Early cretaceous bird-dominated and dinosaur footprint assemblages from the northwestern margin of the Junggar Basin, Xinjiang, China

Here we describe a number of mostly isolated bird and non-avian dinosaur footprints from the Huangyangquan track site in the Lower Cretaceous Lower Layer of the Tugulu Group near Huangyangquan Reservoir in the Wuerhe District, Karamay, Xinjiang Uyghur Autonomous Region, China. Bird footprints at the site pertain to Koreanaornis dodsoni n. ichnosp., Goseongornipes isp., Aquatilavipes isp., and Moguiornipes robusta n. ichnogen. n. ichnosp. Moguiornipes tracks may be the products of a bird with lobed feet. These shorebird-morph tracks are the first solid evidence of birds in the Tugulu Group, and greatly enrich the known faunal diversity of the Lower Layer of the Tugulu Group. The Huangyangquan avian ichnofauna is comparable to those of the Jindong Formation, Korea and the Dakota Formation, USA. Non-avian dinosaur footprints at the site pertain to cf. Jialingpus isp., Asianopodus isp., and Kayentapus isp. This is the first report of Jialingpus from Lower Cretaceous strata. The discoveries of Asianopodus isp. and Kayentapus isp. increase the known ranges of these two ichnogenera.     

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.     

First Thyreophoran Type Tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province,China

Two tridactyl footprints from the Chuanjie Formation (Middle Jurassic) of Yunnan Province, China are morphological characteristics of thyreophoran tracks. They show some similarities to Shenmuichnus, known from the Early Jurassic strata of both Shaanxi and Yunnan provinces, but are somewhat larger, thereby resembling the ichnogenus Stegopodus. Based on their general morphology and size being congruent with this ichnogenus, they are tentatively assigned here to cf. Stegopodus. This is the fourth report of large ornithischian (probably thyreophoran) tracks from the Lower-Middle Jurassic of China that indicates relatively large trackmakers that were likely to be taxonomically distinct from much smaller and gracile Anomoepus trackmakers, also of ornithischian affinity. The larger tracks indicate a hitherto unreported abundance, size range and diversity of track types attributed to this group. The parallel orientation of the two best preserved trackways may indicate gregariousness.     

Unusual deinonychosaurian track morphology (Velociraptorichnus zhangi n. ichnosp.) from the Lower Cretaceous Xiaoba Formation,Sichuan Province,China

A new Lower Cretaceous dinosaur footprint locality named the Mujiaowu tracksite in the Xiaoba Formation, Sichuan Province, has yielded a new assemblage containing the didactyl deinonychosaurian ichnogenus Velociraptorichnus. This is the eleventh report in the global record, the seventh from Asia and the fifth from China. All, except for an isolated report from Europe, occur in Lower Cretaceous deposits. Unlike previous reports of the ichnogenus, the Mujiaowu site has yielded both didactyl Velociraptorichnus tracks and tridactyl tracks which we interpret as a different expression of this same ichnogenus, caused by registration of digit II, either due to special substrate conditions or to less claw retraction. These tridactyl Velociraptorichnus footprints are assigned to the new ichnospecies Velociraptorichnus zhangi. Such tridactyl deinonychosaurid footprint morphology is predictable, but based on current ichnological evidence appears to be the exception rather than the rule.     

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.     

Diverse Dinosaur-Dominated Ichnofaunas from the Potomac Group (Lower Cretaceous) Maryland

Until recently fossil footprints were virtually unknown from the Cretaceous of the eastern United States. The discovery of about 300 footprints in iron-rich siliciclastic facies of the Patuxent Formation (Potomac Group) of Aptian age is undoubtedly one of the most significant Early Cretaceous track discoveries since the Paluxy track discoveries in Texas in the 1930s. The Patuxent tracks include theropod, sauropod, ankylosaur and ornithopod dinosaur footprints, pterosaur tracks, and miscellaneous mammal and other vertebrate ichnites that collectively suggest a diversity of about 14 morphotypes. This is about twice the previous maximum estimate for any known Early Cretaceous vertebrate ichnofauna. Among the more distinctive forms are excellent examples of hypsilophodontid tracks and a surprisingly large mammal footprint. A remarkable feature of the Patuxent track assemblage is the high proportion of small tracks indicative of hatchlings, independently verified by the discovery of a hatchling-sized dinosaur. Such evidence suggests the proximity of nest sites. The preservation of such small tracks is very rare in the Cretaceous track record, and indeed throughout most of the Mesozoic.

This unusual preservation not only provides us with a window into a diverse Early Cretaceous ecosystem, but it also suggests the potential of such facies to provide ichnological bonanzas. A remarkable feature of the assemblage is that it consists largely of reworked nodules and clasts that may have previously been reworked within the Patuxent Formation. Such unusual contexts of preservation should provide intriguing research opportunities for sedimentologists interested in the diagenesis and taphonomy of a unique track-bearing facies.     

Vertebrate Tracksites in the Middle Jurassic-Upper Cretaceous of South Tunisia

Four vertebrate tracksites from the Middle Jurassic and Upper Cretaceous in the Tataouine basin of southern Tunisia are described. Approximately 130 tridactyl footprints distributed over an area of 200 square meters, preserved on Callovian beds exposed at the Beni Ghedir site, represent the oldest evidence of a dinosaur fauna in Tunisia. In addition, three tracksites—Chenini, Ksar Ayaat, and Jebel Boulouha—have been discovered in the Cretaceous beds of the upper Continental Intercalaire, previously considered as a strictly marine depositional sequence. In addition to dinosaur tracks, the Chenini tracksite (late Albian) includes poorly preserved crocodilian tracks, and footprints assigned to a pleurodiran turtle have been recovered at the Ksar Ayaat locality (early Cenomanian). The Jebel Boulouha tracksite is dominated by well-preserved tridactyl tracks referred to small-sized theropods. Depositional settings of each tracksite have been defined on stratigraphic and sedimentologic data, and tracks were ascribed to different ichnocoenoses in relation to their paleoenvironments. This new and differentiated track record gives important information on how the fossil vertebrate fauna changed in southern Tunisia during mid-Jurassic to mid-Cretaceous times. These data provide a unique and useful census of tetrapod associations along the southern margin of the peri-Mediterranean area.     

Theropod and possible ornithopod track assemblages from the Jurassic–Cretaceous boundary Houcheng Formation,Shangyi, northern Hebei,China

Dinosaur track assemblages from the Houcheng Formation in the small continental Shangyi Basin of northern Hebei Province, China bridge a gap in the record of vertebrates from this unit and enrich our knowledge of ichnofaunas from the Jurassic–Cretaceous boundary. Their stratigraphic position between the Middle Jurassic Yan-Liao Biota and the Lower Cretaceous Jehol Biota gives them a special importance. New discoveries allow a re-assessment of theropod and possible ornithopod tracks that are present with several trackways. Seventy-three footprints were examined and documented. Despite their smaller size, the tridactyl mesaxonic theropod tracks show morphological similarities with the ichnogenus Therangospodus known from the Upper Jurassic deposits of North America, Europe, and Central Asia. The possible ornithopod tracks lack an associated manus imprint, suggesting a bipedal trackmaker. These possible ornithopod tracks from the Houcheng Formation provide evidence for the presence of small basal ornithopods or basal Cerapoda in the Upper Jurassic–Lower Cretaceous in this region. The depositional environment was the margin of an extensive shallow lake with fluctuating water levels under seasonally dry climate.     

Probable crocodilian tracks and traces from the Morrison Formation (Upper Jurassic) of eastern Utah

A tetradactyl pes impression and tridactyl manus impression are described as the type specimen of Hatcherichnus sanjuanensis ichnogen. et ichnosp. nov., a probable large crocodilian ichnite from the Salt Wash Member of the Upper Jurassic Morrison Formation in eastern Utah. A similar pes track from the Morrison Formation at Garden Park, Colorado, may also belong to this ichnogenus. The type specimen from Utah consists of plaster replicas of natural casts of a left pes impression and a left manus impression. Associated with the type specimen were possible tail and body drag impressions. The tracks do not appear to be part of a walking trackway and may be swim tracks associated with an animal in shallow water. The tracks occur at a visible contact between slightly fining‐upward channel sandstone units.     

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.     

Behaviour of Dinosaurian Track-Makers in the Winton Formation (Cretaceous,Albian–Cenomanian) at Lark Quarry,Western Queensland,Australia: Running or Swimming?

A remarkable assemblage of dinosaur trackways in the Winton Formation (Albian–Cenomanian) at Lark Quarry, in western Queensland, Australia, has long been regarded as evidence of a stampede involving small theropods, whose tracks were classified in the ichnogenus Skartopus, and small ornithopods, whose tracks represented a second ichnogenus, Wintonopus. However, one recently-published study has claimed that existing interpretation of Lark Quarry is incorrect: it maintains that all the track-makers were ornithopods, that the ichnogenus Skartopus is a variant form of Wintonopus, and that most of the Lark Quarry track-makers were not running but more probably swimming downstream in a current of water. Those iconoclastic claims are investigated here and shown to be untenable. They derive from overgeneralized interpretation of the Lark Quarry track assemblage and invoke ad hoc auxiliary hypotheses which are either untestable or demonstrably incorrect. Closer inspection of the evidence underpinning those claims corroborates the existing interpretation of Lark Quarry as the site of a dinosaurian stampede and confirms the validity of the original distinction between theropod tracks (Skartopus) and ornithopod tracks (Wintonopus).     

New Hettangian tracks from the Causses Basin (Lozère,southern France) complement the poor fossil record of earliest Jurassic crocodylomorphs in Europe

A new vertebrate ichnological assemblage is described from the Hettangian Dolomitic Formation of the Causses Basin, at Le Serre (Lozère, southern France). We report tracks that complement the poor fossil record of lowermost Jurassic crocodylomorphs in Europe. Tetradactyl pes and pentadactyl manus imprints form a unique trackway. Traces are ascribed to Batrachopus isp. and they represent the third occurrence of this ichnogenus in European Hettangian geological formations. This is the first report of Batrachopus in Hettangian deposits of the Causses Basin. The tracks share some similarities with Batrachopus deweyi previously described from the Sinemurian of the Causses Basin. Crocodylomorph tracks co-occur with tridactyl dinosaur traces assigned to Dilophosauripus williamsi and Grallator isp. The sedimentology and palaeoichnology of the tracksite indicate that the depositional environment was a tidal to supratidal flat marsh that was emerged periodically. The ichnological assemblage from Le Serre and a synthesis of ichnotaxa co-occurring with Batrachopus in the European lowermost Jurassic tracksites confirm that crocodylomorphs living in marginal-littoral palaeoenvironments were part of theropod-dominated faunas, together with ornithopods but apparently without sauropods.     

Crocodylian Tracks from Lower Oligocene Flysch deposits of the Barail Group,Manipur, India

A marine tidal delta siltstone from Gelmon locality in Northeast India preserved three crocodylian footprints and an elongate depression that appears to be a tail drag mark. Similar drag marks occur in nearby bedding surfaces. The discovery of crocodylian tracks from the basal part of Laisong Formation, Barail Group (Late Eocene–Early Oligocene age) of Manipur, India is noteworthy because of the age and the geographic location. Crocodylian tracks are rare in Cenozoic formations and they have not previously been reported from Asia. The footprints are herein named as a new ichnogenus and ichnospecies, Indosuchipes manipurensis.     

Crocodile swim tracks from the latest Cretaceous of Europe

Recently discovered evidence of tracks in the continental beds of the Late Cretaceous Tremp Formation in the southern Pyrenees (NE Iberian Peninsula) has been identified as scratch marks made by buoyant crocodiles. The tracks are preserved in two distinct environments and substrates (marly limestones originating in a littoral mud flat and fine‐grained sandstones deposited in fluvial settings). Most of the crocodylian traces are ascribed to ichnogenus Characichnos, whereas a single plantigrade pes track is assigned to ichnogenus cf. Crocodylopodus. The crocodylian swim traces (Characichnos ichnofacies) found in the early and late Maastrichtian co‐occur with Brontopodus ichnofacies attributable to terrestrial tetrapods (titanosaur sauropods, cf. Brontopodus ichnogenus; and hadrosaurid ornithopods, Hadrosauropodus ichnogenus). Analysis of the tracks allows the interpretation of palaeoenvironmental settings and track production. Thus, in lagoonal environments, swim tracks of crocodylians were produced during the rise of the water level in successive tide cycles; in fluvial settings, the swim traces of crocodylians were produced within the channel at the low‐water stage. To date, there are no reports of Late Cretaceous crocodylian tracks in Europe, and the studied evidence represents the first and youngest track record of the group in the latest part of the Cretaceous (C29r) in this continent and probably in the world.     

Scampering,trotting, walking tridactyl bipedal dinosaurs in southern Africa: ichnological account of a Lower Jurassic palaeosurface (upper Elliot Formation,Roma Valley) in Lesotho

In Gondwana, Early Jurassic dinosaur track sites are especially concentrated in Lesotho. Despite intensive investigations during the third quarter of the twentieth century, a limited number of vertebrate track sites of this country have been studied with rigorous ichnological and sedimentological methodology. Here, we present a previously mentioned, but undescribed track site in the upper Elliot Formation (Hettangian?) of Lesotho, located near Roma (at Lephoto dam). Two tridactyl ichnite morphologies, made by bipedal vertebrate trackmakers are recognised. The first can be identified as Grallator-like, an ichnotaxon common in the Lower Jurassic of both Laurasia and Gondwana that can be attributed to small and medium-size theropod dinosaurs. In contrast, the second ichnite type is reminiscent of Trisauropodiscus, which is a rare ichnotaxon that resembles tracks of small birds and is known with certainty in Lesotho from only a few places. We suggest that at our upper Elliot Formation study site, Trisauropodiscus was potentially made by a heterodontosaurid ornithischian dinosaur. Our work provides further evidence that the ichnological record of the Stormberg Group of southern Africa is in a unique position to shed light not only on Early Jurassic biostratigraphy and palaeoenvironments but also on the biodiversity and palaeobiology of early dinosaurs.     

Sauropod and Small Theropod Tracks from the Lower Jurassic Ziliujing Formation of Zigong City,Sichuan, China,with an Overview of Triassic–Jurassic Dinosaur Fossils and Footprints of the Sichuan Basin

A dinosaur footprint assemblage from the Lower Jurassic Ziliujing Formation of Zigong City, Sichuan, China, comprises about 300 tracks of small tridactyl theropods and large sauropods preserved as concave epireliefs (natural molds). The theropod footprints show similarities with both the ichnogenera Grallator and Jialingpus. Three different morphotypes are present, probably related to different substrate conditions and extramorphological variation. A peculiar preservational feature in a morphotype that reflects a gracile trackmaker with extremely slender digits, is the presence of a convex epirelief that occurs at the bottom of the concave digit impressions. It is possibly the result of sediment compaction underweight load when the pes penetrated the substrate, being a resistant residue during exhumation and weathering. The sauropod tracks belong to a trackway with eight imprints consisting of poorly preserved pes and manus tracks and a better preserved set, probably all undertracks. The narrow-gauge trackway pattern resembles the ichnogenus Parabrontopodus well known from the Jurassic but other features such as the minor heteropody are different. The assemblage enriches the dinosaur record from the famous Zigong locality and the evidence from the Lower Jurassic in this area that was restricted to a few skeletal remains and footprints. Furthermore it proves the presence of small theropods, whereas skeletons of the group, well- known from the Middle-Upper Jurassic of Zigong, are of medium to large size only. Remarkable is the dominance of saurischians in these assemblages, which is characteristic of Jurassic dinosaur communities whereas the Cretaceous record shows an increase of ornithopod groups. An overview of the dinosaur trace and body fossil record of the Sichuan Basin supports this view. The paleoenvironment can be designated as a low-latitude tropical freshwater lake as it is indicated by bivalve shells.     

陕西子洲中侏罗世早期恐龙足迹初步研究

陕西子洲县中侏罗统延安组砂岩中,同一地点共厚约1.7 m的5个层面发现4种食肉类恐龙足迹,自上而下分别为:1)第5层面大型三趾型足迹,实雷龙足迹科王氏子洲足迹(新遗迹属、新遗迹种)Zizhoupus wangi ichnogen.et ichnosp,nov.;2)第4和第3层面中型三趾型足迹,实雷龙足迹科龙尾峁张北足迹(新遗迹种)Changpeipus longweimaoensis inchnosp.nov.;3)第2层面小型三趾或四趾型足迹,虚骨龙类小理河陕西足迹(新遗迹种)Shensipus xiaoliheensis inchnosp.nov.;4)第1层面小型三趾型足迹,虚骨龙类的铜川陕西足迹Shensipus tungchuanensis.     

Review of Japanese Dinosaur Track Localities: Implications for Ichnotaxonomy,Paleogeography and Stratigraphic Correlation

Nine dinosaur ichnospecies from the Lower Jurassic to Upper Cretaceous of Japan, including two that are new, are described herein. The new ichnotaxa are Asianopodus pulvinicalx ichnogen. et ichnosp. nov. and Schizograllator otariensis ichnosp. nov. The Japanese ichnotaxa are allied to Lower Jurassic ichnospecies in South China, North America, Western Europe and South Africa, and Upper Jurassic to Lower Cretaceous ichnospecies from Southeast and East Asia. This suggests they were part of a global ichnofauna before continental drift began in the Middle Jurassic, leading to the development of a more endemic dinosaur fauna in the Cretaceous. At least two assemblages, an ornithopod-gracile-toed theropod-dominated community, in northeastern Asia, and a robust theropod- and sauropod-dominated community in the southern part of the continent, existed in the Cretaceous. This parallels North American dinosaur distribution patterns in the Cretaceous and seems to be a reflection of paleolatitudinal controls.     

Dinosaur Tracks from the Cedar Mountain Formation (Lower Cretaceous), Arches National Park,Utah

The seventh and largest known dinosaur tracksite from the Cedar Mountain Formation is reported from two important stratigraphic levels in the Ruby Ranch Member within the boundaries of Arches National Park. Previous reports of sites with a few isolated tracks are of limited utility in indicating the fauna represented by track makers. The Arches site reveals evidence of several theropod morphotypes, including a possible match for the coelurosaur Nedcolbertia and an apparently didactyl Utahraptor-like dromeosaurid. Sauropod tracks indicate a wide-gauge morphotype (cf. Brontopodus). Ornithischian tracks suggest the presence of an iguandontid-like ornithopod and a large ankylosaur. Dinosaur track diversity is high in comparison with other early Cretaceous vertebrate ichnofaunas, and it correlates well with faunal lists derived from skeletal remains, thus providing a convincing census of the known fauna.     

A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal

Mateus, O. & Milàn, J. 2009: A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal. Lethaia, Vol. 43, pp. 245–257. A newly discovered dinosaur track‐assemblage from the Upper Jurassic Lourinhã Formation (Lusitanian Basin, central‐west Portugal), comprises medium‐ to large‐sized sauropod tracks with well‐preserved impressions of soft tissue anatomy, stegosaur tracks and tracks from medium‐ to large‐sized theropods. The 400‐m‐thick Lourinhã Formation consists of mostly aluvial sediments, deposited during the early rifting of the Atlantic Ocean in the Kimmeridgian and Tithonian. The stratigraphic succession shows several shifts between flood‐plain mud and fluvial sands that favour preservation and fossilization of tracks. The studied track‐assemblage is found preserved as natural casts on the underside of a thin bivalve‐rich carbonate bed near the Tithonian–Kimmeridgian boundary. The diversity of the tracks from the new track assemblage is compared with similar faunas from the Upper Jurassic of Asturias, Spain and the Middle Jurassic Yorkshire Coast of England. The Portuguese record of Upper Jurassic dinosaur body fossils show close similarity to the track fauna from the Lourinhã Formation. □Dinosaur tracks, Lusitanian Basin, Portugal, skin impressions, Upper Jurassic.