A new phylogeny of Stegosauria (Dinosauria,Ornithischia)

The stegosaurs are some of the most easily recognizable dinosaurs, but are surprisingly rare as fossils. Consequently much remains unknown about their palaeobiology, and every new stegosaurian find contributes to our understanding of the evolution of the clade. Since the last attempt to examine the evolutionary relationships of Stegosauria, new specimens have come to light, including the most complete individual of Stegosaurus ever found, new taxa have been described and, perhaps most importantly, new methods for analysis of cladistic datasets have been produced. In the light of these new data and technological advances, the phylogenetic relationships of the stegosaurs and basal armoured dinosaurs are investigated. The inclusion of continuous data results in much better resolution than was previously obtained, and the resulting single most parsimonious tree supports re‐erection of the genera Miragaia and Hesperosaurus, which had previously been synonymized with Dacentrurus and Stegosaurus respectively. The recently described genus Alcovasaurus is resolved as a basal thyreophoran, but this is most likely a consequence of a very high degree of missing data and the questionable ontogenetic stage of the specimen. Examination of the effects of continuous data on the analysis suggest that while it contains a phylogenetic signal congruent with that of discrete data and provides better resolution than discrete data alone, it can affect topologies in unpredictable ways, particularly in areas of the tree where there are large amounts of missing data. The phylogeny presented here will form the basis for future work on the palaeobiology of the plated dinosaurs.     

Femoral ontogeny and locomotor biomechanics of Dryosaurus lettowvorbecki (Dinosauria, Iguanodontia)

Ontogenetic changes in femoral morphology and locomotion were analysed in the iguanodontian dinosaur Dryosaurus lettowvorbecki using cross-sectional data and applying principles of beam theory. The results presented here suggest that locomotor ontogeny in D. lettowvorbecki was more complicated than has generally been recognized. The percentage cortical area (a measure of the relative amount of bone) increases abruptly over a relatively short period during early ontogeny and then remains uniform during subsequent increases in body size. Modifications in cross-sectional shape also occur with increasing size, as demonstrated by differences in second moment of area ratios. The patterns of change in these properties indicate that the orientation of mechanical loadings acting on the femur of D. lettowvorbecki differed at various stages of growth and development. It is suggested that the alterations in femoral architecture described here reflect a shift from quadrupedality to bipedality early in the ontogeny of this animal.     

Craniofacial ontogeny in centrosaurine dinosaurs (Ornithischia: Geratopsidae): taxonomic and behavioral implications

Centrosaurine ceratopsians are characterized by well developed nasal horncores or bosses, relatively abbreviated supraorbital horncores or bosses, and adorned parietosquamosal frills. Recent study of several paucispecific (low diversity) bonebed assemblages in Alberta and Montana has contributed greatly to our understanding of ontogenetic and taxonomic variation in the skulls of centrosaurines. Relative age determination of centrosaurines is now possible through examination of ontogenetic change in several characters, including the surface bone morphology of specific skeletal elements. The within-group taxonomy of centrosaurines is based almost entirely on characters of the skull roof, relating particularly to horns and frills. Juvenile and sub-adult centrosaurines are characterized by relatively simple, unadorned skulls compared to their adult counterparts. As in numerous living taxa, the cranial ornaments of centrosaurines developed late in ontogeny, as individuals approached or attained adult size. An important implication arising directly from this study is that juvenile and sub-adult centrosaurines are difficult to distinguish taxonomically at the specific level. Two monospecific genera represented only by immature materials, Brachyceratops montanensis and Monoclonius crassus , cannot be defended and should be considered nomina dubia . The late ontogenetic development and diverse taxonomic variation of horn and frill morphologies support the contention diat these structures are best interpreted as reproductive characters employed in mate competition.     

Cranial anatomy and systematics of Hypacrosaurus altispinus,and a comparative analysis of skull growth in lambeosaurine hadrosaurids (Dinosauria: Ornithischia)

The cranial anatomy of the helmet‐crested lambeosaurine Hypacrosaurus altispinus (Ornithischia: Hadrosauridae) is described, with a focus on ontogenetic and individual variation in phylogenetically significant characters of the cranial crest, braincase, and facial skeleton. Cranial material of H. altispinus represents a relatively complete growth series that includes crestless juveniles of less than half the size of large individuals with fully developed crests. Cranial ontogeny is compared with other lambeosaurines using bivariate morphometrics and through qualitative comparison of a size‐standardized cranial growth series. Bivariate analyses reveal that the relative growth of the skull and cranial crest of H. altispinus and H. stebingeri are similar, and that Hypacrosaurus more closely resembles Corythosaurus than Lambeosaurus. Hypacrosaurus altispinus is systematically revised. The taxon is characterized by five autapomorphies, most of which are concentrated in the skull, as well as an enlarged terminal ischial foot. Maximum parsimony and Bayesian likelihood (Mk+gamma) phylogenetic analyses were conducted to test the monophyly of the genus. Hypacrosaurus monophyly is corroborated in light of new anatomical data. Although H. stebingeri and H. altispinus share few derived characters of the skull, the hypothesis that H. stebingeri is a metaspecies that represents the ancestor of H. altispinus cannot be rejected. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 398–434.     

The skull evolution of oviraptorosaurian dinosaurs: the role of niche partitioning in diversification

Oviraptorosaurs are bird‐like theropod dinosaurs that thrived in the final pre‐extinction ecosystems during the latest Cretaceous, and the beaked, toothless skulls of derived species are regarded as some of the most peculiar among dinosaurs. Their aberrant morphologies are hypothesized to have been caused by rapid evolution triggered by an ecological/biological driver, but little is known about how their skull shapes and functional abilities diversified. Here, we use quantitative techniques to study oviraptorosaur skull form and mandibular function. We demonstrate that the snout is particularly variable, that mandibular form and upper/lower beak form are significantly correlated with phylogeny, and that there is a strong and significant correlation between mandibular function and mandible/lower beak shape, suggesting a form–function association. The form–function relationship and phylogenetic signals, along with a moderate allometric signal in lower beak form, indicate that similar mechanisms governed beak shape in oviraptorosaurs and extant birds. The two derived oviraptorosaur clades, oviraptorids and caenagnathids, are significantly separated in morphospace and functional space, indicating that they partitioned niches. Oviraptorids coexisting in the same ecosystem are also widely spread in morphological and functional space, suggesting that they finely partitioned feeding niches, whereas caenagnathids exhibit extreme disparity in beak size. The diversity of skull form and function was likely key to the diversification and evolutionary success of oviraptorosaurs in the latest Cretaceous.     

Skull ontogeny in Arrhinoceratops brachyops (Ornithischia: Ceratopsidae) and other horned dinosaurs

Disentangling ontogenetic from interspecific variation is key to understanding biodiversity in the fossil record, yet information on growth in the ceratopsid subfamily Chasmosaurinae is sparse. Here, we describe the partial skull of a juvenile chasmosaurine, attributed to Arrhinoceratops brachyops, within the context of more mature specimens of this species, to better understand the ontogenetic transformations therein. We show that as A. brachyops matured, the postorbital horncores became longer and shifted from a posterior to an anterior inclination, the delta‐shaped frill epiossifications became lower and fused to the underlying frill, and the face became more elongate. In these respects, A. brachyops closely resembled Triceratops, suggesting that these ontogenetic changes may have been common to all long‐horned chasmosaurines. However, an event‐paired cladistic analysis of Chasmosaurinae using a standardized matrix of 24 developmental characters reveals that the relative timing of ontogenetic events in Arrhinoceratops was more like that of Chasmosaurus, particularly in the relatively late reduction in scalloping around the frill margins. Thus, the ontogenetic similarities between Arrhinoceratops and Triceratops appear to be plesiomorphic, partly related to the retention of the elongate postorbital horncores, which are primitive for Ceratopsidae. This study elucidates the otherwise contentious evolutionary relationships of Arrhinoceratops, and highlights the importance of ontogenetic data for resolving phylogenies when morphological data from adults alone are inadequate. © 2015 The Linnean Society of London     

Taxonomy,morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs

Heterodontosaurids comprise an important early radiation of small-bodied herbivores that persisted for approximately 100 My from Late Triassic to Early Cretaceous time. Review of available fossils unequivocally establishes Echinodon as a very small-bodied, late-surviving northern heterodontosaurid similar to the other northern genera Fruitadens and Tianyulong. Tianyulong from northern China has unusual skeletal proportions, including a relatively large skull, short forelimb, and long manual digit II. The southern African heterodontosaurid genus Lycorhinus is established as valid, and a new taxon from the same formation is named Pegomastax africanusgen. n., sp. n. Tooth replacement and tooth-to-tooth wear is more common than previously thought among heterodontosaurids, and in Heterodontosaurus the angle of tooth-to-tooth shear is shown to increase markedly during maturation. Long-axis rotation of the lower jaw during occlusion is identified here as the most likely functional mechanism underlying marked tooth wear in mature specimens of Heterodontosaurus. Extensive tooth wear and other evidence suggests that all heterodontosaurids were predominantly or exclusively herbivores. Basal genera such as Echinodon, Fruitadens and Tianyulong with primitive, subtriangular crowns currently are known only from northern landmasses. All other genera except the enigmatic Pisanosaurus have deeper crown proportions and currently are known only from southern landmasses.     

The role of ontogeny on character polarization in early dinosaurs: a new specimen from the Late Triassic of southern Brazil and its implications

Only recently, new ontogenetic series of early dinosaurs and related groups have been described. Here, we present an isolated immature dinosauriform femur from the Late Triassic of southern Brazil and investigate its influence on character polarization. Because the specimen shares a number of synapomorphies with Pampadromaeus barberenai, herein we postulate that it corresponds to a juvenile individual of that taxon. Accordingly, we investigate the morphological variation between juvenile and mature individuals of P. barberenai. Scoring these character states into a published phylogenetic data-set of Dinosauromorpha reveals that morphological variation is higher than that observed among closely-related taxa. Ontogenetic variation thus exerts influence on character polarization. In addition, modification of the scores affected by ontogeny produces different topologies, as noted by the reduction in both the number of most parsimonious trees and number of steps, and increased inclusivity of some clades and reduction of polytomies as well. Our study, together with other recent contributions, sheds light on the morphologic pathways seen during dinosauromorph ontogenetic development, which is crucial to more reliably assess phylogenetic reconstructions and macroevolutionary patterns of this widespread and successful group.     

Aspects of comparative cranial mechanics in the theropod dinosaurs Coelophysis, Allosaurus and Tyrannosaurus

The engineering analysis technique finite element analysis (FEA) is used here to investigate cranial stress and strain during biting and feeding in three phylogenetically disparate theropod taxa: Coelophysis bauri , Allosaurus fragilis and Tyrannosaurus rex . Stress patterns are generally similar in all taxa with the ventral region of the skull tensed whilst the dorsal aspect is compressed, although the skull is not purely behaving as a cantilever beam as there is no discernible neutral region of bending. Despite similarities, stress patterns are not wholly comparable: there are key differences in how certain regions of the skull contain stress, and it is possible to link such differences to cranial morphology. In particular, nasal morphology can be explained by the stress patterns revealed here. Tyrannosaurus models shear and compress mainly in the nasal region, in keeping with the indistinguishably fused and expanded morphology of the nasal bones. Conversely Allosaurus and Coelophysis models experience peak shear and compression in the fronto-parietal region (which is tightly interdigitated and thickened in the case of Allosaurus ) yet in contrast the nasal region is lightly stressed, corresponding to relatively gracile nasals and a frequently patent internasal suture evident in Allosaurus . Such differences represent alternate mechanical specializations between taxa that may be controlled by functional, phylogenetic or mechanical constraints. Creation of finite element models placed in a phylogenetic context permits the investigation of the role of such mechanical character complexes in the cranium of nonavian theropods and the lineage leading towards modern birds.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 309–316.     

A tall-spined ornithopod dinosaur from the Early Cretaceous of Salas de los Infantes (Burgos, Spain)

A tall-spined ornithopod dinosaur from the Pinilla de los Moros Formation (Upper Hauterivian-Lower Barremian) of Salas de los Infantes (Burgos, Spain) is described. The material consists of seven associated axial remains, including five middle dorsal vertebrae, a fragmentary neural spine and a dorsal rib, from a single medium-sized individual. This material was previously referred to Iguanodon cf. fittoni. It is characterised by having a high dorsal neurapophysis that is approximately 4.5 times the height of the centrum. The elongation and vertical orientation of the dorsal neural spines allow it to be distinguished from other ornithopods from the Wealden of Europe, including Hypselospinus and Barilium from the Valanginian, and Iguanodon and Mantellisaurus from the Barremian-Aptian. The material is here referred to Iguanodontia indet. because it is so incomplete, but it is potentially a distinct taxon. Among the ornithopods, only Ouranosaurus and the hadrosaurid Hypacrosaurus possess higher dorsal neural spines.     

The evolution of cranial form and function in theropod dinosaurs: insights from geometric morphometrics

Theropod dinosaurs, an iconic clade of fossil species including Tyrannosaurus and Velociraptor, developed a great diversity of body size, skull form and feeding habits over their 160+ million year evolutionary history. Here, we utilize geometric morphometrics to study broad patterns in theropod skull shape variation and compare the distribution of taxa in cranial morphospace (form) to both phylogeny and quantitative metrics of biting behaviour (function). We find that theropod skulls primarily differ in relative anteroposterior length and snout depth and to a lesser extent in orbit size and depth of the cheek region, and oviraptorosaurs deviate most strongly from the "typical" and ancestral theropod morphologies. Noncarnivorous taxa generally fall out in distinct regions of morphospace and exhibit greater overall disparity than carnivorous taxa, whereas large-bodied carnivores independently converge on the same region of morphospace. The distribution of taxa in morphospace is strongly correlated with phylogeny but only weakly correlated with functional biting behaviour. These results imply that phylogeny, not biting function, was the major determinant of theropod skull shape.     

A molecular phylogeny of Amazona: implications for Neotropical parrot biogeography, taxonomy, and conservation

Amazon parrots (Genus Amazona) are among the most recognizable and imperiled of all birds. Several hypotheses regarding the evolutionary history of Amazona are investigated using a combined phylogenetic analysis of DNA sequence data from six partitions including mitochondrial (COI, 12S, and 16S) and nuclear (beta-fibint7, RP40, and TROP) regions. The results demonstrate that Amazona is not monophyletic with respect to the placement of the Yellow-faced parrot (Amazona xanthops), as first implied by. In addition, the analysis corroborates previous studies suggesting a Neotropical short-tailed parrot genus as sister to Amazona. At a finer level, the phylogeny resolves the Greater Antillean endemic species as constituting a monophyletic group, including the Central American Amazona albifrons, while further revealing a paraphyletic history for the extant Amazon species of the Lesser Antilles. The reconstructed phylogeny provides further insights into the mainland sources of the Antillean Amazona, reveals areas of taxonomic uncertainty within the genus, and presents historical information that may be included in conservation priority-setting for Amazon parrots.     

The first diplodocid from Asia and its implications for the evolutionary history of sauropod dinosaurs

Abstract: An isolated anterior caudal vertebra from the Qingshan (= Ch'ing shan) Formation (Early Cretaceous) of Shandong Province, China, is redescribed and shown to be an advanced diplodocid sauropod. This specimen possesses several derived character states that are typically observed in advanced diplodocoids or diplodocids, including the following: a mildly procoelous centrum; a deep pit‐like pneumatic fossa immediately below the caudal rib; wing‐ or fan‐shaped caudal ribs; and complex lamination of the neural spine. The neural spine is apomorphically short and the centrum is short relative to its height compared to those of other diplodocids, which, when coupled with the specimen’s unique geographical location and stratigraphical age, suggests that it probably represents a new taxon. This caudal vertebra provides the first convincing evidence that diplodocids were present in Asia, perhaps as a result of the dispersal of neosauropod lineages from Europe and/or North America during the Early Cretaceous. The discovery of a member of the Diplodocidae in the Early Cretaceous also indicates that this clade did not become extinct at the Jurassic/Cretaceous boundary as previously supposed.     

A stegosaur vertebra (Dinosauria: Ornithischia) from the Callovian (Middle Jurassic) of Sarthe, western France

A dinosaur vertebra found in the course of road works in the “Chevain Marls” (marine Callovian) of the Vermont outlier, near the village of Béthon (Sarthe, Pays-de-la-Loire, western France) is described and referred to a stegosaur (cf. Lexovisaurus). It is the first record of a dinosaur in the Middle Jurassic of that area. The specimen was probably derived from the floating carcass of an animal that had lived on the emerged areas of the Armorican Massif, near which the Chevain Marls were deposited. This fossil is an addition to the short list of stegosaur remains reported from France.     

Ovule,fruit and seed development in Abolboda (Xyridaceae,Poales): implications for taxonomy and phylogeny

Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.     

Problems of ontogeny and phylogeny in brain-size evolution

Fundamental ambiguities in the interpretation of brain/body allometric trends can only be resolved by analyzing relationships between ontogenetic brain/body growth processes in different groups. The ambiguous concept of adult encephalization confuses at least three distinct types of transformation of a common mammalian growth curve: scalar magnification, total curve didplacement, and changes in proportions of the pre- and postnatal phases of the curve. The conservative ratio between pre- and postnatal growth phases determines the apparent linearity of comparative brain/body allometry and can be explained by assuming that embyological neurogenetic processes ultimately determine both target brain and body size—the first directly and the second indirectly via neurohormonal regulation of somatic growth. Uneven taxonomic distribution of different ontogenetic growth patterns may explain many differences in the allometric trends at different taxonomic levels of analysis. The human brain grows exactly as if it was in a giant ape body; however, because of decoupled growth in different brain regions, it regulates body growth as though it were the size of a chimpanzee brain. Human encephalization exhibits an ontogenetic transformation not found in other mammalian groups.