Theropod Research Articles

Evolution of vision and hearing modalities in theropod dinosaurs.

Owls and nightbirds are nocturnal hunters of active prey that combine visual and hearing adaptations to overcome limits on sensory performance in low light. Such sensory innovations are unknown in nonavialan theropod dinosaurs and are poorly characterized on the line that leads to birds. We investigate morphofunctional proxies of vision and hearing in living and extinct theropods and demonstrate deep evolutionary divergences of sensory modalities. Nocturnal predation evolved early in the nonavialan lineage Alvarezsauroidea, signaled by extreme low-light vision and increases in hearing sensitivity. The Late Cretaceous alvarezsauroid Shuvuuia deserti had even further specialized hearing acuity, rivaling that of today's barn owl. This combination of sensory adaptations evolved independently in dinosaurs long before the modern bird radiation and provides a notable example of convergence between dinosaurs and mammals.

The Role of the Intramandibular Joint, Symphyseal Tissues, and Wrapping Muscles on Theropod Dinosaur Mandibular Function

Sauropsids, unlike mammals, possess an intramandibular joint (IMJ) separating the dentary and postdentary bones, with different lineages either rigidifying (turtles & crocodilians) or maintaining compliance (lepidosaurs & birds) about the joint. IMJ construction and its role on mandibular performance is unclear, impeding our understanding of its function in extinct animals with extreme feeding behaviors like Tyrannosaurus rex. Sauropsids like T. rex pose a particular biomechanical paradox; feeding traces, coprolites, and their robust crania and teeth indicate that they regularly bit through and ingested bone, but their dorsoventrally-tall and mediolaterally-thin hemimandibles, IMJ, and patent mandibular symphyses suggest their mandibles were ill-suited for bone-crushing bites. Extant sauropsids also exhibit wrapping intramandibularis (mIRA) and pterygoideus ventralis (mPTv) muscles, whose effect on mandibular performance and interaction with the IMJ are unknown. Here we model the effect of the IMJ, symphyseal tissue properties, and wrapping muscle orientation on T. rex mandible biomechanical performance in order to give insight into the biomechanical constraints faced by sauropsids, and the link between their mandibular form and function. Joint tissue histology, gross dissection, and iodine contrast data of extant sauropsids were performed to inform model tissue properties, IMJ construction and muscle orientation. We imported a 3D STL model of the T. rex “STAN” and modeled the IMJ by differing material properties of modeled joint tissues to test the influence of linkage materials on mandibular performance. Muscles were mapped onto the model, and their corresponding muscle moments calculated from area centroids, muscle architecture, and volumes. Wrapping mIRA and mPTv muscles were simulated by directing their force vectors to appropriately positioned dummy points using extant sauropsids as a guide. The mandible was then constrained at the jaw joint and in a series of points along the tooth row from rostral to caudal, simulating both unilateral and bilateral bites. We find that while strains are quite high (>6000 µstrains) about the intramandibular joint, mediolateral bending stresses and strain are markedly reduced by the prearticular, which exhibits greater strain than the surrounding bone. Our results suggest the prearticular of T. rex acted as a strain sink to counteract bending about the IMJ and thereby rigidifying the mandible during feeding. We hypothesize that differences in IMJ articulation and prearticular construction may differentially facilitate or impede prearticular streptognathy, and thus intramandibular kinesis about the IMJ, in sauropsids. Forthcoming models will simulate other theropods within and without Tyrannosauridae to test this hypothesis.

Darting towards Storm Shelter: a minute dinosaur trackway from southern Africa

Theropod dinosaurs are considered the main terrestrial carnivores in the Jurassic and Cretaceous. Their rise to dominance has been linked to, among others, body size changes in their early history, especially across the Triassic–Jurassic boundary. However, to qualitatively assess such temporal trends, robust skeletal and trace fossil data sets are needed globally. The richly fossiliferous southern African continental rock record in the main Karoo Basin offers an unparalleled perspective for such investigations. Herein, by documenting a newly discovered Early Jurassic trackway of very small, functionally tridactyl tracks near Storm Shelter (Eastern Cape) in South Africa, the track record can be expanded. Based on ichnological measurements at the ichnosite and digital 3D models, the footprint dimensions (length, width, splay), locomotor parameters (step length, stride, speed), and body size estimates of the trackmaker are presented. In comparison to other similar tracks, these footprints are not only the smallest Grallator-like tracks in the Clarens Formation, but also the most elongated dinosaur footprints in southern Africa to date. The tracks also show that the small-bodied bipedal trackmaker dashed across the wet sediment surface at an estimated running speed of ~12.5 km/h. During the dash, either as a predator or as a prey, the trackmaker’s small feet sunk hallux-deep into the sediment. The tracking surface is overgrown by fossilised microbial mats, which likely enhanced the footprint preservation. Based on track morphometrics and the regional dinosaur skeletal record, the trackmakers are attributed to Megapnosaurus rhodesiensis (formerly Syntarsus rhodesiensis), a small-to-medium-sized, early theropod common in southern Africa.Significance: A newly discovered Early Jurassic theropod trackway in South Africa contains not only the smallest tracks in the Clarens Formation, but also the most elongated dinosaur footprints in southern Africa to date. The tracks show that the small bipedal trackmaker dashed across the wet sediment surface at an estimated running speed of ~12.5 km per hour. During the run, the trackmaker’s feet sunk so deeply into the sediment that even the forwards-directed halluces were impressed. Open data set: https://doi.org/10.6084/m9.figshare.13007240.v1

Tail anatomy of the Alvarezsauria (Theropoda, Coelurosauria), and its functional and behavioural implications

The anatomy of the alvarezsaurian tail has received relatively little attention in the paleontological literature, even though it shows a peculiar combination of anatomical characteristics that are unique among theropod dinosaurs. Nearly complete, informative tails are known from early-branching, intermediate, and late-branching taxa, allowing for robust inferences about their evolution. The alvarezsaurian tail is notable in being the longest among maniraptoran theropods, both in number of caudal vertebrae and proportional length. We examined the comparative anatomy and myology of the tail, and performed a cladistic analysis on a tail-character-based data matrix that provided a general framework for reconstructing alvarezsaurian tail evolution. Our results show that caudal vertebrae of alvarezsaurians have a combination of derived osteological features, intervertebral joint morphology, and inferred musculature, which together suggest that the tail possessed a unique function among theropods. We interpret these features as indicators of an exceptional capacity to change rotational inertia. The form and function of the tail, in combination with the fossorial forelimb, suggests that alvarezsaurians had an ecological niche similar to today's aardvark, pangolins and anteaters.

A detailed osteological description of Xenotarsosaurus bonapartei (Theropoda: Abelisauridae): implications for abelisauroid phylogeny

Xenotarsosaurus bonapartei was the third abelisaurid theropod dinosaur to be named from Argentina. The holotype comprises two partial anterior dorsal vertebrae and a nearly complete right hind limb from the Upper Cretaceous (lower Cenomanian–upper Turonian) Bajo Barreal Formation, central Patagonia, Argentina. The materials display morphological features that undoubtedly position Xenotarsosaurus within Abelisauroidea. Moreover, detailed comparisons with members of that theropod group confirm the close relationship of this taxon to abelisaurids. Here we provide an emended diagnosis of Xenotarsosaurus bonapartei that includes five newly recognized autapomorphies: (1) anterior dorsal vertebrae with large, strongly dorsoventrally developed parapophyses; (2) anterior dorsal vertebrae with well-developed centroprezygapophyseal fossae that are taller dorsoventrally than wide mediolaterally; (3) fibular condyle of femur triangular in shape and projecting posteriorly; (4) well-marked groove on anterolateral corner of proximal fibula; and (5) iliofibularis tubercle of fibula distally interrupted by hook-like concavity. To determine its systematic position within Abelisauroidea, we incorporated Xenotarsosaurus into a phylogenetic analysis, recovering this theropod as a non-carnotaurine abelisaurid more derived than Eoabelisaurus mefi. Xenotarsosaurus displays several plesiomorphic traits when compared with penecontemporaneous abelisaurids from the Neuquén Group. Similarly, other non-avian dinosaur taxa from the Bajo Barreal Formation are frequently postulated as more phylogenetically basal than coeval forms from northern Patagonia. This scenario suggests the potential existence of provincialism in early Late Cretaceous continental vertebrate faunas of southern South America. The present study increases knowledge of abelisaurid systematics, evolution, and paleobiogeography and augments our understanding of the Upper Cretaceous dinosaur assemblage of central Patagonia.

The evolution of pelvic limb muscle moment arms in bird-line archosaurs.

Bipedal locomotion evolved along the archosaurian lineage to birds, shifting from "hip-based" to "knee-based" mechanisms. However, the roles of individual muscles in these changes and their evolutionary timings remain obscure. Using 13 three-dimensional musculoskeletal models of the hindlimbs of bird-line archosaurs, we quantify how the moment arms (i.e., leverages) of 35 locomotor muscles evolved. Our results support two hypotheses: From early theropod dinosaurs to birds, knee flexors' moment arms decreased relative to knee extensors', and medial long-axis rotator moment arms for the hip increased (trading off with decreased hip abductor moment arms). Our results reveal how, from the Triassic Period, bipedal theropod dinosaurs gradually modified their hindlimb form and function, shifting more from hip-based to knee-based locomotion and hip-abductor to hip-rotator balancing mechanisms inherited by birds. Yet, we also discover unexpected ancestral specializations in larger Jurassic theropods, lost later in the bird-line, complicating the paradigm of gradual transformation.

Maniraptoran pelvic musculature highlights evolutionary patterns in theropod locomotion on the line to birds.

Locomotion is a fundamental aspect of palaeobiology and often investigated by comparing osteological structures and proportions. Previous studies document a stepwise accumulation of avian-like features in theropod dinosaurs that accelerates in the clade Maniraptora. However, the soft tissues that influenced the skeleton offer another perspective on locomotory adaptations. Examination of the pelvis for osteological correlates of hind limb and tail musculature allowed reconstruction of primary locomotory muscles across theropods and their closest extant relatives. Additionally, the areas of pelvic muscle origins were quantified to measure relative differences within and between taxa, to compare morphological features associated with cursoriality, and offer insight into the evolution of locomotor modules. Locomotory inferences based on myology often corroborate those based on osteology, although they occasionally conflict and indicate greater complexity than previously appreciated. Maniraptoran pelvic musculature underscores previous studies noting the multifaceted nature of cursoriality and suggests that a more punctuated step in caudal decoupling occurred at or near the base of Maniraptora.

A fast-growing basal troodontid (Dinosauria: Theropoda) from the latest Cretaceous of Europe

A characteristic fauna of dinosaurs and other vertebrates inhabited the end-Cretaceous European archipelago, some of which were dwarves or had other unusual features likely related to their insular habitats. Little is known, however, about the contemporary theropod dinosaurs, as they are represented mostly by teeth or other fragmentary fossils. A new isolated theropod metatarsal II, from the latest Maastrichtian of Spain (within 200,000 years of the mass extinction) may represent a jinfengopterygine troodontid, the first reported from Europe. Comparisons with other theropods and phylogenetic analyses reveal an autapomorphic foramen that distinguishes it from all other troodontids, supporting its identification as a new genus and species, Tamarro insperatus. Bone histology shows that it was an actively growing subadult when it died but may have had a growth pattern in which it grew rapidly in early ontogeny and attained a subadult size quickly. We hypothesize that it could have migrated from Asia to reach the Ibero-Armorican island no later than Cenomanian or during the Maastrichtian dispersal events.

The influence of juvenile dinosaurs on community structure and diversity

Despite dominating biodiversity in the Mesozoic, dinosaurs were not speciose. Oviparity constrained even gigantic dinosaurs to less than 15 kg at birth; growth through multiple morphologies led to the consumption of different resources at each stage. Such disparity between neonates and adults could have influenced the structure and diversity of dinosaur communities. Here, we quantified this effect for 43 communities across 136 million years and seven continents. We found that megatheropods (more than 1000 kg) such as tyrannosaurs had specific effects on dinosaur community structure. Although herbivores spanned the body size range, communities with megatheropods lacked carnivores weighing 100 to 1000 kg. We demonstrate that juvenile megatheropods likely filled the mesocarnivore niche, resulting in reduced overall taxonomic diversity. The consistency of this pattern suggests that ontogenetic niche shift was an important factor in generating dinosaur community structure and diversity.

Not enough room

Paleontology Modern carnivore communities include species that span a range of body sizes. For example, on the African savannah, there are small species (mongooses), medium species (wild dogs), and large species (lions). This variation reflects available prey sources that best suit each group. Carnivorous dinosaur communities, however, were missing species that fall into the middle, or mesocarnivore, group as adults. Schroeder et al. looked across communities, space, and time and found that this absence appears to have been driven by the distinctive biology of dinosaurs, in which giant adults start out as tiny hatchlings. Growing juvenile dinosaurs thus filled the other niches and limited trophic species diversity. Science , this issue p. [941][1] [1]: /lookup/doi/10.1126/science.abd9220

Dinosaur remains from the Upper Cretaceous (Campanian) of the Western Desert, Egypt

Upper Cretaceous dinosaur remains from Afro-Arabia are rare and mainly restricted to pre-Turonian horizons. Consequently, the discovery of new fossils from Upper Cretaceous deposits in the Western Desert, central Egypt is significant because it adds to the meager record of dinosaurs described from this landmass. The oases of Kharga and Dakhla, Western Desert, Egypt, expose the Quseir Formation (Campanian), with the titanosaurian sauropod Mansourasaurus shahinae as the only currently named dinosaur from strata of this age in the entire region. Numerous (~80) other non-avian dinosaur fossils have also been collected from the Quseir Formation, including material that can be referred to Somphospondyli, Titanosauriformes/Titanosauria, and non-avian Theropoda. Among the discoveries from the Dakhla Oasis are the proximal ends of an associated sauropod tibia and fibula and several sauropod caudal vertebrae representing both sub-adult and adult individuals. Middle caudal vertebrae can be referred to Titanosauria on the basis of procoelous centra and anteriorly displaced neural arches. One caudal vertebra even exhibits pneumatic internal structures rarely observed outside of the Upper Cretaceous South American saltasaurines. Dinosaur fossils recovered from the Kharga Oasis include a partial femur and partial cervical vertebra of a sauropod dinosaur and an isolated proximal fibula of a non-avian theropod dinosaur. Taken together, these findings indicate that Late Cretaceous North African ecosystems supported a diversity of non-avian dinosaurs, some demonstrating affinities with South American forms and others (e.g., Mansourasaurus) with Eurasian groups. Additional exploration of the uppermost Cretaceous units in southern Egypt offers promise for the discovery of important fossils to better characterize Afro-Arabian biotas generally while also providing important perspectives on non-avian dinosaur faunas near the close of the Mesozoic Era.

EL MOVIMIENTO DE UN DINOSAURIO DEDUCIDO DE UNA RASTRILLADA TERÓPODA CON ESTRUCTURAS DE INMERSIÓN DE LOS PIES EN BARRO Y DE ARRASTRE DE COLA (FORMACIÓN VILLAR DEL ARZOBISPO. GALVE, TERUEL, ESPAÑA)

En El Cantalar (Galve, Teruel) dentro de la Formación Villar del Arzobispo (Jurásico Superior-Cretácico Inferior), se conocían dos rastrilladas: una de un enorme cocodrilo y otra de un terópodo de andar semiplantígrado. El lavado y erosión del afloramiento ha limpiado la parte conocida y, además, ha puesto al descubierto varias pisadas que añadir al rastro terópodo. En la revisión del yacimiento, se han encontrado, además de las nuevas huellas, estructuras que modifican la primera interpretación del rastro semiplantígrado dada la información adicional que suministran. Las observaciones han permitido: por una parte reconstruir el proceso de formación de huellas en suelo muy blando; por otra, describir un nuevo ejemplo de rastro de cola de dinosaurio, el primero descubierto en Aragón, y, finalmente, aplicar los datos de la rastrillada coordinando el movimiento de balanceo lateral y vertical del cuerpo del animal, con las fases de la pisada para cada autópodo durante su desplazamiento.

The patterns and modes of the evolution of disparity in Mesozoic birds.

The origin of birds from non-avian theropod dinosaurs is one of the greatest transitions in evolution. Shortly after diverging from other theropods in the Late Jurassic, Mesozoic birds diversified into two major clades-the Enantiornithes and Ornithuromorpha-acquiring many features previously considered unique to the crown group along the way. Here, we present a comparative phylogenetic study of the patterns and modes of Mesozoic bird skeletal morphology and limb proportions. Our results show that the major Mesozoic avian groups are distinctive in discrete character space, but constrained in a morphospace defined by limb proportions. The Enantiornithines, despite being the most speciose group of Mesozoic birds, are much less morphologically disparate than their sister clade, the Ornithuromorpha-the clade that gave rise to living birds, showing disparity and diversity were decoupled in avian history. This relatively low disparity suggests that diversification of enantiornithines was characterized in exhausting fine morphologies, whereas ornithuromorphs continuously explored a broader array of morphologies and ecological opportunities. We suggest this clade-specific evolutionary versatility contributed to their sole survival of the end-Cretaceous mass extinction.

BITE MARKS ON AN AETOSAUR (ARCHOSAURIA, SUCHIA) OSTEODERM: ASSESSING LATE TRIASSIC PREDATOR-PREY ECOLOGY THROUGH ICHNOLOGY AND TOOTH MORPHOLOGY

ABSTRACTTrace fossils such as bite marks provide rare, direct evidence of animal behavior, including predator-prey interactions. We present an osteoderm of the aetosaur Typothorax coccinarum from the Late Triassic Chinle Formation of Arizona with several punctures and scores, interpreted here as bite marks, preserved as evidence of predation/scavenging by a large carnivore. The marks include a single bite producing four subparallel fusiform pits on the ventral surface and several additional marks, including striated scores, on the dorsal surface. These traces are described and compared with known contemporaneous carnivorous taxa to determine the source of the bite marks. Some Triassic carnivores, including theropod dinosaurs can be ruled out because of tooth shape and serration densities. Phytosaurs and large paracrocodylomorphs remain as likely candidates based on tooth morphology. Although some phytosaur teeth are too rounded to produce the marks seen in this specimen, we demonstrate that the more lingually flattened teeth typically found in the posterior section of the snout are sufficiently mediolaterally compressed to produce a fusiform pit. A protective function for aetosaur osteoderms cannot be confirmed presently, but the extensive carapace these bones formed would have been a major barrier to both scavengers and active predators and may preserve more feeding/predation traces than previously thought. The bite marks described herein support the hypothesis that aetosaurs were prey items of large archosauromorphs, expanding our understanding of the complex, and seemingly carnivore dominated Late Triassic terrestrial ecosystems of North America.

Late Cretaceous Bauru Group biota from Southern Goiás state, Brazil: history and fossil content

The southern Goiás state region of Central Brazil has an extensive and informative record of fossil biota. Particularly over the last five years, there has been a great increase in fossil finds, which has enabled a greater understanding of this region's fauna and flora during the Late Cretaceous. In this article, we provide an updated synthesis of the biota from the Cretaceous of the southern Goiás state, the record of plants, gastropods, turtles, crocodilians, titanosaurs, and theropod dinosaurs.

A non-averostran neotheropod vertebra (Dinosauria: Theropoda) from the earliest Jurassic Whitmore Point Member (Moenave Formation) in southwestern Utah

ABSTRACT Theropod dinosaurs are minor components of Late Triassic ecosystems in North America, comprising coelophysoids and various non-neotheropods from the Chinle Formation of Arizona, Utah, Colorado, and New Mexico and the Dockum Group of western Texas. By the Sinemurian (Early Jurassic), the coelophysoid “Syntarsus” kayentakatae and the large-bodied non-averostran neotheropod Dilophosaurus wetherilli from the Kayenta Formation were the dominant terrestrial predators. Theropods are virtually unknown from the intervening Rhaetian–Hettangian Moenave Formation, with the exception of two partial coelophysoid pelves from somewhere within the Dinosaur Canyon Member, which includes the Triassic–Jurassic boundary and end-Triassic mass extinction. Here we describe an anterior trunk vertebra from a non-coelophysoid, non-averostran neotheropod from the uppermost Whitmore Point Member of the Moenave Formation in southwestern Utah, which is Hettangian in age. The vertebra has prominent vertebral laminae and associated pneumatic fossae, and anterior and posterior ‘shoulders’ on the neural spine that are similar to those found in Dilophosaurus wetherilli. This vertebra belongs to a theropod that may be as many as 15 million years older than Dilophosaurus wetherilli from the middle of the Kayenta Formation in Arizona. This theropod is associated with Grallator, Eubrontes, and Characichnos theropod traces made on the shores of the Early Jurassic Lake Whitmore that are abundant in the Whitmore Point Member in southwestern Utah. Its occurrence in the Hettangian roughly coincides with the appearance of Eubrontes tracks in North America, indicating that not all contemporaneous theropod traces were made by coelophysoids.

An Upper Triassic terrestrial vertebrate assemblage from the forgotten Kocury locality (Poland) with a new aetosaur taxon

ABSTRACT Since 1990, several localities within the Keuper (upper Middle to Upper Triassic) strata in southern Poland have yielded remains of numerous terrestrial vertebrate species. Here we report a new Upper Triassic vertebrate assemblage from the rediscovered Kocury locality. An incomplete theropod dinosaur fibula named Velocipes guerichi described in 1932 was found there. The site was then forgotten and not explored until our excavations began in 2012, that yielded material of a lungfish, a proterochersid turtle, and a new typothoracin aetosaur Kocurypelta silvestris gen. et sp. nov. The new taxon is characterized by autapomorphies of the maxilla: an elongated edentulous posterior portion longer than 80% of the posterior maxillary process, a short medial shelf restricted to the posterior portion of the bone, an anteriorly unroofed maxillary accessory cavity, and lack of a distinct groove for choanal recess on the anteromedial surface of the bone. These new finds improve our knowledge on the vertebrate diversity of the Germanic Basin in the Late Triassic, evidencing the presence of yet unrecognized taxa. Additionally, the partial cranial aetosaur material emphasizes the issues with the aetosaurian taxonomy that is focused mostly on the osteoderm morphology.

Diverse preserved dinosaur footprint assemblage from Jurassic–Cretaceous transition eolian dune deposits of western Shandong Province, China

Ancient desert, characterized by low biotic diversity as well as low preservation potential, has long been assumed as devoid of biotic activity. However, recent works from Mesozoic trace fossils preserved in the eolianites in North America, South America and East Asia indicate that the ancient desert can also have diverse inhabitants. This paper presents a diversified preserved dinosaur track assemblage from the Jurassic-Cretaceous transition eolian dune deposits in western Shandong Province, China. Based on the ichnological analysis, tracks are assigned to ichnogenus Anomoepus, Eubrontes-like and Ornithopodichnus-like forms, representing the oldest and second example of the Chelichnus ichnofacies reported from China as well as Asia. Sedimentological analysis of the tracksites indicates that tracks occurred in linear dunes under arid climatic conditions. Well-preserved tracks with detailed anatomical information are more likely to be preserved in moist sands, contrary to the dry cohesionless sands. Anomoepus and Ornithopodichnus-like tracks cooccurring with northwestward winds were made during summer wet season, while Eubrontes-like track was present during winter dry season characterized by strong northeastward wind, implying different habits between ornithopod and theropod dinosaurs. The preserved Jurassic-Cretaceous transition dinosaur tracks in eolian dune deposits not only expand the spatial-temporal distribution of dinosaur fauna in North China, but also greatly improve our understanding of the adaptive capacity of dinosaurs in severe living conditions, such as the desert environment.

The easternmost occurrence of Saurornitholestes from the Judith River Formation, Montana, indicates broad biogeographic distribution of Saurornitholestes in the Western Interior of North America

ABSTRACT Small-bodied dromaeosaurid and troodontid theropod dinosaurs are present in most of the terrestrial Late Cretaceous formations of the North American Western Interior, but are represented predominantly by shed teeth and more rarely by isolated skeletal elements and associated skeletons. The eudromaeosaurian dromaeosaurid Saurornitholestes is among the most commonly recovered small theropods from North America, known from abundant shed teeth, isolated cranial and postcranial elements, and several partial or near-complete skeletons from the Campanian aged Two Medicine Formation of Montana and the Dinosaur Park Formation of Alberta. Conversely, few Saurornitholestes specimens beyond shed teeth have been recovered from temporally equivalent units of the Judith River Formation of Montana. This study reports the first cranial material, a near-complete frontal, referable to Saurornitholestes from the Judith River Formation of Montana. This specimen is the easternmost occurrence of this taxon and was recovered approximately 400 km east of the localities which have produced referred Saurornitholestes specimens in the Two Medicine Formation, proximal to the Cordilleran thrust front. As such, this new specimen indicates that Saurornitholestes was biogeographically widespread and occupied both inland and coastal environments across the northern portions of the Late Cretaceous Western Interior of North America.

Epidermal complexity in the theropod dinosaur Juravenator from the Upper Jurassic of Germany

AbstractEpidermal scales among modern reptiles are morphologically diverse and serve a variety of functions ranging from moisture balance to chemoreception. Despite being predominantly squamous‐skinned (scaly), the functional implications of this type of integument have never before been explored for a dinosaur. Re‐examination of the holotype of the theropod Juravenator starki (Kimmeridgian, Late Jurassic) reveals previously unseen diversity in its epidermal covering. In addition to the presence of short monofilaments on the dorsal and ventral parts of the tail, we identify distinctive scale patterns corresponding to discrete longitudinal bands of scutate, tuberculate and ornamented scales along the length of the tail. In modern squamous‐skinned amniotes, variation in scale architecture can be linked to functional differences between scale types. In Juravenator, distinctive circular nodes associated with the ornamented scales have recently been interpreted as integumentary sense organs. We also suggest that it is likely that the snake‐like scutate scales on the underside of the tail fulfilled a role related to mechanical protection rather than locomotory function. Whereas the literature on theropod epidermal structures has been dominated by discussions on the evolution and diversity of feathers, this study highlights the complexity of squamous integument in Juravenator, which is revealed as one of the most complex among known theropods. An exploration of the epidermal structures found in Juravenator and other non‐avian theropods reveals great complexity in the type and distribution of these structures across the clade, suggesting potential for the future inquiry into their taxonomic and biological importance.