Postcranial anatomy of Besanosaurus leptorhynchus (Reptilia: Ichthyosauria) from the Middle Triassic Besano Formation of Monte San Giorgio (Italy/Switzerland), with implications for reconstructing the swimming styles of Triassic ichthyosaurs.

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. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-024-00330-9.

Swiss ichthyosaurs: a review.

Switzerland is an ichthyosaur country: it has a rich record of marine reptile fossils, particularly the fish-shaped ichthyosaurs, and the according research. Here, we provide an overview over the 12 or more genera and at least 13 species plus numerous fragmentary remains of ichthyosaurs from the Triassic to the Cretaceous that have been discovered in twelve cantons thus far, of which four species are based on Swiss holotypes. This wealth of ichthyosaur species can be explained by their abundance in the Middle Triassic conservation deposits (Konservat Lagerstätte) of Monte San Giorgio, as well as occasional discoveries in strata of Middle Triassic to Early Cretaceous age. The moderate abundance of outcrops in reasonable conditions in combination with the long history of palaeontological research in Switzerland explains this good fossil record. In addition to this unique overview, we provide more data for further studies and update the knowledge of these taxa.

Oldest record of Machimosaurini (Thalattosuchia, Teleosauroidea): teeth and scavenging traces from the Middle Jurassic (Bajocian) of Switzerland.

The Jurassic period was a time of major diversification for Mesozoic marine reptiles, including Ichthyosauria, Plesiosauria and thalattosuchian Crocodylomorpha. The latter originated in the Early Jurassic and thrived during the Late Jurassic. Unfortunately, the Middle Jurassic, a crucial time in their evolution, has a poor fossil record. Here, we document the first evidence of macrophagous/durophagous Machimosaurini-tribe teleosauroid thalattosuchians from the late Bajocian (ca 169 Ma) in the form of three robust tooth crowns with conical blunt shapes and anastomosed pattern of thick enamel ridges towards the apex, associated with the skeleton of a large ichthyosaur lacking preserved tooth crowns. The tooth crowns were found on the posterior section of the lower jaw (left angular), a lacrimal and the axis neural arch of the ichthyosaur. In addition, some of the distal sections of the posterior dorsal ribs of the ichthyosaur skeleton exhibit rounded bite marks and some elongated furrows that fit in size and shape with the Machimosaurini teeth. These marks, together with the absence of healing in the rib bone are interpreted here as the indicators of peri- to post-mortem scavenging by a Machimosaurini teleosauroid after the large ichthyosaur carcass settled on the floor of a shallow ocean.

A large new Middle Jurassic ichthyosaur shows the importance of body size evolution in the origin of the Ophthalmosauria.

The Middle Jurassic is an important time period for the evolutionary history of marine reptiles as it represented a transitional phase for many clades. Notably, in ichthyosaurs, many early parvipelvian taxa went extinct. The Middle Jurassic saw the emergence of the derived Ophthalmosauria, ultimately becoming the dominant ichthyosaurian clade by the end of the epoch. Even though this is an important period in the evolutionary history of Ophthalmosauria, our understanding remains limited in terms of morphology and taxonomy due to the scarcity of vertebrate-bearing strata. Here we present a large new ichthyosaur from the Bajocian of Switzerland, represented by an almost complete skull with 3D-preserved bones, the (inter)clavicles and a large portion of the postcranial skeleton. After CT- and surface scanning, we reconstructed the 3D in vivo morphology. Our morphological observations and phylogenetic analyses show that the new taxon named Argovisaurus martafernandezi is nested at the base of the Ophthalmosauria. The holotype and only known specimen of Argovisaurus likely represents an adult individual. Bajocian members of the Ophthalmosauria (Mollesaurus and Argovisaurus) were large-bodied animals, a trait typically associated with the more derived Platypterygiinae. This hints at the importance of a large body size early in ophthalmosaurian evolution.LSID: urn:lsid:zoobank.org:act:C3312628-1544-4B87-BBE3-B12346A30BE3LSID: urn:lsid:zoobank.org:act:23C2BD71-8CF0-4D99-848A-0D631518415B.

Ontogenetic variation in the cranium of Mixosaurus cornalianus, with implications for the evolution of ichthyosaurian cranial development.

Relatively complete ontogenetic series are comparatively rare in the vertebrate fossil record. This can create biases in our understanding of morphology and evolution, since immaturity can represent a source of unrecognized intraspecific variation in both skeletal anatomy and ecology. In the extinct marine reptile clade Ichthyopterygia, ontogenetic series were widely studied only in some Jurassic genera, while the ontogeny of the oldest and most basal members of the clade is very poorly understood. Here, we investigate cranial ontogeny in Mixosaurus cornalianus, from the Middle Triassic Besano Formation of the Swiss and Italian Alps. This small-bodied taxon is represented by a wealth of material from multiple size classes, including fetal material. This allows us to assess ontogenetic changes in cranial morphology, and identify stages in the ontogenetic trajectory where divergence with more derived ichthyosaurs has occurred. Early ontogenetic stages of Mixosaurus show developmental patterns that are reminiscent of the presumed ancestral (early diverging sauropsid) condition. This is prominently visible in the late fetal stage in both the basioccipital, which shows morphology akin to basal tubera, and in the postorbital, which has a triradiate head. The ontogenetic trajectory of at least some of the cranial elements of Mixosaurus is therefore likely still very akin to the ancestral condition, even though the adult cranium diverges from the standard diapsid morphology. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00289-z.

Heads or tails first? Evolution of fetal orientation in ichthyosaurs, with a scrutiny of the prevailing hypothesis.

According to a longstanding paradigm, aquatic amniotes, including the Mesozoic marine reptile group Ichthyopterygia, give birth tail-first because head-first birth leads to increased asphyxiation risk of the fetus in the aquatic environment. Here, we draw upon published and original evidence to test two hypotheses: (1) Ichthyosaurs inherited viviparity from a terrestrial ancestor. (2) Asphyxiation risk is the main reason aquatic amniotes give birth tail-first. From the fossil evidence, we conclude that head-first birth is more prevalent in Ichthyopterygia than previously recognized and that a preference for tail-first birth likely arose in derived forms. This weakens the support for the terrestrial ancestry of viviparity in Ichthyopterygia. Our survey of extant viviparous amniotes indicates that fetal orientation at birth reflects a broad diversity of factors unrelated to aquatic vs. terrestrial habitat, further undermining the asphyxiation hypothesis. We propose that birth preference is based on parturitional mechanics or carrying efficiency rather than habitat.

Ontogenetic variation in the skull of Stenopterygius quadriscissus with an emphasis on prenatal development.

The availability of a large sample size from a range of ontogenetic stages makes Stenopterygius quadriscissus a good model to study ontogenetic variation in a fossil sauropsid. We qualitatively examined pre- and postnatal ontogenetic changes in the cranium of S. quadriscissus. The prenatal ossification sequence is similar to other diapsids, exhibiting delayed chondrocranial ossification compared to the dermatocranium. In the dermatocranium, the circumorbital area is more ossified earlier in development relative to other elements, especially those of the skull roof where ossification is comparatively weaker across prenatal stages. Perinatally all cranial elements are ossified, and many scarf and step joints are already closed. We propose four prenatal and three postnatal stages in S. quadriscissus on the basis of relative ossification, size and qualitative cranial characters pertaining to the jugal, parietal, frontal, pterygoid and surangular. These will provide a basis for determining ontogenetic stages in other ichthyosaurs. Moreover, our postnatal observations aid in refining ontogenetic characters for phylogenetic studies. Lastly, we observed that the antimeric sutures of the midline of the skull roof are open perinatally and that fusion of the midline only appears in the adult stage. We hypothesize that the loose connection of the midline functions as a fontanelle, limiting potential damage during birth.

Modeling tooth enamel in FEA comparisons of skulls: Comparing common simplifications with biologically realistic models.

Palaeontologists often use finite element analyses, in which forces propagate through objects with specific material properties, to investigate feeding biomechanics. Teeth are usually modeled with uniform properties (all bone or all enamel). In reality, most teeth are composed of pulp, dentine, and enamel. We tested how simplified teeth compare to more realistic models using mandible models of three reptiles. For each, we created models representing enamel thicknesses found in extant taxa, as well as simplified models (bone, dentine or enamel). Our results suggest that general comparisons of stress distribution among distantly related taxa do not require representation of dental tissues, as there was no noticeable effect on heatmap representations of stress. However, we find that representation of dental tissues impacts bite force estimates, although magnitude of these effects may differ depending on constraints. Thus, as others have shown, the detail necessary in a biomechanical model relates to the questions being examined.

Cranial anatomy of Besanosaurus leptorhynchus Dal Sasso & Pinna, 1996 (Reptilia: Ichthyosauria) from the Middle Triassic Besano Formation of Monte San Giorgio, Italy/Switzerland: taxonomic and palaeobiological implications.

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.

Cranial morphology of the tanystropheid Macrocnemus bassanii unveiled using synchrotron microtomography.

The genus Macrocnemus is a member of the Tanystropheidae, a clade of non-archosauriform archosauromorphs well known for their very characteristic, elongated cervical vertebrae. Articulated specimens are known from the Middle Triassic of Alpine Europe and China. Although multiple articulated specimens are known, description of the cranial morphology has proven challenging due to the crushed preservation of the specimens. Here we use synchrotron micro computed tomography to analyse the cranial morphology of a specimen of the type species Macrocnemus bassanii from the Besano Formation of Monte San Giorgio, Ticino, Switzerland. The skull is virtually complete and we identify and describe the braincase and palatal elements as well the atlas-axis complex for the first time. Moreover, we add to the knowledge of the morphology of the skull roof, rostrum and hemimandible, and reconstruct the cranium of M. bassanii in 3D using the rendered models of the elements. The circumorbital bones were found to be similar in morphology to those of the archosauromorphs Prolacerta broomi and Protorosaurus speneri. In addition, we confirm the palatine, vomer and pterygoid to be tooth-bearing palatal bones, but also observed heterodonty on the pterygoid and the palatine.