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Relative to large-bodied dinosaurs, the diversity of small-bodied dinosaurs from the Campanian of North America is poorly understood due to a lack of well-preserved skeletons. We document the first articulated remains, as well as the first cranial bones, of non-iguanodontian ornithopods from the Belly River Group of Alberta. The geologically oldest specimen consists of the posterior half of an articulated skeleton from the middle unit of the Oldman Formation and shares many anatomical features with the contemporaneous Orodromeus makelai and the older Oryctodromeus cubicularis. A second, younger specimen from the upper Oldman Formation is distinct from other ornithopods in having a reduced distal portion of the fibula that is fused to the anterior surface of the tibia; it is designated as the type of a new taxon, Albertadromeus syntarsus, gen. et sp. nov. Numerous isolated elements from small ornithopods from the Dinosaur Park Formation are also identified, but cannot be assigned to the generic level with confidence. Although small-bodied ornithopod material is rare, their known postcranial material outnumbers those of taphonomically equivalent and contemporaneous pachycephalosaurs, which are known to be abundant and diverse due to their robust and frequently recovered cranial domes. These findings suggest considerable undiscovered diversity of small-bodied ornithopods, and highlight biases against the preservation of small taxa in this system.
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Although selachian fossil remains have been studied for nearly three centuries, little is known about their evolutionary history. Recent studies have suggested different timings of early diversification events in the Late Triassic, Jurassic, and Early Cretaceous. However, Late Cretaceous selachian diversity has remained little explored despite numerous and diversified selachian assemblages known from this time interval. Sampling standardization, origination/extinction rates, and raw ordinal diversity were examined based on taxonomic occurrences in three data sets representing distinct geographical areas (Anglo-Paris Basin, northwestern Europe, and Western Interior Seaway) spanning the Cenomanian-Campanian interval. This examination allowed the identification of regional diversity events, previously reported for some invertebrate groups, but presented for the first time for a marine vertebrate group. The local mid-Cenomanian diversity drop (Anglo-Paris Basin) is interpreted as a possible consequence of changes in bottom seawater conditions related to the rapid mid-Cenomanian transgression. The Cenomanian/Turonian faunal turnover is likely to be due to various sampling biases (Anglo-Paris Basin and northwestern Europe), but a genuine extinction in the Western Interior Seaway cannot be excluded. The Santonian diversity peak synchronous with a marked global increase in seawater temperatures contrasts with the rapid temperature decrease linked with low diversity in the early Campanian.
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Fossil elements belonging to fishes were recovered from the Pliocene Chiwondo Beds in the Malawi Rift. This is our first knowledge of fishes from Lake Malawi. The Chiwondo Bed fishes can be assigned to living taxa including lungfish (Protopterus), catfishes (Bagridae, Claroteidae and Clariidae), cichlids, and a single tooth referred to tigerfish (Hydrocynus). Although small and taphonomically biased, the assemblage shows several taxonomic differences compared with the modern Lake Malawi fauna. Most of the geographic and taxonomic affinities of the assemblage are with fishes from the African East Coast Province rivers, whereas the modern lake fauna primarily has affinities to the Zambezi River system. The Chiwondo fish assemblage contributes new information on Plio-Pleistocene eastern/southern African biogeography and waterways, and how these may have influenced faunal movements between eastern and southern Africa.
‘Microsaurs’ (Lepospondyli) were a group of tetrapods whose fossil record spanned the Mississippian—Early Permian, and have sometimes been implicated in the origins of lissamphibians (especially the fossorial caecilians). Although common in the well-sampled Permian deposits of Texas and Oklahoma, little is known of conservative recumbirostran ‘microsaurs’ from more northerly localities (Kansas/Nebraska). Data on a new, well-preserved recumbirostran, Huskerpeton englehorni, gen. et sp. nov., from the Eskridge Formation, Nebraska, reveal affinities to the purported gymnarthrid ‘Euryodus’bonneri (here moved to a new genus, Proxilodon) from the Speiser Formation, Kansas. High resolution micro-computed tomography reveals important endocranial data, including an ossified ‘supraoccipital’ that is usually absent in gymnarthrids, a dorsal sinus separating the synotic tectum from the dermal skull roof as in ostodolepids, ventral flanges of the frontals articulating directly with the ascending lamina of the sphenethmoid, and tall, ossified ‘pleurosphenoids,’ thereby providing useful comparison with morphologically derived recumbirostrans. A cladistic analysis of 60 taxa and 227 characters recovered a monophyletic clade of Huskerpeton, Proxilodon, and the recently described Tambaroter within a more inclusive clade that includes gymnarthrids and ostodolepids (both nested members of the Recumbirostra and putative outgroups to caecilians). Whereas other lepospondyls (i.e., lysorophians) are abundant in the region, the records of Huskerpeton and Proxilodon reveal that only two ‘microsaur’ genera are adequately known from the lowermost Permian of Kansas/Nebraska and demonstrate similarities between the Eskridge and Speiser faunas. Moreover, these forms may represent the ground plan from which more specialized fossorial forms (gymnarthrids and ostodolepids) evolved.
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A nearly complete amphibamid skull from the Richards Spur locality in Oklahoma is demonstrated to be a new species of Tersomius. This new species has a mosaic of features seen in other amphibamids, and appears to fall on the Amphibamus side of the amphibamid basal dichotomy of some authors. This specimen also calls into question the validity of the Tersomius assignment of a large specimen from the Texas Red Beds. One feature of note is the presence of a weakly bicuspid fang on one of the vomers. This poorly developed feature is present only on an unattached replacement cusp, and appears to have been rapidly worn away through use. This is the first potential bicuspid palatal fang known from an amphibamid. A second partial skull is assigned to the recently described species Pasawioops mayi. A micro-computed tomographic reconstruction shows a prominent ventral flange of the nasal that meets a counterpart from the prefrontal and palatine to wall the nasal capsule posteriorly. The nasolacrimal canal shows the three openings in the orbit quickly converge to pass rostrally as a single canal except for a mysterious lateral diverticulation that has a superficial exposure on the lacrimal. Both skulls preserve a pars palatina of the premaxilla, in common with batrachians, and the Pasawioops specimen also preserves a prominent palatal process of the premaxilla, a frog synapomorphy. Richards Spur preserves one of the most diverse amphibamid faunas in the world, with three species described so far.
The Albanerpetontidae, small salamander-like tetrapods from the Middle Jurassic-Neogene of Laurasia and northern Africa, are widely considered to be lissamphibians; however, relationships among major lissamphibian clades are unresolved. A recently identified, isolated, and three-dimensionally preserved neurocranium (early Pliocene, Hungary) referred to Albanerpeton pannonicum is described, incorporating information gained from the application of micro-computed tomography. It is revealed that the neurocranium is a robust, box-like structure composed of the coossification of the parasphenoid, otic capsules, and occipital elements. The otic capsule endocast reveals the morphology of the endosseous labyrinth, complete with well-defined endosseous semicircular canals and a modestly developed ventral endosseous auditory region; however, details of the individual auditory organs are not discernable from the endocast. Features of the neurocranium and endosseous labyrinth are consistent with the hypothesis that A. pannonicum, and albanerpetontids in general, were somewhat fossorial. The neurocranium and endosseous labyrinth exhibit a mosaic of anuran, urodele, and apodan traits, thus precluding refinement of the phylogenetic position of albanerpetontids. In general, the neurocranium and endosseous labyrinth appear most similar to urodeles, and similarities with apodans and anurans may be due to convergent evolution resulting from similar habits and responses to inner ear stimulation. This new neurocranium represents the best-known specimen of its kind for albanerpetontids, and the data presented here combined with future comparative studies will contribute to a better understanding of the biology and evolution of this group.
North American Late Cretaceous salamanders are principally known by isolated atlantes and trunk vertebrae. Here I describe a new genus and species of fossil salamander, Paranecturus garbanii, gen. et sp. nov., based on these elements from the lower portions of the Hell Creek Formation (Maastrichtian), Garfield County, northeastern Montana, U.S.A. It is diagnosed by a unique combination of character states that include the presence of an alar-like process of the atlas and a groove on the posterior face of the neural arch and solid dorsal rib-bearers of the trunk vertebrae. My phylogenetic analysis of 13 caudate taxa and 23 atlantal and trunk vertebral characters recovered P. garbanii as a member of the Proteidae. P. garbanii represents the oldest fossil record of the Proteidae and demonstrates that this lineage was present before the Cretaceous-Paleogene mass extinction event.
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A new genus and two new species of mosasaurs are described from five specimens collected during quarrying operations in the ‘Lastame’ lithotype located in the mountains of northern Italy just to the north of Verona (lower Turonian—lower Santonian). These mosasaurs share some anatomical characters with the North American taxon Russellosaurus coheni, but the presence of distinctive features suggests placement into two new species within a new genus (e.g., forked distal tip of suprastapedial process of quadrate; suture between maxilla and premaxilla extends to point above third tooth position [cf. second in Russellosaurus]). We conducted two phylogenetic analyses of the Mosasauroidea (37 ingroup taxa, 131 characters), the first one with equally weighted characters and the second after successive weighting in order to reduce homoplastic noise and amplify the signal in the data. The first analysis (equal weights) resulted in 96 most parsimonious trees, whereas the second produced a single shortest tree. Both analyses consistently placed the new genus as the sister taxon to Russellosaurus, and these two taxa were placed together with Yaguarasaurus, in a clade that is the sister group of the Tethysaurinae (Tethysaurus and Pannoniasaurus). A new clade of mosasaurs, the Yaguarasaurinae, is here defined as comprising the most recent common ancestor of Russellosaurus, Romeosaurus, and Yaguarasaurus, and all of its descendants.
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The Eulert pliosaurid remains (FHSM VP-321) housed at the Sternberg Museum of Natural History (Kansas, U.S.A.) include one of the world's best examples of a Cretaceous pliosaurid plesiosaur skull. The specimen's original assignment to Brachauchenius lucasi was based solely upon the skull (dorsal surface) and left lower jaw (lateral view) because the specimen was embedded in a plaster mount. The history of B. lucasi is similarly problematic, because the type and a referred skull were formerly visible only in ventral and dorsal views, respectively. Further preparation and comparison of these specimens reveal new data about the arrangement of cranial elements. The Eulert pliosaurid bears several distinct autapomorphies as compared with B. Lucasi, including cranial proportions (pretemporal length of palate longer, shorter temporal fenestrae), configuration of skull roof elements (frontals participate in premaxilla-parietal suture, suture occurs further forward), and configuration of the palate (posterior vomers not masked by medial alar extensions of the palatines, caudal vomerian fenestrae positioned further posterior, long slit-like anterior pteryoid vacuity present). Furthermore, FHSM VP-321 possesses double-headed cervical ribs, a feature previously unknown in Cretaceous pliosaurids. This combination of characters merits separation of the Eulert pliosaurid and a referred specimen to a new taxon, Megacephalosauris eulerti. The type and paratype skulls of M. eulerti are 1.5 m and 1.75 m in length, respectively, and thus 50% and 75% larger than the known 1-m-long skulls of B. lucasi, suggesting that M. eulerti may attain larger size than B. lucasi.
Jack L. Conrad, Kirsten Jenkins, Thomas Lehmann, Fredrick K. Manthi, Daniel J. Peppe, Sheila Nightingale, Adam Cossette, Holly M. Dunsworth, William E. H. Harcourt-Smith, Kieran P. McNulty
‘Crocodylus’ pigotti is a relatively small crocodylid from the Miocene of Rusinga Island in Lake Victoria, Kenya. Known only from one relatively complete skull and limited, fragmentary, referred material, ‘Crocodylus‘ pigotti lacks a detailed description. Moreover, recent analyses have shown ‘Crocodylus’ pigotti to be an osteolaemine crocodylid, more closely related to the extant dwarf crocodiles (Osteolaemus) than to true Crocodylus. Here, we describe numerous new remains of ‘Crocodylus’ pigotti recovered from localities within the Fossil Bed Member of the Hiwegi Formation at Kaswanga Point, Rusinga Island. We recovered parts of several individuals and report on previously unknown parts of the anatomy, provide an updated phylogenetic analysis, and reallocate the species ‘Crocodylus’ pigotti to a new genus, Brochuchus.
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Two nearly complete skeletons of the filter-feeding pterodactyloid Pterodaustro guinazui from the Lower Cretaceous of Argentina exhibit clusters of poorly sorted coarse sand to fine gravel inside the abdominal cavity. These stones are interpreted as ingested gastroliths (geogastroliths), which are commonly found in a variety of archosaurs (including birds) but have never before been reported in a pterosaur. The geogastroliths found in these Pterodaustro specimens are interpreted as having assisted in the digestion of hard food items such as ‘shelled’ crustaceans that are abundant in the fossil beds of this pterosaur. One of these specimens with geogastroliths has anterior mandibular teeth that are notably thicker than the posterior teeth and are somewhat procumbent. We suggest that these teeth might have facilitated the apprehension of fine gravel.
Dicynodonts are the most abundant therapsids from the Beaufort Group (Karoo Basin, South Africa), and they have been used as index fossils to define most of the Beaufort biozones. Although speciose during the Late Permian, dicynodont diversity was greatly reduced by the end-Permian mass extinction. The Cynognathus Assemblage Zone, the uppermost biozone of the Beaufort Group, is traditionally known to host only the dicynodonts Kannemeyeria simocephalus and Kombuisia frerensis. Recent biostratigraphic research in this biozone has revealed the presence of two more poorly known dicynodont genera, Angonisaurus and Shansiodon, which are biostratigraphically constrained to the uppermost Cynognathus Assemblage Zone (subzone C). Although these taxa have been reported from subzone C of the Cynognathus Assemblage Zone previously, we provide the first detailed description of the material in question and justifications for the taxonomic identifications. The presence of Angonisaurus in the Cynognathus subzone C allows a direct correlation between this assemblage and that of the Lifua Member of the Manda beds in Tanzania. A correlation with the Antarctic upper Fremouw Formation also may be possible, but is less certain. The presence of Shansiodon allows a direct correlation with the upper Ermaying Formation of China, but we reject the previously proposed global ‘Shansiodon biochron’ because Shansiodon sensu stricto only occurs in China and South Africa. Biogeographically, Angonisaurus appears to have been widespread but rare in southern Gondwana, and the unusual distribution of Shansiodon mirrors that of Diictodon feliceps in the Permian. Southern African dicynodont assemblages display increased provinciality in the Middle Triassic.
Latest Cretaceous (Lancian) mammalian faunas of the Western Interior of North America are mostly known from the northern Great Plains and coastal lowland paleoenvironments. Here, we describe a sample of 143 multituberculate mammal teeth from the Lance Formation of southwestern Wyoming. The specimens, which are from two independent collections made in the 1970s by the University of California Museum of Paleontology and the University of Wyoming Geological Museum, are part of the best-sampled local fauna from the central part of the Western Interior. Deposits of the Lance Formation in this region are on the eastern flank of the Rock Springs Uplift near Black Butte Station. The Black Butte Station local fauna was farther west and possibly paleoenvironmentally distinct from most other Lancian local faunas known. The fossil assemblage preserves eight genera and 11 species of multituberculates. There are many common Lancian taxa, a high relative abundance of Cimexomys, the second published occurrences of Parikimys and Paressonodon, and a new species of Cimolodon. Cluster and ordination analyses of multituberculate abundance data from well-sampled Western Interior local faunas show that the Black Butte Station local fauna is distinct from all other local faunas and that variation among mammalian local faunas in composition is correlated with latitude, though paleoenvironmental, temporal, and taphonomic differences may also be factors. Results highlight that explorations in undersampled regions and paleoenvironments are critical to a more complete understanding of the Cretaceous-Paleogene transition.
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The knowledge of northern South American Glyptodontidae (Mammalia, Xenarthra, Cingulata) is very scarce compared with that of southern South American taxa, which have been systematically studied since the 19th century. Recently, the northern taxa (originally assigned to the Glyptodontidae Propalaehoplophorinae Asterostemma and Propalaehoplophorus) have been reinterpreted as basal Glyptodontinae, belonging to the new genus Boreostemma. In this contribution, we present and describe the most complete Neogene Glyptodontidae from northern South America (middle Miocene of the Honda Group, La Venta, Colombia), and its main taxonomic, paleobiogeographic, and phylogenetic implications. This new material expands the morphological characterization of B. acostae and corroborates differences compared with Glyptodontidae from Miocene southern South America. A cladistic analysis corroborates the monophyly of the Glyptodontinae, that B. acostae and B. venezolensis being the sister group of the remaining taxa of Glyptodontinae. The traditionally recognized genera (e.g., Glyptodon and Glyptotherium) constitute natural groups. Whereas the Miocene seems to represent a diversification period for Glyptodontidae in southern South America, the recorded taxa in northern South America are restricted with certainty only to the Glyptodontinae Boreostemma.
With its short and pointed rostrum, the small fossil dolphin Brachydelphis mazeasi, from late middle to early late Miocene deposits of Chile and Peru, was originally described as an unusual member of the family Pontoporiidae (Cetacea, Odontoceti), presently only represented by the long-snouted Pontoporia blainvillei, a relict species from the eastern coast of South America. However, the phylogenetic relationships of Brachydelphis were debated in subsequent works. Based on a sample of well-preserved specimens from the late Miocene of the Pisco Formation (Tortonian, about 9 Ma) in the Sacaco Basin, southern coast of Peru, we describe a new species of Brachydelphis, B. jahuayensis, sp. nov. Also recorded in Chile, B. jahuayensis differs mostly from the type species in its considerably longer snout and higher tooth count. From a functional standpoint, the new species is interpreted as less specialized for suction feeding than the type species, relying more on its toothed jaws for prey capture. The inclusion of the long-snouted B. jahuayensis in future phylogenies will likely provide further support to the referral of Brachydelphis to the family Pontoporiidae. Finally, a review of the Mio-Pliocene vertebrate levels of the Pisco Formation in the Sacaco Basin clarifies the marine mammal content, succession, and stratigraphic correlation of each level, especially in the Aguada de Lomas locality.
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A new genus and species of true dolphin (Delphinidae), Septidelphis morii, is described on the basis of a partial fossil skeleton (almost complete skull, mandibles, teeth, hyoid bones, 17 vertebrae, several ribs, sternum and sternal ribs), originally belonging to an immature subadult animal, from the late Zanclean—early Piacenzian (Pliocene) marine sands of Piedmont (northern Italy). This new dolphin is characterized by an extreme posterior widening of the dorsal opening of the mesorostral canal and shares with Astadelphis, Etruridelphis, Sousa, and Steno the following combination of characters: (1) long and narrow rostrum; (2) wide premaxillae at the middle of rostrum; and (3) condylobasal length reaching at least 550 mm. A phylogenetic analysis using a molecular scaffold approach placed Septidelphis inside the delphinines in a more apical position than Sousa but basal to the clade formed by all other extant delphinines. Etruridelphis is positioned as the sister taxon of Septidelphis, and Astadelphis is positioned as the sister taxon of a larger grouping that includes this clade and all extant delphinines except Sousa. The phylogenetic position of Septidelphis inside Delphinidae and its stratigraphical age (3.81–3.19 Ma) are consistent with estimated divergence dates reported in a recent molecular study, whereas the first occurrence of Etruridelphis, dated to slightly before 3.98 Ma, could represent a minimal age for the origin of the delphinines and a potential calibration point for molecular clock studies. Septidelphis corroborates the explosive radiation and fast speciation-extinction rate of the delphinids during the Pliocene-Pleistocene.
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