Macroevolutionary drivers of morphological disparity in the avian quadrate.

In birds, the quadrate connects the mandible and skull, and plays an important role in cranial kinesis. Avian quadrate morphology may therefore be assumed to have been influenced by selective pressures related to feeding ecology, yet large-scale variation in quadrate morphology and its potential relationship with ecology have never been quantitatively investigated. Here, we used geometric morphometrics and phylogenetic comparative methods to quantify morphological variation of the quadrate and its relationship with key ecological features across a wide phylogenetic sample. We found non-significant associations between quadrate shape and feeding ecology across different scales of phylogenetic comparison; indeed, allometry and phylogeny exhibit stronger relationships with quadrate shape than ecological features. We show that similar quadrate shapes are associated with widely varying dietary ecologies (one-to-many mapping), while divergent quadrate shapes are associated with similar dietary ecologies (many-to-one mapping). Moreover, we show that the avian quadrate evolves as an integrated unit and exhibits strong associations with the morphologies of neighbouring bones. Our results collectively illustrate that quadrate shape has evolved jointly with other elements of the avian kinetic system, with the major crown bird lineages exploring alternative quadrate morphologies, highlighting the potential diagnostic value of quadrate morphology in investigations of bird systematics.

Origins of slow growth on the crocodilian stem lineage.

Crocodilians grow slowly and have low metabolic rates similar to other living reptiles, but palaeohistology indicates that they evolved from an ancestor with higher growth rates.1,2,3,4,5 It remains unclear when slow growth appeared in the clade due to the sparse data on key divergences among early Mesozoic members of their stem lineage. We present new osteohistological data from a broad sample of early crocodylomorphs, evaluated in a phylogenetic context alongside other pseudosuchians. We find that the transition to slow-growing bone types during mid-late ontogeny occurred around the origin of Crocodylomorpha during the Late Triassic. Earlier-diverging pseudosuchians had high maximum growth rates, as indicated by the presence of woven bone during middle and (sometimes) late ontogeny.6,7,8,9 Large-bodied pseudosuchians in particular exhibit some of the fastest-growing bone types, giving evidence for prolonged, rapid growth. By contrast, early-branching crocodylomorphs, including a new large-bodied taxon, had slow maximum rates of bone deposition, as evidenced by the presence of predominantly parallel-fibered or lamellar bone tissue during middle-late ontogeny. Late Triassic crocodylomorphs show skeletal anatomy consistent with "active" terrestrial habits,10,11,12 and their slow growth rates reject hypotheses linking this transition with sedentary, semiaquatic lifestyles or sprawling posture. Faster-growing pseudosuchian lineages go extinct in the Triassic, whereas slow-growing crocodylomorphs do not. This contrasts with the Jurassic radiation of fast-growing dinosaurs on the bird-stem lineage,13 suggesting that the End-Triassic mass extinction initiated a divergent distribution of growth strategies that persist in present-day archosaurs.

Turtle body size evolution is determined by lineage-specific specializations rather than global trends.

Organisms display a considerable variety of body sizes and shapes, and macroevolutionary investigations help to understand the evolutionary dynamics behind such variations. Turtles (Testudinata) show great body size disparity, especially when their rich fossil record is accounted for. We explored body size evolution in turtles, testing which factors might influence the observed patterns and evaluating the existence of long-term directional trends. We constructed the most comprehensive body size dataset for the group to date, tested for correlation with paleotemperature, estimated ancestral body sizes, and performed macroevolutionary model-fitting analyses. We found no evidence for directional body size evolution, even when using very flexible models, thereby rejecting the occurrence of Cope's rule. We also found no significant effect of paleotemperature on overall through-time body size patterns. In contrast, we found a significant influence of habitat preference on turtle body size. Freshwater turtles display a rather homogeneous body size distribution through time. In contrast, terrestrial and marine turtles show more pronounced variation, with terrestrial forms being restricted to larger body sizes, up to the origin of testudinids in the Cenozoic, and marine turtles undergoing a reduction in body size disparity after the extinctions of many groups in the mid-Cenozoic. Our results, therefore, suggest that long-term, generalized patterns are probably explained by factors specific to certain groups and related at least partly to habitat use.

The influence of fossils in macroevolutionary analyses of 3D geometric morphometric data: A case study of galloanseran quadrates.

In birds and other reptiles, the quadrate acts as a hinge between the lower jaw and the skull and plays an important role in avian cranial kinesis. Though previous studies have qualitatively described substantial variation in quadrate morphology among birds, none have attempted to quantify evolutionary changes in quadrate shape. Here, we investigate geometric evolution of the quadrate in Galloanserae, a major clade of extant birds uniting chicken-like (Galliformes) and duck-like (Anseriformes) fowl. We quantified morphological variation in the quadrate across 50 extant galloanseran species covering all major extant subclades using three-dimensional geometric morphometrics, and performed ancestral shape reconstructions in the context of an up-to-date neornithine phylogeny. We find that our results based only on extant quadrates may overlook plesiomorphic features captured by fossil taxa, resulting in an ancestral state reconstruction for Galloanserae that is seemingly an approximation of the average shape of the extant data set. By contrast, analyses incorporating early fossil galloanseran quadrates (from taxa such as Asteriornis, Presbyornis, and Conflicto) result in ancestral geometric reconstructions more similar to the morphology of extant galliforms, indicating that the quadrate of the last common ancestor of galloanserans may have been more morphologically and functionally similar to those of extant galliforms than to extant anseriforms. These results generally corroborate previous inferences of galloanseran quadrate plesiomorphies and identify several additional plesiomorphic features of the galloanseran quadrate for the first time. Our results illustrate the importance of incorporating fossil taxa into ancestral shape reconstructions and help elucidate important aspects of the morphology and function of the avian feeding apparatus early in crown bird evolutionary history.

Direct quantification of skeletal pneumaticity illuminates ecological drivers of a key avian trait.

Skeletal pneumaticity is a key feature of extant avian structure and biology, which first evolved among the non-flying archosaurian ancestors of birds. The widespread presence of air-filled bones across the postcranial skeleton is unique to birds among living vertebrates, but the true extent of skeletal pneumaticity has never been quantitatively investigated-hindering fundamental insights into the evolution of this key avian feature. Here, we use microCT scans of fresh, frozen birds to directly quantify the fraction of humerus volume occupied by air across a phylogenetically diverse taxon sample to test longstanding hypotheses regarding the evolution and function of avian skeletal pneumatization. Among other insights, we document weak positive allometry of internal air volume with humeral size among pneumatized humeri and provide strong support that humeral size, body mass, aquatic diving, and the presence or absence of pneumaticity all have independent effects on cortical bone thickness. Our quantitative evaluation of humeral pneumaticity across extant avian phylogeny sheds new light on the evolution and ontogenetic progression of an important aspect of avian skeletal architecture, and suggests that the last common ancestor of crown birds possessed a highly pneumatized humerus.

Reconstructing locomotor ecology of extinct avialans: a case study of Ichthyornis comparing sternum morphology and skeletal proportions.

Avian skeletal morphology is associated with locomotor function, including flight style, swimming and terrestrial locomotion, and permits informed inferences on locomotion in extinct taxa. The fossil taxon Ichthyornis (Avialae: Ornithurae) has long been regarded as highly aerial, with flight similar to terns or gulls (Laridae), and skeletal features resembling foot-propelled diving adaptations. However, rigorous testing of locomotor hypotheses has yet to be performed on Ichthyornis, despite its notable phylogenetic position as one of the most crownward stem birds. We analysed separate datasets of three-dimensional sternal shape (geometric morphometrics) and skeletal proportions (linear measurements across the skeleton), to examine how well these data types predict locomotor traits in Neornithes. We then used this information to infer locomotor capabilities of Ichthyornis. We find strong support for both soaring and foot-propelled swimming capabilities in Ichthyornis. Further, sternal shape and skeletal proportions provide complementary information on avian locomotion: skeletal proportions allow better predictions of the capacity for flight, whereas sternal shape predicts variation in more specific locomotor abilities such as soaring, foot-propelled swimming and escape burst flight. These results have important implications for future studies of extinct avialan ecology and underscore the importance of closely considering sternum morphology in investigations of fossil bird locomotion.

Mammalian forelimb evolution is driven by uneven proximal-to-distal morphological diversity.

Vertebrate limb morphology often reflects the environment due to variation in locomotor requirements. However, proximal and distal limb segments may evolve differently from one another, reflecting an anatomical gradient of functional specialization that has been suggested to be impacted by the timing of development. Here, we explore whether the temporal sequence of bone condensation predicts variation in the capacity of evolution to generate morphological diversity in proximal and distal forelimb segments across more than 600 species of mammals. Distal elements not only exhibit greater shape diversity, but also show stronger within-element integration and, on average, faster evolutionary responses than intermediate and upper limb segments. Results are consistent with the hypothesis that late developing distal bones display greater morphological variation than more proximal limb elements. However, the higher integration observed within the autopod deviates from such developmental predictions, suggesting that functional specialization plays an important role in driving within-element covariation. Proximal and distal limb segments also show different macroevolutionary patterns, albeit not showing a perfect proximo-distal gradient. The high disparity of the mammalian autopod, reported here, is consistent with the higher potential of development to generate variation in more distal limb structures, as well as functional specialization of the distal elements.

Comparative anatomy of the passerine carpometacarpus helps illuminate the early fossil record of crown Passeriformes.

The hyper-diverse clade Passeriformes (crown group passerines) comprises over half of extant bird diversity, yet disproportionately few studies have targeted passerine comparative anatomy on a broad phylogenetic scale. This general lack of research attention hinders efforts to interpret the passerine fossil record and obscures patterns of morphological evolution across one of the most diverse clades of extant vertebrates. Numerous potentially important crown passeriform fossils have proven challenging to place phylogenetically, due in part to a paucity of phylogenetically informative characters from across the passerine skeleton. Here, we present a detailed analysis of the morphology of extant passerine carpometacarpi, which are relatively abundant components of the passerine fossil record. We sampled >70% of extant family-level passerine clades (132 extant species) as well as several fossils from the Oligocene of Europe and scored them for 54 phylogenetically informative carpometacarpus characters optimised on a recently published phylogenomic scaffold. We document a considerable amount of previously undescribed morphological variation among passerine carpometacarpi, and, despite high levels of homoplasy, our results support the presence of representatives of both crown Passeri and crown Tyranni in Europe during the Oligocene.

Climatic controls on the ecological ascendancy of dinosaurs.

The ascendancy of dinosaurs to become dominant components of terrestrial ecosystems was a pivotal event in the history of life, yet the drivers of their early evolution and biodiversity are poorly understood.1,2,3 During their early diversification in the Late Triassic, dinosaurs were initially rare and geographically restricted, only attaining wider distributions and greater abundance following the end-Triassic mass extinction event.4,5,6 This pattern is consistent with an opportunistic expansion model, initiated by the extinction of co-occurring groups such as aetosaurs, rauisuchians, and therapsids.4,7,8 However, this pattern could instead be a response to changes in global climatic distributions through the Triassic to Jurassic transition, especially given the increasing evidence that climate played a key role in constraining Triassic dinosaur distributions.7,9,10,11,12,13,14,15,16 Here, we test this hypothesis and elucidate how climate influenced early dinosaur distribution by quantitatively examining changes in dinosaur and tetrapod "climatic niche space" across the Triassic-Jurassic boundary. Statistical analyses show that Late Triassic sauropodomorph dinosaurs occupied a more restricted climatic niche space than other tetrapods and dinosaurs, being excluded from the hottest, low-latitude climate zones. A subsequent, earliest Jurassic expansion of sauropodomorph geographic distribution is linked to the expansion of their preferred climatic conditions. Evolutionary model-fitting analyses provide evidence for an important evolutionary shift from cooler to warmer climatic niches during the origin of Sauropoda. These results are consistent with the hypothesis that global abundance of sauropodomorph dinosaurs was facilitated by climatic change and provide support for the key role of climate in the ascendancy of dinosaurs.

Independent origin of large labyrinth size in turtles.

The labyrinth of the vertebrate inner ear is a sensory system that governs the perception of head rotations. Central hypotheses predict that labyrinth shape and size are related to ecological adaptations, but this is under debate and has rarely been tested outside of mammals. We analyze the evolution of labyrinth morphology and its ecological drivers in living and fossil turtles, an understudied group that underwent multiple locomotory transitions during 230 million years of evolution. We show that turtles have unexpectedly large labyrinths that evolved during the origin of aquatic habits. Turtle labyrinths are relatively larger than those of mammals, and comparable to many birds, undermining the hypothesis that labyrinth size correlates directly with agility across vertebrates. We also find that labyrinth shape variation does not correlate with ecology in turtles, undermining the widespread expectation that reptilian labyrinth shapes convey behavioral signal, and demonstrating the importance of understudied groups, like turtles.

Environmental signal in the evolutionary diversification of bird skeletons.

Characterizing how variation in the tempo and mode of evolution has structured the phenotypic diversity of extant species is a central goal of macroevolution1-3. However, studies are typically limited to a handful of traits4-6, providing incomplete information. We analyse morphological diversification in living birds, an ecologically diverse group7, documenting structural scales from 'pan-skeletal' proportions down to the localized three-dimensional shape changes of individual bones. We find substantial variation in evolutionary modes among avian subgroups and among skeletal parts, indicating widespread mosaicism and possible differences in the structure of the macroevolutionary landscape across Earth's main environments. Water-linked groups, especially Aequorlitornithes (waterbirds), have repeatedly explored a large portion of their total morphospace, emphasizing variation in body proportions and in the shape of bones close to the body core, which are functionally related to the mechanics of locomotion8. By contrast, landbirds (Inopinaves) evolved distinct, group-specific body forms early in the aftermath of the K-Pg and subsequently emphasized local shape variation, especially in the head and distal limb bones, which interact more directly with the environment. Passerines, which comprise more than half of all bird species, show a conservative evolutionary dynamic that resulted in low disparity across all skeletal parts. Evidence for early establishment of the morphospace of living birds is clear for some skeletal parts, including beaks and the combined skeletal morphology. However, we find little evidence for early partitioning of that morphospace, contrary to more specific predictions of 'niche-filling' models1,9. Nevertheless, early divergence among broad environmental types may have caused an early divergence of evolutionary modes, suggesting an important role for environmental divergence in structuring the radiation of crown-group birds.

Rapid growth preceded gigantism in sauropodomorph evolution.

Sauropod dinosaurs include the largest land animals to have walked the earth, mostly weighing 10-70 tons (e.g., Sander et al.1 and Carballido et al.2). Osteohistology suggests that derived physiological traits evolved near the origin of sauropod gigantism, including both rapid and uninterrupted growth from juvenile to adult with little developmental plasticity.1,3,4 This differs from the slower, seasonally interrupted growth of their direct ancestors, as evident in most non-sauropodan sauropodomorphs, which also show developmental plasticity in some groups. Accelerated but seasonally interrupted growth is present in Lessemsauridae, the sister clade to Sauropoda, which also attained giant adult body sizes (>10 tons).5 These observations suggest a correlation between giant size and accelerated growth. However, testing this evolutionary connection has been limited by the incomplete understanding of the growth patterns in some of the closest non-giant relatives of sauropods. We present the osteohistology of two such taxa, Aardonyx celestae and Sefapanosaurus zatronensis. Both exhibit highly vascularized woven-parallel complexes, with fibrolamellar complexes during early to mid-ontogeny, containing regular growth marks. These observations provide strong evidence for rapid but seasonally interrupted growth with limited developmental plasticity (indicated by the regular spacing of growth marks). Combined with our review of early branching sauropodomorph osteohistology, these results show that highly accelerated growth rates originated among smaller, non-sauropodan sauropodomorphs weighing 1 to 2 tons but preceded the origins of giant size (>10 tons). Therefore, the capacity for rapid bone tissue formation, a derived aspect of rapid growth seen in sauropods, did not evolve specifically to enable giant body sizes but may have been a prerequisite for them.

Cranial ecomorphology of turtles and neck retraction as a possible trigger of ecological diversification.

Turtles have a highly modified body plan, including a rigid shell that constrains postcranial anatomy. Skull morphology and neck mobility may therefore be key to ecological specialization in turtles. However, the ecological signal of turtle skull morphologies has not been rigorously evaluated, leaving uncertainties about the roles of ecological adaptation and convergence. We evaluate turtle cranial ecomorphology using three-dimensional geometric morphometrics and phylogenetic comparative methods. Skull shape correlates with allometry, neck retraction capability, and different aquatic feeding ecologies. We find that ecological variables influence skull shape only, whereas a key functional variable (the capacity for neck retraction) influences both shape and size. Ecology and functional predictions from three-dimensional shape are validated by high success rates for extant species, outperforming previous two-dimensional approaches. We use this to infer ecological and functional traits of extinct species. Neck retraction evolved among crownward stem-turtles by the Late Jurassic, signaling functional decoupling of the skull and neck from the shell, possibly linked to a major episode of ecomorphological diversification. We also find strong evidence for convergent ecological adaptations among marine groups. This includes parallel loss of neck retraction, evidence for active hunting, possible grazing, and suction feeding in extinct marine groups. Our large-scale assessment of dietary and functional adaptation throughout turtle evolution reveals the timing and origin of their distinct ecomorphologies, and highlights the potential for ecology and function to have distinct effects on skull form.

Tartarugas tem um plano corpóreo bastante modificado, que inclui um casco rígido que restringe sua anatomia pós-craniana. Portanto, a morfologia craniana e a mobilidade do pescoço devem ser centrais nas especializações ecológicas de tartarugas. No entanto, o sinal ecológico das diferentes morfologias de crânio de tartarugas não foi ainda rigorosamente avaliado, deixando incertezas sobre os papéis de adaptações ecológicas e convergência. Avaliamos a ecomorfologia craniana de tartarugas utilizando morfometria geométrica tridimensional e métodos filogenéticos comparativos. A forma craniana correlaciona com alometria, capacidade de retração do pescoço e diferentes ecologias alimentares aquáticas. Encontramos que variáveis ecológicas influenciam apenas a forma do crânio, enquanto uma importante variável funcional (a capacidade de retração do pescoço) influencia tanto a forma como o tamanho do crânio. Predições ecológicas e funcionais para espécies viventes a partir de formas tridimensionais são validadas com altas taxas de sucesso, superando abordagens bidimensionais. Utilizamos isso para inferir traços ecológicos e funcionais de espécies extintas. A retração do pescoço evoluiu em linhagens extintas mais próximas à origem do grupo-coronal durante o Jurássico Final, indicando uma dissociação funcional entre crânio e pescoço do casco, algo possivelmente ligado a um importante episódio de diversificação ecomorfológica. Também encontramos forte evidência para adaptações ecológicas convergentes em grupos marinhos. Isso inclui a perda paralela da retração do pescoço, evidência de caça ativa, alimentação por sucção, além de possível preferência por plantas aquáticas em grupos marinhos extintos. Nosso estudo de larga-escala sobre adaptações funcionais e relacionadas à dieta ao longo da evolução de tartarugas revela o tempo e origem de suas distintas ecomorfologias, e destaca ainda o potencial de ecologia e função terem efeitos distintos sobre a forma craniana.

Earliest evidence for fruit consumption and potential seed dispersal by birds.

The Early Cretaceous diversification of birds was a major event in the history of terrestrial ecosystems, occurring during the earliest phase of the Cretaceous Terrestrial Revolution, long before the origin of the bird crown-group. Frugivorous birds play an important role in seed dispersal today. However, evidence of fruit consumption in early birds from outside the crown-group has been lacking. Jeholornis is one of the earliest-diverging birds, only slightly more crownward than Archaeopteryx, but its cranial anatomy has been poorly understood, limiting trophic information which may be gleaned from the skull. Originally hypothesised to be granivorous based on seeds preserved as gut contents, this interpretation has become controversial. We conducted high-resolution synchrotron tomography on an exquisitely preserved new skull of Jeholornis, revealing remarkable cranial plesiomorphies combined with a specialised rostrum. We use this to provide a near-complete cranial reconstruction of Jeholornis, and exclude the possibility that Jeholornis was granivorous, based on morphometric analyses of the mandible (3D) and cranium (2D), and comparisons with the 3D alimentary contents of extant birds. We show that Jeholornis provides the earliest evidence for fruit consumption in birds, and indicates that birds may have been recruited for seed dispersal during the earliest stages of the avian radiation. As mobile seed dispersers, early frugivorous birds could have expanded the scope for biotic dispersal in plants, and might therefore explain, at least in part, the subsequent evolutionary expansion of fruits, indicating a potential role of bird-plant interactions in the Cretaceous Terrestrial Revolution.

Birds and plants have a close relationship that has developed over millions of years. Birds became diverse and abundant around 135 million years ago. Shortly after, plants started developing new and different kinds of fruits. Today, fruit-eating birds help plants to reproduce by spreading seeds in their droppings. This suggests that birds and plants have coevolved, changing together over time. But it is not clear exactly how their relationship started. One species that might hold the answers is an early bird species known as Jeholornis. It lived in China in the Early Cretaceous, around 120 million years ago. Palaeontologists have discovered preserved seeds inside its fossilised remains. The question is, how did they get there? Some birds eat seeds directly, cracking them open or grinding them up in the stomach to extract the nutrients inside. Other birds swallow seeds when they are eating fruit. If Jeholornis belonged to this second group, it could represent one of the early steps in plant-bird coevolution. Hu et al. scanned and reconstructed a preserved Jeholornis skull and compared it to the skulls, especially the mandibles, of modern birds, including species that grind seeds, species that crack seeds and species that eat fruits, leaving the seeds whole. The analyses ruled out seed cracking. But it could not distinguish between seed grinding and fruit eating. Hu et al. therefore compared the seed remains found inside Jeholornis fossils to seeds eaten by modern birds. The fossilised seeds were intact and showed no evidence of grinding. This suggests that Jeholornis ate whole fruits for at least part of the year. At around the time Jeholornis was alive, the world was entering a phase called the Cretaceous Terrestrial Revolution, which was characterized by an explosion of new species and an expansion of both flowering plants and birds. This finding opens new avenues for scientists to explore how plant and birds might have evolved together. Similar analyses could unlock new information about how other species interacted with their environments.

Middle Jurassic fossils document an early stage in salamander evolution.

Salamanders are an important group of living amphibians and model organisms for understanding locomotion, development, regeneration, feeding, and toxicity in tetrapods. However, their origin and early radiation remain poorly understood, with early fossil stem-salamanders so far represented by larval or incompletely known taxa. This poor record also limits understanding of the origin of Lissamphibia (i.e., frogs, salamanders, and caecilians). We report fossils from the Middle Jurassic of Scotland representing almost the entire skeleton of the enigmatic stem-salamander Marmorerpeton. We use computed tomography to visualize high-resolution three-dimensional anatomy, describing morphologies that were poorly characterized in early salamanders, including the braincase, scapulocoracoid, and lower jaw. We use these data in the context of a phylogenetic analysis intended to resolve the relationships of early and stem-salamanders, including representation of important outgroups alongside data from high-resolution imaging of extant species. Marmorerpeton is united with Karaurus, Kokartus, and others from the Middle Jurassic-Lower Cretaceous of Asia, providing evidence for an early radiation of robustly built neotenous stem-salamanders. These taxa display morphological specializations similar to the extant cryptobranchid "giant" salamanders. Our analysis also demonstrates stem-group affinities for a larger sample of Jurassic species than previously recognized, highlighting an unappreciated diversity of stem-salamanders and cautioning against the use of single species (e.g., Karaurus) as exemplars for stem-salamander anatomy. These phylogenetic findings, combined with knowledge of the near-complete skeletal anatomy of Mamorerpeton, advance our understanding of evolutionary changes on the salamander stem-lineage and provide important data on early salamanders and the origins of Batrachia and Lissamphibia.

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.

Early Origins of Divergent Patterns of Morphological Evolution on the Mammal and Reptile Stem-Lineages.

The origin of amniotes 320 million years ago signaled independence from water in vertebrates and was closely followed by divergences within the mammal and reptile stem lineages (Synapsida and Reptilia). Early members of both groups had highly similar morphologies, being superficially "lizard-like" forms with many plesiomorphies. However, the extent to which they might have exhibited divergent patterns of evolutionary change, with the potential to explain the large biological differences between their living members, is unresolved. We use a new, comprehensive phylogenetic dataset to quantify variation in rates and constraints of morphological evolution among Carboniferous-early Permian amniotes. We find evidence for an early burst of evolutionary rates, resulting in the early origins of morphologically distinctive subgroups that mostly persisted through the Cisuralian. Rates declined substantially through time, especially in reptiles. Early reptile evolution was also more constrained compared with early synapsids, exploring a more limited character state space. Postcranial innovation in particular was important in early synapsids, potentially related to their early origins of large body size. In contrast, early reptiles predominantly varied the temporal region, suggesting disparity in skull and jaw kinematics, and foreshadowing the variability of cranial biomechanics seen in reptiles today. Our results demonstrate that synapsids and reptiles underwent an early divergence of macroevolutionary patterns. This laid the foundation for subsequent evolutionary events and may be critical in understanding the substantial differences between mammals and reptiles today. Potential explanations include an early divergence of developmental processes or of ecological factors, warranting cross-disciplinary investigation. [Amniote; body size; constraint; phylogeny; rate.].

Subaqueous foraging among carnivorous dinosaurs.

Secondary aquatic adaptations evolved independently more than 30 times from terrestrial vertebrate ancestors1,2. For decades, non-avian dinosaurs were believed to be an exception to this pattern. Only a few species have been hypothesized to be partly or predominantly aquatic3-11. However, these hypotheses remain controversial12,13, largely owing to the difficulty of identifying unambiguous anatomical adaptations for aquatic habits in extinct animals. Here we demonstrate that the relationship between bone density and aquatic ecologies across extant amniotes provides a reliable inference of aquatic habits in extinct species. We use this approach to evaluate the distribution of aquatic adaptations among non-avian dinosaurs. We find strong support for aquatic habits in spinosaurids, associated with a marked increase in bone density, which precedes the evolution of more conspicuous anatomical modifications, a pattern also observed in other aquatic reptiles and mammals14-16. Spinosaurids are revealed to be aquatic specialists with surprising ecological disparity, including subaqueous foraging behaviour in Spinosaurus and Baryonyx, and non-diving habits in Suchomimus. Adaptation to aquatic environments appeared in spinosaurids during the Early Cretaceous, following their divergence from other tetanuran theropods during the Early Jurassic17.

Comment on "The influence of juvenile dinosaurs on community structure and diversity".

Schroeder et al. (Reports, 26 February 2021, p. 941) reported a size gap among predatory dinosaur species. We argue that the supporting dataset is skewed toward Late Cretaceous North America and that the gap was likely absent during other intervals in most geographic regions. We urge broader consideration of this hypothesis, with quantitative evaluation of preservational and dataset biases.