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Echinodermata, Ophiuroidea) Assemblage from the Middle Jurassic of the Northeast Paris Basin

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Echinodermata, Ophiuroidea) Assemblage from the Middle Jurassic of the Northeast Paris Basin B. Thuy Middle Jurassic ophiuroids from the northeast Paris Basin A peri-reefal brittle-star (Echinodermata, Ophiuroidea) assemblage from the Middle Jurassic of the northeast Paris Basin Ben Thuy Musée national d’Histoire naturelle du Luxembourg, section paléontologie 25 rue Münster, L-2160 Luxembourg [email protected] Zusammenfassung Disartikulierte Lateralplatten von Schlangensternen ciliator sp. nov. Letztere stellt den bisher ältesten Vertreter liefern, wie bereits mehrfach gezeigt wurde, ein viel der wiedervereinigten Ophionereididae-Ophiochito- vollständigeres Bild der Paläobiodiversität der Gruppe nidae Gruppe dar. Eine dritte Art wird als neu erfasst, als die seltenen Funde von komplett erhaltenen bleibt jedoch unbenannt. Die neue GattungEnakomusium Skeletten. Nichtsdestotrotz haben Vergesellschaftungen wird für mehrheitlich mesozoische Arten eingeführt, die in Form von disartikulierten Lateralplatten bisher nur eine gewisse Ähnlichkeit mit dem heute noch lebenden wenig Beachtung gefunden. Die vorliegende Arbeit Ophiomsium vorweisen, jedoch die Grenzen der stellt die erste umfassende Bearbeitung einer Schlangen- Gattungsdiagnose sprengen. Es wird gezeigt, dass sich sternvergesellschaftung aus dem Bajocium (Mittleren die mitteljurassische Vergesellschaftung fundamental Jura) dar, die vollständig auf disartikulierten Lateral- auf Familien- und noch stärker auf Gattungsebene von platten basiert. Das untersuchte Material entstammt heutigen flachmarinen, riffnahen Äquivalenten unter- Schlämmresten von zwei Mergelschichten, die an einem Strassenaufschluss nahe Longwy, Frankreich, am scheidet. In der Tat wird sie von typischen modernen nordöstlichen Rand des Pariser Beckens anstehen. Die Tiefseegruppen oder Tiefseevertretern von Flachwas- Mergelschichten werden zur Laeviuscula Zone (unteres sergruppen dominiert. Die hier beschriebene Verge- Bajocium) gestellt und werden als flachmarine, riffnahe sellschaftung bekräftigt somit die Beobachtung, dass Ablagerungen interpretiert. Es wurden insgesamt sieben sich flachmarine Schlangensternvergesellschaftungen Arten identifiziert, zwei davon als neu beschrieben: seit dem Mesozoikum stark in ihrer Zusammensetzung Ophiotitanos aschmannicor sp. nov. und Ophiodoris recon- verändert haben müssen. Abstract Dissociated lateral arm plates of ophiuroids have been and Ophiodoris reconciliator sp. nov. The latter is the oldest shown to produce a much more complete picture of the record of the re-united Ophionereididae-Ophiochi- palaeobiodiversity of the group than the assessment tonidae group. A third species is recorded as new but of the exceptional finds of articulated skeletons. Yet, left unnamed. The new genus Enakomusium gen. nov. is ophiuroid assemblages preserved as dissociated lateral introduced to accommodate mainly Mesozoic species arm plates have received only limited attention so far. which are superficially similar to extant Ophiomusium Here, the first exhaustive survey of an ophiuroid assem- but fail to match the diagnosis of the latter. The Bajocian blage entirely based on dissociated lateral arm plates is ophiuroid assemblage is shown to differ fundamentally presented for the Bajocian (Middle Jurassic). The studied from modern shallow-water peri-reefal equivalents on specimens were retrieved from sieving residues of two marl beds exposed at a roadcut site near Longwy, France, family and, even more, genus level, being dominated on the northeast margin of the Paris Basin. The marl beds by typical extant deep-sea taxa or deep-sea members are dated to the Early Bajocian Laeviuscula Zone and are of otherwise shallow-water groups. The assemblage interpreted as shallow-water peri-reefal deposits. A presented herein thus adds to the growing evidence that total of seven species were identified, two of which are shallow-water ophiuroid communities have undergone described here as new: Ophiotitanos aschmannicor sp. nov. dramatic changes in composition since the Mesozoic. Ferrantia • 71 / 2015 87 B. Thuy Middle Jurassic ophiuroids from the northeast Paris Basin Résumé Les plaques brachiales latérales dissociées d’ophiures, sp. nov.. La dernière représente le plus ancien membre comme l’a été montré à plusieurs reprises, fournissent un du groupe réunifié Ophionereididae-Ophiochitonidae. aperçu beaucoup plus complet de la paléobiodiversité Une troisième espèce est décrite comme nouvelle mais du groupe que les rares trouvailles de squelettes intacts. reste sans nom formel. Le nouveau genre Enakomusium Néanmoins les associations d’ophiures conservées sous est introduit pour les espèces majoritairement mésozo- forme de plaques brachiales latérales dissociées n’ont ïques qui ressemblent superficiellement à l’Ophiomusium reçu que peu d’attention jusqu’à présent. La présente actuel mais qui ne se conforment pas à la diagnose de contribution fournit la première étude exhaustive ce dernier. Il est montré que la population bajocienne d’une population d’ophiures entièrement basée sur des diffère fondamentalement des populations actuelles plaques brachiales dissociées pour le Bajocien (Juras- équivalentes au niveau des familles et, surtout, au niveau sique Moyen). Les spécimens examinés ont été extraits des genres. En effet, la population comporte princi- de résidus de tamisage de deux couches marneuses palement des groupes typiques des mers profondes qui affleurent dans un talus près de Longwy, France, actuelles ou des membres vivant actuellement en mer à la bordure nord-est du Bassin de Paris. Les couches profonde de groupes majoritairement trouvés dans les marneuses sont datées de la zone à Laeviuscula (Bajocien mers peu profondes. Ainsi, la présente étude consolide inférieur) et ont été déposées dans un milieu peu l’hypothèse selon laquelle les populations d’ophiures profond à proximité de récifs de coraux. Au total sept des mers peu profondes auraient subi de profonds espèces ont été identifiées, dont deux nouvelles: Ophio- changements au niveau de leur composition depuis le titanos aschmannicor sp. nov. et Ophiodoris reconiliator Mésozoïque. Introduction palaeoecological and macroevolutionary studies (e.g. Thuy et al. 2012; Thuy & Meyer 2012; Thuy 2013; Thuy et al. 2013). Brittle stars (ophiuroids) are a major component of benthic communities in the modern oceans, Nevertheless, and in spite of fairly numerous abundantly occurring at all latitudes and depths, taxonomic studies (e.g. Hess 1962, 1963, 1964, from the poles to the tropics and from the inter- 1965a, 1965b, 1966, 1972, 1975a, 1975b; Hess & tidal to the hadal trenches (Stöhr et al. 2012). Their Meyer 2008; Kutscher & Hary 1991; Kutscher importance as fossils, however, has long been 1996; Kutscher & Jagt 2000; Kutscher & Villier underestimated. In fact, articulated ophiuroid 2003; Thuy 2005; Thuy 2011; Thuy & Kroh 2011; skeletons, only exceptionally preserved as fossils Thuy 2013; Thuy et al. 2013), the ophiuroid fossil (Ausich 2001), were commonly considered the record is still only patchily known. For some only reliable source of evidence for species stages, only very few ophiuroid species have been identifications. Recent systematic morphological reported so far. This is particularly the case when analyses of previously neglected ophiuroid no exhaustive survey based on dissociated lateral skeletal characters, in particular those pertinent arm plates is available for a given stage. In fact, to the spine-bearing lateral arm plates (Martynov inclusion of lateral arm plate evidence has been 2010; Thuy & Stöhr 2011), however, have set shown to dramatically expand knowledge on the a robust fundament to include disarticulated fossil record of the group (e.g. Thuy 2005, 2013). skeletal plates, abundantly occurring as micro- fossils in most marine sediments, as a promising The Bajocian is one of these insufficiently explored new source of evidence to explore the fossil record parts of the ophiuroid fossil records, with, until of the group. recently, nine species known on the basis of artic- ulated skeletons (Hess 1964; Thuy & Meyer 2012). As a result, the assessment of ophiuroid palaeo- In a recent survey of the family Ophiacanthidae, biodiversity is currently witnessing a remarkable another three species and an unnamed record ascent, with more than one fifth of the accepted were added on the basis of dissociated lateral arm fossil ophiuroid species added during the last three plates (Thuy 2013). years on the basis of dissociated lateral arm plates (Stöhr & O’Hara 2013). In addition, contributions This study provides the first exhaustive survey of a more general scope have begun to explore the of an ophiuroid assemblage entirely based on potential of ophiuroids as a model organism for dissociated lateral arm plates for the Bajocian, 88 Ferrantia • 71 / 2015 B. Thuy Middle Jurassic ophiuroids from the northeast Paris Basin along with the study by Numberger-Thuy and Formation and dated to the early Bajocian Laevi- Thuy in the present volume. The aim of the uscula Zone (Boulvain et al. 2001). The lower part study is to contribute to a more complete picture of the limestone succession includes two distinct of ophiuroid palaeobiodiversity, specifically beds of grey (freshly exposed) to brownish-yellow targeting the species not known from articulated (weathered), clayey, unconsolidated marls, each fossils, and thus to expand the alpha-taxonomical approximately 0,4 m in thickness, separated by basis for further-reaching applications. The study 1,5 m of coarsely-bedded limestone. The samples furthermore attempts a comparison with modern yielding
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