The armoured dissorophid Cacops from the Early Permian of Oklahoma and the exploitation of the terrestrial realm by amphibians

Cacops, one of the most distinctive Paleozoic amphibians, is part of a clade of dissorophoid temnospondyls that diversified in the equatorial region of Pangea during the Late Carboniferous and Early Permian, persisting into the Late Permian in Central Russia and China. Dissorophids were a successful group of fully terrestrial, often spectacularly armoured predators, the only amphibians apparently able to coexist with amniotes when the latter started to dominate terrestrial ecosystems. In this paper, we describe excellent new skulls from the Early Permian of Oklahoma attributed to Cacops, Cacops morrisi sp. nov. and provide for the first time detailed information about this iconic dissorophid. These specimens show anatomical and ontogenetic features that will impact on future studies on the evolution of terrestriality in tetrapods. For example, the large, posteriorly closed tympanic embayment has fine striations on an otherwise smooth surface, documenting the oldest known clear evidence for the presence of a tympanic membrane in the fossil record, a structure that is used for hearing airborne sound in extant tetrapods. The skull of C. morrisi also has several features associated with predatory behaviour, indicating that this dissorophid may have been one of the top terrestrial predators of its time.     

Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.     

The last “pelycosaur”: a varanopid synapsid from the <Emphasis Type="Italic">Pristerognathus</Emphasis> Assemblage Zone,Middle Permian of South Africa

We report on a partial varanopid skull and mandible from the Pristerognathus Assemblage Zone of the Beaufort Group, in the South African Karoo Basin, which is probably latest Middle Permian (Capitanian) in age. This mycterosaurine is not only the youngest known varanopid from the Southern Hemisphere, but it is also the youngest known “pelycosaur” (i.e., non-therapsid synapsid). Like all other members of this clade of hypercarnivores, the teeth are strongly flattened, recurved, and have finely serrated cutting edges. The anterior dentary teeth form a caniniform region, and the splenial features a foramen intermandibularis oralis, the first ever to be described in a “pelycosaur.” The last varanopids were the smallest carnivores of latest Middle Permian continental faunas. Occupation of the small carnivore guild appears to have allowed varanopids to achieve a nearly cosmopolitan distribution throughout the Middle Permian, between the great Early Permian radiation of basal synapsids and the spectacular diversification of therapsid synapsids in the Late Permian and Early Triassic.     

The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids

Little is known about the evolution of large-bodied theropod dinosaurs during the Early to mid Cretaceous in Asia. Prior to this time, Asia was home to an endemic fauna of basal tetanurans, whereas terminal Cretaceous ecosystems were dominated by tyrannosaurids, but the intervening 60 million years left a sparse fossil record. Here, we redescribe the enigmatic large-bodied Chilantaisaurus maortuensis from the Turonian of Inner Mongolia, China. We refer this species to a new genus, Shaochilong, and analyze its systematic affinities. Although Shaochilong has previously been allied with several disparate theropod groups (Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora), we find strong support for a derived carcharodontosaurid placement. As such, Shaochilong is the first unequivocal Asian member of Carcharodontosauridae, which was once thought to be restricted to Gondwana. The discovery of an Asian carcharodontosaurid indicates that this clade was cosmopolitan in the Early to mid Cretaceous and that Asian large-bodied theropod faunas were no longer endemic at this time. It may also suggest that the ascent of tyrannosaurids into the large-bodied dinosaurian predator niche was a late event that occurred towards the end of the Cretaceous, between the Turonian and the Campanian. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Evidence for high taxonomic and morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah

The fossil record of late Campanian tyrannosauroids of western North America has a geographic gap between the Northern Rocky Mountain Region (Montana, Alberta) and the Southwest (New Mexico, Utah). Until recently, diagnostic tyrannosauroids from the Southwest were unknown until the discovery of Bistahieversor sealeyi from the late Campanian of New Mexico. Here we describe an incomplete skull and postcranial skeleton of an unusual tyrannosaurid from the Kaiparowits Formation (Late Cretaceous) of Utah that represents a new genus and species, Teratophoneus curriei. Teratophoneus differs from other tyrannosauroids in having a short skull, as indicated by a short and steep maxilla, abrupt angle in the postorbital process of the jugal, laterally oriented paroccipital processes, short basicranium, and reduced number of teeth. Teratophoneus is the sister taxon of the Daspletosaurus + Tyrannosaurus clade and it is the most basal North American tyrannosaurine. The presence of Teratophoneus suggests that dinosaur faunas were regionally endemic in the west during the upper Campanian. The divergence in skull form seen in tyrannosaurines indicates that the skull in this clade had a wide range of adaptive morphotypes.     

Evidence of earliest human occurrence in Europe: the site of Pirro Nord (Southern Italy)

Some flint lithic artifacts were discovered in the fissure fillings of the well-known Pirro Nord site (Apulia, Southern Italy). The lithic industry, composed by three cores and some flakes, has been found to be associated to an Early Pleistocene vertebrate fossil assemblage. The fossil association contains a wide range of micromammals, including Allophaiomys ruffoi and Episoriculus gibberodon and large mammals including Bison degiulii and Equus altidens together with African elements as the gelada baboon Theropithecus and the saber-toothed cat Megantereon whitei. It defines the latest Villafranchian chronological unit (Pirro Nord Faunal Unit) in the Western European mammal biochronology. The lithic industry of Pirro Nord represents the oldest occurrence of the genus Homo in Europe as it is attributable to a chronological interval between 1.3 and 1.7 Ma. This supports the hypothesis that the genus Homo, with Oldowaian technology, extended its range in Europe, probably from western Asia, during the first half of the Early Pleistocene. The new discovery from Pirro Nord changes the chronology of the first arrival of hominids in Europe and offers new perspectives in the debate about the human dispersal in the Early Pleistocene.     

A molecular phylogenetic analysis of the neotropical dart-poison frog genus Phyllobates (Amphibia: Dendrobatidae)

 A phylogenetic analysis of the Neotropical dart-poison frogs, genus Phyllobates, was performed based on mitochondrial cytochrome b sequences. Members of Phyllobates from South and Central America were found to form each an evolutionary lineage. Among the South American lineage, species with uniform dorsal coloration as adults form a derived monophyletic clade. Received: 26 June 2000 / Accepted in revised form: 8 November 2000     

A new captorhinid reptile, Gansurhinus qingtoushanensis, gen. et sp. nov., from the Permian of China

Captorhinids, a clade of Paleozoic reptiles, are represented by a rich fossil record that extends from the Late Carboniferous into the Late Permian. Representatives of this clade dispersed from the equatorial regions of Laurasia into the temperate regions of Pangea during the Middle and Late Permian. This rich fossil record shows that there was an evolutionary trend from faunivorous to omnivorous and herbivorous feeding habits within this clade. The discovery of well-preserved captorhinid materials in the Middle Permian of China allows us to determine that the new taxon, Gansurhinus qingtoushanensis, gen. et sp. nov, is a member of Moradisaurinae, a clade of captorhinids with multiple tooth rows arranged in parallel. The presence of this moradisaurine in the Middle Permian of south central Asia leads us to suggest that paleogeographic changes during the Permian, with part of what is today China becoming a large peninsula of Pangea, allowed these early reptiles as well as other terrestrial vertebrates to extend their geographic ranges to this region of the Late Paleozoic supercontinent.     

New carnivorous dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid theropods

A nearly complete skeleton of the new abelisaurid Skorpiovenator bustingorryi is reported here. The holotype was found in Late Cenomanian–Early Turonian outcrops of NW Patagonia, Argentina. This new taxon is deeply nested within a new clade of South American abelisaurids, named Brachyrostra. Within brachyrostrans, the skull shortening and hyperossification of the skull roof appear to be correlated with a progressive enclosure of the orbit, a set of features possibly related to shock-absorbing capabilities. Moreover, the development of horn-like structures and differential cranial thickening appear to be convergently acquired within Abelisauridae. Based on the similarities between Skorpiovenator and carcharodontosaurid tooth morphology, we suggest that isolated teeth originally referred as post-Cenomanian Carcharodontosauridae most probably belong to abelisaurids. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos

Spinosaurids are among the largest and most specialized carnivorous dinosaurs. The morphology of their crocodile-like skull, stomach contents, and oxygen isotopic composition of the bones suggest they had a predominantly piscivorous diet. Even if close relationships between spinosaurids and Middle Jurassic megalosaurs seem well established, very little is known about the transition from a generalized large basal tetanuran to the specialized morphology of spinosaurids. Spinosaurid remains were previously known from the Early to Late Cretaceous of North Africa, Europe, and South America. Here, we report the discovery of a new spinosaurid theropod from the late Early Cretaceous Savannakhet Basin in Laos, which is distinguished by an autapomorphic sinusoidal dorsosacral sail. This new taxon, Ichthyovenator laosensis gen. et sp. nov., includes well-preserved and partially articulated postcranial remains. Although possible spinosaurid teeth have been reported from various Early Cretaceous localities in Asia, the new taxon I. laosensis is the first definite record of Spinosauridae from Asia. Cladistic analysis identifies Ichthyovenator as a member of the sub-clade Baryonychinae and suggests a widespread distribution of this clade at the end of the Early Cretaceous. Chilantaisaurus tashouikensis from the Cretaceous of Inner Mongolia, and an ungual phalanx from the Upper Jurassic of Colorado are also referred to spinosaurids, extending both the stratigraphical and geographical range of this clade.     

The earliest record of birch mice from the Early Miocene Nei Mongol,China

The earliest species of birch mouse, Sicista primus sp. nov., was recovered from the 17-Ma-old (Early Miocene) Gashunyinadege locality, central Nei Mongol, China. It is ~9 Ma older than the previous first appearance datum of Sicista in Eurasia. This study indicates that North American Macrognathomys is a synonym of Eurasian Sicista, having 12 shared dental characters. As a result, the biogeography of dipodids indicates that Asian Sicista dispersed to North America as opposed to the hypothesis that Sicista originated from the North American clade. Sicista is one of the few extant rodent genera that originated as early as the Early Miocene.     

New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birds

We report on a new Mesozoic bird, Longirostravis hani, from the Early Cretaceous Jehol Biota of northeastern China. The new taxon has a long, slender rostrum and mandible, and a small number of rostralmost teeth. Postcranial characters such as a furcular ramus wider ventrally than dorsally, a centrally concave proximal margin of the humeral head, and a minor metacarpal that projects distally more than the major metacarpal, support the placement of Longirostravis within euenantiornithine Enantiornithes, the most diverse clade of Mesozoic birds. The morphology of the skull, however, suggests that Longirostravis had a probing feeding behavior, a specialization previously unknown for Enantiornithes. Indeed, this discovery provides the first evidence in support of the existence of such a foraging behavior among basal lineages of Mesozoic birds.     

First Castorid (Mammalia,Rodentia) from the Middle Miocene of Southeast Asia

Today and in the Tertiary, the geographical distribution of castorids is limited throughout all of the northern continents. Fossils of the Castoridae genus Steneofiber are abundant in many localities of Eurasia from the late Oligocene to Pliocene period. Recently, Steneofiber fossils were discovered in two localities of northern Thailand, Mae Moh and Chiang Muan coal mines, in layers of late middle Miocene age. These discoveries represent the first records of castorids from Southeast Asia and correspond to their southernmost known range. The focus of this study is to describe this new Thai species of Steneofiber and to define its wear stages from the molar occlusal surfaces by using micro-CT scan analysis. The CT scan technique permits the analysis of the virtual occlusal surface changes from wear, allowing easier comparison to related species of Steneofiber cheek teeth without destroying the teeth. The new species, Steneofiber siamensis n. sp., can be distinguished from the other species of Steneofiber by several distinct characters, longer mesostriid on p4, presence of premesostria and metastria on P4, which are smaller than most of the other known species. The occurrence of this new castorid also supports a subtropical to tropical paleoclimate for these two localities of northern Thailand.     

A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic

Non-avian theropod dinosaurs attained large body sizes, monopolising terrestrial apex predator niches in the Jurassic–Cretaceous. From the Middle Jurassic onwards, Allosauroidea and Megalosauroidea comprised almost all large-bodied predators for 85 million years. Despite their enormous success, however, they are usually considered absent from terminal Cretaceous ecosystems, replaced by tyrannosaurids and abelisaurids. We demonstrate that the problematic allosauroids Aerosteon, Australovenator, Fukuiraptor and Neovenator form a previously unrecognised but ecologically diverse and globally distributed clade (Neovenatoridae, new clade) with the hitherto enigmatic theropods Chilantaisaurus, Megaraptor and the Maastrichtian Orkoraptor. This refutes the notion that allosauroid extinction pre-dated the end of the Mesozoic. Neovenatoridae includes a derived group (Megaraptora, new clade) that developed long, raptorial forelimbs, cursorial hind limbs, appendicular pneumaticity and small size, features acquired convergently in bird-line theropods. Neovenatorids thus occupied a 14-fold adult size range from 175 kg (Fukuiraptor) to approximately 2,500 kg (Chilantaisaurus). Recognition of this major allosauroid radiation has implications for Gondwanan paleobiogeography: The distribution of early Cretaceous allosauroids does not strongly support the vicariant hypothesis of southern dinosaur evolution or any particular continental breakup sequence or dispersal scenario. Instead, clades were nearly cosmopolitan in their early history, and later distributions are explained by sampling failure or local extinction.     

Middle-Eocene artiodactyls from Shanghuang (Jiangsu Province,Coastal China) and the diversity of basal dichobunoids in Asia

A new assemblage of basal dichobunoid artiodactyls from the middle-Eocene Shanghuang fissure fillings includes the diacodexeid Jiangsudon shanghuangensis gen. and sp. nov., a new species of the lantianine dichobunoid Elaschitotherium, Elaschitotherium crepaturus sp. nov., and an indeterminate suoid which is presently the earliest record of this clade. Diacodexeids are also represented by two forms provisionally referred to cf. Diacodexis sp. and to an indeterminate Diacodexeidae, respectively. The occurrence of diacodexeids in Shanghuang contrasts with the early and earliest middle-Eocene chronological range of the family in Europe and North America and suggests that the stratigraphic range of the family in Asia extends up to the middle Eocene. This may reflect particular habitats in coastal China that may have been relatively stable during the early and middle Eocene, thus preserving forest-dwelling artiodactyls that became extinct in the other Holarctic regions. Compared to other supposedly coeval North American, European, and Asian faunas, the Shanghuang mammalian assemblage is most similar to early Uintan faunas of North America but is also remarkable in recording forms close to taxa that are characteristic of the Wasatchian and Bridgerian North American Land Mammal Ages. The Irdinmanhan age of the Shanghuang fauna is supported by the mammalian assemblage recovered from the fissure D, but an Arshantan age cannot be completely ruled out at this point. Although the Shanghuang assemblage is biased towards preservation of small components of the mammalian fauna, the Shanghuang fauna provide an important and unique window into the Eocene diversity and early evolution of cetartiodactyls in eastern Asia.     

The Triassic dicynodont Kombuisia (Synapsida, Anomodontia) from Antarctica, a refuge from the terrestrial Permian-Triassic mass extinction

Fossils from the central Transantarctic Mountains in Antarctica are referred to a new species of the Triassic genus Kombuisia, one of four dicynodont lineages known to survive the end-Permian mass extinction. The specimens show a unique combination of characters only present in this genus, but the new species can be distinguished from the type species of the genus, Kombuisia frerensis, by the presence of a reduced but slit-like pineal foramen and the lack of contact between the postorbitals. Although incomplete, the Antarctic specimens are significant because Kombuisia was previously known only from the South African Karoo Basin and the new specimens extend the taxon’s biogeographic range to a wider portion of southern Pangaea. In addition, the new finds extend the known stratigraphic range of Kombuisia from the Middle Triassic subzone B of the Cynognathus Assemblage Zone into rocks that are equivalent in age to the Lower Triassic Lystrosaurus Assemblage Zone, shortening the proposed ghost lineage of this taxon. Most importantly, the occurrence of Kombuisia and Lystrosaurus mccaigi in the Lower Triassic of Antarctica suggests that this area served as a refuge from some of the effects of the end-Permian extinction. The composition of the lower Fremouw Formation fauna implies a community structure similar to that of the ecologically anomalous Lystrosaurus Assemblage Zone of South Africa, providing additional evidence for widespread ecological disturbance in the extinction’s aftermath.     

An opilioacarid mite in Cretaceous Burmese amber

A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opilioacarus groehni sp. nov. This ca. 99 Ma record (Upper Cretaceous: Cenomanian) represents only the third fossil example of this putatively basal mite lineage, the others originating from Eocene Baltic amber (ca. 44–49 Ma). Our new record is not only the oldest record of Opilioacarida, but it is also one of the oldest examples of the entire Parasitiformes clade. The presence of Opilioacarida—potentially Opiloacarus—in the Cretaceous of SE Asia suggests that some modern genus groups were formerly more widely distributed across the northern hemisphere, raising questions about previously suggested Gondwanan origins for these mites.     

Diversity in the later Paleogene proboscidean radiation: a small barytheriid from the Oligocene of Dhofar Governorate,Sultanate of Oman

Despite significant recent improvements to our understanding of the early evolution of the Order Proboscidea (elephants and their extinct relatives), geographic sampling of the group’s Paleogene fossil record remains strongly biased, with the first ~30 million years of proboscidean evolution documented solely in near-coastal deposits of northern Africa. The considerable morphological disparity that is observable among the late Eocene and early Oligocene proboscideans of northern Africa suggests that other, as yet unsampled, parts of Afro-Arabia might have served as important centers for the early diversification of major proboscidean clades. Here we describe the oldest taxonomically diagnostic remains of a fossil proboscidean from the Arabian Peninsula, a partial mandible of Omanitherium dhofarensis (new genus and species), from near the base of the early Oligocene Shizar Member of the Ashawq Formation, in the Dhofar Governorate of the Sultanate of Oman. The molars and premolars of Omanitherium are morphologically intermediate between those of Arcanotherium and Barytherium from northern Africa, but its specialized lower incisors are unlike those of other known Paleogene proboscideans in being greatly enlarged, high-crowned, conical, and tusk-like. Omanitherium is consistently placed close to late Eocene Barytherium in our phylogenetic analyses, and we place the new genus in the Family Barytheriidae. Some features of Omanitherium, such as tusk-like lower second incisors, the possible loss of the lower central incisors, an enlarged anterior mental foramen, and inferred elongate mandibular symphysis and diminutive P₂, suggest a possible phylogenetic link with Deinotheriidae, an extinct family of proboscideans whose origins have long been mysterious.     

A new phylogeny for basal Trechnotheria and Cladotheria and affinities of South American endemic Late Cretaceous mammals

The endemic South American mammals Meridiolestida, considered previously as dryolestoid cladotherians, are found to be non-cladotherian trechnotherians related to spalacotheriid symmetrodontans based on a parsimony analysis of 137 morphological characters among 44 taxa. Spalacotheriidae is the sister taxon to Meridiolestida, and the latter clade is derived from a primitive spalacolestine that migrated to South America from North America at the beginning of the Late Cretaceous. Meridiolestida survived until the early Paleocene (Peligrotherium) and early Miocene (Necrolestes) in South America, and their extinction is probably linked to the increasing competition with metatherian and eutherian tribosphenic mammals. The clade Meridiolestida plus Spalacotheriidae is the sister taxon to Cladotheria and forms a new clade Alethinotheria. Alethinotheria and its sister taxon Zhangheotheria, new clade (Zhangheotheriidae plus basal taxa), comprise Trechnotheria. Cladotheria is divided into Zatheria (plus stem taxa, including Amphitherium) and Dryolestida, including Dryolestidae and a paraphyletic array of basal dryolestidans (formerly classified as “Paurodontidae”). The South American Vincelestes and Groebertherium are basal dryolestidans.     

A new Early Oligocene peradectine marsupial (Mammalia)from the Burqin region of Xinjiang,China

Tertiary marsupial records are very scarce in Asia. A new peradectine marsupial, Junggaroperadectes burqinensis gen. et sp. nov., is reported from the Early Oligocene Keziletuogayi Formation in the Burqin region, Xinjiang, China. This new species is based on a single right upper M2. The tooth possesses a straight centrocrista, a characteristic of peradectines. Its main cusps lean buccally, with the paracone being smaller and lower than the metacone. The conules and stylar cusps are weakly developed. These characters distinguish J. burqinensis from Euro-American Tertiary peradectines, but they also imply a close phylogenetic relationship to Siamoperadectes and Sinoperadectes, two Asian Early Miocene peradectines.