Bird-Like Anatomy,Posture, and Behavior Revealed by an Early Jurassic Theropod Dinosaur Resting Trace

Background

Fossil tracks made by non-avian theropod dinosaurs commonly reflect the habitual bipedal stance retained in living birds. Only rarely-captured behaviors, such as crouching, might create impressions made by the hands. Such tracks provide valuable information concerning the often poorly understood functional morphology of the early theropod forelimb.

Methodology/Principal Findings

Here we describe a well-preserved theropod trackway in a Lower Jurassic (∼198 million-year-old) lacustrine beach sandstone in the Whitmore Point Member of the Moenave Formation in southwestern Utah. The trackway consists of prints of typical morphology, intermittent tail drags and, unusually, traces made by the animal resting on the substrate in a posture very similar to modern birds. The resting trace includes symmetrical pes impressions and well-defined impressions made by both hands, the tail, and the ischial callosity.

Conclusions/Significance

The manus impressions corroborate that early theropods, like later birds, held their palms facing medially, in contrast to manus prints previously attributed to theropods that have forward-pointing digits. Both the symmetrical resting posture and the medially-facing palms therefore evolved by the Early Jurassic, much earlier in the theropod lineage than previously recognized, and may characterize all theropods.     

Gigantism among Late Jurassic limulids: New ichnological evidence from the Causses Basin (Lozère,France) and comments on body-size evolution among horseshoe crabs

An abundant ichnological material composed of xiphosuran trackways and isolated traces was discovered in Upper Jurassic limestones from the Causses Basin (Causse Méjean, Lozère, France). The morphology of the imprints supports their identification as Kouphichnium isp. In contrast to the most frequent case, the trackways are composed of omnipresent pusher imprints sometime associated with leg traces, but with no telson mark. We argue that this pattern reflects actual surface traces rather than an incomplete set of undertracks. The size distribution of the sampled ichnites is broadly bimodal. This is best explained by sexual dimorphism, a phenomenon frequently observed in modern xiphosurans. Analysis of the trace fossils further suggests that several growth stages are recorded and that the horseshoe crabs were walking in a protected and flat environment like a lagoon. This area, certainly close to a mating ground, was occasionally affected by a continental influence. The biometric study of the tracks suggests a gigantic size for the trackmakers whose body may have reached 84 cm in length. This discovery complements the few reports on other gigantic horseshoe crabs in the Jurassic of Western Europe, thus casting doubt on the postulated increase in body size from the Palaeozoic to the Recent. Furthermore, a literature review shows that there are still major gaps in the record of limulid body-fossils and tracks. Thus, neither of these archives can be taken at face value for quantifying the body-size evolution of horseshoe crabs.     

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.     

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.     

A tetrapod footprint assemblage with possible swim traces from the Jurassic–Cretaceous boundary,Anning Formation,Konglongshan, Yunnan,China

A small tetrapod footprint assemblage from the Anning Formation (Upper Jurassic–Lower Cretaceous) at Konglongshan Town, Yunnan Province, China, contains possible swim traces attributable to theropod dinosaurs based on their tridactyl and mesaxonic pes morphology. Morphotypes are tentatively assigned to the ichnogenera Characichnos and Wintonopus, a third one is similar to Hatcherichnus. Due to the inherent variability observed in tetrapod swim tracks, the names are used here informally describing footprints that reflect a distinct trackmaker behavior rather than anatomically accurate images of the pes anatomy. Variation of the imprint shape is obviously due to extramorphological effects and does not indicate taxonomic diversity of trackmakers. Elongate, slender impressions associated with these tracks are discussed here as possible tail traces. Trackmakers were possibly buoyant and active swimming individuals touching and scratching the bottom of deeper waterbodies with the distal ends of their digits. The orientation of the traces perpendicular to preserved ripples suggest cross-current movement and activities.     

The oldest record of a boring foraminifer: Early Permian of New Mexico,USA

The bioeroding foraminifer Troglotella incrustans Wernli and Fookes (Bolletino della Societa Paleontologica Italiana 31, 1992, 95), is widely reported from Bajocian?, and Oxfordian to Lower Cenomanian (with a Late Jurassic acme) shallow‐water limestones of the Tethyan realm. A single specimen of a boring foraminifer, assigned to T. incrustans, has now been observed from the Lower Permian (Sakmarian) Community Pit Formation of the Doña Ana Mountains, New Mexico, USA. Surviving the end‐Permian mass extinction, T. incrustans might be a Lazarus taxon that persisted in refuges. This finding represents the oldest record of a foraminifer exhibiting an euendolithic way of life. Boring foraminifera have not been previously recorded from strata older than the Jurassic. Boring traces of potentially foraminiferan origin, however, have been already reported from the Lower Carboniferous (?Ordovician).     

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.     

Late Jurassic footprints reveal walking kinematics of theropod dinosaurs

Avanzini, M., Piñuela, L. & García‐Ramos, J.C. 2011: Late Jurassic footprints reveal walking kinematics of theropod dinosaurs. Lethaia, Vol. 45, pp. 238–252. This study describes a set of theropod footprints collected from the Late Jurassic Lastres Formation (Asturias, N Spain). The footprints are natural casts (tracks and undertracks) grouped into three morphotypes, which are characterized by different size frequency, L/W relationship and divarication angles: ‘Grallatorid’ morphotype, ‘Kayentapus–Magnoavipes’ morphotype, ‘Hispanosauropus’ morphotype. The tracks were produced in firm, stiff and soft sediments. The infills of deep tracks, which are typically formed in soft mud, lack fine anatomical details, but they can reveal the walk kinematics of the trackmaker through the morphology of internal track fills and sinking traces. In all footprints, a horizontal outwardly directed translation movement and rotation are recognizable. The amount and geometry of digit penetration in the ground also show a pronounced difference. It can be inferred from the described sample that different theropoda‐related ichnogenera share common kinematics. □Asturias, dinosaur footprint, late jurassic, theropods, walking kinematics.     

Large theropod trackway from the Lower Jurassic Zhenzhuchong Formation of Weiyuan County,Sichuan Province,China: Review,new observations and special preservation

The well-preserved theropod track Weiyuanpus zigongensis, recently assigned to Eubrontes zigongensis, was not described in exhaustive detail at the time of its original discovery in 2007. Among the morphological details not described was an antero-medially directed hallux seen in five of the six tracks that make up the type trackway. Hallux traces are only rarely reported in large Lower Jurassic theropod tracks such as Eubrontes and Gigandipus, and their presence or absence may be the result of one or both of two factors: track depth and/or differences in hallux configuration in the trackmakers. Here we argue that E. zigongensis is one of the best preserved examples of a eubrontid track, which can be morphologically distinguished from other Eubrontes ichnospecies by the presence of well-defined hallux traces.     

THE FISH AND AMPHIBIAN SWIMMING TRACES UNDICHNA AND LUNICHNIUM, WITH EXAMPLES FROM THE LOWER PERMIAN OF NEW MEXICO, USA

Abstract: The ichnotaxonomy and stratigraphic, geographic and environmental distribution of fish (Undichna) and amphibian (Lunichnium) swimming traces are reviewed. The ichnospecies of Undichna consist of various combinations of sinusoidal waves of differing complexity. Some of the more complex ichnospecies are made up of elements of the simpler forms, and morphological subset relationships between them are presented. Such subset series represent potential taphoseries relationships (i.e. preservational variants that reflect, for example, undertrails), or series of minor behavioural variations. Such a system can be used to highlight that different ichnospecies occurring at a locality may be taphonomic or minor behavioural variants of each other. Caution should, therefore, be exercised before erecting new ichnospecies on the basis of limited material if its morphology is a subset of an existing ichnospecies. However, the naming of such simpler ichnospecies is valid if they represent a recurrent morphology, and it is valid to erect new ichnospecies whose morphology is not a subset of an existing ichnospecies. Specimens that demonstrate intergradation between ichnotaxa can be used to justify their synonomy. Ichnotaxonomic revisions reduce the number of ichnospecies in Undichna from 14 to nine. U. radnicensis, a highly variable ichnospecies, is synonymized with U. britannica on the basis of material from China that demonstrates they can intergrade. U. prava is a partial U. tricosta, which falls within the minimum diagnosis of U. simplicitas. U. gosiutensis is regarded as a subjective junior synonym of U. quina. U. westerbergensis, originally attributed to a ‘crossopterygian’ fish performing a tetrapod‐like gait, is reassigned as a distinct ichnospecies within Lunichnium because it demonstrates the same morphology, representing similar behaviour, albeit by a different producer. L. anceps and L. gracile are synonymized with L. rotterodium. New specimens of U. bina and L. rotterodium are also described from the Lower Permian Robledo Mountains Formation of southern New Mexico, USA.     

A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal

Mateus, O. & Milàn, J. 2009: A diverse Upper Jurassic dinosaur ichnofauna from central‐west Portugal. Lethaia, Vol. 43, pp. 245–257. A newly discovered dinosaur track‐assemblage from the Upper Jurassic Lourinhã Formation (Lusitanian Basin, central‐west Portugal), comprises medium‐ to large‐sized sauropod tracks with well‐preserved impressions of soft tissue anatomy, stegosaur tracks and tracks from medium‐ to large‐sized theropods. The 400‐m‐thick Lourinhã Formation consists of mostly aluvial sediments, deposited during the early rifting of the Atlantic Ocean in the Kimmeridgian and Tithonian. The stratigraphic succession shows several shifts between flood‐plain mud and fluvial sands that favour preservation and fossilization of tracks. The studied track‐assemblage is found preserved as natural casts on the underside of a thin bivalve‐rich carbonate bed near the Tithonian–Kimmeridgian boundary. The diversity of the tracks from the new track assemblage is compared with similar faunas from the Upper Jurassic of Asturias, Spain and the Middle Jurassic Yorkshire Coast of England. The Portuguese record of Upper Jurassic dinosaur body fossils show close similarity to the track fauna from the Lourinhã Formation. □Dinosaur tracks, Lusitanian Basin, Portugal, skin impressions, Upper Jurassic.     

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.”     

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.     

On taxonomy and distribution of fossil Cerophytidae (Coleoptera: Elateriformia) with description of a new Mesozoic species of Necromera Martynov 1926

In this paper the data on fossils of the family Cerophytidae are reviewed. New synonymy on generic names Necromera Martynov 1926 (proposed in composition of the family Oedemeridae); Idiomerus Dolin in Dolin et al. 1980, n. syn. (proposed in composition of Elateridae) and Leptocnemus Hong & Wang 1990, n. syn. (proposed without any family attribution) is established. New materials on this family from Mesozoic deposits of Asia are cited. As a result, it was established that this spreads in deposits of both Upper Jurassic and Lower Cretaceous. Mercata festira Lin 1986 described from Lower Jurassic as member of Silphidae and Abrotus reconditus Dolin in Dolin et al. 1980 described from Upper Jurassic as a member of Elateridae are transferred to Cerophytidae. Diagnoses of the genus Necromera and family Cerophytidae in compression fossils are elaborated. Necromera admiranda n. sp. is described from the Upper Jurassic-Lower Cretaceous in Liaoning, China. The historical development of the family from the Lower Jurassic is discussed.     

Asianopodus-type footprints from the Hekou Group of Honggu District,Lanzhou City,Gansu, China and the “heel” of large theropod tracks

An increasing number of dinosaur tracksites have been reported from the Cretaceous Hekou Group deposits of the Lanzhou-Minhe Basin in the Gansu Province region. These include small sites such as the Huazhuang tracksite, from the Honggu District reported here, the Zhongpu tracksite with multiple track levels but few well-preserved tracks, other small tracksites currently under investigation, and the large and diverse Liujixia National Dinosaur Geopark site at Yanguoxia, where intensive study is ongoing. Collectively these sites reveal that ichnofaunas in the Hekou Group are widespread and diverse. The Huazhuang tracksite yields a small assemblage of moderately well-preserved theropod tracks assigned to Asianopodus. This is the first report of Asianopodus from the Hekou Group. Huazhuang Asianopodus belongs to the Eubrontes morphotype. The large theropod tracks from Lanzhou-Minhe Basin were left by large theropod trackmakers with the same general foot morphology. The specimens are described in detail and compared with other theropod track morphotypes from the Lower Cretaceous of China and elsewhere. In general, although the metatarsophalangeal pads of some Jurassic Eubrontes-type tracks are aligned with the axis of digit III, this feature appears most common in the Early Cretaceous theropod (Eubrontes-type) tracks.     

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.     

The Trace Fossil Diplopodichnus from the Lower Jurassic Lacustrine Sediments of Central China and the Isopod Armadillidium vulgare (Pillbug) Lebensspuren as its Recent Analogue

The trace fossil Diplopodichnus isp. was identified in lacustrine siliciclastics of the Lower Jurassic Anyao Formation in the Henan Province, central China. This is the youngest occurrence of this ichnogenus. It is similar to some variants of recent surface traces of the isopod Armadillidium vulgare (pillbug) crawling on soft mud in temporal puddles. Therefore, isopods are suggested to be producers of the described Diplopodichnus isp. It seems that isopods should be taken into account as important component of deep lacustrine ecosystems by the Early Jurassic.     

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.     

The Fish Swimming Trace Undichna from the Mississippian Mauch Chunk Formation,Eastern Pennsylvania

Undichna, the swimming trace of fish, was recovered from the middle member of the Mauch Chunk Formation in eastern Pennsylvania in strata of Late Mississippian (Visean) age. These traces represent the only evidence of fish known from the Mauch Chunk Formation and the Carboniferous of Eastern Pennsylvania. The Mauch Chunk Formation in the study area is characterized by an ephemeral fluvial depositional environment that yields a diverse invertebrate ichnoassemblage of the Scoyenia ichnofacies and a tetrapod footprint assemblage dominated by the tracks of temnospondyl amphibians. The Undichna specimens are preserved on fine-grained, mudstone-draped, rippled sandstone. Two specimens, assigned to U. Britannica, consist of pairs of well-defined, narrowly incised, sinusoidal wave traces that are out-of-phase. Two specimens, assigned to U. quina, consist of two pairs of in-phase sinusoidal waves intersected by an additional single wave with a greater amplitude. Two specimens, each composed of a single wave, are assigned to U. unisulca.

The slabs of rock with the Undichna traces also preserve tetrapod undertracks assigned to Batrachichnus, Matthewichnus, and Hylopus. This trace fossil association indicates the presence of small fish, some with caudal and/or anal fins and some with pectoral, pelvic, and anal and/or caudal fins, that touched the sediment during periods of intermittent flooding of subaerial surfaces walked upon by early tetrapods. Examples of Undichna from Mississippian or older deposits, especially from a fluvial paleoenvironment, are rare; few have been reported from the Devonian and the Lower Carboniferous.