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We describe new dental material of Eurolagus fontannesi from the Late Miocene (Early Vallesian, MN 9) lignite beds of Bełchatów A (central Poland), which is the northernmost occurrence of this lagomorph. This material considerably increases the sample size of this rare genus and species. We review the systematic position of Eurolagus and argue that it is not an ochotonid, but represents a terminal taxon of an independently evolved stem lineage of Lagomorpha. The enamel of the molars, here studied for the first time, exhibits a relatively thin decussating external and a radial internal layer, and in general resembles the palaeolagine lagomorphs (Palaeolagus) rather than the archaeolagine leporids (Hypolagus) or advanced ochotonids (Ochotona). The dental wear features of Eurolagus fontannesi indicate that the species was a sylvan mixed-feeder. This agrees with the Bełchatów A paleoenvironment, which is best approximated as the kind of mixed mesophytic forest now encountered in the Caucasus, Iran, and India. The extinction of Eurolagus was probably related to the noticeable faunal change following the increased cooling of the European climate during the Late Miocene. The first indicator of this event can be observed in Bełchatów A, marked by the immigration of Microtocricetus and the absence of Neocometes.
Early Eocene mammals from Indo-Pakistan have only recently come under study. Here we describe the first tapiromorph perissodactyls from the subcontinent. Gandheralophus minor gen. et sp. nov. and G. robustus sp. nov. are two species of Isectolophidae differing in size and in reduction of the anterior dentition. Gandheralophus is probably derived from a primitive isectolophid such as Orientolophus hengdongensis from the earliest Eocene of China, and may be part of a South Asian lineage that also contains Karagalax from the middle Eocene of Pakistan. Two specimens are referred to a new, unnamed species of Lophialetidae. Finally, a highly diagnostic M3 and a molar fragment are described as the new eomoropid chalicothere Litolophus ghazijensis sp. nov. The perissodactyls described here, in contrast to most other mammalian groups published from the early Eocene of Indo-Pakistan, are most closely related to forms known from East and Central Asia, where Eocene tapiromorphs are diverse and biochronologically important. Our results therefore allow the first biochronological correlation between early Eocene mammal faunas in Indo-Pakistan and the rest of Asia. We suggest that the upper Ghazij Formation of Pakistan is best correlated with the middle or late part of the Bumbanian Asian Land Mammal Age, while the Kuldana and Subathu Formations of Pakistan and India are best correlated with the Arshantan Asian Land Mammal Age.
Adapisoriculidae are an enigmatic group of small mammals known from the late Cretaceous of India, and from the early Palaeocene to early Eocene of Europe and Africa. Based on their primitive dental morphology, they have been classified as didelphids, nyctitheriids, leptictids, mixodectids, tupaiids, and palaeoryctids. While the latest hypothesis based on dental morphology suggests an affinity with Lipotyphla, postcranial remains indicate a close relationship with Euarchonta. Here, we present new adapisoriculid dental remains from the early Palaeocene locality of Hainin (Belgium). Adapisoriculidae are particularly abundant in Hainin, where they represent about one third of the mammalian fauna, offering new insights into both their specific and generic phylogenetic interrelationships. We describe three new species (Afrodon gheerbranti sp. nov., Bustylus folieae sp. nov. and Proremiculus lagnauxi gen. et sp. nov.) and document the previously unknown lower dentition of Bustylus marandati. The diversity of dental morphologies observed in the Hainin fauna suggests different interrelationships than previously suggested. In particular, the genus Proremiculus is considered morphologically intermediate between Afrodon and Remiculus, and the latter is no longer recognised as the sister group of Adapisoriculus. Although the highest diversity of adapisoriculids occurs in Europe, the oldest and most primitive members of the family were found in India and Africa, respectively. The geographic origin of the family could thus be located in any of these three continents, depending on the importance attributed to each of these factors. The coexistence of primitive and derived adapisoriculids at Hainin might indicate a very quick diversification in Europe, probably starting around the K-T boundary.
Femora referable to metatherians and eutherians recovered from the Bissekty Formation, Dzharakuduk, Kyzylkum Desert, Uzbekistan (90 Mya), are described. Fourteen isolated specimens were sorted based on size and morphology into groups that likely correspond to the species level or higher. Groups were then tentatively assigned to taxa known from teeth, petrosals, and/or other postcrania at these localities. One distal femur of a small arboreal metatherian, and several eutherian distal femora that probably represent zhelestids and/or zalambdalestids were identified. With the exception of one proximal femur that is similar in some aspects to the zalambdalestid Barunlestes, and a previously described multituberculate specimen, all other proximal femora from the Bissekty Formation exhibit a metatherian-like morphology. The dental record currently suggests the presence of twelve eutherian species and only one metatherian at Dzharakuduk, whereas the humeral and crurotarsal evidence supports the presence of at least two or four metatherian species, respectively. Given the sample size of the proximal femora, the morphological diversity present, and the overwhelming presence of eutherians at these localities, it is highly unlikely that the overwhelming majority of proximal femora actually represent metatherians. Therefore, this sample may suggest that the metatherian proximal femoral condition is primitive for Theria and that some eutherian taxa (probably including Zhelestidae, which are dentally most abundant at these localities) retain this condition.
The Early Cretaceous fossil record of large-bodied theropods from Asia is poor, hindering comparison of Asian predatory dinosaur faunas with those from other continents. One of the few large Asian theropod specimens from this interval is a partial skull (maxilla and dentary) from the Lianmugin Formation (?Valanginian—Albian), the holotype of Kelmayisaurus petrolicus. Most authors have either considered this specimen as an indeterminate basal tetanuran or a nomen dubium. We redescribe K. petrolicus and note that it possesses a single autapomorphy (a deep accessory groove on the lateral surface of the anterior dentary), as well as a unique combination of characters that differentiates it from other theropods, affirming its validity. A phylogenetic analysis recovers K. petrolicus as a basal carcharodontosaurid, which is supported by various features: very deep interdental plates (a carcharodontosaurid synapomorphy), fused interdental plates (present in carchardontosaurids and a limited number of other theropods), and the absence of diagnostic features of other clades of large-bodied theropods such as abelisaurids, megalosauroids, and coelurosaurs. As such, Kelmayisaurus is the second known carcharodontosaurid from Asia, and further evidence that this clade represented a global radiation of large-bodied predators during the Early—mid Cretaceous.
In modern terrestrial ecosystems, the population size of large predators is low, and a similar pattern has usually been assumed for dinosaurs. However, fossil finds of monospecific, large theropod accumulations suggest that population dynamics were more complex. Here, we report two Early Cretaceous tracksites dominated by large theropod footprints, in Querulpa Chico (Peru) and Chacarilla (Chile). The two sites correspond to distinct depositional environments—tidal basin/delta (Querulpa Chico) and meandering river (Chacarilla)—with both subject to extensive arid or semiarid palaeoclimatic conditions. Although most trackways show no preferred orientation, a clear relationship between two trackmakers is observed in one instance. This observation, coupled with the high abundance of trackways belonging to distinct large theropods, and the exclusion of tracks of other animals, suggests some degree of grouping behaviour. The presence of freshwater sources in a dry climate and perhaps social behaviour such as pair bonding may have promoted interactions between large carnivores. Further, the occurrence of these two tracksites confirms that large theropod dinosaurs, possibly spinosaurids and/or carcharodontosaurids, existed on the western margin of Gondwana as early as the earliest Cretaceous.
Postcranial skeletal pneumaticity (PSP) is present in a range of basal sauropodomorphs spanning the basal sauropodomorph—sauropod transition. We describe the PSP of five taxa, Plateosaurus engelhardti, Eucnemesaurus fortis, Aardonyx celestae, Antetonitrus ingenipes, and an unnamed basal sauropod from Spion Kop, South Africa (hereafter referred to as the Spion Kop sauropod). The PSP of Plateosaurus is apparently sporadic in its occurrence and has only been observed in very few specimens, in which it is of very limited extent, affecting only the posterior cervical vertebrae and possibly the mid dorsals in one specimen. The PSP of Eucnemesaurus, Aardonyx, Antetonitrus, and the Spion Kop sauropod consists of subfossae (fossa-within-fossa structures) that excavate the vertices of the posterior infradiapophyseal fossae of the posterior dorsal vertebrae. These subfossae range from simple shallow depressions (Eucnemesaurus) to deep, steepsided, internally subdivided and asymmetrically developed chambers (Antetonitrus). The middle and anterior dorsal vertebrae of these taxa lack PSP, demonstrating that abdominal air sacs were the source of the invasive diverticula. The presence of pneumatic features within the infradiapophyseal fossae suggest that the homologous fossae of more basal saurischians and dinosauriforms were receptacles that housed pneumatic diverticula. We suggest that it is probable that rigid non-compliant lungs ventilated by compliant posterior air sacs evolved prior to the origination of Dinosauria.
A partial ornithocheirid, representing a rare example of a pterosaurian body fossil from the Nova Olinda Member of the Crato Formation, NE Brazil, is described from the collections of the State Museum of Natural History, Karlsruhe. While similar in preservation and taphonomy to Arthurdactylus conandoylei, it is distinguished by slight differences in biometric ratios, but the absence of a skull prevents closer identification. Mostly complete body fossils belonging to ornithocheiroid pterosaurs appear to be relatively more abundant in the younger Romualdo Member of the Santana Formation, making the described specimen one of only two well documented ornithocheiroids known from the Nova Olinda Lagerstätte.
The extant venomous Gila monster and beaded lizards, species of Heloderma, live today in southwestern USA and south along the Pacific coastal region into Central America, but their fossil history is poorly understood. Here we report helodermatid osteoderms (dermal ossicles) from the late Miocene-early Pliocene Gray Fossil Site, eastern Tennessee USA. Twenty-three species of mammals are known from the fauna including abundant Tapirus polkensis, as well as fishes, anurans, salamanders, turtles, Alligator, birds, and snakes. Beaded lizards belong to the Monstersauria, a clade that includes PrimadermaParadermaGobiderma Helodermatidae (Estesia, Eurheloderma, Lowesaurus, and Heloderma). Osteoderms of lizards in this clade are unique within Squamata; they typically are circular to polygonal in outline, domed to flat-domed in cross-section, have a vermiculate surface texture, are not compound structures, and do not have imbricate surfaces as on many scincomorph and anguid lizards. We review and characterize the osteoderms of all members of Monstersauria. Osteoderms from the cranium, body, and limbs of Heloderma characteristically have a ring-extension (bony flange) at least partly surrounding the dome. Its presence appears to be a key character distinct to all species of Heloderma, consequently, we propose the presence of a ring-extension to be an apomorphy. Three osteoderms from the Gray Fossil Site range from 1.5 to 3.0 mm in diameter, have the circular shape of helodermatid osteoderms with a domed apical surface, and have the ring-extensions, permiting generic identification. Macrobotanical remains from the Gray Fossil Site indicate an oak-hickory subtropical forest dominated by Quercus (oak) and Carya (hickory) with some conifer species, an understorey including the climbing vines Sinomenium, Sargentodoxa, and Vitis. Plant and mammal remains indicate a strong Asian influence.
The description of a nearly complete skull from the late Albian of northwestern France reveals previously unknown anatomical features of Platypterygius hercynicus, and of European Cretaceous ichthyosaurs in general. These include a wide frontal forming the anteromedial border of the supratemporal fenestra, a parietal excluded from the parietal foramen, and the likely presence of a squamosal, inferred from a very large and deep facet on the quadratojugal. The absence of a squamosal has been considered as an autapomorphy of the genus Platypterygius for more than ten years and has been applied to all known species by default, but the described specimen casts doubt on this putative autapomorphy. Actually, it is shown that all characters that have been proposed previously as autapomorphic for the genus Platypterygius are either not found in all the species currently referred to this genus, or are also present in other Ophthalmosauridae. Consequently, the genus Platypterygius must be completely revised.
A new neotenic salamander, Seminobatrachus boltyschkensis gen. et sp. nov., is described based on 14 skeletons of late Paleocene—early Eocene age preserved on drill core slabs from the Cherkassy Region, central Ukraine. The new taxon is diagnosed by the following unique combination of characters: dorsal process of premaxilla posteriorly elongate and overlaps frontal; maxilla greatly reduced in size; parietal—squamosal contact absent; vomerine tooth row long and parallel to maxillary arcade; pterygoid has long anterior process; quadrate ossified; marginal and palatal teeth pedicellate; trunk vertebrae amphicoelous, each having a subcentral keel, anterior basapophysis, and spinal nerve foramina; ribs bicipital; carpals and tarsals unossified; and phalangeal formulae of 2-2-3-2 and 2-2-3-4-2 for manus and pes, respectively. Phylogenetic analysis nests S. boltyschkensis within Urodela (i.e., crown-clade salamanders), but its exact phylogenetic position is equivocal, resolving in one of three ways: (1) in an unresolved trichotomy with Salamandra and (Ambystomatidae (DicamptodonRhyacotriton)) (results obtained in NONA v. 2.0, with the WINCLADA v. 1.00.08 interface; the parsimony ratchet (island hopper) algorithm), (2) as a sister taxon of (Salamandra (Ambystomatidae (DicamptodonRhyacotriton))) clade (results obtained in TNT v. 1.1; the implicit enumeration search algorithm) or (3) as a sister taxon of Ambystomatidae (results obtained in PAUP v. 4.0b10; the branch-and-bound search algorithm).
The new neopterygian fish taxon Luoxiongichthys hyperdorsalis gen. et sp. nov. is established on the basis of five specimens from the second member of the Guanling Formation (Anisian, Middle Triassic) from Daaozi Quarry, Luoping, Yunnan Province, Southwest China. The new taxon is characterized by the following characters: triangular body outline with a distinct apex located between skull and dorsal fin; free maxilla; slender preopercular almost vertical; three suborbitals; at least eight strong branchiostegals with tubercles and comb-like ornamentation on the anterior margin; clavicles present; two postcleithra; ganoid scales covered by tubercles and pectinate ornamentation on the posterior margin with peg-and-socket structure; hemiheterocercal tail slightly forked. Comparison with basal actinopterygians reveals that the new taxon has parasemionotid-like triangular symplectics, but a semionotid opercular system. Cladistic analysis suggests that this new genus is a holostean, and either a basal halecomorph or basal semionotiform.
New representatives of a stem group Paoliida attributed to family Paoliidae (Insecta: Protoptera) are described from the Upper Carboniferous (Langsettian) sphero-sideritic concretions of the Upper Silesian Coal Basin (USCB) in Poland. Zdenekia silesiensis sp. nov. is based on forewing venation and supplemented by material of isolated hindwing similar in venation pattern. Darekia sanguinea gen. et sp. nov. differs from all other paoliid genera by the presence of a short contact between veins MP and CuA behind the division CuA and CuP. Composition of insect fauna exhibits high abundance of paoliid insects in the early Late Carboniferous ecosystems known also from other European localities such as Hagen Vorhalle in Ruhr Basin (Germany), and South Limbourg (Belgium and the Netherlands). It is the first record of true paoliids from the Polish part of paralic USCB supplementing a single historical record of Stygne roemeri considered as a taxon closely related to Paoliidae. The high abundance of paoliid insects from sphero-sideritic concretions in Sosnowiec and coal deposits previously known from the Czech part of Upper Silesian Coal Basin indicates considerable similarity of both faunas supported as well by their close stratigraphical correlation. Morphology of basal wing parts with remnants of articular sclerites preserved supports neopteran relationships of paoliids. Discovery of the first paoliid immature wing is reported suggesting similar living habitat for larvae and adults.
New species of ostracods from the Tremadocian in the northwest region of Argentina are described. These are among the earliest well-documented records of ostracods, which shed new light onto the early diversification of the group. The described fauna consists of seven species, five of which are new: Saltite uchuy sp. nov., Saltite kuraq sp. nov., Conchoprimitia? iglesiasi sp. nov., Orechina violetae sp. nov., and Orechina catalinae sp. nov. The fauna consists primarily of soanellids, a non-dimorphic family of palaeocopids, and of binodicopids. One factor leading to diversification of the group in this region may have been the complex configuration of the northwest basin, which had a restricted pattern of circulation. The distribution of the first ostracods is largely restricted to Gondwana and peri-Gondwana regions. Accordingly, it is possible to envision that the origin or at least an important radiation of the group was centred in this region. Both the Soanellidae and the genus Orechina would have originated in Gondwana and would have become widespread later during the Middle Ordovician. The Tremadocian fauna located in Argentina show significant affinities with fauna located in the warm-water setting of the east Gondwana, mainly in Australia and China.
Jurassic rhynchonellide brachiopods from the Jordan Valley are herein revised and new taxa are added to the faunal list. In this study of Jurassic rhynchonellides from Wadi Zarqa, northwestern Jordan, we recognize the following taxa: Eurysites rotundus, Cymatorhynchia quadriplicata, Daghanirhynchia triangulata, D. angulocostata, Pycnoria magna, Schizoria elongata, and Schizoria cf. intermedia. The following new taxa are described: Daghanirhynchia susanae sp. nov. and Amydroptychus markowitzi sp. nov. The Middle Jurassic Mughanniyya Formation of northwest Jordan is dominated by limestone beds. The sedimentary environment is interpreted as neritic, light, and nutrient-rich resulting in high faunal diversity. The high rhynchonellide endemism of this fauna is yet another confirmation of pronounced Middle Jurassic endemism along the southern Tethyan margin of the Ethiopian Province. Brachiopods of the Jordanian Mughanniyya Formation can be correlated with the fauna of the Aroussiah Formation in Sinai and the Zohar and Matmor formations in Southern Israel.
A current paradigm accepts the presence of weakly biomineralized animals only, barely above a low metazoan grade of organization in the terminal Neoproterozoic (Ediacaran), and a later, early Cambrian burst of well skeletonized animals. Here we report new assemblages of primarily calcareous shelly fossils from upper Ediacaran (553–542 Ma) carbonates of Spain and Russia (Siberian Platform). The problematic organism Cloudina is found in the Yudoma Group of the southeastern Siberian Platform and different skeletal taxa have been discovered in the terminal Neoproterozoic of several provinces of Spain. New data on the morphology and microstructure of Ediacaran skeletal fossils Cloudina and Namacalathus indicate that the Neoproterozoic skeletal organisms were already reasonably advanced. In total, at least 15 skeletal metazoan genera are recorded worldwide within this interval. This number is comparable with that known for the basal early Cambrian. These data reveal that the terminal Neoproterozoic skeletal bloom was a real precursor of the Cambrian radiation. Cloudina, the oldest animal with a mineralised skeleton on the Siberian Platform, characterises the uppermost Ediacaran strata of the Ust'-Yudoma Formation. While in Siberia Cloudina co-occurs with small skeletal fossils of Cambrian aspect, in Spain Cloudina-bearing carbonates and other Ediacaran skeletal fossils alternate with strata containing rich terminal Neoproterozoic trace fossil assemblages. These finds treated together provide a possibility to correlate transitional Neoproterozoiclower Cambrian strata around the world. Such a correlation concurs with available isotope and radiometric data and indicates that typical Ediacaran shelly fossils have not crossed the Precambrian—Cambrian boundary.
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