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.     

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.     

Track analysis and conservation priorities in the cloud forests of Hidalgo,Mexico

Abstract . A track analysis based on the distributional patterns of 967 species of vascular plant taxa (gymnosperms, angiosperms and pteridophytes) was performed to assess conservation priorities for cloud forests in the state of Hidalgo, Mexico, ranged in the municipalities of Chapulhuacán, Eloxochitlán, Molocotlán, Pisaflores, Tenango de Doria, Tlahuelompa and Tlanchinol, as well as five floristically equivalent areas in the states of Veracruz (Teocelo and Helechales), Tamaulipas (Gómez Farías), Morelos‐México (Ocuilan) and Oaxaca (Huautla de Jiménez). In order to detect generalized tracks we employed a new parsimony method, where clades (considered equivalent to generalized tracks) are defined forbidding homoplasy and acting like a compatibility algorithm. Several generalized tracks were found connecting these areas. Cloud forests of Chapulhuacán were connected according to three different generalized tracks and thus have a higher value, qualifying as a priority area for the conservation of cloud forests in the state of Hidalgo.     

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.     

Sometimes They Come Back: Recovery and Reinterpretation of a Trackway Slab from the Permian Coconino Sandstone of the Southwestern United States

When a fossil vanishes to a private collection, it must be considered lost to science because, frequently, it is no longer available for study. Fortunately some fossils occasionally are regained. We had the opportunity to recoup an interesting footprint-bearing slab that was part of a private collection in Italy. The specimen, found in 1992 near Seligman, Arizona (USA) was described, before disappearing, as one of the best fossil examples of vertebrate (Chelichnus [Laoporus])-on-invertebrate (Octopodichnus) predation. After a careful re-examination of the slab, the primary conclusions of the former describers are demonstrably groundless. The reanalysis of the tracks, as well as peculiar sedimentary structures associated with the tracks, allowed obtaining new information about the depositional environment and the complex interactions between the type of substrate and trackmaker behavior. The re-examination of the specimen also revealed interesting aspects about trackmaker biomechanics.     

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.     

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.     

Assessment of patch quality by ladybirds: role of larval tracks

Gravid females of the two-spot ladybird, Adalia bipunctata (L.), were deterred from ovipositing when kept in petri dishes that had previously contained conspecific larvae but not conspecific adults, or the larvae of another two species of ladybird, Adalia decempunctata (L.) and Coccinella septempunctata L. The deterrent effect was density dependent and mediated via a chloroform-soluble contact pheromone present in the larval tracks. Similarly, gravid females of C. septempunctata were deterred from ovipositing by conspecific larval tracks and chloroform extracts of these tracks, but not by the tracks or extracts of tracks of A. bipunctata larvae. That is, in ladybirds the larvae produce a species-specific oviposition-deterring pheromone. In the field, the incidence of egg cannibalism in ladybirds increases very rapidly with the density of conspecific eggs or larvae per unit area. Thus, in responding to the species specific oviposition deterring pheromone female ladybirds reduce the risk of their eggs being eaten and spread their offspring more equally between patches. Received: 14 March 1997 / Accepted: 26 August 1997     

History and nomenclature of the theropod dinosaur tracks Bueckeburgichnus and Megalosauripus

The most recent account of Bueckeburgichnus maximus Kuhn 1958, a distinctive theropod dinosaur track from the Lower Cretaceous of Germany, is shown to be based on a referred specimen mistakenly identified as the holotype and the correct name of this taxon is deemed to be Megalosauripus maximus (Kuhn 1958). This minor revision has important consequences for nomenclature of the many European, Asian, North American and Australian dinosaur tracks attributed to megalosaurian theropods. Many of those tracks were named Megalosauripus, but that name has a confusing multiplicity of meanings and it should be restricted to the highly characteristic dinosaur track formerly identified as Bueckeburgichnus. Other tracks named "Megalosauripus”; (in its several other senses) will require new nomenclature, despite their extensive and repeated revision since 1996. It is recommended that future revision should adopt conventions of the International Code of Zoological Nomenclature. Although previous revisions expressed an intention to adhere to those conventions, these were not put into practice, with the unfortunate result of multiplying the problems that surround the nomenclature of megalosaur tracks. Introduction of the name Megalosauripus maximus (Kuhn 1958) eliminates those burgeoning problems and permits the introduction of new and objective nomenclature for presumed megalosaur tracks.     

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.     

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

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

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.     

America's Most Famous Human Footprints: History,Context and First Description of Mid-Holocene Tracks from the Shores of Lake Managua,Nicaragua

Human tracks discovered in 1874 at a site named El Cauce or Acahualinca near the shores of Lake Managua, Nicaragua, are the most famous and abundant human tracks in the Americas. They represent a landmark ichnological discovery during the late 19th century that generated much debate regarding their age and origins. Reported dates for the tracks range from 2,120 to 6,500 B.P. The site, which is now situated within the limits of Nicaragua's capital city of Managua, forms the basis of the Acahualinca Tracks Museum (Huellas de Acahualinca) and has been in place since 1953. However, it is still little known and has not been systematically studied, despite being an important window into Meso-American prehistory. Two exposures of a surface of volcanic ash reveal hundreds of human tracks comprising a minimum of 12 clearly defined trackways, and a trampled zone or path representing at least three more individuals, all heading in the same northwesterly direction. Tracks of deer, opossum and at least one bird are also present. Bison tracks collected from another nearby site (El Recreo) that is no longer accessible are on display at the museum. The Acahualinca tracks are noteworthy for the exceptional quality of preservation. We herein present the first maps of the two exposures, which combine to tell a story of human and animal activity along the shores of ancient Lake Managua several thousand years ago.     

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.     

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.     

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.     

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.     

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).     

Grallator-Dominated Fossil Footprint Assemblages and Associated Enigmatic Footprints from the Chinle Group (Upper Triassic), Gateway Area,Colorado

The upper part of the Chinle Group (Late Triassic) of the Gateway area in western Colorado is extraordinarily rich in fossil footprint assemblages. Dominant track types include small Grallator tracks, generally attributed to Coelophysis-like theropods, which often occur in high densities of 50 to 100 per m². Other abundant ichnotaxa that are attributable to dinosaurs or dinosaur-like archosaurs include Pseudotetrasauropus and Tetrasauropus, attributed to prosauropods and sauropods, respectively. Several Pseudotetrasauropus-like tracks appear to be functionally didactyl and may indicate a new ichnotaxon that represents an animal that shows certain unusual features that are convergent with dromeosaurs and certain birds. Such convergence may reflect inherent growth programs as much as functional adaptations. Non-dinosaurian ichnotaxa include Brachychirotherium (probably of aetosaur affinity) and Rhynchosauroides, attributed to a sphenodontid/lizard-like form. Other ichnotaxa include probable therapsid (dicynodont) tracks labeled Pentasauropus sp., mammaloid (non-therian mammal and/or mammal-like reptile) tracks, and the trails of arthropods. Excellent preservation and high track densities mark the Gateway assemblages in a thin stratigraphic interval in the upper part of the Chinle Group (Rock Point Formation). The track assemblages are similar to those reported from the Chinle Group in other parts of the Colorado Plateau and Rocky Mountain region, extending over most of Colorado, Utah, northern Arizona and northern and eastern New Mexico. Some of the Chinle ichnotaxa (Grallator and Brachychirotherium) are found in the overlying Wingate Formation, indicating that it is also Late Triassic in age, at least in the lower part. However, overall the Chinle and Wingate assemblages are quite different, most notably in the rarity of mammaloid/mammal-like tracks in the Chinle Group.