Lyson, T. R. et al. Fossorial origin of the turtle shell. Current Biology 26, 1887–1894 (2016).
Shaffer, H. B., McCartney-Melstad, E., Near, T. J., Mount, G. G. & Spinks, P. Q. Phylogenomic analyses of 539 highly informative loci dates a fully resolved time tree for the major clades of living turtles (Testudines). Molecular Phylogenetics and Evolution 115, 7–15 (2017).
Angielczyk, K. D., Burroughs, R. W. & Feldman, C. R. Do turtles follow the rules? Latitudinal gradients in species richness, body size, and geographic range area of the world’s turtles. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 324, 270–294 (2015).
Lovich, J. E., Ennen, J. R., Agha, M. & Gibbons, J. W. Where have all the turtles gone, and why does it matter? BioScience 68, 771–791 (2018).
Gibbons, J. W. Why do turtles live so long? BioScience 37, 262–269 (1987).
Blake, S. et al. Seed dispersal by Galápagos tortoises. Journal of Biogeography 39, 1961–1972 (2012).
Heppell, S. S. Application of life-history theory and population model analysis to turtle conservation. Copeia 1998, 367–375 (1998).
Lutcavage, M. E. Human impacts on sea turtle survival. In Wyneken, J., Lohmann, K. J. & Musick J. A. (Eds.). The Biology of Sea Turtles Volume I, 387–409pp. (CRC Press, 2017).
Stanford, C. B. et al. Turtles and tortoises are in trouble. Current Biology 30, R721–R735 (2020).
Zangerl, R. The turtle shell. In Gans, C., Bellairs, A.d’A. & Parsons, T. S. (eds). Biology of the Reptilia Vol. 1 Morphology A, 311–339. (Academic Press, London, 1969).
Cherepanov, G. O. Nature of the turtle shell: morphogenetic causes of bone variability and its evolutionary implication. Paleontological Journal 50, 1641–1648 (2016).
Landberg, T., Mailhot, J. D. & Brainerd, E. L. Lung ventilation during treadmill locomotion in a terrestrial turtle, Terrapene carolina. Journal of Experimental Biology 206, 3391–3404 (2003).
Magwene, P. M. & Socha, J. J. Biomechanics of turtle shells: how whole shells fail in compression. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 319, 86–98 (2013).
Congdon, J. D., Dunham, A. E. & van Loben Sels, R. C. Delayed sexual maturity and demographics of Blanding’s turtles (Emydoidea blandingii): implications for conservation and management of long‐lived organisms. Conservation Biology 7, 826–833 (1993).
Congdon, J. D., Reding, D. M. & Bronikowski, A. M. Testing hypotheses of aging in long-lived painted turtles (Chrysemys picta). Experimental Gerontology 38, 765–772 (2003).
Lau, M. & Shi, H. T. Conservation and trade of terrestrial and freshwater turtles and tortoises in the People’s Republic of China. Chelonian Research Monographs 2, 30–38 (2000).
Eisemberg, C. C., Rose, M., Yaru, B. & Georges, A. Demonstrating decline of an iconic species under sustained indigenous harvest — the pig-nosed turtle (Carettochelys insculpta) in Papua New Guinea. Biological Conservation 144, 2282–2288 (2011).
Sigouin, A. et al. Priorities for the trade of less charismatic freshwater turtle and tortoise species. Journal of Applied Ecology 54, 345–350 (2017).
Schneider, L., Ferrara, C. R., Vogt, R. C. & Schaffer, C. Subsistence-level chelonian exploitation on the Rio Negro and one viable alternative. Chelonian Conservation and Biology 15, 36–42 (2016).
Krivoruchko, A. & Storey, K. B. Turtle anoxia tolerance: biochemistry and gene regulation. Biochimica et Biophysica Acta (BBA)-General Subjects 1850, 1188–1196 (2015).
Luiselli, L., Starita, A., Carpaneto, G. M., Segniagbeto, G. H. & Amori, G. A short review of the international trade of wild tortoises and freshwater turtles across the world and throughout two decades. Chelonian Conservation and Biology 15, 167–172 (2016).
IUCN. The IUCN Red List of Threatened Species. https://www.iucnredlist.org (2024).
Hoffmann, M. et al. The impact of conservation on the status of the world’s vertebrates. Science 330, 1503–1509 (2010).
Cox, N. et al. A global reptile assessment highlights shared conservation needs of tetrapods. Nature 605, 285–290 (2022).
Cadotte, M. W. The new diversity: management gains through insights into the functional diversity of communities. Journal of Applied Ecology 48, 1067–1069 (2011).
Devictor, V. et al. Spatial mismatch and congruence between taxonomic, phylogenetic and functional diversity: the need for integrative conservation strategies in a changing world. Ecology Letters 13, 1030–1040 (2010).
Tilman, D. et al. Future threats to biodiversity and pathways to their prevention. Nature 546, 73–81 (2017).
Jones, K. E. et al. PanTHERIA: a species‐level database of life history, ecology, and geography of extant and recently extinct mammals. Ecology 90, 2648 (2009).
Myhrvold, N. P. et al. An amniote life‐history database to perform comparative analyses with birds, mammals, and reptiles. Ecology 96, 3109 (2015).
Vancine, M. H. et al. ATLANTIC AMPHIBIANS: a data set of amphibian communities from the Atlantic Forests of South America. Ecology 99, 1692 (2018).
Wang, Y. et al. A dataset on the life-history and ecological traits of Chinese birds. Biodiversity Science 29, 1149–1153 (2021).
Meiri, S. SquamBase—A database of squamate (Reptilia: Squamata) traits. Global Ecology and Biogeography 33, e13812 (2024).
Itescu, Y., Karraker, N. E., Raia, P., Pritchard, P. C. & Meiri, S. Is the island rule general? Turtles disagree. Global Ecology and Biogeography 23, 689–700 (2014).
Regis, K. W. & Meik, J. M. Allometry of sexual size dimorphism in turtles: a comparison of mass and length data. PeerJ 5, e2914 (2017).
Colston, T. J., Kulkarni, P., Jetz, W. & Pyron, R. A. Phylogenetic and spatial distribution of evolutionary diversification, isolation, and threat in turtles and crocodilians (non-avian archosauromorphs). BMC Evolutionary Biology 20, 1–16 (2020).
Rodríguez-Caro, R. C. et al. Anthropogenic impacts on threatened species erode functional diversity in chelonians and crocodilians. Nature Communications 14, 1542 (2023).
Oskyrko, O., Mi, C., Meiri, S. & Du, W. ReptTraits: a comprehensive dataset of ecological traits in reptiles. Scientific Data 11, 243 (2024).
Meiri, S. et al. Different solutions lead to similar life history traits across the great divides of the amniote tree of life. Journal of Biological Research-Thessaloniki 28, 1–17 (2021).
Lovich, J. E. & Gibbons, W. Turtles of the world: A guide to every family. (Princeton University Press, New Jersey, 2021).
Swingland, I. R. & Klemens, M. W. The conservation biology of tortoises Vol. 5 (IUCN, 1989).
Uetz, P. The reptile database. https://reptile-database.reptarium.cz/ (2024).
Turtle Taxonomy Working Group (TTWG). (Rhodin, A. G. K., et al). Turtles of the World Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (9th Ed.). Chelonian Research Monographs 8, 1–472 (2021).
Gong, S., Fritz, U., Vamberger, M., Gao, Y. & Farkas, B. Disentangling the Pelodiscus axenaria complex, with the description of a new Chinese species and neotype designation for P. axenaria (Zhou, Zhang & Fang, 1991). Zootaxa 5125, 131–143 (2022).
Xiao, F., Lin, Z., Wang, J. & Shi, H. T. Shell shape-habitat correlations in extant turtles: A global-scale analysis. Global Ecology and Conservation 46, e02543 (2023).
Rhodin, A. G. et al. Global conservation status of turtles and tortoises (order Testudines). Chelonian Conservation and Biology 17, 135–161 (2018).
Myers, P. et al. The Animal Diversity Web (online). https://animaldiversity.org (2023).
Smith, G. R. & Iverson, J. B. Diel activity patterns of the turtle assemblage of a northern Indiana lake. The American Midland Naturalist 152, 156–164 (2004).
Butterfield, T. G., Scoville, A., García, A. & Beck, D. D. Habitat use and activity patterns of a terrestrial turtle (Rhinoclemmys rubida perixantha) in a seasonally dry tropical forest. Herpetologica 74, 226–235 (2018).
Hjort Toms, A., Browning, L. V. T., Paterson, J. E., Angoh, S. Y. J. & Davy, C. M. Night moves: nocturnal movements of endangered spotted turtles and Blanding’s turtles. Journal of Zoology 316, 40–48 (2022).
Djordjevic, S. et al. Sexual body size and body shape dimorphism of Testudo hermanni in central and eastern Serbia. Amphibia-Reptilia 32, 445–458 (2011).
Pritchard, P. C. H. Encyclopedia of turtles. (TFH Publications, New Jersey, 1979).
Georges, A., Guarino, F. & Bito, B. Freshwater turtles of the TransFly region of Papua New Guinea: notes on diversity, distribution, reproduction, harvest and trade. Wildlife Research 33, 373–384 (2006).
Nemesházi, E. & Bókony, V. HerpSexDet: the herpetological database of sex determination and sex reversal. Scientific Data 10, 377 (2023).
Olson, D. M. et al. Terrestrial ecoregions of the world: a new map of life on earth. BioScience 51, 933–938 (2001).
CITES Trade Database https://trade.cites.org/ (2024).
NCBI Sequence Read Archive https://www.ncbi.nlm.nih.gov/ (2024).
Wang, J. et al. CheloniansTraits: a comprehensive trait database of global turtles and tortoises, figshare, https://doi.org/10.6084/m9.figshare.28828241 (2025).
