Bird and Mammal Footprints From the Tertiary of Navarre (Western Pyrenees)

A rich variety of vertebrate footprints is known from a number of Upper Eocene to Lower Miocene localities of Navarre (western Pyrenees). The sediments were deposited in a wide range of depositional environments, from marginal marine to diversified terrestrial. Abundant bird tracks have been found in the coastal deposits of the Upper Eocene Liedena Sandstone of the Yesa and Itzagaondoa areas. Ciconiiformes-like (Leptoptilostipus pyrenaicus) and Charadriiformes-like (Charadriipeda ichnospp.) footprints have been recognized. Mammal ichnites have been discovered in the Oligocene and Lower Miocene deposits of Navarre. Equoid perissodactyl ichnites similar to those of Plagiolophustipus occur in the Oligocene fluviatile rocks of the Mués Sandstone of Olexoa and the Rocaforte Sandstone near Oibar and Sada. Trackways of entelodontids (Entelodontipus) are known in fluviatile-palustrine beds of the Oligocene Mués Sandstone of Olkotz. Additionally, bird (Charadriiformes-like) tracks are known in fluviatile-palustrine floodplain deposits of the Lower Miocene Ujué Formation of Los Arcos. In the same area, the Desoio and Los Arcos outcrops have also yielded perissodactyl trackways of possible Equoidea. Trackways of rhinocerotids (?) and artiodactyls (possibly Pecoripeda) are described from the Lower Miocene (Ramblian) palustrine limestones marginal to the Lerín Formation of Kaparroso and from alluvial fan deposits of the Uncastillo-Perdón Formation of Altzorritz, respectively.     

A NEW WIDE-GAUGE SAUROPOD TRACK SITE FROM THE LATE CRETACEOUS OF MENDOZA, NEUQUÉN BASIN, ARGENTINA

Abstract:  Agua del Choique is a new Late Cretaceous sauropod track site from Mendoza Province, Neuquén Basin, Argentina. It is situated in the Loncoche Formation, late Campanian – early Maastrichthian in age, and is one of the youngest sauropod tracks site recorded in the world. Agua del Choique represents a lake setting and river-dominated delta deposits, and comprises at least 160 well-preserved tracks, located on a calcareous sandstone bed. A new ichnotaxon, Titanopodus  mendozensis ichnogen. et ichnosp. nov., is erected for the footprints of this track site. Titanopodus  mendozensis exhibits the following association of features: (1) wide-gauge trackway (manus and pes trackway ratios of 18–22 and 26–31 per cent respectively), (2) pronounced heteropody (manus-pes area ratio of 1:3), (3) outer limits of trackway defined, in some cases, by the manus tracks, and (4) manus impression with an asymmetrical crescent contour and acuminate external border. Titanopodus  mendozensis is an excellent case study of the wide-gauge style of locomotion produced by Late Cretaceous derived titanosaurs that have no impression of manual phalanges. These features, and the fossil record from the Loncoche Formation, suggest that the trackmakers were, probably, middle size saltasaurine or aeolosaurine titanosaurs (14–16 m long).     

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 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.     

Magnetostratigraphy of the Sherwood Sandstone Group (Lower and Middle Triassic), south Devon, UK: detailed correlation of the marine and non-marine Anisian

The Sherwood Sandstone Group in south Devon consists of the Budleigh Salterton Pebble Beds and the Otter Sandstone Formation. The Otter Sandstone Formation comprises four lithostratigraphic sub-units, from the base labelled A-D, and contains tetrapods corresponding to the Perovkan land vertebrate faunachron, and is consequently a key European non-marine Anisian succession. A total of 181 palaeomagnetic specimens from 92 horizons were measured to determine the palaeomagnetic signal. The magnetic properties and characteristic remanence is predominantly carried by haematite, most of which probably formed early in the depositional and diagenetic history. The lower part of the Sherwood Sandstone Group (Budleigh Salterton Pebble Beds and Otter Sandstone Formation, unit A, lower part of unit B) has predominantly normal polarity (magnetozones BS1, BS2 and BS3n) with three thin reversed polarity magnetozones. The lower part of unit B to the lower part of unit C of the Otter Sandstone Formation is predominantly reverse polarity (magnetozones BS3r, BS4 and BS5), but includes three short normal polarity intervals. The upper ca. 50 m of unit C shows a mix of normal and reverse polarity (BS6, BS7) with normal dominating. The ca. 15-m-thick unit D is entirely of normal polarity (BS8n). The magnetostratigraphy allows unambiguous comparison with conodont and ammonite calibrated magnetostratigraphies from elsewhere in Europe. These show that the BS2 and BS3n magnetozones of the Otter Sandstone Formation are lower Aegean to middle Pelsonian (lower to middle Anisian). The BS3r to BS5r magnetozone interval, is middle Pelsonian to lower Illyrian (middle to upper Anisian), and the BS6 to BS8n magnetozone interval is middle Illyrian to lower Fassanian. The top of unit D is correlated to an interval in the middle of the Nevadites Zone (close to the Anisian-Ladinian boundary). Consequently, the Otter Sandstone Formation correlates with the Muschelkalk in the Polish Basin, but is equivalent to only part of the Germanic Basin Muschelkalk. The continuous magnetostratigraphy from the Sherwood Sandstone Group, provides additional details of the polarity pattern through the Pelsonian to Illyrian interval, which appears to be lacking in discontinuous marine sections. A synthesis of adjacent European sedimentary basins in combination with the magnetostratigraphy, suggests that the Budleigh Salterton Pebble Beds are probably Late Spathian to lower Aegean.     

The oldest evidence of a sauropod dinosaur in the western united states and other important vertebrate trackways from grand staircase‐escalante national monument,Utah

Recent discoveries of abundant fossil footprints from the new Grand Staircase‐Escalante National Monument of southern Utah, have important implications for the spatial and temporal distribution of Mesozoic vertebrates in Triassic and Jurassic time. Since the monument's creation in 1996, fossil footprints have been reported from at least seven formations in the Mesozoic (Triassic‐Cretaceous) within the monument. By far the most significant of these discoveries are sauropod and theropod tracks from the upper part of the Middle Jurassic Entrada Sandstone and a large Apatopus trackway from the Late Triassic Chinle Formation. Tracks in the Entrada Sandstone are found at the same stratigraphic level as those in the Moab megatracksite, and so considerably extend this large ichnological complex. A wide‐gauge sauropod trackway (cf. Brontopodus) from this unit represents the first reported from the Entrada Sandstone, and so is the oldest known from the western United States. This trackway also reveals a tail trace, which is the first reliable record of a sauropod tail trace.     

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.     

Large Carnivore Footprints from the Late Pleistocene of Argentina

The aim of the present contribution is to describe large felipedid footprints from a new ichnological site from the Late Pleistocene of Buenos Aires Province, Argentina. The prints are referred as the new ichnospecies Felipeda miramarensis nov. ichnosp. Based on size and morphology, this new ichnotaxon may have been produced by the large machairodontine felid Smilodon populator. Track analysis indicates that the producer of the tracks had fully retractile claws, a plantigrade feet, and lacked strong supination capabilities on pes and manus. The size and depth differences between manus and pes prints indicate that the producer had notably robust anterior limbs. If correctly assigned, the new ichnospecies reinforces the idea that Smilodon was an ambush predatory mammal.     

Las cerradicas tracksite (Berriasian,Galve, Spain): Growing evidence for quadrupedal ornithopods

Dinosaur footprints occur in shallow marine sedimentary units of the Berriasian Villar del Arzobispo Formation in the Aliaga basin, NE Spain. Las Cerradicas is a small outcrop (25 m²), near Galve, with four dinosaur trackways. Three tridactyl trackways indicate bipedal animals, and have parallel orientation. A fourth or‐nithopod trackway indicates a quadrupedal animal and is the smallest among any reported in the literature (L = 23 cm, W = 23 cm of pes tracks). It has manus prints that are oval‐shaped impressions. These and other recent discoveries shed much light on the abundance of quadrupedal ornithopods during the early Cretaceous.     

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.     

Eocene Bird and Mammal Tracks from the Karaj Formation,Tarom Mountains,Northwestern Iran

A long continuous trackway of a shorebird (cf. Charadriipeda) and a suite of possible Pantodonta (?Corophydon) or Dinocerata tracks are reported from the Eocene Karaj Formation. Fossil footprints are still rare in Iran, described only from two Jurassic and three Tertiary sites. Thus the current record, the third in the Teritary, adds to a sparse record and increases the diversity of known track types.     

Dinosaur Footprints from the Lower Cretaceous Sarten Member of the Mojado Formation at Cerro de Cristo Rey,Dona Ana County,New Mexico

Dinosaur tracks and swimming traces have been discovered at three localities in the latest Albian Sarten Member of the Mojado Formation, Bisbee Group (= “Anapra Sandstone”), at Cerro de Cristo Rey in Sunland Park, southernmost Dona Ana County, New Mexico. These localities preserve footprints of ornithopod (Caririchnium) and theropod (Magnoavipes) dinosaurs, ?reptilian swimming traces and possible tracks of an ankylosaurian dinosaur. The Sarten Member is of the latest Albian age, so the Cerro de Cristo Rey tracks are slightly younger than the well-known late Albian tracksites of northeastern New Mexico. At Cerro de Cristo Rey, the dominance of ornithopod tracks and absence of sauropod tracks fit regional patterns of late Albian-early Cenomanian track distribution consistent with North American extirpation of sauropods before the end of Albian time. The deltaic/coastal plain depositional setting of the Sarten Member is also remarkably similar to the track-bearing late Albian-Cenomanian sandstones of NE New Mexico, Oklahoma, Nebraska, and SE Colorado, which also have a tetrapod footprint ichnofacies dominated by ornithopod (Caririchnium) and theropod (Magnoavipes) tracks throughout the so-called “dinosaur freeway.”     

Middle Pennsylvanian Ichnofauna from Eastern Oklahoma,USA

The Keota site, in the Middle Pennsylvanian (Desmoinesian) Keota Sandstone Member, McAlester Formation (Krebs Group) of Haskell County, Oklahoma, yields an extensive ichnofossil assemblage of arthropod trackways, insect resting traces, arthropod feeding and grazing traces, fish swimming traces, and tetrapod footprints. This ichnofossil assemblage occurs in a 0.8 to 1.5-m-thick unit of thinly laminated and ripple-laminated sandstone at the base of the Keota Sandstone Member that is interpreted as a tidal flat sandstone. The arthropod traces are assigned to the ichnotaxa Diplichnites gouldi Gevers types A and B, Diplopodichnus biformis Brady, Paleohelcura tridactyla Gilmore, Tonganoxichnus buildexensis Mángano, Buatois, Maples and Lanier, Gordia marina Emmons, cf. Cochlichnus sp. and Treptichnus bifurcus Miller. The fish swimming traces are assigned to Undichna britannica Higgs, and the tetrapod tracks to Notalacerta ichnosp. and Pseudobradypus ichnosp. The Keota ichnofossil assemblage thus documents the first North American Pennsylvanian record of Paleohelcura tridactyla and the first record of Pseudobradypus from the western United States. This ichnofossil assemblage also provides compelling evidence of a freshwater habitat in a tidal flat setting during deposition of the fossil-bearing sandstone at the Keota site. Indeed, the Keota ichnofossil assemblage is characteristic of a Carboniferous-Permian Tonganoxichnus assemblage that identifies nonmarine tidal flat settings.     

记河北赤城上侏罗统-下白垩统土城子组窄足龙-巨齿龙(兽脚类恐龙)足迹组合及其古生态学意义(英文)

记述了河北省赤城县落凤沟化石点163个窄足龙足迹(Therangospodus isp.)。当该地区洪水泛滥时,同样的造迹者留下了5个游泳迹。一个异常大的足迹和一道疑似尾迹被归入巨齿龙足迹(Megalosauripus isp.)。土城子组7个主要足迹化石点表明,兽脚类的似鹬龙足迹型(Grallator morphotype)占据主导地位,这些足迹型的大小在各个足迹化石点都有着特定的范围。如果这些大小不同的足迹属于同一种造迹者,则暗示着同年龄的成员在特定年龄时会共同栖息在一起,在动物行为学上类似现生的一些蜥蜴和短吻鳄;如果这些足迹由不同的造迹者组成,则暗示着不同大小的种(主要表现在小型动物)可能拥有自己的领地,并特意避免与其他动物(特别是大型动物)相接触,在动物行为学上类似现生的肉食性哺乳动物。     

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.     

The Mississippian Tetrapod Footprint Ichnogenus Palaeosauropus: Extramorphological Variation and Ichnotaxonomy

Palaeosauropus primaevus is a tetrapod footprint ichnotaxon first described from the Upper Mississippian (Visean) Mauch Chunk Formation near Pottsville, Pennsylvania, United States. Our relocation of the type locality and stratigraphic horizon of P. primaevus, a long-available but unstudied collection of tetrapod footprints from these strata, and our new collections allow a much fuller characterization of this ichnotaxon and the range of extramorphological variation encompassed by it. P. primaevus is characterized as the footprints of a quadruped with a pentadactyl pes and a tetradactyl manus, in which the pes frequently oversteps the manus and with which tail drags are common. In the manus, all digits are relatively broad and have rounded tips, digit III is longest, and digit IV is more widely separated from digit III than the other digits are from each other. The pes has five digits that are also wide and blunt-tipped, digit IV is longest, and digit V projects nearly laterally. P. primaevus is the track of a relatively large temnospondyl (～400 mm gleno-acetabular length) and documents the Mississippian presence of such large amphibians long before their body fossil record. Palaeosauropus also occurs in Mississippian strata in Indiana and is distinguished from the geologically younger but similar temnospondyl footprint ichnogenus Limnopus by its relatively narrower manus and pes that lack broad and rounded sole impressions.     

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.     

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.     

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.     

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.