Cecily S. C. Nicholl

Crocodylomorph diversity in Europe peaked during the Late Cretaceous, with eusuchians dominating the fossil record. However, ziphodont forms, including the enigmatic Doratodon, are increasingly recognized from multiple Santonian–Maastrichtian European localities. A new occurrence from the earliest Campanian Villaggio del Pescatore site in northeastern Italy represents the most complete notosuchian from the latest Cretaceous Mediterranean carbonate platforms. High-resolution micro-computed tomographic imaging reveals key dental features, including alternate, asymmetric tooth replacement and varied tooth morphologies lacking denticulated carinae. The preserved neurovascular architecture of the specimen also provides new data on trigeminal innervation, suggesting high tactile sensitivity. Comparisons with both eastern and western European taxa support the hypothesis of biogeographical partitioning between the western Ibero-Armorican region, the Adriatic–Dinaric Carbonate Platform (ADCP), and the remaining eastern European archipelago. The Villaggio del Pescatore occurrence increases the faunal diversity from the site and strengthens the biogeographical links between Italian and other eastern European faunas, while also emphasizing evolutionary divergence from western forms, such as Doratodon ibericus. This specimen bridges a gap in the geographical distribution of Doratodon-like forms and suggests intermittent faunal exchanges across ADCP landmasses, underscoring the pivotal role of the ADCP in the persistence, diversification, and dispersal of ziphodont crocodylomorphs and other vertebrates across the Late Cretaceous European Archipelago.

Sebecid notosuchians are the only terrestrial crocodyliforms to survive the Cretaceous–Palaeogene extinction, 66 Ma, which eliminated large-bodied species (above approximately 5 kg) in terrestrial ecosystems. Early sebecid evolution is unclear due to the scarcity of remains from both sides of the boundary. We present the stratigraphically earliest post-extinction notosuchian record, from the lower Palaeocene Salamanca Formation of Patagonia. Tewkensuchus salamanquensis n. gen. n. sp. has unique features, including a skull roof with elevated lateral margins, and an accessory peg and socket articulation between the postorbital and posterior palpebral. Our phylogenetic analysis allies Tewkensuchus with a clade of predatorial crocodyliforms from the Eocene of Europe (and possibly of Africa, as Eremosuchus may also belong to this clade). This clade forms the sister taxon of South American sebecids. We name Sebecoidea for this more inclusive clade of Eurogondwanan notosuchians and suggest that its spatial distribution reflects earlier diversification and dispersal events, which are only partially known. We estimate a body mass of around 300 kg for Tewkensuchus , one of the largest known notosuchians. Phylogenetic optimization of notosuchian body size change reconstructs a Cretaceous–Palaeogene boundary-crossing sebecoidean lineage with an estimated mass between 332 and 443 kg. This provides the first support for the survival of a large-bodied terrestrial vertebrate lineage across the K–Pg boundary.

Notosuchia is a large and diverse clade of Mesozoic crocodyliforms that thrived in continental environments especially during the Cretaceous of Gondwana. This clade has been the focus of several phylogenetic studies that led to the consensus of two main topological arrangements, mostly differing in the position of the clade Sebecidae: the sebecosuchian and sebecian hypotheses. In this manuscript we review the main phylogenetic definitions of several clades of notosuchians in the context of these hypotheses. As a result, besides the clade names previously established under the ICPN/Phylocode (including registration in RegNum) by Ruiz et al. (2021) (Notosuchia, Xenodontosuchia, Sphagesauria, Sphagesauridae) and Darlim et al. (2021a) (Baurusuchia, Baurusuchidae, Baurusuchinae, Pissarrachampsinae), eleven clade names have been registered and are defined in this contribution. Ten of these represent converted clade names (Uruguaysuchidae, Mahajangasuchidae, Peirosauridae, Itasuchidae, Ziphosuchia, Sphagesaurinae, Caipirasuchinae, Sebecia, Sebecosuchia, and Sebecia), while one represents a new clade name (Peirosauria). Furthermore, the definition of Xenodontosuchia is emended (unrestricted emendation) so that the name can be used in the context of a wider array of topologies.

The open‐source programming language ‘R' has become a standard tool in the palaeobiologist's toolkit. Its popularity within the palaeobiological community continues to grow, with published articles increasingly citing the usage of R and R packages. However, there are currently a lack of agreed standards for data preparation and available frameworks to support the implementation of such standards. Consequently, data preparation workflows are often unclear and not reproducible, even when code is provided. Moreover, due to a lack of code accessibility and documentation, palaeobiologists are often forced to ‘reinvent the wheel’ to find solutions to issues already solved by other members of the community. Here, we introduce palaeoverse, a community‐driven R package to aid data preparation and exploration for quantitative palaeobiological research. The package is freely available and has three core principles: (1) streamline data preparation and analyses; (2) enhance code readability; and (3) improve reproducibility of results. To develop these aims, we assessed the analytical needs of the broader palaeobiological community using an online survey, in addition to incorporating our own experiences. In this work, we first report the findings of the survey, which shaped the development of the package. Subsequently, we describe and demonstrate the functionality available in palaeoverse and provide usage examples. Finally, we discuss the resources we have made available for the community and our future plans for the broader Palaeoverse project. palaeoverse is a community‐driven R package for palaeobiology, developed with the intention of bringing palaeobiologists together to establish agreed standards for high‐quality quantitative research. The package provides a user‐friendly platform for preparing data for analysis with well‐documented open‐source code to enhance transparency. The functionality available in palaeoverse improves code reproducibility and accessibility, which is beneficial for both the review process and future research.

Notosuchians are an extinct clade of terrestrial crocodyliforms with a particularly rich record in the late Early to Late Cretaceous (approx. 130–66 Ma) of Gondwana. Although much of this diversity comes from South America, Africa and Indo-Madagascar have also yielded numerous notosuchian remains. Three notosuchian species are currently recognized from the early Late Cretaceous (approx. 100 Ma) Kem Kem Group of Morocco, including the peirosauridHamadasuchus rebouli. Here, we describe two new specimens that demonstrate the presence of at least a fourth notosuchian species in this fauna.Antaeusuchus taouzensisn. gen. n. sp. is incorporated into one of the largest notosuchian-focused character-taxon matrices yet to be compiled, comprising 443 characters scored for 63 notosuchian species, with an increased sampling of African and peirosaurid species. Parsimony analyses run under equal and extended implied weighting consistently recoverAntaeusuchusas a peirosaurid notosuchian, supported by the presence of two distinct waves on the dorsal dentary surface, a surangular which laterally overlaps the dentary above the mandibular fenestra, and a relatively broad mandibular symphysis. Within Peirosauridae,Antaeusuchusis recovered as the sister taxon ofHamadasuchus. However, it differs fromHamadasuchuswith respect to several features, including the ornamentation of the lateral surface of the mandible, the angle of divergence of the mandibular rami, the texture of tooth enamel and the shape of the teeth, supporting their generic distinction. We present a critical reappraisal of the non-South American Gondwanan notosuchian record, which spans the Middle Jurassic–late Eocene. This review, as well as our phylogenetic analyses, indicate the existence of at least three approximately contemporaneous peirosaurid lineages within the Kem Kem Group, alongside other notosuchians, and support the peirosaurid affinities of the ‘trematochampsid’Miadanasuchus oblitafrom the Maastrichtian of Madagascar. Furthermore, the Cretaceous record demonstrates the presence of multiple lineages of approximately contemporaneous notosuchians in several African and Madagascan faunas, and supports previous suggestions regarding an undocumented pre-Aptian radiation of Notosuchia. By contrast, the post-Cretaceous record is depauperate, comprising rare occurrences of sebecosuchians in north Africa prior to their extirpation.

Once a much more globally widespread crocodylian clade, Tomistominae is today represented by just one species, Tomistoma schlegelii (the false gharial), restricted to south-east Asia. Although tomistomine fossil occurrences are recognized from the early Eocene (∼55 Ma) onwards, their remains are often incomplete, making appropriate taxonomic classification within the group problematic. This is especially pertinent to several taxa from the Miocene of Europe, which were historically erected from fragmentary remains. Here we re-examine and describe four approximately contemporaneous taxa from Malta and Italy to determine their taxonomy and phylogenetic affinities: Melitosaurus champsoides, Tomistoma calaritanum, Tomistoma gaudense and Tomistoma lyceense. We place them into a phylogenetic analysis for the first time, comprising 70 taxa scored for 244 characters, several of which are revised or novel, and apply a number of character weighting strategies. Whereas ‘Tomistoma lyceense’ is deemed to be an indeterminate tomistomine, a unique combination of features confirms Melitosaurus champsoides, Tomistoma calaritanum and Tomistoma gaudense as three distinct species. These three taxa are recovered as derived tomistomines, with characters such as a posterior maxillary process between the lacrimal and nasal, large supratemporal fenestrae that are wider than long, and the posteromedial alignment of the last three premaxillary teeth, suggesting a close relationship with the approximately contemporaneous European taxa, Tomistoma lusitanica and Gavialosuchus eggenburgensis. It is unlikely that any of these species belong to Tomistoma, with the possibility that they can all be classified under Melitosaurus and Gavialosuchus. However, we retain them in open nomenclature pending reassessment of the remaining European Miocene tomistomines. Our taxonomic and phylogenetic revision helps to elucidate past tomistomine diversity in the Miocene of the Mediterranean region, prior to the group’s extirpation, and is an important first step in resolving the complicated history of European tomistomine systematics.