Paul Upchurch | University College London (original) (raw)

Papers by Paul Upchurch

Proceedings of the National Academy of Sciences of the United States of America, Feb 8, 2011

Palaeontology, Mar 1, 2004

The Middle Jurassic (Bathonian) Ardley trackway site in Oxfordshire, UK is described in detail. T... more The Middle Jurassic (Bathonian) Ardley trackway site in Oxfordshire, UK is described in detail. The track site is extensive, containing over 40 more-or-less continuous theropod and sauropod trackways preserved together on a single bedding plane with some trackways up to 180 m in length. The trackways display reasonable preservation, with claw marks discernible. Sauropod trackways are by far the most abundant and are representative of taxa that exhibit both narrow and wide-gauge styles of locomotion. Theropod trackways are represented by large tridactyl prints and claw impressions that display exceptionally low pace angulation values during their walking phase. One theropod trackway is unique in that it reveals a gait transition associated with a temporary increase of speed. These dinosaurs were walking over a lime-mud firmground to hardground that was emergent, or locally very shallowly submerged. GPS (Global Positioning Satellite) data reveal that the tracks, preserved during a short time interval, trend in a northeasterly direction. Multi-herding behaviour is suggested for the sauropods of differing size. The discovery of this site adds significantly to our knowledge of the taxonomic composition of Middle Jurassic ecosystems. Evidence concerning locomotor styles employed by saurischian dinosaurs, footprint identification, potential trackmaker, social interactions and observations concerning estimates of Middle Jurassic faunal diversity, as well as aspects of dinosaur locomotor evolution are discussed.

Proceedings of the National Academy of Sciences of the United States of America, May 3, 2011

PLOS ONE, May 22, 2012

Background: The most primitive ornithischian dinosaurs were small bipeds, but quadrupedality evol... more Background: The most primitive ornithischian dinosaurs were small bipeds, but quadrupedality evolved three times independently in the clade. The transition to quadrupedality from bipedal ancestors is rare in the history of terrestrial vertebrate evolution, and extant analogues do not exist. Constraints imposed on quadrupedal ornithischians by their ancestral bipedal bauplan remain unexplored, and consequently, debate continues about their stance and gait. For example, it has been proposed that some ornithischians could run, while others consider that none were cursorial. Methodology/Principal Findings: Drawing on biomechanical concepts of limb bone scaling and locomotor theory developed for extant taxa, we use the largest dataset of ornithischian postcranial measurements so far compiled to examine stance and gait in quadrupedal ornithischians. Differences in femoral midshaft eccentricity in hadrosaurs and ceratopsids may indicate that hadrosaurs placed their feet on the midline during locomotion, while ceratopsids placed their feet more laterally, under the hips. More robust humeri in the largest ceratopsids relative to smaller taxa may be due to positive allometry in skull size with body mass in ceratopsids, while slender humeri in the largest stegosaurs may be the result of differences in dermal armor distribution within the clade. Hadrosaurs are found to display the most cursorial morphologies of the quadrupedal ornithischian cades, indicating higher locomotor performance than in ceratopsids and thyreophorans. Conclusions/Significance: Limb bone scaling indicates that a previously unrealised diversity of stances and gaits were employed by quadrupedal ornithischians despite apparent convergence in limb morphology. Grouping quadrupedal ornithischians together as a single functional group hides this disparity. Differences in limb proportions and scaling are likely due to the possession of display structures such as horns, frills and dermal armor that may have affected the center of mass of the animal, and differences in locomotor behaviour such as migration, predator escape or home range size.

Butler, RJ and Upchurch P, N and DB, P and J, C (2006) A biogeographical analysis of the ornithis... more Butler, RJ and Upchurch P, N and DB, P and J, C (2006) A biogeographical analysis of the ornithischian dinosaurs; pp. In: Barrett, PM and Evans, SE, (eds.) (Proceedings) Ninth International Symposium on Mesozoic Terrestrial Ecosystems and Biota. (pp. 13 - 16). The Natural History Museum: London. ... Full text not available from this repository. ... A biogeographical analysis of the ornithischian dinosaurs; pp.

Geological Society, London, Special Publications, 2011

PLOS Biology, May 6, 2014

Large-scale adaptive radiations might explain the runaway success of a minority of extant vertebr... more 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.

Communications biology, Feb 10, 2022

Discovered in 1852, the scaly skin belonging to Haestasaurus becklesii was the first to be descri... more Discovered in 1852, the scaly skin belonging to Haestasaurus becklesii was the first to be described in any non-avian dinosaur. Accordingly, it has played a crucial role in the reconstruction of sauropod integument and dinosaurs more broadly. Here, we reassess this historic specimen using Laser-Stimulated Fluorescence (LSF), revealing extensive, previously unknown regions of skin that augment prior interpretations of its integumentary morphology and taphonomy. Under white light, polygonal-subrounded, convex scales are visible on one side of the block ('side A'), but LSF reveals extensive smaller and more flattened scales, which are diagenetically fragmented, on the reverse block surface ('side B'). Contrary to the prior interpretation that the visible scales are the epidermal undersides, the presence of convex, intrascale papilliform textures on side A suggests that the external skin surface is exposed. We define intrascale papillae and provide a review of sauropod skin morphology, which clarifies that intrascale papillae are unique to and widespread across stem Neosauropoda, and likely have an evolutionary origin in the Early Jurassic. Intrascale papillae may ultimately have been integral to the evolution of gigantism in this charismatic clade.

PLOS ONE, May 15, 2013

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robus... more Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail's morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail's aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tailfanning capacity in the earliest pygostylian birds.

Global Ecology and Biogeography, Dec 22, 2011

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially... more 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.

UCL logo UCL Discovery. ...

Acta Palaeontologica Polonica, 2011

The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, ... more The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non−avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of fea− tures that are transitional between parvicursorine and non−parvicursorine alvarezsauroids, but detailed comparisons also re− veal that some characters had a more complex distribution. We also use event−based tree−fitting to perform a quantitative analysis of alvarezsauroid biogeography incorporating several recently discovered taxa. The results suggest that there is no statistical support for previous biogeographic hypotheses that favour pure vicariance or pure dispersal scenarios as explana− tions for the distributions of alvarezsauroids across South America, North America and Asia. Instead, statistically significant biogeographic reconstructions suggest a dominant role for sympatric (or "within area") events, combined with a mix of vicariance, dispersal and regional extinction. At present the alvarezsauroid data set is too small to completely resolve the biogeographic history of this group: future studies will need to create larger data sets that encompass additional clades.

American Museum of Natural History, 2020

American Museum of Natural History Research Library, 2020

Zoological Journal of the Linnean Society, 2019

Biological Reviews, Jul 28, 2014

Non-avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of... more Non-avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of a large bolide (comet or asteroid) during an interval of massive volcanic eruptions and changes in temperature and sea level. There has long been fervent debate about how these events affected dinosaurs. We review a wealth of new data accumulated over the past two decades, provide updated and novel analyses of long-term dinosaur diversity trends during the latest Cretaceous, and discuss an emerging consensus on the extinction's tempo and causes. Little support exists for a global, long-term decline across non-avian dinosaur diversity prior to their extinction at the end of the Cretaceous. However, restructuring of latest Cretaceous dinosaur faunas in North America led to reduced diversity of large-bodied herbivores, perhaps making communities more susceptible to cascading extinctions. The abruptness of the dinosaur extinction suggests a key role for the bolide impact, although the coarseness of the fossil record makes testing the effects of Deccan volcanism difficult.

Nature Reviews Earth & Environment

Representation and discrimination of those with mobility and sensory impairments is a serious pro... more Representation and discrimination of those with mobility and sensory impairments is a serious problem in academia, especially in the Earth sciences. Non-disabled academics must fight prejudice and increase the support to, and visibility of, physically impaired colleagues, states Paul Upchurch.

Proceedings of the National Academy of Sciences of the United States of America, Feb 8, 2011

Palaeontology, Mar 1, 2004

The Middle Jurassic (Bathonian) Ardley trackway site in Oxfordshire, UK is described in detail. T... more The Middle Jurassic (Bathonian) Ardley trackway site in Oxfordshire, UK is described in detail. The track site is extensive, containing over 40 more-or-less continuous theropod and sauropod trackways preserved together on a single bedding plane with some trackways up to 180 m in length. The trackways display reasonable preservation, with claw marks discernible. Sauropod trackways are by far the most abundant and are representative of taxa that exhibit both narrow and wide-gauge styles of locomotion. Theropod trackways are represented by large tridactyl prints and claw impressions that display exceptionally low pace angulation values during their walking phase. One theropod trackway is unique in that it reveals a gait transition associated with a temporary increase of speed. These dinosaurs were walking over a lime-mud firmground to hardground that was emergent, or locally very shallowly submerged. GPS (Global Positioning Satellite) data reveal that the tracks, preserved during a short time interval, trend in a northeasterly direction. Multi-herding behaviour is suggested for the sauropods of differing size. The discovery of this site adds significantly to our knowledge of the taxonomic composition of Middle Jurassic ecosystems. Evidence concerning locomotor styles employed by saurischian dinosaurs, footprint identification, potential trackmaker, social interactions and observations concerning estimates of Middle Jurassic faunal diversity, as well as aspects of dinosaur locomotor evolution are discussed.

Proceedings of the National Academy of Sciences of the United States of America, May 3, 2011

PLOS ONE, May 22, 2012

Background: The most primitive ornithischian dinosaurs were small bipeds, but quadrupedality evol... more Background: The most primitive ornithischian dinosaurs were small bipeds, but quadrupedality evolved three times independently in the clade. The transition to quadrupedality from bipedal ancestors is rare in the history of terrestrial vertebrate evolution, and extant analogues do not exist. Constraints imposed on quadrupedal ornithischians by their ancestral bipedal bauplan remain unexplored, and consequently, debate continues about their stance and gait. For example, it has been proposed that some ornithischians could run, while others consider that none were cursorial. Methodology/Principal Findings: Drawing on biomechanical concepts of limb bone scaling and locomotor theory developed for extant taxa, we use the largest dataset of ornithischian postcranial measurements so far compiled to examine stance and gait in quadrupedal ornithischians. Differences in femoral midshaft eccentricity in hadrosaurs and ceratopsids may indicate that hadrosaurs placed their feet on the midline during locomotion, while ceratopsids placed their feet more laterally, under the hips. More robust humeri in the largest ceratopsids relative to smaller taxa may be due to positive allometry in skull size with body mass in ceratopsids, while slender humeri in the largest stegosaurs may be the result of differences in dermal armor distribution within the clade. Hadrosaurs are found to display the most cursorial morphologies of the quadrupedal ornithischian cades, indicating higher locomotor performance than in ceratopsids and thyreophorans. Conclusions/Significance: Limb bone scaling indicates that a previously unrealised diversity of stances and gaits were employed by quadrupedal ornithischians despite apparent convergence in limb morphology. Grouping quadrupedal ornithischians together as a single functional group hides this disparity. Differences in limb proportions and scaling are likely due to the possession of display structures such as horns, frills and dermal armor that may have affected the center of mass of the animal, and differences in locomotor behaviour such as migration, predator escape or home range size.

Butler, RJ and Upchurch P, N and DB, P and J, C (2006) A biogeographical analysis of the ornithis... more Butler, RJ and Upchurch P, N and DB, P and J, C (2006) A biogeographical analysis of the ornithischian dinosaurs; pp. In: Barrett, PM and Evans, SE, (eds.) (Proceedings) Ninth International Symposium on Mesozoic Terrestrial Ecosystems and Biota. (pp. 13 - 16). The Natural History Museum: London. ... Full text not available from this repository. ... A biogeographical analysis of the ornithischian dinosaurs; pp.

Geological Society, London, Special Publications, 2011

PLOS Biology, May 6, 2014

Large-scale adaptive radiations might explain the runaway success of a minority of extant vertebr... more 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.

Communications biology, Feb 10, 2022

Discovered in 1852, the scaly skin belonging to Haestasaurus becklesii was the first to be descri... more Discovered in 1852, the scaly skin belonging to Haestasaurus becklesii was the first to be described in any non-avian dinosaur. Accordingly, it has played a crucial role in the reconstruction of sauropod integument and dinosaurs more broadly. Here, we reassess this historic specimen using Laser-Stimulated Fluorescence (LSF), revealing extensive, previously unknown regions of skin that augment prior interpretations of its integumentary morphology and taphonomy. Under white light, polygonal-subrounded, convex scales are visible on one side of the block ('side A'), but LSF reveals extensive smaller and more flattened scales, which are diagenetically fragmented, on the reverse block surface ('side B'). Contrary to the prior interpretation that the visible scales are the epidermal undersides, the presence of convex, intrascale papilliform textures on side A suggests that the external skin surface is exposed. We define intrascale papillae and provide a review of sauropod skin morphology, which clarifies that intrascale papillae are unique to and widespread across stem Neosauropoda, and likely have an evolutionary origin in the Early Jurassic. Intrascale papillae may ultimately have been integral to the evolution of gigantism in this charismatic clade.

PLOS ONE, May 15, 2013

Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robus... more Theropod dinosaurs show striking morphological and functional tail variation; e.g., a long, robust, basal theropod tail used for counterbalance, or a short, modern avian tail used as an aerodynamic surface. We used a quantitative morphological and functional analysis to reconstruct intervertebral joint stiffness in the tail along the theropod lineage to extant birds. This provides new details of the tail's morphological transformation, and for the first time quantitatively evaluates its biomechanical consequences. We observe that both dorsoventral and lateral joint stiffness decreased along the non-avian theropod lineage (between nodes Theropoda and Paraves). Our results show how the tail structure of non-avian theropods was mechanically appropriate for holding itself up against gravity and maintaining passive balance. However, as dorsoventral and lateral joint stiffness decreased, the tail may have become more effective for dynamically maintaining balance. This supports our hypothesis of a reduction of dorsoventral and lateral joint stiffness in shorter tails. Along the avian theropod lineage (Avialae to crown group birds), dorsoventral and lateral joint stiffness increased overall, which appears to contradict our null expectation. We infer that this departure in joint stiffness is specific to the tail's aerodynamic role and the functional constraints imposed by it. Increased dorsoventral and lateral joint stiffness may have facilitated a gradually improved capacity to lift, depress, and swing the tail. The associated morphological changes should have resulted in a tail capable of producing larger muscular forces to utilise larger lift forces in flight. Improved joint mobility in neornithine birds potentially permitted an increase in the range of lift force vector orientations, which might have improved flight proficiency and manoeuvrability. The tail morphology of modern birds with tail fanning capabilities originated in early ornithuromorph birds. Hence, these capabilities should have been present in the early Cretaceous, with incipient tailfanning capacity in the earliest pygostylian birds.

Global Ecology and Biogeography, Dec 22, 2011

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially... more 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.

UCL logo UCL Discovery. ...

Acta Palaeontologica Polonica, 2011

The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, ... more The alvarezsauroid theropod Linhenykus monodactylus from the Upper Cretaceous of Inner Mongolia, China is the first known monodactyl non−avian dinosaur, providing important information on the complex patterns of manual evolution seen in alvarezsauroids. Here we provide a detailed description of the osteology of this taxon. Linhenykus shows a number of fea− tures that are transitional between parvicursorine and non−parvicursorine alvarezsauroids, but detailed comparisons also re− veal that some characters had a more complex distribution. We also use event−based tree−fitting to perform a quantitative analysis of alvarezsauroid biogeography incorporating several recently discovered taxa. The results suggest that there is no statistical support for previous biogeographic hypotheses that favour pure vicariance or pure dispersal scenarios as explana− tions for the distributions of alvarezsauroids across South America, North America and Asia. Instead, statistically significant biogeographic reconstructions suggest a dominant role for sympatric (or "within area") events, combined with a mix of vicariance, dispersal and regional extinction. At present the alvarezsauroid data set is too small to completely resolve the biogeographic history of this group: future studies will need to create larger data sets that encompass additional clades.

American Museum of Natural History, 2020

American Museum of Natural History Research Library, 2020

Zoological Journal of the Linnean Society, 2019

Biological Reviews, Jul 28, 2014

Non-avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of... more Non-avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of a large bolide (comet or asteroid) during an interval of massive volcanic eruptions and changes in temperature and sea level. There has long been fervent debate about how these events affected dinosaurs. We review a wealth of new data accumulated over the past two decades, provide updated and novel analyses of long-term dinosaur diversity trends during the latest Cretaceous, and discuss an emerging consensus on the extinction's tempo and causes. Little support exists for a global, long-term decline across non-avian dinosaur diversity prior to their extinction at the end of the Cretaceous. However, restructuring of latest Cretaceous dinosaur faunas in North America led to reduced diversity of large-bodied herbivores, perhaps making communities more susceptible to cascading extinctions. The abruptness of the dinosaur extinction suggests a key role for the bolide impact, although the coarseness of the fossil record makes testing the effects of Deccan volcanism difficult.

Nature Reviews Earth & Environment

Representation and discrimination of those with mobility and sensory impairments is a serious pro... more Representation and discrimination of those with mobility and sensory impairments is a serious problem in academia, especially in the Earth sciences. Non-disabled academics must fight prejudice and increase the support to, and visibility of, physically impaired colleagues, states Paul Upchurch.