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First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny

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Abstract

A new fossil amniote from the Fossil Forest of Chemnitz (Sakmarian-Artinskian transition, Germany) is described as Ascendonanus nestleri gen. et sp. nov., based on five articulated skeletons with integumentary preservation. The slender animals exhibit a generalistic, lizard-like morphology. However, their synapsid temporal fenestration, ventrally ridged centra and enlarged iliac blades indicate a pelycosaur-grade affiliation. Using a renewed data set for certain early amniotes with a similar typology found Ascendonanus to be a basal varanopid synapsid. This is the first evidence of a varanopid from Saxony and the third from Central Europe, as well as the smallest varanopid at all. Its greatly elongated trunk, enlarged autopodia and strongly curved unguals, along with taphonomical observations, imply an arboreal lifestyle in a dense forest habitat until the whole ecosystem was buried under volcanic deposits. Ascendonanus greatly increases the knowledge on rare basal varanopids; it also reveals a so far unexpected ecotype of early synapsids. Its integumentary structures present the first detailed and soft tissue skin preservation of any Paleozoic synapsid. Further systematic results suggest a varanodontine position for Mycterosaurus, the monophyly of South African varanopids including Anningia and the distinction of a skeletal aggregation previously assigned to Heleosaurus, now renamed as Microvaranops parentis gen. et sp. nov.

Kurzfassung

Basierend auf fünf artikulierten Skeletten mit Hauterhaltung wird ein neuer, fossiler Amniot aus dem Versteinerten Wald von Chemnitz (Sakmarium–Artinskium-Grenzbereich, Deutschland) beschrieben als Ascendonanus nestleri gen. et sp. nov. Die schlanken Tiere sind von generalistischer, echsenhafter Gestalt. Demgegenüber zeigen die synapsiden Schläfenfenster, ventral gekantete Zentren und vergrößerte Iliumblätter eine Zugehörigkeit zur Pelycosaurier-Stufe an. Unter Anwendung eines erneuerten Datensatzes für ausgewählte frühe Amnioten ähnlicher Typologie wird Ascendonanus zu basalen Varanopiden gestellt. Damit liegt der erste Nachweis eines Varanopiden aus Sachsen und der dritte aus Mitteleuropa vor, zudem der kleinste Varanopide überhaupt. Sein besonders verlängerter Rumpf, vergrößerte Autopodien und stark gekrümmte Krallen sowie taphonomische Beobachtungen legen eine arboreale Lebensweise inmitten eines dichten Waldhabitats nahe, bis das gesamte Ökosystem von vulkanischen Ablagerungen verschüttet wurde. Ascendonanus erweitert die Kenntnis der seltenen basalen Varanopiden enorm, zumal er einen bei frühen Synapsiden bisher unerwarteten Ökotyp aufdeckt. Die Integumentstrukturen stellen die ersten detaillierten und durch Weichteile erhaltenen Hautfunde aller paläozoischen Synapsiden dar. Weitergehende systematische Ergebnisse deuten an: eine varanodontine Position für Mycterosaurus, die Monophylie südafrikanischer Varanopiden einschließlich Anningia, sowie die Unterscheidung einer vormals zu Heleosaurus gestellten Skelettaggregation, nun benannt als Microvaranops parentis gen. et sp. nov.

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Acknowledgements

We gratefully acknowledge the support by the Chemnitz excavation team including Ralph Kretzschmar and Mathias Merbitz for professional fieldwork, saving of the finds and many fruitful discussions and Dr. Thorid Zierold for encouraging the project management. We express special thanks to Georg Sommer, Schleusingen, for excellent preparation of the skeletons. We are further indebted to Maibrit Scheibel, Annika Buitink and Thomas Israel. This research was supported by the Deutsche Forschungsgemeinschaft (DFG grants RO 1273-3/1 to R.R. and SCHN 408/20 to J.W.S.) and Volkswagen Foundation (Az: I/84638), as well as Raimund Albersdörfer and Michael Völker (Dinosaurier Freiluftmuseum Altmühltal). The manuscript benefited greatly from the constructive reviews provided by Sean P. Modesto and Neil Brocklehurst.

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Authors and Affiliations

  1. Dinosaurier-Park Altmühltal, Dinopark 1, 85095, Denkendorf, Germany

    Frederik Spindler

  2. Naturhistorisches Museum Schloss Bertholdsburg, Burgstraße 6, 98553, Schleusingen, Germany

    Ralf Werneburg

  3. Geological Institute, TU Bergakademie Freiberg, Bernhard von Cotta-Straße 2, 09599, Freiberg, Germany

    Joerg W. Schneider & Ronny Rößler

  4. Kazan Federal University, 420008, Kazan, Russia

    Joerg W. Schneider

  5. Museum für Naturkunde, Moritzstraße 20, 09111, Chemnitz, Germany

    Ludwig Luthardt, Volker Annacker & Ronny Rößler

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  1. Frederik Spindler
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Correspondence to Frederik Spindler.

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Spindler, F., Werneburg, R., Schneider, J.W. et al. First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny. PalZ 92, 315–364 (2018). https://doi.org/10.1007/s12542-018-0405-9

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