Abstract

Terrestrial ecosystems during the Pennsylvanian (late Carboniferous) and Cisuralian (early Permian) are usually described in the literature as being dominated by synapsids, the mammal-line amniotes. The pelycosaurs (a paraphyletic grouping of synapsid families) have been considered more speciose, abundant, and ecologically diverse than contemporary reptile-line amniotes. However, this dominance has never been subjected to quantitative testing accounting for sampling bias. Moreover, in recent years the amniote phylogeny has undergone numerous revisions, with suggestions that varanopids and recumbirostran microsaurs fall within reptiles, and that diadectomorphs may be pelycosaurian-grade synapsids. An examination of local species richness (alpha diversity) of synapsids and reptiles during the Pennsylvanian and Cisuralian at different spatial scales shows that these taxonomic revisions have substantial impacts on relative diversity patterns of synapsids and reptiles. Synapsids are only found to be consistently more diverse through the early Permian when using the “traditional” taxonomy. The recent taxonomic updates produce diversity estimates where reptile diversity is consistent with, or in some cases higher than that of synapsids. Moreover, biases in preservation may affect patterns. Where preservation favors smaller vertebrates, e.g., Richards Spur, South Grandfield, reptiles overwhelmingly dominate. If smaller vertebrates are expected to make up the bulk of amniote diversity, as they do in the present day, such lagerstätten may be more representative of true diversity patterns. Therefore, the dominance of pelycosaurs during this interval should be reconsidered, and this interval may be considered the First Age of Reptiles.

Highlights

  • Amniotes, the vertebrates with the amniotic egg that gives their reproduction complete independence from water, first appear in the fossil record about 315 million years ago (Carroll, 1964) and rapidly diversified into a great diversity of species and ecologies (Sahney et al, 2010; Dunne et al, 2018) that is seen in the present day

  • We show how incorporating the novel positions suggested for varanopids, recumbirostrans and lysorophians into analyses of early amniote diversity produces estimates of reptile diversity more consistent with those of synapsids, challenging the assumption that synapsids dominated during the Pennsylvanian and Cisuralian

  • The recent revisions of amniote phylogeny will doubtless be subject to further debate and modification, and there may well be other clades whose position will come under scrutiny

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Summary

INTRODUCTION

The vertebrates with the amniotic egg that gives their reproduction complete independence from water, first appear in the fossil record about 315 million years ago (Carroll, 1964) and rapidly diversified into a great diversity of species and ecologies (Sahney et al, 2010; Dunne et al, 2018) that is seen in the present day. There has been debate surrounding the relationships within the two lineages (e.g., Müller and Reisz, 2006; Benson, 2012; Brocklehurst et al, 2016; Laurin and Piñeiro, 2017; MacDougall et al, 2018) and individual species have sometimes been moved between synapsids and reptiles (e.g., Reisz and Modesto, 2007; Reisz et al, 2010; Ford and Benson, 2019; Mann et al, 2020), but on the whole the clade, Synapsida or Reptilia, to which the Pennsylvanian and Cisuralian lineages have been assigned has been consistent. We show how incorporating the novel positions suggested for varanopids, recumbirostrans and lysorophians into analyses of early amniote diversity (species richness) produces estimates of reptile diversity more consistent with those of synapsids, challenging the assumption that synapsids dominated during the Pennsylvanian and Cisuralian. Consideration of the preservation biases and the effect of lagerstätten suggest that any apparent dominance of synapsids during this time may well be artifactual

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