New information on Bobosaurus forojuliensis (Reptilia: Sauropterygia): implications for plesiosaurian evolution

Preparation of the holotype specimen of Bobosaurus forojuliensis, a large sauropterygian from the lower Carnian of northeastern Italy, revealed new morphological data relevant in establishing its phylogenetic affinities among pistosauroid taxa and its relationships with plesiosaurians. Inclusion of B. forojuliensis in two phylogenetic analyses focusing, respectively, on sauropterygians and pistosauroids agreed in placing the Italian taxon as closer to plesiosaurians than to other pistosauroids. The phylogenetic interpretation of Bobosaurus was not biased by assumptions on character weighting, is consistent with its relatively younger age compared to most pistosauroids, extends the fossil record of the plesiosaurian basal lineage back to the Carnian and supports the earliest diversification of the clade during the Late Triassic in agreement with the record of several distinct lineages of rhomaleosaurids, plesiosauroids and pliosauroids in the lowermost Jurassic. Bobosaurus shows that the evolution of the plesiosaurian body plan from the ancestral pistosauroid grade was a step-wise process, and that some of the vertebral and appendicular specialisations of Jurassic and Cretaceous plesiosaurians had already developed in the earliest Late Triassic.     

The internal cranial anatomy of the Plesiosauria (Reptilia, Sauropterygia): evidence for a functional secondary palate

In the late 19th Century, the choanae (or internal nares) of the Plesiosauria were identified as a pair of palatal openings located rostral to the external nares, implying a rostrally directed respiratory duct and air path inside the rostrum. Despite obvious functional shortcomings, this idea was firmly established in the scientific literature by the first decade of the 20th Century. The functional consequences of this morphology were only re-examined by the end of the 20th Century, leading to the conclusion that the choanae were not involved in respiration but instead in underwater olfaction, the animals supposedly breathing with the mouth agape. Re-evaluation of the palatal and internal cranial anatomy of the Plesiosauria reveals that the traditional identification of the choanae as a pair of fenestrae situated rostral to the external nares appears erroneous. These openings more likely represent the bony apertures of ducts that lead to internal salt glands situated inside the maxillary rostrum. The 'real' functional choanae (or caudal interpterygoid vacuities), are situated at the caudal end of the bony palate between the sub-temporal fossae, as was suggested in the mid-19th Century. The existence of a functional secondary palate in the Plesiosauria is therefore strongly supported, and the anatomical, physiological, and evolutionary implications of such a structure are discussed.     

A new genus of pliosaur (Reptilia: Sauropterygia) from the Lower Jurassic of Holzmaden,Germany

Abstract: The exquisitely preserved holotype of the pliosaur ‘Rhomaleosaurus’victor (SMNS 12478) is described from the Toarcian Posidonien‐Schiefer (Upper Lias, Lower Jurassic) of Holzmaden (Baden‐Württemberg), Germany. The specimen presents a novel combination of synapomorphies and unique morphometric proportions separating it from Rhomaleosaurus sensu stricto and warranting the erection of a new genus, Meyerasaurus gen. nov. Historically, the name ‘Thaumatosaurus’ has been interchangeable with Rhomaleosaurus and is frequently associated with SMNS 12478 in the literature. However, this is an invalid taxon and cannot be reinstated. The anatomy of Meyerasaurus victor is compared in detail with other pliosaurs, and its taxonomic affinity is reviewed. M. victor belongs to the family Rhomaleosauridae and shares several anatomical characters with Rhomaleosaurus including a short and robust premaxillary rostrum (length‐to‐width ratio c. 1.0), parallel premaxilla–maxilla sutures anterior to the nares, vomers contacting the maxillae posterior to the internal nares, and c. 28 cervical vertebrae minus the atlas–axis. The known geographical distribution of Rhomaleosaurus, which previously extended across the German and English palaeobiogeographical zones, is reduced to the English zone as a consequence of the referral of SMNS 12478 to a new genus. This is significant because it contributes to an ongoing trend of increasing generic separation between the German and English zones, while increasing the generic diversity within the German zone itself.     

A new specimen of Yunguisaurus (Reptilia; Sauropterygia) from the Ladinian (Middle Triassic) Zhuganpo Member,Falang Formation,Guizhou, China and the restudy of Dingxiaosaurus

A new articulated juvenile specimen of Yunguisaurus liae (GMPKU-P-1528) from the Ladinian (Middle Triassic) Zhuganpo Member, Falang Formation, Xingyi, Guizhou, China presents new morphological information on the relatively complete atlas and axis. The comparison of the all four Y. liae skeletons with the holotype (DMK 8) of Dingxiaosaurus luyinensis reveals that there is no morphological difference between the two species. The similarities include the similar orientation of the tarsals, the same value of the ratio of the distal width to the proximal width of the tibia and the ratio of the fibula length to the tibia length. The taxonomy of D. luyinensis, however, remains controversial. It cannot be distinguished from other pistosauroid taxa due to the lack of comparative portions of the skeletons preserved (e.g., Chinchenia, Kwangsissaurus). It may turn out that D. luyinensis and Y. liae stand for the same species when more complete materials become available.     

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.     

Cranial anatomy,taxonomic implications and palaeopathology of an Upper Jurassic Pliosaur (Reptilia: Sauropterygia) from Westbury,Wiltshire, UK

Abstract: Complete skulls of giant marine reptiles of the Late Jurassic are rare, and so the discovery of the 1.8‐m‐long skull of a pliosaur from the Kimmeridge Clay Formation (Kimmeridgian) of Westbury, Wiltshire, UK, is an important find. The specimen shows most of the cranial and mandibular anatomy, as well as a series of pathological conditions. It was previously referred to Pliosaurus brachyspondylus, but it can be referred reliably only to the genus Pliosaurus, because species within the genus are currently in need of review. The new specimen, together with another from the same locality, also referred to P. brachyspondylus, will be crucial in that systematic revision, and it is likely that the genus Pliosaurus will be found to include several genera. The two Westbury Pliosaurus specimens share many features, including the form of the teeth, but marked differences in the snout and parietal crest suggest sexual dimorphism; the present specimen is probably female. The large size of the animal, the extent of sutural fusion and the pathologies suggest this is an ageing individual. An erosive arthrotic condition of the articular glenoids led to prolonged jaw misalignment, generating a suite of associated bone and dental pathologies. Further damage to the jaw joint may have been the cause of death.     

Taxonomy of the proterosuchid archosauriforms (Diapsida: Archosauromorpha) from the earliest Triassic of South Africa,and implications for the early archosauriform radiation

Proterosuchidae is one of the first clades of Archosauriformes (archosaurs and closely related species) to appear in the fossil record, with the richest sample of the group coming from the Lystrosaurus Assemblage Zone (earliest Triassic) of South Africa. Four nominal proterosuchid species were described from South Africa during the twentieth century (Proterosuchus fergusi, Chasmatosaurus vanhoepeni, Chasmatosaurus alexanderi and Elaphrosuchus rubidgei), but interpretations of their taxonomy have been widely disparate. The most recent taxonomic revision concluded that P. fergusi is the only valid species and that the other nominal species are junior subjective synonyms of this taxon. This proposal was based on the interpretation that anatomical differences between the nominal species could be explained as a result of ontogenetic changes and/or post‐mortem deformation. The recent discoveries of multiple new South African proterosuchid specimens provide an impetus to revisit their taxonomy. Based upon a comprehensive re‐examination of all known specimens, as well as examination of other proterosuchid taxa in collections worldwide, we conclude that the holotype of Proterosuchus fergusi is undiagnostic. As a result, we propose a neotype (RC 846) for the species. ‘Chasmatosaurus vanhoepeni’ and ‘Elaphrosuchus rubidgei’ are considered subjective junior synonyms of P. fergusi. ‘Chasmatosaurus’ alexanderi is considered a valid species, for which we propose the new combination P. alexanderi comb. nov. A third species, P. goweri sp. nov., is erected on the basis of a single specimen (NMQR 880). All three species recognized here are taxonomically distinct from a previously described archosauriform maxilla from the lower Lystrosaurus AZ. As a result, we recognize a minimum of four archosauriform species following the Permo‐Triassic mass extinction in South Africa. Our results suggest a greater species richness of earliest Triassic archosauriforms than previously appreciated, but that archosauriform morphological disparity remained low and did not expand until the late Early Triassic – early Mid‐Triassic.     

A new gomphodont cynodont (Traversodontidae) from the Middle–Late Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence,Brazil

The tetrapod faunas from the terrestrial Middle–Late Triassic basins of Africa and South America are among the richest in the world, especially in non‐mammalian cynodonts. Despite the great abundance of cynodont specimens found in these basins, there are few known taxa that exhibit interbasinal distributions. Here we describe a new species of traversodontid cynodont of the genus Scalenodon from the Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence, from the state of Rio Grande do Sul, Brazil. Scalenodon ribeiroae sp. nov. is based on a partial skull that possesses a combination of features not observed in any other South American traversodontid: ellipsoid upper postcanines with the transverse crest formed by three cusps, lacking a mesiobuccal accessory cusp, and with lingual cusp projected lingually creating a concave lingual surface on the upper postcanines; the paracanine fossa is positioned medially to the upper canine, and jugal lacks a suborbital process. A phylogenetic analysis places the new taxon in a basal position within the Family Traversodontidae, with the African Scalenodon angustifrons as sister‐taxon. The new specimen of Scalenodon represents the first record of this genus outside of the Manda Beds of Tanzania, and reinforces the biostratigraphical and biogeographical connection between Gondwanan Middle–Late Triassic tetrapod faunas. Although recent advances have been made, our current knowledge of these faunas is limited by the lack of absolute dates for most units and by uncertainties in the taxonomy and stratigraphical provenance of key fossils.     

云南罗平中三叠世利齿滇东龙(鳍龙超目)—新材料

记述了产自云南省罗平县中三叠世安尼期关岭组上段一件近乎完整的利齿滇东龙(Diandongosaurus acutidentatus)新标本。根据该标本所提供的新信息,讨论了部分前人研究中不确定或者存在争议的骨骼特征,并重新厘定了滇东龙的鉴定特征。分支系统学分析表明滇东龙属于始鳍龙类,且与真鳍龙亚目的亲缘关系较近。     

Phylogenetics of tribe Sabiceeae (Ixoroideae,Rubiaceae) revisited,with a new subgeneric classification for Sabicea

Tribe Sabiceeae (Ixoroideae, Rubiaceae) has undergone recent taxonomical changes with the incorporation of the related genera Ecpoma, Pseudosabicea and Stipularia into the type genus Sabicea. We use phylogenetic analysis and morphological data to verify the relationships among members of the tribe, including the most comprehensive taxon sampling of the tribe to date with 74 of 145 species. Sequence data from the nuclear internal transcribed spacer (ITS) and three plastid markers (petD, rps16, trnT–F) were used to infer relationships among the members of the tribe. Individual analyses using maximum likelihood, parsimony and Bayesian approaches reveal several supported clades: the former genus Stipularia is resolved as a monophyletic unit, but Ecpoma is monophyletic only if Sabicea urbaniana and Sabicea xanthotricha are included (corresponding to Sabicea subgenus Stipulariopsis sensu Wernham). Pseudosabicea is biphyletic, with one clade corresponding to section Anisophyllae of Hallé (1964) and the other one to the other sections (Floribundae and Sphaericae) of the genus. Eleven morphological characteristics were recorded for all species studied and seven have been mapped onto the phylogenetic tree to study their evolution in the group and assess their value for the classification of Sabicea s.l. Finally, our study shows that a combination of diagnostic characteristics should be used to differentiate each group and we propose to recognise four subgenera in Sabicea.     

Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii,Saurichthyidae)

Viviparity has evolved independently at least 12 times in ray‐finned fishes. However, the fossil record of actinopterygian viviparity is poor, with only two documented occurrences. Both of these are from the non‐teleost actinopterygian Saurichthys, and include S. curionii and S. macrocephalus from the Middle Triassic Meride Limestone (Monte San Giorgio, Switzerland). Here, we present new data on the reproductive biology of these species, giving unprecedented insights into their life‐history. Based on positional and preservational criteria, six specimens were identified as unambiguously gravid. Embryos were positioned dorsal to the gastrointestinal tract, parallel to the axial skeleton and to each other, in the posterior two‐thirds of the abdominal region. A minimum of 16 embryos are preserved in the most fecund females and, based on the largest preserved embryos and smallest preserved neonates, birth must have occurred at 7–12% of maternal fork length. Embryonic crania and teeth are relatively well‐ossified, however ossification of the parietal region is delayed. In the postcranium, the median scale rows and lepidotrichia are ossified, but not the lateral scale rows. Ossified squamation and gradual allometric growth suggests that neonates did not undergo metamorphosis and were relatively precocial. When considered in a phylogenetic context, neither live birth nor internal fertilization appears to represent the primitive state for saurichthyid fishes.     

Elasmosaur (Reptilia: Sauropterygia) neck flexibility: implications for feeding strategies

Elasmosaurs were extremely long-necked, aquatic reptiles that used four flippers for locomotion. Their distinctive long neck distinguishes them from all other Mesozoic forms, yet the potential uses and constraints of this structure are poorly understood, particularly with regard to feeding. Several associated series of elasmosaurian cervical vertebrae were used to measure ranges of potential flexion. Two-dimensional models, based on a complete specimen of the Late Cretaceous elasmosaur Aphrosaurus furlongi, were created to measure mobility in both vertical and horizontal planes. Accuracy of the models was assessed through comparative analyses with currently extant vertebrate analogues (e.g. snake, turtle, seal). Results suggest that the elasmosaurian neck was capable of a 75-177 degrees ventral, 87-155 degrees dorsal, and 94-176 degrees lateral range of movement depending upon the thickness of cartilage reconstructed between each vertebra. Neck postures such as a 'swan-like' S-shape are shown to be implausible because they require >360 degrees vertical flexion. However, maintenance of a straight neck while swimming, together with considerable lateral and/or ventral movement during prey capture and feeding are feasible.     

Elasmosaur (Reptilia: Sauropterygia) neck flexibility: implications for feeding strategies.

Elasmosaurs were extremely long-necked, aquatic reptiles that used four flippers for locomotion. Their distinctive long neck distinguishes them from all other Mesozoic forms, yet the potential uses and constraints of this structure are poorly understood, particularly with regard to feeding. Several associated series of elasmosaurian cervical vertebrae were used to measure ranges of potential flexion. Two-dimensional models, based on a complete specimen of the Late Cretaceous elasmosaur Aphrosaurus furlongi, were created to measure mobility in both vertical and horizontal planes. Accuracy of the models was assessed through comparative analyses with currently extant vertebrate analogues (e.g. snake, turtle, seal). Results suggest that the elasmosaurian neck was capable of a 75-177 degrees ventral, 87-155 degrees dorsal, and 94-176 degrees lateral range of movement depending upon the thickness of cartilage reconstructed between each vertebra. Neck postures such as a 'swan-like' S-shape are shown to be implausible because they require >360 degrees vertical flexion. However, maintenance of a straight neck while swimming, together with considerable lateral and/or ventral movement during prey capture and feeding are feasible.     

Taphonomy and palaeoecology of Late Triassic (Carnian) ammonoid concentrations from the Taurus Mountains,Turkey

The deposits of the Carnian Kas?mlar Formation within the Taurus Platform Units of south‐western Turkey represent an important archive of a Late Triassic ecosystem. New palaeontological information was obtained by analysing the Kasimlarceltites mass occurrence, located within the Kas?mlar Formation and named after the Lower Carnian (Julian) ammonoid genus Kasimlarceltites. This is the dominant taxon (> 94%) within the mass occurrence: nearly 775 million ammonoids and 50 million gastropods were extrapolated for the whole extension (at least 5 km²) of the Kasimlarceltites beds. This calculation is one of the main findings within this study, as it is the first time that such a fossil mass occurrence was quantified. Additionally, orientation measurements of the planispiral ammonoids and the helical gastropods enabled reconstructing the history of the mass occurrence and interpreting the underlying transport mechanisms. Further taphonomic aspects (e.g. biofabric, preservation, bioerosion or genetic classification) as well as comparisons with samples of the same acme zone from different localities near A?a?iyaylabel (AS IV, KA I‐II) point to a two‐phased genetic history. Accordingly, local mass mortality within the Kasimlarceltites fauna due to oxygen fluctuations or methane degassing may have initially led to a primary accumulation. These deposits were then reworked and redeposited basinward by gravity flows to create the present‐day secondary allochthonous concentrations.     

A combined morphological,ultrastructural, molecular,and biochemical study of the peculiar family Gomontiellaceae (Oscillatoriales) reveals a new cylindrospermopsin‐producing clade of cyanobacteria

Members of the morphologically unusual cyanobacterial family Gomontiellaceae were studied using a polyphasic approach. Cultured strains of Hormoscilla pringsheimii, Starria zimbabweënsis, Crinalium magnum, and Crinalium epipsammum were thoroughly examined, and the type specimen of the family, Gomontiella subtubulosa, was investigated. The results of morphological observations using both light microscopy and transmission electron microscopy were consistent with previous reports and provided evidence for the unique morphological and ultrastructural traits of this family. Analysis of the 16S rRNA gene confirmed the monophyletic origin of non‐marine repre‐sentatives of genera traditionally classified into this family. The family was phylogenetically placed among other groups of filamentous cyanobacterial taxa. The presence of cellulose in the cell wall was analyzed and confirmed in all cultured Gomontiellaceae members using Fourier transform infrared spectroscopy and fluorescence microscopy. Evaluation of toxins produced by the studied strains revealed the hepatotoxin cylindrospermopsin (CYN) in available strains of the genus Hormoscilla. Production of this compound in both Hormoscilla strains was detected using high‐performance liquid chromatography in tandem with high resolution mass spectrometry and confirmed by positive PCR amplification of the cyrJ gene from the CYN biosynthetic cluster. To our knowledge, this is the first report of CYN production by soil cyanobacteria, establishing a previously unreported CYN‐producing lineage. This study indicates that cyanobacteria of the family Gomontiellaceae form a separate but coherent cluster defined by numerous intriguing morphological, ultrastructural, and biochemical features, and exhibiting a toxic potential worthy of further investigation.     

Eocene Pachynolophinae (Perissodactyla,Palaeotheriidae) from China,and their palaeobiogeographical implications

The Eocene perissodactyl family Palaeotheriidae has traditionally been considered to be a nearly endemic European group within Equoidea, but a few palaeotheres have been reported from Asia. Here, I reanalyse a maxilla containing M1–3 from the Lunan Basin, Yunnan Province, China. This element was initially assigned to a new tapiromorph species, Lophialetes yunnanensis, but is here placed in a new genus Lophiohippus within Pachynolophinae based mainly on the absence of mesostyles, the strongly oblique metalophs, the strong development of lophodonty, parastyles overlapping metastyles of preceding teeth and situated mesial to the paracone, and the fact that M3 is longer than wide and has a large and buccally deflected metastyle. Lophiohippus differs from European Anchilophus and Paranchilophus in that the parastyles are situated mesial or even slightly lingual, rather than mesiobuccal, to the paracones, and M3 is markedly relatively larger than M1. I further reanalyse Qianohippus magicus from the Shinao Basin of Guizhou Province, China, in which the complete dentition is known. Qianohippus is characterized by a molariform P2 and non‐molariform P3–4; a relatively high degree of lophodonty; the absence of mesostyles; an angular bending in the protoloph on P3‐M3 and the metaloph on M1–3 at the paraconule and metaconule, respectively; and weakly developed ‘metastylid’ on the lower cheek teeth. A cladistic analysis supports a close relationship between Lophiohippus yunnanensis and Paranchilophus, and suggests that Qianohippus is closely related to some derived pachynolophs. The appearance of the pachynolophins Lophiohippus and Qianohippus in China supports the existence of a biogeographical connection between Europe and Asia in the Middle‐Late Eocene, and the dispersal route was probably along the Tethyan microcontinents in the south.     

Increases in sampling support the southern Gondwanan hypothesis for the origin of dinosaurs

Dinosaurs were ubiquitous in terrestrial ecosystems through most of the Mesozoic and are still diversely represented in the modern fauna in the form of birds. Recent efforts to better understand the origins of the group have resulted in the discovery of many new species of early dinosaur and their closest relatives (dinosauromorphs). In addition, recent re‐examinations of early dinosaur phylogeny have highlighted uncertainties regarding the interrelationships of the main dinosaur lineages (Sauropodomorpha, Theropoda and Ornithischia), and questioned the traditional hypothesis that the group originated in South Gondwana and gradually dispersed over Pangaea. Here, we use an historical approach to examine the impact of new fossil discoveries and changing phylogenetic hypotheses on biogeographical scenarios for dinosaur origins over 20 years of research time, and analyse the results in the light of different fossil record sampling regimes. Our results consistently optimize South Gondwana as the ancestral area for Dinosauria, as well as for more inclusive clades including Dinosauromorpha, and show that this hypothesis is robust to increased taxonomic and geographic sampling and divergent phylogenetic results. Our results do not find any support for the recently proposed Laurasian origin of dinosaurs and suggest that a southern Gondwanan origin is by far the most plausible given our current knowledge of the diversity of early dinosaurs and non‐dinosaurian dinosauromorphs.