Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove,Victoria

Two thin‐toed tridactyl tracks in a fluvial sandstone bed of the Eumeralla Formation (Albian) at Dinosaur Cove (Victoria, Australia) were likely made by avian trackmakers, making these the oldest known fossil bird tracks in Australia and the only Early Cretaceous ones from Gondwana. These tracks, which co‐occur on the same surface with a slightly larger nonavian theropod track, are distinguishable by their anisodactyl form, hallux impressions and wide digit II–IV divarication angles. A lengthy hallux impression and other deformational structures associated with one track indicate foot movement consistent with an abrupt stop, suggesting its tracemaker landed after either flight or a hop. The single nonavian theropod track is similar to other tracks described from the Eumeralla Formation at another locality. The avian footprints are larger than most Early Cretaceous avian tracks recorded worldwide, indicating sizeable enantiornithine or ornithurine species in formerly polar environments of Australia. The avian tracks further supplement scant body fossil evidence of Early Cretaceous birds in southern Australia, which includes a furcula from the Wonthaggi Formation. Because of this discovery, Dinosaur Cove, previously known for its vertebrate body fossils, is added to a growing list of Early Cretaceous vertebrate tracksites in southern Australia.     

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.     

The Fossil Record of Vertebrate Tracks in Mexico

The fossil record of vertebrate tracks in Mexico is taxonomically rich and geographically diverse, and ranges in age from Middle Jurassic to Pleistocene and possibly Holocene. Middle Jurassic saurischian dinosaur tracks from Oaxaca represent the oldest record of vertebrate tracks in Mexico. Tracks attributed to Late Jurassic theropods and ornithopods are known from Michoacán. Theropod tracks of supposed Jurassic age are known from Durango. Lower Cretaceous Saurischian and ornithopod tracks are known from southern Puebla. Central Mexico has yielded Late Cretaceous hadrosaurid and sauropod tracks (Puebla), and tracks belonging to Theropoda and Ornithopoda (El Aguaje, Michoacan). In Coahuila several tracksites occur within the Cerro del Pueblo Formation and include tracks of mesoreptiles, pterosaurs, small to large theropods, bipedal and quadrupedal ornithopods, birds and possibly a mammal. A tracksite in the Olmos Formation includes tracks of turtles, crocodilians, medium-sized theropods, small ornithopods, and birds. Bird tracks of supposed Eocene age are known from Sinaloa. Two important Pleistocene ichnofaunas occur in the states of Puebla (Tepexi de Rodríguez) and Jalisco (San Juan de los Lagos), and include tracks of camelids, artiodactyls, small ungulates, elephants, large felids and birds. Pleistocene human tracks occur in Coahuila and Jalisco. Mexico's track record provides important insights into vertebrate diversity, paleoecology, and paleoenvironments. Given the rate of new discoveries since 1998, there is no doubt that new sites await to be found.     

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.     

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.     

Small Theropod Track Assemblages from Middle Jurassic Eolianites of Eastern Utah: Paleoecological Insights from Dune Ichnofacies in a Transgressive Sequence

New discoveries show that very small theropod tracks (cf. Wildeichnus) are abundant in the upper part of the Moab Member, recently assigned to the Curtis Formation (formerly considered part of the Entrada Formation) in the Mid-?Late Jurassic of eastern Utah. The tracks represent a distinct small-theropod ichnofacies associated with eolian dune deposits that is easily differentiated from the water-lain beds of the overlying Megalosauripus-Therangospodus ichnofacies, which comprises the single-surface Moab megatracksite. Pterosaur track assemblages, representing the Pteriachnus ichnofacies, are found a few meters above the megatracksite surface in the upper tongue of the Summerville Formation.

The small theropod ichnofacies is reminiscent of other early Mesozoic dune facies ichnofaunas from the Wingate and Navajo formations (Late Triassic and Early Jurassic) where small theropod tracks occur in association with other small tetrapod footprints. All such examples evidently represent a recurrent dune facies ecosystem dominated by diminutive vertebrates. Because the small theropod ichnofacies is one of three ichnofacies found in a thin stratigraphic sequence (<20 m) that contains no body fossils, it is clear that vertebrate tracks play an important role in providing insight into the paleoecology of units previously considered devoid of any useful fossil evidence. The three successive ichnofacies represent a transgressive transition from sand dunes, through sandy shoreline to shallow marine environments, each with its quite distinct vertebrate fauna.     

An enigmatic early mesozoic dinosaur trackway from Zimbabwe

A trackway from Zimbabwe of probably the smallest dinosaur footprints recorded in Africa, is described and tentatively assigned to the Early Jurassic. The footprints are possibly those of a theropod and show strong negative (outward) rotation of the pes and are associated with manus prints. The shape of the footprints, unusual negative rotation, posterior curvature of digit IV and curious positioning of the manus prints in relation to the pes are enigmatic but somewhat reminiscent of Atreipus. Although a number of propositions are considered the most likely is that the animal was an immature dinosaur using a quadrupedal gait. A second trackway of slightly larger footprints of a bipedal theropod dinosaur is also recorded along with other diminutive tracks that suggest an early dinosaur assemblage, possibly dating from near the Trias‐sic‐Jurassic boundary.     

Los Cayos Dinosaur Tracksite: An Overview on the Lower Cretaceous Ichno-Diversity of the Cameros Basin (Cornago,La Rioja Province,Spain)

The Los Cayos dinosaur tracksite is located at the eastern sector of the Cameros Basin (Cornago Township, La Rioja province), NE Spain. The sediments consist of interbedded terrigenous siliciclastics and carbonates belonging to the Enciso Group (Early Cretaceous, Aptian in age). The sedimentological and faunal evidence suggests that these sediments were deposited in a low gradient lacustrine environment. Los Cayos constitutes a relatively wide area with at least 6 localities that have yielded dinosaur tracks. More than 2,000 dinosaur tracks have been discovered to date. Medium to large theropod dinosaurs constitute about 95% of the ichnofauna. Ornithopod dinosaur tracks have been reported from only one of the outcrops (Los Cayos D). Los Cayos S has yielded theropod and sauropod tracks. One sauropod trackway shows a narrow-gauge locomotion pattern and a manus print morphology suggesting that the trackmaker was a titanosaurid, or at least a titanosauriform sauropod. Some pterosaur manus impressions, avian-like footprints of small size and possible turtle tracks complete the assemblage of one of the most impressive and best-preserved dinosaur tracksites of the European Lower Cretaceous.     

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.     

Dinosaur footprints from the Sao Khua Formation (Early Cretaceous) of Phu Kao National Park,Northeast Thailand

The first documented dinosaur footprints from Southeast Asia were theropod footprints found in Thailand at Phu Luang Wildlife Sanctuary. Since then, Mesozoic vertebrate footprints have been discovered in twelve different localities in Thailand. In 2000, dinosaur footprints were found in Phu Kao National Park in Northeastern Thailand, however, they have never been studied in detail. The following study presents the first map, the complete measurements, the geology, and stratigraphy of the site and surrounding area. The study discusses the nature of the trackmaker and the digitigrade character of the footprints. This discovery represents the first documented occurrence of dinosaur footprints inside the Early Cretaceous Sao Khua Formation from Thailand.     

Early Cretaceous pterosaur tracks from a “buried” dinosaur tracksite in Shandong Province,China

Here we describe the pterosaur and tridactyl dinosaur footprint assemblage from a new tracksite in the Early Cretaceous Qugezhuang Formation near the Wenxiyuan Community in Jimo County-level City, Qingdao City, Shandong Province, East China. The construction of a community building has, since the discovery, covered the majority of tracks, but a few specimens on abandoned building stones constitute the first pterosaur track record in eastern China. The pterosaur tracks are assigned to Pteraichnus isp. and were probably made by a small to medium-sized pterodactyloid. The new pterosaur trackway contributes to the growing database of pterosaurian ichnites in Asia. The Wenxiyuan tridactyl dinosaur tracks are morphologically similar to those of theropods, but they are too poorly preserved to be referred to any particular ichnotaxon.     

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.     

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.     

The Lower Cretaceous Patuxent Formation Ichnofauna of Virginia

The vertebrate fauna from the Lower Cretaceous Patuxent Formation of Virginia is composed of a single partial fish impression from the James River at Dutch Gap and a diverse tetrapod ichnofauna from near Fredericksburg that includes trace fossils made by frogs, turtles, theropods, sauropods, ankylosaurs, and ornithopods. The footprints occur on overbank deposits preserved locally within a fluvial braided-stream sequence that formed near the western border of the Early Cretaceous Atlantic Coastal Plain.     

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.     

Tetrapod Footprint Biostratigraphy and Biochronology

Tetrapod footprints have a fossil record in rocks of Devonian-Neogene age. Three principal factors limit their use in biostratigraphy and biochronology (palichnostratigraphy): invalid ichnotaxa based on extramorphological variants, slow apparent evolutionary turnover rates and facies restrictions. The ichnotaxonomy of tetrapod footprints has generally been oversplit, largely due to a failure to appreciate extramorphological variation. Thus, many tetrapod footprint ichnogenera and most ichnospecies are useless phantom taxa that confound biostratigraphic correlation and biochronological subdivision. Tracks rarely allow identification of a genus or species known from the body fossil record. Indeed, almost all tetrapod footprint ichnogenera are equivalent to a family or a higher taxon (order, superorder, etc.) based on body fossils. This means that ichnogenera necessarily have much longer temporal ranges and therefore slower apparent evolutionary turnover rates than do body fossil genera. Because of this, footprints cannot provide as refined a subdivision of geological time as do body fossils. The tetrapod footprint record is much more facies controlled than the tetrapod body fossil record. The relatively narrow facies window for track preservation, and the fact that tracks are almost never transported, redeposited or reworked, limits the facies that can be correlated with any track-based biostratigraphy.

A Devonian-Neogene global biochronology based on tetrapod footprints generally resolves geologic time about 20 to 50 percent as well as does the tetrapod body fossil record. The following globally recognizable time intervals can be based on the track record: (1) Late Devonian; (2) Mississippian; (3) Early-Middle Pennsylvanian; (4) Late Pennsylvanian; (5) Early Permian; (6) Late Permian; (7) Early-Middle Triassic; (8) late Middle Triassic; (9) Late Triassic; (10) Early Jurassic; (11) Middle-Late Jurassic; (12) Early Cretaceous; (13) Late Cretaceous; (14) Paleogene; (15) Neogene. Tetrapod footprints are most valuable in establishing biostratigraphic datum points, and this is their primary value to understanding the stratigraphic (temporal) dimension of tetrapod evolution.     

Late Cretaceous Mammal Tracks from North America

Mammal tracks from the latest Cretaceous (Campanian-Maastrichtian) of Colorado are described as Schadipes crypticus ichnogen. et ichnosp. nov., on the basis of material from the Laramie Formation in Golden, eastern Colorado. This ichnospecies, and a closely related form (Schadipes sp.) from the Mesaverde Group of western Colorado, represent the only mammal tracks so far identified from the Upper Cretaceous. A possible mammal track from North Africa (Agadirichnus elegans) was originally attributed to a lizard/lacertilian. Other purported Lower Cretaceous mammal tracks are based on isolated specimens of materials that are dubious or as yet undescribed in detail. Morphologically, Late Cretaceous mammal tracks resemble those of some modern rodents. However, based on the dominant mammalian elements of faunas at that time they are probably of marsupial or multituberculate affinity.     

First Record of Theropod and Ornithopod Tracks and Detailed Description of Sauropod Trackways from the Tacuarembó Formation (Late Jurassic-?Early Cretaceous) of Uruguay

The record of dinosaur tracks and trackways from the Tacuarembó Formation (Late Jurassic-?Early Cretaceous) is described in detail: two trackways and four isolated tracks. Ichnites are preserved in sandstones of interdune facies of the Batoví (lower) Member. The interpretation herein presented indicates a sauropod affinity of the trackways, while the isolated prints are considered to have been made by an ornithopod and small theropods. The fossil record for this formation was primarily integrated by fresh-water organisms. The presence of new groups (sauropods and ornithopods) has specially contributed to the reconstruction of the terrestrial fauna, represented up until this time only by theropods.     

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.     

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.