A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia

Although the group played an important role in the evolution of Late Mesozoic terrestrial ecosystems, the early evolutionary history of the ornithischian dinosaurs remains poorly understood. Here, we report on a new primitive ornithischian, Eocursor parvus gen. et sp. nov., from the Late Triassic (?Norian) Lower Elliot Formation of South Africa. Eocursor is known from a single specimen comprising substantial cranial and postcranial material and represents the most complete Triassic member of Ornithischia, providing the earliest evidence for the acquisition of many key ornithischian postcranial characters, including an opisthopubic pelvis. A new phylogenetic analysis positions this taxon near the base of Ornithischia, as the sister taxon to the important and diverse clade Genasauria. The problematic clade Heterodontosauridae is also positioned basal to Genasauria, suggesting that an enlarged grasping manus may represent a plesiomorphic ornithischian condition. This analysis provides additional phylogenetic support for limited ornithischian diversity during the Late Triassic, and suggests that several major ornithischian clades may have originated later than generally believed. There are few morphological differences between Late Triassic and Early Jurassic ornithischians, supporting previous suggestions that the Early Jurassic ornithischian radiation may simply represent the filling of vacant ecological space following Late Triassic terrestrial extinctions.     

Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65–75 cm in total body length and 0.5–0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.     

Tooth function and replacement in early Mesozoic ornithischian dinosaurs: implications for aestivation

Thulborn (1978, Leihaia II ) suggests that ornithischian dinosaurs of the upper Stormberg Series (Late Triassic-Early Jurassic) of southern Africa underwent aestivation during an annual dry season. His argument, based on an interpretation of tooth function and replacement in heterodontosaurids, is: (1) unequivocal evidence of tooth replacement is not seen, and (2) piecemeal replacement of the dentition would be incompatible with maintenance of a fore-aft grinding function of the teeth; therefore, the entire dentition must have been rapidly replaced as a unit during periods of non-feeding, i.e. during aestivation. However, study of tooth wear patterns in Lanasaurus, Lycorhinus , and Heterodontosaurus show that jaw movements during mastication were orthal (open-and-close) and lateral to medial, not forwards and backwards. Differences in degree of tooth wear would not interfere with masticatory movements. Patterns of differential wear indicate that tooth replacement was not periodic but continuous, as in other reptiles. Zahnreihen , with a Z-spacing of about 3.0, are recognizable. Replacement ceased in mature individuals. The dentition shows adaptations for prolonging its effective life despite heavy wear. Differential tooth wear is incompatible with the idea of replacement of the entire dentition as a unit during an hypothesized period of aestivation. Thulborn's suggestion of aestivation in fabrosaurid ornithischians is also shown to be unlikely.     

Reconsidering the status and affinities of the ornithischian dinosaur Tatisaurus oehleri Simmons, 1965

The early Mesozoic fossil fauna collected from the Lower Lufeng Formation of Yunnan Province, China, has attracted considerable interest and attention since its discovery in the late 1930s. Its importance reflected a combination of its comparatively remote geographical position and, more particularly, the similarities of its fauna compared with approximately contemporary discoveries from Europe, North and South America, and southern Africa. The fragmentary and poorly preserved Lufeng ornithischian dinosaur Tatisaurus oehleri was described in 1965 and proved taxonomically and systematically enigmatic from the start. Originally assigned, with some noted ambivalence, to the basal ('primitive') group of ornithischians known as hypsilophodontids, since 1965 Tatisaurus has been variously ignored, assigned to a more rigorously defined Hypsilophodontidae, referred to both of the armoured (thyreophoran) ornithischian dinosaur clades (Stegosauria and Ankylosauria), or referred to a more basal position within the thyreophoran lineage. In 1996 the holotype of Tatisaurus was renamed Scelidosaurus oehleri , and the genus Scelidosaurus was proposed as an index fossil of the ' Scelidosaurus biochron' with the potential to be used for the global stratigraphic correlation of Early Jurassic (early Sinemurian) rocks. Because of this chequered history Tatisaurus oehleri Simmons, 1965 has been re-examined and is redescribed so that its taxonomic status and systematic position could be reassessed. Tatisaurus is identified as a basal thyreophoran (armoured ornithischian dinosaur); there is no basis for amalgamating it in synonymy with the genus Scelidosaurus , and the proposed creation of a ' Scelidosaurus biochron' for the purposes of biostratigraphic correlation of Lower Jurassic outcrops has no utility whatever. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 865–874.     

A new basal hadrosauroid dinosaur from the Late Cretaceous of Uzbekistan and the early radiation of duck-billed dinosaurs

Levnesovia transoxiana gen. et sp. nov., from the Late Cretaceous (Middle–Late Turonian) of Uzbekistan, is the oldest well-documented taxon referable to Hadrosauroidea sensu Godefroit et al. It differs from a somewhat younger and closely related Bactrosaurus from Inner Mongolia (China) by a tall sagittal crest on the parietals and the absence of club-shaped dorsal neural spines in adult specimens. Levnesovia, Bactrosaurus and possibly Gilmoreosaurus represent the earliest radiation of Hadrosauroidea, which took place during the Cenomanian–Turonian and possibly in North America. The second, Santonian-age radiation of Hadrosauroidea included Aralosaurus, Hadrosauridae and lineages leading to Tanius (Campanian) and Telmatosaurus (Maastrichtian). Hadrosauridae appears to be monophyletic, but Hadrosaurinae and Lambeosaurinae originated in North America and Asia, respectively.     

The Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton & Charig, 1962: cranial anatomy,functional morphology,taxonomy, and relationships

The cranial anatomy of the Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton & Charig, 1962 is described in detail for the first time on the basis of two principal specimens: the holotype (SAM‐PK‐K337) and referred skull (SAM‐PK‐K1332). In addition several other specimens that have a bearing on the interpretation of the anatomy and biology of Heterodontosaurus are described. The skull and lower jaw of Heterodontosaurus are compact and robust but perhaps most notable for the heterodont dentition that merited the generic name. Details of the cranial anatomy are revealed and show that the skull is unexpectedly specialized in such an early representative of the Ornithischia, including: the closely packed, hypsodont crowns and ‘warping’ of the occlusal surfaces (created by progressive variation in the angulation of wear on successive crowns) seen in the cheek dentition; the unusual sutural relationships between the bones along the dorsal edge of the lower jaw; the very narrow, deeply vaulted palate and associated structures on the side wall of the braincase; and the indications of cranial pneumatism (more commonly seen in basal archosaurs and saurischian dinosaurs). Evidence for tooth replacement (which has long been recognized, despite frequent statements to the contrary) is suggestive of an episodic, rather than continuous, style of tooth replacement that is, yet again, unusual in diapsids generally and particularly so amongst ornithischian dinosaurs. Cranial musculature has been reconstructed and seems to conform to that typically seen in diapsids, with the exception of the encroachment of M. adductor mandibulae externus superficialis across the lateral surface of the temporal region and external surface of the lower jaw. Indications, taken from the unusual shape of the occlusal surfaces of the cheek dentition and jaw musculature, are suggestive of a novel form of jaw action in this dinosaur. The taxonomy of currently known late Karoo‐aged heterodontosaurids from southern Africa is reviewed. Although complicated by the inadequate nature of much of the known material, it is concluded that two taxa may be readily recognized: H. tucki and Abrictosaurus consors. At least one additional taxon is recognized within the taxa presently named Lanasaurus and Lycorhinus; however, both remain taxonomically problematic and their status needs to be further tested and may only be resolved by future discoveries. The only other named taxon, Geranosaurus atavus, represents an invalid name. The recognition of at least four distinct taxa indicates that the heterodontosaurids were speciose within the late Karoo ecosystem. The systematics of Heterodontosaurus and its congeners has been analysed, using a restricted sample of taxa. A basal (nongenasaurian) position within Ornithischia is re‐affirmed. There are at least four competing hypotheses concerning the phylogenetic placement of the Heterodontosauridae, so the evidence in support of the various hypotheses is reviewed in some detail. At present the best‐supported hypothesis is the one which places Heterodontosauridae in a basal (non‐genasaurian) position; however, the evidence is not fully conclusive and further information is still needed in respect of the anatomy of proximate outgroups, as well as more complete anatomical details for other heterodontosaurids. Heterodontosaurids were not such rare components of the late Karoo ecosystem as previously thought; evidence also suggests that from a phylogenetic perspective they occupied a potentially crucial position during the earliest phases of ornithischian dinosaur evolution. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011.     

Footprints pull origin and diversification of dinosaur stem lineage deep into Early Triassic

The ascent of dinosaurs in the Triassic is an exemplary evolutionary radiation, but the earliest phase of dinosaur history remains poorly understood. Body fossils of close dinosaur relatives are rare, but indicate that the dinosaur stem lineage (Dinosauromorpha) originated by the latest Anisian (ca 242-244 Ma). Here, we report footprints from the Early-Middle Triassic of Poland, stratigraphically well constrained and identified using a conservative synapomorphy-based approach, which shifts the origin of the dinosaur stem lineage back to the Early Olenekian (ca 249-251 Ma), approximately 5-9 Myr earlier than indicated by body fossils, earlier than demonstrated by previous footprint records, and just a few million years after the Permian/Triassic mass extinction (252.3 Ma). Dinosauromorph tracks are rare in all Polish assemblages, suggesting that these animals were minor faunal components. The oldest tracks are quadrupedal, a morphology uncommon among the earliest dinosauromorph body fossils, but bipedality and moderately large body size had arisen by the Early Anisian (ca 246 Ma). Integrating trace fossils and body fossils demonstrates that the rise of dinosaurs was a drawn-out affair, perhaps initiated during recovery from the Permo-Triassic extinction.     

First Jurassic dinosaur tracksite from Guizhou Province,China: morphology,trackmaker and paleoecology

Abstract

Sauropod tracks from the Early Jurassic Maanshan Member of the Ziliujing Formation in the Dazhuanwan area of Guizhou represent the first Jurassic dinosaur track record for this province. The best preserved and longest trackway is narrow gauge (Brontopodus type), and indicates a relatively small trackmaker (footprint length ~35.0 cm). In conjunction with sauropodomorph skeletal remains these tracks suggest that basal sauropodomorphs and primitive sauropods coexisted in this region during the Early Jurassic. This pattern is same in the Lufeng and Sichuan basins. Thus, sauropod tracks from southwest China are diverse in the Early Jurassic, and include narrow gauge Parabrontopodus, wide gauge Brontopodus-type, and basal sauropodomorph tracks.     

A Middle Jurassic abelisaurid from Patagonia and the early diversification of theropod dinosaurs

Abelisaurids are a clade of large, bizarre predatory dinosaurs, most notable for their high, short skulls and extremely reduced forelimbs. They were common in Gondwana during the Cretaceous, but exceedingly rare in the Northern Hemisphere. The oldest definitive abelisaurids so far come from the late Early Cretaceous of South America and Africa, and the early evolutionary history of the clade is still poorly known. Here, we report a new abelisaurid from the Middle Jurassic of Patagonia, Eoabelisaurus mefi gen. et sp. nov., which predates the so far oldest known secure member of this lineage by more than 40 Myr. The almost complete skeleton reveals the earliest evolutionary stages of the distinctive features of abelisaurids, such as the modification of the forelimb, which started with a reduction of the distal elements. The find underlines the explosive radiation of theropod dinosaurs in the Middle Jurassic and indicates an unexpected diversity of ceratosaurs at that time. The apparent endemism of abelisauroids to southern Gondwana during Pangean times might be due to the presence of a large, central Gondwanan desert. This indicates that, apart from continent-scale geography, aspects such as regional geography and climate are important to reconstruct the biogeographical history of Mesozoic vertebrates.     

云南早侏罗世一新具嵴冠兽脚类恐龙

报道了具嵴冠的兽脚类恐龙一新属种,安龙堡双柏龙(Shuangbaisaurus anlongbaoensis gen.et sp.nov.).双柏龙发现于云南省楚雄彝族自治州双柏县下侏罗统冯家河组中,保存了部分头骨带下颌.双柏龙沿两侧眼眶背缘向上有嵴冠发育,这在其他兽脚类中未曾报道过.与其他早侏罗世体型较大且具一对矢状嵴冠的兽脚类(双嵴龙属和中国龙属)相比,双柏龙还独具一些特征组合,如相对较高的前颌骨体、抬高的前颌骨腹缘、后腹向延伸的轭骨后突及较小的上颞孔.比较研究表明,尽管中国“双嵴龙”(“Dilophosaurus”sinensis)可能应归入中国龙属(Sinosaurus),但未必属于模式种.双柏龙的发现将有助于研究基干兽脚类的演化,尤其是各类头骨骨饰在其中的作用.     

准噶尔盆地克拉美丽地区的鸟脚类

本文记述了一个小型鸟脚类——五彩湾工部龙(新种)Gongbusaurus wucaiwanensis sp.nov.,讨论了它们的分类位置和产出地层时代。     

A new basal ornithopod dinosaur (Frenchman Formation,Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America

A small, articulated basal ornithopod skeleton from the Frenchman Formation (late Maastrichtian) of Saskatchewan (RSM P 1225.1), previously referred to the taxon Thescelosaurus, differs from both recognized species of this taxon (Thescelosaurus neglectus and Thescelosaurus garbanii). The differences are taxonomically informative and we recognize this specimen as the holotype of a new species, Thescelosaurus assiniboiensis sp. nov. , diagnosed by the presence of two autapomorphies, and displaying plesiomorphic traits more similar to those of Parksosaurus, than to those of the other Thescelosaurus species. The Frenchman Formation also harbours an intriguing faunal assemblage in which Thescelosaurus represents one of the most abundant dinosaur taxa, and preserves a relatively high proportion of small (putatively juvenile and subadult) specimens of many dinosaur taxa. Further work that increases the faunal sample from this formation, and that permits quantitative comparisons with contemporary formations, will determine whether or not these differences are well supported, and will determine their ultimate palaeobiological significance. Identification of a third species of Thescelosaurus from the late Maastrichtian of North America suggests that this taxon was more diverse than previously recognized, and shows an increase in diversity from the Campanian through the late Maastrichtian, contrasting the trends seen in most other ornithischian clades. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1157–1198.     

The dinosaur tracksite of Loulle (early Kimmeridgian; Jura,France)

More than 1500 sauropod and theropod tracks have been discovered and excavated on the floor of a disused 3800 m² quarry, near the village of Loulle (French Jura). These levels correspond to a tidal-flat environment, from intertidal to supratidal zone, dated from the earliest Kimmeridgian (∼157 M.a.). Eighteen sauropod trackways have been recognized, ranging from 3.8 m to 51.5 m in length. These trackways do not correspond to a unique herd passage, because trackways are diversely oriented, most of them are crossing others and prints are more or less deeply marked, depending of different stages of substratum competence. According to the trackway gauges, the three types (narrow-, medium and wide-gauge) are represented. Some trackways could be referred to the ichnogenus Brontopodus Farlow et al., 1989, and others to Parabrontopodus Lockley et al., 1994a. Based on footprint size, the body-sizes of these 18 sauropod trackways range from tiny (Pes Length < 25 cm) to large individuals (PL > 75 cm), showing a regular continuum of sizes. Several biometrical ratios classically used in sauropod ichnology appear to be closely related to size (e.g., the smallest individuals apparently moved faster than the largest; large individuals are all narrow-gauge while small individuals are all wide-gauge; large specimens are characterized by a high heteropody ratio when the smallest show a low heteropody ratio). Associated with these sauropod trackways, four theropod trackways have also been recorded. Three small to medium-sized theropod are referred to the ichnogenus Carmelopodus Lockley et al., 1998a, while a large individual (PL = 77 cm) is tentatively referred to Megalosauripus Lessertisseur, 1955.     

Changes in the ammonite taxonomical diversity gradient during the late Jurassic–early Cretaceous

Aim Changes in the latitudinal gradient of taxonomic diversity of ammonites have been studied in four time slices of the late Jurassic to early Cretaceous (1) to assess the correspondence of the distribution with the general idea of a simple gradient of diversity decreasing polewards, and (2) to offer tentative interpretations of these patterns with respect to palaeoclimatic conditions. Locations The localities selected lie within the Boreal palaeohemisphere and most are concentrated between 10° and 45° N. Methods The distributions of genus‐level palaeobiodiversity data have been plotted on palaeogeographical maps. Localities were selected on the basis of criteria that emphasize the accuracy of the rock dating. Palaeolatitudinal positions were obtained using software developed by one of us (BV). Data are largely derived from the literature and data sets have been built for each of the four time slices selected. Information on the palaeoenvironmental context has been included, with observations divided between four palaeoenvironmental systems: epicontinental platforms, intracratonic basins, transitional areas and distal epioceanic basins. Results The distribution plots demonstrate changes in the steepness of the latitudinal gradient of palaeobiodiversity through time. In general the lowest diversity values are recorded in mid‐ to high‐palaeolatitudes and in epicontinental platform settings of all latitudes, whereas the highest diversities typically correspond to low‐latitude basins. However, in most cases it is difficult to determine whether the poleward decrease of diversity values is a reflection of the palaeoenvironmental context or simply conforms to the diversity gradient model. All patterns are influenced by sampling, and no data were available for equatorial and high palaeolatitudes. Main conclusions Continental plate movements did not affect the shaping of latitudinal patterns of ammonite diversity and the palaeoenvironmental context is the most important controlling factor. For similar latitudes, the highest diversities occur in basin settings, suggesting that depth is an important influence on diversity distribution. Provincialism was enhanced by the sea‐level fall at the end of the Jurassic that fragmented and restricted ammonite biotas. The effects of climatic fluctuations on the steepness of diversity gradients are probably significant but are currently difficult to demonstrate.     

Geographic control on phenotype expression. The case of Hybonoticeras mundulum (Oppel) from the Mexican Altiplano

Hybonoticeras mundulum (Oppel) (m) s.s. is reported from the Mexican Altiplano on the basis of material collected bed-by-bed in sections in the States of Durango and Zacatecas. The known range of this species in the Mexican Altiplano is interpreted to be uppermost Kimmeridgian (upper to uppermost Beckeri Zone) to lowermost Tithonian (basal to lower Hybonotum Zone). The Mexican specimens studied are the most complete and valuable collection of H. mundulum (Oppel) s.s. known from a given area, and reveal that Mexican populations show phenotypic features different from European ones. Vicariant events accord with data available about the areal and biostratigraphic distribution of this species, as well as with the combination of allocyclic and autocyclic factors influencing Mexican seas, the breaking of populations rather than colonization events, and the impoverished ammonite assemblages showing endemic traits within the stratigraphic interval studied.     

Evolutionary patterns of Clavatoraceae (Charophyta) in the mesogean basins analysed according to environmental change during Malm and lower cretaceous

Evolutionary patterns of Clavatoraceae during the Malm and the Lower Cretaceous can be understood by considering how palaeoecological constraints of these charophytes were affected by palaeoenvironmental change. Speciation of Clavatoraceae reached maxima in the Tithonian and especially in the Lower Barremian, coinciding with an important areal extension and environmental diversification of freshwater swamps. Extinction reached a peak in the Upper Barremian, coinciding with the marine flooding of freshwater environments in Mesogea, and continued through the Aptian and Albian due to substitution of carbonatic freshwater swamps by terrigenous deltaic environments and probably by development of highly competitive aquatic flora of angiosperms. Anagenetic change within species attained maxima during the Berriasian and Lower Barremian, when freshwater environments became extensively developed. Absence of change (stasis) was marked during the Valanginian and Hauterivian, in a geological context of environmental stability and areal reduction of the freshwater environments.     

A tyrannosauroid dinosaur from the Upper Jurassic of Portugal

Fragmentary theropod remains from the Upper Jurassic (Kimmeridgian) of Guimarota, Portugal, represent a new taxon of theropod dinosaurs, Aviatyrannis jurassica gen. et sp. nov. Together with Stokesosaurus from the Morrison Formation of North America, Aviatyrannis represents the oldest known tyrannosauroid, indicating that tyrannosauroid origins may be found in the Middle–Late Jurassic of Europe/North America. Furthermore, current evidence suggests that early tyrannosaurs were rather small animals, which is in general accordance with their origin amongst the generally rather small coelurosaurs.     

Longevity and growth rate estimates for a polar dinosaur: a Pachyrhinosaurus (Dinosauria: Neoceratopsia) specimen from the North Slope of Alaska showing a complete developmental record

Our knowledge of growth dynamics in large ceratopsian dinosaurs is very poor, in part, due to the paucity of quantifiable age markers such as growth lines in their bones. We sought marker-based, osteohistological evidence for ceratopsid age structure from high Arctic paleolatitudes based on the observations that: (1) extant mammals from high latitudes better express growth lines in their hard tissues than those from lower latitudes, and (2) the occurrence of accentuated growth banding in teeth from Arctic dinosaurs. We examined the long bones in the specimens of Pachyrhinosaurus sp. from the early Maastrichtian of northern Alaska, and found conspicuous osseous banding. Histological analysis of the spacing, structure, pattern and numbers of these bands in the femur of a very large specimen suggests that they are not a taphonomic artefact, rather they appear to reflect annual growth cycling. Counts and measurements of the growth zones suggest that the animal showed rapid linear growth early in ontogeny, sexual maturity in perhaps the 9th year of life, and that it died in the 19th year of life. Our data adds to a growing body of evidence that a genetically distinct northern dinosaurian fauna existed at high paleolatitudes in Alaska during the Late Cretaceous.