Taxonomy and paleobiogeographic implication of Glabrobournonia Morris and Skelton (Hippuritida,Radiolitidae) from the Late Cretaceous Yigeziya Formation,southwestern Tarim Basin

A re-examination of the specimens that were identified as Biradiolites minor Pojarkova from the late Campanian to early Maastrichtian middle member of the Yigeziya Formation of southwestern Tarim Basin reveals that they should be assigned to the genus Glabrobournonia Morris and Skelton. Glabrobournonia is a genus of radiolitids characterized by indented radial bands, salient ridges on the shell margins and absence of fine ribs on the surface of the right valve. Apart from southwestern Tarim Basin, Glabrobournonia minor (Pojarkova) has also been recorded from the late Campanian of Fergana and Alai basins. The central Asian, late Campanian to early Maastrichtian G. minor differs from the late Campanian to Maastrichtian, eastern Arabian type species Glabrobournonia arabica Morris and Skelton in the flat left valve and an additional fourth ridge on the junction of the dorsal and posterior sides of the right valve. Biradiolites ingens (Des Moulins) could be the direct ancestor of Glabrobournonia. The paleogeographic distribution of Glabrobournonia suggests that this genus dispersed to central Asia from the late Campanian time, becoming widely distributed in the eastern Tethyan region rather than endemic to eastern Arabia. Correspondingly, specimens belonging to Gyropleura yielded from the same bed as G. minor in southwestern Tarim Basin, are similar to the specimens which were attributed to the eastern Arabian Gyropleura sp.; Campanian to early Maastrichtian Osculigera specimens described from the Yigeziya Formation are comparable with those known from the Campanian–Maastrichtian of Iran, Afghanistan and eastern Arabia. The similarity of the rudist assemblages between central Asia and eastern Arabia suggests a faunal connection and affinity between the north and south margins of the eastern Tethyan realm during Campanian to early Maastrichtian times.     

Intra- and interspecific variability in allodaposuchid crocodylomorphs and the status of western European taxa

The genus Allodaposuchus is an endemic eusuchian from the Late Cretaceous of Europe. This genus was erected in 1928 by Baron Franz Nopcsa based on Allodaposuchus precedens from the Maastrichtian of Romania. Fragmentary skulls recovered from France and Spain were later referred to A. precedens, but three new species of Allodaposuchus have since been described: A. subjuniperus, A. palustris and A. hulki. A set of remains from Velaux, France, was recently interpreted as an ontogenetic series of A. precedens, prompting the argument that other species referred to Allodaposuchus are synonyms of A. precedens. Here, we review intra- and interspecific variability among allodaposuchids. Diagnostic characters for different allodaposuchids are outside the ranges of variation for modern species. Ontogenetic (intraspecific) variation observed in the allodaposuchid from Velaux is not in conflict with the presence of at least four taxa in the European Archipelago during the Late Cretaceous.     

New Crocodyliforms from Southwestern Europe and Definition of a Diverse Clade of European Late Cretaceous Basal Eusuchians

The late Campanian-early Maastrichtian site of Lo Hueco (Cuenca, Spain) has provided a set of well-preserved crocodyliform skull and lower jaw remains, which are described here and assigned to a new basal eusuchian taxon, Lohuecosuchus megadontos gen. et sp. nov. The reevaluation of a complete skull from the synchronous site of Fox-Amphoux (Department of Var, France) allows us to define a second species of this new genus. Phylogenetic analysis places Lohuecosuchus in a clade exclusively composed by European Late Cretaceous taxa. This new clade, defined here as Allodaposuchidae, is recognized as the sister group of Hylaeochampsidae, also comprised of European Cretaceous forms. Allodaposuchidae and Hylaeochampsidae are grouped in a clade identified as the sister group of Crocodylia, the only crocodyliform lineage that reaches our days. Allodaposuchidae shows a vicariant distribution pattern in the European Late Cretaceous archipelago, with several Ibero-Armorican forms more closely related to each other than with to Romanian Allodaposuchus precedens.     

A new late Cretaceous azhdarchid (Pterosauria,Azhdarchidae) from the Volga Region

A new azhdarchid genus and species, Volgadraco bogolubovi gen. et sp. nov., is described based on an anterior fragment of the mandibular symphysis (mandibular beak) and some postcranial elements from the Rybushka Formation (Upper Cretaceous, Lower Campanian) of the Shirokii Karamysh 2 locality, Saratov Region. The new taxon is intermediate in size and vascularization of the mandibular beak between medium-sized Turonian-Santonian azhdarchids (Azhdarcho, Bakonydraco) and the giant Maastrichtian azhdarchid Quetzalcoatlus.     

Osteology of Pseudochampsa ischigualastensis gen. et comb. nov. (Archosauriformes: Proterochampsidae) from the Early Late Triassic Ischigualasto Formation of Northwestern Argentina

Proterochampsids are crocodile-like, probably semi-aquatic, quadrupedal archosauriforms characterized by an elongated and dorsoventrally low skull. The group is endemic from the Middle-Late Triassic of South America. The most recently erected proterochampsid species is “Chanaresuchus ischigualastensis”, based on a single, fairly complete skeleton from the early Late Triassic Ischigualasto Formation of northwestern Argentina. We describe here in detail the non-braincase cranial and postcranial anatomy of this species and revisit its taxonomy and phylogenetic relationships. The phylogenetic analysis recovered ‘Chanaresuchus ischigualastensis’ as part of a trichotomy together with Gualosuchus reigi and Chanaresuchus bonapartei. Accordingly, “Chanaresuchus ischigualastensis” can be potentially more closely related to Gualosuchus reigi, or even Rhadinosuchus gracilis, than to Chanaresuchus bonapartei. In addition, after discussing previously claimed synapomorphies of Chanaresuchus, we could not find unambiguous support for the monophyly of the genus. As a result, we propose here the erection of the new genus Pseudochampsa for ‘Chanaresuchus ischigualastensis’, which results in the new combination Pseudochampsa ischigualastensis. The information provided here about the anatomy and taxonomy of Pseudochampsa ischiguaslastensis will be useful for future quantitative analyses focused on the biogeography and macroevolutionary history of proterochampsids.     

Diversity,Relationships, and Biogeography of the Lambeosaurine Dinosaurs from the European Archipelago,with Description of the New Aralosaurin Canardia garonnensis

We provide a thorough re-evaluation of the taxonomic diversity, phylogenetic relationships, and historical biogeography of the lambeosaurine hadrosaurids from the European Archipelago. Previously published occurrences of European Lambeosaurinae are reviewed and new specimens collected from upper Maastrichtian strata of the south-central Pyrenees are described. No support is found for the recognition of European saurolophines in the available hadrosaurid materials recovered so far from this area. A new genus and species of basal lambeosaurine, Canardia garonnensis, is described on the basis of cranial and appendicular elements collected from upper Maastrichtian strata of southern France. C. garonnensis differs from all other hadrosaurids, except Aralosaurus tuberiferus, in having maxilla with prominent subrectangular rostrodorsal flange; it differs from A. tuberiferus in a few maxillary and prefrontal characters. Together with A. tuberiferus, C. garonnensis integrates the newly recognized tribe Aralosaurini. Inference of lambeosaurine interrelationships via maximum parsimony analysis indicates that the other three known European lambeosaurines are representatives of two additional subclades (tribes) of these hadrosaurids: Tsintaosaurini (Pararhabdodon isonensis) and Lambeosaurini (the Arenysaurus ardevoli-Blasisaurus canudoi clade). The tribes Aralosaurini, Tsintaosaurini, Lambeosaurini, and Parasaurolophini are formally defined and diagnosed for the first time. Three event-based quantitative methods of ancestral range reconstruction were implemented to infer the historical biogeography of European lambeosaurines: Dispersal-Vicariance Analysis, Bayesian Binary MCMC, and Dispersal-Extinction-Cladogenesis. The results of these analyses, coupled with the absence of pre-Maastrichtian lambeosaurines in the Mesozoic vertebrate fossil record of Europe, favor the hypothesis that aralosaurins and tsintaosaurins were Asian immigrants that reached the Ibero-Armorican island via dispersal events sometime during the Maastrichtian. Less conclusive is the biogeographical history of European lambeosaurins; several scenarios, occurring sometime during the Maastrichtian, are possible, from vicariance leading to the splitting of Asian or North American from European ranges to a dispersal event from North America to the European Archipelago.     

Phylogeny,Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary

Background

Rhabdodontid ornithopod dinosaurs are characteristic elements of Late Cretaceous European vertebrate faunas and were previously collected from lower Campanian to Maastrichtian continental deposits. Phylogenetic analyses have placed rhabdodontids among basal ornithopods as the sister taxon to the clade consisting of Tenontosaurus, Dryosaurus, Camptosaurus, and Iguanodon. Recent studies considered Zalmoxes, the best known representative of the clade, to be significantly smaller than closely related ornithopods such as Tenontosaurus, Camptosaurus, or Rhabdodon, and concluded that it was probably an island dwarf that inhabited the Maastrichtian Haţeg Island.

Methodology/Principal Findings

Rhabdodontid remains from the Santonian of western Hungary provide evidence for a new, small-bodied form, which we assign to Mochlodon vorosi n. sp. The new species is most similar to the early Campanian M. suessi from Austria, and the close affinities of the two species is further supported by the results of a global phylogenetic analysis of ornithischian dinosaurs. Bone histological studies of representatives of all rhabdodontids indicate a similar adult body length of 1.6–1.8 m in the Hungarian and Austrian species, 2.4–2.5 m in the subadults of both Zalmoxes robustus and Z. shqiperorum and a much larger, 5–6 m adult body length in Rhabdodon. Phylogenetic mapping of femoral lengths onto the results of the phylogenetic analysis suggests a femoral length of around 340 mm as the ancestral state for Rhabdodontidae, close to the adult femoral lengths known for Zalmoxes (320–333 mm).

Conclusions/Significance

Our analysis of body size evolution does not support the hypothesis of autapomorhic nanism for Zalmoxes. However, Rhabdodon is reconstructed as having undergone autapomorphic giantism and the reconstructed small femoral length (245 mm) of Mochlodon is consistent with a reduction in size relative to the ancestral rhabdodontid condition. Our results imply a pre-Santonian divergence between western and eastern rhabdodontid lineages within the western Tethyan archipelago.     

A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia

Although the group played an important role in the evolution of Late Mesozoic terrestrial ecosystems, the early evolutionary history of the ornithischian dinosaurs remains poorly understood. Here, we report on a new primitive ornithischian, Eocursor parvus gen. et sp. nov., from the Late Triassic (?Norian) Lower Elliot Formation of South Africa. Eocursor is known from a single specimen comprising substantial cranial and postcranial material and represents the most complete Triassic member of Ornithischia, providing the earliest evidence for the acquisition of many key ornithischian postcranial characters, including an opisthopubic pelvis. A new phylogenetic analysis positions this taxon near the base of Ornithischia, as the sister taxon to the important and diverse clade Genasauria. The problematic clade Heterodontosauridae is also positioned basal to Genasauria, suggesting that an enlarged grasping manus may represent a plesiomorphic ornithischian condition. This analysis provides additional phylogenetic support for limited ornithischian diversity during the Late Triassic, and suggests that several major ornithischian clades may have originated later than generally believed. There are few morphological differences between Late Triassic and Early Jurassic ornithischians, supporting previous suggestions that the Early Jurassic ornithischian radiation may simply represent the filling of vacant ecological space following Late Triassic terrestrial extinctions.     

Phytogeographical Implication of Bridelia Will. (Phyllanthaceae) Fossil Leaf from the Late Oligocene of India

Background

The family Phyllanthaceae has a predominantly pantropical distribution. Of its several genera, Bridelia Willd. is of a special interest because it has disjunct equally distributed species in Africa and tropical Asia i.e. 18–20 species in Africa-Madagascar (all endemic) and 18 species in tropical Asia (some shared with Australia). On the basis of molecular phylogenetic study on Bridelia, it has been suggested that the genus evolved in Southeast Asia around 33±5 Ma, while speciation and migration to other parts of the world occurred at 10±2 Ma. Fossil records of Bridelia are equally important to support the molecular phylogenetic studies and plate tectonic models.

Results

We describe a new fossil leaf of Bridelia from the late Oligocene (Chattian, 28.4–23 Ma) sediments of Assam, India. The detailed venation pattern of the fossil suggests its affinities with the extant B. ovata, B. retusa and B. stipularis. Based on the present fossil evidence and the known fossil records of Bridelia from the Tertiary sediments of Nepal and India, we infer that the genus evolved in India during the late Oligocene (Chattian, 28.4–23 Ma) and speciation occurred during the Miocene. The stem lineage of the genus migrated to Africa via “Iranian route” and again speciosed in Africa-Madagascar during the late Neogene resulting in the emergence of African endemic clades. Similarly, the genus also migrated to Southeast Asia via Myanmar after the complete suturing of Indian and Eurasian plates. The emergence and speciation of the genus in Asia and Africa is the result of climate change during the Cenozoic.

Conclusions

On the basis of present and known fossil records of Bridelia, we have concluded that the genus evolved during the late Oligocene in northeast India. During the Neogene, the genus diversified and migrated to Southeast Asia via Myanmar and Africa via “Iranian Route”.     

A new genus for a rare African vespertilionid bat: insights from South Sudan

A new genus is proposed for the strikingly patterned African vespertilionid “Glauconycteris” superba Hayman, 1939 on the basis of cranial and external morphological comparisons. A review of the attributes of a newly collected specimen from South Sudan (a new country record) and other museum specimens of “Glauconycteris” superba suggests that “Glauconycteris” superba is markedly distinct ecomorphologically from other species classified in Glauconycteris and is likely the sister taxon to Glauconycteris sensu stricto. The recent capture of this rarely collected but widespread bat highlights the need for continued research in tropical sub-Saharan Africa and in particular, for more work in western South Sudan, which has received very little scientific attention. New country records for Glauconycteris cf. poensis (South Sudan) and Glauconycteris curryae (Gabon) are also reported.     

A new turtle from the La Colonia Formation (Campanian–Maastrichtian), Patagonia,Argentina, with remarks on the evolution of the vertebral column in turtles

Abstract: Patagoniaemys gasparinae gen. et sp. nov. is a new stem turtle found in central Patagonia, Chubut Province, Argentina, in outcrops of the La Colonia Formation (Campanian–Maastrichtian). This is a turtle of relatively large size (carapace length c. 70 cm), and the preserved remains of the holotype consist of skull fragments and several postcranial elements including a nearly complete vertebral column. A phylogenetic analysis shows Patagoniaemys gasparinae gen. et sp. nov. forming a monophyletic group with Otwayemys cunicularius and Mongolochelys efremovi, as a sister group to Meiolaniidae. A comprehensive review confirms that formed cervical vertebrae appeared independently several times during turtle evolution: in the clade that includes Patagoniaemys gasparinae gen. et sp. nov. and Meiolaniidae, in some baenids, in the total group Pleurodira and in crown group Cryptodira. Likewise, formed caudal vertebrae appeared several times in turtle evolution.     

Reassessment of the taxonomic position of Iranocypris typhlops Bruun & Kaiser, 1944 (Actinopterygii,Cyprinidae)

The Iranian cave barb (Iranocypris typhlops Bruun & Kaiser, 1944) is a rare and endemic species of the family Cyprinidae known from a single locality in the Zagros Mountains, western Iran. This species is “Vulnerable” according to the IUCN Red List and is one of the top four threatened freshwater fish species in Iran. Yet, the taxonomic position of I. typhlops is uncertain. We examined phylogenetic relationships of this species with other species of the family Cyprinidae based on the mitochondrial cytochrome b gene. Our results show that I. typhlops is monophyletic and is sister taxon of a cluster formed by Garra rufa (Heckel, 1843) and Garra barreimiae (Fowler & Steinitz, 1956) within a clade that includes other species of the genus Garra. Based on previous molecular and morphological studies, as well as our new results, we recommend that I. typhlops should be transferred to the genus Garra Hamilton, 1822.     

A new kogaionid multituberculate mammal from the Maastrichtian of the Transylvanian Basin,Romania

The Latest Cretaceous (Maastrichtian) terrestrial sedimentary sequences of the Haţeg Basin in Transylvania are well known for the so-called “Haţeg Island” vertebrate faunas, which evolved in endemic (insular?) conditions. In addition to frogs, lizards, turtles, crocodilians, birds and dinosaurs, peculiar multituberculate mammals have been recorded, all belonging to the family Kogaionidae. Here, a new species of the genus Barbatodon is reported from the Maastrichtian Şard Formation in the Transylvanian Basin (Alba County, Romania). Barbatodon oardaensis n. sp. is characterized by M1 cusp formula 3:4:2 and is much smaller than the two other Maastrichtian kogaionids from Transylvania, Barbatodon transylvanicus and Kogaionon ungureanui. The origin and paleobiogeography of kogaionids are discussed.     

A Review of Shamosuchus and Paralligator (Crocodyliformes,Neosuchia) from the Cretaceous of Asia

The crocodyliform Shamosuchus is known from numerous Late Cretaceous localities in southern and eastern Mongolia and fragmentary remains from Uzbekistan. Seven species of Shamosuchus have been named from six localities in Mongolia and three in Uzbekistan. Six species originally described as Paralligator were later referred to Shamosuchus. Only the type species, Shamosuchus djadochtaensis has been examined in detail. Many of the named species of Shamosuchus show striking similarity in size and cranial morphology but most are based on partial remains suggesting that the true species diversity is overestimated. A review of all species referred to Shamosuchus recognizes three valid taxa: Shamosuchus djadochtaensis, S. gradilifrons, and S. major. Shamosuchus sungaricus, S. borealis, and S. karakalpakensis are nomena dubia, whereas S. ancestralis, S. ulgicus, S. tersus, and S. ulanicus are junior subjective synonyms of S. gradilifrons. Phylogenetic analysis of 318 phenotypic characters recovers a Paralligatoridae clade consisting of Shamosuchus, Rugosuchus, Batrachomimus, Glen Rose Form, and Wannchampsus. Shamosuchus is non-monophyletic: S. djadochtaensis is near the base of Paralligatoridae whereas S. gradilifrons + S. major are the most deeply nested. The name Paralligator is resurrected for this clade. Rugosuchus and Batrachomimus are sister taxa to Paralligator. Paralligatoridae is closely related to Theriosuchus, hylaeochampsids and a speciose Allodaposuchus clade, which together are the sister group of Borealosuchus plus Crocodylia. These results support the presence of a diverse clade in eastern Asia and western North America throughout the Cretaceous with origins in the Late Jurassic.     

A new species of Phoenicopsis (Leptostrobales) from the Maastrichtian–Danian of Chukotka,Russia

A new species, Phoenicopsis anadyrensis Nosova, is described based on the leaf morphology and cuticle features from the Maastrichtian–Danian Rarytkin Formation, Chukotka, North-East of Russia. It is characterized by hypostomatic leaves with stomatal bands on the abaxial side, undulate anticlinal epidermal cell walls and papillae on the epidermal cells and on the subsidiary cells of the stomata. The new species is the youngest reliable representative of the genus Phoenicopsis. This record extends the stratigraphic range of Phoenicopsis from its previously known latest records in the early Campanian up to Maastrichtian–Danian. The persistence of the genus Phoenicopsis up to the Cretaceous–Paleogene boundary is most likely related with vicinity to mountain floras associated with Late Cretaceous and Paleogene volcanic uplands.     

A new species of Acynodon (Crocodylia) from the upper cretaceous (Santonian–Campanian) of Villaggio del Pescatore,Italy

Abstract: The new species Acynodon  adriaticus is described on the basis of remains from the Santonian–Campanian of Villaggio del Pescatore (Trieste, NE Italy). This species differs in several cranial features from Acynodon  iberoccitanus, the only other Acynodon species whose cranial osteology is known in detail. The absence of maxillary and dentary caniniform teeth coupled with the presence of enlarged molariform teeth suggests that Acynodon probably fed on slowly moving hard‐shelled prey. Moreover, the new materials reveal for the first time the morphology of some postcranial elements of Acynodon: in particular, medial‐most paravertebral osteoderms that are characterized by two keels. A new cladistic phylogenetic analysis resolves the previously reported polytomy among the basal Globidonta: Acynodon is recognized as the most primitive globidontan. This genus may represent the geologically oldest known globidontan. The fact that Acynodon has been found only in Europe and that the outgroup of Globidonta, the Diplocynodontinae, is mainly known from Europe, suggests that globidontans may have originated in Europe and not in North America as previously supposed.     

Premiere decouverte en Europe Sud-occidentale de Praeovibos priscus (Mammalia,Artiodactyla, Ovibovinae) dans le gisement pleistocene moyen ante-risseien de la Caune de l'Arago (Tautavel,Pyrenees-Orientales,France)

Teeth, cranial and postcranial skeletton of Praeovibos were sampled from the sediments of the Caune de l'AragoCave. This material is enough to recognize the anatomical relationships between Praeovibos and Ovibos. Praeovibos is discovered for the first time in France and in South Western Europe also. The problem of the correlation of the whole European Middle Pleistocene is approached from a different point of view.     

Fossil and Genetic Evidence for the Polyphyletic Nature of the Planktonic Foraminifera "Globigerinoides", and Description of the New Genus Trilobatus

Planktonic foraminifera are one of the most abundant and diverse protists in the oceans. Their utility as paleo proxies requires rigorous taxonomy and comparison with living and genetically related counterparts. We merge genetic and fossil evidence of “Globigerinoides”, characterized by supplementary apertures on spiral side, in a new approach to trace their “total evidence phylogeny” since their first appearance in the latest Paleogene. Combined fossil and molecular genetic data indicate that this genus, as traditionally understood, is polyphyletic. Both datasets indicate the existence of two distinct lineages that evolved independently. One group includes “Globigerinoides” trilobus and its descendants, the extant “Globigerinoides” sacculifer, Orbulina universa and Sphaeroidinella dehiscens. The second group includes the Globigerinoides ruber clade with the extant G. conglobatus and G. elongatus and ancestors. In molecular phylogenies, the trilobus group is not the sister taxon of the ruber group. The ruber group clusters consistently together with the modern Globoturborotalita rubescens as a sister taxon. The re-analysis of the fossil record indicates that the first “Globigerinoides” in the late Oligocene are ancestral to the trilobus group, whereas the ruber group first appeared at the base of the Miocene with representatives distinct from the trilobus group. Therefore, polyphyly of the genus "Globigerinoides" as currently defined can only be avoided either by broadening the genus concept to include G. rubescens and a large number of fossil species without supplementary apertures, or if the trilobus group is assigned to a separate genus. Since the former is not feasible due to the lack of a clear diagnosis for such a broad genus, we erect a new genus Trilobatus for the trilobus group (type species Globigerina triloba Reuss) and amend Globoturborotalita and Globigerinoides to clarify morphology and wall textures of these genera. In the new concept, Trilobatus n. gen. is paraphyletic and gave rise to the Praeorbulina / Orbulina and Sphaeroidinellopsis / Sphaeroidinella lineages.     

The marine diapsid reptile Endennasaurus from the Upper Triassic of Italy

Abstract:  The marine reptile Endennasaurus from the Upper Triassic Zorzino Limestone of northern Italy is redescribed and reassessed. New details of the skull and postcranial skeleton are revealed, confirming the attribution of this genus to the diapsid reptile clade Thalattosauriformes. Phylogenetic analysis suggests that Endennasaurus was related to the European genus Askeptosaurus and the Chinese Anshunsaurus . Despite a rather conservative postcranial morphology, Endennasaurus clearly occupied a highly specialized dietary niche as it combined a slender tapering premaxillary rostrum with a complete absence of either marginal or palatal teeth.