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The dinosaur record from the stratigraphically-lowest portions of Upper Triassic deposits of western North America is poor and only consists of a handful of fragmentary hind limb elements. Here we present an articulated segment of the ankle region of a dinosaur that consists of the distal ends of the tibia and fibula and a complete astragalocalcaneum. Additionally, we suggest that an isolated femur and maxilla from the same locality may belong to the same taxon. Using the most comprehensive analysis of early theropod relationships currently available, we determined that the new specimen pertains to a coelophysoid neotheropod (i.e., more closely related to Coelophysis bauri than to Allosaurus fragilis). The stratigraphic position of the locality where the new specimen was discovered is equivalent to the famous Otis Chalk localities and this set of localities likely predates the rest of the Dockum Group and possibly the entirety of the fossiliferous portion of the Chinle Formation on the Colorado Plateau. Therefore, the new specimen represents one of the oldest neotheropods. Accordingly, neotheropods were present at or just after the onset of both the Chinle Formation and Dockum Group deposits.
Here we describe and illustrate specimens of hyaenodont mammals from two early Eocene localities of Southern France: Fournes (Minervois) and Fordones (Corbières). Some of these specimens were previously described as cf. Hyracolestes sp. (Cimolesta, Sarcodontidae), a taxon only known from Asia, but new arguments allow their referring to the small proviverrine hyaenodont Parvagula palulae which was previously only recorded in Palette (Provence). The material notably includes the oldest p4 ever recorded for the European endemic subfamily Proviverrinae. This new material shows that, by the beginning of the early Eocene, proviverrines already displayed their typical combination of dental features characterized by the presence of a large paraconid and entoconid on p4. The comparison between the earliest European proviverrines and sinopines (a mostly North American radiation) supports the divergence of the two subfamilies by this time. Moreover, the early proviverrines are morphologically similar to the African hyaenodont Tinerhodon (late Paleocene). Consequently, the history of the European proviverrines is likely rooted in Africa. Finally, the new specimens support a similar age for the localities of Palette and Fordones, and a younger age for Fournes. Due to the ages of these localities Parvagula palulae must be regarded as the oldest proviverrine.
Saddle oysters (Anomiidae) attach themselves to calcareous hard substrates by means of a calcified byssus that etches an attachment structure, referred to as ichnospecies Centrichnus eccentricus. Examination of rich material from the Late Cretaceous of central Europe extends the fossil record of this ichnotaxon and revealed a set of previously unrecognised morphological features which appear to be typical for this time period and the respective anomiid trace maker. Excellent preservation of a large number of trace fossil specimens with a complete set of morphological characters allowed a biometrical analysis and additional observations indicating a distinct substrate preference for belemnite rostra, a strong intra- and interspecific competition for settlement space, as well as interactions with durophaguous predators. Further implications for anomiid palaeobiology and palaeoecology arise from allometric shell growth and an etched outline suture in the substrate along the dorsal, lateral and ventral shell margins. These features enhanced a firm attachment and increased shear resistance, and thus are interpreted as an effective defence mechanism against shell-crushing enemies under the intensified predation pressure in marine environments in the Late Cretaceous.
The Chinchilla Sand is a formally defined stratigraphic sequence of Pliocene fluviatile deposits that comprise interbedded clay, sand, and conglomerate located in the western Darling Downs, south-east Queensland, Australia. Vertebrate fossils from the deposits are referred to as the Chinchilla Local Fauna. Despite over a century and a half of collection and study, uncertainties concerning the taxa in the Chinchilla Local Fauna continue, largely from the absence of stratigraphically controlled excavations, lost or destroyed specimens, and poorly documented provenance data. Here we present a detailed and updated study of the vertebrate fauna from this site. The Pliocene vertebrate assemblage is represented by at least 63 taxa in 31 families. The Chinchilla Local Fauna is Australia's largest, richest and best preserved Pliocene vertebrate locality, and is eminently suited for palaeoecological and palaeoenvironmental investigations of the late Pliocene.
The International Code of Zoological Nomenclature recommends providing the author and year of original description of any taxon, when first mentioned in a publication, which is useful for both taxonomic and nomenclatural reasons. Here we show that the authorship of the extinct hyaenid taxon Hyaena brevirostris, type species of the genus Pachycrocuta, has been incorrectly attributed to “Aymard, 1846” for more than a century on the basis of a publication that never existed. Our bibliographic research indicates that the first published work in which the name was employed—Gervais' (1848–1852) Zoologie et Paléontologie françaises, the part dealing with carnivorans being issued in 1850—fulfils the criteria of availability. Given that Gervais did not attribute the description to Aymard, but merely employed the latter's name in reference to a museum label/record (which does not constitute a valid indication), the correct name with authorship for this taxon is Pachycrocuta brevirostris (Gervais, 1850).
In this paper, we describe a previously unknown species of the glirid Eliomys from the Late Miocene and Early Pliocene Cabriel, Alcoy and Granada basins of southeastern Spain. Eliomys yevesi sp. nov. is characterized by its relative small size, narrow lingual wall and common presence of two centrolophs in the upper molars, and well-developed centrolophids in the lower molars. The new species is the probable ancestor of E. intermedius, which in turn represents the ancestor of the extant E. quercinus. According to its morphologic and biometric features, the origin of E. yevesi sp. nov. is likely to be found in some population of E. truci from the Late Miocene. Based on these affinities, we propose the lineage E. truci—E. yevesi sp. nov.—E. intermedius—E. quercinus, in which there is a trend towards the development of centrolophs, as well as the reduction of accessory crests.
Seriemas (Cariamidae) include two extant species, Cariama cristata and Chunga burmeisteri, which live in semi-open plains of South America and have a poorly documented evolutionary history. One of the earliest fossil representatives of Cariamidae is the recently described Noriegavis santacrucensis from the early Miocene Santa Cruz Formation (Argentina). So far, however, this species was only known from a cranium and tentatively referred distal tibiotarsi, and its phylogenetic assignment has been questioned. Here we describe a well-preserved partial skeleton from the Santa Cruz Formation, which substantiates the classification of Noriegavis in Cariamidae. Plesiomorphic features show N. santacrucensis to be outside crown group Cariamidae, but the species is nevertheless very similar to its modern relatives and documents that the osteology of seriemas underwent only few changes during the past 16 million years.
Therians (marsupials and placentals), archosaurs, and chameleons are remarkable in that they evolved postures and gaits with inturned forelimbs. However, recent studies have indirectly recognized that, unlike fully pronated therian and chameleon forearms, dinosaur forearms were mechanically constrained by semi-pronated (misaligned) joints. This has led to the hypothesis that quadrupedal dinosaurs mitigated this constraint via proximal migration of the radius, indirectly forming a more pronated, tubular manus distally. To test this hypothesis, a standardized pose was used to examine the forearm pronation of ornithischian dinosaurs that were obligatory quadrupeds and facultative bipeds. Results show that only restructuring of the distal, not the proximal radius, causes additional pronation of the pre-axial edge of the carpus, but also unexpectedly reveal that the ulna may help form a tubular manus by supinating the post-axial edge. Thus, relative to the plane of the elbow joint the wrist and finger joints remain wholly semi-pronated. These findings do not support the hypothesis that a tubular cross-section evolved in dinosaurs to pronate the manus further to allow the finger joints to participate in locomotion. Instead these results indicate that quadrupedal dinosaurs tended to abandon propulsive use of their wrist and finger joints by converting their carpus metacarpus into a vertical stilt-like extension of the forearm. Prior studies have overlooked that this divergent path to parasagittal forelimb kinematics had its phylogenetic basis in the retention of the semi-pronated forearm joint alignment that is plesiomorphic to tetrapods. Thus, this test provides the first functional explanation for the convergent responses of quadrupedal archosaurs to their misaligned forearm joints, and provides a foundation for elucidating why the quadrupedal evolution of archosaur forelimbs diverged from those of therians and chameleons.
The dinosaur record in the South Pyrenees Basin is diverse and rich. A total of 142 theropod teeth were studied for this paper, which constitutes one of the richest samples for these remains in Europe. Eight upper Campanian to upper Maastrichtian outcrops from the Pyrenees produced six non-avian theropod taxa (Theropoda indet., Coelurosauria indet., ?Richardoestesia, ?Dromaeosauridae indet., ?Pyroraptor olympius, ?Paronychodon). These six taxa are added to two previously described theropods (a Richardoestesia-like form and a possible ornithomimosaurid), indicating that there was considerable theropod diversity on the Iberian Peninsula during the Late Cretaceous.
Theropod teeth are particularly abundant in the fossil record and frequently reported in the literature. Yet, the dentition of many theropods has not been described comprehensively, omitting details on the denticle shape, crown ornamentations and enamel texture. This paucity of information has been particularly striking in basal clades, thus making identification of isolated teeth difficult, and taxonomic assignments uncertain. We here provide a detailed description of the dentition of Megalosauridae, and a comparison to and distinction from superficially similar teeth of all major theropod clades. Megalosaurid dinosaurs are characterized by a mesial carina facing mesiolabially in mesial teeth, centrally positioned carinae on both mesial and lateral crowns, a mesial carina terminating above the cervix, and short to well-developed interdenticular sulci between distal denticles. A discriminant analysis performed on a dataset of numerical data collected on the teeth of 62 theropod taxa reveals that megalosaurid teeth are hardly distinguishable from other theropod clades with ziphodont dentition. This study highlights the importance of detailing anatomical descriptions and providing additional morphometric data on teeth with the purpose of helping to identify isolated theropod teeth in the future.
Here we describe two instances of pathological vertebral fusion in two genera of sauropodomorph dinosaurs from the Early Jurassic Lufeng Formation in Yunnan, China. The first is a specimen, of Lufengosaurus huenei with two fused cervical vertebrae, and the other is a specimen of the Lufeng basal sauropod, with two fused caudal vertebrae. Both pathologies are consistent with spondyloarthropathy and represent the earliest known occurrence of that disease in dinosaurs. The two specimens affirm that early dinosaurs suffered from the same bone diseases as living vertebrates. Spondyloarthropathy in these dinosaurs may have been induced by long-term mechanical stress, such as weight bearing, and/or limited motion at the joint that would otherwise have inhibited such remodeling. In both cases, surface remodeling suggests that the animals survived well beyond the initiation of spondyloarthropathy.
The lifestyles of all pterosaurs are contentious, but those of the pterodactyloid clade Azhdarchidae are particularly debated. A 2008 review of the functional morphology of azhdarchid pterosaurs concluded that they were probably terrestrial foragers, as evidenced by their long limbs, generalised skull construction, the arthrological limitations of their cervical series, trackway data indicating terrestrial proficiency, a strong continental skew in the depositional context of their fossils, and several additional lines of corroborating evidence. This hypothesis was recently challenged on three counts: (i) azhdarchid fossils routinely occur in aquatic deposits; (ii) terrestrially-foraging pterosaurs were highly vulnerable to predation and (iii), aerial “water trawling”, where the mandible is pulled though water to catch food in a distended throat pouch, is a more likely foraging strategy. Pelican-like jaw mechanics were suggested for azhdarchids because of the asymmetrical jaw joints in these pterosaurs, which permit lateral deflection of the mandibular rami during jaw extension. We evaluate these three claims and conclude that all are flawed. The frequent occurrence of azhdarchid fossils in aquatic sedimentary systems is not significant with regard to ecology or behaviour, since these provide the overwhelming mechanism for the preservation of all fossil terrestrial animals. Likely pterosaur takeoff abilities and the ubiquitous nature of modern, terrestrially-foraging birds indicate that predation risks on ground-foraging pterosaurs are probably overstated. The kinematics of pterosaur jaws are entirely different to those of pelicans, which are highly specialised compared to those of all other tetrapods, and there are no indications from azhdarchid jaw anatomy that azhdarchids indulged in pelican- like foraging behaviour. The estimated amount of jaw expansion present in azhdarchids was minimal compared to that of pelicans, even when the asymmetrical jaw joints of azhdarchids are taken into account. Moreover, the widespread occurrence of asymmetrical jaw joints in other reptiles demonstrates that they are not related to any specific feeding habits. We conclude that terrestrial foraging remains the most parsimonious habit for azhdarchid pterosaurs.
Teleosauridae was a group of largely marine Mesozoic crocodylomorphs, typically considered as akin to “marine gavials” due to their elongate, tubular, polydont rostra that are indicative of a piscivorous diet. Here we show that these extinct crocodylomorphs were more anatomically, and perhaps ecologically, varied than previously thought. We report the first evidence of denticles in a teleosaurid tooth, revealed by scanning electron microscopic (SEM) analysis of a tooth from the holotype of “Steneosaurus” obtusidens. These denticles are cryptic, because they are microscopic, not contiguous along the carinae (instead forming short series), and are detectable only using SEM. This incipient denticle morphology is similar to that recently discovered in a closely related group of marine crocodylomorphs, the Metriorhynchidae. In particular, the denticulation morphology of “Steneosaurus” is similar to that of the geosaurin metriorhynchid Torvoneustes, indicating that these two taxa may have employed similar feeding styles and that “S.”obtusidens may have been a nearshore ecological analogue to the more offshore, fast-swimming geosaurins. Previous authors have considered “S.”obtusidens and Machimosaurus to be durophagous, but the discovery of denticulated teeth indicates that they had a more varied diet and feeding style, and included flesh slicing as part of their feeding toolkit. It is currently unknown how extensive denticulate carinae may be in Teleosauridae, and we hypothesise that cryptic denticles may also be present in other marine crocodylomorphs once they are subjected to SEM study.
Isolated elements comprising a set of ziphodont teeth and osteoderms from the middle Eocene karst deposit of Lissieu, France are assigned to either Iberosuchus sp. or Bergisuchus sp., poorly known crocodylomorphs of possible sebecosuchian affinities. A general survey of dorsal osteoderm variability among Crocodylomorpha points to the similarity between the osteoderms from Lissieu and those of some Cretaceous notosuchians, notably the sebecosuchian Baurusuchus. Such isolated osteoderms represent a useful tool for identifying non-eusuchian crocodylomorphs in post-Cretaceous deposits. Relying on the distinctive morphology of these osteoderms might help augmenting the extremely scant fossil record of Paleogene sebecosuchians in Europe, thus improving the spatiotemporal resolution of this group. At least three types of predators coexisted in the ecosystem of Lissieu, which indicates that in the middle Eocene of Europe, carnivoran mammals were not the exclusive predators in terrestrial habitats and shared resources with a ground bird and a sebecosuchian. This faunal composition is comparable to contemporaneous ecosystems in South America.
Keichousaurus hui is a small pachypleurosaur (Reptilia: Sauropterygia) from the Triassic of China. Many specimens of various growth stages are known, making them ideal for ontogenetic research. We report 22 new specimens from the Middle Triassic of Xingyi (Guizhou, south China), and combined their skeletal measurements with those from 85 published specimens to analyze the ontogenetic trajectory of sexual dimorphism. An Exploratory Factor Analysis suggests that the largest factors behind morphological disparity within the species are body size followed by gender. Sexual dimorphism is most clearly reflected in selected skeletal ratios that are more pronounced in males than in females. We found that the relative length of femur to body size was useful in gender identification, in addition to three ratios that are traditionally used, namely a distal expansion of the humerus relative to its shaft, humerus length relative to body size, and humerus length relative to femur length. Two distinctive patterns exist in allometric changes of these four ratios. The distal expansion of the humerus is exceptional in that it is equally pronounced in juvenile and adult males and therefore must have been fully established during embryonic growth. The other three features are not pronounced at birth size and subsequently become pronounced during postembryonic growth. However, males and females already show different growth trajectories at birth size even in these three. Therefore, the fate of sexually dimorphic features seems to have already been set during embryonic growth in K. hui.
Honania complicidentata and Tsiyuania simplicidentata are pareiasaur taxa based on material from the Shangshihezi Formation of Jiyuan, Henan Province, China that were earlier designated as nomina vana. Based on the study of new material, and the reexamination of old specimens, we determine that the pareiasaur material from Jiyuan represents a single species that differs from all known species from other localities. Thus, we resurrect the name H. complicidentata for the material from Jiyuan. H. complicidentata is characterized by maxillary teeth with high crowns, dentary teeth slightly posteriorly inclined compared to the dentary dorsal margin, nearly all preserved marginal teeth have a cusped cingulum on the lingual surface, and humerus without an ectepicondylar foramen. Phylogenetic analysis shows Honania is more basal than Shansisaurus and Shihtienfenia from the Sunjiagou Formation of China.
Fishes of the subfamily Dussumieriinae, also known as round herrings, are a small subgroup of the family Clupeidae inhabiting tropical and subtropical marine coastal waters. A new genus and species of round herring, Trollichthys bolcensis gen. et sp. nov., is described from the lower Eocene micritic limestone of Monte Bolca, northern Italy. This new clupeid taxon is based on seven partially complete articulated skeletons that exhibit a unique combination of features, including: two supramaxillae, edentulous jaw and palate bones, 41–42 preural vertebrae and 22–24 pleural ribs, pleural ribs-preural vertebrae ratio ranging 0.52–0.57, five or six supraneural bones, dorsal-fin origin located at about mid-length of the body, dorsal fin with about 16 rays, two postcleithra, pelvic-fin insertion slightly behind the dorsal-fin origin, and pelvic fin with eight rays. Trollichthys bolcensis shares several features with the extant round herring genus Spratelloides. However, because of its unique combination of features, Trollichthys bolcensis cannot be confidently assigned to any of the extant dussumieriine lineages and present evidence does not favour any particular sister-group relationship.
Anatomical, systematic, and paleobiogeographical data on the Devonian antiarchs from Belgium are reviewed, updated and completed thanks to new data from the field and re-examination of paleontological collections. The material of Bothriolepis lohesti is enhanced and the species redescribed in more detail. An undetermined species of Bothriolepis is recorded from the Famennian of Modave (Liège Province), one species of Asterolepis redescribed from the Givetian of Hingeon and another one described from the Givetian of Mazy (Namur Province). Grossilepis rikiki sp. nov. is recorded from the Famennian tetrapod-bearing locality of Strud (Namur Province) and from the Famennian of Moresnet (Liège Province). It is the first occurrence of Grossilepis after the Frasnian and on the central southern coast of the Euramerican continent. Its occurrence in the Famennian of Belgium may be the result of a late arrival from the Moscow Platform and the Baltic Depression, where the genus is known from Frasnian deposits. Remigolepis durnalensis sp. nov. is described from the Famennian of Spontin near Durnal (Namur Province). Except for the doubtful occurrence of Remigolepis sp. in Scotland, this is the first record of this genus in Western Europe. Its occurrence in Belgium reinforces the strong faunal affinities between Belgium and East Greenland and the hypothesis of a hydrographical link between the two areas during the Late Devonian.
A prioniodontid conodont Arianagnathus jafariani gen. et sp. nov. from the late Llandovery part of the Niur Formation of the Derenjal Mountains, East Central Iran had an apparatus bearing 3 pairs of P elements. Pa elements of its apparatus are closest to those of Icriodella sandersi (Llandovery—Wenlock boundary interval, Wales, Great Britain) in the weak development of an icrion. Due to the small sample size not all S-elements have been identified but those present are similar to those described in the Icriodella and Icriognathus apparatuses. Based on similarities with previously described apparatus Notiodella we suggest that Arianagnathus jafariani gen. et sp. nov. probably had an apparatus of 17 elements. Arianagnathus is therefore an important additional example that has potential for aiding the future revision of the palaeobiological arrangement of elements within and the phylogeny of conodont apparatuses with 3 P elements, one of which is icrion bearing. The completely known apparatus of associated Ozarkodina derenjalensis sp. nov. shows similarity to some unnamed Ozarkodina from Wales, Great Britain. Many of the conodonts found in the Llandovery part of the studied section are cosmopolitan; the new conodont species seem to have their possible closest relatives in Avalonia.
A new, unusual rhynchonellide brachiopod Jafarirhynchus alatus assigned to the newly established family Jafarirhynchidae is described from the Silurian (Telychian) of the Boghu Mountains in east-central Iran. It forms a low diversity association with the spiriferide Striispirifer? ocissimus, which exhibits well preserved calcified brachial supports. A strophic shell, well-developed ventral interarea and liberosessile mode of life make this taxon unique among Palaeozoic rhynchonellide brachiopods. In spite of a superficial similarity to spiriferides and the atrypide family Davidsonioidea, Jafarirhynchus retains the typical rhynchotrematoid cardinalia with a septalium supported by the median septum, a septiform cardinal process and long, raduliform crura. It is considered as an offshoot of the local lineage, which includes two successive species of Stegocornu (family Rhynchotrematidae) which proliferated in Central Iran and adjacent Afghanistan during Aeronian time.
Kerberellus marcouensis Devaere, Holmer, and Clausen gen. et sp. nov., originally described as Dictyonina? sp., from the Terreneuvian of northern Montagne Noire (France) is re-interpreted as the oldest relative to or member of mickwitziidlike stem-group brachiopods. We extracted 170 partial to complete phosphatic internal moulds of two types of adult and one type of juvenile disarticulated valves, rarely externally coated with phosphates, from the calcareous Heraultia Member of the Marcou Formation. They correspond to microbially infested, ventribiconvex, inequivalved, bivalved shells. The ventral interarea is bisected by a triangular sinus. The shell, most probably dominantly organic in origin, is orthogonally pierced throughout its entire thickness by radially-aligned, smooth-walled, cylindrical to hour-glass shaped canals except for the sub-apical planar field (interarea). The through-going canals of K. marcouensis are compared with brachiopods endopunctae and with canals of mickwitziid brachiopods. The absence of striations on K. marcouensis canal walls, typical of mickwitziids, implies that (i) the tubes could have been depleted of setae or; (ii) traces of the microvilli were not preserved on the tube wall (taphonomic bias) or, (iii) the tubes could have been associated with an outer epithelial follicle.
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