The first record of a nyctiphruretid parareptile from the Early Permian of North America,with a discussion of parareptilian temporal fenestration

The Richards Spur Locality of Oklahoma, USA, long known for its highly diverse Early Permian terrestrial tetrapod assemblage, is particularly interesting for the presence of many endemic taxa. The parareptilian component of the assemblage, rare members of other Early Permian communities, is especially diverse at Richards Spur, consisting of six species. The newest parareptile, A byssomedon williamsi gen. et sp. nov. , consists of an articulated left jaw and various disarticulated cranial and postcranial elements. A new phylogenetic analysis of parareptiles, based on an updated modified data matrix revealed that Ab . williamsi is a member of the small clade Nyctiphruretidae. This makes Ab . williamsi the first and oldest nyctiphruretid, a clade of parareptiles otherwise known from the Middle and Late Permian of Russia, extending the age of the clade back into the Early Permian. This discovery also raises the possibility that nyctiphruretids may have dispersed from western Laurasia to eastern Laurasia. The characteristic jugal morphology of Ab . williamsi shows that it would have possessed a slender, deep, temporal emargination. The current topology of Parareptilia indicates that there was considerable variability in the patterns of lateral temporal openings amongst the various members of this clade, suggesting that there may have been multiple, independent modifications of this region of the skull. © 2014 The Linnean Society of London     

A new Early Permian reptile and its significance in early diapsid evolution

The initial stages of evolution of Diapsida (the large clade that includes not only snakes, lizards, crocodiles and birds, but also dinosaurs and numerous other extinct taxa) is clouded by an exceedingly poor Palaeozoic fossil record. Previous studies had indicated a 38 Myr gap between the first appearance of the oldest diapsid clade (Araeoscelidia), ca 304 million years ago (Ma), and that of its sister group in the Middle Permian (ca 266 Ma). Two new reptile skulls from the Richards Spur locality, Lower Permian of Oklahoma, represent a new diapsid reptile: Orovenator mayorum n. gen. et sp. A phylogenetic analysis identifies O. mayorum as the oldest and most basal member of the araeoscelidian sister group. As Richards Spur has recently been dated to 289 Ma, the new diapsid neatly spans the above gap by appearing 15 Myr after the origin of Diapsida. The presence of O. mayorum at Richards Spur, which records a diverse upland fauna, suggests that initial stages in the evolution of non-araeoscelidian diapsids may have been tied to upland environments. This hypothesis is consonant with the overall scant record for non-araeoscelidian diapsids during the Permian Period, when the well-known terrestrial vertebrate communities are preserved almost exclusively in lowland deltaic, flood plain and lacustrine sedimentary rocks.     

A new Lower Permian trematopid (Temnospondyli: Dissorophoidea) from Richards Spur,Oklahoma

A new trematopid amphibian, Acheloma dunni, is reported based on excellently preserved cranial and postcranial elements recovered from the Lower Permian fissure fill deposits of the Dolese Brothers Co. limestone quarry near Richards Spur, Oklahoma. The new taxon is characterized by lateral exposures of the palatine (l.e.p.) and ectopterygoid (l.e.e.), which are clearly visible externally and completely enclosed within the suborbital elements. This large, terrestrial carnivore may represent the top predator of the Richards Spur assemblage. A phylogenetic analysis including 12 taxa and 53 cranial characters yielded a single most parsimonious tree, placing Ach. dunni within the monophyletic Trematopidae as the sister taxon to Acheloma cumminsi. Furthermore, the analysis includes the enigmatic Ecolsonia and Actiobates within Trematopidae, forming a clade with the Upper Pennsylvanian Anconastes and the Lower Permian Tambachia. The study comprehensively analyses all valid and aberrant forms of Trematopidae. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 789–815.     

Comparative anatomy and osteohistology of hyperelongate neural spines in the sphenacodontids Sphenacodon and Dimetrodon (Amniota: Synapsida)

Osteohistological investigations of hyperelongate vertebral spinous processes (neural spines) are presented to elucidate previously unknown aspects of dorsal sail form and function in two, closely related genera of “sail‐backed” synapsids: Sphenacodon and Dimetrodon. Although recent and classic surveys of bone histology in extinct vertebrates have sampled the genus Dimetrodon, new sectioning of Sphenacodon material allows a comparative analysis of these structures among Sphenacodontidae for the first time. Variability within the histological organization of the neural spine is assessed by examining multiple regions along its length, and implications for soft tissue correlates, growth and mechanics are considered here. Both genera exhibit extensive parallel‐fibered and fibrolamellar bone, in addition to lamellar bone. Several features vary along the length of the spine in each species. Muscle scars and extensive Sharpey's fibers are present at the base of the spine; no scars and fewer fibers are manifested ～55–60 mm above the zygapophyses in mature individuals. The distal cortex of the spine does not exhibit greater vascularity than the proximal region in either genus. However, both genera manifest distinct vascular grooves of variable size along the distal periosteal surface, some of which become incorporated into the distal cortex. The observed histovariability appears to record the transition from the proximal (epaxial muscle embedded) to the distally protruding portion of the spine. These observations and independent pathological evidence support the existence of a short dorsal crest in Sphenacodon and possibly other basal sphenacodontids. Although the thermoregulatory capacity of such a crest remains uncertain, developmental and mechanical features are readily interpretable and are discussed with respect to the origins and early evolution of the dorsal sail in sphenacodontid synapsids. J.Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Early Miocene gastropod and ectothermic vertebrate remains from the Lesvos Petrified Forest (Greece)

The Lesvos Petrified Forest (western Lesvos, Greece) has long been famous for its plant fossils. Recently, one proboscidean (from the Gavathas locality) and seven micromammalian species (from the Lapsarna locality) were described; these were the first animals to be found in the Early Miocene subtropical forest. For the first time, a fauna of gastropods and ectothermic vertebrates from the Lapsarna locality is now available. This fauna derives from lacustrine sediments under the pyroclastic material that contains the petrified plants. Based on fragmented mollusc remains, isolated fish pharyngeal teeth and utricular otoliths (lapilli), fragmented amphibian vertebrae and a tooth-bearing element, and reptile fragmented dentaries, teeth, osteoderms and vertebrae, the presence of eight freshwater and three terrestrial gastropod species, three freshwater cyprinid species, and two amphibian and five reptile taxa has been confirmed. Stratigraphical and radiometric data suggest an age older than 18.4 ± 0.5 Ma (latest Early Miocene), in good agreement with the faunal composition. This paper is the first report of the concurrent presence of three cyprinid fish species in a Greek Early Miocene locality, as well as the first documentation of an Early Miocene proteid amphibian in southeastern Europe. The present findings represent one of the best-documented Early Miocene gastropod and fish faunas in the Aegean/southern Balkans, thus adding to our knowledge of Early Miocene amphibians and reptiles from that region and providing valuable information on the local subtropical ecosystem.     

The cranial anatomy and relationships of Secodontosaurus, an unusual mammal-like reptile (Synapsida: Sphenacodontidae) from the early Permian of Texas

The cranial anatomy of the Early Permian sphenacodontid synapsid Secodontosaurus is redescribed. There is no evidence for recognition of more than one species of Secodontosaurus, and S. willistoni is declared a junior subjective synonym of S. obtusidens. Numerous derived characters (autapomorphies), mainly related to the unusually slender and slightly elongated skull, distinguish Secodontosaurus. The cranial specializations are interpreted as an adaptation toward a feeding strategy which involved preying upon small tetrapods that attempted to avoid capture by hiding in crevices and burrows. A cladistic analysis supports the following hypotheses of relationships: (a) the Sphenacodontidae is the nearest sister taxon of Therapsida; (b) Secodontosaurus, Sphenacodon, Ctenospondylus and Dimetrodon share a more recent common ancestor with one another than any of them do with Haptodus, and (c) Secodontosaurus shares a more recent common ancestor with Dimetrodon than with either Sphenacodon or Ctenospondylus. The latter hypothesis suggests that the sphenacodont subfamily 'Sphenacodontinae' is paraphyletic and, therefore, invalid.     

Late Maastrichtian small-sized herpetofauna from Valencia province,eastern Spain

A Late Maastrichtian microvertebrate assemblage which includes amphibian remains was recovered from continental deposits of the palaeontological site of La Solana, Valencia Province, Spain. This site is composed of variegated mudstones, pedogenically modified, interbedded with fluvial sand bodies and freshwater limestones lenses, and has also yielded plant debris, freshwater and oligohaline invertebrates, abundant fish remains (isolated bones and scales), turtle plates and archosaur bones. This fossil assemblage, dominated by aquatic forms, also includes semiaquatic and terrestrial elements, and may be interpreted as the palaeofauna of a wetland environment with terrestrial environs. The new material described here consists of fragmentary remains of an indeterminate albanerpetontid, a salamandrid and two anuran taxa (an alytid and a palaeobatrachid). The amphibians from La Solana are typical Laurasiatic taxa. This faunal association shows broad similarities to other coeval faunas of the Iberian Peninsula and contrasts with the Upper Campanian–Lower Maastrichtian sites where Gondwanan elements are frequent.     

A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes?

Living species of mammals, crocodiles and most species of birds exhibit parental care, but evidence of this behaviour is extremely rare in the fossil record. Here, we present a new specimen of varanopid 'pelycosaur' from the Middle Permian of South Africa. The specimen is an aggregation, consisting of five articulated individuals preserved in undisturbed, close, lifelike, dorsal-up, subparallel positions, indicating burial in 'life position'. Two size classes are represented. One is 50% larger than the others, is well ossified, has fused neurocentral sutures and is distinguished by a coat of dermal ossifications that covers the neck and shoulder regions. We regard this individual to be an adult. The remaining four skeletons are considered to be juveniles as they are approximately the same size, are poorly ossified, have open neurocentral sutures and lack dermal ossifications. Aggregates of juvenile amniotes are usually siblings. Extant analogues of adult and juvenile groupings suggest that the adult is one of the parents, leading us to regard the aggregation as a family group. The Late Middle Permian age of the varanopid family predates the previously known oldest fossil evidence of parental care in terrestrial vertebrates by 140 Myr.     

An Early Devonian arthropod fauna from the Windyfield cherts, Aberdeenshire, Scotland

New terrestrial and freshwater arthropods are described from the Windyfield cherts, a suite of silicified sinters deposited 700m north‐east of the Rhynie cherts and part of the same Early Devonian hot‐spring complex. The diverse assemblage consists of Heterocrania rhyniensis (Hirst and Maulik, 1926a), here recognized as a euthycarcinoid; scutigeromorph centipede material assigned to Crussolum sp.; the crustacean Lepidocaris; trigonotarbid arachnids; a new arthropod of myriapod affinities named Leverhulmia mariae gen. et sp. nov.; and the distinctively ornamented arthropod cuticle of Rhynimonstrum dunlopi gen. et sp. nov. The Leverhulmia animal preserves gut content identifying it as an early terrestrial detritivore. Abundant coprolites of similar composition and morphology to the gut contents of the euthycarcinoid crowd the matrix. Chert texture, faunal associations, and study of modern analogues strongly suggest that the terrestrial arthropods were ubiquitous Early Devonian forms with no particular special adaptation to localized conditions around the terrestrial hot‐spring vents. The aquatic arthropods represent biota from ephemeral cool‐water pools in the vicinity of the hot‐spring vents.     

Mio-Pliocene rodent assemblages from Alcoi Forn (Alcoy Basin,Eastern Spain). Biostratigraphical and palaeoclimatical inferences

In this work, we describe four new micromammal sites in the northern side of the Gormaget ravine, in the Alcoy Basin (Spain): AF-1’06, AF-1’07, AF-1A and AF-2. Based on the study of the faunal remains from these localities, we infer a latest Turolian-earliest Ruscinian age for AF-1’06 and AF-1A, and an earliest Ruscinian age for AF-1’07 and AF-2. The species assemblage of the locality AF-1’06, the only one which have yielded a sufficient number of remains to perform a palaecological analysis, shows warm and dry conditions near the Mio-Pliocene boundary in the Alcoy Basin. These data show a reduction of dry and warm indicators from older to younger localities in the Alcoy Basin, suggesting a change to colder and more humid conditions during the Early Pliocene.     

The oldest Eurasian hominoid.

Engelswies is an early Miocene vertebrate locality in southern Germany with a rich assemblage of terrestrial mammals, invertebrates and fossil plants. It is dated to 16.5-17.0 Ma based on magnetostratigraphy, biostratigraphy and lithostratigraphy, and includes among the faunal remains a hominoid upper molar fragment, the oldest hominoid so far identified from Europe. The evidence from Engelswies suggests that hominoids arrived in Eurasia about 17 Ma, roughly contemporaneously with pliopithecoids and Deinotherium, and before the last marine transgression to isolate Eurasia from Africa. Thick enamel and low dentine penetrance may have been key adaptations that contributed to the success of hominoids of dentally modern aspect in western Eurasia and ultimately to their ability to spread to eastern Eurasia and Africa in the middle and late Miocene.     

The oldest rhynchosaur from Argentina: a Middle Triassic rhynchosaurid from the Chañares Formation (Ischigualasto–Villa Unión Basin,La Rioja Province)

Rhynchosaurs first appear in the Early Triassic fossil record and flourished during the late Carnian as the dominant members of several assemblages worldwide. In Argentina, the rhynchosaur record is currently restricted to the Ischigualasto Formation of late Carnian–earliest Norian age. Recent fieldwork in the new locality of Brazo del Puma, in the lowermost levels of the Chañares Formation, yielded three rhynchosaur tooth-bearing bones, which were collected five metres above the contact with the underlying Tarjados Formation. The most complete specimen is the posterior end of the alveolar region of a left dentary. The dentary possesses densely packed tooth rows on the lingual surface and medial half of the occlusal surface, showing longitudinal Zahnreihen. The teeth of the occlusal surface are worn flat and those of the lingual surface are organized in multiple rows, supporting the referral of the specimen to Rhynchosauridae. In addition, the dentary teeth are conical to mesiodistally compressed, resembling the condition observed in hyperodapedontines. The rhynchosaur remains reported here are the oldest collected in Argentina and among the oldest in South America, together with an unnamed form from Brazil. The new rhynchosaur specimens come from levels in which dicynodonts are numerically dominant, whereas cynodonts are considerably less abundant. Accordingly, the specimens reported here bolster faunal differences within the Chañares Formation and add a new faunal component to this already diverse vertebrate assemblage.     

Aubenas-les-Alpes (S-E France). Part III – Last and final part of the mammalian assemblage with some comments on the palaeoenvironment and palaeobiogeography

The study of the new material from Aubenas-les-Alpes makes it possible to add to the faunal list taxa hitherto unknown in this locality (Neurogymnurus cayluxi, Issiodoromys minor, Eomys minor, Pseudocricetodon cf. hausi, Pseudocricetodon aff. philippi). New specimens of the shrew Srinitium marteli increase our knowledge of this rare insectivore. The characteristics of the theridomorph I. minor confirm, as does the whole rodent fauna, the correlation with the MP25 level (Late Rupelian). The assemblage of Aubenas-les-Alpes indicates a rather open and dry environment, in accordance with the Early Oligocene climatic context, but some differences compared to other localities of the same age are also noticed. Finally the mammalian assemblage suggests an ongoing biogeographic differentiation at European scale during the Late Rupelian.     

Tetrapod postural shift estimated from Permian and Triassic trackways

Abstract:  The end-Permian mass extinction, 252 million years (myr) ago, marks a major shift in the posture of tetrapods. Before the mass extinction, terrestrial tetrapods were sprawlers, walking with their limbs extended to the sides; after the event, most large tetrapods had adopted an erect posture with their limbs tucked under the body. This shift had been suspected from the study of skeletal fossils, but had been documented as a long process that occupied some 15–20 myr of the Triassic. This study reads posture directly from fossil tracks, using a clear criterion for sprawling vs erect posture. The track record is richer than the skeletal record, especially for the Early and Middle Triassic intervals, the critical 20 myr during which period the postural shift occurred. The shift to erect posture was completed within the 6 myr of the Early Triassic and affected both lineages of medium to large tetrapods of the time, the diapsids and synapsids.     

A second varanopseid skull from the Upper Permian of South Africa: implications for Late Permian 'pelycosaur' evolution

Late Permian terrestrial faunas of South Africa and Russia are dominated taxonomically and ecologically by therapsid synapsids. On the basis of a single specimen from the Upper Permian of South Africa, the varanopseid Elliotsmithia longiceps is the sole basal synapsid ('pelycosaur') known from Gondwana. Recent fieldwork in the Upper Permian of South Africa has produced a second varanopseid specimen that is referrable to Elliotsmithia . Data from both this specimen and the holotype suggest that Elliotsmithia forms a clade with Mycterosaurus from the Lower Permian of North America and Mesenosaurus from the Upper Permian of Eastern Europe. That postulate is supported by the three most parsimonious trees discovered in a new analysis of varanopseid phylogeny. However, the available data cannot resolve the interrelationships of these three genera. The new phylogenetic results contrast with earlier work identifying Elliotsmithia as the basal member of a clade that includes the North American taxa Aerosaurus , Varanops , and Varanodon . The new trees reduce the stratigraphic debt required by the latter scenario, and the one with the least stratigraphic debt identifies Elliotsmithia and Mesenosaurus as sister taxa. Two new taxa are erected, Mycterosaurinae and Varanodontinae, for the two varanopseid subclades.     

Late Carboniferous-Early Permian Tetrapod Ichnofauna from the Khenifra Basin, Central Morocco

In terms of cumulative thickness and areal extent, the Khenifra Basin is one of the most important outcrops of Late Palaeozoic red-beds in central Morocco. Macro- and microfloral remains near the centre of the 1800 m-thick succession of interbedded conglomerates, sandstones, and mudstones are considered to be of middle to late Early Permian age. Here we give the first comprehensive analysis of the vertebrate ichnofossil record from the study area, based on 17 specimens of isolated footprints and incomplete step cycles collected at three localities that are lithostratigraphically equivalent to the plant-bearing horizons. The tetrapod ichnofauna comprises tracks of the plexus Batrachichnus Woodworth - Limnopus Marsh, Ichniotherium sphaerodactylum (Pabst), Dimetropus Romer and Price, and Dromopus Marsh which can be referred to temnospondyl, diadectomorph, synapsid (“pelycosaurian”) and early sauropsid trackmakers. This clearly Euramerican footprint assemblage, including the first occurrences of Ichniotherium and Dimetropus from outside Europe and North America, indicates a Late Carboniferous to Early Permian age of the fossiliferous strata. Judging from the relatively diverse ichnofauna and flora, the Khenifra Basin must have represented a well-established terrestrial ecosystem during that period. Its habitat could be specially important for the understanding of the phylogeny and dispersal of early tetrapods, inasmuch as we are able to report on an extremely rare type of diadectomorph footprint hitherto known only from the Early Permian of central Germany.     

Evolution of the Permian and Triassic tetrapod communities of Eastern Europe

The Permo-Triassic terrestrial and freshwater tetrapod communities of Eastern Europe are reconstructed as food-webs. The Late Permian theriodont-dinocephalian community (Ocher, Mezen, Isheyevo) changes to a latest Permian theriodont-pareiasaur community (North Dvina, Vyazniki). After a major extinction, the Triassic thecodontian-dicynodont communities appear, a lystrosaurid one in the Early Triassic (Lower and ?Upper Vetluga), and a kannemeyerid one in the later Early Triassic (?Yarenga) and the Mid Triassic (Donguz, Bukobay). Similar stages are represented in the evolution of aquatic communities: the Late Permian temnospondyl community (Ocher, Isheyevo), the latest Permian chroniosuchian one (North Dvina, Vyazniki), the Lower and Middle Triassic new temnospondyl one (from Vetluga to Bukobay). The faunal changes in Eastern Europe are mirrored in other parts of the world, although there are some endemic Russian forms.     

A new record of Gobius jarosi (Teleostei,Gobiidae) from the Early Miocene of Poland with inference to paleogeography and palaeoecology of the Carpathian Basin

Abstract

Gobies (family Gobiidae) were in the past and now important components of marine ecosystems as an essential part of the food chain. However, the early fossil record of this group is relatively meager, with only scarce skeletal remains. The oldest known representative of the genus Gobius has been recently described from the Early Miocene of Czech Republic as Gobius jarosi P?ikryl & Reichenbacher, 2018. Here we present a detailed study of a well-preserved goby skeleton of the same age from the Harta locality (Poland). This specimen is assumed as belonging to Gobius jarosi based on its almost complete morphological and meristic identity with the type material from the Vá?any nad Litavou locality. Some aspects of the paleogeography and paleoecology of the Early Miocene fish assemblage from Harta with special reference to the Carpathian Basin are also discussed in this paper.     

Early Permian bioevent in the fusulinacean fauna of South China

Analysis of a large database of the stratigraphic distribution of fusulinacean Foraminifera reveals an Early Permian event of significant decline of species diversity in South China. Data from Late Carboniferous to Early Permian sections without apparent unconformity in southwest China were evaluated to determine if the apparent pattern of species disappearance was caused by bias in fossil preservation associated with Early Permian sea-level changes. Statistical analysis suggests that the Early Permian event started in the Late Sakmarian with a significant drop of species diversity in the Robustoschwagerina ziyunensis Zone and continued through the Pamirina darvasica Zone of the Artinskian and into the Brevaxina dyhrenfurthi Zone of the Early Kungarian, resulted in a total loss of about 40% species diversity in the fusulinacean fauna. The Early Permian event is the most extensive bioevent in the history of fusulinacean Foraminifera at the species level although it is less significant at the generic level. Because a similar faunal change has been found among the fusulinacean assemblages in North America and in various regions of Tethys, this event may represent a major faunal turn-over in response to the Early Permian changes in sea level and could be of a global nature. Previous recognition of this event was hampered by Early Permian unconformities in North America and other regions of Tethys.