Search

Abstract Besanosaurus leptorhynchus Dal Sasso & Pinna, 1996 was originally described on the basis of a single complete fossil specimen excavated near Besano (Italy). However, a recent taxonomic revision and re-examination of the cranial osteology allowed for the assignment of five additional specimens to the taxon. Here, we analyse, describe and discuss the postcranial anatomy of Besanosaurus leptorhynchus in detail. The size of the specimens examined herein ranged from slightly more than one meter to eight meters. Overall, several diagnostic character states for this taxon are proposed, demonstrating a mosaic of plesiomorphic and derived features. This is best exemplified by the limbs, which show very rounded elements in the forelimbs, and pedal phalanges with retained rudimentary shafts. We suggest that the widely spaced phalanges in the forefins of Besanosaurus leptorhynchus were embedded in a fibrocartilage-rich connective tissue, like in modern cetaceans. We also review the similarities of Besanosaurus with Pessopteryx and Pessosaurus, allowing us to conclude that Besanosaurus is not a junior synonym of either of the two taxa. Lastly, to test the swimming capabilities of Besanosaurus leptorhynchus, we expanded on a previously published study focussing on reconstructing the swimming styles of ichthyosaurs. Besanosaurus leptorhynchus was found to possess a peculiar locomotory mode, somewhat intermediate between anguilliform swimmers, such as Cymbospondylus and Utatsusaurus, and some shastasaur-grade (e.g., Guizhouichthyosaurus) and early-diverging euichthyosaurian (e.g., Californosaurus) ichthyosaurs. Based on our results, we furthermore suggest that mixosaurids acquired their characteristic body profile (dorsal fin and forefins that are distinctly enlarged compared to the hindfins) independently and convergently to the one that later appeared in Parvipelvia. Moreover, the different swimming styles inferred for Cymbospondylus, Mixosauridae, and Besanosaurus strengthen the earlier hypothesis of niche partitioning among these three distinct ichthyosaur taxa from the Besano Formation.

Abstract Chaohusaurus is an early ichthyosauriform represented by three species known from the Early Triassic of Chaohu, Anhui Province, China, with a fourth species—Chaohusaurus zhangjiawanensis—known from the Nanzhang-Yuan’an region of Hubei Province. In contrast to the Chaohusaurus species from Chaohu, Chaohusaurus zhangjiawanensis remains poorly known, hindering our understanding of early ichthyosauriform evolution. Here, we report a new specimen of Chaohusaurus zhangjiawanensis, which provides new information on its dentition. The new specimen confirms that Chaohusaurus zhangjiawanensis had heterodont dentition consisting of pointed anterior teeth and robust, rounded posterior teeth, indicating a generalist diet. The posterior teeth of Chaohusaurus zhangjiawanensis are more robust (broader and larger) than the posterior dentition of Chaohusaurus chaoxianensis and Chaohusaurus brevifemoralis from Chaohu. This suggests differences in hard-shelled prey preference between species of Chaohusaurus from Chaohu and Nanzhang-Yuan’an, with Chaohusaurus zhangjiawanensis likely capable of feeding on harder and larger prey than Chaohusaurus brevifemoralis and Chaohusaurus chaoxianensis. In turn, this probably reflects differences in durophagous prey availability between the shallow-marine palaeoecosystem of Nanzhang-Yuan’an and the deeper, slope-basin palaeoecosystem of Chaohu. The posterior dentition and forefin of Chaohusaurus zhangjiawanensis are strikingly similar to those of Chaohusaurus geishanensis, the rarest species of Chaohusaurus from the Chaohu fauna. The scarcity of Chaohusaurus geishanensis in the Chaohu fauna, and its morphological similarity to Chaohusaurus zhangjiawanensis, possibly indicate that Chaohusaurus geishanensis was closely related with Chaohusaurus zhangjiawanensis and that it was also a shallow-marine species that was not a typical component of the Chaohu fauna. It probably occasionally wandered out into the deeper waters of Chaohu from a nearby coastal environment.

Ichthyosaurs, an iconic lineage of Mesozoic marine reptiles, were an important component of recovering ecosystems after the Permo-Triassic Mass Extinction event. Mixosauridae, a clade of small, early-diverging ichthyosaurs, were of particular significance for this process, being abundant predators in Middle Triassic shallow seas. Despite the abundance of well-preserved mixosaurid specimens from South China, Mixosaurus panxianensis remains the only comprehensively described species, hindering our understanding of the variability, taxonomy and diversity of mixosaurids from this re-gion. Here, we report a new species of Mixosaurus, Mixosaurus luxiensis, from Luxi County, Yunnan Province, South China. The wider postorbital skull portion differentiates the new species from Mixosaurus cornalianus and Mixosaurus kuhnschnyderi from central Europe. The non-durophagous dentition, composed of tiny piercing mesial teeth and robust but pointed distal teeth, resembles the dentition of M. cornalianus. However, the distal teeth of M. luxiensis sp. nov. are twice the size of the mesial ones, in contrast to M. cornalianus, in which the mesial and distal teeth are approximately equal in size. The forelimb exhibits a unique morphology, including a proportionally narrow radius, the presence of a peripheral notch on the ulna, and a large metacarpal V. A preliminary phylogenetic analysis suggests a close affinity of the new taxon with M. cornalianus from Western Tethys. Our study introduces important, new anatomical information on Mixosaurus from South China, useful for future studies of mixosaurid diversity.

Further evidence for an explosive marine reptile diversification event in the Early Triassic is presented via morphological analysis of a new Sclerocormus specimen.

Marine reptiles from the Upper Jurassic of Central Europe are rare and often fragmentary, which hinders their precise taxonomic identification and their placement in a palaeobiogeographic context. Recent fieldwork in the Kimmeridgian of Krzyżanowice, Poland, a locality known from turtle remains originally discovered in the 1960s, has reportedly provided additional fossils thought to indicate the presence of a more diverse marine reptile assemblage, including giant pliosaurids, plesiosauroids, and thalattosuchians. Based on its taxonomic composition, the marine tetrapod fauna from Krzyżanowice was argued to represent part of the “Matyja-Wierzbowski Line”—a newly proposed palaeobiogeographic belt comprising faunal components transitional between those of the Boreal and Mediterranean marine provinces. Here, we provide a detailed re-description of the marine reptile material from Krzyżanowice and reassess its taxonomy. The turtle remains are proposed to represent a “plesiochelyid” thalassochelydian (Craspedochelys? sp.) and the plesiosauroid vertebral centrum likely belongs to a cryptoclidid. However, qualitative assessment and quantitative analysis of the jaws originally referred to the colossal pliosaurid Pliosaurus clearly demonstrate a metriorhynchid thalattosuchian affinity. Furthermore, these metriorhynchid jaws were likely found at a different, currently indeterminate, locality. A tooth crown previously identified as belonging to the thalattosuchian Machimosaurus is here considered to represent an indeterminate vertebrate. The revised taxonomy of the marine reptiles from Krzyżanowice, as well as the uncertain provenance of the metriorhynchid specimen reported from the locality, cast doubt on the palaeobiogeographic significance of the assemblage.

Besanosaurus leptorhynchus Dal Sasso & Pinna, 1996 was described on the basis of a single fossil excavated near Besano (Italy) nearly three decades ago. Here, we re-examine its cranial osteology and assign five additional specimens to B. leptorhynchus, four of which were so far undescribed. All of the referred specimens were collected from the Middle Triassic outcrops of the Monte San Giorgio area (Italy/Switzerland) and are housed in various museum collections in Europe. The revised diagnosis of the taxon includes the following combination of cranial characters: extreme longirostry; an elongate frontal not participating in the supratemporal fenestra; a prominent ‘triangular process’ of the quadrate; a caudoventral exposure of the postorbital on the skull roof; a prominent coronoid (preglenoid) process of the surangular; tiny conical teeth with coarsely-striated crown surfaces and deeply-grooved roots; mesial maxillary teeth set in sockets; distal maxillary teeth set in a short groove. All these characters are shared with the holotype of Mikadocephalus gracilirostris Maisch & Matzke, 1997, which we consider as a junior synonym of B. leptorhynchus. An updated phylogenetic analysis, which includes revised scores for B. leptorhynchus and several other shastasaurids, recovers B. leptorhynchus as a basal merriamosaurian, but it is unclear if Shastasauridae form a clade, or represent a paraphyletic group. The inferred body length of the examined specimens ranges from 1 m to about 8 m. The extreme longirostry suggests that B. leptorhynchus primarily fed on small and elusive prey, feeding lower in the food web than an apex predator: a novel ecological specialisation never reported before the Anisian in a large diapsid. This specialization might have triggered an increase of body size and helped to maintain low competition among the diverse ichthyosaur fauna of the Besano Formation.

Lagomorpha is a clade of herbivorous mammals nested within Euarchontoglires, one of the major placental groups represented today. It comprises two extant families with markedly different body plans: the long-eared and long-limbed Leporidae (hares and rabbits) and the short-eared and short-limbed Ochotonidae (pikas). These two lagomorph lineages diverged probably during the latest Eocene/early Oligocene, but it is unclear whether the last common ancestor of crown lagomorphs was more leporid- or more ochotonid-like in morphology. Palaeolagus, an early lagomorph dominant in western North America from the late Eocene to Oligocene is of particular importance for addressing this controversy. Here, we present new and comprehensive data on the cranial anatomy of Palaeolagus haydeni, the type species for the genus, based on micro-computed tomography (μCT). Our μCT data allow us to confirm, revise and score for the very first time the states of several leporid-like and ochotonid-like characters in the skull of Palaeolagus. This mixed cranial architecture differentiates Palaeolagus from the crown groups of Lagomorpha and supports its phylogenetic status as a stem taxon.

Sauropterygia was a taxonomically and ecomorphologically diverse clade of Mesozoic marine reptiles spanning the Early Triassic to the Late Cretaceous. Sauropterygians are traditionally divided into two groups representing two markedly different body plans – the short-necked, durophagous Placodontia and the long-necked Eosauropterygia – whereas Saurosphargidae, a small clade of armoured marine reptiles, is generally considered as the sauropterygian sister-group. However, the early evolutionary history of sauropterygians and their phylogenetic relationships with other groups within Diapsida are still incompletely understood. Here, we report a new saurosphargid from the Early Triassic (Olenekian) of South China – Prosaurosphargis yingzishanensis gen. et sp. nov. – representing the earliest known occurrence of the clade. An updated phylogenetic analysis focussing on the interrelationships among diapsid reptiles recovers saurosphargids as nested within sauropterygians, forming a clade with eosauropterygians to the exclusion of placodonts. Furthermore, a clade comprising Eusaurosphargis and Palatodonta is recovered as the sauropterygian sister-group within Sauropterygomorpha tax. nov. The phylogenetic position of several Early and Middle Triassic sauropterygians of previously uncertain phylogenetic affinity, such as Atopodentatus, Hanosaurus, Majiashanosaurus, and Corosaurus, is also clarified, elucidating the early evolutionary assembly of the sauropterygian body plan. Finally, our phylogenetic analysis supports the placement of Testudines and Archosauromorpha within Archelosauria, a result strongly corroborated by molecular data, but only recently recovered in a phylogenetic analysis using a morphology-only dataset. Our study provides evidence for the rapid diversification of sauropterygians in the aftermath of the Permo-Triassic mass extinction event and emphasises the importance of broad taxonomic sampling in reconstructing phylogenetic relationships among extinct taxa.