An early-diverging iguanodontian (Dinosauria: Rhabdodontomorpha) from the Late Cretaceous of North America.

Intensifying macrovertebrate reconnaissance together with refined age-dating of mid-Cretaceous assemblages in recent decades is producing a more nuanced understanding of the impact of the Cretaceous Thermal Maximum on terrestrial ecosystems. Here we report discovery of a new early-diverging ornithopod, Iani smithi gen. et sp. nov., from the Cenomanian-age lower Mussentuchit Member, Cedar Mountain Formation of Utah, USA. The single known specimen of this species (NCSM 29373) includes a well-preserved, disarticulated skull, partial axial column, and portions of the appendicular skeleton. Apomorphic traits are concentrated on the frontal, squamosal, braincase, and premaxilla, including the presence of three premaxillary teeth. Phylogenetic analyses using parsimony and Bayesian inference posit Iani as a North American rhabdodontomorph based on the presence of enlarged, spatulate teeth bearing up to 12 secondary ridges, maxillary teeth lacking a primary ridge, a laterally depressed maxillary process of the jugal, and a posttemporal foramen restricted to the squamosal, among other features. Prior to this discovery, neornithischian paleobiodiversity in the Mussentuchit Member was based primarily on isolated teeth, with only the hadrosauroid Eolambia caroljonesa named from macrovertebrate remains. Documentation of a possible rhabdodontomorph in this assemblage, along with published reports of an as-of-yet undescribed thescelosaurid, and fragmentary remains of ankylosaurians and ceratopsians confirms a minimum of five, cohabiting neornithischian clades in earliest Late Cretaceous terrestrial ecosystems of North America. Due to poor preservation and exploration of Turonian-Santonian assemblages, the timing of rhabdodontomorph extirpation in the Western Interior Basin is, as of yet, unclear. However, Iani documents survival of all three major clades of Early Cretaceous neornithischians (Thescelosauridae, Rhabdodontomorpha, and Ankylopollexia) into the dawn of the Late Cretaceous of North America.

Estimating ancient biogeographic patterns with statistical model discrimination.

The geographic ranges in which species live is a function of many factors underlying ecological and evolutionary contingencies. Observing the geographic range of an individual species provides valuable information about these historical contingencies for a lineage, determining the distribution of many distantly related species in tandem provides information about large-scale constraints on evolutionary and ecological processes generally. We present a linear regression method that allows for the discrimination of various hypothetical biogeographical models for determining which landscape distributional pattern best matches data from the fossil record. The linear regression models used in the discrimination rely on geodesic distances between sampling sites (typically geologic formations) as the independent variable and three possible dependent variables: Dice/Sorensen similarity; Euclidean distance; and phylogenetic community dissimilarity. Both the similarity and distance measures are useful for full-community analyses without evolutionary information, whereas the phylogenetic community dissimilarity requires phylogenetic data. Importantly, the discrimination method uses linear regression residual error to provide relative measures of support for each biogeographical model tested, not absolute answers or p-values. When applied to a recently published dataset of Campanian pollen, we find evidence that supports two plant communities separated by a transitional zone of unknown size. A similar case study of ceratopsid dinosaurs using phylogenetic community dissimilarity provided no evidence of a biogeographical pattern, but this case study suffers from a lack of data to accurately discriminate and/or too much temporal mixing. Future research aiming to reconstruct the distribution of organisms across a landscape has a statistical-based method for determining what biogeographic distributional model best matches the available data.

Citizen Science in Postsecondary Education: Current Practices and Knowledge Gaps.

Citizen science involves the public in science to investigate research questions. Although citizen science facilitates learning in informal educational settings, little is known about its use or effects in postsecondary (college or university) settings. Using a literature review and a survey, we describe how and why citizen science is being used in postsecondary courses, as well as the impacts on student learning. We found that citizen science is used predominantly in biologically related fields, at diverse types of institutions, to improve student engagement and expose students to authentic research. Considerable anecdotal evidence supporting improved student learning from these experiences exists, but little empirical evidence exists to warrant any conclusion. Therefore, there is a need to rigorously assess the relationship between citizen science participation and postsecondary student learning. We highlight considerations for instructors planning to incorporate citizen science and for citizen science projects wanting to facilitate postsecondary use.

A new brachylophosaurin (Dinosauria: Hadrosauridae) from the Upper Cretaceous Menefee Formation of New Mexico.

Brachylophosaurini is a clade of hadrosaurid dinosaurs from the Campanian of western North America. Although well-known from northern localities in Montana and Alberta, including abundant material of Brachylophosaurus canadensis and Maiasaura peeblesorum and the holotypes of Acristavus gagslarsoni and Probrachylophosaurus bergei, material from southern localities in Utah and Colorado is restricted to a partial skull referred to A. gagslarsoni and several indeterminate specimens. Here we describe Ornatops incantatus gen. et sp. nov., a new brachylophosaurin known from a partial skeleton from the Allison Member of the Menefee Formation in New Mexico. Ornatops is the first brachylophosaurin reported from New Mexico and the southernmost occurrence of the clade. Ornatops shares with Probrachylophosaurus and Brachylophosaurus a caudally expanded nasofrontal suture on the frontals, but also exhibits an autapomorphic nasofrontal suture morphology, with a horizontal rostral region and elevated caudal region with two prominent parasagittal bumps, which is different from other brachylophosaurin specimens, including juvenile and adult Brachylophosaurus. A phylogenetic analysis places Ornatops in a trichotomy with Probrachylophosaurus and Brachylophosaurus, with Maiasaura and Acristavus as successive outgroups.

Description and rediagnosis of the crested hadrosaurid (Ornithopoda) dinosaur Parasaurolophus cyrtocristatus on the basis of new cranial remains.

For nearly 60 years, skulls of Parasaurolophus species have been differentiated primarily on the basis of crest shape rather than on unique morphologic characters of other cranial elements. Complicating matters is the fact that crests dramatically change shape throughout ontogeny. Without a complete growth series, it has become difficult to assess the taxonomic distinctness of each species through the lens of allometric growth. Parasaurolophus cyrtocristatus has proven to be especially troublesome to assess because of the poorly preserved nature of the type and only skull. A new, partial skull from the Fossil Forest Member of the Fruitland Formation-the same geologic unit as the type specimen-is the first opportunity to re-diagnose this species as well as redefine the genus with many new traits. An undescribed, short-crested subadult skull from the Kaiparowits Formation of Utah previously assigned to cf. P. cyrtocristatus allows detailed comparisons to be made between the unnamed Utah taxon and the material of this species from the type locality. We find that several characteristics of the squamosal, supraoccipital, and premaxilla shared between the referred skull and the type skull are unique to P. cyrtocristatus (senso stricto) within the genus, irrespective of the overall crest shape. A phylogenetic analysis that includes six new characters posits that P. cyrtocristatus and P. tubicen are sister taxa, and that the latter does not share a closest common ancestor with the long-crested P. walkeri as previously hypothesized. This result helps to explain why both taxa are found in northeastern New Mexico, USA and in sequential geologic units (Fruitland Formation and Kirtland Formation, respectively). Additionally, the exquisitely preserved new skull provides the first opportunity to unequivocally identify the osteological make-up of the Parasaurolophus cranial crest. Unlike in previous reconstructions, the crest composition in Parasaurolophus follows what is seen in other lambeosaurines such as Corythosaurus, where the dorsal process of the premaxilla dominates the crest, with the nasal forming 80% of the ventral paired tubes, and the lateral premaxillary process acting a lateral cover between the dorsal and ventral tubes. The skull of P. cyrtocristatus is still incompletely known, so more complete material will likely reveal new features that further differentiate this species and aid in determining the pace of ornamental crest evolution.

Diminutive fleet-footed tyrannosauroid narrows the 70-million-year gap in the North American fossil record.

To date, eco-evolutionary dynamics in the ascent of tyrannosauroids to top predator roles have been obscured by a 70-million-year gap in the North American (NA) record. Here we report discovery of the oldest Cretaceous NA tyrannosauroid, extending the lineage by ~15 million years. The new taxon-Moros intrepidus gen. et sp. nov.-is represented by a hind limb from an individual nearing skeletal maturity at 6-7 years. With a ~1.2-m limb length and 78-kg mass, M. intrepidus ranks among the smallest Cretaceous tyrannosauroids, restricting the window for rapid mass increases preceding the appearance of colossal eutyrannosaurs. Phylogenetic affinity with Asian taxa supports transcontinental interchange as the means by which iconic biotas of the terminal Cretaceous were established in NA. The unexpectedly diminutive and highly cursorial bauplan of NA's earliest Cretaceous tyrannosauroids reveals an evolutionary strategy reliant on speed and small size during their prolonged stint as marginal predators.

A new microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain Formation: insights into the paleobiodiversity and paleobiogeography of early Late Cretaceous ecosystems in western North America.

The vertebrate fauna of the Late Cretaceous Mussentuchit Member of the Cedar Mountain Formation has been studied for nearly three decades, yet the fossil-rich unit continues to produce new information about life in western North America approximately 97 million years ago. Here we report on the composition of the Cliffs of Insanity (COI) microvertebrate locality, a newly sampled site containing perhaps one of the densest concentrations of microvertebrate fossils yet discovered in the Mussentuchit Member. The COI locality preserves osteichthyan, lissamphibian, testudinatan, mesoeucrocodylian, dinosaurian, metatherian, and trace fossil remains and is among the most taxonomically rich microvertebrate localities in the Mussentuchit Member. To better refine taxonomic identifications of isolated theropod dinosaur teeth, we used quantitative analyses of taxonomically comprehensive databases of theropod tooth measurements, adding new data on theropod tooth morphodiversity in this poorly understood interval. We further provide the first descriptions of tyrannosauroid premaxillary teeth and document the earliest North American record of adocid remains, extending the appearance of this ancestrally Asian clade by 5 million years in western North America and supporting studies of pre-Cenomaninan Laurasian faunal exchange across Beringia. The overabundance of mesoeucrocodylian remains at the COI locality produces a comparatively low measure of relative biodiversity when compared to other microvertebrate sites in the Mussentuchit Member using both raw and subsampling methods. Much more microvertebrate research is necessary to understand the roles of changing ecology and taphonomy that may be linked to transgression of the Western Interior Seaway or microhabitat variation.

A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia.

We describe a new iguanodontian ornithopod, Choyrodon barsboldi gen. et sp. nov. from the Albian-aged Khuren Dukh Formation of Mongolia based on several partial skeletons interpreted to represent a subadult growth stage based on osteohistological features. This new taxon is diagnosed by many autapomorphies of the maxilla, nasal, lacrimal, opisthotic, predentary, and surangular. Choyrodon displays an unusual combination of traits, possessing an open antorbital fenestra (a primitive ornithopod trait) together with derived features such as a downturned dentary and enlarged narial fenestra. Histological imaging suggests that the type specimen of Choyrodon would have been a subadult at the time of death. Phylogenetic analysis of two different character matrices do not posit Choyrodon to be the sister taxon or to be more primitive than the iguanodontian Altirhinus kurzanovi, which is found in the same formation. The only resolved relationship of this new taxon is that it was hypothesized to be a sister-taxon with the North American species Eolambia caroljonesa. Though discovered in the same formation and Choyrodon being smaller-bodied than Altirhinus, it does not appear that the former species is an ontogimorph of the latter. Differences in morphology and results of the phylogenetic analyses support their distinction although more specimens of both species will allow better refinement of their uniqueness.

Anatomy, taphonomy, and phylogenetic implications of a new specimen of Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation, Utah, USA.

BACKGROUND: Eolambia caroljonesa is the most abundant dinosaur in the lower Cenomanian Mussentuchit Member of the Cedar Mountain Formation of Utah, and one of the most completely known non-hadrosaurid iguanodontians from North America. In addition to the large holotype and paratype partial skulls, copious remains of skeletally immature individuals, including three bonebeds, have been referred to E. caroljonesa. Nevertheless, aspects of the postcranial anatomy of this taxon, particularly the pelvic girdle, have remained ambiguous due to the lack of associated postcranial material of larger, more mature individuals. METHODOLOGY/PRINCIPAL FINDINGS: Here we describe a recently discovered associated partial postcranial skeleton of a large Eolambia caroljonesa. This specimen, FMNH PR 3847, provides new anatomical data regarding the vertebral column and pelvic girdle, supplementing previous diagnoses and descriptions of E. caroljonesa. A new phylogenetic analysis incorporating information from FMNH PR 3847 places E. caroljonesa as a basal hadrosauromorph closely related to Protohadros byrdi from the Cenomanian Woodbine Formation of Texas. Histological analysis of FMNH PR 3847 reveals that it represents a subadult individual eight to nine years of age. Taphonomic analysis indicates that FMNH PR 3847 was preserved in a crevasse splay deposit, along with an unusual abundance of small crocodylomorph material. CONCLUSIONS/SIGNIFICANCE: FMNH PR 3847 provides a wealth of new morphological data, adding to the anatomical and systematic characterization of Eolambia caroljonesa, and histological data, revealing new information on growth history in a basal hadrosauromorph. Taphonomic characterization of FMNH PR 3847 and associated vertebrate material will allow comparison with other vertebrate localities in the Mussentuchit Member of the Cedar Mountain Formation.

Bony cranial ornamentation linked to rapid evolution of gigantic theropod dinosaurs.

Exaggerated cranial structures such as crests and horns, hereafter referred to collectively as ornaments, are pervasive across animal species. These structures perform vital roles in visual communication and physical interactions within and between species. Yet the origin and influence of ornamentation on speciation and ecology across macroevolutionary time scales remains poorly understood for virtually all animals. Here, we explore correlative evolution of osseous cranial ornaments with large body size in theropod dinosaurs using a phylogenetic comparative framework. We find that body size evolved directionally toward phyletic giantism an order of magnitude faster in theropod species possessing ornaments compared with unadorned lineages. In addition, we find a body mass threshold below which bony cranial ornaments do not originate. Maniraptoriform dinosaurs generally lack osseous cranial ornaments despite repeatedly crossing this body size threshold. Our study provides novel, quantitative support for a shift in selective pressures on socio-sexual display mechanisms in theropods coincident with the evolution of pennaceous feathers.

Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria).

Definitive therizinosaurid cranial materials are exceptionally rare, represented solely by an isolated braincase and tooth in the North American taxon Nothronychus mckinleyi, the remarkably complete skull of the Asian taxon Erlikosaurus andrewsi, and the lower hemimandibles of Segnosaurus galbinensis. To date, comprehensive descriptions of the former taxa are published; however, the mandibular materials of S. galbinensis have remained largely understudied since their initial description in 1979. Here we provide a comprehensive description of the well-preserved hemimandibles and dentition of S. galbinensis (MPC-D 100/80), from the Upper Cretaceous Bayanshiree Formation, Gobi Desert, Mongolia. The subrectangular and ventrally displaced caudal hemimandible, extreme ventral deflection of the rostral dentary, and edentulism of the caudal dentary of S. galbinensis are currently apomorphic among therizinosaurians. Unique, unreported dental traits including lingually folded mesial carinae, development of a denticulated triangular facet on the distal carinae near the cervix, and extracarinal accessory denticles, suggest a highly specialized feeding strategy in S. galbinensis. The presence of triple carinae on the distalmost lateral tooth crowns is also unique, although may represent an abnormality. Contrasted with the simplistic dentition of the contemporaneous therizinosaurid E. andrewsi, the dentition of S. galbinensis is indicative of niche partitioning in food acquisition, processing, or resources among known therizinosaurids inhabiting Asian ecosystems in the Late Cretaceous. Although not quantitatively correlated with diet, this suite of specializations is otherwise unique among theropod dinosaurs and supports derived inferences of facultative or obligate herbivory in therizinosaurids, ultimately adding novel information to our understanding of ecomorphology in theropods.

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.

Mountain building triggered late cretaceous North American megaherbivore dinosaur radiation.

Prior studies of Mesozoic biodiversity document a diversity peak for dinosaur species in the Campanian stage of the Late Cretaceous, yet have failed to provide explicit causal mechanisms. We provide evidence that a marked increase in North American dinosaur biodiversity can be attributed to dynamic orogenic episodes within the Western Interior Basin (WIB). Detailed fossil occurrences document an association between the shift from Sevier-style, latitudinally arrayed basins to smaller Laramide-style, longitudinally arrayed basins and a well substantiated decreased geographic range/increased taxonomic diversity of megaherbivorous dinosaur species. Dispersal-vicariance analysis demonstrates that the nearly identical biogeographic histories of the megaherbivorous dinosaur clades Ceratopsidae and Hadrosauridae are attributable to rapid diversification events within restricted basins and that isolation events are contemporaneous with known tectonic activity in the region. SymmeTREE analysis indicates that megaherbivorous dinosaur clades exhibited significant variation in diversification rates throughout the Late Cretaceous. Phylogenetic divergence estimates of fossil clades offer a new lower boundary on Laramide surficial deformation that precedes estimates based on sedimentological data alone.