Small-bodied Theropods Research Articles

A review of small-bodied theropod dinosaurs from the Upper Cretaceous of India, with description of new cranial remains of a noasaurid (Theropoda: Abelisauria)

ABSTRACT Small-bodied theropod dinosaurs are rare on southern landmasses but have been known from India for a century. Excavations by Charles Matley and Durgansankar Bhattacharji in uppermost Cretaceous sediments at Bara Simla, central India in 1917–1919 recovered small theropod vertebral and limb elements originally interpreted as coelurosaurians and separated into at least three species (Compsosuchus solus, Laevisuchus indicus, Jubbulpuria tenuis) based on features that can now be attributed to their serial position in the vertebral column. The comparatively recent discoveries of Noasaurus leali and Masiakasaurus knopfleri from similar-aged rocks in South America and Madagascar, respectively, and advances in basal theropod systematics led to a revised interpretation of most small-bodied Indian theropods as noasaurid abelisauroids. Here we review and redescribe Laevisuchus, Jubbulpuria, and Compsosuchus, including several elements that until now were thought lost, and describe a new partial noasaurid dentary from central India. The dentary bears the characteristic procumbent dentition of Masiakasaurus, which apparently is absent in Noasaurus. Likewise, cervical vertebrae of Laevisuchus more closely resemble those of Masiakasaurus than those of Noasaurus. Despite these similarities, phylogenetic analyses indicate that the balance of character data supports the Indian noasaurid species outside the sister-taxon pairing of South American and Malagasy species. Bones of small-bodied theropods have been recovered exclusively from the youngest Mesozoic localities in India (e.g., Pisdura, Bara Simla); to date they have not been reported from the slightly older localities in western and central India, from southern Indian sites in the Cauvery Basin, nor from the Vitakri Formation of Pakistan.

The fast and the frugal: Divergent locomotory strategies drive limb lengthening in theropod dinosaurs

Limb length, cursoriality and speed have long been areas of significant interest in theropod paleobiology, since locomotory capacity, especially running ability, is critical in the pursuit of prey and to avoid becoming prey. The impact of allometry on running ability, and the limiting effect of large body size, are aspects that are traditionally overlooked. Since several different non-avian theropod lineages have each independently evolved body sizes greater than any known terrestrial carnivorous mammal, ~1000kg or more, the effect that such large mass has on movement ability and energetics is an area with significant implications for Mesozoic paleoecology. Here, using expansive datasets that incorporate several different metrics to estimate body size, limb length and running speed, we calculate the effects of allometry on running ability. We test traditional metrics used to evaluate cursoriality in non-avian theropods such as distal limb length, relative hindlimb length, and compare the energetic cost savings of relative hindlimb elongation between members of the Tyrannosauridae and more basal megacarnivores such as Allosauroidea or Ceratosauridae. We find that once the limiting effects of body size increase is incorporated there is no significant correlation to top speed between any of the commonly used metrics, including the newly suggested distal limb index (Tibia + Metatarsus/ Femur length). The data also shows a significant split between large and small bodied theropods in terms of maximizing running potential suggesting two distinct strategies for promoting limb elongation based on the organisms' size. For small and medium sized theropods increased leg length seems to correlate with a desire to increase top speed while amongst larger taxa it corresponds more closely to energetic efficiency and reducing foraging costs. We also find, using 3D volumetric mass estimates, that the Tyrannosauridae show significant cost of transport savings compared to more basal clades, indicating reduced energy expenditures during foraging and likely reduced need for hunting forays. This suggests that amongst theropods, hindlimb evolution was not dictated by one particular strategy. Amongst smaller bodied taxa the competing pressures of being both a predator and a prey item dominant while larger ones, freed from predation pressure, seek to maximize foraging ability. We also discuss the implications both for interactions amongst specific clades and Mesozoic paleobiology and paleoecological reconstructions as a whole.

First elaphrosaurine theropod dinosaur (Ceratosauria: Noasauridae) from Australia — A cervical vertebra from the Early Cretaceous of Victoria

Elaphrosaurinae is an enigmatic clade of gracile ceratosaurian theropod dinosaurs known from the Late Jurassic of Africa (Elaphrosaurus bambergi) and Asia (e.g., Limusaurus inextricabilis), and the early Late Cretaceous of Argentina (Huinculsaurus montesi). Elaphrosaurinae is often placed within Noasauridae as the sister taxon to Noasaurinae, a clade of small-bodied theropods that lived in South America, Africa, Madagascar and India throughout much of the Cretaceous. Herein, we report the first evidence of Elaphrosaurinae from Australia: a nearly complete middle cervical vertebra from the upper Lower Cretaceous (lower Albian) Eumeralla Formation of Cape Otway, Victoria, Australia. The fact that this site would have been situated at ~76°S towards the end of the Early Cretaceous (~110–107 Ma) implies that elaphrosaurines were capable of tolerating near-polar palaeoenvironments, whereas its age indicates that elaphrosaurines persisted in Australia until at least the late Early Cretaceous. The new Australian elaphrosaurine, in tandem with the recently described Huinculsaurus montesi from the Cenomanian–Turonian of Argentina, implies that the spatiotemporal distribution of Elaphrosaurinae has heretofore been greatly underestimated. Historic confusion of elaphrosaurines with coelurosaurs, especially ornithomimosaurs, coupled with our generally poor understanding of noasaurid evolution, might explain the apparent dearth of fossils of this theropod clade worldwide.

Bite marks on the frill of a juvenile Centrosaurus from the Late Cretaceous Dinosaur Provincial Park Formation, Alberta, Canada.

Bite marks on bones can provide critical information about interactions between carnivores and animals they consumed (or attempted to) in the fossil record. Data from such interactions is somewhat sparse and is hampered by a lack of records in the scientific literature. Here, we present a rare instance of feeding traces on the frill of a juvenile ceratopsian dinosaur from the late Campanian Dinosaur Park Formation of Alberta. It is difficult to determine the likely tracemaker(s) but the strongest candidate is a small-bodied theropod such as a dromaeosaur or juvenile tyrannosaur. This marks the first documented case of carnivore consumption of a juvenile ceratopsid, but may represent scavenging as opposed to predation.

Osteology and bone microstructure of new, small theropod dinosaur material from the early Late Cretaceous of Morocco

The ‘Kem Kem beds’ of Morocco have yielded abundant material of large-bodied theropod dinosaurs, but remains of small theropod taxa are rare. Here, we describe two femora that provide additional information on the diversity of small-bodied theropods in the Gondwanan mid-Cretaceous. An almost complete femur (ROM 64666) represents a noasaurid theropod based on the presence of an elongate anteromedial flange that arises from the distal shaft and terminates proximal to the distal end of the bone; osteohistological analysis indicates that it was from a juvenile individual. It is possible that this femur represents a juvenile Deltadromeus, which is recovered as a putative noasaurid in some phylogenetic analyses. Nevertheless, as the affinities of Deltadromeus are debated, this femur currently represents the first definitive record of a noasaurid from northern Africa and one of the few records of this clade from the early Late Cretaceous. Moreover, if this specimen is not a juvenile Deltadromeus then it probably represents a new addition to the Kem Kem theropod assemblage. A second partial femur (ROM 65779) can be identified only as an indeterminate averostran theropod. It is similar in size to ROM 64666, but the presence of an external fundamental system (EFS) indicates that it pertained to a small-sized adult individual. These observations indicate that the two femora described herein are from different theropod taxa, thereby demonstrating that at least one additional small-bodied theropod taxon was present in the Kem Kem fauna, adding to the already high theropod species-richness of this unit.

Theropod fauna from southern Australia indicates high polar diversity and climate-driven dinosaur provinciality.

The Early Cretaceous fauna of Victoria, Australia, provides unique data on the composition of high latitude southern hemisphere dinosaurs. We describe and review theropod dinosaur postcranial remains from the Aptian–Albian Otway and Strzelecki groups, based on at least 37 isolated bones, and more than 90 teeth from the Flat Rocks locality. Several specimens of medium- and large-bodied individuals (estimated up to ∼8.5 metres long) represent allosauroids. Tyrannosauroids are represented by elements indicating medium body sizes (∼3 metres long), likely including the holotype femur of Timimus hermani, and a single cervical vertebra represents a juvenile spinosaurid. Single specimens representing medium- and small-bodied theropods may be referrable to Ceratosauria, Ornithomimosauria, a basal coelurosaur, and at least three taxa within Maniraptora. Thus, nine theropod taxa may have been present. Alternatively, four distinct dorsal vertebrae indicate a minimum of four taxa. However, because most taxa are known from single bones, it is likely that small-bodied theropod diversity remains underestimated. The high abundance of allosauroids and basal coelurosaurs (including tyrannosauroids and possibly ornithomimosaurs), and the relative rarity of ceratosaurs, is strikingly dissimilar to penecontemporaneous dinosaur faunas of Africa and South America, which represent an arid, lower-latitude biome. Similarities between dinosaur faunas of Victoria and the northern continents concern the proportional representatation of higher clades, and may result from the prevailing temperate–polar climate of Australia, especially at high latitudes in Victoria, which is similar to the predominant warm–temperate climate of Laurasia, but distinct from the arid climate zone that covered extensive areas of Gondwana. Most dinosaur groups probably attained a near-cosmopolitan distribution in the Jurassic, prior to fragmentation of the Pangaean supercontinent, and some aspects of the hallmark ‘Gondwanan’ fauna of South America and Africa may therefore reflect climate-driven provinciality, not vicariant evolution driven by continental fragmentation. However, vicariance may still be detected at lower phylogenetic levels.

Intra-guild competition and its implications for one of the biggest terrestrial predators, Tyrannosaurus rex

Identifying tradeoffs between hunting and scavenging in an ecological context is important for understanding predatory guilds. In the past century, the feeding strategy of one of the largest and best-known terrestrial carnivores, Tyrannosaurus rex, has been the subject of much debate: was it an active predator or an obligate scavenger? Here we look at the feasibility of an adult T. rex being an obligate scavenger in the environmental conditions of Late Cretaceous North America, given the size distributions of sympatric herbivorous dinosaurs and likely competition with more abundant small-bodied theropods. We predict that nearly 50 per cent of herbivores would have been within a 55–85 kg range, and calculate based on expected encounter rates that carcasses from these individuals would have been quickly consumed by smaller theropods. Larger carcasses would have been very rare and heavily competed for, making them an unreliable food source. The potential carcass search rates of smaller theropods are predicted to be 14–60 times that of an adult T. rex. Our results suggest that T. rex and other extremely large carnivorous dinosaurs would have been unable to compete as obligate scavengers and would have primarily hunted large vertebrate prey, similar to many large mammalian carnivores in modern-day ecosystems.

Dinosaur eggs and nesting behaviors: A paleobiological investigation

Although dinosaur eggs were first discovered and identified in the late 1800s, limited attention was given to the scientific value of oological fossils in contrast to observations based on skeletal features. Here, we offer a review of Mesozoic saurischian egg materials, in comparison with extant crocodilians and avians, and their paleobiological interpretation based either on the presence of embryos in ovo or brooding adults on egg clutches. Our study focuses on the eggs of the oviraptorid Citipati osmolka (Mongolia), the troodontid Troodon formosus (North America), the theropod oospecies Macroelongatoolithus xixiaensis (China), the ornithothoracine bird (Argentina), an indeterminate theropod (Thailand), and titanosaurs (Argentina). Results show that (1) many oological characters and reproductive behaviors associated with modern birds are rooted among non-avian theropods, (2) there is a reproductive evolutionary cline from crocodilians to modern birds with (3) a noticeable pattern of coeval development between the accretion of eggshell layers, origination and size increased of larger air cells (inferred from egg polar asymmetry), and brooding/incubating behaviors. Most of these pre-adaptations are grouped in two main clades of the saurischian cladogram: one at the level of Oviraptorosauridae and the other at Troodontidae. Although undeniably these two theropod taxa seem to represent two important phases for the evolution of avian reproduction, the phylogenetic distance between these clades and Titanosauria cannot be ignored. As such, the reproductive features that appeared in concert in oviraptorids might have gradually evolved across more basal theropod clades. Although Troodon formosus by its egg shape and nesting behavior seems to be in this study the precursors of modern avian reproduction, the importance of small-bodied theropods such as those who laid the Phu Phok eggs cannot be dismissed and the eggs of such dinosaurs could suggest a closer phylogenetic ties to Aves than troodontids. At a higher level of inferences, there is a strong possibility that the evolution of these reproductive features is concurrent with profound physiological and metabolic changes that occurred in saurischian dinosaurs throughout their evolution.