Using linear measurements to diagnose the ecological habitat of Spinosaurus.

Much of the ecological discourse surrounding the polarising theropod Spinosaurus has centred on qualitative discussions. Using a quantitative multivariate data analytical approach on size-adjusted linear measurements of the skull, we examine patterns in skull shape across a range of sauropsid clades and three ecological realms (terrestrial, semi-aquatic, and aquatic). We utilise cluster analyses to identify emergent properties of the data which associate properties of skull shape with ecological realm occupancy. Results revealed terrestrial ecologies to be significantly distinct from both semi- and fully aquatic ecologies, the latter two were not significantly different. Spinosaurids (including Spinosaurus) plotted away from theropods in morphospace and close to both marine taxa and wading birds. The position of nares and the degree of rostral elongation had the greatest effect on categorisation. Comparisons of supervised (k-means) and unsupervised clustering demonstrated categorising taxa into three groups (ecological realms) was inappropriate and suggested instead that cluster division is based on morphological adaptations to feeding on aquatic versus terrestrial food items. The relative position of the nares in longirostrine taxa is associated with which skull bones are elongated. Rostral elongation is observed by either elongating the maxilla and the premaxilla or by elongating the maxilla only. This results in the nares positioned towards the orbits or towards the anterior end of the rostrum respectively, with implications on available feeding methods. Spinosaurids, especially Spinosaurus, show elongation in the maxilla-premaxilla complex, achieving similar functional outcomes to elongation of the premaxilla seen in birds, particularly large-bodied piscivorous taxa. Such a skull construction would bolster "stand-and-wait" predation of aquatic prey to a greater extent than serving other proposed feeding methods.

Early Jurassic origin of avian endothermy and thermophysiological diversity in dinosaurs.

A fundamental question in dinosaur evolution is how they adapted to long-term climatic shifts during the Mesozoic and when they developed environmentally independent, avian-style acclimatization, becoming endothermic.1,2 The ability of warm-blooded dinosaurs to flourish in harsher environments, including cold, high-latitude regions,3,4 raises intriguing questions about the origins of key innovations shared with modern birds,5,6 indicating that the development of homeothermy (keeping constant body temperature) and endothermy (generating body heat) played a crucial role in their ecological diversification.7 Despite substantial evidence across scientific disciplines (anatomy,8 reproduction,9 energetics,10 biomechanics,10 osteohistology,11 palaeobiogeography,12 geochemistry,13,14 and soft tissues15,16,17), a consensus on dinosaur thermophysiology remains elusive.1,12,15,17,18,19 Differential thermophysiological strategies among terrestrial tetrapods allow endotherms (birds and mammals) to expand their latitudinal range (from the tropics to polar regions), owing to their reduced reliance on environmental temperature.20 By contrast, most reptilian lineages (squamates, turtles, and crocodilians) and amphibians are predominantly constrained by temperature in regions closer to the tropics.21 Determining when this macroecological pattern emerged in the avian lineage relies heavily on identifying the origin of these key physiological traits. Combining fossils with macroevolutionary and palaeoclimatic models, we unveil distinct evolutionary pathways in the main dinosaur lineages: ornithischians and theropods diversified across broader climatic landscapes, trending toward cooler niches. An Early Jurassic shift to colder climates in Theropoda suggests an early adoption of endothermy. Conversely, sauropodomorphs exhibited prolonged climatic conservatism associated with higher thermal conditions, emphasizing temperature, rather than plant productivity, as the primary driver of this pattern, suggesting poikilothermy with a stronger dependence on higher temperatures in sauropods.

Using your head - cranial steering in pterosaurs.

The vast majority of pterosaurs are characterized by relatively large, elongate heads that are often adorned with large, elaborate crests. Projecting out in front of the body, these large heads and any crests must have had an aerodynamic effect. The working hypothesis of the present study is that these oversized heads were used to control the left-right motions of the body during flight. Using digital models of eight non-pterodactyloids ("rhamphorhyncoids") and ten pterodactyloids, the turning moments associated with the head + neck show a close and consistent correspondence with the rotational inertia of the whole body about a vertical axis in both groups, supporting the idea of a functional relationship. Turning moments come from calculating the lateral area of the head (plus any crests) and determining the associated lift (aerodynamic force) as a function of flight speed, with flight speeds being based on body mass. Rotational inertias were calculated from the three-dimensional mass distribution of the axial body, the limbs, and the flight membranes. The close correlation between turning moment and rotational inertia was used to revise the life restorations of two pterosaurs and to infer relatively lower flight speeds in another two.

New information on paleopathologies in non-avian theropod dinosaurs: a case study on South American abelisaurids.

Studies on pathological fossil bones have allowed improving the knowledge of physiology and ecology, and consequently the life history of extinct organisms. Among extinct vertebrates, non-avian dinosaurs have drawn attention in terms of pathological evidence, since a wide array of fossilized lesions and diseases were noticed in these ancient organisms. Here, we evaluate the pathological conditions observed in individuals of different brachyrostran (Theropoda, Abelisauridae) taxa, including Aucasaurus garridoi, Elemgasem nubilus, and Quilmesaurus curriei. For this, we use multiple methodological approaches such as histology and computed tomography, in addition to the macroscopic evaluation. The holotype of Aucasaurus shows several pathognomonic traits of a failure of the vertebral segmentation during development, causing the presence of two fused caudal vertebrae. The occurrence of this condition in Aucasaurus is the first case to be documented so far in non-tetanuran theropods. Regarding the holotype of Elemgasem, the histology of two fused vertebrae shows an intervertebral space between the centra, thus the fusion is limited to the distal rim of the articular surfaces. This pathology is here considered as spondyloarthropathy, the first evidence for a non-tetanuran theropod. The microstructural arrangement of the right tibia of Quilmesaurus shows a marked variation in a portion of the outer cortex, probably due to the presence of the radial fibrolamellar bone tissue. Although similar bone tissue is present in other extinct vertebrates and the cause of its formation is still debated, it could be a response to some kind of pathology. Among non-avian theropods, traumatic injuries are better represented than other maladies (e.g., infection, congenital or metabolic diseases, etc.). These pathologies are recovered mainly among large-sized theropods such as Abelisauridae, Allosauridae, Carcharodontosauridae, and Tyrannosauridae, and distributed principally among axial elements. Statistical tests on the distribution of injuries in these theropod clades show a strong association between taxa-pathologies, body regions-pathologies, and taxa-body regions, suggesting different life styles and behaviours may underlie the frequency of different injuries among theropod taxa.

The longevity bottleneck hypothesis: Could dinosaurs have shaped ageing in present-day mammals?

The evolution and biodiversity of ageing have long fascinated scientists and the public alike. While mammals, including long-lived species such as humans, show a marked ageing process, some species of reptiles and amphibians exhibit very slow and even the absence of ageing phenotypes. How can reptiles and other vertebrates age slower than mammals? Herein, I propose that evolving during the rule of the dinosaurs left a lasting legacy in mammals. For over 100 million years when dinosaurs were the dominant predators, mammals were generally small, nocturnal, and short-lived. My hypothesis is that such a long evolutionary pressure on early mammals for rapid reproduction led to the loss or inactivation of genes and pathways associated with long life. I call this the 'longevity bottleneck hypothesis', which is further supported by the absence in mammals of regenerative traits. Although mammals, such as humans, can evolve long lifespans, they do so under constraints dating to the dinosaur era.

Soft tissues influence nasal airflow in diapsids: Implications for dinosaurs.

The nasal passage performs multiple functions in amniotes, including olfaction and thermoregulation. These functions would have been present in extinct animals as well. However, fossils preserve only low-resolution versions of the nasal passage due to loss of soft-tissue structures after death. To test the effects of these lower resolution models on interpretations of nasal physiology, we performed a broadly comparative analysis of the nasal passages in extant diapsid representatives, e.g., alligator, turkey, ostrich, iguana, and a monitor lizard. Using computational fluid dynamics, we simulated airflow through 3D reconstructed models of the different nasal passages and compared these soft-tissue-bounded results to similar analyses of the same airways under the lower-resolution limits imposed by fossilization. Airflow patterns in these bony-bounded airways were more homogeneous and slower flowing than those of their soft-tissue counterparts. These data indicate that bony-bounded airway reconstructions of extinct animal nasal passages are far too conservative and place overly restrictive physiological limitations on extinct species. In spite of the diverse array of nasal passage shapes, distinct similarities in airflow were observed, including consistent areas of nasal passage constriction such as the junction of the olfactory region and main airway. These nasal constrictions can reasonably be inferred to have been present in extinct taxa such as dinosaurs.

How smart dinosaurs?

This commentary discusses the main points made in Reiner's article on the prospect that some theropod dinosaurs could have given rise to a lineage that achieved a human level of intelligence, and those made in Herculano-Houzel's article on the potentially monkey-like numbers of neurons in the pallium of large theropods, and the implications of this for their intelligence.

Could theropod dinosaurs have evolved to a human level of intelligence?

Noting that some theropod dinosaurs had large brains, large grasping hands, and likely binocular vision, paleontologist Dale Russell suggested that a branch of these dinosaurs might have evolved to a human intelligence level, had dinosaurs not become extinct. I offer reasons why the likely pallial organization in dinosaurs would have made this improbable, based on four assumptions. First, it is assumed that achieving human intelligence requires evolving an equivalent of the about 200 functionally specialized cortical areas characteristic of humans. Second, it is assumed that dinosaurs had an avian nuclear type of pallial organization, in contrast to the mammalian cortical organization. Third, it is assumed that the interactions between the different neuron types making up an information processing unit within pallium are critical to its role in analyzing information. Finally, it is assumed that increasing axonal length between the neuron sets carrying out this operation impairs its efficacy. Based on these assumptions, I present two main reasons why dinosaur pallium might have been unable to add the equivalent of 200 efficiently functioning cortical areas. First, a nuclear pattern of pallial organization would require increasing distances between the neuron groups corresponding to the separate layers of any given mammalian cortical area, as more sets of nuclei equivalent to a cortical area are interposed between the existing sets, increasing axon length and thereby impairing processing efficiency. Second, because of its nuclear organization, dinosaur pallium could not reduce axon length by folding to bring adjacent areas closer together, as occurs in cerebral cortex.

Theropod dinosaurs had primate-like numbers of telencephalic neurons.

Understanding the neuronal composition of the brains of dinosaurs and other fossil amniotes would offer fundamental insight into their behavioral and cognitive capabilities, but brain tissue is only rarely fossilized. However, when the bony brain case is preserved, the volume and therefore mass of the brain can be estimated with computer tomography; and if the scaling relationship between brain mass and numbers of neurons for the clade is known, that relationship can be applied to estimate the neuronal composition of the brain. Using a recently published database of numbers of neurons in the telencephalon of extant sauropsids (birds, squamates, and testudines), here I show that the neuronal scaling rules that apply to these animals can be used to infer the numbers of neurons that composed the telencephalon of dinosaur, pterosaur, and other fossil sauropsid species. The key to inferring numbers of telencephalic neurons in these species is first using the relationship between their estimated brain and body mass to determine whether bird-like (endothermic) or squamate-like (ectothermic) rules apply to each fossil sauropsid species. This procedure shows that the notion of "mesothermy" in dinosaurs is an artifact due to the mixing of animals with bird-like and squamate-like scaling, and indicates that theropods such as Tyrannosaurus and Allosaurus were endotherms with baboon- and monkey-like numbers of telencephalic neurons, respectively, which would make these animals not only giant but also long-lived and endowed with flexible cognition, and thus even more magnificent predators than previously thought.

"Dragons" on the landscape: Modeling the abundance of large carnivorous dinosaurs of the Upper Jurassic Morrison Formation (USA) and the Upper Cretaceous Dinosaur Park Formation (Canada).

Counts of the number of skeletal specimens of "adult" megaherbivores and large theropods from the Morrison and Dinosaur Park formations-if not biased by taphonomic artifacts-suggest that the big meat-eaters were more abundant, relative to the number of big plant-eaters, than one would expect on the basis of the relative abundance of large carnivores and herbivores in modern mammalian faunas. Models of megaherbivore population density (number of individuals per square kilometer) that attempt to take into account ecosystem productivity, the size structure of megaherbivore populations, and individual megaherbivore energy requirements, when combined with values of the large theropod/megaherbivore abundance ratio, suggest that large theropods may have been more abundant on the landscape than estimates extrapolated from the population density versus body mass relationship of mammalian carnivores. Models of the meat production of megaherbivore populations and the meat requirements of "adult" large theropods suggest that herbivore productivity would have been insufficient to support the associated number of individuals of "adult" large theropods, unless the herbivore production/biomass ratio was substantially higher, and/or the large theropod meat requirement markedly lower, than expectations based on modern mammals. Alternatively, or in addition to one or both of these other factors, large theropods likely included dinosaurs other than megaherbivores as significant components of their diet.

Iyuku raathi, a new iguanodontian dinosaur from the Early Cretaceous Kirkwood Formation, South Africa.

We name and describe a new iguanodontian dinosaur from the Early Creteceous Kirkwood Formation, Eastern Cape Province, South Africa. This dinosaur is one of only two ornithopod dinosaurs known from the Cretaceous of southern Africa, and is unique in being represented primarily by hatchling to young juvenile individuals as demonstrated by bone histological analysis. All of the juvenile material of this new taxon comes from a single, laterally-restricted bonebed and specimens were primarily recovered as partial to complete single elements, although rare articulated materials and one partial skeleton were found. Sedimentology of the bonebed suggests that this horizon heralds a change in environment upsection to a drier and more seasonal climate. This accumulation of bones is interpreted as seasonal mortality from a nesting site or nesting grounds and may be linked to this environmental shift.

Estimating bite force in extinct dinosaurs using phylogenetically predicted physiological cross-sectional areas of jaw adductor muscles.

I present a Bayesian phylogenetic predictive modelling (PPM) framework that allows the prediction of muscle parameters (physiological cross-sectional area, A Phys) in extinct archosaurs from skull width (W Sk) and phylogeny. This approach is robust to phylogenetic uncertainty and highly versatile given its ability to base predictions on simple, readily available predictor variables. The PPM presented here has high prediction accuracy (up to 95%), with downstream biomechanical modelling yielding bite force estimates that are in line with previous estimates based on muscle parameters from reconstructed muscles. This approach does not replace muscle reconstructions but one that provides a powerful means to predict A Phys from skull geometry and phylogeny to the same level of accuracy as that measured from reconstructed muscles in species for which soft tissue data are unavailable or difficult to obtain.

Comment on "The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization".

Hanson et al. (Research Articles, 7 May 2021, p. 601) claim that the shape of the vestibular apparatus reflects the evolution of reptilian locomotion. Using biomechanics, we demonstrate that semicircular canal shape is a dubious predictor of semicircular duct function. Additionally, we show that the inference methods used by Hanson et al. largely overestimate relationships between semicircular canal shape and locomotion.

Response to Comment on "The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization".

David et al. claim that vestibular shape does not reflect function and that we did not use phylogenetic inference methods in our primary analyses. We show that their claims are countered by comparative and direct experimental evidence from across Vertebrata and that their models are empirically unverified. We did use phylogenetic methods to test our hypotheses. Moreover, their phylogenetic correction attempts are methodologically inappropriate.

Comparison of vertebrate skin structure at class level: A review.

The skin is a barrier between the internal and external environment of an organism. Depending on the species, it participates in multiple functions. The skin is the organ that holds the body together, covers and protects it, and provides communication with its environment. It is also the body's primary line of defense, especially for anamniotes. All vertebrates have multilayered skin composed of three main layers: the epidermis, the dermis, and the hypodermis. The vital mission of the integument in aquatic vertebrates is mucus secretion. Cornification began in apmhibians, improved in reptilians, and endured in avian and mammalian epidermis. The feather, the most ostentatious and functional structure of avian skin, evolved in the Mesozoic period. After the extinction of the dinosaurs, birds continued to diversify, followed by the enlargement, expansion, and diversification of mammals, which brings us to the most complicated skin organization of mammals with differing glands, cells, physiological pathways, and the evolution of hair. Throughout these radical changes, some features were preserved among classes such as basic dermal structure, pigment cell types, basic coloration genetics, and similar sensory features, which enable us to track the evolutionary path. The structural and physiological properties of the skin in all classes of vertebrates are presented. The purpose of this review is to go all the way back to the agnathans and follow the path step by step up to mammals to provide a comparative large and updated survey about vertebrate skin in terms of morphology, physiology, genetics, ecology, and immunology.

The first occurrence of an avian-style respiratory infection in a non-avian dinosaur.

Other than repaired fractures, osteoarthritis, and periosteal reaction, the vertebrate fossil record has limited evidence of non-osseous diseases. This difficulty in paleontological diagnoses stems from (1) the inability to conduct medical testing, (2) soft-tissue pathologic structures are less likely to be preserved, and (3) many osseous lesions are not diagnostically specific. However, here reported for the first time is an avian-style respiratory disorder in a non-avian dinosaur. This sauropod presents irregular bony pathologic structures stemming from the pneumatic features in the cervical vertebrae. As sauropods show well-understood osteological correlates indicating that respiratory tissues were incorporated into the post-cranial skeleton, and thus likely had an 'avian-style' form of respiration, it is most parsimonious to identify these pathologic structures as stemming from a respiratory infection. Although several extant avian infections produce comparable symptoms, the most parsimonious is airsacculitis with associated osteomyelitis. From actinobacterial to fungal in origin, airsacculitis is an extremely prevalent respiratory disorder in birds today. While we cannot pinpoint the specific infectious agent that caused the airsacculitis, this diagnosis establishes the first fossil record of this disease. Additionally, it allows us increased insight into the medical disorders of dinosaurs from a phylogenetic perspective and understanding what maladies plagued the "fearfully great lizards".

The dinosaur tracks of Tyrants Aisle: An Upper Cretaceous ichnofauna from Unit 4 of the Wapiti Formation (upper Campanian), Alberta, Canada.

The Wapiti Formation of northwest Alberta and northeast British Columbia, Canada, preserves an Upper Cretaceous terrestrial vertebrate fauna that is latitudinally situated between those documented further north in Alaska and those from southern Alberta and the contiguous U.S.A. Therefore, the Wapiti Formation is important for identifying broad patterns in vertebrate ecology, diversity, and distribution across Laramidia during the latest Cretaceous. Tracksites are especially useful as they provide a range of palaeoecological, palaeoenvironmental, and behavioural data that are complementary to the skeletal record. Here, we describe the Tyrants Aisle locality, the largest in-situ tracksite known from the Wapiti Formation. The site occurs in the lower part of Unit 4 of the formation (~72.5 Ma, upper Campanian), exposed along the southern bank of the Redwillow River. More than 100 tracks are documented across at least three distinct track-bearing layers, which were deposited on an alluvial floodplain. Hadrosaurid tracks are most abundant, and are referable to Hadrosauropodus based on track width exceeding track length, broad digits, and rounded or bilobed heel margins. We suggest the hadrosaurid trackmaker was Edmontosaurus regalis based on stratigraphic context. Tyrannosaurids, probable troodontids, possible ornithomimids, and possible azhdarchid pterosaurs represent minor but notable elements of the ichnofauna, as the latter is unknown from skeletal remains within the Wapiti Formation, and all others are poorly represented. Possible social behaviour is inferred for some of the hadrosaurid and small theropod-like trackmakers based on trackway alignment, suitable spacing and consistent preservation. On a broad taxonomic level (i.e., family or above), ichnofaunal compositions indicate that hadrosaurids were palaeoecologically dominant across Laramidia during the late Campanian within both high-and low-latitude deposits, although the role of depositional environment requires further testing.

The neuroanatomy of Zulmasuchus querejazus (Crocodylomorpha, Sebecidae) and its implications for the paleoecology of sebecosuchians.

The endocranial structures of the sebecid crocodylomorph Zulmasuchus querejazus (MHNC 6672) from the Lower Paleocene of Bolivia are described in this article. Using computed tomography scanning, the cranial endocast, associated nerves and arteries, endosseous labyrinths, and cranial pneumatization are reconstructed and compared with those of extant and fossil crocodylomorphs, representative of different ecomorphological adaptations. Z. querejazus exhibits an unusual flexure of the brain, pericerebral spines, semicircular canals with a narrow diameter, as well as enlarged pharyngotympanic sinuses. First, those structures allow to estimate the alert head posture and hearing capabilities of Zulmasuchus. Then, functional comparisons are proposed between this purportedly terrestrial taxon, semi-aquatic, and aquatic forms (extant crocodylians, thalattosuchians, and dyrosaurids). The narrow diameter of the semicircular canals but expanded morphology of the endosseous labyrinths and the enlarged pneumatization of the skull compared to other forms indeed tend to indicate a terrestrial lifestyle for Zulmasuchus. Our results highlight the need to gather new data, especially from altirostral forms in order to further our understanding of the evolution of endocranial structures in crocodylomorphs with different ecomorphological adaptations.