Biostratinomic alterations of an Edmontosaurus "mummy" reveal a pathway for soft tissue preservation without invoking "exceptional conditions".

Removal or protection from biostratinomic agents of decomposition, such as predators and scavengers, is widely seen as a requirement for high-quality preservation of soft tissues in the fossil record. In this context, extremely rapid burial is an oft-cited mechanism for shielding remains from degradation, but not all fossils fit nicely into this paradigm. Dinosaurian mummies in particular seemingly require two mutually exclusive taphonomic processes to preserve under that framework: desiccation and rapid burial. Here we present a recently prepared Edmontosaurus mummy that reveals an alternate fossilization pathway for resistant soft tissues (e.g., skin and nails). While the skin on this specimen is well-preserved in three dimensions and contains biomarkers, it is deflated and marked by the first documented examples of injuries consistent with carnivore activity on dinosaurian soft tissue during the perimortem interval. Incomplete scavenging of the carcass provided a route for the gases, fluids, and microbes associated with decomposition to escape, allowing more durable soft tissues to persist through the weeks to months required for desiccation prior to entombment and fossilization. This pathway is consistent with actualistic observations and explains why dinosaurian skin, while rare, is more commonly preserved than expected if extreme circumstances were required for its preservation. More broadly, our assumptions guide specimen collection and research, and the presence of soft tissues and biomolecules in fossils that demonstrably were not rapidly buried, such as this mummy, suggests that such types of evidence may be substantially more common than previously assumed.

The phylogenetic nomenclature of ornithischian dinosaurs.

Ornithischians form a large clade of globally distributed Mesozoic dinosaurs, and represent one of their three major radiations. Throughout their evolutionary history, exceeding 134 million years, ornithischians evolved considerable morphological disparity, expressed especially through the cranial and osteodermal features of their most distinguishable representatives. The nearly two-century-long research history on ornithischians has resulted in the recognition of numerous diverse lineages, many of which have been named. Following the formative publications establishing the theoretical foundation of phylogenetic nomenclature throughout the 1980s and 1990s, many of the proposed names of ornithischian clades were provided with phylogenetic definitions. Some of these definitions have proven useful and have not been changed, beyond the way they were formulated, since their introduction. Some names, however, have multiple definitions, making their application ambiguous. Recent implementation of the International Code of Phylogenetic Nomenclature (ICPN, or PhyloCode) offers the opportunity to explore the utility of previously proposed definitions of established taxon names. Since the Articles of the ICPN are not to be applied retroactively, all phylogenetic definitions published prior to its implementation remain informal (and ineffective) in the light of the Code. Here, we revise the nomenclature of ornithischian dinosaur clades; we revisit 76 preexisting ornithischian clade names, review their recent and historical use, and formally establish their phylogenetic definitions. Additionally, we introduce five new clade names: two for robustly supported clades of later-diverging hadrosaurids and ceratopsians, one uniting heterodontosaurids and genasaurs, and two for clades of nodosaurids. Our study marks a key step towards a formal phylogenetic nomenclature of ornithischian dinosaurs.

The systematic relationships and biogeographic history of ornithischian dinosaurs.

The systematic relationships of taxa traditionally referred to as 'basal ornithopods' or 'hypsilophodontids' remain poorly resolved since it was discovered that these taxa are not a monophyletic group, but rather a paraphyletic set of neornithischian taxa. Thus, even as the known diversity of these taxa has dramatically increased over the past two decades, our knowledge of their placement relative to each other and the major ornithischian subclades remained incomplete. This study employs the largest phylogenetic dataset yet compiled to assess basal ornithischian relationships (255 characters for 65 species level terminal taxa). The resulting strict consensus tree is the most well-resolved, stratigraphically consistent hypothesis of basal ornithischian relationships yet hypothesized. The only non-iguanodontian ornithopod (=basal ornithopod) recovered in this analysis is Hypsilophodon foxii. The majority of former 'hypsilophodontid' taxa are recovered within a single clade (Parksosauridae) that is situated as the sister-taxon to Cerapoda. The Parksosauridae is divided between two subclades, the Orodrominae and the Thescelosaurinae. This study does not recover a clade consisting of the Asian taxa Changchunsaurus, Haya, and Jeholosaurus (=Jeholosauridae). Rather, the former two taxa are recovered as basal members of Thescelosaurinae, while the latter taxon is recovered in a clade with Yueosaurus near the base of Neornithischia.The endemic South American clade Elasmaria is recovered within the Thescelosaurinae as the sister taxon to Thescelosaurus. This study supports the origination of Dinosauria and the early diversification of Ornithischia within Gondwana. Neornithischia first arose in Africa by the Early Jurassic before dispersing to Asia before the late Middle Jurassic, where much of the diversification among non-cerapodan neornithischians occurred. Under the simplest scenario the Parksosauridae originated in North America, with at least two later dispersals to Asia and one to South America. However, when ghost lineages are considered, an alternate dispersal hypothesis has thescelosaurines dispersing from Asia into South America (via North America) during the Early Cretaceous, then back into North America in the latest Cretaceous. The latter hypothesis may explain the dominance of orodromine taxa prior to the Maastrichtian in North America and the sudden appearance and wide distribution of thescelosaurines in North America beginning in the early Maastrichtian. While the diversity of parksosaurids has greatly increased over the last fifteen years, a ghost lineage of over 40 myr is present between the base of Parksosauridae and Cerapoda, indicating that much of the early history and diversity of this clade is yet to be discovered. This new phylogenetic hypothesis provides a comprehensive framework for testing further hypotheses regarding evolutionary patterns and processes within Ornithischia.

Insight on the anatomy, systematic relationships, and age of the Early Cretaceous ankylopollexian dinosaur Dakotadon lakotaensis.

Knowledge regarding the early evolution within the dinosaurian clade Ankylopollexia drastically increased over the past two decades, in part because of an increase in described taxa from the Early Cretaceous of North America. These advances motivated the recent completion of extensive preparation and conservation work on the holotype and only known specimen of Dakotadon lakotaensis, a basal ankylopollexian from the Lakota Formation of South Dakota. That specimen (SDSM 8656) preserves a partial skull, lower jaws, a single dorsal vertebra, and two caudal vertebrae. That new preparation work exposed several bones not included in the original description and revealed that other bones were previously misidentified. The presence of extensive deformation in areas of the skull is also noted that influenced inaccuracies in prior descriptions and reconstructions of this taxon. In addition to providing an extensive re-description of D. lakotaensis, this study reviews previously proposed diagnoses for this taxon, identifies two autapomorphies, and provides an extensive differential diagnosis. Dakotadon lakotaensis is distinct from the only other ankylopollexian taxon known from the Lakota Formation, Osmakasaurus depressus, in the presence of two prominent, anteroposteriorly oriented ridges on the ventral surfaces of the caudal vertebrae, the only overlapping material preserved between these taxa. The systematic relationships of D. lakotaensis are evaluated using both the parsimony and posterior probability optimality criteria, with both sets of analyses recovering D. lakotaensis as a non-hadrosauriform ankylopollexian that is more closely related to taxa from the Early Cretaceous (e.g., Iguanacolossus, Hippodraco, and Theiophytalia) than to more basally situated taxa from the Jurassic (e.g., Camptosaurus, Uteodon). This taxonomic work is supplemented by field work that relocated the type locality, confirming its provenance from unit L2 (lower Fuson Member equivalent) of the Lakota Formation. Those data, combined with recently revised ages for the members of the Lakota Formation based on charophyte and ostracod biostratigraphy, constrain the age of this taxon to the late Valanginian to early Barremian.

Methods for the quantitative comparison of molecular estimates of clade age and the fossil record.

Approaches quantifying the relative congruence, or incongruence, of molecular divergence estimates and the fossil record have been limited. Previously proposed methods are largely node specific, assessing incongruence at particular nodes for which both fossil data and molecular divergence estimates are available. These existing metrics, and other methods that quantify incongruence across topologies including entirely extinct clades, have so far not taken into account uncertainty surrounding both the divergence estimates and the ages of fossils. They have also treated molecular divergence estimates younger than previously assessed fossil minimum estimates of clade age as if they were the same as cases in which they were older. However, these cases are not the same. Recovered divergence dates younger than compared oldest known occurrences require prior hypotheses regarding the phylogenetic position of the compared fossil record and standard assumptions about the relative timing of morphological and molecular change to be incorrect. Older molecular dates, by contrast, are consistent with an incomplete fossil record and do not require prior assessments of the fossil record to be unreliable in some way. Here, we compare previous approaches and introduce two new descriptive metrics. Both metrics explicitly incorporate information on uncertainty by utilizing the 95% confidence intervals on estimated divergence dates and data on stratigraphic uncertainty concerning the age of the compared fossils. Metric scores are maximized when these ranges are overlapping. MDI (minimum divergence incongruence) discriminates between situations where molecular estimates are younger or older than known fossils reporting both absolute fit values and a number score for incompatible nodes. DIG range (divergence implied gap range) allows quantification of the minimum increase in implied missing fossil record induced by enforcing a given set of molecular-based estimates. These metrics are used together to describe the relationship between time trees and a set of fossil data, which we recommend be phylogenetically vetted and referred on the basis of apomorphy. Differences from previously proposed metrics and the utility of MDI and DIG range are illustrated in three empirical case studies from angiosperms, ostracods, and birds. These case studies also illustrate the ways in which MDI and DIG range may be used to assess time trees resultant from analyses varying in calibration regime, divergence dating approach or molecular sequence data analyzed.

The cranial anatomy of the neornithischian dinosaur Thescelosaurus neglectus.

Though the dinosaur Thescelosaurus neglectus was first described in 1913 and is known from the relatively fossiliferous Lance and Hell Creek formations in the Western Interior Basin of North America, the cranial anatomy of this species remains poorly understood. The only cranial material confidently referred to this species are three fragmentary bones preserved with the paratype, hindering attempts to understand the systematic relationships of this taxon within Neornithischia. Here the cranial anatomy of T. neglectus is fully described for the first time based on two specimens that include well-preserved cranial material (NCSM 15728 and TLAM.BA.2014.027.0001). Visual inspection of exposed cranial elements of these specimens is supplemented by detailed CT data from NCSM 15728 that enabled the examination of otherwise unexposed surfaces, facilitating a complete description of the cranial anatomy of this species. The skull of T. neglectus displays a unique combination of plesiomorphic and apomorphic traits. The premaxillary and 'cheek' tooth morphologies are relatively derived, though less so than the condition seen in basal iguanodontians, suggesting that the high tooth count present in the premaxillae, maxillae, and dentaries may be related to the extreme elongation of the skull of this species rather than a retention of the plesiomorphic condition. The morphology of the braincase most closely resembles the iguanodontians Dryosaurus and Dysalotosaurus, especially with regard to the morphology of the prootic. One autapomorphic feature is recognized for the first time, along with several additional cranial features that differentiate this species from the closely related and contemporaneous Thescelosaurus assiniboiensis. Published phylogenetic hypotheses of neornithischian dinosaur relationships often differ in the placement of the North American taxon Parksosaurus, with some recovering a close relationship with Thescelosaurus and others with the South American taxon Gasparinisaura, but never both at the same time. The new morphological observations presented herein, combined with re-examination of the holotype of Parksosaurus, suggest that Parksosaurus shares a closer relationship with Thescelosaurus than with Gasparinisaura, and that many of the features previously cited to support a relationship with the latter taxon are either also present in Thescelosaurus, are artifacts of preservation, or are the result of incomplete preparation and inaccurate interpretation of specimens. Additionally, the overall morphology of the skull and lower jaws of both Thescelosaurus and Parksosaurus also closely resemble the Asian taxa Changchunsaurus and Haya, though the interrelationships of these taxa have yet to be tested in a phylogenetic analysis that includes these new morphological data for T. neglectus.

Crocodyliform feeding traces on juvenile ornithischian dinosaurs from the Upper Cretaceous (Campanian) Kaiparowits Formation, Utah.

Crocodyliforms serve as important taphonomic agents, accumulating and modifying vertebrate remains. Previous discussions of Mesozoic crocodyliform feeding in terrestrial and riverine ecosystems have often focused on larger taxa and their interactions with equally large dinosaurian prey. However, recent evidence suggests that the impact of smaller crocodyliforms on their environments should not be discounted. Here we present direct evidence of feeding by a small crocodyliform on juvenile specimens of a 'hypsilophodontid' dinosaur from the Upper Cretaceous (Campanian) Kaiparowits Formation of southern Utah. Diagnostic crocodyliform bite marks present on a left scapula and a right femur, as well as a partial probable crocodyliform tooth crown (ovoid in cross-section) preserved within a puncture on the right femur, comprise the bulk of the feeding evidence. Computed tomography scans of the femoral puncture reveal impact damage to the surrounding bone and that the distal tip of the embedded tooth was missing prior to the biting event. This is only the second reported incidence of a fossil crocodyliform tooth being found embedded directly into prey bone. These bite marks provide insight into the trophic interactions of the ecosystem preserved in the Kaiparowits Formation. The high diversity of crocodyliforms within this formation may have led to accentuated niche partitioning, which seems to have included juvenile dinosaurian prey.

Exploring the effects of phylogenetic uncertainty and consensus trees on stratigraphic consistency scores: a new program and a standardized method.

The stratigraphic record of first appearances provides an independent source of data for evaluating and comparing phylogenetic hypotheses that include taxa with fossil histories. However, no standardized method exists for calculating these metrics for polytomous phylogenies, restricting their applicability. Previously proposed methods insufficiently deal with this problem because they skew or restrict the resulting scores. To resolve this issue, we propose a standardized method for treating polytomies when calculating these metrics: the Comprehensive Polytomy approach (ComPoly). This approach accurately describes how phylogenetic uncertainty, indicated by polytomies, affects stratigraphic consistency scores. We also present a new program suite (Assistance with Stratigraphic Consistency Calculations) that incorporates the ComPoly approach and simplifies the calculation of absolute temporal stratigraphic consistency metrics. This study also demonstrates that stratigraphic consistency scores calculated from strict consensus trees can be overly inclusive and those calculated from less-than-strict consensus trees inaccurately describe the phylogenetic signal present in the source most-parsimonious trees (MPTs). Therefore, stratigraphic consistency scores should be calculated directly from the source MPTs whenever possible to ensure their accuracy. Finally, we offer recommendations for standardizing comparisons between molecular divergence dates and the stratigraphic record of first appearances, a promising new application of these methods. © The Willi Hennig Society 2010.

Paleogene equatorial penguins challenge the proposed relationship between biogeography, diversity, and Cenozoic climate change.

New penguin fossils from the Eocene of Peru force a reevaluation of previous hypotheses regarding the causal role of climate change in penguin evolution. Repeatedly it has been proposed that penguins originated in high southern latitudes and arrived at equatorial regions relatively recently (e.g., 4-8 million years ago), well after the onset of latest Eocene/Oligocene global cooling and increases in polar ice volume. By contrast, new discoveries from the middle and late Eocene of Peru reveal that penguins invaded low latitudes >30 million years earlier than prior data suggested, during one of the warmest intervals of the Cenozoic. A diverse fauna includes two new species, here reported from two of the best exemplars of Paleogene penguins yet recovered. The most comprehensive phylogenetic analysis of Sphenisciformes to date, combining morphological and molecular data, places the new species outside the extant penguin radiation (crown clade: Spheniscidae) and supports two separate dispersals to equatorial (paleolatitude approximately 14 degrees S) regions during greenhouse earth conditions. One new species, Perudyptes devriesi, is among the deepest divergences within Sphenisciformes. The second, Icadyptes salasi, is the most complete giant (>1.5 m standing height) penguin yet described. Both species provide critical information on early penguin cranial osteology, trends in penguin body size, and the evolution of the penguin flipper.