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Tanidromites maerteni sp. nov. urn:lsid:zoobank.org:act: 6AF2E747-B3FE-4417-A4D9-3B94B2A5693A Tanidromites maerteni Fraaije, Van Bakel, Guinot & Jagt, 2013: 250, fig. 1 [unavailable]. Type material. The holotype, MNHN.F.A47612 (leg. L. Maerten), is deposited in the collections of the Muséum National d’Histoire Naturelle, Département Histoire de la Terre (Paris). The type specimen originates from the town quarry of the Commune de Maizet, Calvados (northwest France) from a level which is locally referred to as the “Oolithes ferrugineuses de Bayeux”, the age of which is early late Bajocian (Niortense ammonite Zone) (Gauthier et al. 1996). Etymology. In honour of Lionel Maerten (Ver-sur-Mer, France), who collected the holotype and kindly donated it to the Muséum National d’Histoire Naturelle collections. Diagnosis. A tanidromitid with lateral epibranchial spine; slightly concave portion of cervical groove at central posterior margin of mesogastric region; two faint, yet clearly visible, V-shaped grooves on central part of mesogastric region; relatively deep and long postcervical grooves; pustulate ornament on cardiac, epibranchial and epigastric regions (in accordance with Fraaije et al. 2013: 252). Remarks. A full description and figures of Tanidromites maerteni sp. nov. are supplied by Fraaije et al. (2013).

Tanidromites maerteni sp. nov. urn:lsid:zoobank.org:act: 6AF2E747-B3FE-4417-A4D9-3B94B2A5693A Tanidromites maerteni Fraaije, Van Bakel, Guinot & Jagt, 2013: 250, fig. 1 [unavailable]. Type material. The holotype, MNHN.F.A47612 (leg. L. Maerten), is deposited in the collections of the Muséum National d’Histoire Naturelle, Département Histoire de la Terre (Paris). The type specimen originates from the town quarry of the Commune de Maizet, Calvados (northwest France) from a level which is locally referred to as the “Oolithes ferrugineuses de Bayeux”, the age of which is early late Bajocian (Niortense ammonite Zone) (Gauthier et al. 1996). Etymology. In honour of Lionel Maerten (Ver-sur-Mer, France), who collected the holotype and kindly donated it to the Muséum National d’Histoire Naturelle collections. Diagnosis. A tanidromitid with lateral epibranchial spine; slightly concave portion of cervical groove at central posterior margin of mesogastric region; two faint, yet clearly visible, V-shaped grooves on central part of mesogastric region; relatively deep and long postcervical grooves; pustulate ornament on cardiac, epibranchial and epigastric regions (in accordance with Fraaije et al. 2013: 252). Remarks. A full description and figures of Tanidromites maerteni sp. nov. are supplied by Fraaije et al. (2013)., Published as part of Fraaije, René H. B., Van Bakel, Barry W. M., Guinot, Danièle & Jagt, John W. M., 2023, Validation of Tanidromites maerteni Fraaije, Van Bakel, Guinot & Jagt, a species of crab (Crustacea, Decapoda, Brachyura) from Middle Jurassic (Bajocian) deposits of Maizet, Calvados, northwest France, pp. 437-438 in Zootaxa 5318 (3) on page 437, DOI: 10.11646/zootaxa.5318.3.9, http://zenodo.org/record/8167015, {"references":["Fraaije, R. H. B., Van Bakel, B. W. M., Guinot, D. & Jagt, J. W. M. (2013) A new Middle Jurassic (Bajocian) homolodromioid crab from northwest France; the earliest record of the Tanidromitidae. Boletin de la Sociedad Geologica Mexicana, 65, 249 - 254. https: // doi. org / 10.18268 / BSGM 2013 v 65 n 2 a 6","Gauthier, H., Rioult, M. & Trevisan, M. (1996) Repartition biostratigraphique des ammonites dans l'Oolithe ferrugineuse de Bayeux (Bajocien) a Feuguerolles-sur-Orne (Calvados). Elements nouveaux pour une revision des Garantianinae. Geologie de la France, 2, 27 - 67."]}

Paguristes timoni n. sp. (Fig. 2) Zoobank: urn:lsid:zoobank.org:act: C4F34FA0-7A19-46B4-91BC-C93C621CD977 Diagnosis. Keeled merus with row of small teeth on distal margin; moveable finger corneous; fixed finger covered with rows of alveolate tubercles, decreasing in size on distal end. Upper margin with tubercles; cutting edge with several teeth proximally. Walking legs with keeled edge and row of tubercles. Type material. The holotype, and sole specimen known to date, is CMM-I-4600. Etymology. Named after the legendary misanthrope and hermit, Timon of Athens, who was popularized in the play ‘Timon of Athens’ by William Shakespeare (1564–1616). Locality and stratigraphy. Driftwood Beach, Calvert County, Maryland, from the upper Miocene (Tortonian) Little Cove Point Member of the St. Marys Formation in a silty lens within Bed E (Ward & Andrews 2008). Kidwell et al. (2015) identified this level as belonging to “SM-C,” assigned to Shattuck zones 22–23 and part of dinocyst zone 8. Description. CMM-I-4600, preserved inside gastropod shell (Busycon sp.), length 25 mm, greatest width 17 mm. Manus missing from major (left) cheliped, carpus and merus preserved. Keeled merus with row of small teeth on distal margin. Minor (right) cheliped comprising complete propodus, lodged in gastropod aperture, measuring 7 mm length, 2 mm maximum width. Moveable finger corneous, fixed finger covered with rows of alveolate tubercles decreasing in size on distal end; most tubercles with alveoli, indicative of setal insertions; upper margin covered with tubercles; cutting edge with several teeth proximally; distal side not visible. Carapace fragment preserved but covered; small portion visible lacking any specific details; Single fragment of walking leg preserved showing keeled edge with row of tubercles. Several other fragments preserved, but unidentifiable due to weathering of specimen. Remarks. In this novel form, the left cheliped is larger than the right one, which is diagnostic feature of diogenid and annuntidiogenid hermit crabs (McLaughlin 2003; Fraaije 2014). The Maryland specimen is remarkably similar to the extant Paguristes candelae De Matos-Pita & Ramil, 2015, from Mauritania (see below). Discussion. Cheliped shape and ornament in P. timoni n. sp. compare closely with those of P. candelae, in that both have rows of tubercles and setae and tubercles decrease in size toward the distal end. The dorsomesial margin in P. candelae has three large tubercles; these are absent from P. timoni n. sp. The dorsal cheliped surface in the latter appears to be less convex in comparison with that of P. candelae, while the teeth along the cutting edge appear to be larger in the extinct form. The walking legs in both taxa show a keeled ridge with a row of spines, although the one in P. timoni n. sp. seems more acute.

Paguristes timoni n. sp. (Fig. 2) Zoobank: urn:lsid:zoobank.org:act: C4F34FA0-7A19-46B4-91BC-C93C621CD977 Diagnosis. Keeled merus with row of small teeth on distal margin; moveable finger corneous; fixed finger covered with rows of alveolate tubercles, decreasing in size on distal end. Upper margin with tubercles; cutting edge with several teeth proximally. Walking legs with keeled edge and row of tubercles. Type material. The holotype, and sole specimen known to date, is CMM-I-4600. Etymology. Named after the legendary misanthrope and hermit, Timon of Athens, who was popularized in the play ‘Timon of Athens’ by William Shakespeare (1564–1616). Locality and stratigraphy. Driftwood Beach, Calvert County, Maryland, from the upper Miocene (Tortonian) Little Cove Point Member of the St. Marys Formation in a silty lens within Bed E (Ward & Andrews 2008). Kidwell et al. (2015) identified this level as belonging to “SM-C,” assigned to Shattuck zones 22–23 and part of dinocyst zone 8. Description. CMM-I-4600, preserved inside gastropod shell (Busycon sp.), length 25 mm, greatest width 17 mm. Manus missing from major (left) cheliped, carpus and merus preserved. Keeled merus with row of small teeth on distal margin. Minor (right) cheliped comprising complete propodus, lodged in gastropod aperture, measuring 7 mm length, 2 mm maximum width. Moveable finger corneous, fixed finger covered with rows of alveolate tubercles decreasing in size on distal end; most tubercles with alveoli, indicative of setal insertions; upper margin covered with tubercles; cutting edge with several teeth proximally; distal side not visible. Carapace fragment preserved but covered; small portion visible lacking any specific details; Single fragment of walking leg preserved showing keeled edge with row of tubercles. Several other fragments preserved, but unidentifiable due to weathering of specimen. Remarks. In this novel form, the left cheliped is larger than the right one, which is diagnostic feature of diogenid and annuntidiogenid hermit crabs (McLaughlin 2003; Fraaije 2014). The Maryland specimen is remarkably similar to the extant Paguristes candelae De Matos-Pita & Ramil, 2015, from Mauritania (see below). Discussion. Cheliped shape and ornament in P. timoni n. sp. compare closely with those of P. candelae, in that both have rows of tubercles and setae and tubercles decrease in size toward the distal end. The dorsomesial margin in P. candelae has three large tubercles; these are absent from P. timoni n. sp. The dorsal cheliped surface in the latter appears to be less convex in comparison with that of P. candelae, while the teeth along the cutting edge appear to be larger in the extinct form. The walking legs in both taxa show a keeled ridge with a row of spines, although the one in P. timoni n. sp. seems more acute., Published as part of Wallaard, Jonathan J. W., Fraaije, René H. B., Van Bakel, Barry W. M., Nance, John R., Lindholm, Adam & Jagt, John W. M., 2023, New hermit crab species (Anomura, Paguroidea) from the upper Miocene St Marys Formation of Maryland (USA), preserved in their host shells, pp. 389-397 in Zootaxa 5227 (3) on pages 393-394, DOI: 10.11646/zootaxa.5227.3.7, http://zenodo.org/record/7518846, {"references":["Ward, L. W. & Andrews, G. W. (2008) Stratigraphy of the Calvert, Choptank, and St. Mary's formations (Miocene) in the Chesapeake Bay area, Maryland and Virginia. Virginia Museum of Natural History Memoir, 9, 1 - 60.","Kidwell, S. M., Powars, D. S., Edwards, L. E. & Vogt, P. R. (2015) Miocene stratigraphy and paleoenvironments of the Calvert Cliffs, Maryland. Bulletin of the Geological Society of America, 40, 231 - 279.","McLaughlin, P. A. (2003) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirsof Museum Victoria, 60 (1), 111 - 144.","Fraaije, R. H. B. (2014) Diverse Late Jurassic anomuran assemblages from the Swabian Alb and evolutionary history of paguroids based on carapace morphology. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen, 273 (2), 121 - 145.","De Matos-Pita, S. S. & Ramil, F. (2015) Hermit crabs (Decapoda: Crustacea) from deep Mauritanian waters (NW Africa) with the description of a new species. Zootaxa, 3926 (2), 151 - 190."]}

Genus Pagurus Fabricius, 1775 Type species. Cancer bernhardus Linnaeus, 1758, by monotypy. Species included. For data on extinct forms, reference is made to lists provided by Schweitzer et al. (2010) and to subsequent records by Beschin et al. (2012), De Angeli & Caporiondo (2017) and Polkowsky & Fraaije (2019). Extant species have recently been discussed by Lemaitre & McLaughlin (2021a)., Published as part of Wallaard, Jonathan J. W., Fraaije, René H. B., Van Bakel, Barry W. M., Nance, John R., Lindholm, Adam & Jagt, John W. M., 2023, New hermit crab species (Anomura, Paguroidea) from the upper Miocene St Marys Formation of Maryland (USA), preserved in their host shells, pp. 389-397 in Zootaxa 5227 (3) on page 390, DOI: 10.11646/zootaxa.5227.3.7, http://zenodo.org/record/7518846, {"references":["Fabricius, J. C. (1775) Systema entomologiae: sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Officina Libraria Kortii, Flensbergi et Lipsiae, xxx + 832 pp.","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundumclasses, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. Editio 10. Reformata. Laurentius Salvius, Holmiae, 824 pp.","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.","Beschin, C., De Angeli, A., Checchi, A. & Zarantonello, G. (2012) Crostacei del giacimento eocenico di Grola presso Spagnago di Cornedo Vicentino (Vicenza, Italia settentrionale) (Decapoda, Stomatopoda, Isopoda). Museo di Archeologia e Scienze Naturali \" G. Zannato \", Montecchio Maggiore, Vicenza, 101 pp.","De Angeli, A. & Caporiondo, F. (2017) I granchi eremiti (Crustacea, Decapoda, Anomura, Paguroidea) dell'Eocene medio di cava \" Main \" di Arzignano (Vicenza, Italia settentrionale). Studi Trentini di Scienze Naturali, 96, 11 - 32. [http: // www. muse. it / it / Editoria-Muse / Studi-Trentini-Storia-Naturale / Documents / STSN _ 95 - 2016. aspx]","Polkowsky, S. & Fraaije, R. H. B. (2019) A new Oligocene hermit crab (Decapoda, Anomura, Paguroidea) from the erratic ' Sternberger Gestein', northern Germany. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen, 291 (1), 61 - 63. https: // doi. org / 10.1127 / njgpa / 2019 / 0789","Lemaitre, R. & McLaughlin, P. (2021 a) World Paguroidea & Lomisoidea database. Pagurus J. C. Fabricius, 1775. World Register of Marine Species. Available from: http: // www. marinespecies. org / aphia. php? p = taxdetails & id = 106854 (accessed 15 March 2021)"]}

Pagurus hazenorum n. sp. (Fig. 1) Zoobank: urn:lsid:zoobank.org:act: 9E06A12D-8371-4F45-B75F-60E3032C4847 Diagnosis. Major carpus (right) covered with small, randomly scattered tubercles, increasing in size toward outer margin, there becoming setose. On minor carpus (left), tubercles arranged in rows and slightly increasing in size toward distal end. Both claws densely covered with large granules, decreasing in size toward outer margin. Outer margins of propodus and dactylus covered with blunt teeth, smaller on dactylus. Outer lateral surface of dactylus with longitudinal, medially elevated surface. Walking legs covered with row of teeth, decreasing in size toward distal end. Dactylus of walking leg only covered by blunt teeth and surface with some longitudinal grooves. Type material. The holotype, and sole specimen known to date, is CMM-I-4785. It is preserved inside its gastropod host shell (Buccinofusus parilis Conrad, 1832), which is severely damaged and allows the hermit crab inside to be observed in more detail. The right and left chelipeds are preserved within the gastropod aperture. A relatively complete walking leg, consisting of carpus, propodus and dactylus, is seen posterior of that aperture. A second walking leg, of which only the carpus is preserved, is found anterior of the aperture, while several fragments of other legs are present just behind the chelipeds. The carapace should have been situated here, but this part of the individual has suffered considerable damage and there is no trace of a carapace. Etymology. In honor of Dr Robert M. Hazen, senior staff scientist at the Carnegie Institution for Science, and his wife, Margaret Hazen, writer and historian. Locality and stratigraphy. Driftwood Beach, Calvert County, Maryland, from the upper Miocene (Tortonian) Little Cove Point Member of the St. Marys Formation in a silty lens within Bed E (Ward & Andrews 2008). Kidwell et al. (2015) identified this level as belonging to “SM-C,” assigned to Shattuck zones 22–23 and part of dinocyst zone 8. Description. Chelipeds stout, broad, with propodus of major (right) claw measuring 21 mm by 14 mm; that of minor (left) claw measuring 16 mm by 12 mm. Entire surface of carpus of major claw covered with small, randomly scattered tubercles, increasing in size and setation toward outer margin. Carpus of minor claw with tubercles arranged in rows and slightly increasing in size toward distal end. Both left and right propodi densely covered with large granules, decreasing in size toward outer margin. Outer margin of propodus arcuate; that of dactylus almost straight. Outer margin of both propodus and dactylus covered with blunt teeth, smaller on latter. Outer lateral surface of dactylus covered with longitudinal, medially elevated ridge. Walking legs covered with row of teeth, decreasing in size toward distal end. Dactylus of walking leg with blunt teeth; surface with some longitudinal grooves. Remarks. The new species compares fairly well with the extant Pagurus impressus (Benedict, 1892), from the west coast of Florida (Provenzano 1959), as well as with Diacanthurus rubricatus Henderson, 1888, from the coast of New Zealand and P. bernhardus (Linnaeus, 1758) from the eastern North Atlantic. Discussion. The assignment of the Maryland material to the genus Pagurus is based on several morphological features which P. hazenorum n. sp. has in common with extant congeners, as described below. Although P. hazenorum n. sp. is preserved in situ within its gastropod shell, the carapace appears to be missing, most likely as a result of the damage to the host shell. Overall, the shape of the dactylus is more elongated and bears a closer cover of granules in P. impressus, whereas that of P. hazenorum n. sp. is stout, with blunt teeth along the outer edge and a cover of large granules. Chelipeds of the present-day D. rubricatus bear small spines, in equal density as in P. hazenorum n. sp. Pagurus bernhardus is comparable as well; this has a coarse ornament which, however, is less dense than that of P. hazenorum n. sp. (see Hyžný & Dulai 2021: fig. 35.8). Our comparison of P. hazenorum n. sp. with both extant and extinct species has yielded numerous forms with closely comparable anatomical features. Molecular and genetic research carried out recently on extant representatives of the genus Pagurus has shown that this is in fact a wastebasket taxon, comprising forms with closely comparable morphologies, but widely divergent genetic structures (e.g., Olguin & Mantelatto 2013; Sultana et al. 2018). Naturally, genetic research cannot be carried out on extinct forms, which makes any workable subdivisions of the genus Pagurus even more difficult. This morphological similarity amongst genetically diverse species is most likely a reflection of functional morphology. Particularly in paguroids, the shell has a marked impact on cheliped shape, and most hermit crabs inhabit comparable mollusks, which explains the closely comparable morphology of the chelipeds. In the fossil record, isolated paguroid chelipeds (mostly propodi) are quite common, whereas carapaces are extremely rare. In view of this, it is highly unlikely that the difficulties surrounding the ‘lump’ taxon Pagurus can be resolved on the basis of extinct forms., Published as part of Wallaard, Jonathan J. W., Fraaije, René H. B., Van Bakel, Barry W. M., Nance, John R., Lindholm, Adam & Jagt, John W. M., 2023, New hermit crab species (Anomura, Paguroidea) from the upper Miocene St Marys Formation of Maryland (USA), preserved in their host shells, pp. 389-397 in Zootaxa 5227 (3) on pages 391-392, DOI: 10.11646/zootaxa.5227.3.7, http://zenodo.org/record/7518846, {"references":["Conrad, T. A. (1832) Fossil shells of the Tertiary formations of North America, illustrated by figures drawn on stone by T. A. Conrad, 1 (2), 21 - 28. [reprinted by Harris, G. D., 1893 and by the Paleontological Research Institution, Ithaca, New York, 1963]","Ward, L. W. & Andrews, G. W. (2008) Stratigraphy of the Calvert, Choptank, and St. Mary's formations (Miocene) in the Chesapeake Bay area, Maryland and Virginia. Virginia Museum of Natural History Memoir, 9, 1 - 60.","Kidwell, S. M., Powars, D. S., Edwards, L. E. & Vogt, P. R. (2015) Miocene stratigraphy and paleoenvironments of the Calvert Cliffs, Maryland. Bulletin of the Geological Society of America, 40, 231 - 279.","Provenzano Jr., A. J. (1959) The shallow-water hermit crabs of Florida. Bulletin of Marine Science, 9 (4), 349 - 420.","Henderson, J. R. (1888) Report on the Anomura collected by H. M. S. \" Challenger \" during the years 1873 - 76. Scientific results of the exploratory voyage of HMS Challenger, Zoology, 27, i - xi + 1 - 221 pp., 21 pls.","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundumclasses, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. Editio 10. Reformata. Laurentius Salvius, Holmiae, 824 pp.","Hyzny, M. & Dulai, A. (2021) Badenian decapods of Hungary, GeoLitera Publishing House, Institute of Geosciences, University of Szeged, Szeged, 300 pp.","Olguin, N. & Mantelatto, F. L. (2013) Molecular analysis validates of some informal morphological groups of Pagurus (Fabricius, 1775) (Anomura: Paguridae) from South America. Zootaxa, 3666 (4), 436 - 448.","Sultana, Z., Asakura, A., Kinjo, S., Nozawa, M., Nakano, T. & Ikeo, K. (2018) Molecular phylogeny of ten intertidal hermit crabs of the genus Pagurus inferred from multiple mitochondrial genes, with special emphasis on the evolutionary relationship of Pagurus lanuginosus and Pagurus maculosus. Genetica, 146 (4), 369 - 381."]}

Genus Paguristes Dana, 1851 Type species. Pagurus weddellii H. Milne Edwards, 1848. Included species. For a listing of fossil taxa, reference is made to Schweitzer et al. (2010), Gagnaison (2012), Beschin et al. (2016, 2018), De Angeli & Caporiondo (2017), Karasawa & Fudouji (2018), Jakobsen et al. (2020), Marangon & De Angeli (2020) and Wallaard et al. (2020). For extant forms, reference is made to Lemaitre & McLaughlin (2021b)., Published as part of Wallaard, Jonathan J. W., Fraaije, René H. B., Van Bakel, Barry W. M., Nance, John R., Lindholm, Adam & Jagt, John W. M., 2023, New hermit crab species (Anomura, Paguroidea) from the upper Miocene St Marys Formation of Maryland (USA), preserved in their host shells, pp. 389-397 in Zootaxa 5227 (3) on page 392, DOI: 10.11646/zootaxa.5227.3.7, http://zenodo.org/record/7518846, {"references":["Dana, J. D. (1851) Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wilkes e classe reipublicae foederata educe, lexit et descripsit. Paguridea. Proceedings of the Academy of Natural Sciences of Philadelphia, 5, 267 - 272.","Milne Edwards, H. (1848) Note sur quelques nouvelles especes du genre Pagure. Annales des Sciences naturelles, Serie 3, Zoologie, 10, 59 - 64.","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.","Gagnaison, C. (2012) Des bernard-l'hermites dans les faluns miocenes de Channay-sur-Lathan (Indre-et-Loire, France). Cossmanniana, 14, 67 - 72.","Beschin, C., Busulini, A., Tessier, G. & Zorzin, R. (2016) I crostacei associati a coralli nell'Eocene inferiore dell'area di Bolca (Verona e Vicenza, Italia nordorientale). In: Memorie del Museo Civico di Storia naturale di Verona. Vol. 2. Sezione Scienze della Terra, 9, pp. 13 - 189.","Beschin, C., Busulini, A., Fornaciari, E., Papazzoni, C. A. & Tessier, G. (2018) La fauna di Crostacei associati a coralli dell'Eocene superiore di Campolongo di Val Liona (Monti Berici, Vicenza, Italia nordorientale). Bollettino del Museo di Storia naturale di Venezia, 69, 129 - 215.","De Angeli, A. & Caporiondo, F. (2017) I granchi eremiti (Crustacea, Decapoda, Anomura, Paguroidea) dell'Eocene medio di cava \" Main \" di Arzignano (Vicenza, Italia settentrionale). Studi Trentini di Scienze Naturali, 96, 11 - 32. [http: // www. muse. it / it / Editoria-Muse / Studi-Trentini-Storia-Naturale / Documents / STSN _ 95 - 2016. aspx]","Karasawa, H. & Fudouji, Y. (2018) Two new species of hermit crabs (Decapoda: Anomura) from the Paleogene Kishima Group, Saga Prefecture, Japan. Bulletin of the Mizunami Fossil Museum, 44, 23 - 28.","Jakobsen, S. L., Fraaije, R. H. B., Jagt, J. W. M. & Van Bakel, B. W. M. (2020) New early Paleocene (Danian) paguroids from deep-water coral / bryozoan mounds at Faxe, eastern Denmark. Geologija, 63, 47 - 56. https: // doi. org / 10.5474 / geologija. 2020.005","Marangon, S. & De Angeli, A. (2020) New hermit crabs (Crustacea, Anomura, Paguroidea) from the Lower Oligocene of the Ligure Piemontese Basin, northwest Italy. In: Jagt, J. W. M., Fraaije, R. H. B., Van Bakel, B. W. M., Donovan, S. K., Mellish, C. & Schweigert, G. (Eds.), A lifetime amongst fossil crustaceans: a tribute to Joseph S. H. Collins (1927 - 2019). Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen, 286 (1 - 2), pp. 157 - 165. https: // doi. org / 10.1127 / njgpa / 2020 / 0895","Wallaard, J. J. W., Fraaije, R. H. B., Jagt, J. W. M., Klompmaker, A. A. & Van Bakel, B. W. M. (2020) The first record of a paguroid shield (Anomura, Annuntidiogenidae) from the Miocene of Cyprus. Geologija, 63 (1), 37 - 43. https: // doi. org / 10.5474 / geologija. 2020.004","Lemaitre, R. & McLaughlin, P. (2021 b) World Paguroidea & Lomisoidea database. Paguristes Dana, 1851. World Register of Marine Species. Available from: http: // www. marinespecies. org / aphia. php? p = taxdetails & id = 106844 (accessed 18 November 2021)"]}

Pagurus hazenorum n. sp. (Fig. 1) Zoobank: urn:lsid:zoobank.org:act: 9E06A12D-8371-4F45-B75F-60E3032C4847 Diagnosis. Major carpus (right) covered with small, randomly scattered tubercles, increasing in size toward outer margin, there becoming setose. On minor carpus (left), tubercles arranged in rows and slightly increasing in size toward distal end. Both claws densely covered with large granules, decreasing in size toward outer margin. Outer margins of propodus and dactylus covered with blunt teeth, smaller on dactylus. Outer lateral surface of dactylus with longitudinal, medially elevated surface. Walking legs covered with row of teeth, decreasing in size toward distal end. Dactylus of walking leg only covered by blunt teeth and surface with some longitudinal grooves. Type material. The holotype, and sole specimen known to date, is CMM-I-4785. It is preserved inside its gastropod host shell (Buccinofusus parilis Conrad, 1832), which is severely damaged and allows the hermit crab inside to be observed in more detail. The right and left chelipeds are preserved within the gastropod aperture. A relatively complete walking leg, consisting of carpus, propodus and dactylus, is seen posterior of that aperture. A second walking leg, of which only the carpus is preserved, is found anterior of the aperture, while several fragments of other legs are present just behind the chelipeds. The carapace should have been situated here, but this part of the individual has suffered considerable damage and there is no trace of a carapace. Etymology. In honor of Dr Robert M. Hazen, senior staff scientist at the Carnegie Institution for Science, and his wife, Margaret Hazen, writer and historian. Locality and stratigraphy. Driftwood Beach, Calvert County, Maryland, from the upper Miocene (Tortonian) Little Cove Point Member of the St. Marys Formation in a silty lens within Bed E (Ward & Andrews 2008). Kidwell et al. (2015) identified this level as belonging to “SM-C,” assigned to Shattuck zones 22–23 and part of dinocyst zone 8. Description. Chelipeds stout, broad, with propodus of major (right) claw measuring 21 mm by 14 mm; that of minor (left) claw measuring 16 mm by 12 mm. Entire surface of carpus of major claw covered with small, randomly scattered tubercles, increasing in size and setation toward outer margin. Carpus of minor claw with tubercles arranged in rows and slightly increasing in size toward distal end. Both left and right propodi densely covered with large granules, decreasing in size toward outer margin. Outer margin of propodus arcuate; that of dactylus almost straight. Outer margin of both propodus and dactylus covered with blunt teeth, smaller on latter. Outer lateral surface of dactylus covered with longitudinal, medially elevated ridge. Walking legs covered with row of teeth, decreasing in size toward distal end. Dactylus of walking leg with blunt teeth; surface with some longitudinal grooves. Remarks. The new species compares fairly well with the extant Pagurus impressus (Benedict, 1892), from the west coast of Florida (Provenzano 1959), as well as with Diacanthurus rubricatus Henderson, 1888, from the coast of New Zealand and P. bernhardus (Linnaeus, 1758) from the eastern North Atlantic. Discussion. The assignment of the Maryland material to the genus Pagurus is based on several morphological features which P. hazenorum n. sp. has in common with extant congeners, as described below. Although P. hazenorum n. sp. is preserved in situ within its gastropod shell, the carapace appears to be missing, most likely as a result of the damage to the host shell. Overall, the shape of the dactylus is more elongated and bears a closer cover of granules in P. impressus, whereas that of P. hazenorum n. sp. is stout, with blunt teeth along the outer edge and a cover of large granules. Chelipeds of the present-day D. rubricatus bear small spines, in equal density as in P. hazenorum n. sp. Pagurus bernhardus is comparable as well; this has a coarse ornament which, however, is less dense than that of P. hazenorum n. sp. (see Hyžný & Dulai 2021: fig. 35.8). Our comparison of P. hazenorum n. sp. with both extant and extinct species has yielded numerous forms with closely comparable anatomical features. Molecular and genetic research carried out recently on extant representatives of the genus Pagurus has shown that this is in fact a wastebasket taxon, comprising forms with closely comparable morphologies, but widely divergent genetic structures (e.g., Olguin & Mantelatto 2013; Sultana et al. 2018). Naturally, genetic research cannot be carried out on extinct forms, which makes any workable subdivisions of the genus Pagurus even more difficult. This morphological similarity amongst genetically diverse species is most likely a reflection of functional morphology. Particularly in paguroids, the shell has a marked impact on cheliped shape, and most hermit crabs inhabit comparable mollusks, which explains the closely comparable morphology of the chelipeds. In the fossil record, isolated paguroid chelipeds (mostly propodi) are quite common, whereas carapaces are extremely rare. In view of this, it is highly unlikely that the difficulties surrounding the ‘lump’ taxon Pagurus can be resolved on the basis of extinct forms.

Family PAGUROPSIDAE n. fam. urn:lsid:zoobank.org:act: 88399E62-3864-4B66-8D89-FC32C562F5A9 TYPE GENUS. — Paguropsis Henderson, 1888. OTHER GENERA INCLUDED. — Eopaguropsis Fraaije, Krzemiński, Van Bakel. Krzemińska & Jagt, 2012 and Paguropsina Lemaitre, Rahayu & Komai, 2018 (Fig. 11). ETYMOLOGY. — The name is derived from the type genus. DIAGNOSIS. — Prominent subtriangular rostrum, considerably exceeding lateral projections.Cervical and branchial grooves subparallel, encompassing small branchial regions (= lateral lobes) or forming broad, sinuous groove. Shield well calcified, subtriangular or subrectangular; dorsal surface slightly vaulted with incomplete midline crest; welldelineated massetic region covered with numerous setal pits; posterior carapace less calcified, delineated cardiac region; uropods and telson symmetrical. Chelipeds subequal, similar in armature and setation.

Fraaije, René H. B., Van Bakel, Barry W. M., Jagt, John W. M., Charbonnier, Sylvain, Schweigert, Guenter, Garcia, Géraldine, Valentin, Xavier (2022): The evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-the-art-report. Geodiversitas 44 (1): 1-16, DOI: 10.5252/geodiversitas2022v44a1

Bucculentum plettenbergense, n. sp. Fig. 1G, 7 Diagnosis. Metabranchial region large, more than half as long as carapace, and a little wider than hepatic region; augenrest round, slightly concave, shifted more dorsally than in other species of genus, and flanked laterally by long, crescent ridge; suborbital spine undivided, curved upwards; epibranchial region very narrow. Etymology. The new species is named after its type locality. Material examined. Holotype MAB 3576, Germany: Plettenberg quarry n. Balingen (N 48 o 21’, E 8 o 80’), coll. RHB Fraaije, Late Oxfordian: Planula Zone (Schweigert & Callomon 1997; Jantschke & Schweigert 2020). Description. The carapace is 8.43 mm long. The rostrum and centro-anterior portion have been split off. Left augenrest complete, large, round, shifted to dorsal side at an angle c. 50 o to the surface of carapace and delimited laterally by a narrow, crescent-like ridge. The suborbital spine is directed upwards (Fig. 7A, B, arrows). The epibranchial region is very narrow (less than 10% of the total lateral length of carapace), with two symmetric swellings over the cardiac region. The cardiac region has three tubercles arranged axially, the distalmost one being the largest, and also the highest point of the carapace. The metabranchial region is characteristically large, a little bit wider than the hepatic region, and more than half as long as the carapace. Remarks. Bucculentum plettenbergense, n. sp., is distinguished because of having the largest metabranchial region amongst all bucculentids, combined with the narrowest epibranchial region. Such proportions are not met in any other congener., Published as part of Krzemińska, Ewa, Starzyk, Natalia, Fraaije, René H. B., Schweigert, Günter & Lukeneder, Alexander, 2021, Jurassic brachyurans of the genus Bucculentum, pp. 395-410 in Zootaxa 5032 (3) on pages 402-403, DOI: 10.11646/zootaxa.5032.3.5, http://zenodo.org/record/5497036, {"references":["Schweigert, G. & Callomon, J. H. (1997) Der bauhini - Faunenhorizont und seine Bedeutung fur die Korrelation zwischen tethyalem und subborealem Oberjura. Stuttgarter Beitrage zur Naturkunde (B), 247, 1 - 69.","Jantschke, H. & Schweigert, G. (2020) The semimammatum and semiarmatum ammonite biohorizons (Late Oxfordium, Hypselum Zone) in the Upper Jurassic of SW Germany. Palaeodiversity, 13, 89 - 130. https: // doi. org / 10.18476 / pale. v 13. a 9"]}

Bucculentum bucculentum (Wehner, 1988) Fig. 1A, B, 2, 3 1988 Nodoprosopon bucculentum Wehner, p. 53, pl. 4, figs. 1–2, 6. 2009 Bucculentum bucculentum (Wehner, 1988) in: Schweitzer & Feldmann, 2009a: p. 79, fig. 3.3–3.4. Diagnosis. After Schweitzer & Feldmann (2009), amended. Carapace from longer than wide to almost quadrangular, widest across hepatic regions; rostrum tri-lobed, armoured with spines, central lobe being longest and depressed; augenrest positioned perpendicular to body long axis, laterally flanked by short ridge; suborbital spine strong, single, with several sharp terminal spines; cardiac region with three tubercles arranged along body long axis, distalmost tubercle being largest. Age and distribution. From at least Middle Oxfordian (Poland) to Early Kimmeridgian: Platynota-Divisum zones (Germany); the specimen SMNS 66654 described here as „ Bucculentum cf. bucculentum ” would extend the range of this species to the Late Kimmeridgian Pseudomutabilis Zone. Materials examined. Holotype SNS̲-̲SPG 1980 XXX 1255 (Fig. 2A, B), Biburg (Germany), age Early Kimmeridgian: Planula Zone (Schairer 1982; Schweitzer & Feldmann 2009a). Paratype SNS̲-̲SPG 1980 XXX 1254 (Fig. 2C), same locality and age. Specimen SNS̲-̲SPG 1987 I 55 (Fig. 2D, E), Unterwilflingen (Germany), age Early Kimmeridgian: Platynota-Divisum zones (Wehner 1988). Specimens from the Polish Jura Chain described and illustrated by Starzyk et al. (2012) were re-examined for this study; they come from two quarries: Ogrodzieniec (Lower and Middle Oxfordian) and Niegowonice (from Cordatum to Transversarium zones, except discontinuous Mariae Zone; Głowniak 2006). New material: MAB 3572 (Fig. 3 E, F), Geisingen (Germany; N 47 o 93’, E 8 o 67’), age Late Kimmeridgian: Mutabilis/Acanthicum Zone (Van Bakel et al. 2008); MAB 3577, MAB 3579 (Fig. 3G), MAB 3582a (positive, Fig. 3A) and MAB 3582b (negative, Fig. 3B, C, D), Plettenberg quarry n. Balingen, Germany; all coll. R. H. B. Fraaije), age Early Kimmeridgian, Planula Zone (Schweigert & Callomon 1997; Jantschke & Schweigert 2020); I-F/MP/6210/1578/11 (ISEA) and I-F/MP/4648/1534/08 (ISEA), Wysoka n. Łazy (N 50 o 25’44’’, E 19 o 21’13’’), Polish Jura Chain, age Middle Oxfordian (Głowniak 2002). Additional description. The rostrum is tri-lobed, the central lobe being the longest and directed downwards while the two flanking lobes are raised; together, they form a kind of a scoop armoured with terminal rows of sharp spines. The entire rostrum, or almost so, is preserved only in the specimens MAB 3572 from Plettenberg and I-F/ MP/6210/1578/11 from Poland (Fig. 3E, H, respectively), but it is fragmentarily present in several other specimens, including the holotype (Fig. 2A, B) and specimen SNSB-BSPG 1987 I 55 (Fig. 2D, E). The augenrest is convex and directed frontally, i.e, at an angle c. 90 o to the body long axis. The suborbital spine is single, massive, and oval in cross section. The distal portion of this spine is armoured with few teeth (traces of four teeth are visible in two specimens, Fig. 2B and 3G). The epibranchial region of the holotype (Fig. 2A) and MAB 3572 has six ridges ending in rows of tubercles, fan-like spreading just over the cardiac region; in other specimens this feature is weakly expressed or absent. Along the lateral sides there are strong spines (Fig. 3D, F, G). Remarks. Noteworthy is the specimen MAB 3852, preserved in positive (convex) and negative (concave) parts (Fig. 3A, B, resp.), which supplement each other with details; the concave mould of the rostrum with white depressions left by the warts is completely preserved in the negative part (Fig. 3C); this part retained also the traces of lateral spines along the metabranchial region (Fig. 3D), and almost complete cover of white cuticle, visible from the inner side. The specimen MAB 3572 (Fig. 3F) shows that the pattern of tubercles is preserved both on the cuticle and on the surface of the mould beneath., Published as part of Krzemińska, Ewa, Starzyk, Natalia, Fraaije, René H. B., Schweigert, Günter & Lukeneder, Alexander, 2021, Jurassic brachyurans of the genus Bucculentum, pp. 395-410 in Zootaxa 5032 (3) on pages 396-400, DOI: 10.11646/zootaxa.5032.3.5, http://zenodo.org/record/5497036, {"references":["Wehner, G. (1988) Uber die Prosopiden (Crustacea, Decapoda) des Jura. Inaugural-Dissertation zur Erlangung des Doktorgrades der Fakultat fur Geowissenschaften der Ludwig-Maximilians-Universitat zu Munchen, Munchen, 154 pp.","Schairer, G. & Yamani, S. - A. (1982) Die Schwammkalke von Biburg bei Weissenburg / Bayern (Oberoxford, Sudliche Frankenalb). Allgemeine Ubersicht. Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und historische Geologie, 22, 9 - 17.","Glowniak, = E. (2006) Correlation of the zonal schemes at the Middle-Upper Oxfordian (Jurassic) boundary in the Submediterranean Province: Poland and Switzerland. Acta Geologica Polonica, 56, 33 - 50.","Bakel, B. W. M. van, Fraaije, R. H. B., Jagt, J. W. M. & Artal, P. (2008) An unexpected diversity of Late Jurassic hermit crabs (Crustacea, Decapoda, Anomura) in Central Europe. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 250, 137 - 156. https: // doi. org / 10.1127 / 0077 - 7749 / 2008 / 0250 - 0137","Schweigert, G. & Callomon, J. H. (1997) Der bauhini - Faunenhorizont und seine Bedeutung fur die Korrelation zwischen tethyalem und subborealem Oberjura. Stuttgarter Beitrage zur Naturkunde (B), 247, 1 - 69.","Jantschke, H. & Schweigert, G. (2020) The semimammatum and semiarmatum ammonite biohorizons (Late Oxfordium, Hypselum Zone) in the Upper Jurassic of SW Germany. Palaeodiversity, 13, 89 - 130. https: // doi. org / 10.18476 / pale. v 13. a 9","Glowniak, E. (2002) The ammonites of the family Perisphinctidae from the Plicatilis Zone (lower Middle Oxfordian) of the Pol- ish Jura Chain (Central Poland); their taxonomy, phylogeny and biostratigraphy. Acta Geologica Polonica, 52, 307 - 364."]}

Genus Bucculentum Schweitzer & Feldmann, 2009 Diagnosis. After Schweitzer & Feldmann (2009), amended. Carapace roughly rectangular; rostrum tri-lobed, projected well beyond frontal margin of carapace; orbit appearing to be placed under rostrum, augenrest on hepatic region of dorsal carapace, bounded by spines or ridges; hepatic region strongly inflated; groove (named here augenrest groove) leading from mid dorsal margin of augenrest to cervical groove; cervical and branchio-cardiac grooves deep; urogastric region not well defined, depressed below level of metagastric and cardiac regions; cardiac region elevated, its dominant tubercle making highest point of carapace. Type species: Nodoprosopon bucculentum Wehner, 1988., Published as part of Krzemińska, Ewa, Starzyk, Natalia, Fraaije, René H. B., Schweigert, Günter & Lukeneder, Alexander, 2021, Jurassic brachyurans of the genus Bucculentum, pp. 395-410 in Zootaxa 5032 (3) on page 396, DOI: 10.11646/zootaxa.5032.3.5, http://zenodo.org/record/5497036, {"references":["Wehner, G. (1988) Uber die Prosopiden (Crustacea, Decapoda) des Jura. Inaugural-Dissertation zur Erlangung des Doktorgrades der Fakultat fur Geowissenschaften der Ludwig-Maximilians-Universitat zu Munchen, Munchen, 154 pp."]}

Bucculentum horstkuscheli, n. sp. Fig. 1E, F, 6 Diagnosis. Hepatic regions large, much wider than metabranchial regions; metabranchial region not expanded laterally; augenrest directed laterad at c. 60 o to body long axis and aligned with proximal portions of hepatic region; suborbital spine wide, directed outwards and downwards, and ending in two short, triangular teeth; cardiac region with one distalmost dominant tubercle accompanied anteriorly by row of small tubercles. Etymology. Trivial name is dedicated to Horst Kuschel (Faurndau), who found and donated the holotype of this species. Material examined. Holotype: # SMNS 70607 (leg. H. Kuschel), Germany: Bad Überkingen-Oberböhringen, near Geislingen a. d. Steige. Age: Late Kimmeridgian: Acanthicum Zone (Fraaije et al. 2017). Description. The holotype represents an internal mould, with only the traces of the white cuticle in the grooves. It is excellently preserved and symmetrical, lacking only the rostrum. The hepatic regions are very wide, c. 1.4x wider than the mesobranchial region and 1.2x wider than the metabranchial region. The frontal portions containing the augenrests and anterior portions of hepatic regions are aligned, obliquely set to the long axis at an angle of 57 o, symmetrical on both sides. Augenrests are almost perpendicular to the plane of carapace, only very slightly shifted dorsally (Fig. 6B), at an angle of c. 80 o to the upper surface of the carapace; they are laterally flanked by single, broad, triangular tubercles. The suborbital spine is very broad, supporting most of the lower margin of the augenrest, directed down- and outwards, and terminating in two short, triangular teeth (Fig. 6B, C, arrows). The epibranchial region is narrow; the cervical groove is almost straight. The cardiac region is equipped at its distal end with a large tubercle (which is the highest point of the carapace) and a median row of small tubercles towards the mesobranchial region. The metabranchial regions are narrow, almost rectangular; the distal end is broad, stretched over the entire distal portion of the metabranchial region and upturned. Remarks. The new species resembles B. bachmayeri in having a large hepatic region, a divided suborbital spine, and a cardiac region with one large tubercle, but differs in having the augenrests directed outwards and positioned obliquely to body long axis. The suborbital spine is divided into two short, triangular lobes, unlike the long, thin prongs in B. bachmayeri.

Bucculentum horstkuscheli, n. sp. Fig. 1E, F, 6 Diagnosis. Hepatic regions large, much wider than metabranchial regions; metabranchial region not expanded laterally; augenrest directed laterad at c. 60 o to body long axis and aligned with proximal portions of hepatic region; suborbital spine wide, directed outwards and downwards, and ending in two short, triangular teeth; cardiac region with one distalmost dominant tubercle accompanied anteriorly by row of small tubercles. Etymology. Trivial name is dedicated to Horst Kuschel (Faurndau), who found and donated the holotype of this species. Material examined. Holotype: # SMNS 70607 (leg. H. Kuschel), Germany: Bad Überkingen-Oberböhringen, near Geislingen a. d. Steige. Age: Late Kimmeridgian: Acanthicum Zone (Fraaije et al. 2017). Description. The holotype represents an internal mould, with only the traces of the white cuticle in the grooves. It is excellently preserved and symmetrical, lacking only the rostrum. The hepatic regions are very wide, c. 1.4x wider than the mesobranchial region and 1.2x wider than the metabranchial region. The frontal portions containing the augenrests and anterior portions of hepatic regions are aligned, obliquely set to the long axis at an angle of 57 o, symmetrical on both sides. Augenrests are almost perpendicular to the plane of carapace, only very slightly shifted dorsally (Fig. 6B), at an angle of c. 80 o to the upper surface of the carapace; they are laterally flanked by single, broad, triangular tubercles. The suborbital spine is very broad, supporting most of the lower margin of the augenrest, directed down- and outwards, and terminating in two short, triangular teeth (Fig. 6B, C, arrows). The epibranchial region is narrow; the cervical groove is almost straight. The cardiac region is equipped at its distal end with a large tubercle (which is the highest point of the carapace) and a median row of small tubercles towards the mesobranchial region. The metabranchial regions are narrow, almost rectangular; the distal end is broad, stretched over the entire distal portion of the metabranchial region and upturned. Remarks. The new species resembles B. bachmayeri in having a large hepatic region, a divided suborbital spine, and a cardiac region with one large tubercle, but differs in having the augenrests directed outwards and positioned obliquely to body long axis. The suborbital spine is divided into two short, triangular lobes, unlike the long, thin prongs in B. bachmayeri., Published as part of Krzemińska, Ewa, Starzyk, Natalia, Fraaije, René H. B., Schweigert, Günter & Lukeneder, Alexander, 2021, Jurassic brachyurans of the genus Bucculentum, pp. 395-410 in Zootaxa 5032 (3) on page 402, DOI: 10.11646/zootaxa.5032.3.5, http://zenodo.org/record/5497036, {"references":["Fraaije, R. H. B., Schweigert, G. & Van Bakel, B. W. M. (2017) New Late Jurassic (Kimmeridgian) paguroids (Crustacea: Decapoda: Anomura) from the Swabian Alb, southern Germany. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 285 (1), 117 - 121. https: // doi. org / 10.1127 / njgpa / 2017 / 0673"]}

Bucculentum plettenbergense, n. sp. Fig. 1G, 7 Diagnosis. Metabranchial region large, more than half as long as carapace, and a little wider than hepatic region; augenrest round, slightly concave, shifted more dorsally than in other species of genus, and flanked laterally by long, crescent ridge; suborbital spine undivided, curved upwards; epibranchial region very narrow. Etymology. The new species is named after its type locality. Material examined. Holotype MAB 3576, Germany: Plettenberg quarry n. Balingen (N 48 o 21’, E 8 o 80’), coll. RHB Fraaije, Late Oxfordian: Planula Zone (Schweigert & Callomon 1997; Jantschke & Schweigert 2020). Description. The carapace is 8.43 mm long. The rostrum and centro-anterior portion have been split off. Left augenrest complete, large, round, shifted to dorsal side at an angle c. 50 o to the surface of carapace and delimited laterally by a narrow, crescent-like ridge. The suborbital spine is directed upwards (Fig. 7A, B, arrows). The epibranchial region is very narrow (less than 10% of the total lateral length of carapace), with two symmetric swellings over the cardiac region. The cardiac region has three tubercles arranged axially, the distalmost one being the largest, and also the highest point of the carapace. The metabranchial region is characteristically large, a little bit wider than the hepatic region, and more than half as long as the carapace. Remarks. Bucculentum plettenbergense, n. sp., is distinguished because of having the largest metabranchial region amongst all bucculentids, combined with the narrowest epibranchial region. Such proportions are not met in any other congener.

Bucculentum bachmayeri Schweitzer & Feldmann, 2009 Fig. 1C, D, 5 2009a Bucculentum bachmayeri in: Schweitzer & Feldmann, Ann. Naturhist. Mus. Wien 110 A: 81, fig. 1.4, 3.1, 3.2. Diagnosis. Carapace widest across hepatic regions which are more expanded laterally than in Bucculentum bucculentum, but flattened (in frontal view); augenrest positioned perpendicular to body long axis, bounded by small tubercles on outer-orbital edge; suborbital spine bifid, its two prongs being thin and long; cardiac region small, ovate, with one dominant, sharp tubercle. Material examined. Holotype, NHMW 1990 /0041/3376, near Dörfles, Vienna, Ernstbrunn Limestone, Tithonian. Specimen SMNS 70606: Bad Überkingen-Oberböhringen near Geislingen an der Steige, southern Germany (N 48 o 61’, E 9 o 79’); Upper Kimmeridgian, Pseudomutabilis Zone (leg. H. Kuschel). Additional description. The frontal view of the holotype exposes a very characteristic feature of this species: the hepatic regions are broad and flattened when compared to B. bucculentum, so that the distance between the upper margin of the augenrest and the surface of the hepatic region over it is very short (marked with an arrow in Fig. 5A); the same character is visible in the specimen SMNS 70606 (Fig. 5C, arrow). The area of the augenrest is flat and smaller than in B. bucculentum; the plane of the augenrest is perpendicular to the long body axis, similar to B. bucculentum. The suborbital spine in the holotype is bifid, and its two prongs are thin (Fig. 5B); in the specimen SMNS 70606 these spines are not preserved. The lateral sides of carapace are endowed with strong, short spurs partially preserved in SMNS 70606 (Fig. 5E, F). The known specimens delimit the range of this species to Late Kimmeridgian-Tithonian., Published as part of Krzemińska, Ewa, Starzyk, Natalia, Fraaije, René H. B., Schweigert, Günter & Lukeneder, Alexander, 2021, Jurassic brachyurans of the genus Bucculentum, pp. 395-410 in Zootaxa 5032 (3) on pages 401-402, DOI: 10.11646/zootaxa.5032.3.5, http://zenodo.org/record/5497036

Fraaije, René H. B., Van Bakel, Barry W. M., Jagt, John W. M., Machalski, Marcin (2015): A new hermit crab (Anomura, Paguroidea) from the upper Albian (Cretaceous) of Annopol, Poland. Zootaxa 3955 (4): 588-594, DOI: http://dx.doi.org/10.11646/zootaxa.3955.4.9

Family Retroplumidae Gill, 1894, Published as part of Artal, Pedro, Van Bakel, Barry W. M., Fraaije, Ren�� H. B. & Jagt, John W. M., 2013, New retroplumid crabs (Crustacea, Brachyura, Retroplumidae Gill, 1894) from the Eocene of Huesca (Arag��n, Spain), pp. 343-352 in Zootaxa 3652 (3) on page 344, DOI: 10.11646/zootaxa.3652.3.3, http://zenodo.org/record/221749

Gaudipluma n. gen. Type species. Gaudipluma bacamortensis n. gen., n. sp. Diagnosis. Carapace large; subrectangular, wider than long, maximum width posteriorly; carapace strongly compressed dorso-ventrally; orbits large, supraorbital margins markedly sinuous; front narrow, relatively long; lateral margins arched, with 2 marked raised lateral nodes; posterior margin convex, wider than orbitofrontal margin; dorsal surface flat in both directions, with 3 subtle, low ridges; thoracic sternum with sternites 3, 4 trapezoidal, sternites 5–7 subrectangular, sternites 6, 7 with prominent median line; sternite 8 inclined, reduced; vulvae subcircular, at extremities of sutures 5 / 6; male, female abdomens weakly differentiated, narrow; chelipeds subequal; P 2 –P 4 long, with numerous spines; P 5 conspicuously reduced; dorsal surface with numerous large pits. Etymology. In honour of the Catalan architect Antoni Gaudí (1852–1926), in allusion to the shape and ornament of the new taxon which is defined by sinuous lines, reminiscent of his works, plus the ending - pluma, which refers to the main character of the family. Remarks. The new genus can be differentiated from all others genera in the family Retroplumidae (see De Grave et al. 2009: 44) by a few unique dorsal and ventral characters. The dorsal ridges are fairly attenuated, lateral margins have two conspicuous lateral nodes, male and female abdomens are particularly narrow, the male abdominal segment 6 is long, sternite 7 is unusually large for the family, and distinctly long. Distinguishing features also include a dorsal surface which is densely covered by large pits and P 2 –P 4 with numerous spines, characters never seen in any other retroplumid genera. Gaudipluma bacamortensis n. sp. (Figs. 3–4) Diagnosis. As for genus. Etymology. From Bacamorta, the name of the village from where the material originates. Material examined. MGSB 75283, holotype; MGSB 75284, 75285 a, b and 75286 a, b, paratypes; MAB k. 3282–3283, paratypes, all from the vicinity of Bacamorta (Huesca), of early Eocene (Ypresian) age. In the holotype, the maximum carapace width and length (in millimetres) are 39 and 30, respectively. Description. Carapace large (maximum carapace width about 40 mm); subrectangular in outline; wider than long, widest posteriorly, at level of metabranchial region. Dorsal surface flat in both directions, densely covered by large pits. Orbits large, supraorbital margins sinuous, median sinus projected anteriorly; supraorbital margins terminating in long outer-orbital spines. Front slender, directed anteriorly, relatively long. Lateral carapace margins broadly arched, with 2 conspicuously raised nodes, the first at level of epibranchial region, the second at metabranchial region; posterior margin convex, wider than orbitofrontal margin. Dorsal carapace surface crossed by 3 subtle ridges; anterior ridge continuous, sinuous; median ridge inclined (oblique), interrupted by gastric grooves; posterior ridge continuous. Mesobranchial region subcircular, bounded by subtle grooves. Epibranchial regions slightly swollen, inclined (oblique), interrupted by shallow gastric grooves. Cardiac region weakly defined, slightly raised, forming continuous faint ridge with swollen metabranchial region. Intestinal region depressed. Buccal frame large, subrectangular, mxp 3 with large, robust endopodite, narrow exopodite. Thoracic sternum wider than long, sternites 1, 2 subtriangular, small, sternites 3, 4 subtrapezoidal; strong, notable notch between sternites 3, 4; sternite 4 with conspicuous lateral projection; sternites 5–7 subrectangular, wide; sternite 7 longer than 5, 6; sternites 6, 7 with prominent median line; sternite 8 inclined, conspicuously reduced; vulvae subcircular, at extremities of sutures 5 / 6. Male, female abdomens narrow, sexual dimorphism weak, telson short, small in both sexes; female abdomen subelliptical to subtriangular, with rounded lateral margins, all segments separated; male abdomen subtriangular, with straight, inclined lateral margins; abdominal segment 6 conspicuously long, with clear lateral extensions; male abdominal segments 3–5 fused. Abdominal holding mechanism clearly indicated by lateral projections in sternite 5, lateral extensions of abdominal segment 6. Female vulvae at extremity of suture 5 / 6 (Fig. 4 F). P 5 subequal, dactyli with numerous small denticles. P 2 –P 4 long, flattened dorso-ventrally, lateral sides with numerous spines. P 5 conspicuously reduced (Fig. 34 D, E). Dorsal carapace surface densely covered by large pits. Remarks. The main features of Gaudipluma bacamortensis n. gen., n. sp. match those attributed to other retroplumids (de Saint Laurent 1989: 111–112). Diagnostic features of the new taxon include the general shape of the carapace, large orbits with sinuous margins, narrow front, general shape of the thoracic sternum, with subtrapezoidal sternites 3, 4, rectangular sternites 5–7, an inclined and reduced sternite 8, female vulvae at the extremity of sutures 5 / 6, P 2 –P 4 long and flattened; P 5 conspicuously reduced (Fig. 4 D, E). The similarly sized Retrocypoda almelai has a different dorsal surface, with more numerous and better defined dorsal ridges, much wider abdomens, with marked sexual dimorphism, a shorter abdominal segment 6 and longer telson (Via 1969: fig. 41). The female abdomen in Retrocypoda is conspicuously large, occupying a large portion of the ventral surface (Via 1969: pl. 38, fig. 4 b). Similar features are seen in Retropluma (see Beschin et al. 1996) as far as male and female abdomens are concerned. The dorsal carapace surface in Retropluma has more strongly marked, acute ridges. The male and female abdomens of Gaudipluma n. gen. are markedly narrow, and sexual dimorphism is weak. The male abdominal segment 6 is unusually long. The male and female telsons are short in the new genus. Serrablopluma n. gen., Loerentheya and Loerenthopluma all differ mainly in carapace size (smaller), general outline, the construction of the orbits, and all the specific features indicated in the discussion of Serrablopluma n. gen. The female abdomen in Serrablopluma n. gen., Loerenthopluma and Retrocypoda is fairly broad, with distinct sexual dimorphism (see Beschin et al. 1996). Costacopluma is characterised by a wide geographical distribution (America, Africa) and a long stratigraphic range (Coniacian–Eocene; see Beschin et al. 1996; Schweitzer et al. 2010). The ten species currently assigned to this genus exhibit a wide range of morphological features, with variable carapace sizes and shapes (subcircular to subelliptical to widely sub-rectangular or elongate). The main distinguishing features of Costacopluma are the stronger dorsal carapace areolation, shorter orbitofrontal margins, wider abdomens and more accentuated sexual dimorphism. The dorsal surfaces of the carapace are more convex and marginal angles are more rounded. The dorsal carapace regions are clearly separated; moreover, the front is much wider and the orbitofrontal margins much shorter (Schweitzer et al. 2003: fig. 16).

Genus Rogueus Berglund & Feldmann, 1989 Rogueus Berglund & Feldmann, 1989: 70. Type species. Rogueus orri Berglund & Feldmann, 1989, by monotypy. Species included. Rogueus orri, and Rogueus robustus Collins & Jakobsen, 1996. Remarks. The genus has so far been recorded from the Paleocene of Denmark and the lower middle Eocene of Vancouver Island, British Columbia, Canada. Rogueus sp. (sensu Ludvigsen & Beard 1998: 125; Trent River Formation, Upper Cretaceous, Vancouver Island, Canada), has subsequently been described as a new genus and species, Bicornisranina bocki, by Nyborg & Fam (2008: 689, figs. 3, 4t, 5, 6). Bicornisranina should be placed in the subfamily Raninoidinae Lőrenthey in Lőrenthey & Beurlen, 1929 (see below), while Rogueus is retained in Lyreididae., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 84, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Berglund, R. E. & Feldmann, R. M. (1989) A new crab, Rogueus orri n. gen. and sp. (Decapoda: Brachyura), from the Lookingglass Formation (Ulatisian Stage: lower Middle Eocene) of southwestern Oregon. Journal of Paleontology, 63, 69 - 73.","Collins, J. S. H. & Jakobsen, S. L. (1996) A new crab, Rogueus robustus, from the Middle Palaeocene of Denmark. Bulletin of the Mizunami Fossil Museum, 22 (1995), 61 - 65.","Ludvigsen, R. & Beard, G. (1998) West Coast fossils: A Guide to the Ancient Life of Vancouver Island, 194 pp. Whitecap Books, Vancouver.","Nyborg, T. G. & Fam, J. (2008) Bicornisranina bocki, n. gen., n. sp. (Decapoda: Raninidae) from the Cretaceous of Vancouver Island, British Columbia, Canada. Journal of Crustacean Biology, 28, 686 - 694.","Lorenthey, E. & Beurlen, K. (1929) Die fossilen Dekapoden der Lander der Ungarischen Krone. Geologica Hungarica, Series Palaeontologica, 3, 1 - 420, 16 pls."]}

Genus Pseudorogueus Fraaye, 1995 Pseudorogueus Fraaye, 1995: 66. Type species. Pseudorogueus rangiferus Fraaye, 1995, by original designation. Material examined. Holotype, and only specimen so far known, a partly decorticated carapace (MAB k. 1040). Locality and stratigraphy were not included in the original description; the specimen was recovered from a roadcutting along the road Ager-Tremp on Serra del Pi, 1 km northwest of the village of La Baronia, northern Spain, from the top of the Ei2 Member of the ‘Gresos deltaics’, of early Eocene age (Rosell & Llompart 1988). Remarks. Pseudorogueus rangiferus (see Fraaye 1995: 66, figs. 1, 2) should be included in the subfamily Raninoidinae. The frontal characters do not match those of Rogueus Berglund & Feldmann, 1989, or any other member of Lyreididae for that matter. According to Tucker (1998: 359) ‘the specimen described by Fraaye (1995) is more closely related to the Raninoidinae clade, not the Lyreidinae which includes Rogueus ’. Pseudorogueus was synonymised with Raninoides by Tucker (1998: 359), but considered valid by Nyborg & Fam (2008: 689), the latter decision we agree with. The wide carapace and strong anterolateral spine with multiple anteriorly directed spinules warrant generic separation of Pseudorogueus and Raninoides., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 100-101, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Fraaye [sic], R. H. B. (1995) A new raninid crab, Pseudorogueus rangiferus (Decapoda, Crustacea), from the Eocene of Spain. Estudios Geologicos, 51, 65 - 67.","Rosell, J. & Llompart, C. (1988) Guia Geologica del Montsec i de la Vall d'Ager, 168 pp., Martin C. E. C., Barcelona.","Berglund, R. E. & Feldmann, R. M. (1989) A new crab, Rogueus orri n. gen. and sp. (Decapoda: Brachyura), from the Lookingglass Formation (Ulatisian Stage: lower Middle Eocene) of southwestern Oregon. Journal of Paleontology, 63, 69 - 73.","Tucker, A. B. (1998) Systematics of the Raninidae (Crustacea: Decapoda: Brachyura), with accounts of three new genera and two new species. Proceedings of the Biological Society of Washington, 111, 320 - 371.","Nyborg, T. G. & Fam, J. (2008) Bicornisranina bocki, n. gen., n. sp. (Decapoda: Raninidae) from the Cretaceous of Vancouver Island, British Columbia, Canada. Journal of Crustacean Biology, 28, 686 - 694."]}

Bournelyreidus carlilensis (Feldmann & Maxey, 1980) n. comb. Raninella carlilensis Feldmann & Maxey, 1980: 858. Remarks. Raninella carlilensis (see Feldmann & Maxey 1980: 858, text-figs. 1–3), as based on four specimens from the middle Turonian of Kansas (U.S.A.), is here reassigned to Bournelyreidus n. gen. The ventral sides of all specimens is poorly preserved, but the large mxp3 coxae, intercalated between the thoracic sternum and pterygostome, are well visible (Feldmann & Maxey 1980: 859, text-fig. 1A). There is a narrow junction between sternite 4 and the pterygostome, despite the fact that this is not sufficiently well-preserved in the type series. The rostrum is excavated; its tip and the outer orbital corners may be incompletely preserved in the type series. The pterygostome is tumid, ‘possessing two fairly deep sulci’ (Feldmann & Maxey 1980: 859; see also their text-fig. 1C), and the carapace bears two slender anterolateral spines. Raninella carlilensis, R. tridens and R. oaheensis appear to have a similar rostral region (Feldmann & Maxey 1980: 860, 861)., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 79, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Feldmann, R. M. & Maxey, M. (1980) Raninella carlilensis, a new raninid crab from the Carlle Shale (Turonian) of Kansas. Journal of Paleontology, 54, 858 - 861."]}

Cenomanocarcinus cantabricus n. sp. (Fig. 17E, F) Diagnosis. Carapace of medium size for genus, dorsal surface weakly convex in both directions; anterolateral margins arched, with 4 teeth of similar size, additional larger tooth corresponding to epibranchial region; posterolateral margins with 2 teeth, anteriormost close to epibranchial tooth; dorsal grooves weakly marked; dorsal regions with few, coarse, blunt tubercles; hepatic region without clear tubercles. Derivation of name. In reference to Cantabria, the region in northern Spain from where specimens of the new species were collected. Material examined. Holotype, a complete carapace with the majority of cuticle preserved (MGSB75431) (ex Manuel Díaz Collection); paratype (MGSB75423) (ex Pedro Artal Collection), a carapace with fragments of cuticle preserved, both from the Cabo de Ajo sea cliff, within the township of Bareyo (Cantabria) from levels considered to be of Albian age (Baron-Szabo & Fernández-Mendiola 1997). Description. Carapace medium sized for genus, subcircular in outline, wider than long; maximum width at epibranchial spine; front advanced, trilobate, with 2 additional blunt teeth at inner orbital corner; orbits small, directed forwards, with 2 short fissures in supraorbital margin, outer orbital node robust, fairly salient; anterolateral margin long, arched, with 5 teeth (excluding outer orbital node, including epibranchial spine), anterior 4 stout, fairly salient, of similar size; epibranchial tooth not completely preserved, appears to have been strong; posterolateral margin longer, converging backwards, sharp edged, with 2 small teeth, anterior one close to epibranchial node, second near posterior corner; posterior margin axially concave, defined by fine rim behind a narrow groove, slightly wider than orbitofrontal margin; dorsal carapace regions weakly vaulted, defined by shallow grooves, with coarse, blunt tubercles; protogastric region with 2 small tubercles; hepatic region barely defined with indistinct inflations, no evident tubercles present; mesogastric region with 2 elongated tubercles; epibranchial region with faint ridge, with large medial tubercle, additional barely inflated lateral tubercle; postbranchial regions with 2 longitudinally positioned tubercles, anterior one larger, rounded, weakly raised, posterior one close to end of lateral margin; cardiac region relatively broad, defined by elongated tubercle, bounded by shallow lateral grooves; intestinal region small, somewhat depressed, bounded by 2 small tubercles; cervical groove weakly marked, even axially, but clearly notching anterolateral margin; branchiocardiac groove well defined at inner portion of epibranchial region. Ventral parts or appendages not preserved. Dorsal carapace surface with dense, small granules. Remarks. The new species is assigned to Cenomanocarcinus on the basis of the curvature and armature of the anterolateral margin, the strong epibranchial spine, the narrow orbitofrontal margin, triangular five-spined rostrum, division of coarse tubercles on the dorsal carapace, and possession of three ridges on the posterior carapace. The new species can be clearly distinguished from congeners on account of the shallow, weakly defined dorsal grooves; the longitudinal and transverse ridges being only weakly salient; the dorsal surface with few dorsal tubercles; the lack of tubercles in the hepatic region; the posterior half of the carapace with large, blunt tubercles; and the anterior tooth of the posterolateral margin being positioned close to the epibranchial spine. The only other European species, C. inflatus, shows more marked transverse and longitudinal ridges with more numerous, evenly spaced tubercles and more convex branchial ridges. There is also a strong posterolateral tooth at mid-length, instead of being closer to the epibranchial tooth; there is an additional anterolateral tooth and the hepatic region bears several clear tubercles. Of the American species, C. beardi exhibits a more subhexagonal carapace, with more divergent posterolateral margins, more salient longitudinal and transverse ridges, stronger dorsal tubercles, a longer epibranchial and posterior anterolateral spine and more distinct H-shaped grooves in the cardiac region. Cenomanocarcinus vanstraeleni exhibits a more subhexagonal carapace; more salient transverse and longitudinal ridges, with more numerous tubercles; the antero- and posterolateral margins with numerous small denticles. Cenomanocarcinus oklahomensis is characterised by stronger longitudinal and transverse ridges, the H-shaped groove pattern in the cardiac region being strongly marked; C. pierrensis exhibits a more distinct, continuous cervical groove; the dorsal tubercles are more raised and conical; the branchial ridges are more salient; the tubercles in the posterior branchial region are not lined up. Cenomanocarcinus pierrensis exhibits a mixed set of characters, which make it difficult to ascribe this species with certainty to Cenomanocarcinus or Necrocarcinus. More completely preserved material is needed to decide in this matter; for the time being, it is retained in the former genus. Two species from Nigeria were recently described by Collins (2010). Cenomanocarcinus tenuicarinatus, of early Turonian age, has a more elongated carapace; posterolateral margins lack strong teeth and clearly tuberculate hepatic ridges; the axial carina is more salient and continuous, less tuberculate, whereas C. dissimilis, from the lower Cenomanian, exhibits a wider carapace with more distinct longitudinal ridges, clear hepatic tubercles and a narrower cardiac region., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 54-55, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Baron-Szabo, R. C. & Fernandez-Mendiola, P. A. (1997) Cretaceous scleractinian corals from the Albian of Cabo de Ajo (Cantabria Province, N-Spain). Palaontologische Zeitschrift, 71, 35 - 50.","Collins, J. S. H. (2010) New species of crabs (Crustacea, Decapoda), one from the Middle Danian of Denmark, and three new species from the Upper Cretaceous of Nigeria. Bulletin of the Mizunami Fossil Museum, 36, 13 - 19."]}

Symethoides monmouthorum n. sp. Notopocorystes cf. N. tridens — Landman et al. 2007, p. 29, fig. 15N, O. (Fig. 33A–D) Diagnosis. As for genus. Derivation of name. In honour of the Monmouth Amateur Paleontologist's Society (New Jersey, U.S.A.), of which Ralph Johnson is a member, for bringing this specimen to our attention. Material examined. One partly decorticated carapace from the top of the lower Danian Tinton Formation, Manasquan River Basin, Monmouth County, New Jersey, U.S.A. (AMNH 50421). Description. Carapace small, fusiform, about twice as long as broad, widest at mid-length, strongly convex in transverse cross section, slightly convex in longitudinal cross section, uniformly covered with granules, coarsest towards orbitofrontal, anterolateral regions. Orbitofrontal margin narrow. Broad base of rostrum with prominent ridge extending posteriorly about one-third of total carapace length, bounded by 2 deep grooves effacing posteriorly. Convex anterolateral margin with diminutive, anteriorly directed spine at approximately one-fifth of total carapace length from front. Subhepatic region with blunt crest. Posterior margin slightly wider than orbitofrontal margin. Branchiocardiac grooves curved, situated slightly posterior of mid-length. Pterygostome elongated, convex, covered by evenly spaced, fine granules; buccal margins concave, with smooth rim; posterior corner of pterygostome recessed for mxp3 coxa. Branchiostegite low posteriorly, with pronounced border. Remarks. Despite the superficial resemblance to Lyreidinae, Symethoides n. gen. differs markedly from Lyreidus, Lysirude and Macroacaena in having an anterior axial ridge bounded by two grooves and a completely granular carapace, and in lacking long anterolateral spines (few exceptions in Lyreidus). It differs also from Macroacaena in lacking a posterior ridge and a narrower orbitofrontal region. The latest Cretaceous Lyreidina Fraaye & Van Bakel, 1998, shows an anterior axial ridge, but differs by its pyriform shape, carapace areolation and ornament. From all currently known raninids, the new genus is distinguished by a unique combination of carapace outline, frontal region and ornamentation. The fusiform carapace, small, forwardly directed anterolateral spine and narrow orbitofrontal region support assignment to the subfamily Symethinae., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 106-107, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Landman, N. H., Johnson, R. O., Garb, M. P., Edwards, L. E. & Kyte, F. T. (2007) Cephalopods from the Cretaceous / Tertiary boundary interval on the Atlantic Coastal Plain, with a description of the highest ammonite zones in North America. Part III. Manasquan River Basin, Monmouth County, New Jersey. Bulletin of the American Museum of Natural History, 303, 1 - 122.","Fraaye [sic], R. H. B. & Van Bakel, B. W. M. (1998) New raninid crabs (Crustacea, Decapoda, Brachyura) from the late Maastrichtian of the Netherlands. Geologie en Mijnbouw, 76, 293 - 299."]}

Subfamily Lyreidinae Guinot, 1993 Lyreidinae Guinot, 1993b: 1326. Type genus. Lyreidus De Haan, 1841. Genera included. Bournelyreidus n. gen., Heus Bishop & Williams, 2000, Lyreidus De Haan, 1841, Lysirude Goeke, 1986, Macroacaena Tucker, 1998, and Rogueus Berglund & Feldmann, 1989. Diagnosis. Carapace longer than wide, elongated, generally narrow, pyriform or fusiform, with long, narrow anterior portion, or with wide, sinuous anterior margin. Anterolateral margin lacking tooth or with 1 or 2 teeth, with long spine (Lysirude, Macroacaena) or single, strong bifurcated spine (Rogueus). Posterolateral margin long, generally rimmed. Posterior margin short, concave. Cervical groove absent; branchiocardiac groove short. Carapace smooth, regions indistinct; cuticle microstructure with pits and upright nodes. Rostrum broadly triangular or subtrapezoidal. Orbitofrontal margin from narrow, less than one-third of maximum carapace width, to wide, threequarters of maximum carapace width. Supraorbital margin generally unarmed, with single fissure, or with few teeth (Macroacaena). Antennules, antennae about same size and general shape; long, slender, modified in connection with respiratory currents. Antennule not folded, basal article expanded, somewhat concave internally so that when apposed both antennules form conduit for the exhalant current. Proepistome narrow. Epistome anteriorly produced between basal articles of antennules. Subantennary lobe of pterygostome developed, produced far in front of mandibles, fused over a long stretch with epistome edges. Mxp3 long, narrow, slender, oxystomian condition; merus longer than ischium. Milne-Edwards openings absent. Pterygostome elongated, non-areolated, tumid. Pleurites 5‒7 partially exposed, not excavated, forming flat area, not overhung by edge of branchiostegite. Sternum/pterygostome junction moderately developed; junction sternum/pleurites well developed between P1, P2, less so between P2, P3. Thoracic sternum long, narrow, strongly deflected behind sternite 7; sternite 3 variously crown shaped; sternite 4 not much expanded, flat; sutures 4/5 reduced, crescent shaped; sternite 5 relatively wide; sternite 6 only slightly narrower; sternite 7 narrowing posteriorly; sternite 8 narrow, elongated, perpendicular to preceding ones. Medial line along sternites 7, 8. Pair of strong, elongated, hook-like projections arise from episternite 5, recurved at tip, distally with double peg that firmly fits into pair of deep sockets in latero-posterior extended corners of abdominal somite 6; locking may be effective in ovigerous females, even with large egg mass, but becoming obsolete in larger females. Socket on abdominal somite 6 long, limited by thickening. Two small spermathecal apertures face each other on opposite sides of depression (sunken pit) on sternite 7, separated by vertical medial wall, marked externally by medial line. Chelipeds homochelous, homodontous. Basis-ischium immoveably fused with long merus. Propodus of variable length, flattened; its upper margin unarmed or with single spine; lower margin armed with few sharp spines; dactylus long, smooth on dorsal border, bent against anterior border of palm; fixed finger much inflated; prehensile borders of both fingers with staggered, low teeth. P2‒P4 rather slender. Merus rather long, slender. Propodus, dactylus flattened, compressed. P2 propodus short, broad, dactylus slightly spatulate. P3 propodus longer, dactylus elongated, styliform, externally ridged. P4 carpus, propodus, dactylus variously lobate. P4 coxae subdorsally located. P5 more dorsal, much reduced, filiform, ending in small, flattened, elliptical dactylus. Sterno-abdominal depression present posteriorly, entirely covered by male abdomen. Abdomen freely articulated (6 articles plus small telson), narrow, fixed; somites 1‒3 dorsal, in straight line with carapace, rest completely folded; sharp flexure at level of somite 4 which bears a strong spine; somite 5 may also bear single spine; somite 6 longer, ventrally with developed sockets fitting double peg of the hook-like projections of thoracic sternite 5 for abdominal locking mechanism. Sexual dimorphism indistinct. Posterior branchial orifices or water conduits on the flanks of carapace absent; instead, with post-frontal modifications for respiratory current., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 77-78, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Guinot, D. (1993 b) Donnees nouvelles sur les Raninoidea de Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l'Academie des Sciences (Paris), Sciences de la vie, (3) 316, 1324 - 1331.","Bishop, G. A. & Williams, A. B. (2000) Fossil crabs from Tepee Buttes, submarine seeps of the Late Cretaceous Pierre Shale, South Dakota and Colorado, U. S. A. Journal of Crustacean Biology, 20 (Special Number, 2), 286 - 300.","Goeke, G. D. (1986) Decapod Crustacea: Raninidae. In: Forest, J. (Ed.), Resultats des Campagnes MUSORSTOM I et II. Philippines, Tome 2. Memoires du Museum d'Histoire naturelle (Paris), nouvelle serie, A. Zoologie, 133 (1985), 205 - 228.","Tucker, A. B. (1998) Systematics of the Raninidae (Crustacea: Decapoda: Brachyura), with accounts of three new genera and two new species. Proceedings of the Biological Society of Washington, 111, 320 - 371.","Berglund, R. E. & Feldmann, R. M. (1989) A new crab, Rogueus orri n. gen. and sp. (Decapoda: Brachyura), from the Lookingglass Formation (Ulatisian Stage: lower Middle Eocene) of southwestern Oregon. Journal of Paleontology, 63, 69 - 73."]}

Genus Macroacaena Tucker, 1998 Macroacaena Tucker, 1998: 325. Carinaranina Tucker, 1998: 334. Type species. Lyreidus succedanus Collins & Wienberg Rasmussen, 1992, by original designation. Type species, by original designation, of Carinaranina is Eumorphocorystes naselensis Rathbun, 1926a. Species included. Macroacaena alseana (Rathbun, 1932), M. bispinulata (Collins & Wienberg Rasmussen, 1992), M. chica Schweitzer, Feldmann, Fam, Hessin, Hetrick, Nyborg & Ross, 2003, M. franconica Schweigert, Feldmann & Wulf, 2004, M. fudoujii (Karasawa, 2000), M. johnsoni (Rathbun, 1935), M. leucosiae (Rathbun, 1932), M. marionae (Tucker, 1998), M. naselensis (Rathbun, 1926), M. rosenkrantzi (Collins & Wienberg Rasmussen, 1992), M. schencki (Rathbun, 1932), M. succedana (Collins & Wienberg Rasmussen, 1992), and M. venturai Vega, Nyborg, Fraaye & Espinosa, 2007. Remarks. In Tucker’s PhD thesis (1995), which was distributed in printed form, the new genus Macracaena (1995: 113) was attributed to the Lyreididae. This name, however, is preoccupied by Macracaena Common, 1958, a genus of moth in the family Gelechiidae Stainton, 1854. The generic name of the crab was formally introduced as Macroacaena by Tucker (1998). Schweitzer et al. (2003a: 29) synonymised Carinaranina and Macroacaena on the basis of similarities of the dorsal carapace (see also Schweitzer et al. 2010: 71). The type species of both genera retain ventral details and a re-examination is called for in order to verify the synonymy. Tucker (1998: 325) noted that Macroacaena ‘possibly’ had the ‘processes to lock the abdomen in the sternum’ and attributed the genus to Lyreidinae. Material of the various species of Macroacaena needs to be re-evaluated to document the presence of hook-like projections., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 84, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Tucker, A. B. (1998) Systematics of the Raninidae (Crustacea: Decapoda: Brachyura), with accounts of three new genera and two new species. Proceedings of the Biological Society of Washington, 111, 320 - 371.","Collins, J. S. H. & Wienberg Rasmussen, H. (1992) Upper Cretaceous-Lower Tertiary decapod crustaceans from West Greenland. Bulletin fra Gronlands geologiske Undersogelse, 162, 1 - 46.","Rathbun, M. J. (1926 a) The fossil stalk-eyed Crustacea of the Pacific slope of North America. Bulletin of the United States National Museum, 138, 1 - 155.","Rathbun, M. J. (1932) New species of fossil Raninidae from Oregon. Journal of the Washington Academy of Science, 22: 239 - 242.","Schweitzer, C. E., Feldmann, R. M., Fam, J., Hessin, W. A., Hetrick, S. W., Nyborg, T. G. & Ross, R. L. M. (2003 a) Cretaceous and Eocene Decapod Crustaceans from Southern Vancouver Island, British Columbia, Canada, 66 pp. NRC Research Press, Ottawa.","Schweigert, G., Feldmann, R. M. & Wulf, M. (2004) Macroacaena franconica n. sp. (Crustacea: Brachyura: Raninidae) from the Turonian of S Germany. Zitteliana, A 44, 61 - 65.","Common, I. F. B. (1958) A revision of the pink bollworms of cotton (Pectinophoras Busck, Lepidoptera: Gelechiidae) and related genera in Australia. Australian Journal of Zoology, 6, 268 - 306.","Stainton, H. T. (1854) Insecta Britannica. Lepidoptera: Tineina, viii + 313 pp., 10 pls. L. Reeve, London."]}

Orithopsis tricarinata (Bell, 1863) (Fig. 22A–F) Necrocarcinus tricarinatus Bell, 1863: 21. Orithopsis bonneyi Carter, 1872: 530. Material examined. Holotype of Orithopsis bonneyi Carter, 1872, upper Albian (upper Greensand), Lyme Regis, Dorset, UK (SM B58557); female, carapace and abdomen, Cambridge Greensand, Cambridge, UK (SM B58557); upper Albian, Cambridge Greensand, Cambridge, England, partial carapace with partial thoracic sternum (IRScNB unnumbered (Van Straelen Collection, see also Guinot et al. 2008: fig. 9f); carapace, upper Albian, ‘Gault’, Folkestone, England (NHM BM 59808); lectotype, designated by Wright & Collins (1972: 67), carapace, upper Greensand (probably uppermost Albian according to Wright & Collins 1972: 67), Wiltshire, England (NHM BM 59519); female, carapace with sternum, abdomen and bases of pereiopods, Albian, ‘Gault’, Folkestone, England (NHM In. 30297). Emended description. Subhepatic region narrow, with single, blunt, granular crest. Pleural line distinct, raised, granular. Pterygostome tumid, with blunt crest parallel to subhepatic crest, becoming more acute anteriorly; second blunt crest less well-defined, parallel to margin of P1 coxa. Buccal cavity wide, buccal margin weakly concave, with broad, smooth buccal collar; posterior corner of pterygostome simple. Branchiostegite developed, tumid. Sternites 1‒4 exposed: sternites 1, 2 on lower level; sternite 1 small, oval; sternite 2 trapezoidal; sternite 3 diamond shaped with apex pointing downwards; separated from sternite 4 by deep lateral grooves; sternite 4 large, trapezoidal, anterior margin slightly wider than sternite 3, lateral sides concave, surface with deep axial gutter; episternite 4 large, wide, gynglyme for P1 large, visible in dorsal view. P1–P4 coxae large, on same level, slightly decreasing in size posteriorly; P5 (sub)dorsal, reduced. Female abdomen with all somites free, covering thoracic sternum in width, thus in contact with coxae of pereiopods. Abdominal somite 1 not preserved; somites 2–5 progressively increasing in width, tricarinate, with strong, widened, thorn-like tubercles, strongest on somite 4; somite 6 long, widening towards telson, surface with central spine, anterior corners swollen. Telson incompletely preserved, granular. Remarks. Discovery of a specimen with well-preserved ventral characters (NHM In. 30297) necessitates an amendment of the description; the dorsal carapace was described by Wright & Collins (1972: 67). Orithopsis tricarinata (see Bell 1863: 21, pl. 4, figs. 9–11), from the upper Aptian-lower Cenomanian of southern England, northern Spain, plus the lower Cenomanian of Mangyshlak, Kazakhstan (Ilyin 2005, as Necrocarcinus tricarinatus) and?upper Albian of Angola, of which O. bonneyi Carter, 1872 (upper Aptian-lower Cenomanian, southern England) is a junior synonym, was previously known mainly from the dorsal carapace. Guinot et al. (2008: 32) stated that, ’ Orithopsis tricarinata has remained an insufficiently known species and, moreover, lacks preserved ventral structures, except for the trituberculate (?male) abdominal segments described by Wright & Collins (1972: 68) ’. Specimen NHM In. 30297 shows well-preserved features of the thoracic sternum and abdomen for the first time., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 69, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Bell, T. (1863) A monograph of the fossil malacostracous Crustacea of Great Britain. Part II. Crustacea of the Gault and Greensand. Monograph of the Palaeontographical Society (London), 14, vii + 40 pp., pls. 1 - 11.","Carter, J. (1872) On Orithopsis Bonneyi, a new fossil crustacean. Geological Magazine, 9, 529 - 532, pl. 13, fig. 1.","Guinot, D., Vega, F. J. & Van Bakel, B. W. M. (2008) Cenomanocarcinidae n. fam., a new Cretaceous podotreme family (Crustacea, Brachyura, Raninoidia), with comments on related families. Geodiversitas, 30, 681 - 719.","Wright, C. W. & Collins, J. S. H. (1972) British Cretaceous crabs. Monograph of the Palaeontographical Society (London), 126 (533), 1 - 114, 22 pls.","Ilyin, I. V. (2005) Melovye i paleogenovye desiatinogie rakoobraznye (Crustaceamorpha, Decapoda) zapadnoii chasti Severnoj Evrazii, 295 + i pp., pls. 1 - 16. Izdatel'stvo Moskovskogo Universiteta, Moskva."]}

Genus Symethoides n. gen. Type species. Symethoides monmouthorum n. sp. Diagnosis. Small fusiform carapace; narrow orbitofrontal region, pronounced axial ridge on anterior third of carapace; diminutive, forwardly directed anterolateral spine; cuticle granular. Derviation of name. As for Symethis., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 105, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640

Subfamily Cyrtorhininae Guinot, 1993 Cyrtorhininae Guinot, 1993b: 1330. Type genus. Cyrtorhina Monod, 1956. Genera included. Antonioranina n. gen.; Cyrtorhina Monod, 1956. Diagnosis. Carapace ovate, variously wide, convex in both directions. Cervical groove obsolete, branchiocardiac groove indistinct. Posterolateral margins straight, anterolateral margins rounded; margins rimmed or lined by granules. Anterolateral margin with 2, 3 teeth, subhepatic spine may be present. Cuticle microstructure diverse: upright nodes, fungiform nodes, inclined nodes, pits. Front short, widely triangular, weakly to strongly sulcate, with sharp, anteriorly directed tooth on each side. Supraorbital margin with 2 fissures, inner fissure proportionally short, outer fissure longer, separating extraorbital teeth; outer extraorbital tooth robust, falciform. Orbits small; eyestalk short, of 3 elements: proximal article reduced, only ventrally visible; second article triangular, elongated; third dorsally exposed, calcified, strongly arched, with fungiform granules; cornea small. Antennary fossae situated posterior to antennulary fossae. Antennule deeply inserted, completely hidden by antenna, basal article notched distally, following articles much smaller; flagellum short. Proepistome distinct. Antennae massive, meeting medially, with external surface granular; articles 1, 2 fused, urinal opening at inner angle; article 3 expanded, with spatulate dorsal lobe, pointed ventral lobe; article 4 with salient anterodorsal angle; article 5 short; flagellum developed. Epistome triangular, medially raised, keeled. Subantennary lobe of pterygostome not delimited. Mxp3 operculiform; endopod long, blunt anteriorly, merus shorter than ischium; palp short, concealed. Exopod short, wider than endopod, anteriorly pointed. Pterygostome granular, ridged, with rows of dense setae, completely setiferous posteriorly. Exposed pleurites 5–7 forming large, unexcavated plate; branchiostegite developed, high, overhanging exposed pleurites, limited by P5 apposed along branchiostegal margins. Sternum/pterygostome junction very narrow, recessed. Sternum/exposed peurite connections large between P1, P2, narrower between P2, P3. Thoracic sternum narrow all along, thus P1‒P5 coxae approximated. A strong deflection at level of sternite 8. Sternite 1, 2 in a lower level, concealed; sternite 3 exposed, as small, V-shaped, horizontally extended strip; sternite 4 narrow, not laterally expanded except the slender, recessed anterolateral extensions forming sternum/pterygostome junction; sternites 4, 5 expanded laterally between P1, P2; sternites 6, 7 conspicuously narrow, linear; sternite 8 perpendicular to sternite 7. Medial line along sternites 5–8. Spermathecal apertures recessed, small, contiguous. Chelipeds homochelous, homodontous. Basis-ischium short, immoveably fused with massive merus. Propodus longer than wide, narrow, upper, lower margins smooth, except for acute superodistal spine; fingers long, acicular, crossing at acute tips; dactylus deflected, proximally with long acute spine, prehensile border unarmed; fixed finger only slightly deflected, prehensile border with 3 acute spines proximally. P2–P4 moderately stout; carpus short, modified. P2–P4 dactylus spatulate on P2, sickle shaped on P3, P4. P5 dorsal, moderately reduced, granular, lying along posterolateral carapace margin, adapted to branchiostegal edge; dactyl narrow, elongated. Abdomen proportionally small but rather wide in male, freely articulated, 6 articles plus telson, incompletely folded, first 4 articles dorsal, first somite as wide as posterior carapace margin. Female abdomen only slightly enlarged, thus sexual dimorphism indistinct. Uropods or sockets absent. Large posterior branchial orifices present., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 107-108, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Guinot, D. (1993 b) Donnees nouvelles sur les Raninoidea de Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l'Academie des Sciences (Paris), Sciences de la vie, (3) 316, 1324 - 1331.","Monod, T. (1956) Hippidea et Brachyura ouest-africains. Memoires de l'Institut francaise d'Afrique Noire, 45, 1 - 674."]}

Bournelyreidus teodorii n. sp. (Fig. 24A) Diagnosis. Carapace large, oval, elongated, widest at one-third of total carapace length from front. Posterolateral margin with divergent posterior portion; anterolateral margin shorter, with 2 small teeth. Orbitofrontal margin tridentate, occupying about half of maximum carapace width. Outer orbital corners prominent, supraorbital borders with 2 orbital notches. Rostrum triangular, bifid, excavated. Carapace surface with minute pits and granules. Derivation of name. Named in honour of Mr Dominique Téodori (Pechbonnieu, France), who collected and donated the specimen. Material examined. Holotype (MNHN.F A38533) collected at working quarry southeast of the village of Saint-Loup-en-Comminges (Haute-Garonne, Midi-Pyrénées, southwest France; September 2009), from a bluegrey marly limestone bed at the base of a series assigned to the ‘Marnes bleues de Saint-Loup’. A second specimen in the private collection of D. Teodori has also been examined; this retains a better-preserved anterolateral and frontal margin, so that features of the holotype could be checked and confirmed. Co-occurring macrofaunal assemblages are rich in holasteroid echinoids, ostreid bivalves, fishes, cephalopods (e.g., the scaphitid ammonite Hoploscaphites constrictus (Sowerby, 1817) and other unidentified crustaceans. The thickness of such limestone levels varies between 0.1 and 0.3 m; they alternate with softer marls. As noted by Kennedy et al. (1986: 1004, 1005, fig. 2) and Peybernès et al. (1998), the ‘Marnes bleues de St. Loup’ in the Blajan-Bazordan anticline are of late Maastrichtian age. Description. Carapace large (maximum length 66 mm, width 40 mm), oval, elongated, widest behind the lateral spines at about at one-third of total carapace length from front. Orbitofrontal margin tridentate, occupying about half of maximum carapace width. Base of rostrum about one-third of orbitofrontal region. Rostrum triangular, bifid, excavated. Large subcircular orbits with prominent outer orbital spines. Supraorbital borders with 2 relatively deep orbital fissures. Posterolateral margin with diverging posterior portion; anterolateral margin shorter, with 2 rather small, forwardly directed spines. Branchiocardiac grooves, impressions of epimeral adductor muscle scars faint. Carapace surface with minute pits, granules. Granules significantly coarser along lateral margins. Both chelipeds similar in size, form; merus, carpus, propodus of comparable length, width, covered with granules; propodus distinctly more flattened. Remarks. Despite the fact that the carapace has suffered from sediment compaction, the morphology of the orbitofrontal and anterolateral margins allows assignment of this form to Bournelyreidus n. gen. It is distinguished from B. carlilensis n. comb. and B. ? oaheensis n. comb. in attaining a markedly larger size and in having a less broad orbitofrontal region and a much larger carpus. Bournelyreidus tridens n. comb. has a much shorter anterolateral margin, and a more projected orbitofrontal region; its posterolateral margins are more concave and less divergent than the new species. Bourneylyreidus manningi n. comb. has much coarser ornamentation and a more clearly areolated carapace. The new species differs from B. laevis n. comb. in having a broader base to the rostrum and subcircular orbits rather than near-rectangular ones; it can also be distinguished from B. eysunesensis n. comb. by its less concave anterolateral margin between the outer orbital and first anterolateral spine.

Genus Heus Bishop & Williams, 2000 Heus Bishop & Williams, 2000: 290. Type species. Heus foersteri Bishop & Williams, 2000. Remarks. Only the carapace of Heus foersteri (see Bishop & Williams 2000: 290, fig. 4), from the upper Campanian of South Dakota and Colorado, U.S.A., is preserved. This reveals a unique set of characters: rostrum broad and narrow, strongly sulcate, with a keel that extending onto the carapace; oval swellings on epigastric and protogastric regions; cervical groove discernible; lateral margins with but a single, diminutive spine. These characters distinguish it easily from Bournelyreidus n. gen. Re-examination of the type series of B. ? oaheensis (see above) is needed to decide if it might also be better accommodated in Heus. The wide carapace, with clearly concave posterior (divergent) portions of the posterolateral margins, weakly armoured anterolateral margin, and swellings on the anterior carapace surface are somewhat similar to the configuration in Symethis Weber, 1795 (see below). The orbitofrontal construction, however, makes its placement in the Lyreidinae more likely., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 83, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Bishop, G. A. & Williams, A. B. (2000) Fossil crabs from Tepee Buttes, submarine seeps of the Late Cretaceous Pierre Shale, South Dakota and Colorado, U. S. A. Journal of Crustacean Biology, 20 (Special Number, 2), 286 - 300.","Weber, F. (1795) Nomenclator entomologicus secundum entomologiam systematicum ill. Fabricii adjectis speciebus recens detectis et varietatibus, viii + 171 pp., C. E. Bohn, Chilonii / Hamburgi."]}

Genus Marylyreidus n. gen. Type species. Notopocorystes punctatus Rathbun, 1935b. Diagnosis. As for subfamily. Derivation of name. In honour of the late Mary J. Rathbun, in recognition of her highly valuable work on fossil and extant crabs. Gender masculine. Remarks. Notopocorystes punctatus (see Rathbun 1935b: 8, pl. 12, figs. 14–16; Haj & Feldmann 2002: 473, figs. 3, 4.1–4.6), which is a senior synonym of both Notopocorystes parvus (see Rathbun 1935b: 48, pl. 12, figs. 11–13) and Raninella mucronata (see Rathbun 1935b: 50, pl. 14, figs. 32, 33), from the upper Albian–lower Cenomanian of Texas, was originally referred to Raninidae. It has typically been attributed to Cretacoranina (e.g., Haj & Feldmann 2002; Waugh et al. 2009) or Raninella (e.g., Rathbun 1935b; Feldmann & Maxey 1980; Tucker 1995; Bishop & Williams 2000). Haj & Feldmann (2002: 478) mentioned that the species was distinguished from other members of Cretacoranina by the fainter carapace areolation and weaker lateral and frontal spines. Waugh et al. (2009: 33) noted that the thoracic sternum and frontal arrangement of the present species did not conform to that of Cretacoranina, but that the cuticle microstructure (fungiform nodes) was remarkably similar. Marylyreidus n. gen. is erected herein to accommodate Notopocorystes punctatus, and placed in Lyreididae, as the only member of Marylyreidinae n. subfam. The new genus can easily be distinguished from Cretacoranina by those characters that exclude it from Palaeocorystoidea (see above). Characters distinguishing Marylyreidus punctatus n. comb. from members of Raninella are a carapace that is much less wide, anterolateral margin with one small spine (two strong, flattened spines in Raninella), posterolateral margin rounded (sharp, with distinct rim in Raninella), cuticle with fungiform nodes (inclined nodes in Raninella); pterygostome areolated (evenly swollen in Raninella), sternite 4 narrow, not connected to pterygostome (clearly widened anteriorly, junction sternum-pterygostome in Raninella), and sternite 5 devoid of medial line (medial line present in Raninella). The presence of a single anterolateral spine, the rounded nature of the posterolateral margin and cuticle microstructure serve to distinguish the carapace of Marylyreidus n. gen. from Bournelyreidus n. gen. The condition of the pterygostome is remarkably similar in these genera. The unique condition in Marylyreidus n. gen. of sternite 4 (i.e., not widened anteriorly, no junction sternum-pterygostome) and sternite 5 (extremely wide between P1 and P2), however, is absent in Bournelyreidus n. gen.

Cenomanocarcinus beardi Schweitzer, Feldmann, Fam, Hessin, Hetrick, Nyborg & Ross, 2003 (Figs. 17A–D; 18F) Material examined. MAB k. 2969: cast of paratype (GSC 124821), provided by C.E. Schweitzer, March 2007 (see Schweitzer et al. 2003a: 35, fig. 12.1); MAB k. 2544, k. 2880, k. 2916, k. 2917, Woodbine Formation, Cenomanian, Creek in Wagner Park, southwest Freud Prairie, Texas (U.S.A.). Emended description. Pterygostome large, with blunt, granular crest, buccal margin concave, rimmed. Mxp3: exopod long, slender, lateral margin gently arched; endopod ischium slightly longer than exopod, medially weakly excavated. Thoracic sternum narrow, elongated, with medial sterno-abdominal depression; sternite 3 small, crown shaped, sternite 4 trapezoidal, rather short; episternites 4 wide, large, subtriangular, anteriorly with distinct gynglyme for P1; sternites 5–8 equal in width, with flat, undivided medial surface, raised, posterolaterally directed episternites; sutures 4/5–6/7 sinuous, directed forwards; suture 7/8 long, straight, terminating in oval spermathecal aperture. Remarks. Discovery of a specimen with preserved ventral characters necessitates an amendment of the description; dorsal carapace was described by Schweitzer et al. (2003a: 38). Guinot et al. (2008: 38) described the morphology of a specimen (MAB k. 2544) referred to as Cenomanocarcinus aff. beardi. It has spermathecae (aperture preserved at one side), hence confirming the podotreme status of Cenomanocarcinidae. This particular specimen is here assigned beyond doubt to C. beardi. All features of the dorsal carapace of the examined Texas material compared well with the type series. The lack of a lateral tooth between the fourth anterolateral tooth and the epibranchial spine, the strongly produced epibranchial spine, the rounded cross section of the posterolateral margin, which is devoid of spinules or teeth (except near the posterior corner), and the tuberculation of the branchial crests distinguish specimens of the new species from both C. inflatus and C. vanstraeleni. The record of C. beardi from the Cenomanian of Texas marks the oldest occurrence of the species and a southerly range extension. Ventral features are described for the first time for this species, which thus far represents the only example of a spermatheca in Cenomanocarcinidae., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 54, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Schweitzer, C. E., Feldmann, R. M., Fam, J., Hessin, W. A., Hetrick, S. W., Nyborg, T. G. & Ross, R. L. M. (2003 a) Cretaceous and Eocene Decapod Crustaceans from Southern Vancouver Island, British Columbia, Canada, 66 pp. NRC Research Press, Ottawa.","Guinot, D., Vega, F. J. & Van Bakel, B. W. M. (2008) Cenomanocarcinidae n. fam., a new Cretaceous podotreme family (Crustacea, Brachyura, Raninoidia), with comments on related families. Geodiversitas, 30, 681 - 719."]}

Campylostoma matutiforme Bell, 1858 (Fig. 16A–F) Material examined. Partial carapace with mxp3, London Clay, Ypresian, lower Eocene, Herne Bay, Kent, England (NHM In. 32654/32655); syntype, carapace with sternite 4, P1 coxae and mxp3, London Clay, Ypresian, lower Eocene, Isle of Sheppey, England (NHM In. 7314); partial carapace with sternite 4, abdomen and bases of pereiopods, same provenance (NHM In. 35231); 2 carapaces, same provenance (NHM In. 39424, In. 59083). Emended description. Pleural suture distinct, running just below anterolateral, posterolateral margins. Pterygostome large, anteriorly tumid, with blunt granular crest, extended anteriorly into prominent subantennary lobe; buccal cavity narrow, margins weakly concave, rimmed. Mxp3: coxae large, flabelliform, inserted in front of P1; exopod relatively short, wedge shaped with narrow distal end; endopod elongated, slender, basis-ischium, merus equally long, merus strongly grooved. Sternites 3‒6 preserved; sternite 3 small, incompletely preserved, separated from sternite 4 by oblique lateral grooves; sternite 4 trapezoidal, lateral sides weakly raised, lateral margins slightly concave, nearly vertical groove at level of P1 gynglyme separating large raised episternite 4; episternite 5 raised, oblique, at least one prominence present for abdominal holding; sutures 4/5, 5/6 similarly shaped, only lateral, crescent shaped, medial part vertical, demarcating flat, undivided medial floor of sternum. Abdomen curvature in prolongation with carapace, all somites free, somite 1 short, wide, somite 3 narrowest, somites 3‒5 with raised medial ridge, which is strong, acute in somite 4, somite 6 longer, telson unknown. P1‒P 3 in the same level, P4 weakly tilted, slightly reduced, P5 subdorsal, strongly reduced. Remarks. Study of the material examined necessitates an amendment of the description; the carapace was described by Bell (1858: 23) and Carter (1898: 30). Campylostoma matutiforme (see Bell 1858: 23, pl. 3, figs. 8– 10) was deemed a podotreme crab, possibly a cenomanocarcinid, by Guinot et al. (2008: 21). The re-examination of several specimens in the NHM collections (Fig. 16A–F) confirm this placement. The much elongated mxp3 with large, flabelliform coxae, sculpture of the pterygostome with pronounced, pointed subantennary lobes, crownshaped sternite 3, subtrapezoidal sternite 4, match the cenomanocarcinid configuration. The carapace does not show the strong branchial crests that are typical of Cenomanocarcinus spp., and the orbits are much wider. The presence of an abdominal holding system on episternite 5 is confirmed in one specimen, where a single, sharp tubercle is visible; because of poor preservation, it is unclear if a more posteriorly situated peg was present., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 57-58, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Bell, T. (1858) A monograph of the fossil malacostracous Crustacea of Great Britain. Part I. Crustacea of the London Clay. Monographs of the Palaeontographical Society (London), 10 (40) (1856), viii + 44 pp., pls. 1 - 11.","Carter, J. (1898) A contribution to the palaeontology of the decapod Crustacea of England. Quarterly Review of the Geological Society of London, 54, 15 - 44, pls. 1 - 2.","Guinot, D., Vega, F. J. & Van Bakel, B. W. M. (2008) Cenomanocarcinidae n. fam., a new Cretaceous podotreme family (Crustacea, Brachyura, Raninoidia), with comments on related families. Geodiversitas, 30, 681 - 719."]}

Eosymethis aragonensis n. sp. (Fig. 32A–C) Diagnosis. As for genus. Derivation of name. From the region of Aragón, northern Spain. Material examined. Holotype, and only specimen so far known, a near-complete carapace with remains of the pereiopods, preserved as a natural mould, from the marls of the ‘ upper Ilerdian’ (lowermost Ypresian, Eocene) Roda Formation (Cuevas-Gozalo et al. 1985), approximately 7 km northwest of the village of Isabena (Huesca, Aragón) (MGSB75293; ex José Enrique Ortega Collection). Description. Carapace small, subovally elongated in outline, longer than wide; maximum width at confluence of lateral margins, posterior of mid-length. Dorsal surface slightly concave in both directions. Orbitofrontal margin narrow, markedly advanced; orbits small, directed anteriorly; front strongly projected with triangular tip. Anterolateral margins long, with notable angular hepatic node after concave first portion; posterior portion nearly straight, diverging posteriorly. Posterolateral margins shorter, from straight to slightly concave, rimmed, convergent to the rear. Posterior margin fairly concave, much broader than orbitofrontal margin. Dorsal regions poorly defined; postfrontal depression present; gastric regions weakly defined; branchial regions with notable oblique branchial scars, lateratelly disposed; cardiac region large, elongated, bounded by marked branchiocardiac grooves, with angular node at lateral upper portion; anterior portion separated from gastric regions by subtle groove. Dorsal surface covered by large, dense pits, uniformly distributed over entire carapace; some granules in posterior portion of posterolateral margins. Pereiopods partially preserved, showing short, flattened articles with falciform dactylus. Remarks. The species exhibits features that permit placement in Symethinae: the carapace has a narrow, projected anterior portion of the carapace and a post-frontal depression; the dorsal surface shows an isolated, lateral, acute groove; the anterolateral margin has only a diminutive node; and falciform dactyli of the pereiopods. Symethinae so far consists of a single genus, Symethis, with only three extant species (Ng et al. 2008: 43). Eosymethis n. gen. can be distinguished from Symethis on the basis of a smoother, less ‘eroded’ anterior carapace and in having the cuticle microstructure consisting of large perforations or setal pits rather than fungiform nodes (Waugh et al. 2009). Eosymethis aragonensis n. sp. originates from what appears to have been a fairly muddy-bottom habitat., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 104, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Cuevas-Gozalo, M., Donselaar, M. E. & Nio, S. D. (1985) Eocene clastic tidal deposits in the Tremp-Graus Basin (Prov. of Lerida and Huesca). 6 th European Regional Meeting IAS, Lerida, Guidebook Excursion no. 6, 215 - 266.","Ng, P. K. L., Guinot, D. & Davie, P. J. F. (2008) Systema Brachyurorum: Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology, Supplement no. 17, 1 - 286.","Waugh, D. A., Feldmann, R. M. & Schweitzer, C. E. (2009) Systematic evaluation of raninid cuticle microstructure. Bulletin of the Mizunami Fossil Museum, 35, 15 - 41."]}

Genus Eosymethis n. gen. Type species. Eosymethis aragonensis n. sp. Diagnosis. Small-sized carapace, elongatedly suboval, weakly convex in both directions. Orbitofrontal margin extremely narrow, well projected, orbits small, front strongly advanced. Anterolateral margins with a notable angular node at level of hepatic region. Dorsal regions weakly defined; post-frontal depression present; arched branchiocardiac grooves, oblique branchial scars. Dorsal surface densely covered by large pits. Pereiopods with short, flattened articles, dactyli falciform. Derivation of name. Indicating an ancestral form of Symethis.

Bournelyreidus tridens (Roberts, 1962) n. comb. Raninella tridens Roberts, 1962: 187. (Fig. 23B) Material examined. Casts of GAB 37-832 (ventral) and GAB 37-833 (dorsal) (Bishop 1983b: fig. 3l; table 1) (MAB k.0214). Remarks. Raninella tridens (see Roberts 1962: 187, pl. 88, figs. 5, 6), from the Campanian of New Jersey and Delaware, U.S.A. and the lower Maastrichtian of Mississippi (Bishop 1983b), is here transferred to Bournelyreidus n. gen. The carapace is small, elongated, and the cuticle shows fine granules (upright nodes). The orbital margin is horizontal, branchiocardiac grooves are weak, the median third of the carapace has vertical margins, the posterior portions of the posterolateral margin being divergent and slightly concave. Sternite 3 is crown shaped, sternite 4 is long, with strongly concave lateral margins and it joins the pterygostome anteriorly. The posterior corner of the pterygostome is recessed for the rather small mxp3 coxa. Episternites 4 small, separated by distinct, crescentshaped sutures. Sternite 5 is comparatively large, widened anteriorly and with two small depressions posteriorly; the episternites show rather short, curved projections. No pegs are discernible, which may be the result of incomplete preservation, or the specimen represents a mature female in which the holding was lost. The projections are curved outwards, and seem to ‘rest’ on the surface of sternite 6; such configuration suggests the specimen to be a mature female. Only the anterior portion of sternite 6 is preserved; it is strongly excavated. In Lyreidinae the surface in between these projections and the anterior portion of sternite 6 is typically excavated as a kind of sternoabdominal depression. No medial line is present on sternites 1‒6, a characteristic of the genus. Exposed pleurites 5 and 6 are preserved; they are rather low, and notably granular., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 83, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Roberts, H. B. (1962) The Upper Cretaceous decapod crustaceans of New Jersey and Delaware. In: Richards, H. G., Ramsdell R. C., Miller, A. K., Garner, H. F. & Reeside, J. B. (Eds.), The Cretaceous fossils of New Jersey. Paleontology Series, New Jersey Geology Survey, 61,163 - 191, pls. 80 - 89.","Bishop, G. A. (1983 b) Fossil decapod Crustacea from the Late Cretaceous Coon Creek Formation, Union County, Mississippi. Journal of Crustacean Biology, 3, 417 - 430."]}

Genus Campylostoma Bell, 1858 Campylostoma Bell, 1858: 23. Type species. Campylostoma matutiforme Bell, 1858, by monotypy. Remarks. The genus was tentatively included in Cenomanocarcinidae by Guinot et al. (2008: 20). Material examined here corroborates this assignment; Campylostoma represents the youngest (Paleogene, Cenozoic) member of the family., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 57, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Bell, T. (1858) A monograph of the fossil malacostracous Crustacea of Great Britain. Part I. Crustacea of the London Clay. Monographs of the Palaeontographical Society (London), 10 (40) (1856), viii + 44 pp., pls. 1 - 11.","Guinot, D., Vega, F. J. & Van Bakel, B. W. M. (2008) Cenomanocarcinidae n. fam., a new Cretaceous podotreme family (Crustacea, Brachyura, Raninoidia), with comments on related families. Geodiversitas, 30, 681 - 719."]}

Eosymethis aragonensis n. sp. (Fig. 32A–C) Diagnosis. As for genus. Derivation of name. From the region of Aragón, northern Spain. Material examined. Holotype, and only specimen so far known, a near-complete carapace with remains of the pereiopods, preserved as a natural mould, from the marls of the ‘ upper Ilerdian’ (lowermost Ypresian, Eocene) Roda Formation (Cuevas-Gozalo et al. 1985), approximately 7 km northwest of the village of Isabena (Huesca, Aragón) (MGSB75293; ex José Enrique Ortega Collection). Description. Carapace small, subovally elongated in outline, longer than wide; maximum width at confluence of lateral margins, posterior of mid-length. Dorsal surface slightly concave in both directions. Orbitofrontal margin narrow, markedly advanced; orbits small, directed anteriorly; front strongly projected with triangular tip. Anterolateral margins long, with notable angular hepatic node after concave first portion; posterior portion nearly straight, diverging posteriorly. Posterolateral margins shorter, from straight to slightly concave, rimmed, convergent to the rear. Posterior margin fairly concave, much broader than orbitofrontal margin. Dorsal regions poorly defined; postfrontal depression present; gastric regions weakly defined; branchial regions with notable oblique branchial scars, lateratelly disposed; cardiac region large, elongated, bounded by marked branchiocardiac grooves, with angular node at lateral upper portion; anterior portion separated from gastric regions by subtle groove. Dorsal surface covered by large, dense pits, uniformly distributed over entire carapace; some granules in posterior portion of posterolateral margins. Pereiopods partially preserved, showing short, flattened articles with falciform dactylus. Remarks. The species exhibits features that permit placement in Symethinae: the carapace has a narrow, projected anterior portion of the carapace and a post-frontal depression; the dorsal surface shows an isolated, lateral, acute groove; the anterolateral margin has only a diminutive node; and falciform dactyli of the pereiopods. Symethinae so far consists of a single genus, Symethis, with only three extant species (Ng et al. 2008: 43). Eosymethis n. gen. can be distinguished from Symethis on the basis of a smoother, less ‘eroded’ anterior carapace and in having the cuticle microstructure consisting of large perforations or setal pits rather than fungiform nodes (Waugh et al. 2009). Eosymethis aragonensis n. sp. originates from what appears to have been a fairly muddy-bottom habitat.

Family Raninidae De Haan, 1839 emend. Raninidae De Haan, 1839: 102. Included subfamilies. Cyrtorhininae Guinot, 1993b (see below), Notopodinae Serène & Umali, 1972, Raninoidinae Lőrenthey in Lőrenthey & Beurlen, 1929, Ranininae De Haan, 1839, and Symethinae Goeke, 1981 (see below). Type genus. Ranina Linnaeus, 1758. Diagnosis. Carapace elongated to oval, rounded, widened anteriorly; convex in transverse direction; may be roof shaped, with medial carina. Dorsal surface smooth, pitted, granular, inclined or fungiform nodes, eroded, scabrous or terraced; cervical groove generally indistinct, branchiocardiac grooves present. Anterolateral margin armed with 1‒3 teeth, which may be broad, or long, bifurcate, spinose, or small, blunt. Posterolateral margin long, often rimmed. Front generally triangular or subtrapezoidal, pointed, may be trifid, or replaced by V-shaped emargination. Orbitofrontal margin narrow to conspicuously wide. Supraorbital margin with 1 or 2 closed or open fissures, lined with granules, spinules or armed with teeth; extraorbital tooth generally produced, may be broad, bilobate. Orbits directed anteriorly or anterolaterally, narrow to wide. Eyestalks short to long, composed of 1‒3 calcified articles, bent at angle to one another when retracted into orbit. Antennules, antennae rather stout; antenna not folded, inserted somewhat below antennule; antennules, antennae modified in connection with respiratory currents. Proepistome concealed; epistome short; endostome large, produced; strongly excavated. Pterygostome large, tumid, subantennary lobe well defined. Buccal frame large, with broad collar. Mxp3 strongly elongated, operculiform, endopod often grooved. Thoracic sternum wider anteriorly, conspicuously narrow posteriorly, strongly deflected at different levels. Sternite 3 crown shaped (compressed in Cyrtorhininae); sternite 4 usually wide, flat (narrow in Cyrtorhininae); suture 4/5 crescent shaped; sternite 5 expanded laterally; sternite 6 smaller, wide anteriorly, restricted between P3 coxae; sternites 6‒8 conspicuously narrow. Medial line present along posterior sternites, running up to sternite 5 or 6. Sternum/pterygostome junction present, narrow to large. Junction sternum/exposed pleurites present between P1, P2 and between P2, P3, or only between P1, P2 (Notopodinae). Pleurites 5‒7 partially exposed (gymnopleurity), calcified, forming polished surface, generally excavated, concave. Spermathecal apertures various: opening anteriorly, within sternite 7; generally close to each other, recessed in medial depression (more superficial, wider, hooded in Symethinae). Absence of sterno-abdominal cavity. Abdomen short, incompletely folded, 6 freely articulated somites plus small telson; somites smooth; first somites dorsal, in prolongation with carapace, relatively wide in male, not much enlarged in female, thus sexual dimorphism relatively indistinct. Uropods or sockets absent; absence of abdominalholding structures. Chelipeds homochelous, homodontous; arthrodial cavities large, placed laterally. Basis-ischium short, immoveably fused with long merus; propodus short, inflated, flattened, armed with long, sharp spines on upper and lower margins, or elongated, narrow, unarmed; fingers usually with teeth of prehensile borders joining alternatively, unarmed (several long, sharp spines in Cyrtorhininae); fixed finger short to long, may be strongly deflected. Arthrodial cavities of P2‒P4 almost ventral; those of P5 only, or both P4, P5 subdorsal. Carpus, propodus, dactylus flattened; dactylus variously modified: flattened, paddle shaped, lanceolated, or falciform. P5 weakly to strongly reduced, dorsal or subdorsal in position, usually fitting posterolateral border of carapace. Respiratory mechanism highly specialised. Absence of Milne-Edwards openings; strongly modified frontal region, antennules, antennae modified for inhalant current; posterior branchial orifices generally present, or absent (Notopodinae). Remarks. Included in Raninidae are those raninoidean crabs that are markedly modified by their burying behaviour, thus with peculiar respiratory properties and having lost abdominal holding, in contrast to Lyreididae. The Cyrtorhininae and Symethinae are here tentatively placed in Raninidae awaiting the revision of the Raninoidea by M. Tavares., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on pages 87-89, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Guinot, D. (1993 b) Donnees nouvelles sur les Raninoidea de Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l'Academie des Sciences (Paris), Sciences de la vie, (3) 316, 1324 - 1331.","Serene, R. & Umali, A. F. (1972) The family Raninidae and other new and rare species of brachyuran decapods from the Philippines and adjacent regions. The Philippine Journal of Science, 99 (1970), 21 - 105, pls. 1 - 9.","Lorenthey, E. & Beurlen, K. (1929) Die fossilen Dekapoden der Lander der Ungarischen Krone. Geologica Hungarica, Series Palaeontologica, 3, 1 - 420, 16 pls.","Goeke, G. D. (1981) Symethinae, new subfamily, and Symethis garthi, new species, and the transfer of Raninoides ecuadorensis to Notosceles (Raninidae: Brachyura: Gymnopleura). Proceedings of the Biological Society of Washington, 93, 971 - 981.","Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (Editio decima), 1, 824 pp. Laurentii Salvii, Holmiae.","Rathbun, M. J. (1935 b) Fossil Crustacea of the Atlantic and Gulf Coastal Plain. Geological Society of America, Special Papers, 2, viii + 1 - 160, pls. 1 - 26."]}

Genus Camarocarcinus Holland & Cvancara, 1958 Camarocarcinus Holland & Cvancara, 1958: 499. Type species. Camarocarcinus arnesoni Holland & Cvancara, 1958, by original designation. Species included. Camarocarcinus arnesoni Holland & Cvancara, 1958, C. obtusus Jakobsen & Collins, 1979, and C. quinquetuberculatus Collins & Wienberg Rasmussen, 1992. Material examined. Camarocarcinus arnesoni: well-preserved carapace, Paleocene, Cannonball Formation, Burleigh County, North Dakota (U.S.A.) (MGSB unregistered; Fig. 15E–G); C. quinquetuberculatus: holotype, carapace with thoracic sternum and mxp3, Paleocene, Turritellakløft Member, Agatdal Formation, central Nûgssuaq, West Greenland (MGUH 21.609; Fig. 15A–D)., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 49, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Holland Jr, F. D. & Cvancara, A. M. (1958) Crabs from the Cannonball Formation (Paleocene) of North Dakota. Journal of Paleontology, 32, 495 - 505, pl. 74.","Jakobsen, S. L. & Collins, J. S. H. (1979) Decapod Crustacea from the Palaeocene of Zealand, Denmark. Proceedings of the Geologists' Association, 90, 61 - 64.","Collins, J. S. H. & Wienberg Rasmussen, H. (1992) Upper Cretaceous-Lower Tertiary decapod crustaceans from West Greenland. Bulletin fra Gronlands geologiske Undersogelse, 162, 1 - 46."]}

Genus Cristafrons Feldmann, Tshudy & Thomson, 1993 Cristafrons Feldmann, Tshudy & Thomson, 1993: 31. Type species. Cristafrons praescientis Feldmann, Tshudy & Thomson, 1993, by monotypy. Remarks. Cristafrons shows an unusual set of characters. Feldmann et al. (1993: 31) remarked that the general form was similar to species of Notopocorystes. Cristafrons can be readily distinguished from Palaeocorystidae by the prescence of only a single lateral spine and a narrow, non-bifurcate rostrum. The well-defined cervical groove and the hepatic and postfrontal areolations, however, are unusual for Raninoidea, whereas those features are commonly found in Palaeocorystoidea. The thoracic sternum of Cristafrons is ‘not cleft longitudinally’, i.e., the medial line is absent (sternites 1‒5 are described), members of Raninoidinae typically having a medial line up to sternite 5 (Fig. 44C, D). It is unlikely that Cristafrons belongs to Ranininae, which has wider carapaces and two wide lateral teeth. Exposed pleurites are not described in the type series: they may be concealed by matrix or were overlooked in the original description, which states that the type specimens ‘are deposited in the Natural History Museum, London’ (Feldmann et al. 1993: 33), where only two specimens could be traced by us. The majority of the type series appears to have been forwarded to the British Antarctic Survey (BAS), under ‘ Cretiscalpellum praescientis ’ (see also http://www.antarctica.ac.uk/bas_research/data/access/fossildatabase/filter.php?taxonKeyword= CRUSTACEA). A re-examination of the type series is called for in order to resolve the precise taxonomic placement of Cristafrons., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 100, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Feldmann, R. M., Tshudy, D. M. & Thomson, M. R. A. (1993) Late Cretaceous and Paleocene decapod crustaceans from James Ross Basin, Antarctic Peninsula. The Paleontological Society Memoir, 28, 1 - 41."]}

Genus Silvacarcinus Collins & Smith, 1993 Silvacarcinus Collins & Smith, 1993: 263. Type species. Silvacarcinus laurae Collins & Smith, 1993, by original designation.

Genus Silvacarcinus Collins & Smith, 1993 Silvacarcinus Collins & Smith, 1993: 263. Type species. Silvacarcinus laurae Collins & Smith, 1993, by original designation., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 69, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Collins, J. [S. H.] & Smith, R. (1993) Ypresian (Lower Eocene) crabs (Decapoda, Crustacea) from Belgium. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Sciences de la Terre, 63, 261 - 270, pl. 1."]}

Cenocorystes bretoni n. sp. (Fig. 2A–D) Diagnosis. Carapace small, subhexagonal in outline, slightly longer than wide, maximum width at epibranchial spine one-third distant from front; posterior margin rimmed, strongly concave, shorter than orbitofrontal margin; dorsal regions weakly differentiated; cervical groove weakly defined, V-shaped; branchiocardiac grooves welldefined; epibranchial region with median oblique scar; cuticle granular. Pterygostome large, inflated, buccal margins concave, rimmed. Thoracic sternum narrow, elongated; sternites 1, 2 narrow, triangular; sternites 3, 4 separated by oblique grooves. Mxp3 elongated, coxae large, basis-ischium fused, grooved. Derivation of name. In honour of Gérard Breton (Université de Rennes I, Rennes, France), who has substantially contributed to our knowledge of fossil decapods from France. Material examined. Holotype: MAB k. 2899 (indeterminate sex); upper Cenomanian, precise level unknown; Vibraye, near Le Mans (Sarthe, northern France). Description. Carapace small, subhexagonal in outline, slightly longer than wide (in absence of orbitofrontal margin, length measured from basis of extraorbital spine), widest approximately one-third of total carapace length from front; dorsal surface fairly convex in both directions. Orbitofrontal margin broad, front not preserved. Anterolateral margins arched, with 3 broadly based spines; 1 hepatic plus 2 epibranchial, decreasing in size posteriorly. Posterolateral margins converging backwards, sinuous, slightly concave anteriorly, slightly convex posteriorly, margin rounded; with small spine anteriorly. Posterior margin fairly concave, rimmed, narrower than orbitofrontal margin. Dorsal regions of carapace poorly differentiated. Cervical groove weakly defined, discernible at lateral margins, axially; medially interrupted by small gastric pits. Epibranchial region exhibits oblique scar in medial portion, arched muscle scars posteriorly running subparallel to cervical groove. Cardiac region large, bounded by rather deep branchiocardiac grooves. Dorsal surface of carapace densely, uniformly granular. Pterygostome large, inflated; buccal margin concave, rimmed. Pleural suture well-defined as thin line. Thoracic sternum narrow, not connected to pterygostome, narrowing backwards, sternites 1, 2 narrowly triangular, situated at lower level; sternite 3 crown shaped, well separated from sternite 4 by oblique grooves; sternite 4 trapezoidal, episternite 4 suboval, robust, slightly extending laterally; suture 4/5 crescent-shape; sternite 5 without lateral depression; suture 5/6 crescent-shape. P4 arthrodial cavity tilted; P5 reduced, (sub)dorsal. Abdomen narrow, first somite broader. Mxp3 elongated, with large, flabelliform coxa; endopod basis-ischium elongated, merus shorter, strongly depressed axially. Remarks. The new species presents the following features that allow placement in Cenocorystes: reduced length of the carapace, with length roughly equalling width; large orbits; weakly defined cervical groove; arched anterolateral margin with four spines directed forwardly and outwardly, gently rounded posterolateral margins. In absence of a completely preserved front in both species of the genus, the total length was taken from the base of the extraorbital spine, which is preserved in the holotypes of both. Cenocorystes bretoni n. sp. differs from the type species, C. fournieri, in having a more tumid dorsal carapace surface and in lacking clearly defined grooves around the hepatic anterolateral spine (Collins & Breton 2009: fig. 6) as well as four protogastric tubercles, and in having a more concave posterior margin. In addition, C. bretoni n. sp. has more distinct branchiocardiac furrows, whereas the intestinal region is better defined in C. fournieri. The new species, with partial ventral side preserved, reveals the thoracic sternum of Cenocorystes for the first time. It matches the general sternal configuration of Palaeocorystidae; differences with other genera are the rather strong grooves separating sternites 3 and 4, sternite 5 lacking a lateral depression, and sutures are more closed.

Genus Eosymethis n. gen. Type species. Eosymethis aragonensis n. sp. Diagnosis. Small-sized carapace, elongatedly suboval, weakly convex in both directions. Orbitofrontal margin extremely narrow, well projected, orbits small, front strongly advanced. Anterolateral margins with a notable angular node at level of hepatic region. Dorsal regions weakly defined; post-frontal depression present; arched branchiocardiac grooves, oblique branchial scars. Dorsal surface densely covered by large pits. Pereiopods with short, flattened articles, dactyli falciform. Derivation of name. Indicating an ancestral form of Symethis., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 104, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640

Cenomanocarcinus cantabricus n. sp. (Fig. 17E, F) Diagnosis. Carapace of medium size for genus, dorsal surface weakly convex in both directions; anterolateral margins arched, with 4 teeth of similar size, additional larger tooth corresponding to epibranchial region; posterolateral margins with 2 teeth, anteriormost close to epibranchial tooth; dorsal grooves weakly marked; dorsal regions with few, coarse, blunt tubercles; hepatic region without clear tubercles. Derivation of name. In reference to Cantabria, the region in northern Spain from where specimens of the new species were collected. Material examined. Holotype, a complete carapace with the majority of cuticle preserved (MGSB75431) (ex Manuel Díaz Collection); paratype (MGSB75423) (ex Pedro Artal Collection), a carapace with fragments of cuticle preserved, both from the Cabo de Ajo sea cliff, within the township of Bareyo (Cantabria) from levels considered to be of Albian age (Baron-Szabo & Fernández-Mendiola 1997). Description. Carapace medium sized for genus, subcircular in outline, wider than long; maximum width at epibranchial spine; front advanced, trilobate, with 2 additional blunt teeth at inner orbital corner; orbits small, directed forwards, with 2 short fissures in supraorbital margin, outer orbital node robust, fairly salient; anterolateral margin long, arched, with 5 teeth (excluding outer orbital node, including epibranchial spine), anterior 4 stout, fairly salient, of similar size; epibranchial tooth not completely preserved, appears to have been strong; posterolateral margin longer, converging backwards, sharp edged, with 2 small teeth, anterior one close to epibranchial node, second near posterior corner; posterior margin axially concave, defined by fine rim behind a narrow groove, slightly wider than orbitofrontal margin; dorsal carapace regions weakly vaulted, defined by shallow grooves, with coarse, blunt tubercles; protogastric region with 2 small tubercles; hepatic region barely defined with indistinct inflations, no evident tubercles present; mesogastric region with 2 elongated tubercles; epibranchial region with faint ridge, with large medial tubercle, additional barely inflated lateral tubercle; postbranchial regions with 2 longitudinally positioned tubercles, anterior one larger, rounded, weakly raised, posterior one close to end of lateral margin; cardiac region relatively broad, defined by elongated tubercle, bounded by shallow lateral grooves; intestinal region small, somewhat depressed, bounded by 2 small tubercles; cervical groove weakly marked, even axially, but clearly notching anterolateral margin; branchiocardiac groove well defined at inner portion of epibranchial region. Ventral parts or appendages not preserved. Dorsal carapace surface with dense, small granules. Remarks. The new species is assigned to Cenomanocarcinus on the basis of the curvature and armature of the anterolateral margin, the strong epibranchial spine, the narrow orbitofrontal margin, triangular five-spined rostrum, division of coarse tubercles on the dorsal carapace, and possession of three ridges on the posterior carapace. The new species can be clearly distinguished from congeners on account of the shallow, weakly defined dorsal grooves; the longitudinal and transverse ridges being only weakly salient; the dorsal surface with few dorsal tubercles; the lack of tubercles in the hepatic region; the posterior half of the carapace with large, blunt tubercles; and the anterior tooth of the posterolateral margin being positioned close to the epibranchial spine. The only other European species, C. inflatus, shows more marked transverse and longitudinal ridges with more numerous, evenly spaced tubercles and more convex branchial ridges. There is also a strong posterolateral tooth at mid-length, instead of being closer to the epibranchial tooth; there is an additional anterolateral tooth and the hepatic region bears several clear tubercles. Of the American species, C. beardi exhibits a more subhexagonal carapace, with more divergent posterolateral margins, more salient longitudinal and transverse ridges, stronger dorsal tubercles, a longer epibranchial and posterior anterolateral spine and more distinct H-shaped grooves in the cardiac region. Cenomanocarcinus vanstraeleni exhibits a more subhexagonal carapace; more salient transverse and longitudinal ridges, with more numerous tubercles; the antero- and posterolateral margins with numerous small denticles. Cenomanocarcinus oklahomensis is characterised by stronger longitudinal and transverse ridges, the H-shaped groove pattern in the cardiac region being strongly marked; C. pierrensis exhibits a more distinct, continuous cervical groove; the dorsal tubercles are more raised and conical; the branchial ridges are more salient; the tubercles in the posterior branchial region are not lined up. Cenomanocarcinus pierrensis exhibits a mixed set of characters, which make it difficult to ascribe this species with certainty to Cenomanocarcinus or Necrocarcinus. More completely preserved material is needed to decide in this matter; for the time being, it is retained in the former genus. Two species from Nigeria were recently described by Collins (2010). Cenomanocarcinus tenuicarinatus, of early Turonian age, has a more elongated carapace; posterolateral margins lack strong teeth and clearly tuberculate hepatic ridges; the axial carina is more salient and continuous, less tuberculate, whereas C. dissimilis, from the lower Cenomanian, exhibits a wider carapace with more distinct longitudinal ridges, clear hepatic tubercles and a narrower cardiac region.

Cenomanocarcinus inflatus Van Straelen, 1936 (Fig. 18A, D) Material examined. Dorsal carapace, upper Cenomanian, Butte de Gazonfier, Le Mans, France (MNHN J08587 [see Van Straelen 1936: pl. 4, fig. 8]; MAB k. 2968 represents a cast); dorsal carapace, same provenance (MNHN R05504, the original of A. Milne-Edwards in Guillier 1886: 244); dorsal carapaces, same provenance (MHN LM 3804, MHN LM 3806). Remarks. Specimen MNHN J08587 (Fig. 18A, D) was illustrated by Van Straelen (1936: pl. 4, fig. 8) as ‘topotype’. The name proposed by A. Milne-Edwards in Guillier (1886), was a nomen nudum because a diagnosis was not given. Van Straelen (1936: 37, 38) stated that the type (A. Milne-Edwards’ original) was probably in the MNHN palaeontological collections. This specimen was designated lectotype by Breton & Collins (2007: 18). MNHN J08587 is, however, a carapace from the Hébert Collection (Collections de la Sorbonne), whereas the original of A. Milne-Edwards bears registration number MNHN R05504. The latter is considered here the holotype, of C. inflatus (see ICZN 1999, Art. 73.1.1), and Van Straelen (1936) as the author of the taxon, because he was the first to present a diagnosis, retaining the name suggested earlier by A. Milne-Edwards in Guillier (1886). The lectotype designation by Breton & Collins (2007) is therefore invalid., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 56, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Van Straelen, V. (1936) Crustaces decapodes nouveaux ou peu connus de l'epoque cretacique. Bulletin du Musee royal d'Histoire naturelle de Belgique, 12 (45), 1 - 50, pls. 1 - 4.","Guillier, A. (1886) Geologie du Departement de la Sarthe, 430 pp. Monnoyer, Le Mans.","Breton, G. & Collins, J. S. H. (2007) Decapod fauna from the Cenomanian stratotype. In: Garassino, A., Feldmann, R. M. & Teruzzi, G. (Eds.), 3 rd Symposium on Mesozoic and Cenozoic decapod crustaceans, Museo di Storia Naturale di Milano, May 23 - 25, 2007. Memorie della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 35, 17 - 20.","ICZN (1999) International Code of Zoological Nomenclature (Fourth edition), xxix + 306 pp. The International Trust for Zoological Nomenclature, London."]}

Infraorder Brachyura Latreille, 1802 Remarks. Guinot et al. (in press) is followed for the authorship of the infraorder Brachyura., Published as part of Van Bakel, Barry W. M., Guinot, Danièle, Artal, Pedro, Fraaije, René H. B. & Jagt, John W. M., 2012, A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata) 3215, pp. 1-216 in Zootaxa 3215 (1) on page 13, DOI: 10.11646/zootaxa.3215.1.1, http://zenodo.org/record/5248640, {"references":["Latreille, P. A. (1802) Histoire naturelle, generale et particuliere, des Crustaces et des Insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie du Cours complet d'Histoire naturelle redige par C. S. Sonnini, membre de plusieurs Societes savantes, 3, xii + 467 pp. Dufart, Paris."]}