No abstract

Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently proposed synonymy of both Spinosaurus maroccanus and Sigilmassasurus brevicollis with Spinosaurus aegyptiacus from the Cenomanian of Egypt, as there are clear differences between the vertebrae of these taxa, and they do not share any derived character that is not found in other spinosaurids. Together with a comparison with other spinosaurid vertebral material from the Kem Kem, this suggests that more than one taxon of spinosaurid was present in the Kem Kem assemblage of Morocco, so the referral of non-overlapping material from this unit to a single taxon should be regarded with caution.

Brachiosauridae is a clade of titanosauriform sauropod dinosaurs that includes the well-known Late Jurassic taxa Brachiosaurus and Giraffatitan. However, there is disagreement over the brachiosaurid affinities of most other taxa, and little consensus regarding the clade’s composition or inter-relationships. An unnamed partial sauropod skeleton was collected from middle–late Oxfordian (early Late Jurassic) deposits in Damparis, in the Jura department of eastern France, in 1934. Since its brief description in 1943, this specimen has been informally known in the literature as the ‘Damparis sauropod’ and ‘French Bothriospondylus’, and has been considered a brachiosaurid by most authors. If correctly identified, this would make the specimen the earliest known titanosauriform. Coupled with its relatively complete nature and the rarity of Oxfordian sauropod remains in general, this is an important specimen for understanding the early evolution of Titanosauriformes. Full preparation and description of this specimen, known from teeth, vertebrae and most of the appendicular skeleton of a single individual, recognises it as a distinct taxon: Vouivria damparisensis gen. et sp. nov. Phylogenetic analysis of a data matrix comprising 77 taxa (including all putative brachiosaurids) scored for 416 characters recovers a fairly well resolved Brachiosauridae. Vouivria is a basal brachiosaurid, confirming its status as the stratigraphically oldest known titanosauriform. Brachiosauridae consists of a paraphyletic array of Late Jurassic forms, with Europasaurus, Vouivria and Brachiosaurus recovered as successively more nested genera that lie outside of a clade comprising (Giraffatitan + Sonorasaurus) + (Lusotitan + (Cedarosaurus + Venenosaurus)). Abydosaurus forms an unresolved polytomy with the latter five taxa. The Early Cretaceous South American sauropod Padillasaurus was previously regarded as a brachiosaurid, but is here placed within Somphospondyli. A recent study contended that a number of characters used in a previous iteration of this data matrix are ‘biologically related’, and thus should be excluded from phylogenetic analysis. We demonstrate that almost all of these characters show variation between taxa, and implementation of sensitivity analyses, in which these characters are excluded, has no effect on tree topology or resolution. We argue that where there is morphological variation, this should be captured, rather than ignored. Unambiguous brachiosaurid remains are known only from the USA, western Europe and Africa, and the clade spanned the Late Jurassic through to the late Albian/early Cenomanian, with the last known occurrences all from the USA. Regardless of whether their absence from the Cretaceous of Europe, as well as other regions entirely, reflects regional extinctions and genuine absences, or sampling artefacts, brachiosaurids appear to have become globally extinct by the earliest Late Cretaceous.

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.