ADDITIONAL SPECIMENS OF SUDAMERICID (GONDWANATHERIA) MAMMALS FROM THE EARLY PALEOCENE OF ARGENTINA

Abstract: The extinct, Cretaceous–Paleogene Gondwanatherians have previously been considered to be early xenarthrans, multituberculates and more recently Mammalia incertae sedis. However, the phylogenetic relationships of Gondwanatheria have yet to be resolved. In this paper, additional dental specimens of the gondwanatherian Sudamerica ameghinoi from the Early Paleocene Salamanca Formation of Argentina are described. These specimens provide additional information on Gondwanatheria affinities, sudamericid morphology and help support earlier hypotheses on Sudamerica dental formula and tooth categories. Sudamericid dental functional morphology and body mass estimates, based on measurements of isolated teeth, are inferred. Dental morphology such as hypsodonty, enamel microstructure and crown features do support a robust clade for Sudamericidae.     

Squatiniformes (Chondrichthyes,Neoselachii) from the Late Cretaceous of southern England and northern France with redescription of the holotype of Squatina cranei Woodward, 1888

Abstract: Bulk sampling of phosphate‐rich horizons within the Late Cretaceous of the Anglo‐Paris Basin yielded numerous teeth of members of the Squatiniformes. Along with isolated tooth remains, two museum specimens comprising partial articulated encoskeletal remains including the holotype of the species Squatina cranei Woodward, 1888a are described, and a new subgenus Cretascyllium is proposed for species of the genus Squatina with high degree of heterodonty and triangular anterior teeth. The species Squatina (Cretascyllium) cranei comb. nov. and Squatina (Cretascyllium) hassei comb. nov. are referred to this subgenus. The genus Parasquatina Herman, 1982 previously erected on a single tooth is valid, and two new species P. justinensis sp. nov. and P. jarvisi sp. nov. are described along with a third taxon Parasquatina sp. An enigmatic tooth referred to ?Neoselachii incertae sedis is also reported. The palaeoecology of these taxa is discussed.     

SHARKS OF THE ORDER CARCHARHINIFORMES FROM THE BRITISH CONIACIAN, SANTONIAN AND CAMPANIAN (UPPER CRETACEOUS)

Abstract: Bulk sampling of phosphate‐rich horizons within the British Coniacian to Campanian (Upper Cretaceous) yielded very large samples of shark and ray teeth. All of these samples yielded teeth of diverse members of the Carcharhiniformes, which commonly dominate the fauna. The following species are recorded and described: Pseudoscyliorhinus reussi ( Herman, 1977 ) comb. nov., Crassescyliorhinus germanicus ( Herman, 1982 ) gen. nov., Scyliorhinus elongatus ( Davis, 1887 ), Scyliorhinus brumarivulensis sp. nov., ?Palaeoscyllium sp., Prohaploblepharus riegrafi ( Müller, 1989 ) gen. nov., ?Cretascyliorhinus sp., Scyliorhinidae incertae sedis 1, Scyliorhinidae incertae sedis 2, Pteroscyllium hermani sp. nov., Protoscyliorhinus sp., Leptocharias cretaceus sp. nov., Palaeogaleus havreensis Herman, 1977 , Paratriakis subserratus sp. nov., Paratriakis tenuis sp. nov., Paratriakis sp. indet. and ?Loxodon sp. Taxa belonging to the families ?Proscylliidae, Leptochariidae and Carcharhinidae are described from the Cretaceous for the first time. The evolutionary and palaeoecological implications of these newly recognised faunas are discussed.     

The Phylogenetic Affinities of the Enigmatic Mammalian Clade Gondwanatheria

Gondwanatheria is a group of extinct mammals known from the Cretaceous and Paleogene of Gondwana. Resolution of the phylogenetic affinities of gondwanatherians has proven problematical, with the group currently considered Mammalia incertae sedis. We briefly review the morphology of known gondwanatherians, and argue that isolated upper premolars and a partial dentary preserving a blade-like p4 originally referred to the ferugliotheriid gondwanatherian Ferugliotherium windhauseni but subsequently identified as Multituberculata incertae sedis do indeed belong to F. windhauseni. We also suggest that the recently described ?cimolodontan multituberculate Argentodites coloniensis, based on an isolated lower premolar, may in fact be an unworn p4 of Ferugliotherium or a closely related taxon. We present the first phylogenetic analyses to include gondwanatherians, using maximum parsimony and Bayesian methods. Both methods place Ferugliotherium and sudamericid gondwanatherians in a clade with cimolodontan and “plagiaulacidan” multituberculates, although relationships within this clade are largely unresolved. The Gondwanatheria + Multituberculata clade supported here may reflect the convergent evolution of similar dental features, but it is the best supported hypothesis based on currently available data. However, denser sampling of multituberculate taxa and the discovery of more complete gondwanatherian fossils will be required to clarify the precise relationship between gondwanatherians and multituberculates, specifically to determine whether or not gondwanatherians are members of Multituberculata. We hypothesize that the anterior molariforms of sudamericid gondwanatherians evolved from blade-like precursors similar to the p4 of Ferugliotherium, possibly in response to the appearance of grasses in Gondwana during the Cretaceous.     

First mammal evidence from the Late Cretaceous of India for biotic dispersal between India and Africa at the KT transition

The Late Cretaceous record of mammals from India assumes great significance in view of the fact that it is the only Gondwanan landmass that has yielded definitive eutherian mammals. These mammals have variously been assigned to palaeoryctids, archontans or Eutheria incertae sedis. Well preserved lower molars recovered from a new mammal-yielding Deccan intertrappean site near Kisalpuri village, Dindori District, Madhya Pradesh (state), India, are described here under a new species Deccanolestes narmadensis sp. nov. The new fossil material indicates close phylogenetic relationship between Deccanolestes from India and Afrodon (Adapisoriculidae) from the Late Palaeocene of Africa and Europe. In view of older age and more primitive state of Deccanolestes teeth, it is inferred that Deccanolestes represents an ancestral morphotype from which the African/European adapisoriculid Afrodon may have been derived. This is the first compelling terrestrial fossil evidence for an early dispersal between India and Africa. Such a dispersal possibly involved an East African contact with India at the KT transition.     

Elasmobranchs from the Lower Triassic Sulphur Mountain Formation near Wapiti Lake (BC, Canada)

Hybodontoid and nonhybodontoid sharks are described from the Lower Triassic Vega‐Phroso Siltstone Member of Sulphur Mountain Formation on the basis of newly discovered material. The age of the classic fossil site ‘Wapiti Lake’ in the Canadian Rocky Mountains is discussed on the basis of new field data and one conodont found in association. Preliminary results suggest that these elasmobranch remains are between early Smithian and Spathian in age. Apart from the enigmatic genus Listracanthus and previously reported edestoids, the shark fauna consists of at least one hybodont, at least two questionable hybodontoid genera and an elasmobranch of enigmatic affinities, represented by peculiar denticles only and described as ‘genus A’incertae sedis. The presence of the only previously reported hybodont genus, cf. Palaeobates, is erroneous. The largest specimen represents the most complete Early Mesozoic shark known. The heterodonty of its dentition, fin spine morphology and the short, robust body shape imply it represents a member of a new family of shark, Wapitiodidae fam. nov. , and is described here as Wapitiodus aplopagus gen. et sp. nov. The unique dental morphology shows affinities to Polyacrodus but clearly differs in the complete lack of side cusps. Wapitiodus gen. nov. possesses a primitive fin spine structure. The tooth crowns are entirely blunt in the distal (posterior) tooth files, and are acuminate‐unicuspid in several anterior files. Tooth morphology, the shape of the basal cartilages, the proximal insertion of the fin spines and the pectoral fin structure are interpreted as diagnostic characters for this new genus, and possibly for the Wapitiodidae fam. nov. The majority of observed characters appear to be primitive and are reminiscent of Palaeozoic sharks, however, and these features include dorsal fin spine morphology and gross skull anatomy. A second species, provisionally placed in the same genus, is described as Wapitiodus homalorhizo sp. nov. Wapitiodus homalorhizo sp. nov. can be distinguished from W. aplopagus gen. et sp. nov. by the proportions of the fin spines, tooth morphology and possibly the body shape. Several isolated teeth and other fragmentary material are referred to either Wapitiodus gen. nov. sp. indet. or to ?Polyacrodus sp. (Polyacrodontidae gen. et sp. indet.). A third genus of elasmobranch (incertae sedis) is described as ‘Genus A’ and is recognized by its peculiar scales. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149 , 309–337.     

Paranthodon africanus (broom) a stegosaurian dinosaur from the Lower Cretaceous of South Africa

The holotype of Paranthodon africanus (Broom) from the Kirkwood Formation (Lower Cretaceous) of the Algoa Basin, Cape Province, South Africa consists of a partial skull, the bones of which are very similar to those of Stegosaurus. Both sides of the maxillary tooth crowns have a bulbous cingulum and a very prominent central vertical ridge above the large apical denticle that is bordered anteriorly and posteriorly by four to seven smaller denticles. Diagnostically stegosaurian material is also known from the Lower Cretaceous of England (Craterosaurus) and China (Wuerhosaurus) and the Upper Cretaceous of India (Dravidosaurus).     

A new scaphognathid pterosaur from western Liaoning,China

A partial skeleton of a new pterosaur, Jianchangnathus robustus gen. et sp. nov. from western Liaoning, China, is described. The specimen (IVPP V16866) was collected near Linglongta, Jianchang County, whose deposits have a disputed age that range from Middle Jurassic to Early Cretaceous. The new species shares several features with the non-pterodactyloid Scaphognathus from the Late Jurassic deposits of southern Germany, such as a deep anterior end of the lower jaw, a piriform lower temporal fenestra with the ventral margin broader than the dorsal one and the interalveolar spacing of the maxillary teeth about three alveolar spaces, allowing its allocation to the Scaphognathidae. The main diagnostic features of J. robustus include the large maxillary process of the jugal, the convex alveolar margin of the lower jaw and the procumbent disposition of the first three pairs of dentary teeth. The new Chinese taxon also differs from Fenghuangopterus lii, which comes from the same deposit and is here regarded as Scaphognathidae incertae sedis, mainly by the lower number of teeth and several proportions of the wing elements. The discovery of J. robustus demonstrates a larger diversity in the pterosaur fauna of the Linglongta region so far dominated by the non-pterodactyloid clade Wukongopteridae.     

Sauropod teeth from the Upper Cretaceous Bissekty Formation of Uzbekistan

The isolated adult teeth of titanosaurian sauropods from the Upper Cretaceous Bissekty Formation at Dzharakuduk, Uzbekistan, differ little in overall structure but show considerable variation in enamel sculpturing and wear patterns. The crown shape of unworn juvenile teeth ranges from lanceolate to conical. Most specimens have enamel texture resembling crumpled paper or completely smooth enamel. Longitudinal grooves along the mesial and distal edges are present on only a few tooth crowns and might be developed on both the labial and lingual sides. Among 252 worn tooth crowns there are eight variants of wear patterns, all possible combinations of 0–2 apical and 0–2 lateral wear facets. The most common is wear pattern A1L0 (one apical facet, no lateral facets; 62.7%). The next most common variant has two apical and no lateral facets (A2L0, 12.3%). These apical wear facets include the primary wear facets, which are produced by an opposing functional tooth, and secondary wear facets, which are produced by a replacing upper tooth coming into contact with the functional lower tooth at a late wear stage. The relative abundance of tooth crowns with two apical wear facets possibly suggests incipient development of a tooth battery in the Bissekty titanosaur.     

Fabrosauridae,the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda)

In fabrosaurids the upper jaw is flat and the lower jaw is slender so the ’cheek’ teeth are marginal and not inset as is the case in all other ornithischian dinosaurs. The ’cheek’ teeth of fabrosaurids have anteroposteriorly expanded crowns but lack wear surfaces formed by tooth to tooth contact. Two genera are recognized from the Triassic-Jurassic boundary of Lesotho with good material previously referred toFabrosaurus as a new genus that represents the most conservative ornithopod described to date. The anatomy ofNanosaurus (Upper Jurassic, U.S.A.) andEchinodon (Jurassic-Cretaceous boundary, England) is redescribed; in both genera the tooth bearing bone of the lower jaw is deepened posteriorly and inEchinodon there is a true canine tooth in the upper jaw.     

Reorganising the order Bacillales through phylogenomics

Bacterial classification at higher taxonomic ranks such as the order and family levels is currently reliant on phylogenetic analysis of 16S rRNA and the presence of shared phenotypic characteristics. However, these may not be reflective of the true genotypic and phenotypic relationships of taxa. This is evident in the order Bacillales, members of which are defined as aerobic, spore-forming and rod-shaped bacteria. However, some taxa are anaerobic, asporogenic and coccoid. 16S rRNA gene phylogeny is also unable to elucidate the taxonomic positions of several families incertae sedis within this order. Whole genome-based phylogenetic approaches may provide a more accurate means to resolve higher taxonomic levels. A suite of phylogenomic approaches were applied to re-evaluate the taxonomy of 80 representative taxa of eight families (and six family incertae sedis taxa) within the order Bacillales. This showed several anomalies in the current family and order level classifications including the existence of four Bacillaceae and two Paenibacillaceae “family” clades. Furthermore, the families Staphylococcaceae and Listeriaceae belong to the sister order Lactobacillales. Finally, we propose a consensus phylogenomic approach which may diminish algorithmic biases associated with single approaches and facilitate more accurate classification of a broad range of taxa at the higher taxonomic levels.     

New genera and species of coelacanths from the Bear Gulch Limestone (Lower Carboniferous) of Montana (U.S.A.)

Four new species of coelacanths (Actinistia) from the Namurian E2b marine Bear Gulch Limestone of Montana are described, and a fifth species, originally described as an actinopterygian, is redescribed. Two new suborders are diagnosed. The taxonomy is: Suborder Coelacanthocdei, family Rhabdodermatidae, Caridosuctor populosum nov. sp., suborder Hadronectoroidei nov., family Hadronectoridae nov., Hadronector donbairdi nov. sp., Polyosteorhynchus simplex nov. sp. and Allenypterus montanusMelton. Lochmocercus aciculiodontus nov. sp. is incertae sedis.     

A new chinchilloid (Rodentia,Hystricognathi) from the early Miocene of the localities of Bryn Gwyn and Gran Barranca (Patagonia,Argentina)

A new genus and species of chinchilloid rodent is described here. It was found in Colhuehuapian levels (early Miocene) of the localities of Bryn Gwyn and Gran Barranca, Chubut Province, Argentina. The new taxon shows a unique combination of characters (e.g., protohypsodont teeth, upper molars trilophodont, m1 and m2 tri- or tetralophodont with the second crest in position of variable development, and m3 trilophodont, cement absent) that make it different from any other known chinchilloids. We performed a phylogenetic analysis to corroborate the relationships of the new taxon within the Chinchilloidea. Our results indicate that the new taxon is best classified as Chinchilloidea incertae sedis, together with Incamys, Garridomys, and Scotamys. These species form the basal stock that leads to the modern lineage Chinchillidae (chinchillas and viscachas). The presence of the new taxon in these localities increases the diversity of chinchilloids during the early Miocene and reduces the dissimilarity between the faunas found in Bryn Gwyn and Gran Barranca.     

Hyostragulum simplex n. sp. (Incertae sedis) du dévonien du massif armoricain intérêt paléobiogéographique et systématique

In the western part of the “Synclinorium median armoricain (Rade de Brest), the presence of Hyostragulum simplex n. sp. in the lower part of the Troaon formation (Upper Emsian) is a supplementary element of hercynian fauna in Brittany during lower devonian time.For the new species, characterised by the lack of a septum on the base of the corallite, a fibrous microstructure, perhaps trabecular, is shown. The systematic position of the genus is discussed but ends up with a suggestion to let it remain incertae sedis..     

Phylogenetic reassessment of the five forewing radial-veined tribes of Riodininae (Lepidoptera: Riodinidae)

The relationships among the genera and tribal groupings of Riodininae with five forewing radial veins, and between these and tribes with four forewing radial veins, were examined using a phylogenetic analysis. Using the type species from all sixteen genera in the tribal groupings Eurybiini, Mesosemiini and incertae sedis (a presumed paraphyletic group of loosely related genera), and representatives from the four forewing radial‐veined riodinine tribes, thirty‐five new and traditional characters were coded from adult ecology, wing venation and pattern, the adult head and body, male and female genitalia, and early stage ecology and morphology. The majority of characters are illustrated. Phylogenetic analysis of these data produced five equally most parsimonious cladograms using equal weights and after successive weighting. The strict consensus of these confirms the monophyly of Eurybiini and Mesosemiini as currently conceived, but also indicates several higher‐level relationships not previously hypothesized. Mesosemiini is here more broadly defined to also include the entire incertae sedis section, and the tribe is divided into Mesosemiina, for the previously delimited Mesosemiini plus Eunogyra and Teratophthalma, and Napaeina, subtr.n. for the incertae sedis section minus these two genera. The following hypothesis of relationships is tentatively proposed for the basal clades of Riodininae: Mesosemiini + (Eurybiini + remainder of Riodininae). These new hypotheses, and the characters supporting them, are discussed and compared with those previously proposed.     

Pathologic tooth deformities in modern and fossil chondrichthians: a consequence of feeding-related injury

Deformed teeth are found as rare components of the dentitions of both modern and fossil chondrichthians. Tooth deformities occur as bent or twisted tooth crowns, missing or misshaped cusps, atypical protuberances, perforations, and abnormal root structures. Deformed tooth files consisting of unusually overlapped or small teeth, or teeth misaligned in the jaw also occur in modern forms, but deformed tooth files generally are not recognizable in fossils due to post-mortem dissociation of teeth and jaws. A survey of 200 modern lamniform and carcharhiniform sharks as well as literature sources indicate that such deformities are produced by feeding-related injury to the tooth-forming tissue of the jaws, particularly by impaction of chondrichthian and teleost fin and tail spines. Tooth counts for several late Cretaceous genera, based on material recovered from coastal plain sites from New Jersey to Alabama, suggest that the frequency of occurrence of deformed teeth in a species varies from about 0.015% in Squalicorax kaupi to about 0.36% in Paranomotodon sp. Tooth counts for modern lamniform and carcharhiniform sharks yield a comparable range in frequency of tooth deformities. Variation in frequency of tooth deformity may reflect interspecific differences in feeding behavior and dietary preferences. There is no suggestion in our data of any strong patterns of temporal variation in tooth deformity frequency, or of patterns ­reflecting chondrichthian phylogenetic history and evolution. Skeletal components of the probable prey of the Cretaceous species are preserved in the same horizons as the deformed teeth, and also are found within co-occurring chondrichthian coprolites.     

Lamniform Shark Teeth from the Late Cretaceous of Southernmost South America (Santa Cruz Province,Argentina)

Here we report multiple lamniform shark teeth recovered from fluvial sediments in the (Campanian-Maastrichtian) Cerro Fortaleza Formation, Santa Cruz Province, Argentina. This small tooth assemblage is compared to various lamniform sharks possessing similar dental morphologies, including Archaeolamna, Cretalamna, Dwardius, Dallasiella, and Cretodus. Although the teeth share numerous morphological features with the genus Archaeolamna, including a developed neck that maintains a relatively consistent width along the base of the crown, the small sample size and incomplete nature of these specimens precludes definitive taxonomic assignment. Regardless, the discovery of selachian teeth unique from those previously described for the region broadens the known diversity of Late Cretaceous South American sharks. Additionally, the discovery of the teeth in fluvial sandstone may indicate a euryhaline paleobiology in the lamniform taxon or taxa represented by this tooth assemblage.     

“South American” Marsupials from the Late Cretaceous of North America and the Origin of Marsupial Cohorts

Newly described marsupial specimens of Judithian (late Campanian) and Lancian (Maastrichtian) age in the western interior of North America (Wyoming to Alberta) have dental morphologies consistent with those expected in comparably aged sediments in South America (yet to be found). Three new Lancian species are referable to the didelphimorphian Herpetotheriidae, which suggests that the ameridelphian radiation was well under way by this time. The presence of a polydolopimorphian from Lancian deposits with a relatively plesiomorphic dental morphology and an additional polydolopimorphian taxon from Judithian deposits with a more derived molar form indicate that this lineage of typically South American marsupials was diversifying in the Late Cretaceous of North America. This study indicates that typical South American lineages (e.g. didelphimorphians and polydolopimorphians) are not the result of North American peradectian progenitors dispersing into South America at the end of the Cretaceous (Lancian), or at the beginning of the Paleocene (Puercan), and giving rise to the ameridelphian marsupials. Instead, these lineages, and predictably others as well, had their origins in North America (probably in more southerly latitudes) and then dispersed into South America by the end of the Cretaceous. Geophysical evidence concerning the connections between North and South America in the Late Cretaceous is summarized as to the potential for overland mammalian dispersal between these places at those times. Paleoclimatic reconstructions are considered, as is the dispersal history of hadrosaurine dinosaurs and boid snakes, as to their contribution to an appraisal of mammalian dispersals in the Late Cretaceous. In addition, we present a revision of the South American component of the Marsupialia. One major outcome of this process is that the Polydolopimorphia is placed as Supercohort Marsupialia incertae sedis because no characteristics currently known from this clade securely place it within one of the three named marsupial cohorts.