Introduction to Dinosaurs: New Ideas from Old Bones papers honoring Peter Dodson.

This special volume of The Anatomical Record honors the life-long commitment to anatomy and paleontology by Professor Peter Dodson (Emeritus, University of Pennsylvania). Peter's legacy is not only rooted in his own research interests, but also in the wealth of former students he mentored over his career, many of whom have made their own new contributions to the fields of anatomy and paleontology through original scientific investigation. Within these 18 scientific papers, which cover multiple taxa, multiple continents, and multiple methodologies, each of the contributors brought to this volume their own unique work that can be traced back to some form of inspiration by the honoree.

Iyuku raathi, a new iguanodontian dinosaur from the Early Cretaceous Kirkwood Formation, South Africa.

We name and describe a new iguanodontian dinosaur from the Early Creteceous Kirkwood Formation, Eastern Cape Province, South Africa. This dinosaur is one of only two ornithopod dinosaurs known from the Cretaceous of southern Africa, and is unique in being represented primarily by hatchling to young juvenile individuals as demonstrated by bone histological analysis. All of the juvenile material of this new taxon comes from a single, laterally-restricted bonebed and specimens were primarily recovered as partial to complete single elements, although rare articulated materials and one partial skeleton were found. Sedimentology of the bonebed suggests that this horizon heralds a change in environment upsection to a drier and more seasonal climate. This accumulation of bones is interpreted as seasonal mortality from a nesting site or nesting grounds and may be linked to this environmental shift.

Computed tomographic analysis of the dental system of three Jurassic ceratopsians and implications for the evolution of tooth replacement pattern and diet in early-diverging ceratopsians.

The dental system of ceratopsids is among the most specialized structure in Dinosauria by the presence of tooth batteries and high-angled wear surfaces. However, the origin of this unique dental system is poorly understood due to a lack of relevant knowledge in early-diverging ceratopsians. Here, we study the dental system of three earliest-diverging Chinese ceratopsians: Yinlong and Hualianceratops from the early Late Jurassic of Xinjiang and Chaoyangsaurus from the Late Jurassic of Liaoning Province. By micro-computed tomographic analyses, our study has revealed significant new information regarding the dental system, including no more than five replacement teeth in each jaw quadrant; at most one replacement tooth in each alveolus; nearly full resorption of the functional tooth root; and occlusion with low-angled, concave wear facets. Yinlong displays an increase in the number of maxillary alveoli and a decrease in the number of replacement teeth during ontogeny as well as the retention of functional tooth remnants in the largest individual. Chaoyangsaurus and Hualianceratops have slightly more replacement teeth than Yinlong. In general, early-diverging ceratopsians display a relatively slow tooth replacement rate and likely use gastroliths to triturate foodstuffs. The difference in dietary strategy might have influenced the tooth replacement pattern in later-diverging ceratopsians.

A New Taxon of Basal Ceratopsian from China and the Early Evolution of Ceratopsia.

Ceratopsia is one of the best studied herbivorous ornithischian clades, but the early evolution of Ceratopsia, including the placement of Psittacosaurus, is still controversial and unclear. Here, we report a second basal ceratopsian, Hualianceratops wucaiwanensis gen. et sp. nov., from the Upper Jurassic (Oxfordian) Shishugou Formation of the Junggar Basin, northwestern China. This new taxon is characterized by a prominent caudodorsal process on the subtemporal ramus of the jugal, a robust quadrate with an expansive quadratojugal facet, a prominent notch near the ventral region of the quadrate, a deep and short dentary, and strongly rugose texturing on the lateral surface of the dentary. Hualianceratops shares several derived characters with both Psittacosaurus and the basal ceratopsians Yinlong, Chaoyangsaurus, and Xuanhuaceratops. A new comprehensive phylogeny of ceratopsians weakly supports both Yinlong and Hualianceratops as chaoyangsaurids (along with Chaoyangsaurus and Xuanhuaceratops), as well as the monophyly of Chaoyangosauridae + Psittacosaurus. This analysis also weakly supports the novel hypothesis that Chaoyangsauridae + Psittacosaurus is the sister group to the rest of Neoceratopsia, suggesting a basal split between these clades before the Late Jurassic. This phylogeny and the earliest Late Jurassic age of Yinlong and Hualianceratops imply that at least five ceratopsian lineages (Yinlong, Hualianceratops, Chaoyangsaurus + Xuanhuaceratops, Psittacosaurus, Neoceratopsia) were present at the beginning of the Late Jurassic.

Turning semicircular canal function on its head: dinosaurs and a novel vestibular analysis.

Previous investigations have correlated vestibular function to locomotion in vertebrates by scaling semicircular duct radius of curvature to body mass. However, this method fails to discriminate bipedal from quadrupedal non-avian dinosaurs. Because they exhibit a broad range of relative head sizes, we use dinosaurs to test the hypothesis that semicircular ducts scale more closely with head size. Comparing the area enclosed by each semicircular canal to estimated body mass and to two different measures of head size, skull length and estimated head mass, reveals significant patterns that corroborate a connection between physical parameters of the head and semicircular canal morphology. Head mass more strongly correlates with anterior semicircular canal size than does body mass and statistically separates bipedal from quadrupedal taxa, with bipeds exhibiting relatively larger canals. This morphologic dichotomy likely reflects adaptations of the vestibular system to stability demands associated with terrestrial locomotion on two, versus four, feet. This new method has implications for reinterpreting previous studies and informing future studies on the connection between locomotion type and vestibular function.

New horned dinosaurs from Utah provide evidence for intracontinental dinosaur endemism.

BACKGROUND: During much of the Late Cretaceous, a shallow, epeiric sea divided North America into eastern and western landmasses. The western landmass, known as Laramidia, although diminutive in size, witnessed a major evolutionary radiation of dinosaurs. Other than hadrosaurs (duck-billed dinosaurs), the most common dinosaurs were ceratopsids (large-bodied horned dinosaurs), currently known only from Laramidia and Asia. Remarkably, previous studies have postulated the occurrence of latitudinally arrayed dinosaur "provinces," or "biomes," on Laramidia. Yet this hypothesis has been challenged on multiple fronts and has remained poorly tested. METHODOLOGY/PRINCIPAL FINDINGS: Here we describe two new, co-occurring ceratopsids from the Upper Cretaceous Kaiparowits Formation of Utah that provide the strongest support to date for the dinosaur provincialism hypothesis. Both pertain to the clade of ceratopsids known as Chasmosaurinae, dramatically increasing representation of this group from the southern portion of the Western Interior Basin of North America. Utahceratops gettyi gen. et sp. nov.-characterized by short, rounded, laterally projecting supraorbital horncores and an elongate frill with a deep median embayment-is recovered as the sister taxon to Pentaceratops sternbergii from the late Campanian of New Mexico. Kosmoceratops richardsoni gen. et sp. nov.-characterized by elongate, laterally projecting supraorbital horncores and a short, broad frill adorned with ten well developed hooks-has the most ornate skull of any known dinosaur and is closely allied to Chasmosaurus irvinensis from the late Campanian of Alberta. CONCLUSIONS/SIGNIFICANCE: Considered in unison, the phylogenetic, stratigraphic, and biogeographic evidence documents distinct, co-occurring chasmosaurine taxa north and south on the diminutive landmass of Laramidia. The famous Triceratops and all other, more nested chasmosaurines are postulated as descendants of forms previously restricted to the southern portion of Laramidia. Results further suggest the presence of latitudinally arrayed evolutionary centers of endemism within chasmosaurine ceratopsids during the late Campanian, the first documented occurrence of intracontinental endemism within dinosaurs.

A basal alvarezsauroid theropod from the early Late Jurassic of Xinjiang, China.

The fossil record of Jurassic theropod dinosaurs closely related to birds remains poor. A new theropod from the earliest Late Jurassic of western China represents the earliest diverging member of the enigmatic theropod group Alvarezsauroidea and confirms that this group is a basal member of Maniraptora, the clade containing birds and their closest theropod relatives. It extends the fossil record of Alvarezsauroidea by 63 million years and provides evidence for maniraptorans earlier in the fossil record than Archaeopteryx. The new taxon confirms extreme morphological convergence between birds and derived alvarezsauroids and illuminates incipient stages of the highly modified alvarezsaurid forelimb.

A Jurassic ceratosaur from China helps clarify avian digital homologies.

Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156-161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.

A basal ceratopsian with transitional features from the Late Jurassic of northwestern China.

Although the Ceratopsia and Pachycephalosauria, two major ornithischian groups, are united as the Marginocephalia, few synapomorphies have been identified due to their highly specialized body-plans. Several studies have linked the Heterodontosauridae with either the Ceratopsia or Marginocephalia, but evidence for these relationships is weak, leading most recent studies to consider the Heterodontosauridae as the basal member of another major ornithischian radiation, the Ornithopoda. Here, we report on a new basal ceratopsian dinosaur, Yinlong downsi gen. et. sp. nov. from the Late Jurassic upper part of the Shishugou Formation of Xinjiang, China. This new ceratopsian displays a series of features transitional between more derived ceratopsians and other ornithischians, shares numerous derived similarities with both the heterodontosaurids and pachycephalosaurians and provides strong evidence supporting a monophyletic Marginocephalia and its close relationship to the Heterodontosauridae. Character distributions along the marginocephalian lineage reveal that, compared to the bipedal Pachycephalosauria, which retained a primitive post-cranial body-plan, the dominantly quadrupedal ceratopsians lost many marginocephalian features and evolved their own characters early in their evolution.

A basal tyrannosauroid dinosaur from the Late Jurassic of China.

The tyrannosauroid fossil record is mainly restricted to Cretaceous sediments of Laurasia, although some very fragmentary Jurassic specimens have been referred to this group. Here we report a new basal tyrannosauroid, Guanlong wucaii gen. et sp. nov., from the lower Upper Jurassic of the Junggar Basin, northwestern China. G. wucaii is the oldest known tyrannosauroid and shows several unexpectedly primitive pelvic features. Nevertheless, the limbs of G. wucaii share several features with derived coelurosaurs, and it possesses features shared by other coelurosaurian clades. This unusual combination of character states provides an insight into the poorly known early radiation of the Coelurosauria. Notably, the presumed predatory Guanlong has a large, fragile and highly pneumatic cranial crest that is among the most elaborate known in any non-avian dinosaur and could be comparable to some classical exaggerated ornamental traits among vertebrates.

A Middle Jurassic 'sphenosuchian' from China and the origin of the crocodylian skull.

The skull of living crocodylians is highly solidified and the jaw closing muscles are enlarged, allowing for prey capture by prolonged crushing between the jaws. Living species are all semi-aquatic, with sprawling limbs and a broad body that moves mainly from side-to-side; however, fossils indicate that they evolved from terrestrial forms. The most cursorial of these fossils are small, gracile forms often grouped together as the Sphenosuchia, with fully erect, slender limbs; their relationships, however, are poorly understood. A new crocodylomorph from deposits in northwestern China of the poorly known Middle Jurassic epoch possesses a skull with several adaptations typical of living crocodylians. Postcranially it is similar to sphenosuchians but with even greater adaptations for cursoriality in the forelimb. Here we show, through phylogenetic analysis, that it is the closest relative of the large group Crocodyliformes, including living crocodylians. Thus, important features of the modern crocodylian skull evolved during a phase when the postcranial skeleton was evolving towards greater cursoriality, rather than towards their current semi-aquatic habitus.