Rise of Dinosaurs Reveals Major Body-Size Transitions Are Driven by Passive Processes of Trait Evolution

A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252–201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian–Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle–early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted ‘rule’ that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates

Biology, not environment, drives major patterns in maximum tetrapod body size through time

Abiotic and biological factors have been hypothesized as controlling maximum body size of tetrapods and other animals through geological time. We analyse the effects of three abiotic factors—oxygen, temperature and land area—on maximum size of Permian–Jurassic archosauromorphs and therapsids, and Cenozoic mammals, using time series generalized least-squares regression models. We also examine maximum size growth curves for the Permian–Jurassic data by comparing fits of Gompertz and logistic models. When serial correlation is removed, we find no robust correlations, indicating that these environmental factors did not consistently control tetrapod maximum size. Gompertz models—i.e. exponentially decreasing rate of size increase at larger sizes—fit maximum size curves far better than logistic models. This suggests that biological limits such as reduced fecundity and niche space availability become increasingly limiting as larger sizes are reached. Environmental factors analysed may still have imposed an upper limit on tetrapod body size, but any environmentally imposed limit did not vary substantially during the intervals examined despite variation in these environmental factors

New partial dentaries of amphitheriid mammal Palaeoxonodon ooliticus from Scotland, and posterior dentary morphology in early cladotherians

We describe two partial dentaries of mammals from the Middle Jurassic of Scotland. They belong to the early cladotherian Palaeoxonodon ooliticus. These dentaries comprise the first specimen of P. ooliticus ever found—although its significance was initially unrecognised so it remained undescribed until now—and the most recently discovered specimen, found during fieldwork in 2017. The new specimen preserves part of the coronoid process of the dentary, previously unknown for P. ooliticus, demonstrating the presence of a deep masseteric fossa, with a prominent crest enclosing the fossa anteriorly, and a masseteric foramen, located in the masseteric fossa on the buccal surface of the dentary. On the lingual surface, the mandibular foramen is offset from the Meckel’s sulcus, and positioned below the alveolar plane. These morphologies allow an updated analysis of the phylogenetic position of P. ooliticus, confirming a sister-taxa relationship between Palaeoxonodon and Amphitherium. The position of the mandibular foramen, and the slight extension of the masseteric fossa into the body of the dentary are new autapomorphies for Palaeoxonodon

The postcranial skeleton of monolophosaurus jiangi (dinosauria: Theropoda) from the Middle Jurassic of Xinjiang, China, and a review of Middle Jurassic Chinese theropods

The Middle Jurassic was a critical time in the evolution of theropod dinosaurs, highlighted by the origination and radiation of the large-bodied and morphologically diverse Tetanurae. Middle Jurassic tetanurans are rare but have been described from Europe, South America and China. In particular, China has yielded a number of potential basal tetanurans, but these have received little detailed treatment in the literature. Here we redescribe the postcranial skeleton of one of the most complete Chinese Middle Jurassic theropods, Monolophosaurus. Several features confirmthe tetanuran affinities of Monolophosaurus, but the possession of ‘primitive’ traits such as a double-faceted pubic peduncle of the ilium and a hood-like supracetabular crest suggest a basal position within Tetanurae. This conflicts with most published cladistic analyses that place Monolophosaurus in a more derived position within Allosauroidea.We review the Middle Jurassic record of Chinese theropods and compare Monolophosaurus to other Middle Jurassic theropods globally. These comparisons suggest that Monolophosaurus and Chuandongocoelurus formed an endemic theropod clade limited to the Middle Jurassic of Asia. Other Middle Jurassic Chinese theropods deserve further study

The Role of Legal Services in the Antipoverty Program

Large-scale adaptive radiations might explain the runaway success of a minority of extant vertebrate clades. This hypothesis predicts, among other things, rapid rates of morphological evolution during the early history of major groups, as lineages invade disparate ecological niches. However, few studies of adaptive radiation have included deep time data, so the links between extant diversity and major extinct radiations are unclear. The intensively studied Mesozoic dinosaur record provides a model system for such investigation, representing an ecologically diverse group that dominated terrestrial ecosystems for 170 million years. Furthermore, with 10,000 species, extant dinosaurs (birds) are the most speciose living tetrapod clade. We assembled composite trees of 614-622 Mesozoic dinosaurs/birds, and a comprehensive body mass dataset using the scaling relationship of limb bone robustness. Maximum-likelihood modelling and the node height test reveal rapid evolutionary rates and a predominance of rapid shifts among size classes in early (Triassic) dinosaurs. This indicates an early burst niche-filling pattern and contrasts with previous studies that favoured gradualistic rates. Subsequently, rates declined in most lineages, which rarely exploited new ecological niches. However, feathered maniraptoran dinosaurs (including Mesozoic birds) sustained rapid evolution from at least the Middle Jurassic, suggesting that these taxa evaded the effects of niche saturation. This indicates that a long evolutionary history of continuing ecological innovation paved the way for a second great radiation of dinosaurs, in birds. We therefore demonstrate links between the predominantly extinct deep time adaptive radiation of non-avian dinosaurs and the phenomenal diversification of birds, via continuing rapid rates of evolution along the phylogenetic stem lineage. This raises the possibility that the uneven distribution of biodiversity results not just from large-scale extrapolation of the process of adaptive radiation in a few extant clades, but also from the maintenance of evolvability on vast time scales across the history of life, in key lineages

A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning

Aim  Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially driven by climate. Although this latitudinal biodiversity gradient (LBG) also characterizes the marine invertebrate fossil record, distributions of ancient terrestrial faunas are poorly understood. This study utilizes data on the dinosaur fossil record to examine spatial patterns in terrestrial biodiversity throughout the Mesozoic.\ud Location  We compiled data on fossil occurrences across the globe.\ud Methods  We compiled a comprehensive dataset of Mesozoic dinosaur genera (738), including birds. Following the utilization of sampling standardization techniques to mediate for the uneven sampling of the fossil record, we constructed latitudinal patterns of biodiversity from this dataset.\ud Results  The dominant group of Mesozoic terrestrial vertebrates did not conform to the modern LBG. Instead, dinosaur diversity was highest at temperate palaeolatitudes throughout the 160 million year span of dinosaurian evolutionary history. Latitudinal diversity correlates strongly with the distribution of land area. Late Cretaceous sauropods and ornithischians exhibit disparate LBGs.\ud Main conclusions  The continuity of the palaeotemperate peak in dinosaur diversity indicates a diminished role for climate on the Mesozoic LBG; instead, dinosaur diversity may have been driven by the amount of land area among latitudinal belts. There is no evidence that the tropics acted as a cradle for dinosaur diversity. Geographical partitioning among major clades of herbivorous dinosaurs in the Late Cretaceous may result from the advanced stages of continental fragmentation and/or differing responses to increasing latitudinal climatic zonation. Our results suggest that the modern-day LBG on land was only established 30 million years ago, following a significant post-Eocene recalibration, potentially related to increased seasonality

The apparent exponential radiation of Phanerozoic land vertebrates is an artefact of spatial sampling biases

There is no consensus about how terrestrial biodiversity was assembled through deep time, and in particular whether it has risen exponentially over the Phanerozoic. Using a database of 60 859 fossil occurrences, we show that the spatial extent of the worldwide terrestrial tetrapod fossil record itself expands exponentially through the Phanerozoic. Changes in spatial sampling explain up to 67% of the change in known fossil species counts, and these changes are decoupled from variation in habitable land area that existed through time. Spatial sampling therefore represents a real and profound sampling bias that cannot be explained as redundancy. To address this bias, we estimate terrestrial tetrapod diversity for palaeogeographical regions of approximately equal size. We find that regional-scale diversity was constrained over timespans of tens to hundreds of millions of years, and similar patterns are recovered for major subgroups, such as dinosaurs, mammals and squamates. Although the Cretaceous/Palaeogene mass extinction catalysed an abrupt two- to three-fold increase in regional diversity 66 million years ago, no further increases occurred, and recent levels of regional diversity do not exceed those of the Palaeogene. These results parallel those recovered in analyses of local community-level richness. Taken together, our findings strongly contradict past studies that suggested unbounded diversity increases at local and regional scales over the last 100 million years