Research
A full list of my 41 publications, including first-author work and papers published by my students or with collaborators, is available at Google Scholar.
Themes
Macroevolution of tetrapod morphology
The key theme underlying all my research is the study of anatomical evolution. I study the evolution of morphological variety (disparity) over large geological timescales. This includes disparity dynamics during evolutionary radiations, the evolution of ecological innovations and extinction events. So far, my research has focused on insightful case studies, mainly in marine reptiles, crocodiles, dinosaurs and lizards. In my current postdoctoral research, I am exploring large-scale patterns of morphological change inclusively across all tetrapods (amphibians, reptiles, birds, mammals), covering both living (~580 families) and extinct (>1000 families) animals in a universal framework. I will look at patterns of body size evolution and changes to skeletal morphology and physiology, and link this to the rise and fall of tetrapods over >300 million years.
The rise and fall of crocodile diversity
I have always been interested in the rise and fall of biodiversity through time. Why are some groups very diverse and others not? How well does the current biodiversity of animal groups represent their extinct ancestors? Crocodiles and their ancestors are an incredible group for exploring these ideas. Today there are only 26 crocodiles, which is a very low number when compared to other groups like birds (10,000 species). However, the 230-million-year fossil record of crocodiles and their ancestors reveals a lost world of innovation, including hundreds of species, some of which had very different lifestyles to modern species, including land-dwelling herbivores and ocean-going predators. Together with Dr Stephanie Pierce, Dr Phil Anderson and Prof. Emily Rayfield, I have shown that rapid evolution in some extinct crocodile groups, including dolphin-like thalattosuchians and small land-dwelling notosuchians, allowed them to expand into niches today occupied by other groups, particularly mammals. Whilst the modern families, the crocodiles, alligators and gharials, have evolved more slowly for the last 80 million years – limiting their ecological variety. I am now supervising new upcoming research on the evolution of teeth and bite force, and collaborating on projects about limb bone disparity in crocodile-line reptiles with Dr Davide Foffa and Dr Masaya Iijima.
Stubbs, T. L., Pierce, S. E., Elsler, A., Anderson, P. S. L., Rayfield, E. J. Benton, M. J. Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation. Proceedings of the Royal Society B: Biological Sciences 288, 20210069 (2021). (link)
Stubbs, T. L., Pierce, S. E., Rayfield, E. J., Anderson, P. S. Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B: Biological Sciences 280, 20131940 (2013). (link)
Stubbs, T. L., Pierce, S. E., Elsler, A., Anderson, P. S. L., Rayfield, E. J. Benton, M. J. Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation. Proceedings of the Royal Society B: Biological Sciences 288, 20210069 (2021). (link)
Stubbs, T. L., Pierce, S. E., Rayfield, E. J., Anderson, P. S. Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B: Biological Sciences 280, 20131940 (2013). (link)
Evolution of marine tetrapods
Tetrapods have made 12–18 independent transitions from land to sea (e.g. ichthyosaurs, turtles, mosasaurs, penguins, whales, plesiosaurs). Expanding within a new environment represents an exceptional ecological opportunity - driving diversification and evolutionary novelty. My PhD focused on exploring patterns of morphological evolution during the expansion and extinctions of Mesozoic marine reptiles, and I have worked on other related projects with collaborators Dr Ben Moon and Dr Davide Foffa, as well as multiple students. I continue to lead and collaborate on upcoming papers looking at the evolution of the skeleton in living aquatic animals, comparing expansions of marine mammals (whales, seals etc) and birds over the last 66 million years to Mesozoic reptiles, the evolution of body size and the hydrodynamics of bizarre body plans in some marine tetrapods (such as the hugely long-necked plesiosaurs).
Gutarra, S., Stubbs, T. L., Moon, B. C., Heighton, B. H., & Benton, M. J. The locomotor ecomorphology of Mesozoic marine reptiles. Palaeontology, 66(2), e12645 (2023). (link)
Liu, Q. L., Cheng, L., Stubbs, T. L., Moon, B. C., Benton, M. J., Yan, C. B., & Tian, L. Rapid neck elongation in Sauropterygia (Reptilia: Diapsida) revealed by a new basal pachypleurosaur from the Lower Triassic of China. BMC Ecology and Evolution, 23(1), 44 (2023). (link)
Laboury, A., Scheyer, T. M., Klein, N., Stubbs, T. L., & Fischer, V. High phenotypic plasticity at the dawn of the eosauropterygian radiation. PeerJ, 11, e15776 (2023). (link)
Gutarra, S., Stubbs, T.L., Moon, B.C. Palmer, C. Benton. M. J. Large size in aquatic tetrapods compensates for high drag caused by extreme body proportions. Communications Biology 5, 380 (2022). (link)
Cross, S. R. R., Moon, B. C., Stubbs, T. L., Rayfield, E. J., Benton, M. J. Climate, competition, and the rise of mosasauroid ecomorphological disparity. Palaeontology. doi: 10.1111/pala.12590 (2022). (link)
Moon, B. C., Stubbs, T. L. Early high rates and disparity in the evolution of ichthyosaurs. Communications biology 3, 1-8 (2020). (link)
Reeves, J. C., Moon, B. C., Benton, M. J., Stubbs, T. L. Evolution of ecospace occupancy by Mesozoic marine tetrapods. Palaeontology (2020) (link)
Driscoll, D. A., Dunhill, A. M., Stubbs, T. L., Benton, M. J. The mosasaur fossil record through the lens of fossil completeness. Palaeontology 62, 51-75 (2019). (link)
Foffa, D., Young, M. T., Stubbs, T. L., Dexter, K. G., Brusatte, S. L. The long-term ecology and evolution of marine reptiles in a Jurassic seaway. Nature Ecology & Evolution 2, 1548-1555 (2018). (link)
Stubbs, T. L., Benton, M. J. Ecomorphological diversifications of Mesozoic marine reptiles: the roles of ecological opportunity and extinction. Paleobiology 42, 547-573 (2016). (link)
Gutarra, S., Stubbs, T. L., Moon, B. C., Heighton, B. H., & Benton, M. J. The locomotor ecomorphology of Mesozoic marine reptiles. Palaeontology, 66(2), e12645 (2023). (link)
Liu, Q. L., Cheng, L., Stubbs, T. L., Moon, B. C., Benton, M. J., Yan, C. B., & Tian, L. Rapid neck elongation in Sauropterygia (Reptilia: Diapsida) revealed by a new basal pachypleurosaur from the Lower Triassic of China. BMC Ecology and Evolution, 23(1), 44 (2023). (link)
Laboury, A., Scheyer, T. M., Klein, N., Stubbs, T. L., & Fischer, V. High phenotypic plasticity at the dawn of the eosauropterygian radiation. PeerJ, 11, e15776 (2023). (link)
Gutarra, S., Stubbs, T.L., Moon, B.C. Palmer, C. Benton. M. J. Large size in aquatic tetrapods compensates for high drag caused by extreme body proportions. Communications Biology 5, 380 (2022). (link)
Cross, S. R. R., Moon, B. C., Stubbs, T. L., Rayfield, E. J., Benton, M. J. Climate, competition, and the rise of mosasauroid ecomorphological disparity. Palaeontology. doi: 10.1111/pala.12590 (2022). (link)
Moon, B. C., Stubbs, T. L. Early high rates and disparity in the evolution of ichthyosaurs. Communications biology 3, 1-8 (2020). (link)
Reeves, J. C., Moon, B. C., Benton, M. J., Stubbs, T. L. Evolution of ecospace occupancy by Mesozoic marine tetrapods. Palaeontology (2020) (link)
Driscoll, D. A., Dunhill, A. M., Stubbs, T. L., Benton, M. J. The mosasaur fossil record through the lens of fossil completeness. Palaeontology 62, 51-75 (2019). (link)
Foffa, D., Young, M. T., Stubbs, T. L., Dexter, K. G., Brusatte, S. L. The long-term ecology and evolution of marine reptiles in a Jurassic seaway. Nature Ecology & Evolution 2, 1548-1555 (2018). (link)
Stubbs, T. L., Benton, M. J. Ecomorphological diversifications of Mesozoic marine reptiles: the roles of ecological opportunity and extinction. Paleobiology 42, 547-573 (2016). (link)
Impacts of extinction events
Extinction events have punctuated Earth’s history leading to major turnovers in biodiversity and the disappearance of once successful groups. My research has explored the impacts of extinction events on morphological evolution, focusing on the loss of anatomical diversity and selectivity during the Permo-Triassic and Triassic-Jurassic extinctions. This work has broader implications for global change as we seek to understand the dynamic between species extinction and the loss of morphological and ecological variety.
Xue, C., Yuan, D. X., Chen, Y., Stubbs, T. L., Zhao, Y., & Zhang, Z. Morphological innovation after mass extinction events in Permian and Early Triassic conodonts based on Polygnathacea. Palaeogeography, Palaeoclimatology, Palaeoecology, 112149. (2024) (link)
Junyu, W., Foster, W. J., Tian, L., Stubbs, T. L., Benton, M. J., Xincheng, Q., Yuana, A. Decoupling of morphological disparity and taxonomic diversity during the end-Permian mass extinction. Paleobiology, 1-16 (2021). (link)
Wu, H., Zhang, Y., Stubbs, T. L., Sun, Y. Changhsingian brachiopod communities along a marine depth gradient in South China and their ecological significance in the end‐Permian mass extinction. Lethaia (2020). (link)
Wu, H., Zhang, Y., Stubbs, T. L., Liu, J., Sun, Y. A new Changhsingian (Lopingian) brachiopod fauna of the shallow‐water clastic shelf facies from Fujian Province, south‐eastern China. Papers in Palaeontology (2020). (link)
Allen, B. J., Stubbs, T. L., Benton, M. J., Puttick, M. N. Archosauromorph extinction selectivity during the Triassic–Jurassic mass extinction. Palaeontology 62, 211-224 (2019). (link)
Smithwick, F. M., Stubbs, T. L. Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events. Evolution 72, 348-362 (2018). (link)
Stubbs, T. L., Benton, M. J. Ecomorphological diversifications of Mesozoic marine reptiles: the roles of ecological opportunity and extinction. Paleobiology 42, 547-573 (2016). (link)
Xue, C., Yuan, D. X., Chen, Y., Stubbs, T. L., Zhao, Y., & Zhang, Z. Morphological innovation after mass extinction events in Permian and Early Triassic conodonts based on Polygnathacea. Palaeogeography, Palaeoclimatology, Palaeoecology, 112149. (2024) (link)
Junyu, W., Foster, W. J., Tian, L., Stubbs, T. L., Benton, M. J., Xincheng, Q., Yuana, A. Decoupling of morphological disparity and taxonomic diversity during the end-Permian mass extinction. Paleobiology, 1-16 (2021). (link)
Wu, H., Zhang, Y., Stubbs, T. L., Sun, Y. Changhsingian brachiopod communities along a marine depth gradient in South China and their ecological significance in the end‐Permian mass extinction. Lethaia (2020). (link)
Wu, H., Zhang, Y., Stubbs, T. L., Liu, J., Sun, Y. A new Changhsingian (Lopingian) brachiopod fauna of the shallow‐water clastic shelf facies from Fujian Province, south‐eastern China. Papers in Palaeontology (2020). (link)
Allen, B. J., Stubbs, T. L., Benton, M. J., Puttick, M. N. Archosauromorph extinction selectivity during the Triassic–Jurassic mass extinction. Palaeontology 62, 211-224 (2019). (link)
Smithwick, F. M., Stubbs, T. L. Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events. Evolution 72, 348-362 (2018). (link)
Stubbs, T. L., Benton, M. J. Ecomorphological diversifications of Mesozoic marine reptiles: the roles of ecological opportunity and extinction. Paleobiology 42, 547-573 (2016). (link)
Ecomorphological radiation of lepidosaurs
Lepidosaurs, the lizards, snakes and tuatara, include over 10,000 living species and a 200-million-year fossil record. They are an incredible resource for understanding how biodiversity changes through time and the links between morphology and ecology. Together with my former PhD student, Dr Jorge Herrera-Flores, collaborator Dr Arnau Bolet and Prof. Mike Benton, I have published work on the evolutionary relationships, morphology and ecology of the group. We are working on more upcoming papers about exceptional fossils and a hidden radiation deep in the group's history.
Bolet, A., Stubbs, T. L., Herrera-Flores, J. A., Benton, M. J. The Jurassic rise of squamates: insights from lepidosaur disparity and evolutionary rates. eLIFE 11, e66511 (2022) (link).
Herrera-Flores, J., Stubbs, T. L., Sour-Tovar, F. Redescription of the type specimens for the Late Jurassic rhynchocephalian Opisthias rarus and a new specimen of Theretairus antiquus from Quarry 9, Morrison Formation, Wyoming, USA. Acta Palaeontologica Polonica 67 (2022) (link).
Herrera-Flores, J. A., Elsler, A., Stubbs, T. L., Benton, M. J. Slow and fast evolutionary rates in the history of lepidosaurs. Palaeontology. pala.12579 (2021) (link).
Herrera-Flores, J. A., Stubbs, T. L., Benton, M. J. Ecomorphological diversification of squamates in the Cretaceous. Royal Society Open Science 8, 201961 (2021). (link)
Herrera-Flores, J. A., Stubbs, T. L., Elsler, A., Benton, M. J. Taxonomic reassessment of Clevosaurus latidens Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference. Journal of Paleontology 92, 734-742 (2018). (link)
Herrera‐Flores, J. A., Stubbs, T. L., Benton, M. J. Macroevolutionary patterns in Rhynchocephalia: is the tuatara (Sphenodon punctatus) a living fossil? Palaeontology 60, 319-328 (2017). (link)
Bolet, A., Stubbs, T. L., Herrera-Flores, J. A., Benton, M. J. The Jurassic rise of squamates: insights from lepidosaur disparity and evolutionary rates. eLIFE 11, e66511 (2022) (link).
Herrera-Flores, J., Stubbs, T. L., Sour-Tovar, F. Redescription of the type specimens for the Late Jurassic rhynchocephalian Opisthias rarus and a new specimen of Theretairus antiquus from Quarry 9, Morrison Formation, Wyoming, USA. Acta Palaeontologica Polonica 67 (2022) (link).
Herrera-Flores, J. A., Elsler, A., Stubbs, T. L., Benton, M. J. Slow and fast evolutionary rates in the history of lepidosaurs. Palaeontology. pala.12579 (2021) (link).
Herrera-Flores, J. A., Stubbs, T. L., Benton, M. J. Ecomorphological diversification of squamates in the Cretaceous. Royal Society Open Science 8, 201961 (2021). (link)
Herrera-Flores, J. A., Stubbs, T. L., Elsler, A., Benton, M. J. Taxonomic reassessment of Clevosaurus latidens Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference. Journal of Paleontology 92, 734-742 (2018). (link)
Herrera‐Flores, J. A., Stubbs, T. L., Benton, M. J. Macroevolutionary patterns in Rhynchocephalia: is the tuatara (Sphenodon punctatus) a living fossil? Palaeontology 60, 319-328 (2017). (link)
Morphology, function and ecology
An underlying theme of much of my work, collaborations and student supervision is the link between morphology, function and ecology. We often use variation in anatomy as a proxy for ecological differences, particularly in jaw and skull shape, but also in the limbs and post cranial skeleton. In some work, we specifically test these links mathematically.
Hill, J. J., Puttick, M. N., Stubbs, T. L., Rayfield, E. J., Donoghue, P. C. Evolution of jaw disparity in fishes. Palaeontology 61, 847-854 (2018). (link)
Foffa, D., Young, M. T., Stubbs, T. L., Dexter, K. G., Brusatte, S. L. The long-term ecology and evolution of marine reptiles in a Jurassic seaway. Nature Ecology & Evolution 2, 1548-1555 (2018). (link)
Stubbs, T. L., Pierce, S. E., Rayfield, E. J., Anderson, P. S. Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B: Biological Sciences 280, 20131940 (2013). (link)
Hill, J. J., Puttick, M. N., Stubbs, T. L., Rayfield, E. J., Donoghue, P. C. Evolution of jaw disparity in fishes. Palaeontology 61, 847-854 (2018). (link)
Foffa, D., Young, M. T., Stubbs, T. L., Dexter, K. G., Brusatte, S. L. The long-term ecology and evolution of marine reptiles in a Jurassic seaway. Nature Ecology & Evolution 2, 1548-1555 (2018). (link)
Stubbs, T. L., Pierce, S. E., Rayfield, E. J., Anderson, P. S. Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B: Biological Sciences 280, 20131940 (2013). (link)
Dinosaur herbivore evolution
During my first post-doc (2015-2017) I worked on multiple projects exploring the morphological diversity of herbivorous dinosaurs during the Cretaceous period (145-66 million years ago). Together with collaborator Dr Albert Prieto-Marquez and students, we quantified morphological variation through time and the speed of evolution, to better understand the success of Late Cretaceous dinosaurs, such as hadrosaurs. This work had a strong ‘eco-morphological’ link, focusing on teeth variety and dietary evolution. We also explored how the famous flamboyant hadrosaur crests evolved, and compared this to other aspects of their anatomy.
Stubbs, T. L., Benton, M. J., Elsler, A., Prieto-Márquez, A. Morphological innovation and the evolution of hadrosaurid dinosaurs. Paleobiology 45, 347-362 (2019). (link)
Nordén, K. K., Stubbs, T. L., Prieto-Márquez, A., Benton, M. J. Multifaceted disparity approach reveals dinosaur herbivory flourished before the end-Cretaceous mass extinction. Paleobiology 44, 620-637 (2018). (link)
Strickson, E., Prieto-Márquez, A., Benton, M. J., Stubbs, T. L. Dynamics of dental evolution in ornithopod dinosaurs. Scientific Reports 6, 28904 (2016). (link)
Stubbs, T. L., Benton, M. J., Elsler, A., Prieto-Márquez, A. Morphological innovation and the evolution of hadrosaurid dinosaurs. Paleobiology 45, 347-362 (2019). (link)
Nordén, K. K., Stubbs, T. L., Prieto-Márquez, A., Benton, M. J. Multifaceted disparity approach reveals dinosaur herbivory flourished before the end-Cretaceous mass extinction. Paleobiology 44, 620-637 (2018). (link)
Strickson, E., Prieto-Márquez, A., Benton, M. J., Stubbs, T. L. Dynamics of dental evolution in ornithopod dinosaurs. Scientific Reports 6, 28904 (2016). (link)
Measuring morphology - tools and methods
My research is very numerically driven, often using large datasets describing anatomical differences or shape variation. I am an experienced modeller and have expertise in time series analyses, comparative phylogenetic methods and geometric morphometrics, in software such as R, BayesTraits and MrBayes. I am interested in the technicalities of how we accurately measure morphological differences between different species, across different animals groups and over long time intervals.
Lautenschlager, S., Figueirido, B., Cashmore, D. D., Bendel, E.-M., Stubbs, T. L. Morphological convergence obscures functional diversity in sabre-toothed carnivores. Proceedings of the Royal Society B 287, 20201818 (2020). (link)
Schaeffer, J., Benton, M. J., Rayfield, E. J., Stubbs, T. L. Morphological disparity in theropod jaws: comparing discrete characters and geometric morphometrics. Palaeontology 63, 283-299 (2020). (link)
Hu, K., et al. Ontogenetic endocranial shape change in alligators and ostriches and implications for the development of the non‐avian dinosaur endocranium. The Anatomical Record (2020). (link)
Flannery Sutherland, J. T., Moon, B. C., Stubbs, T. L., Benton, M. J. Does exceptional preservation distort our view of disparity in the fossil record? Proceedings of the Royal Society B 286, 20190091 (2019). (link)
Navarro, C. A., Martin-Silverstone, E., Stubbs, T. L. Morphometric assessment of pterosaur jaw disparity. Royal Society Open Science 5, 172130 (2018). (link)
Lautenschlager, S., Figueirido, B., Cashmore, D. D., Bendel, E.-M., Stubbs, T. L. Morphological convergence obscures functional diversity in sabre-toothed carnivores. Proceedings of the Royal Society B 287, 20201818 (2020). (link)
Schaeffer, J., Benton, M. J., Rayfield, E. J., Stubbs, T. L. Morphological disparity in theropod jaws: comparing discrete characters and geometric morphometrics. Palaeontology 63, 283-299 (2020). (link)
Hu, K., et al. Ontogenetic endocranial shape change in alligators and ostriches and implications for the development of the non‐avian dinosaur endocranium. The Anatomical Record (2020). (link)
Flannery Sutherland, J. T., Moon, B. C., Stubbs, T. L., Benton, M. J. Does exceptional preservation distort our view of disparity in the fossil record? Proceedings of the Royal Society B 286, 20190091 (2019). (link)
Navarro, C. A., Martin-Silverstone, E., Stubbs, T. L. Morphometric assessment of pterosaur jaw disparity. Royal Society Open Science 5, 172130 (2018). (link)
Triassic vertebrate expansion
The Triassic was a time of exceptional expansion in vertebrate evolution. In multiple papers with collaborators and students, we have provided new insights into the radiations of tetrapods, fish and invertebrates, showing expansions in morphology and ecology during this interval.
Singh, S. A., Elsler, A., Stubbs, T. L., Rayfield, E. J., & Benton, M. J. Predatory synapsid ecomorphology signals growing dynamism of late Palaeozoic terrestrial ecosystems. Communications Biology, 7(1), 201 (2024). (link).
Singh. S., Elsler, A., Stubbs, T. L., Bond, R., Rayfield, E. J., Benton, M. J. Niche partitioning shaped herbivore macroevolution through the early Mesozoic. Nature Communications 12, 1-13.(2021). (link)
Moon, B. C., Stubbs, T. L. Early high rates and disparity in the evolution of ichthyosaurs. Communications biology 3, 1-8 (2020). (link)
Smithwick, F. M., Stubbs, T. L. Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events. Evolution 72, 348-362 (2018). (link)
Singh, S. A., Elsler, A., Stubbs, T. L., Rayfield, E. J., & Benton, M. J. Predatory synapsid ecomorphology signals growing dynamism of late Palaeozoic terrestrial ecosystems. Communications Biology, 7(1), 201 (2024). (link).
Singh. S., Elsler, A., Stubbs, T. L., Bond, R., Rayfield, E. J., Benton, M. J. Niche partitioning shaped herbivore macroevolution through the early Mesozoic. Nature Communications 12, 1-13.(2021). (link)
Moon, B. C., Stubbs, T. L. Early high rates and disparity in the evolution of ichthyosaurs. Communications biology 3, 1-8 (2020). (link)
Smithwick, F. M., Stubbs, T. L. Phanerozoic survivors: Actinopterygian evolution through the Permo‐Triassic and Triassic‐Jurassic mass extinction events. Evolution 72, 348-362 (2018). (link)
