identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
2F1AC921FFC6FFC5F2F0FA3FFCB65BD3.text	2F1AC921FFC6FFC5F2F0FA3FFCB65BD3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Holopodidae Zittel 1879	<div><p>Family  Holopodidae Zittel, 1879</p><p>Remarks. Sieverts-Doreck (1951) believed that some ossicles and deformed  Eugeniacrinus cups figured by Remeš (1902: pl. 19, figs. 11 a–c, 12–15) from the Štramberk Tithonian could belong to an Upper Jurassic species of  Cyathidium . However, the figure is too imprecise to confirm an attribution to  Holopodidae . The mere inclusion of  Cyathidium depressum in a long list of cyrtocrinids cited from Lower Cretaceous Štramberk-type limestones (Salamon &amp; Gorzelak 2010: 871) cannot be taken into account without a critical analysis of the material concerned. Moreover, the attribution to  C. senessei (Valette in Lambert &amp; Valette, 1934) of a very poorly preserved axillary from the Santonian of Poland by Salamon and Gorzelak (2011: 311, fig. 2C, D) is highly questionable. Here, we consider that the appearance of  Holopodidae prior to the Cenomanian has not been demonstrated.</p><p>It appears that the H/Wd ratio of IBrax is the best quantitative character to discriminate between the two genera in the family (&gt; 0.7 in  Holopus, &lt;0.7 in  Cyathidium) (Fig. 7).</p><p>Stratigraphic distribution. Upper Cenomanian–Recent.</p></div>	https://treatment.plazi.org/id/2F1AC921FFC6FFC5F2F0FA3FFCB65BD3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFC1FFC2F2F0FF7CFCB6599D.text	2F1AC921FFC1FFC2F2F0FF7CFCB6599D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cyathidium Steenstrup 1847	<div><p>Genus  Cyathidium Steenstrup in Michaelis &amp; Scherk, 1847</p><p>Type-species.  Cyathidium holopus Steenstrup in Michaelis &amp; Scherk, 1847, by monotypy.</p><p>Synonymy.  Cyathidium (Steenstrup 1847) in Michaelis &amp; Scherk, 1847: 119, 150;  Micropocrinus Michelin, 1851: 93; Sieverts, 1931:173;  Pseudocupressocrinus Valette in Lambert &amp; Valette, 1934: 59;  Cyathidium Sieverts-Doreck, 1938: 29;  Micropocrinus Biese &amp; Sieverts-Doreck, 1939: 117;  Cyathidium Rasmussen, 1961: 238–239; 1978: T839.</p><p>Remarks. Žítt et al. (2014) thoroughly described the wide range of shape variation in a lower Turonian population of 136 aboral cups attributed to  Cyathidium aff. depressum . The ratio H/D varies from 0.2 to c. 1.0 and usually increases during growth (Fig. 7). The cup curvature is also greatly variable with H/h from 1 to 2.7. Details of characters discriminating the extant species such as interradial morphology and fulcral ridge arrangement (Améziane et al. 1999) are only preserved in a few specimens, and only parts of the external surface show growth lines and peculiar ornamentation. Therefore, attribution to a new species or to previously described species merely based on aboral cups requires well-preserved specimens and discriminating qualitative characters. Intact fossil specimens with a retracted crown are known exclusively in  C. holopus (Baumiller &amp; Gaździcki 1996; Donovan &amp; Jakobsen 2004; Hess &amp; Messing 2011; Nielsen 1913; Rasmussen 1961). The vault covering the distal arm series is composed solely of IIBr1, their height being substantially greater than their width. IIBr have not yet been described from the large species  Cyathidium vlieksi Jagt, 1986, but are known (Jagt pers. comm.). This species has an aboral cup seemingly very similar to some specimens of  C. holopus . Intraspecific variation of the aboral cup in  C. holopus and  C. vlieksi show two extreme morphotypes: one very depressed, like in  C. depressum, and another as high or taller than wide with conspicuous growth lines. The lower Santonian species,  C. senessei, differs from all other known congeners in having a stout and tumid curved IBrax, with relatively small distal articular facets facing inwards and subperpendicular to the upper plan of the IBr circlet (Améziane et al. 1999; Heinzeller et al. 1996; Rasmussen 1961). The central space provided by the IBrax circlet (Rasmussen 1961, pl. 34 fig. 7c) probably houses IIBr1 keystone plates, but these have not yet been described. Žítt et al. (2014) figured the main discriminating characters of dissociated brachials of  C. aff. depressum from the lower Turonian. Unfortunately, IIBr were not available in the original description of  C. depressum from the upper Cenomanian but IBr1 were illustrated (Rasmussen 1961). The presence of granulation on the aboral cup appears highly variable among fossil assemblages and cannot be used as a specific discriminating character except when there are sufficient ossicles to demonstrate the stability of this trait. Another early Campanian form from southern England and north-west France will be published soon (Gale &amp; Jagt, in press).</p><p>Included fossil species and occurrence.  Cyathidium chiampoensis sp. nov. (lower Lutetian of Chiampo valley, north-eastern Italy),  C. depressum including  C. aff. depressum first described by Žítt 1996 (upper Cenomanian–lower Turonian, Germany and Bohemia),  C. gastaldii (Middle Miocene near Turin, northwestern Italy),  C. holopus (upper Maastrichtian of Netherlands, Danian of Denmark and Sweden, lower Eocene of Seymour Island, Antarctica),  C. senessei (lower Santonian of Bugarach, southern France),  C. vlieksi (Maastrichtian of Belgium, Netherlands), C. sp. nov. (Gale &amp; Jagt, in press; lower Campanian, north-western France and southern England).</p><p>Stratigraphic distribution. Upper Cenomanian–Recent.</p></div>	https://treatment.plazi.org/id/2F1AC921FFC1FFC2F2F0FF7CFCB6599D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFC1FFC3F2F0F9CAFC495E81.text	2F1AC921FFC1FFC3F2F0F9CAFC495E81.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cyathidium chiampoensis Martinez-Soares & Roux & Giusberti & Gatto & Eléaume & Améziane 2024	<div><p>Cyathidium chiampoensis sp. nov.</p><p>Figures 8A–C; Table 4</p><p>Synonymy.  Cyathidium sp. Frisone et al., 2020: 306.</p><p>Type material.  Two aboral cups including the holotype (MGP-PD 33229, Fig. 8 A – B), one well-preserved paratype (MGP-PD 33230, Fig. 8C) .</p><p>Etymology. From Chiampo valley, Vicenza province, north-eastern Italy.</p><p>Diagnosis. Small species (D&lt;3 mm), aboral cup shape more or less depressed (H/D 0.42 to 1.03), conspicuous growth lines interrupting numerous perpendicular ridges, subcircular adoral face with coarse, short interradial processes. Brachials unknown.</p><p>Type stratum. Lower Lutetian tuffite horizon.</p><p>Type locality.  Cengio dell’Orbo quarry (Cava Boschetto) in the Chiampo Valley, north-eastern Italy.</p><p>Description of type series. Aboral cups belonging to small specimens with cup diameter up to 2.72 mm, H equal or substantially lower than D (H/D 0.42), usually D greater than Db. Quantitative characters of the type series listed in Table 4. The holotype, which is the largest specimen, has a curved aboral cup and an external surface showing 8–9 conspicuous parallel growth lines interrupting numerous perpendicular ridges; base smaller than D straight on one side (Fig. 8A) and displaying a coarse groove on the other side probably corresponding to the attachment on an irregular substrate (Fig. 8B); subcircular adoral face without asymmetry despite the marked cup curvature, external border wide and rounded having coarse outer interradial processes; wide central calycinal cavity. Main characters similar in paratype 1, except cup base more regular than in holotype.</p><p>Remarks. The ornamentation consisting of regular growth bands and perpendicular ridges is unknown in congeners.  Cyathidium depressum and  C. aff. depressum both differ in usually having granulation and more irregular growth bands (Sieverts 1931; Žítt 1996; Žítt et al. 2014).  Cyathidium holopus differs in having variable growth banding either with relatively wide bands (Rasmussen, 1961: pl. 35, fig. 6) or only numerous very serrated lines (Donovan &amp; Jakobsen, 2004: fig. 2 f, h), granulation (Donovan &amp; Jakobsen, 2004: fig. 2 d) or an almost smooth external surface in the specimen from Seymour Island (Baumiller &amp; Gaździcki 1996). The two latter species and  C. vlieksi (Jagt, 1986: fig. 1) do not show ridges perpendicular to growth lines.</p><p>Occurrence. Chiampo Valley (north-eastern Italy), lower Lutetian.</p></div>	https://treatment.plazi.org/id/2F1AC921FFC1FFC3F2F0F9CAFC495E81	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFC3FFC0F2F0FCAAFB345BA2.text	2F1AC921FFC3FFC0F2F0FCAAFB345BA2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cyathidium gastaldii (Michelin 1851)	<div><p>Cyathidium gastaldii (Michelin, 1851)</p><p>Figures 8F, G; Table 6</p><p>Synonymy.  Micropocrinus gastaldii Michelin, 1851: 93–94; Michelotti, 1861: 354; Jaekel, 1891: 621; Sieverts, 1931: 173; Biese &amp; Sieverts-Doreck, 1939: 11;  Cyathidium gastaldii Manni, 2005: 214, pl. 1 fig. 8, text fig. 4</p><p>Non  Cyathidium gastaldii Manni &amp; Pacioni, 2021: 68–69, fig. 3.</p><p>Type material. A  single aboral cup (holotype) from the Miocene of the Superga Hill near Turin and two plaster casts are housed in the palaeontological collection of the MNHN in Paris (MNHN-F-R68284). Several casts of this holotype were sent to collections outside of France. Two of them are housed in the Palaeontological Museum of Turin University (Manni &amp; Pacioni 2021). The cast of an aboral cup (MPURLS NS 6/800) having nearly the same size and belonging to the Palaeontological Museum of Rome (Manni 2005) is also a cast of the holotype .</p><p>Emended diagnosis. Aboral cup rounded subpentagonal, diameter up to 14.2 mm, ratio of height to diameter 0.7, dense regular granulation developed especially on upper part of external surface, interradial processes conspicuous, no marked growth bands.</p><p>Description of the holotype. Aboral cup rounded subpentagonal, with granulated external surface as indicated in Michelin’s diagnosis. Quantitative characters listed in Table 6. Wall relatively thick (2.5–2.7 mm); interradial processes conspicuous but irregularly preserved; synarthrial articulation of radials worn, aboral ligament pit inconspicuous (Fig. 8G). Lateral view asymmetrical (H/h 1.6); side of maximum height with a convex subconical growth discontinuity of smooth surface related to the undulating attachment surface (Fig. 8F). Dense regular granulation especially well developed on the upper part of the external surface and extended very close to the fulcral ridge of radial articulation, no marked growth bands.</p><p>Other specimen. Manni (2005) figured a small aboral cup (quantitative characters listed in Table 6) with a granulated external surface belonging to the Michelotti Collection housed in the Museum of Palaeontology of “La Sapienza” University of Rome (MPURLS 2534). He attributed it to a juvenile of  C. gastaldii corresponding to the specimen cited by Michelotti (1861) from the Miocene near Turin.</p><p>Remarks. Jaekel (1891) considered  Micropocrinus gastaldii to be very similar to the Cretaceous  C. holopus and deemed that the difference in granulation was a result of better preservation. Rasmussen (1961, p. 246) pointed out that  C. gastaldii is “very similar to large specimens of  C. depressum and to specimens of  C. holopus with low theca and a broad base”. Manni and Pacioni (2021) attributed to  C. gastaldii three poorly preserved specimens from Cava Boschetto (= Cengio dell’Orbo quarry, Chiampo Valley, Lutetian), specimens here maintained in open nomenclature (see above). However, at the species rank, all these presumed affinities are not well supported by discriminating characters. Additional new material is required to confirm the validity of  C. gastaldii .</p><p>Occurrence. Serravallian (Middle Miocene) of  Superga Hill near Turin, northwestern Italy.</p></div>	https://treatment.plazi.org/id/2F1AC921FFC3FFC0F2F0FCAAFB345BA2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFDDFFDEF2F0FE53FCC45FEC.text	2F1AC921FFDDFFDEF2F0FE53FCC45FEC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Holopus Orbigny 1837	<div><p>Genus  Holopus Orbigny, 1837</p><p>Type-species.  Holopus rangii Orbigny, 1837, by monotypy.</p><p>Remarks. For main discriminating characters between  Holopus and  Cyathidium, see Table 3.  H . fabianii  sp. nov. and  H. spileccense are the only fossil species of  Holopus for which both the aboral cup and brachials are described.  H. plaziati is known only from brachials, and the Campanian species in Jagt et al. (2010) is known exclusively from a single aboral cup. We here present the first description of well-differentiated keystone plates as well as distal brachials in a fossil  Holopus species.</p><p>Included fossil species and occurrences.  Holopus fabianii sp. nov. (lower Lutetian of Chiampo Valley, north-eastern Italy),  H. plaziati (lower Ypresian, Corbières, southern France),  H. spileccense (lower Ypresian, Spilecco Hills near Bolca, north-eastern Italy),  Holopus sp. in Jagt et al. (2010) (upper Campanian, Kronsmoor, northern Germany).</p><p>Stratigraphic distribution. Upper Campanian–Recent.</p></div>	https://treatment.plazi.org/id/2F1AC921FFDDFFDEF2F0FE53FCC45FEC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFDDFFD8F2F0FC18FBC25D28.text	2F1AC921FFDDFFD8F2F0FC18FBC25D28.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Holopus fabianii Martinez-Soares & Roux & Giusberti & Gatto & Eléaume & Améziane 2024	<div><p>Holopus fabianii sp. nov.</p><p>Figs. 7, 8H–J, 9–11; Tables 7–10</p><p>Synonymy.  Holopus spileccense Manni &amp; Pacioni, 2021: 69–73, figs. 4–6; Frisone et al., 2020: 306 (non Schlüter 1878).</p><p>Type material.  Eight aboral cups including the holotype (Fig. 8 H – I), one figured paratype (paratype 1, Fig. 8J), six non-figured paratypes (paratypes 2–7); 17 figured brachials including 12 IBrax (paratypes 8–19, Fig. 10A–L) and five IIBr (paratypes 20–24, Fig. 11A–H); 14 non-figured IBrax (paratypes 46–59), 21 non-figured IIBr (paratypes 25–45) including three keystone plates and three IIBr from distal series .</p><p>Other material examined. 17 IBrax, 58 IIBr of Wp&lt;2.8 mm, 21 IIBr of Wp&gt; 2.8 mm.</p><p>Etymology. Species dedicated to Ramiro Fabiani (1879–1954) for his contributions to stratigraphy and palaeontology of the Paleogene from north-eastern Italy.</p><p>Diagnosis. Cup with conspicuous constriction in the lower half; upper part of cup with rounded pentagonal cross-section; pronounced radial ribs and marked interradial ribs; adoral face without external interradial process; calycinal cavity unknown; estimated maximum cup size: H 21.0 mm, D 14.8 mm; IBrax about as high as wide, with Y-shaped ridge, upper part of ridge (V-shape) irregularly doubled, usually with outer branches reaching angles of maximum brachial width (Wd) and inner branches corresponding to radial axis of brachial ramifications; small IBrax with only a few tubercles or none at all; large IBrax with several tubercles especially in the proximal half; external ornamentation of proximal IIBr consisting of X-shaped positioned tubercles, muscular synarthries with fulcral ridge shorter than maximum brachial width axis, adoral border forming an obtuse angle giving relatively narrow V-shape section; keystone plates with distal muscular synarthry roughly triangular and rounded, proximal synarthry symmetrical with fulcral ridge as long as Wp, external face smooth, shaped like an isosceles triangle, four conspicuous parallel symplexial crenulations on each side; distal IIBr series higher than broad, muscular synarthries with inconspicuous fulcral ridge, marked aboral ligament pit and axial nervous canal, conspicuous adoral muscle pits.</p><p>Type stratum. Lower Lutetian, tuffite horizon.</p><p>Type locality.  Cengio dell’Orbo quarry in the Chiampo Valley, north-eastern Italy.</p><p>Description of type series. Among the 8 aboral cups of the type series (quantitative characters listed in Table 7), the holotype was selected as the best preserved aboral cup (Fig. 8H, I). Holotype medium sized (H 3.83 mm, D 2.75 mm), with a strong constriction (Db/Dcs 1.68) located in the aboral half and separating adoral two-thirds of cup from a broad basal part with flat attachment surface with a greater diameter than the adoral side (D/Db 0.88); adoral part with rounded pentagonal cross-section demarcated by five conspicuous radial ribs with intermediate ribs more pronounced on one side (Fig. 8H) than on the other (Fig. 8I); adoral face without external interradial process, calycinal cavity filled with recrystallized sediment making it impossible to identify any possible internal interradial processes, if present. Other aboral cups of type series are of variable sizes (H 1.72 to 6.55 mm, D 1.76 to 4.62 mm), usually height greater than adoral diameter (H/D 1.21 to 1.48) except in small paratype 7 (H/D 0.98), H/D increasing from nearly 1 to more than 1.4 with size (Fig. 9), similar characters as the holotype, more or less conspicuous, attachment surface often smaller than adoral diameter and very irregular, sometimes appearing as a mold of a subcylindrical substrate (Fig. 8J), cup with moderate curvature (H/h~1.3) only in paratype 3.</p><p>Type series of IBrax (paratypes 8 to 19) illustrate changes in form and ornamentation with size (Fig. 10, Table 8). Smallest IBrax nearly as high as wide; external surface (Fig. 10A–E) with a more or less thick Y-shaped ridge, distal part of ridge (V-shape) irregularly doubled with usually outer branches reaching angles of maximum brachial width (Wp) and inner branches corresponding to radial axis of brachial ramifications; proximal external surface without tubercles on either side of axial ridge. Larger IBrax about as wide as high except paratypes 10 and 18 which show H/ Wd 0.82 and 0.84, respectively; external ornamentation (Fig. 10I–L) showing Y-shaped ridge composed of a short lower segment and predominantly longer V-shaped segments corresponding to radial axis of brachial ramifications, 10 to 20 large tubercles developed irregularly over the entire external surface, sometimes transforming former ridges into rows of tubercles (Fig. 10K–L). Asymmetric paratype 17 possibly belongs to a lateral trivium plate (Fig. 10J). Aboral interradial process moderately developed.</p><p>Type series of IIBr, proximal series to keystone plate (paratypes 20 to23 and 25 to 39) with external ornamentation predominantly X-shaped, always with a more or less marked axial ridge and few tubercles (Fig. 11D–E). Ratios H/h, H/Wp and Wp/Wd independent of size (Table 9). Height asymmetry mostly due to oblique articulation with IBrax, 12 cases with H/h&lt;1.3, 4 cases with 1.5&lt;H/h&lt;1.7 (Fig. 11E) and 3 cases with 1.95&lt;H/h&lt;2.05 (Fig. 11D). Lateral faces with 4 to 6 parallel symplexial crenulations, sometimes restricted to the highest one. Muscular synarthries with fulcral ridge shorter than maximum brachial width (W). Two types of muscular synarthries: one symmetrical with equal fulcral ridge segments parallel to W (Fig. 11A), another asymmetrical with unequal fulcral ridge segments oblique to W (Fig. 11B); most IIBr with one synarthry asymmetrical and the other symmetrical (IIBr1 or IIBr2?), other IIBr with two articular facets asymmetrical (IIBr1) or symmetrical (IIBr2 or 3?). Four IIBr nearly as high as wide and asymmetrical, assigned to IIBr1 of a bivium (Fig. 11E). Articulations with adoral border forming an obtuse angle, ambulacral groove shallow, giving the whole ossicle a relatively flat V-shaped section.</p><p>Keystone plates of type series (paratypes 24 and 40–42, Table 10) with distal muscular synarthry roughly triangular and rounded (Fig. 11F); proximal synarthry symmetrical, with fulcral ridge as long as Wp (Fig. 11H); external face smooth, shaped as an isosceles triangle (Fig. 11G), four conspicuous parallel symplexial crenulations on each side (Fig. 11F).</p><p>A few brachials from the distal IIBr series (paratypes 43–45, Table 10) higher than broad, muscular synarthries similar to the distal articulation of the keystone plates with inconspicuous fulcral ridge, marked aboral ligament pit and axial nervous canal, conspicuous adoral muscle pits.</p><p>Complementary data from material not included into the type series. The additional material allows us to investigate in more detail the range of variation of quantitative characters (Supplementary Tables 1–2) and ratios in the main types of brachials. All material examined including the types-series: 8 aboral cups, 43 IBrax, 46 IIBr of the proximal series (including 14 with Wp&gt; 2.8 mm) and 60 IIBr with Wp&lt;2.8 mm (distal series plus a few keystone plates poorly preserved). Ratio of IBrax to aboral cups 5.4, very close to expected ratio of 5; ratio of IBrax to proximal IIBr 1.07, far from the expected ratio of 4 or 6 indicating a high IIBr deficit. Height asymmetry (H/h) of the proximal IIBr is independent of their size. In 50% of the proximal IIBr 1.1&lt;H/h&lt;1.3 (possibly IIBr2 or IIBr3), in 34.6% 1.4&lt;H/ h&lt;1.75 (possibly IIBr1 or IIBr2) and in 15.4% 1.75&lt;H/h&lt;2.3 (possibly IIBr1). Quantitative characters of IBrax attaining maximum values with H 4.78 mm, Wd 5.30 mm, Wp 4.15 mm and Wm 3.25 mm (Appendix Table 1).</p><p>Remarks. The high number of isolated ossicles of  Holopus fabianii sp. nov. from the type locality provides a unique insight into intraspecific character variation in the genus  Holopus . To date, no comparable data have been published for extant species. However, the absence of a crown preserved with connected brachials makes it impossible to establish the precise number of IIBr in the arm’s proximal series. The combination of H/h asymmetry, which appears to increase proximally in extant species, and the fact that 50% of the IIBr have a H/h close to 1 suggest that there might have been at least 2–3 IIBr preceding the keystone plate. Like in the extant species  H. alidis, the position of the keystone plate could have varied depending on the arm and the developmental stage (Bourseau et al. 1991).</p><p>The recrystallized sediment within the calycinal cavity masks possible inner interradial processes, whose presence and characters would have been useful to describe and compare to extant species. These structures might have been absent as in  H. spileccense (see below).</p><p>The maximum size of the aboral cup and IBrax of  H. fabianii sp. nov. can be estimated by comparing the greatest width of the facets corresponding to the same articulation joining two adjacent ossicles. The proximal articulation of the largest IIBr (paratype 20) is 1.6 times wider (Wp 5.06 mm) than the largest distal articulation of IBrax (Wm 3.25 mm). The largest proximal articulation of IBrax is 2 times wider (Wp 4.15 mm) than the largest distal articulation of the available aboral cups (paratype 2, Wr 2.03 mm). Thus, the estimated size of the individual to which paratype 20 brachial belonged was 3.2 times larger than paratype 2. Therefore, the aboral cup of  H. fabianii sp. nov. could possibly be as large as H 21.0 mm and D 14.8 mm, a size intermediate between the known maximum aboral cup sizes of  H. alidis (H 13 mm, D 9 mm) and  H. rangii (H 30 mm, D 25 mm).</p><p>Holopus fabianii sp. nov. displays closer affinities with  H.alidis than with  H.rangii,as both possess a differentiated keystone plate beyond IIBr2 separating proximal and distal arm series (Figs. 5C–H and Fig. 11F–H). Both species also have an aboral cup with rather marked radial ribs. Yet,  H. alidis differs in having a distinctly granulated external cup and brachials, including the keystone plates, with a predominantly tuberculated ornamentation unlike the nearly smooth keystone plates found for  H. fabianii sp. nov. (Fig. 5C, Fig. 5 G, Bourseau et al. 1991, pl. 12). However, judging from our observation on  Cyathidium, the relevance of the latter character might be called into question. The IBrax ornamentation with Y- or V-shaped ridges common in  H. fabianii sp. nov. is restricted to juveniles of  H. alidis (Fig 4B, C) and tends to disappear under the tuberculate ornamentation in older specimens. Moreover, the adoral side of the fulcral ridge of the proximal-most brachials often conspicuously crenulated in  H. alidis (Fig. 5A–B) is not found in more distal synarthries (Fig. 5E). This character appears to be absent in  H. fabianii sp. nov. (Fig. 11A–B). Furthermore, the IIBr fulcral ridges in  H. alidis (Fig. 5B) and  H. rangii (Fig. 6D, F) are as long as the brachial is wide, which contrasts significantly with what is observed in  H. fabianii sp. nov. (Fig. 11A–C). This appears to be due to a lateral aboral displacement of the adoral surfaces of the synarthries, “opening” the articulation and thus giving the brachial a rather flat V-shape. The brachials of  H. alidis (and  H. rangii) are thus far thicker than those of  H. fabianii sp. nov., and, in IBrax, exhibit a more pronounced distal interradial process.</p><p>Holopus plaziati, known exclusively from isolated brachials (7 IBrax + 1 proximal IIBr), comes from the lower Eocene of the Corbières in southern France (Roux et al. 2021). Its IBrax differ from those of  H. fabianii sp. nov. and  H. spileccense in being generally smaller with a smaller H/Wd (Fig. 7), and in having muscular synarthries with usually conspicuous crenulation on the adoral side of the fulcral ridge and sometimes on the adoral border of the muscular area. While syntype 2 of  H. plaziati appears similar to the IBrax of  H. fabianii sp. nov., IBrax of syntypes 3 and 7 have adoral articulation facets larger than in  H. fabianii sp. nov. In  H. plaziati this increases the size of the distal interradial process and the general thickness of the brachial, resulting in an ambulacral groove that is more deeply carved into the ossicle. The conspicuous lateral symplexial crenulations in  H. plaziati are variable (parallel to undulated or branched). Its aboral surface displays irregular tuberculate ornamentation without formation of interradial or Y-shaped ridges.</p><p>Manni and Pacioni (2021) attributed 30 brachials from Cava Boschetto (= Cengio dell’Orbo quarry, the type-locality of  H. fabianii sp. nov.) to  H. spileccense (Schlüter), a taxon known only from cups found in the Ypresian strata of Spilecco Hill (see below). These 30 brachials are housed in the MGP-PD collections. Manni and Pacioni (2021) figured the ornamentation of 5 IBrax and 6 IIBr from the proximal series but did not identify keystone plates. The variation of X-shaped ridges on IIBr was well-illustrated and corresponds to the range observed in the material here ascribed to  H. fabianii sp. nov. The quantitative characters of these brachials also fall within the variation range of those of  H. fabianii sp. nov. (except IBrax MGP-PD 31442 which presents the largest Wd, 5.3 mm) and they should thus be attributed to this species.</p><p>Occurrence. Middle Eocene (Lutetian) of Chiampo Valley, north-eastern Italy.</p></div>	https://treatment.plazi.org/id/2F1AC921FFDDFFD8F2F0FC18FBC25D28	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
2F1AC921FFDAFFD5F2F0FF7CFB2159D0.text	2F1AC921FFDAFFD5F2F0FF7CFB2159D0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Holopus spileccense (Schluter 1878)	<div><p>Holopus spileccense (Schlüter, 1878)</p><p>Figs. 7, 9, 12–14; Tables 11–13</p><p>Synonymy.  Cyathidium spileccense Schlüter, 1878: 54, 65, pl. 3, figs. 11–15; Carpenter, 1884: 212; Jaekel, 1891: 614–616, fig. 14; Jaekel, 1907: 293;  Holopus (Cyathidium) spileccense Jaekel, 1891: 567,  Holopus spileccense Jaekel, 1891: 607, 615–616;  Holopus (Cyathidium) spileccensis Jaekel, 1891: 592;  Holopus spileccensis Jaekel, 1891: 591, 619–620;  Cyathidium spileccense Nielsen, 1913: 17, 40, 58; Wanner, 1929: 319;  Holopus spileccensis Sieverts, 1931: 174–175;  Holopus spileccense Biese &amp; Sieverts-Doreck, 1939: 117; Manni, 2005: 216–217, pl. 1 figs. 9–10, text-fig. 5a–b.</p><p>Non  Holopus spileccense Manni &amp; Pacioni, 2021: 69–73, figs. 4–6.</p><p>Material examined. Nine aboral cups, 12 IBr, 4 IIBr</p><p>Diagnosis. Cup with strong constriction in the aboral half; adoral part with markedly pentagonal cross-section; marked radial ridge without other ornamentation; frequently slight interradial depression; calycinal cavity without inner interradial process; measured maximum cup size: H 8.1 mm, D 6 mm; IBrax wider than high; Y-shaped ridge coarse, usually simple (not doubled) with branches corresponding to radial axis of brachial ramifications; no other ornamentation; small IBrax with more marked and doubled ridge; external ornamentation of proximal IIBr with coarse X-shaped and discrete mid-radial ridge; articular facets with fulcral ridge slightly shorter than maximum brachial width; no prominent muscular or ligament pits; more distal IIBr and keystone plates unknown.</p><p>Type stratum.  Lower Ypresian .</p><p>Type locality. Spilecco Hill near Bolca, north-eastern Italy.</p><p>Description of material examined. Quantitative characters of aboral cup relatively variable (Table 11), H/D from 1.02 to 1.63 independent of size (D used as growth index; Fig. 9). Constriction at mid-height conspicuous (D/Dcs up to 1.73), upper adoral cup and adoral face markedly pentagonal with each angle forming a radial ridge (Fig. 12E), no additional ridges or granulation on external surface, no trace of interradial process and inner process masked by recrystallized sediment (Fig. 12D), attachment surface flat to slightly concave (Fig. 12F), cup curvature usually marked, especially in the two largest specimens (H/h 1.5 to 1.6) (Fig. 12G).</p><p>Primibrachials pentagonal except for a single anomalous quadrangular specimen MGP-PD 33245 Br7 (Fig. 12C). Pentagonal IBrax broader than high (0.75&lt;H/Wd&lt;0.86), in smallest (youngest) specimens nearly as wide as high (Fig. 13A–B), external face showing coarse Y-shaped ribs, no other ornamentation (Fig. 13A–B), Y-pattern more conspicuous in small specimen (Fig. 13A) than in larger ones (Fig. 13E–F) (quantitative characters listed in Table 12). The coarse Y-shaped rib identifies the quadrangular specimen MGP-PD 33245 Br7 as a primibrachial (probably corresponding to a bivium). Articulations poorly preserved.</p><p>IIBr belonging to proximal series slightly wider than high with H/h 1.12 to 1.42 (Table 13), no keystone plate identified possibly due to the small number of IIBr found; external face showing a coarse X-shaped rib with a discrete mid-radial ridge (Fig. 12A). Distal IIBr articulations rather symmetrical (Fig. 12B). The articular facets appear to present a fulcral ridge that is slightly shorter than the maximum brachial width and there are no prominent muscular or ligament pits.</p><p>Remarks. The maximum sizes of the aboral cup and IBrax of  H. spileccense have been estimated using the same method as for  H. fabianii sp. nov. (see above) and are nearly the same as the maximum sizes measured. However, this estimation might be biased due to the small number of brachials available.</p><p>Schlüter (1878) figured four aboral cups without describing them in the text. The current whereabouts of these are unknown to us. Three of the four are rather straight without significant curvature. The fourth is markedly curved (H/h 1.56). It also appears to be the best-preserved specimen and its adoral face and side view are figured enlarged, showing the main characters of the species (Fig. 14; Schlüter 1878, pl. 3, figs 11, 12). Its cup is pentagonal, higher than wide (H/D 1.22), with a conspicuous constriction (D/Dsc 2.09) in the lower part (H/ht 1.33), marked radial ribs, slight interradial depressions with an outer interradial process, and symmetric, rounded muscular synarthries. While not explicitly shown in the drawing, the calycinal cavity is described in the text as being partly filled with sediment. We here designate this drawn aboral cup as the lectotype of  H. spileccense . The side views of the three paratypes show variable general shape (1.37&lt;H/D&lt;1.78, 1.4&lt;D/Dcs&lt;2.0). Jaekel (1891: 616, fig. 14) figured an aboral cup (sent to him by Schlüter) that is very similar to the lectotype except that it is not curved (H/D 1.26, H/h 1.03, H/ht 1.45, D/Dcs 1.98). As Schlüter (1878) and Jaekel (1891) do not give precise measurements, we can only deduce ratios with some confidence from the figures. Measures taken from a printed copy of Schlüter (1878) are listed in Table 11. Manni (2005) described and figured a single aboral cup from the Michelotti Collection (H/D 1.0, H/h 1.04, H/ht 1.92, D/Dcs 1.29). With the 9 additional aboral cups described here, we now have 15 cups from Spilecco allowing us to assess the variation of quantitative and qualitative characters in  H. spileccense . A comparison with data for  H. fabianii sp. nov., based on eight aboral cups, indicates a similar range of variation in the two species (Table 14).  Holopus spileccense differs in having the adoral part of the cup more markedly pentagonal and having five usually well-marked radial ribs as well as interradial depressions that can be more or less pronounced.</p><p>A comparison of the quantitative characters of the brachials of  H. spileccense and  H. fabianii sp. nov. reveals further differences (Tables 15 and 16). The former species differs in having IBrax that are wider than high (H/Wd up to 0.93, mean 0.84) and IIBr are less asymmetrical (H/h up to 1.43, mean 1.19) tending to be almost as high as wide (mean H/Wp 0.91). The brachials of both species differ further in their ornamentation. While the number of brachials available is limited, the ornamentation patterns remain consistent: coarse ribs without granulation in  H. spileccense versus finer ridges with pronounced derived granulation in  H. fabianii sp. nov. Nevertheless, both species share the same Y-shaped ridge pattern in IBrax and X-shaped ridges in IIBr. The two species are likely closely related as both might lack the inner interradial processes in the aboral cup and share keystone plates, which have not yet been found in  H. spileccense . The presence of a well-developed keystone plate would put both these species closer to the extant  H. alidis than to the Carribean congeners. However, if the drawings of  H. spileccense by Schlüter are accurate, its rounded radial articulations with an interradial depression and the outer interradial process are reminiscent of  H. rangii .</p><p>The brachials from the Lutetian of the Chiampo Valley attributed to  H. spileccense by Manni and Pacioni (2021) are instead here referred to  H. fabianii sp. nov. (see above). Thus, to date, the stratigraphic distribution of  H. spileccense appears confined to the Ypresian.</p><p>Occurrence. Lower Eocene (lower Ypresian) of Spilecco Hill near Bolca, north-eastern Italy.</p></div>	https://treatment.plazi.org/id/2F1AC921FFDAFFD5F2F0FF7CFB2159D0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Martinez-Soares, Pablo;Roux, Michel;Giusberti, Luca;Gatto, Roberto;Eléaume, Marc;Améziane, Nadia	Martinez-Soares, Pablo, Roux, Michel, Giusberti, Luca, Gatto, Roberto, Eléaume, Marc, Améziane, Nadia (2024): New Eocene species of the crinoid genera Holopus and Cyathidium (Cyrtocrinida: Holopodidae) from north-eastern Italy. Zootaxa 5541 (4): 401-437, DOI: 10.11646/zootaxa.5541.4.1, URL: http://dx.doi.org/10.11646/zootaxa.5360.3.8
