A Middle Jurassic dinosaur trackway site from Oxfordshire, UK

The Middle Jurassic (Bathonian) Ardley trackway site in Oxfordshire, UK is described in detail. The track site is extensive, containing over 40 more-or-less continuous theropod and sauropod trackways preserved together on a single bedding plane with some trackways up to 180 m in length. The trackways display reasonable preservation, with claw marks discernible. Sauropod trackways are by far the most abundant and are representative of taxa that exhibit both narrow and wide-gauge styles of locomotion. Theropod trackways are represented by large tridactyl prints and claw impressions that display exceptionally low pace angulation values during their walking phase. One theropod trackway is unique in that it reveals a gait transition associated with a temporary increase of speed. These dinosaurs were walking over a lime-mud firmground to hardground that was emergent, or locally very shallowly submerged. GPS (Global Positioning Satellite) data reveal that the tracks, preserved during a short time interval, trend in a north-easterly direction. Multi-herding behaviour is suggested for the sauropods of differing size. The discovery of this site adds significantly to our knowledge of the taxonomic composition of Middle Jurassic ecosystems. Evidence concerning locomotor styles employed by saurischian dinosaurs, footprint identification, potential trackmaker, social interactions and observations concerning estimates of Middle Jurassic faunal diversity, as well as aspects of dinosaur locomotor evolution are discussed.     

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.     

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.     

A theropod tooth from the Late Triassic of southern Africa

An isolated, large recurved and finely serrated tooth found associated with the prosauropodEuskelosaurus from the Late Triassic part of the Elliot Formation is described here. It is compared to the Triassic thecodonts and carnivorous dinosaurs and its possible affinity is discussed. The tooth possibly belongs to a basal theropod and shows some features similar to the allosauroids. This tooth is of significance, as dinosaur remains except for some footprints and trackways, are poorly known in the Late Triassic horizons of southern Africa.     

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。     

Lavinipes Cheminii Ichnogen., Ichnosp. nov., A Possible Sauropodomorph Track from the Lower Jurassic of the Italian Alps

A new ichnotaxon is described from the Lower Jurassic (Upper Hettangian-Lower Sinemurian) carbonate tidal flats on the central-eastern Italian Alps. The narrow-gauge trackway is that of a large quadrupedal dinosaur. The pes is functionally tetradactyl with three rounded antero-medially directed digits, and the manus is pentadactyl. This quadrupedal form is close to Otozoum and Pseudotetrasauropus jaquesi both traditionally related to sauropodomorph trackmakers. The similarity with Otozoum is so marked that Lavinipes and Otozoum could be cogeneric. But the overall evidence today is that the Otozoum trackmaker was generally bipedal, whereas the trackmaker of L. cheminii is fully quadrupedal. The manual prints of L. cheminii show five short clawless digits and are different from the tetradactyl slender toed manual prints of Otozoum. The possible sauropodomorph affinity of the L. cheminii trackmaker is here discussed with an attempt to a revision of the Late Triassic-Jurassic tracks which have been traditionally related to sauropod and prosauropod.     

A dinosaur ichnocoenosis from the middle jurassic of Yorkshire,UK

An assemblage of dinosaur tracks from the undersurface of a sandstone bed in the Saltwick Formation (Middle Jurassic) of Yorkshire shows a range of morphological types, preservational variants and behavioral styles. The tracks are combinations of transmitted prints and underprints and include three distinct trackways. One trackway was made by an animal walking on exposed damp sand, another was left by an animal swimming diagonally across a current and being swept down current, while the third may have been made by an animal either running or pushing off with its feet as it drifted down current. Environmental conditions that existed during the formation of the trackways varied between crevasse splay floods and exposed damp sand. The morphology of the swimming tracks is sufficiently distinctive to warrant the erection of a new ichnogenus and ichnospecies—Characichnos tridactylus. Previously described material from Kansas, Wyoming and New Mexico can be attributed to this new ichnogenus, while specimens from Germany and Utah are only provisionally referred to it. This indicates a known range from Triassic to Cretaceous.     

Two new dinosaur tracksites from the Lower Cretaceous Jiaguan Formation of Sichuan Basin,China: specific preservation and ichnotaxonomy

Two new dinosaur tracksites are reported from the Lower Cretaceous Jiaguan Formation in the Sichuan Basin, Qijiang District of Chongqing. These are the Gaoqing-Yongsheng and the Huibu tracksites, which represent the 13th and 14th reports from this formation. The Gaoqing-Yongsheng tracksite reveals the trackway of a large biped (ornithopod) in association with isolated sauropod tracks and large indeterminate undertracks with radial cracks. These features are preserved as natural casts with pebble infillings in a coarse, cross bedded and very thick bedded sandstone sequence. The Huibu tracksite reveals isolated theropod tracks and ornithopod tracks, the latter having a quadripartite, Caririchnium-like morphology, preserved in a thin bedded sandstone sequence with intercalated mudstone.     

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.     

Tracking a Legend: An Early Cretaceous Sauropod Trackway from Zhaojue County,Sichuan Province,Southwestern China

Cretaceous tetrapod (dinosaur and pterosaur) tracks from Zhaojue County in Sichuan Province are locally very abundant. Large scale quarrying operations at the Sanbiluoga Copper Mine site have produced extensive exposures, and track material for detailed study. However, natural track-bearing outcrops also occur at a site in Jiefang Township. The traditions of the local Yi people, indigenous to the area, attribute such tracks to Zhigealu, a central creator hero-ancestor, who made the footprints while riding his heavenly steed through the area. Through seeing tracks exposed by quarrying the local people offered these legend-based interpretations, and reported the Jiefang site which was previously unknown to scientists from outside the area. Thus, it is important to pay attention to local legends about track makers since they may lead directly to significant fossil footprint discoveries. Thereby paleontology and ichnological research can benefit largely from archeological sciences as well as from oral narratives from the local people. The recently discovered sauropod trackway from Jiefang is an excellent example. It comprises 16 pes-manus sets arranged in a narrow-gauge pattern. A peculiarity is the combination of this feature with morphological characteristics known from typical wide-gauge Brontopodus trackways suggesting a tentative assignment to cf. Brontopodus. The discovery enlarges the distribution and diversity of Brontopodus-like trackways and their producers in the Cretaceous Sichuan Basin.     

A diversified vertebrate ichnite fauna from the Dasheng Group (Lower Cretaceous) of southeast Shandong Province,China

The Qingquan dinosaur tracksite, from the Lower Cretaceous Dasheng Group, Shandong Province, China adds to the growing record of saurischian-dominated ichnofaunas of the region. The site reveals the presence of avian theropods (Koreanaornis) and non-avian theropods tentatively referred to Jialingpus. Sauropod tracks are referred to Brontopodus. One site shows evidence of extensive trampling attributable to sauropods and theropods that moved in the same westerly direction, though not necessarily at exactly the same time. This site is reminiscent of the famous Davenport Ranch site in Texas which has provoked much debate about the herding behavior of sauropods as a defense strategy against theropod predators.     

Reconstruction of Middle Jurassic Dinosaur-Dominated Communities from the Vertebrate Ichnofauna of the Cleveland Basin of Yorkshire,UK

Globally, skeletal remains of dinosaurs are particularly rare throughout much of the Middle Jurassic. Thus, other sources of evidence, and most importantly ichnofaunas, are important indicators of the contemporary terrestrial vertebrate communities. The outcrops of the Ravenscar Group (Aalenian—Bajocian) within the Cleveland Basin of Yorkshire, UK, which have recently been recognised as a megatracksite of global significance, provide one such major source of ichnofaunal information of this age. A comprehensive database on the variety and occurrence of dinosaur and other vertebrate traces within the Ravenscar Group has been built from a long-term and detailed study of the sequence. Thirty different and distinct morphotypes of vertebrate traces have been recognised and are being analysed and further differentiated morphometrically. Some of the morphotypes represent behavioural, preservational and perhaps ontogenetic variants of other morphotypes, but nevertheless the range of quadrupedal and bipedal prints allows an overall fauna of sauropod, stegosaurian, ornithopod and theropod dinosaurs along with crocodiles, pond turtles and fish to be reconstructed. The distribution and abundance of prints and print types within the succession shows evidence of environmental control on the behaviour and distribution of the vertebrates. Case studies highlight both the advantages and disadvantages of this type of data in reconstructing palaeocommunities.     

Telling Apart Ornithopod and Theropod Trackways: A Closer Look at a Large,Late Jurassic Tridactyl Dinosaur Trackway at Serwah,Republic of Yemen

A large bipedal tridactyl dinosaur trackway from the Late Jurassic of Serwah, near Madar, Arhab district, Republic of Yemen, has been attributed to an ornithopod trackmaker. As the distinction between theropod and ornithopod dinosaurs can pose a challenge, we present additional data to support and reconfirm the previous attribution.     

The First Iguanodontid Dinosaur Tracks from the Swiss Alps (Schrattenkalk Formation,Aptian)

Until now dinosaur tracks from Switzerland were only known from Triassic and Late Jurassic strata. We report here for the first time the occurrence of ornithopod tracks from the Schrattenkalk Formation (Late Aptian) from the Swiss Central Alps. The locality is situated in an abandoned quarry on the shore of Lake Lucerne close to the village of Beckenried. The steeply inclined surface has more than 50 tracks (in three trackways) of ornithopod dinosaurs that are attributed to iguanodontids. Three trackways can be followed for distances of 25 to 35 m. The lengths the footprints (mean: 30 cm) point to animals ranging in size of from 4 to 6 m, with estimated hip heights between 1.8 and 2 m (hip height 6 FL) and 1.4 to 1.7 m (hip height 5 FL). One of the trackways shows two succesive manus impressions, indicating facultative quadrupedal gait. The track bearing layer consists of shallow water micrites with traces of emersion, and it is overlain by bioclastic grainstones. Previously the Upper Schrattenkalk Member in the Helvetic realm was thought to have formed on a large shelf far away from any continents. The present discovery will shed new light on the paleogeographic position of the Helvetic nappes.     

A palaeoequatorial ornithischian and new constraints on early dinosaur diversification

Current characterizations of early dinosaur evolution are incomplete: existing palaeobiological and phylogenetic scenarios are based on a fossil record dominated by saurischians and the implications of the early ornithischian record are often overlooked. Moreover, the timings of deep phylogenetic divergences within Dinosauria are poorly constrained owing to the absence of a rigorous chronostratigraphical framework for key Late Triassic–Early Jurassic localities. A new dinosaur from the earliest Jurassic of the Venezuelan Andes is the first basal ornithischian recovered from terrestrial deposits directly associated with a precise radioisotopic date and the first-named dinosaur from northern South America. It expands the early palaeogeographical range of Ornithischia to palaeoequatorial regions, an area sometimes thought to be devoid of early dinosaur taxa, and offers insights into early dinosaur growth rates, the evolution of sociality and the rapid tempo of the global dinosaur radiation following the end-Triassic mass extinction, helping to underscore the importance of the ornithischian record in broad-scale discussions of early dinosaur history.     

Unusually thick dinosaur eggshell fragments from the Spanish Late Cretaceous

Fieldwork carried out recently in the southeastern branch of the Iberian Range (Valencia Province, Spain) has led to the collection of a large volume of dinosaur eggshell fragments of unusual thickness. These specimens, up to 4.9 mm thick, were recovered from palustrine grey marls of the upper Campanian-lower Maastrichtian Sierra Perenchiza Formation, which comprises a wetland paleoenvironment deposit. These eggshell fragments have a characteristic compactituberculate ornamentation, dinosauroid-spherulitic organisation, and exhibit a complex canaliculate respiratory system. The external tuberculate surface of the shell as well as the internal microstructure enable referral to Megaloolithus aff. siruguei, the most common megaloolithid oospecies known from the Iberian Peninsula and southern France. The biostratigraphic range of M. siruguei matches the temporal distribution of titanosaurid dinosaurs across the Iberian Range, tentatively considered to be potential producers.     

The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru,Tanzania

Theropod dinosaurs from the Late Jurassic of Gondwana are still poorly known, with Elaphrosaurus bambergi Janensch, 1920, from the late Kimmeridgian of Tendaguru, Tanzania, being the only taxon represented by more than isolated remains from Africa. Having long been considered a coelurosaurian, more specifically an ornithomimosaur, Elaphrosaurus is currently regarded as a basal ceratosaur. Here, we revise the osteology and phylogenetic position of this important taxon. Elaphrosaurus shows many unusual osteological characters, including extremely elongated and constricted cervical vertebrae, an expansive shoulder girdle with strongly modified forelimbs, a relatively small ilium, and elongate hindlimbs with a very small ascending process of the astragalus that is fused to the tibia. We found this taxon to share many derived characters with noasaurids, such as: strongly elongate cervical and dorsal vertebrae; low, rectangular neural spines in the mid‐caudal vertebrae; presence of only an anterior centrodiapophyseal lamina in anterior caudal vertebrae; presence of a wide, U–shaped notch between the glenoid and the anteroventral hook in the coracoid; a laterally flared postacetabular blade of the ilium; a flat anterior side of the distal tibia; and a reduced shaft of metatarsal II. Our analysis placed Elaphrosaurus within a dichotomous Noasauridae as part of a Jurassic subclade, here termed Elaphrosaurinae, that otherwise includes taxa from eastern Asia. These results underscore the long and complex evolutionary history of abelisauroids, which is still only beginning to be understood.     

The non-avian theropod track Jialingpus from the Cretaceous of the Ordos Basin,China, with a revision of the type material: Implications for ichnotaxonomy and trackmaker morphology

Deposits from the Ordos Basin of mid-western China are rich in body fossils and ichnofossils of Early Cretaceous vertebrates. Thousands of Early Cretaceous sauropod, theropod and bird tracks described since 1958 have been found at several localities in the basin. We report two new sites (Dijiaping and Bawangzhuang) in the Luohe Formation of the Ordos Basin, Shaanxi Province, which contain small theropod footprints that are here referred to the ichnogenus Jialingpus. The assignment is based on pad configurations including (1) the large metatarsophalangeal area positioned in line with the axis of digit III, (2) the subdivision of this part into a small pad behind digit II, which in some specimens is close to the general position of the hallux (digit I), and a large metatarsophalangeal pad behind digit IV, and (3) a distinct inter-pad space between metatarsophalangeal pads and proximal phalangeal pads of digits II and III. We re-describe the type material of the type ichnospecies Jialingpus yuechiensis from the Upper Jurassic Penglaizhen Formation of Sichuan Province, proposing a largely amended diagnosis for this ichnotaxon. The presence of a digit I trace in the holotype, indicating a relatively long hallux, and the large metatarsophalangeal area positioned in line with digit III distinguishes Jialingpus from the ichnogenus Grallator and similar tracks that all lack these features. The only difference between Jialingpus specimens from the Cretaceous of the Ordos Basin and those of the Jurassic Penglaizhen Formation is the larger digit divarication in the Cretaceous taxon. This is the fourth record of Jialingpus in China and the second in Cretaceous strata, with the first being those from the Huangyangquan locality in Xinjiang, China.     

New Dinosaur Footprints from the Upper Cretaceous of Poland in the Light of Paleogeographic Context

The Late Cretaceous track assemblage of southeastern Poland includes intriguing ichnotaxa with Asian or North American affinities. New finds of the ceratopsian-like ichnite and small hadrosauroid track refer directly to this problem, whereas the re-evaluation of foraminifer taxa estimates the stratigraphic positions of the track-bearing horizon and allowed to understand paleogeographic context of the studied and adjacent areas. The results demonstrate that during the Late Cretaceous times, migration of dinosaurs was possible from Asia through the East European Land to Kukeritz Island and the islands at peri-Tethys area.     

Theropod and Ornithischian Footprints from the Middle Jurassic Yanan Formation of Zizhou County,Shaanxi, China

New tracksites reported from the Zizhou area elucidate the nature of the Early-Middle Jurassic dinosaurian ichnofaunas in Shaanxi Province. The assemblage is composed of footprints and trackways of medium- to large-sized theropods that show similarities with both the ichnogenera Kayentapus and Eubrontes and of small bipedal ornithischians that are referred to AS Anomoepus isp. Additionally tracks of a quadruped are present and assigned to Deltapodus isp. that may be related to a stegosaurian. Anomoepus isp. is similar to the holotype of Shensipus tungchuanensis which is, although apparently lost, re-assigned here and considered to be a subjective junior synonym of Anomoepus. It is therefore placed in new combination as Anomoepus tungchuanensis comb. nov. Identical tracks have been reported from well-preserved trackways both in the Zizhou and Shenmu areas, where they also co-occur with theropod tracks (Kayentapus and Grallator) and tracks of quadrupedal ornithischians (Shenmuichnus and Deltapodus). Thus, it appears that the carbonaceous (coal-bearing) facies of the region reveal ichnofaunas with both relatively abundant saurischian (theropod) and ornithischian tracks. This is in contrast with many areas where the ichnofaunas are heavily or exclusively saurischian (theropod) dominated.