taxonID	type	description	language	source
5B0C164C663C882441B9D788D840F9CF.taxon	description	(Figs 1 D, E, 2 A – E, 3 – 9)	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C663C882441B9D788D840F9CF.taxon	diagnosis	Diagnosis: Frontal furrow deepest on oral side of test. Peristome at posterior end of deep furrow, facing forwards, not visible from below. Labral plate small, asymmetrical, separated from sternal plate by two, three, or even four asymmetrical pairs of plates (Fig. 1 D, E). No rostral plate. All gonopore-bearing plates in contact. Marginal fasciole absent.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C663C882441B9D788D840F9CF.taxon	discussion	Remarks: The family Pourtalesiidae is restricted to include only the genera Pourtalesia and Cystocrepis. These genera are characterized by the plastron plating of the type VI: a small asymmetrical labral plate is separated from the sternal plate by two to four asymmetrical pairs of plates (two or three pairs of ambulacrals and, sometimes, one pair of interambulacrals); the intercalating ambulacral plates lack intra-plate lines; the labral plate is often not visible. This morphological synapomorphy of pourtalesiids (type VI plastron plating) is well-supported by the molecular data and clearly differentiates them from the representatives of the families Ceratophysidae and Galeasteridae. Apart from the plastron plating, the other diagnostic features of the family Pourtalesiiadae, listed in the revised diagnosis, do not depart from those previously used. The characters of the apical system are in accordance with the diagnosis of Smith and Kroh (2011), with some simplifications. The characters of the frontal furrow, peristome, and fascioles follow those listed by Smith (2004) for the family Pourtalesiidae as represented by the genus Pourtalesia. The genus Pourtalesia differs from the monotypic Cystocrepis in having a supramarginal periproct, subanal rostrum, subanal fasciole, and longer valves of the ophicephalous pedicellariae. Among 12 Pourtalesia species described to date, P. aurorae Koehler 1926 is morphologically closest to Cystocrepis. In this species, the subanal rostrum is very small, and the periproct is more marginal than supramarginal. The species here referred to as Pourtalesia (?) sp. A has an even smaller subanal rostrum that takes the inframarginal position below the marginal periproct (Fig. 4 C, D). This set of features makes the species transitional in morphology between the two genera. Pourtalesia (?) sp. A thus might represent a new genus of pourtalesiids. Molecular data provide evidence that Pourtalesia (?) sp. A represents a separate phylogenetic lineage that is sister to the group comprised of Cystocrepis setigera and all other representatives of Pourtalesia (Fig. 3). However, the statistical support of these groupings is weak in one of the two analyses (namely, the ML). Apart from the unusual test shape, no significant differences (e. g. in external test appendages) between Pourtalesia (?) sp. A and the species of the genus Pourtalesia are found at the moment. Additional morphological and molecular data are needed to determine whether Pourtalesia (?) sp. A deserves establishing a new genus to accommodate the species. This problem will be addressed in the future. Hereinafter, this putative new species will be provisionally referred to the genus Pourtalesia. Pourtalesia in its current state is the most species-rich among the genera of Urechinina. Apart from the 12 species described to date (Mironov 1995 b), representatives of the five more yet undescribed species are available at the authors’ disposal.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C663B8826419AD3D7D96CF926.taxon	description	(Figs 1 F – H, 2 F – I, 3, 5 – 7, 8, 10) ZooBank LSID: urn: lsid: zoobank. org: act: FFBDFC 0 D- 66 AF- 45 E 4 - 8593 - D 3 EA 5 FBCD 941.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C663B8826419AD3D7D96CF926.taxon	diagnosis	Diagnosis: Frontal furrow deepest on oral side of test. Peristome at posterior end of deep furrow, facing forwards or slightly upwards, not visible from below. Labral plate large, symmetrical, separated from sternal plate by one or two symmetrical pairs of plates (Fig. 1 F – H). No rostral plate. All gonopore-bearing plates in contact. Marginal fasciole absent.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C663B8826419AD3D7D96CF926.taxon	discussion	Remarks: The establishment of the family Ceratophysidae is well supported by both molecular and morphological data (Fig. 3). Representatives of the family demonstrate peculiar arrangement of plastron plating: well-developed labrum is always separated from the sternum by one or two symmetrical pairs of elongated ambulacral plates. This feature allows ceratophysids to be unambiguously distinguished from both pourtalesiids and galeasterids. Tuberculation of the plastronal area is different in Ceratophysidae and Pourtalesiidae. The intercalating ambulacral plates of the ceratophysids are densely covered with tubercles along the ventral midline. Dense tuberculation then stretches posteriorly onto the sternal plate and episternals. Intercalating ambulacral plates of the pourtalesiids lack dense tuberculation, which is observed along the ventral midline only on the sternal plate and episternals. Furthermore, the family Ceratophysidae is characterized by the prevalence of tridentate pedicellariae over other types, whereas in most pourtalesiid species, rostrate pedicellariae are the most common. Based on morphological features, Mironov (2008) previously distinguished two groups among genera now belonging to the family Ceratophysidae. Each group is characterized by a unique type of plastron plating. In the first group (Ceratophysa, Echinocrepis, Echinogutta, Echinosigra, and Helgocystis) only one pair of ambulacral plates separates the labrum from the sternum (plastron plating type IV; Fig. 1 F, G). The intercalating ambulacral plates carry long intra-plate lines. These lines extend meridionally and resemble the sutures between adjacent plates, but typically do not reach the plate border at one or both ends (Fig. 1 F, G). According to Mironov (2008), these structures might either be original elements related to the strengthening of the plates, or traces indicating incomplete plate coalescence. The anterior part of the test in the genera of the first group usually forms a so-called ‘ neck’ (Fig. 2 G). The ‘ neck’ is indistinguishable only in Ceratophysa and Echinocrepis, characterized by the broadest tests. Miliary spines are curved in their distal part in this group (with the exception of Helgocystis carinata). In the second group (Solenocystis, Spatagocystis and, probably, Rictocystis), the labrum is separated from the sternum by two pairs of ambulacral plates (plastron plating type V; Fig. 1 H). The latter lack long intra-plate lines. The ‘ neck’ is absent in this group. Since Rictocystis is known only by test fragments, the type V plastron plating is inferred for this genus based on morphological similarity of its spines and pedicellariae with those of Solenocystis and Spatagocystis. Miliary spines in all three genera of this group are curved in the median part. Molecular phylogenetic reconstructions reveal two major sub-clades within Ceratophysidae (Fig. 3); however, the genera characterized by a differentiated ‘ neck’ and plastron, disrupted by one pair of ambulacral plates, are found within both of them. Each of these sub-clades includes a genus, characterized by the miliary spines curved in the median part (either Solenocystis or Helgocystis). No morphological features distinguishing these sub-clades from each other were found. However, only one genus with two pairs of intercalating ambulacral plates (Solenocystis) was included in the analysis. Molecular data on two other genera, Spatagocystis and Rictocystis, are needed to test the monophyletic status of this morphological grouping. The rank of the subgenus Echinosigra (Echinogutta) is raised to a genus rank. Phylogenetic reconstructions show that the genera Echinosigra and Echinogutta belong to different, very distant lineages within the family Ceratophysidae (Fig. 3). Although Echinosigra was represented in the analysis only by the type species Echinosigra phiale, we propose that the three other species with a long ‘ neck’ and two gonopores (diagnostic features of the nominal subgenus) should remain within the genus. These species are Echinosigra mortenseni, Echinosigra porrecta, and Echinosigra vityazi. The genus Echinogutta thus should accommodate the species with the short ‘ neck’ and four gonopores (Echinogutta amphora, Echinogutta antarctica, Echinogutta fabrefacta, Echinogutta valvaedentata, and, provisionally, Echinogutta sp. A). In the phylogenetic reconstructions presented here, the putative new species Echinogutta sp. A and Echinogutta amphora belong to the same lineage, but do not group together (Fig. 3). This indicates the possibility that Echinogutta is paraphyletic. Further reconstructions with broader taxon sampling are needed to clarify the phylogeny and taxonomic composition of Echinosigra and Echinogutta. The only species of the genus Rodocystis, R. rosea, was initially described as a Pourtalesia species (Agassiz 1879, 1881). The majority of subsequent studies attributed it to the genus Cerathophysa based on the presence of the prominent subanal rostrum and umbrella-shaped ophicephalous pedicellaria (Clark 1917, 1925, Mortensen 1950, McCauley and Carey 1967, Smith and Kroh 2011, Kroh and Mooi 2023 b). The subgenus Pourtalesia (Rodocystis) was erected by Lambert and Thiéry in 1924 to accommodate this species. Mironov (1975, 1978 a) promoted the subgenus to a genus level and noted substantial differences in apical disc structure and fascioles between Rodocystis and pourtalesiids. The monotypic genus Rodocystis was recently transferred to the family Calymnidae based on the presence of the marginal fasciole (Mironov et al. 2015). The presence of the marginal fasciole, passing over plates 5. a. 4 / 5. b. 5 beneath the periproct, is the main diagnostic feature of the family Calymnidae (Smith 2004). Among all recent Urechinina, the marginal fasciole is recorded only in the genera Calymne, Rodocystis, and Sternopatagus. Molecular evidence supports the grouping of genera Rodocystis and Sternopatagus (Fig. 3). However, molecular data on Calymne are needed to confirm that these genera should be attributed to the family Calymnidae with certainty.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
5B0C164C6639882741ADD3EBDC05F9F0.taxon	description	(Fig. 1 A) ZooBank LSID: urn: lsid: zoobank. org: act: 557 DEC 84 - 05 B 3 - 4 C 7 C-B 5 B 1 - 81091 BE 59436. Diagnosis: Frontal furrow deepest at ambitus. Peristome at posterior end of furrow, facing slightly downwards, visible from below. Labral plate not separated from sternal plate. No rostral plate. Posterior gonopore-bearing plates disjunct from anterior gonopore-bearing plate by pair of intercalating plates; or all gonopore-bearing plates in contact. Marginal fasciole absent. Genera included: Galeaster Seunes 1889, Upper Cretaceous (Maastrichtian) to Palaeocene, from Europe to Kazakhstan. Remarks: The fossil genus Galeaster has been attributed by Poslavskaya and Moskvin (1960) to the family Pourtalesiidae. This status was widely accepted in later studies (Poslavskaya and Solovjev 1964, Solovjev 1994, 2005, Saucède et al. 2004, Smith 2004, Kroh and Mooi 2023 a). The morphological differences discerning Galeasteridae from both Pourtalesiidae and Ceratophysidae are more substantial than the differences between the two latter families. Among these three families, Galeasteridae are unique in the orientation of the peristome, the shape of the frontal groove, and in plastron plating, while Ceratophysidae differ from Pourtalesiidae in plastron plating only. In the family Galeasteridae, two morphological features show much greater variation than in both Pourtalesiidae and Ceratophysidae. The opening of the posterior gonopores is extremely variable in Galeaster. In the species Galeaster carinatus Ravn 1927 (Danian of Crimea, Mangyshlak, Ustyurt, and Denmark) and Galeaster dagestanensis Poslavskaya and Moskvin 1960 (Upper Danian — Montian of Crimea, Dagestan, and Mangyshlak), posterior gonopores can open either on the genital plates 1 and 4 (typical condition for Galeaster) or on the anterior paired ocular plates (Poslavskaya and Moskvin 1960: fig. 9 d; Solovjev 1994: fig. 4 d). Although in most Galeaster species the labral and sternal plates remain in full contact, in G. carinatus they are almost separated by the ambulacral plates Ia. 2 and Vb. 2 wedging in (Poslavskaya and Moskvin 1960). Smith and Jeffery (2000) indicated that the differences in plastron plating might represent growth variability and proposed to synonymize all Galeaster species under the type species Galeaster bertrandi Seunes 1889. Further detailed morphological comparisons are needed to confirm this synonymy. The preceding stage of plastron disjunction recorded in G. carinatus Ravn 1927 and a migration of the posterior gonopores onto the anterior paired oculars in G. carinatus and G. dagestanensis Poslavskaya and Moskvin 1960 result in their morphological similarity to the Pourtalesiidae and Ceratophysidae. In the two latter families, posterior gonopore-bearing plates are pierced by ocular pores and adjacent posterolaterally to the anterior genital plate (s). Saucède et al. (2004) and Smith (2004) interpret the apical system of pourtalesiids and ceratophysids as a result of anterior migration of the posterior gonopores.	en	Minin, Kirill V., Mironov, Alexandr N., Petrov, Nikolay B., Vladychenskaya, Irina P. (2024): Evolutionary and biogeographic patterns in the deep-sea echinoid families Pourtalesiidae Agassiz 1881 and Ceratophysidae fam. nov. (Echinoidea). Zoological Journal of the Linnean Society 202 (4): 1-30, DOI: 10.1093/zoolinnean/zlae034, URL: https://doi.org/10.1093/zoolinnean/zlae034
