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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).

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"]}

Jurapecten dhondtae sp. nov. urn:lsid:zoobank.org:act: 3CEF9EA0-184A-4352-87FA-CC5F76FED11A Fig. 8A–H, J–V benthopectinid sp. 1 (? spp.) – Jagt 2000: 393, pl. 6 figs 1–4. Cheiraster ? sp. – Blake & Jagt 2005: 190, pl. 5 figs 1–7. Diagnosis Jurapecten in which the marginal ossicles possess a coarsely rugose sculpture and the inferomarginals bear a single, large, laterally directed spine base. Etymology Named after Annie V. Dhondt (1942–2006), specialist of Cretaceous bivalves and close friend. Material examined The inferomarginal illustrated here (Fig. 8F) is the holotype (NHMM JJ 10490a), while the other figured ossicles in lot NHMM JJ 10490 are paratypes. Additional material includes around 50 isolated ossicles (lot NHMM JJ 9591). All material is from the upper Maastrichtian (Maastricht Formation, Emael Member, Lava Horizon) at the CBR-Romontbos Quarry, Eben Emael (Liège, northeast Belgium). Description Superomarginals (Fig. 8A–D) block-like, nearly equidimensional, elongating distally. Outer face with sculpture of dense rugosities of similar size and single, crater-rimmed spine base positioned towards abactinal-distal margin. Inferomarginals (Fig. 8E–H, J–K) asymmetrical, proximal margins broader than distal margins; inter-inferomarginal articulation facet close to actinal surface of plate (Fig. 8J), forming short projection. External face of inferomarginal with sculpture of coarse, rounded rugosities and single, large crater-rimmed spine base, positioned centrally, or slightly towards abactinal margin. Ambulacrals (Fig. 8Q–V) with elongated, bar-like ambh, broad, flat ambb, with large, flat surface for dadam and ada3, padam on short wing.Abactinal ridge and inferomarginal articulation absent (Fig. 8V). Adambulacrals (Fig. 8L–M) with concave abactinal face, actinal face with single subambulacral spine base. Enlargement of dadam (Fig. 8P) shows irregularly ridged region, similar to that seen on extant Pectinaster filholi (Perrier, 1885) (Fig. 8I). Remarks The marginal ossicles (Fig. 8A–H, J–K) are closely similar to those of Pontaster tenuispinus (von Düben & Koren, 1846) (compare with Fig. 4J–M, O–Q) in shape, sculpture and spine base development, but the ambulacrals (Fig. 8Q–V) possess elongated heads and lack abactinal ridges and inferomarginal articulation structures, and compare better with those of Jurapecten hessi and J. infrajurensis sp. nov. (see Figs 6J–M, 7M–O). The partially preserved specimen (NHMM MD 4105), described and illustrated by Blake & Jagt (2005), may be conspecific, but recrystallisation of all ossicle types precludes detailed comparison of the sculpture of infero- and superomarginal ossicles and ambulacral ossicles are too poorly preserved. The type material of J. dhondtae sp. nov., from the middle Emael Member, is ca 200 000 years younger than NHMM MD 4105, from the basal Gronsveld Member (compare Keutgen 2018).

Alkaidia sumralli Blake & Reid, 1998 Figs 16A–C, 17B–E, H, K Alkaidia sumralli Blake & Reid, 1998: 529, fig. 9/1–14. Alkaidia sumralli – Ewin & Gale 2020: 13, figs 10.1–10.2, 13.3, 13.8, 13.11–13.12. Diagnosis Alkaidia in which the primary radial ossicles are elongated and the terminal ossicle is not deeply notched on its proximal margin. Material examined The holotype (Texas Memorial Museum, number 1786 TX1) is from the Grayson Formation (lower Cenomanian) at the Waco shale pit (Waco, Mclennan County, Texas, USA).Additional material comprises a magnificent individual (NHMUK PI EE 15225), collected by Frank Holterhoff from the Grayson Formation (lower Cenomanian) of Dottie Lynn, Fort Worth, Texas and illustrated here (Fig. 16A–C), as well as numerous dissociated ossicles from the same locality (NHMUK PI EE 18005–18007, 18009). Remarks The affinities of A. sumralli have recently been discussed in some detail by Ewin & Gale (2020) and the evidence for its inclusion in the Forcipulatida (Zorocallina) and the family Terminasteridae can be summarised briefly as follows: the presence of abundant, straight ‘duck-billed’ forcipulate pedicellariae is a characteristic of the Zorocallina (Fig. 16D); the construction of the abactinal surface is closely similar to that of zoroasterids and terminasterids, which also have large, Y-shaped first superomarginals and a row of robust, lobed, quadrangular radial ossicles which imbricate proximally and each carry a centrally placed conical spine (Ewin & Gale 2020). Additionally, the morphology of the adambulacrals and ambulacrals, and the nature of their articulation is similar in zoroasterids, Terminaster and Alkaidia (Fig. 17). Ada1a and ada2 are concave on the adamulacrals (Fig. 17A–C) and positioned on a process on the ambulacrals (Fig. 17G–H, J–K). The dadam and padam facets are subequal in size, broad and short (Fig. 17A–C). The abactinal construction is never seen in extant benthopectinids, in which the abactinal ossicles in the arms are small and parapaxilliform or very small, and never imbricate. Additionally, in benthopectinids the abactinal surface is invariably flat, and the arm section is not subcylindrical.

Henricia ? venturana Durham & Roberts, 1948 Henricia ? venturana Durham & Roberts, 1948: 437, pl. 66 figs 1, 3. Remarks The unique holotype, from the Upper Cretaceous Chico Formation of the North Fork of Matilija Creek, Ventura County (California, USA), is contained in the palaeontological collections of the University of California. Part (number 4866B) and counterpart (4866A) show the abactinal and actinal surfaces, respectively. The type specimen is an external mould of a near-complete asteroid with moderately long, narrow arms and a small disc, which has lost the abactinal ossicles and exposes the ambulacrals on the abactinal surface. Unfortunately, parts of the moulds have been worn by erosion and few details are well preserved. The ambulacral heads are elongated, and the adambulacral ossicles are clearly visible, as are probable small marginal ossicles bearing spines. The groove is very wide. Although Blake (1984) assigned the taxon to the Benthopectinidae, there does not seem to be any compelling evidence for its taxonomic affinity, and it is considered here to be an indeterminate asteroid., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 186, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Durham J. W. & Roberts W. A. 1948. Cretaceous asteroids from California. Journal of Paleontology 22: 432 - 439.","Blake D. B. 1984. The Benthopectinidae (Aseroidea: Echinodermata) of the Jurassic of Switzerland. Eclogae geologicae Helvetiae 77 (3): 631 - 647."]}

Genus Nearchaster Fisher, 1911 Nearchaster Fisher, 1911: 91. Type species Acantharchaster aciculosus Fisher, 1910, by original designation., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 160, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Fisher W. K. 1911. Asteroidea of the North Pacific and adjacent waters. Part 1. Phanerozonia and Spinulosa. Bulletin of the United States National Museum 76: 1 - 420. https: // doi. org / 10.5479 / si. 03629236.76. i"]}

Nearchaster spinosus (Blake, 1973) comb. nov. Mistia spinosa Blake, 1973: 48, pl. 16 figs 30–44, pl. 17 figs 1–21, 35–36. Brisingid (?) – Zullo et al. 1964: 334. Material examined UCMP A-5018 (holotype no. 10675) is the type and only specimen; it exposes a partly disarticulated abactinal surface showing the disc and proximal portions of four arms. Large marginal spines and smaller abactinal ones are visible. Marginals, adambulacrals, ambulacrals and abactinal ossicles of the holotype were figured individually by Blake (1973). Occurrence Keasey Formation (Lower Oligocene) near Mist, Oregon (USA). Description The abactinal surface of UCMP A-5018, embedded in matrix (Blake 1973: pl.16 fig. 44), shows part of the disc and four proximal arms. Although the outline is retained, the ossicles are jumbled and largely dissociated, such that adambulacrals and ambulacrals are visible on the actinal surface. The marginal spines, largely in place, are elongated and tapering. The abactinal spines are much smaller, perhaps one-fifth the size of those on the marginals. The adambulacrals are well preserved, subrectangular, with 2–3 large subadambulacral spine bases, and the concave inner (abactinal) surface and ridge bearing ada2 and ada3, characteristic of benthopectinids. The ambulacrals have the typical hourglass shape of benthopectinids, and asymmetry of the interambulacral muscles (P1 small, P2 large) is seen. The marginals are longer than broad with a convex, mound-like outer surface which carries 1–2 large spine bases and a number of sparsely scattered smaller ones. The inner surface of the marginals is flat. The abactinal ossicles are parapaxillae, with centrally placed, single spine bases, surrounded by a ring of smaller spines. Remarks As recognised by Blake (1973), the distinctive characters of the ambulacral, adambulacral and marginal ossicles place this form firmly in the Benthopectinidae. Comparison with extant benthopectinid species studied here indicates that Mistia spinosa shares important characters with the Recent Pacific genus Nearchaster, including the following: 1. Adambulacrals are nearly identical in shape to those of Nearchaster aciculosus, and both carry 2–3 bases for subambulacral spines. 2. Marginals are closely similar to those of N. aciculosus in both shape and distribution of spine bases. The overall form of the body, with large marginal spines, and shorter abactinal spines on the disc is broadly similar to the development in the genera Benthopecten, Nearchaster and Myonotus Fisher, 1911 (see Fisher 1911: pls 22–26). The proportionate sizes and distributions of spines in Mistia spinosa are closest to those in Benthopecten claviger Fisher, 1910, Myonotus intermedius (Fisher, 1910) and Nearchaster aciculosus (Fisher, 1910) (see Fig. 2E–F herein). The abactinal parapaxillae of Mistia spinosa are very close in structure to those of N. aciculosus, with a central spine base surrounded by a ring of smaller ones. In conclusion, Mistia spinosa is a benthopectinid which has remarkably detailed similarities of ossicle morphology to the present-day Pacific species Nearchaster aciculosus and it is therefore provisionally placed in that genus. The genus thus has a history in the Pacific Ocean of at least 33 million years., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on pages 160-161, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Blake D. B. 1973. Ossicle morphology of some Recent asteroids and description of some West American fossil asteroids. University of California Publications in Geological Sciences 104: 1 - 59.","Zullo V. A., Kaar R. F., Durham J. W. & Allison E. C. 1964. The echinoid genus Salenia in the eastern Pacific. Palaeontology 7: 331 - 349.","Fisher W. K. 1911. Asteroidea of the North Pacific and adjacent waters. Part 1. Phanerozonia and Spinulosa. Bulletin of the United States National Museum 76: 1 - 420. https: // doi. org / 10.5479 / si. 03629236.76. i"]}

Alkaidia Blake & Reid, 1998 Alkaidia Blake & Reid, 1998: 529, fig. 8. Type species Alkaidia sumralli Blake & Reid, 1998, by original designation. Diagnosis Terminasteridae with elongated rhombic superomarginals directed obliquely towards radials; inferomarginals with tall, distally swollen central spine bases and prominent groove for spine attachment; radials (except primary radial) rhombic in outline. Assigned species In addition to the type species, A. megaungula Ewin & Gale, 2020. Remarks Alkaidia ranges from the Barremian to Cenomanian and appears to be restricted to the western Tethys.

Chrispaulia wrightorum sp. nov. urn:lsid:zoobank.org:act: DC13434F-62E3-47A9-9F62-327F77141670 Fig. 13F–G ? Benthopecten sp. – Spencer & Wright in Moore 1966: U48. Diagnosis Chrispaulia with smooth marginal ossicles, in which the distal superomarginals possess a single large spine pit close to their distal, abactinolateral border. Etymology After the late brothers C.W. and E.V. Wright, who collected the specimen. Material examined The arm fragment (NHMUK PI EE 17997) from the Albian Red Chalk (Hunstanton Formation) at Speeton, Yorkshire (United Kingdom) is the holotype and single specimen known to date. The specimen was originally articulated; it was subsequently reconstructed on plasticene by the Wright brothers. Description Arm elongated, narrow; supero- and inferomarginals opposed, shortening rapidly distally (Fig. 13F–G). Superomarginals bearing single, distally directed, crater-like spine base on distal, abactinolateral margin. Supero- and inferomarginals smooth, grooves for cribriform organs between infero-/superomarginal pairs poorly defined. Remarks Chrispaulia wrightorum sp. nov. differs from all congeners in the smooth marginal ossicles, lacking rugosities., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 175, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970

Genus Jurapecten Gale, 2011 Jurapecten Gale, 2011a: 84, pl. 19. Type species Jurapecten hessi Gale, 2011, by original designation. Diagnosis Benthopectinids which possess strongly rugose marginals; rugosities conjoined by thin radiating strips of imperforate stereom; ambulacrals lack abactinal ridges and inferomarginal articulation. Assigned species In addition to the type species, J. infrajurensis sp. nov. and J. dhondtae sp. nov., both described below. Remarks Jurassic–Cretaceous benthopectinids are locally common among isolated ossicles in washed residues. All share the same distinctive sculpture type of prominent imperforate rugosities, conjoined by radiating strips of stereom (e.g., Figs 6N–O, 7G, I–J), absent on extant genera. In the extant genera Pontaster and Cheiraster, the rugosities on the superomarginals are smaller and more widely spaced (e.g., Fig. 4J–M, Q), although the inferomarginal sculpture is similar to that seen in Jurapecten (Fig. 6C–E). Jurapecten also lacks a number of characters seen in all extant taxa, including an abactinal ridge on the ambulacrals (compare Fig. 7N–O with Fig. 5I, K–L, Q), and there is no ambulacral articulation surface with the inferomarginal (compare Fig. 7N–O with Fig. 5I, K, Q). Additionally, the ambulacral heads are more elongated in Jurapecten (e.g., Figs 6J, L, 8Q–U). The absence of the abactinal ridge, to which the longitudinal arm muscles attach in all living genera (Clark 1981), is a plesiomorphic feature of Jurapecten., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 161, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Gale A. S. 2011 a. The phylogeny of post-Palaeozoic Asteroidea (Neoasteroidea, Echinodermata). Special Papers in Palaeontology 85: 1 - 112.","Clark A. M. 1981. Notes on Atlantic and other Asteroidea. 1. Family Benthopectinidae. Bulletin of the British Museum of Natural History (Zoology) 41: 91 - 135."]}

Chrispaulia spinosa sp. nov. urn:lsid:zoobank.org:act: 4C003A9F-1F91-44BC-91AB-2910B8EA756A Fig. 13D–E, H–O Diagnosis Chrispaulia in which distal inferomarginals bear an oblique row of large, bifid spine attachment sites, which carried long, flattened spines. Etymology Latin for ‘bearing spines’, in reference to the row of spine attachment sites on distal inferomarginals. Material examined A distal arm fragment (Nds LH 105.107), comprising five marginal pairs, with articulating spines preserved, is the holotype; it is from the lower Hauterivian (Endemoceras amblygonium ammonite Zone) at Engelbostel near Hannover (northern Germany). Paratypes (NHMUK PI EE 17998–18004) are 35 marginal ossicles and a single oral ossicle from the upper 3 metres of the Tealby Clay (Hauterivian) at Nettleton, Lincolnshire (United Kingdom). Description Arms elongated, narrow, tapering slowly (Fig. 13D–E); proximal superomarginals tall, block-like, bearing a single, abactinally directed, crater-like spine base. Central region of external face narrow, poorly defined, lateral surfaces broad with small rugosities for attachment of tiny cribriform spines (Fig. 13H–I, N; spines still articulated in Fig. 13D). Distal superomarginals with or without an abactinally situated, distally directed large spine base (Fig. 13I–J), rugose (Fig. 13D) or smooth (Fig. 13I–J). Distal infero- and superomarginals thin, imbricating proximally; distal inferomarginals with oblique row of large, bifid spine bases, which bore flattened, lanceolate spines (Fig. 13D). Sharply defined grooves for cribriform organs between each infero-/superomarginal pair. Oral ossicle (Fig. 13O) with broad actinal face, bearing large rugosities for attachment of sos; large elongate oradm, low, broad apophyse. Remarks Chrispaulia spinosa sp. nov. differs from its congeners in its possession of 3–4 bifid spine pits on distal inferomarginals.

Genus Chrispaulia Gale, 2005 Chrispaulia Gale, 2005: 2, fig. 4a–c. Type species Nymphaster radiatus Spencer, 1905, by original designation (see Fig. 13A–C). Diagnosis Arms long, narrow; disc small, with rounded interbrachial arcs; superomarginals meeting over radius along length of arm; distal marginals imbricate (emended from Gale 2005). Assigned species In addition to the type species, Chrispaulia jurassica Gale, 2011 (Gale 2011a), C. wrightorum sp. nov. and C. spinosa sp. nov. Remarks The record in the Treatise of Invertebrate Paleontology (Part U; Spencer & Wright 1966) of a benthopectinid from the “Albian of England ” is based on an arm fragment from the Albian Red Chalk of Yorkshire, United Kingdom (C.W. Wright, pers. comm. to ASG, 1978). We have examined this specimen (NHMUK PI EE 17997), which comprises five marginal ossicles reconstructed on plasticene. The shape of the ossicles indicates that this specimen belongs to the genus Chrispaulia; it is here described as a new species, C. wrightorum sp. nov. (see below). In addition, we record another species from the Hauterivian (Lower Cretaceous) of northeast England and northern Germany., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on pages 174-175, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Gale A. S. 2005. Chrispaulia, a new genus of mud star (Asteroidea, Goniopectinidae) from the Cretaceous of England. Geological Journal 40: 383 - 397. https: // doi. org / 10.1002 / gj. 1019","Gale A. S. 2011 a. The phylogeny of post-Palaeozoic Asteroidea (Neoasteroidea, Echinodermata). Special Papers in Palaeontology 85: 1 - 112.","Spencer W. K. & Wright C. W. 1966. Asterozoans. In: Moore R. C. (ed.) Treatise on Invertebrate Paleontology, Part U, Echinodermata 3: U 4 - U 107. The Geological Society of America and The University of Kansas Press, Boulder, CO and Lawrence, KS."]}

Family Terminasteridae Gale, 2011 Terminasteridae Gale, 2011a: 92, fig. 8. Diagnosis Zorocallinids with extra-axial arm constructed of seven rows of ossicles organised with one row of radials, two rows of adradials, two rows of superomarginals and two rows of infromarginals; both marginal rows extend to arm tip. Assigned genera Alkaidia Blake & Reid, 1998 and Terminaster Hess, 1974. Remarks The family Terminasteridae is sister group to the Eocene–Recent Zoroasteridae Sladen, 1889, which is widespread in bathyal to abyssal depths of the present-day oceans., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on pages 182-183, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Gale A. S. 2011 a. The phylogeny of post-Palaeozoic Asteroidea (Neoasteroidea, Echinodermata). Special Papers in Palaeontology 85: 1 - 112.","Blake D. B. & Reid R. III. 1998. Some Albian (Cretaceous) asteroids (Echinodermata) from Texas and their paleobiological implications. Journal of Paleontology 72 (3): 512 - 532. https: // doi. org / 10.1017 / S 002233600002429 X"]}

Benthopectinidae Verrill, 1894 Fig. 6A–B, F ? Benthopectinidae Verrill, 1894: 217. Material Seventeen marginal ossicles; the inferomarginal figured (Fig. 6A) is MHI 2183/7; the other inferomarginal and oral ossicle illustrated here are MHI 2183/8 and 9. All are from the Maantang Formation (Carnian, Upper Triassic; sample C30) at Jiancougou, Sechuan Province, China. Description The inferomarginal and oral ossicles are strongly reminiscent in shape and external sculpture of those of benthopectinids; compare Fig. 6A–B with benthopectinid marginals (Fig. 6C–E), and the oral (Fig. 6F) with those of benthopectinids (Fig. 6G–I). The inferomarginals share a number of similarities to those of benthopectinids (Fig. 6A–E), notably the coarse, longitudinally arranged rows of rugosities, the presence of a large spine (or pedicellaria) base close to the abactinal margin of the plate, and the marked asymmetry of the distal inferomarginals, in which the distal height is less than the proximal one. The oral ossicles (Fig. 6F) are remarkably similar to those of benthopectinids in the evenly convex actinal margin, the distally angled apophyse and the shape of the adambulacral articulation (Fig. 6G–I). However, the material available to date is too limited, and in the absence of ambulacrals and adambulacrals, this referral is very tentative., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 174, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Verrill A. E. 1894. Descriptions of new species of starfishes and ophiurans, with a revision of certain species formerly described; mostly from the collections made by the United States Commission of Fish and Fisheries. Proceedings of the United States National Museum 17 (1000): 245 - 297. https: // doi. org / 10.5479 / si. 00963801.1000.245"]}

Genus Punkaster gen. nov. urn:lsid:zoobank.org:act: D63466E2-7CA4-4896-BFD2-4D156A280AA8 Type species Punkaster spinifera gen. et sp. nov. Diagnosis Highly derived form in which the marginal ossicles are very elongated, proximal infero-and superomarginal pairs occasionally fused, and marginals may possess 1–2 large rounded bases for attachment of conical spines. Adambulacrals with deep rounded notch to allow extension of tube feet and very large furrow spines. Etymology From the similarity of the marginal spination to the 1980s punk hairdos. Assigned species In addition to the type species, P. ruegenensis gen. et sp. nov. (see below). Remarks The highly unusual marginal ossicles of this new genus (Fig. 9E, I–J, M, P) have been known for over 60 years, but they remained undescribed and it has hitherto not been possible to assign them to any family. The new material includes ambulacral and adambulacral ossicles, which demonstrate a likely affinity with benthopectinids in the elongated, bar-like ambulacral heads and, especially, in the nature of the ambulacral/adambulacral articulation. This is typically benthopectinid, in the abradial position of ada3, the position of padam on a short wing-like process and the presence of ada2 on a steep ridge. The abactinal ossicles are closely similar to parapaxillae of modern benthopectinids. The presence of an abactinal ridge on the ambulacrals of P. spinifera gen. et sp. nov. may indicate that the species possessed longitudinal muscles in the arms. The genus perhaps represents a highly specialised offshoot from the mainline benthopectinids., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 168, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970

Plesiastropecten hallovensis Peyer, 1944 Fig. 14 Plesiastropecten hallovensis Peyer, 1944: 320, pls 7–8. Plesiastropecten hallovensis – Blake 1984: 633, fig. 1a–h. Diagnosis As for genus. Material examined The holotype (MZA L13a) is a complete asteroid in a claystone matrix, comprising part and counterpart, from the Lower Jurassic (Hettangian, liasicus ammonite Zone) of Hallau, Canton Schaffhausen, Switzerland (Peyer1944). It is contained in the collections of the Museum zu Aller Heiligen, Schaffhausen, and has suffered from considerable over-preparation. Additional material includes a suite of specimens from the same locality and horizon, most notably, specimen MZA L 13b/32a-e, which is a partially dissociated individual of which one arm is well preserved (Blake 1984: fig. 1a–g). This specimen is coated with a glue-like substance which obscures much detail and requires specialist preparation. The present description is based on these two individuals. Description The overall form is well shown by the type specimen (Fig. 14A); the arms are moderately long and tapering, the interbrachial angles acute (R:r = approximately 6:1). The arms bear an even fringe of closely spaced marginal spines. The marginals are numerous, narrow and short with a three-tiered structure, resembling tiny ‘cottage loaves’ of bread (Fig. 14F). They are clearly visible in MZA L 13b/32a-e, and are robustly paxilliform, bearing a single large spine pit centrally. The abactinal ossicles are conspicuous and relatively large, convex to flat and carry four to six, lobe-like lateral projections. Each abactinal has a single, large and centrally placed crater-like spine pit (Fig. 14B, D). The ossicles imbricate, and the lobes are notched on their inside (actinal) surfaces for contact with adjacent ossicles. These ossicles are more or less radially symmetrical in the disc, but elongated in the arm, where the abactinal ossicles are arranged in three rows comprising radials and two adradials (Fig. 14B). In the distal arm, the large radial and adradials are separated from adjacent ossicles of the same row by smaller inset ossicles. The adambulacrals were prepared in a small part of specimen MZA L 13b/32b. They are very broad and short (3:1), and carry 5–6 large, horseshoe-shaped spine bases arranged in a single transverse row (Fig. 14E). The adambulacrals of opposing rows are slightly angled (150°) in a proximal direction. The ambulacrals are only seen in abactinal view, and the ambh forms a conspicuous, elongated triangular proximal wing which overlaps the more proximal adjacent ambulacral (Fig. 14D). The ambb is oval and symmetrical. The marginal spines are conspicuous, forming an even comb-like fringe to the starfish. Each marginal plate carries a single tapering spine with a unique construction. The cross section is a shallow U-shape, and the abactinal surface bears a groove. The rounded actinal surface is made up of 4–6, length-parallel coalescing rods of trabecular stereom. The lateral margins of the spines carry outwardly directed short barbs, probably lost on the holotype through over-preparation. The base of the spine is swollen and rounded, and a simple socket on the base articulates with a boss on the marginal. The abactinal spines are shorter, and round in cross section; these are also composed of elongated trabeculae. The 5–6 adambulacral spines articulate with horseshoe-shaped bases, and are long and gently curved. Remarks The robust paxilliform construction of the marginals is not found in any benthopectinid asteroid, but is characteristic of modern solasterids and the Jurassic genus Plumaster, and each marginal carries only a single spine. We cannot agree with Blake (1984) that these resemble marginals of benthopectinids, other than in that they carry large spines. The large, oval or rounded, imbricating abactinal ossicles which are alternately large and spine-bearing and small and spineless in the arm are quite different from the parapaxillae of benthopectinids. The construction of the marginal and abactinal spines, with elongated trabeculae running along the length of the spines and a semicircular, concavo-convex cross section bearing two rows of lateral thorns, are dissimilar to benthopectinid spines, which are conical, cylindrical and carry numerous, irregular, distally directed thorns. The concavo-convex construction is otherwise seen only in the multi-armed Early Jurassic Plumaster ophiuroides (Wright, 1863). The transversely broad, short adambulacrals, carrying 5–6 large curved adambulacral spines, are quite unlike adambulacrals of benthopectinids which are narrow and rather elongated, but are similar to those of Plumaster (Gale 2011b: figs 13–14). The elongated, imbricating proximal flanges of the ambulacral heads are not found in any benthopectinid, where the ambulacral heads are short, upright and do not imbricate, as in all paxillosidans (Gale 2011a), but are similar to those of Plumaster (Gale 2011b: fig. 14b–c). Other than a superficial similarity in shape and the presence of elongated marginal and abactinal spines (also found in other asteroids), Plesiastropecten hallovensis does not show any of the characteristics of the Benthopectinidae, but bears a close similarity to the Pliensbachian–Aalenian multiarmed genus Plumaster, with which it shares broad, short adambulacrals with 5–6 large transversely arranged, hyaline spines and the flanged, imbricating abactinal ossicles (Gale 2011b)., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on pages 178-179, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Peyer B. 1944. Beitrage zur Kenntnis von Rhat und Lias. Eclogae geologicae Helvetiae 36: 303 - 326.","Blake D. B. 1984. The Benthopectinidae (Aseroidea: Echinodermata) of the Jurassic of Switzerland. Eclogae geologicae Helvetiae 77 (3): 631 - 647.","Gale A. S. 2011 b. Asteroidea (Echinodermata) from the Oxfordian (Late Jurassic) of Savigna, Department [sic] du Jura, France. In: Meyer C. A. & Costeur L. (eds) Special Issue: Echinoderms - from the early","Gale A. S. 2011 a. The phylogeny of post-Palaeozoic Asteroidea (Neoasteroidea, Echinodermata). Special Papers in Palaeontology 85: 1 - 112."]}

Plesiastropecten Peyer, 1944 Plesiastropecten Peyer, 1944: 320, pls 7–8. Type species Plesiastropecten hallovensis Peyer, 1944, by monotypy. Diagnosis Five-rayed plumasterid, in which the abactinal ossicles are stellate and which bears a fringe of elongated marginal spines., Published as part of Gale, Andy S. & Jagt, John W. M., 2021, The fossil record of the family Benthopectinidae (Echinodermata, Asteroidea), a reappraisal, pp. 149-190 in European Journal of Taxonomy 755 on page 178, DOI: 10.5852/ejt.2021.755.1405, http://zenodo.org/record/5032970, {"references":["Peyer B. 1944. Beitrage zur Kenntnis von Rhat und Lias. Eclogae geologicae Helvetiae 36: 303 - 326."]}