Track Assemblage Research Articles

A diverse saurischian (theropod–sauropod) dominated footprint assemblage from the Lower Cretaceous Jiaguan Formation in the Sichuan Basin, southwestern China: A new ornithischian ichnotaxon, pterosaur tracks and an unusual sauropod walking pattern

A sample of fallen blocks of fluvial sandstone from the Lower Cretaceous Jiaguan Formation of Sichuan Province yielded an assemblage of dinosaur and pterosaur tracks preserved as natural impressions and casts. Collectively the assemblage reveals 132 tracks representing at least 30 trackways of tridactyl and didactyl theropods, sauropods, ornithopods and pterosaurs. Ichnotaxonomically, the trackways of small tridactyl theropods (pes lengths 7–18 cm) are indeterminable, whereas the trackway of a small didactyl dromaeosaur (pes length up to 7.5 cm) is tentatively assigned to cf. Velociraptorichnus. The sauropod trackways are assigned to cf. Brontopodus based on the medium to nearly wide-gauge pattern. Other characteristics are the U-shaped manus and strong heteropody. One sauropod trackway shows a peculiar pattern with a lack of left manus imprints, and an unusual position and rotation of right manus imprints. Different scenarios and explanations for this phenomenon are discussed. Ornithopod trackways are the most abundant in the sample and characterized by pes imprints of a small biped that are assigned here to the new ichnospecies Caririchnium liucixini. It is characterized by an unusual broad shape and weak mesaxony. Bivariate analysis of different Caririchnium ichnospecies reveals increasing mesaxony toward the larger forms, a trend that is the reverse of typical theropod ichnotaxa, where large imprints have weak mesaxony. Three isolated, small pterosaur tracks (two manus, one pes) are visible on a single surface. They show strong similarities to the widespread ichnogenus Pteraichnus. This is the ninth report of tetrapod tracks from the Jiaguan Formation in recent years and represents one of the most diverse assemblages recorded to date. It is also rare evidence of typical didactyl dromaeosaur tracks and the co-occurrence of sauropod and ornithopod tracks in a fluvial depositional environment representing arid climate conditions.

Chirotherium trackways and feeding traces on seismic-influenced carbonate intertidals of the Middle Triassic of Central Europe: global food chain reactions onto horseshoe crab reproduction mud flat beaches of the Germanic Basin

Newly excavated Middle Triassic reptile Chirotherium, Rhynchosauroides and horseshoe crab Kouphichnium imprints from the uppermost Pelsonian megatracksite Bernburg, Germany (Central Europe) represent different behavioural trackways. The Chirotherium/Rhynchosauroides/Kouphichnium track assemblage of the lower intertidal carbonate flats is dominated by well-known horseshoe crab tracks of various subaquatic benthic movements which were all left on mud-cracked biolaminites in reproduction coastal zones. Most abundant reptile Rhynchosauroides trackways (50 % of reptile tracks) are oriented more beach-parallel (to Rhenish Massif Island) as a result of food searching along the intertidal seismic-influenced coast (16 slickensided biolaminate layers within 2 m biolaminates due to Alpine fold belt and Germanic intracratonic Basin tectonics), most probably for feeding on high abundant horseshoe crab eggs resulting from mating events in the lower intertidal. Large basal archosaur predators (?Ticinosuchus) produced large Chirotherium trackways (25 % of reptile tracks) which are represented in the new material with exceptional unusual trackways (slipping and side steps), but in directions to and from the beach to the Rhenish Massif main land, indicating predation on small reptiles. Finally, medium-sized Chirotherium trackways of subadult animals are present (25 % of reptile tracks). One trackway has a unique “feeding place” area, indicating the unexpected either predation, playing or feeding of a basal archosaur on a horseshoe crab. The horseshoe crab migrations from the North Tethys onto the extended mating intertidal mud flat beaches of the Germanic Basin, and recently known even into Chinese lagoons migrating limulid populations of different horseshoe crab species must have caused a global and seasonal food chain reaction, whereas Macrocnemus reptiles (Rhynchosauroides trackmaker) might have fed only on Millions of their eggs.

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.

New Dinosaur Footprints from the Upper Cretaceous of Poland in the Light of Paleogeographic Context

The Late Cretaceous track assemblage of southeastern Poland includes intriguing ichnotaxa with Asian or North American affinities. New finds of the ceratopsian-like ichnite and small hadrosauroid track refer directly to this problem, whereas the re-evaluation of foraminifer taxa estimates the stratigraphic positions of the track-bearing horizon and allowed to understand paleogeographic context of the studied and adjacent areas. The results demonstrate that during the Late Cretaceous times, migration of dinosaurs was possible from Asia through the East European Land to Kukeritz Island and the islands at peri-Tethys area.

Cretaceous saurischian tracksites from southwest Sichuan Province and overview of Late Cretaceous dinosaur track assemblages of China

Dinosaur tracks from Upper Cretaceous deposits of Sichuan Province, China have proved of great significance for two reasons: they (1) provide unambiguous evidence of dinosaurs from this epoch in this region, and (2) demonstrate that the track bearing units of the Leidashu Formation are not Paleocene-Eocene in age as previously claimed by some authors. We herein describe five small sites, from the Zhaojue area and the nearby Xide areas. Two sites are dominated by theropod tracks, including one with small tracks of Eubrontes-like morphology, and another with deep tracks revealing long metatarsal impressions, probably indicating a soft substrate. Three sites yield only sauropod tracks. Combined data from these five tracksites document a typical saurischian dominated ichnofauna, and based on present evidence these assemblages are less diverse than those from Lower Cretaceous sites in the region. However a larger sample is necessary to test this suggestion.

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.

Recent vertebrate tracks in sandy substrates and their paleoenvironmental implications: examples from coastal Lithuania

A diverse suite of vertebrate footprints documented along the coast of Lithuania offers insight into the distribution and preservation potential of tracks in sandy substrates. Whereas ichnotaxonomy is not conventionally applied to modern tracks, a case is made for using footprint classification based on Vialov’s (1966) nomenclature to both observed and predicted track distribution based on extramorphological characteristics of the tracemaker. Along a barrier spit (Curonian Spit) and mainland shoreline (Būtingė) sites, two general types of avian footprints (Avipeda sp), as well as three carnivore (Bestiopeda), four ungulate (Pecoripeda, Suipeda, Hippipeda), and two rodent (Rodentipeda) track morphotypes are described. The habitat preferences result in a predictable distribution of track assemblages with a potential of differentiating vertebrate ichnofacies (lagoon shoreline, open sea beach, dunes). The moisture content, possibility of freezing and rapid aeolian burial, and lithological anomalies (e.g., heavy-mineral concentrations) enhance the preservation potential of tracks. Once preserved, the attitude of trackways in respect to bedding surfaces determines their integrity and here a model of potential regional track distribution is proposed. Track formation and preservation in modern settings provides a useful guide to locating and studying vertebrate footprints and other traces in sand-dominated coastal and aeolian sequences.

A review of dinosaur track occurrences from the Morrison Formation in the type area around Dinosaur Ridge

Abstract Although the type section of the Morrison Formation, near Denver Colorado, now also well-known as Dinosaur Ridge, has since 1877 been world famous as the source of iconic Late Jurassic dinosaurs like Stegosaurus , Apatosaurus , and Diplodocus , little detailed information has been published on fossil footprints from the formation in this area. Late Jurassic (Morrison) footprints were not reported from the type section at Dinosaur Ridge until the early 1990s. Since then, other footprints have been reported, and their precise stratigraphic occurrences documented. To date, diagnostic examples of theropod, sauropod, and thyreophoran (stegosaur) tracks have been documented from at least three different locations. The quality of preservation of these tracks ranges from moderately good to quite poor. Nevertheless, more than 25 individual footprints, including some undertracks, have been identified unequivocally, including several manus–pes sets of quadrupedal trackmakers. The track assemblages are typical of many small Morrison Formation ichnofaunas and are consistent with what is known of the body fossil assemblages. The track assemblage is reviewed and compared with summary data from the 64 Morrison Formation tracksites currently known.

Reanalysis of Wupus agilis (Early Cretaceous) of Chongqing, China as a Large Avian Trace: Differentiating between Large Bird and Small Non-Avian Theropod Tracks.

Trace fossils provide the only records of Early Cretaceous birds from many parts of the world. The identification of traces from large avian track-makers is made difficult given their overall similarity in size and tridactyly in comparison with traces of small non-avian theropods. Reanalysis of Wupus agilis from the Early Cretaceous (Aptian-Albian) Jiaguan Formation, one of a small but growing number of known avian-pterosaur track assemblages, of southeast China determines that these are the traces of a large avian track-maker, analogous to extant herons. Wupus, originally identified as the trace of a small non-avian theropod track-maker, is therefore similar in both footprint and trackway characteristics to the Early Cretaceous (Albian) large avian trace Limiavipes curriei from western Canada, and Wupus is reassigned to the ichnofamily Limiavipedidae. The reanalysis of Wupus reveals that it and Limiavipes are distinct from similar traces of small to medium-sized non-avian theropods (Irenichnites, Columbosauripus, Magnoavipes) based on their relatively large footprint length to pace length ratio and higher mean footprint splay, and that Wupus shares enough characters with Limiavipes to be reassigned to the ichnofamily Limiavipedidae. The ability to discern traces of large avians from those of small non-avian theropods provides more data on the diversity of Early Cretaceous birds. This analysis reveals that, despite the current lack of body fossils, large wading birds were globally distributed in both Laurasia and Gondwana during the Early Cretaceous.

Hints of the early Jehol Biota: important dinosaur footprint assemblages from the Jurassic-Cretaceous boundary Tuchengzi Formation in Beijing, China.

New reports of dinosaur tracksites in the Tuchengzi Formation in the newly established Yanqing Global Geopark, Beijing, China, support previous inferences that the track assemblages from this formation are saurischian-dominated. More specifically, the assemblages appear theropod-dominated, with the majority of well-preserved tracks conforming to the Grallator type (sensus lato), thus representing relatively small trackmakers. Such ichnofaunas supplement the skeletal record from this unit that lacks theropods thus far, proving a larger diversity of dinosaur faunas in that region. Sauropods are represented by medium to large sized and narrow and wide-gauge groups, respectively. The latter correspond with earlier discoveries of titanosauriform skeletons in the same unit. Previous records of ornithischian tracks cannot be positively confirmed. Purported occurrences are re-evaluated here, the trackways and imprints, except of a single possible specimen, re-assigned to theropods. Palecologically the Tuchengzi ichnofauna is characteristic of semi-arid fluvio-lacustrine inland basins with Upper Jurassic-Lower Cretaceous deposits in northern China that all show assemblages with abundant theropod and sauropod tracks and minor components of ornithopod, pterosaur and bird tracks.

Tracking the yellow dragons: Implications of China's largest dinosaur tracksite (Cretaceous of the Zhucheng area, Shandong Province, China)

Surfaces with more than 2200 dinosaur footprints from the Huanglonggou (Yellow Dragon Valley) site near Zhucheng, in Shandong Province, were excavated for scientific study and with a view to future development as an educational site suitable for further research and tourism. Although geographically close to spectacular and historically famous Upper Cretaceous sites yielding vast bone assemblages, representing giant hadrosaurs and other dinosaurs from the Wangshi Group, the tracksite is in the Lower Cretaceous Yangjiazhuang Formation (equivalent in part to the Longwangzhuang Formation), represents an entirely different dinosaurian fauna, dominated by small theropods. In contrast to a recent pre-excavation study of a localized outcrop which identified only three theropod track morphotypes, in a sample of 135 tracks, the present study has identified at least 2000 additional tracks including those of sauropods and turtles. It is therefore possible to present a more complete interpretation of the site based on the larger and more diverse track assemblage presently exposed. Three theropod track morphotypes are identified as grallatorid morphotypes A and B, with the latter assigned to Grallator yangi comb. nov., and Corpulentapus lilasia . Tracks have been identified from at least 5 levels, of which level 4 exhibits the vast majority in an excellent state of preservation. Other recent studies, which we re-evaluate, suggest the tracks help define an ENE–WSW shoreline with the lake center to the SSE. In terms of number of tracks documented the Huanglonggou site is one of the largest dinosaur tracksites in China, or indeed in the world. • The largest mapped dinosaur tracksite in China • Five trackway morphotypes, most with exquisite preservation • Shore line micro-paleogeography mapped in fine detail • Unique track superimposition patterns are identified and explained.

A manus dominated pterosaur track assemblage from Gansu, China: implications for behavior

The Yangouxia dinosaur tracksites are well known for a diverse assemblage of tetrapod tracks preserved as natural impressions (concave epireliefs) on large bedding planes, representing a locally widespread surface marking the transition from a sand- to a mud-dominated sequence in the Hekou Group. Previous ichnological studies at these large sites have focused on the morphology and ichnotaxonomy of the tracks, including a single trackway representing the first pterosaur tracks reported from China. Here, we report a distinctly different assemblage associated with minor sandstones in the mud-dominated sequence 20m above the main tracksite level. This assemblage consists of at least 20 pterosaur manus track casts attributed to a single ichnotaxon (Pteraichnus). No pes tracks have been identified. These tracks mostly occur in random orientations, although one possible trackway segment is inferred, to represent walking progression. Manus-only pterosaur track assemblages are common and likely reflect differential registration depths of manus and pes and/or sub optimal preservation conditions. The tracks are associated with distinctive invertebrate traces including Cochlichnus, Spongeliomorpha and Paleophycus and suggest the pterosaurs were likely feeding on the invertebrate tracemakers.

TRACKWAYS OF A GREGARIOUS, DUNEFIELD-DWELLING, EARLY JURASSIC THERAPSID IN THE AZTEC SANDSTONE OF SOUTHERN NEVADA

ABSTRACT We describe and interpret a tracksite in the Lower Jurassic Aztec Sandstone in Valley of Fire State Park, southern Nevada. The site contains approximately one hundred tracks of the ichnogenus Brasilichnium, arranged in twelve, subparallel trackways, all on the same foreset bedding plane. The Brasilichnium trackmaker was most probably a fossorial, tritylodontid therapsid. Sedimentologicial analyses indicate that the trackway surface is a wind-ripple horizon with a primary dip of about 25°, and that the animals climbed straight up the slip face of the dune. A combination of features leads us to conclude that the footprints were impressed into a crust of moist, cohesive sand, leaving two modes of preserved tracks: (1) shallow, well-defined tracks without associated sand crescents, and (2) deeper, less well defined tracks with associated sand crescents. We interpret this assemblage of tracks to record gregarious behavior in a mixed-age group of tritylodontid therapsids. In the correlative Navajo Sand...

Upper Cretaceous dinosaur track assemblages and a new theropod ichnotaxon from Anhui Province, eastern China

Several noteworthy Cretaceous tracksites are known in the Huizhou District (Huangshan City) in the Yangtzee valley of southern Anhui Province. These include some that have been known since the late 1970s but have not been studied in detail until now. The footprints described here occur in siliciclastic fluvial deposits in three distinct horizons. The Xiaohutian tracksite in the Upper Cretaceous Xiaoyan Formation is the most interesting, being situated at a historically famous location used as a Taoist and Buddhist shrine. The Xiaohutian tracksite yields an assemblage with three different morphotypes of non-avian theropod tracks including the new ichnotaxon Paracorpulentapus zhangsanfengi that can be attributed to a theropod with relatively short “fleshy” toes showing convergence with the footprints of small ornithopods. A further diagnostic feature is the trackway pattern with relatively short steps. Associated ichnofossils are invertebrate traces that can be assigned to eurybathic forms such as Palaeophycus and ?Planolites or ?Scoyenia. Thus far skeletal remains from the Xiaoyan Formation have proved the pachycephalosaur Wannanosaurus and indeterminate sauropods. The ichnoassemblages enlarge the known dinosaur fauna by small- to medium-sized theropods that are identified here as the trackmakers and that are otherwise rare in Upper Cretaceous deposits of eastern China. The Shangshangen locality is another significant tracksite which has yielded small bird tracks (cf. Koreanaornis) in association with small tracks of non-avian theropods.

New Early Cretaceous Pterosaur‐Bird Track Assemblage from Xinjiang, China: Palaeoethology and Palaeoenvironment

AbstractA pterosaur‐bird track assemblage from a sandstone‐siltstone‐mudstone sequence of the Lower Cretaceous Tugulu Group of Xinjiang comprises the first pterosaur track record from this province and the largest specimen thus far known from China. The pterosaur tracks are assigned to the ichnogenus Pteraichnus based on the triangular overall‐shape, the four elongate digit traces and the robust manual digit trace III. Supposed trackmakers were dsungaripterid pterodactyloids whose skeletal remains are well known from the Tugulu Group. The bird tracks that occur on the same surface, are those of typical shorebirds, known from different other localities in southeast Asia. The congruence with Koreanaornis dodsoni described from the same stratigraphic level justifies an assignment to this ichnospecies. This is a further evidence of the co‐occurrence of pterosaurs and birds in a typical lakeshore environment with possible seasonal alteration of water supply and aerial exposure indicated by wave ripples, mudcracks and repeated cycles of coarse to fine sediment. Pterosaurs and birds frequented the shoreline and may have fed also on the numerous invertebrates such as the Scoyenia tracemaker that left abundant burrows.

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.

Diverse dinosaur ichnoassemblages from the Lower Cretaceous Dasheng Group in the Yishu fault zone, Shandong Province, China

New dinosaur track assemblages were discovered recently in the Tianjialou Formation of the Lower Cretaceous Dasheng Group in Shandong Province, China. Theropods are represented by the trackways of two different medium-sized groups: (1) tridactyl tracks with a typical mesaxonic shape; (2) functionally didactyl tracks attributed to deinonychosaurian theropods. The latter report, the third from the Cretaceous of Shandong Province, enlarges the global record of didactyl theropod tracks, until now sparsely documented from only a few locations in Asia, North America and Europe. A number of features in the dromaeosaur trackway suggest the assignment to cf. Dromaeosauripus. Several medium-sized trackways resemble the narrow-gauge, small manus ichnogenus Parabrontopodus, and one large trackway is characterised by a wide-gauge and large manus, similar to Brontopodus. This suggests the co-occurrence of two different sauropod groups. A further component in these ichnoassemblages is a tetradactyl morphotype and trackways of ornithischian affinity that are tentatively attributed to psittacosaurs.

Lark Quarry revisited: a critique of methods used to identify a large dinosaurian track-maker in the Winton Formation (Albian–Cenomanian), western Queensland, Australia

Thulborn, R.A., 2013. Lark Quarry revisited: a critique of methods used to identify a large dinosaurian track-maker in the Winton Formation (Albian–Cenomanian), western Queensland, Australia. Alcheringa, http://dx.doi.org/10.1080/03115518.2013.748482 A remarkable assemblage of dinosaur tracks in the Winton Formation (Albian–Cenomanian) at Lark Quarry, a site in western Queensland, Australia, has long been regarded as evidence of a dinosaurian stampede. However, one recently published study has claimed that existing interpretation of Lark Quarry is incorrect because the largest track-maker at the site was misidentified and could not have played a pivotal role in precipitating a stampede. That recent study has identified the largest track-maker as an ornithopod (bipedal plant-eating dinosaur) similar or identical to Muttaburrasaurus and not, as formerly supposed, a theropod (predaceous dinosaur) resembling Allosaurus. Those iconoclastic claims are examined here and are shown to be groundless: they are based partly on misconceptions and partly on fabricated data that have been assessed uncritically using quantitative measures of questionable significance. Such ill-founded claims do not reveal any substantial flaw in the existing interpretation of the Lark Quarry dinosaur tracks.

A new early cretaceous dinosaur track assemblage and the first definite non-avian theropod swim trackway from China

The trackway of a swimming theropod (ichnogenus Characichnos) is reported from the Lower Cretaceous Feitianshan Formation of Sichuan, China. These swim tracks help confirm that non-avian theropods were capable of forging moderately deep bodies of water. The trackway occurs on the same surface as a typical walking trackway of a sauropod (ichnogenus Brontopodus). Both occurrences are the first reported from the Cretaceous of Sichuan, and the swim tracks are the first well-preserved example of a Characichnos trackway from China. Additionally, a theropod walking trackway and several ornithopod walking trackways (similar to the ichnogenus Caririchnium) occur in the same horizon. The ornithopod trackways show a parallel orientation, suggesting gregarious behavior of the trackmakers, which may have been iguanodontiforms and/or hadrosauriforms. The co-occurrence of theropod swim tracks and theropod walking tracks suggests a fluctuation of water depth within a distinct time span.

Late Triassic Tetrapod-Dominated Ichnoassemblages from the Argana Basin (Western High Atlas, Morocco)

Diverse tetrapod track assemblages with Scoyenia invertebrate traces were discovered in the Triassic Timezgadiouine and Bigoudine formations of the Argana Basin (Western High Atlas, Morocco). The ichnofossils occur in alluvial plain sandstones and mudstones of the Irohalène Member (T5) and Tadart Ouadou Member (T6) considered Carnian-Norian in age by vertebrate remains and palynomorphs. Tetrapod footprints are assigned to Apatopus, Atreipus-Grallator, Eubrontes isp., Parachirotherium, cf. Parachirotherium postchirotherioides, Rhynchosauroides ispp., and Synaptichnium isp. They can be referred to lepidosauromorph/ archosauromorph, basal archosaur, and dinosauromorph trackmakers. Apatopus, represented by 11 tracks of a more than 4 m long trackway, is recorded for the first time outside of North America and Europe. The assemblage concurs with the proposed Late Triassic age of the track-bearing beds by the occurrence of Apatopus, Atreipus-Grallator, and Eubrontes. If this is accepted, the stratigraphic range of Synaptichnium and Parachirotherium, hitherto known only from Early or Middle Triassic deposits, has to be extended to the Carnian-Norian. The occurrence of Eubrontes in the Irohalene Member (T5) provides further evidence for large theropods in pre-Jurassic strata. All assemblages are referred to the Scoyenia ichnofacies indicating continental environments with alternating wet and dry conditions.