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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).

Dinocarcinus velauciensis gen. nov., sp. nov. urn:lsid:zoobank.org:act: A4528EE1-0899-46E7-AB72-A1D763D8A4D1 Dinocarcinus velauciensis Van Bakel, Hyžný, Valentin & Robin in Robin, Van Bakel, Hyžný, Cincotta, Garcia, Charbonnier, Godefroit & Valentin, 2019: 4, figs. 1–3 [unavailable]. Type material. HOLOTYPE: MMS /VBN.00.004; PARATYPES 1–4: MMS /VBN.02.94, 09.132d, 12.A.003, 12.A.006 (see Robin et al. 2019: table 1), housed in the palaeontological collections of Velaux municipal institutions (Musée du Moulin Seigneurial /Velaux-La Bastide Neuve). Etymology. From the type locality: Velaux-La Bastide Neuve, Bouches-du-Rhône, southern France. Diagnosis. As for genus. Remarks. A full description and figures of Dinocarcinus velauciensis gen. nov., sp. nov. is given in Robin et al. (2019).

Van Bakel, Barry W. M., Hyžný, Matúš, Valentin, Xavier, Robin, Ninon (2023): Validation of Dinocarcinus velauciensis Van Bakel, Hyžný, Valentin & Robin, a fossil crab (Crustacea, Decapoda, Brachyura) from Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity, southern France. Zootaxa 5315 (5): 483-484, DOI: 10.11646/zootaxa.5315.5.5, URL: http://dx.doi.org/10.11646/zootaxa.5315.5.5

Dinocarcinus gen. nov. urn:lsid:zoobank.org:act: 96EBB8A3-15BE-4E6D-9461-981A163ECED6 Dinocarcinus Van Bakel, Hyžný, Valentin & Robin in Robin, Van Bakel, Hyžný, Cincotta, Garcia, Charbonnier, Godefroit & Valentin, 2019: 2, figs. 1–3 [unavailable]. Etymology. Denoting the actual association with dinosaur (ornithopodan) remains. Type species. Dinocarcinus velauciensis gen. nov., sp. nov. Diagnosis. Chelae large and massive. Fingers gaping, arched, with strong teeth, proximal tooth molariform. Fixed finger dorsal surface with single ‘pitted groove’, palm surface smooth, articulation with dactylus oblique, prominent (in accordance with Van Bakel, Hyžný, Valentin & Robin in Robin et al. 2019)., Published as part of Van Bakel, Barry W. M., Hyžný, Matúš, Valentin, Xavier & Robin, Ninon, 2023, Validation of Dinocarcinus velauciensis Van Bakel, Hyžný, Valentin & Robin, a fossil crab (Crustacea, Decapoda, Brachyura) from Upper Cretaceous (Campanian) continental deposits of Velaux and vicinity, southern France, pp. 483-484 in Zootaxa 5315 (5) on page 483, DOI: 10.11646/zootaxa.5315.5.5, http://zenodo.org/record/8142411, {"references":["Robin, N., Van Bakel, B. W. M., Hyzny, M., Cincotta, A., Garcia, G., Charbonnier, S., Godefroit, P. & Valentin, X. (2019) The oldest freshwater crabs: claws on dinosaur bones. Scientific Reports, 9, 20220. https: // doi. org / 10.1038 / s 41598 - 019 - 56180 - w"]}

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.

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)"]}

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., Published as part of Fraaije, Ren�� H. B., Van Bakel, Barry W. M., Jagt, John W. M., Charbonnier, Sylvain, Schweigert, Guenter, Garcia, G��raldine & Valentin, Xavier, 2022, The evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-the-art-report, pp. 1-16 in Geodiversitas 44 (1) on page 12, DOI: 10.5252/geodiversitas2022v44a1, http://zenodo.org/record/5834191, {"references":["HENDERSON J. R. 1888. - Scientific results of the exploratory voyage of HMS Challenger (Zoology), in Report on the Anomura collected by H. M. S. Challenger during the years 1873 - 1876 27: 1 - 221. HMSO, Edinburgh. https: // www. biodiversitylibrary. org / page / 2007958","LEMAITRE R., RAHAYU D. L. & KOMAI T. 2018. - A revision of \" blanket-hermit crabs \" of the genus Paguropsis Henderson, 1888, with the description of a new genus and five new species (Crustacea, Anomura, Diogenidae). ZooKeys 752: 17 - 97. https: // doi. org / 10.3897 / zookeys. 752.23712"]}

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

Family PROBEEBEIDAE Boone, 1926, emended herein Probeebeidae Boone, 1926b: 73. TYPE GENUS. ��� Probeebei (Boone, 1926) by monotypy. OTHER GENERA INCLUDED. ��� Labidochirus Benedict, 1892, Tisea Morgan & Forest, 1991 and Tylaspis Henderson, 1885. DIAGNOSIS. ��� Shield (excluding rostrum) strongly convex, well calcified, width equal to length or width exceeding length, with distinct bulges (i.e., keraial and massetic regions) laterally, strong to weaker spinose ornament. Rostrum well developed, exceeding lateral projections. Cervical and branchial grooves subparallel, encompassing small branchial regions. Posterolateral margins spinose. Posterior carapace well calcified, broadly inflated, with dense spinose ornamentation. Well-developed cardiac grooves encompassing cardiac region. REMARKS. ��� The bulges on the shield of extant probeebeids (Fig. 10) are quite similar to those of the most basal paguropsid, Eopaguropsis nidiaquilae (see Fraaije et al. 2012c: figs 2c; 3), which also has a spinose ornamentation on the gastric and lateral parts of the shield., Published as part of Fraaije, Ren�� H. B., Van Bakel, Barry W. M., Jagt, John W. M., Charbonnier, Sylvain, Schweigert, Guenter, Garcia, G��raldine & Valentin, Xavier, 2022, The evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-the-art-report, pp. 1-16 in Geodiversitas 44 (1) on page 11, DOI: 10.5252/geodiversitas2022v44a1, http://zenodo.org/record/5834191, {"references":["BOONE L. 1926 b. - A new family of Crustacea. Preliminary technical description. New York Zoological Society Bulletin 29: 73.","BENEDICT J. E. 1892. - Preliminary descriptions of thirty-seven new species of hermit crabs of the genus Eupagurus in the U. S. National Museum. Proceedings of the United States National Museum 15: 1 - 26. https: // doi. org / 10.5479 / si. 00963801.15 - 887.1","MORGAN G. J. & FOREST J. 1991. - A new genus and species of hermit crab (Crustacea, Anomura, Diogenidae) from the Timor Sea, north Australia. Bulletin du Museum national d'histoire naturelle, 4 eme serie, Section A: Zoologie, biologie et ecologie animales 13 (1 - 2). https: // www. biodiversitylibrary. org / page / 59053247","HENDERSON J. R. 1885. - Diagnoses of new species of Galatheidae collected during the \" Challenger \" expedition. Annals and Magazine of Natural History (ser. 5) 16: 407 - 421.","FRAAIJE R. H. B., KRZEMINSKI W., VAN BAKEL B. W. M., KRZEMINSKA E. & JAGT J. W. M. 2012 c. - The earliest record of a diogenid hermit crab from the Late Jurassic of the southern Polish Uplands, with notes on paguroid carapace terminology. Acta Palaeontologica Polonica 57: 655 - 660. https: // doi. org / 10.4202 / app. 2011.0052"]}

Os camarões-alevinos (Decapoda: Caridea: Alpheidae) constituem um dos grupos-modelo para inferências que visam compreender a evolução de características estruturais, comportamentais e ecológicas complexas entre invertebrados marinhos bentônicos. Apesar de ser um táxon super-diversificado com uma ampla distribuição geográfica, o registro fóssil de alpheid ainda é pouco conhecido. No entanto, os dados aqui apresentados mostram que as pontas dos dedos fortemente calcificadas das garras de agarramento do alpheid não são incomuns no registro fóssil e devem ser consideradas um novo tipo de mesofossil. Os vestígios cenozóicos aqui analisados ​​representam uma correspondência estrutural convincente com espécies existentes de Alpheus Baseado na presença de vários morfotipos distintos de garras de dedos, a principal radiação das linhagens de Alpheus é Estima-se que tenha ocorrido já aos 18 anos. Além disso, o registro fóssil mais antigo de alpheids em geral pode agora ser confirmado para o Oligoceno Superior (27-28 mya), proporcionando assim uma nova idade mínima para todo o grupo, bem como o primeiro ponto de calibração confiável para inferências filogenéticas profundas. Alpheid snapping shrimps (Decapoda: Caridea: Alpheidae) constitute one of the model groups for inferences aimed at understanding the evolution of complex structural, behavioural, and ecological traits among benthic marine invertebrates. Despite being a super-diverse taxon with a broad geographical distribution, the alpheid fossil record is still poorly known. However, data presented herein show that the strongly calcified fingertips of alpheid snapping claws are not uncommon in the fossil record and should be considered a novel type of mesofossil. The Cenozoic remains analysed here represent a compelling structural match with extant species of Alpheus. Based on the presence of several distinct snapping claw-fingertip morphotypes, the major radiation of Alpheus lineages is estimated to have occurred as early as 18 mya. In addition, the oldest fossil record of alpheids in general can now be confirmed for the Late Oligocene (27–28 mya), thus providing a novel minimum age for the entire group as well as the first reliable calibration point for deep phylogenetic inferences.

Basadromia n. gen. Type species. Basadromia longifrons n. sp. Diagnosis. Small carapace, longitudinally subelliptical in outline, slightly longer than wide, maximum width at position of epibranchial region; markedly convex in both directions; front fairly projecting beyond orbits, narrow, with notable V-shaped notch and deep groove, bearing four long, subtriangular teeth plus short rostral tooth situated in lower plane; orbits small, anterolaterally disposed, supraorbital margin with strong supraorbital (inner orbital) tooth, anterior portion of supraorbital margin nearly vertical; lateral margins of carapace broadly arched, with short, subtle teeth; posterior margin narrow, being narrower than orbitofrontal margin; dorsal surface strongly areolated, uniformly granular; dorsal regions very well defined and individualised by swellings and grooves; epibranchial regions large, elongated, posterior portion distinctly swollen; mesogastric region transversely subelliptical, swollen, with subtle axial depression, broad subtriangular anterior extension; epibranchial region large, divided into 2 portions, axial portion strongly swollen; cardiac region large, markedly swollen, inverted subpentagonal in shape; cervical, branchial grooves clearly marked. Dorsal surface of carapace, chelipeds densely, uniformly granular. Etymology. From Basa, the name of the valley in the province of Huesca (Aragón, Spain) from where the new form was recovered, and - dromia, the common suffix for members of the family and superfamily. Remarks. The new genus differs from all extinct and extant members assigned to the Dromioidea (Guinot and Tavares, 2003; Ng et al., 2008; Karasawa et al., 2011; McLay, 2001 a, b; Schweitzer et al., 2012) in having four strong, conspicuously long, frontal teeth, a narrow posterior carapace margin, the posterior portion of the protogastric region being nearly hemispherical and the axial portion of the epibranchial region strongly swollen, subelliptical in shape.

Basadromia longifrons n. sp. (Fig. 1���4) Diagnosis. As for genus. Etymology. From the Latin longus, meaning long and frons, front, in allusion to the fairly projected front, with long subtriangular teeth. Material examined. The holotype, MPZ 2011.185, and six paratypes, MPZ 2013.73��� MPZ 2013.78, all from the Upper Eocene (Priabonian) in the Yebra de Basa area, Huesca. Maximum carapace width and length in the holotype are 14.1 and 16.4 mm, respectively. Description. Carapace transversely elliptical in outline; slightly longer than wide, widest anteriorly, at level of epibranchial region; dorsal surface strongly convex in both directions. Front strongly projected, narrow, with noticeable V-shaped notch, deep groove; frontal margin defined by 4 long, subtriangular teeth, short rostral tooth in lower plane, not visible in dorsal view. Orbits small, somewhat rimmed, directed anteriorly, laterally; supraorbital margin nearly vertical in inner portion (Fig. 1). Anterolateral margin short, arched, densely granular, with small, subtle teeth. Posterolateral margin longer than anterolateral, broadly arched. Posterior margin slightly concave, notably reduced, shorter than orbitofrontal margin. Dorsal carapace surface very well defined by numerous, strong swellings, fairly marked grooves. Epigastric regions small, slightly swollen, separated by frontal groove. Protogastric regions large, elongated, posterior portion nearly circular, strongly swollen. Mesogastric region transversely subelliptical, swollen, with axial depression; anterior extension large, broadly subtriangular. Hepatic region slightly swollen, bounded by notable cervical, hepatogastric grooves. Urogastric region tranversely subrectangular, bounded by lateral grooves, divided into 2 portions by short axial depression. Epibranchial region notably large, divided into 2 portions by longitudinal groove; axial portion large, subelliptical, strongly swollen. Posterior branchial regions large, broadly swollen. Cardiac region large, gently swollen, subpentagonal in shape, with apex pointed downwards. Intestinal region small, flat. Cervical groove well marked, continuous from side to side, notching lateral margins. Branchial groove deep, oblique in lateral portions, converging abruptly towards rear in posterior portion of cardiac region. Dorsal carapace surface densely, uniformly covered by small granules. Chelipeds not completely preserved, short, robust, strongly granular; moveable finger thin, delicate, strongly curved (Fig. 3). Remarks. The dorsal carapace of Basadromia longifrons n. gen., n. sp. exhibits the main characters of the family Dromiidae (sensu Guinot and Tavares, 2003; Ng et al., 2008; Guinot et al., 2013), including general carapace outline, shape and distribution of dorsal regions, presence of well-defined cervical, branchial, and branchiocardiac grooves, a narrow and projected front with two inner orbital teeth, two marked median teeth, and a rostral tooth situated in a lower plane. Following recent revisions and general classifications (Guinot and Tavares, 2003; Schweitzer and Feldmann, 2010; Karasawa et al., 2011; Schweitzer et al., 2012; Guinot et al., 2013), these main features, as well as the distribution and shape of dorsal regions in Basadromia n. gen., are also seen in members of various subfamilies. The epigastric, protogastric, mesogastric, urogastric, hepatic, epibranchial, and cardiac regions, either weak to strongly marked, are similar in general shape and distribution in extinct forms assigned to the Dromiinae, Basinotopinae, Sphaerodromiinae and Dynomenidae (compare Schweitzer et al., 2012). With regard to dorsal carapace grooves (frontal, cervical, branchial, and branchiocardiac) are shared by representatives of the various subfamilies as well. The mesogastric region (of fairly constant shape amongst dromioids) in Basadromia n. gen. resembles that of other members of the superfamily. This region, which is either weakly to strongly divided, with two somewhat swollen portions separated by either a faint or deep longitudinal depression, usually with the anterior extension broadly triangular, shows rather constant features in extinct genera such as Dromilites, Ferricorda Schweitzer & Feldmann, 2010, Pseudodromilites Beurlen, 1928, Basinotopus, and even Kromtitis M��ller, 1984 (Schweitzer & Feldmann, 2010; Karasawa et al., 2011; Schweitzer et al., 2012). The new form yet exhibits a set of distinctive features that set it apart from all known fossil and extant dromioids, warranting the erection of a new genus. The front is distinctly projected, with four long teeth, the two median ones being longer, the two inner orbital ones shorter; the rostral tooth, situated in a lower plane, is short, not visible in dorsal view; the anterolateral margin is broadly arched, densely granular, bearing small teeth or protuberances; the posterolateral margins are broadly arched, the posterior margin is fairly narrow; the posterior portion of the protogastric region presents a near-circular, subhemispherical swelling; the epibranchial region is divided into two portions by a longitudinal groove, and the axial portion is strongly swollen and subelliptical in shape. This set of features differentiates the new taxon from all known extinct and extant forms. Basadromia n. gen. appears to be close to the extant genus Petalomera Stimpson, 1858, and the extinct Pseudodromilites. The former is characterised by a granular dorsal surface of the carapace, well-defined regions with numerous swellings and grooves (similar to the new taxon), granular anterolateral margins, usually without long spines or noticeable teeth, and a relatively projected front with a single rostral tooth in lower plane, two lateral median teeth and supraorbital (inner orbital) lobes (McLay & Ng, 2007). Petalomera, however, presents a less projected front than Basadromia n. gen., with the rostral tooth clearly visible in dorsal view and the inner orbital teeth (supraorbital) are less salient, the outer orbital nodes less pronounced. For describing orbitofrontal construction different terms have been used in the literature to capture the same morphology. Here we use ���frontal margin, front, rostral tooth, lateral median teeth, and inner orbital teeth���. Collins & Jakobsen (2004) also used the terms ���front��� and ���inner orbital���; ���produced front���slender and slightly diverging inner orbital spines��� (Collins & Jakobsen, 2004: 69). These authors considered dromioids to have usually a frontal margin with one rostral tooth and two lateral median teeth isolated (i.e., differentiated) from the inner orbital teeth or lobes, all the lobes from weakly to strongly marked in the different genera, but to some degree always present (Collins & Jakobsen, 2004: 70). Guinot & Tavares (2003) usually described the front as ���front with median rostrum and two pseudorostral teeth, one at each side of the rostrum��� and ���supraorbital teeth well developed��� (Guinot & Tavares, 2003: 49, 50). McLay (2007: 109) employed ���rostrum���, rather than ���front���, ���lateral teeth��� instead of ���median frontal teeth��� (or ���pseudorostral teeth���) and ���supraorbital teeth��� instead of ���inner orbital teeth���; ���rostrum tridentate, median tooth deflexed, lateral teeth separated by a V-shaped sinus���well developed supraorbital tooth���. In the absence of well-preserved ventral features, we tentatively assign the new taxon to the Dromiinae. Pseudodromilites, included in the Dromiidae by Schweitzer & Feldmann (2010), presents a dorsal surface densely covered by granules; the distribution and shape of dorsal regions is similar to that seen in Basadromia n. gen. Pseudodromilites, however, has a subpentagonal carapace shape, with near-straight portions of posterolateral margins and the posterior margin is longer; the dorsal regions are more uniformly swollen, less protuberant and subdivided, not clearly differentiated as in Basadromia n. gen. The frontal margin presents only two median teeth and nearly undifferentiated inner orbital lobes. The anterolateral margin in Pseudodromilites was described as having triangular teeth by Schweitzer et al. (2010: 422), due to the appearance conveyed in drawings of the original descriptions. A good illustration of a complete specimen (Beschin et al., 2012: 32, fig. 25; pl. 4 / 2) confirms that the anterolateral margin is slightly notched, with two granular protuberances, but lacking triangular teeth. Basadromia n. gen. appears to be morphologically close to some fossil species that have recently been assigned to the subfamily Sphaerodromiinae Guinot & Tavares, 2003 (see Schweitzer & Feldmann, 2010). Dromilites bucklandii H. Milne Edwards, 1837, the type species of the genus, also presents a frontal margin with one rostral tooth, two notable median teeth and two supraorbital teeth, thus five frontal teeth in total. Dromilites nevertheless presents a less salient front, with smaller frontal teeth, the two inner orbital ones being fairly smaller, the lateral margins of the carapace have salient teeth, and the dorsal carapace surface shows different dorsal regions, being more individualised, nearly hemispherical. Dromilites simplex Quayle & Collins, 1981 presents similar characters. The frontal margin is less projecting, without pronounced teeth or spines, and in general the dorsal regions are weakly defined, less swollen and separate. This species exhibits a typical marked groove, subparallel, between the cervical and branchial grooves (Schweitzer et al., 2010: fig. 4). Basadromia n. gen. shares with Dromilites vicensis Barnolas, 1973, the two fairly long median teeth in the front, and the deep, broadly V-shaped notch (Barnolas, 1973, figs. 2, 4). Despite that the dorsal regions and grooves are weakly pronounced in D. vicensis, some characters, such as the shape of the mesogastric, urogastric and cardiac regions, appear to be closey similar to those of the new species. The main differences in D. vicensis are the absence of marked supraorbital spines (teeth or spines in the orbitofrontal corner) and the presence of salient teeth along the anterolateral margins. In spite of sharing some diagnostic features such as the general carapace outline, the produced front and dorsal regions and grooves in D. pastoris Via, 1959, differ from Basadromia n. gen. There are notable lobes along the lateral carapace margins, the front is axially produced, the rostral tooth being the more projected one, and the four adjacent few marked, notably small; dorsal regions are more uniformly swollen, less individualised; there is a subparallel dorsal groove between the cervical and branchial grooves, the post-cervical groove. This groove is very characteristic in some dromioids, being notably marked in genera such as Pseudodromilites, Basinotopus, and Ferricorda. A cast of the holotype of D. pastoris, illustrated by Schweitzer et al. (2010: fig. 3) is either broken or incomplete; the original specimen, housed in the MGSB collections, has recently been reexamined by the senior author and does exhibit a complete front. The holotype of D. pastoris actually possesses a very complete front, showing the rostral tooth to be the more produced (Via, 1959; Via, 1969: fig. 10; pl. 4). The Italian specimen assigned to D. pastoris also exhibits the same features, as clearly indicated in the text and illustrations (Beschin et al., 2012: 34, fig. 26; pl. 4). Quinquerugatus, assigned to Dromiidae (Schweitzer et al., 2010), presents numerous characters that are also present in D. pastoris, as mentioned by Beschin et al. (2012: 34), despite the fact that the two genera were assigned to two different subfamilies, Dromiinae and Sphaerodromiinae, respectively (sensu Schweitzer & Feldmann, 2010). The carapace is subpentagonal in outline in these two species, the dorsal regions are weakly marked and the frontal margin has a median spine that in both species is the more projected in the front. Dromilites pastoris was assigned to Sphaerodromiinae despite the produced rostral tooth which is utterly different from all other assigned members, whereas Dromilites was diagnosed as having a clearly ���bilobed rostrum��� (Schweitzer et al., 2010: 418). All data mentioned above clearly indicate that much more work is needed to make assignment of the various dromiid species clearer. The Basinotopinae is a recently erected subfamily that is characterised by a subtriangular carapace shape, being notably longer than wide, a triangular front with three lobes (spines), the rostral one being the longer, and lateral carapace margins with clearly marked spines (Karasawa et al., 2011: 539). Basinotopus has the strongly swollen carapace regions and the produced front in common with Basadromia n. gen. The dorsal regions present a general aspect and similar distribution, are numerous and well divided by numerous grooves in both genera. Basinotopus lamarckii and B. tricornis Collins & Jakobsen, 2004 are easily distinguished from Basadromia longifrons n. gen., n. sp. in that both have a subtriangular front with a markedly long rostral spine, notably more projecting than the two inner orbital spines; extremely long spines along the anterolateral and posterolateral margins of carapace; a longer posterior carapace margin, at least of the same size of the orbitofrontal margin. Schweitzer & Feldmann (2010: 422) mentioned that Dromilites alpina Glaessner, 1929, and Dromilites lothi F��rster & Mundlos, 1982 should be assigned to Kromtitis. A close reading of the text and a careful examination of the images for D. alpina confirms that this species exhibits a subtriangular carapace, with typical strong, long lateral spines and cervical and branchial grooves characteristic of the configuration in Basinotopus. Collins & Jakobsen (2004: 70) also mentioned that this species should be assigned to Basinotopus. Dromilites lothi presents a similar carapace outline, lateral spines along the lateral carapace margins, distribution of dorsal regions and shape of cervical, branchial and branchiocardiac grooves (F��rster & Mundlos, 1982: 155, figs. 5, 6). Despite the fact that the frontal margin appears to be different, we consider this species to belong to Basinotopus, but for now we retain it in Dromilites until better-preserved material becomes available. Kromtitis is characterised by a subquadrate carapace outline, a broad and near-straight frontal margin in dorsal view, dorsal regions that are subdivided into numerous portions, presenting a range of small swellings and absence of cervical and branchial grooves that are typical of genera such as Dromilites, Basinotopus, or Pseudodromilites (Beschin et al., 2007: pl. 3; Schweitzer et al., 2012: fig. 21). Reasons to exclude B. alpina and D. lothi from Kromtitis were discussed by Van Bakel et al. (2009: 49, 50)., Published as part of Artal, Pedro, Van Bakel, Barry W. M., Dom��nguez, Jos�� L. & G��mez, Guillermo, 2016, A new dromiid crab (Crustacea, Brachyura, Dromioidea) from the Upper Eocene of Huesca (Arag��n, northern Spain), pp. 438-446 in Zootaxa 4061 (4) on pages 440-445, DOI: 10.11646/zootaxa.4061.4.8, http://zenodo.org/record/270404, {"references":["Guinot, D. & Tavares, M. (2003) A new subfamilial arrangement for the Dromiidae de Haan, 1833, with diagnoses and descriptions of new genera and species (Crustacea, Decapoda, Brachyura). Zoosystema, 25 (1), 43 - 129.","Guinot, D., Tavares, M. & Castro, P. (2013) Significance of sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa. Zootaxa, 3665 (1), 1 - 414. http: // dx. doi. org / 10.11646 / zootaxa. 3665.1.1","Karasawa, H., Schweitzer, C. E. & Feldmann, R. M. (2011) Phylogenetic analysis and revised classification of podotrematous brachyurans (Decapoda) including extinct and extant families. Journal of Crustacean Biology, 31 (3), 523 - 565. http: // dx. doi. org / 10.1651 / 10 - 3423.1","Schweitzer, C. E., Feldmann, R. M. & Karasawa, H. (2012) Part R, Revised, Volume 1, Chapter 8 M: Systematic decriptions: Infraorder Brachyura, Section Dromiacea. Treatise Online, 51, 1 - 43.","McLay, C. L. & Ng, P. K. L. (2007) Revision of the Indo-West Pacific sponge crabs of the genus Petalomera Stimpson, 1858 (Decapoda: Brachyura: Dromiidae). The Raffles Bulletin of Zoology, 55 (1), 119 - 132.","Collins, J. S. H. & Jakobsen, S. L. (2004) New crabs (Crustacea, Decapoda) from the Eocene (Ypresian / Lutetian) Lillebaelt Clay Formation of Jutland, Denmark. Bulletin of the Mizunami Fossil Museum, 30 (2003), 63 - 96.","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 Naturale \" G. Zannato \", Montecchio Maggiore, Vicenza, 100 pp.","Quayle, W. J. & Collins, J. S. H. (1981) New Eocene crabs from the Hampshire Basin. Palaeontology, 24, 733 - 758.","Barnolas, A. (1973) Dromilites vicensis n. sp. nuevo braquiuro del Eoceno marino de Cataluna. Instituto de Investigaciones Geologicas, 28, 5 - 13.","Via, L. (1959) Decapodos fosiles del Eoceno espanol. Boletin del Instituto Geologico y Minero de Espana, 70, 331 - 402.","Via, L. (1969) Crustaceos decapodos del Eoceno espanol. Pirineos, 91 - 94, 1 - 469.","Forster, R. & Mundlos, R. (1982) Krebse aus dem Alttertiar von Helmstedt and Handorf (Niedersachsen). Palaeontographica, A 179, 148 - 184.","Beschin, C., Busulini, A., De Angeli, A. & Tessier, G. (2007) I decapodi dell'Eocene inferiore di Contrada Gecchelina (Vicenza, Italia settentrionale) (Anomura e Brachyura). Museo di Archeologia e Scienze Naturale \" G. Zannato \", Montecchio Maggiore, Vicenza, 76 pp.","Van Bakel, B. W. M., Artal, P., Fraaije, R. H. B. & Jagt, J. W. M. (2009) A new Early Oligocene crab (Decapoda, Brachyura, Dromiacea) from northwest Belgium, with comments on its palaeobiology. Geologica Belgica, 12, 45 - 57."]}

Artal, Pedro, Van Bakel, Barry W. M., Domínguez, José L., Gómez, Guillermo (2016): A new dromiid crab (Crustacea, Brachyura, Dromioidea) from the Upper Eocene of Huesca (Aragón, northern Spain). Zootaxa 4061 (4): 438-446, DOI: http://doi.org/10.11646/zootaxa.4061.4.8

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

Paguristes liwinskii n. sp. (Fig. 1 C) Diagnosis. Cross section of right cheliped transversely oval; dorsal and ventral sides convex, with regular, dense cover of forwardly directed, fine tubercles, largest arranged in row along dorsal outer side and along central proximal inner side; fixed finger short, stout and triangular; dactylus short and triangular, about twice length of fixed finger. Etymology. Named in honour of the mayor of Annopol, Mr Wiesław Liwiński, who facilitated large-scale excavations conducted in this area in recent years by a team led by the last author (for details, see Machalski & Kennedy 2013 and Popov & Machalski 2014). Type. The holotype, and sole specimen known (ZPAL Cr. 9 / 43), is a near-complete right palm with fixed finger and dactylus, measuring 10 mm in length, and 7 mm in greatest width. Locality and stratigraphy. Kopiec near Annopol, lower part of unit 3 (Fig. 1 A), upper upper Albian. Unit 3, c. 50 cm thick, is composed of quartz sand with abundant glauconite, passing gradually upwards into quartzglauconitic marl. A distinct bed of phosphatic clasts and fossils occurs in the upper part of unit 3. Its upper boundary is sharp, whereas the lower boundary is diffuse due to bioturbation (Marcinowski & Walaszczyk 1985). The phosphatic clasts are c. 10 mm in diameter, but in the upper part of the layer they commonly form larger aggregates, up to 120 mm in diameter. Unit 3 of the Annopol succession is dated on the basis of specifically indeterminate specimens of the ammonite Mortoniceras (Subschloenbachia), preserved as attachment scars on the left valves of some oysters (Machalski and Kennedy 2013). These specimens are closest to Mortoniceras (Subschloenbachia) that characterise the upper upper Albian Mortoniceras perinflatum Zone. Description. Globose right cheliped, cross section transversely oval; convex dorsal and ventral sides; all sides with regular, dense cover of forwardly directed, fine tubercles, largest in a row along dorsal outer and along central proximal inner sides. Fixed finger short, stout, curved, with a row of about four large setal pores on the inner and outer cutting edge and numerous irregular, longitudinal rows of finer pores on central and ventral parts; tip spatulate; cutting edge near-straight with concave central part. Dactylus short, triangular, curved, about twice the length of fixed finger; covered with irregular rows of setal pores smallest in dorsal half; tip pointed; strongly concave dorsal side here considered to be a preservational artefact, possibly resulting from unsuccessful predatory attack as some regeneration cuticle is visible on the proximal dorsal part of dactylus and the distal part of dorsal palm. Discussion. The generic placement of fossil paguroid chelae is difficult in that claws and, occasionally, meri and carpi are all that is preserved. The specimen studied has been assigned to Paguristes because the claws of Paguristes santamartaensis are remarkably similar and members of Paguristes tend to be isochelous, whereas most other diogenid genera comprise species that are predominantly heterochelous with the left chela considerably larger. We have to keep in mind that the lefthandedness of most diogenids probably is the result of gastropod shell inhabitation, the great majority of marine snails being dextral. Fraaije (2003) noted a shift in molluscan inhabitation from ammonites to gastropods during the Cretaceous Period; this affected the morphology of hermit claws that ���blocked��� the aperture. It is also highly probable that Cretaceous species of Paguristes had isochelous chelae and that left heterochely mainly occurred subsequent to the Cretaceous/Paleogene boundary in relation to this shift in inhabitation. To date, about 120 extant species of Paguristes are on record (McLaughlin et al. 2010; Komai et al. 2015), illustrating that this genus is the most successful member of the Diogenidae by far. From the fossil record, Schweitzer et al. (2010) recorded a total of 15 species. Here, we list 17 species (Tab. 1), inclusive of the new form, which extends the stratigraphic range of Paguristes downwards into the late Albian by about 25 myr. Four of the listed species (P. santamartaensis, P. ouachitensis, P. whitteni and P. florae) are of late Late Cretaceous, eight (P. baldoensis, P. extentus, P. prealpinus, P. wheeleri, P. mexicanus, P. johnsoni, P. hokoensis and P. lineatuberculatus) of Paleogene, two (P. oligotuberculata and P. cserhatensis) of Neogene and two (P. cf. lymani and P. cf. syrtensis) of Quaternary age. Only one of these records, i.e., that of P. cf. syrtensis, is based on a fragmentary dorsal shield (see Garassino et al. 2014). Species Stratigraphic/geographic occurrence P. liwinskii sp. nov. upper Albian, Poland P. santamartaensis Feldmann, Tshudy & Thomson, 1993 upper Santonian/lower Campanian, Antarctica P. ouachitensis Rathbun, 1935 upper Campanian, USA P. whitteni Bishop, 1983 Maastrichtian, USA P. florae Collins, Fraaye & Jagt, 1995 upper Maastrichtian, the Netherlands ��� Belgium P. baldoensis Garassino, De Angeli & Pasini, 2009 Lower Eocene, Italy P. extentus Beschin, Busulini, De Angeli & Tessier, 2007 Lower Eocene, Italy P. prealpinus Beschin, De Angeli, Checchi & Zarantonello, 2005 Middle Eocene, Italy P. wheeleri Blow & Manning, 1996 Middle Eocene, USA P. mexicanus Vega, Cosma, Couti��o, Feldmann, Nyborg, Schweitzer & Waugh, 2001 Middle Eocene, Mexico P. johnsoni Rathbun, 1935 Eocene, USA P. hokoensis Schweitzer & Feldmann, 2001 Upper Eocene, USA P. oligotuberculatus M��ller & Collins, 1991 Upper Eocene, Hungary P. lineatuberculatus Beschin, De Angeli, Checchi & Mietto, 2006 Upper Eocene���Oligocene, Italy P. cserhatensis M��ller, 1984 Miocene, Hungary P. cf. lymani A. Milne-Edwards & Bouvier, 1893 Pleistocene, Jamaica P. cf. syrtensis de Saint Laurent, 1971 Pleistocene, Italy In overall shape and dense uniform ornamentation, Paguristes liwinskii sp. nov. most closely resembles P. santamartaensis, although the latter differs in having a smooth inner surface, a flattened dorsal surface and coarser tuberculation. The new species can be differentiated from P. ouachitaensis in that the palm has a convex inner side and transverse rows of tubercles are lacking. Paguristes whitteni has a much coarser ornament and a longer, strongly deflected fixed finger, while P. f l or a e is less globose, has fingers with a lesser curvature and the ornamentation is more subdued. Paguristes baldoensis and P. extensus are less convex, have straighter dorsal and ventral sides and much longer fingers, whereas P. prealpinus has a smoother inner surface and a much longer and downturned fixed finger. Paguristes lineatuberculatus has a different ornamentation and the distal tip of the fixed finger is directed downwards, whereas P. wheeleri shows a strongly curved occlusal surface and much coarser tubercles and spines on both sides. The ornamentation in P. mexicanus is less uniform, with a reticulate outer surface near the upper margin. Paguristes johnsoni can be differentiated in having a less globose palm, a longer fixed finger, and less dense and coarser ornamentation, while P. hokoensis has a significantly longer, downturned fixed finger and fewer, yet larger tubercles, arranged in longitudinal rows on the outer side of the manus. Paguristes cserhatensis is less globose, and has fewer, yet much larger tubercles arranged in longitudinal rows; in addition, the fixed finger has a lesser curvature. The outer surface of the palm in P. oligotuberculatus has only few rows of coarse tubercles and the dorsal margin is straight and spinose, while P. cf. lymani differs in showing a smoother inner surface, straighter dorsal and ventral margins. Moreover, it does not have a uniform ornamentation. Rathbun (1926: 101) described the preservation of the type and only known specimen of P. subequalis as follows, ��� [.....] so crumbly that no more can be uncovered without destroying it. Only the form of the fingers is seen, and in the case of the propodal finger, nothing but the impression.��� Paguristes chipolensis was erected by Rathbun (1935) on the basis of a single, incomplete dactylus and propodus. The preservation of both P. subequalis and P. chipolensis is so poor that a detailed comparison is impossible; we propose that they are best regarded as nomina dubia and recommend removal from decapod crustacean species lists., Published as part of Fraaije, Ren�� 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, pp. 588-594 in Zootaxa 3955 (4) on pages 589-592, DOI: 10.11646/zootaxa.3955.4.9, http://zenodo.org/record/239809, {"references":["Machalski, M. & Kennedy, W. J. (2013) Oyster-bioimmured ammonites from the Upper Albian of Annopol, Poland: stratigraphic and palaeobiogeographic implications. Acta Geologica Polonica, 63, 545 - 554. http: // dx. doi. org / 10.2478 / agp- 2013 - 0024","Popov, E. V. & Machalski, M. (2014) Late Albian chimeroid fishes (Chimaeroidei) from Annopol, Poland. Cretaceous Research, 47, 1 - 18. http: // dx. doi. org / 10.1016 / j. cretres. 2013.09.011","Marcinowski, R. & Walaszczyk, I. (1985) Middle Cretaceous deposits and biostratigraphy of the Annopol section, Central Polish Uplands. Osterreichische Akademie der Wissenschaften, Schriftenreiche der Erdwissenschaftlichen Komisssionen, 7, 27 - 41.","McLaughlin, P. A., Komai, T., Lemaitre, R. & Rahayu, D. L. (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea). Part I - Lithodoidea, Lomisoidea and Paguroidea. The Raffles Bulletin of Zoology, 23 (Supplement), 5 - 107.","Komai, T., Reshmi, R. & Kumar, A. B. (2015) A new species of the hermit crab genus Paguristes Dana, 1851 (Crustacea: Decapoda: Anomura: Diogenidae) from southwestern India. Zootaxa, 3937 (3), 517 - 532. http: // dx. doi. org / 10.11646 / zootaxa. 3937.3.5","Schweitzer, C. E., Feldmann, R. M., Garassino, A., Karasawa, H. & Schweigert, G. (2010) Systematic list of fossil decapod crustacean species. Crustaceana Monographs, 10, i - viii + 1 - 221. http: // dx. doi. org / 10.1163 / ej. 9789004178915. i- 222","Garassino, A., Pasini, G., De Angeli, A. & Hyzny, M. (2014) The decapod fauna (Axiidea, Anomura, Brachyura) from the Late Pleistocene of Trumbaca, Reggio Calabria (Calabria, southern Italy). Natural History Sciences, Atti della Societa italiana di Scienze naturali del Museo civico di Storia naturale di Milano, 1 (2), 119 - 130. http: // dx. doi. org / 10.4081 / nhs. 2014.60","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.","Rathbun, M. J. (1935) Fossil Crustacea of the Atlantic and Gulf Coastal Plain. Geological Society of America, Special Paper, 2, 1 - 160. http: // dx. doi. org / 10.1130 / SPE 2 - p 1","Bishop, G. (1983) Fossil decapod Crustacea from the Late Cretaceous Coon Creek Formation, Union County, Mississippi. Journal of Crustacean Biology, 3, 417 - 430. http: // dx. doi. org / 10.2307 / 1548142","Collins, J. S. H., Fraaye, R. H. B. & Jagt, J. W. M. (1995) Late Cretaceous anomurans and brachyurans from the Maastrichtian type area. Acta Palaeontologica Polonica, 40, 165 - 210.","Garassino, A., De Angeli, A. & Pasini, G. (2009) In situ hermit crab (Crustacea, Anomura, Paguroidea from the Early Eocene (Ypresian) of NE Italy. Atti della Societa italiana di Scienze naturali del Museo civico di Storia naturale di Milano, 150 (II), 229 - 238.","Beschin, C., Busulini, A., De Angeli, A. & Tessier G. (2007) I decapodi dell'Eocene inferiore di Contrada Gecchelina (Vicenza - Italia settentrionale) (Anomura e Brachiura). Museo di Archeologia e Scienze Naturali ' G. Zannato ', Montecchio Maggiore (Vicenza), 2007, 9 - 76.","Beschin, C., De Angeli, A., Checchi, A. & Zarantonello, G. (2005) Crostacei eocenici di Grola di Cornedo Vicentino presso Spagnago (Vicenza, Italia settentrionale). Studi e Ricerche - Associazione Amici Museo civico ' G. Zannato', Montecchio Maggiore (Vicenza), 12, 5 - 35.","Blow, W. C. & Manning, R. B. (1996) Preliminary descriptions of 25 new decapod crustaceans from the Middle Eocene of the Carolinas, U. S. A. Tulane Studies in Geology and Paleontology, 29, 1 - 26.","Vega, F. J., Cosma, T., Coutino, M. A., Feldmann, R. M., Nyborg, T. G., Schweitzer, C. E. & Waugh, D. A. (2001) New Middle Eocene decapods (Crustacea) from Chiapas, Mexico. Journal of Paleontology, 75, 929 - 946. http: // dx. doi. org / 10.1666 / 0022 - 3360 (2001) 075 % 3 C 0929: NMEDCF % 3 E 2.0. CO; 2","Muller, P. & Collins, J. S. H. (1991) Late Eocene coral-associated decapods (Crustacea) from Hungary. Contributions to Tertiary and Quaternary Geology, 28, 47 - 92.","Beschin, C., De Angeli, A., Checchi, A. & Mietto, P. (2006) Crostacei del Priaboniano di Priabona (Vicenza - Italia settentrionale). Lavori - Societa Veneziana di Scienze Naturali, Venezia, 31, 95 - 112.","Muller, P. (1984) Decapod Crustacea of the Badenian. Geologica Hungarica, Series Palaeontologica, 42, 1 - 317.","Milne-Edwards, A. & Bouvier, E. L. (1893) Description des Crustaces de la famille des Paguriens recueillis pendant l'expedition. Reports of the results of dredging under the supervision of Alexander Agassiz in the Gulf of Mexico (1877 - 78) and along the Atlantic coast of the United States (1880) by the U. S. Survey steamer \" Blake, \" Lieut. - Com. S. D. Sigsbee and Commander J. R. Bartlett commanding. Memoirs of the Museum of Comparative Zoology, 14, 1 - 172. [Harvard]","Saint Laurent, M. de (1971) Paguristes syrtensis, espece nouvelle de cotes tunisiennes (Crustacea Decapoda Diogenidae). Bulletin du Museum national d'Histoire naturelle de Paris, Series 2, 42 (for 1970), 1009 - 1107.","Rathbun, M. J. (1926) The fossil stalk-eyed Crustacea of the Pacific Slope of North America. United States National Museum Bulletin, 138, i - vii + 1 - 155. http: // dx. doi. org / 10.5479 / si. 03629236.138. i"]}

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 Gaud�� (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)., 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 pages 348-351, DOI: 10.11646/zootaxa.3652.3.3, http://zenodo.org/record/221749

Serrablopluma n. gen. Type species. Serrablopluma diminuta n. sp. Diagnosis. Small size carapace, subtrapezoidal in outline, much wider than long, widest anteriorly, maximum width at position of lateral spines; anterior margin wide, broadly sinuous, terminating in fairly widely projecting lateral spines; front conspicuously reduced, narrow; orbits wide, with sub-orbital spine; lateral margins straight, inclined, converging posteriorly; posterior margin wide, broadly convex; dorsal carapace surface flat in both directions, with 3 strongly marked ridges; anterior ridge continuous, intersecting anterior margin; medium ridge interrupted only by gastric grooves; posterior ridge continuous, straight. Thoracic sternum broad; sternites 3, 4 trapezoidal, sternites 5���7 subrectangular; vulvae large, subcircular, situated below extremity of sutures 5 / 6; abdomen subtriangular in both sexes; female abdomen much broader than males; thoracic sternites, abdominal somites notably ridged; thoracic sternite 5, abdominal somite 6 with distinct abdominal holding mechanism; chelipeds equal, robust, relatively large; P 2 ���P 4 long, flattened; P 5 conspicuously reduced. Etymology. From Serrablo, the mediaeval name of the area (Pyrenees, Huesca) and the ending pluma, which refers to the main character of the family. Remarks. The new genus differs from all others currently included in the Retroplumidae (see De Grave et al. 2009: 41) in having a wide, broadly arched anterior margin, a conspicuously reduced front and an anterior dorsal ridge that intersects the anterior margin at the position of long, fairly projecting lateral spines. Serrablopluma diminuta n. sp. (Figs. 1���2) Diagnosis. As for genus. Etymology. From the Latin diminutus, in allusion to the rather reduced size. Material examined. MGSB 75287, holotype; MGSB 75289, MGSB 75290 a, b, MGSB 78334 a, b, and MGSB 78335 a, b, c, paratypes; MAB k. 3279���3281, MAB k. 3293, MAB k. 3295, paratypes, all from Fanlillo, municipality of Yebra de Basa, Huesca, and of late Eocene (Priabonian) age. In the holotype, the maximum carapace width and length (in millimetres) are 12 and 9, respectively. Description. Carapace small (maximum carapace width 12 mm); subtrapezoidal in outline; wider than long, widest anteriorly at level of lateral spines; dorsal surface flat in both directions. Anterior margin sinuous, broadly arched, with projecting lateral spines. Front conspicuously reduced, narrow, directed forwards; orbits wide, with sub-orbital spine. Lateral margins straight, inclined, converging posteriorly. Posterior margin convex, wide, only slightly shorter than orbitofrontal margin. Dorsal surface with 3 ridges; anteriormost intersecting anterior margins laterally; median ridge defined by epibranchial swellings, interrupted by gastric grooves; posterior ridge straight, continuous, interrupted only by small indentations of subtle branchiocardiac grooves. Dorsal surface smooth except for horizontal ridges, shallow grooves. Mesogastric region defined by 2 faint swellings, bounded by weak posterior grooves. Cardiac region large, gently swollen, sub-rhomboidal in shape. Intestinal region flat. Thoracic sternum much broader than long, sternites with subtle ridges; sternites 1,2 fused, small, subtriangular; sternites 3, 4 subtrapezoidal; sternite 4 with pronounced lateral extensions; small, weak indentation between sternites 3, 4; sternites 5���7 subrectangular, with subtle inclined ridges; sternite 8 conspicuously reduced, oblique; vulvae large, subcircular, situated below extremity of suture 5 / 6. Male abdomen subtriangular, with segments 3���5 fused, all segments with pronounced horizontal ridge; female abdomen much broader, all segments separated, with marked ridges. Abdominal holding mechanism with press-button on sternite 5, and lateral extensions of abdominal segment 6. P 1 subequal, robust, relatively large, strong, inner portion of dactyli with numerous denticles. P 2 ���P 4 long, flattened. P 5 conspicuously reduced. Remarks. Serrablopluma diminuta n. gen., n. sp. shows the plesiomorphic characters of Retroplumidae (de Saint Laurent 1989: 111���112), namely a sinuous anterior margin and a narrow front; a set of dorsal transverse ridges; a thoracic sternum that is much broader than long; subtrapezoidal sternites 3, 4, with pronounced lateral extensions in sternite 4 behind a distinct notch in sternite 3; distinctly wide, sub-rectangular sternites 5���7; a conspicuously reduced, oblique sternite 8; distinct sexual dimorphism for abdomen; P 2 ���P 4 long, flattened; P 5 conspicuously reduced, sub-dorsal. The thoracic sternum is typical of Retroplumidae (see Guinot & Quenette 2005: fig. 29 A). Genera that resemble Serrablopluma n. gen. are Loerentheya (see Glaessner 1969: fig. 338, 5) and Loerenthopluma, both of which have a sub-trapezoidal carapace, with the maximum width anteriorly and a wide anterior margin. Serrablopluma n. gen., however, shows distinct characters such as the anterior carapace ridge that intersects the anterior margin, a markedly long lateral spine and lateral margins that are uniformly straight. The extinct genera Retropluma and Retrocypoda differ (see Beschin et al. 1996: figs 4, 5) in having a subrectangular carapace outline, a much narrower anterior margin, lateral margins broadly arched, not diverging posteriorly and an anterior dorsal carapace ridge without lateral spines. The carapace outline is sub-trapezoidal in Serrablopluma n. gen., the anterior margin conspicuously wide, clearly broader than the posterior margin;moreover, the anterior margins terminate in a long lateral spine intersecting the first anterior dorsal ridge, while lateral margins are straight and continuous and converge posteriorly. Species of Costacopluma and Archaeopus, which were included in the Retroplumidae by De Grave et al. (2009: 41), exhibit much more inflated carapaces, with markedly convex dorsal surfaces. The dorsal surface in Archaeopus has in addition swollen regions (Rathbun 1908: pl. 47, fig. 5), rather than the thin ridges that define Serrablopluma n. gen., 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 pages 344-348, DOI: 10.11646/zootaxa.3652.3.3, http://zenodo.org/record/221749

Van Bakel, Barry W. M. (2013): Preservation of internal pleurites in a new palaeocorystid crab (Crustacea, Brachyura, Raninoidia) from the Cenomanian (Upper Cretaceous) of Poitou-Charentes, France. Zootaxa 3701 (3): 322-328, DOI: http://dx.doi.org/10.11646/zootaxa.3701.3.2

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."]}