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Metriorhynchid thalattosuchians represent the most extreme archosaurian adaptation to the marine realm. Metriorhynchids possess aquatic adaptations throughout the skeleton. These adaptations were so extensive that some have suggested that they lost the ability to move on land, yet their evolutionary timing remains unresolved. The closest relatives of the metriorhynchoids, the teleosauroids, lack these aquatic adaptations, and the earliest metriorhynchoids are known exclusively from cranial material. Here I describe a partial skull with associated forelimb elements of a new marine crocodylomorph, Zoneait nargorum, gen. et sp. nov., of Aalenian—Bajocian age from the Snowshoe Formation of east-central Oregon. Phylogenetic analysis identifies Zoneait as the sister taxon to Metriorhynchidae. It possesses a derived skull with orbits that are more laterally directed and prefrontals that are more expanded than in other basal metriorhynchoids. The preserved forelimb elements are less derived. The humerus is elongate in comparison with that of other metriorhynchoids. The ulna is slightly reduced in length and flattened but resembles the teleosauroid condition more so than the plate-like element of metriorhynchids. This suggests that marine adaptations in metriorhynchoids were acquired in mosaic fashion, with modifications of the skull preceding forelimb reduction, with this forelimb reduction occurring first in the zeugopodial elements, prior to reduction of the humerus. This evolutionary timing has important implications for the transition from nearshore ambush predation to pelagic open-marine predation in Thalattosuchia, suggesting that adaptations related to prey detection and capture preceded the locomotor adaptations that allowed these organisms to fully invade the oceans.
A new species of Sinamia is described from fluvial-lacustrine deposits in southern Gansu Province, China. The type material consists of multiple specimens that were recovered in three large blocks, two containing over 60 individuals preserved on a single bedding plane. In addition, material from two other localities is referred to this taxon. One of these contains disarticulated remains of numerous individuals preserved in a single bedding plane. Material from the other locality, from a different region of Gansu, was collected years previously and consists of a block with multiple specimens previously attributed to Sinamia zdanskyi. We interpret the preservation of specimens from all three localities as resulting from the fish becoming isolated in a restricted area, possibly made seasonally inhabitable during flooding, that then was cut off from the main channel forming a small temporary lake or pond that eventually dried out, leaving the fish to be concentrated in a steadily shrinking water body. The covering sediments that preserved these fish were possibly deposited by a subsequent flood not long after the mass-death event. The type locality includes only Sinamia, but the other two localities include remains of a small turtle. The locality containing disarticulated remains also includes isolated crocodile and dinosaur teeth, suggesting that this locality is a crevasse splay. A phylogenetic analysis including the new species supports the monophyly of Sinamiidae, but not the monophyly of Sinamia.
We describe a new species of Lower Jurassic (Hettangian/Sinemurian—Pliensbachian) ichthyosaur, Ichthyosaurus anningae, sp. nov., from west Dorset, England, U.K. The holotype of I. anningae (DONMG:1983.98), at least a subadult, is from the lower Pliensbachian Stonebarrow Marl Member (Charmouth Mudstone Formation). It is the most complete ichthyosaur known from this time interval worldwide. The species is assigned to Ichthyosaurus on the basis of humerus, forefin, and pectoral girdle morphologies. Diagnostic features of the species include a short, robust humerus with prominent processes; a femur in which the proximal width is almost as large as the distal width; and a very small femur relative to the humerus (humerus/femur ratio >1.7). Four other specimens, at least three of which are juveniles, are referred to this species. The new species may display sexual dimorphism in humeral morphology, but this cannot be confirmed due to a lack of stratigraphic information. With the recognition of I. anningae, at least three and possibly as many as five ichthyosaur species, representing three genera, are known from the Pliensbachian.
Astrapotheria is an order of extinct South American herbivores recorded throughout the continent, from the late Paleocene to middle Miocene. Here we describe Hilarcotherium castanedaii, gen. et sp. nov., an Uruguaytheriinae astrapothere from sediments of La Victoria Formation (middle Miocene) in the Tolima Department, Upper Magdalena Valley, Colombia. H. castanedaii, represented by a partial skull, mandible, and some postcranial remains, is characterized by (1) unique dental formula, with 0/3i, 1/1c, 1/1p, and 3/3 m; and (2) lower canines with subtriangular transverse section at the base. Hilarcotherium differs from the equatorial Uruguaytheriinae genera Xenastrapotherium and Granastrapotherium in (1) having three lower incisors; (2) the diagonal implantation of the lower canines; (3) lower molars with lingual cingulid; (4) the presence of the hypocone in the third upper molar; and (5) the presence of an anterolingual pocket in the fourth upper premolar. Our phylogenetic analysis supports the monophyly of the subfamilies Astrapotheriinae and Uruguaytheriinae. Within the latter, we confirm the monophyly of the neotropical clade (Hilarcotherium, Xenastrapotherium, and Granastrapotherium). H. castanedaii shows some plesiomorphic features such as the aforementioned presence of the i3 and the developed hypocone in the last upper molar. Its estimated body mass (1303 kg) is intermediate among Astrapotheriidae.
The question of what role differential jaw mechanics may have played in facilitating dietary niche partitioning among Late Cretaceous megaherbivorous dinosaurs from Laramidia is examined, using the fossil assemblage of the Dinosaur Park Formation as a test case. We use phylogenetic inference to reconstruct the mandibular adductor musculature of these animals, which we then apply to the construction of biomechanical lever models of the mandible to estimate relative bite forces. Our findings reveal predictably weak bite forces in ankylosaurs, and comparatively high bite forces in ceratopsids and hadrosaurids, both of which possessed a mechanical advantage that produced bite forces 2–3 times higher than those forces exerted by the adductor musculature. The impressive jaw mechanism shared by the last two taxa evolved in a stepwise fashion, independently in each lineage. There is tentative evidence to suggest that nodosaurids had more powerful bites than ankylosaurids, but the overall mechanical diversity among megaherbivores from the Dinosaur Park Formation is low, suggesting that differential jaw mechanics could have played only a subsidiary role in niche partitioning. Such mechanical conservatism may have may have been selected for, or it may simply reflect the limits imposed by evolutionary constraints. Regardless, mechanical disparity patterns remained stable throughout the ∼1.5 Ma evolution of the Dinosaur Park Formation megaherbivore chronofauna.
The Tithonian record from northwestern Patagonia (Neuquén Basin) documents a complete succession of lower, middle, and late Tithonian marine reptiles. From the late Tithonian—early Berriasian of Patagonia, three marine crocodyliforms have been recorded: Dakosaurus andiniensis, Cricosaurus araucanensis, and Purranisaurus potens. Although P. potens includes the first metriorhynchid skull described from Patagonia, no detailed description and/or illustrations have been published. Since the mid-20th century, several authors have discussed the validity of P. potens. Recent preparation of the type material allowed a detailed description as well as exploration of its phylogenetic relationships. We consider P. potens as a valid taxon characterized by a unique combination of characters, such as medial and posterolateral processes of the frontal forming an angle of 60°; supratemporal fossae reaching the minimum interorbital distance, with the dorsal margin of the supratemporal arch being slightly concave in lateral view; thin bony lamina projecting from the lateral and medial alveolar margins of the maxilla; occipital surface ventral to occipital condyle parallel to transverse plane of skull; well- developed crest that extends along the entire height of the supraoccipital; and orbital process of the quadrate very conspicuous and acute, horizontal, and without bony attachment. Phylogenetic analysis recovers P. potens as deeply nested within Geosaurini and, contrary to previous proposals, not closely related to ‘Metriorhynchus’ casamiquelai and ‘M.’ westermanni. Internal relationships within Geosaurini are unresolved. Purranisaurus, as a monotypic genus, is restricted to the late Tithonian—early Berriasian.
The relevant legislation regarding fossil protection in the People's Republic of China is reviewed, with particular emphasis on the aspects of illegal excavation and export of vertebrate fossils contained within the 1982 Cultural Relics Protection Law and in the current legislation as of 2010–2013. Guidelines for how to follow procedures to stay within the law are given for the assistance of international researchers wishing to work on Chinese specimens with a collaborating Chinese institution.
Gomphochelys nanus, new genus and species, is described from the earliest Wasatchian (biohorizon Wa 0; ∼55.8 Ma) of the southeastern Bighorn Basin, Washakie County, Wyoming. The new taxon represents the only known dermatemydid from the Paleocene—Eocene Thermal Maximum (PETM) interval and extends the lineage back from previous records by approximately 2 million years. Gomphochelys nanus has a thick tricarinate carapace and differs from other dermatemydids in attaining a smaller adult body size, having reduced plastral features, a posteriorly situated gular—humeral sulcus, an acarinate pygal, and thick shortened peripherals. Reexamination of previously described fossil dermatemydids suggests that the taxa Baptemys tricarinata and Kallistira costilata are junior synonyms of the middle—late Wasatchian Notomorpha garmanii, and Baptemys fluviatilis is likely a junior synonym of Baptemys wyomingensis. Gomphochelys nanus is a stem dermatemydid that is similar to N. garmanii but differs in possessing symplesiomorphies with the Late Cretaceous—Paleocene genera Agomphus and Hoplochelys. Aspects of shell morphology suggest that G. nanus was a commensurate swimmer and bottom-walker like extant Dermatemys and Staurotypus. The presence of a dermatemydid (a tropically distributed clade) in the southeastern Bighorn Basin during the PETM (when global temperatures increased by 5°C–10°C over a period of ∼60 ky) further supports the hypothesis that climate was megathermal in the region during this interval and is consistent with previously documented geographic range changes in both plants and animals. Dermatemydids disappear from the fossil record at the end of the PETM and don't reemerge until the next warming event, Eocene Thermal Maximum 2.
Fossil fishes were first collected from deposits of the Sangkarewang Formation of the Ombilin Basin in Sumatra, Indonesia, in the 1870s, but a comprehensive study of these fishes was not published until almost 50 years later. New material from these deposits was collected in 2009, which included a small anabantoid fish. This fish is not conspecific with any of the material described previously and is here named as a new genus and species. A phylogenetic analysis indicates that the new anabantoid is closely related to Osphronemus among the taxa studied, and we place it in the family Osphronemidae. However, the new fish does not appear to belong to any of the named modern subfamilies, so we leave it incertae sedis in the family. Additionally, the osphronemid subfamilies are not recovered as a monophyletic group. The anabantoid named here and another previously described from the same locality are the only fossil anabantoids known. However, the age of the Sangkarewang Formation is not confirmed and has been variously attributed to Cretaceous, Palaeocene, Eocene, and Miocene, with Eocene being the presently favored estimate. Based on the presence of an anabantoid in these deposits, an Eocene age is reasonable and is not contradicted by the known fish fauna.
This study describes new remains of Hegetotheriidae (Notoungulata), including a new species, from the Deseadan (late Oligocene) of Quebrada Fiera, Mendoza Province, Argentina. The assemblage is composed of four hegetotheriines, Prohegetotherium cf. P. sculptum, Prohegetotherium sp., Prohegetotherium schiaffinoi, and Prohegetotherium malalhuense, sp. nov., and the pachyrukhine Propachyrucos cf. P. simpsoni. The presence of Prosotherium cannot be totally discounted because lower molariforms are rather similar between the two pachyrukhine genera. The new species Prohegetotherium malalhuense, sp. nov., differs from all previously described hegetotheriines by having a lingually projecting, sharp parastyle and marked parastyle groove on the ectoloph of M2–3; talonid of m1–2 posterolabially projected; talonid of m3 with marked posterolabial groove; and the smaller size. Its phylogenetic affinities are not well resolved. Prohegetotherium is paraphyletic, with P. sculptum sister taxon to the remaining hegetotheres, and the new taxon more closely related to Hegetotherium mirabile than to P. schiaffinoi. The recognition of P. schiaffinoi and Prohegetotherium cf. P. sculptum emphasizes that the fauna from Quebrada Fiera shares elements with roughly contemporaneous Deseadan faunas from northern and southern latitudes, but important faunal particularities distinguish the region as well. The record of pachyrukhines at Quebrada Fiera more closely resembles Deseadan faunas in Patagonia than temporally correlative faunas from Bolivia and Uruguay, and indicates the presence of suitable habitats in mid-latitudes of Argentina for this hypselodont clade. Faunal affinities together with particular taxa from Quebrada Fiera appear to support a significant faunal provinciality in South America during the late Oligocene.
A new genus and species of selenosteid arthrodire is described from the Late Devonian of Morocco. Driscollaspis pankowskiorum, gen. nov. sp. nov., is defined as a selenosteid with a shallow preorbital plate embayment of the central plate, a paranuchal plate embayment of the central plate as a deep embayment determined by the lateral and posterior lobe, a central plate expanded at the contact with the pineal plate as transverse anterior border, and a suborbital plate overlapping the postorbital plate. The dermal ornamentation is tubercular, forming patches of reticular ridges clustered around sensory-line canal junctions in plate centers. The sensory-line canals are distinctly raised just above the level of the dermal ornamentation, a unique character not previously recognized in any arthrodire but seen in some ptyctodontids. A new phylogenetic hypothesis supports the monophyly of the Selenosteidae within which this new taxon is resolved, but emphasizes also unresolved relationships among aspinothoracid arthrodires. The paleogeographic distribution of the Frasnian vertebrates from Morocco and especially the seleonsteids on the western margin of Gondwana and Laurussia are discussed, and the indication for a contact of both continents during the late Frasnian is emphasized.
A new turtle from the Early Cretaceous (Aptian) Jiufotang Formation of western Liaoning, China, Perochelyslamadongensis, gen. et sp. nov., represents the first species of soft-shelled turtle from the Jehol Biota. The new taxon is diagnosed by the combination of the following characters: nuchal bone about five times wider than long; preneural absent; reversal of the orientation in the neural series at neural V; neural series fully separates costal series; costal VIIIs reduced; plastral callosities poorly developed and poorly sculpted; postorbital bar narrow, around one-fourth of orbit diameter; jugal contacting squamosal; foramen jugulare posterius separated from fenestra postotica; neural spines weakly developed on anterior cervicals; and phalangeal formula for pes 2-3-3-4-?. High levels of homoplasy make the phylogenetic relationships of the new taxon difficult to assess, and the possibility therefore exists that Perochelys lamadongensis either represents a stem or a crown trionychid. This phylogenetic uncertainty confirms that the skeletal morphology of trionychids has remained virtually unchanged for the last 120 million years.
We report on tanystropheids from the Late Triassic (middle Norian) Hayden Quarry of northern New Mexico (Chinle Formation, Hayden Quarry). These elements, consisting of isolated vertebrae and appendicular bones, represent the first unambiguously identified tanystropheid from western North America and likely the latest occurrence of the group, postdating Tanytrachelos in the eastern United States. A new phylogenetic analysis of early saurians identifies synapomorphies of tanystropheid subclades, which are recognized in the recovered vertebrae and a calcaneum. The femora are consistent with referral to Tanystropheidae. There is no clear association of the remains, however, so we refrain from erecting a new taxon. The analysis also indicates that the Hayden Quarry tanystropheid fossils belong to a newly recognized clade including the Late Triassic taxa Langobardisaurus and Tanytrachelos. Because most tanystropheid specimens are twodimensionally crushed skeletons, the Hayden Quarry tanystropheid fossils provide valuable insights into the threedimensional osteology of derived tanystropheids. The most striking feature of the Hayden vertebrae is a rugose, flattened expansion of the neural spines in the dorsal, sacral, and caudal regions, probably linked to a ligamentous bracing system. These fossils and others from Late Triassic sites in the American West suggest that tanystropheids underwent a previously unrecognized radiation in North America just prior to their extinction.
A new ginglymodian fish Beiduyu qijiangensis, gen. et sp. nov., is described from the freshwater Upper Jurassic Suining Formation of Chongqing, China. Although the fish is not completely preserved, it is most similar to Lepidotes and Scheenstia, which were once considered to be members of the Semionotiformes but are currently considered to be basal members of the Lepisosteiformes. The new fish lacks the strong dorsal ridge scales found in the Semionotiformes but has other features such as ganoid scales with anterodorsal and anteroventral processes but a reduced dorsal peg found in these basal lepisosteiform genera. With the description of this new taxon, there are three distinct ginglymodian fishes now known from southern China, all of them from Sichuan Province.
Terror birds constitute the most outstanding group of the South American Cenozoic avifauna. Considered as apex predators, their hunting skills have recently been examined, but their diversity is still unresolved. Here we report a new terror bird from the late Pliocene of Argentina, represented by the most complete articulated skeleton of one yet found. Our phylogenetic analysis places this taxon among derived phorusrhacids (Mesembriornithinae). One of the most striking cranial features of the new species is the suppression of intracranial kinesis due to the presence of an independent ossified bone that increases the structural link between the lacrimal and jugal bars, and the absence of both palatal hinges. The new species possesses ossified tracheal rings and a tracheobronchial syrinx, as well as sclerotic ossicles to adjust the shape of the cornea during its diurnal vision, and reveals a mean hearing sensitivity (∼2300 Hz) below the average for living species. The discovery of this new species provides new insights for studying the anatomy and phylogeny of phorusrhacids and a better understanding of this group's diversification.
Two new species of Batropetes are described from the Lower Rotliegend of the Saar-Nahe Basin in western Germany. Batropetes palatinus, sp. nov., is characterized by a narrow, anteriorly elongated prefrontal, a slender postfrontal without an anterolateral process, a trapezoid postorbital, and a large quadrate in exterior view. Batropetes appelensis, sp. nov., is characterized by a high angular, ‘Z’-shaped angular-dentary suture, no indentation between stem and plate of interclavicle, and scapulocoracoid without ossified coracoid. The genus Batropetes is characterized by an average length of almost 8 cm, a short trunk with 17 or 19 presacrals depending on the species, large orbit with a raised rim, a large pineal foramen, three pits on the frontal, tricuspid teeth, a toothless palate, four short robust limbs, and completely ossified pectoral and pelvic girdles (except for the coracoids of B. appelensis and B. fritschi). Batropetes belongs to the Brachystelechidae, together with Carrolla and Quasicaecilia. Cladistic analysis reveals for the monophyletic genus Batropetes that B. appelensis forms a sister group to the remaining three species in the genus, and B. niederkirchensis is the sister group to B. palatinus and B. fritschi. The functional interpretation of the appendicular skeleton and the skull morphology suggest that all Batropetes species were terrestrial, living near the lakes in which they were preserved. They appear to have fed on small arthropods in the leaf litter or in the upper zone of the soil.
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