A new scanilepiform from the Lower Triassic of northern Gansu Province,China, and phylogenetic relationships of non‐teleostean Actinopterygii

A new scanilepiform, Beishanichthys brevicaudalis gen. et sp. nov. , is named and described based on fossils from the Lower Triassic lake deposits exposed in Beishan area, Gansu Province, China. The discovery documents a new record of this group, which is significantly older than other known scanilepiforms from China, and is slightly younger than Evenkia from the Lowest Triassic of Central Siberia. Although the Beishan beds were previously interpreted as Late Permian in age, based on megaplant fossils, this new discovery supports the reinterpretation of the deposits as Early Triassic in age, based on vertebrate fossils from the same locality and horizon. Phylogenetic analysis was conducted to resolve the relationships of Scanilepiformes with other actinopterygian clades, and the inter‐relationships within Scanilepiformes. Contrary to previous thought that scanilepiforms are closely related to the Amiidae, the phylogenetic results of this study recognize the Scanilepiformes as stem‐group neopterygians. Relationships of the Scanilepiformes and Australosomus with other neopterygians remain unresolved. With a characteristic long‐based dorsal fin, scanilepiforms represent a small group that emerged in Early Triassic freshwater environments, inhabited Eurasia and North America during the Middle–Late Triassic, briefly invaded the marine environment by the Late Triassic in Europe, and became extinct at the end of Triassic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 595–612.     

Redescription of the phytosaurs Paleorhinus (‘Francosuchus’) angustifrons and Ebrachosuchus neukami from Germany,with implications for Late Triassic biochronology

Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile‐like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long‐range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well‐preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy‐based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London     

The braincase of a specimen of <Emphasis Type="Italic">Proterochampsa</Emphasis> Reig (Archosauriformes: Proterochampsidae) from the Late Triassic of Argentina

The proterochampsids are a Triassic group of superficially crocodile-like forms belonging to the Archosauriformes. In the present contribution, we present new information regarding the braincase of the proterochampsid Proterochampsa Reig 1959, from the Ischigualasto Formation (Carnian) of Argentina, and discuss its phylogenetic considerations. Some unique neurocranial features of Proterochampsa are described, including: the prominence and thickness of the V-shaped ridge that surrounds the basisphenoidal fossa; the medially concave lateral arms of the same ridge; and the semilunar depression on the parabasisphenoid ventrolaterally exposed. Other features are only shared with likely unrelated archosauriforms, including: the great lateral development of the basipterygoid processes and caudal development of its distal end; an eight-shaped metotic foramen; laterally directed basipterygoid processes; and rostral boundary of the basisphenoidal recess V-shaped. Proterochampsa differs in many other aspects from the archosauriform Chanaresuchus, including: a proportionally shorter basioccipital basal tubera; cultriform process ovoid in cross-section; longitudinal sulcus dorsal to the basipterygoid process; deep basisphenoidal recess; and the absence of a prominent intertuberal plate. In many braincase features, Proterochampsa is more similar to archosaurs than to Euparkeria, erythrosuchids and Proterosuchus. They include a reduced semilunar depression. A ventral border of the basioccipital forming a wide convexity and a dorsoventrally thin paroccipital process likely represents a feature shared with Chanaresuchus, but not with Doswellia and other basal archosauriforms.     

Osteology and phylogenetic relationships of Tehuelchesaurus benitezii (Dinosauria,Sauropoda) from the Upper Jurassic of Patagonia

The diversification and early evolution of neosauropod dinosaurs is mainly recorded from the Upper Jurassic of North America, Europe, and Africa. Our understanding of this evolutionary stage is far from complete, especially in the Southern Hemisphere. A partial skeleton of a large sauropod from the Upper Jurassic Cañadón Calcáreo Formation of Patagonia was originally described as a ‘cetiosaurid’ under the name Tehuelchesaurus benitezii. The specimen is here redescribed in detail and the evidence presented indicates that this taxon is indeed a neosauropod, thus representing one of the oldest records of this clade in South America. A complete preparation of the type specimen and detailed analysis of its osteology revealed a great number of features of phylogenetic significance, such as fully opisthocoelous dorsal vertebrae, the persistence of true pleurocoels up to the first sacral vertebra, associated with large camerae in the centrum and supraneural camerae, and an elaborate neural arch lamination, including two apomorphic laminae in the infradiapophyseal fossa. The phylogenetic relationships of this taxon are tested through an extensive cladistic analysis that recovers Tehuelchesaurus as a non‐titanosauriform camarasauromorph, deeply nested within Neosauropoda. Camarasauromorph sauropods were widely distributed in the Late Jurassic, indicating a rapid evolution and diversification of the group. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 605–662.     

Two new species of Saurichthys (Actinopterygii: Saurichthyidae) from the Middle Triassic of Monte San Giorgio,Switzerland, with implications for character evolution in the genus

Saurichthys, characterized by a long slender body and an elongated rostrum, is one of the most iconic genera of Late Paleozoic–Early Mesozoic fishes. The genus was particularly speciose in the Triassic, with a global distribution in both marine and freshwater habitats. Here, we describe two new species from the Middle Triassic Besano Formation of Monte San Giorgio, Switzerland, Saurichthys breviabdominalis sp. nov. and Saurichthys rieppeli sp. nov. S. breviabdominalis is characterized by a proportionately long operculum, short abdominal region and rib‐like mid‐lateral scales, whereas S. rieppeli is divergent from other Middle Triassic saurichthyids in the block‐like haemal arches, fringing fulcra on the pelvic and unpaired fins, and reduction of the squamation to a single row in the abdominal region. Phylogenetic analysis places S. rieppeli in a basal position relative to congeners from the Alpine Triassic, and supports previous hypotheses regarding the convergent evolution of reduced squamation within saurichthyids. S. breviabdominalis forms a monophyletic group with species from the same locality, suggesting divergence in sympatry. This finding has implications for our understanding of disparity and character evolution in saurichthyid fishes, as well as ecomorphological divergence and resource partitioning between closely related fishes in Triassic marine ecosystems. © 2015 The Linnean Society of London     

A new species of the enigmatic archosauromorph Doswellia from the Upper Triassic Bluewater Creek Formation,New Mexico,USA

Abstract: Doswellia sixmilensis is a new species of the doswelliid archosauromorph genus Doswellia named for an incomplete skeleton from the Upper Triassic Bluewater Creek Formation of the Chinle Group in west‐central New Mexico, USA. D. sixmilensis differs from D. kaltenbachi Weems, the type and only other known species of Doswellia, in its larger size, higher tooth count and greater heterodonty, possession of keels on the cervical centra and the presence of discrete knobs or spikes on some osteoderms. The holotype of D. sixmilensis is the fourth occurrence of Doswellia and only the second occurrence of a Doswellia skull, which includes the previously unknown premaxilla and maxilla (and therefore the best dentition) and has the best‐preserved cervical vertebrae. Although it adds to our knowledge of the anatomy of Doswellia, this new information does not alter previous concepts of the phylogenetic relationships of the doswelliid genera, largely because they are so poorly known anatomically. The genus Doswellia is known from the Newark Supergroup in Virginia, and the Chinle Group in Texas, New Mexico and Utah, in strata of Otischalkian–Adamanian age. The type locality of D. sixmilensis is c. 43 m stratigraphically below a bed from which U‐Pb dating of detrital zircons yields a maximum depositional age of c. 220 Ma, so this is a reasonable approximate numerical age for D. sixmilensis.     

Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms

Titanosauriforms represent a diverse and globally distributed clade of neosauropod dinosaurs, but their inter‐relationships remain poorly understood. Here we redescribe Lusotitan atalaiensis from the Late Jurassic Lourinhã Formation of Portugal, a taxon previously referred to Brachiosaurus. The lectotype includes cervical, dorsal, and caudal vertebrae, and elements from the forelimb, hindlimb, and pelvic girdle. Lusotitan is a valid taxon and can be diagnosed by six autapomorphies, including the presence of elongate postzygapophyses that project well beyond the posterior margin of the neural arch in anterior‐to‐middle caudal vertebrae. A new phylogenetic analysis, focused on elucidating the evolutionary relationships of basal titanosauriforms, is presented, comprising 63 taxa scored for 279 characters. Many of these characters are heavily revised or novel to our study, and a number of ingroup taxa have never previously been incorporated into a phylogenetic analysis. We treated quantitative characters as discrete and continuous data in two parallel analyses, and explored the effect of implied weighting. Although we recovered monophyletic brachiosaurid and somphospondylan sister clades within Titanosauriformes, their compositions were affected by alternative treatments of quantitative data and, especially, by the weighting of such data. This suggests that the treatment of quantitative data is important and the wrong decisions might lead to incorrect tree topologies. In particular, the diversity of Titanosauria was greatly increased by the use of implied weights. Our results support the generic separation of the contemporaneous taxa Brachiosaurus, Giraffatitan, and Lusotitan, with the latter recovered as either a brachiosaurid or the sister taxon to Titanosauriformes. Although Janenschia was recovered as a basal macronarian, outside Titanosauria, the sympatric Australodocus provides body fossil evidence for the pre‐Cretaceous origin of titanosaurs. We recovered evidence for a sauropod with close affinities to the Chinese taxon Mamenchisaurus in the Late Jurassic Tendaguru beds of Africa, and present new information demonstrating the wider distribution of caudal pneumaticity within Titanosauria. The earliest known titanosauriform body fossils are from the late Oxfordian (Late Jurassic), although trackway evidence indicates a Middle Jurassic origin. Diversity increased throughout the Late Jurassic, and titanosauriforms did not undergo a severe extinction across the Jurassic/Cretaceous boundary, in contrast to diplodocids and non‐neosauropods. Titanosauriform diversity increased in the Barremian and Aptian–Albian as a result of radiations of derived somphospondylans and lithostrotians, respectively, but there was a severe drop (up to 40%) in species numbers at, or near, the Albian/Cenomanian boundary, representing a faunal turnover whereby basal titanosauriforms were replaced by derived titanosaurs, although this transition occurred in a spatiotemporally staggered fashion. © 2013 The Linnean Society of London     

Phylogenetic analysis of the leafhopper genus Apogonalia (Insecta: Hemiptera: Cicadellidae) and comments on the biogeography of the Caribbean islands

A phylogenetic analysis of the leafhopper genus Apogonalia was conducted based on a matrix of 40 terminal taxa and 147 morphological characters. The analysis yielded 1391 equally most‐parsimonious trees, which do not support the monophyly of Apogonalia in the strict consensus. A successive weighting procedure yielded 62 trees in which the genus appeared as a monophyletic group. The strict consensus of these 62 trees is almost entirely dichotomous, showing only two polytomies. The test of phylogenetic integrity was applied for distinct variations of three species: A. germana, A. sanguinipes, and A. histrio. Only for the first species was the conjecture that its variations belong to the same entity corroborated. The best‐supported clade within Apogonalia, which has several synapomorphies and high branch support indices, comprises nine Antillean endemic species. This distributional pattern probably was originated by vicariance in the Late Cretaceous, when the Proto‐Antillean archipelago was pushed north‐eastward by the Caribbean Plate to become the modern Greater Antilles. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 548–570.     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

A new species of basal rhynchosaur (Diapsida: Archosauromorpha) from the early Middle Triassic of South Africa,and the early evolution of Rhynchosauria

Rhynchosauria was an important clade of herbivorous archosauromorph reptiles during the Triassic, with a worldwide distribution. We describe a new genus and species of early rhynchosaur, E ohyosaurus wolvaardti gen. et sp. nov. , from the early Middle Triassic (early Anisian) Cynognathus Assemblage Zone (Subzone B) of the Karoo Supergroup, South Africa. Eohyosaurus wolvaardti is known from a single skull, and is recovered as the sister taxon of Rhynchosauridae in a new phylogenetic analysis. Cynognathus Subzone B has previously yielded the stratigraphically oldest well‐understood rhynchosaur species, Mesosuchus browni and Howesia browni. Eohyosaurus wolvaardti increases the rhynchosaur diversity within this stratigraphical horizon to three species. Intriguingly, all currently confirmed rhynchosaur occurrences from the Early Triassic to earliest Middle Triassic are from South Africa. This may suggest a relatively restricted palaeogeographical distribution for early rhynchosaurs, followed by a global dispersal of rhynchosaurids during the Middle Triassic. © 2015 The Linnean Society of London     

Cranial anatomy of the stem salamander Kokartus honorarius (Amphibia: Caudata) from the Middle Jurassic of Kyrgyzstan

The cranial anatomy of the stem salamander Kokartus honorarius, from the Middle Jurassic of Kyrgyzstan, is redescribed in detail on the basis of all available material using conventional methods and high‐resolution computed microtomography. In contrast to previous interpretations, the skull is characterized by posteriorly positioned external nostrils, the presence of an internarial fenestra between the premaxillae, the maxillae are not shortened, the overlapping of the frontal by the nasal and the parietal by the frontal, a dentate vomer, a dentate pterygoid with a long, strongly arcuate and anteromedially orientated anterior process, the absence of a fontanelle between frontals and pineal pit on the ventral surface of the frontals, and an inferred unique attachment of the adductor mandibulae internus to the lateral border and ventral ridge of the frontals and parietals. Kokartus is closely related to Karaurus and both these taxa constitute the monophyletic taxon Karauridae, which forms the sister group of crown‐group salamanders (Urodela). Karaurids were neotenic forms and neoteny was probably the ancestral life history strategy of salamanders. The morphological analysis of Kokartus supports the current phylogenetic hypothesis that Permian amphibamids (including branchiosaurids) are the closest relatives of salamanders amongst known non‐lissamphibian temnospondyls. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 816–838.     

Molecular systematics of flyingfishes (Teleostei: Exocoetidae): evolution in the epipelagic zone

The flyingfish family Exocoetidae is a diverse group of marine fishes that are widespread and abundant in tropical and subtropical seas. Flyingfishes are epipelagic specialists that are easily distinguished by their enlarged fins, which are used for gliding leaps over the surface of the water. Although phylogenetic hypotheses have been proposed for flyingfish genera based on morphology, no comprehensive molecular studies have been performed. In the present study, we describe a species‐level molecular phylogeny for the family Exocoetidae, based on data from the mitochondrial cytochrome b gene (1137 bp) and the nuclear RAG2 gene (882 bp). We find strong support for previous morphology‐based phylogenetic hypotheses, as well as the monophyly of most currently accepted flyingfish genera. However, the most diverse genus Cheilopogon is not monophyletic. Using our novel flyingfish topology, we examine previously proposed hypotheses for the origin and evolution of gliding. The results support the progressive transition from two‐wing to four‐wing gliding. We also use phylogenetic approaches to test the macroecological effects of two life history characters (e.g. egg buoyancy and habitat) on species range size in flyingfishes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 161–174.     

Umbonal musculature and relationships of the Late Triassic filibranch unionoid bivalves

Exceptionally well‐preserved Late Triassic unionoids from Silesia, Poland, show prominently ornamented juvenile shells and umbonal muscle attachments that are similar to Margaritifera, which are anatomically the least derived among extant unionoids. Their phosphatized (originally chitinous and impregnated with calcium phosphate) gill supports lacked transverse connections, and occasionally some of them were separated from others, being thus at the filibranch grade, like their trigonioid ancestors. Several separate small foot elevator attachments in these unionoids indicate Trigonodidae adaptation to marine or brackish water, in which the original trigonioid strong single attachment was already split into two in the Early Triassic. The ribbing of juvenile shells suggests a change to deeper infaunal life for juvenile stages, and generally less efficient near‐surface locomotion, with a wedge‐like foot, in adults. Much later the unionoids became eulamellibranchial, which promoted the brooding of the fish that their larvae parasitize. To accomodate the classification of the order under this scenario of evolutionary changes, a new suborder Silesunionina is proposed for its filibranch members. It includes the Silesunionidae fam. nov. , with the location of umbonal muscles similar to that in the extant underived unionoids, and the Unionellidae fam. nov. , with umbonal muscles attached to the external wall of the umbonal cavity. The early Late Triassic (Carnian) Silesunio parvus gen. et sp. nov. and latest Triassic (Rhaetian) Tihkia(?) silesiaca sp. nov. are proposed. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 863–883.     

Origin and relationships of the Ictidosauria to non-mammalian cynodonts and mammals

Abstract

Ictidosaurian genera are allocated to two families, Tritheledontidae and Therioherpetidae. This paper provides a diagnosis for Ictidosauria. The previously named family Brasilodontidae is shown to be a junior synonym of a family, Therioherpetidae. It is concluded that Ictidosauria originated from Late Permian procynosuchid non-mammalian cynodonts rather than from Middle Triassic probainognathid non-mammalian cynodonts. The structure of the skull and jaws of a derived traversodontid Ischignathus sudamericanus from the early Late Triassic of Argentina supports an earlier view that tritylodontids are more closely related to traversodontid than probainognathid non-mammalian cynodonts. Tritylodontids should not be included in Ictidosauria, nor should they considered to be a sister group to mammaliaforms.     

A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods

Megalosaurus bucklandii (Dinosauria: Theropoda), the oldest named dinosaur taxon, from the Bathonian (Middle Jurassic) of England, is a valid taxon diagnosed by a unique character combination of the lectotype dentary. Abundant referred material is described and several autapomorphies are identified: ventral surfaces of first and third to fifth sacral centra evenly rounded, ventral surface of second sacral centrum bearing longitudinal, angular ridge; dorsally directed flange around midheight on the scapular blade; an array of posterodorsally inclined grooves on the lateral surface of the median iliac ridge; anteroposteriorly thick ischial apron with an almost flat medial surface; and complementary groove and ridge structures on the articular surfaces between metatarsals II and III. A new phylogenetic analysis focuses on basal tetanurans and includes 41 taxa, six of which have never been included in a cladistic analysis, and 213 characters, 29 of which are new. This is the first phylogenetic analysis to focus on basal tetanuran relationships, and it reveals several new results. Megalosauroidea (= Spinosauroidea) includes two clades, basal to the traditional content of Megalosauridae + Spinosauridae. These comprise Xuanhanosaurus, Marshosaurus, Condorraptor + Piatnitzkysaurus and Chuandongocoelurus + Monolophosaurus. Almost all large‐bodied Middle Jurassic theropods are megalosauroids, but Poekilopleuron is an allosauroid. Megalosauroids show geographical differentiation among clades, indicating the development of endemic theropod faunas across Pangaea during the Middle Jurassic. Notably, megalosaurids are not known from outside of Europe during this epoch. Megalosauroids are less diverse and abundant during the Late Jurassic, when most theropods are neotetanurans and allosauroids dominate the large‐bodied predator niche. This indicates faunal turnover between the Middle and Late Jurassic. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 882–935.     

An investigation into the cladistic relationships and monophyly of therocephalian therapsids (Amniota: Synapsida)

A comprehensive phylogenetic investigation was performed to elucidate the cladistic relationships and possible monophyly of therocephalian therapsids (Amniota: Synapsida). The phylogenetic positions of 30 therapsid taxa were examined under maximum parsimony, including 23 therocephalian genera. The analysis incorporated 110 cranial and postcranial characters in order to assess the interrelationships of basal therocephalians and eutherocephalians and their relationships to Cynodontia, representing the most complete review of therocephalian phylogeny to date. The analysis supports the hypothesis that Therocephalia represents the monophyletic sister taxon to Cynodontia, with as many as 15 morphological synapomorphies, in contrast with other recent analyses of lesser taxon sampling. The results also support the hypothesis that Scylacosauridae is more closely related to Eutherocephalia than to the basal therocephalian family Lycosuchidae, supporting a ‘Scylacosauria’ clade. The taxa suggested here to be neotenic forms (e.g. Ictidosuchoides and Ictidosuchops) are positioned near the base of a monophyletic Baurioidea. Neotenic development of the therocephalian feeding apparatus and evolutionary parallelism with cynodonts are suggested to have been important trends in the early evolution of baurioid therocephalians into the Late Permian and Early Triassic.