Palaeoecology and palaeoenvironment of Discosauriscus Kuhn, 1933 with exceptional preservation of skin and intestine molds from the early Permian of Franchesse (Allier, France)

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Christopher Beard (University of Kansas, Lawrence) Paléoanthropologie/Palaeoanthropology Aurélien Mounier (CNRS/Muséum national d’Histoire naturelle, Paris) Archéologie préhistorique (Paléolithique et Mésolithique)/Prehistoric archaeology (Palaeolithic and Mesolithic) Nicolas Teyssandier (CNRS/Université de Toulouse, Toulouse) Archéologie préhistorique (Néolithique et âge du bronze)/Prehistoric archaeology (Neolithic and Bronze Age) Marc Vander Linden (Bournemouth University, Bournemouth) référés / reviewers : https://sciencepress.mnhn.fr/fr/periodiques/comptes-rendus-palevol/referes-du-journal couverture / cover : Made from the Figures of the article. Comptes Rendus Palevol est indexé dans / Comptes Rendus Palevol is indexed by: – Cambridge Scientific Abstracts – Current Contents® Physical – Chemical, and Earth Sciences® – ISI Alerting Services® – Geoabstracts, Geobase, Georef, Inspec, Pascal – Science Citation Index®, Science Citation Index Expanded® – Scopus®. Les articles ainsi que les nouveautés nomenclaturales publiés dans Comptes Rendus Palevol sont référencés par / Articles and nomenclatural novelties published in Comptes Rendus Palevol are registered on: – ZooBank® (http://zoobank.org) 489 COMPTES RENDUS PALEVOL • 2025 • 24 (24) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr Antoine LOGGHE Centre de Recherche en Paléontologie – Paris (CR2P), Muséum national d’Histoire naturelle, CNRS-MNHN-SU, CP38, 8 rue Buffon, F-75005 Paris (France) [email protected] (corresponding author) Arnaud REBILLARD Leibniz-Institut für Evolutionsund Biodiversitätsforschung, Museum für Naturkunde, Invalidenstraße, 43, 10115 Berlin (Germany) and Institut für Biologie Humboldt-Universität zu Berlin, Invalidenstraße, 42, 10115 Berlin (Germany) and Centre de Recherche en Paléontologie – Paris (CR2P), Muséum national d’Histoire naturelle, CNRS-MNHN-SU, CP38, 8 rue Buffon, F-75005 Paris (France) Guillaume HOUÉE Alicia SÁNCHEZ GIMENO Centre de Recherche en Paléontologie – Paris (CR2P), Muséum national d’Histoire naturelle, CNRS-MNHN-SU, CP38, 8 rue Buffon, F-75005 Paris (France) Erwan COURVILLE Biogéosciences, Université de Bourgogne, CNRS, 6 Boulevard Gabriel, F-21000 Dijon (France) Lazare ELBAZ Department of Herpetology, STBio post-graduation program, Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, Ipiranga, São Paulo 04263-000, SP (Brazil) Paloma-Maria TARABA Mairie de Saint-Yrieix La Perche, 45 boulevard de l’hôtel de ville, F-87500 Saint-Yrieix La Perche (France) Rodolphe PASO 1 avenue du recteur Pineau, F-86000 Poitiers (France) Max HERDE Leibniz-Institut für Evolutionsund Biodiversitätsforschung, Museum für Naturkunde, Invalidenstraße, 43, 10115 Berlin (Germany) and Institut für Biologie, Humboldt-Universität zu Berlin, Invalidenstraße, 42, 10115 Berlin (Germany) Stanislav ŠTAMBERG Museum of Eastern Bohemia, Hradec Králové Eliščino nábřeží 465, 500 01 Hradec Králové (Czech Republic) Palaeoecology and palaeoenvironment of Discosauriscus Kuhn, 1933 with exceptional preservation of skin and intestine molds from the early Permian of Franchesse (Allier, France) 490 COMPTES RENDUS PALEVOL • 2025 • 24 (24) Logghe A. et al. urn:lsid:zoobank.org:pub:DBE4A506-5D02-4AFF-846E-3E367D472281 Logghe A., Rebillard A., Houée G., Sánchez Gimeno A., Courville E., Elbaz L., Taraba P.-M., Paso R., Herde M., Štamberg S., Pouillon J.-M. & Steyer J.-S. 2025. — Palaeoecology and palaeoenvironment of Discosauriscus Kuhn, 1933 with exceptional preservation of skin and intestine molds from the early Permian of Franchesse (Allier, France). Comptes Rendus Palevol 24 (24): 489-504. https://doi.org/10.5852/cr-palevol2025v24a24 ABSTRACT The Carboniferous-Permian is a key period concerning the emergence and diversification of major tetrapod clades such as amphibians and amniotes. This is particularly the case of the Seymouriamorpha Watson, 1917. Some seymouriamorphs are well-known, such as Seymouria Broili, 1904 from North America and Germany, but the ecology of most taxa remains debated. This is the case of Discosauriscus Kuhn, 1933 from the early Permian of Europe. Here we provide a preliminary study of new specimens from the early Permian Lagerstätte of Franchesse (Allier, France) whose exceptionally well-preserved soft tissues including skin and intestinal casts provide valuable insights into the palaeoecology of Discosauriscus. We describe for the first time the complete skin external and histological features of a seymouriamorph indicating that juvenile discosauriscids already displayed epidermal scalation. This suggests a skin adapted towards terrestrial lifestyle in juvenile Discosauriscus. The recovery of an in situ cololite facilitates the identification of seymouriamorph coprolites found ex situ, and thus, future interpretations of the specific diet of this taxon. In order to better understand how these soft tissues have been preserved, we also conducted a taphonomic study including a stratigraphic and palaeoenvironmental analysis of the fossil layer deposits. Our results suggest that pronounced volcanic activity at work during the early Cisuralian of the Bourbon-l’Archambault Basin may be the cause of the multiple Permian mass mortality assemblages found at Franchesse, as well as the exceptional preservation of the fossil remains from this site. RÉSUMÉ Paléoécologie et paléoenvironnement de Discosauriscus Kuhn, 1933 avec préservation exceptionnelle des moules de peau et d’intestin du Permien précoce de Franchesse (Allier, France). Le Carbonifère-Permien est une période clé en ce qui concerne l’émergence et la diversification des principaux clades de tétrapodes tels que les amphibiens et les amniotes. C’est notamment le cas des Seymouriamorpha Watson, 1917. Certains seymouriamorphes sont bien connus, comme Seymouria Broili, 1904 d’Amérique du Nord, mais l’écologie de la plupart des taxons reste débattue. C’est le cas de Discosauriscus Kuhn, 1933 du Permien inférieur d’Europe. Ici, nous étudions de manière préliminaire de nouveaux spécimens provenant du Lagerstätte du Permien inférieur de Franchesse (Allier, France) dont les tissus mous exceptionnellement bien conservés, y compris la peau et le moulage des intestins, permettent d’en apprendre davantage sur la paléoécologie de Discosauriscus. Nous décrivons pour la première fois la morphologie externe et interne de la peau d’un seymouriamorphe, indiquant que les discosauriscidés juvéniles présentaient des écailles épidermiques. Cela suggère une peau adaptée à un mode de vie terrestre chez les Discosauriscus juvéniles. La découverte d’un cololithe insitu facilite l’identification des coprolithes de seymouriamorphes trouvés exsitu et ainsi de futures interprétations sur le régime alimentaire de ce taxon. Afin de mieux comprendre comment ces tissus mous ont été préservés, nous avons également mené une étude taphonomique comprenant une analyse stratigraphique et paléoenvironnementale des dépôts des couches fossilifères. Nos résultats suggèrent qu’une activité volcanique marquée au cours du Cisuralien inférieur dans le bassin de Bourbon-l’Archambault pourrait être à l’origine des multiples assemblages de mortalité massive trouvés à Franchesse, ainsi que de la préservation exceptionnelle des restes fossiles de ce site. Jean-Marc POUILLON Rhinopolis, 179 rue des Plattières, F-38300 Nivolas Vermelle (France) J.-Sébastien STEYER Centre de Recherche en Paléontologie – Paris (CR2P), Muséum national d’Histoire naturelle, CNRS-MNHN-SU, CP38, 8 rue Buffon, F-75005 Paris (France) Submitted on 4 February 2025 | Accepted on 20 June 2025 | Published on 6 November 2025 KEY WORDS Permian, Seymouriamorpha, skin, cololite, palaeoecology. MOTS CLÉS Permien, Seymouriamorpha, peau, cololithe, paléoécologie. 491 Palaeoecology and palaeoenvironment of Discosauriscus COMPTES RENDUS PALEVOL • 2025 • 24 (24) INTRODUCTION Complementary to the study of mineralised tissues in fossil vertebrates, new palaeoecological traits can be inferred from fossilised soft tissues, such as skin (e.g. Vincent etal. 2017; Hendrickx etal. 2022; Jagielska etal. 2024), and bromalites (Hoffmann etal. 2020; Halaçlar etal. 2023; Qvarnström etal. 2024). Indeed, the skin, as first barrier between the organism and the external environment, shows several adaptations such as protection against mechanical stress, UV-radiation or predatory attacks (Vickaryous & Sire 2009). It can also prevent desiccation (Alibardi 2003), favour thermoregulation (Lindgren etal. 2014), and is a main component of behavioural displays such as camouflage and/or sexual display, for example through colouration (e.g. McNamara etal. 2021). Fossilised digestive remains, preserved either within (e.g. consumulites) or outside the body (e.g. coprolites, regurgitalites), also provide valuable information regarding palaeoecology (e.g. environmental factors; Hunt 1992; Northwood 2005) or palaeobiodiversity (some floral and faunal elements are only preserved in bromalites; Byrne et al. 2022). Especially, consumulites such as gastrolites or cololites are the most compelling evidence in the fossil record of the predator-prey interactions (e.g. Qvarnström etal. 2024). Furthermore, fossilised gut content is precious evidence allowing to directly link a predator with its prey and environment, at a relatively precise temporal and spatial resolution (Kriwet etal. 2008; Mondini 2023). However, as both soft tissues and bromalites fossilise more rarely than mineralised tissues, these key aspects of the palaeoecology of Palaeozoic tetrapods are poorly known (e.g. Eliason etal. 2017; Hunt & Lucas 2021). The Discosauriscidae Romer, 1947 is a clade of Seymouriamorpha Watson, 1917. Their palaeoecology is debated between mainly aquatic or more amphibious/terrestrial lifestyle (Klembara 1995; Malakhov 2000; Kriloff etal. 2008; Sanchez etal. 2008; Calábková etal. 2022). Mainly, external gills are frequently preserved in discosauriscids (Ivakhnenko 1987; Klembara 1995; Bulanov 2003). Cololites have been described in association with discosauriscid remains, suggesting putative cannibalism (Klembara & Meszároš 1992). Dermal coverage has been described but only assigned to ossified dermal scales, without palaeoecological implications (Špinar 1952; Klembara & Bartík 1999). Thus, almost no skin soft tissue is known for this clade, and even for all Seymouriamorpha (Olson 1965). The Permian Lagerstätte of Franchesse (Allier, France), discovered by Prof. Roger-Paul Dechambre (MNHN) in 1964, yielded hundreds of sub-complete to complete specimens of Discosauriscus Kuhn, 1933, preserving soft-tissue remains (Steyer etal. 2012: fig. 2). From 2007, one of us (J.-S.S.) organised regular systematic excavations at the Les Charbonnières site in Franchesse, under a working convention with Epona Society, owner of the site. These fieldworks led to the discovery and description of a very rich flora and fauna including numerous aeduellids, one partial Bohemiacanthus Schneider, 1996 shark, numerous exquisite Discosauriscus Kuhn, 1933 specimens, and a larval eryopoid temnospondyl (Steyer etal. 2012). Then A.L. co-organised further fieldworks (2021, 2022, 2024) in order to collect more palaeontological data, to better understand the soft tissue preservation and to precise the stratigraphic log of the locality (Houée etal. 2022a, b; Logghe etal. 2023a). Here we describe new Discosauriscus specimens from the latest fieldworks, with exceptionally preserved complete skin coverage identified as epidermal skin, and a cololite recovered in situ within a complete skeleton. We also precise the stratigraphic log of the locality. These new discoveries allow: 1) the identification of seymouriamorph skin soft tissues and its comparison with the cutaneous coverage of other Palaeozoic tetrapods; 2) the study of a seymouriamorph cololite; 3) the discussion of these new data in terms of palaeoecology (e.g. skin adaptation to an aquatic or amphibious/terrestrial life), and palaeoenvironment; and 4) to better understand the mass mortality of Discosauriscus from Franchesse and the exceptional quality of preservation of the described soft-tissue and cololite remains. GEOLOGICAL SETTING The Les Charbonnières site in Le Pontet, 4 km near the village of Franchesse, is located in the north of the Bourbonl’Archambault Basin (Allier, Massif Central, France), approx. 10 km away from the early Permian historic site of La Queuede-l’Étang (de Saint-Seine 1949; Heyler 1969). More precisely, this site belongs to the Souvigny sub-basin (Steyer etal. 2012; Fig. 1A). As with the other Carboniferous-Permian basins of Autun, Blanzy-Le Creusot and Decize-La Machine, the Bourbon-l’Archambault Basin is located in the northern part of the Massif Central (Debriette 1992), belonging to the Variscan Belt of the Central Pangea (Châteauneuf & Farjanel 1989). The fossil-bearing horizons of the Les Charbonnières site come from the upper-most shale sequence of the Renière B Member of the Renière Formation (Steyer etal. 2012). The Renière Formation has been identified from the site of Buxières-les-Mines, south of the Aumance sub-basin of the Bourbon-l’Archambault Basin (Steyer etal. 2000, 2012; Kaulfuss 2003; Fig. 1B). The underlying Buxières Formation may be of early Artinskian age (288 ± 4 Ma; Kaulfuss 2003 according to the International Chronostratigraphic Chart 2024-12) but recent U-Pb dating in the Buxières Formation rather proposed a lowermost Asselian age (Cisuralian, lower Permian) of 298.592 ± 0.075/0.15/0.35 Ma (Mercuzot etal. 2023). An absolute age of the Les Charbonnières site has not been calculated so far, but a Sakmarian age has been proposed based on the previously assigned Sakmarian age of the Buxières Formation (Steyer etal. 2012, based on the ICC 2010-09). An older age of the Les Charbonnières site is in this way possible regarding the new radiometric ages of the Buxières Formation. Valli (2007) conducted the first stratigraphic survey of the Les Charbonnières site (Steyer etal. 2022: fig. 2A) and observed a strong horizontal variability of the layers in the span a of few meters, with changes in both sedimentology and fossil remain contents. Our recent fieldworks (excavations 492 COMPTES RENDUS PALEVOL • 2025 • 24 (24) Logghe A. et al. fig. 1. — Geological context of the Les Charbonnières site near Franchesse, Bourbon-l’Archambault Basin (Allier, France) with locations of the main fossiliferous deposits of La Queue-de-l’Étang, Buxières-les-Mines and the early Permian Lagerstâtte of Franchesse: A, simplified map of the Carboniferous and Permian Bourbon-l’Archambault Basin (modified after Paquette & Feys 1989); B, log of the main localities, from bottom to top: Buxières-les-Mines, Franchesse, La Queuede-l’Étang; red star, stratigraphic position of the Les Charbonnières site (modified after Debriette 1992); C, detailed stratigraphic log of the SM sector of the Les Charbonnières site near Franchesse. post-Permian rocks lower Permian upper Carboniferous Crystalline basement Gipcy-Boubon ridge ? A BC 100 m 5 km N Franchesse Franchesse Bourbon-l'Archambault Bourbonl'Archambault Aumance sub-basin Souvigny sub-basin Buxieres Buxieresles-Mines Allier river Murat fault Renière Formation Renière B At At At At At At Renière A Permian Cisuralian Buxières Formation Supra Buxières Infra Buxières Basal conglo. Inférieure Formation Carboniferous Pennsylvanian Sector SM Crystalline rock Argilites Sandstone Oil-shale beds Ash-tuff (At) Soil Clay Claystone Siltstone Cinerite Gypsum Discosauriscus Kuhn, 1933 Plant fossil Actinopterygians 493 Palaeoecology and palaeoenvironment of Discosauriscus COMPTES RENDUS PALEVOL • 2025 • 24 (24) and detailed stratigraphy; Fig. 1C) have been realised in the sector SM (see Steyer etal. 2022: fig. 2A), which showed a high potential for recovering Discosauriscus specimens through drillings (Valli 2007; Steyer etal. 2022). The stratigraphy of this sector is composed of, from bottom to top (Fig. 1C): an alternance (15 cm) of crystallised gypsum, a possible cinerite and shales; an alternance (4 cm) of claystones, clays and a dark indurated shale level (3 cm) with numerous well-preserved Discosauriscus remains of various sizes. From this layer, rare plant fragments and four slabs containing (sub)complete Discosauriscus – their taphonomy will be the subject of another article – were sampled. Above, a thin (0.5 cm) orange cinerite, sampled for absolute age investigations, is associated with crystallised gypsum and white to grey clay. It is topped by a dark indurate claystone (10 cm) rich in plant fragments, recovered in association with another possible cinerite layer (1 cm, also sampled). This plant layer is topped by an alternance of clay (1 cm) with crystallised gypsum, paper shale layers (5 cm), then a darker thin claystone layer (4 cm) with numerous actinopterygian remains. Above this “fish”-layer, numerous specimens of Discosauriscus have been found in a grey shale level (10 cm). They are exceptionally well-preserved and abundant, with several complete specimens exposed side by side (e.g. Steyer etal. 2012, 2022; Houée etal. 2022a; Logghe etal. 2023a). The material described here comes from this layer. It has been recovered in association with an fig. 2. — New specimens of Discosauriscus sp. from the Permian of Franchesse: A, RH-FR-2022-01, complete and articulated juvenile individual (only the right forelimb is missing) preserving dermal soft-tissue alongside the whole body; B, RH-FR-2022-07, sub-complete individual of poor cranial and soft-tissue preservation but yielding an in situ mold of the intestine; C, RH-FR-2022-03, sub-complete individual preserving dermal soft-tissue alongside the whole body. Scale bars: A, B, 5 cm; C, 3 cm. Credits: Lilian Cazes (CR2P). A B C 494 COMPTES RENDUS PALEVOL • 2025 • 24 (24) Logghe A. et al. orange dusty layer, possibly another cinerite (2 cm), sampled as well to precise its nature and run absolute dating analyses. Above was observed a grey-yellowish claystone (17 cm) with mostly well-preserved actinopterygians, rare chondrichthyan spines and some large and sub-complete Discosauriscus at the base. Finally, this layer is topped by dark and weathered claystones (20 cm) with isolated small Discosauriscus fragments and numerous more or less complete actinopterygians, and then the grass soil. MATERIAL AND METHODS We sampled four specimens of articulated Discosauriscus (RH-FR-2022-01, RH-FR-2022-02, RH-FR-2022-03 and RH-FR-2022-07; Figs 2; 3) whose soft tissues are preserved, including a specimen containing a cololite found in situ in the abdomen (RH-FR-2022-07; Fig. 2B). All these specimens were collected from the Permian of Franchesse (see the Geological Setting section for more details) and are part of the collections of the Rhinopolis Association, which aims to promote and preserve the palaeontological heritage of the Département de l’Allier (Allier county, France). All specimen images were taken using a Nikon D800 camera (apn full-frame 36 Mpx) with a lens Makro-Planar Zeiss 50 mm. For macro, we used a Canon 5DSR camera (apn full-frame 50Mpx) with a Canon MP-E65 mm lens mounted on micrometer rail and illuminated using cold light on fiber. The images were processed in Focus Stacking using Helicon Focus (v. 8.3.0). The images were taken and processed by Lilian Cazes (CR2P). The specimens are stored in the collections of the Rhinopolis Association in Varennes-sur-Allier (Allier, France). IDENTIFICATION SyStematicS The studied specimens show an exquisite preservation of the soft tissues but a very poor preservation of the skeletal elements, especially those from the skull: this renders their systematic identification very difficult at the species level. The aim of this paper is not to give a detailed systematic palaeontological description of the whole skeletal elements. However, some skeletal elements allow an identification at the family and genus level: 1) all the specimens have relatively short trunks and stout limbs. RH-FR-2022-02 preserves at least 24 presacral vertebrae (Fig. 2C), and RH-FR-2022-01 small ribs at the level of the cervical vertebrae and a left foot with a phalangeal formula of 2-3-4-5-3 (Fig. 2A). Also, RH-FR-2022-03 shows swollen neural arches, large cylindrical pleurocentra, and haemal arches at the level of the tail (Fig. 3): these characters are often seen in Discosauriscidae (e.g. Klembara & Bartík 1999; Laurin 2010). 2) these specimens have relatively short snouts, rounded orbits lying anteriorly to the middle of the skull length, and a relatively high skull in the region of the otic notches when preserved: these characters are typical of the genus Discosauriscus, e.g. Klembara 1997. Unfortunately, as mentioned above, the peculiar preservation of these specimens does not allow to identify the species: pending a more thorough systematic study of the whole material of Franchesse (composed of hundreds of specimens), we therefore prefer to stay prudent and to refer the studied specimens as to Discosauriscus sp. This identification is consistent with that of Steyer etal. (2012, 2022). fig. 3. — New specimen (RH-FR-2022-02) of Discosauriscus sp. from the Permian of Franchesse, preserving dermal soft-tissues: posterior body region (left), and anterior body region (right) of a probable unique individual. Scale bar: 5 cm. Credits: Lilian Cazes (CR2P). 495 Palaeoecology and palaeoenvironment of Discosauriscus COMPTES RENDUS PALEVOL • 2025 • 24 (24) Growth StaGe The high ossification degree of the preserved long bones, ribs and vertebrae; the well-developed ornamentation pattern of the dermal bones; the fact that the orbits are relatively anteriorly located within the skull; the relatively long postorbital region; and the robust mandibles preserving well-developed teeth are typical for a late juvenile ontogenetic stage (Klembara 1997; Steyer 2000; Sanchez etal. 2008). However, according to Klembara (2009) who distinguished ontogenetic stages based on skull length only, these specimens might be younger in somatic age. Considering all the characters together, we prefer to stay prudent and to give an “average” juvenile age for the studied specimens. DESCRIPTION OF THE SOFT TISSUES Putative ePidermal ScaleS (Figs 4, 5) Several rows of scales are well-preserved on RH-FR-2022-01 (Fig. 4A, B). These scales seem to be positioned on the flanks and/or dorsal side of the specimen as they do not appear to be below the vertebral column but rather constrained around it. These scaly skin remains are mostly preserved on the body region of the other Discosauriscus skeletons, and less detailed skin remains are visible around the tails. Skin remains from the shoulder girdles and pelvic regions are seldomly visible (see RH-FR-2022-03; Fig. 4C), and overall appear smoother than in the thoracic region, and deprived of scales. These skin remains present a blackish coloration, consistent with that of the rest of the skeleton and with other tetrapod skin remains recovered from the Franchesse locality, and contrasting with the grey colour of the substrate. They are preserved in thee-dimensions (Fig. 4B), and are therefore not the result of the impression of the skin on the substrate. The scales are mostly triangular (Fig. 4B), although some seem rather quadrangular (Fig. 4C). They measure between 1 and 1.5-1.7 mm. No concentric rings seem to be preserved within the scales. However, they seem to preserve ridges, that follow the overall shape of the scales (Fig. 5A). Some kind of tubercule-like ornamentation is also seen on some scales, however without definite A B C D fig. 4. — Close-ups on dermal soft-tissues of Discosauriscus sp. specimens from the Permian of Franchesse: A, RH-FR-2022-01, showing arrangement in longitudinal and transversal rows; B, macrophotography of the scales of RH-FR-2022-01 showing the triangular (to quadrangular in some regions) shape of the scales. The arrangement in longitudinal and transversal rows is well visible here; C, RH-FR-2022-02, showing a similar arrangement of the scales as in the other specimens, preserving however a more quadrangular shape of the scales; D, RH-FR-2022-03, showing the longitudinal and transversal arrangement of the scales. The rows seem less straight than those in RH-FR-2022-01 but display a more ‘wavy’ arrangement alongside the body. Scale bars: A, D, 2 cm; B, 2 mm; C, 1 cm. 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