early
Campanian (C) from R. C. Scotese (2014a & b).
<figure><img src="image_3.png" /><figcaption>C</figcaption></figure>

<figure><img src="image_4.png" /><figcaption>B</figcaption></figure>

<figure><img src="image_5.png" /><figcaption>D</figcaption></figure>

<figure><img src="image_6.png" /><figcaption>E</figcaption></figure>

<figure><img src="image_7.png" /><figcaption>F</figcaption></figure>

<figure><img src="image_8.png" /><figcaption>G</figcaption></figure>

<figure><img src="image_9.png" /><figcaption>H</figcaption></figure>

<figure><img src="image_10.png" /></figure>
<figure><img src="image_11.png" /><figcaption>B</figcaption></figure>

<figure><img src="image_12.png" /><figcaption>C</figcaption></figure>

<figure><img src="image_13.png" /><figcaption>D</figcaption></figure>

<figure><img src="image_14.png" /><figcaption>E</figcaption></figure>

<figure><img src="image_15.png" /><figcaption>F</figcaption></figure>

<figure><img src="image_16.png" /></figure>
<figure><img src="image_17.png" /><figcaption>B</figcaption></figure>

<figure><img src="image_18.png" /><figcaption>C</figcaption></figure>

<figure><img src="image_19.png" /><figcaption>D</figcaption></figure>

<figure><img src="image_20.png" /><figcaption>E</figcaption></figure>

<figure><img src="image_21.png" /></figure>
<figure><img src="image_22.png" /><figcaption>B</figcaption></figure>

<figure><img src="image_23.png" /><figcaption>C</figcaption></figure>

<figure><img src="image_24.png" /><figcaption>D</figcaption></figure>

<figure><img src="image_25.png" /><figcaption>E</figcaption></figure>

<figure><img src="image_26.png" /><figcaption>F</figcaption></figure>

<figure><img src="image_27.png" /><figcaption>G</figcaption></figure>

<figure><img src="image_28.png" /><figcaption>H</figcaption></figure>

<figure><img src="image_29.png" /><figcaption>I</figcaption></figure>

<figure><img src="image_30.png" /><figcaption>J</figcaption></figure>

<figure><img src="image_31.png" /></figure>
<figure><img src="image_32.png" /></figure>

007766
A

<figure><img src="image_33.png" /></figure>

B

<figure><img src="image_34.png" /></figure>

C

<figure><img src="image_35.png" /></figure>
<figure><img src="image_36.png" /></figure>

<figure><img src="image_37.png" /></figure>

<figure><img src="image_38.png" /></figure>
A

Early Cretaceous

Late Cretaceous

<table><thead><tr><th>stages</th><th>Ber.</th><th>Val.</th><th>Haut.</th><th>Bar.</th><th>Apt.</th><th>Alb.</th><th>Cen.</th><th>Tur.</th><th>Con.</th><th>Sant.</th><th>Camp.</th><th>Maas.</th></tr></thead><tbody><tr><td rowspan="5">Europe</td><td>1<br>P. occitana</td><td>4<br>P. loryi</td><td>7<br>E. glaessneri</td><td>10<br>E. augustobonae</td><td></td><td>-13<br>P. sussexiensis</td><td>13<br>P. leachi</td><td>13<br>E. seitzii</td><td></td><td>20<br>S. aft. armata</td><td></td><td></td></tr><tr><td>2<br>P. victori</td><td>5<br>T. tithonius</td><td>8<br>E. sulcumat</td><td>9<br>P. colossea</td><td></td><td>16<br>E. vocontii</td><td>22<br>P. tuberculata</td><td>22<br>E. leachi</td><td>22<br>E. seitzii</td><td>22<br>S. aft. armata</td><td>22<br>E. seitzii</td><td></td></tr><tr><td>3<br>S. delphinensis</td><td></td><td></td><td></td><td></td><td>17<br>P. tuberculata</td><td>21<br>E. seitzii</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>22<br>E. gigantea</td><td>23<br>P. trisulcata</td><td></td><td>25<br>S. triglypta</td><td>27<br>E. tepeyacensis</td><td>31<br>E. minor</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>20<br>E. wintani</td><td>24<br>P. walkeri</td><td></td><td></td><td>20<br>P. kimzeyi</td><td>32<br>E. tumimana</td></tr><tr><td>North America</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td rowspan="4">other regions</td><td></td><td>6<br>P. spinulata</td><td>1<br>E. nippon</td><td>14<br>P. sussexiensis</td><td></td><td></td><td>25<br>P. cretacea</td><td></td><td></td><td></td><td>26<br>S. armata</td><td></td></tr><tr><td></td><td></td><td>12<br>P. foersteri</td><td>15<br>P. terraoreginae</td><td></td><td></td><td></td><td></td><td></td><td></td><td>20<br>E. collignoni</td><td></td></tr><tr><td></td><td></td><td></td><td>13<br>P. terraoreginae</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>15<br>P. tenuidigitatus</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

B

<figure><img src="image_39.png" /></figure>

Middle Albian
106 Ma

C

<figure><img src="image_40.png" /></figure>

Early Campanian
80.3 Ma
<table><thead><tr><th>TAXA</th><th>EXAMINED MATERIAL</th><th>ANATOMY</th><th>AGES</th><th>LOCALITIES</th></tr></thead><tbody><tr><td><strong>Eryma Meyer, 1840</strong><br><em>Eryma vocontii</em> n. sp.</td><td>holotype MNHN.F.A57457, paratype MNHN.F.A57458.</td><td>carapace, P1 chela</td><td>Albian</td><td>France</td></tr><tr><td colspan="5"><strong>Enoploclytia M'Coy, 1849</strong></td></tr><tr><td><em>Enoploclytia collignoni</em> Secrétan, 1964</td><td>—holotype MNHN.F.R03925, 43 paratypes.<br>—16 additional specimens MNHN.F.A33041, A33058, A33079, A33137, A33162, A33164, A33167, A33178, A33186, A33187, A33193, A33229, A33332, A33362, A33469, A33472.<br>—lectotype NHMUK 5636, 2 paralectotypes NHMUK 5623, In.61020.</td><td>carapaces, P1 chelae, pleons</td><td>Campanian</td><td>Madagascar</td></tr><tr><td><em>Enoploclytia leachi</em> (Mantell, 1822)</td><td>—86 additional specimens NHMUK 5600, 5606, 5609, 5610, 5611, 5615, 5616, 5634, 34404, 59825, I.193, I.1977, I.1980, I.2190, In.27682; BM 007759, 007760, 007762, 007763, 007764, 007765, 007766, 016987, 016989; MNHN.F.A60030, A66890, A66894, B16580, B16612, R03372; NM O1261, O3500, O4002, O4042, O4043, O4062, O4063, O4915, O6084, O6085, O6086, O7123, O7124, O7125, MV 1-MV 11, MV 17-MV 24, MV 26-MV 31, MV 39-MV 45, MV 53-MV 56, MV 62-MV 68.</td><td>carapaces, P1 chelae, pleons</td><td>Cenomanian - Campanian</td><td>France, Czech Republic, United Kingdom</td></tr><tr><td><em>Enoploclytia seitzi</em> Glaessner, 1932</td><td>—holotype STIPB-Schlüter-124 of <em>E. granulicauda</em> Schlüter, 1879.<br>lectotype BGR X 00266, 1 paralectotype BGR X 00267.<br>1 additional specimen NHMUK 5918.</td><td>carapace, P1 chela, pleon<br>carapaces<br>carapace, P1 chela, pleon</td><td>Campanian<br>Albian<br>Cenomanian</td><td>Germany<br>Germany<br>United Kingdom</td></tr><tr><td><em>Enoploclytia tumimanus</em> Rathbun, 1935</td><td>holotype USNM PAL 73799, 8 paratypes USNM PAL 336023 A, 336023 B, 336023 C, 336023 D, 336023 E, 336023 F, 336023 G, 336023 H.</td><td>carapaces, P1 chelae</td><td>Maastrichtian</td><td>United States</td></tr><tr><td><em>Enoploclytia tepeyacensis</em> Vega, Garassino &amp; Jaime, 2013</td><td>holotype MUZ-824, 1 paratype MUZ-825.</td><td>carapaces</td><td>Campanian</td><td>Mexico</td></tr><tr><td colspan="5"><strong>Palaeastacus Bell, 1850</strong></td></tr><tr><td><em>Palaeastacus kimzeyi</em> Rathbun, 1935</td><td>holotype USNM PAL 73797, 2 paratypes USNM PAL 335146, 73841.<br>—lectotype NHMUK 5601, 13 paralectotypes NHMUK 5024, 5584, 5586, 5589, 5600, 5602, 5608, 5613, 5618, 5624, 5626, 5629, 10760.<br>—37 additional specimens NHMUK In.27682, In.36768, I.62, I.2190, I.2628, 5685, 35124, 44343, 024318, 026316, 46782, 59668, 59824, 59824; BAS KG.103.134; BM 002548, 002549, 007751, 007752, 007753, 007754, 007755, 007758, 008632, 008633, 008635, 008636, 008637, 008659, 016988, 024317, 024318, 026316; MNHN.F.A58710, A66893, B14182, B17969. 3 syntypes SM B8884, B8885, B8886 of <em>Enoploclytia brevimana</em> M'Coy 1849.</td><td>carapace, P1 chelae</td><td>Campanian</td><td>United States</td></tr><tr><td><em>Palaeastacus sussexiensis</em> (Mantell, 1824)</td><td>8 syntypes SM B8892, B8893, B8894, B8895, B8896, B8897, B8898, B8899 of <em>Enoploclytia imagei</em> M'Coy 1849.<br>— lectotype NHMUK 41938, 2 paralectotypes NHMUK 39377, 39380 of <em>Hoploparia scabra</em> Bell, 1863.<br>—4 additional specimens NHMUK In.27684, In.61022, In.27685, In. 27686 of <em>Hoploparia scabra</em> Bell, 1863.<br>—lectotype BM 007750, paralectotype NHMUK I.1916, of <em>Palaeastacus dixoni</em> Bell, 1850.<br>—1 additional specimen BM 007757 of <em>Palaeastacus dixoni</em> Bell, 1850.<br>2 syntypes SM B22448, B22449 of <em>Phlyctisoma granulatum</em> Bell, 1863.</td><td>carapaces, P1 chelae, pleons<br>P1 chelae<br>carapace, P1 pereiopods<br>carapaces and P1 pereiopods<br>carapaces, P1 chelae, pleon<br>carapaces</td><td>Cenomanian - Turonian<br>Cenomanian<br>Cenomanian<br>Albian<br>Cenomanian<br>Albian</td><td>United Kingdom<br>United Kingdom<br>United Kingdom<br>United Kingdom<br>United Kingdom<br>United Kingdom</td></tr><tr><td colspan="5"><strong>Pustulina Quenstedt, 1857</strong></td></tr><tr><td><em>Pustulina cretacea</em> (Roger, 1946)</td><td>2 syntypes MNHN.F.B18903, MNHN.F.B18904.<br>—11 additional specimens MNHN.F.A30680, A30681, A30682, A30683, A30684, A30685, A30686, A30687, A38526, A38527, A38528.</td><td>carapaces, P1 chelae, pleons</td><td>Cenomanian</td><td>Lebanon</td></tr><tr><td><em>Pustulina trisulcata</em> (Schweitzer &amp; Feldmann, 2001) nov. comb.</td><td>holotype USNM 512150.</td><td>carapace, pleon</td><td>Cenomanian</td><td>United States</td></tr><tr><td colspan="5"><strong>Stenodactylina Beurlen, 1928</strong></td></tr><tr><td><em>Stenodactylina armata</em> (Secrétan, 1964)</td><td>—holotype MNHN.F.R03913, 3 paratypes MNHN.F.A33209, A33464, R03912.<br>—additionnal specimen MNHN.F.A33331</td><td>P1 chelae</td><td>Campanian</td><td>Madagascar</td></tr><tr><td><em>Stenodactylina cf. armata</em> (Secrétan, 1964)</td><td>MNHN.F.A66891.</td><td>P1 pereiopod</td><td>Santonian</td><td>France</td></tr><tr><td><em>Stenodactylina triglypta</em> (Stenzel, 1945)</td><td>lectotype NPL TMM-BEG 21127; 2 paralectotypes NPL TMM-BEG 21124, 21125.</td><td>carapaces, P1 chela</td><td>Coniacian</td><td>United States</td></tr><tr><td colspan="5"><strong>Non erymid species</strong></td></tr><tr><td><em>Eryma americanum</em> Rathbun, 1923</td><td>holotype USNM PAL 31899, 2 paratypes USNM PAL 31900, 31901.</td><td>carapaces, P1 chela</td><td>Late Cretaceous</td><td>United States</td></tr><tr><td><em>Eryma flectum</em> Rathbun, 1926</td><td>holotype USNM PAL 73115.</td><td>P1 chela</td><td>Maastrichtian</td><td>United States</td></tr><tr><td><em>Eryma stantoni</em> Rathbun, 1935</td><td>holotype USNM PAL 73790.</td><td>P1 chela</td><td>Maastrichtian</td><td>United States</td></tr><tr><td><em>Enoploclytia sculpta</em> Rathbun, 1926</td><td>holotype USNM PAL 73119.</td><td>P1 pereiopod, pleon</td><td>Maastrichtian</td><td>United States</td></tr><tr><td><em>Palaeastacus selmaensis</em> Rathbun, 1935</td><td>holotype USNM PAL 73848.</td><td>unidentifiable fragment</td><td>Late Cretaceous</td><td>United States</td></tr></tbody></table>
LATE CRETACEOUS

<table><thead><tr><th>Europe</th><th>AGES</th><th>OCCURENCES</th></tr></thead><tbody><tr><td>Enoploclytia leachi (Mantell, 1822)</td><td>Cenomanian to Campanian</td><td>France, United Kingdom, Germany, Belgium, Czech Republic</td></tr><tr><td>Enoploclytia seitzi Glaessner, 1932</td><td>Albian to Cenomanian</td><td>United Kingdom, Germany</td></tr><tr><td>Palaeastacus sussexiensis (Mantell, 1824)</td><td>Aptian to Turonian</td><td>France, United Kingdom, Germany, Antarctic</td></tr><tr><td>Stenodactylina cf. armata (Secrétan, 1964)</td><td>Santonian</td><td>France</td></tr><tr><td colspan="3"><strong>North America</strong></td></tr><tr><td>Enoploclytia minor Woodward, 1900</td><td>Maastrichtian</td><td>Canada</td></tr><tr><td>Enoploclytia tepeyacensis Vega, Garassino &amp; Jaime, 2013</td><td>Campanian</td><td>Mexico</td></tr><tr><td>Enoploclytia tumimana Rathbun, 1935</td><td>Maastrichtian</td><td>USA</td></tr><tr><td>Palaeastacus kimzeyi Rathbun, 1935</td><td>Campanian</td><td>USA</td></tr><tr><td>Pustulina dawsoni (Woodward, 1900)</td><td>Campanian to Maastrichtian</td><td>Canada</td></tr><tr><td>Pustulina trisulcata (Schweitzer &amp; Feldmann, 2001) nov. comb.</td><td>Cenomanian</td><td>USA</td></tr><tr><td>Stenodactylina triglypta (Stenzel, 1945)</td><td>Coniacian</td><td>USA</td></tr><tr><td colspan="3"><strong>Other regions</strong></td></tr><tr><td>Enoploclytia collignoni Secrétan, 1964</td><td>Campanian</td><td>Madagascar</td></tr><tr><td>Pustulina cretacea (Roger, 1946)</td><td>Cenomanian</td><td>Lebanon</td></tr><tr><td>Stenodactylina armata (Secrétan, 1964)</td><td>Campanian</td><td>Madagascar</td></tr></tbody></table>

EARLY CRETACEOUS

<table><thead><tr><th>Europe</th><th>AGES</th><th>OCCURENCES</th></tr></thead><tbody><tr><td>Eryma glaessneri Van Straelen, 1936</td><td>Hauterivian</td><td>France</td></tr><tr><td>Eryma sulcatum Harbort, 1905</td><td>Hauterivian</td><td>United Kingdom, Germany</td></tr><tr><td>Eryma vocontii Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Albian</td><td>France</td></tr><tr><td>Enoploclytia augustobonae Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Barremian</td><td>France</td></tr><tr><td>Enoploclytia seitzi Glaessner, 1932</td><td>Albian to Cenomanian</td><td>United Kingdom, Germany</td></tr><tr><td>Palaeastacus loryi (Van Straelen, 1923)</td><td>Valanginian</td><td>France</td></tr><tr><td>Palaeastacus sussexiensis (Mantell, 1824)</td><td>Aptian to Turonian</td><td>France, United Kingdom, Germany, Antarctic</td></tr><tr><td>Pustulina colossea Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Hauterivian</td><td>France</td></tr><tr><td>Pustulina occitana Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Berriasian</td><td>France</td></tr><tr><td>Pustulina tuberculata (Bell, 1863)</td><td>Albian</td><td>United Kingdom</td></tr><tr><td>Pustulina victori Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Berriasian</td><td>France</td></tr><tr><td>Stenodactylina delphinensis (Moret, 1946)</td><td>Berriasian</td><td>France</td></tr><tr><td>Tethysastacus tithonius (Van Straelen, 1936)</td><td>Valanginian</td><td>France</td></tr><tr><td colspan="3"><strong>North America</strong></td></tr><tr><td>Enoploclytia gigantea Devillez, Charbonnier, Hyžný &amp; Leroy, 2016</td><td>Albian</td><td>USA</td></tr><tr><td>Enoploclytia wintoni Stenzel, 1845</td><td>Albian</td><td>USA</td></tr><tr><td>Palaeastacus walkeri (Whitfield, 1883)</td><td>Albian</td><td>USA</td></tr><tr><td colspan="3"><strong>Other regions</strong></td></tr><tr><td>Eryma nippon Karasawa, Ohara &amp; Kato, 1936</td><td>Barremian</td><td>Japan</td></tr><tr><td>Palaeastacus foersteri Taylor, 1979</td><td>Barremian</td><td>Antarctic</td></tr><tr><td>Palaeastacus terraereginae (Etheridge Jr., 1914)</td><td>Barremian</td><td>Antarctic, Argentina, Australia</td></tr><tr><td>Palaeastacus tenuidigitatus (Woods, 1957)</td><td>Aptian</td><td>Australia</td></tr><tr><td>Pustulina spinulata Secrétan, 1964</td><td>Valanginian to Hauterivian</td><td>Madagascar</td></tr></tbody></table>