A revision of the sauropod dinosaur genus ‘Bothriospondylus’ with a redescription of the type material of the Middle Jurassic form ‘B. madagascariensis’

Abstract: The sauropod dinosaur ‘Bothriospondylus’, originally named on the basis of Late Jurassic remains from England, is demonstrated to be invalid, and the characters used to diagnose it are shown to be obsolescent features which are widespread throughout Sauropoda. Material referred to this genus spans a temporal range from the Middle Jurassic until the early Late Cretaceous and has been described from five different countries, across three continents. These remains represent a wide array of sauropod groups, comprising non‐neosauropod eusauropods, a macronarian, titanosauriforms (including at least one definite brachiosaurid) and a rebbachisaurid. The type material of the Middle Jurassic ‘B. madagascariensis’ represents a derived non‐neosauropod eusauropod and possesses two potential autapomorphies. However, as a result of the fragmentary nature of the material and the uncertainty surrounding its association, a new taxon is not erected. Of the numerous specimens referred to ‘Bothriospondylus’, however, several remains are considered diagnostic: Ornithopsis hulkei (Early Cretaceous, UK), Lapparentosaurus madagascariensis (Middle Jurassic, Madagascar) and Nopcsaspondylus alarconensis (early Late Cretaceous, Argentina). At least three types of sauropod were present in the Bathonian (Middle Jurassic) of north‐west Madagascar, with a basal eusauropod (Archaeodontosaurus), a more derived eusauropod (‘B. madagascariensis’) and a titanosauriform (Lapparentosaurus) all approximately contemporaneous. Palaeocontinental reconstructions suggest that Middle Jurassic Madagascan sauropods would still have been capable of global biotic interchange, and this is perhaps reflected in their diverse assemblage. Re‐evaluation of these Malagasy forms has shed new light on this important time period in sauropod evolution.     

Temnospondyl amphibians from the Jurassic of the Southern Junggar Basin (NW China)

For the first time, remains of post-Triassic temnosponyls are described from the southern Junggar Basin (NW China). Middle Jurassic material from the uppermost part of the Toutunhe Formation consists of several isolated but well preserved skull bones and intercentra from all regions of the body that can be attributed to a brachyopid. The material in part shares important autapomorphies withGobiops desertus Shishkin, 1991, from the Upper Jurassic of Outer Mongolia, and is consequently referred to this taxon, partially with reservation. This represents the first record of this genus from China. It also yields new anatomical data and permits an emended diagnosis.Ferganobatrachus from the Middle Jurassic of Kirghisia is probably, as also shown by the new material, congeneric withGobiops. Two fragmentary specimens from the lower part of the Upper Jurassic Qigu Formation are tentatively referred to the Brachyopoidea because of features of the skull, mandible and vertebral column. They probably both represent a single taxon that was larger and more massively built thanGobiops desertus but closely related to this form.      

Sauropod dinosaurs from the Lower Cretaceous of eastern Asia: taxonomic and biogeographical implications

Sauropod dinosaurs are poorly represented in the Lower Cretaceous of eastern Asia. Here, we describe a number of isolated sauropod teeth from the Kuwajima Formation (?Berriasian–?Hauterivian) of Shiramine, Japan. The mosaic of shared derived characters and symplesiomorphies displayed by the teeth indicate that they are referable to a basal member of the titanosauriform radiation. A taxonomic review of previously described sauropod specimens from eastern and south–eastern Asia reveals that a diversity of sauropods (including a titanosaurian, a basal titanosauriform and a ?euhelopodid, as well as several forms of indeterminate systematic position) was present in this region in the Early Cretaceous. This diversity conflicts with previous suggestions that eastern Asia was biogeographically isolated from the rest of Laurasia until the late Early Cretaceous and that the sauropod fauna was limited to the endemic East Asian clade Euhelopodidae. The presence of titanosauriform sauropods in the basal Cretaceous of Japan and Thailand indicate that the proposed faunal isolation of eastern Asia ended approximately 20 myr earlier than usually suggested.     

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.     

The ‘Giant of Ksour’, a Middle Jurassic sauropod dinosaur from Algeria

Continental strata of Early and Middle Jurassic age are seldom-exposed, and little is known of the history of sauropod dinosaurs prior to the neosauropod radiation of the end of the Middle Jurassic. Here, we report, in the Middle Jurassic of the Occidental Saharan Atlas (Algerian High Atlas), the discovery of a skeleton, including cranial material, of a new cetiosaurid sauropod. Chebsaurus algeriensis n. g., n. sp. represents the most complete Algerian sauropod available to date, only few remains were found before. To cite this article: F. Mahammed et al., C. R. Palevol 4 (2005).     

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 New Basal Sauropod Dinosaur from the Middle Jurassic of Niger and the Early Evolution of Sauropoda

Background

The early evolution of sauropod dinosaurs is poorly understood because of a highly incomplete fossil record. New discoveries of Early and Middle Jurassic sauropods have a great potential to lead to a better understanding of early sauropod evolution and to reevaluate the patterns of sauropod diversification.

Principal Findings

A new sauropod from the Middle Jurassic of Niger, Spinophorosaurus nigerensis n. gen. et sp., is the most complete basal sauropod currently known. The taxon shares many anatomical characters with Middle Jurassic East Asian sauropods, while it is strongly dissimilar to Lower and Middle Jurassic South American and Indian forms. A possible explanation for this pattern is a separation of Laurasian and South Gondwanan Middle Jurassic sauropod faunas by geographic barriers. Integration of phylogenetic analyses and paleogeographic data reveals congruence between early sauropod evolution and hypotheses about Jurassic paleoclimate and phytogeography.

Conclusions

Spinophorosaurus demonstrates that many putatively derived characters of Middle Jurassic East Asian sauropods are plesiomorphic for eusauropods, while South Gondwanan eusauropods may represent a specialized line. The anatomy of Spinophorosaurus indicates that key innovations in Jurassic sauropod evolution might have taken place in North Africa, an area close to the equator with summer-wet climate at that time. Jurassic climatic zones and phytogeography possibly controlled early sauropod diversification.     

The origin and early evolution of dinosaurs

The oldest unequivocal records of Dinosauria were unearthed from Late Triassic rocks (approximately 230 Ma) accumulated over extensional rift basins in southwestern Pangea. The better known of these are Herrerasaurus ischigualastensis, Pisanosaurus mertii, Eoraptor lunensis, and Panphagia protos from the Ischigualasto Formation, Argentina, and Staurikosaurus pricei and Saturnalia tupiniquim from the Santa Maria Formation, Brazil. No uncontroversial dinosaur body fossils are known from older strata, but the Middle Triassic origin of the lineage may be inferred from both the footprint record and its sister‐group relation to Ladinian basal dinosauromorphs. These include the typical Marasuchus lilloensis, more basal forms such as Lagerpeton and Dromomeron, as well as silesaurids: a possibly monophyletic group composed of Mid‐Late Triassic forms that may represent immediate sister taxa to dinosaurs. The first phylogenetic definition to fit the current understanding of Dinosauria as a node‐based taxon solely composed of mutually exclusive Saurischia and Ornithischia was given as “all descendants of the most recent common ancestor of birds and Triceratops”. Recent cladistic analyses of early dinosaurs agree that Pisanosaurus mertii is a basal ornithischian; that Herrerasaurus ischigualastensis and Staurikosaurus pricei belong in a monophyletic Herrerasauridae; that herrerasaurids, Eoraptor lunensis, and Guaibasaurus candelariensis are saurischians; that Saurischia includes two main groups, Sauropodomorpha and Theropoda; and that Saturnalia tupiniquim is a basal member of the sauropodomorph lineage. On the contrary, several aspects of basal dinosaur phylogeny remain controversial, including the position of herrerasaurids, E. lunensis, and G. candelariensis as basal theropods or basal saurischians, and the affinity and/or validity of more fragmentary taxa such as Agnosphitys cromhallensis, Alwalkeria maleriensis, Chindesaurus bryansmalli, Saltopus elginensis, and Spondylosoma absconditum. The identification of dinosaur apomorphies is jeopardized by the incompleteness of skeletal remains attributed to most basal dinosauromorphs, the skulls and forelimbs of which are particularly poorly known. Nonetheless, Dinosauria can be diagnosed by a suite of derived traits, most of which are related to the anatomy of the pelvic girdle and limb. Some of these are connected to the acquisition of a fully erect bipedal gait, which has been traditionally suggested to represent a key adaptation that allowed, or even promoted, dinosaur radiation during Late Triassic times. Yet, contrary to the classical “competitive” models, dinosaurs did not gradually replace other terrestrial tetrapods over the Late Triassic. In fact, the radiation of the group comprises at least three landmark moments, separated by controversial (Carnian‐Norian, Triassic‐Jurassic) extinction events. These are mainly characterized by early diversification in Carnian times, a Norian increase in diversity and (especially) abundance, and the occupation of new niches from the Early Jurassic onwards. Dinosaurs arose from fully bipedal ancestors, the diet of which may have been carnivorous or omnivorous. Whereas the oldest dinosaurs were geographically restricted to south Pangea, including rare ornithischians and more abundant basal members of the saurischian lineage, the group achieved a nearly global distribution by the latest Triassic, especially with the radiation of saurischian groups such as “prosauropods” and coelophysoids.     

A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan,China fills a gap in the track record

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based on well-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected to accommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination of features, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum, others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features, the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basal sauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from the tracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects the diversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.     

A SPALACOLESTINE SPALACOTHERIID (MAMMALIA,TRECHNOTHERIA) FROM THE EARLY CRETACEOUS (BARREMIAN) OF SOUTHERN ENGLAND AND ITS BEARING ON SPALACOTHERIID EVOLUTION

Abstract: Bulk screening of Early Cretaceous (Barremian) Wealden Group strata of the Wessex Formation exposed on the south‐west and south‐east coasts of the Isle of Wight, southern England, has resulted in the recovery of fragmentary remains pertaining to a new spalacolestine spalacotheriid mammal, Yaverlestes gassoni gen. et sp. nov. These represent the first European record of the Spalacolestinae. The remains comprise a dentulous incomplete dentary and isolated upper and lower molariforms, the former representing the most substantial mammal remains yet recovered from the Wealden Group. Hitherto, six species of spalacotheriid mammal were known from the Lower Cretaceous of Europe. All are referred to the genus Spalacotherium but in the case of taxa diagnosed on the basis of isolated lower teeth and other specimens where the post‐canine dentition is incompletely known, it is now evident that these referrals should be treated with caution. Furthermore, the new Wessex Formation spalacotheriid and recently described spalacotheriids from the ?Barremian of Japan, and the Barremian and Aptian of China exhibit combinations of characters that suggest that spalacotheriids were more diverse and that their evolution was more complex than previously recognized. The systematic position of an isolated tooth from the basal Cretaceous Lourinhã Formation of Portugal is discussed and the tooth reassigned to the Spalacotheriidae. Together with the new Wessex Formation taxon, eight species are now known from the Lower Cretaceous of Europe. The discovery of a spalacolestine in the Barremian Wessex Formation supports the concept of faunal interchange between Europe, Asia and North America during the Early Cretaceous. It also supports derivation of North American spalacotheriids from a European or Eurasian ancestor.     

A new juvenile specimen of Yunnanosaurus robustus (Dinosauria: Sauropodomorpha) from Early to Middle Jurassic of Chuxiong Autonomous Prefecture,Yunnan Province,China

An almost complete skeleton with partial cranial material (ZMNH-M8739) is recovered from the Early or Middle Jurassic of southwest China. ZMNH-M8739 is identified as a juvenile individual of basal sauropodomorph dinosaur, Yunnanosaurus robustus Young, 1951. The revised diagnoses are as follows: absence of anteroposterior expansion on the medial end of astragalus and dorsoventrally compressed medium shaft of the metatarsal IV. Unfused neural arch and finely grooved long bone surface texture indicate that this individual is in the immature growth stage. ZMNH-M8739 possesses the tooth–tooth wear facet on its mesial maxillary and dentary teeth. However, the distal maxillary teeth have coarse serrations. Such a characteristic dentition could represent a unique feeding mechanism of this animal. Finally, ZMNH-M8739 constitutes a monophyletic group with Y. robustus (holotype), and Y. huangi is nesting this clade in the phylogenetic tree of the present analysis. Comparison of juvenile and adult specimen reveals distinctive growth changes of Y. robustus. This clade is positioned in an unnamed clade at a sister taxon of Sauropoda. Finally, some members of the so-called prosauropod dinosaurs constitute a monophyletic group in the present result.     

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.     

Considerations on the age of the Tiouaren Formation (Iullemmeden Basin,Niger, Africa): Implications for Gondwanan Mesozoic terrestrial vertebrate faunas

Pre-Aptian mid-Mesozoic terrestrial vertebrates from the African continent are still very poorly known. In Niger, the Tiouaren Formation in the Iullemmeden Basin has yielded dinosaur and other vertebrate remains, and this unit has been dated as Early Cretaceous, most probably pre-Aptian, on the basis of its fish fauna and geological relations to other units in the basin. A review of the fish fauna and invertebrates from this formation does not provide any evidence for such an age, and the geological relations only help to constrain the upper limit for the age of the formation (Aptian). In contrast, the described dinosaur taxa are phylogenetically nested with late Middle Jurassic to Early Late Jurassic taxa from other localities, and thus indicate a pre-Kimmeridgian, probably late Middle Jurassic age for the Tiouaren Formation. Under the assumption of such an age, the dinosaur fauna of this formation provides new insights into dinosaur faunal provincialism during the latest Middle Jurassic. Northern Gondwanan faunas of that time seem to have been different from southern Gondwanan faunas, and show closer affinities to Eurasian faunas than to the latter. A possible explanation for this might be a climatically controlled geographic barrier due to pronounced arid conditions and thus desert environments in central Gondwana during this time.     

Basal titanosauriform (Dinosauria, Sauropoda) teeth from the Lower Cretaceous Yixian Formation of Liaoning Province, China

The Yixian Formation of Liaoning Province, People's Republic of China has yielded a diverse fauna of non-avian dinosaurs, but is dominated by small-bodied taxa. Here, we describe a series of isolated teeth from the Lujiatun Beds of the formation that are referable to a basal titanosauriform sauropod. Some of the teeth possess a distinctive circular boss on the lingual surface, which suggests that they are referable to cf. Euhelopus sp. This identification provides some additional support for biostratigraphical correlations between the Jehol Group and the Mengyin Formation of Shandong Province that suggest an Early Cretaceous age for the latter unit. Moreover, the titanosauriform affinities of the teeth provide further evidence for the dominance of this sauropod clade in eastern Asia during the Cretaceous.     

The first record of Anomoepus tracks from the Middle Jurassic of Henan Province,Central China

Small, gracile mostly tridactyl tracks from the Middle Jurassic of Henan Province represent the first example of the ichnogenus Anomoepus from this region. They represent a growing number of reports (at least eight) of this ichnogenus from the Jurassic of China. In conjunction with Changpeipus and Eubrontes, they appear characteristic of known global footprint biochrons. Anomoepus indicates the presence of ornithischian dinosaurs that are often scarce or unknown from skeletal remains in coeval deposits. When first discovered, these tracks were informally referred to as bird tracks. This interpretation reflects convergence between small Jurassic Anomoepus and avian theropod tracks that are hitherto known only from the Cretaceous and the Cenozoic. However, most Anomoepus are larger and more robust than any hitherto known Mesozoic avian tracks.     

A New Basal Sauropodomorph Dinosaur from the Lower Jurassic Navajo Sandstone of Southern Utah

Background

Basal sauropodomorphs, or ‘prosauropods,’ are a globally widespread paraphyletic assemblage of terrestrial herbivorous dinosaurs from the Late Triassic and Early Jurassic. In contrast to several other landmasses, the North American record of sauropodomorphs during this time interval remains sparse, limited to Early Jurassic occurrences of a single well-known taxon from eastern North America and several fragmentary specimens from western North America.

Methodology/Principal Findings

On the basis of a partial skeleton, we describe here a new basal sauropodomorph dinosaur from the Lower Jurassic Navajo Sandstone of southern Utah, Seitaad ruessi gen. et sp. nov. The partially articulated skeleton of Seitaad was likely buried post-mortem in the base of a collapsed dune foreset. The new taxon is characterized by a plate-like medial process of the scapula, a prominent proximal expansion of the deltopectoral crest of the humerus, a strongly inclined distal articular surface of the radius, and a proximally and laterally hypertrophied proximal metacarpal I.

Conclusions/Significance

Phylogenetic analysis recovers Seitaad as a derived basal sauropodomorph closely related to plateosaurid or massospondylid ‘prosauropods’ and its presence in western North America is not unexpected for a member of this highly cosmopolitan clade. This occurrence represents one of the most complete vertebrate body fossil specimens yet recovered from the Navajo Sandstone and one of the few basal sauropodomorph taxa currently known from North America.     

The early evolution of titanosauriform sauropod dinosaurs

Titanosauriformes was a globally distributed, long‐lived clade of dinosaurs that contains both the largest and smallest known sauropods. These common and diverse megaherbivores evolved a suite of cranial and locomotory specializations perhaps related to their near‐ubiquity in Mesozoic ecosystems. In an effort to understand the phylogenetic relationships of their early (Late Jurassic–Early Cretaceous) members, this paper presents a lower‐level cladistic analysis of basal titanosauriforms in which 25 ingroup and three outgroup taxa were scored for 119 characters. Analysis of these characters resulted in the recovery of three main clades: Brachiosauridae, a cosmopolitan mix of Late Jurassic and Early Cretaceous sauropods, Euhelopodidae, a clade of mid‐Cretaceous East Asian sauropods, and Titanosauria, a large Cretaceous clade made up of mostly Gondwanan genera. Several putative brachiosaurids were instead found to represent non‐titanosauriforms or more derived taxa, and no support for a Laurasia‐wide clade of titanosauriforms was found. This analysis establishes robust synapomorphies for many titanosauriform subclades. A re‐evaluation of the phylogenetic affinities of fragmentary taxa based on these synapomorphies found no body fossil evidence for titanosaurs before the middle Cretaceous (Aptian), in contrast to previous reports of Middle and Late Jurassic forms. Purported titanosaur track‐ways from the Middle Jurassic either indicate a substantial ghost lineage for the group or – more likely – represent non‐titanosaurs. Titanosauriform palaeobiogeographical history is the result of several factors including differential extinction and dispersal. This study provides a foundation for future study of basal titanosauriform phylogeny and the origins of Titanosauria. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 624–671.     

A Re-evaluation of Small Tetrapods from the Middle Triassic Otter Sandstone Formation of Devon, England

Material of small sauropsids from the Otter Sandstone Formation of east Devon (Sherwood Sandstone Group; Middle Triassic; Anisian) includes remains that were formerly attributed to a primitive procolophonid. In the light of new specimens, this material is instead found to contain remains of a diapsid and a procolophonine procolophonid. Among these fossils, the medium-sized procolophonine, Kapes bentoni sp. nov., is the first record of this Russian genus in the British Triassic. Coartaredens isaaci gen. et sp. nov. is a small diapsid tentatively assigned to Lepidosauromorpha. The heterodont lower dentition of Coartaredens comprises a row of large, conical posterior teeth and tightly packed, procumbent incisiforms. Two additional specimens are distinguished on the basis of distinctive dentary remains. One of these is of possible procolophonid affinity, while the dentition of the second resembles that of the aberrant Early Triassic parareptilian genus Sclerosaurus .