New information on Hauffiosaurus (Reptilia, Plesiosauria) based on a new species from the Alum Shale Member (Lower Toarcian: Lower Jurassic) of Yorkshire

An almost complete, three-dimensionally preserved plesiosaurian from the Hildoceras bifrons Zone of the Alum Shale Member (Whitby Limestone Formation; Lower Toarcian) of Yorkshire, UK, is described in detail. This represents a new species of Hauffiosaurus, H. tomistomimus, distinguished from H. zanoni (Harpoceras serpentinum Zone, Lower Toarcian, Germany) by the proportionally shorter neck and strongly concave preaxial margin of the tibia. It differs from H. longirostris (previously ‘Macroplata’ longirostris; Har. serpentinum Zone, Yorkshire) by the absence of prominent midline ridges on the dorsal surface of the premaxillae and ventral surface of the mandibular symphysis, and the absence of midline pterygoid contact ventral to the basioccipital. Several synapomorphies support a monophyletic Hauffiosaurus: broad longitudinal troughs occupy the dorsolateral surface of the maxilla and the posterior half of the lateral surface of the dentary; basicranial fontanelle bounded laterally by posterolaterally elongate projections of an undetermined ossification; and the neural arch contacts the rib facet in all postaxial cervical vertebrae. However, the systematic position of Hauffiosaurus, as a pliosauroid or basal plesiosauroid, remains uncertain. There is little evidence for geographic differentiation of Lower Toarcian plesiosaurian faunas in the United Kingdom and Germany as minor differences between abundant taxa may arise from temporal offset of fossils from these regions, and marked taxonomic differences are confined to rare taxa whose absence in one or other area may be attributable to incomplete sampling. Lack of consensus on the relationships of Lower Jurassic plesiosaurians requires further detailed description of Lower Jurassic taxa

Data from: A new pliosaurid (Sauropterygia, Plesiosauria) from the Oxford Clay Formation (Middle Jurassic, Callovian) of England: evidence for a gracile, longirostrine grade of Early–Middle Jurassic pliosaurids

A partial skeleton from the Sigiloceras enodatum ammonite Subzone (early Callovian, Middle Jurassic) of the Oxford Clay Formation of Quest Pit, near Stewartby, Bedfordshire, UK respresents one of the basalmost Middle Jurassic pliosaurids. Marmornectes candrewi gen. et sp. nov. possesses seven autapomorphies and a longirostrine snout. Reassessment of pliosauroid relationships demonstrates that Thalassiodracon hawkinsii, Hauffiosaurus spp. and Attenborosaurus conybeari are successively more derived basal representatives of Pliosauridae. This indicates that a longirostrine snout was acquired among Early Jurassic pliosaurids (such as Hauffiosaurus), but that these early taxa had a small body size, long neck, and gracile skull and limb bones relative to the contemporaneous rhomaleosaurids, which were robust, large-bodied macropredators. The pliosaurid lineage only acquired its ‘characteristic’ large size, robust skull and short neck in the late Middle Jurassic after the extinction of Rhomaleosauridae

Data from: A new pliosaurid (Sauropterygia, Plesiosauria) from the Oxford Clay Formation (Middle Jurassic, Callovian) of England: evidence for a gracile, longirostrine grade of Early–Middle Jurassic pliosaurids

A partial skeleton from the Sigiloceras enodatum ammonite Subzone (early Callovian, Middle Jurassic) of the Oxford Clay Formation of Quest Pit, near Stewartby, Bedfordshire, UK respresents one of the basalmost Middle Jurassic pliosaurids. Marmornectes candrewi gen. et sp. nov. possesses seven autapomorphies and a longirostrine snout. Reassessment of pliosauroid relationships demonstrates that Thalassiodracon hawkinsii, Hauffiosaurus spp. and Attenborosaurus conybeari are successively more derived basal representatives of Pliosauridae. This indicates that a longirostrine snout was acquired among Early Jurassic pliosaurids (such as Hauffiosaurus), but that these early taxa had a small body size, long neck, and gracile skull and limb bones relative to the contemporaneous rhomaleosaurids, which were robust, large-bodied macropredators. The pliosaurid lineage only acquired its ‘characteristic’ large size, robust skull and short neck in the late Middle Jurassic after the extinction of Rhomaleosauridae

Global interrelationships of Plesiosauria (Reptilia, Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses

Previous attempts to resolve plesiosaurian phylogeny are reviewed and a new phylogenetic data set of 66 taxa (67% of ingroup taxa examined directly) and 178 characters (eight new) is presented. We recover two key novel results: a monophyletic Plesiosauridae comprising Plesiosaurus dolichodeirus, Hydrorion brachypterygius, Microcleidus homalospondylus, Occitanosaurus tournemirensis and Seeleyosaurus guilelmiimperatoris; and five plesiosaurian taxa recovered outside the split between Plesiosauroidea and Pliosauroidea. These taxa are Attenborosaurus conybeari, ‘Plesiosaurus’ macrocephalus and a clade comprising Archaeonectrus rostratus, Macroplata tenuiceps and BMNH 49202. Based on this result, a new name, Neoplesiosauria, is erected for the clade comprising Plesiosauroidea and Pliosauroidea. Taxon subsamples of the new dataset are used to simulate previous investigations of global plesiosaurian relationships. Based on these simulations, most major differences between previous global phylogenetic hypotheses can be attributed to differences in taxon sampling. These include the position of Leptocleididae and Polycotylidae and the monophyly or paraphyly of Rhomaleosauridae. On this basis we favour the results recovered by our, larger analysis. Leptocleididae and Polycotylidae are sister taxa, forming a monophyletic clade within Plesiosauroidea, indicating that the large-headed, short-necked ‘pliosauromorph’ body plan evolved twice within Plesiosauria. Rhomaleosauridae forms the monophyletic sister taxon of Pliosauridae within Pliosauroidea. Problems are identified with previous phylogenetic definitions of plesiosaurian clades and new, stem-based definitions are presented that should maintain their integrity over a range of phylogenetic hypotheses. New, rank-free clade names Cryptoclidia and Leptocleidia are erected to replace the superfamilies Cryptoclidoidea and Leptocleidoidea. These were problematic as they were nested within the superfamily Plesiosauroidea. The incongruence length difference test indicates no significant difference in levels of homoplasy between cranial and postcranial characters

The cranial anatomy and taxonomy of Peloneustes philarchus (Sauropterygia, Pliosauridae) from the Peterborough Member (Callovian, Middle Jurassic) of the United Kingdom

Peloneustes philarchus is the most abundant pliosaurid from the Peterborough Member of the Oxford Clay Formation (Callovian) of the UK. It is a valid taxon possessing a unique character combination, including a single autapomorphy: the interdentary symphysis is raised dorsally on a narrow platform. Twenty-one specimens can be positively referred to P. philarchus. However, other specimens previously referred to Peloneustes, from the Peterborough Member near Peterborough, and the lower Oxfordian strata of Marquise, northern France, represent distinct, unnamed taxa. The skull of P. philarchus is described in detail, including new information from an uncrushed, three-dimensionally preserved specimen and a specimen with a well-preserved palate. Well-preserved material clearly indicates that P. philarchus lacked nasals, but possessed a lacrimal. A previously unrecognised ‘palpebral’ forms part of the dorsal orbit margin adjacent to the prefrontal. The number of maxillary (30–31) and dentary (36–44) alveoli, the number of dentary alveoli adjacent to the mandibular symphysis (13–15), the number of foramina on the frontal and jugal, the breadth of the parasphenoid, breadth of the anterior (narial) process of the palatine and the presence of a lappet of the angular that extends onto the posterodorsal surface of the retroarticular process vary among individuals but are not considered sufficient to justify the recognition of new taxa. The presence of an open palpebral–prefrontal suture, the size of the ventral midline tubercle of the basioccipital and the presence of an anterior interpterygoid vacuity seem to vary with body size

A new pliosaurid (Sauropterygia, Plesiosauria) from the Oxford Clay Formation (Middle Jurassic, Callovian) of England: evidence for a gracile, longirostrine grade of Early-Middle Jurassic pliosaurids

A partial skeleton from the Sigiloceras enodatum ammonite Subzone (lower Callovian, Middle Jurassic) of the Oxford Clay Formation of Quest Pit, near Stewartby, Bedfordshire, UK, represents one of the basalmost Middle Jurassic pliosaurids. Marmornectes candrewi gen. et sp. nov. possesses seven autapomorphies and a longirostrine snout. Reassessment of pliosauroid relationships demonstrates that Thalassiodracon hawkinsii, Hauffiosaurus spp. and Attenborosaurus conybeari are successively more derived basal representatives of Pliosauridae. This indicates that a longirostrine snout was acquired among Early Jurassic pliosaurids (such as Hauffiosaurus) but that these early taxa had a small body size, long neck, and gracile skull and limb bones relative to the contemporaneous rhomaleosaurids, which were robust, large-bodied macropredators. The pliosaurid lineage only acquired its ‘characteristic’ large size, robust skull and short neck in the late Middle Jurassic after the extinction of Rhomaleosauridae

A new leptocleidid (Sauropterygia, Plesiosauria) from the Vectis Formation (Early Barremian–early Aptian; Early Cretaceous) of the Isle of Wight and the evolution of Leptocleididae, a controversial clade

Among the most enigmatic and controversial plesiosaurian clades is the Early Cretaceous Leptocleididae, a small group of (mostly) short-necked taxa with ‘intermediate' or ‘pliosauromorph' body proportions. Leptocleidids have often been interpreted as basal members of Pliosauroidea, and their presence in marginal marine and even freshwater facies has led to suggestions that they might represent late-surviving relicts, perhaps related to the Lower Jurassic rhomaleosaurids. We describe a new leptocleidid, Vectocleidus pastorum gen. et sp. nov., from the late Barremian part of the Cowleaze Chine Member (Vectis Formation), Isle of Wight, UK, and undertake a detailed reassessment of leptocleidid anatomy and relationships. New data on the long-necked Brancasaurus gives extra support to a monophyletic Leptocleididae with taxa of ‘intermediate' body plan and robust skulls, Leptocleidus superstes, Leptocleidus capensis, Nichollssaura and Vectocleidus. Thus, leptocleidids adopted a range of body plans on the pliosauromorph–plesiosauromorph spectrum. Support for a placement of Leptocleididae within Pliosauroidea is weak, and most proposed synapomorphies fail the test of similarity. However, numerous synapomorphies, including many new observations, support a derived position within Plesiosauroidea. Thus, the ‘intermediate' body plan of many leptocleidids is not plesiomorphic, and plesiosaurian body plan evolution was complex and highly plastic. We also summarize the anatomy of ‘Cimoliasaurus' valdensis, a short-necked Early Cretaceous taxon. ‘C.' valdensis is a valid taxon for which we erect the new monotypic genus Hastanectes. Hastanectes shows evidence of possible pliosaurid affinities. Thus, multiple plesiosaurian clades may have invaded margin and freshwater environments

ESM2 Full Phylogenetic dataset

Nexus file with most parsimonious trees include

ESM3: Taxon ages for full phylogenetic dataset

Numerical; per stage or substag