identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03B387E30B67FFABFF39FDF6FA6118E1.text	03B387E30B67FFABFF39FDF6FA6118E1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster Jagt 2000	<div><p>Genus  Haccourtaster Jagt, 2000</p><p>Type species.  Haccourtaster aemstelensis Jagt, 2000, by original designation.</p><p>Diagnosis. A small, pentagonal goniasterid in which two superomarginals and two inferomarginals are typically present on each side of the arm. A narrow, depressed margin is present on all marginals, and the raised central region bears coarse granule pits. Blunt internal intramarginal cavities are present between the marginals.</p><p>Other included species.  Haccourtaster hrbac Žítt, 2005,  Haccourtaster berryensis sp. nov.,  Haccourtaster liticola sp. nov. and  Haccourtaster nattestadae sp. nov.</p><p>Remarks.  Haccourtaster is a distinctive small goniasterid, which typically occurs in marginal marine Upper Cretaceous facies. Although two inferomarginals are typically present on each side of the arm a small, distal inferomarginal is found on some specimens. The internal intramarginal cavities (Žítt, 2005, fig. 11A) represent short, narrow extensions of the coelom into the marginal frame and are of unknown function. Species are characterised by distinctive marginal morphologies and external sculptures. In the present paper, we describe three new species of the genus.</p><p>Haccourtaster probably evolved from a species of  Metopaster, by retaining four marginals on each side of the disc, either developing paedomorphically from a species with more marginals, or from an ancestor which also possessed four marginals (see below; Discussion). It developed the distinctive intramarginal cavities in the interior of the marginals, and the ultimate inferomarginal became enlarged to correspond with the ultimate superomarginal.</p></div>	https://treatment.plazi.org/id/03B387E30B67FFABFF39FDF6FA6118E1	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
03B387E30B67FFA9FF39FAA5FE131E14.text	03B387E30B67FFA9FF39FAA5FE131E14.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster berryensis Gale & Jagt 2025	<div><p>Haccourtaster berryensis sp. nov.</p><p>urn:lsid:zoobank.org:act: D71A59CE-ABD9-4469-92CE-0D39455B15A3</p><p>Fig. 1G–I</p><p>1987a  Metopaster thoracifer (Geinitz, 1872): Gale, p. 37, text-fig. 12.</p><p>Type material.  Associated ossicles from a single individual, including 14 superomarginals and 6 inferomarginals, from the  lower Turonian chalk at Berry Cliff, Branscombe (Devon, UK) (NHMUK PI EE 18203)  .</p><p>Diagnosis. A small  Haccourtaster (R = 4–5 mm) in which the trapezoidal median superomarginals narrow towards the lateral border. Central regions of marginals sharply demarcated from a narrow, depressed rim. Median superomarginals carry a row of centrally placed large granule pits; inferomarginals smooth.</p><p>Description. The median superomarginals (Fig. 1G, I) are trapezoidal and narrow towards the lateral margin; the breadth is approximately twice the length. The slightly convex central area is sharply demarcated from a narrow, depressed rim. The median superomarginals bear a single, median row of granule pits and a few scattered pits towards the inner margin. The ultimate superomarginals (Fig. 1G) are equilaterally triangular, gently tumid and the lateral margin is convex. A group of 4–5 granule pits are present on the lateral-distal margin. The median inferomarginals (Fig. 1H, I) are rectangular and slightly broader than the corresponding superomarginal (Fig. 1I). They possess a narrow, depressed margin and entirely lack granule pits. The distal inferomarginals (Fig. 1H) have the outline of right triangles in actinal view and correspond precisely with the enlarged ultimate superomarginals. Large intramarginal cavities are present (Fig. 1I).</p><p>Remarks.  Haccourtaster berryensis sp. nov. differs from all other species of the genus in its possession of trapezoidal superomarginals bearing very sparse granule pits.Although the holotype specimen might be an immature individual, it differs significantly in morphology from material of  H. hrbac of similar dimensions (compare Fig. 1G with Fig. 1J). In small  H. hrbac, the superomarginals are nearly square in abactinal view, whereas they are proportionately broad and trapezoidal in  H. berryensis sp. nov.</p><p>The succession at Berry Cliff, 2 km west of Branscombe Mouth in Devon was described by Rowe (1904, p. 26), who found chalks containing abundant asteroid ossicles and cidarid echinoid debris, which he ascribed to the zone of  Rhynchonella cuvieri (=  Mytiloides mytiloides inoceramid Zone, lower Turonian). The asteroids include numerous ossicles of  Metopaster thoracifer (see Spencer, 1913), and evidently characterise a shallower environment than that represented elsewhere in southern England. The locality lies 4 km east of the western margin of preserved Turonian chalks in the United Kingdom.</p></div>	https://treatment.plazi.org/id/03B387E30B67FFA9FF39FAA5FE131E14	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
03B387E30B65FFA9FF39FDF0FE551AB0.text	03B387E30B65FFA9FF39FDF0FE551AB0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster liticola Gale & Jagt 2025	<div><p>Haccourtaster liticola sp. nov.</p><p>urn:lsid:zoobank.org:act: EA485F5E-310D-457E-8FBD-2DBD95D3EAE7</p><p>Fig. 1A–F</p><p>Type material.   The median superomarginal (Fig. 1C) is the holotype (NHMUK PI EE 18204); the other figured marginals are paratypes (NHMUK PI EE 18205–18224).  Upper lower Campanian of Ivö Klack (Skåne, southern Sweden)  .</p><p>Diagnosis. A relatively large  Haccourtaster (R = 10–12 mm; r = 8–10 mm) in which the surfaces of the marginals are evenly convex and bear densely and evenly spaced, impressed granule pits and oval pedicellariae.</p><p>Additional material. Over 100 isolated marginals ossicles from the type locality and horizon (NHMUK, A.S. Gale Collection) .</p><p>Etymology. Latin, litus, shore, - icola, inhabitant, in allusion to the shallow-water, rocky shoreline habitat of the species.</p><p>Description. The median superomarginals (Fig. 1A, C, F) are trapezoidal in abactinal outline, narrowing to the lateral border, and the breadth is twice the length. The abactinal surface is evenly convex and carries slightly impressed granule pits and short oval bases for pedicellariae (Fig. 1C). The ultimate superomarginals are equilaterally triangular (Fig. 1D) and abactinal and lateral surfaces are well demarcated. The median inferomarginals (Fig. 1B, E, F) are rectangular and have a breadth which is twice the length. The external surface is evenly tumid and bears numerous, shallow, evenly sized and spaced granule pits. The enlarged 2 nd inferomarginals (Fig. 1B) are triangular and taper distally (Fig. 1B, left). In some cases, a facet for a short 3 rd inferomarginal is present (Fig. 1B, right). The intramarginal cavities (Fig. 1F) extend close to the lateral borders of the ossicles.</p><p>Remarks.  Haccourtaster liticola sp. nov. differs from all other species of the genus in its evenly tumid marginals which possess dense, evenly spaced granule pits of equal dimensions. The locality of Ivö Klack represents an upper lower Campanian rocky shoreline, where marginal sediments rest on Proterozoic gneiss. Boulders encrusted with zoned epifauna (Surlyk &amp; Christensen 1974; Surlyk &amp; Sørensen 2010) were buried by bioclastic shell sands and gravels, deposited in less than 30 metres of water. These yield an abundant and highly diverse calcitic fauna of bryozoans, echinoderms, bivalves, serpulids, brachiopods and cirripedes (Surlyk &amp; Sørensen 2010; Sørensen &amp; Surlyk, 2010, 2011a, b; Sorensen et al. 2012; Gale &amp; Sørensen, 2015). Asteroids were described by Gale (1987 a, 2025) including a number of species of  Metopaster that are endemic to the Kristianstad Basin. These include, for instance,  M. bromleyi Gale, 1987a, morphologically close to the older  M. thoracifer which is widely associated with species of  Haccourtaster .</p></div>	https://treatment.plazi.org/id/03B387E30B65FFA9FF39FDF0FE551AB0	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
03B387E30B65FFAFFF39F894FBE71900.text	03B387E30B65FFAFFF39F894FBE71900.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster nattestadae Gale & Jagt 2025	<div><p>Haccourtaster nattestadae sp. nov.</p><p>urn:lsid:zoobank.org:act: 74BB38E5-CFFA-45EF-9A49-0988DE70E146</p><p>Fig. 2L–O</p><p>Type material. A group of associated and partly articulated ossicles (MGUH 34867, holotype) from the Arnager Limestone (lower Coniacian) of the coastal cliff southwest of Arnager (Bornholm, Denmark; compare Kennedy &amp; Christensen 1991; Tröger &amp; Christensen 1991; Christensen &amp; Schulz 1997).</p><p>Diagnosis. Medium-sized  Haccourtaster (R approximately 8 mm) in which the superomarginals are strongly convex and have a central ridge. Marginals possess evenly spaced granule pits some of which are large and deeply impressed on the median ridges of the superomarginals.</p><p>Etymology. Named after local non-professional palaeontologist, Marianne Falbe Nattestad, who collected this and other material of early Coniacian echinoderms from the cliffs near Arnager, Bornholm.</p><p>Description. The holotype comprises numerous ossicles, some articulated, in a small piece of chalk. The median superomarginals (Fig. 2L–O) are rectangular in abactinal view, and twice as broad as long. The abactinal surface is tumid (Fig. 2N, O); in some ossicles this is symmetrical (N), whereas in others the distal margin is sharply depressed (O). The central area of the median superomarginals is sharply defined by a narrow ridge. The central area carries rather evenly spaced granule pits, some of which are deeply impressed (Fig. 2N); in one case (Fig. 2O) larger pits are present on the lateral-abactinal part of the tumidity. The ultimate superomarginals are triangular with a steep lateral surface which carries evenly sized granule pits (Fig. 2O); the abactinal face bears large, widely spaced crater-like pits. The inferomarginals are gently convex and carry evenly sized and evenly spaced granule pits. The median inferomarginal (Fig. 2M, left) is rectangular in outline, the enlarged penultimate inferomarginal (Fig. 2M, right) tapers distally. A very small 3 rd inferomarginal is present distal to the enlarged 2 nd ossicles. Intramarginal cavities are present (Fig. 2L, upper right).</p><p>Remarks.  Haccourtaster nattestadae sp. nov. differs from its closest relative,  H. liticola sp. nov. in the presence of a raised median ridge on the superomarginals which bears deeply impressed, crater-like granule pits. Additionally, the inferomarginals are less broad (compare Fig. 1B with Fig. 2M) and  H. liticola sp. nov. lacks the narrow ridge surrounding the central area on the superomarginals in  H. nattestadae sp. nov. The Arnager Limestone was interpreted by Svennevig &amp; Surlyk (2018) to represent a relatively deep chalk environment, dominated by hexactinellid sponges and the other invertebrate fauna is of very low diversity. The only other asteroid described previously from the Arnager Limestone is  Metopaster cf. thoracifer (Wienberg Rasmussen 1950; Svennevig &amp; Surlyk 2018). However, recent collecting from the cliff southwest of Arnager by Ms Nattestad has yielded additional material of what Wienberg Rasmussen (1950) referred to as  M. cf. thoracifer, inclusive of (semi-articulated) specimens. These, and comparative material of late Cenomanian–early Turonian  M. thoracifer from the Dresden area (eastern Germany) and the Bohemian Cretaceous Basin (Czech Republic) as described by Geinitz (1871) and Niebuhr &amp; Seibertz (2016) casts doubt over this specific assignment. This matter will be discussed in full in a forthcoming paper.</p><p>The Arnager locality is less than 2 km from outcropping Palaeozoic basement (Svennevig &amp;Surlyk 2018, fig. 1c), so it is possible that the two currently known asteroid taxa, both characteristically nearshore species, were washed into the region from a coastal environment, perhaps a rocky shoreline.</p></div>	https://treatment.plazi.org/id/03B387E30B65FFAFFF39F894FBE71900	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
03B387E30B63FFAFFF39FAC4FBA61B82.text	03B387E30B63FFAFFF39FAC4FBA61B82.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster hrbac Zitt 2005	<div><p>Haccourtaster hrbac Žítt, 2005</p><p>Fig. 1J–M</p><p>2005  Haccourtaster hrbac Žítt: p. 226, figs 3–7, 11A.</p><p>Type material. The holotype is a median superomarginal, figured by Žítt (2005, fig. 4A–E; NMP O 6367); the other ossicles illustrated by Žítt (2005) are paratypes (NMP O 6368, 6370–6394). The material is from the uppermost portion of the Korycany Member (Peruc-Korycany Formation), lower Turonian ( Whiteinella archaeocretacea foram Zone) as exposed at Velim-Václav, Czech Republic.</p><p>Diagnosis. A  Haccourtaster of medium size (R = 7–8 mm) in which the median superomarginals have a lateral, abactinal swelling which bears a cluster of deeply impressed granule pits. A row of similar pits is present on the lateral margin of the ultimate superomarginal.</p><p>Remarks.  Haccourtaster hrbac is distinguished by the distinctive tumidity, bearing deeply impressed granule pits, on the lateral part of the median superomarginals, and the narrow ridge along the lateral margin of the ultimate superomarginals which carries a row of similar pits. The species occurs in sediments deposited on a rocky shoreline and is associated with common  Metopaster thoracifer (T. Kočí, pers. comm., 2004).</p></div>	https://treatment.plazi.org/id/03B387E30B63FFAFFF39FAC4FBA61B82	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
03B387E30B60FFACFF39FA52FC751BF9.text	03B387E30B60FFACFF39FA52FC751BF9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haccourtaster aemstelensis Jagt 2000	<div><p>Haccourtaster aemstelensis Jagt, 2000</p><p>Fig. 2A–K</p><p>2000  Haccourtaster aemstelensis Jagt: p. 408, pl. 27, figs 1–6.</p><p>2005  Haccourtaster aemstelensis Jagt: Žítt, p. 226.</p><p>Type material. An ultimate superomarginal figured by Jagt (2000, pl. 27, fig. 1) is the holotype (NHMM MB 1044- 6/c) and other marginals illustrated by Jagt (2000, pl. 27, figs 2–5) are paratypes (NHMM MB 1044-6d–f), basal Zeven Wegen Member (Gulpen Formation), lowermost upper Campanian, Haccourt, northeastern Belgium.</p><p>Diagnosis. A small  Haccourtaster (R estimated to have been 5 mm) in which the external surfaces of the marginals bear deeply impressed, large, closely spaced granule pits separated by rugosities on the abactinal surfaces of the superomarginals and the actinal surfaces of the inferomarginals.</p><p>Additional material. Eight additional marginals from the same horizon and locality as the type material (NHMM JJ 16614) .</p><p>Remarks.  Haccourtaster aemstelensis differs from all other species of the genus in its possession rugose marginal ossicles and deeply impressed granule pits of variable size.</p></div>	https://treatment.plazi.org/id/03B387E30B60FFACFF39FA52FC751BF9	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	Gale, Andrew S.;Jagt, John W. M.	Gale, Andrew S., Jagt, John W. M. (2025): New species of the asteroid genus Haccourtaster Jagt, 2000 (Echinodermata, Goniasteridae) from the Upper Cretaceous of Denmark, Sweden and the United Kingdom. Zootaxa 5632 (2): 325-336, DOI: 10.11646/zootaxa.5632.2.6, URL: https://doi.org/10.11646/zootaxa.5632.2.6
