SciCombinator

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Concept: Dinosaur

789

The population extinction pulse we describe here shows, from a quantitative viewpoint, that Earth’s sixth mass extinction is more severe than perceived when looking exclusively at species extinctions. Therefore, humanity needs to address anthropogenic population extirpation and decimation immediately. That conclusion is based on analyses of the numbers and degrees of range contraction (indicative of population shrinkage and/or population extinctions according to the International Union for Conservation of Nature) using a sample of 27,600 vertebrate species, and on a more detailed analysis documenting the population extinctions between 1900 and 2015 in 177 mammal species. We find that the rate of population loss in terrestrial vertebrates is extremely high-even in “species of low concern.” In our sample, comprising nearly half of known vertebrate species, 32% (8,851/27,600) are decreasing; that is, they have decreased in population size and range. In the 177 mammals for which we have detailed data, all have lost 30% or more of their geographic ranges and more than 40% of the species have experienced severe population declines (>80% range shrinkage). Our data indicate that beyond global species extinctions Earth is experiencing a huge episode of population declines and extirpations, which will have negative cascading consequences on ecosystem functioning and services vital to sustaining civilization. We describe this as a “biological annihilation” to highlight the current magnitude of Earth’s ongoing sixth major extinction event.

Concepts: Evolution, Biology, Species, Mammal, Extinction, Dinosaur, Extinction event, Timeline of extinctions

555

Birds still share many traits with their dinosaur ancestors, making them the best living group to reconstruct certain aspects of non-avian theropod biology. Bipedal, digitigrade locomotion and parasagittal hindlimb movement are some of those inherited traits. Living birds, however, maintain an unusually crouched hindlimb posture and locomotion powered by knee flexion, in contrast to the inferred primitive condition of non-avian theropods: more upright posture and limb movement powered by femur retraction. Such functional differences, which are associated with a gradual, anterior shift of the centre of mass in theropods along the bird line, make the use of extant birds to study non-avian theropod locomotion problematic. Here we show that, by experimentally manipulating the location of the centre of mass in living birds, it is possible to recreate limb posture and kinematics inferred for extinct bipedal dinosaurs. Chickens raised wearing artificial tails, and consequently with more posteriorly located centre of mass, showed a more vertical orientation of the femur during standing and increased femoral displacement during locomotion. Our results support the hypothesis that gradual changes in the location of the centre of mass resulted in more crouched hindlimb postures and a shift from hip-driven to knee-driven limb movements through theropod evolution. This study suggests that, through careful experimental manipulations during the growth phase of ontogeny, extant birds can potentially be used to gain important insights into previously unexplored aspects of bipedal non-avian theropod locomotion.

Concepts: Bird, Knee, Extinction, Locomotion, Dinosaur, Theropoda, Bipedalism, Archosaur

506

Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian-Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

Concepts: Continent, Pacific Ocean, Cretaceous, Antarctica, Sauropoda, Dinosaur, Titanosaur, Macronaria

412

Relationships between non-avian theropod dinosaurs and extant and fossil birds are a major focus of current paleobiological research. Despite extensive phylogenetic and morphological support, behavioural evidence is mostly ambiguous and does not usually fossilize. Thus, inferences that dinosaurs, especially theropods displayed behaviour analogous to modern birds are intriguing but speculative. Here we present extensive and geographically widespread physical evidence of substrate scraping behavior by large theropods considered as compelling evidence of “display arenas” or leks, and consistent with “nest scrape display” behaviour among many extant ground-nesting birds. Large scrapes, up to 2 m in diameter, occur abundantly at several Cretaceous sites in Colorado. They constitute a previously unknown category of large dinosaurian trace fossil, inferred to fill gaps in our understanding of early phases in the breeding cycle of theropods. The trace makers were probably lekking species that were seasonally active at large display arena sites. Such scrapes indicate stereotypical avian behaviour hitherto unknown among Cretaceous theropods, and most likely associated with terrirorial activity in the breeding season. The scrapes most probably occur near nesting colonies, as yet unknown or no longer preserved in the immediate study areas. Thus, they provide clues to paleoenvironments where such nesting sites occurred.

Concepts: Bird, Reptile, Fossil, Dinosaur, Saurischia, Theropoda, Maniraptora, Avialae

360

The fossil record of ceratopsid dinosaurs between the occurrence of their proximate sister taxa in the Turonian and the beginning of their well-documented radiation from the late Campanian of North America onwards (approximately 90 and 77 Ma) is poor, with only seven taxa described from this early period in their evolution. We describe a new taxon of a highly adorned basal centrosaurine, Wendiceratops pinhornensis gen. et sp. nov., from the lower part of the Oldman Formation (middle Campanian, approximately 78-79 Ma), Alberta, Canada. Over 200 bones derived from virtually all parts of the skeleton, including multiple well-preserved specimens of the diagnostic parietosquamosal frill, were collected from a medium-density monodominant bonebed, making the new taxon one of the best-represented early ceratopsids. The new taxon is apomorphic in having epiparietals at loci 2 and 3 developed as broad-based, pachyostotic processes that are strongly procurved anterodorsally to overhang the posterior and lateral parietal rami, and an ischium with a broad, rectangular distal terminus. Although the morphology of the nasal is incompletely known, Wendiceratops is inferred to have a large, upright nasal horn located close to the orbits, which represents the oldest occurrence of this feature in Ceratopsia. Given the phylogenetic position of the new taxon within Centrosaurinae, a enlarged nasal horn is hypothesized to have arisen independently at least twice in ceratopsid evolution.

Concepts: Phylogenetics, Cladistics, Fossil, Paleontology, Dinosaur, Taxon, Ceratopsidae, Albertaceratops

354

The running ability of Tyrannosaurus rex has been intensively studied due to its relevance to interpretations of feeding behaviour and the biomechanics of scaling in giant predatory dinosaurs. Different studies using differing methodologies have produced a very wide range of top speed estimates and there is therefore a need to develop techniques that can improve these predictions. Here we present a new approach that combines two separate biomechanical techniques (multibody dynamic analysis and skeletal stress analysis) to demonstrate that true running gaits would probably lead to unacceptably high skeletal loads in T. rex. Combining these two approaches reduces the high-level of uncertainty in previous predictions associated with unknown soft tissue parameters in dinosaurs, and demonstrates that the relatively long limb segments of T. rex-long argued to indicate competent running ability-would actually have mechanically limited this species to walking gaits. Being limited to walking speeds contradicts arguments of high-speed pursuit predation for the largest bipedal dinosaurs like T. rex, and demonstrates the power of multiphysics approaches for locomotor reconstructions of extinct animals.

Concepts: Locomotion, Dinosaur, Carnivore, Tyrannosaurus, Tyrannosauridae, Allosaurus

345

Diplodocidae are among the best known sauropod dinosaurs. Several species were described in the late 1800s or early 1900s from the Morrison Formation of North America. Since then, numerous additional specimens were recovered in the USA, Tanzania, Portugal, and Argentina, as well as possibly Spain, England, Georgia, Zimbabwe, and Asia. To date, the clade includes about 12 to 15 nominal species, some of them with questionable taxonomic status (e.g., ‘Diplodocus’ hayi or Dyslocosaurus polyonychius), and ranging in age from Late Jurassic to Early Cretaceous. However, intrageneric relationships of the iconic, multi-species genera Apatosaurus and Diplodocus are still poorly known. The way to resolve this issue is a specimen-based phylogenetic analysis, which has been previously implemented for Apatosaurus, but is here performed for the first time for the entire clade of Diplodocidae. The analysis includes 81 operational taxonomic units, 49 of which belong to Diplodocidae. The set of OTUs includes all name-bearing type specimens previously proposed to belong to Diplodocidae, alongside a set of relatively complete referred specimens, which increase the amount of anatomically overlapping material. Non-diplodocid outgroups were selected to test the affinities of potential diplodocid specimens that have subsequently been suggested to belong outside the clade. The specimens were scored for 477 morphological characters, representing one of the most extensive phylogenetic analyses of sauropod dinosaurs. Character states were figured and tables given in the case of numerical characters. The resulting cladogram recovers the classical arrangement of diplodocid relationships. Two numerical approaches were used to increase reproducibility in our taxonomic delimitation of species and genera. This resulted in the proposal that some species previously included in well-known genera like Apatosaurus and Diplodocus are generically distinct. Of particular note is that the famous genus Brontosaurus is considered valid by our quantitative approach. Furthermore, “Diplodocus” hayi represents a unique genus, which will herein be called Galeamopus gen. nov. On the other hand, these numerical approaches imply synonymization of “Dinheirosaurus” from the Late Jurassic of Portugal with the Morrison Formation genus Supersaurus. Our use of a specimen-, rather than species-based approach increases knowledge of intraspecific and intrageneric variation in diplodocids, and the study demonstrates how specimen-based phylogenetic analysis is a valuable tool in sauropod taxonomy, and potentially in paleontology and taxonomy as a whole.

Concepts: Biology, Sauropoda, Dinosaur, Diplodocus, Supersaurus, Jurassic, Brachiosaurus, Camarasaurus

335

Approximately 40% of a skeleton including cranial and postcranial remains representing a new genus and species of basal neotheropod dinosaur is described. It was collected from fallen blocks from a sea cliff that exposes Late Triassic and Early Jurassic marine and quasi marine strata on the south Wales coast near the city of Cardiff. Matrix comparisons indicate that the specimen is from the lithological Jurassic part of the sequence, below the first occurrence of the index ammonite Psiloceras planorbis and above the last occurrence of the Rhaetian conodont Chirodella verecunda. Associated fauna of echinoderms and bivalves indicate that the specimen had drifted out to sea, presumably from the nearby Welsh Massif and associated islands (St David’s Archipelago). Its occurrence close to the base of the Blue Lias Formation (Lower Jurassic, Hettangian) makes it the oldest known Jurassic dinosaur and it represents the first dinosaur skeleton from the Jurassic of Wales. A cladistic analysis indicates basal neotheropodan affinities, but the specimen retains plesiomorphic characters which it shares with Tawa and Daemonosaurus.

Concepts: United Kingdom, Wales, Dinosaur, Jurassic, Cardiff, Early Jurassic, Hettangian, Lias Group

314

Most carnivorous mammals can pulverize skeletal elements by generating tooth pressures between occluding teeth that exceed cortical bone shear strength, thereby permitting access to marrow and phosphatic salts. Conversely, carnivorous reptiles have non-occluding dentitions that engender negligible bone damage during feeding. As a result, most reptilian predators can only consume bones in their entirety. Nevertheless, North American tyrannosaurids, including the giant (13 metres [m]) theropod dinosaur Tyrannosaurus rex stand out for habitually biting deeply into bones, pulverizing and digesting them. How this mammal-like capacity was possible, absent dental occlusion, is unknown. Here we analyzed T. rex feeding behaviour from trace evidence, estimated bite forces and tooth pressures, and studied tooth-bone contacts to provide the answer. We show that bone pulverization was made possible through a combination of: (1) prodigious bite forces (8,526-34,522 newtons [N]) and tooth pressures (718-2,974 megapascals [MPa]) promoting crack propagation in bones, (2) tooth form and dental arcade configurations that concentrated shear stresses, and (3) repetitive, localized biting. Collectively, these capacities and behaviors allowed T. rex to finely fragment bones and more fully exploit large dinosaur carcasses for sustenance relative to competing carnivores.

Concepts: Dinosaur, Theropoda, Carnivore, Coelurosauria, Tyrannosauroidea, Tyrannosaurus, Tyrannosauridae, Allosaurus

313

Predator-prey dynamics are an important evolutionary driver of escalating predation mode and efficiency, and commensurate responses of prey [1-3]. Among these strategies, camouflage is important for visual concealment, with countershading the most universally observed [4-6]. Extant terrestrial herbivores free of significant predation pressure, due to large size or isolation, do not exhibit countershading. Modern predator-prey dynamics may not be directly applicable to those of the Mesozoic due to the dominance of very large, visually oriented theropod dinosaurs [7]. Despite thyreophoran dinosaurs' possessing extensive dermal armor, some of the most extreme examples of anti-predator structures [8, 9], little direct evidence of predation on these and other dinosaur megaherbivores has been documented. Here we describe a new, exquisitely three-dimensionally preserved nodosaurid ankylosaur, Borealopelta markmitchelli gen. et sp. nov., from the Early Cretaceous of Alberta, which preserves integumentary structures as organic layers, including continuous fields of epidermal scales and intact horn sheaths capping the body armor. We identify melanin in the organic residues through mass spectroscopic analyses and observe lighter pigmentation of the large parascapular spines, consistent with display, and a pattern of countershading across the body. With an estimated body mass exceeding 1,300 kg, B. markmitchelli was much larger than modern terrestrial mammals that either are countershaded or experience significant predation pressure as adults. Presence of countershading suggests predation pressure strong enough to select for concealment in this megaherbivore despite possession of massive dorsal and lateral armor, illustrating a significant dichotomy between Mesozoic predator-prey dynamics and those of modern terrestrial systems.

Concepts: Predation, Crypsis, Reptile, Mammal, Cretaceous, Dinosaur, Carnivore, Ankylosauria