How many sabertooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae

LITERATURE CITED

• Akaike, H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19:716–723.
   Web of Science ®Google Scholar
• Albright, L. B., M. O. Woodburne, T. J. Fremd, C. C. Swisher, B. J. MacFadden, and G. R. Scott. 2008. Revised chronostratigraphy and biostratigraphy of the John Day formation (Turtle Cove and Kimberly Members), Oregon, with implications for updated calibration of the Arikareean North American land mammal age. Journal of Geology 116:211–237.
   Web of Science ®Google Scholar
• Andersson, K., D. Norman, and L. Werdelin. 2011. Sabretoothed carnivores and the killing of large prey. PLoS ONE 6:8–13.
   Web of Science ®Google Scholar
• Antoine, P. O., J. L. Welcomme, L. Marivaux, I. Baloch, M. Benammi, and P. Tassy. 2003. First record of Paleogene Elephantoidea (Mammalia, Proboscidea) from the Bugti Hills of Pakistan. Journal of Vertebrate Paleontology 23:977–980.
   Web of Science ®Google Scholar
• Antón, M. 2013. Sabertooth. Indiana University Press, Bloomington, IN, pp.
   Google Scholar
• Antón, M., and À. Galobart. 1999. Neck Function and Predatory Behavior in the Scimitar Toothed Cat Homotherium latidens (Owen). Journal of Vertebrate Paleontology 19:771–784.
   Web of Science ®Google Scholar
• Antón, M., M. J. Salesa, J. F. Pastor, I. M. Sánchez, S. Fraile, and J. Morales. 2004. Implications of the mastoid anatomy of larger extant felids for the evolution and predatory behaviour of sabretoothed cats (Mammalia, Carnivora, Felidae). Zoological Journal of the Linnean Society 140:207–221.
   Web of Science ®Google Scholar
• Averianov, A., E. Obraztsova, I. Danilov, P. Skutschas, and J. Jin. 2016. First nimravid skull from Asia. Scientific Reports 6:1–8.
   PubMed Web of Science ®Google Scholar
• Barnett, R., M. V. Westbury, M. Sandoval-Velasco, F. G. Vieira, S. Jeon, G. Zazula, M. D. Martin, S. Y. W. Ho, N. Mather, S. Gopalakrishnan, J. Ramos-Madrigal, M. de Manuel, M. L. Zepeda-Mendoza, A. Antunes, A. C. Baez, B. De Cahsan, G. Larson, S. J. O’Brien, E. Eizirik, W. E. Johnson, K.-P. Koepfli, A. Wilting, J. Fickel, L. Dalén, E. D. Lorenzen, T. Marques-Bonet, A. J. Hansen, G. Zhang, J. Bhak, N. Yamaguchi, and M. T. P. Gilbert. 2020. Genomic Adaptations and Evolutionary History of the Extinct Scimitar-Toothed Cat, Homotherium latidens. Current Biology 1–8.
   Web of Science ®Google Scholar
• Barnett, R., I. Barnes, M. J. Phillips, L. D. Martin, C. R. Harrington, J. A. Leonard, and A. Cooper. 2005. Evolution of the extinct Sabretooths and the American cheetah-like cat. Current Biology 15:R589–R590.
   PubMed Web of Science ®Google Scholar
• Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658.
   PubMed Web of Science ®Google Scholar
• Baskin, J. A. 1981. Barbourofelis (Nimravidae) and Nimravides (Felidae), with a Description of Two New Species from the Late Miocene of Florida. Journal of Mammalogy 62:122–139.
   Web of Science ®Google Scholar
• Baskin, J. A. 2005. Carnivora from the Late Miocene Love Bone Bed of Florida. Bulletin of the Florida Museum of Natural History 45:413–434.
   Google Scholar
• de Beaumont, G. 1964. Remarques sur la classification des Felidae. Eclogae Geologicae Helvetiae 57:837–845.
   Google Scholar
• Blainville, H. M. D. 1843. Ostéographie ou description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles pour servir de base à la zoologie et à la géologie. Mammifères. Carnassiers: des Felis. J. B. Baillière & Fils, Paris, 196 pp.
   Google Scholar
• Bouckaert, R., J. Heled, D. Künert, T. G. Vaughan, C.-H. Wu, D. Xie, M. A. Suchard, A. Rambaut, and A. J. Drummond. 2014. BEAST2: a software platform for Bayesian evolutionary analysis. PLoS Computational Biology 10:e1003537.
   PubMed Web of Science ®Google Scholar
• Brown, J. G. 2014. Jaw function in Smilodon fatalis: A reevaluation of the canine shear-bite and a proposal for a new forelimb-powered class 1 lever model. PLoS ONE 9:1–14.
   Web of Science ®Google Scholar
• Bryant, H. N. 1988. The anatomy, phylogenetic relationships and systematics of the Nimravidae (Mammalia: Carnivora). PhD Thesis, University of Toronto, 414 pp.
   Google Scholar
• Bryant, H. N. 1991. Phylogenetic relationships and systematics of the Nimravidae (Carnivora). Journal of Mammalogy 72:56–78.
   Web of Science ®Google Scholar
• Bryant, H. N. 1996. Nimravidae; pp. 453–475 in D. R. Prothero and R. J. Emry (eds.), The Terrestrial Eocene-Oligocene Transition in North America. Cambridge University Press, Cambridge [England]; New York.
   Google Scholar
• Christiansen, P. 2006. Sabertooth characters in the clouded leopard (Neofelis nebulosa Griffiths 1821). Journal of Morphology 267:1186–1198.
   PubMed Web of Science ®Google Scholar
• Christiansen, P. 2013. Phylogeny of the sabertoothed felids (Carnivora: Felidae: Machairodontinae). Cladistics 29:543–559.
   PubMed Web of Science ®Google Scholar
• Clarke, J. A., and K. M. Middleton. 2008. Mosaicism, modules, and the evolution of birds: results from a Bayesian approach to the study of morphological evolution using discrete character data. Systematic Biology 57:185–201.
   PubMed Web of Science ®Google Scholar
• Cope, E. D. 1878. On some of the characters of the Miocene fauna of Oregon. Proceedings of the American Philosophical Society 18:63–78.
   Google Scholar
• Didiera, G., M. Royer-Carenzib, and M. Laurinc. 2012. The reconstructed evolutionary process with the fossil record. Journal of Theoretical Biology 315:26–37.
   PubMed Web of Science ®Google Scholar
• Dupin, J., N. J. Matzke, T. Särkinen, S. Knapp, R. G. Olmstead, L. Bohs, and S. D. Smith. 2017. Bayesian estimation of the global biogeographical history of the Solanaceae. Journal of Biogeography 44:887–899.
   Web of Science ®Google Scholar
• Egi, N., T. Tsubamoto, M. Saneyoshi, K. Tsogtbaatar, M. Watabe, B. Mainbayar, T. Chinzorig, and P. Khatanbaatar. 2016. Taxonomic revisions on nimravids and small feliforms (Mammalia, Carnivora) from the Upper Eocene of Mongolia. Historical Biology 28:105–119.
   Web of Science ®Google Scholar
• Fan, Y., R. Wu, M. H. Chen, L. Kuo, and P. O. Lewis. 2011. Choosing among partition models in Bayesian phylogenetics. Molecular Biology and Evolution 28:523–532.
   PubMed Web of Science ®Google Scholar
• Filhol, H. 1890. Études sur les mammifères fossiles de Sansan. Bibliothèque Des Hautes Études Section Des Sciences Naturelles 37:1–319.
   Google Scholar
• Flower, W. H. 1869. On the value of the characters of the base of the cranium in the classification of the order Carnivora, and on the systematic position of Bassaris and other disputed forms. Proceedings of the Zoological Society of London 4–37.
   Google Scholar
• Flynn, J. J., and H. Galiano. 1982. Phylogeny of early Tertiary Carnivora, with a description of a new species of Protictis from the middle Eocene of northwestern Wyoming. American Museum Novitates 2725:1–64.
   Google Scholar
• Flynn, J. J., J. A. Finarelli, S. Zehr, J. Hsu, and M. A. Nedbal. 2005. Molecular phylogeny of the Carnivora (Mammalia): assessing the impact of increased sampling on resolving enigmatic relationships. Systematic Biology 54:317–337.
   PubMed Web of Science ®Google Scholar
• Gavryushkina, A., D. Welch, T. Stadler, and A. J. Drummond. 2014. Bayesian inference of sampled ancestor trees for epidemiology and fossil calibration. PLoS Computational Biology 10:e1003919.
   PubMed Web of Science ®Google Scholar
• Gavryushkina, A., T. A. Heath, D. T. Ksepka, T. Stadler, D. Welch, and A. J. Drummond. 2017. Bayesian total-evidence dating reveals the recent crown radiation of penguins. Systematic Biology 66:57–73.
   PubMed Web of Science ®Google Scholar
• Geraads, D., and E. Güleç. 1997. Relationships of Barbourofelis piveteaui (Ozansoy, 1965), a late Miocene nimravid (Carnivora, Mammalia) from Central Turkey. Journal of Vertebrate Paleontology 17:370–375.
   Web of Science ®Google Scholar
• Ginsburg, L. 1961. La faune des carnivores miocènes de Sansan (Gers). Mémoires Du Muséum National d’Histoire Naturelle, Sér. C - Sciences de La Terre 9:1–190.
   Google Scholar
• Goloboff, P. A., and S. A. Catalano. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32:221–238.
   PubMed Web of Science ®Google Scholar
• Goswami, A., J. B. Smaers, C. Soligo, and P. D. Polly. 2014. The macroevolutionary consequences of phenotypic integration: from development to deep time. Philosophical Transactions of the Royal Society B 369:20130254.
   PubMed Web of Science ®Google Scholar
• Griffin, R. H. 2018. btw: Run BayesTraitsV3 from R. R package version 2.0. http://www.randigriffin.com/projects/btw.html.
   Google Scholar
• Harzhauser, M., A. Kroh, O. Mandic, W. E. Piller, U. Göhlich, M. Reuter, and B. Berning. 2007. Biogeographic responses to geodynamics: a key study all around the Oligo-Miocene Tethyan Seaway. Zoologischer Anzeiger 246:241–256.
   Web of Science ®Google Scholar
• Hunt, R. M. 1974. The auditory bulla in Carnivora: an anatomical basis for reappraisal of carnivore evolution. Journal of Morphology 143:21–75.
   PubMed Web of Science ®Google Scholar
• Hunt, R. M. 1987. Evolution of the aeluroid Carnivora: significance of auditory structure in the nimravid cat Dinictis. American Museum Novitates 2886:1–74.
   Google Scholar
• Hunt, R. M. 1989. Evolution of the aeluroid Carnivora: significance of the ventral promontorial process of the petrosal, and the origin of basicranial patterns in the living families. American Museum Novitates 2930:1–32.
   Google Scholar
• Hunt, R. M. 1998. Evolution of the aeluroid Carnivora: diversity of the earliest aeluroids from Eurasia (Quercy, Hsanda-Gol) and the origin of felids. American Museum Novitates 3252:1–65.
   Google Scholar
• Hunt, R. M. 2001. Basicranial anatomy of the living linsangs Prionodon and Poiana (Mammalia, Carnivora, Viverridae), with comments on the early evolution of aeluroid carnivorans. American Museum Novitates 3330:1–24.
   Google Scholar
• Ivanoff, D. V. 2019. Composition of the canid auditory bulla and a new look at the evolution of carnivoran entotympanics. Organisms Diversity and Evolution 19:363–375.
   Google Scholar
• Ivanoff, D. V. 2001. Partitions in the carnivoran auditory bulla: their formation and significance for systematics. Mammal Review 31:1–16.
   Web of Science ®Google Scholar
• Joeckel, R. M., S. Peigné, R. M. Hunt Jr., and R. I. Skolnick. 2002. The auditory region and nasal cavity of Oligocene Nimravidae (Mammalia: Carnivora). Journal of Vertebrate Paleontology 22:830–847.
   Web of Science ®Google Scholar
• Kass, R. E., and A. E. Raftery. 1995. Bayes Factors. Journal of the American Statistical Association 90:773–795.
   Web of Science ®Google Scholar
• Katoh, K., J. Rozewicki, and K. D. Yamada. 2019. MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20:1160–1166.
   PubMed Web of Science ®Google Scholar
• Landis, M. J., N. J. Matzke, B. R. Moore, and J. P. Huelsenbeck. 2013. Bayesian analysis of biogeography when the number of areas is large. Systematic Biology 62:789–804.
   PubMed Web of Science ®Google Scholar
• Lanfear, R., X. Hua, and D. L. Warren. 2016a. Estimating the effective sample size of tree topologies from Bayesian phylogenetic analyses. Genome Biology and Evolution 8:2319–2332.
   PubMed Web of Science ®Google Scholar
• Lanfear, R., P. B. Frandsen, A. M. Wright, T. Senfeld, and B. Calcott. 2016b. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses . Molecular Biology and Evolution 34:772–773.
   Web of Science ®Google Scholar
• Lee, M., P. Oliver, and M. Hutchinson. 2009. Phylogenetic uncertainty and molecular clock calibrations: a case study of legless lizards (Pygopodidae, Gekkota). Molecular Phylogenetics and Evolution 50:661–666.
   PubMed Web of Science ®Google Scholar
• Lewis, P. O. 2001. A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology 50:913–925.
   PubMed Web of Science ®Google Scholar
• Liow, L. H., and J. A. Finarelli. 2014. A dynamic global equilibrium in carnivoran diversification over 20 million years. Proceedings of the Royal Society B: Biological Sciences 281:20132312.
   PubMed Web of Science ®Google Scholar
• Lucas, S. G., and O. G. Bendukidze. 1997. Proboscidea (Mammalia) from the Early Miocene of Kazakhstan. Neues Jahrbuch Fur Geologie Und Palaontologie - Monatshefte 4:659–673.
   Google Scholar
• Martín-Serra, A., B. Figueirido, and P. Palmqvist. 2017. Non-decoupled morphological evolution of the fore- and hindlimb of sabretooth predators. Journal of Anatomy 231:532–542.
   PubMed Web of Science ®Google Scholar
• Martin, L. D. 1980. Functional morphology and the evolution of cats. Transactions of the Nebraska Academy of Sciences 8:141–154.
   Google Scholar
• Martin, L. D. 1998a. Nimravidae; pp. 228–235 in C. M. Janis, K. M. Scott, and L. L. Jacobs (eds.), Evolution of Tertiary Mammals of North America: Volume 1. Cambridge University Press, Cambridge, UK.
   Google Scholar
• Martin, L. D. 1998b. Felidae; pp. 236–242 in C. M. Janis (ed.), Evolution of Tertiary Mammals of North America. Cambridge University Press, Cambridge, UK.
   Google Scholar
• Matzke, N. J. 2013. Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing. Frontiers of Biogeography 5.
   PubMedGoogle Scholar
• Matzke, N. J. 2014. Model selection in historical biogeography reveals that founder-event speciation is a crucial process in island clades. Systematic Biology 63:951–970.
   PubMed Web of Science ®Google Scholar
• Matzke, N. J. 2015. BioGeoBEARS example script. PhyloWiki BioGeoBEARS website. Available at http://phylo.wikidot.com/biogeobears#script.
   Google Scholar
• Matzke, N. J., and A. Wright. 2016. Inferring node dates from tip dates in fossil Canidae: The importance of tree priors. Biology Letters 12.
   Web of Science ®Google Scholar
• McKenna, M. C., and S. K. Bell. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, pp.
   Google Scholar
• Meachen-Samuels, J. A. 2012. Morphological convergence of the prey-killing arsenal of sabertooth predators. Paleobiology 38:1–14.
   Web of Science ®Google Scholar
• Meachen-Samuels, J. A., and B. Van Valkenburgh. 2010. Radiographs reveal exceptional forelimb strength in the sabertooth cat, Smilodon fatalis. PLoS ONE 5:e11412.
   PubMed Web of Science ®Google Scholar
• Meade, A., and M. Pagel. 2019. BayesTraits V3.0.2. Available at http://www.evolution.rdg.ac.uk/BayesTraitsV3.0.2/BayesTraitsV3.0.2.html.
   Google Scholar
• Miller, M. A., W. Pfeiffer, and T. Schwartz. 2011. The CIPRES science gateway: a community resource for phylogenetic analyses. TG ‘11: Proceedings of the 2011 TeraGrid Conference: Extreme Digital Discovery 1–8.
   Google Scholar
• Morales, J., and M. Pickford. 2018. A new barbourofelid mandible (Carnivora, Mammalia) from the Early Miocene of Grillental-6, Sperrgebiet, Namibia. Communications of the Geological Survey of Namibia 18:113–123.
   Google Scholar
• Morales, J., M. Pickford, M. Salesa, and D. Soria. 2000. The systematic status of Kelba, Savage, 1965, Kenyalutra, Schmidt-Kittler, 1987 and Ndamathaia, Jacobs et al., 1987, (Viverridae, Mammalia) and a review of Early Miocene mongoose-like carnivores of Africa. Annales de Paléontologie 86:243–251.
   Google Scholar
• Morales, J., M. J. Salesa, M. Pickford, and D. Soria. 2001. A new tribe, new genus and two new species of Barbourofelinae (Felidae, Carnivora, Mammalia) from the early Miocene of East Africa and Spain. Transactions of the Royal Society of Edinburgh, Earth Sciences 92:97–102.
   Google Scholar
• Morlo, M. 2006. New remains of Barbourofelidae (Mammalia, Carnivora) from the Miocene of Southern Germany: implications for the history of barbourofelid migrations. Beiträge Zur Paläontologie 30:339–349.
   Google Scholar
• Morlo, M., S. Peigné, and D. Nagel. 2004. A new species of Prosansanosmilus: implications for the systematic relationships of the family Barbourofelidae new rank (Carnivora, Mammalia). Zoological Journal of the Linnean Society 140:43–61.
   Web of Science ®Google Scholar
• Neff, N. A. 1983. The basicranial anatomy of the Nimravidae (Mammalia: Carnivora): character analyses and phylogenetic inferences. City University of New York, 642 pp.
   Google Scholar
• Nyakatura, K., and O. Bininda-Emonds. 2012. Updating the evolutionary history of Carnivora (Mammalia): a new species-level supertree complete with divergence time estimates. BMC Biology 10:1–31.
   PubMed Web of Science ®Google Scholar
• O’Connor, A., and M. A. Wills. 2016. Measuring Stratigraphic Congruence Across Trees, Higher Taxa, and Time. Systematic Biology 65:792–811.
   PubMed Web of Science ®Google Scholar
• O’Keefe, F. R., E. V Fet, and J. M. Harris. 2009. Compilation, calibration, and synthesis of faunal and floral radiocarbon dates, Rancho La Brea, California. Contributions in Science (Los Angeles) 518:1–16.
   Google Scholar
• O’Leary, M. A., J. I. Bloch, J. J. Flynn, T. J. Gaudin, A. Giallombardo, N. P. Giannini, S. L. Goldberg, B. P. Kraatz, Z. Luo, J. Meng, X. Ni, M. J. Novacek, F. a Perini, Z. S. Randall, G. W. Rougier, E. J. Sargis, M. T. Silcox, N. B. Simmons, M. Spaulding, P. M. Velazco, M. Weksler, J. R. Wible, and A. L. Cirranello. 2013. The placental mammal ancestor and the post–K-Pg radiation of placentals. Science 339:662–667.
   PubMed Web of Science ®Google Scholar
• Okonechnikov, K., O. Golosova, M. Fursov, A. Varlamov, Y. Vaskin, I. Efremov, O. G. German Grehov, D. Kandrov, K. Rasputin, M. Syabro, and T. Tleukenov. 2012. Unipro UGENE: A unified bioinformatics toolkit. Bioinformatics 28:1166–1167.
   PubMed Web of Science ®Google Scholar
• Paijmans, J. L. A., R. Barnett, M. T. P. Gilbert, M. L. Zepeda-mendoza, J. W. F. Reumer, J. de Vos, G. Zazula, D. Nagel, G. F. Baryshnikov, J. A. Leonard, N. Rohland, M. V Westbury, A. Barlow, and M. Hofreiter. 2017. Evolutionary history of saber-toothed cats based on ancient mitogenomics. Current Biology 27:3330–3336.
   PubMed Web of Science ®Google Scholar
• Palmqvist, P., V. Torregrosa, J. A. Pérez-Claros, B. Martínez-Navarro, and A. Turner. 2007. A re-evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairodontinae) and the problem of species identification in extinct carnivores. Journal of Vertebrate Paleontology 27:160–175.
   Web of Science ®Google Scholar
• Paterson, R. S., N. Rybczynski, N. Kohno, and H. C. Maddin. 2020. A total evidence phylogenetic analysis of pinniped phylogeny and the possibility of parallel evolution within a monophyletic framework. Frontiers in Ecology and Evolution 7:1–16.
   Web of Science ®Google Scholar
• Peigné, S. 2003. Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32:199–229.
   Web of Science ®Google Scholar
• Piras, P., D. Silvestro, F. Carotenuto, S. Castiglione, A. Kotsakis, L. Maiorino, M. Melchionna, A. Mondanaro, G. Sansalone, C. Serio, V. A. Vero, and P. Raia. 2018. Evolution of the sabertooth mandible: A deadly ecomorphological specialization. Palaeogeography, Palaeoclimatology, Palaeoecology 496:166–174.
   Web of Science ®Google Scholar
• Prevosti, F. J., G. F. Turazzini, and M. Amelia Chemisquy. 2010. Morfología craneana en tigres dientes de sable: alometría, función y filogenia. Ameghiniana 47:239–256.
   Web of Science ®Google Scholar
• Pyron, R. A. 2011. Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia. Systematic Biology 60:466–481.
   PubMed Web of Science ®Google Scholar
• R Core Team. 2019. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.
   Google Scholar
• Rambaut, A., A. J. Drummond, D. Xie, G. Baele, and M. A. Suchard. 2018. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67:901–904.
   Google Scholar
• Ree, R. H., and S. A. Smith. 2008. Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Systematic Biology 57:4–14.
   PubMed Web of Science ®Google Scholar
• Revell, L. J. 2012. phytools: An R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution 3 217–223.
   Web of Science ®Google Scholar
• Rieux, A., and F. Balloux. 2016. Inferences from tip-calibrated phylogenies: a review and a practical guide. Molecular Ecology 25:1911–1924.
   PubMed Web of Science ®Google Scholar
• Robles, J. M., D. M. Alba, J. Fortuny, S. De Esteban-Trivigno, C. Rotgers, J. Balaguer, R. Carmona, J. Galindo, S. Almecija, J. V Berto, and S. Moya-Sola. 2013. New craniodental remains of the barbourofelid Albanosmilus jourdani (Filhol, 1883) from the Miocene of the Valles-Penedes Basin (NE Iberian Peninsula) and the phylogeny of the Barbourofelini. Journal of Systematic Palaeontology 11:993–1022.
   Web of Science ®Google Scholar
• Rögl, F. 1999. Mediterranean and Paratethys. Facts and Hypotheses of an Oligocene to Miocene paleogeography (short overview). Geologica Carpathica 50:339–349.
   Web of Science ®Google Scholar
• Ronquist, F. 1997. Dispersal-vicariance analysis: A new approach to the quantification of historical biogeography. Systematic Biology 46:195–203.
   Web of Science ®Google Scholar
• Ronquist, F., S. Klopfstein, L. Vilhelmsen, S. Schulmeister, D. L. Murray, and A. P. Rasnitsyn. 2012. A total-evidence approach to dating with fossils, applied to the early radiation of the Hymenoptera. Systematic Biology 61:973–999.
   PubMed Web of Science ®Google Scholar
• Rothwell, T. P. 2003. Phylogenetic systematics of North American Pseudaelurus (Carnivora, Felidae). American Museum Novitates 3403.
   Web of Science ®Google Scholar
• Salesa, M. J., M. Antón, A. Turner, and J. Morales. 2005. Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bit. Zoological Journal of the Linnean Society 144:363–377.
   Web of Science ®Google Scholar
• Salesa, M. J., M. Anton, A. Turner, and J. Morales. 2010a. Functional anatomy of the forelimb in Promegantereon* ogygia (Felidae, Machairodontinae, Smilodontini) from the Late Miocene of Spain and the origins of the sabre-toothed felid model. Journal of Anatomy 216:381–396.
   PubMed Web of Science ®Google Scholar
• Salesa, M. J., M. Antón, A. Turner, L. Alcalá, P. Montoya, and J. Morales. 2010b. Systematic revision of the late Miocene sabre-toothed felid Paramachaerodus in Spain. Palaeontolgy 53:1369–1391.
   Web of Science ®Google Scholar
• Sallan, L. C., and M. Friedman. 2012. Heads or tails: staged diversification in vertebrate evolutionary radiations. Proceedings of the Royal Society B 279:2025–2032.
   PubMed Web of Science ®Google Scholar
• Sanders, W. J., and E. R. Miller. 2002. New proboscideans from the early miocene of wadi moghara, Egypt. Journal of Vertebrate Paleontology 22:388–404.
   Web of Science ®Google Scholar
• Schultz, C. B., M. R. Schultz, and L. D. Martin. 1970. A new tribe of saber-toothed cats (Barbourofelini) from the Pliocene of North America. Bulletin of the University of Nebraska State Museum 9:1–31.
   Google Scholar
• Slater, G. J., and B. Van Valkenburgh. 2008. Long in the tooth: evolution of sabertooth cat cranial shape. Paleobiology 34:403–419.
   Web of Science ®Google Scholar
• Solé, F., R. Smith, T. Coillot, E. de Bast, and T. Smith. 2014. Dental and tarsal anatomy of Miacis latouri and a phylogenetic analysis of the earliest carnivoraforms (Mammalia, Carnivoramorpha). Journal of Vertebrate Paleontology 34:1–21.
   Web of Science ®Google Scholar
• Solé, F., T. Smith, E. de Bast, V. Codrea, and E. Gheerbrant. 2016. New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia). Journal of Vertebrate Paleontology 36:e1082480.
   Web of Science ®Google Scholar
• Spaulding, M., and J. J. Flynn. 2012. Phylogeny of the Carnivoramorpha: the impact of postcranial characters. Journal of Systematic Palaeontology 10:653–677.
   Web of Science ®Google Scholar
• Stadler, T. 2010. Sampling-through-time in birth-death trees. Journal of Theoretical Biology 267:396–404.
   PubMed Web of Science ®Google Scholar
• Stadler, T., D. Künert, S. Bonhoeffer, and A. J. Drummond. 2013. Birth-death skyline plot reveals temporal changes of epidemic spread in HIV and hepatitis C virus (HCV). Proceedings of the National Academy of Sciences of the United States of America 110:228–233.
   PubMed Web of Science ®Google Scholar
• Tedford, R. H. 1978. History of dogs and cats: a view from the fossil record; pp. 1–10 in Nutrition and Management of Dogs and Cats. Ralston Purina Co., St. Louis.
   Google Scholar
• Therrien, F. 2005. Feeding behaviour and bite force of sabretoothed predators. Zoological Journal of the Linnean Society 145:393–426.
   Web of Science ®Google Scholar
• Tomiya, S., and Z. J. Tseng. 2016. Whence the beardogs? Reappraisal of the middle to late eocene ‘Miacis’ from Texas, USA, and the origin of amphicyonidae (Mammalia, Carnivora). Royal Society Open Science 3:160518.
   PubMed Web of Science ®Google Scholar
• Towns, J., T. Cockerill, M. Dahan, I. Foster, K. Gaither, A. Grimshaw, V. Hazlewood, S. Lathrop, D. Lifka, G. D. P. R. Roskies, J. R. Scott, and N. Wilkins-Diehr. 2014. XSEDE: Accelerating Scientific Discovery. Computing in Science & Engineering 16:62–74.
   Web of Science ®Google Scholar
• Tseng, Z. J., G. T. Takeuchi, and X. Wang. 2010. Discovery of the upper dentition of Barbourofelis whitfordi (Nimravidae, Carnivora) and an evaluation of the genus in California. Journal of Vertebrate Paleontology 30:244–254.
   Web of Science ®Google Scholar
• Wagenmakers, E.-J., and S. Farrell. 2004. AIC model selection using Akaike weights. Psychonomic Bulletin & Review 11:192–196.
   PubMed Web of Science ®Google Scholar
• Wagner, P. J., and D. H. Erwin. 1995. Phylogenetic Patterns as Tests of Speciation Models; pp. 87–122 in D. H. Erwin and R. L. Anstey (eds.), New Approaches to Speciation in the Fossil Record. Columbia University Press, New York.
   Google Scholar
• Wang, X., S. C. White, and J. Guan. 2020. A new genus and species of sabretooth, Oriensmilus liupanensis (Barbourofelinae, Nimravidae, Carnivora), from the middle Miocene of China suggests barbourofelines are nimravids, not felids. Journal of Systematic Palaeontology 18:783–803.
   Web of Science ®Google Scholar
• Werdelin, L., and M. E. Lewis. 2001. A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132:147–258.
   Web of Science ®Google Scholar
• Werdelin, L., N. Yamaguchi, W. E. Johnson, and S. J. O’Brien. 2010. Phylogeny and evolution of cats (Felidae); pp. 59–82 in D. W. Macdonald and A. J. Loveridge (eds.), Biology and Conservation of Wild Felids. Oxford University Press, New York.
   Google Scholar
• Wesley-Hunt, G. D., and J. J. Flynn. 2005. Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of “Miacoidea” relative to Carnivora. Journal of Systematic Palaeontology 3:1–28.
   Web of Science ®Google Scholar
• Wills, M. A. 1999. Congruence between phylogeny and stratigraphy: randomization tests and the gap excess ratio. Systematic Biology 48:559–580.
   Web of Science ®Google Scholar
• Wozencraft, W. C. 2005. Order Carnivora; pp. 532–628 in D. E. Wilson and D. M. Reeder (eds.), Mammal species of the world. A taxonomic and geographic reference, 3rd. editi. Johns Hopkins University Press, Baltimore.
   Google Scholar
• Wyss, A. R., and J. J. Flynn. 1993. A Phylogenetic Analysis and Definition of the Carnivora; pp. in F. S. Szalay, M. J. Novacek, and M. C. McKenna (eds.), Mammal Phylogeny: Placentals, Volume 2. Springer-Verlag, New York.
   Google Scholar
• Zhou, Y., S.-R. Wang, and J.-Z. Ma. 2017. Comprehensive species set revealing the phylogeny and biogeography of Feliformia (Mammalia, Carnivora) based on mitochondrial DNA. PLoS ONE 12:e0174902.
   PubMed Web of Science ®Google Scholar