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.     

Lunulichnus Tuberosus Ichnogen. and Ichnosp. Nov. from the Early Eocene Wasatch Formation,Fossil Butte National Monument,Wyoming: An Arthropod-Constructed Trace Fossil Associated with Alluvial Firmgrounds

A new trace fossil, Lunulichnus tuberosus, is described from fluvial deposits of the Wasatch Formation (early Eocene) at Fossil Butte National Monument, southwestern Wyoming, USA. L. tuberosus are straight, vertical to obliquely oriented, unlined cylindrical burrows with pronounced crescent-shaped wall sculptings. In situ examples of these trace fossils are most commonly preserved as sand-filled casts emanating from the erosional bases of fluvial channel sandstone bodies into underlying floodplain mudstone/siltstone beds. L. tuberosus is interpreted as the dwelling trace of a stream-dwelling decapod crustacean. Excellent preservation of fine detail, particularly their diagnostic crescent-shaped wall sculptings, support the hypothesis that L. tuberosus were excavated in firm substrata subjacent to fluvial erosional surfaces. As such, they are interpreted as constituents of alluvial Glossifungites trace fossil assemblages.     

A Suite of Dermestid Beetle Traces on Dinosaur Bone from the Upper Jurassic Morrison Formation,Wyoming, USA

Most studies of insect traces on fossil bone deal with one or two trace morphs found on isolated bone fragments, making it difficult to identify the trace-maker and its behavior. We report the discovery of a suite of insect traces on an articulated Camptosaurus dinosaur skeleton that permits the identification of the trace-maker and interpretations of its behavior. The traces include mandible marks, pits, and shallow bores on cortical bone, and deep, meandering furrows and tunnels (borings) on articular surfaces. The interiors of bones are intensely mined, and the cavities and borings are filled with fine bone fragments (insect frass). The distinctive mandible marks consist of opposing sets of parallel grooves, indicating the maker had two apical teeth set on symmetrical mandibles and that all of the traces were made by a single taxon. Comparison of the fossils with the mandible morphology and bone traces of extant insects indicates dermestid beetles made the traces. Based on extant dermestid behavior, soft tissues were likely absent and the bones were lipid-laden when the traces were made. Examination of more than 5,000 bones from the Morrison and Cedar Mountain formations shows insect traces on bone are common but overlooked and that many bones are substantially damaged by insect mining. The key to the recognition of these important yet subtle traces is a search model and an intense, oblique light source.     

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.     

Dinosaur Tracks from the Kilmaluag Formation (Bathonian,Middle Jurassic) of Score Bay,Isle of Skye,Scotland, UK

Tracks of a juvenile theropod dinosaur with footprint lengths of between 2 and 9 cm as well as adults of the same ichnospecies with footprints of about 15–25 cm in length were found in the Bathonian (Middle Jurassic) Kilmaluag Formation of Score Bay, northwestern Trotternish Peninsula, Isle of Skye, Scotland, UK. Two footprint sizes occur together on the same bedding plane in the central portion of Score Bay, both in situ and on loose blocks. Another horizon containing footprints above this was also identified. The footprints from the lowest horizon were produced in a desiccated silty mud that was covered with sand. A close association of both adults and juveniles with similar travel direction indicated by the footprints may suggest post-hatching care in theropod dinosaurs. Other footprints, produced on a rippled sandy substrate, have been found on the slightly higher bedding plane at this locality. Loose blocks found 130 m to the northeast in the central part of Score Bay have not been correlated with any in situ sediments, but were preserved in a similar manner to those from the higher bedding plane. These tracks represent the youngest dinosaur remains yet found in Scotland.     

Ammonite eggs and ammonitellae from the Kimmeridge Clay Formation (Upper Jurassic) of Dorset, England

Eight clusters of small spherical and subspherical objects, some isolated and some associated with shells of perisphinctid ammonites, have been recovered from the Lower and Upper Kimmeridge Clay (Upper Jurassic) of the Dorset coast, England. They have been interpreted as ammonite egg sacs and represent the freshest and best-preserved examples known so far. Their structures and the ecological framework in which they occur are discussed. The parents are thought to be members of the two eudemic genera Aulacostephanus and Pectinatites that dominate the biostratigraphy of the ammonites in the range of the Kimmeridge Clay in which they occur. Isolated nuclei of ammonitellae have also been recovered.     

Middle and Upper Jurassic bivalve associations from the Iberian Range (Spain)

The four sections richest in bivalves from the Middle and Upper Jurassic of the Iberian Range (Spain) were selected for a quantitative palaeoecological analysis of the bivalve fraction of the macrobenthos. Five bivalve associations and two assemblages were recognized with the help of a Q-mode hierarchical cluster analysis (Ward method). The main environmental factors controlling bivalve associations are thought to be substrate, water energy and distribution of organic matter. The bivalves exhibit a distinct spatial and temporal distribution pattern within the Aragonian Branch of the Iberian Range. Four of the bivalve associations occur in the Upper Oxfordian (Sot de Chera Fm) and one association in the Lower Callovian (Chelva Fm). In the Sot de Chera and Loriguilla formations, the abundance of bivalves decreases from NW to SE i.e., from relatively close to the shore line towards the distal-most part of the carbonate platform. In the Chelva Fm, bivalves are abundant in the Ariño region, interpreted as a palaeogeographic high. The spatial distribution of bivalves might have been largely controlled by the availability of nutrients.     

Fossil coniferous wood from the Middle Jurassic of Liaoning Province, China

Silicified coniferous wood was collected from the Lanqi Formation (late Middle Jurassic in age) at Shebudaigou Village, Liaoning Province, China. Three taxa are identified, namely Pinoxylon dacotense Knowlton, Xenoxylon phyllocladoides Gothan, and Araucariopitys sp. Based on these new data, and those of other fossil plants reported previously from the same formation, we consider the climate during the deposition of the Lanqi Formation was subtropical, humid and seasonal. In this respect the Lanqi flora differs from the coeval Shimengou and Longmen floras from North China. The Longmen flora was deposited during more humid, subtropical conditions, while the Shimengou Formation indicates that the climate was warm temperate and dry. Our data would suggest that the Late Jurassic climatic pattern was initiated as early as the late Middle Jurassic.     

Toarcian and Aalenian (Jurassic) ammonites from the Shemshak Formation of the Jajarm area (eastern Alborz, Iran)

In the Jajarm area (eastern Alborz Mountains, NE Iran) the ?Upper Triassic — Lower Bajocian siliciclastic Shemshak Formation is up to 2000 m thick. Whereas the lower third of the formation is nearly exclusively non-marine, the upper two-thirds are fully marine. The middle part is characterized by several levels containing a relatively diverse and well preserved assemblage of ammonites of the Toarcian to Aalenian stages. Two sections of the ammonite-bearing strata, spaced 20 km, are presented and correlated by means of ammonite biostratigraphy. The ammonite fauna consists of 27 taxa, some of which are recorded for the first time from the Alborz Mountains. The ammonites are briefly described and their palaeobiogeographic context is reviewed. The ammonite fauna of the Shemshak Formation at Jajarm, as elsewhere in North and Central Iran, is exclusively Northwestern Tethyan in character and is closely related to the faunas of Northwestern and Central Europe.     

An ammonite–fish association from the Kimmeridgian (Upper Jurassic) of La Rochelle, western France

A partially disarticulated actinopterygian fish preserved in a large three-dimensional ammonite body chamber is described from the Kimmeridgian of western France. Taphonomic observations on the degree of preservation of the fish and the development of epibiont organisms on the inner wall of the shell indicate a rather long time interval before sediment totally filled the body chamber. The fish, referred to an indeterminate Macrosemiidae, probably used this empty ammonite ( Rasenioides , Aulacostephanidae) shell as a refuge, or possibly for spawning and/or brooding. It can be assumed that ammonite shells may have constituted common shelters for demersal fishes living in an open-marine shelf environment, near to a muddy bottom devoid of rocks.     

New skull of Lesothosaurus (Dinosauria: Ornithischia) from the Upper Elliot Formation (Lower Jurassic) of southern Africa

A new ornithischian skull from the Elliot Formation of southern Africa is described. The specimen is compared in detail with the fabrosaurid Lesothosaurus diagnosticus. It actually shares many characters with specimens of the syntypes of this species or specimens referred to it. It is nevertheless not identical to any of these specimens and it is, moreover, remarkably larger than them. The possibility of attributing this specimen to a so far undescribed ‘large fabrosaur’ from the same formation is discussed. It is concluded that the specimen in question in this paper, while being ascribable to the genus Lesothosaurus, cannot be determined to a specific level until the existence of two fabrosaurid species in the ‘Stormberg Group’ is demonstrated and their range of morphological and size variation is properly appraised.     

The appendicular skeleton of Suuwasseaemilieae (Sauropoda: Flagellicaudata) from the Upper Jurassic Morrison Formation of Montana (USA)

Appendicular elements of the sauropod dinosaur Suuwasseaemilieae, from the Upper Jurassic Morrison Formation of Montana, USA, display a peculiar mix of autapomorphic and plesiomorphic features. While more similar in overall morphology to Apatosaurus than other flagellicaudatans, the coracoid of Suuwassea lacks the quadrangular shape of Apatosaurus. The humerus of Suuwassea bears a pronounced proximal tuberculum, a feature seen elsewhere only in saltasaurine titanosaurian sauropods. The rectangular proximal articular surface of the tibia is proportioned neither like Diplodocus nor Apatosaurus type specimens, although this region may be intraspecifically variable. The pes of Suuwassea possesses plesiomorphically elongate phalanges and a small, uncompressed ungual, unlike other flagellicaudatans except Dyslocosaurus. The localization of tooth marks on the pedal elements suggests that sauropod feet may have been singled out by scavengers, as has been noted for elephants.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Contribution to the scleractinian corals of Hanifa Formation,Upper Jurassic,Jabal Al-Abakkayn,central Saudi Arabia

Eighteen scleractinian coral species belonging to 13 genera, 8 families and 4 suborders have been identified from the lower and upper parts of the Upper Jurassic (Oxfordian) Hanifa Formation at Jabal Al-Abakkayn, central Saudi Arabia. Actinastrea bernensis, A. crassoramosa, Coenastraea hyatti, Stylina kachensis, Cryptocoenia slovenica, C. wegeneri, Isastrea hemisphaerica, I. bernensis, Montlivaltia cornutiformis, M. frustriformis, Collignonastraea jumarensis, Ovalastrea michelini and Vallimeandropsis davidsoni are believed to be recorded for the first time from the Jurassic rocks of central Arabia. Most corals have massive hemispherical and globular forms, and few corals have dendroid and conical growth forms. They occur as small, isolated patches, about 0.5 m thick and about 10–30 m wide, in argillaceous reefal limestones. The identified corals show Africa, north America, northern, southern and western Europe, and southern or eastern Asia corals. The low diversity and abundance as well as the small size of colonies are attributed to inimical palaeoecological factors throughout the reefoids formation such as muddy substratum, water turbidity, high rate of sedimentation.     

Trace Fossils in Eolian Facies of the Upper Triassic-Lower Jurassic Dinosaur Canyon Member,Moenave Formation,Northern Arizona

The Dinosaur Canyon Member of the Moenave Formation, of Late Triassic to Early Jurassic (Rhaetian-Hettangian) age, consists predominantly of sandstones and mudstones deposited by sheet and channelized flow and by eolian processes. We document several previously undescribed biogenic structures in these strata. At one location, eolian dune sandstone hosts the lined, smooth-walled burrow Palaeophycus heberti (Saporta); these may be the nesting burrows of small sand wasps, or alternatively, the burrows of sand-treader camel crickets. Eolian dune facies that host abundant rhizoliths also are typically bioturbated. Circular markings that are convex in epirelief are assigned to Pustulichnus gregarious Ekdale and Picard. Unlined, non-ornamented burrows are unassigned. We speculate that both of these also were formed by the burrowing activity of one of the many species of sand wasp. Rhizoliths are particularly prominent at the tops of eolian dune sandstone beds and locally may host termite nests. Stabilization of the landscape by plants permitted pedogenic alteration of the sediments.     

First report of Apatosaurus (Diplodocidae: Apatosaurinae) from the Cleveland-Lloyd Quarry in the Upper Jurassic Morrison Formation of Utah: Abundance,distribution, paleoecology,and taphonomy of an endemic North American sauropod clade

A cervical vertebra preserved at the famous and productive Cleveland-Lloyd Dinosaur Quarry in the Upper Jurassic Morrison Formation of Utah is that of an Apatosaurus, a sauropod dinosaur genus not previously recognized at the site and the first new dinosaur taxon identified at the site in years. The presence of Apatosaurus at a mudstone site dominated by other taxa, both theropod and sauropod, suggests a pattern of preservation within the Morrison Formation in which sites in fine-grained sediments yield dramatically uneven relative abundances of dinosaurs, with variable dominant taxa by site, compared with more time-averaged and attritional coarse-grained channel sandstone deposits. In addition, the continued demonstration of the wide-spread occurrence and abundance of Apatosaurus within the Morrison Formation, and the absence of its clade among diplodocid faunas on other continents, suggest that this group may have been endemic to North America during the Late Jurassic and that it may have originated there, though this is far from clear.     

Sunosuchus (Archosauria, Crocodyliformes) from the Toutunhe Formation (Middle Jurassic) of the Southern Junggar Basin (Xinjiang, NW-China)

A new specimen of the widespread Middle Jurassic to Lower Cretaceous Asian gonipholidid crocodilian genus Sunosuchus is described on the basis of a partial skeleton from the Upper Toutunhe Formation (Middle Jurassic, ?Bathonian-Callovian) of Liuhonggou, SW of Urumqi, Xinjiang Uygur Autonomous Region, People’s Republic of China. The specimen is represented by a partial mandible, teeth, vertebrae, limb and girdle bones and osteoderms. It can be distinguished from other nominal species of the genus by a unique combination of characters: slightly heterodontous dentition, strongly sculptured posteroventral part of the mandible, short fenestra mandibularis, convex dorsal surface of the retroarticular process, keeled cervical vertebral centra and ventral osteoderms with a distinctive sculpture of wide pits and narrow ridges. The heterodontous dentition is a potential autapomorphy of this form. The new specimen is closest in morphology to material described recently from the Callovian of Kirghisia as Sunosuchus sp. It represents the second Middle Jurassic record of the genus, the first crocodile from the Toutunhe Formation, the first substantial crocodile find from the Mesozoic of the Southern Junggar Basin, and the first Middle Jurassic record of Sunosuchus from China. This extends both the paleobiogeographical distribution of the genus in Asia and its stratigraphic distribution in China considerably.     

A new Xinjiangchelyid turtle (Testudines, Eucryptodira) from the Jurassic Qigu Formation of the southern Junggar Basin, Xinjiang, north-west China

A new eucryptodiran turtle, Xinjiangchelys qiguensis sp. nov. from the Upper Jurassic (Oxfordian — ?Kimmeridgian) Qigu Formation of the southern Junggar Basin (north‐west China) is described. The type material consists of a partial skeleton, including the complete carapace, plastron, nearly all cervical vertebrae, both scapulae, the pelvis and one ulna. It is clearly identifiable as a basal eucryptodire since it lacks the mesoplastron. It is distinguished from other species of Xinjiangchelys by several autapomorphies of the carapace and plastron, such as the first and fifth vertebrals extending on the peripherals, the plastron with three pairs of gulars, and an intergular which does not contact the hyoplastron. In the postcranium, the scapula with a long acromial and a small scapular process, the pelvis with a short ilial shaft and the elongated cervical vertebrae are characteristic. A new phylogenetic analysis of the in‐group phylogeny of the Xinjiangchelyidae is proposed and discussed, resulting in a new classification of the family. Xinjiangchelys (Toxocheloides) narynensis is regarded as a nomen dubium. Shartegemys is referred to Xinjiangchelys, whereas the holotypes of ‘Plesiochelys’chungkingensis and ‘P’. latimarginalis are excluded from the genus Xinjiangchelys but included in the Xinjiangchelyidae.