Research Program & Publications
My research examines palaeoenvironments, palaeoecology, and the life history and evolution of vertebrates in greenhouse systems.
I use a multi-scale, systems approach to disentangle complex interactions between environments and ecosystems in deep time.
(A full list of my publications can be found further down this page)
I use a multi-scale, systems approach to disentangle complex interactions between environments and ecosystems in deep time.
(A full list of my publications can be found further down this page)
Primary Research
From Organisms to Ecosystems: multi-scale approaches to palaeobiology & environments
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Cretaceous Community Dynamics & Environmental Reconstruction
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Analyzing trends in vertebrate community structure to test the response of species and ecosystems to environmental perturbation. Combination of sedimentary palaeoenvironment data & analyses of fossil assemblages from microsites.
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Applying stable isotope analyses to reconstruct diet, environment, trophic interactions, and habitat preferences in extant and ancient coastal floodplain forest systems.
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Quantifying the variability present in multiple growth proxies found in bone microstructure and reconstructing the growth curves and body size evolution of different dinosaur clades.
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Secondary Research
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Theropod macroevolution & biodiversity, amniote dental histology, the evolution of sexual dimorphism in carnivorans (and particularly pinnipeds)
PDFs are available upon request. Google Scholar page is here
ORCID: 0000-0002-4261-1323
ORCID: 0000-0002-4261-1323
Peer-Reviewed Publications
26. Thompson, M., Cullen, TM., Shröder-Adams, C., Ryan, MJ. (Submitted). Multi-proxy palaeoenvironmental and palaeoecological reconstruction of the Foremost Formation (Upper Cretaceous; Campanian) of Alberta. Submitted to Palaeo-3.
25. Wilkinson, RD., Cullen, TM., Avrahami, H., Zanno, L., Evans, DC. (Submitted). Paleontology and sedimentology of the Acheroraptor holotype locality, a multi-taxic bonebed in the Hell Creek Formation of Montana, USA. Submitted to Palaeo-3.
24. Cullen, TM., Zhang, S., Spencer, J., Cousens, B. (Submitted). Sr-C-O isotope signatures reveal herbivore niche-partitioning in a Cretaceous greenhouse ecosystem. Submitted to Palaeontology.
23. Nanglu, K., Cullen, TM. (In Review). Across space and time: an empirical investigation and review of sampling and analytical biases in fossil data across macroecological scales. Under review at Global Ecology and Biogeography.
22. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Huang, L., Evans, DC. (In Review). Dinosaur enamel 13C-enrichment reflects non-analogue environmental conditions in a ‘vanished world’. Under review at Proceedings of the Royal Society B.
21. Cullen, TM., Brown, CM., Chiba, K., Brink, KS., Makovicky, PJ., Evans, DC. (Accepted). Growth variability, dimensional scaling, and the interpretation of osteohistological data. Biology Letters. (MS#: RSBL-2021-0383)
20. Benson, R., Brown, CM., Campione, N., Cullen, TM., Evans, DC., Zanno, L. (Accepted). Comment on the influence of juvenile dinosaurs on community structure and diversity. Science. (MS#: abj5976)
19. Cullen, TM., Dunn, R., Suarez, C., Chiba, K. (Accepted). Chp. 2.5: Studying Ecosystems. In The Complete Dinosaur, 3rd Edition. Edited by L. Zanno, T. Holtz, and V. Arbour. Indiana University Press. [invited book chapter contribution]
18. Cullen, TM. (In Press). Stable isotopic analyses of living and extinct crocodylians: implications for understanding their ecology, environments, and physiology. In Crocodylian Biology and Paleobiology. Edited by H Woodward and J Farlow. Indiana University Press. [invited book chapter contribution]
17. Cullen, TM., Zanno, L., Larson, DW., Todd, E., Currie, PJ., Evans, DC. (2021). Anatomical, morphometric, and stratigraphic review of theropod biodiversity in the Dinosaur Park Formation (Late Cretaceous; Campanian) of Alberta. Canadian Journal of Earth Sciences 58(9): 870-884.
16. Cullen, TM., Canale, JI., Apesteguía, S., Smith, NS., Hu, D., Makovicky, PJ. (2020). Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proceedings of the Royal Society B 287: 20202258. http://dx.doi.org/10.1098/rspb.2020.2258.
15. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Huang, L., Fanti, F., Goodwin, MB., Ryan, MJ., Evans, DC. (2020). Large-scale stable isotope characterization of a Late Cretaceous dinosaur-dominated ecosystem. Geology 48(6): 546-551. (Featured Article & subject of GSA blog)
14. Cullen, TM., Simon, DJ., Benner, EKC., Evans, DC. (2020). Morphology and osteohistology of a large-bodied caenagnathid (Theropoda: Oviraptorosauria) from the Hell Creek Formation (Montana): implications for size-based classifications and growth estimation in theropods. Papers in Palaeontology. https://doi.org/10.1002/spp2/1302
13. Arbour, VM., Evans, DC., Simon, DJ., Cullen, TM., Braman, D. (2019). Cretaceous flora and fauna of the Sustut Group near the Sustut River, northern British Columbia, Canada. Canadian Journal of Earth Sciences. https://doi.org/10.1139/cjes-2019-0031.
12. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Goodwin, MB., Huang, L., and Evans, DC. (2019). Stable isotopic characterization of a coastal floodplain forest community: a case-study for isotopic reconstruction of Mesozoic vertebrate assemblages. Royal Society Open Science 6: 181210. http://dx.doi.org/10.1098/rsos.181210.
11. McFeeters, B., Ryan, MJ., and Cullen, TM. (2018). Response to Brownstein (2018) 'Rebuttal of McFeeters, Ryan and Cullen, 2018'. Vertebrate Anatomy Morphology Palaeontology 6: 73-74. dx.doi.org/10.18435/vamp29343
10. McFeeters, B., Ryan, MJ., and Cullen, TM. (2018). Positional variation in pedal unguals of North American ornithomimids (Dinosauria, Theropoda): a response to Brownstein (2017) . Vertebrate Anatomy Morphology Palaeontology 5. doi: 10.18435/vamp29283
9. LeBlanc, ARH., Brink, K., Cullen, TM., and Reisz, R. (2017). Evolutionary implications of tooth attachment versus tooth implantation: a case study using dinosaur, crocodilian, and mammal teeth. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2017.1354006
8. Evans, DC., Cullen, TM., Larson, DW., and Rego, A. (2017). A new species of troodontid theropod (Dinosauria: Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta, Canada. Canadian Journal of Earth Sciences. doi: 10.1139/cjes-2017-0034 (Postprint available on PaleorXiv: https://paleorxiv.org/b8mqe/)
7. Cullen, TM. and Evans, DC. (2016). Palaeoenvironmental drivers of vertebrate community composition in the Belly River Group (Campanian) of Alberta, Canada, with implications for dinosaur biogeography. BMC Ecology. doi: 10.1186/s12898-016-0106-8 (Editor's Pick and BMC Ecology top 10 highlight for 2016, and subject of a BMC Series blog)
6. McFeeters, B., Ryan, MJ., Schröder-Adams, C., and Cullen, TM. (2016). A new ornithomimid theropod from the Dinosaur Park Formation of Alberta, Canada. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2016.1221415
5. Cullen, TM., Fanti, F., Capobianco, C., Ryan, MJ., and Evans, DC. (2016). A vertebrate microsite from a marine-terrestrial transition in the Foremost Formation (Campanian) of Alberta, Canada, and the use of faunal assemblage data as a palaeoenvironmental indicator. Palaeogeography, Palaeoclimatology, Palaeoecology 444: 101-114. doi:10.1016/j.palaeo.2015.12.015 (Postprint available on PaleorXiv: https://paleorxiv.org/43j7z/)
4. Cullen, TM., Ryan, MJ., Currie, PJ., Kobayashi, Y., and Evans, DC. (2014). Osteohistological variation in growth marks and osteocyte lacunar density in a theropod dinosaur (Coelurosauria: Ornithomimidae). BMC Evolutionary Biology. doi: 10.1186/s12862-014-0231-y. (Editor’s Pick and a BMC series highlight for November 2014, and included in list of the 12 best figures in BMC Evolutionary Biology for 2014)
3. Evans, DC., Larson, DW., Cullen, TM., & Sullivan, RM. (2014). "Saurornitholestes" robustus is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences 51(7): 730-734. doi: 10.1139/cjes-2014-0073
2. Cullen, TM., Fraser, D., Rybczynski, N., and Schroder-Adams, C. (2014). Early evolution of sexual dimorphism and polygyny in Pinnipedia. Evolution 68-5: 1469–1484. (Subject of the May 2014 cover of Evolution) doi: https://dx.doi.org/10.1111/evo.12360 (Postprint available on PaleorXiv: https://paleorxiv.org/zgh62/)
1. Cullen TM., Ryan MJ, Schröder-Adams C, Currie PJ, Kobayashi Y. (2013). An Ornithomimid (Dinosauria) Bonebed from the Late Cretaceous of Alberta, with Implications for the Behavior, Classification, and Stratigraphy of North American Ornithomimids. PLoS ONE 8(3): e58853. doi:10.1371/journal.pone.0058853
25. Wilkinson, RD., Cullen, TM., Avrahami, H., Zanno, L., Evans, DC. (Submitted). Paleontology and sedimentology of the Acheroraptor holotype locality, a multi-taxic bonebed in the Hell Creek Formation of Montana, USA. Submitted to Palaeo-3.
24. Cullen, TM., Zhang, S., Spencer, J., Cousens, B. (Submitted). Sr-C-O isotope signatures reveal herbivore niche-partitioning in a Cretaceous greenhouse ecosystem. Submitted to Palaeontology.
23. Nanglu, K., Cullen, TM. (In Review). Across space and time: an empirical investigation and review of sampling and analytical biases in fossil data across macroecological scales. Under review at Global Ecology and Biogeography.
22. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Huang, L., Evans, DC. (In Review). Dinosaur enamel 13C-enrichment reflects non-analogue environmental conditions in a ‘vanished world’. Under review at Proceedings of the Royal Society B.
21. Cullen, TM., Brown, CM., Chiba, K., Brink, KS., Makovicky, PJ., Evans, DC. (Accepted). Growth variability, dimensional scaling, and the interpretation of osteohistological data. Biology Letters. (MS#: RSBL-2021-0383)
20. Benson, R., Brown, CM., Campione, N., Cullen, TM., Evans, DC., Zanno, L. (Accepted). Comment on the influence of juvenile dinosaurs on community structure and diversity. Science. (MS#: abj5976)
19. Cullen, TM., Dunn, R., Suarez, C., Chiba, K. (Accepted). Chp. 2.5: Studying Ecosystems. In The Complete Dinosaur, 3rd Edition. Edited by L. Zanno, T. Holtz, and V. Arbour. Indiana University Press. [invited book chapter contribution]
18. Cullen, TM. (In Press). Stable isotopic analyses of living and extinct crocodylians: implications for understanding their ecology, environments, and physiology. In Crocodylian Biology and Paleobiology. Edited by H Woodward and J Farlow. Indiana University Press. [invited book chapter contribution]
17. Cullen, TM., Zanno, L., Larson, DW., Todd, E., Currie, PJ., Evans, DC. (2021). Anatomical, morphometric, and stratigraphic review of theropod biodiversity in the Dinosaur Park Formation (Late Cretaceous; Campanian) of Alberta. Canadian Journal of Earth Sciences 58(9): 870-884.
16. Cullen, TM., Canale, JI., Apesteguía, S., Smith, NS., Hu, D., Makovicky, PJ. (2020). Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proceedings of the Royal Society B 287: 20202258. http://dx.doi.org/10.1098/rspb.2020.2258.
15. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Huang, L., Fanti, F., Goodwin, MB., Ryan, MJ., Evans, DC. (2020). Large-scale stable isotope characterization of a Late Cretaceous dinosaur-dominated ecosystem. Geology 48(6): 546-551. (Featured Article & subject of GSA blog)
14. Cullen, TM., Simon, DJ., Benner, EKC., Evans, DC. (2020). Morphology and osteohistology of a large-bodied caenagnathid (Theropoda: Oviraptorosauria) from the Hell Creek Formation (Montana): implications for size-based classifications and growth estimation in theropods. Papers in Palaeontology. https://doi.org/10.1002/spp2/1302
13. Arbour, VM., Evans, DC., Simon, DJ., Cullen, TM., Braman, D. (2019). Cretaceous flora and fauna of the Sustut Group near the Sustut River, northern British Columbia, Canada. Canadian Journal of Earth Sciences. https://doi.org/10.1139/cjes-2019-0031.
12. Cullen, TM., Longstaffe, FJ., Wortmann, UG., Goodwin, MB., Huang, L., and Evans, DC. (2019). Stable isotopic characterization of a coastal floodplain forest community: a case-study for isotopic reconstruction of Mesozoic vertebrate assemblages. Royal Society Open Science 6: 181210. http://dx.doi.org/10.1098/rsos.181210.
11. McFeeters, B., Ryan, MJ., and Cullen, TM. (2018). Response to Brownstein (2018) 'Rebuttal of McFeeters, Ryan and Cullen, 2018'. Vertebrate Anatomy Morphology Palaeontology 6: 73-74. dx.doi.org/10.18435/vamp29343
10. McFeeters, B., Ryan, MJ., and Cullen, TM. (2018). Positional variation in pedal unguals of North American ornithomimids (Dinosauria, Theropoda): a response to Brownstein (2017) . Vertebrate Anatomy Morphology Palaeontology 5. doi: 10.18435/vamp29283
9. LeBlanc, ARH., Brink, K., Cullen, TM., and Reisz, R. (2017). Evolutionary implications of tooth attachment versus tooth implantation: a case study using dinosaur, crocodilian, and mammal teeth. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2017.1354006
8. Evans, DC., Cullen, TM., Larson, DW., and Rego, A. (2017). A new species of troodontid theropod (Dinosauria: Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta, Canada. Canadian Journal of Earth Sciences. doi: 10.1139/cjes-2017-0034 (Postprint available on PaleorXiv: https://paleorxiv.org/b8mqe/)
7. Cullen, TM. and Evans, DC. (2016). Palaeoenvironmental drivers of vertebrate community composition in the Belly River Group (Campanian) of Alberta, Canada, with implications for dinosaur biogeography. BMC Ecology. doi: 10.1186/s12898-016-0106-8 (Editor's Pick and BMC Ecology top 10 highlight for 2016, and subject of a BMC Series blog)
6. McFeeters, B., Ryan, MJ., Schröder-Adams, C., and Cullen, TM. (2016). A new ornithomimid theropod from the Dinosaur Park Formation of Alberta, Canada. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2016.1221415
5. Cullen, TM., Fanti, F., Capobianco, C., Ryan, MJ., and Evans, DC. (2016). A vertebrate microsite from a marine-terrestrial transition in the Foremost Formation (Campanian) of Alberta, Canada, and the use of faunal assemblage data as a palaeoenvironmental indicator. Palaeogeography, Palaeoclimatology, Palaeoecology 444: 101-114. doi:10.1016/j.palaeo.2015.12.015 (Postprint available on PaleorXiv: https://paleorxiv.org/43j7z/)
4. Cullen, TM., Ryan, MJ., Currie, PJ., Kobayashi, Y., and Evans, DC. (2014). Osteohistological variation in growth marks and osteocyte lacunar density in a theropod dinosaur (Coelurosauria: Ornithomimidae). BMC Evolutionary Biology. doi: 10.1186/s12862-014-0231-y. (Editor’s Pick and a BMC series highlight for November 2014, and included in list of the 12 best figures in BMC Evolutionary Biology for 2014)
3. Evans, DC., Larson, DW., Cullen, TM., & Sullivan, RM. (2014). "Saurornitholestes" robustus is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences 51(7): 730-734. doi: 10.1139/cjes-2014-0073
2. Cullen, TM., Fraser, D., Rybczynski, N., and Schroder-Adams, C. (2014). Early evolution of sexual dimorphism and polygyny in Pinnipedia. Evolution 68-5: 1469–1484. (Subject of the May 2014 cover of Evolution) doi: https://dx.doi.org/10.1111/evo.12360 (Postprint available on PaleorXiv: https://paleorxiv.org/zgh62/)
1. Cullen TM., Ryan MJ, Schröder-Adams C, Currie PJ, Kobayashi Y. (2013). An Ornithomimid (Dinosauria) Bonebed from the Late Cretaceous of Alberta, with Implications for the Behavior, Classification, and Stratigraphy of North American Ornithomimids. PLoS ONE 8(3): e58853. doi:10.1371/journal.pone.0058853
Non-Refereed Publications
4. Cullen, T. January 10, 2017. Dinosaur distributions in the Belly River. Blog for BMC Series blog. http://blogs.biomedcentral.com/bmcseriesblog/2017/01/10/dinosaur-distributions-belly-river/
3. Cullen, T. July 9, 2016. Microsites and Macroinferences. Blog for 'Valley of the Last Dinosaurs' project of Denver Museum of Nature & Science and the Howard Hughes Medical Institute. http://lastdinos.livesci.org/field-notebook/7/9/microsites-and-macroinferences
2. Cullen, T. July 7, 2016. Field Camps in Many Shapes and Sizes. Blog for 'Valley of the Last Dinosaurs' project of Denver Museum of Nature & Science and the Howard Hughes Medical Institute. http://lastdinos.livesci.org/field-notebook/7/7/field-camps-in-many-shapes-and-sizes
1. Cullen, T. November 19, 2014. Photography in the Field: equal parts business & pleasure. Royal Ontario Museum blogs. https://www.rom.on.ca/en/blog/photography-in-the-field-equal-parts-business-pleasure
3. Cullen, T. July 9, 2016. Microsites and Macroinferences. Blog for 'Valley of the Last Dinosaurs' project of Denver Museum of Nature & Science and the Howard Hughes Medical Institute. http://lastdinos.livesci.org/field-notebook/7/9/microsites-and-macroinferences
2. Cullen, T. July 7, 2016. Field Camps in Many Shapes and Sizes. Blog for 'Valley of the Last Dinosaurs' project of Denver Museum of Nature & Science and the Howard Hughes Medical Institute. http://lastdinos.livesci.org/field-notebook/7/7/field-camps-in-many-shapes-and-sizes
1. Cullen, T. November 19, 2014. Photography in the Field: equal parts business & pleasure. Royal Ontario Museum blogs. https://www.rom.on.ca/en/blog/photography-in-the-field-equal-parts-business-pleasure
Funding, Support, and Collaboration
I would like to thank the following agencies and institutions for their funding and/or support of my research (detailed amounts and further information in my CV PDF):
Natural Sciences and Engineering Research Council of Canada (Alexander Graham Bell Canada Graduate Scholarships & Postdoctoral Fellowship)
Government of Ontario (Ontario Graduate Scholarship; Queen Elizabeth II Scholarship for Science and Technology)
Dinosaur Research Institute (project grants, travel funding)
Carleton University (internal scholarships and RA support)
University of Toronto (internal scholarships)
University of Western Ontario Laboratory for Stable Isotope Science (support, training, and use of analytical equipment)
Canadian Museum of Nature (access to collections, permissions to destructively sample materials)
Royal Ontario Museum (access to collections, use of laboratory facilities, permissions to destructively sample materials)
Royal Tyrrell Museum (access to collections, permissions to destructively sample materials)
Field Museum of Natural History (access to collections, permissions to destructively sample materials)
Kenneth C. Griffin Charitable Fund (support for FMNH postdoctoral position)
Friends of the North Carolina Museum of Natural Sciences (support for NCMNS / NCSU postdoctoral position)
Government of Ontario (Ontario Graduate Scholarship; Queen Elizabeth II Scholarship for Science and Technology)
Dinosaur Research Institute (project grants, travel funding)
Carleton University (internal scholarships and RA support)
University of Toronto (internal scholarships)
University of Western Ontario Laboratory for Stable Isotope Science (support, training, and use of analytical equipment)
Canadian Museum of Nature (access to collections, permissions to destructively sample materials)
Royal Ontario Museum (access to collections, use of laboratory facilities, permissions to destructively sample materials)
Royal Tyrrell Museum (access to collections, permissions to destructively sample materials)
Field Museum of Natural History (access to collections, permissions to destructively sample materials)
Kenneth C. Griffin Charitable Fund (support for FMNH postdoctoral position)
Friends of the North Carolina Museum of Natural Sciences (support for NCMNS / NCSU postdoctoral position)