Despite their profound adaptations to the aquatic realm and their apparent success throughout the Triassic and the Jurassic, ichthyosaurs became extinct roughly 30 million years before the end-Cretaceous mass extinction. Current hypotheses for this early demise involve relatively minor biotic events, but are at odds with recent understanding of the ichthyosaur fossil record. Here, we show that ichthyosaurs maintained high but diminishing richness and disparity throughout the Early Cretaceous. The last ichthyosaurs are characterized by reduced rates of origination and phenotypic evolution and their elevated extinction rates correlate with increased environmental volatility. In addition, we find that ichthyosaurs suffered from a profound Early Cenomanian extinction that reduced their ecological diversity, likely contributing to their final extinction at the end of the Cenomanian. Our results support a growing body of evidence revealing that global environmental change resulted in a major, temporally staggered turnover event that profoundly reorganized marine ecosystems during the Cenomanian.

Ichthyosauria is a diverse clade of marine amniotes that spanned most of the Mesozoic. Until recently, most authors interpreted the fossil record as showing that three major extinction events affected this group during its history: one during the latest Triassic, one at the Jurassic–Cretaceous boundary (JCB), and one (resulting in total extinction) at the Cenomanian-Turonian boundary. The JCB was believed to eradicate most of the peculiar morphotypes found in the Late Jurassic, in favor of apparently less specialized forms in the Cretaceous. However, the record of ichthyosaurs from the Berriasian–Barremian interval is extremely limited, and the effects of the end-Jurassic extinction event on ichthyosaurs remains poorly understood. Based on new material from the Hauterivian of England and Germany and on abundant material from the Cambridge Greensand Formation, we name a new ophthalmosaurid, Acamptonectes densus gen. et sp. nov. This taxon shares numerous features with Ophthalmosaurus, a genus now restricted to the Callovian–Berriasian interval. Our phylogenetic analysis indicates that Ophthalmosauridae diverged early in its history into two markedly distinct clades, Ophthalmosaurinae and Platypterygiinae, both of which cross the JCB and persist to the late Albian at least. To evaluate the effect of the JCB extinction event on ichthyosaurs, we calculated cladogenesis, extinction, and survival rates for each stage of the Oxfordian–Barremian interval, under different scenarios. The extinction rate during the JCB never surpasses the background extinction rate for the Oxfordian–Barremian interval and the JCB records one of the highest survival rates of the interval. There is currently no evidence that ichthyosaurs were affected by the JCB extinction event, in contrast to many other marine groups. Ophthalmosaurid ichthyosaurs remained diverse from their rapid radiation in the Middle Jurassic to their total extinction at the beginning of the Late Cretaceous.

A new ophthalmosaurid ichthyosaur, Sveltonectes insolitus, gen. et sp. nov., is described from a sub-complete and three-dimensionally preserved specimen from the late Barremian of western Russia. This new taxon is supported by 11 cranial, dental, and postcranial autapomorphies, and is also characterized by features previously considered as autapomorphic for some other Ophthalmosauridae, such as a processus narialis on the prefrontal and relatively long hind fins with pre- and postaxial accessory digits. We conducted a new phylogenetic analysis of Thunnosauria, which supports a ‘Stenopterygius’ origin for Ophthalmosauridae. Sveltonectes is regarded as the sister taxon of Aegirosaurus, which shares a similar skull roof construction. Contrary to most other Cretaceous ichthyosaurs, Sveltonectes is characterized by delicate and sharply pointed teeth, confirming that the ophthalmosaurids were ecologically highly diversified during the Early Cretaceous.

Ophthalmosaurinae is a recently recognized clade of derived ichthyosaurs (marine reptiles) ranging from the Bajocian (Middle Jurassic) to the late Albian (late Early Cretaceous). Whereas the Middle–Late Jurassic ophthalmosaurineOphthalmosaurusis often regarded as a hyperspecialized deep diver, very little is known about the anatomy, evolutionary history and ecology of Cretaceous ophthalmosaurines because of the scarcity of the fossils and the lack of well-preserved skull material. Here, we describe the skull of a new basal ophthalmosaurine ichthyosaur,Leninia stellansgen. et sp. nov., from the lower Aptian of western Russia, and compare the ocular characteristics of ophthalmosaurids.Leniniais recovered as a basal ophthalmosaurine; it possesses unique traits such as a star-shaped frontal–parietal suture as well as features previously thought to be unique toOphthalmosaurussuch as a supratemporal–stapes contact. A large sclerotic aperture – significantly larger than in platypterygiine ophthalmosaurids and similar to that of the largest-eyed modern animals (giant and colossal squids) – and reduced dentition appear widespread within ophthalmosaurines. This conservatism suggests ophthalmosaurine ophthalmosaurids occupied similar ecological niche(s) throughout their long evolutionary history.

Ichthyosaurs are reptiles that inhabited the marine realm during most of the Mesozoic. Their Cretaceous representatives have traditionally been considered as the last survivors of a group declining since the Jurassic. Recently, however, an unexpected diversity has been described in Upper Jurassic–Lower Cretaceous deposits, but is widely spread across time and space, giving small clues on the adaptive potential and ecosystem control of the last ichthyosaurs. The famous but little studied English Gault Formation and ‘greensands’ deposits (the Upper Greensand Formation and the Cambridge Greensand Member of the Lower Chalk Formation) offer an unprecedented opportunity to investigate this topic, containing thousands of ichthyosaur remains spanning the Early–Late Cretaceous boundary. To assess the diversity of the ichthyosaur assemblage from these sedimentary bodies, we recognized morphotypes within each type of bones. We grouped these morphotypes together, when possible, by using articulated specimens from the same formations and from new localities in the Vocontian Basin (France); a revised taxonomic scheme is proposed. We recognize the following taxa in the ‘greensands’: the platypterygiines ‘Platypterygius’ sp. and Sisteronia seeleyi gen. et sp. nov., indeterminate ophthalmosaurines and the rare incertae sedis Cetarthrosaurus walkeri. The taxonomic diversity of late Albian ichthyosaurs now matches that of older, well-known intervals such as the Toarcian or the Tithonian. Contrasting tooth shapes and wear patterns suggest that these ichthyosaurs colonized three distinct feeding guilds, despite the presence of numerous plesiosaur taxa. Western Europe was a diversity hot-spot for ichthyosaurs a few million years prior to their final extinction. By contrast, the low diversity in Australia and U.S.A. suggests strong geographical disparities in the diversity pattern of Albian–early Cenomanian ichthyosaurs. This provides a whole new context to investigate the extinction of these successful marine reptiles, at the end of the Cenomanian.