Extending the footprint record of Pareiasauromorpha to the Cisuralian: earlier appearance and wider palaeobiogeography of the group

Abstract

Pareiasauromorpha is one of the most important tetrapod groups of the Permian. Skeletal evidence suggests a late Kungurian origin in North America, whereas the majority of occurrences come from the Guadalupian and Lopingian of South Africa and Russia. However, Pareiasauromorpha footprints include the ichnogenus Pachypes, which is unknown from strata older than late Guadalupian. A revision of several Pachypes-like footprints from the Cisuralian-Guadalupian of Europe and North America confirm the occurrence of this ichnogenus and of the ichnospecies Pachypes ollieri comb. nov. beginning in the Artinskian. This is the earliest known occurrence of

A captorhinid-dominated assemblage from the palaeoequatorial Permian of Menorca (Balearic Islands, western Mediterranean)

“…Remarks. MT01-13 can be assigned to Erpetopus because of the following combination of characters (Marchetti et al 2014): (1) pes slightly wider than long; (2) slender, curved digit imprints with clawed tips; (3) digit imprints with overlapping digit bases, typical of parareptiles (Marchetti et al 2020b); (4) short digit V imprint; (5) high IV-V divergence; (6) high I-V divergence; and (7) small size, less than 20 mm. The most probable trackmakers for Erpetopus are acleistorhinid and nyctiphruretid parareptiles, and this ichnogenus has been found from the middle Artinskian…”

Hyloidichnus trackways with digit and tail drag traces from the Permian of Gonfaron (Var, France): New insights on the locomotion of captorhinomorph eureptiles

“…Parareptiles such as pareiasaurs (trackmakers of large Pachypes dolomiticus) are too large to fit with Hyloidichnus, whereas smaller pareiasauromorphs (trackmakers of Pachypes ollieri) have closelypacked digits and overlapping metatarsals and metacarpals, which are inconsistent with the radiating digit imprints and the large digit divarication angle of Hyloidichnus (Ma rchetti et al, 2020a). Also, their tails were short, whereas continuous tail impressions are commonly observed in trackwa ys of Hyloidichnus.…”

“…This includes valuable data on, for example, functional morphology and locomotion of the trackmakers (Voigt et al, 2007;Romano et al, 2016;Ma rchetti et al, 2017;Buchwitz and Voigt, 2018;Mujal and Ma rchetti, 2020;Mujal and Schoch, 2020;Mujal et al, 2020). Tetrapod footprints are usually compared with the morphology of the appendicular skeletons of contemporary taxa (Ga nd, 1987;Voigt, 2005;Ga nd and Durand, 2006;Voigt et al, 2007;Ma rchetti et al, 2019aMa rchetti et al, , 2020aMujal and Ma rchetti, 2020;Mujal and Schoch, 2020). To render these studies possible, wellpreserved tracks and trackwa ys are needed: these exhibit morphological features directly related to anatomy, without 'extramorphologies' (i.e., morphological features that hide anatomical traits of the producer; Peabody, 1948;Haubold et al, 1995;Ma rchetti et al, 2019b).…”

Evolutionary Change in Locomotion Close to the Origin of Amniotes Inferred From Trackway Data in an Ancestral State Reconstruction Approach

“…Since the approaches of Haubold (1970Haubold ( , 1971 and Fichter (1983a;1983b), track-trackmaker relationships have been clarified for various groups of early amniotes and amniote relatives from the Carboniferous and early Permian (e.g., Voigt, 2005;Voigt et al, 2007;Marchetti et al, 2017Marchetti et al, , 2019aMarchetti et al, , 2020c and provide a robust framework for the biostratigraphic interpretation of tracks (Voigt and Lucas, 2018;Marchetti et al, 2019b;Schneider et al, 2020;Lucas et al, 2021). Some track types with a tetradactyl manus as in extant amphibians, such as Batrachichnus and Limnopus, have been assigned to temnospondyl producers (Haubold et al, 1995;Voigt, 2005), whereas the trackway pattern in Matthewichnus points to a lepospondyl trackmaker (Voigt and Lucas, 2015).…”

“…tracks (Wilson, 2005). The idea that tetrapod footprint ichnotaxa can be sorted phylogenetically and assigned to certain clades of a skeletal-morphology-based phylogenetic tree according to the occurrence of certain diagnostic features in both, track and trackmaker morphology, has been termed "synapomorphy-based track-trackmaker correlation" by Carrano and Wilson (2001) and is discussed in several recent approaches on Paleozoic tracks (e.g., Voigt et al, 2007;Marchetti et al, 2017Marchetti et al, , 2019aMarchetti et al, , 2020cBuchwitz and Voigt, 2018;. Mapping features on a phylogenetic tree and inferring character states in common ancestors of terminal taxa, whose character states are known, represents a standard way of addressing character evolution and has been introduced to paleontology together with cladistic methods.…”