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A new troodontid dinosaur, Hypnovenator matsubaraetoheorum gen. et sp. nov., is described based on an articulated postcranial skeleton recovered from the fluvial deposits of the Albian Ohyamashimo Formation of the Sasayama Group in Tambasasayama City, Hyogo Prefecture, Japan. Hypnovenator is distinguished from other troodontids by four autapomorphies and a combination of additional features. Our phylogenetic analysis positions Hypnovenator as the oldest and one of the most basal troodontines, forming a clade with Gobivenator mongoliensis. The discovery of Hypnovenator suggests that small-bodied maniraptorans with a sleeping posture were common not only in environments with volcanic and eolian events or alluvial systems but also in fluvial systems. Geometric morphometric analysis of manual ungual phalanges shows that manual ungual phalanges I and III of Hypnovenator exhibit considerable morphological variation but are functionally similar, which differs from those of non-troodontine troodontids, reflecting the transition of manual motion within Troodontinae. Hypnovenator also has mosaic features in the pes related to cursoriality. This study reveals that asymmetrical arctometatarsus occurred by the Albian, and some morphological changes, such as shorter digit IV than digit III and non-ungual phalanges of digits III with roller joints and digit IV with weakly ginglymoid articulation, arose during the early Late Cretaceous.

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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.

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The noasaurid ceratosaur Kiyacursor longipes gen. et sp. nov. is described based on a fragmentary skeleton including cervical vertebra, pectoral girdle, humerus and hind limbs from the Lower Cretaceous (Aptian) Ilek Formation at Shestakovo 1 locality in Western Siberia, Russia. This is the first ceratosaur from the Early Cretaceous of Asia, extending the stratigraphic range of Ceratosauria by 40 Myr on that continent. Kiyacursor shares unique hind limb proportions with Elaphrosaurus and Limusaurus, suggesting improved cursorial ability. These taxa show an ostrich-like specialization of the pes, with a large third metatarsal and greatly reduced second metatarsal. By contrast, all other fast running non-avian theropod dinosaurs have an arctometatarsalian pes, with the third metatarsal strongly reduced proximally. The new taxon lived in the Early Cretaceous ecosystem containing a number of other Jurassic relics, such as stem salamanders, protosuchian and shartegosuchid crocodyliforms, tritylodontid synapsids and docodontan mammaliaforms.

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The evolution of flight is a rare event in vertebrate history, and one that demands functional integration across multiple anatomical/physiological systems. The neuroanatomical basis for such integration and the role that brain evolution assumes in behavioural transformations remain poorly understood. We make progress by (i) generating a positron emission tomography (PET)-based map of brain activity for pigeons during rest and flight, (ii) using these maps in a functional analysis of the brain during flight, and (iii) interpreting these data within a macroevolutionary context shaped by non-avian dinosaurs. Although neural activity is generally conserved from rest to flight, we found significant increases in the cerebellum as a whole and optic flow pathways. Conserved activity suggests processing of self-movement and image stabilization are critical when a bird takes to the air, while increased visual and cerebellar activity reflects the importance of integrating multimodal sensory information for flight-related movements. A derived cerebellar capability likely arose at the base of maniraptoran dinosaurs, where volumetric expansion and possible folding directly preceded paravian flight. These data represent an important step toward establishing how the brain of modern birds supports their unique behavioural repertoire and provide novel insights into the neurobiology of the bird-like dinosaurs that first achieved powered flight.

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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.

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The Carnian (Upper Triassic) rocks of the Candelária Sequence present a rich record of dinosaurs, including some of the oldest known dinosaurs worldwide. In this contribution we describe the first unequivocal dinosaur from the Pivetta site, located in the Restinga Sêca municipality, Southern Brazil. The specimen CAPPA/UFSM 0373 is an isolated but well-preserved left ilium. A thorough examination of the specimen's anatomy and a phylogenetic analysis provides evidence that CAPPA/UFSM 0373 belongs to the Herrerasauria. We were able to identify several similarities with potential non-herrerasaurid herrerasaurians (e.g., Tawa hallae, "Caseosaurus crosbyensis"), which were previously only known from North American deposits. In contrast, herrerasaurids (e.g., Herrerasaurus ischigualastensis) are almost exclusively known from South America. Our results support the nesting of CAPPA/UFSM 0373 as an early-diverging herrerasaurian. Furthermore, this is potentially the first record of a non-herrerasaurid herrerasaurian in unambiguous Carnian beds, suggesting a hidden diversity of dinosaurs in the Carnian rocks of the Candelária Sequence, which can be revealed even by fragmentary specimens.

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Background: Range of motion in the forelimb of the Upper Cretaceous theropod dinosaur Mononykus olecranus, a member of the family Alvarezsauridae, has previously been investigated. However, the method used to investigate range of motion at the shoulder in M. olecranus did not follow the standardized procedure used in subsequent studies. The latter procedure yields more reliable results, and its standardization provides that its results are directly comparable to the results of similar studies in other species. I therefore reinvestigated the range of motion at the shoulder in M. olecranus, using the latter procedure. Methods: Casts of the left scapula and coracoid of M. olecranus were posed on a horizontal surface, supported from beneath with modeling clay, with the medial surface of the scapula facing toward the horizontal surface. A cast of the left humerus was posed at the limits of motion through the transverse and parasagittal planes. Photos of the poses in orthal views were superimposed and used to measure range of motion, which was measured as the angle between lines drawn down the long axis of the humerus in each position. Results: Through the transverse plane, the humerus of M. olecranus could be elevated to a subhorizontal position and depressed to a subvertical position. It could move through the parasagittal plane from a subvertical position at full protraction to a position above the horizontal at full retraction. These results correct the previous mischaracterization of shoulder motion in M. olecranus as restricted to a small arc with the arms held in a permanent sprawl. The range of humeral motion in M. olecranus is much greater than that found by the previous method and allowed the animal to tuck its arms in at the sides, in addition to allowing them to sprawl so as to orient the palm downward. The wide range of humeral motion allowed M. olecranus to forage for insects by employing hook-and-pull digging at surfaces with a wider range of orientations than the previous study showed to be possible.

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Aucasaurus garridoi is an abelisaurid theropod from the Anacleto Formation (lower Campanian, Upper Cretaceous) of Patagonia, Argentina. The holotype of Aucasaurus garridoi includes cranial material, axial elements, and almost complete fore- and hind limbs. Here we present a detailed description of the axial skeleton of this taxon, along with some paleobiological and phylogenetic inferences. The presacral elements are somewhat fragmentary, although these show features shared with other abelisaurids. The caudal series, to date the most complete among brachyrostran abelisaurids, shows several autapomorphic features including the presence of pneumatic recesses on the dorsal surface of the anterior caudal neural arches, a tubercle lateral to the prezygapophysis of mid caudal vertebrae, a marked protuberance on the lateral rim of the transverse process of the caudal vertebrae, and the presence of a small ligamentous scar near the anterior edge of the dorsal surface in the anteriormost caudal transverse process. The detailed study of the axial skeleton of Aucasaurus garridoi has also allowed us to identify characters that could be useful for future studies attempting to resolve the internal phylogenetic relationships of Abelisauridae. Computed tomography scans of some caudal vertebrae show pneumatic traits in neural arches and centra, and thus the first reported case for an abelisaurid taxon. Moreover, some osteological correlates of soft tissues present in Aucasaurus and other abelisaurids, especially derived brachyrostrans, underscore a previously proposed increase in axial rigidity within Abelisauridae.

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Tooth-marked bones provide important evidence for feeding choices made by extinct carnivorous animals. In the case of the dinosaurs, most bite traces are attributed to the large and robust osteophagous tyrannosaurs, but those of other large carnivores remain underreported. Here we report on an extensive survey of the literature and some fossil collections cataloging a large number of sauropod bones (68) from the Upper Jurassic Morrison Formation of the USA that bear bite traces that can be attributed to theropods. We find that such bites on large sauropods, although less common than in tyrannosaur-dominated faunas, are known in large numbers from the Morrison Formation, and that none of the observed traces showed evidence of healing. The presence of tooth wear in non-tyrannosaur theropods further shows that they were biting into bone, but it remains difficult to assign individual bite traces to theropod taxa in the presence of multiple credible candidate biters. The widespread occurrence of bite traces without evidence of perimortem bites or healed bite traces, and of theropod tooth wear in Morrison Formation taxa suggests preferential feeding by theropods on juvenile sauropods, and likely scavenging of large-sized sauropod carcasses.

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It has long been discussed whether non-avian dinosaurs were physiologically closer to ectotherms or endotherms, with the internal nasal structure called the respiratory turbinate present in extant endotherms having been regarded as an important clue for this conundrum. However, the physiological function and relevance of this structure for dinosaur physiology are still controversial. Here, we found that the size of the nasal cavity relative to the head size of extant endotherms is larger than those of extant ectotherms, with that of the dromaeosaurid Velociraptor being below the extant endotherms level. The result suggests that a large nasal cavity accommodating a well-developed respiratory turbinate is primarily important as a thermoregulation apparatus for large brains characteristic of endothermic birds and mammals, and the nasal cavity of Velociraptor was apparently not large enough to carry out this role required for an endothermic-sized brain. In addition, a hypothesis that the enlargement of the nasal cavity for brain cooling has been associated with the skull modification in the theropod lineage toward modern birds is proposed herein. In particular, the reduction of the maxilla in derived avialans may have coincided with acquisition of the avian-like cephalic thermoregulation system.

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The digital reconstruction of neurocranial endocasts has elucidated the gross brain structure and potential ecological attributes of many fossil taxa, including Irritator, a spinosaurine spinosaurid from the "mid" Cretaceous (Aptian) of Brazil. With unexceptional hearing capabilities, this taxon was inferred to integrate rapid and controlled pitch-down movements of the head that perhaps aided in the predation of small and agile prey such as fish. However, the neuroanatomy of baryonychine spinosaurids remains to be described, and potentially informs on the condition of early spinosaurids. Using micro-computed tomographic scanning (µCT), we reconstruct the braincase endocasts of Baryonyx walkeri and Ceratosuchops inferodios from the Wealden Supergroup (Lower Cretaceous) of England. We show that the gross endocranial morphology is similar to other non-maniraptoriform theropods, and corroborates previous observations of overall endocranial conservatism amongst more basal theropods. Several differences of unknown taxonomic utility are noted between the pair. Baryonychine neurosensory capabilities include low-frequency hearing and unexceptional olfaction, whilst the differing morphology of the floccular lobe tentatively suggests less developed gaze stabilisation mechanisms relative to spinosaurines. Given the morphological similarities observed with other basal tetanurans, baryonychines likely possessed comparable behavioural sophistication, suggesting that the transition from terrestrial hypercarnivorous ancestors to semi-aquatic "generalists" during the evolution of Spinosauridae did not require substantial modification of the brain and sensory systems.

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Megaraptora is a group of enigmatic, carnivorous non-avian theropod dinosaurs from the Cretaceous of Asia, Australia, and especially South America. Perhaps the most striking aspect of megaraptoran morphology is the large, robustly constructed forelimb that, in derived members of the clade, terminates in a greatly enlarged manus with hypertrophied, raptorial unguals on the medialmost two digits and a substantially smaller ungual on digit III. The unique forelimb anatomy of megaraptorans was presumably associated with distinctive functional specializations; nevertheless, its paleobiological significance has not been extensively explored. Here we draw from observations of the pectoral girdle and forelimb skeletons of Megaraptora and myological assessments of other archosaurian taxa to provide a comprehensive reconstruction of the musculature of this anatomical region in these singular theropods. Many muscle attachment sites on megaraptoran forelimb bones are remarkably well developed, which in turn suggests that the muscles themselves were functionally significant and important to the paleobiology of these theropods. Furthermore, many of these attachments became increasingly pronounced through megaraptoran evolutionary history, being substantially better developed in derived taxa such as Australovenator wintonensis and especially Megaraptor namunhuaiquii than in early branching forms such as Fukuiraptor kitadaniensis. When considered alongside previous range of motion hypotheses for Australovenator, our results indicate that megaraptorans possessed a morphologically and functionally specialized forelimb that was capable of complex movements. Notable among these were extensive extension and flexion, particularly in the highly derived manus, as well as enhanced humeral protraction, attributes that very probably aided in prey capture.

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Despite nearly 200 years of scientific collecting and study, none of the extinct, bipedal, predatory, theropod dinosaurs have been reliably shown to exceed 12 m in length. Using digital 3D models of theropods with lengths spanning 80 cm to 12 m, their body masses were found to scale to the 3.5 power of body lengths. The lateral area of the pelvis and the cross-sectional area of the tail base of these animals corresponds to the cross-sectional areas of key muscle groups important for balance and locomotion, and both scale to the 2.4 power of body length. Body accelerations in the lateral and forward directions are, using F = ma, given by dividing muscle area (force proxy) by body mass. Plotting these acceleration estimates against body length shows them to decrease exponentially. The largest theropods with body lengths of 10-12 m have less than 10% of the acceleration capacity of the smaller forms. The distinct lack of fossil remains of theropods demonstrably longer than 12 m suggests that the theropod body plan had an upper size limit based on a minimum acceleration threshold. Rotational inertia of the theropod body was found to be proportional to body length raised to the 5.5 power, and with increasing length, the capacity for agility would rapidly diminish. The tight relationship between theropod pelvic area and body length allows for the estimation of body lengths of specimens lacking complete axial skeletons, and this is done for four, large, well-preserved pelves.

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A set of fragmentary bones excavated from the Inversand Company Pit at Sewell, Gloucester County, NJ, contains portions of broken hollow femur bone that display unusual interior structure. Two hypotheses are considered; (1) the lumina represents the distinctive physical features of medullary bone as described by M. Schweitzer in a series of papers; or (2) the interior bone growth is a pathology. The specimen is attributed to the Theropoda on the basis of the possession of a pneumatic foramen in one of the bones. This specimen is from the upper part of the New Egypt Formation, just below the K/T boundary in the base of the overlying Hornerstown Formation. This stratigraphic horizon is the same as the type specimen of Dryptosaurus aquiluguis which was excavated from a marl pit nearby.

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Dromaeosaurids (Theropoda: Dromaeosauridae), a group of dynamic, swift predators, have a sparse fossil record, particularly at the end of the Cretaceous Period. The recently described Dineobellator notohesperus, consisting of a partial skeleton from the Upper Cretaceous (Maastrichtian) of New Mexico, is the only diagnostic dromaeosaurid to be recovered from the latest Cretaceous of the southwestern United States. Reinterpreted and newly described material include several caudal vertebrae, portions of the right radius and pubis, and an additional ungual, tentatively inferred to be from manual digit III. Unique features, particularly those of the humerus, unguals, and caudal vertebrae, distinguish D. notohesperus from other known dromaeosaurids. This material indicates different physical attributes among dromaeosaurids, such as use of the forearms, strength in the hands and feet, and mobility of the tail. Several bones in the holotype exhibit abnormal growth and are inferred to be pathologic features resulting from an injury or disease. Similar lengths of the humerus imply Dineobellator and Deinonychus were of similar size, at least regarding length and/or height, although the more gracile nature of the humerus implies Dineobellator was a more lightly built predator. A new phylogenetic analysis recovers D. notohesperus as a dromaeosaurid outside other previously known and named clades. Theropod composition of the Naashoibito Member theropod fauna is like those found in the more northern Late Cretaceous North American ecosystems. Differences in tooth morphologies among recovered theropod teeth from the Naashoibito Member also implies D. notohesperus was not the only dromaeosaurid present in its environment.

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Giant carnivorous dinosaurs such as Tyrannosaurus rex and abelisaurids are characterized by highly reduced forelimbs that stand in contrast to their huge dimensions, massive skulls, and obligate bipedalism.1,2 Another group that follows this pattern, yet is still poorly known, is the Carcharodontosauridae: dominant predators that inhabited most continents during the Early Cretaceous3-5 and reached their largest sizes in Aptian-Cenomanian times.6-10 Despite many discoveries over the last three decades, aspects of their anatomy, especially with regard to the skull, forearm, and feet, remain poorly known. Here we report a new carcharodontosaurid, Meraxes gigas, gen. et sp. nov., based on a specimen recovered from the Upper Cretaceous Huincul Formation of northern Patagonia, Argentina. Phylogenetic analysis places Meraxes among derived Carcharodontosauridae, in a clade with other massive South American species. Meraxes preserves novel anatomical information for derived carcharodontosaurids, including an almost complete forelimb that provides evidence for convergent allometric trends in forelimb reduction among three lineages of large-bodied, megapredatory non-avian theropods, including a remarkable degree of parallelism between the latest-diverging tyrannosaurids and carcharodontosaurids. This trend, coupled with a likely lower bound on forelimb reduction, hypothesized to be about 0.4 forelimb/femur length, combined to produce this short-armed pattern in theropods. The almost complete cranium of Meraxes permits new estimates of skull length in Giganotosaurus, which is among the longest for theropods. Meraxes also provides further evidence that carchardontosaurids reached peak diversity shortly before their extinction with high rates of trait evolution in facial ornamentation possibly linked to a social signaling role.

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While the terrestrial fossil record of the mid-Cretaceous interval (Aptian to Cenomanian) in North America has been poorly studied, the recent focus on fossil localities from the western United States has offered a more detailed picture of vertebrate diversity, ecosystem dynamics and faunal turnover that took place on the western landmass of Laramidia. This is in stark contrast to the terrestrial record from the eastern landmass of Appalachia, where vertebrate fossils are rare and consist mostly of isolated and fragmentary remains. However, a detailed understanding of these fossil communities during this interval is necessary for comparison of the faunal patterns that developed during the opening of the Western Interior Seaway (WIS). The Woodbine Group of Texas is a Cenomanian age (95-100 mya) deposit consisting of shallow marine, deltaic, and terrestrial communities, which were only recently separated from their western counterparts. These deposits have yielded a wealth of vertebrate remains, yet non-avian theropods are still largely unknown. Recently, multiple localities in the Lewisville Formation of the Woodbine Group have yielded new non-avian theropod material, including numerous isolated teeth and postcranial remains. While largely fragmentary, this material is sufficiently diagnostic to identify the following taxa: a large-bodied carcharodontosaur, a mid-sized tyrannosauroid, a large ornithomimosaur, a large dromaeosaurine, a small dromaeosaurid, a small troodontid, and a small coelurosaur. Some of these groups represent the first occurrence for Appalachia and provide a broader understanding of a newly expanded faunal diversity for the Eastern landmass. The Lewisville Formation theropod fauna is similar in taxonomic composition to contemporaneous deposits in Laramidia, confirming that these groups were widespread across the continent prior to extension of the WIS. The Lewisville Formation documents the transitional nature of Cenomanian coastal ecosystems in Texas while providing additional details on the evolution of Appalachian communities shortly after WIS extension.

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Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

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The adult ankle of early reptiles had five distal tarsal (dt) bones, but in Dinosauria, these were reduced to only two: dt3 and dt4, articulated to metatarsals (mt) mt3 and mt4. Birds have a single distal tarsal ossification center that fuses to the proximal metatarsals to form a new adult skeletal structure: the composite tarsometatarsus. This ossification center develops within a single large embryonic cartilage, but it is unclear if this cartilage results from fusion of earlier cartilages. We studied embryos in species from four different bird orders, an alligatorid, and an iguanid. In all embryos, cartilages dt2, dt3, and dt4 are formed. In the alligatorid and the iguanid, dt2 failed to ossify: only dt3 and dt4 develop into adult bones. In birds, dt2, dt3, and dt4 fuse to form the large distal tarsal cartilage; the ossification center then develops above mt3, in cartilage presumably derived from dt3. During the entire dinosaur-bird transition, a dt2 embryonic cartilage was always formed, as inferred from the embryology of extant birds and crocodilians. We propose that in the evolution of the avian ankle, fusion of cartilages dt3 and dt2 allowed ossification from dt3 to progress into dt2, which began to contribute bone medially, while fusion of dt3 to dt4 enabled the evolutionary loss of the dt4 ossification center. As a result, a single ossification center expands into a plate-like unit covering the proximal ends of the metatarsals, that is key to the development of an integrated tarsometatarsus.

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Dinosaurs dominated Mesozoic terrestrial ecosystems globally. However, whereas a pole-to-pole geographic distribution characterized ornithischians and theropods, sauropods were restricted to lower latitudes. Here, we evaluate the role of climate in shaping these biogeographic patterns through the Jurassic-Cretaceous (201-66 mya), combining dinosaur fossil occurrences, past climate data from Earth System models, and habitat suitability modeling. Results show that, uniquely among dinosaurs, sauropods occupied climatic niches characterized by high temperatures and strongly bounded by minimum cold temperatures. This constrained the distribution and dispersal pathways of sauropods to tropical areas, excluding them from latitudinal extremes, especially in the Northern Hemisphere. The greater availability of suitable habitat in the southern continents, particularly in the Late Cretaceous, might be key to explaining the high diversity of sauropods there, relative to northern landmasses. Given that ornithischians and theropods show a flattened or bimodal latitudinal biodiversity gradient, with peaks at higher latitudes, the closer correspondence of sauropods to a subtropical concentration could hint at fundamental thermophysiological differences to the other two clades.

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