• Simpson, G. G. Major Features of Evolution (Columbia Univ. Press, 1953).

  • Simpson, G. G. Tempo and Mode in Evolution (Columbia Univ. Press, 1944).

  • Vermeij, G. J. Adaptation, versatility, and evolution. Syst. Zool. 22, 466–477 (1973).

    Article 

    Google Scholar
     

  • Dumont, E. R. et al. Morphological innovation, diversification and invasion of a new adaptive zone. Proc. R. Soc. B 279, 1797–1805 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Hunter, J. P. & Jernvall, J. The hypocone as a key innovation in mammalian evolution. Proc. Natl Acad. Sci. USA 92, 10718–10722 (1995).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Mitter, C., Farrell, B. & Wiegmann, B. The phylogenetic study of adaptive zones: has phytophagy promoted insect diversification? Am. Nat. 132, 107–128 (1988).

  • Liem, K. F. Evolutionary strategies and morphological innovations: cichlid pharyngeal jaws. Syst. Zool. 22, 425–441 (1973).

    Article 

    Google Scholar
     

  • Van Valkenburgh, B. Déjà vu: the evolution of feeding morphologies in the Carnivora. Integr. Comp. Biol. 47, 147–163 (2007).

    Article 
    PubMed 

    Google Scholar
     

  • Martin, L. D. In Carnivore Behavior, Ecology, and Evolution (ed Gittleman, J. L.) 536–568 (Cornell Univ. Press, 1989).

  • Werdelin, L. In Carnivore Behavior, Ecology, and Evolution (ed Gittleman, J. L.) 582–624 (Cornell Univ. Press, 1996).

  • Polly, P. D. In Mammalian Evolutionary Morphology (eds Sargis, E. J. & Dagosto, M.) 167–196 (Springer, 2008).

  • Slater, G. J. Topographically distinct adaptive landscapes for teeth, skeletons, and size explain the adaptive radiation of Carnivora (Mammalia). Evolution 76, 2049–2066 (2022).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Tseng, Z. J. Testing adaptive hypotheses of convergence with functional landscapes: a case study of bone-cracking hypercarnivores. PLOS ONE 8, e65305 (2013).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Balisi, M. A. & Van Valkenburgh, B. Iterative evolution of large-bodied hypercarnivory in canids benefits species but not clades. Commun. Biol. 3, 461 (2020).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Rasmussen, G. S. A., Gusset, M., Courchamp, F. & Macdonald, D. W. Achilles’ heel of sociality revealed by energetic poverty trap in cursorial hunters. Am. Nat. 172, 508–518 (2008).

    Article 
    PubMed 

    Google Scholar
     

  • Holliday, J. A. & Steppan, S. J. Evolution of hypercarnivory: the effect of specialization on morphological and taxonomic diversity. Paleobiology 30, 108–128 (2004).

    Article 

    Google Scholar
     

  • Van Valkenburgh, B., Wang, X. & Damuth, J. Cope’s rule, hypercarnivory, and extinction in North American canids. Science 306, 101–104 (2004).

    Article 
    PubMed 

    Google Scholar
     

  • Barrett, P. Z., Hopkins, S. S. B. & Price, S. A. How many sabretooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae. J. Vertebr. Paleontol. 41, e1923523 (2021).

  • Coca-Ortega, C. & Pérez-Claros, J. A. Characterizing ecomorphological patterns in hyenids: a multivariate approach using postcanine dentition. PeerJ 6, e6238 (2019).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Tarquini, S. D., Chemisquy, M. A. & Prevosti, F. J. Evolution of the carnassial in living mammalian carnivores (Carnivora, Didelphimorphia, Dasyuromorphia): diet, phylogeny, and allometry. J. Mamm. Evol. 27, 95–109 (2020).

    Article 

    Google Scholar
     

  • Hansen, T. F. Stabilizing selection and the comparative analysis of adaptation. Evolution 51, 1341–1351 (1997).

    Article 
    PubMed 

    Google Scholar
     

  • McNab, B. K. The influence of food habits on the energetics of eutherian mammals. Ecol. Monogr. 56, 1–19 (1986).

    Article 

    Google Scholar
     

  • Friscia, A. R., Van Valkenburgh, B. & Biknevicius, A. R. An ecomorphological analysis of extant small carnivorans. J. Zool. 272, 82–100 (2007).

    Article 

    Google Scholar
     

  • Sacco, T. & Van Valkenburgh, B. Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). J. Zool. 263, 41–54 (2004).

    Article 

    Google Scholar
     

  • Van Valkenburgh, B. & Koepfli, K.-P. Cranial and dental adaptations to predation in canids. Symp. Zool. Soc. Lond. 65, 15–37 (1993).


    Google Scholar
     

  • Van Valkenburgh, B. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology 17, 340–362 (1991).

    Article 

    Google Scholar
     

  • Meachen-Samuels, J. & Van Valkenburgh, B. Craniodental indicators of prey size preference in the Felidae. Biol. J. Linn. Soc. 96, 784–799 (2009).

    Article 

    Google Scholar
     

  • Hopkins, S. S. B., Price, S. A. & Chiono, A. J. Influence of phylogeny on the estimation of diet from dental morphology in the Carnivora. Paleobiology 48, 324–339 (2022).

  • Waldman, E., Gonzalez, Y., Flynn, J. J. & Tseng, Z. J. Dental topographic proxies for ecological characteristics in carnivoran mammals. J. Anat. 242, 627–641 (2023).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Pineda-Munoz, S., Lazagabaster, I. A., Alroy, J. & Evans, A. R. Inferring diet from dental morphology in terrestrial mammals. Methods Ecol. Evol. 8, 481–491 (2017).

    Article 

    Google Scholar
     

  • Barrett, P. Z. & Hopkins, S. S. B. Adaptive zones of feliforms and evolutionary regimes within terrestrial mammalian carnivores. figshare https://doi.org/10.6084/m9.figshare.30689975 (2026).

  • Slater, G. J., Harmon, L. J. & Alfaro, M. E. Integrating fossils with molecular phylogenies improves inference of trait evolution. Evolution 66, 3931–3944 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Harmon, L. J. et al. Early bursts of body size and shape evolution are rare in comparative data. Evolution 64, 2385–2396 (2010).

    PubMed 

    Google Scholar
     

  • Slater, G. J. Iterative adaptive radiations of fossil canids show no evidence for diversity-dependent trait evolution. Proc. Natl Acad. Sci. USA 112, 4897–4902 (2015).

    Article 
    CAS 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Barrett, P. Z. & Hopkins, S. S. B. Mosaic evolution underlies feliform morphological disparity. Proc. R. Soc. B 291, 20240756 (2024).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Slater, G. J. & Friscia, A. R. Hierarchy in adaptive radiation: a case study using the Carnivora (Mammalia). Evolution 73, 524–539 (2019).

    Article 
    PubMed 

    Google Scholar
     

  • Carbone, C., Teacher, A. & Rowcliffe, J. M. The costs of carnivory. PLOS Biol. 5, 363–368 (2007).

    Article 
    CAS 

    Google Scholar
     

  • Carbone, C., Mace, G. M., Roberts, S. C. & Macdonald, D. W. Energetic constraints on the diet of terrestrial carnivores. Nature 402, 286–288 (1999).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Van Valen, L. A new evolutionary law. Evol. Theory 1, 1–30 (1973).


    Google Scholar
     

  • Figueirido, B. et al. Constraint and adaptation in the evolution of carnivoran skull shape. Paleobiology 37, 490–518 (2011).

    Article 

    Google Scholar
     

  • Andersson, K. I. Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores. Zool. J. Linn. Soc. 142, 91–104 (2004).

    Article 

    Google Scholar
     

  • Andersson, K. & Werdelin, L. The evolution of cursorial carnivores in the Tertiary: implications of elbow-joint morphology. Proc. R. Soc. B 270, S163–S165 (2003).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Meachen-Samuels, J. & van Valkenburgh, B. Forelimb indicators of prey-size preference in the felidae. J. Morphol. 270, 729–744 (2009).

    Article 
    PubMed 

    Google Scholar
     

  • Figueirido, B., Martín-Serra, A., Tseng, Z. J. & Janis, C. M. Habitat changes and changing predatory habits in North American fossil canids. Nat. Commun. 6, 7976 (2015).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Fabre, A. C. et al. Getting a grip on the evolution of grasping in musteloid carnivorans: a three-dimensional analysis of forelimb shape. J. Evol. Biol. 26, 1521–1535 (2013).

    Article 
    PubMed 

    Google Scholar
     

  • Slater, G. J., Dumont, E. R. & Van Valkenburgh, B. Implications of predatory specialization for cranial form and function in canids. J. Zool. 278, 181–188 (2009).

    Article 

    Google Scholar
     

  • Tseng, Z. & Wang, X. Do convergent ecomorphs evolve through convergent morphological pathways? Cranial shape evolution in fossil hyaenids and borophagine canids (Carnivora, Mammalia). Paleobiology 37, 470–489 (2011).

    Article 

    Google Scholar
     

  • Figueirido, B., Tseng, Z. J. & Martín-Serra, A. Skull shape evolution in durophagous carnivorans. Evolution 67, 1975–1993 (2013).

    Article 
    PubMed 

    Google Scholar
     

  • Tseng, Z. J. Connecting Hunter–Schreger Band microstructure to enamel microwear features: new insights from durophagous carnivores. Acta Palaeontol. Pol. 57, 473–484 (2012).

    Article 

    Google Scholar
     

  • Penrose, F., Kemp, G. J. & Jeffery, N. Scaling and accommodation of jaw adductor muscles in Canidae. Anat. Rec. 299, 951–966 (2016).

    Article 

    Google Scholar
     

  • Werdelin, L., Yamaguchi, N., Johnson, W. E. & O’Brien, S. J. In Biology and Conservation of Wild Felids (eds Macdonald, D. W. & Loveridge, A. J.) 59–82 (Oxford Univ. Press, 2010).

  • Salesa, M. J., Gamarra, Je., Siliceo, G., Antón, M. & Morales, J. Unraveling the diversity of early felines: a new genus of Felinae (Carnivora, Felidae) from the Middle Miocene of Madrid (Spain). J. Vertebr. Paleontol. https://doi.org/10.1080/02724634.2023.2288924 (2023).

  • Robles, J. M. et al. New Pseudaelurus and Styriofelis remains (Carnivora: Felidae) from the Middle Miocene of Abocador de Can Mata (Vallès-Penedès Basin). C. R. Palevol. 12, 101–113 (2013).

    Article 

    Google Scholar
     

  • Poust, A. W., Barrett, P. Z. & Tomiya, S. An early nimravid from California and the rise of hypercarnivorous mammals after the middle Eocene climatic optimum. Biol. Lett. 18, 20220291 (2022).

    Article 
    PubMed Central 

    Google Scholar
     

  • Hawkins, C. E. & Racey, P. A. Food habits of an endangered Carnivore, Cryptoprocta ferox, in the dry deciduous forests of western Madagascar. J. Mammal. 89, 64 (2008).

    Article 

    Google Scholar
     

  • Sunquist, M. & Sunquist, F. Wild Cats of the World (Univ. Chicago Press, 2002).

  • Christiansen, P. Evolutionary convergence of primitive sabertooth craniomandibular morphology: the clouded leopard (Neofelis nebulosa) and Paramachairodus ogygia compared. J. Mamm. Evol. 15, 155–179 (2008).

    Article 

    Google Scholar
     

  • Law, C. J. et al. Effects of diet on cranial morphology and biting ability in musteloid mammals. J. Evol. Biol. 31, 1918–1931 (2018).

    Article 
    PubMed 

    Google Scholar
     

  • Slater, G. J. & Van Valkenburgh, B. Allometry and performance: the evolution of skull form and function in felids. J. Evol. Biol. 22, 2278–2287 (2009).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Chatar, N., Fischer, V. & Tseng, Z. J. Many-to-one function of cat-like mandibles highlights a continuum of sabre-tooth adaptations. Proc. R. Soc. B 289, 20221627 (2022).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Chatar, N., Michaud, M., Tamagnini, D. & Fischer, V. Evolutionary patterns of cat-like carnivorans unveil drivers of the sabertooth morphology. Curr. Biol. 34, 2460–2473 (2024).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Figueirido, B., Tucker, S. & Lautenschlager, S. Comparing cranial biomechanics between Barbourofelis fricki and Smilodon fatalis: is there a universal killing-bite among saber-toothed predators? Anat. Rec. https://doi.org/10.1002/ar.25451 (2024).

  • Price, S. A. & Hopkins, S. S. B. The macroevolutionary relationship between diet and body mass across mammals. Biol. J. Linn. Soc. 115, 173–184 (2015).

    Article 

    Google Scholar
     

  • Christiansen, P. What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)?. Ann. Zool. Fennici 36, 93–102 (1999).


    Google Scholar
     

  • Soibelzon, L. H. & Schubert, B. W. The largest known bear, Arctotherium angustidens, from the early pLeistocene Pampean region of Argentina: with a discussion of size and diet trends in bears. J. Paleontol. 85, 69–75 (2011).

    Article 

    Google Scholar
     

  • Sorkin, B. Ecomorphology of the giant bear-dogs Amphicyon and Ischyrocyon. Hist. Biol. 18, 375–388 (2006).

    Article 

    Google Scholar
     

  • Peigné, S., Salesa, M. J., Antón, M. & Morales, J. A new amphicyonine (Carnivora: Amphicyonidae) from the Upper Miocene of Batallones-1 Madrid, Spain. Palaeontology 51, 943–965 (2008).

    Article 

    Google Scholar
     

  • Morales, J., Abella, J., Sanisidro, O. & Valenciano, A. Ammitocyon kainos gen. et sp. nov., a chimerical amphicyonid (Mammalia, Carnivora) from the late Miocene carnivore traps of Cerro de los Batallones (Madrid, Spain). J. Syst. Palaeontol. 19, 393–415 (2021).

    Article 

    Google Scholar
     

  • Morales, J. & Pickford, M. Carnivores from the middle Miocene Ngorora formation (13-12 MA), Kenya. Estud. Geol. 61, 271–284 (2005).

    Article 

    Google Scholar
     

  • Morlo, M. et al. The apex of amphicyonid hypercarnivory: solving the riddle of Agnotherium antiquum Kaup, 1833 (Mammalia, Carnivora). J. Vertebr. Paleontol. 39, e1705848 (2019).

  • Müller, N. F. & Bouckaert, R. R. Adaptive metropolis-coupled MCMC for BEAST 2. PeerJ 8, e9473 (2020).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Bouckaert, R. et al. BEAST 2: a software platform for Bayesian evolutionary analysis. PLOS Comput. Biol. 10, e1003537 (2014).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Stadler, T., Künert, D., Bonhoeffer, S. & Drummond, A. J. Birth-death skyline plot reveals temporal changes of epidemic spread in HIV and hepatitis C virus (HCV). Proc. Natl Acad. Sci. USA 110, 228–233 (2013).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Gavryushkina, A., Welch, D., Stadler, T. & Drummond, A. J. Bayesian inference of sampled ancestor trees for epidemiology and fossil calibration. PLOS Comput. Biol. 10, e1003919 (2014).

    Article 
    PubMed 
    PubMed Central 

    Google Scholar
     

  • Jones, K. E. et al. PanTHERIA: a species-level database of life history, ecology, and geography of extant and recently extinct mammals. Ecology 90, 2648–2648 (2009).

  • Van Valkenburgh, B. In Body Size in Mammalian Paleobiology: Estimation and Biological Implications (eds Damuth, J. & MacFadden, B. J.) 181–205 (Cambridge Univ. Press, 1990).

  • Werdelin, L. & Solounias, N. The Hyaenidae: taxonomy, systematics and evolution. Fossils Strata 30, 104 (1991).


    Google Scholar
     

  • Christiansen, P. & Harris, J. M. Body size of Smilodon (Mammalia: Felidae). J. Morphol. 266, 369–384 (2005).

    Article 
    PubMed 

    Google Scholar
     

  • Wheeler, H. T. & Jefferson, G. T. Panthera atrox: body proportions, size, sexual dimorphism, and behavior of the cursorial lion of the North American Plains. Mus. North. Ariz. Bull. 65, 423–444 (2009).


    Google Scholar
     

  • Palmqvist, P. et al. The giant hyena Pachycrocuta brevirostris: modelling the bone-cracking behavior of an extinct carnivore. Quat. Int. 243, 61–79 (2011).

    Article 

    Google Scholar
     

  • Venables, W. N. & Ripley, B. D. Modern Applied Statistics with S (Springer, 2002).

  • R Core Team R: A Language and Environment for Statistical Computing (R Core Team, 2020).

  • Wilman, H. et al. EltonTraits 1.0: species-level foraging attributes of the world’s birds and mammals. Ecology 95, 2027 (2014).

    Article 

    Google Scholar
     

  • Pérez-Claros, J. A. An ecomorphological characterization of the percrocutoid hyaenids: a multivariate approach using postcanine dentition. J. Vertebr. Paleontol. 42, e2197972 (2022).

    Article 

    Google Scholar
     

  • Pennell, M. W. et al. Geiger v2.0: an expanded suite of methods for fitting macroevolutionary models to phylogenetic trees. Bioinformatics 30, 2216–2218 (2014).

    Article 
    CAS 
    PubMed 

    Google Scholar
     

  • Clavel, J., Escarguel, G. & Merceron, G. mvMORPH: an R package for fitting multivariate evolutionary models to morphometric data. Methods Ecol. Evol. 6, 1311–1319 (2015).

    Article 

    Google Scholar
     

  • Beaulieu, J. M., Jhwueng, D. C., Boettiger, C. & O’Meara, B. C. Modeling stabilizing selection: expanding the Ornstein–Uhlenbeck model of adaptive evolution. Evolution 66, 2369–2383 (2012).

    Article 
    PubMed 

    Google Scholar
     

  • Barrett, P. Data from: Mosaic evolution underlies feliform morphological disparity. Zenodo https://doi.org/10.5281/zenodo.11221809 (2024).

  • Revell, L. J. phytools: An R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3, 217–223 https://doi.org/10.1111/j.2041-210X.2011.00169.x (2012).