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Interrelationships, palaeobiogeography and early evolution of Stereospondylomorpha (Tetrapoda: Temnospondyli)

  • Estevan EltinkEmail author
  • Rainer R. Schoch
  • Max C. Langer
Research Paper

Abstract

The stereospondylomorph temnospondyls form a diverse group of early tetrapods that survived the Permian–Triassic extinction event and radiated during the Triassic. They encompass Carboniferous and Permian taxa from central and eastern Europe, such as ‘archegosauroids’, and early-divergent Gondwanan forms, such as rhinesuchids. By the Early Triassic, the group reached a worldwide distribution, with stereospondyls experiencing an impressive diversification and becoming major aquatic predators in fresh water, brackish, and even marine ecosystems. The origin of Stereospondyli dates back into the Permian, but the phylogenetic relationships of stereospondylomorphs, including ‘archegosauroids’ and the first stereospondyls remain unclear, representing the focus of the present study. Incorporating new records of the group to a broad revision of the different phylogenic hypotheses, a 37-taxon sample was scored for 221 morphological characters revised from previous works. The parsimony analysis resulted three most parsimonious trees (MPTs) of 737 steps. Their strict consensus tree depicts Sclerocephalidae and Intasuchidae as early-diverging stereospondylomorphs, ‘Archegosauroidea’ as paraphyletic array of taxa and Konzhukoviidae as the sister-group of a monophyletic Stereospondyli. An early-diverging and monophyletic Rhinesuchidae is divided into Australerpetinae and Rhinesuchinae, as the sister-group of the clade containing the Permian Peltobatrachus pustulatus, Arachana nigra, and a newly named clade (Superstes = ‘survivors’) of Triassic stereospondyls, formed by Lydekkerinidae and Neostereospondyli (Capitosauria and Trematosauria). Likelihood ancestral area reconstructions and time-ranging distributions along phylogeny provided a comprehensive description of early Stereospondylomorpha palaeobiogeography history. The initial evolution of the group took place in Laurasian areas (central and eastern Europe) during the Cisuralian and Guadalupian (early-mid Permian), with a broader Pangaean distribution for Platyoposauridae and Konzhukoviidae. Stereospondyls have Africa as their ancestral area, followed by dispersions to other Gondwanan regions during the Guadalupian and Lopingian (mid-late Permian), revealing a remarkable diversity previous to the P-Tr extinction. In the Triassic, Superstes greatly expanded across the Pangaea, highlighting another significant event in the evolution of Stereospondylomorpha.

Keywords

Temnospondyli Stereospondylomorpha Permian Systematics Biogeography 

Resumen

Los temnospóndilos estereospondilomorfos forman un grupo diverso de primeros tetrápodos que sobrevivieron al evento de la extinción Pérmico-Triásica y radiaron durante el Triásico. Ellos abarcan taxones del Carbonífero y el Pérmico de Centro y Este de Europa, tales como los “archegosauroides”, y las primeras formas Gondwánicas divergentes, como los rhinesúquidos. Durante el Triásico Inicial, el grupo alcanzó una distribución mundial, con estereospóndilos experimentando una diversificación impresionante y convirtiéndose en grandes depredadores acuáticos en ecosistemas de agua dulce, salobre e incluso marinos. El origen de Stereospondyli data del Pérmico, pero las relaciones filogenéticas de los esterespondilomorfos, incluyendo “archegosauroides” y los primeros estereospóndilos permanece poco clara, representando el foco del presente estudio. Incorporando nuevos registros del grupo a una amplia revisión de las diferentes hipótesis filogenéticas, una muestra de 37 taxones fue codificada para 221 caracteres morfológicos revisados de trabajos previos. El análisis de parsimonia resultó en tres arboles más parsimoniosos (MPTs en sus siglas en inglés) de 737 pasos. El árbol de consenso estricto representa Sclerocephalidae y Intasuchidae como los primeros estereospondilomorfos divergentes, “Archegosauroidea” como un grupo parafilético de taxones y Konzhukoviidea como el grupo hermano del monofilético grupo Stereospondyli. Rhinesuchidae es monofilético, divergió tempranamente y se divide en Australerpetinae y Rhinesuchinae, como el grupo hermano del clado que contiene el taxón Pérmico Peltobatrachus pustulatus, Arachana nigra, y el nuevo clado aquí nombrado (Superestes=`supervivientes’) de estereospóndilos Triásicos, formado por Lydekkerinidae y Neostereospondyli (Capitosauria y Trematosauria). La reconstrucción del área ancestral potencial y distribución de rangos temporales a lo largo de la filogenia proporcionan una descripción exhaustiva de la historia paleobiogeográfica de Stereospondylomorpha. La evolución inicial del grupo tuvo lugar en áreas de Laurasia (Europa Central y del Este) durante el Cisuraliense y Guadalupiense (Pérmico inicial-medio), con una amplia distribución en Pangea de Platyoposauridae y Konzhukoviidae. Los esterespóndilos tuvieron África como su área ancestral, seguido de dispersiones en otras regiones Gondwánicas durante el Guadalupiense y Lopingiense (Pérmico medio-tardío), revelando una remarcable diversidad previa a la extinción Pérmico-Triásica. En el Triásico, Superstes se expandió enormemente a través de Pangea, destacando otro evento significativo en la evolución de Stereospondylomorpha.

Palabras clave

Temnospondyli Stereospondylomorpha Pérmico Sistemática Biogeografía 

Notes

Acknowledgements

We thank Rodrigo Rocha Machado (DGM), Cesar Leandro Schultz (UFRGS), Philipe Havlik (GPIT), Florian Witzmann (MB), Mathew Lowe (CAMZM), Lorna Steel and Andrew Milner (UKNHM), Yuri Gubin and Mikhail Shishkin (PIN), Bernhard Zipfel and Bruce Rubidge (BPI), Sheena Kaal (IZIKO), Heidi Fourie (TM), and Carl Mehling (AMNH) to provide access to studied specimens in the research. EE was supported by foundations: FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo) [grant number 2009/54656-9]; and CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) [grant number 290092-2011-6]. This contribution used TNT v.1.1, a program made freely available by the Willi Hennig Society. Thorough reviews by Florian Witzmann, anonymous reviewer, and Josep Fortuny greatly improved the final manuscript.

Supplementary material

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Supplementary material 1 (DOCX 44 kb)
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© Universidad Complutense de Madrid 2019

Authors and Affiliations

  1. 1.Universidade Federal do Vale do São Francisco, Colegiado de EcologiaBahiaBrazil
  2. 2.Staatliches Museum für Naturkunde StuttgartStuttgartGermany
  3. 3.Universidade de São Paulo, FFCLRP, Laboratório de PaleontologiaRibeirão PretoBrazil

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