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We describe newly recovered cranial material of Bunostegos akokanensis, a pareiasaurian reptile known from the Upper Permian Moradi Formation of northern Niger. Bunostegos is highly autapomorphic, with diagnostic cranial features including two or three hemispherical bosses located above and between the external nares; laterally projecting supraorbital ‘horn’ formed by an enlarged postfrontal; large foramen present on ventral surface of postfrontal; and hemispherical supratemporal boss located at posterolateral corner of skull roof. We addressed the phylogenetic position of Bunostegos by incorporating it into a cladistic analysis of 29 parareptilian taxa (including all 21 currently valid pareiasaurs) and 127 cranial and postcranial characters. The results of this analysis place Bunostegos as more derived than middle Permian forms such as Bradysaurus and as the sister taxon to the clade including Deltavjatia plus Velosauria. Certain characters, such as the pattern of cranial ornamentation and the size and placement of the tabulars, appear to be more similar to more derived pareiasaurs such as Elginia from Scotland and Arganaceras from Morocco, but the most parsimonious tree topology indicates that these features were evolved independently in the Nigerien form. The lack of both dicynodont herbivores and Glossopteris, combined with the presence of a giant herbivorous captorhinid, indicates a markedly different community structure in the Permian of Niger compared with those for contemporaneous southern Pangean basins (i.e., Karoo, Luangwa, Ruhuhu). The endemic tetrapod fauna of Niger supports the theory that central Pangea was biogeographically isolated from the rest of the supercontinent by desert-like conditions during Late Permian times.
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Based upon fragmentary remains of dermal armor, a new form of arandaspid fish, Ritchieichthys nibili, gen et. sp. nov., is described from subsurface core material from the Katian (Late Ordovician) Nibil Formation of the Canning Basin, Western Australia. Ritchieichthys nibili represents the first documented record of a fish from the Ordovician of the Canning Basin. Allied to the previous descriptions of arandaspsids from the Amadeus and Warburton basins of the Northern Territory and New South Wales, respectively, this record extends the paleogeographic range of arandaspids across the hypothetical Ordovician Larapintine Seaway and increases the stratigraphic range of the Order Arandaspidiformes into the Katian. The hard tissue histology of Ritchieichthys nibili confirms the presence of a cellular dentine forming the bulk of the dermal armor ornament in arandaspids, a tissue that had not been directly observed previously, and confirms the presence of largely unconnected osteocytes within the dermal bone that forms the majority of the armor.
Here we present for the first time a detailed taxonomic study of a diverse chondrichthyan fauna from the Middle Triassic of the Iberian Range (Spain). The assemblage consists of isolated remains of seven species of five non-neoselachian shark genera (Palaeobates, Hybodus, Pseudodalatias, Prolatodon, gen. nov., and Lissodus), including a new species of hybodontiform shark, Hybodus bugarensis, sp. nov. In addition, a new homalodontid genus, Prolatodon, sp. nov., is erected for the taxa ‘Polyacrodus’ bucheri and ‘Polyacrodus’ contrarius. The chondrichthyans of the Iberian Range represent a heterogeneous group from a paleogeographic point of view made up of common components of Middle Triassic shark faunas of northern Europe (Hybodus plicatilis and Palaeobates angustisimus) together with species only known previously from North America and China (Prolatodon bucheri, comb. nov., and Prolatodon contrarius, comb. nov.), as well as several ‘endemic’ taxa (Pseudodalatias henarejensis, Hybodus bugarensis, sp. nov., and Lissodus aff. L. lepagei). This fauna demonstrated adaptation for a wide diversity of feeding strategies, implying that non-neoselachian sharks dominated among the predator community of Middle Triassic coastal ecosystems of Iberia. The co-occurrence with bivalves, ammonoids, and conodonts allows us to date the chondrichthyan assemblage as ‘Longobardian’ (upper Ladinian).
Representative fossils of three extant lanternfish (Myctophidae) genera (Bolinichthys, Lampanyctus, and Myctophum) from the collections of Natural History Museum of Los Angeles County were identified, based on a combination of osteological and photophore characters. These specimens come from the upper Miocene Modelo Formation (Mohnian-Delmontian) of California, and represent the first skeletal records of Bolinichthys (otherwise identified as otoliths from the Late Miocene of Italy) and Myctophum body fossils in North America, and confirm that both modern myctophid subfamilies exhibited a cosmopolitan distribution by the late Miocene.
The Miocene deposits of the Agnevo Formation cropping out along the western coast of Sakhalin Island, Russia, are widely known for the well-preserved fossil remains of marine organisms, including a diverse assemblage of primarily benthic and demersal teleost fishes. †Sakhalinia multispinata, gen et sp. nov., a new cottoid fish from the Miocene (Serravallian-Tortonian) finely laminated sandstone of the Agnevo Formation, is described herein based on a single, nearly complete specimen in part and counterpart. Several morphological features indicate that †Sakhalinia multispinata is a basal member of the cottoid family Hexagrammidae. Within hexagrammids, †Sakhalinia multispinata exhibits a unique combination of features and is primarily typified by slender and recurved jaw teeth, 44 (22 22) vertebrae, 14 (6 8) principal caudal fin rays, a moderate consolidation of the caudal skeleton, an entire dorsal fin with 25 thick spines and 13 soft rays, anal fin with four strong spines and 12 soft rays, and ctenoid scales. The elongated, slender, and recurved jaw teeth displayed by †Sakhalinia multispinata suggest that this hexagrammid fish was specialized to feed mainly on small crustaceans.
Hugues-Alexandre Blain, Jordi AgustÍ, Juan Manuel López-García, Hamid Haddoumi, Hassan Aouraghe, Kamal El Hammouti, Alfredo Pérez-González, Maria Gema Chacón, Robert Sala
The fossil amphibians and squamate reptiles from the late Miocene (Vallesian) of Guefaït-1 are described for the first time. The herpetofaunal assemblage is composed of Discoglossinae indet., Lacertidae indet., Dopasia sp. (Anguidae), Colubridae s. 1. indet., and Naja cf. antiqua (Elapidae). This herpetofaunal assemblage is less diverse than the Moroccan herpetofauna from the middle and early late Miocene mainly because of the absence of Afro-tropical taxa, probably due to increasing aridity; likewise, it is less diverse than the Plio-Pleistocene herpetofauna, which was enriched by the entry of some European taxa during the Messinian Crisis. The presence of a discoglossine frog, different from all existing European and North African genera, in the late Miocene of Guefaït-1 suggests that the diversity of this group in the Miocene of North Africa may have been greater than previously thought. The occurrence of an anguid lizard of the genus Dopasia in the Miocene of Morocco is confirmed and may constitute the earliest record of the genus for Africa. The range of the cobra Naja antiqua, until now only known from the middle Miocene (Mellalesian) of Beni Mellal, Morocco, is extended; this constitutes the latest record of the species.
Utatsusaurus hataii is an early member of Ichthyopterygia from the Osawa Formation (Lower Triassic) of Miyagi Prefecture, Japan. Although the holotype of the species lacks a well-preserved cranium, UHR 30691 preserves a near-complete skull and may be referred to Utatsusaurus based on a similar geologic occurrence and, most importantly, the following shared features with the holotype: subthecodont tooth implantation; the presence of distomesially compressed tooth crowns; a humerus that is longer than wide; and an ulnar facet on the humerus that is as wide as the radial facet. The first complete description of the skull of UHR 30691 and the incorporation of data derived from it into a phylogenetic analysis retrieves a monophyletic Ichthyopterygia that is supported by 18 unambiguous synapomorphies. The updated analysis recovers Utatsusaurus in a basal clade with Parvinatator (united by one unambiguous feature), which collectively form the sister group to all other ichthyopterygians.
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In 1989, TMP 89.127.3, a basal ichthyopterygian, was collected as a counterpart slab from the Vega-Phroso Siltstone Member (Lower Triassic) of the Sulphur Mountain Formation in east-central British Columbia, Canada. Previous investigations of its postcranial skeleton resulted in ambiguous taxonomic affinities, with it being identified as either Grippia cf. longirostris or Parvinatator wapitiensis. More recently, the part slab of TMP 89.127.3 was collected and prepared, revealing important new cranial data that permits its systematic relationships to be more thoroughly assessed and previously proposed taxonomic affinities to be tested. By integrating both cranial and postcranial data, phylogenetic analysis recovers TMP 89.127.3 and Grippia as members of Grippidia, nested within Eoichthyosauria. TMP 89.127.3 can be distinguished from Grippia by possessing the following autapomorphies: metacarpal I with a notched peripheral shaft; cylindrical anterior teeth with a crown shape index >5.0; and a postfrontal-frontal contact at the rostral limit of the anterior margin of the supratemporal terrace. A new taxon, Gulosaurus helmi, is erected to accommodate this suite of features. The placement of Gulosaurus helmi within Grippidia reaffirms the Early Triassic distribution of this clade to include western and northern Laurasia, which is represented by outcrop in present-day Canada and Norway (Spitzbergen), respectively.
Many recent studies have detailed the morphology of archosaurian endocrania. However, the outgroup to Archosauria, Phytosauria, has yet to be studied with modern techniques that would allow reconstruction of their internal anatomy. Pseudopalatus mccauleyi is a derived phytosaur from the Upper Triassic Chinle Formation, Arizona. A skull of P. mccauleyi, USNM 15839, was imaged using computed tomography in order to create the first high-quality, digitally reconstructed phytosaur endocast. Pseudopalatus mccauleyi exhibits overall endocast morphology that is similar to that of an extant crocodylian. These clades, phytosaurs and extant crocodylians, exhibit convergent Baupläne and similar inferred ecologies. A notable difference between the endocasts of the two clades is a considerable dural expansion in P. mccauleyi that denotes a large pineal body. This expansion, and the overall morphology of the endocast, is consistent with the historic endocranial reconstructions of the phytosaurs Pseudopalatus buceros, Smilosuchus gregorii, and Parasuchus hislopi. A comparison with phylogenetically diverse archosaurian endocasts reveals that endocast morphologies are highly conserved within Pseudosuchia, regardless of Bauplan or ecology. This conservatism is in contrast to the diversity of endocast morphology observed within Theropoda and Sauropodomorpha, or between members of those clades and Pseudosuchia. The most pronounced variability in pseudosuchian endocast morphology is a trend in size reduction of the pineal region, from a large basal condition to a reduced derived condition wherein the pineal region is indistinguishable from the rest of the endocast. A similar trend in pineal reduction is also seen in theropods and sauropods.
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A rostrum fragment from the marginal marine upper Lunz Formation of Austria (early Late Triassic: late Carnian) was previously identified as a new species of phytosaurian archosauriform, ‘Francosuchus’ trauthi. ‘Francosuchus’ trauthi was subsequently synonymized with the non-phytosaurid phytosaur Paleorhinus, and this synonymy was used as evidence to correlate the ‘Paleorhinus biochron’ and the Otischalkian land-vertebrate faunachron to the marine timescale. Here, I provide a redescription of ‘Francosuchus’ trauthi and document anatomical features that differ substantially from all known species of non-phytosaurid phytosaur. There is no evidence to support synonymy of ‘Francosuchus’ trauthi with Paleorhinus, and no unambiguous features to support a phytosaurian identification. However, ‘Francosuchus’ trauthi possesses a unique combination of characters that distinguish it from all other Triassic tetrapods, and necessitates referral of the species to a new genus, Dolerosaurus, gen. nov. Rejection of the proposed synonymy between ‘Francosuchus’ trauthi and Paleorhinus means that the ‘Paleorhinus biochron’ cannot be tied to the marine late Carnian as previously suggested, and provides further evidence of the problems inherent in attempting to correlate the terrestrial Triassic to the marine timescale.
Noasauridae is a clade of ceratosaurian theropods that evolved small body size independently of other nonavian theropods. The best-preserved and most complete noasaurid is Masiakasaurus knopfleri from the Maastrichtian-aged Maevarano Formation in Madagascar. An abundance of skeletal material from several individuals spanning a wide range of ontogeny makes Masiakasaurus an ideal candidate for the analysis of growth. We histologically sampled a growth series of elements consisting of four femora and three tibiae. Bright-field and circularly polarized light microscopy were used to distinguish between slowly and rapidly growing forms of bone. To simultaneously estimate age at death and reconstruct growth trajectories, we measured the perimeters of growth lines in each specimen and fitted models to these data using a novel application of mixed-effects regression. Our histological results show an external fundamental system in the largest tibial specimen and confirm that Masiakasaurus grew determinately, matured at small body size, and is not the juvenile form of a larger-bodied theropod. Parallel-fibered bone is unusually prominent and suggests relatively slow growth. Moreover, our quantitative analysis shows that the average individual took about 8-10 years to get to the size of a large dog. Although Masiakasaurus grew 40% faster than crocodylians, it grew about 40% slower than comparably sized non-avian theropods. Slowed growth may have evolved as a means to minimize structural and maintenance costs while living in a semiarid and seasonally stressful environment. Dimorphism does not appear related to asymptotic size or growth rate but seems to reflect the degree of skeletal maturity.
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Liang Bua, a limestone cave on the Indonesian island of Flores, has a depositional sequence that spans the last 95,000 years and includes well-preserved faunal remains. Birds are well represented throughout the stratigraphic sequence at Liang Bua. Here, we present the results of the first comprehensive study of avian remains retrieved from Sector XI, a 2 m by 2 m archaeological excavation along the east wall of the cave. A total of 579 specimens were identified as avian, with 244 belonging to at least 26 non-passerine taxa in 13 families. The late Pleistocene assemblage (23 taxa) includes the first recorded occurrence of vultures in Wallacea, as well as kingfishers, snipes, plovers, parrots, pigeons, and swiftlets. Together, these taxa suggest that during this time the surrounding environment was floristically diverse and included several habitat types. Two of these taxa, the giant marabou Leptoptilos robustus and the vulture Trigonoceps sp., are extinct. Eight taxa were identified in the Holocene assemblage, and five of these were also present in the late Pleistocene. Imperial pigeons Ducula sp. and the Island Collared Dove Streptopelia cf. bitorquata appear only in the Holocene assemblage. The differences in faunal composition between the late Pleistocene and Holocene assemblages may reflect a change in avifaunal composition due to climatic and environmental changes near the Pleistocene-Holocene transition, possibly amplified by impacts associated with the arrival of modern humans; however, the small Holocene sample prevents a firm conclusion about faunal turnover from being made.
We describe a new avian species, Qianshanornis rapax, gen. et sp. nov., from the middle Paleocene Wanghudun Formation of the Qianshan Basin in Anhui Province, China. The holotype consists of an incomplete articulated foot and a few associated bones, mainly of the leg, wing, and pectoral girdle. Qianshanornis rapax is characterized by a derived foot morphology and may have had a hyperextendible second toe, which has so far only been reported for some Mesozoic taxa. The new species is markedly different from all other known Cenozoic birds and is here classified in the new taxon Qianshanornithidae. The leg bones, especially the distal end of the tibiotarsus, most closely resemble those of Strigogyps (Ameghinornithidae) from the Eocene, and possibly Oligocene, of Europe, but unlike the latter, the much smaller Q. rapax appears to have had well-developed flight capabilities.
The giant Miocene erinaceid Deinogalerix from the Gargano paleoisland, southern Italy, represents one of the most spectacular cases of insular adaptation among terrestrial mammals. The type species of this genus, Deinogalerix koenigswaldi, was described for the first time early in the 1970s based on a nearly complete skeleton and the limits and composition of the genus were subsequently defined based on a large number of disarticulated remains. Recent excavations in the Gargano area led to the discovery of a second partially complete skeleton of Deinogalerix koenigswaldi. This material, which is described herein, consists of a well-preserved nearly complete skull and a large portion of the postcranial skeleton. As a result of its excellent preservation, this material allowed the description of previously unrecognized morphological features of the braincase, tympanic region, and forelimb. Finally, based on a comparative analysis of the skeletal features of Deinogalerix, its affinities within the Erinaceidae have been discussed. Several features observed on the skeleton documented in this paper conclusively demonstrate that this giant hedgehog is a member of the subfamily Galericinae and, more particularly, of the extinct tribe Galericini.
Isolated cheek teeth of Sciuridae (Rodentia, Mammalia) from nine late Miocene localities in central Anatolia (Turkey) are described. The teeth represent at least 12 different species, five of which belong to the ground squirrel genus Tamias, two to the ground squirrel genus Spermophilinus, one to the flying squirrel genus Hylopetes, and two to the flying squirrel genus Pliopetaurista. One species, Tamias anatoliensis (type locality Altintaş 1), is new. An unknown genus and species of giant tree or ground squirrel is represented by one tooth. Two teeth probably form the oldest record in western Eurasia of the tree squirrel genus Sciurus. Seven of the localities that yielded Sciuridae are lacustrine deposits, two are karst fissure fills. Their estimated ages range from MN 9 to MN 13. The majority of late Miocene Sciuridae from Anatolia show affinities with European sciurids of the same period.
A new species of cricetid rodent, Megacricetodon vandermeuleni, sp. nov., is described from the local zone Db, middle Aragonian, from the Calatayud-Montalbán Basin and the Loranca Basin (Spain). The large-sized Megacricetodon is characterized by the elongated anteroconid and anterolophulid on ml and the robust and swollen cusps. Comparisons with other species of similar size and morphology are discussed. We consider M. bezianensis a valid species, based on the morphological differences with M. gersii. We propose a closely related group of Megacricetodon species that includes M. aff. collongensis, M. bavaricus, M. aff. bavaricus, M. bezianensis, M. lappi, M. aunayi, and M. vandermeuleni, nov. sp. The new species is of importance for Spanish biostratigraphy and the proposed Megacricetodon group is very useful for European faunal correlations and timing of migration events.
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A partial postcranial skeleton (vertebrae and ribs) of an indeterminate sirenian is described from Selminum Tem cave in the Hindenburg Range, Western Province of Papua New Guinea. It was derived from a section of the Darai Limestone dating to the Burdigalian-Serravallian (early-middle Miocene) and representing shallow platform carbonates. The thoracic vertebrae are remarkably small, being comparable in size to the vertebrae of Nanosiren garciae and implying small body size, although it is uncertain whether the specimen represents a diminutive adult or juvenile individual. These fossils represent the geologically earliest mammal recorded from the island of New Guinea and the earliest evidence of Sirenia in Australasia. Thus, this fossil evidence provides a minimum date (∼11.8 Ma) for the earliest presence of sirenians in Australasian coastal marine ecosystems, as well as their primary food source, seagrasses.
‘Arctocyonids’ are generally considered as including some of the most primitive ‘ungulates’ from the Paleocene. Although more than 15 genera are known from North America, European members of this order are less common and mainly belong to derived genera such as Arctocyon. However, one species of primitive arctocyonid, Prolatidens waudruae, was described from the early Paleocene of Hainin, Mons Basin, Belgium. Here we describe new dental positions of this small taxon, including for the first time upper molars and upper fourth premolar. Morphological comparisons confirm the position of P. waudruae among primitive ‘ungulates,’ with the closest North American arctocyonids being Prothryptacodon furens and Oxyprimus galadrielae. Oxyprimus galadrielae features slightly more primitive morphological traits than both other species. Apheliscids share several characters with Prolatidens, but the latter lacks the apomorphies defining the family. Among ‘arctocyonids,’ Prolatidens shares with only Protungulatum and Oxyprimus the incomplete lingual cingulum at the base of the protocone of M1. Prolatidens waudruae is unique among Procreodi in its combination of primitive and derived characters. The cladistic analysis places P. waudruae close to the base of the ingroup, indicating that this species is among the most primitive members of the Paleocene ‘ungulates.’ However, the lower nodes of the trees are not well supported and definitive conclusions should await more complete specimens and analysis. Apheliscids are situated relatively far from Prolatidens, suggesting that the resemblances between them are better considered as convergences. Based on the morphological comparisons, the arctocyonid from Hainin correlates best with North American Torrejonian taxa.
A new species of the genus Aceratherium, A. porpani sp. nov., from the Tha Chang sand pits in Nakhon Ratchasima Province, northwestern Thailand, is described. It is a mid-sized rhinocerotid in the subfamily Aceratheriinae, and represents the first discovery of Aceratherium in Thailand. The material includes a well-preserved skull and mandible. A. porpani has broadly separated parietal crests, slightly expanded zygomatic arches, a straight nuchal crest, moderate supraorbital tuberosities, a flat skull roof, a deep nasal notch above the P4/M1 boundary, a moderately wide mandibular symphysis with a posterior border at the p3/p4 boundary, a short diastema between 12 and p2, absence of DP1 and dp1, strong crochets, constricted molar protocones, and long metalophs. This new species has a mixture of primitive and derived characters that differ from the known species of Aceratherium, A. incisivum, and A. depereti. The evolutionary stage of A. porpani is consistent with the latest Miocene age of the associated fauna and flora in the Tha Chang sand pits.
A new genus and species of Ziphiidae, Notoziphius bruneti, gen. et sp. nov., from the late Miocene of Patagonia, is described on the basis of a well-preserved skull and partial left and right dentaries. It can be diagnosed by large, triangular, and markedly asymmetric nasals that strongly point anteroventrally, the ascending process of the maxilla not expanded posteriorly, supraoccipital strongly sloped posteroventrally; elliptical fossa on the nasal process of the premaxilla, small and anterolaterally directed premaxillary crest, and the presence of well-defined alveoli in the maxilla. A phylogenetic analysis including 25 ziphiid genera and 31 characters shows Notoziphius as nested within Ziphiidae in a basal clade with Aporotus, Beneziphius, Messapicetus, and Ziphirostrum. This clade is diagnosed by medial fusion of the premaxillae that closes the mesorostral groove, lateral margin of prenarial basin formed by a thick strip of maxilla, premaxillary crest anterolaterally directed, and reduced contact between nasal and premaxillary crest. The presence of Notoziphius in the Miocene of Patagonia increases our knowledge of ziphiid diversity in South America. Notoziphius bruneti and other Miocene records of ziphiids reinforce the idea that during the Miocene ziphiids were widely distributed and diverse.
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