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4. Arbeitstreffen deutschsprachiger Echinodermenforscher Wien, Oktober 08

4. Arbeitstreffen deutschsprachiger Echinodermenforscher, Wien, Okt. 08 4th Workshop of German & Austrian Research

4. Arbeitstreffen deutschsprachiger Echinodermenforscher

4th Workshop of German & Austrian Echinoderm Research

24.-26. October 2008

ABSTRACTS

Andreas Kroh & Brigitta Schmid Naturhistorisches Museum Wien

4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Impressum

Eigentümer, Herausgeber und Verleger: © 2008 Natuhistorisches Museum Wien Alle Rechte vorbehalten. Für den Inhalt der Beiträge sind die Autoren verantwortlich.

Redaktion: Naturhistorisches Museum Wien Burgring 7, A-1010 Wien, Österreich (Austria) Tel.: +43 (1) 521 77 / 576 (Kroh); +43 (1) 521 77 / 564 (Schmid) Fax: +43 (1) 521 77 / 459 e-mail: [email protected] e-mail: [email protected]

Redaktionelle Bearbeitung (EDV, Layout, Grafik): Andreas Kroh Umschlagentwurf: Andreas Kroh

Titelbild: Coelopleurus (Keraiophorus) exquisitus COPPARD & SCHULTZ, 2006;

Neukaledonien (NHMW 2005z0277/0016; Photo von Alice SCHUMACHER, NHM Wien).

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Inhalt

Encrinus sp. cf. E. robustus (Crinoidea, Encrinidae) aus dem Unteren Muschelkalk von Niedersachsen (Poster) Ulrich Bielert, Friedrich Bielert & Hans Hagdorn...... 1 Classification of pre- and postmortem ossicular modifications of the Cupressocrinitid skeletons (Crinoidea, ) (Poster) Jan Bohatý ...... 3 Dental insights into ophiuroids: Feeding mechanisms and life style (Poster) Karin Boos...... 14 Ökophänotypbildung der Muschelkalk-Crinoide Encrinus liliiformis - Vergleich zweier Populationen aus Nord-Württemberg (Vortrag) Janina F. Dynowski & James H. Nebelsick...... 15 Stereomdifferenzierung in verschiedenen Seeigeln und ihr bionisches Potenzial (Vortrag) Nils Großmann & James H. Nebelsick...... 17 Die Stereome diverser regulärer Seeigelstachel (Poster) Nils Großmann & James H. Nebelsick...... 19 30 Years of Research on / 30 Jahre Crinoidenforschung (Keynote) Hans Hess ...... 20 Pattern formation in starfish: arm stumps, regeneration models, and evolution (Poster) Frederick H.C. Hotchkiss...... 21 Bdellacoma in the Hunsrück Slate (Lower Devonian, Germany): reidentification of Urasterella verruculosa (Asteroidea, Bdellacomidae) (Poster) Frederick H.C. Hotchkiss & Alexander Glass...... 23 Index of Living and Fossil Echinoids: 1971 - present (Poster) Andreas Kroh...... 25 Crinoids from the Tithonian of the Klippenbelt (, Austria) (Poster) Andreas Kroh & Alexander Lukeneder...... 26

iv 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Temperature effect on feed consumption, absorption and assimilation efficiencies and production in the Strongylocentrotus intermedius (Poster) John M. Lawrence, Xuebin Cao, Yaqing Chang, Ping Wang, Y. Yu, Addison L. Lawrence, Stephen A. Watts ...... 28 Amount of arm loss and rate of arm regeneration by Luidia clathrata (Echinodermata: Asteroidea) (Poster) J.M. Lawrence & C.M. Pomory ...... 29 Seastars (Asteroidea) from Madagascar (Vortrag) Horst Moosleitner ...... 30 Morphologie und Faziesverteilung des indo-pazifischen Seeigels Jacksonaster depressum (Vortrag) James Nebelsick & Atef Abd El-Hamied A. S. Elattaar...... 32 Taphonomie von Jacksonaster depressum (Echinoidea: Clypeasteroida) aus dem Roten Meer (Poster) James Nebelsick & Atef Abd El-Hamied A. S. Elattaar...... 33 Skelett-Endosymbionten fossiler und rezenter Echinodermen: Von Bioklaustrationen zu komplexen Strukturen (Vortrag) Christian Neumann ...... 34 Echinoids from the Neogene of Portugal mainland: Systematics, Palaeoecology, Palaeobiogeography (Vortrag) Pedro Pereira ...... 36 Phylogenetic relationships within Echinodermata based on mitochondrial sequence data (Vortrag) Marleen Perseke, Guido Fritzsch, Detlef Bernhard, Peter F. Stadler & Martin Schlegel ...... 38 New insights into early diversification of sea cucumbers as inferred from calcareous ring material (/, Baltoscandia) (Vortrag) Mike Reich...... 39 A nearly articulated holothurian from the Santonian () of England (Poster) Mike Reich...... 40

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Unusual holothurians (Echinodermata) from the Late Ordovician of Sweden (Poster) Mike Reich...... 42 Signs of predation on Middle crinoids form Poland (Poster) Mariusz A. Salamon & Przemysław Gorzelak ...... 43 Are that sea urchins ???? - The strange morphology of the echinoid family Pourtalesiidae (Vortrag) Heinke Schultz ...... 45 The early history of : new insights from disarticulated early and middle ossicles (Keynote) Andrew B Smith ...... 48 Rhaetian ophiuroids from the Netherlands: a preliminary report (Poster) Ben Thuy, Adiël A. Klompmaker & John W. M. Jagt...... 49 Irregular echinoids from the Turonian Seewen Limestone of the Allgäu area (S. Germany) (Vortrag) Frank Wiese & Nils Schlüter ...... 51 Antiquity and Longevity of a deep-sea oyster/ association (Azores Archipelago) (Vortrag) Max Wisshak, Christian Neumann, Matthias López Correa, Serge Gofas, Carmen Salas, Marco Taviani, Joachim Jakobsen, André Freiwald ...... 53 Bromierte Anthrachinon-Pigmente aus der rezenten Seelilie Proisocrinus ruberrimus (Vortrag) Klaus Wolkenstein, Wolfgang Schoefberger, Norbert Müller & Tatsuo Oji...... 55

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4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Encrinus sp. cf. E. robustus (Crinoidea, Encrinidae) aus dem Unteren Muschelkalk von Niedersachsen (Poster)

ULRICH BIELERT1, FRIEDRICH BIELERT2 & HANS HAGDORN3

1 Rheinblick 4, 69226 Nußloch, Deutschland; e-mail: [email protected]

2 Am Goldgraben 21 37073 Göttingen, Deutschland; e-mail: [email protected]

3 Muschelkalkmuseum, Schloss-Str. 11, 74653 Ingelfingen, Deutschland; e-mail: encrinus@t- online.de

Auf zwei oolithischen Gesteinsstücken aus den Terebratelbänken (Unterer Muschelkalk, Jena-Formation; Anisium, Pelsonium) von Königslutter am Elm (Niedersächsisches Landesmuseum Hannover 50004 und 52040, aus Slg. O. Klages) sind artikulierte Reste von Kronen und Stielen eines großwüchsigen Encrinus erhalten. Die Gattung ist im Unteren Muschelkalk mit mehreren nominalen Arten vertreten, die jedoch ungenau diagnostiziert sind, weshalb die Zuordnung von Einzelfunden oft Schwierigkeiten bereitet. Die drei vorliegenden Individuen (Kronenlänge bis 50 mm) sind gekennzeichnet durch einen stumpf kegelförmigen Kelch mit langen, mit ca. 50o gegen die Längsachse geneigten, in Seitenansicht sichtbaren Basalia (B), die distal tief in die interradiale Sutur eingreifen. Dorsal sind Basalia und Radialia (R) schwach gewölbt und zeigen keine Buckel oder sonstige Ornamente; die Suturen liegen nicht vertieft. Die Artikulation der Kelchplatten ist zygosynostosial. Die Arme sind proximal relativ breit, werden nach distal jedoch rasch schmaler. Das erste Primibrachiale (IBr1) ist relativ lang, flach quergewölbt und grenzt

24.-26. October 2008 1 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research mit langer Sutur gegen die Nachbarglieder. Auch das Primaxillare (IBr2Ax) ist relativ lang und hat wie das IBr1 keine dorsalen Buckel. Zusammen mit den beiden ersten Sekundibrachialia (IIBr1, IIBr2) wirken Kelch und proximale Arme ballonförmig aufgetrieben. Ab dem dritten Sekundibrachiale (IIBr3) setzt abrupt echte Zweizeiligkeit der Armglieder ein. Die Armzweige sind zunächst dorsal noch schwach quergewölbt und haben scharf abgegrenzte Seitenflächen, werden ab dem IIBr3 aber schmaler und dorsal gerundet. Die Brachialia sind dorsal annähernd glatt und unornamentiert. Die Pinnularia sind schlecht erhalten. Am Stiel fällt auf, dass auch die proximalen, zirrenlosen Nodalia kaum länger und dicker sind als die Internodalia und flache Seiten ohne Randwulst haben. Die Nähte zwischen den durchweg runden Columnalia sind kaum vertieft. Wie bei Populationen von E. liliiformis, wo die dorsale Ornamentierung von Kelch und Armgliedern als ökophänotypisches Merkmal erheblich schwankt, ist der diagnostische Wert des Kelches mit den langen Basalia höher einzuschätzen als die dorsale Ornamentierung von Kelch- und Armgliedern. Nur die altersgleichen

E. brahli aus Rüdersdorf (BEYRICH 1857) und E. robustus aus Oberschlesien

(ASSMANN 1926) und Thüringen (BIELERT & BIELERT 2000) haben einen flachkonischen Kelch und Armglieder ohne dorsale Skulptur. Den halbkugelförmigen dorsalen Auftreibungen der Primaxillaria beim schlesischen und thüringischen Material von E. robustus messen wir nur geringe diagnostische, sondern ökophänotypische Bedeutung zu. Der Status von E. brahli, dessen Typusmaterial (subadulte Individuen) verschollen ist, bedarf weiterer Untersuchungen. Die hier vorgestellten Encriniden werden als Encrinus sp. cf. E. robustus ASSMANN, 1925 bezeichnet.

Literatur

ASSMANN, P., 1926. Die Fauna der Wirbellosen und Diploporen der ober- schlesischen Trias mit Ausnahme der Brachiopoden, Lamellibranchiaten, Gastropoden und Korallen. – Jahrbuch der preußischen Geologischen Landes-Anstalt für 1925, 46: 504-527.

BEYRICH, E., 1857. Über die Crinoiden des Muschelkalks. – Abhandlungen der Akadademie der Wissenschaften Berlin, 1: 1-49.

BIELERT, U. & BIELERT, F., 2000. Gemeinsames Auftreten von zwei Arten der Gattung Encrinus im Unteren Muschelkalk am Nordrand des Thüringer Beckens. – Beiträge zur Geologie von Thüringen, Neue Folge 7: 137-145.

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Classification of pre- and postmortem ossicular modifications of the Cupressocrinitid skeletons (Crinoidea, Devonian) (Poster)

JAN BOHATÝ

Institut für Geologie und Mineralogie der Universität zu Köln, Zülpicher Str. 49a, 50674 Köln, Germany; e-mail: [email protected]

Skeletal anomalies on Devonian cupressocrinitids (Crinoidea, Cladida) are herein classified as: 1, growth anomalies without external influences; 2, growth anomalies without classifiable causes; 3, premortem ossicle anomalies as a reaction of external interferences; 4, pre- and postmortem borings and bite marks; and 5, pre- and postmortem epizoan encrusting.

1. Growth anomalies without recognizable external influences – "generic" abnormalities (Figs. 1-2):

Growth anomalies without recognizable external influences are predominantly distinguished by the reduction of thecal or brachial-ossicles respectively by additional intermediary plates. These abnormalities could not be attributed to injuries or involved regeneration and are obviously "genetically modified anomalies" (BOHATÝ 2001). Most common are variances of the columnal axial canal (Fig. 1), which occurs at the rate of ~1: 30 compared with regular grown axial canals (~1500 skeletons analyzed). Further, individuals with additional or a reduced number of ossicles are recognized. Cupressocrinitids with a developed quadrangular or hexagonal symmetry (Figs. 1-2) are relatively rare and occur at several localities with an average rate of ~1: 70 compared with regularly developed skeletons (~700 aboral cups and ~300 crowns analyzed). Due to the abundance of anomalously grown axial canals or symmetry aberrations within one fossil-horizon, the genetic basis of these interferences is assumed. In this case, the appropiative rates of detectable growth anomalies compared with normal individuals, could be higher than above-mentioned.

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2. Growth anomalies without classifiable causes – without indications of external influences (Fig. 3):

In some cases it is not possible to determine a cause for a growth anomaly. The individuals with one additional or missing plate (Fig. 3), with an inexplicable ossicular-swelling, or a modified exobrachial layer are not recognizable as regeneration of the skeleton, "wound healings", or as documented "generic" abnormalities. No direct evidence of predatory influences like borings or bite marks can be recognized. Therefore, these modifications are summarized as growth anomalies without classifiable causes – without indications of external influences.

3. Premortem ossicular anomalies as a reaction of external interferences – "wound healing" and skeletal regeneration of thecal or brachial injuries (Figs. 4-5):

"Wound healing": Different sized anomalies in numerous small ossicles were recognized on ~5 % of the studied cupressocrinitids (~700 aboral cups and ~300 crowns analyzed). These anomalies are obviously "wound healings" of nonlethal injured individuals. Possible causes of these anomalies could be injuries caused by predators or possibly by impact-injuries with suspended clastic material. The affected regions may be small or large (Fig. 4). The maxim observed injury affects up to 80 % of the surface of the cup.

Regeneration: Regenerations of echinoderm skeletons was recently reconsidered by MOZZI et al. (2006), exemplified by the regenerative processes of the

"Mediterranean Featherstar" Antedon mediterranea (LAMARCK, 1816). AMEMIYA &

OJI (1992) described the crinoid regeneration processes. The regeneration in fossil crinoids was also discussed by GAHN & BAUMILLER (2005). For example, they showed arm regeneration of Rhodocrinites kirbyi (WACHSMUTH & SPRINGER, 1889) and Dichocrinus cinctus MILLER & GURLEY, 1890. Direct interconnections between the increase of shell-breaking predators and the number of observed arm regenerations of nonlethal injured crinoids were recognized (GAHN & BAUMILLER

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2005, p. 151-164). Further, WEISSMÜLLER (1998) discussed arm regeneration of the Muschelkalk-crinoid Encrinus liliiformis LAMARCK, 1801 as did MEYER & OJI (1993) for several metacrinitids. Arm regeneration in Devonian crinoids is recognized by the conditions specified by GAHN & BAUMILLER. At the juncture of the injury, the regenerated skeleton has either 1, the insertion of particularly small arms; or 2, the abrupt change in the magnitude of the arm-ossicles (2005, p. 156). The arms

24.-26. October 2008 5 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research recognized as regenerated were all smaller than regularly developed arms (Fig. 5). Nevertheless, the arms of the relevant individuals are enclosed in the typical cupressocrinitid-like resting or avoidance posture, whereas the adjoining, normal longer arms closed about the smaller one and are tangent distally above the regenerated arm.

4. Pre- and postmortem borings and bite marks (Figs. 6-7):

Postmortem multi-borings: Almost 90 % of ~50 analyzed skeletons of C. elongatus were covered by borings (SIEVERTS-DORECK 1963; BOHATÝ 2001, p. 8; 2006, pl. 10, figs. 1-3) [Fig. 6]. More infrequently, specimens with multiple borings were identified on the crowns of C. crassus (2006, pl. 10, fig. 8b). Both are covered by a thin and monolamellar exoplacoid layer, which apparently offered less resistance against boring organisms, in contrast to the multilamellar layers of Abbreviatocrinites. Generally, these borings were restricted to the non-embedded side of the relevant skeletons and trend in inordinated lines from the cup (or also from the preserved stem) and over one or several arms. Presumably, the borings occurred soon after death. The skeletons are articulated and covered by the unsheared exoplacoid layer on the one hand, but on the other, the borings are restricted to the non-embedded side of the crown. Platyceratid gastropods were discussed as a possible causer of the borings (SIEVERTS-DORECK 1963). This theory cannot be verified.

Pre- and postmortem incurred single borings: Single borings are present on the ossicles of A. abbreviatus abbreviatus, A. geminatus, and Robustocrinites n. sp. In abbreviatocrinitids, they are normally restricted to the plates with sheared exoplacoid layers and, therefore, most likely occurred postmortem. The single boring of an unknown organism at the surface of the monolamellar exoplacoid layer, observed in one affected robustocrinitid, is filled by a trepostome bryozoan (?Eostenopora sp.). Because the boring is positioned on the non-embedded side of the crown and runs across several plate boundaries, it is assumed to have occurred postmortem.

BAUMILLER & MACURDA (1995) and BAUMILLER (1990, 1993) documented borings on Paleozoic blastoids and crinoids. Also in this case, platyceratid gastropods were discussed as the possible borers. A significant bit of evidence for

6 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research this theory is perhaps documented in the combined fossil evidence of a borehole, positioned next to a gastropod valve (BAUMILLER 1990). SEM-observations of thin cross-sections of the multilamellar exoplacoid layer of A. geminatus exhibits potentially premortem microendolithic borings. These meandering single borings have an averag proportion of 20 µm in width to 300 µm length. They were presumably lined with biogenous matter and ultimately resulted in a secondary sulphide-ion surplus. Through this, the borings are lined with marcasite-crystal agglomerates (FeS2). Microendolithic borings could be observed in ~70 % of the studied multilamellar exoplacoid layers, but in less than 20 % of the basal, radial, or brachial plates (30 thin sections analyzed). Fig. 7 presumably has a deep, oval boring(?) on the basal plate of A. abbreviatus. The visible stereomatic reaction in the form of an annulus-like swelling indicates that the single-boring occurred most likely premortem. But isolated placoderm teeth from the same location also permit the assumption that this trace may to the bite of a larger predator instead of a boring organism, like a gastropod with specialised radula.

Premortem bite marks: Bite marks at cupressocrinitids are rare and could be observed in less than 3 % of the studied individuals (~1500 skeletons analyzed). They are possibly attributed to cephalopods, placoderms or arthropods. Premortem bite marks are recognized as nonlethal injuries, because the bite marks are accompanied by "wound healings".

5. Pre- and postmortem incurred epizonal encrusting (Fig. 8):

The epibiontic encrusting of Devonian crinoids, exemplified by Upper Eifelian columnals, was recently discussed by Gluchowski (2005). Bryozoa, Microconchida, Crinoidea, , Rugosa, and Stromatoporida are also identified on the crown-ossicles of cupressocrinitids.

Bryozoa: "Cyclostome bryozoans" (Hederella sp.) apparently preferentially encrusted the crown-ossicles of Abbreviatocrinites nodosus (Fig. 8). Unlike other cupressocrinitids, nearly 95 % of the observed A. nodosus-skeletons from the Klausbach and Nims members bear encrustings (~80 skeletons analyzed). The growth of the hederellids most likely occurred instantaneously postmortem,

24.-26. October 2008 7 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research because some articulated crowns retain unsheared exoplacoid layers. In contrast, the "cyclostome bryozoans" settled beyond the primary movable ossicle boundaries of the endoskeleton. Presumably, the hederellids had a rapid rate of growth. Gluchowski (2005, figs. 4F-H) also documented the hederellid-encrusting of Upper Eifelian crinoid columnals. Hederella is presumably not a true bryozoan (A. Ernst, personal commun.; also see WILSON & TAYLOR 2001). TAYLOR & WILSON (2007) favored a close relationship with phoronids, tentatively interpreting hederelloids as colonial, phoronid-like invertebrates with retractable lophophores. Along with microconchids and cornulitids, hederelloids may have been part of a mid- Paleozoic acme of lophophorate "worms".

Trepostome bryozoans: One brachial of a completely preserved A. nodosus crown, one cup of an also entire Abbreviatocrinites schreueri crown, and one theca of P. gracilis were encrusted postmortem by trepostome bryozoans (?Eostenopora sp.). The boring trace of an affected Robustocrinites arm is also populated by (?)Eostenopora sp.; in this case, the colony settled in a non- exposed position.

Fenestrate bryozoans: Especially within the uppermost Ahbach Formation (lowermost Lower Givetian) of the "Wotan" quarry (Hillesheim syncline, Eifel, Rhenish Massif, Germany), several longer stems of A. geminatus and P. gracilis were found encrusted by fenestrate bryozoans. The length of the overgrown columnals as well as some observed embedding patterns of bryozoans located underneath the attached stem, allows the presumption of a premortem settlement (compare BOHATÝ 2005, fig. 3B). In contrast, some shorter stem fragments or other disarticulated cupressocrinitid-ossicles were usually encrusted postmortem. This assumption is based on the entire enclosure of some skeletal elements. Similarly holdfasts of most likely rhomboporid bryozoans attached to the columnals of Schyschcatocrinus creber DUBATOLOVA, 1975, as reported by

GLUCHOWSKI (2005, figs. 3A-B). Gluchowski indicated that the bryozoans lived attached to the fragmented dead stems that lay horizontally on the sea floor.

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Crinoidea: One observed cup of A. abbreviatus abbreviatus as well as one isolated radial and arm plate of A. geminatus were encrusted by the holdfasts of other cladid crinoids (?P. gracilis). The roots settled postmortem on the shearing of the multilamelar exoplacoid layer, at the interior of a radial plate, or at the multilamellar exobrachial layer of an isolated brachial. This association was observed on less than 3 % of the studied crinoids (~1500 skeletons analyzed).

GLUCHOWSKI (2005, p. 322) documented the postmortem encrusting of several small crinoid holdfasts attached to Upper Eifelian crinoid columnals. Various attachments of crinoid juveniles to living or dead adults are known from the Silurian to the Mississippian (see MEYER & AUSICH 1983). Coiling stems, modified discoid holdfasts on the columns of crinoid hosts, as well as dendritic holdfasts distributed on all sides of the column were reported from Silurian strata by FRANZÉN (1977) and PETERS & BORK (1998).

Lophophorata (?) - Microconchida (Fig. 8): Microconchids with unstructured or ornamented valves encrusted the ossicles of nearly 40 % of the studied cupressocrinitids (~1500 skeletons analyzed). It is remarkable, that larger individuals are rare and isolated (compare Bohatý 2006, pl. 5, fig. 8), whereas numerous smaller microconchids encrusted the crinoids. Supposeably, the colonization occurred immediately postmortem, before shearing of the exoplacoid layers and ossicle disarticulation. The single-species encrusting of microconchids on the columnals of

Tantalocrinus scutellus LE MENN, 1985 and Schyschcatocrinus creber DUBATOLOVA,

1975, represent additional settlement examples (GLUCHOWSKI 2005, p. 323, figs. 5I-L).

Anthozoa - Tabulata: The predominantly postmortem settlement of tabulate was recognized on 5-10 % of the studied cupressocrinitids (~1500 skeletons analyzed). The most common epibiontic tabulates were auloporids like

Aulopora cf. A. serpens minor (GOLDFUSS, 1829) and A. cf. A. s. serpens

(GOLDFUSS, 1829), settling on isolated crown-ossicles, completely preserved cups of A. geminatus, and on the crowns of A. nodosus. Furthermore, one cup of A. a. abbreviatus with an encrusting favositid (Favosites cf. F. goldfussi D'ORBIGNY, 1850) was found within the lower part of the Loogh Formation (Lower Givetian) in the "Wotan" quarry (Hillesheim

24.-26. October 2008 9 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research syncline). GLUCHOWSKI (2005) documented small colonies of Favosites sp. attached to Pentagonostipes petaloides MOORE & JEFFORDS, 1968 and

Tantalocrinus scutellus LE MENN, 1985 and discussed the possible growth along the axis of the upright stalk of a living host. Favositids that lived attached to living crinoid hosts have also been reported from the Upper Silurian (HALLECK

1973; BRETT & ECKERT 1982; PETERS & BORK 1998), Lower Devonian (GALLE 1978;

GALLE & PROKOP 2000), and Lower (compare MEYER & AUSICH 1983). Other tabulate corals (e.g. Antholites, Cladochonus, and Emmonsia) associated with living crinoids are known from Devonian–Mississippian strata

(GLUCHOWSKI 2005, p. 319; also see MEYER & AUSICH 1983; POWERS & AUSICH 1990 and DONOVAN & LEWIS 1999).

Rugosa: Within the Ahbach and Loogh formations (Eifelian/Givetian threshold) in the "Wotan" quarry (Hillesheim syncline), rugose corals settled on disarticulated cupressocrinitid stems and isolated ossicles, including Glossophyllum soetenicum

(SCHLÜTER, 1895) and Thamnophyllum caespitosum (GOLDFUSS, 1826). The additional recovery of a completely overgrown theca (stromatoporoid suffusions, see below) documents a further epibiontic settlement by an indeterminable juvenile stadium of a rugose coral. All settlements occurred postmortem.

GLUCHOWSKI (2005, p. 317-319) detected the premortem encrustings of the rugose coral (?)Adradosia sp. on Schyschcatocrinus creber by the stereomic response of the crinoid.

Porifera(?) - Stromatoporida: Some non-disarticulated cups of A. a. abbreviatus were completely encrusted by indeterminable stromatoporoid suffusions. These encrustings could be settled again by chaetetids, tabulate and rugose corals, microconchids, and bryozoans.

References:

AMEMIYA, S. & OJI, T. 1992. Regeneration in sea lilies. Nature, 357: 546-547.

BAUMILLER, T.K. 1990. Non-predatory drilling of Mississippian crinoids by platyceratid gastropods. Palaeontology, 33/3: 743-748.

BAUMILLER, T.K. 1993. Boreholes in Devonian blastoids and their implication for boring by platyceratids. Lethaia, 26: 41-47.

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BAUMILLER, T.K. & MACURDA, D.B. JR. 1995. Borings in Devonian and Mississippian Blastoids (Echinodermata). Journal of Paleontology, 69/6: 1084-1089.

BOHATÝ, J. 2001. Wachstumsanomalien mitteldevonischer Crinoidenkelche aus den Kalkmulden der Eifel (Rheinisches Schiefergebirge). Greifswalder Geowissenschaftliche Beiträge, 9: 7-9.

BOHATÝ, J. 2005. Doppellagige Kronenplatten: Ein neues anatomisches Merkmal paläozoischer Crinoiden und Revision der Familie Cupressocrinitidae (Devon). Paläontologische Zeitschrift, 79/2: 201-225.

BOHATÝ, J. 2006. Neue Cupressocrinitidae (Crinoidea) aus den mitteldevonischen Kalkmulden der Eifel (linksrheinisches Schiefergebirge, Deutschland). Senckenbergiana lethaea, 86/2: 151-189.

BRETT, C.E. & ECKERT, J.D. 1982. Paleoecology of a well-preserved crinoid colony from the Silurian Rochester Shale in Ontario. Life Science Contribution, Royal Ontario Museum, 131: 1-20.

DONOVAN, S.K. & LEWIS, D.N. 1999. An epibiont and the functional morphology of the column of a platycrinitid crinoid. Proceedings of the Yorkshire Geological Society, 52: 321-323.

DUBATOLOVA, YU. A. 1975. Devonian crinoids of the Minusin Basin [in Russian]. Trudy Instituta Geologii i Geofiziki AN SSSR, Sibirskoe Otdelenie, 272: 1-58.

FRANZÉN, C. 1977. Crinoid holdfasts from the Silurian of Gotland. Lethaia, 10: 219–234.

GAHN, F.J. & BAUMILLER, T.K. 2005. Arm regeneration in Mississippian crinoids: evidence of intense predation pressure in the Paleozoic?. Paleobiology, 31/1: 151-164.

GALLE, A. 1978. Favositidae (Tabulata) from the Devonian of Bohemia. Sbornik Geologických Věd, Paleontologie 20: 33-62.

GALLE, A. & PROKOP, R.J. 2000. Complex parasitism and symbiosis of crinoid, subepidermal parasite, and tabulate coral. Lower Devonian (Pragian), Barrandian, Czech Republic. Věstnik Českého Geologického Ústavu, 75: 441-444.

GLUCHOWSKI, E. 2005. Epibionts on upper Eifelian crinoid columnals from the Holy Cross Mountains, Poland. Acta Palaeontologica Polonica, 50/2: 315-328.

GOLDFUSS, G.A. 1826-44. Petrefacta Germaniae tam ea, quae in museo universitatis regiae Borussicae Fridericiae Wilhelmiae Rhenanae servantur, quam alia quaecunque in Museis Hoeninghusiano, Muensteriano aliisque extant iconibus et descriptionibus illustrate (Abbildungen und Beschreibungen der Petrefacten

Deutschlands und der angrenzenden Länder, unter Mitwirkung des Herrn GRAFEN

GEORG ZU MÜNSTER, herausgegeben von Dr. AUG. GOLDFUSS) – 1 (1826-33): Divisio secunda: Radiariorum Reliquiae – Strahlenthiere der Vorwelt, p. 115-221. Arnz and Co, Düsseldorf.

24.-26. October 2008 11 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

HALLECK, M.S. 1973. Crinoids, hardgrounds, and community succession: The Silurian Waldron−Laurel contact in southern Indiana. Lethaia, 6: 239-252.

LAMARCK, J.B.P.A. de 1801. Systême des animaux sans vertèbres, ou tableau général des classes, des ordres et des genres de ces animaux; présentant leurs caractères essentiels et leur distribution, d'après la considération de leurs rapports naturels et de leur organisation, et suivant l'arrangement établi dans les galeries du Muséum d'Hist. Naturelle, parmi leurs dépouilles conservées; précédé du discours d'ouverture du Cours de Zoologie, donné dans le Muséum National d'Histoire Naturelle l'an 8 de la République. Lamarck & Deterville, Paris, viii + 432 p.

LAMARCK, J.B.P.A. de 1816. Histoire naturelle des animaux sans vertèbres, présentant les caractères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation des principales espèces qui s'y rapportent; précédée d'une introduction offrant la détermination des caractères essentiels de l', sa distinction du végétal et des autres corps naturels, enfin, l'exposition des principes fondamentaux de la zoologie, Vol. 2. Verdière, Paris, 568 p.

LE MENN, J. 1985. Les Crinoides du Dévonien inferieur et moyen du Massif armoricain. Mémoires de la Societé géologique et minéralogique de Bretagne, 30: 1-268.

MEYER, D.L. & AUSICH, W.I. 1983. Biotic interactions among recent and among fossil

crinoids, p. 377–427. In M.J.S. TEVESZ & P.L. MCCALL (eds.), Biotic Interactions in Recent and Fossil Benthic Communities. Plenum Press, New York.

MEYER, D.L. & OJI, T. 1993. Eocene crinoids from Seymour Island, Antarctic Peninsula: paleobiogeographic and paleoecologic implications. Journal of Paleontology, 67/2: 250-257.

MILLER, S.A. & GURLEY, W.F.E. 1890. Description of some new genera and species of Echinodermata from the Coal Measures and Subcarboniferous rocks of Indiana, Missouri, and Iowa. Journal Cincinnati Society of Natural History, 13/1: 3-25.

MOORE, R.C. & JEFFORDS, R.M. (1968): Classification and nomenclature of fossil crinoids based on studies of dissociated parts of their columns. The University of Kansas Paleontological Contributions, 46: 1-86.

MOZZI, D., DOLMATOV, I.Y., BONASORO, F., & CARNEVALI, M.D.C. 2006. Visceral regeneration in the crinoid Antedon mediterranea: basic mechanisms, tissues and cells involved in gut regrowth. Central European Journal of Biology, 1/4: 609-635.

PETERS, S.E. & BORK, K.B. 1998. Secondary tiering on crinoids from the Waldron Shale (Silurian: Wenlockian) of Indiana. Journal of Paleontology, 72: 887-893.

POWERS, B.G. & AUSICH, W.I. 1990. Epizoan associations in a Lower Mississippian Paleocommunity (Borden Group, Indiana, U.S.A.). Historical Biology, 4: 245-265.

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SCHLÜTER, C. 1885. Über einige neue Anthozoen aus dem Devon. Verhandlungen des naturhistorischen Vereins der preussischen Rheinlande und Westphalens, 44: 114- 151.

SIEVERTS-DORECK, H. 1963. Über Missbildungen bei Cupressocrinus elongatus aus dem Mitteldevon der Eifel. Decheniana, 115/2: 239-244.

TAYLOR, P.D.T. & WILSON, M.A. 2007. Morphology and affinities of hederelloid "bryozoans", p. 88. In: S. J. Hageman and F. K. McKinney (eds.), 14th Meeting of the International Bryozoology Association, 02.-07.07.2007, Abstracts with Program. Boone, North Carolina, 2007, (IBA Website, p. 88).

WACHSMUTH, C. & SPRINGER, F. 1889. Crinoids. In S. A. MILLER (ed.), North American geology and paleontology. Western Methodist Book Concern, Cincinnati, 664 p.

WILSON, M. A. & TAYLOR, P.D.T. 2001. "Pseudobryozoans" and the problem of encruster diversity in the Paleozoic. PaleoBios, 21/ 2 addendum: 134-135.

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Dental insights into ophiuroids: Feeding mechanisms and life style (Poster)

KARIN BOOS

Biologische Anstalt Helgoland/ Alfred Wegener Institute for Polar and Marine Research, PO Box 180, 27483 Helgoland; e-mail: [email protected]

Ophiuroid echinoderms are highly specific towards different habitats reflecting lifestyles and feeding mechanisms. While previous studies have considered ophiuroids to be generally omnivorous macro- or microphageous feeders, different feeding mechanisms have evolved according to an epibenthic or infaunal lifestyle, e.g. predation, scavenging, deposit feeding or filter feeding. Most ophiuroids have been reported to show more than one feeding mechanism along with their main feeding habit. The presence and morphology of teeth, oral and dental papillae from ophiuroids performing different lifestyles (epibenthic, infaunal or cryptic) was compared and discussed in relation to reported feeding mechanisms and diets.

14 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Ökophänotypbildung der Muschelkalk-Crinoide Encrinus liliiformis - Vergleich zweier Populationen aus Nord-Württemberg (Vortrag)

JANINA F. DYNOWSKI1 & JAMES H. NEBELSICK2

1 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Deutschland; e- mail: [email protected]

2 Institut für Geowissenschaften, Eberhard Karls Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Deutschland; e-mail: [email protected]

Die vorliegende Arbeit untersucht die Ökophänotypbildung von Encrinus liliiformis aus dem süddeutschen Muschelkalk (Mitteltrias) von Nord-Württemberg. Wie schon von früheren Bearbeitern beobachtet wurde, scheinen sich Kronen aus flacheren Meeresbereichen in ihrer Morphologie von solchen aus tieferen Bereichen zu unterscheiden. Um diese Unterschiede in der Morphologie der Kronen zu erfassen, wurden zwei Populationen von Encrinus liliiformis miteinander verglichen, die aus unterschiedlichen Gebieten des Germanischen Beckens stammen: Crailsheim stellt einen Lebensraum auf einer flach einfallenden Karbonatrampe in geringer Wassertiefe dar, während die Population aus Neckarwestheim aus einem Lebensraum in tieferen Beckenbereichen stammt. Die Vermessung der Kronen und die darauf folgende statistische Auswertung der Daten führte zu einer Differenzierung von zwei verschiedenen Ökophänotypen: 1) Kronen mit kürzeren Armen und weniger stark ornamentierter Oberfläche aus geringerer Wassertiefe; 2) Kronen mit längeren Armen und stärker ornamentierter Oberfläche aus größeren Wassertiefen.

Diese Unterschiede können als Anpassungen an Prädationsdruck und unterschiedliche Strömungsverhältnisse interpretiert werden. Der Prädationsdruck ist im Allgemeinen im flacheren Wasser höher als in größeren Tiefen, weswegen hier eine Verstärkung des Körperbaus als Schutz vor Fressfeinden erwartet wird. Während die Encrinus-Kronen aus Crailsheim allerdings eine weniger stark ausgeprägte Ornamentierung zeigen als die Kronen

24.-26. October 2008 15 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research aus Neckarwestheim, sind die Arme der Crinoiden in Crailsheim kürzer als in Neckarwestheim. Letzteres kann demnach durchaus in Zusammenhang mit einem Unterschied im Prädationsdruck stehen. Die Länge der Arme kann auch ein Hinweis auf die Strömungsstärke sein, die in flacheren Meeresbereichen wie Crailsheim höher ist als in tieferen Regionen. Damit sind Unterschiede im Nahrungsangebot verbunden, das in tieferen Bereichen mit weniger Strömung niedriger ist als in küstennahen, flacheren Lebensräumen. Bei geringerer Strömungsenergie wären für Crinoiden als passive Suspensionsfresser längere Arme, und somit ein größerer Filtrationsfächer, von Vorteil, um mehr Nahrungspartikel aus dem Wasser filtern zu können. Hier könnte auch die stärkere Ornamentierung einen Einfluss haben, zum Beispiel durch eine Änderung im Strömungsregime in direkter Nähe der Krone, um das Wasser effektiver zu filtern.

16 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Stereomdifferenzierung in verschiedenen Seeigeln und ihr bionisches Potenzial (Vortrag)

NILS GROßMANN1 & JAMES H. NEBELSICK2

1, 2 Institut für Geowissenschaften, Eberhard Karls Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Deutschland; e-mail: [email protected], [email protected]

Die Biomimetik (Bionik), die als relativ junges interdisziplinäres Forschungsgebiet gilt, beschäftigt sich mit der Nutzbarkeit biologischer Phänomene bzw. Eigenschaften für eine alltägliche technische Umsetzung. Als Beispiel dient der Lotus-Effekt, welcher u. a. in der Automobilbranche Anwendung findet. Das Projekt „Neue Materialien für leichte, stoffdurchlässige Einschlagschutzsysteme - Seeigel als Modellsystem”, unter finanzieller Leitung der Landesstiftung Baden-Württemberg, wird in Kooperation mit der AG Angewandte Mineralogie (Universität Tübingen) und dem Institut für Textil- und Verfahrenstechnik Denkendorf durchgeführt. Bisher wurden die morphologischen Unterschiede in den Festigkeiten der einzelnen Stereomtypen untereinander (innerhalb der Arten und der Gruppen) und bezüglich ihres Habitats, also hinsichtlich Temperatur, Bewegungsverhalten, Größe und Funktionen der Stacheln, miteinander verglichen. Die Stacheln, wie auch die Schalen von Echinodermen weisen eine poröse Mikrostruktur (Stereom) aus Mg-Calcit mit geringen Einlagerungen (unter 1%) von organischem Material auf. Diese sehr feste und trotzdem leichte Struktur ist außerordentlich stabil und charakteristisch für die Gruppe der Echinodermen. In Verbindung mit den sehnigen Fasern an der Basis der Stachel wird eine zusätzliche Stabilität erreicht. Zur Durchführung des Projekts werden mehrere Seeigelarten aus den Familien Echinidae, Echinometridae und Cidaridae in eigens dafür geschaffenen Aquarien unter realen Umweltbedingungen (Salzgehalt 36‰; 25°C etc.) gehalten. Anhand von CT- und REM-Aufnahmen wird die Stereomstruktur in den Stacheln dokumentiert, auch die Kritische-Punkt-Trocknung wird verwendet. Diese Datensätze können inzwischen bei Druck- und Biegeversuchen zur Interpretation des strukturellen Versagens herangezogen werden. In weiteren Untersuchungen

24.-26. October 2008 17 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research wurden die Magnesium- und Calciumgehalte an verschiedenen Positionen in den Stacheln gemessen und es zeigte sich, dass diese unterschiedlich sind. Stacheln von Heterocentrotus mammillatus dienen dem Verkeilen im Riffgestein, während sie bei anderen Seeigeln, wie bspw. Eucidaris metularia oder Plococidaris verticillata der Abwehr dienen. Ob die dabei festgestellte Unterschiedlichkeit in der Mikrostruktur in einem direkten Zusammenhang mit der Aufgabe der Stacheln steht, bleibt noch zu prüfen. Das Stereom ist im zentralen Bereich der Stacheln großporig und bei Heterocentrotus mammillatus zusätzlich labyrinthisch, bei Plococidaris verticillata gleichmäßig angeordnet. Die Kritische-Punkt-Trocknung zeigt innerhalb des Stereoms eine geringe Anzahl von Weichteilen, die meist aus verschiedenen Zelltypen und Fasern bestehen. Auch gibt es Hinweise, dass die Porenräume mit einer dünnen organischen Schicht ausgekleidet sind. Anhand der Computertomographie-Aufnahmen werden besonders bei Heterocentrotus mammillatus die Wachstumsringe deutlich. Bei ersten Bruchversuchen deutet sich an, dass diese Verdickungen zur Stabilität beitragen können. Im weiteren Projektverlauf werden auch etwaige Unterschiede in der Morphologie der Stacheln bzw. Schalen zwischen Warm- und Kaltwasserarten untersucht.

18 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Die Stereome diverser regulärer Seeigelstachel (Poster)

NILS GROßMANN1 & JAMES H. NEBELSICK2

1, 2 Institut für Geowissenschaften, Eberhard Karls Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Deutschland; e-mail: [email protected], [email protected]

Das Forschungsziel ist eine detaillierte Darstellung der Stereomstrukturen ganzer Seeigelstachel. Es werden Stacheln von diversen Arten der Familien Echinidae, Echinometridae und Cidaridae in Bezug auf ihre Mikrostrukturen untersucht und Habitatsfaktoren wie Größe der Stacheln, Funktionsweise und Umgebungstemperatur verglichen. Hierbei eignen sich besonders Vergleiche zwischen Arten, die entweder Riffrand- oder Riffdachbereiche bewohnen. Als Methoden werden REM-Aufnahmen und zusätzlich das Verfahren Kritische-Punkt- Trocknung sowie Computer-tomografie verwendet. Unterschiede zeigen sich bisher nicht nur zwischen den einzelnen Familien, sondern auch zwischen den Arten. So weisen Heterocentrotus mammillatus und Echinometra mathaei unterschiedliche Ausprägungen von Wachstumsringen auf, während Phyllacanthus imperialis trotz ähnlichem Habitat wie Heterocentrotus mammillatus eher gleichmäßig angeordnete Strukturen und keine Wachstumsringe im Stereom besitzt. Insgesamt weisen die Stacheln wie auch die Schalen von Echinodermen, eine poröse Mikrostruktur (Stereom) aus Mg-Calcit mit geringen Einlagerungen (unter 1%) von organischem Material auf. Weiterhin erfolgen Untersuchungen in der Morphologie der Stacheln bzw. Schalen zwischen Warm- und Kaltwasserarten.

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30 Years of Research on Crinoids / 30 Jahre Crinoidenforschung (Keynote)

HANS HESS

Naturhistorisches Museum Basel; Im Gerstenacker 8, 4102 Binningen, Schweiz; e-mail: [email protected]

1978 erschienen die drei Bände von Part T, Echinodermata 3 (Crinoidea) des Treatise on Invertebrate Paleontology. Die Revision unter Leitung von W. I.

AUSICH ist im Gang, und das Manuskript von Band 3 (Articulata, Mesozoikum - heute) liegt nun vor. Die wesentlichen Fortschritte und Aenderungen in der Systematik werden kurz vorgestellt und mit einigen Beispielen, insbesondere von Trias-Crinoiden, illustriert.

The three volumes of Part T, Echinodermata 3 (Crinoidea) of the Treatise on Invertebrate Paleontology were published in 1978. The revision under the

Editorship of W. I. AUSICH is ongoing, and the manuscript of the revised vol. 3 (Articulata, Mesozoic-Recent) is now available. Significant progress in systematics has been made and the major changes are briefly presented, illustrated by examples of Triassic crinoids and others.

20 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Pattern formation in starfish: arm stumps, regeneration models, and evolution (Poster)

FREDERICK H.C. HOTCHKISS

Marine and Paleobiological Research Institute, PO Box 1016, Vineyard Haven, MA 02568 USA; e- mail: [email protected]

Objective: To interpret a Linckia laevigata and an Asterias rubens that have an arm stump that healed and did not regenerate the missing arm (specimens from nature). Method: Interpret using positional information theory and regeneration models from amphibians, insects, and planarians. Observations: Wound closure was by symmetrical midline joining through distal folding of both sides of the arms. In contrast, in specimens that regenerate the arm, wound closure is by downward sliding of the aboral surface. First finding: Failure of these arm stumps to regenerate indicates symmetrical positional information in the starfish arm and indicates wound closure and healing along the line of symmetry. Thereby no positional value gaps/confrontations were formed, and intercalary gap-filling regeneration was not stimulated. Second finding: The best fit model for starfish arm regeneration is the distalization followed by intercalation model of AGATA et al. (2003, 2007).

Proposals: It is proposed that arm regeneration is initiated by formation of a distalized signaling center just behind the regenerated terminal plate. The signal induces distal intercalary regeneration. It is further proposed that distal intercalary gap-filling growth is the normal growth mechanism of the arm; thus regeneration reestablishes the growth mechanism of intact arms. According to this idea, there is always a positional gap between the last-formed section of arm and the distalized signaling center located just behind the terminal plate; this gap is never bridged, and arm growth is indeterminate. Evo-Devo Extrapolation: Mutations could cause the signaling center behind the terminal plate to induce formation of additional or fewer series of plates in the arm. This may help to explain the sudden origins and losses of novel plate series such as the sublateral podial plates of Paleozoic Eopentaroida. Once the organism possesses this type

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of signaling mechanism, then mutations could cause it to function in more places than just behind the terminal plate. Thus plate series that intercalate in other parts of the body wall could have this type of origin (such as asteroid actinal intermediate plates, intermarginal plates, dorsolateral plates, etc.). Mosaic evolution and recurrent appearance/loss of intercalary plates within and between echinoderm lineages are expected under this proposal.

References

AGATA, K., T. TANAKA, C. KOBAYASHI, K. KATO & Y. SAITOH, 2003. Intercalary regeneration in planarians. Developmental Dynamics 226: 308-316.

AGATA, K., Y. SAITO & E. NAKAJIMA, 2007. Unifying principles of regeneration I: Epimorphosis versus morphallaxis. Development, Growth and Differentiation 49: 73-78.

22 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Bdellacoma in the Hunsrück Slate (Lower Devonian, Germany): reidentification of Urasterella verruculosa (Asteroidea, Bdellacomidae) (Poster)

FREDERICK H.C. HOTCHKISS1 & ALEXANDER GLASS2

1 Marine and Paleobiological Research Institute, PO Box 1016, Vineyard Haven, MA 02568 USA; e- mail: [email protected]

2 Division of Earth and Ocean Sciences, Box 90227, Duke University, Durham, North Carolina 27708

The Problem: Occurrence of Bdellacoma SALTER, 1857, in the Hunsrück Slate was mentioned by W. K. SPENCER (1940), but no specimens were cited. On the other hand, no specialist on the fauna of the Hunsrück Slate has listed Bdellacoma. The Solution: In the Natural History Museum, London, are specimens BMNH E13627 and E13630 from the Hunsrück Slate labeled Bdellacoma. The specimens have highly distinctive bursulella-type pedicellariae that confirm identification as Bdellacoma sp. Next Problem: The BMNH specimens also match the Hunsrück

Slate asteroid Urasterella verruculosa LEHMANN, 1957. The Solution: Examination of the holotype of U. verruculosa confirmed the presence of bursulella-type pedicellariae and establishes that it is a species of Bdellacoma. The madreporite is of asteroid type: aboral, interradial, convex, large, and with madreporiform markings. The Hunsrück Slate asteroid Urasterella verruculosa is reidentified as Bdellacoma verruculosa n. comb. (Asteroidea, Bdellacomidae). The fossil record of Bdellacoma, including isolated pedicellariae reported as “Bursulella” ostracod microfossils, ranges from Early Silurian to Early Carboniferous, with localities in England, Gotland, Poland, Germany and USA. Improved understanding of

Bdellacoma (especially SUTTON et al. 2005) permits identifying specimens that do not have pedicellariae, namely unpublished Bdellacomidae sp. collected by

THAYER (1972) from the Upper Devonian Genesee Group, Cortland County, New York: Yale Peabody Museum 5-rayed YPM 211365 with bursulella-type pedicellariae evident; YPM 204172-204186 with pedicellariae not evident but otherwise resembling YPM 211365; and the remarkable YPM 211610 with 16 or 17 rays and preserving at least one bursulella-type pedicellaria. Occurrence of Bdellacoma in the Lower Silurian (Llandovery) Gutterford Burn Starfish Bed,

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Pentland Hills, Scotland, was mentioned by SPENCER (1940: 529), but no specimens were cited. Significance: Bdellacoma is confirmed present in the

Hunsrück Slate. Assignment of Bdellacoma to the Asteroidea by SUTTON et al. (2005) is supported, except that future research should investigate the sublateral plates described by SPENCER (1940) in Bd. vermiformis; sublaterals are not yet recognized in Bd. verruculosa. Acknowledgements: The importance of papers by

BOCZAROWSKI (2001) and by SUTTON et al. (2005) cannot be overstated. We thank

D. N. LEWIS (BMNH), C. MACCLINTOCK (YPM), M. SANDER (Institute for

Paläontologie, Rheinische Friedrich-Wilhelms Universität, Bonn), and A. B. SMITH

(BMNH) for loans/visits; D. CLAIN, M. CLAIN, A. P. HOTCHKISS, M. SGAN and G.

SGAN for support; and D. NICHOLLS for composing the poster. The poster was prepared for the 12th IEC, August 2006, Durham, New Hampshire; contribution No. 23 of Project Nahecaris; contribution No. 2 of MPRI; this abstract is new for this workshop.

References

BOCZAROWSKI, A. 2001. Isolated sclerites of Devonian non-pelmatozoan echinoderms. Palaeontologia Polonica No. 59: 1-219.

SPENCER, W. K. 1940. A monograph of the British Palaeozoic Asterozoa, Part. X, pp. 495-540, pls. 33-37, Palaeontographical Society for 1940.

SUTTON, M. D, D. E. G. BRIGGS, DAVID J. SIVETER, DEREK J. SIVETER, & D. J. GLADWELL. 2005. A starfish with three-dimensionally preserved soft parts from the Silurian of England. Proceedings of the Royal Society, Series B 272: 1001-1006.

THAYER, C. W. 1972. Marine paleoecology of the Upper Devonian Genesee Group of New York. Ph.D. dissertation, Yale University.

24 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Index of Living and Fossil Echinoids: 1971 - present (Poster)

ANDREAS KROH

Naturhistorisches Museum Wien, Geologisch-Paläontologische Abteilung, Burgring 7, 1010 Wien, Österreich; e-mail: [email protected]

In 1978 Porter M. KIER & Mary H. LAWSON published their "Index of Living and Fossil Echinoids 1924-1970" which listed all species- and genus-level taxa erected since the publication of the “Essai de Nomenclature Raisonnée des

Echinides” by Jules LAMBERT & Paul THIÉRY (1909-25). Both books have been an outstanding resource for taxonomic and (palaeo-)biodiversity research and are among the most consulted volumes by echinoid specialists. KIER & LAWSON (1978) in the introduction of their Index announced a supplement for the years 1971-75 but this, apparently, has never been published. Data collection for such an supplement was started in 1999 by the present author, while doing research on the biogeography of Cenozoic echinoids, but soon was extended to include all epochs. The list was prepared by culling the recent echinoderm literature for new taxa and by cross checking this list with the Zoological Record and various other offline and online sources. Each paper included was consulted in the original. Citations and taxa before 1971 were included if they were absent in LAMBERT & THIÉRY (1909-25) and KIER & LAWSON

(1978). In contrast to KIER & LAWSON (1978) the present index also includes detailed locality data and repository of the types. With the increasing availability of taxonomic databases such as the Zoological Record or the Paleobiology Database, the need for published indices may seem obsolete. Many of these resources, however, are incomplete and/or accessible for subscribers only. Despite the benefits of an online database, there are many instances where an old-fashioned catalogue is quite handy, be it in the storage area of a museum or during field work in remote places.

KIER, P.M. & LAWSON, M.H. 1978. Index of living and fossil echinoids 1924-1970. – Smithsonian Contributions to Paleobiology, 34: 1-182.

LAMBERT, J. & THIÉRY, P. 1909-1925. Essai de Nomenclature Raisonnée des Echinides. – iii+607 pp. Chaumont (L. Ferrière).

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Crinoids from the Tithonian of the Klippenbelt (Jurassic, Austria) (Poster)

ANDREAS KROH1 & ALEXANDER LUKENEDER2

1, 2 Naturhistorisches Museum Wien, Geologisch-Paläontologische Abteilung, Burgring 7, 1010 Wien, Österreich; e-mail: [email protected] 1, [email protected] 2

The Nutzhof section, 5 km north of Hainfeld, comprises an 18 metre long succession of strongly tilted, inverse, well-bedded marls and limestones of the Blassenstein Formation. At the base marls and marly limestone bands dominate, being replaced by increasingly pure limestones towards the top. Based on preliminary data from ammonites, palaeomagnetics, nannofossils and calpionellids (LUKENEDER, in press; PRUNER et al., in press; REHÁKOVÁ et al., in press) the section comprises a continuous succession of Lower Tithonian to Middle Berriasian strata. Microfacies analysis of the more strongly lithified parts of the section revealed high abundances of crinoidal remains, particularly in the lower part. Based on cross section shape these fragments could be tentatively assigned to saccocomid crinoids. In order to obtain three-dimensional specimens of the crinoids, as well as other microfossils commonly observed in the thin sections (namely foraminifera, ostracods, rhyncholiths, small aptychi, ophiuroid remains, etc.) the marly parts of the succession (i.e. the lower 8 m) were sampled intensely. Employing a combination of traditional (hydrogene-superoxide) as well as specialized agents (the tenside Rewoquat) it was possible to disaggregate the bulk samples and clean the microfossils. A first survey of the residues shows high abundances of the pelagic crinoid

Saccocoma tenella (GOLDFUSS, 1831) in the lower 6 metres of the section. The crinoids are represented by isolated radial and brachial ossicles mainly. The former being characterized by their arrow-head like shape with serrated edges and coarse reticulate sculpture, the latter by its disc-like wings (in the proximal brachials). Above, calyxes of the cyrtocriid Phyllocrinus belbekenis ARENDT, 1974 appear, while saccocomid remains become rare and vanish. Additionally, juvenile columnals tentatively referred to the Margocrinus/Balanocrinus-group sporadically occur in the lower part of the succession.

26 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

In the upper part of the section bulk sampling was unsuccessful due to low clay content and high lithification of the rocks. Thin sections, however, show that the saccocomid-rich microfacies is replaced by pure calpionellid limestone, that can be interpreted as more distally situated pelagic facies. Similar successions have been documented from other parts of Austria, Czech Republic and Poland as well. It is likely that the observed changes in lithology and microfauna are related to changes of current patterns, possibly caused by geodynamically induced palaeogeographic changes and basinal deepening.

24.-26. October 2008 27 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Temperature effect on feed consumption, absorption and assimilation efficiencies and production in the sea urchin Strongylocentrotus intermedius (Poster)

JOHN M. LAWRENCE1,*, XUEBIN CAO2, YAQING CHANG2, PING WANG2, Y. YU2, ADDISON L.

LAWRENCE3, STEPHEN A. WATTS4

1 Department of Biology, University of South Florida, Tampa Florida 33620, U.S.A; e-mail: [email protected]

2 Key Laboratory of Mariculture and Biotechnology, Dalian Fisheries University, 52 Heishijiao Street, Dalian, Liaoning Province, 116023, People’s Republic of China

3 Texas Agrilife Research, Texas A&M System, 1300 Port Street, Port Aransas, Texas 78373, U.S.A.

4 Department of Biology, University of Alabama at Birmingham, Birmingham, Alabama 35294, U.S.A.

Strongylocentrotus intermedius is one of the most economically important species of sea urchins and has great potential for aquaculture. It is essential to understand how temperature affects utilization of nutrients and production as controlled temperature may be an important expense in land aquaculture and affect optimization of production. We maintained large (5 cm test diameter) S. intermedius in the laboratory and fed them a formulated feed from 28 February when the gonads were small (gonad index, GI, ~10) and sea water temperature was low (7 °C) until 27 June when sea water temperature was high (24 °C) under five temperature treatments (environmental temperature, 7, 12, 17 and 22 °C. Temperature had little effect on rate of consumption of food or absorption efficiency. Temperature had little effect on assimilation efficiency except for a significant decrease at 22 °C. The gonad index was consistently least at 22 °C. Organic matter production in the gonads was greatest and earliest (April) at 12 and 17 °C. Because S. intermedius has a great ability to acclimate to temperature, temperature should not be a great concern in land aquaculture. Under the conditions of this experiment, S. intermedius in culture produced gonads of marketable size (gonad index >15) within one month.

28 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Amount of arm loss and rate of arm regeneration by Luidia clathrata (Echinodermata: Asteroidea) (Poster)

J.M. LAWRENCE1 & C.M. POMORY2

1 Department of Biology, University of South Florida, Tampa, Florida 33620, U.S.A.; e-mail: [email protected]

2 Department of Biology, University of West Florida, Pensacola, Florida 32514, U.S.A.

Three contiguous arms of Luidia. clathrata were amputated. The initial mean lengths (±SE) of the two intact arms were 57.36 ± 2.13 and 57.38 ± 1.97 mm. The initial mean lengths of the amputated arm stumps were 12.18 ± 0.59 (proximal), 25.91 ± 1.67 (medial) and 42.25 ± 1.37 (distal) mm. Buds appeared on all amputated arms after approximately 8 days. After 54 days proximal, medial and distal regenerated arms had mean lengths (±SE) of 10.96 ± 0.90, 7.69 ± 0.61 and 3.99 ± 0.38 mm, mean dry weights (±SE) of 41.75 ± 6.79, 20.95 ± 3.18, 8.07 ± 1.13 mg, and mean amounts of organic matter (pooled samples) of 6, 3 and 1 mg, respectively. All three arm positions are statistically different from one another (P<0.001). Appearance of arm buds is the first phase of arm regeneration and is independent of level of amputation. Growth of regeneration is the second phase and is dependent on level of amputation. Studies have reported rate of growth of intact arms and of arms regenerating from the disc also declines as the asymptotic length of the arm is reached. This suggests similar mechanisms of control of growth occur in all three situations, all depending on the relative position of the regenerating arm tip.

24.-26. October 2008 29 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Seastars (Asteroidea) from Madagascar (Vortrag)

HORST MOOSLEITNER

Organismische Biologie, Universität Salzburg, Hellbrunnerstr. 34, 5020 Salzburg, Austria; e-mail: [email protected]

About a small collection of shallow water seastars from NW-Madagascar

The inspection of a small collection of seastars, Mr. Hinterkircher (Munich) had sampled 2007 in NW-Madagascar, showed astonishing results:

The collection included not only seastars known from that region (LORIOL 1885), like Protoreaster linckii (DE BLAINVILLE, 1834), Fromia milleporella (LAMARCK,

1816) and Monachaster sanderi (MEISSNER, 1892), but some species not known from there, too.

One of them was Fromia indica (PERRIER, 1869). Its discovery is astonishing because this species was hitherto known only from places east of the

Maldives and the Lakkadives (CLARK & ROWE 1971, MOOSLEITNER 1997, SASTRY 2007), and these islands lie more than 3000 km away from Madagascar. One specimen of Aquilonastra sp. could not be classified as it did not confirm to any species known. It was sent to Mr. O’LOUGHLYN (Australia), a specialist for that genus, for inspection. The most astonishing result was the discovery of the seastar Ophidiaster granifer, LÜTKEN 1872. The determination first seemed to be impossible, as the seastar did not confirm to the description of any species known from the Indian

Ocean. Frank ROWE gave me the crucial tip to include Ophidiaster granifer and O. armatus into my investigations. He was right, the first of the two species was applicable. The seastar from Madagascar fits not only to the key for that species, but also in all important details to O. granifer. Exact details are given in the presentation. O. granifer was hitherto known from the easternmost part of the Indian Ozean and from the western–central Pacific (to Japan und to the island of the South Pacific). This finding expands the range of occurrence across the whole Indian Ocean to western Madagascar.

30 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Ophidiaster granifer

References

CLARK, A.M. & F.W.E. ROWE, 1971. Monograph of Shallow-Water Indo-West Pacific Echinoderms. Trusteet of the British Museum (Nat. Hist.), London. 238 pp.

DE LORIOL P., 1885. Catalogue Raisonne des Echinodermes recueillis par M.V.de ROBILLARD a l’ile Maurice, II Stellerides. Mémoires de la Société de Physique et d'Histoire Naturelle de Genève 29/4: 1-84.

MOOSLEITNER, H., 1997. Shallow water Asteroidea (Echinodermata) from the coral reefs of the Maldive Islands. Revue française d'Aquariologie 24: 3-14.

SASTRY, D.R.K., 2007. Echinodermata of India: An annotated list. Records of the Zoological Survey India, Occassional Paper 271: 1-387.

24.-26. October 2008 31 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Morphologie und Faziesverteilung des indo-pazifischen Seeigels Jacksonaster depressum (Vortrag)

JAMES NEBELSICK1 & ATEF ABD EL-HAMIED A. S. ELATTAAR2 1 Institut für Geowissenschaften, Universität Tübingen, Sigwartstrasse 10, 72076 Tübingen, Deutschland; e-mail: [email protected] 2 Geology Department, Sohag Faculty of Science, Sohag University, Sohag, P. O. Box 82524, Egypt; e-mail: [email protected]

Jacksonaster depressum ist ein irregulärer indopazifischer Seeigel der Ordnung Clypeasteroida. Das bearbeitete Material stammt aus der Nördlichen Bucht von Safaga, Rotes Meer, Ägypten, wo dieser Seeigel am häufigsten in schlammigen Sanden in Tiefen von 20 – 50 Meter anzutreffen ist. Der Seeigel erreicht Größen von bis zu 3 cm und zeigt die für diese Ordnung charakteristische abgeflachte Schale mit internen Stützelementen, die in dieser Gattung auf den äußeren Rand des Tieres beschränkt sind. Diese Seeigel sind kaum untersucht worden und leben wahrscheinlich infaunal als Detritivor.

Wir haben die Skelettmorphologie, Taphonomie und ökologische Verteilung von zahlreichen Exemplaren innerhalb des Untersuchungsgebietes studiert. Hierbei wurden sowohl lebende wie auch tote Schalen inkludiert, die durch verschiedene Methoden aufgesammelt worden sind. Vermessungen von spezifischen morphologischen Merkmalen wurden durchgeführt, z.B. die allgemeinen Schalenparameter, Apikalsystem, Petalodium, Lage und Ausmaß des Peristoms und Periprokts usw.. Außerdem wurden REM - Aufnahmen von Stacheln, Pedicellarien, Tuberkeln wie auch von den Poren gemacht. Zusätzlich wurden Differenzierungen im Stereom notiert. Die Variation dieser morphologische Parameter zwischen lebenden und toten Tiere wurde analysiert, und zwar innerhalb der Gesamtpopulation, wie auch zwischen unterschiedlichen Probenpunkten und Faziestypen.

32 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Taphonomie von Jacksonaster depressum (Echinoidea: Clypeasteroida) aus dem Roten Meer (Poster)

JAMES NEBELSICK1 & ATEF ABD EL-HAMIED A. S. ELATTAAR2 1 Institut für Geowissenschaften, Universität Tübingen, Sigwartstrasse 10, 72076 Tübingen, Deutschland; e-mail: [email protected] 2 Geology Department, Sohag Faculty of Science, Sohag University, Sohag, P. O. Box 82524, Egypt; e-mail: [email protected]

Jacksonaster depressum ist ein irregulärer indopazifischer Seeigel der Ordnung Clypeasteroida. Das bearbeitete Material stammt aus der Nördlichen Bucht von Safaga, Rotes Meer, Ägypten, wo dieser Seeigel am häufigsten in schlammigen Sanden in Tiefen von 20 – 50 Meter anzutreffen ist. Dieser Seeigel zeigt die für diese Ordnung charakteristische abgeflachte Schale mit internen Stützelementen, die für die Erhaltung signifikant sind. Die Taphonomie dieser Seeigel ist bis jetzt kaum untersucht worden. Die taphonomische Zerstörung der Schale könnte sowohl an „kompletten“ als auch an Fragmenten erfolgen. Die taphonomische Merkmale können anhand von der Veränderungen von Oberflächenmerkmalen wie Tuberkulation oder Stereomtypen charakterisiert werden. Eine Vielzahl von taphonomischen Prozessen konnte beobachtet werden, u.a. Disartikulation, Fragmentation, Inkrustation wie auch Bioerosion. Angriffe durch Räuber spielen auch eine wichtige Rolle bei der Zerstörung der Schale und der Produktion von Schalenfragmenten. Eine Reihe von Fragen stellen sich bezüglich der Taphonomie von Jacksonaster: 1) Wie beeinflusst die Morphologie von Jacksonaster die beobachteten taphonomischen Signaturen? 2) Welche taphonomischen Prozesse sind für die Zerstörung wie auch Erhaltung dieser Gattung von Bedeutung, und 3) wie ist das Erhaltungspotential dise Seeigel in Vergleich zu anderen Clypeasteroiden des Roten Meeres, wie Clypeaster, Echinodiscus, Fibularia und Echinocyamus? Charakteristisch für Jacksonaster ist z.B. ihre Erhaltung als ringförmige Schalenreste, ein Erhaltungsform, die bei den anderen oben aufgeführten Seeigeln kaum in Erscheinung tritt.

24.-26. October 2008 33 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Skelett-Endosymbionten fossiler und rezenter Echinodermen: Von Bioklaustrationen zu komplexen Strukturen (Vortrag)

CHRISTIAN NEUMANN

Museum für Naturkunde, Humboldt Universität Berlin, Invalidenstrasse 43, D-10115 Berlin, Deutschland; e-mail: [email protected]

Als Symbiose wird das Zusammenleben von Organismen unterschiedlicher Artzugehörigkeit bezeichnet. Evolutionsbiologisch sind Symbiosen sehr interessant, denn sie erfordern in der Regel einen hohen Grad an Spezialisierung mindestens bei einem der beiden Partner. Verschiedene Fragen tun sich auf: Wie sind diese oft komplexen Wirt/Symbiont-Systeme entstanden? Wie haben sich Symbiosen über geologische Zeiträume verändert? Inwiefern sind dabei Prozesse der Koevolution beteiligt? Fragen, die bisher schwer zu beantworten waren, denn Symbiosen sind im Fossilbericht kaum überliefert. Ausnahmen sind solche Interaktionen, die Spuren an den erhaltungsfähigen Hartteilen (Schalen, Skelette) hinterließen. Echinodermen bieten sich modellhaft für die Untersuchung fossiler Symbiosen an, denn bei ihrem Innenskelett handelt es sich um ein lebendes Gewebe, welches direkt und plastisch auf Irritationen (wie etwa einen Parasitenbefall) reagieren kann. Diese Gewebeveränderungen sind fossil erhaltungsfähig und somit vom Paläontologen interpretierbar. So versuchen Stachelhäuter oft, unerwünschte Fremdorganismen, die sich auf ihrer Körperoberfläche festgesetzt haben, mit ihrem Skelettgewebe zu überwachsen (Neoplasie, Callus-Bildung). Diese Abwehrreaktion machen sich manche Symbionten zu Nutze: Das „Überwuchert-werden“ durch das Kalzitskelett des Wirtes bietet Vorteile, wie etwa einen besseren Schutz vor Fressfeinden. So sind Crinoiden bevorzugte Wirte von Symbionten. Als passive Filtrierer besitzen sie einen Stiel, der eine günstige Position des Fressapparates innerhalb der Wasserströmung gewährleistet. Für andere filtrierende Organismen bilden die Stiele eine willkommene Struktur, um sich darauf festzusetzen und so „kostenlos“ in die Meeresströmung zu gelangen(„secondary tiering“). Als Abwehrreaktion versucht die Seelilie oftmals, die unerwünschten Untermieter mit ihrem Skelett geschwulstartig zu überwachsen. Bereits im Paläozoikum passen

34 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research sich Vertreter unterschiedlichster taxonomischer Zugehörigkeit an diese Wirtsreaktion an und gehen zu einer endosymbiontischen Lebensweise über. Um eine vollständige Einkapselung durch das Wirtsgewebe zu verhindern, bedarf es gewisser Anpassungen von Seiten des Symbionten. So entstanden schließlich aus einfachen Skelett-Einbettungen (Bioklaustrationen) hochkomplexe gallenartige Strukturen, welche physiologischen Prozesse des Symbionten unterstützen. Die Bildungs- und Funktionsweise solcher Gallen lässt sich beispielhaft an Gallen parasitischer Schnecken (Fam. Eulimidae) in den Stacheln von Lanzenseeigeln(Cidaroida) untersuchen: Hier nimmt der Parasit wesentlichen Einfluss auf die Art und Weise der Gewebereaktion seines Wirtes. Je nach Schneckenart sind die Gallen in ihrer Struktur höchst unterschiedlich und oftmals hochkomplex. Es ist anzunehmen, dass die Morphologie der Gallen vom Genom der Schnecken kontrolliert wird, obwohl die Galle selbst eine Bildung des Wirtes ist („extended phenotype“). Sollte diese Hypothese zutreffen, offenbart sich darin die Chance, anhand fossiler Gallen die Phylogenie ihrer nicht überlieferten Bewohner zu rekonstruieren und schließlich durch den Vergleich mit der Phylogenie ihrer Wirte koevolutionäre Prozesse (z.B. Kospeziation) nachzuweisen. Mit Hilfe Computer-tomographischer Methoden wurden dreidimensionale Einblicke in die Internstruktur des Skeletts befallener fossiler und rezenter Echinodermen gewonnen, ohne dass dabei die Objekte zerstört werden mussten. Dabei wurde sowohl Neutronen- als auch hochauflösende Synchrotronstrahlung eingesetzt und spektakuläre Einblicke in das verborgene Innenleben längst vergangener Lebensgemeinschaften gewonnen.

24.-26. October 2008 35 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Echinoids from the Neogene of Portugal mainland: Systematics, Palaeoecology, Palaeobiogeography (Vortrag)

PEDRO PEREIRA

Universidade Aberta, Rua Fernão Lopez, 9, 1° Esa, 1000-132 Lisboa, Portugal; e-mail: [email protected]

The Portuguese Neogene echinoid fauna is reviewed in order to establish a taxonomically standardised echinoid database. 41 taxa, including one new genus and new species, from the Burdigalian and Middle Miocene of the Lower Tagus Basin and Middle Miocene of Algarve and two taxa from the Middle Pliocene of Mondego Basin are described. 25 taxa are reported for the first time from Portugal mainland, while 23 of the previously reported Miocene species were excluded (for different reasons) from the accepted species list. This study uses echinoid diversity and the present geographic distribution of genera that already existed during Neogene to assess climatic changes in the Neogene of Portugal. The presently available data suggest tropical conditions for the Burdigalian (Neogene Cycle I) and Lower Langhian (Early Neogene Cycle II) of the Lower Tagus Basin, followed by a temperature drop, coincident with the end of the Upper Burdigalian-Lower Langhian Climatic Optimum. This temperature drop is reflected by the reduction of the echinoid diversity, the disappearance of several species, particularly among clypeasteroids and cassiduloids, and by the appearance and following dominance of euechinoids (mainly, Psammechinus) and Echinocyamus. In Algarve the echinoid data suggest, for the Middle Miocene (Neogene Cycle II), tropical conditions for that region and, for the Upper Tortonian (Neogene Cycle III), probably subtropical conditions. During the Neogene Cycle II there was a clear temperature gradient between Algarve (warmer) and Lower Tagus Basin (less warm) which is obviously related to the more southerly position of Algarve. For the Middle Pliocene (Neogene Cycle III) of the in Mondego Basin the echinoid data are two few to allow a palaeoclimate interpretation, but gastropod data suggest subtropical conditions.

36 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

The echinoid fauna of Portugal mainland is closely related to that of the Mediterranean region. In fact, the biogeographic investigation of the Portuguese echinoid fauna shows that a major part of the Portuguese species is composed by immigrants from the Mediterranean area (42.9% of the fauna in the Burdigalian and 60.9% in the Middle Miocene). Endemism is low during Miocene, with endemic species not exceeding 25% of total Portuguese echinoid fauna.

24.-26. October 2008 37 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Phylogenetic relationships within Echinodermata based on mitochondrial sequence data (Vortrag)

MARLEEN PERSEKE1, GUIDO FRITZSCH2, DETLEF BERNHARD1, PETER F. STADLER2 & MARTIN

SCHLEGEL1

1 Institute of Biology II, Molecular Evolution and Animal Systematics, University of Leipzig, Talstrasse 33, 04103 Leipzig, Deutschland; e-mail: [email protected]

2 Interdisciplinary Center for Bioinformatics, University of Leipzig, Haertelstrasse 16-18, 04107 Leipzig, Deutschland

Reconstruction of phylogenetic relationships within Echinodermata seems to be difficult using mitochondrial datasets. Especially the ophiuroids are strongly different compared to the other echinoderm groups. However, only two complete mitochondrial gene orders of ophiuroids are known. In our analyses this dataset was expanded to a total of six mitochondrial genomes of ophiuroids with 3 new families: Ophiocomidae, Amphiuridae and Gorgonocephalidae. With these new mitochondrial gene orders, a consensus gene order of Ophiuroidea becomes apparent with only moderate rearrangement events within this group. Additionally, new complete mitochondrial genomes from other echinoderm groups were determined: an irregular echinoid and a further holothuroid. Both mitochondrial genomes show no gene order rearrangements compared to the known gene order of echinoids.

Using the amino acid sequences of this extended dataset for phylogenetic analyses, the Echinodermata and all five echinoderm groups are clearly recovered as monophyletic groups. Within the Echinodermata only the relationships between Echinoidea, Asteroidea, and Holothuroidea are unstable depending on the analyses method used. Most analyses show Holothuroidea as sistergoup of a clade consisting of Echinoidea and Asteroidea. However, the basal branching of Ophiuroidea within echinoderms persists in all analyses with strong support and keeps the main problem within the mitochondrial dataset.

38 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

New insights into early diversification of sea cucumbers as inferred from calcareous ring material (Ordovician/Silurian, Baltoscandia) (Vortrag)

MIKE REICH

Geowissenschaftliches Zentrum der Universität Göttingen, Museum, Sammlungen & Geopark, Goldschmidtstr. 1-5, 37077 Göttingen, Deutschland; e-mail: [email protected]

Compared to other modern Echinodermata, the early evolutionary history of holothurians is poorly understood. In part, this is due to their disjunct endoskeleton with ossicles and calcareous ring elements, which are released following decomposition of the surrounding tissue. Newly sampled Silurian sections on Gotland, Sweden, yield well-preserved echinozoan echinoderms, including more than 1000 isolated calcareous ring elements. The studied material is Telychian to Ludfordian (Early to Late Silurian) in age and reveals the presence of several new taxa of arthrochirotid (extinct) and apodid sea cucumbers as well as stem group representatives. The calcareous pharyngeal ring (CR) is a defining feature of sea cucumbers and is possessed by nearly all extant species. The CR supports the pharynx, the water vascular system and tentacle ampullae in part, as well as providing insertion points for the muscles. Therefore calcareous rings are distinct for systematics and phylogeny, and important evolutionary structures. The new material shows, that the Holothuroidea diversified significantly through Late Ordovician and Silurian times.

24.-26. October 2008 39 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

A nearly articulated holothurian from the Santonian (Cretaceous) of England (Poster)

MIKE REICH

Geowissenschaftliches Zentrum der Universität Göttingen, Museum, Sammlungen & Geopark, Goldschmidtstr. 1-5, 37077 Göttingen, Deutschland; e-mail: [email protected]

With the exception of the Maastrichtian, only scarce data exist on fossil holothurians from the Upper Cretaceous, especially the Santonian. Up to now, only two records have been mentioned worldwide: (1) from the Münsterland area, Germany, and (2) from the Montsec area, Spain; both referred to the Apodida and Dendrochirotida. Here I report the discovery of a nearly articulated holothuroid specimen from the “Upper Chalk” (Santonian; BMNH E 48764) of England (unfortunately exact locality unknown). The unique specimen shows two different types of ossicles: (A) tables and (B) buttons, demonstrating aspidochirote affinities, very likely to the family Holothuriidae. Concerning the morphology of the buttons the new find share characte- ristics with tube feet and papillae ossicles of Holothuria (Mertensiothuria), H. (Microthele), and H. (Semperothuria) and H. (Theelothuria). According to the decomposition of the surrounding tissue after death of the animal, only a part of the former body skeleton is preserved; the anterior part of the skeleton (calcareous ring) is missing. Due to the well-preserved body-wall ossicles and the formation of the ossicles of the new find, I can exclude a digestion and excretion of the sea cucumber by any predator. After comparison with modern members of the Holothuriidae, we can probably speculate that the body of our new Santonian species was also cylindrical, elongate and scattered with numerous podia over the entire body, dorsally as papillae, ventrally as cylindrical tube feet. A wider distribution of the new genus / species can also be assumed, because comparable long button-shaped sclerites were also known from Early and Late Maastrichtian chalk sediments of the islands of Møn (Denmark) and Rügen (NE Germany) as well as the Baltic Sea.

40 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Holothuriid sea cucumbers are globally distributed and primarily deposit-feeders. Modern members of the Holothuriidae inhabit the tropical eulittoral, usually coral reefs and adjacent sandy areas; only a few species were known from temperate shallows and deeper waters (bathyal depths). Considering how important aspidochirote sea cucumbers are today (nearly ¼ of all known modern species), their fossil record is meagre and poorly understood. The phylogeny of the family Holothuriidae was recently investigated by KERR et al. (2005) and SAMYN et al. (2005), unfortunately without detailed consideration to the fossil record. The origin and early diversification of the Holothuriidae can be located in the Late Palaeozoic. There is still a hidden Mesozoic diversity since detailed investigations are missing or sparse.

References

KERR, A.M., JANIES, D.A., CLOUSE, R.M., SAMYN, Y., KUSZAK, J. & KIM, Junhyong, 2005. Molecular Phylogeny of Coral-Reef Sea Cucumbers (Holothuriidae: Aspidochirotida) Based on 16S Mitochondrial Ribosomal DNA Sequence. Marine Biotechnology 7/1: 53-60.

SAMYN, Y., APPELTANS, W. & KERR, A.M., 2005. Phylogeny of Labidodemas and the Holothuriidae (Holothuroidea: Aspidochirotida) as inferred from morphology. Zoological Journal of the Linnean Society 144/1: 103-120.

24.-26. October 2008 41 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Unusual holothurians (Echinodermata) from the Late Ordovician of Sweden (Poster)

MIKE REICH

Geowissenschaftliches Zentrum der Universität Göttingen, Museum, Sammlungen & Geopark, Goldschmidtstr. 1-5, 37077 Göttingen, Deutschland; e-mail: [email protected]

Compared to other modern echinoderms, the early evolutionary history of sea cucumbers is poorly understood. In part, this is due to their disjunct endoskeleton with ossicles and calcareous ring elements, which are released following decomposition of the surrounding tissue. Newly sampled öjlemyr flints from the northwestern part of Gotland, Sweden, yield well-preserved echinozoan echinoderms, including holothurian ossicles. The studied material is Ashgill (upper Pirgu stage) in age and reveals the presence of several new or poorly known taxa of elasipodid, aspidochirote, and apodid holothurians or stem group representatives. The minute ossicles, around 70–400 µm in length / diameter, exhibit an impressive morphology with arms, spires, perforations, teeth etc. The new material differs from previous described Palaeozoic, Mesozoic and modern material by unusual symmetry. This study shows that the Holothuroidea diversified significantly through Late Ordovician times.

42 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Signs of predation on Middle Triassic crinoids form Poland (Poster)

MARIUSZ A. SALAMON1 & PRZEMYSŁAW GORZELAK2

1, 2 University of Silesia, Faculty of Earth Sciences, Department of Palaeontology & Biostratigraphy, Bedzinska Str. 60, 41-200 Sosnowiec, Poland; e-mail: [email protected] , [email protected]

Observations of both extant and fossil crinoids indicate that the range of their natural predators includes fishes, arthropods, asteroids and echinoids (Baumiller et al., 2008 and the references cited therein). Using large samples of disarticulated fossil crinoids from numerous Middle Triassic localities, we looked for evidence of bite marks produced by predators. A large proportion of columnals possessed characteristic scratches and pits imprinted in stereom. Some of these signs are very similar to those found on and Late

Cretaceous crinoids and ascribed to predation (comp. JAGT 1999, pl. 35, figs. 5,

10; JAGT & SALAMON 2007, pl. 3, fig. 4; GORZELAK & SALAMON in press, fig. 2). These scratches are relatively shallow (0.1-1.3 mm deep), but clean, cuts in the stereom of the ossicle surface. They are 0.1-1.1 mm wide and up to 2.1 mm long. The pits are usually perpendicular to the surface of the ossicle and their outline is circular or slightly oval. Their diameter varies form 0.15 to 0.7 mm.

A,B – bite marks (dotted line) and C – traces of regeneration (arrow) on Middle Triassic crinoids from Poland [Gogolin Beds (Lower Anisian, Muschelkalk); Upper Silesia area). Scale bars equal 1mm.

24.-26. October 2008 43 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

They are relatively shallow (0.2-1.4 mm) and taper towards the bottom. In most ossicles these marks occur on the external part of the lateral faces of well preserved columnals/pluricolumnals which are distict from abrasion suggesting that the crinoid material was fresh and probably live, when the marks were made and we treat it as evidence of predation. This is also consistent with the fact that certain ossicles, which possess bite marks, comprise also some pathological anomalies and regeneration traces. It has been argued that predation increases over geological time should result in increases in defensive adaptations in prey taxa. It seems that most of the major functional "experiments" in post-Paleozoic articulate crinoids that occurred during the Triassic can be described in terms of predation including planktonic microcrinoids (roveacrinids), pseudoplanktonic stalked crinoids (traumatocrinids and pentacrinids), strongly anti-predatory specialised cyrtocrinids, stalk and arm shedding benthic crinoids (isocrinids and holocrinids), free moving crinoids (paracomatulids). The high bite mark frequency on Early Mesozoic crinoids from Poland proves that the evolution of morphologies and behaviors of the above mentioned groups was stimulated by their interactions with predators.

References

BAUMILLER, T.K., MOOI, R. & MESSING, C.G. (2008): Urchins in the meadow: paleobiological and evolutionary implications of cidaroid predation on crinoids. - Paleobiology, 34/1: 22-34.

GORZELAK, P. & SALAMON, M.A. (in press): Signs of benthic predation on Late Jurassic stalked crinoids; preliminary data. - Palaios.

JAGT, J.W.M. (1999): Late Cretaceous-Early Palaeogene echinoderms and the K/T boundary in the southeast Netherlands and northeast Belgium – Part 2: Crinoids. - Scripta Geologica, 116: 59-255.

JAGT, J.W.M. & SALAMON, M.A. (2007): Late Cretaceous bourgueticrinid crinoids from southern Poland – preliminary observations. - Scripta Geologica, 134: 61-76.

44 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Are that sea urchins ???? - The strange morphology of the echinoid family Pourtalesiidae (Vortrag)

HEINKE SCHULTZ

Dorfstr. 32, 25485 Hemdingen, Deutschland; e-mail: [email protected]

The family consists of species with very strangely transformed architectures looking like a bottle or a vase rather than a pentameral echinoid. At least seven extant genera belong to the pourtalesiids, living in the deep-sea. During the Challenger-Expedition 1873 – 1876 most of them were dredged for the first time and were described by A. AGASSIZ (1881). LOVÉN (1883) and

MORTENSEN (1907, 1950) covered the pourtalesiids exhaustively – and all studies based on a very small number or even on a single specimen. Russian research expeditions have brought new specimens of abyssal species, which were described by A. MIRONOV in the seventies and nineties. This year (2008) he wrote a publication on echinoids from the Mid-Atlantic Ridge, which were dredged at depths of about 620 to 3460 m. The typical features for all species are a deep peristomial groove anteriorly, a disjunct apical system and on the oral side an interrupted interambulacrum IA 5. The strange shapes are not expressed in juveniles, but become progressively more and more obvious during ontogeny. These changing plate patterns made explanations on plate homologies rather problematic.

The Extraxial-axial Theory (EAT) by MOOI et al. (1994), DAVID & MOOI (1996, 1998), including the Ocular Plate Rule, was the base for new interpretations.

Two areas on the surface of pourtalesiids are of special interest:

1. The apical system on the aboral side: The holasteroid apical system is typically elongated, but compact. Behind the single ocular plate OIII follow towards the posterior two genital plates (G2 + G3), then the two anterior oculars (OIV and OII), the two posterior genital plates (G1 + G4) and the posterior oculars (OI and OV).

24.-26. October 2008 45 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

This scheme is modified in the members of the pourtalesiids: In the genus Echinosigra there are only gonopores in the two anterior genital plates, but the posterior genitals shift towards the posterior. They are not perforated by a pore and they stay in contact with the posterior oculars. In the genus Pourtalesia the adult specimens display another “model” of apical pattern: There are four gonopores, two in the anterior genital plates and two perforating the following ocular plates. The posterior genital plates are separated and without function, and the posterior ocular plates are again separated from the genitals and from each other.

2. The plastron on the oral side: The holasteroid plastron (interambulacrum IA 5) is usually composed by a labral plate, followed by a single sternal plate and then two episternal plates. In very young pourtalesiids this pattern is still present, but during ontogeny the adjacent ambulacral plates shift towards the midline separating the sternum from the labrum. In adult Pourtalesia (and Cystocrepis) the ambulacral plates of each column are positioned one in front of the other pushing the labrum towards the anterior into the peristomial groove.

Detailed and illustrated descriptions of the pourtalesiid species will be displayed in the book: Sea urchins of the deep-sea (in prep.).

References (more references in the book)

AGASSIZ, A., 1881. Report on the Challenger-Expedition. Zool. 3(9): 1-321.

DAVID, B. 1987. Dynamics of plate growth in the deep-sea echinoid Pourtalesia miranda Agassiz: a new architectural interpretation. Bulletin of Marine Science 40(1): 29- 47.

LOVÉN, S., 1883. On Pourtalesia. A genus of Echinoidea. Kongl. Svenska Vetenskaps- Akademiens Handlingar 19/7: 1-95.

MIRONOV, A.N., 2008. Pourtalesiid sea urchins of the northern Mid-Atlantic Ridge. Marine Biology Research 4: 3-24.

MOOI, R. & DAVID, B., 1996. Phylogenetic analysis of extreme morphologies: deep-sea holasteroid echinoids. Journal of Natural History 30: 913-953.

MOOI, R. & DAVID, B. 1998. Evolution within a bizarre phylum: Homologies of the first echinoderms. American Zoologist 38: 965-974

46 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

MOOI, R., DAVID, B. & MARCHAND, D., 1994. Echinoderm skeletal homologies.: Classical morphology meets modern phylogenetics. Proc. 18th Intern. Echinoderm Conference, Dijon 1994. A.A. Balkema, Rotterdam.

MORTENSEN, T., 1907. The Danish Ingolf-Expedition. Vol. IV Echinoidea (Part II). - 200 pp., Copenhagen.

MORTENSEN, T., 1950. A Monograph of the Echinoidea. V, 1. Spatangoida I. Protosternata, Meridosternata, Amphisternata I. Palæopneustidæ, Palæostomatidæ, Aëropsidæ, Toxasteridæ, Micrasteridæ, Hemiasteridæ. – 432 pp., Copenhagen (C. A. Reitzel).

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The early history of echinoderms: new insights from disarticulated early and middle Cambrian ossicles. (Keynote)

ANDREW B SMITH

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK; e-mail: [email protected]

Our current view of the early history of echinoderms is based largely on articulate specimens from Cambrian deposits of North America and Europe. However, Cambrian deposits in Australia and North Africa that are rich in echinoderm debris suggests this is a very incomplete picture and provide important glimpses into an unseen diversity. Furthermore this material is often exquisitely preserved, allowing the detailed study of skeletal histology and reconstruction of the soft-tissue anatomy of primitive echinoderms. Middle Cambrian columnals show unexpected diversity and provide information on the ligamental structure of the stem, while Australian material includes various elements of a pelmatozoan with uniserial arms. These finds are leading to a reinterpretation of distinction between and Blastozoa and the earliest history of the Pelmatozoa.

48 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Rhaetian ophiuroids from the Netherlands: a preliminary report (Poster)

BEN THUY1, ADIËL A. KLOMPMAKER2 & JOHN W. M. JAGT3

1, 2 Institut für Geowissenschaften, Eberhard Karls Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Deutschland; e-mail: [email protected]

3 Natuurhistorisch Museum Maastricht, de Bosquetplein 6-7, NL-6200 AW Maastricht, The Netherlands; e-mail: [email protected]

For the first time, ophiuroids are recorded from the quarry complex at Winterswijk, eastern Netherlands. Strata excavated at the quarry comprise mainly Lower Muschelkalk (Anisian, Middle Triassic) limestones, renowned for their abundant vertebrate body fossils and traces. Recently, a subrosion pipe fill in the Muschelkalk (OOSTERINK et al. 2006) yielded dark shales being dated as early to middle Rhaetian on bivalve and palynological evidence (KLOMPMAKER et al. in prep.). Also, similar, early to middle Rhaetian dark shales occur in situ in the northern part of the quarry complex (HERNGREEN et al. 2005; KLOMPMAKER et al. in prep.). The ophiuroid faunule presented herein was collected from both shale appearances of the quarry by splitting the soft slabs. Specimens generally are rare and occur individually or in groups of three to four individuals. The material consists of more than 15 complete and articulated specimens and disc fragments, plus some 30 arm fragments. Specimen size (disc diameter) ranges from about 1 to 6 mm. Most specimens are at least partially pyritised. Compaction and recrystallisation, however, tend to blur morphological details of many specimens. To make matters worse, small gypsum needles and encrustations formed some weeks after recovery of the slabs, preferably on the ophiuroid remains. However, in thoroughly pyritised specimens, brief exposure to a 20% H2O2-solution turned out to be succesfull - disc and arm fragments could be freed from the matrix and from thin clayey encrustations. This preparation method leaves specimens prone to fragmentation but it does render them perfectly exposed and suitable for SEM-examination. The prepared specimens exhibit enough morphological detail to be taxonomically assessed. First observations suggest that the faunule is monospecific. There are affinities with species of the genus Aplocoma D’ORBIGNY, 1852, and ongoing investigations aim at a precise taxonomic assignment and

24.-26. October 2008 49 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research comparison with other Rhaetian ophiuroids from Europe and especially from the lithologically comparable Penarth Group in Britain (HESS 1965; SWIFT & MARTILL 1999).

References

HERNGREEN, G.F.W., VAN KONIJNENBURG-VAN CITTERT, J.H.A. & OOSTERINK, H.W., 2005. New geological data (Middle Triassic, Rhaetian-Liassic and Oligocene) of the Winterswijk quarry, the eastern Netherlands. Netherlands Journal of Geosciences 84: 409-413.

HESS, H., 1965. Trias-Ophiuren aus Deutschland, England, Italien und Spanien. Mitteilungen aus der Bayerischen Staatssammlung für Paläontologie und historische Geologie 5: 151-177.

KLOMPMAKER, A.A., HERNGREEN, G.F.W. & OOSTERINK, H., (in prep.). Uppermost Triassic (Rhaetic) sediments and subrosion pipe age and stratigraphy at Winterswijk, the eastern Netherlands.

OOSTERINK, H.W., SIMON, T., HAGDORN, H. & WINKELHORST, H., 2006. A subrosion pipe fill in the Lower Muschelkalk, Winterswijk Quarry, Eastern Netherlands. Netherlands Journal of Geosciences 85: 293-297.

SWIFT, A. & MARTILL, D. M., 1999. Fossils of the Rhaetian Penarth Group. Palaeontological Association, Field Guides to Fossils, 9, 312 pp. London.

50 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Irregular echinoids from the Turonian Seewen Limestone of the Allgäu area (S. Germany) (Vortrag)

FRANK WIESE1 & NILS SCHLÜTER2

1, 2 Institut für Geologische Wissenschaften, Fachrichtung Paläontologie, FU Berlin, Malteserstr. 74- 100, Gebäude Haus D, 12249 Berlin, Deutschland; e-mail: [email protected]

The pelagic Seewen Limestone of the Burgberg quarry near Sonthofen, Allgäu area (Turonian-?Middle Coniacian, Alpide Helvetic Zone = north Tethyan shelf; c- dinocyst dominated biosedmentary system) is generally regarded as poorly macrofossiliferous. Nevertheless ca. 30 specimens of irregular echinoids belonging to 7 species were collected during a field campaign for isotope stratigraphic studies. Most of the taxa come from a stratigraphically limited interval (Echinoid Level, EL, 1), where Infulaster excentricus FORBES and

Sternotaxis plana (MANTELL) are dominant. In addition, (?)Sternotaxis icaunensis

(COTTEAU), Conulus subrotundus (MANTELL) and Echinocorys gravesi (DESOR) occur. EL 2 consists of two Micraster leseki (DESMOULINS), and M. ex gr. precursor/normanniae have collected from the top of the exposed rocks (EL 3). δ13C stratigraphy dates EL 1 as terminal Mid-Turonian to lowermost Upper Turonian, EL 2 as middle Upper Turonian (level Hitchwood Event = reussianum Fauna of England & Germany) and EL 3 as Lower Coniacian, also confirmed by associated inoceramids. Specifically the assemblage of AL 1 consist of taxa which are characristic of more Boreal areas such as the Anglo Paris Basin, NW Germany or – in parts – intermediate regions like northern Spain. Although single elements of the assemblage are known to occur in southern areas (e.g.

Infulaster: Calcareous Alps, Austria: KROH & WAGREICH 2007; Scaglia, Italy; Betic Cordillera, Spain) the assemblage as a whole spreads a decent flair of Boreal influence on the distal south Tethyan shelf. Although southward migration of northern faunas in the course of ocean water cooling is recorded from the Upper

Turonian onwards (WIESE & VOIGT 2002), the Middle Turonian shows still well- developed biogeographic units among ammonites and echinoids of the Central

European shelf seas and a clear N-S temperature gradient (HART 2007). Thus, the somehow disjunct position of AL 1 from the areas of its main occurrence may not be expression of a cooler water zone in the Helvetian shelf seas. Given the

24.-26. October 2008 51 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research variable facies types in which the representatives of AL 2 occur (e.g.Chalk, Pläner Limestones of NW Germany, marl/limestone alternations, marls), it may rather be argued that nutrient suppy in context with a distinct diet may have triggered the development of this assemblage.

AL 2 & 3 cannot be interpreted due to the very low number of specimens.

References

HART, M. B. 2007. Late Cretaceous climates and foraminiferid distributions. In: WILLIAMS,

M., HAYWOOD, A.M., GREGORY, F.J. & SCHMIDT, D.N. (eds), Deep-Time Perspectives on Climate Change: Marrying the Signal from Computer Models and Biological Proxies, The Micropalaeontological Society, Special Publications. The Geological Society, London, 235-250.

KROH, M. & WAGREICH, M. 2007. Infulaster (Echinodermata, Echinoidea) from the Turonian of the Northern Calcareous Alps. - Austrian Journal of Earth Sciences 100: 154- 161.

WIESE, F. & VOIGT, S. 2002. Late Turonian (Cretaceous) climate cooling in Europe: faunal response and possible causes. – Geobios 35: 65-77.

52 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Antiquity and Longevity of a deep-sea oyster/crinoid association (Azores Archipelago) (Vortrag)

MAX WISSHAK1, CHRISTIAN NEUMANN2, MATTHIAS LÓPEZ CORREA1, SERGE GOFAS3,

CARMEN SALAS3, MARCO TAVIANI4, JOACHIM JAKOBSEN5, ANDRÉ FREIWALD1

1 GeoZentrum Nordbayern, Erlangen University, Loewenichstr. 28, D-91054 Erlangen, Deutschland, e-mail: [email protected]

2 Museum für Naturkunde, Humboldt University Berlin, Invalidenstrasse 43, D-10115 Berlin, Deutschland

3 Departamento de Biología Animal, Universidad de Málaga, E-29071 Málaga, Spain

4 CNR-Istituto di Scienze Marine, Via Gobetti 101, I-40129 Bologna, Italy.

5 Rebikoff-Niggeler Foundation, Rocha Vermelha, 9900-451 Horta, Faial, Azores, Portugal

From the steep submarine slope of the southern Faial Channel (Azores Archipelago), a conspicuous archibenthic community of two ‘living fossils’, comprising the up to 30 cm sized deep-sea oyster Neopycnodonte zibrowii and the sessile cyrtocrinid Cyathidium foresti, is reported. Both taxa thrive in particularly high densities concealed underneath volcanic bedrock overhangs in 420-500 m water depth under stable environmental conditions. Radiocarbon dating of multi-generation oyster stacks (an effective measure to optimise shell stability with minimal biomineralisation effort) reveals that individuals reach an impressive lifespan of one to more than four centuries, placing them among the longest-lived molluscs known to date. This pronounced longevity in concert with a distinct incremental growth pattern, exhibiting a proposed reproductive cyclicity of few years (conchiolin-rich growth breaks), and an annual cyclicity potentially triggered by seasonal varying nutrient flux (distinct Mg and S fluctuations in the shell calcite), make N. zibrowii a promising geochemical archive (stable isotopes and trace element signatures) for the upper bathyal. Submersible investigations showed that the direct association of C. foresti and N. zibrowii is not obligatory but that the crinoid is preferentially settling on oysters, which provide benefits for the crinoid in terms of substrate availability and/or participation in the oysters’ active feeding current. The upside down

24.-26. October 2008 53 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research position of both species provides protection from background sedimentation and possibly shelter from predators. Closely related ancestors of both species are also found as fossils more than 60 million years old in the Danian (Paleocene) of Faxe in Denmark – associated with deep-water coral limestone. There Cyathidium holopus is found settling in impressive numbers settling on various substrata, such as scleractinian corals or, analogous to the Azorean occurrence, on pycnodontine oysters. This remarkable resemblance suggests that this association and semi- cryptic habitat preference was already established in the early Palaeogene. This stands in contrast to the Cretaceous (Cenomanian to Maastrichtian) records of Cyathidium which were exclusively reported from shallow-water palaeoenvironments. We thus assume a habitat-shift of these cyrtocrinids toward deeper waters during the transition of the Mesozoic and Cenozoic erathems. This reflects the escape from enhanced predation pressure (particularly teleost fishes) during the ‘Mesozoic Marine Revolution’. Sessile crinoids almost completely vanished from shallow waters and were replaced by mobile and often strictly nocturnal comatulids. The Cyathidium/(Neo)pycnodonte-association did not only survive the Cretaceous/Palaeogene mass , but the whole Cenozoic and can today be appreciated as a ‘living-fossil community’.

54 24.-26. October 2008 4. Arbeitstreffen deutschsprachiger Echinodermenforscher 4th Workshop of German & Austrian Echinoderm Research

Bromierte Anthrachinon-Pigmente aus der rezenten Seelilie Proisocrinus ruberrimus (Vortrag)

KLAUS WOLKENSTEIN1, WOLFGANG SCHOEFBERGER1, NORBERT MÜLLER1 & TATSUO OJI2

1 Institut für Analytische Chemie, Johannes Kepler Universität Linz, Altenbergerstraße 69, 4040 Linz, Österreich; e-mail: [email protected]

4 Department of Earth and Planetary Science, University of Tokyo, Hongo, Tokyo 113-0033, Japan

Die rezente Seelilie Proisocrinus ruberrimus ist für ihre intensive orangerote

Färbung bekannt (CLARK 1910). Bis heute gab es über die chemische Natur ihrer Pigmente jedoch keine Informationen. Die Entdeckung von P. ruberrimus in etwa 1800 m Meerestiefe im Okinawa-Trog (Japan) mit dem Tauchboot „Shinkai 6500“

(OJI & KITAZAWA 2008) ermöglichte nun die chemische Untersuchung eines Rohextrakts dieser seltenen Seelilie. Die färbenden Verbindungen eines wässrigen Extrakts von P. ruberrimus konnten durch Festphasenextraktion aufkonzentriert und gereinigt und die Hauptverbindungen durch semipräparative Hochleistungsflüssigkeits- chromatographie (HPLC) isoliert werden. Mittels hochauflösender Massenspektrometrie konnten die Summenformeln der Pigmente bestimmt werden (C18H12Br3NaO9S bis C20H15Br4NaO9S). Die charakteristischen Isotopenmuster der Verbindungen bestätigten dabei das Vorkommen von drei bzw. vier Bromatomen. Die vollständige Aufklärung der Strukturen durch ein- und zweidimensionale Kernresonanzspektroskopie ermöglichte schließlich die Zuordnung der neuen Verbindungen zur Gruppe der Anthrachinon-Pigmente. Anthrachinone wurden bisher in zahlreichen Comatuliden nachgewiesen, nicht jedoch bei Seelilien. Soweit bekannt, handelt es sich bei den Pigmenten um die ersten bromierten Anthrachinone natürlicher Herkunft. Zudem besteht eine Verwandtschaft der Pigmente aus P. ruberrimus mit den ebenfalls bromierten Gymnochromen (Phenanthroperylenchinon-Pigmente) aus der Cyrtocrinide

Gymnocrinus richeri (DE RICCARDIS et al. 1991).

Literatur

CLARK, A.H., 1910. Proisocrinus, a new genus of recent crinoids. Proceedings of the United States National Museum 38: 387-390.

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DE RICCARDIS, F., IORIZZI, M., MINALE, L., RICCIO, R., RICHER DE FORGES, B. & DEBITUS, C., 1991. The gymnochromes: Novel marine brominated phenanthroperylenequinone pigments from the stalked crinoid Gymnocrinus richeri. Journal of Organic Chemistry 56: 6781-6787.

OJI, T. & KITAZAWA, K., 2008. Discovery of two rare species of stalked crinoids from Okinawa Trough, southwestern Japan, and their systematic and biogeographic implications. Zoological Science 25: 115-121.

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