Saurischian (theropod–sauropod) track assemblages from the Jiaguan Formation in the Sichuan Basin,Southwest China: ichnology and indications to differential track preservation

Saurischian (theropod and sauropod) tracks and trackways from the Jiaguan Formation (Lower Cretaceous) of the Sichuan Basin are exposed as natural casts with associated undertrack or transmitted print casts. The theropod tracks (cf. Eubrontes and Grallator) were left by differently sized trackmakers. This is a further example for the occurrence of characteristic Lower Jurassic ichnotaxa in the Cretaceous that obviously had a more extended stratigraphic range in East Asia. The sauropod trackway is tentatively assigned to cf. Brontopodus based on imprint morphology and (nearly wide) gauge. The tracks, however, allow a detailed study of their formation and the taphonomic processes under different substrate conditions. Differential preservation and erosion of primary sedimentary structures, and post-burial deformation structures, give insight into a complex preservational history during a low energy phase interrupting the deposition of a sequence of thick high energy sandstones. This is the sixth report of dinosaur tracks from the Jiaguan Formation and the fifteenth report from the Lower Cretaceous of Sichuan Province. Thus, the tetrapod ichnological record in this region is rapidly becoming of major importance for our knowledge of dinosaur faunas in south-western China.     

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 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.     

Sauropod Trackway Reflecting an Unusual Walking Pattern from the Early Cretaceous of Shandong Province,China

The trackway of a quadrupedal dinosaur from the Early Cretaceous (Albian) Qingquan tracksite (Tancheng, Shandong Province) is redescribed, and the trackmaker is identified as a sauropod. The trackway makes a slight turn towards the northwest and is characterized by an extremely narrow gauge pattern and an unusual configuration, i.e., a conspicuous difference between the position of the left and right manus tracks with respect to the position of the preceding pes track. Left manus tracks are located on the inside of the trackway, very close (and sometimes even in connection) to the opposite right pes tracks. So far, the Qingquan trackway is possibly the only extremely narrow-gauge sauropod trackway known from China. However, it is not clear to what extent this extremely narrow gauge pattern is related to the turning or a special behavior, or even linked to an injury (“limping trackway”). We tentatively attribute the Qingquan trackway to cf. Parabrontopodus, even though it has a rather low heteropody that is significantly lower than in Parabrontopodus and not typical for narrow-gauge sauropod trackways, but occurs in the wide-gauge ichnotaxon Brontopodus. Because of this discrepancy, the Qingquan trackway cannot readily be attributed to a more basal sauropod, which is generally considered the producer of narrow-gauge trackways. Therefore, the identification of a distinct sauropod group is not possible presently. The only skeletal remains of sauropods from the Lower Cretaceous of Shandong Province belong to the large titanosauriform, Euhelopus zdanskyi.     

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.     

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.     

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.     

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.     

Hominid Ichnotaxonomy: An Exploration of a Neglected Discipline

Although more than 60 ancient hominid track sites ranging in age from 3.7 million to less than 500 B. P. are recorded from all continents except Antarctica, no ichnotaxonomic names have ever been formally proposed for hominid tracks. There is no prohibition to the naming of fossil footprints of species that created tracks and trackways similar to those of living species. On the contrary, there is precedent for the naming of ichnotaxa corresponding to the dominant extant vertebrates classes: mammals = Mammalipedia and birds = Avipeda. The hominid track site sample includes only about a dozen sites where footprint preservation is good enough to show details of diagnostic foot morphology and typical trackway morphology. We infer that the Acahualinca Footprint Museum site in Nicaragua represents the most important ancient hominid track site that combines accessibility, a large sample of well-preserved trackways and reliable dating. For this reason, we select the Nicaraguan tracks as the type sample for the new ichnotaxon Hominipes modernus ichnogen., and ichnsp. et ichnosp. nov., which we infer to represent fully modern Homo sapiens. Our preliminary investigations of other track sites suggest that the majority also yield H. modernus. However, at many sites preservation is insufficient to make an ichnotaxonomic designation at the species level or to infer that the trackmaker was H. sapiens. Thus, at many sites including the famous Laetoli site, we apply the more general label of Hominipes isp. indet.     

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.     

Trackway Ratio: A New Look at Trackway Gauge in the Analysis of Quadrupedal Dinosaur Trackways and its Implications for Ichnotaxonomy

A new parameter, the Trackway Ratio (TR), is proposed to supplement the previously used trackway gauge to describe and quantify the relative width of trackways in dinosaur quadrupedal gaits. It is expressed as the ratio of the width of the tracks relative to the total width of the trackway (both measured perpendicular to the long axis of the trackway). The ratio may be used with either pes (PTR) or manus (MTR) tracks. The PTR range of values for wide-, medium- and narrow-gauge trackways of previous authors are provisionally suggested to be ≤35%, 36–49% and ≥50%, respectively. The application of such a ratio would permit a more consistent ichnotaxonomy to be adopted where both track morphology and trackway parameters are used to define ichnotaxa.

Determination of the TR, as well as other parameters, will be affected by track preservation quality. Recent experiments on track simulation in the laboratory have shed further light on observations made in the field concerning the value of track measurements (in particular track length and width) recorded from below the surface on which the maker was moving. Experimental track simulations in the laboratory have shown that the dimensions of transmitted tracks preserved below the surface on which the foot was impressed may vary from 65% to 135% of the true dimensions of the indenter. Two case studies are presented that quantify the errors that may be made on calculating TR and the size, gait and speed of the maker, from trackways if the preservation of the tracks are not fully understood.

It is shown that in individual trackways the PTR may vary along the length of the trackway; so that part of the trackway may be classified as wide-gauge and other parts medium-gauge. There is a relationship between variation in PTR and that of pace angulation along the length of a single trackway. An analysis of 42 trackways, principally sauropod, shows a temporal distribution that does not agree closely with previous suggestions relating to narrow- and wide-gauge trackways.     

A New Pterosaur Tracksite from the Jurassic Summerville Formation,Near Ferron,Utah

Pterosaur tracks (cf. Pteraichnus) from the Summerville Formation of the Ferron area of central Utah add to the growing record of Pteraichnus tracksites in the Late Jurassic Summerville Formation and time-equivalent, or near time-equivalent, deposits. The site is typical in revealing high pterosaur track densities, but low ichnodiversity suggesting congregations or “flocks” of many individuals. Footprint length varies from 2.0 to 7.0 cms. The ratio of well-preserved pes:manus tracks is about 1:3.4. This reflects a bias in favor of preservation of manus tracks due to the greater weight-bearing role of the front limbs, as noted in other pterosaur track assemblages. The sample also reveals a number of well-preserved trackways including one suggestive of pes-only progression that might be associated with take off or landing, and another that shows pronounced lengthening of stride indicating acceleration.

One well-preserved medium-sized theropod trackway (Therangospodus) and other larger theropod track casts (cf. Megalosauripus) are associated with what otherwise appears to be a nearly monospecific pterosaur track assemblage. However, traces of a fifth pes digit suggest some tracks are of rhamphorynchoid rather than pterodactyloid origin, as usually inferred for Pteraichnus. The tracks occur at several horizons in a thin stratigraphic interval of ripple marked sandstones and siltstones. Overall the assemblage is similar to others found in the same time interval in the Western Interior from central and eastern Utah through central and southern Wyoming, Colorado, northeastern Arizona, and western Oklahoma. This vast “Pteraichnus ichnofacies,” with associated saurischian tracks, remains the only ichnological evidence of pre-Cretaceous pterosaurs in North America and sheds important light on the vertebrate ecology of the Summerville Formation and contiguous deposits.     

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.     

Intra-Trackway Morphological Variations Due to Substrate Consistency: The El Frontal Dinosaur Tracksite (Lower Cretaceous,Spain)

An ichnological and sedimentological study of the El Frontal dinosaur tracksite (Early Cretaceous, Cameros basin, Soria, Spain) highlights the pronounced intra-trackway variation found in track morphologies of four theropod trackways. Photogrammetric 3D digital models revealed various and distinct intra-trackway morphotypes, which reflect changes in footprint parameters such as the pace length, the track length, depth, and height of displacement rims. Sedimentological analyses suggest that the original substrate was non-homogenous due to lateral changes in adjoining microfacies. Multidata analyses indicate that morphological differences in these deep and shallow tracks represent a part of a continuum of track morphologies and geometries produced by a gradient of substrate consistencies across the site. This implies that the large range of track morphologies at this site resulted from similar trackmakers crossing variable facies. The trackways at the El Frontal site present an exemplary case of how track morphology, and consequently potential ichnotaxa, can vary, even when produced by a single trackmaker.     

Middle Triassic horseshoe crab reproduction areas on intertidal flats of Europe with evidence of predation by archosaurs

A systematically excavated track site in a 243.5 Myr old Middle Triassic (Karlstadt Formation, Pelsonian, middle Anisian) intertidal carbonate mud‐flat palaeoenvironment at Bernburg (Saxony‐Anhalt, central Germany) has revealed extensive horseshoe crab trackways attributable to the Kouphichnium Nopsca, 1923 ichnogenus. The exposed track bed of a Germanic Basin‐wide spanned intertidal megatrack site is a mud‐cracked biolaminate surface on which detailed tracks have been preserved because of rapid drying and cementation as a result of high temperatures, followed by rapid covering with a protective layer of arenitic storm or tsunami sediments. The different trackway types and their orientations have allowed a tidal sequence to be reconstructed, with the initial appearance of swimming horseshoe crabs followed by half‐swimming/half‐hopping limulids under the shallowest water conditions. The Bernburg trackways, which have mapped lengths of up to 40 m, were all produced by adult animals and exhibit a variety of shapes and patterns that reflect a range of subaquatic locomotion behaviour more typical of mating than of feeding activities. The closest match to the proportions and dimensions of the horseshoe crab tracks at Bernburg is provided by the largest known Middle Triassic limulid Tachypleus gadeai, which is known from the north‐western Tethys in Spain. The horseshoe crab body fossils recognized in the German Mesozoic intertidal zones, instead, are from juveniles. The uniformly adult size indicated by the trackways therefore suggests that they may record the oldest intertidal reproductive zones of horseshoe crabs known from anywhere in the world, with the track‐makers having possibly migrated thousands of kilometres from shallow marine areas of the north‐western Tethys to reproduce in the intertidal palaeoenvironments of the Germanic Basin. Chirotherium trackways of large thecodont archosaurs also appeared on these flats where they appear to have fed on the limulids. With the tidal ebb, smaller reptiles such as Macrocnemus (Rhynchosauroides trackways) appeared on the dry intertidal flats, probably feeding on marine organisms and possibly also on horseshoe crab eggs. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 76–105.     

A Diverse Dinosaur-Bird Footprint Assemblage from the Lance Formation,Upper Cretaceous,Eastern Wyoming: Implications for Ichnotaxonomy

A diverse assemblage of dinosaur and bird tracks from Niobrara County, Wyoming, represents the first vertebrate ichnofauna reported from the bone-rich Lance Formation (Maastrichtian, Upper Cretaceous). The ichnofauna includes a hadrosaur track with skin impressions; three theropod track types, including the tetradactyl track Saurexallopus zerbsti (ichnosp. nov.); a tridactyl dinosaur footprint with a fusiform digit III; possible Tyrannosaurus tracks; four distinctive avian ichnites; and invertebrate traces. The footprints are generally well-preserved and so offer a unique insight into the ecology of a small river valley during the Maastrichtian.

Saurexallopus zerbsti ichnosp. nov. from the Lance is similar to Saurexallopus lovei recently reported from the Maastrichtian, Harebell Formation, of northwestern Wyoming, but is represented by much better material, facilitating amendment of the ichnogenus. Skeletal equivalents for Saurexallopus are not currently known. Similarly, the tridactyl track with fusiform digit III is similar to footprints reported from the coeval Laramie Formation of Colorado and may also be similar to ichnogenus Ornithomimipus from the Edmonton Group of Alberta (though not necessarily of ornithomimid affinity). The hadrosaurian track with the skin impression is reminiscent of a similar ichnite reported from the Maastrichtian, St. Mary River Formation in Alberta, which is herein named Hadrosauropodus langstoni as part of a reassessment of Cretaceous ornithopod track ichnotaxonomy. Such correlations demonstrate the utility of tracks for local or regional biostratigraphy (palichnostratigraphy) in western North America. It is also clear that tracks add to our knowledge of the composition and distribution of dinosaurian and avian components of Maastrichtian faunas. In particular the bird tracks indicate a diversity of at least four species, one of which was a semi-palmate form, hitherto unknown in the ichnological record and named Sarjeantichnus semipalmatus.     

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).     

Complex In-Substrate Dinosaur (Sauropoda,Ornithopoda) Foot Pathways Revealed by Deep Natural Track Casts from the Lower Cretaceous Xiagou and Zhonggou Formations,Gansu Province,China

Several new Early Cretaceous tracksites from the Lower Cretaceous Xiagou Formation of Gansu Province (China) with tracks of large sauropods and ornithopods are described. Previously reported bird tracks were missing due to human negligence. The studied specimens are preserved as impressions and shallow and deep natural track casts. These dinosaur tracks are first reported from the Jiuquan area in the Changma Basin, matching well with the skeletal record of diverse non-avian dinosaur-bird faunas of this region. Moreover, they add new data to the dinosaur ichnofaunas of the Lanzhou-Minhe Basin (Gansu Province) and indicate a wide distribution of dinosaur-bird assemblages in the Early Cretaceous. Regarding morphology, sauropod, and ornithopod tracks from the Lanzhou-Minhe Basin and the Jiuquan area are very similar to each other. Titanosauriform trackmakers are assumed for the sauropod tracks and possibly iguanodontids have left the large, tridactyl ornithopod tracks. Of particular interest are well-preserved, deep natural track casts of large ornithopods and sauropods preserving ridges and grooves as well as striation marks on the lateral sides of the casts that allow the reconstruction of complex pathways of the foot within the substrate. One particular sauropod pes–manus track cast even indicates lateral and vertical sliding within the sediment because of the presence of “double impressions of digits” on the bottom.