New Latest Cretaceous and Earliest Paleogene Asteroids (Echinodermata) from the Netherlands and Denmark and Their Palaeobiological Significance

Home , Calliderma (sea star), Dipsacaster, Geon (geology), Mediaster

BULLETIN DE L’INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE, 75: 183-200, 2005 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN, 75: 183-200, 2005

New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) from The Netherlands and Denmark and their palaeobiological significance

by Daniel B. BLAKE & John W.M. JAGT

B l a k e , D.B & J a g t , 2005. — New latest Cretaceous and famille alors qu’aujourd'hui elle est limitée aux mers profondes. Elle earliest Paleogene asteroids (Echinodermata) from The Netherlands appartient aux Neobenthopectininae, ce qui démontre la présence au and Denmark and their palaeobiological significance.Bulletin de Mésozoïque de benthopectinidés dérivés. l ’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 75: 183-200, 5 pis; Bruxelles-Brussel, March 31, 2005 - ISSN Mots-clefs: Asteroidea, Crétacé, Paléogène, Pays-Bas, Danemark, 0374-6291. taxionomie.

A bstract Introduction

Three new starfish (Skiaster vikingr n. gen., n. sp., Betelgeusia exposita Although the Asteroidea includes many heavily skeleto n. sp., and Aldebarania taberna n. sp.), and the first fossil occurrence of nized species, specimens are rare among marine inverte Cheiraster? sp., are recorded from Maastrichtian (Late Cretaceous) and Danian (Early Paleogene) rocks of The Netherlands and Denmark. brate fossils, and as a result overall history of the class Skiaster vikingr, a member of the goniasterid subfamily Pseudarchas- and its phylogeny are poorly understood. The Cretaceous terinae, adds to the known diversity and apparent significance of that and Paleogene chalks of northwest and central Europe subfamily.Betelgeusia exposita is the second Cretaceous species of the Radiasteridae to be described; together, the two species suggest that have provided one of the more extensive ancient asteroid this now infrequently encountered deep-water family was of greater faunas, and one that has been studied by generations of significance in the past. Morphology and occurrences of Betelgeusia dedicated palaeontologists. Nevertheless, fossils are in suggest niches similar to those now occupied by Astropecten. Aldebar ania taberna (Astropectinidae) is similar to A. arenitea from eastern frequently encountered, and most occurrences consist of North America, suggesting communication across a narrower North disarticulated marginals, which, although important, pro Atlantic. Cheiraster? sp. (Benthopectinidae) provides added support vide only an incomplete history. Mesozoic and Cenozoic for shelf occurrence during the Cretaceous of this now deep-water events leading to the emergence of the contemporary family; it is a member of the Neobenthopectininae, indicating Meso zoic presence of derived benthopectinids. Reflecting earlier work, the fauna have received only limited attention. present study suggests gradual emergence o f the modem asteroid fauna Four recently discovered chalk specimens representing during the Mesozoic and Cenozoic without major terminal Cretaceous all three orders of valvatacean asteroids ( sensu B l a k e , extinction. 1987) document a bit of the status of this important group Key-words: Asteroidea, Cretaceous, Paleogene, The Netherlands, near the end of the Mesozoic and beginning of the Ce Denmark, taxonomy. nozoic eras. These fossils, in conjunction with others previously described (e.g., B lake & Reid, 1998;V i l l i e r et al., 2004), suggest a gradual emergence of familial R ésum é groups established in the Mesozoic and absence of abrupt faunal change at the end of the Cretaceous. Trois nouvelles astéries (Skiaster vikingr n. gen., n. sp., Betelgeusia exposita n. sp., et Aldebarania taberna n. sp.) et la première présence fossile de Cheiraster? sp. ont été reconnues sur la marge continentale dans des dépôts d’âge Maastrichtien (Crétacé supérieur) et Danien Geographic and stratigraphie setting (Paléocène) des Pays-Bas et du Danemark.Skiaster vikingr de la sous-famille goniastéride Pseudarchasterinae étend la diversité connue et la signification apparente de cette sous-famille.Betelgeusia exposita Of the four taxa here described, three have been collected est le second taxon crétacé des Radiasteridae à être décrit. A eux deux from units assigned to the Maastricht Formation [late ces deux espèces laissent supposer que cette famille typique, pour les Maastrichtian, Belemnitella junior and Belemnella (Neo- dépôts profonds, mais rare aujourd’hui, fût plus importante dans le passé. La morphologie et les occurrences de Betelgeusia suggèrent des belemnella) kazimiroviensis zones], as exposed in the niches similaires à celles occupées parAstropecten. Aldebarania ta type area of the Maastrichtian Stage (SE Netherlands, berna (Astropectinidae) est proche de A. arenitea de l’Est de l’Amér NE Belgium). In recent years, the basal portion of the ique du Nord - probablement grâce à un Océan Atlantique nord plus étroit. Cheiraster? sp. (Benthopectinidae) ajoute un argument en faveur Gronsveld Member at the ENCI-Maastricht bv quarry (St de la présence de cette famille sur le shelf pendant le Crétacé de cette Pietersberg, Maastricht) has yielded numerous well-pre 184 Daniel B. BLAKE & John W.M. JAGT

served bourgueticrinine crinoids ( J a g t , 1999b), with as occurrence (e.g., B l a k e , 1988; V i l l i e r et al., 2004). sociated, albeit rarer, comatulids, as well as ophiuroids The poor fossil record of asteroids has not demon ( J a g t , 2000a) and dissociated asteroid ossicles ( J a g t , strated significant extinction within the group at the 2000b). This assemblage, which includes Cheiraster? end of the Cretaceous; instead, implications such as sp., accumulated under storm conditions followed by those discussed below suggest more subtle change in near-absence of bioturbation.L ie b a u (1978) typified this faunal make-up through the Mesozoic and Cenozoic. part of the Maastricht Formation as a mid-sublittoral Discussion below treats the Pseudarchasterinae as ba setting, with subtropical temperatures and characterised sal within the Goniasteridae; this approach follows the by the occurrence of seagrass communities. phylogeny of extant families ofB l a k e (1987, 1990b), Aldebarania taberna n. sp. is from the upper portion of which in turn was based on publications of many sys- the Nekum Member, which at the CBR-Romontbos quar tematists, including V e r r i l l (1899), discussed below. ry (Eben Emael, Liège), comprises a discontinuous tab Pseudarchaster is known from the Portlandian (Upper ular flint level from which numerous echinoids and am- Jurassic; see B l a k e , 1986), which is fairly close to but monoids have been collected recently. This level is close somewhat younger than the age of the oldest-known to the Kanne Horizon, an undulating erosion surface goniasterids. This chronological inconsistency might re overlain by very coarse bioclastic sand. The type of sult from the poor fossil record (V illier & Kutscher, Betelgeusia exposita n. sp. was collected from the over- 1999), or pseudarchasterines might remain unrecognised lying unit, the Meerssen Member, at the ENCI-Maastricht among known Jurassic fossils, or phylogenetic inference bv quarry, from a level that has yielded numerous ophiur based on the morphology of extant taxa might be incor oids ( J a g t , 2000a) and decapod crustacean remains. Ac rect; these are hypotheses to be tested by future research. cording to Liebau (1978), the Nekum and Meerssen No phylogenetic analysis is attempted here. A pseu- members are high-energy deposits, being characterised darchasterine-like morphology was selected as a basal by a high production of carbonate detritus that led to the configuration for crown-group asteroid morphology establishment of a broad, shallow, well-lit and warm ( B l a k e , 1990b) and discussions here do not affect that carbonate platform with a rich phytal association. There interpretation. Efforts have been made to isolate a stem- is a continuous shallowing trend with the Meerssen Mem group branch basal to the crown group (Blake & Hag ber, but deposition of the highest unit (IVf-6) must have d o r n , 2003; Blake & Hotchkiss, 2004), but candidate occurred in deeper water. genera do not appear close enough to pseudarchasterines Skiaster vikingr n. gen., n. sp., is from Stevns Klint, to provide a reliable outgroup for phylogenetic recon north of Hojerup Kirke, having been collected from the struction of the Pseudarchasterinae. Discussions below, early Paleocene (Danian)Tylocidaris abildgaardi Zone e.g. on the position of Skiaster n. gen., therefore use a bioherms (Gravesen, 1993). The matrix is bryozoan traditional descriptive notion of morphological interme limestone, which B r o m l e y (1979) considered to have diacy. been deposited in shallow water, below wave base. Skiaster vikingr n. gen., n. sp. (Valvatida) is a member Associated goniasterid asteroids include the key index of the Pseudarchasterinae S l a d e n , 1889. V e r r i l l (1899) taxa Metopaster spencerii Brünnich Nielsen, 1943, found this subfamily to be morphologically intermediate M. kagstrupensis Brünnich Nielsen,1943, and Cratera- between more typical goniasterids and certain astropecti- ster anchylus (Brünnich Nielsen,1943). nids; he specifically citedPlutonaster S l a d e n , 1889 but Tethyaster S l a d e n , 1889 and other genera could be listed as well. The fossil record includes examples of seemingly Palaeobiological significance intermediate morphology; for instance, the Jurassic astropectinid Pentasteria V a l e t t e , 1929 is suggestive of as- Fossils described here document a small part of status of tropectinids such as Plutonaster as well as of goniasterids crown-group asteroid occurrence near the Mesozoic-Ce- (e.g., H e s s , 1960, 1973, 1987;Hess & Blake, 1995; nozoic transition. The fossil record of asteroids is poor K utscher & Röper, 1999). and provides only limited information on the post-Pa- V e r r i l l (1899) carried evaluation of inferred transi laeozoic history of the class(V illier & K utscher, 1999). tional types a step further in noting thate.g., Mediaster These authors nevertheless were able to recognise two is intermediate between pseudarchasterines and more char stages in crown-group diversification, consisting of a acteristic goniasterids. The more characteristic goniaster Jurassic radiation followed by a long period of compara ids typically have non-fasciolate, more equidimensional tive quiescence. The crown group is first known from the (rather than tabular) marginals. In addition, abactinals are Triassic ( Z a r d i n i, 1973; Blake & Hagdorn, 2003) but tabular, closely abutted, and both marginals and abactinals the fossil record from this interval is extremely sketchy, are incompletely granulate in some taxa. Apparent dom even by the standards of asteroids. Significant asteroid inance of goniasterid assemblages by varied non-pseu- diversification has been documented from Jurassic strata darchasterines in the Cretaceous chalk faunas (e.g., S l a [e.g., W r i g h t , 1863-1880; Schöndorf, 1910; H e s s , d e n , 1891, 1893; S p e n c e r , 1905, 1907, 1908, 1913; 1972, 1975; B l a k e , 1984, 1990a] including representa Schulz & W eitschat, 1971, 1975, 1981;B r e t o n , 1979, tion of many surviving families. Other families, some of 1992; G a l e , 1987a, b;J a g t , 2000b) documents significant fragile construction, have a Cretaceous earliest-known diversification of goniasterids during Cretaceous time. New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 185

Pseudarchaster and Skiaster n. gen. are joined by RGM - Nationaal Natuurhistorisch Museum, Leiden (formerly Rijks museum van Geologie en Mineralogie); USNM - United States Na another Mesozoic pseudarchasterine, Formalhautia tional Museum, Washington DC. B lake & Reid, 1998, which exhibits a third, distinctive pseudarchasterine morphology (inFormalhautia, a com Class Asteroidea de B lainville, 18 3 0 paratively small disk and long, slender arms reminiscent Order Valvatida P e r r i e r , 1 8 8 4 of those of certain species of Astropecten). The morpho Family GoniasteridaeF o r b e s , 1841 logical differences among the three suggest a diversity of Subfamily Pseudarchasterinae S l a d e n , 1 8 8 9 pseudarchasterines during the Mesozoic and perhaps the early Cenozoic, as well as significant convergence be tween pseudarchasterines and other goniasterids. R e m a r k s Betelgeusia exposita (Paxillosida; Radiasteridae) is Skiaster n. gen. exhibits robust, tabular, closely appressed only the second member of its family to be described, marginals that lack significantly enlarged spines, and joining the mid-Cretaceous B. reidi B lake & Reid, 1998. stout, paxilliform abactinal ossicles. These characters Betelgeusia is similar to extant Radiaster, which is indicate the subfamily Pseudarchasterinae within the fa known from a comparatively few widely separated mily Goniasteridae. Further, superomarginals abutted at deep-water settings, whereas Betelgeusia in contrast is the dorsal arm midline are only known in goniasterids. now known from Cretaceous shallow-water, perhaps tur Ossicular form is also broadly suggestive of certain bulent sediments from both sides of the North Atlantic. members of the Astropectinidae although abactinals are Betelgeusia is suggestive of Astropecten in marginal and more gracile in this family and marginals less uniform fasciolar development as well as in the arrangement of and tabulate. ambulacral-adambulacral muscle flanges, which were Genera included in this subfamily arePseudarchaster interpreted (Blake, 1981) as suited for twisting of arms, S l a d e n , 1 8 8 9 , and Paragonaster S l a d e n , 1 8 8 9 , reported useful in predation or righting behaviour. Given the lim from both fossil and Recent occurrences; Gephyreaster ited fossil record, Betelgeusia distribution in time and F is h e r , 1 9 1 0 , and Perissogonaster F is h e r , 1 9 1 3 , reported space suggests past success in shallow-water niches si only from Recent occurrences; andFormalhautia B l a k e milar to those occupied by Astropecten and Luidia today. & R e id , 1 9 9 8 , and Skiaster n. gen., known exclusively Blake (1987) assigned Radiaster to a basal position from fossil occurrences. within the Paxillosida, and its Cretaceous occurrence is consistent with this interpretation. D e s c r ip t io n Aldebarania taberna (Paxillosida; Astropectinidae) is Abactinal ossicles robust, paxilliform, more or less similar to the type species, A. arenitea Blake & Stur columnar. Crowns convex, rounded or elliptical; in larger geon, 1995from eastern North America, demonstrating ossicles, re-entrant present below crown, column distinct. occurrence on both sides of the then-smaller North Atlan Base enlarged, lobate, providing passageway for papulae. tic. Both Aldebarania species are of low profile and Arrangement such that about six abactinals cluster about constructed of stout primary ossicles with sturdy gran another abactinal, all aligned in rows. Carinal series ules, and share an overall form similar to those of certain enlarged. modern and Cretaceous species found in shallow, turbu Marginals tabular, moderately large, wide compared to lent sedimentary settings(Blake & Kues, 2002) that are length, closely appressed, deeply fasciolated, paired. at least similar to the Aldebarania occurrences. Other than in Skiaster n. gen., superomarginals rarely Cheiraster? sp. (Notomyotida) represents the family abutted across arm midline, and then only close to arm Benthopectinidae, which is today a largely deep-water tips. Outer surfaces of marginals arched; however, not family. In contrast,Cheiraster ? sp. and other fossil strongly so as to yield a bulbous appearance; outer benthopectinids have been collected from shallow-water surfaces not flattened. Lateral shoulders of marginals sediments (e.g., Blake, 1984; B lake & Reid, 1998; Jagt, rounded rather than angular, re-entrants beneath 2000b) and thus the Benthopectinidae is among families shoulders (above shoulders in inferomarginals) at most once but no longer common in shallow-water settings weakly developed. Superomarginals closely granulated; (B ottjer & Jablonski, 1988). Blake (1984)recognised inferomarginals of some species with several somewhat the Paleobenthopectininae for certain Jurassic species; enlarged spines whereas in other species, inferomarginals Cheiraster? sp., a member of the living Neobenthopecti- evenly granulated. ninae, helps to constrain the divergence of crown-group Adambulacrals robust, furrow margins angular, in life benthopectinids. providing partial separation between subsequent tube feet; angular margins serving to narrow furrow, espe cially distally. Furrow spines curved or divergent, sub- T axonom y ambulacral spines either in longitudinal rows or clustered. Actinal series numerous in many species; ossicles angu Terminological usage follows Spencer & W right (1966) andB l a k e & lar, arranged in columns parallel to the ambulacral fur H a g d o r n (2003). The following abbreviations are used to denote the rows. Specialized spinules but not true pedicellariae can repository of material referred to in the text: CAS - California Acad emy of Sciences, San Francisco; NHMM - Natuurhistorisch Museum be adjacent to ambulacrals; pedicellariae found only Maastricht (K - M.M.M. Kuypers Colin; MD - M.J.M. Deckers Colin); rarely on actinal, marginal ossicles ofParagonaster. 186 Daniel B. BLAKE & John W.M. JAGT

Mouth angle ossicles large, prominent, with elongate by small granules occur between abactinals, marginals, spinules both on ventral surfaces and at ossicular margins. and actinals of many goniasterids; these grooves typically In some species or individuals, an enlarged, unpaired are quite large and well defined in pseudarchasterines and median spine is present at proximal end of ossicles. smaller in other goniasterids. Accessories consist of clo sely spaced granules in pseudarchasterines, and spines D is c u s s io n can occur on inferomarginals. Among other genera, ossi Sladen (1889) assigned both Pseudarchaster and Aph cular surfaces bare of granules and even raised relative to roditaster Sladen, 1889,to the then-new Pseudarchaster the granulate area are present, and spines are lacking. inae, but he provided no diagnosis. Since that time, the Abactinals are paxilliform in pseudarchasterines concept has received scattered recognition in the litera whereas in other goniasterids they are more nearly tabu ture but little discussion. The most comprehensive avail lar, broader, and less uniform, although in some (e.g., able treatment of the subfamily is that ofV errill (1899). Mediaster), tabulae approach a paxilliform condition. V errill (1899) did not formally list assigned genera but The paxilliform dorsals of the pseudarchasterines are he included species of Pseudarchaster, Aphroditaster, separated by passageways near their dorsal ends whereas Paragonaster, and Rosaster Perrier, 1894, and he in abactinals of the other genera tend to be closely fitted and cluded the type species for all but Paragonaster, which hence of polygonal outline. was only subsequently designated byFisher (1919). Pre V e r r i l l (1 8 9 9 ) found pseudarchasterine abactinals sumablyV e r r i l l ’s approach was intended to encompass and marginals and their accessories to be similar to those genera rather than only the cited species. of the goniasterid Mediaster and its allies but differing in The above description was largely rephrased from the angular furrow margins of the adambulacrals, the lack V errill (1899), although it has not been comprehen of bivalved pedicellariae, and the divergent furrow spi sively checked against component species, and it might nules. Fie noted the presence of fascioles, spines on the need to be altered further. Marginal and abactinal form actinals and inferomarginals, and the unpaired oral spine are given greater emphasis here than that accorded by provide further distinguishing characters. V e r r i l l . Because pseudarchasterine ancestry and a com The recommended synonymy Aphroditaster of with prehensive phylogeny are not available, apomorphic Pseudarchaster (M ortensen, 1 9 2 7 ) is accepted here, characters cannot now be recognised. and Rosaster is not included because its marginal and Spencer & W right (1966) also recognised the sub abactinal morphology are not pseudarchasterine. Para family, and they includedParagonaster, Perissogona gonaster and Perissogonaster are similar to Pseudarcha ster, Pseudarchaster and Aphroditaster. Aphroditaster ster, and the difference between Perissogonaster and was considered to be a synonym ofPseudarchaster by Paragonaster is limited to presence of an odd interbra- M ortensen (1927), an interpretation subsequently chial marginal inPerissogonaster. Formalhautia B l a k e embraced by A.M. C lark (1993) and followed here. & R e id , 1 9 9 8 , shares the essential marginal and abactinal Gephyreaster also is included here. form of Pseudarchaster, as does Skiaster n. gen., insofar Marginal and abactinal form indicate alignment of as can be determined. Skiaster n. gen. withPseudarchaster whereas the super omarginals, abutted across the midline of the arm, might Genus Skiaster n. gen. suggest affinities with any of the numerous non-pseudarchasterine goniasterids similarly developed. Non- T y p e s p e c ie s pseudarchasterine goniasterid genera with abutted super Skiaster vikingr n. sp., by monotypy. omarginals were surveyed here, and their marginal and abactinal form found to be distinct from that of the D e r iv a t io n o f n a m e Pseudarchasterinae. In the pseudarchasterines, marginals From Gr. skia, meaning shadow. Superomarginals are tabular (their width, or dimension normal to the arm abutted across the midline of the arms shadows develop margin, is significantly greater than their length) whereas ment in typical goniasterid genera, but presence in a in the other genera, marginals are proportionately nar pseudarchasterine indicates a separate lineage and there rower, even becoming square in dorsal (or ventral, for by a shadowing of the pattern evolved in other goni inferomarginals) outline. The outer face of the marginal is asterids. approximately parallel to the overall dorsal (or ventral) surface near the abactinals (or actinals), and it curves to D ia g n o s is perpendicular to the body plane at the ambitus, where the Monospecific, see below. surface joins that of the paired marginal. In pseudarchas terines, curvature is approximately even, typically yield Skiaster vikingr n. gen., n. sp. ing a rounded shoulder, whereas in the other genera, the (PI. 1, Figs. 1, 2 ; PI. 2 , Figs. 1 -8 ) shoulder tends to be quite abrupt, in many genera with a re-entrant at the juncture of the superomarginals and T y p e inferomarginals. Fascioles between subsequent infero Holotype and only known specimen is NHMM 2 0K 8 6 , marginals are present in pseudarchasterines but absent from the Lower Danian Bryozoan Limestone(Tylocidaris from the other genera. Narrow, sharp grooves bordered abildgaardi Zone), north of Hojerup Kirke, Stevns Klint New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 187

(Sjælland, Denmark). Specimen incomplete, exposed in No other ossicular type clearly exposed. dorsal view, consisting of the disc and one incomplete arm. Radius of the arm remnant approximately 17 mm, R e m a r k s interbrachia 10 and 11 mm. The marginal frame is par Body shape, including the pentagonal disc and apparently tially collapsed and ossicles are displaced; the dorsal elongate arms with marginals abutted along the arm mid surface is collapsed into the disk displacing abactinal line, is indicative of the Goniasteridae. Both arm shape ossicles. Recrystallization partially obscures ossicular de and the conformable fit of the marginals along the arm tail, and dorsal ossicles are further obscured by remnants midline indicate that the ossicles were abutted in life of accessory ossicles. rather than collapsed together taphonomically. Abactinals are similar to those of Pseudarchaster or perhaps D e r iv a t io n o f n a m e Mediaster (Goniasteridae) but not greatly unlike those From Old Norse vikingr, or Norseman, in allusion to its of e.g., Tethyaster, Blakiaster P e r r i e r , 1881, and Pati- geographic provenance; grammar: noun, gender; mascu gaster F is h e r , 1906 (Astropectinidae). line. Skiaster n. gen. is the only pseudarchasterine with marginals abutted at the dorsal arm midline. Among D ia g n o s is goniasterids with marginals abutted at the midline, it is Pseudarchasterine with pentagonal disc and supero suggestive of Nymphaster S l a d e n , 1889 (see PI. 1, Figs. 4, marginals abutted at dorsal arm midlines, the abutted 6 here), from which it differs in shape of both marginals arrangement extending from the proximal end of the and abactinals. Probably for both taphonomic and taxo ann. Abactinals paxilliform, slender and columnar to ro nomic reasons, abactinal ossicles have not been well bust with a flaring base and short re-entrant below crown. illustrated in the literature, but among the examples avail able to the authors, the abactinal morphology of the D e s c r ip t io n Cretaceous (late Campanian) species Nymphaster Five-armed asteroid, disc proportionately large, penta studlandensis (Schulz & W eitschat, 1975) is most gonal. Arms slender, robust, length unknown. suggestive of that of the pseudarchasterines in that these Abactinals paxilliform, up to approximately 1 mm in ossicles are both tali and slender and thereby suggestive height, 1 mm in diameter at base. Crown hemispherical, of paxillae (see PI. 1, Figs. 9-11). In N. studlandensis, column short, stout, situated medially on base; base con however, abactinals are closely fitted and columnar, with ical, circular or equidimensional and polygonal in outline subdued papular pores and deep lateral depressions (for with facets for papulae on some ossicles. Enlarged abact connective tissues?), which are characteristic of non- inals (e.g., primary circlet, carinal series) not detected on pseudarchasterine goniasterid abactinals in spite of their largely disrupted and debris-obscured surface. paxillae-like height. Among non-pseudarchasterines, Inferomarginals incompletely exposed, but marginals Skiaster n. gen. and other pseudarchasterines are quite of two series generally similar insofar as can be ascer similar in marginal and abactinal shape to such genera as tained. Marginals tabular, width of interbrachial super- Mediaster (see PI. 1, Figs. 5, 8), from which they differ in omarginal about 2 mm, length about 1 mm, height about a number of characters as outline byV e r r i l l (1899; see 1.5 mm; superomarginals at arm base of width about 3 above). Specifically, Mediaster lacks abutted supero mm, length 1.5 mm. Dorsally directed portion of super marginals. omarginals quite wide, then evenly rounded approaching Attempts to further expose the ventral skeleton were lateral side faces (i.e., ossicular transverse section not unsuccessful due to preservational constraints. angular). Longitudinal profile of adradial part of interbrachial superomarginals rounded. Adjacent superomar Order Paxillosida P e r r i e r , 1884 ginals sharing very regular, straight-sided boundary fur Family Radiasteridae F is h e r , 1916 row approximately 0.2 mm wide. Outer face finely and Genus Betelgeusia Blake & Reid, 1998 evenly pitted, pits clearly separated, some ossicular deb ris suggests an even covering of granules; no indication of T y p e s p e c ie s differential sizes of pits to indicate differentiation of Betelgeusia reidi Blake & Reid, 1998, by original des accessories. Interossicular grooves smooth. Side faces ignation. comparatively large, bordered by a low, rounded ridge, remainder of surfaces slightly sunken, flattened medially. Marginals from adjacent sides of arm abutted at arm Betelgeusia exposita n. sp. midline beginning at disc. Adradial-ventral comer of (PI. 3, Figs. 1-5) superomarginal and adradial-dorsal comer of inferomarginal bearing a rounded notch that would have formed a v. 2000b astropectinid sp. nov. J a g t , p. 387 (partim), pi. 5, broad interior groove in the living individual. Intermar figs. 1-5. ginal face of superomarginal apparently concave, that of inferomarginal flattened. Inferomarginals probably T y p e fasciolate, ossicular debris suggests presence of enlarged Holotype and only known specimen is NHMM K 3364, spinelets. from the higher Meerssen Member (Maastricht Forma- 188 Daniel B. BLAKE & John W.M. JAGT tion), upper Upper Maastrichtian [Belenniella (Neobe- surfaces. Ventral surface of inferomarginals and dorsal lemnella) kazimiroviensis Zone], ENCI-Maastricht BV surfaces of inferomarginals and superomarginals bearing quarry, Maastricht. The incomplete specimen is exposed apparently uniform, closely spaced cylindrical spinelets in ventral view. It consists of about one-half of the disc, about 0.3 mm in length, similar to but perhaps a bit more one complete arm, and part of a second. Arm radii robust than paxillary spinelets. Inferomarginals with a approximately 15 mm, interbrachial radius about 8 mm. dense, marginal, laterally directed fringe of somewhat A fragment from the proximal right side of the incom more robust spinelets approximately 0.5 mm in length. plete arm is separate from the remainder of the specimen. Spinelets circular or perhaps somewhat flattened in cross The fragment is about 6 mm in length and includes 7 section. marginal pairs; both dorsal and ventral surfaces are ex Actinals arranged in distinct radial series separated by posed. The specimen was preserved largely undistorted; well-developed fasciolar grooves; 19 to 22 series of however, recrystallization and some accessories partially actinals correspond to about 12 proximalmost inferomar obscure detail. ginals; beyond that interval, inferomarginals abut adambulacrals. Actinals poorly exposed; ossicles probably D e r iv a t io n o f n a m e massive, nearly 15 ossicles present in longer series near From Latin expositus, -a, because the specimen was interbrachia, becoming reduced to a single ossicle at found on the moss-covered bedding plane of a well- series termination. exposed slab of rock at the ENCI-Maastricht bv quarry. Ambulacral-adambulacral series approximately 33 to 35 between mouth angle ossicle and terminal; distalmost D ia g n o s is 3 or 4 ossicular pair extremely tiny, partially enclosed by Betelgeusia with a rounded body margin that largely the much-enlarged terminal. Ambulacrals very similar to results from a rounded margin to the superomarginals. those of Astropecten, with a nearly square adradial inter Actinal and inferomarginal series not corresponding; val bearing robust cross-furrow articular structures. mouth angle pair robust, forming a keel-shaped, broad, Ossicles robust, with lateral (rather than overlapping) angular surface. Prominent ventral spines not recognised. longitudinal articular surfaces. Podial pores large. Ambu lacral-adambulacral articulation flanges asymmetrical, D e s c r ip t io n the proximal offset adradially and hook-shaped; distal Arm spacing indicating a five-armed asteroid; disc of flange J-shaped and prolonged abradially. moderate size; interbrachia rounded but not broad; arms Adambulacrals incompletely exposed, upright, rectan triangular, broad at base, straight-sided, tapering evenly. gular in outline, transversely elongate with long axes Abactinals paxilliform, delicate, with a low, expanded inclined to the arm axis. Like inferomarginals, abradial base; column slender, grading evenly into an expanded ends of ossicles inclined distally. Proximalmost adambu- crown; crown not differentiated from column by distinct lacral somewhat enlarged relative to subsequent ossicles, re-entrant. Paxillae small, about four in one mm in trans its adradial side abutting mouth angle ossicles. No differ verse series on proximal arm fragment. Paxillae aligned entiation of furrow series spine bases from subambulacral in transverse and probably inclined longitudinal series. bases can be recognised; any differentiation must have Crown bearing cluster of small, cylindrical spinelets. No been comparatively minor. Remaining spines few, differentiation of abactinals recognised on available very essentially similar to other ventral-surface spines: short, small fragment of dorsal surface, which does not include cylindrical, simple. either mid-arm interval (that would reveal possible car Mouth angle ossicles robust, broad, keel-like, outline ináis) nor central disc (that would reveal a differentiated of pair rounded rectangular to broadly elliptical, adjacent primary circlet). Madreporite not exposed. Terminal pairs not closely abutted in life. Furrow sides bearing robust, square, prominent. uniform pustules extending well up sides of ossicle. Marginals paired, approximately 16 between midline Dorsal articular flange apparently large, broad, blade of interbrachia and terminal; more proximal marginals like, extended strongly towards oral-aboral axis. Circu- radially arranged, more distally becoming inclined to arm moral (i.e., first ambulacral) similar to ambulacrals in axis. Marginals of two series similar but inferomarginals ventral view, but more robust. Odontophore partially larger than superomarginals; proximal inferomarginal visible between mouth angle ossicles; ossicle robust, approximately 1.25 mm in width, corresponding super closely fitted between MAO. omarginal approximately 0.75 mm, superomarginal set back from arm margin such that a portion of inferomar R e m a r k s ginal is visible in dorsal view. Marginals uniformly tab Recrystallization and the sandy carbonate sediment ren ular but with prominent lateral articular ridges (or perhaps der determination of detail under about 1 mm difficult. disjunct articular facets); fasciolar groove large. Articular The new species is similar to the type species,Betelgeu ridges set back from abradial margin of both marginal sia reidi, in all essential aspects, differing only in com series forming a well-defined fasciolar abradial notch on paratively minor details. The margin of the body was margins of arms. Fascioles parallel sided, aligned or more angular in B. reidi, with the superomarginal cuneate slightly offset from corresponding grooves separating in outline (Blake & Reid, 1998, figs. 8.6, 8.7, 8.15), adjacent actinal and abactinal series on ventral and dorsal whereas the margin of B. exposita n. sp. is more rounded. New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 189

Actinal series approximately correspond in number with Two disc fragments, NHMM MB 377-23/a, b, both inferomarginals in the type species, but series are more from the Meerssen Member (base IVf-3; Maastricht For numerous in the new species. The mouth angle pair is mation) at Blom quarry (Berg en Terblijt), might belong more robust, rounded, and keel-like in the new species. to the new species, but granular preservation does not The ventral surface of the single specimen of the new allow unequivocal assessment. Although the fragments species has been almost entirely denuded of spines, but are included here they are not considered a part of the those remaining do not suggest presence of the elongate hypodigm on which the species is based and they are not spines of the type, and the dorsal ossicles appear to be specifically encompassed in the description below, more delicate in the new species. The proportionately although they are not known to be inconsistent with this greater width of the marginals of the type species might description. be a function of specimen size. Betelgeusia exposita n. sp. appears similar in form to D e r iv a t io n o f n a m e Dipsacaster jadeti B r e t o n et al., 1995, from the Maas From Latin taberna, -a, hut, inn (grammar: noun, gender: trichtian of Petites-Pyrénées (France). The latter species feminine); the species name, a play on words, is in honour is also known from a single specimen, unfortunately (for of Mr Ludo Indeherberge of Zonhoven, Belgium, who comparison purposes) with the dorsal rather than the collected these (and other) important fossils and has made ventral surface exposed. Dipsacaster (Astropectinidae) them available for study. is similar to Betelgeusia in overall form but the two species of concern here differ in some details. The mar ginal ossicles of D. jadeti (the inferomarginals only in G e n e r ic a n d f a m il ia l a s s ig n m e n t completely exposed) appear robust in comparison with The new species shares all of the diagnostic characters of those of Betelgeusia exposita n. sp. The somewhat deli Aldebarania B l a k e & S t u r g e o n , 1995, including stellate cate ambulacral ossicles (largely obscured by abactinals) overall shape and rectangular marginals with deep fas and the delicate, aligned series of apparently paxilliform cioles. B l a k e & S t u r g e o n (1995) discussed ordinal and abactinals both appear similar to those of Betelgeusia, familial assignment of the genus, which is not repeated and reflect the difficulty of taxonomic assessment of here. scanty and incompletely exposed asteroid material. Un fortunately, many asteroid taxa are known only from such D ia g n o s is material. Paxillae low, stout, with a subpolygonal crown; paxillae arranged in regular series. Accessories consist of closely Family AstropectinidaeG r a y , 1840 fitted short spinelets. Superomarginals weakly inset from Genus Aldebarania B l a k e & S t u r g e o n , 1995 edge of inferomarginals. Medial, raised ridge on infero marginals lacking. T y p e s p e c ie s Aldebarania arenitea B l a k e & S t u r g e o n , 1995, by ori ginal designation. D e s c r ip t io n Similar to the type and only other known species,A. Aldebarania taberna n. sp. arenitea B l a k e & S t u r g e o n , 1995, in form and paxillary (PI. 4, Figs. 1-6) arrangement. Paxillae uniform over surface except those adjacent to interbrachial superomarginals and distally on v. 2000b Aldebarania sp. nov. Jagt, p. 392, pi. 4, figs. 8- arm are progressively reduced in size; no indication of 10; pi. 5, figs. 6-12 (?pl. 7, figs. 5, 6). enlarged carinals nor primary circlet in limited available material. Paxillae stout, low, some at least with papulary T y p e s facets; crowns subpolygonal forming tightly fitted dorsal The five specimens included in the hypodigm are all from surface. Ossicles arranged in intersecting radial, long the Nekum Member (Maastricht Formation); the holotype itudinal series with one or two series reaching terminal. (NHMM 1999015-1) and paratypes (NHMM 1999015-2, Madreporite not located. Terminal enlarged relative to -3, NHMM JJ 2787) come from the CBR-Romontbos adjacent ossicles, elongate, outline polygonal with distal quarry. All three NHMM 1999015 specimens are external tip somewhat rounded, surface rounded, covered by short moulds in flint preserving both surfaces and all were spinelets. buried nearly intact with only limited ossicular dis Marginals similar to those of A. arenitea in form and ruption, although arm tips have been lost. In NHMM arrangement. Superomarginals weakly inset from edge of 1999015-1, R of most complete arm is 22 mm with a superomarginals, no raised ridges present on inferomar few mm missing; r = 13 mm; in NHMM 1999015-2, ginals. Inferomarginal fringe spinelets elongate, spinelets R = 14 mm, r = 6 mm, and in NHMM 1999015-3, of ventral surface uniform. R = 6 mm, r = 4 mm. NHMM JJ 2787 is a single well- Actinal fields collapsed and obscured taphonomically, preserved distal superomarginal and paratype RGM 428 but similar to those of A. arenitea insofar as can be 072 (from the ENCI-Maastricht BV quarry) is a well- ascertained except actinal ossicle articular flanges rela preserved medial superomarginal. tively subdued. Mouth angle ossicular pair prominent, 190 Daniel B. BLAKE & John W.M. JAGT broadly rounded. MAO accessories including scattered Cheiraster? sp. spinelets but preservation incomplete; other mouth frame (PI. 5, Figs. 1-7) characters unavailable. Ambulacrals, adambulacrals very incompletely exposed, form typical of astropectinids and M a t e r ia l similar to those of A. arenitea. Ambulacrals nearly The sole known specimen is NHMM MD 4105 from the bilateral with robust cross-furrow articular structures; basal Gronsveld Member (Maastricht Formation), Upper adambulacrals upright, strongly overlapping, with furrow Maastrichtian, ENCI-Maastricht bv quarry. The speci series of curved, flattened spines; subambulacrals men, exposed in dorsal view, consists of most of the straight, conical, pointed, similar to those of the furrow ventral portion of the disc and proximal intervals of three series. arms. Arm remnant radii approximately 18, 17 and 15 mm, interbrachia 4 and 5 mm. Dorsal disc ossicles have R e m a r k s been nearly lost, and ossicles of the arms have suffered Differentiation of the two known species is subtle in part some displacement, although overall arm form is re because of similarities, in part because of limited material tained. Recrystallization and some accessories partially available for each, and in part due to taphonomic disrup obscure ossicular detail. tion. The two species differ in the form of the inferomar ginals, and more subtly, in the nature of the abactinals. D e s c r ip t io n Arm spacing indicating a five-armed asteroid; disc small; Order Notomyotida L u d w ig , 1910 arms slender, columnar; length unknown. A few small Family BenthopectinidaeV e r r i l l , 1899 (approx. 0.5 mm), conical abactinals remain on the disc Subfamily NeobenthopectininaeB l a k e , 1984 and proximal intervals of the arms; these are too few to determine original numbers or arrangement. Madreporite R e m a r k s not recognised. The rectangular to subelliptical form of the marginals, Marginals in a double series; probably slightly offset, with a bulbous outer face and a single spine base, the elliptical in outline, elongate parallel to arm axis, outer small, conical abactinals, and the rectangular form of face bulbous (arched), inferomarginals overlapping, ambulacral ossicles are typical of modern benthopecti superomarginals abutting, intermarginal faces flat. Outer nids ( B l a k e , 1984), indicating affinities with the Neo face bearing one(?) enlarged, circular spine base, remain benthopectininae of the Benthopectinidae. der of face bearing small, separated pustules. Ambulacrals preserved in approximate life positions, Genus Cheiraster S t u d e r , 1883 with dorsal and adradial portions exposed but ventral and abradial portions largely obscured. Ambulacrals robust, R e m a r k s length of proximal ossicles approximately 1 mm, width Characterisation of modem benthopectinid genera approximately 2 mm. Dorsal outline hourglass shaped, (Clark & Downey, 1992) stresses characters not readily with large ampullar basins. Ossicles approximately bilat available in fossil material in general and in the present eral (although incompletely exposed and some apparently specimen in particular. The fossil has robust, blocky truncated taphonomically). Adambulacral articular marginals; among modern benthopectinid genera,C l a r k flanges large, broad. Adradial part of ossicle large, robust, & D o w n e y (1992) identified three with similar enlarged lateral ambulacral-ambulacral articular surfaces robust, marginals, Cheiraster, Acontiaster Döderlein, 1921 and abutted laterally (rather than overlapping), cross-furrow Pectinaster P e r r i e r , 1885. Unfortunately, blocky marginarticular structures robust. Dorsal ridge well defined als perhaps are plesiomorphic at the familial level be medially but both adradial and abradial ends of ossicle cause they are suggestive of marginals of other valvata- flattened. Other ossicular types not clearly exposed. cean families and unlike the distinctive flatter marginals with prominent spine bases that typify most benthopecti R e m a r k s nid genera. The enlarged marginals of this specimen are similar to those Species assignments among Pontaster, Pectinaster of Cheiraster echinulatus (see PI. 5, Fig. 8), which, like the and Cheiraster have been uncertain (see C l a r k , 1989), fossil, has comparatively small marginal spine bases; debris testifying to the uncertainty of the generic concepts. associated with the fossil also suggests small spines. Presence of prominent spines serve to separateAcontia Ambulacral ossicles in the fossil are typical of neo- ster. Similtarities between the fossil and Cheiraster echi benthopectinids in general; unfortunately, these ossicles nulatus ( P e r r i e r , 1875) suggest affinities (see below) are not well enough known among benthopectinid genera with that genus, and C. echinulatus is the type species to be used in identification. of the subgenus Barbadosaster C l a r k , 1981, indicating correct assignment of C. echinulatus at the generic level. Final taxonomic assessment of the present specimen, Acknowledgements however, must await re-evaluation of modem species with more comprehensive treatment of their ossicular We thank M.J.M. Deckers, L. Indeherberge and M.M.M. Kuypers for morphology. donation of material, D.L. Pawson, Cynthia Ahem and authorities of New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 191

the U.S. National Museum (Washington DC) who made facilities fully ments of CBR and ENCI-Maastricht bv (Heidelberg Cementgroup) for available and allowed access to collections in their care, L. Villier for permission to do fieldwork at their quarries over recent years, commenting on an earlier version of the typescript, and the manage-

References

B la in ville , H.M. de , 1830. Zoophytes. Dictionnaire des triel de la Société géologique du Normandie et Amis du Mu Sciences Naturelles, F.G. Levrault, Strasbourg, 60 p. séum du Havre, 82: 35-42.

B la k e , D.B., 1981. The new Jurassic sea star Eokainaster and B r o m l e y , R.G., 1979. Chalk and bryozoan limestone: facies, comments on life habits and the origins of the modem Aster sediments and depositional environments.In\ B ir k e l u n d , T. & oidea. Journal o f Paleontology, 55: 33-46. B r o m l e y , R.G. (Editors), Cretaceous-Tertiary Boundary Events Symposium, I. The Maastrichtian and Danian of Den B la k e , D .B ., 1984. The Benthopectinidae (Asteroidea: Echi nodermata) of the Jurassic of Switzerland. Eclogae geologicae mark. University of Copenhagen, pp. 16-32. Helvetiae, 77: 631-647. B r ü n n ic h N ie l s e n , K., 1943. The asteroids of the Senonian and Danian deposits of Denmark.Biologiske Skrifter fra det danske B la k e , D.B., 1986. Some new post-Paleozoic sea stars (Aster oidea: Echinodermata) and comments on taxon endurance. Videnskabernes Selskab, (2)5: 3-68. Journal o f Paleontology, 60: 1103-1119. C l a r k , A.M., 1989. Notes on Atlantic and other Asteroidea, 1. Family Benthopectinidae.Bulletin of the British Museum of B la k e , D .B ., 1987. A classification and phylogeny of post- Palaeozoic sea stars (Asteroidea: Echinodermata). Journal o f Natural History (Zoology), 41: 91-135. Natural History, 21: 481-528. C l a r k , A.M., 1989. An index of names of Recent Asteroidea - J a n g o u x , B l a k e , D.B., 1988. A first fossil member of the Ctenodiscidae Part 1: Paxillosida and Notomyotida. In: M. & (Asteroidea; Echinodermata). Journal of Paleontology, 62: L a w r e n c e , J.M. (Editors), Echinoderm Studies 3. A.A. Balk- 626-631. ema, Rotterdam, pp. 225-349. C l a r k , A.M., 1993. An index of names of Recent Asteroidea - B la k e , D.B., 1990a. Hettangian Asteriidae (Echinodermata: Asteroidea) from southern Germany: taxonomy, phylogenyPart 2: Valvatida. In: J a n g o u x , M. & L a w r e n c e , J.M. (Edi and life habits. Paläontologische Zeitschrift, 64: 103-123. tors), Echinoderm Studies 4. A.A. Balkema, Rotterdam, pp. 187-366. B la k e , D.B., 1990b. Adaptive Zones of the Class Asteroidea (Echinodermata). Bulletin o f Marine Science, 46: 701-718. C l a r k , A.M. & D o w n e y , M.E.D., 1992. Starfishes of the Atlantic. Chapman & Haii, London, 794 pp. B la k e , D.B. & H a gdorn , H., 2003. The Asteroidea (Echino dermata) of the Muschelkalk (Triassic of Germany). Paläonto D ö d e r l e in , L., 1921. Die Asteriden der Siboga-Expedition i: logische Zeitschrift, 77: 23-58. Porcellanasteridae, Astropectinidae, Benthopectinidae. Siboga- Expeditie Monograph, 46(1): 1-47. B la k e , D.B. & H o tch kiss , F.R.C., 2004. Recognition of the asteroid (Echinodermata) crown group: implications of the D ü b e n , M.W. & K o r e n , J., 1846. Öfversigt af Skandinaviens ventral skeleton. Journal o f Paleontology, 78: 359-370. Echinodermer. Kungliga Svenska Vetenskapsakademiens Handlingar, 1844: 229-238. B la k e , D.B. & K u es , B.S., 2002. Homeomorphy in the Aster oidea (Echinodennata); a new Late Cretaceous genus and spe F is h e r , W.K., 1906. The starfishes of the Hawaiian Islands. cies from Colorado. Journal o f Paleontology, 76: 1007-1013. Bulletin o f the United States Fish Commission, 23: 987-1130. B lake, D.B. & Reid, R., III., 1998. Some Albian (Cretaceous) F is h e r , W.K., 1910. New genera of starfishes. Annals and asteroids (Echinodermata) from Texas and their paleobiological Magazine o f Natural History, 5: 171-173. Implications. Journal o f Paleontology, 72: 512-532. F is h e r , W.K., 1913. Four new genera and fifty-eight new species of starfishes from the Philippine Islands, Celebes and B la k e , D.B. & Stu rg eo n , K., 1995.Aldebarania arenitea, a new genus and species of Astropectinidae (Asteroidea; Echi the Moluccas. Proceedings o f the United States National Mu nodermata) from the Maastrichtian (Upper Cretaceous) Peedee seum, 46: 599-648. Formation o f North Carolina. Journal o f Paleontology, 69: 376- F is h e r , W.K., 1916. Notes on the systematic position of certain 380. genera and higher groups of starfishes. Proceedings of the Biological Society o f Washington, 29: 1-6. B ottjer , D.J. & J a blonski , D., 1988. Paleoenvironmental patterns in the evolution of post-Paleozoic benthic marine in F is h e r , W.K., 1919. Starfishes of the Philippine Seas and vertebrates. Palaios, 3: 540-560. adjacent waters. Bulletin o f the United States National Museum,

B reton , G., 1979. Les astéries du Crétacé de Normandie. 100: 1-712. Bulletin trimestriel de la Société géologique de Normandie Feto r b e s , E., 1841. A history of British starfish and other animals Amis du Muséum du Havre, 65: 5-85. of the class Echinodermata. John Van Voorst, London, 162 pp.

B r e t o n , G., 1992. Les Goniasteridae (Asteroidea, Echinoder G a l e , A.S., 1987a. Goniasteridae (Asteroidea, Echinodermata) mata) jurassiques et crétacés de France: taphonomie, systéma from the Late Cretaceous of north-west Europe. Part 1. Intro tique, biostratigraphie, paléobiogéographie, évolution. Bulletin duction. The genera Metopaster and Recurvaster. Mesozoic trimestriel de la Société géologique de Normandie et Amis duResearch, 1: 1-69.

Muséum du Havre, Hors-série, Supplément 78: 1-590. G a l e , A.S., 1987b. Goniasteridae (Asteroidea, Echinodermata) B reton , G., B ilo tte , M. & S ig ro , G., 1995.Dipsacaster jadeti from the Late Cretaceous of north-west Europe. 2. The genera sp. nov., Astropectinidae (Asteroidea, Echinodermata) du Calliderma, Crateraster, Nymphaster and Chomataster. Meso Maastrichtien des Petites-Pyrénées (France).Bulletin trimes zoic Research, 1: 151-186. 192 Daniel B. BLAKE & John W.M. JAGT

G ra vesen , P., 1993 Early Danian species of the echinoid genus P e r r ie r , E., 1884. Mémoire sur les étoiles de mer recueillis Tylocidaris (Cidaridae, Psychocidarinae) from eastern Den dans la Mer des Antilles et le Golfe de Mexique. Nouvelles mark. Contributions to Tertiary and Quaternary Geology, 30: Archives du Muséum d'Histoire Naturelle Paris, 6: 127-276. 41-73. P e r r ie r , E., 1885. Première note préliminaire sur les échino- G r a y, J.E., 1840. A synopsis of the genera and species of the dermes recueillis durant les campagnes de dragages sous-mar- class Hypostoma Asterias( Linnaeus).Annals and Magazine of ins du Travailleur et du Talisman. Annales des Sciences natur Natural History, 20: 193-204. elles, (6)19: 1-72.

H e s s , H., 1960. Pentasteria (Archastropecten) procera n. sp. P e r r ie r , E., 1894. Expéditions scientifiques du Travailleur et (Asteroidea, Astropectinidae) aus dem Bajocien von Chelten du Talisman: échinodermes. G. Masson, Paris, 431 pp. ham (England). Eclogae geologicae Helvetiae, 53: 331-334. S c h ö n d o r f , F., 1910. Die Asteriden der deutschen Trias. H e s s , H., 1972. Eine Echinodermen-Fauna aus dem mittleren Jahresbericht des Niedersächsischen Geologischen Vereins, Dogger des Aargauer Juras. Schweizerische Paläontologische 1910: 90-115. Abhandlungen, 92: 1-87. S c h u l z , M.-G. & W e it s c h a t , W., 1971. Asteroideen aus der Hess, H., 1973. Neue Echinodermenfunde aus dem mittleren Schreibkreide von Lägerdorf (Holstein) und Hemmoor (Nord- Dogger des Aargauer Juras. Eclogae geologicae Helvetiae, 66: Niedersachsen). Mitteilungen aus dem geologisch-paläontolo- 625-656. gischen Institut der Universität Hamburg, 40: 107-130. H e s s , H., 1975. Die fossilen Echinodermen des Schweizer S c h u l z , M.-G. & W e it s c h a t , W., 1975. Phylogenie und Strati Juras. Veröffentlichungen aus dem Naturhistorischen Museum graphie der Asteroideen der nordwestdeutschen Schreibkreide. Basel, 8, 1-130. Teil I: Metopaster/Recurvaster- und Calliderma/Chomataster- H e s s , H., 1987. Neue Seestemfunde aus dem Dogger des Gruppe. Mitteilungen aus dem geologisch-paläontologischen Schweizer Juras. Eclogae geologicae Helvetiae, 80: 907-918. Institut der Universität Hamburg, 44: 249-284.

H e s s , H. & B l a k e , D.B., 1995.Coulonia platyspina n. sp., a S c h u l z , M.-G. & W e it s c h a t , W., 1981. Phylogenie und Strati new astropectinid sea star from the Lower Cretaceous of graphie der Asteroideen der nordwestdeutschen Schreibkreide. Morocco. Eclogae geologicae Helvetiae, 88: 777-788. Teil II: Crateraster/Teichaster-Gruppe und Gattung O p h ia Jagt, J.W.M., 1999a. Late Cretaceous-Early Paleogene echi- ster. Mitteilungen aus dem geologisch-paläontologischen noderms and the K/T boundary in the southeast Netherlands and Institut der Universität Hamburg, 51: 27-42. northeast Belgium - Part 1: Introduction and stratigraphy. S l a d e n , W.P., 1885. Asteroidea. In: W y v il l e T h o m s o n , C. & Scripta Geológica, 116: 1-57. M u r r a y , J., Report on the Scientific Results of the voyage of Jagt, J.W.M., 1999b. Late Cretaceous-Early Palaeogene echi H.M.S. Challenger during the years 1873-76, Narrative 1(2): noderma and the K/T boundary in the southeast Netherlands and 607-617. northeast Belgium - Part 2: Crinoids. Scripta Geológica, 116: S l a d e n , W.P., 1889. The Asteroidea. Report of the Scientific 59-255. Results, Voyage o f the Challenger (Zoology), 30: 1-935. J a g t , J.W.M., 2000a. Late Cretaceous-Early Palaeogene echi- S l a d e n , W.P., 1891. A monograph on the British fossil Echi noderms and the K/T boundary in the southeast Netherlands and nodermata from the Cretaceous formations, Volume second. northeast Belgium - Part 3: Ophiuroids, with a chapter on: The Asteroidea, Part First. Monograph o f the Palaeontographi- Early Maastrichtian ophiuroids from Rügen (northeast Ger cal Society o f London, 1891: ii + 1-28. many) and Mon (Denmark) by Manfred Kutscher and John W.M. Jagt. Scripta Geológica, 121: 1-179. S l a d e n , W.P., 1893. A monograph on the British fossil Echi nodermata from the Cretaceous formations, Volume Second. Jagt, J.W.M., 2000b. Late Cretaceous-Early Palaeogene echi- The Asteroidea, Part Second. Monograph o f the Palaeontogra- noderms and the K/T boundary in the southeast Netherlands and phical Society o f London, 1893: 29-66. northeast Belgium - Part 5: Asteroids. Scripta Geológica, 121: 181-375. S p e n c e r , W.K., 1905. A monograph on the British fossil Echi nodermata from the Cretaceous formations, Volume second. K u t s c h e r , M. & R ö p e r , M., 1999. Die Asteroidea (Seesteme) aus den “ Papierschiefem” von Hienheim und Ried (Malm The Asteroidea, Part Third. Monograph o f the palaeontogra- Zeta 3, Untertithonium). Archaeopteryx, 17: 1-13. phical Society o f London, 1905: 67-90. S p e n c e r , L ieba u , A., 1978. Paläobathymetrische und paläoklimatische W.K., 1907. A monograph on the British fossil Echi Veränderungen im Mikrofaunenbild der Maastrichter Tuff- nodermata from the Cretaceous formations, Volume second. kreide. Neues Jahrbuch für Geologie und Paläontologie TheAb Asteroidea and Ophiuroidea, Part Fourth. Monograph of handlungen, 157: 233-237. the Palaeontographical Society o f London, 1907: 91-132.

L u d w ig , H., 1910. Notomyota, eine neue Ordnung der See S p e n c e r , W.K., 1908. A monograph on the British fossil Echi steme. Sitzungsberichte der Königlich Preussischen Akademie nodermata from the Cretaceous formations, Volume second. der Wissenschaften, 23: 435-466. The Asteroidea and Ophiuroidea, Part Fifth. Monograph o f the Palaeontographical Society o f London, 1908: 133-138. M o r t e n s e n , T., 1927. Echinoderms of the British Isles. Oxford University Press, Oxford, 269 pp. S p e n c e r , W.K., 1913. The evolution of the Cretaceous Aster oidea. Philosophical Transactions o f the Royal Society o f Lon P e r r ie r , E., 1875-1876. Révision de la collection de stellérides du Muséum d’Histoire Naturelle de Paris. Archives de Zoologie don, B214: 99-177. expérimentale et générale, 4(1875): 265-540; 5(1876): 1-104, S p e n c e r , W.K. & W r ig h t , C.W., 1966. Asterozoans.In: M o o r e , 209-309. R.C. (Editor), Treatise on Invertebrate Paleontology, Part U, Echinodermata 3. Geological Society of America and University P e rrier, E., 1881. Description sommaire des espèces nouvelles d’astéries. Bulletin o f the Museum o f Comparative Zoology, 9: of Kansas Press, Boulder and Lawrence, pp. U4-U107. 1-31. S t im p s o n , W., 1857. On the Crustacea and Echinodermata of New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 193

the Pacific shores of North America. Boston Journal o f natural W r ig h t , T., 1863-1880. British Fossil Echinodermata of History, 6: 444-532. the Oolitic formations: vol. 2, Asteroidea and Ophiuroidea. S tu d e r , T., 1883. Bericht über die Asteriden welche während Monograph of the Palaeontographical Society of London, der Reise SMS ‘Gazelle’ um die Erde gesammelt wurden. 1861(1863): 1-130; 1864(1866): 131-154; 1880(1880): 155- Sitzungsberichte der Gesellschaft Naturforschender Freunde 203. zu Berlin, 1883: 128-132. Z a r d in i , R., 1973. Fossili di Cortina; atlante degii echinodermi S t u d e r , T., 1884. Verzeichniss der während der Reise SMS cassiani. Trias Medio Superiore della Regione Dolomitica At- ‘Gazelle’ um die Erde 1874-76 gesammelten Asteriden und tomo a Cortina d’Ampezzo. Foto Ghedina, Cortina d’Ampezzo, Euryaliden.Physikalische Abhandlungen der Königlich-Preus- 29 pp. sichen Akademie der Wissenschaften zu Berlin, 1884(3): 1-64.

V a l e t t e , A., 1929. Note sur quelques stellérides jurassiques du Laboratoire de Géologie de la Faculté des Sciences de Lyon. Daniel B. B l a k e Travaux du Laboratoire de Géologie de la Faculté des Sciences Department of Geology, de Lyon, 16: 1-39. Room 245 Natural History Building V errill, A., 1899. Revision of certain genera and species of University of Illinois starfishes with description of new forms. Transactions of the 1301 West Green Street Connecticut Academy, 10: 145-234. Urbana, IL 61820 V il l ie r , L., B l a k e , D.B, J a g t , J.W.M. & K u t s c h e r , M., 2004. USA A preliminary phylogeny of the Pterasteridae (Echinodermata, E-mail: [email protected] Asteroidea) and the first fossil record: Late Cretaceous of Germany and Belgium.Paläontologische Zeitschrift, 78: 281 - John W.M. J a g t 300. Natuurhistorisch Museum Maastricht

V il l ie r , L. & K u t s c h e r , M., 1999. How dissociated ossicles de Bosquetplein 6 can further our understanding of the origins of the Neoaster- NL-6211 KJ Maastricht oidea. An example from the Toarcian of Western Europe. In: The Netherlands C a n d ía C a r n e v a l i , M.D. & B o n a s o r o , F. (Editors), Echino E-mail: [email protected] derm Research 1998, Proceedings of the Fifth European Con ference on Echinoderms, Milan/Italy/7-12 September 1998. Typescript submitted: July 1, 2004 A.A. Balkema, Rotterdam/Brookfield, pp. 417-422. Revised typescript received: October 15, 2004 194 Daniel B. BLAKE & John W.M. JAGT

Explanation of Plates

P late 1

Comparative morphology of Pseudarchasterinae (Figs. 1, 2) and non-pseudarchasterine goniasterids (Figs. 3-11). Scale bar equals 5 mm in 1-5, 1 mm in 6-11.

Figs. 1,2 — Skiaster vikingr n. gen., n. sp., holotype (NHMM K 2086) in inclined dorsal and full dorsal views, respectively; marginals are abutted across the midline of the arm. Figs. 3, 7 — Pseudarchaster parelii DÜBEN & Ko r en , 1846; inclined dorsal view (CAS 137908; Recent, Aleutian Islands, Alaska) and lateral view of abactinal ossicles (CAS 117915; Recent, off Oregon); tabular marginals and paxilliform abactinals such as those of Pseudarchaster indicate pseudarchasterinae affinities of Skiaster vikingr n. gen., n. sp. Figs. 4, 6 — Nymphaster moebii (S t u d er , 1884); Muséum national d’Histoire naturelle collections, Paris (unnumbered); Recent, 12° 55’ S, 44u 57’ E; inclined dorsal view and lateral view of abactinals. Species is similar toS. vikingr but marginals are too angular with an intermarginal re-entrant and abactinals are not paxilliform; compare with Figs. 9-11. Figs. 5, 8 — Mediaster aequalis S tim p s o n , 1857; University of Illinois collections, unnumbered specimen; Recent, off California. V errill (1899) recognised Mediaster as intermediate between pseudarchasterines and other gonias terids (see text). Abactinals are superficially similar to those ofPseudarchaster, but they are not paxilliform. Marginals are more rounded, although differences are subtle. Figs. 9-11 — Nymphaster studlandensis (S chulz & W eitsch a t , 1975); 9, NHMM 1994640b, ambulacrals and lower surfaces of abactinals to right; 10, 11, NHMM 1994640a. This fossil is close to the pseudarchasterines but the abactinals, although tail, are facetted and closely fitted rather than paxilliform (see PI. 2, Figs. 2, 5-8).

P late 2

Skiaster vikingr n. gen., n. sp., holotype (NHMM K 2086). Scale bar equals 1 mm.

Fig. 1 — Marginals at base of arm are closely fitted, indicating that they abutted at the arm midline in life. Fig. 2 — Central disc, marginal series encloses paxillae. Fig. 3 — Superomarginals in slightly inclined dorsal view. Fig. 4 — Superomarginal in lateral view. Fig. 5 — Interbrachial superomarginals with paxillae to right. Fig. 6 — Superomarginals (centre), inferomarginal (lower left), and abactinals (right), with smaller abactinals below. Figs. 7, 8 — Paxilliform abactinal with re-entrant below crown; papular facets to left of right photograph; compare PI. 1, Figs. 6-11.

P late 3

Betelgeusia exposita n. sp., holotype and only known specimen (NHMM K 3364). Scale bar in 1-3 equals 3 mm, and 1 mm in 4, 5.

Fig. 1 — Ventral view showing general configuration. Fig. 2 — Alignment of actinals between marginals and adambulacrals, with distinct fasciolar grooves between series of ossicles. Fasciolar grooves are not precisely aligned between ossicular types. Two mouth angle pair to right. Fig. 3 — Arrangement of actinals, adambulacrals; ambulacrals as seen to left are similar to those ofB. reidi as well as those of Astropecten (see B l a k e & R e id , 1998) suggesting functional parallels. Fig. 4 — Ventral view of terminal to right partially enclosing distalmost adambulacrals; small distal marginals border adambulacrals. Fig. 5 — Delicate dorsals bordered by marginal series. Slots between the marginals are fascioles. New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 195

P late 4

Aldebarania taberna n. sp. Scale bar equals 5 mm.

Figs. 1,2,4 — Holotype (NHMM 199901-1); 1, 2, dorsal and ventral views, respectively, showing general configuration; 4, ventral view of distal arm showing simple spinelets on inferomarginals, lacking raised areas ofA. arenitea, the type species. Figs. 3, 6 — Paratype (NHMM 199901-3); 3, dorsal view with uniform, small paxillae (collapsed towards centre), terminals on upper arms, form of marginals; 6, ventral view, inferomarginals similar to superomarginals, central disc is disrupted, showing ambulacrals, actinal arrangement. Fig. 5 — Paratype (NHMM 199901-2); ventral view, inferomarginals, mouth angle pair preserved in interbrachium to left, actinals disrupted but partially exposed in all interbrachia; ventral surfaces of paxillae visible to right of mouth angle pair.

P late 5

Figs. 1-7 Cheiraster? sp., (NHMM MD 4105). Scale bar equals 3 mm in 1, 8, and 1 mm in 2-7.

Fig. 1 — Overall dorsal view of specimen. Fig. 2 — Lateral view of four medial ambulacrals from the left side of the lower arm (see Fig. 1). Fig. 3 — Dorsal-lateral view of four ambulacrals, the two distal partially obscured by bulbous marginals, from proximal right side of the upper arm (see Fig. 1). Fig. 4 — Dorsal view of ambulacrals with one marginal to upper right and several below, right proximal side of medial arm (see Fig. 1). Fig. 5 — Dorsal view of ambulacral series bordered by marginals, upper arm (see Fig. 1). Figs. 6, 7 — Two inclined views of right side of upper left arm showing form and arrangement of ambulacrals along midline of arm, and marginals. Fig. 8 — Cheiraster echinulatus (P errier, 1875), USNM E23500, Recent, off Venezuela. Inclined dorsal view of arm; abactinal, marginal form and spine development are similar to those of NHMM MD 4105 but similarities are too generalized to finalise fossil assignment (see text). 196 Daniel B. BLAKE & John W.M. JAGT

P l a t e 1 New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 197

P l a t e 2 198 Daniel B. BLAKE & John W.M. JAGT

P l a t e 3 New latest Cretaceous and earliest Paleogene asteroids (Echinodermata) 199

P l a t e 4 200 Daniel B. BLAKE & John W.M. JAGT

P l a t e 5