The appendicular skeleton of Suuwasseaemilieae (Sauropoda: Flagellicaudata) from the Upper Jurassic Morrison Formation of Montana (USA)

Appendicular elements of the sauropod dinosaur Suuwasseaemilieae, from the Upper Jurassic Morrison Formation of Montana, USA, display a peculiar mix of autapomorphic and plesiomorphic features. While more similar in overall morphology to Apatosaurus than other flagellicaudatans, the coracoid of Suuwassea lacks the quadrangular shape of Apatosaurus. The humerus of Suuwassea bears a pronounced proximal tuberculum, a feature seen elsewhere only in saltasaurine titanosaurian sauropods. The rectangular proximal articular surface of the tibia is proportioned neither like Diplodocus nor Apatosaurus type specimens, although this region may be intraspecifically variable. The pes of Suuwassea possesses plesiomorphically elongate phalanges and a small, uncompressed ungual, unlike other flagellicaudatans except Dyslocosaurus. The localization of tooth marks on the pedal elements suggests that sauropod feet may have been singled out by scavengers, as has been noted for elephants.     

Cranial biomechanics underpins high sauropod diversity in resource-poor environments

High megaherbivore species richness is documented in both fossil and contemporary ecosystems despite their high individual energy requirements. An extreme example of this is the Late Jurassic Morrison Formation, which was dominated by sauropod dinosaurs, the largest known terrestrial vertebrates. High sauropod diversity within the resource-limited Morrison is paradoxical, but might be explicable through sophisticated resource partitioning. This hypothesis was tested through finite-element analysis of the crania of the Morrison taxa Camarasaurus and Diplodocus. Results demonstrate divergent specialization, with Camarasaurus capable of exerting and accommodating greater bite forces than Diplodocus, permitting consumption of harder food items. Analysis of craniodental biomechanical characters taken from 35 sauropod taxa demonstrates a functional dichotomy in terms of bite force, cranial robustness and occlusal relationships yielding two polyphyletic functional ‘grades’. Morrison taxa are widely distributed within and between these two morphotypes, reflecting distinctive foraging specializations that formed a biomechanical basis for niche partitioning between them. This partitioning, coupled with benefits associated with large body size, would have enabled the high sauropod diversities present in the Morrison Formation. Further, this provides insight into the mechanisms responsible for supporting the high diversities of large megaherbivores observed in other Mesozoic and Cenozoic communities, particularly those occurring in resource-limited environments.     

Taphonomy of Sheridan College Quarry 1, Buffalo, Wyoming: Implications for reconstructing historic dinosaur localities including Utterback's 1902-1910 Morrison dinosaur expeditions

The Upper Jurassic Morrison Formation has yielded some of the most important, voluminous and diverse dinosaur bonebeds in western North America, yet many of its historic sites were excavated during the celebrated period of vertebrate paleontology in western North America referred to as the first and second “Great Dinosaur Rush” (1870-1910s). Because of the large quantity of fossils collected during this era, a considerable amount of data pertaining to patterns of sedimentation, preservation, and paleoecology across broad portions of the Morrison Formation (and indeed many other Mesozoic and Cenozoic units) is still poorly understood. This paper critically re-evaluates the Sheridan College Quarry 1 dinosaur bonebed which lies along the western rim of the Powder River Basin in the region of localities excavated during Utterback's expeditions in the 1900s. Sedimentologically, the bonebed is interpreted as having been formed by episodic flooding events affecting the proximal floodplain depocenter of a meandering river system. Limited evidence of bone abrasion or rounding and progressive upsection changes in bone orientations suggest that minimal transport occurred, but that at least four episodes of overbank flooding resulted in the concentration and burial of attritional, time averaged vertebrate skeletal material that accumulated in topographic lows on the floodplain. Taphonomic analysis indicates that multiple unassociated to partially associated fossil elements excavated represent at least three taxa of sauropod dinosaurs, whereas isolated elements from the site indicate the presence of several other small vertebrate taxa. This work provides significant new information not only about the Sheridan College Quarry 1, but also about local sedimentary and taphonomic conditions that were likely influential to burial and preservation of other nearby Morrison dinosaur localities in the Big Horn Mountains, most notably those excavated during the famous Utterback expeditions. This study highlights the research potential in reconstructing lost data for historic dinosaur localities.     

A theropod–sauropod track assemblage from the ?Middle–Upper Jurassic Shedian Formation at Shuangbai,Yunnan Province,China, reflecting different sizes of trackmakers: Review and new observations

A dinosaur tracksite at Hemenkou (Shuangbai County, Yunnan Province) in the ?Middle–Upper Jurassic Shedian Formation that consists mainly of gray-purple feldspathic quartz sandstones was previously reported incorrectly as being in the Lower Cretaceous Puchanghe Formation. The previous assignment is also inconsistent with two regional geological maps. Although mostly yielding poorly preserved tracks, the site nevertheless indicates a diversity of theropod and sauropod trackmakers partly consistent with the Late Jurassic body fossils from the region. Purported ornithopod are re-evaluated here as those of theropods. The theropod tracks and trackways show distinct similarities to those of the Grallator–Eubrontes plexus and can be subdivided into three morphotypes that may reflect different pes anatomy and/or substrate conditions. Two sizes of tracks (small, large) indicate the presence of different size classes or species in this area in the Late Jurassic. Similarly, the sauropod trackways document three differently sized trackmakers (small–medium–large) showing a typical wide-gauge (Brontopodus) pattern. The track record is the first evidence of theropods in the ?Middle–Late Jurassic of central Yunnan, whereas the sauropod tracks suggest a relation to the coeval basal eusauropods known from this region by skeletal remains.     

The palaeoecology of the dinosaurs of the Wessex Formation (Wealden Group, Early Cretaceous), Isle of Wight, Southern England

The Wessex Formation on the Isle of Wight yields an Early Cretaceous dinosaur fauna. Sedimentological evidence shows that this represents a mosaic of fluvial, floodplain and lacustrine environments within a relatively narrow east-west oriented valley. The vegetational cover on the alluvial plain had a savannah-or chaparral-like aspect, probably of low productivity. The relative scarcity of small aquatic vertebrates, absence of coals, abundance of oxidixed sediments and the presence of immature calcretes indicate seasonal water supply. The dinosaur taxa compising the Wessex Formation faunal assemblage represent a single palaeocommunity which inhabited the local alluvial plain, although some species may have been transient. The fauna had a relatively low diversity and this is attributed to the low productivity of the local vegetation. Iguanodontids and Hypsilophodon were the dominant elements in the fauna. In contrast to Late Jurassic dinosaur faunas, sauropods are less abundant in the Wessex Formation, although they remain taxonomically diverse. It is concluded that climatic changes which took place in the Late Jurassic or Early Cretaceous resulted in the appearance of low productivity vegetation and that this was incapable of supporting large sauropod populations.     

Sauropod tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province and the pre-Cretaceous sauropodomorph trackways from China

The Xixipo dinosaur tracksite in the Chuanjie Formation of Yunnan Province is one of the 14 Chinese sites yielding sauropod tracks from between the Triassic–Jurassic and Jurassic–Cretaceous boundaries, but is only one of the two that represent the Middle Jurassic. Although it is a small site, it adds incrementally to the overall track record of the region and allows comparison with the body fossil record and classification of the Chuanjie Formation as a Type 3b or Type 4b deposit in which both the body fossil and trace fossil record, in this case representing sauropodomorphs, are similar in composition and frequency of occurrence. We argue that the sauropod trace and body fossil records, while based on different categories of evidence, are very important. Integrating and correlating all available data from both records increases our understanding of sauropod communities, and both are equally valuable for this. In addition, we also discuss narrow to wide gauge, coeval sauropod trackways from China, and the relationship between the potential trackmaker of China's Jurassic Brontopodus-type trackways and mamenchisaurids and, beginning from the Late Jurassic, representatives of this type and titanosauriform sauropods.     

New dinosaur tracksites from the Middle Jurassic of Madagascar: ichnotaxonomical,behavioural and palaeoenvironmental implications

Abstract: New dinosaur tracksites are described from the Bajocian–Bathonian Bemaraha Formation of western Madagascar. Two track‐bearing surfaces can be followed over a distance of at least 4 km, suggesting the existence of a hitherto unrecognized megatracksite. The track assemblage is theropod dominated, but sauropod tracks also occur at one site. Qualitative and quantitative analysis of the abundant theropod track material suggests that most, if not all, theropod footprints are attributable to a single trackmaker and are referred to Kayentapus isp. Although this ichnogenus, originally described from the Lower Jurassic of North America, has never been recorded from Gondwana nor from the Middle Jurassic, track morphology strongly suggests this attribution. Palaeogeographical, sedimentological and ichnological data suggest that the dinosaur tracks formed in an intertidal to supratidal setting where the coastline influenced the preferred walking direction of the animals.     

A New Sauropod Dinosaur from the Middle Jurassic of the United Kingdom

A new record of a sauropodomorph dinosaur is here described from the Middle Jurassic (Aalenian) Saltwick Formation of Whitby (Yorkshire), UK. A single caudal vertebra represents an early sauropodomorph and signifies the earliest recognised eusauropod dinosaur from the United Kingdom. The absence of pleurocoels and a narrow, dorsoventrally deep, but craniocaudally short centrum, suggests a primitive sauropodomorph. Distinct spinopostzygopophyseal laminae rise from the lateral margins of the postzygapophyses and pass caudally along what remains of the neural spine, a character unique to a subgroup of sauropods that includes Barapasaurus, Omeisaurus and other neosauropods and eusauropods. The lack of phylogenetically robust characters in sauropod caudal vertebrae usually makes it difficult to establish affinities, but the absence of mild procoely excludes this specimen from both Diplodocoidea and Lithostrotia. The vertebra cannot be further distinguished from those of a wide range of basal sauropods, cetiosaurids and basal macronarians. However, this plesiomorphic vertebra still signifies the earliest stratigraphic occurrence for a British sauropod dinosaur.     

Thyreophoran vertebrae from the Callovian (Middle Jurassic) of the Vaches Noires cliffs (Normandy,France), with remarks on the dinosaur assemblage from the Vaches Noires

Two associated incomplete thyreophoran dorsal vertebrae from the Callovian Marnes de Dives Formation of the Vaches Noires cliffs, on the Normandy coast, are referred to an indeterminate stegosaur that appears to be different from Lexovisaurus, previously reported from the Callovian of western France. These vertebrae are the first evidence of thyreophorans from the Vaches Noires and complement the dinosaur assemblage from this locality, which hitherto consisted of several theropod taxa and an indeterminate sauropod. The dinosaur record from the Vaches Noires is heavily dominated by theropods and this imbalance is difficult to explain. A possible explanation may be that the dinosaur sample from the Vaches Noires is too small to be statistically significant and representative of the original faunal assemblage from which it is derived.     

Caudal Pneumaticity and Pneumatic Hiatuses in the Sauropod Dinosaurs Giraffatitan and Apatosaurus

Skeletal pneumaticity is found in the presacral vertebrae of most sauropod dinosaurs, but pneumaticity is much less common in the vertebrae of the tail. We describe previously unrecognized pneumatic fossae in the mid-caudal vertebrae of specimens of Giraffatitan and Apatosaurus. In both taxa, the most distal pneumatic vertebrae are separated from other pneumatic vertebrae by sequences of three to seven apneumatic vertebrae. Caudal pneumaticity is not prominent in most individuals of either of these taxa, and its unpredictable development means that it may be more widespread than previously recognised within Sauropoda and elsewhere in Saurischia. The erratic patterns of caudal pneumatization in Giraffatitan and Apatosaurus, including the pneumatic hiatuses, show that pneumatic diverticula were more broadly distributed in the bodies of the living animals than are their traces in the skeleton. Together with recently published evidence of cryptic diverticula—those that leave few or no skeletal traces—in basal sauropodomorphs and in pterosaurs, this is further evidence that pneumatic diverticula were widespread in ornithodirans, both across phylogeny and throughout anatomy.     

Aging,Maturation and Growth of Sauropodomorph Dinosaurs as Deduced from Growth Curves Using Long Bone Histological Data: An Assessment of Methodological Constraints and Solutions

Information on aging, maturation, and growth is important for understanding life histories of organisms. In extinct dinosaurs, such information can be derived from the histological growth record preserved in the mid-shaft cortex of long bones. Here, we construct growth models to estimate ages at death, ages at sexual maturity, ages at which individuals were fully-grown, and maximum growth rates from the growth record preserved in long bones of six sauropod dinosaur individuals (one indeterminate mamenchisaurid, two Apatosaurus sp., two indeterminate diplodocids, and one Camarasaurus sp.) and one basal sauropodomorph dinosaur individual (Plateosaurus engelhardti). Using these estimates, we establish allometries between body mass and each of these traits and compare these to extant taxa. Growth models considered for each dinosaur individual were the von Bertalanffy model, the Gompertz model, and the logistic model (LGM), all of which have inherently fixed inflection points, and the Chapman-Richards model in which the point is not fixed. We use the arithmetic mean of the age at the inflection point and of the age at which 90% of asymptotic mass is reached to assess respectively the age at sexual maturity or the age at onset of reproduction, because unambiguous indicators of maturity in Sauropodomorpha are lacking. According to an AIC-based model selection process, the LGM was the best model for our sauropodomorph sample. Allometries established are consistent with literature data on other Sauropodomorpha. All Sauropodomorpha reached full size within a time span similar to scaled-up modern mammalian megaherbivores and had similar maximum growth rates to scaled-up modern megaherbivores and ratites, but growth rates of Sauropodomorpha were lower than of an average mammal. Sauropodomorph ages at death probably were lower than that of average scaled-up ratites and megaherbivores. Sauropodomorpha were older at maturation than scaled-up ratites and average mammals, but younger than scaled-up megaherbivores.     

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.     

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 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.     

Large sauropod and theropod tracks from the Middle Jurassic Chuanjie Formation of Lufeng County,Yunnan Province and palaeobiogeography of the Middle Jurassic sauropod tracks from southwestern China

Tracks of large theropods and a single sauropod footprint are reported from red beds at Beikeshan locality in the Middle Jurassic Chuanjie Formation, of Lufeng County, near the large World Dinosaur Valley Park complex. The Chuanjie theropod tracks are assigned to the ichnogenus Eubrontes and the large sauropod track is given the provisional label Brontopodus. All occur as isolated tracks, i.e., trackways are not preserved. Saurischian dominated ichnofaunas are relatively common in the Jurassic of China. The producers of the Chuanjie tracks may have been similar to the basal tetanuran theropod Shidaisaurus and to mamenchisaurid sauropods, which were widely distributed throughout China, during the Jurassic, and are known from skeletal remains found in the same unit. Other potential sauropod trackmakers include titanosauriforms or as-yet-unknown basal eusauropods. The ichno- and skeletal records from the Jurassic of the Lufeng Basin are largely consistent, and both document the presence of middle-large sized theropods and sauropods.     

重庆晚侏罗世两处兽脚类足迹组合与四川盆地侏罗纪地层(英文)

重庆市已报道了4处侏罗纪的恐龙足迹点,包括下侏罗统珍珠冲组的大足化石点,该地点发现了中国迄今最古老的蜥脚类行迹;另外两处化石点(南岸和金鸡)的层位曾被视为中或上侏罗统,这里首次将其确认为上侏罗统上沙溪庙组;第4处化石点(成渝), 本文全部作者目前都未曾观察,这里不做描述。南岸化石点是南岸重庆足迹(Chongqingpus nananensis)模式标本的所在地,位于重庆市中心,曾发现过大量兽脚类足迹,如今已消逝在城市化进程中。幸运的是,这批标本被保存于重庆自然博物馆,本文作者曾前后两次对其进行研究,并得出文中的结论。南岸重庆足迹为中型足迹(平均长约29 cm), 最可能被卡岩塔足迹(Kayentapus)所囊括,这批标本中有一些保存着边界不清的拇趾迹。该化石点的其他足迹被归于似异样龙足迹(cf. Anomoepus)。其他较小的、来自重庆市周边中侏罗统中部的足迹此前已被归于跷脚龙足迹(Grallator)。金鸡化石点保存了一条孤立的兽脚类行迹,因其粗壮的特征而被暂时归于似窄足龙足迹(cf. Therangospodus)。金鸡点的行迹也保存了非连续出现的拇趾迹。虽然重庆市和四川盆地及其周边更广泛地区之侏罗纪地层发现的兽脚类足迹日渐增多,但要明确其确切地质年龄及有效的足迹分类依然需要进一步工作。这是因为侏罗纪的恐龙足迹尽管丰富,但显示出连续变化的形态学特征,而保存状况的变化更使其难以在时间与空间上做出定义与鉴别。     

Giants on the landscape: modelling the abundance of megaherbivorous dinosaurs of the Morrison Formation (Late Jurassic,western USA)

The ecosystem impact of megaherbivorous dinosaurs of the Morrison Formation would have depended on their abundance (number of animals per unit of habitat area) on the landscape. We constrain Morrison megaherbivore abundance by modelling dinosaur abundance in terms of carrying capacity (K), average body mass (ABM) and animal's energy needs. Two kinds of model are presented: ‘demand-side’ models that estimate K in terms of the aggregate energy demand of the dinosaur community, and ‘supply-side’ models that estimate K in terms of retrodicted primary productivity. Baseline values of K, ABM and energy needs for the models are further derived from comparisons with modern large herbivores, and from the composition of the megaherbivore fauna from a particular stratigraphic interval of the Morrison, but in all models a broad range of fractions and multiples of these baseline parameters are considered. ‘Best-guess’ estimates of Morrison megaherbivore abundance suggest an upper limit of a few hundred animals across all taxa and size classes per square kilometre, and up to a few tens of individuals of large subadults and adults.     

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.