Colonization of the ocean floor by jawless vertebrates across three mass extinctions.

BACKGROUND: The deep (> 200 m) ocean floor is often considered to be a refugium of biodiversity; many benthic marine animals appear to share ancient common ancestry with nearshore and terrestrial relatives. Whether this pattern holds for vertebrates is obscured by a poor understanding of the evolutionary history of the oldest marine vertebrate clades. Hagfishes are jawless vertebrates that are either the living sister to all vertebrates or form a clade with lampreys, the only other surviving jawless fishes. RESULTS: We use the hagfish fossil record and molecular data for all recognized genera to construct a novel hypothesis for hagfish relationships and diversification. We find that crown hagfishes persisted through three mass extinctions after appearing in the Permian ~ 275 Ma, making them one of the oldest living vertebrate lineages. In contrast to most other deep marine vertebrates, we consistently infer a deep origin of continental slope occupation by hagfishes that dates to the Paleozoic. Yet, we show that hagfishes have experienced marked body size diversification over the last hundred million years, contrasting with a view of this clade as morphologically stagnant. CONCLUSION: Our results establish hagfishes as ancient members of demersal continental slope faunas and suggest a prolonged accumulation of deep sea jawless vertebrate biodiversity.

A juvenile bird with possible crown-group affinities from a dinosaur-rich Cretaceous ecosystem in North America.

BACKGROUND: Living birds comprise the most speciose and anatomically diverse clade of flying vertebrates, but their poor early fossil record and the lack of resolution around the relationships of the major clades have greatly obscured extant avian origins. RESULTS: Here, I describe a Late Cretaceous bird from North America based on a fragmentary skeleton that includes cranial material and portions of the forelimb, hindlimb, and foot and is identified as a juvenile based on bone surface texture. Several features unite this specimen with crown Aves, but its juvenile status precludes the recognition of a distinct taxon. The North American provenance of the specimen supports a cosmopolitan distribution of early crown birds, clashes with the hypothesized southern hemisphere origins of living birds, and demonstrates that crown birds and their closest relatives coexisted with non-avian dinosaurs that independently converged on avian skeletal anatomy, such as the alvarezsaurids and dromaeosaurids. CONCLUSIONS: By revealing the ecological and biogeographic context of Cretaceous birds within or near the crown clade, the Lance Formation specimen provides new insights into the contingent nature of crown avian survival through the Cretaceous-Paleogene mass extinction and the subsequent origins of living bird diversity.

A late-surviving phytosaur from the northern Atlantic rift reveals climate constraints on Triassic reptile biogeography.

BACKGROUND: The origins of all major living reptile clades, including the one leading to birds, lie in the Triassic. Following the largest mass extinction in Earth's history at the end of the Permian, the earliest definite members of the three major living reptile clades, the turtles (Testudines), crocodylians and birds (Archosauria), and lizards, snakes, amphisbaenians, and Tuatara (Lepidosauria) appeared. Recent analyses of the Triassic reptile fossil record suggest that the earliest diversifications in all three of these clades were tightly controlled by abrupt paleoclimate fluctuations and concordant environmental changes. Yet, this has only been preliminarily tested using information from evolutionary trees. Phytosauria consists of superficially crocodylian-like archosaurs that either form the sister to the crown or are the earliest divergence on the crocodylian stem and are present throughout the Triassic, making this clade an excellent test case for examining this biogeographic hypothesis. RESULTS: Here, I describe a new phytosaur, Jupijkam paleofluvialis gen. et sp. nov., from the Late Triassic of Nova Scotia, Canada, which at that time sat in northern Pangaea near the northern terminus of the great central Pangean rift. As one of the northernmost occurrences of Phytosauria, J. paleofluvialis provides critical new biogeographic data that enables revised estimations of phytosaur historical biogeography along phylogenies of this clade built under multiple methodologies. Reconstructions of phytosaur historical biogeography based on different phylogenies and biogeographic models suggest that phytosaurs originated in northern Pangaea, spread southward, and then dispersed back northward at least once more during the Late Triassic. CONCLUSIONS: The results presented in this study link phytosaur biogeography to major changes to Triassic global climate and aridity. Together with the earliest dinosaurs and several other reptile lineages, phytosaur diversification and migration appear to have been restricted by the formation and loss of arid belts across the Pangean supercontinent.

Phylogenetics and the Cenozoic radiation of lampreys.

The development of a movable jaw is one of the most important transitions in the evolutionary history of animals.1 Jawed vertebrates rapidly diversified after appearing approximately 470 million years ago. Today, only lampreys and hagfishes represent the once dominant jawless grade2,3,4 and comprise less than 1% of living vertebrate species. Their relationship to other vertebrates ranks among the more contentious problems in animal phylogenetics.5,6,7,8,9,10,11,12 Further, the phylogenetic relationships within lampreys and hagfishes remain unclear,13,14,15 and the ages of their living lineages are largely unexplored.16,17 Because of their importance for the genomic and developmental changes that prefigured jawed vertebrate diversity,18,19,20,21 the evolutionary history of lampreys and hagfishes is a major frontier of organismal biology. Of these two clades, lampreys22 are more ecologically diverse, exhibiting freshwater, anadromous, and fully marine forms, as well as parasitic and nonparasitic species.23,24 Here, we present a new phylogeny and historical biogeographic reconstruction of all living lampreys. Whereas the early diversification of this clade tracks Pangaean fragmentation, lampreys also rapidly radiated in the northern hemisphere during the mid-Cretaceous and directly after the Cretaceous-Paleogene extinction. These radiations mirrored concurrent ones in other animals and plants and coincided with changes to lamprey ecology and feeding behavior. Our results suggest that 80% of living lamprey clades appeared in the last 20 million years of Earth history. Rather than gradually accumulating since the oldest stem-group forms appeared in the early Paleozoic, living lamprey biodiversity results from diversifications extending from the Cretaceous to present.

Species delimitation and coexistence in an ancient, depauperate vertebrate clade.

BACKGROUND: A major challenge to understanding how biodiversity has changed over time comes from depauperons, which are long-lived lineages with presently low species diversity. The most famous of these are the coelacanths. This clade of lobe-finned fishes occupies a pivotal position on the vertebrate tree between other fishes and tetrapods. Yet only two extant species and fewer than 100 extinct forms are known from the coelacanth fossil record, which spans over 400 million years of time. Although there is evidence for the existence of additional genetically isolated extant populations, a poor understanding of morphological disparity in this clade has made quantifying coelacanth species richness difficult. RESULTS: Here, we quantify variation in a sample of skulls and skeletons of the Triassic eastern North American coelacanth †Diplurus that represents the largest assemblage of coelacanth individuals known. Based on the results of these quantitative comparisons, we identify a diminutive new species and show that multiple lacustrine ecosystems in the Triassic rift lakes of the Atlantic coastline harbored at least three species of coelacanths spanning two orders of magnitude in size. CONCLUSIONS: Conceptions about the distribution of species diversity on the tree of life may be fundamentally misguided when extant diversity is used to gauge signals of extinct diversity. Our results demonstrate how specimen-based assessments can be used to illuminate hidden biodiversity and show the utility of the fossil record for answering questions about the hidden richness of currently species-poor lineages.

Giant gar from directly above the Cretaceous-Palaeogene boundary suggests healthy freshwater ecosystems existed within thousands of years of the asteroid impact.

The Cretaceous-Palaeogene (K-Pg) mass extinction was responsible for the destruction of global ecosystems and loss of approximately three-quarters of species diversity 66 million years ago. Large-bodied land vertebrates suffered high extinction rates, whereas small-bodied vertebrates living in freshwater ecosystems were buffered from the worst effects. Here, we report a new species of large-bodied (1.4-1.5 m) gar based on a complete skeleton from the Williston Basin of North America. The new species was recovered 18 cm above the K-Pg boundary, making it one of the oldest articulated vertebrate fossils from the Cenozoic. The presence of this freshwater macropredator approximately 1.5-2.5 thousand years after the asteroid impact suggests the rapid recovery and reassembly of North American freshwater food webs and ecosystems after the mass extinction.

High morphological disparity in a bizarre Paleocene fauna of predatory freshwater reptiles.

BACKGROUND: The consequences of the K-Pg mass extinction are reflected across present biodiversity, but many faunas that appeared immediately after the extinction event were very different from current ones. Choristodera is a clade of reptiles of uncertain phylogenetic placement that have an extremely poor fossil record throughout their 150-million-year history. Yet, choristoderes survived the K-Pg event and persisted until the Miocene. RESULTS: I describe the skulls and skeletons of two new choristoderes from a single Paleocene ecosystem in western North America that reveal the hidden Cenozoic diversity of this reptile clade. Despite their similar size, the new species deviate dramatically in morphology. Kosmodraco magnicornis gen. et sp. nov. possesses an extremely short snout and extensive cranial ornamentation. The sacrum of K. magnicornis bears enlarged muscle attachment sites and other modifications reminiscent of some giant crocodylians. In contrast, Champsosaurus norelli sp. nov. is a longirostrine species with an uninflated and ventrally divergent postorbital skull. Together with a North American choristodere previously classified in the European genus Simoedosaurus, K. magnicornis substantiates a new clade of giant, short-snouted taxa endemic to the Americas. C. norelli is found to be an early-diverging member of the genus Champsosaurus from the Cretaceous-Paleogene of the northern hemisphere. This suggests the presence of several ghost lineages of champsosaurid that crossed the K-Pg boundary. CONCLUSIONS: The new taxa greatly increase Cenozoic choristodere richness and strengthen the evidence for the existence of distinctive freshwater faunas in Paleogene Eurasia and North America, where this clade diversified to exploit newly available macropredatory niches in the aftermath of the asteroid impact. The new choristoderes also reveal the distinct ecological context in which extant freshwater predators of the Americas like alligatoroids and gars have their origins: Paleocene fluviolacustrine ecosystems in North America displayed high large predator diversity and morphological disparity relative to modern ones.

New records of theropods from the latest Cretaceous of New Jersey and the Maastrichtian Appalachian fauna.

The faunal changes that occurred in the few million years before the Cretaceous-Palaeogene extinction are of much interest to vertebrate palaeontologists. Western North America preserves arguably the best fossil record from this time, whereas terrestrial vertebrate fossils from the eastern portion of the continent are usually limited to isolated, eroded postcranial remains. Examination of fragmentary specimens from the American east, which was isolated for the majority of the Cretaceous as the landmass Appalachia, is nonetheless important for better understanding dinosaur diversity at the end of the Mesozoic. Here, I report on two theropod teeth from the Mount Laurel Formation, a lower-middle Maastrichtian unit from northeastern North America. One of these preserves in detail the structure of the outer enamel and resembles the dentition of the tyrannosauroid Dryptosaurus aquilunguis among latest Cretaceous forms in being heavily mediolaterally compressed and showing many moderately developed enamel crenulations. Along with previously reported tyrannosauroid material from the Mt Laurel and overlying Cretaceous units, this fossil supports the presence of non-tyrannosaurid tyrannosauroids in the Campanian-Maastrichtian of eastern North America and provides evidence for the hypothesis that the area was still home to relictual vertebrates through the end of the Mesozoic. The other tooth is assignable to a dromaeosaurid and represents both the youngest occurrence of a non-avian maniraptoran in eastern North America and the first from the Maastrichtian reported east of the Mississippi. This tooth, which belonged to a 3-4 m dromaeosaurid based on size comparisons with the teeth of taxa for which skeletons are known, increases the diversity of the Maastrichtian dinosaur fauna of Appalachia. Along with previously reported dromaeosaurid teeth, the Mt Laurel specimen supports the presence of mid-sized to large dromaeosaurids in eastern North America throughout the Cretaceous.

Description of Arundel Clay ornithomimosaur material and a reinterpretation of Nedcolbertia justinhofmanni as an "Ostrich Dinosaur": biogeographic implications.

The fossil record of dinosaurs from the Early Cretaceous of Eastern North America is scant, especially since a few stratigraphic units from the east are fossiliferous. Among these stratigraphic units, the Arundel Clay of the eastern seaboard has produced the best-characterized dinosaur faunas known from the Early Cretaceous of Eastern North America. The diverse dinosaur fauna of the Arundel Clay has been thoroughly discussed previously, but a few of the dinosaur species originally described from the Arundel Clay are still regarded as valid genera. Much of the Arundel material is in need of review and redescription. Among the fossils of dinosaurs from this stratigraphic unit are those referred to ornithomimosaurs. Here, the researcher describes ornithomimosaur remains from the Arundel Clay of Prince George's County, Maryland which may be from two distinct ornithomimosaur taxa. These remains provide key information on the theropods of the Early Cretaceous of Eastern North America. Recent discoveries of small theropod material from the Arundel Clay possibly belonging to ornithomimosaurs are also reviewed and described for the first time. The description of the Arundel material herein along with recent discoveries of basal ornithomimosaurs in the past 15 years has allowed for comparisons with the coelurosaur Nedcolbertia justinhofmanni, suggesting the latter animal was a basal ornithomimosaur rather than a "generalized" coelurosaur as it was originally described. Comparisons between the Arundel ornithomimosaur material and similar Asian and European specimens suggest that both extremely basal ornithomimosaurs and more intermediate or derived forms may have coexisted throughout the northern hemisphere during the Early Cretaceous.