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Stuttgarter Beiträge Zur Naturkunde S^5 ( © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zobodat.at Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) Herausgeber: Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart Stuttgarter Beitr. Naturk. Ser. B Nr. 278 175 pp., 4pls., 54figs. Stuttgart, 30. 12. 1999 Comparative osteology oi Mastodonsaurus giganteus (Jaeger, 1828) from the Middle Triassic (Lettenkeuper: Longobardian) of Germany (Baden-Württemberg, Bayern, Thüringen) By Rainer R. Schoch, Stuttgart With 4 plates and 54 textfigures Abstract Mastodonsaurus giganteus, the most abundant and giant amphibian of the German Letten- keuper, is revised. The study is based on the excellently preserved and very rieh material which was excavated during road construction in 1977 near Kupferzeil, Northern Baden- Württemberg. It is shown that there exists only one diagnosable species of Mastodonsaurus, to which all Lettenkeuper material can be attributed. All finds from other horizons must be referred to as Mastodonsauridae gen. et sp. indet. because of their fragmentary Status. A sec- ond, definitely diagnostic genus of this family is Heptasaurus from the higher Middle and Upper Buntsandstein. Finally a diagnosis of the family Mastodonsauridae is provided. Ä detailed osteological description of Mastodonsaurus giganteus reveals numerous un- known or formerly inadequately understood features, yielding data on various hitherto poor- ly known regions of the skeleton. The sutures of the skull roof, which could be studied in de- tail, are significantly different from the schemes presented by previous authors. The endocra- nium and mandible are further points of particular interest. The palatoquadrate contributes a significant part to the formation of the endocranium by an extensive and complicated epi- pterygoid. The neurocranium is dominated by the exoccipital, otic, and several portions of the sphenethmoid. For the first time the stapes is found in articulation and uncompressed. The mandible is slender and bears a massive postglenoid area, and the symphyseal area is extend- ed to accommodate giant tusks. The prearticular forms a pronounced process anterior to the glenoid facet. The axial skeleton is strongly ossified. It consists of massive, cylindrical intercentra that tend to be lower towards the tail and which have a dorsal incisure for the chorda, further it contains paired pleurocentra which are set in facets of the neural arch and intercentrum, and finally it bears low neural arches possessing posteriorly inclined, short and unfinished Proces- sus spinosi and broad, laterally extended transverse processes. The costal apparatus is wide, bearing broadened ribs in the anterior and middle part of the trunk which are strcngthened by two different uncmate processes. While in the trunk these processes disappear tailwards, the tail base and anterior tail proper again bear wide ribs with markcd processes. Doctoral dissertation Univ. Tübingen 1998. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zobodat.at 2 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Ser. B, Nr. 278 The girdles and appendages are well represented in the Kupferzell material, with the excep- tion of the pes; because of their disarticulated State their original arrangement has to be recon- structed. The dermal bones of the pectoral girdle consist of broad and flattened claviculae and a wide, rhomboidal interclavicle which has, as is typical of capitosaurs, a stilette-like anterior process with a broadened tip. The ontogeny of the cleithrum and scapulocoracoid is docu- mented in detail. Scapula and coracoid are separate at first, but later in development fuse to give a Compound ossification, whose coracoid portion tends to increase in area. In addition the geometry and surface of the glenoid facet changes during ontogeny. The humerus is rela- tively slender and long, yet poorly differentiated with respect to most other temnospondyls. Only in the largest specimens are there definite epicondyles and a faint supinator process. Ra- dius and ulna are well differentiated and elongate, and the manus consists mainly of slender metacarpals and phalanges. In the pelvic girdle the ilium is slender and low, with an acetabu- lum that enlarged during later development. The ischium is small and remains poorly differ- entiated. In larger specimens it appears to fuse with the pubis which is still unknown as isolat- ed element. The femur is slender, elongate, and differentiated, whereas the lower leg gains less than half the length of the upper leg and has a rudimentary structure. Finally open questions and problems regarding the reconstruction of the whole skeleton are discussed. This involves a clarification of the methods used and a discussion of the lines of evidence. A complete yet still partially provisional restoration of the skeleton of Mastodon- saurus giganteus gives an animal similar to a gigantic Salamander. Mastodonsaurus was most similar to cryptobranchid Salamanders in its bodily proportions such as the relative length of the trunk and tail, as well as the considerable breadth and flattness of the rib cage. Zusammenfassung Mastodonsaurus giganteus, das häufigste und größte Amphib der europäischen Mitteltrias, wird eingehend revidiert. Die Untersuchung geht aus von dem hervorragend erhaltenen und sehr umfangreichen Material, das beim Autobahnbau 1977 bei Kupferzell geborgen wurde. Im Lettenkeuper existiert nur eine diagnostizierbare Art von Mastodonsaurus, während die fragmentarischen Funde aus anderen Formationen lediglich der Familie Mastodonsauridae zugeordnet werden können. Die zweite, gegenwärtig faßbare Gattung dieser Familie ist Hep- tasaurus, die auf den höheren Mittleren und Oberen Buntsandstein beschränkt bleibt. Die ta- xonomische Revision schließt mit einer Diagnose der Mastodonsauridae. Im Rahmen einer detaillierten osteologischen Untersuchung von Mastodonsaurus gigante- us werden zahlreiche neue Merkmale beschrieben und viele, bisher unzureichend verstandene Regionen des Skeletts vorgestellt und rekonstruiert. Die Suturen im Schädeldach können dank der guten Erhaltung der Kupferzeller Funde erstmals genau erfaßt werden; sie weichen in einigen Punkten erheblich von den Schemata früherer Bearbeiter ab. Das Endocranium und die Mandibel von Mastodonsaurus bilden weitere Schwerpunkte. Das Palatoquadratum ist mit einem ausgedehnten und kompliziert gebauten Epipterygoid wesentlich an der Gestal- tung des Endocranium beteiligt. Im knöchernen Neurocranium dominieren das Exoccipitale, das Oticum sowie verschiedene Portionen des Sphenethmoid; der Stapes wird erstmals un- verdrückt und in Artikulation vorgefunden und vorgestellt. Im Unterkiefer, der sehr niedrig ist und eine massive, langgestreckte Postglenoid-Region besitzt, dominieren riesige Fangzäh- ne in der verbreiterten Symphyse. Ein hoher, allometrisch positiv wachsender Fortsatz des Praearticulare bildet eine knöcherne Sperre unmittelbar vor dem Unterkiefergelenk. Das Axialskelett ist stark verknöchert. Es besteht aus massiven, kurz-zylindrischen Inter- centra, die zur Cauda hin niedriger werden und eine Inzisur für die Chorda aufweisen, wei- terhin enthält es paarige Pleurocentra, die in Facetten des Neuralbogens und des Intercentrum eingelassen sind, und schließlich die Bögen selbst, die niedrig sind, mit nach hinten geneigten Dornfortsätzen und breit nach lateral vorgreifenden Transversalfortsätzen. Der Rippenkorb ist breit, im vorderen und mittleren Rumpf aus kräftigen Rippen bestehend, die mehrere Un- cinat-Fortsätze tragen können. Diese werden im hinteren Rumpfabschnitt kürzer und ver- schwinden schließlich ganz, während im Schwanzskelett die Rippen erneut größer sind und auch dort blatt- bis stilettförmige Uncinat-Fortsätze tragen. Die Gürtel und Extremitäten sind bis auf das Fußskelett gut dokumentiert, müssen jedoch weitgehend anhand disartikulierter Reste rekonstruiert werden. Der hautknöcherne Schulter- gürtel besteht aus breiten, großflächigen Claviculae und einer breitrhombischen Interclavicu- © Biodiversity SCHOCH,Heritage Library,MASTODONSAURUS http://www.biodiversitylibrary.org/;GIGANTEUS www.zobodat.at 3 la, die den für Capitosaurier typischen, vorne verbreiterten anterioren Fortsatz trägt. Die On- togenese des Cleithrum sowie der ersatzknöchernen Elemente läßt sich im Detail verfolgen. Die Scapula und das Coracoid sind anfangs getrennt und verschmelzen später suturlos, wobei sich der Coracoid-Anteil stark medial ausdehnt. Auch die Glenoidfacette erweitert und ver- kompliziert sich im Zuge der späteren Ontogenese. Der Humerus ist anfangs verhältnismäßig schlank, insgesamt relativ lang und bis in das hohe Alter von rudimentärer Gestalt verglichen mit den meisten anderen Temnospondylen; erst bei den größten Individuen bilden sich die Condyli an seinem Distalende sowie ein schwacher Supinatorfortsatz. Radius und Ulna sind lang und differenziert, und das Handskelett besteht aus überwiegend schlanken Metacarpalia und Phalangen. Im Beckengürtel findet sich ein niedriges und schlankes Ilium, dessen Anteil am Acetabulum sich im Alter vergrößert. Das Ischium ist klein und bleibt zeitlebens wenig differenziert; bei größeren Tieren scheint es mit dem Pubis, das bisher isoliert nicht bekannt ist, verschmolzen zu sein. Das Femur ist schlank, langgestreckt und differenziert, während Ti- bia und Fibula weniger als halb so lang werden und rudimentär bleiben. Die Untersuchung schließt mit der Besprechung offener Fragen und Probleme bei der Re- konstruktion des Gesamtskeletts.
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