THE EVOLUTIONARY HISTORY OF THE FELIFORMIA: CONTINGENCY, CONSTRAINT, DISPARITY by PAUL ZACHARY BARRETT A DISSERTATION Presented to the Department of Earth Sciences and the Division of Graduate Studies of the University of Oregon in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 2022 DISSERTATION APPROVAL PAGE Student: Paul Zachary Barrett Title: The Evolutionary History of the Feliformia: Contingency, Constraint, Disparity This dissertation has been accepted and approved in partial fulfillment of the requirements for the Doctor of Philosophy degree in the Department of Earth Sciences by: Prof. Samantha S. B. Hopkins Chairperson/Advisor Prof. Edward B. Davis Core Member Prof. Thomas Giachetti Core Member Prof. Stephen Frost Institutional Representative and Krista Chronister Vice Provost for Graduate Studies Original approval signatures are on file with the University of Oregon Division of Graduate Studies. Degree awarded June 2022 ii © 2022 Paul Z. Barrett This work is licensed under a Creative Commons Attribution-NonCommercial (United States) License. iii DISSERTATION ABSTRACT Paul Zachary Barrett Doctor of Philosophy Department of Earth Sciences June 2022 Title: The Evolutionary History of the Feliformia: Contingency, Constraint, Disparity Constraint is a fundamental concept in evolutionary theory. The possibilities are not endless for what shape an organism can take, nor ecology of one generation to the next. Cat-like carnivorans (Feliformia) offer a unique opportunity in investigating aspects of evolutionary constraint, given several of their constituent clades are purported to experience this phenomenon in terms of limited evolutionary potential, but also in the presence of some clades with extreme durophagous (bone-crushing) and sabertooth morphology. I investigated the evolutionary history of feliforms by considering their phylogeny, ecology and cranial morphology. I recover results that suggest there are three distinct ecospaces within which feliforms reside. The first is occupied by those small in overall size, covering a wide dietary and ecological spectrum, up to but not including hypercarnviory. These broad ecologies are facilitated by a narrow band of similar cranial shapes, evocative of “one to many mapping”. The second ecopace is occupied by soft- flesh specialists, such as felids (cats) and nimravids. These hypercarnivores possess distinct (non-overlapping) crania from the first ecospace, optimized for bite force and large gape angles. The evolutionary end member of this ecospace are those taxa possessing sabertooth morphology and the most disparate cranial shapes of all. The third ecospace is that of hyaenids, with a diverse ecological breadth optimized for dental toolkit and body mass, channeled by postcranial cursorial adaptations. The evolutionary end member of this ecospace are those taxa possessing iv durophagous morphology. In all three ecospaces, constraint exists in what cranial morphology and ecological transitions are possible. Large size nor hypercarnivory are achievable within the small feliform ecospace, and thus require transitions into the other two zones. Soft-flesh specialization is a road of no return that does not facilitate ecospace transitions. The hyaenid ecospace does not facilitate transitions into the ‘soft flesh’ ecospace, nor by extension sabertooth morphology. These limitations are a result of phylogenetic baggage and functional challenges, but fruitfully channel extreme cranial shapes (soft-flesh specialist) and diverse ecologies (hyaenids) that couldn’t exist without these constraints. v CURRICULUM VITAE NAME OF AUTHOR: Paul Zachary Barrett GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED: University of Oregon, Eugene, Oregon, USA South Dakota School of Mines & Technology, Rapid City, South Dakota, USA Idaho State University, Pocatello, Idaho, USA DEGREES AWARDED: Doctor of Philosophy, Earth Sciences, 2022, University of Oregon Master of Science, Paleontology, 2015, South Dakota School of Mines & Technology Bachelor of Science, Geology, 2010, Idaho State University AREAS OF SPECIAL INTEREST: Paleontology Phylogenetic Systematics Paleoecology Morphology PROFESSIONAL EXPERIENCE: Graduate Employee, University of Oregon, 2016-2022 Staff Research Associate I, University of California, Davis, 2017 Field Paleontologist/Preparator/Curator, Geoscientists-in-the-Parks, Wind Cave National Park, 2015 Paleontology Monitor, United States Bureau of Reclamation, 2010 Curatorial Assistant, Idaho Museum of Natural History, 2009-2011 vi GRANTS, AWARDS, AND HONORS: Department of Earth Sciences Research Excellence Award, University of Oregon, 2022 Open Access Article Processing Charge Award Fund, University of Oregon, 2021 Baldwin Scholarship, University of Oregon Department of Earth Sciences 2017, 2018, 2019, 2020 Annual Meeting Grant, European Association of Vertebrate Palaeontology, 2019 Special "Opps" Travel and Research Award, University of Oregon, 2019 Graduate Student Research Award, Geological Society of America, 2018 J.P. and Virginia Gries Memorial Fellowship, South Dakota School of Mines & Technology, 2013 Idaho Promise Scholarship, Idaho State University, 2006, 2007 PUBLICATIONS: Barrett, P. Z. 2021. The largest hoplophonine and a complex new hypothesis of nimravid evolution. Scientific Reports 11:1–9. Barrett, P. Z., S. S. B. Hopkins, and S. A. Price. 2021. How many sabretooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae. Journal of Vertebrate Paleontology 41(1): e1923523. Miller, S. A., P. Z. Barrett, W. N. F. McLaughlin, and S. S. B. Hopkins. 2020. Endemism and migration in the Kochkor Basin? Identification and description of Adcrocuta eximia (Mammalia: Carnivora: Hyaenidae) and c.f. Paramachaerodus (Mammalia: Carnivora: Felidae) fossils at the Miocene locality of Ortok, Kyrgyzstan. Palaeontologia Electronica 23(3): a45. Barrett, P. Z., L. Finkelman, G. Perdue, W. N. F. McLaughlin, D. M. Reuter and S. S. B. Hopkins. 2020. Small carnivoran fauna of the Mascall Formation, Crooked River Basin, central Oregon. Journal of Vertebrate Paleontology 39(5): e1717506. Retallack, G. J., J. M. Theodor, E. B. Davis, S. S. B. Hopkins and P. Z. Barrett. 2018. First dinosaur (Ornithopoda) from Early Cretaceous (Albian) of Oregon, USA. Journal of Vertebrate Paleontology 38(4): (1)-(5). Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658. vii E. Welsh, C. A. Boyd, K. D. Spearing, and P. Z. Barrett. 2015. Stratigraphic and taxonomic revision of a North American false saber-toothed cat cub. Proceedings of the South Dakota Academy of Science 94:141-153. viii ACKNOWLEDGMENTS I wish to thank my parents who fostered my interest in paleontology since I was four years old. To my committee, who provided countless hours of engagement and discourse that brought this project to fruition. Of special thanks, my advisor Dr. Samantha Hopkins, who facilitated my growth on evolutionary theory leading up to my comprehensive exams, and supplied much of the funds and connections to collect the data for this dissertation once I passed. Dr. Stephan Frost, who shaped the way I view the evolution of morphology, the dark arts of imputation and rigor in my science. I also thank the many curators and collection managers that steward the natural history of the world. Through them I engaged with the evolutionary history of our planet and learned the best places to get a bite to eat and things to do on the weekend. Dr. Suzanne Gascoyne, who helped me become a better human. You immeasurably aided me in navigating the emotional trials of a Ph.D., resulting in a more confident, objective and hopefully, introspective person. To my partner, Sophie Miller, you showed me the path to my PhD was more than the research it contains, the transformative experience which outshines the paper on the wall or any occupation that paper may bring. Thank you for being patient, listening to me drone on about exciting as well as drab ideas. You are my most aggressive cheerleader. Finally, to my future self, you should be proud. This journey was worth it. My investigation was supported in part by the Ewart Baldwin Scholarship and Special “Opps” Travel and Research Award from the University of Oregon, from the Graduate Research Grant of the Geological Society of America and by a grant from the National Science Foundation, number DEB- 1256897. ix For those of whom the old bones come alive and join the pageant of the living. x TABLE OF CONTENTS Chapter I.......................................................................................................................................... 1 Chapter II ........................................................................................................................................ 5 Introduction ................................................................................................................................. 5 Methods ....................................................................................................................................... 6 Results ....................................................................................................................................... 13 Discussion ................................................................................................................................. 18 Chapter III ..................................................................................................................................... 28 Introduction ............................................................................................................................... 28 Methods ..................................................................................................................................... 31 Results ....................................................................................................................................... 39 Discussion ................................................................................................................................. 50 Chapter IV ..................................................................................................................................... 65 Introduction ............................................................................................................................... 65 Methods ..................................................................................................................................... 68 Results ....................................................................................................................................... 78 Discussion ................................................................................................................................. 88 Chapter V .................................................................................................................................... 102 Appendix A ................................................................................................................................. 110 Appendix B ................................................................................................................................. 115 xi Settings .................................................................................................................................... 115 Appendix C ................................................................................................................................. 118 Appendix D ................................................................................................................................. 144 Appendix E ................................................................................................................................. 195 Appendix F.................................................................................................................................. 217 Appendix G ................................................................................................................................. 223 Appendix H ................................................................................................................................. 259 Appendix I .................................................................................................................................. 287 References ................................................................................................................................... 297 xii LIST OF FIGURES Figure 1. Skyline plot of assessed changes in speciation (Re) and extinction (δ) rates through time of the Feliformia. ........................................................................................................... 11 Figure 2. Maximum clade credibility tree of Feliformia. ............................................................. 14 Figure 3. Measurements taken for ecomorphological traits. ......................................................... 32 Figure 4. Distribution of ecological proxy variables used in the presented analysis, natural log of the cube root of body mass in grams, left, and the relative blade length of the lower m1, right. ....................................................................................................................................... 41 Figure 5. Latitudinal boundary shift of tropical and boreal climate zones from the Eocene to the present .................................................................................................................................... 53 Figure 6. Position of the landmarks taken on the cranium for 3DGM analyses (69 landmarks) to quantify variation in shape of feliform crania. ...................................................................... 69 Figure 7. Pruned MCC phylogeny of Chapter II to include only those feliforms analyzed in the present cranial morphology chapter. ..................................................................................... 77 Figure 8. PCA of all cranial feliform specimens.. ........................................................................ 79 Figure 9. Relationship between feliform lineage age and disparity (Procrustes variance). Disparity has been adjusted for allometry. ............................................................................ 83 Figure 10. Relationship between feliform cranial integration (Eigenvalue dispersion) and disparity (Procrustes variance) .............................................................................................. 84 Figure 11. Relationship between feliform cranial integration (between developmental module) and disparity (Procrustes variance). ....................................................................................... 85 Figure 12. Relationship between feliform size and shape evolutionary rates, and morphological disparity (Procrustes variance). ............................................................................................. 87 xiii Figure 13. Log centroid size versus the regression scores of shape on size for each specimen (A). Idealized allometric trajectories predicted values for each clade, highlighting similarities in evolutionary allometric slopes (B). ....................................................................................... 89 Figure 14. Relationship between degree of carnivory and feliform cranial shape (PGLS). ......... 92 Figure 15. Hypothesis of terrestrial carnivoran evolution with regards to ecology and cranial morphology. ......................................................................................................................... 104 xiv LIST OF TABLES Table 1. Best supported substitution models recovered in PartitionFinder2 using the Bayesian information criterion (BIC). .................................................................................................... 8 Table 2. Summary of marginal log-likelihoods and Bayes factor (BF) support for differential substitution models and partitioning schemes of the morphology data................................. 10 Table 3. Ecomorphological traits of analysis with abbreviations and constituent measurements.. ............................................................................................................................................... 33 Table 4. Priors for bayou analyses. *= Calculated using ‘cdpois’ option in bayou. ..................... 36 Table 5. Median results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2. .............................................. 43 Table 6. Median Felidae results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2 .................................... 44 Table 7. Median Nimravidae results from macroevolutionary model fits to 500 random pruned trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2 ............. 47 Table 8. Median Hyaenidae results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2 ............................... 48 Table 9. Median small feliform results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2 ...................... 49 Table 10. Median optimal body mass (in kg) values from model fits of 500 random trees from the stable Bayesian posterior of the phylogenetic analysis for hyaenids of the present analysis and that of canids in Slater (Slater, 2015). ............................................................... 59 Table 11. Definition of anatomical landmarks used in the three-dimensional geometric morphometric analyses. ......................................................................................................... 70 xv Table 12. Definitions of dietary categories for analysis of cranial shape. .................................... 74 Table 13. Allometry and effect of relative canine size (RCS) on cranial shape as determined by ANOVA of shape (Procrustes coordinates) ~ log(centroid size)+RCS for each clade. ........ 80 Table 14. Evolutionary allometry ANCOVA. .............................................................................. 81 Table 15. Cranial disparity (Procrustes variance adjusted for size) per dietary category. ............ 82 Table 16. Pairwise cranial shape evolution rates between dietary ecologies in upper triangle, with p-value significance in lower triangle. .................................................................................. 88 Table 17. Evolutionary allometry pANCOVA. ............................................................................ 90 Table 18. Pair-wise homogeneity of slope test summary of p values for distinct evolutionary allometric trajectories ............................................................................................................ 91 xvi Chapter I Introduction Variation supplies the raw material by which natural selection operates, “variation supposes, while selection disposes.” However, numerous studies have acknowledged the importance of understanding that, unlike Darwin’s original formulation, variation generated by organisms is not isotropic (Olson and Miller, 1958; Vermeij, 1973; Gould, 2002; Goswami et al., 2011). Possibilities are not limitless, as evident by the non-equal occupation of morphospace and ecology of all lineages. What obstructs the opportunities for some lineages and perhaps facilitates diversity and disparity for others remain central questions in biology and evolutionary theory (Dobzhansky, 1951; Hutchinson, 1959; Gould, 2002). Hypotheses have existed since Galton (1869) that there may be minimally stable states or positions in both morphology and ecology, where evolution proceeds less like a rolling billiard ball of isotropic generational variation, and more analogous to a polyhedron that flips facets of stable ecomorphology. In other words, maybe a reasonable amount of isotropic variation is created in each generation, but only a limited amount is tenable in being a successful organism. From these proposals rose ideas of channelized evolution (Haeckel, 1866; Eimer, 1890; Hyatt, 1897; Goldschmidt, 1940) that held promise in predicting the course of shape and niche change if the framework could be understood. Evolutionary history or deep homology may play a truly integral part in determining what shapes and occupations an organism can achieve, either through lack of genetic raw material or variation, or developmental correlation of parts that will not allow certain changes to happen. However, these same correlations may also provide an axis of exploitation, where a greater magnitude of change may be possible within a certain trajectory 1 compared to equally probable random change in any direction (Goswami et al., 2014). This fruitful channeling could then be viewed as a positive aspect of constraint leading to more extreme shapes and, by extension, ecologies than would be possible under Darwin’s initial hypothesis. Areas of limited variation are often ascribed to specialists. With the concept of one-to-one mapping of ecology to shape, one would expect that fewer niches or a more limited ecology would correlate with less morphological variation. Among the carnivoran clades, low amounts of cranial-dental variation have been described in literature for “cat-like” feliform carnivores such as felids and nimravids (Radinsky, 1981a, 1982; Van Valkenburgh, 1991; Holliday and Steppan, 2004). This reduced raw material is argued to relate to increased specialization towards a hypercarnivorous diet, one dominated by a predominance of vertebrate prey. Felids and nimravids both exhibit simplified dentition that efficiently slices soft flesh but lacks the grinding surfaces found in more omnivorous carnivores (Evans and Pineda-Munoz, 2018). The loss of grinding surfaces on teeth seems to be an evolutionary road of no return, resulting in an ecomorphological constraint that did not allow these specialists to evolve additional ecologies (Solé and Ladevèze, 2017; Brocklehurst, 2019). Consequently, feliforms offer a unique opportunity to investigate aspects of evolutionary constraint, not only for the perceived lack of variation inherent to their specialist clades, but also the extreme morphologies acquired by extinct sabertooth and bone-crushing members. Was constraint an aspect of feliform evolution that limited possibilities for certain lineages, or did that same constraint fruitfully channel the extreme morphologies that did evolve? Additionally, since all clades did not enter into the hypercarnivorous and presumably specialized realms of cats, nimravids and hyenas, what gave these latter groups the push to get there in the first place? 2 I investigated the evolutionary history of feliforms by considering their phylogeny, ecology and cranial morphology across the following chapters. In Chapter II, I construct a hypothesis of feliform evolutionary relationships using total-evidence Bayesian techniques. This methodology allowed me to incorporate molecular (nuclear and mitochondrial genome), dental and osteological morphology and stratigraphic occurrence data in a single analysis. This phylogeny serves as a framework for all following analyses, while also assessing relationships of several purported feliform taxa for the first time, e.g., percrocutids, and holistically estimating major clade originations and relative speciation and extinction rates by combining the fossil and living record. Chapter III assesses ecological evolution of feliforms in terms of their diet, dental toolkit and body mass. I used hypotheses of adaptive radiation, as facilitated by tectonic and climatic events, as a framework to assess different clades’ ecological trajectory. However, I compared models of adaptive radiation to other potential processes to best explain the patterns observed in feliform natural history. These models included random walk, trended evolution, optimal body mass and toolkit per dietary category, and clade-based regime shifts. Using the results of Chapter II, I was also able to incorporate phylogenetic uncertainty for all analyses by virtue of sampled trees over the course of the Bayesian analysis tree search. Chapter IV explores the generating forces of feliform cranial shape through 3D geometric morphometric analysis. I assessed which feliform clades possess the greatest cranial disparity, such do clades with broad ecological occupation have greater variance compared to specialists? I consider multiple plausible sources of cranial disparity such as allometry as a line of least evolutionary resistance, integration of the developmental units of the cranium, the geologic time a clade has existed to evolve disparate shapes, and dietary ecology. 3 Chapter V synthesizes the above analyses into a narrative of feliform evolution and offers predictions for all terrestrial carnivorans. This narrative finds support for constraint in the evolutionary history of feliforms with a finite number of possible ecospaces. Certain transitions are not possible from one ecospace to another, but these same constraints fruitfully channel the extreme endmembers of cranial evolution and ecology that otherwise are inaccessible to other groups, i.e., sabertooth and durophagous morphology. Similar patterns are further observed for caniforms and suggestive of a general process at work in the evolution of terrestrial carnivores. 4 Chapter II Phylogeny of the Feliformia Introduction Current hypotheses of feliform relationships are derived from primarily molecular- or morphological-only datasets (though see Barrett et al., 2021). Molecular hypotheses (e.g. Nyakatura and Bininda-Emonds, 2012; Paijmans et al., 2017; Zhou et al., 2017; Slater and Friscia, 2019; Hassanin et al., 2021) differ extensively from morphological ones (e.g. Wesley- Hunt and Flynn, 2005; Spaulding and Flynn, 2012), such as the placement of the Hyaenidae (hyenas) as distantly related to the Felidae (cats) in molecular phylogenies, while most closely related to the Felidae in morphological phylogenies. An additional issue is the cited morphological studies, including the preliminary total-evidence analysis of Barrett et al. (2021), sample extinct taxa from across the Feliformia, but contain typically one representative operational taxonomic unit (OTU) per family, usually extant. These living OTUs are commonly derived members of their respective families and thus may generate long-branch attraction in phylogenetic hypotheses (Bergsten, 2005). Another explanation for this discrepancy is that differing topologies in molecular trees result from the inability to include most extinct taxa, while all multifamily morphological trees are depauperate in the extant and fossil taxa that they do contain. Furthermore, several extinct clades of feliform (e.g., Percrocutidae, Lophocyonidae, Machairodontinae) have rarely been included in broad feliform phylogenetic analyses, so their placement is poorly known. Combined, these challenges have resulted in uneven knowledge of feliform relationships. Extant or recently extinct clades with molecular data have returned 5 consistent relationships, but necessarily devoid of the diversity of extinct clades, while taxa known only from morphology suffer from lack of analyses, limited taxon sampling or both. Is there a contradiction in the evolutionary relationships of feliforms depending on what datasets one employs, or is this an artifact of methodology and taxon sampling? Here I present a broad investigation of the phylogenetic relationships of feliform taxa by examining the relationships of all major families, and numerous OTUs within each family, with extensive morphological and molecular datasets. I address the conflict of past analyses by employing total-evidence techniques (Lee et al., 2009; Pyron, 2011; Ronquist et al., 2012) which use a Bayesian framework to create a probabilistic model of evolution, inclusive of molecular data, fossil data, and stratigraphic information in a single analysis. Specifically, I implement a total-evidence analysis with a Sampled Ancestor, Birth-Death Skyline with Serial Sampling (SABDSKY) model (Stadler et al., 2013; Gavryushkina et al., 2014) which allows the identification of direct ancestors and changing speciation, extinction and sampling rates through time. This analysis yields a dated phylogeny of extinct and living taxa combining data previously studied in separate morphology-only and molecular-only analyses. Furthermore, this analysis places several taxa/clades in novel arrangements allowing de novo assessment of historical patterns of morphological and ecological evolution. Methods Taxa Selection I chose feliform taxa that captured the cranial disparity and ecological diversity of their given clade, while providing even taxonomic coverage. All feliform taxa with published aDNA (n=7) 6 were also incorporated into this analysis to provide greater resolution on phylogenetic relationships along with molecular data from all extant taxa. The fossil record of feliform carnivorans is not symmetric across clades, with collections and literature favoring large-bodied taxa, e.g. felids, hyaenids and nimravids. However, I included well-preserved small-bodied fossil viverrids, herpestids and stem feloids/viverroids whenever possible. The final phylogenetic analysis includes 124 taxa, plus five caniform outgroups, with approximately two-thirds of all taxa being extinct. Molecular Data I downloaded three nuclear loci and complete mitochondrial genomes from Genbank for the forty-five extant and seven extinct (aDNA) taxa of the analysis (Appendix A). However, where complete mitochondrial genomes were unavailable, I used individual mitochondrial genes. I aligned Nucleotide sequences one gene at a time using MAFFT v.7 online server (Katoh et al., 2019), with the G-INS-I and ‘Leave gappy regions’ parameters selected. All sites containing missing data were removed. This necessitated the loss of codon-position information for all save one mitochondrial gene (ND4L). I deleted the stop codons and manually optimized these aligned sequences in UGENE v. 35.1 (Okonechnikov et al., 2012) to maintain codon positions when possible and trimmed excess bases of tail data that had no comparative sequences. I then concatenated the aligned sequences in SequenceMatrix (Vaidya et al., 2011) and analyzed them for partitioning schemes utilizing PartitionFinder version 2.1.1 (Lanfear et al., 2016) on XSEDE (Towns et al., 2014) through the CIPRES Science Gateway v. 3.3 (Miller et al., 2011). I designated twenty data blocks for the dataset, such that only genes IRBP and ND4L were separated into three blocks based on codon positions. The remainder I gave a single block per 7 molecular sequence. I defined linked branch lengths and investigated all evolutionary models available to BEAST 2 (40 models) using a ‘greedy’ partitioning strategy to compare partitions and differential substitution models under the Bayesian information criterion (BIC). A 10- partition scheme was best supported (Appendix B, Table 1). Table 1. Best supported substitution models recovered in PartitionFinder2 using the Bayesian information criterion (BIC). No. of partitions Partition Model 1 ND4L_1stpos, COX3, ATP6, COX2, COX1 HKY+I+G+X 2 ND4, CYB, ND1, ATP8, ND5, ND3 GTR+I+G+X 3 TTR, CHRNA_1 SYM+G 4 IRBP_1stpos TRN+G+X 5 IRBP_2ndpos TRNEF+I 6 IRBP_3rdpos HKY+G+X 7 ND2 HKY+I+G+X 8 ND4L_2ndpos HKY+I+G+X 9 ND4L_3rdpos HKY+I+G+X 10 ND6 HKY+G+X 8 Morphology Data I scored all taxa from personal observation of specimens or literature as listed in Appendix C for 325 morphological characters derived from prior carnivoran/feliform phylogenetic analyses (Van Valkenburgh et al., 1990; Werdelin and Solounias, 1991; Salles, 1992; Bryant, 1996; Peigné, 2003; Rothwell, 2003; Gaubert et al., 2005; Wesley-Hunt and Flynn, 2005; Tseng and Wang, 2007; Christiansen, 2008a; Salesa et al., 2010; Sakamoto and Ruta, 2012; Salesa et al., 2012; Spaulding and Flynn, 2012; Christiansen, 2013; Robles et al., 2013a; Barrett, 2016; Morales et al., 2019; Barrett, 2021; Barrett et al., 2021) and several novel characters. When possible, multiple individuals of a given taxon were scored to assess polymorphic conditions. Characters and their associated descriptions can be found in Appendix D, while the resultant morphology character matrix is Appendix E. I assessed six competing schemes of morphological character partitioning and evolution by Bayes Factor via a Generalized Stepping Stone (GSS; Fan et al., 2011) analysis (Table 2). These schemes partitioned characters based upon anatomical association (cranial/post-cranial), number of states a character contains (n-states), or a combination of the two. I additionally wrote custom evolutionary rate matrices for specific multistate characters (Appendix F) that included ordered, multipath and Dollo (irreversible) characters to compare to null unordered character evolution. Each scheme was assessed by GSS with 10 steps, a chain length of five million and 50% burnin. 9 Table 2. Summary of marginal log-likelihoods and Bayes factor (BF) support for differential substitution models and partitioning schemes of the morphology data. Log BF values are reported as the support of the best model (model 6) over the inline model. P=partition used, * = best supported model. Scheme n-state Cranial/Post- Complex Evo. Marginal log- Log BF vs partition cranial part. models likelihood Best Model 1 P -16804.22 4058.26 2 P P -14672.40 1926.44 3 P P -12984.09 238.13 4 P -12974.56 228.60 5 P P P -12752.54 6.58 6* P P -12745.96 - Total-evidence Analysis I jointly estimated tree topology, branch lengths and evolutionary parameters in a Bayesian framework using BEAST2, version 2.6.3 (Bouckaert et al., 2014) with a Sampled Ancestor, Birth-Death Skyline with Serial Sampling (SABDSKY) model as a tree prior (Stadler et al., 2013; Gavryushkina et al., 2014). A SABDSKY model allows analyzed taxa to be direct ancestors (zero length branches) to other taxa, while serial skyline models facilitate changes in extinction, speciation or sampling rates through time, factors which are likely non-trivial to the approximately 50 million year history of feliform evolution (Liow and Finarelli, 2014; Pires et al., 2017). I assessed the number of speciation and extinction rate shifts (sampling standard deviation was found to approximately zero, thus rejecting the presence of rate variation and justifying the use of a single sampling parameter) by first arbitrarily testing ten evenly spaced 10 shifts at 5 Ma increments. I visualized/assessed the proposed shifts in Tracer version 1.7.1 (Rambaut et al., 2018) and the ‘bdskytools’ R package (Plessis, 2016). From these results, I reduced the number of shifts to six with boundaries correlating to the rate inflection points in Figure 1. Speciation (Re) and extinction (δ) were given uniform priors [0,10], while sampling was given a uniform [0,1]. The origin parameter was given a lognormal distribution, offset of 47.4 Ma (being the FAD of the oldest taxon in the analysis), mean=1.0, standard deviation=1.25 and ‘mean in real space’ parameter selected. Figure 1. Skyline plot of assessed changes in speciation (Re) and extinction (δ) rates through time of the Feliformia. I set each morphological partition to evolve under the Lewis Mkv model, or those best supported by the GSS analysis, scheme 6. All partitions were set to accommodate ascertainment bias and ambiguities in the character matrix with a substitution rate that varied across characters 11 according to a Gamma distribution shared by all partitions. I modelled the shared morphological clock with an uncorrelated relaxed clock model with log-normal distributed rates. For molecular data, I applied a single lognormal relaxed clock model with partitions and evolutionary models following the results of the PartitionFinder2 analysis. I derived the distributions (with provided justifications) of tip dates from the information in Appendix G. Early runs of the analysis displayed a lack of convergence between the morphological and molecular data, thus I constructed a reasonable time-scaled starting tree in R version 4.0.3 (R Core Team, 2020) using the ‘DatePhylo’ function in the ‘strap’ package (Bell and Lloyd, 2014). I set a relatively uninformed “equal” set of branch lengths, calculated across a feliform tree derived from relationships in literature (e.g. Turner et al., 2008; Christiansen, 2013; Slater, 2015; Slater and Friscia, 2019; Barrett, 2021; Barrett et al., 2021) using the First Appearance Data (FAD) as described in Appendix G. These branches I further scaled using the ‘minBranchLength’ function in ‘paleotree’ (Bapst, 2012) with a setting of 1.5 to approximate divergence dates of major clades following molecular hypotheses as listed in the aforementioned literature. I finally specified a minimal set of topology constraints separating the ingroup from the outgroup taxa, as well as relationships within the outgroup Caniformia as ascertained from phylogenetic analyses in the prior listed literature. I ran Markov Chain Monte Carlo (MCMC) chains for 100 million generations, sampling trees and parameters every 10,000 generations, until satisfactory sampling of parameters had been reached as assessed by Tracer v. 1.7.1. Thus, I discarded 10% burn-in and estimated topology and posterior probabilities from the remaining generations in the form of a maximum clade credibility (MCC) tree with median node heights. 12 I assessed morphological synapomorphies via TNT v. 1.5 (Goloboff and Catalano, 2016) on the MCC tree of the above analysis. TNT only reports synapomorphies supported by both ACCTRAN and DELTRAN optimization and thus forms a conservative assessment of shared derived character states. Results The MCC tree is presented in Figure 2. Overall, broad (family-level) relationships are consistent to those recovered in molecular analyses, with felids and prionodontids sister to the viverroids (e.g. Zhou et al., 2017; Slater and Friscia, 2019; Hassanin et al., 2021). Similarly, relationships within families containing extant taxa largely follow those of molecular analyses, though differences do exist. Euplerids follow relationships of Hassanin et al. (2021), but differ in the placement of the fossa, Cryptoprocta ferox, relative to previously referenced studies. The Herpestinae (solitary mongooses) lineage differs amongst all cited studies, while the clade Mungotinae (eusocial mongooses) is recovered in the same arrangement amongst all cited studies. The poor resolution of the Herpestinae is reflected in the relatively poor support metrics amongst these studies and the present one, indicating need for future work with this clade. Similarly, certain feline relationships differ amongst all studies, while pantherine felids were recovered in the same arrangement. The recovered internal relationships of the remaining extant clades were identical to those of the literature previously cited. 13 Figure 2. Maximum clade credibility tree of Feliformia. Values adjacent to nodes are posterior probability support ≥0.5. Extinct taxa with analyzed aDNA are indicated by the double-helix symbol. Silhouettes are taken from phylopic.org. I recovered extinct taxa in several novel arrangements, though the overall pattern of nimravids as sister to the feloids with Tapocyon as a stem-feliform was recovered in recent analyses (Tomiya and Tseng, 2016; Barrett et al., 2021) and in the present analysis with 1.0 posterior probability 14 (PP) for each clade. The Nimravidae exhibits a basal split consisting of the Hoplophoninae and Nimravinae of Barrett (2021), though the taxa Dinictis and Pogonodon are recovered as basal members of the Hoplophoninae (0.74 PP), while Maofelis and unnamed specimen MA-PHQ 348 as basal members of the Nimravinae (0.12 PP). The remaining differences in relationships of the Nimravidae are to regions of the tree where both the present study and Barrett (2021) recovered poor resolution. The African palm civet, Nandinia binotata, is recovered in a novel relationship nested amongst the Stenoplesictidae (0.53 PP), late Oligocene stem feloids. This clade further includes Proailurus, Stenogale and Palaeoprionodon, similar to previous morphology only studies, minus the inclusion of Nandinia (Wesley-Hunt and Flynn, 2005; Spaulding and Flynn, 2012; Solé et al., 2014, 2016). The felids are split amongst the Machairodontinae (0.94 PP, seven synapomorphies) and Felinae + Pantherinae (1.0 PP, eight synapomorphies) with Pseudaelurus validus as stem to both (1.0 PP). The Machairodontinae contains the Smilodontini, Homotherini and a series of stem “Metailurini” taxa which diverged from the remaining living felids at approximately 15.17 Ma, comparable to other molecular-only analyses, ranging from 14-22 Ma (Paijmans et al., 2017; Slater and Friscia, 2019; Barnett et al., 2020). The Homotherini of the present study contains Homotherium, Xenosmilus and Amphimachairodus taxa (0.54 PP, three synapomorphies) diverging from the Smilodontini at 10.78 Ma, much shallower than the molecular aDNA analysis of Paijmans et al. (2017) at ~18 Ma, though comparable to Slater and Friscia (2019), 11.78 Ma. The Homotherium species H. latidens and H. serum were also assessed to have an extremely shallow divergence (~0.46 Ma), supporting a conspecific relationship previously hypothesized in analyses with narrower taxonomic scope (Antón et al., 2014; Paijmans et al., 2017). The 15 Smilodontini of the present study contain Smilodon and Megantereon (0.99 PP, seven synapomorphies). As previously mentioned the remaining machairodontines form a grade of stem “metailurins”, resembling the relationships presented in Christiansen (2013) most closely. However, the taxa Nimravides and Machairodus were recovered as well supported (0.72-0.86 PP) stem members of Smilodontini + Homotherini clade, while the remainder recovered poorer support at the base of the Machairodontinae. Extinct lions, Panthera atrox and Panthera leo spelaea, are recovered in the same relationships of previous molecular studies, with the Eurasian cave lion sister to the North American lion and the living African lion stem to both, but at slightly deeper divergence dates, 1.58-2.31 Ma (Barnett et al., 2009, 2016). The North American cheetah-like cat, Miracinonyx, was recovered as sister to Puma concolor (1.0 PP), diverging approximately 3.63 Ma, comparable to past molecular analyses (Barnett et al., 2005; Slater and Friscia, 2019). Hyaenids experienced perhaps the most disparate changes in relationships compared to past analyses which generally described an evolutionary trajectory of ‘civet-like’ to extremely durophagous forms (Werdelin and Solounias, 1991; Turner et al., 2008; Coca-Ortega and Pérez- Claros, 2019). The present study recovers up to three independent lineages evolving durophagous morphology. However, similar to Werdelin and Solounias (1991), the civet- and mongoose-like Protictitherium, Plioviverrops and Tungurictis were recovered in basal positions. Tip ward of these taxa, the first durophagous clade is recovered of Tongxinictis and Allohyaena (0.5 PP). This is followed by the divergence of the ‘ictithere’, Hyaenotheriini + Ictitheriinae of Semenov (2008), taxa including Ictitherium viverrinum, Hyaenictitherium hyaenoides and Hyaenotherium wongii (0.96 PP, four synapomorphies). Subsequently to diverge is the second clade of durophagous morphs, including Belbus, Palinhyaena, and Lycyaena (0.58 PP). 16 However, this clade’s precise position within the greater Hyaenidae is not robustly supported with adjacent nodes yielding only 0.37-0.54 PP values. The final major clade (0.58 PP) of the Hyaenidae contains the crown group plus a sister-group of cursorial meat and bone specialists and durophagous taxa (sensu Coca-Ortega and Pérez-Claros, 2019). The non-crown clade contains the taxa Chasmaporthetes, and a novel Adcrocuta + Pachycrocuta clade (0.94 PP, five synapomorphies). Pachycrocuta has been closely connected to the Hyaena genus, both taxonomically and cladistically (see Liu et al., 2021), but the present study finds a more distant relationship. The crown clade contains the living and recently extinct taxa (i.e. Crocuta crocuta spelaea) in the relationship of molecular hypotheses (Koepfli et al., 2006; Slater and Friscia, 2019; Westbury et al., 2020; Hassanin et al., 2021), diverging from a common ancestor at ~7.7 Ma. Basal to these taxa lie the entirety of the ‘percrocutid’ (sensu Werdelin and Solounias, 1991) species of this study. Percrocutids have long been attributed to a separate family (Schmidt- Kittler, 1976; Chen and Schmidt-Kittler, 1983; Werdelin and Solounias, 1991), based upon derived dentition and their early geologic occurrence. However, the present study recovers them nested far within the Hyaenidae, immediately basal to the living taxa (0.68 PP, nine synapomorphies). The taxa Dinocrocuta gigantea and Percrocuta carnifex are recovered in a sister-group relationship (0.97 PP), while the remaining Percrocuta taxa form a grade below the extant species. The last feliform family assessed was the Lophocyonidae, whose highly supported internal relationships (0.96-1.0 PP, seven synapomorphies) are found to be identical to those of Morales et al. (2019). However, placement of this clade within the Viverroidea was not well resolved by the present analysis. The aforementioned analysis of Morales et al. (2019) hypothesized a relationship sister to the hyaenids, to the exclusion of viverrids, but did not include herpestid 17 taxa. The present study recovered lophocyonids between viverrids and the hyaneid + euplerid + herpestid clade, but posterior probability of placement only ranged from 0.09-0.27 PP for adjacent nodes. This high uncertainty is likely related to the equally uncertain placement of other historically enigmatic feliform taxa: Herpestides antiquus, Kichechia zamanae and Kanuites lewisae (0.05-0.25 PP) in this analysis. Discussion The recovered feliform phylogeny presents new insights into relationships of several clades, as well as aspects of overall diversification and rates of speciation for the clade. Compared to the most recent Nimravidae analysis (Barrett, 2021), Maofelis and MA-PHQ 348 were recovered as stem Nimravines, sister to the European taxa Dinailurictis, Quercylurus and Eofelis. This is not unreasonable, given all these Eurasian taxa share an apomorphic triple rooted P3, though the support for this clade is low, 0.25 PP. Likewise, the clade of Dinictis and Pogonodon was also recovered in a different part of the tree, as stem hoplophonines, but in contrast, this relationship had substantial support, 0.74 PP. These differences in topology to Barrett (2021) may relate to the differential model choice (FBD versus BDSKY) of the two analyses, given the morphology character matrices contain identical scorings for the shared characters. Of course, the present analysis also contains one hundred more characters than Barrett (2021), which may have reconstructed different models of character evolution even when most of these characters were non-parsimony informative for nimravids. Regardless, the results of this analysis suggest that the stem members of each nimravid subfamily are somewhat unstable, needing additional characters and/or newly discovered taxa to aid in their resolution. However, the current phylogeny is informative on the oldest occurrences of purported nimravids. These specimens are from the 18 Irdinmanhan Asian land mammal age of China, and represented by fragmentary upper canines (Chow, 1958; Ding et al., 1977; Zheng et al., 1978). Problematically, these canine fragments are referred to Eusmilus (a genus otherwise not known until the late Eocene), based upon their degree of compression and presence of serrations. These teeth are more derived than the geologically younger (late Eocene) Maofelis cantonensis (Averianov et al., 2016) for which a whole cranium is known. The present phylogenetic analysis suggests that an Irdinmanhan or older age (i.e., ≳43 Ma) for the basal nimravid node is highly improbable (Median = 41.33 Ma; 95% credible interval = [38.76, 43.94] Ma). Combined, these early records are more likely those of machaeroidine oxyaenodonts, another group of sabertooth carnivores known more securely from the Irdinmanhan of China (Zack, 2019a; Zack et al., 2022). A notable long branch of this study’s phylogeny is located on the stem of the Feloidea. This region of the tree has received little phylogenetic work in literature, with groups such as the palaeogalids and stenoplesictids rarely incorporated in evolutionary trees. Palaeogalids are small viverroid-like feliforms first known from the late Eocene of North America, whereby they achieve a Holarctic distribution in the Oligocene and early Miocene (Wang and Zhang, 2015). The exact phylogenetic placement of this family has long been disputed, but the only analysis thus far places the clade at the base of the Feliformia (Wang and Zhang, 2015). However, given the geologic age of the earliest nimravids and lack of temporal context in the aforementioned analysis, a future Bayesian tip-dated phylogeny may shed light on the placement of this clade. Furthermore, a recently described Palaeogale specimen (JODA 13221) from the Turtle Cove Member of the John Day Formation features a complete cranium with intact and fully ossified auditory bullae (Famoso and Orcutt, 2022). Basal nimravids and all stenoplesictids lack a fully ossified auditory bulla, which implies that palaeogalids independently acquired this condition, as 19 seen in Miocene nimravids, or may nest higher within crown Feliformia. Inclusion of the John Day specimen in any subsequent phylogenetic analysis will go far in unraveling the evolutionary history of this group. Stenoplesictids also offer a fruitful avenue of investigation given numerous publications describing their anatomy, but lacking in phylogenetic analysis (e.g. Hunt, 1989, 1998a; Peigné, 1999; Peigné and De Bonis, 1999; Hunt, 2001a; Hans-Volker et al., 2007). A sample of stenoplesictids was analyzed in the presented phylogeny, suggesting that the living African palm civet Nandinia may represent the last surviving descendant of this once hypo- to hypercarnivorous clade. Nandinia has been found has the most-basal member of the extant Feliformia in several molecular analyses (Flynn et al., 2005; Nyakatura and Bininda-Emonds, 2012; Paijmans et al., 2017; Zhou et al., 2017; Slater and Friscia, 2019; Hassanin et al., 2021), with frequent commentary on its apparently plesiomorphic partially-ossified auditory bullae in morphological analyses (Hunt, 1987, 1989, 1998a; Wible and Spaulding, 2013). The inclusion of Nandinia in the Stenoplesictidae fits well with the molecular divergence estimates of the current and prior studies, as well as the bullar condition of representative stenoplesictids of the latest Oligocene (Hunt, 1998a; Peigné and De Bonis, 1999). Thus, the evolution of an ossified bulla amongst feliforms may have occurred on three independent occasions: nimravids, palaeogalids and crown Feliformia minus stenoplesictids. Machairodontines were recovered with two, not three, well-supported tribes. However, it is probably no coincidence that the purported third tribe, Metailurini, was also the only tribe lacking aDNA data. Felids are generally believed to be conservative in their hypercarnivourous morphology (Holliday and Steppan, 2004; Van Valkenburgh, 2007; Chamoli and Wroe, 2011), creating challenges in the available diversity of morphological characters when the identified fossil material is limited in most taxa to craniodental remains. The most recent machairodontine 20 analysis (Werdelin and Flink, 2018) was also unable to recover these traditional tribes, even with a slightly expanded ingroup compared to this study’s phylogeny (18 versus 14). However, the presented phylogeny did provide support for some disputed machairodontine hypotheses. The genus Nimravides has been suggested to be a taxon stem to Pantherinae + Felinae which convergently became sabertoothed given its relatively less derived dental morphology, geologic age and geographic context (Werdelin et al., 2010; Piras et al., 2018). This hypothesis cannot be ruled out by the tree of Werdelin and Flink (2018), given its unstable position as a stem feline or machairodontine depending on the settings of that study. However, this analysis supports the taxon as a machairodontine (0.94 PP) located near the base of the subfamily, before the split of smilodontins and homotherins. Nimravides is a New World taxon, with at least one species (Nimravides catocopis) suggested to be a member of the Old World homotherin Machairodus lineage (Antón et al., 2013). The presented results cast doubt on that hypothesis. However, no Old World Machairodus taxa were included in this analysis, so assessment of immigration events and tribe allocation for Machairodus taxa will require an expanded taxa set, though recent phylogenetic analyses suggest Old World Machairodus may not be a homotherin either (Christiansen, 2013; Werdelin and Flink, 2018). Additionally, Promegantereon was not recovered as a stem smilodontin, a relationship historically hypothesized, but not demonstrated via phylogenetic analysis (Christiansen, 2013; Werdelin and Flink, 2018). Instead, this taxon was recovered in a poorly supported clade (0.24 PP) at the base of the machairodontines reminiscent of hypothesized metailurins (see Christiansen, 2013). Much like Nimravides, this region of the machairodontine tree would benefit from the addition of further taxa. Recent revisions and anatomical descriptions offer abundant content to include a potential clade or grade of taxa thought to be convergent on felines and pantherines (Werdelin and Lewis, 2001; Spassov and 21 Geraads, 2015; Li and Spassov, 2017; de Bonis et al., 2018; Piras et al., 2018). One final area of the felid tree that would benefit from increased scrutiny, is the ancestry and descendant relationships of Pseudaelurus and Hyperailurictis with other felids. Numerous stem felids have historically been lumped into the Pseudaelurus genus, but renewed interest in a hypothesis of differential lineages of felid evolution has gained ground in recent years. Old World Pseudaelurus has been hypothesized to be a stem member of the Machairodontinae, while New World Hyperailurictis stem to all felids, or even a basal felid offshoot that gave rise to an endemic Nimravides (Kretzoi, 1929; Werdelin et al., 2010; Browne and Reynolds, 2015; Piras et al., 2018). However, even with the wealth of fossil material and publications on these early felids (e.g. Hunt, 1998a; Rothwell, 2001, 2003; Robles et al., 2013b), no analysis has given them broad context to support the splitting of these stem taxa into distinct lineages. Future work will go a long way towards our understanding of the evolution of the felids, how many lineages converged upon sabertooth, feline and pantherine morphologies, and biogeographic history of the entire family. Perhaps the most novel result of this phylogeny is the recovery of percrocutids as derived hyaenids (1.0 PP). As previously mentioned, the derived dentition of these taxa has been the main evidence for their exclusion from the hyaenids. Specifically, in the dp4 a metaconid that is medially located within the trigonid complex and clearly separated from the talonid has been argued to be synapomorphic for hyaenids, while an inline, posteriorly placed metaconid connected with the talonid is the pattern among percrocutids (Schmidt-Kittler, 1976; Chen and Schmidt-Kittler, 1983; Werdelin and Solounias, 1991). However, in the work of Schmidt-Kittler (1976) it is suggested that this percrocutid condition could be the result of rapid evolution towards a hypercarnivorous morphology, losing the crushing talonid in favor of the slicing 22 trigonid. In the same work, the author describes the percrocutid condition in a variety of other feloids and nimravids (see Xiong, 2019 for a more detailed discussion) suggesting that it may have evolved multiple times. In fact, this morphology is present in the aardwolf, Proteles, and formed the basis for its removal to a separate Protelidae, inclusive of Percrocuta and Dinocrocuta (Baryshnikov and Averianov, 1993; Averyanov and Baryshnikov, 1996). The presented phylogeny used this character as part of its analysis, but still recovered all percrocutids as derived hyaenids, a hypothesis proposed, but untested in Xiong (2019) based upon basicranial morphology in Dinocrocuta. Furthermore, these taxa were recovered basal to the living hyaenids, but this relationship is challenged by the well-supported hypothesis that Proteles is the most basal living hyaenid (Zhou et al., 2017; Slater and Friscia, 2019; Westbury et al., 2020; Hassanin et al., 2021). Thus, any phylogenetic analysis relying on morphology (especially dental characters) to connect extinct taxa to the extant will be hampered by Proteles’ near total lack of postcanine dentition and derived insectivorous lifestyle. This is reflected in the nodal support of the extant taxa to the most closely related “percrocutid” (0.46 PP). However, support does increase through progressive root-ward nodes (0.58-0.75 PP). This resolution of relationships does suggest rapid evolution of the aardwolf worlds insectivory given its most recent common ancestors were well adapted for a durophagous lifestyle. Several internal nodes of the Hyaenidae were recovered with low to moderate support. For example, the relationship between the civet- and mongoose-like taxa Protictitherium and Plioviverrops only received 0.36 PP. This may be related to the species chosen if these genera are well represented in the fossil record, but later occurring than other congeners (Turner et al., 2008; Gracia, 2015). Addition of more of the currently thirteen recognized species of these genera (Turner et al., 2008) will likely increase support and resolution of this region of the 23 hyaenid tree by reducing long branch attraction and maximizing inclusion of transitional morphology. This general suggestion of increased taxon sampling will likely benefit all poorly or moderately supported nodes of the presented phylogeny, especially given the large diversity of extinct hyaenids, approximately one-third of which was sampled for this study (Werdelin and Solounias, 1991; Turner et al., 2008). One of these regions that still contains several evolutionary and taxonomic questions is that of the ‘ictitheres’. Representing a diverse group (~13 taxa) of jackal or wolf-like taxa, ‘ictitheres’ may represent a unique clade characterized by a combination of five synapomorphies: a more anteriorly located bullar partition, the presence of second molars in both upper and lower dentition, enlarged m1 talonid and m2 surface area, an alisphenoid canal, and the lack of an enlarged frontal sinus (Semenov, 1989, 2008). However, all of the above features are ancestral for viverroids (save the bullar partition character) and thus unable to diagnose a clade with a recent common ancestor distinct from living hyaenids. Indeed, Ictitherium viverrinum, namesake of the ‘ictitheres’, was recovered in a position within typical ‘hyaenotheres’ (sensu Semenov, 1989), casting doubt within this analysis of a monophyletic clade for these canid-like taxa. Finally, the last region of the recovered phylogeny that would benefit from increased sampling and investigation is that of stem viverroids. Of those analyzed, the lophocyonids formed the most cohesive group with highly supported internal relationships, but poorly constrained broader placement within Viverroidea. Lophocyonids are feliforms that evolved to be herbivorous browsers with highly complex lophate dentition (Morales et al., 2019). This dental complexity provides numerous morphological characters to resolve intraclade relationships, but basicranial morphology is unknown, creating challenges in determining the most recent common ancestor within the viverroids. Additional taxa such as Lophocyon paraskevaidisi (Koufos et al., 1995) 24 with known rostrum and near complete dentaries may help refine these relationships, but knowledge of the bullar condition from future specimens would likely contribute much more in broader relationships given the synapomorphic disparity of viverroid families for this feature (Hunt, 1987, 1989, 1991). Conversely, a taxon which does present known bullar morphology, could not be confidently resolved. Kanuites lewisae has a mosaic of basicranial features found in both herpestids and viverrids. K. lewisae possesses the viverrid oblique bullar partition, but herpestid + euplerid carotid artery caudal entotympanic posterior entrance (Werdelin, 2019). Furthermore, the temporal occurrence of K. lewisae (14.0-13.4 Ma) is as expected for both stem herpestids and viverrids. However, one illuminating feature that may be determined from CT scanning is the condition of the fossa for the tensor tympani in the petrosal, character 34. In both herpestids and euplerids this fossa forms a delicate tube, compared to a general depression or absence in all other feliforms. This feature may then help resolve the placement of this relatively well-known taxon and inform on greater mosaic evolution of a set of families known for their convergence (Gaubert et al., 2005). Kichechia zamanae was also tentatively placed in the viverroids. Despite substantial fossil material, this hypothesized basal paradoxurine resolved outside both the Viverridae and Herpestidae. Without known basicranial morphology, clade allocation relies upon dental morphology. However, the prominent bunodont cusps of this taxon are known from other viverrid and herpestid taxa such as Hemigalus, Bdeogale and possibly Atilax, while the prominent cusps of the m1 trigonid and talonid including hypoconid and entoconid is ancestral for all viverroids. Additionally, the temporal occurrence of 20.0-17.0 Ma for this taxon is deeper than the inferred split of the Paradoxurinae and Viverrinae (sensu Gaubert and Veron, 2003) at 15.7 Ma. As already discussed, the main issue with placement of all of this study’s stem 25 viverroids is lack of information on non-convergent morphological features (i.e., basicranial anatomy). Unfortunately, what remains is homoplastic or ancestral dental morphology that creates uncertainty in the precise placement of Kichechia zamanae and likely the other stem viverroids such as Herpestides and Kanuites. The phylogenetic history of the Feliformia is relatively recent compared to the deeper Eocene roots of many Caniformia clades (e.g. Hunt, 1996; Nyakatura and Bininda-Emonds, 2012; Paterson et al., 2020) with diversification receiving three main pulses, the end Eocene, end Oligocene, and possibly a third within the last five million years (Figure 1). Even so, feliforms acquired broad morphology and ecology, from likely folivores, to durophages, and sabertooth morphs, the latter of which is not found within caniforms. Furthermore, feliforms, in contrast to caniforms, can be recognized by a distinctive auditory bulla morphology (often distinct per family) and a pattern of ontogenetic development that remains stable over a significant time interval in the middle and late Cenozoic. This basicranial morphology is fruitful ground for assessment of phylogenetic relationships as in the present study, even in the face of rapid evolution and dentognathic convergence of many clades. The present phylogeny used this information to generate a novel set of relationships among extinct taxa, while combining advances in molecular phylogenetic methods to produce the most expansive hypothesis to date. While not comprehensive, this phylogeny does inform on the relationships of “percrocutids” as derived hyaenids, the ancestry of the living African palm civet, Nandinia within the stenoplesictids, the origin of the earliest nimravids, and details of machairodontine tribal evolution. From these inferences, a benchmark for subsequent analyses is possible, but perhaps more importantly, a framework within which to assess aspects of the timing and rates of feliform trait evolution across a time-scaled phylogeny, one that includes thousands of posterior sampled 26 trees to incorporate phylogenetic uncertainty. To put it simply, we have barely begun to scratch the cat tree. 27 Chapter III Ecological Evolution in the Feliformia Introduction The uneven distribution of niche occupation across the tree of life has been a central question of evolutionary biologists (Dobzhansky, 1951; Hutchinson, 1959). Adaptive radiations, in which a key adaptive feature allows a clade to speciate rapidly, have often been used to explain the dominance of members of a single lineage in a particular ecological role (Simpson, 1944, 1953; Schluter, 2000). Opportune events are thought to create high rates of evolution for a clade early in its history, evolutionary rates that quickly diminish as an ecospace is saturated. This pattern is typified by “early burst” models of evolution (Harmon et al., 2010), one among many growing analytical tools to assess evolutionary patterns. However, recent literature suggests early burst evolution is primarily observed within higher taxonomic levels while other forces operate upon lower ones, such as selection towards optimal phenotypic values or Brownian motion over geologic time (Slater, 2015; Law et al., 2018a; Slater and Friscia, 2019). Yet, even within an adaptive radiation there appears to be limitations on what niches are filled/created. For example, why are there not more grazing ecologies amongst bovids and cervids following their late Miocene radiations, particularly with the spread of C4 grasslands (Cantalapiedra et al., 2013)? Horses famously developed traits that facilitated their grazing ecology through the Cenozoic (Matthew, 1926), including tall-crowned teeth (hypsodonty) with complex grinding surfaces, and a single toe (monodactyly) on an elongate limb. Though these traits may not be directly caused by expanding Cenozoic grasslands (Hansen, 1997; McHorse et 28 al., 2019), the comparative dearth of bovids and cervids with similar morphology and grazing ecology raises the question of why these clades did not radiate into this ecospace. One possible explanation for biased ecospace exploitation relates to diet. It has been shown that there is a dietary transition bias associated with mammals, likely related to physiological challenges of switching to certain food types and energetic constraints of a given diet and body mass (Carbone et al., 1999; Price et al., 2012; Pineda-Munoz et al., 2016; Reuter, 2021). For example, the transition from herbivore to omnivore is far more common than omnivore to carnivore, and non-specialized insectivory is only feasible for an organism up to approximately 21 kg (Carbone et al., 2007). This latter point relates to a general decrease in dietary breadth at larger body size, where energetic constraints prevent exploitation of certain resources beyond a given size (McNab, 1986). Furthermore, phylogenetic baggage may not allow specific morphologies to efficiently utilize a given portion of ecospace, or at all. Even if a hypothetical phenotype can be produced it may not be actualized through functional restrictions, such as non- overlapping whorls in the shells of bivalves to produce a working hinge between the valves (Raup, 1966). Amongst carnivorans, Law (2021) found patterns in head-body elongation ratios showed best- support for clade-based optimality shifts, and not models grounded in ecological traits, such as diet, locomotion or hunting behavior. Overall, living carnivorans have similar values in body shape, but certain clades possess clade-based radiations into stout (hyaenids and ursids) or elongate forms (weasels, genets and seals). This is suggestive of contingent opportunities leading to access of new ecospace with its own optimal trait value, and not that of rapid/broad occupation of ecospace that quickly decelerates. Conversely, Slater and Friscia (2019) found support for early burst patterns across living Carnivora for certain dental traits associated with 29 food processing, but not for body mass nor metrics of mechanical advantage in craniodental morphology. Together, this supports a decoupling, or differential selection, of size and shape evolution across Carnivora, and the importance of phylogenetic history In this study, I test for the presence of adaptive radiations in body size and dental toolkit amongst living and fossil feliforms. The carnivoran clade Feliformia is an example of a group that has experienced a presumed bias in ecospace exploitation (Van Valkenburgh, 2007). Specifically, felids and nimravids seem to acquire their characteristic ‘cat-like’ morphology (e.g. foreshortened face, reduced dentition specializing in soft-flesh, retention of supinating/pronating forelimbs) in the earliest fossil forms and remain relatively unchanged until the present, or until their extinction, in the case of nimravids (Werdelin et al., 2010; Barrett, 2021). Why felids and nimravids did not come to occupy additional ecospace (save sabertooth ecomorphology), such as omnivory or even durophagy (bone-cracking as in hyenas) has remained a question many evolutionary biologists and paleontologists have asked (Radinsky, 1982; Holliday and Steppan, 2004; Van Valkenburgh, 2007; Chamoli and Wroe, 2011). The present study seeks to assess the evolutionary patterns of the Feliformia, and well-represented fossil clades. Were felids and nimravids examples of adaptive radiations that quickly occupied their adaptive zones, and if so, how did this differ from other feliform groups, such as hyenas, which occupied a far greater set of dietary and morphological ecologies? Furthermore, do small feliforms exhibit constraint in their evolution or facilitation because of the metabolic and functional discussion above? Below, I assess the patterns of living and extinct feliform ecological evolution with comparisons to differential evolutionary models. These include adaptive radiation, stabilizing selection, clade- based shifts and stochastic evolution. Finally, I offer a process-based framework for these patterns as a narrative of feliform evolutionary history. 30 Methods Data Collection I collected morphological measurements from fossil and living feliform specimens, as well as published values, from museums and literature as listed in Appendix C. Measurements were taken using Mitutoyo digital calipers to 0.01mm precision. Where possible, only wild-caught specimens were measured, though dental measurements were taken from some zoo animals for certain taxa to increase sample sizes. From these measurements, I calculated four ecomorphological traits (Figure 3, Table 3) that have been shown to describe morphology associated with food acquisition and processing in previous studies (Friscia et al., 2007; Meachen-Samuels and Van Valkenburgh, 2009; Slater, 2015; Slater and Friscia, 2019). Body masses (in grams) for living taxa were taken from the panTHERIA database (Jones et al., 2009), while I estimated body masses for fossil taxa from lower m1 lengths, using Van Valkenburgh’s (1990) family-specific regression equations. Fossil hyenas posed their own unique problems, for the small number of living taxa do not express the entire ecological or morphological diversity of extinct members of the family (Werdelin and Solounias, 1991). For these extinct ecomorphs I used the ecological classification of Coca-Ortega and Pérez-Claros (2019) and analogous family- specific regression equations (e.g. canid, viverrid, bone-crushing hyaenid) of Van Valkenburgh (1990). Finally, for certain extinct taxa, robust reconstructions of body mass are available (Christiansen and Harris, 2005; Wheeler and Jefferson, 2009; Palmqvist et al., 2011), in which case, I used these values with source cited in Appendix C. I then converted mass estimates to a linear scale by taking their cube roots and subsequently natural log-transformed them for analysis. This was done to avoid negative values and conflict with R functions throughout the analysis. 31 Figure 3. Measurements taken for ecomorphological traits. A: occlusal view of dentary and teeth. B: lateral view of dentary. C: occlusal view of upper dentition. PMW, lower fourth premolar width; PML, lower fourth premolar length; BL, blade length of lower first molar (trigonid); M1L, lower first molar length; dL, dentary length; C1, upper canine length. Modified from Slater and Friscia (2019: Fig. 1). 32 Table 3. Ecomorphological traits of analysis with abbreviations and constituent measurements. Abbreviation Meaning C1 Compression of the upper canine, measured at the dentine-enamel junction as the mediolateral width divided by anteroposterior length P4S Robustness of the lower fourth premolar, measured as the maximum mediolateral width divided by maximum anteroposterior length RBL Relative blade length of the lower carnassial, measured as the anteroposterior ratio of trigonid length to overall m1 length M1BS m1 blade size relative to jaw length, measured as m1 trigonid length divided by length of dentary from anterior symphysis to posterior most portion of articular process The natural logarithm of the cube root of mass, in grams I classified each feliform species to one of five dietary/ecological categories (hypocarnivore, mesocarnivore, hypercarnivore, durophage, or sabertooth) using linear discriminant analysis (LDA) implemented in the ‘MASS’ (Venables and Ripley, 2002) package for R v. 4.0.3 (R Core Team, 2020). The training set consisted of the four ecomorphological traits listed in Table 3 (C1, P4S, RBL, M1BS) measured for 134 extant taxa (Appendix H). Non-feliform species were included because fossil feliforms exhibit more extreme adaptions to hypo- and mesocarnivory than is realized in the extant radiation. Extant caniform data was supplemented with extinct species from Slater and Friscia (Slater and Friscia, 2019). The first three dietary categories were 33 assigned to extant taxa using the cutoffs of Van Valkenburgh (Van Valkenburgh, 2007) and the Elton Traits dataset (Wilman et al., 2014) checked against the recent carnivoran ecological analysis of Hopkins et al. (2021). Hypercarnivorous feliforms were considered as having ≥70% vertebrate material in the diet, mesocarnivore = 50-69% and hypocarnivore ≤40% vertebrate material, modified from ≤30% vert. of Van Valkenburgh (2007). Crocuta, Hyaena, and Dinocrocuta were assigned to durophages, as were “transitional and fully developed bone- crackers” from Coca-Ortega and Pérez-Claros (2019). Dinocrocuta has much literature justifying a durophagous ecology (Tseng, 2009; Tseng and Binder, 2010), while the Percrocuta taxa of this study do not and were thus classified using the canonical variates analysis of Coca-Ortega and Pérez-Claros (2019). Scimitar- and dirk-tooth taxa were assigned as sabertooth in the training set, while transitional scimitar- or dirk-tooth taxa were assigned by the LDA. Due to the fragmentary nature of much fossil material, I performed a series of discriminant analyses, iteratively reducing the number of variables to maximize species coverage. Classifications for highly fragmentary taxa were consistent with expectations based on congeners. The predicted categories for extinct taxa are available in Appendix H. Analytical Methods To understand the dynamics of body mass and carnassial evolution, I fit models of continuous trait evolution to a random sample of 500 trees from the post-burnin posterior distribution of the phylogenetic analysis of Chapter II. Constant-rate Brownian motion, temporally accelerating/decelerating rate (early burst), and trended random walk models were fitted using the ‘fitContinuous’ function in geiger (Pennell et al., 2014). I additionally fit two multipeak Ornstein-Uhlenbeck (OU) models. The first was an ecological OU model using the mvMORPH package (Clavel et al., 2015), while the second was a phylogenetic regime shift model using the 34 bayou package (Uyeda and Harmon, 2014). I limited the assessed OU evolutionary models to those possessing constant α (strength of pull) and σ2 (evolutionary rate) terms; even though variable rate models do exist, simulations have shown unreliable inference for these models with phylogenies with a limited number of tips (Beaulieu et al., 2012). Measurement error was accounted for by including trait standard error in all evolutionary models. Because sample sizes were small for some species, I supplemented values for living taxa from Slater and Friscia (2019), while for poorly sampled extinct taxa I assigned each species a standard error of 0.0345, following Harmon et al. (2010). This was also the case for body mass standard error for all species, given that information is not included in the panTHERIA database. Relative model fit was assessed by computing small sample corrected Akaike Weights, AICcW. This was done for the entirety of the Feliformia, and subset analyses for the Felidae, Nimravidae, Hyaenidae and small feliforms. Bayou uses a reversible-jump Markov chain Monte Carlo (MCMC) to fit multipeak OU models to estimate the placement and magnitude of regime shifts along lineages. This differs from the mvMORPH OU model by not assessing convergent evolution, but each shift is considered a unique adaptive regime (Uyeda and Harmon, 2014). Priors (Table 4) differed amongst the clades analyzed given the morphological breadth observed for body mass and RBL values. 35 Table 4. Priors for bayou analyses. Model parameter α = strength of pull, σ2 = evolutionary rate, θ = optimal value. *= Calculated using ‘cdpois’ option in bayou. RBL = relative blade length of the lower m1. Model Feliformia body Feliformia RBL Felidae body Felidae RBL Nimravidae body Nimravidae RBL Parameter mass mass mass α Half-cauchy Half-cauchy Half-cauchy Half-cauchy Half-cauchy with Half-cauchy with with scale factor with scale factor with scale factor with scale factor scale factor 1 scale factor 1 1 1 1 1 σ2 Half-cauchy Half-cauchy Half-cauchy Half-cauchy Half-cauchy with Half-cauchy with with scale factor with scale factor with scale factor with scale factor scale factor 0.1 scale factor 0.1 0.1 0.1 0.1 0.1 θ Normal Beta distribution Normal Beta distribution Normal Beta distribution distribution with with shape 1 = 5 distribution with with shape 1 = 5 distribution with with shape 1 = 5 standard and shape 2 = 1 standard and shape 2 = 1 standard and shape 2 = 1 deviation = deviation = deviation = 0.5, 1.3*sd of mass 0.75*sd of mass mean = mean of data, mean = data, mean = mass data mean of mass mean of mass data data 36 Table 4. Continued for Feliformia, Felidae and Nimravidae. Model Parameter Feliformia body Feliformia RBL Felidae body Felidae RBL Nimravidae Nimravidae RBL mass mass body mass Number of shifts Fixed at one Fixed at one Fixed at one Fixed at one Fixed at one Fixed at one per branch Branch-wise Uniform Uniform Uniform Uniform Uniform Uniform shift probability Number of shifts Conditional Conditional Conditional Conditional Conditional Conditional Poisson Poisson Poisson Poisson Poisson Poisson distribution* distribution* distribution* distribution* distribution* distribution* with mean = with mean = with mean = with mean = with mean = with mean = 0.1*number of 0.1*number of 0.1*number of 0.1*number of 0.1*number of 0.1*number of edges on edges on edges on edges on edges on edges on phylogeny and phylogeny and phylogeny and phylogeny and phylogeny and phylogeny and maximum = number of tips of maximum = maximum = maximum = maximum = maximum = number of tips of number of tips of number of tips of phylogeny 2*number of phylogeny 2*number of phylogeny phylogeny edges of edges of phylogeny -2 phylogeny -2 Location of shift Uniform Uniform Uniform Uniform Uniform Uniform along branch 37 Table 4. Continued for Hyaenidae and small feliforms. Model Hyaenidae body mass Hyaenidae RBL Small feliform body mass Small feliform RBL Parameter α Half-cauchy with scale Half-cauchy with scale Half-cauchy with scale factor Half-cauchy with scale factor 1 factor 1 1 factor 1 σ2 Half-cauchy with scale Half-cauchy with scale Half-cauchy with scale factor Half-cauchy with scale factor 0.1 factor 0.1 0.1 factor 0.1 θ Normal distribution Beta distribution with Normal distribution with Normal distribution with with standard deviation shape 1 = 5 and shape 2 = standard deviation = 0.75*sd standard deviation = = 0.3, mean = mean of 1.5 of mass data, mean = mean 0.75*sd of RBL data, mean mass data of mass data = mean of RBL data Number of Fixed at one Fixed at one Fixed at one Fixed at one shifts per branch Branch-wise Uniform Uniform Uniform Uniform shift probability Number of Conditional Poisson Conditional Poisson Conditional Poisson Conditional Poisson shifts distribution* with distribution* with mean = distribution* with mean = 6 distribution* with mean = mean = 0.1*number of 0.1*number of edges on and maximum = number of 0.1*number of edges on edges on phylogeny phylogeny and maximum tips of phylogeny and maximum = and maximum = = 2*number of edges of phylogeny. 2*number of edges of number of tips of phylogeny -2. phylogeny -2. phylogeny. Location of Uniform Uniform Uniform Uniform shift along branch 38 I ran two independent MCMC chains with 1.5 million generations, each sampled every 1,000 generations for the full Feliformia phylogeny. These family-level phylogenies ran for 500,000 generations, with sampling at every 500. Convergence was assessed using Gelman and Rubin’s R statistic via the ‘gelman.R’ function in bayou (Uyeda and Harmon, 2014). R values <1.1 were considered to have reached convergence, and samples prior to this were discarded as burn-in. Effective sample sizes were greater than 200 for all analyses, and frequently over 1,000. Only evolutionary shifts with a posterior probability (PP) above 0.5 using parameters averaged from the two chains were considered in subsequent analyses. To compare the results of the bayou model to that of the likelihood-based geiger and mvMORPH models, I converted the bayou output using the ‘bayou2OUwie’ function (Uyeda and Harmon, 2014). Clades identified via bayou for adaptive shifts were then assessed by a comparable OUM model in mvMORPH. The relative AICc weights of all models were then compared for best fit. Results Feliformia The distribution of ecological variables can be seen in Figure 4. An Ornstein–Uhlenbeck (OU) for multiple dietary optima is the best-fitting evolutionary model for both body mass and RBL for the Feliformia, (Table 5, median AICcW = 0.49 for body mass and 1.0 for RBL). Body mass sees a general increase in optimal value progressing from hypocarnivore to sabertooth ecology, with the largest jump between hypercarnivore and durophagous. Carnivores are predicted to switch to hunting of large vertebrate prey at around 14.5-21 kg (Carbone et al., 2007), with prey of equal or larger mass than their own at a subsequent 21.5-25 kg (Carbone et al., 1999). The 39 hypercarnivore optimum is here inferred to be 11.32 kg. This was smaller than that observed by Slater (Slater, 2015) for living and extinct canids (median = 20.7 kg). A smaller optimal mass of 1.28 kg was estimated for hypocarnivory, but a larger mass of 16.55 kg for mesocarnivory (Table 5). However, both durophagous and sabertooth ecologies returned comparable and very large optimal body masses, with 173.62 kg for the former, and 179.11 kg for the latter. The α-parameter of OU models describes the strength of attraction to the associated optima, with higher values indicating a stronger pull. This value can be reparametrized in terms of phylogenetic half-life, t1/2= ln(2)/α, which describes the time required for adaptation to a new selective regime to outpace implied constraints of an ancestral regime (Hansen, 1997). The phylogenetic half-life of the median estimated α-parameter for body mass is approximately 10 million years (My). This implies strong and rapid selective pressure towards an optimal body mass given a dietary category. Relative blade length of the m1 is also best explained by an adaptive peak model (median AICcW = 1.0, Table 5). This model predicts a median optimal relative blade length of 0.50 for hypocarnivorous feliforms, while mesocarnivorous feliforms are attracted to a slightly greater relative blade lengths of 0.54. Hypercarnivorous, durophagous, and sabertooth feliforms are all attracted to relative blade lengths of approximately 1.0. These latter two dietary ecologies in fact returned optimal values greater than 1.0. Though biologically impossible, the assessed OUM model could not be constrained for possible optimal ranges, and thus suggests even stronger attraction to total slicing component of the m1 than is observed in hypercarnivorous feliforms that do not have durophagous or sabertooth morphology. 40 Figure 4. Distribution of ecological proxy variables used in the presented analysis, natural log of the cube root of body mass in grams, left, and the relative blade length of the lower m1, right. Ancestral states for both body mass and relative m1 blade length were estimated using the ‘fastANC’ function found in ‘phytools’ (Revell, 2012) on the MCC tree of Chapter II. Silhouettes taken from phylopic.org or created by the author. 41 Likewise, to optimal body mass values, the median estimated α-parameter (0.040) for relative m1 blade length implies strong selection for these values, but not as strong as bodymass, t1/2= 17.3 My. However, this value does approximate the divergence of major feloid clades, 15-31 Ma, suggesting the primary pull of RBL selection began at the clade’s appearance and has been sustained since. Felidae In contrast to overall feliform evolution, felid body mass was best explained by a model of Brownian motion (AICcW = 0.62). This was approximately three times higher in weight than the next best supported model, trended evolution, AICcW = 0.20, while an early burst model received only slightly less support than this, AICcW = 0.18. Both OUM models were the poorest supported with no node returned 0.5 or greater PP for the bayou analysis. RBL evolution was best explained by an early burst model (AICcW = 1.0). This suggests that initial high rates of evolution for a clade exponentially decay over time as a new niche becomes available, but then is quickly saturated (Blomberg et al., 2003; Harmon et al., 2010). The scalar value for this model (r = -13.98) describes the deceleration of evolution, as expected by a hypothesis of adaptive radiation. The very large and negative value gives a feel for the rapidity of this burst, one that is several orders of magnitude greater than found in estimates of overall feliform RBL evolution under an early burst model, and indeed in large clades of other analyses (Harmon et al., 2010). 42 Table 5. Median results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter II. Mass has been converted to kgs. OUMbayou did not recover a shift at any location with over 0.5 PP for RBL. Trait Model lnLk AICc AICcW σ2 Scalar Hypo- Meso- Hyper- Durophagous Sabertooth carnivore carnivore carnivore BM -33.76 71.62 0.02 0.013 - - - - - - ACDC -33.32 72.85 0.13 0.019 -0.012 - - - - - Trend -33.98 74.16 0.01 0.013 0.002 - - - - - OUMdiet -22.80 62.89 0.49 0.018 0.067 1.28 16.55 11.32 173.62 179.11 OUMbayou -31.96 76.66 0.34 0.014 0.018 - - - - - RBL BM -33.76 71.62 <0.01 0.013 - - - - - - ACDC 145.17 -284.12 <0.01 7.5-4 -0.013 - - - - - Trend 145.79 -285.37 <0.01 5.2-4 0.002 - - - - - OUMdiet 159.32 -301.29 1.0 0.018 0.040 0.50 0.54 0.97 1.02 1.09 OUMbayou - - - - - - - - - - 43 Table 6. Median Felidae results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter II. Mass has been converted to kgs. OUMbayou did not recover a shift at any location with over 0.5 PP for body mass. Trait Model lnLk AICc AICcW σ2 Scalar Hypercarnivore Sabertooth BM -8.00 20.43 0.62 0.014 - - - ACDC -8.05 22.98 0.18 0.011 0.016 - - Trend -8.06 23.00 0.20 0.014 -0.002 - - OUMdiet -8.21 38.97 <0.01 0.017 0.030 5.63 1788.15 OUMbayou - - - - - - - RBL BM -7.98 20.39 <0.01 0.014 - - - ACDC 6808.59 -13610.30 1.0 7.6-218 -13.98 - - Trend 5213.70 -10420.50 <0.01 7.1-218 -2.5-18 - - OUMdiet 297.69 -572.83 <0.01 0.016 0.112 1.0 1.0 OUMbayou 462.74 -913.09 <0.01 1.8-33 0.112 - - 44 Nimravidae Similar to felids, the best supported evolutionary models for nimravid body mass and RBL was Brownian motion and early burst respectively, Table 7. However, the relative support of a Brownian motion model for body mass over the next best supported model was not as great as in the Felidae, BM AICcW = 0.52, ACDC = 0.32. Additionally, the sigma squared parameter, which can be interpreted as the net rate of evolution (Harmon et al., 2010) was the same as in both nimravid and felid analyses. This suggests similar overall rates of body mass evolution even over different persistence times for each clade (Nimravidae = 34.3 Ma, Felidae = 18.9 Ma). RBL evolution for nimravids is also best explained by an early burst model (AICcW = 1.0). The rate parameter for this model is even greater than that inferred for felid evolution (r = -16.68), implying a very rapid decrease in evolutionary rates beyond an initial adaptive radiation. Hyaenidae Hyaenid ecological evolution returned similar results to the overall Feliformia, such that body mass and RBL evolution was best supported by an OU model for multiple dietary optima (Table 8), median AICcW = 0.95 for body mass and 0.48 for RBL). Body mass also sees an increase in optimal value progressing from hypocarnivore to durophagous ecology compared to overall Feliformia. The hypercarnivore optimum for hyaenids is inferred to be 25.46 kg. This is quite similar to that observed by Slater (Slater, 2015) for canids as mentioned previously, but the additional optimal values for hypo- and mesocarnivorous hyaenids (median optima = 3.19 kg, 16.11 kg respectively) are also quite similar to that observed by Slater for living and fossil canids, (hypo. = 3.2 kg and meso. = 4.7 kg). Durophagous hyaenids returned the largest optimal mass at 64.98 kg, similar to living Crocuta but less than that estimated in the overall feliform analysis. The phylogenetic half-life of the α-parameter for body mass returned the shortest value 45 of all analyses at 1.96 My. This implies extremely strong selective pressure towards an optimal body mass given a hyaenid dietary category. Relative blade length of the m1 is also best explained by an adaptive peak model (median AICcW = 0.48, Table 8), but only moderately better supported than the next best model, early burst (AICcW = 0.27). The adaptive peak model predicts a median optimal relative blade length of 0.69 for hypocarnivorous hyaenids, while mesocarnivorous hyaenids are attracted to substantially smaller relative blade lengths of 0.44. Hypercarnivorous and durophagous hyaenids are all attracted to relative blade lengths closer to 1.0 (hyper. = 0.78, duro. = 0.86), but substantially less in value than seen in all other analyses. The α-parameter for RBL values also displayed a high attractive force (median = 0.348), t1/2= 1.99 My compared to body mass estimates, but slightly smaller. Small feliforms Unique amongst all analyses, small feliforms found best support for Brownian motion in both RBL and body mass evolution, Table 9. The relative support of these models was about twice as great as the next best models, ACDC for the former and trended evolution for the latter. The sigma squared parameter for body mass evolution (0.008) was less than that in both nimravid and felid analyses, about half as large. This suggests that body mass evolution occurs at a slower pace amongst small feliforms, though expected given their overall amount of variance in this variable. The sigma squared value for RBL evolution (0.008) is about half of that inferred for other analyzed clades, also implying relatively slower rates of evolution for the dental toolkit. 46 Table 7. Median Nimravidae results from macroevolutionary model fits to 500 random pruned trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter II. Mass has been converted to kgs. OUMbayou did not recover a shift at any location with over 0.5 PP for body mass or RBL. Trait Model lnLk AICc AICcW σ2 Scalar Hypercarnivore Sabertooth BM -3.82 12.06 0.52 0.014 - - - ACDC -3.76 14.37 0.32 0.018 -0.023 - - Trend -3.64 14.14 0.15 0.014 0.013 - - OUMdiet -1.19 24.64 <0.01 0.023 0.139 3.89 61.44 OUMbayou - - - - - - - RBL BM -3.82 12.06 <0.01 0.014 - - - ACDC 938.19 -1869.38 1.0 2.0-71 -16.677 - - Trend 55.36 -103.73 <0.01 1.3-4 0.005 - - OUMdiet 54.22 -98.71 <0.01 0.023 0.034 - 1.0 OUMbayou - - - - - - - 47 Table 8. Median Hyaenidae results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2. Mass has been converted to kgs. OUMbayou did not recover a shift at any location with over 0.5 PP for body mass or RBL. Trait Model lnLk AICc AICcW σ2 Scalar Hypo- Meso- Hyper- Durophagous carnivore carnivore carnivore BM -3.38 11.36 0.03 0.012 - - - - - ACDC -3.52 14.29 0.01 0.017 -0.041 - - - - Trend -2.79 12.85 0.01 0.011 0.021 - - - - OUMdiet 7.06 12.16 0.95 0.023 0.353 3.19 16.11 25.46 64.98 OUMbayou - - - - - - - - - RBL BM -3.38 11.36 0.25 0.012 - - - - - ACDC 24.41 -41.41 0.27 0.003 -0.135 - - - - Trend 23.03 -38.66 <0.01 0.001 0.006 - - - - OUMdiet 34.36 -40.73 0.48 0.023 0.348 0.69 0.44 0.78 0.86 OUMbayou - - - - - - - - - 48 Table 9. Median small feliform results from macroevolutionary model fits to 500 random trees from the stable Bayesian posterior of the phylogenetic analysis in Chapter 2. Mass has been converted to kgs. OUMbayou did not recover a shift at any location with over 0.5 PP for body mass or RBL. Trait Model lnLk AICc AICcW σ2 Scalar Hypo- Meso- Hyper- carnivore carnivore carnivore BM -8.29 20.96 0.51 0.008 - - - - ACDC -8.39 23.58 0.29 0.008 -3.0-4 - - - Trend -7.52 21.84 0.18 0.008 -0.011 - - - OUMdiet -4.18 30.13 0.01 0.015 0.094 1.73 19.85 1.83 OUMbayou - - - - - - - - RBL BM -8.29 20.96 0.46 0.008 - - - - ACDC 30.23 -53.66 0.22 0.001 0.0134 - - - Trend 31.02 -55.25 0.30 0.001 -0.004 - - - OUMdiet 35.62 -49.48 0.03 0.015 0.142 0.59 0.65 0.79 OUMbayou - - - - - - - - 49 Discussion Overall, feliform ecological evolution is pulled towards optima related to dietary category. However, clade-specific patterns suggest far more nuance in evolutionary history. Both nimravids and felids experienced an adaptive radiation (early burst) with entrance into hypercarnivore and sabertooth ecologies, while body mass evolution of these clades is best supported by a random walk over geologic time. The processes behind adaptive radiations are often hypothesized to relate to the development of ‘key innovations’ or contingent historical events that facilitate diversification by creating new ecological opportunities (Schaeffer, 1948; Van Valen, 1971; Gould, 1989; Sanderson and Donoghue, 1994; Hunter and Jernvall, 1995; Slater, 2015). Though it may be more straightforward to correlate the acquisition of a given morphologic feature with an adaptive radiation, the nature of contingent events implies chance opportunities such as sweepstakes dispersals or climatic change that generates a land bridge due to falling sea levels. These are events that if we were to “rewind the tape” (Gould, 1989) may not happen again. However, there are theoretical frameworks that, although unable to predict a given climatic or tectonic event, offer utility in predicting the biotic response to such a perturbation. For example, under Habitat Theory (Vrba, 1992) most speciation and extinction events are correlated to global tectonic and/or climatic changes, while common climatic fluctuations typically drive geographic habitat tracking as species move with their biome if possible. Furthermore, at times of global warming with a strong latitudinal thermal gradient, Habitat Theory predicts higher speciation rates at lower latitudes and higher extinction rates at higher latitudes. This principle is related to the habitat preference of specialist clades where resources would tend to disappear during recurrent environmental extremes. Though challenged for certain African Pliocene taxa (Bobe et al., 2002; Werdelin and Lewis, 2005; Frost, 2007; Faith and 50 Behrensmeyer, 2013), Habitat Theory may still provide a useful framework for global dynamics of the Cenozoic. One assumption in the application of Habitat Theory to the fossil record is that clades, generally speaking, have maintained habit preference, or in other words, that the fundamental habit of a clade has changed minimally over geologic time. Most living feliforms, including felids, hyenas, mongooses, civets, and genets are found in tropical regions, while most terrestrial caniforms reside within temperate or seasonal climatic regions (Pickford and Morales, 1994; Hunt, 1996). This differential climatic signal is even found in the cranial shape of feliforms versus caniforms, where mean precipitation and temperature are significant predictors of cranial shape in feliforms, but not caniforms (Tseng and Flynn, 2018). The divergence time of the most recent common ancestor (MRCA) of nimravids for the phylogeny presented in Chapter II is suggested to be at approximately 41.3 Ma. Even the earliest nimravids are known to possess hypercarnivorous dentition, but based on phylogenetic analysis, and the results of the presented chapter, quickly derived this dental condition from more generalist feliforms present in the middle to early Eocene (Solé, 2014; Tomiya and Tseng, 2016; Barrett et al., 2021). The above MRCA date occurs slightly after the Middle Eocene Climatic Optimum (Zachos et al., 2001). This period of global warming is associated with an increase in hypercarnivory amongst mammals in North America (Tomiya et al., 2021), but also in the earliest Asian nimravid material (Chow, 1958; Peigné et al., 2000; Averianov et al., 2016). Nimravids likely originated in Asia, as did the earliest feliforms (Hunt, 1996, 1998a). If nimravids were also tropical specialists, the spread of this clade into North America would be predicted by Habitat Theory. As the thermal gradient extended northward during the Middle Eocene Climatic Optimum (Figure 5), nimravids would have found dispersal easier via northern passages into North America. Tropical specialists would 51 also benefit from increased speciation rates during this time, as seen in Figure 1, though it is unclear if this provided the impetus for the rapid development of hypercarnivory within this clade and non-carnivorans. Hypercarnivory amongst mammals appears to have experienced a process of slow assembly within the Cenozoic. One that didn’t develop the familiar secant dentition and reduced tooth count until the Early to Middle Eocene in oxyaenodontan, hyaenodontan and carnivoramorphan taxa (Borths et al., 2016; Solé and Ladevèze, 2017). However, hypercarnivorous members of the above clades all seem to have rapidly acquired this dental toolkit during this epoch. Nimravids arrival in North America is also associated with rapid acquisition of sabertooth morphology compared to Asian ancestors, e.g., Maofelis. Compressed and serrated canines are known from the Hancock mammal quarry of Oregon (~40 Ma, Hanson, 1996), and a c.f. Hoplophoneus maxilla (San Diego Natural History Museum 60554) from the Pomerado conglomerate of San Diego (ca. 38-37 Ma). These first nimravids would have encountered large hyaenodont and mesonychid hypercarnivores (e.g. Hyaenodon, Hemipsalodon, Harpagolestes). Perhaps the rapid acquisition of sabertooth morphology allowed nimravids a way to access a greater variety of prey resources while staying relatively small in the presence of potentially competing non-carnivoramorphan hypercarnivores. What’s more, following the loss of sabertooth machaeroidines prior to the Middle Eocene Climatic Optimum (Zack et al., 2022), an opportunity may have presented itself for a new lineage to create or occupy a similar ecology in North America. 52 Figure 5. Latitudinal boundary shift of tropical and boreal climate zones from the Eocene to the present. Climate gradient modified from Pickford and Morales (1994), Fig. 5. Global climate data from Zachos et al. (2008), where absolute temperatures were converted from δ18O using the equation of Epstein et al. (1953). Climatic events are annotated above temperature curve while biogeographic events below curve. Note global geography depicts the modern arrangement, approximately Miocene to present. 53 Though unanalyzed at the clade level for ecological evolution, stenoplesictids are the earliest known feloids (save perhaps palaeogalids with a Chadronian occurence, Famoso and Orcutt, 2022) with a Late Eocene occurrence in Mongolia (Dashzeveg, 1996; Hunt, 1998a). The diversity and morphology of this clade is poorly known until the start of the Oligocene when a diversity of forms arise and spread into Europe. In fact, the earliest known feloids of the Oligocene are found at Quercy (France) at the Eo-Oligocene transition along with nimravids (Hunt, 1998a). This sudden European appearance is likely related to the Grande Coupure, an event with substantial faunal turnover and associated immigration of Asian taxa into Europe. The Grande Coupure was during a time of global cooling associated with Antarctic ice sheet formation (Zachos et al., 2001). Costa et al. (2011) summarize several theories for this event and the replacement of European taxa, but sea level drop with the formation of the Antarctic icesheet likely facilitated a land bridge to Europe to aid the dispersal of Asian taxa. Stenoplesictids are also known in Mongolia post-Grande Coupure, along with palaeogalids and nimravids, though there is a distinct lag (~2 Ma) between the first Oligocene geographic radiation and following feloid diversification (Dashzeveg, 1996; Hunt, 1998a). The latitude of both France and Mongolia are approximately at 45 degrees north, being substantially outside the tropics of today, combined with the rapid global cooling at the formation of Antarctic ice sheets, it would be expected under Habitat Theory to see a retraction of tropical specialists to the equator during this time, while biome generalists would be relatively unaffected. However, recent data suggests that high- latitude temperatures may have been almost as warm as those pre-Antarctic glaciation (O’Brien et al., 2020). This is hypothesized to be in part related to the lack of a northern ice sheet. Additionally, the East Asian record of mid-late Eocene and early Oligocene carnivoramorpha is quite poor (Ducrocq et al., 1995; Wang et al., 2007; Böhme et al., 2013). This suggests the taxa 54 that did immigrate into Europe during the Grande Coupure were relatively unencumbered by the global cooling at this period, either to being biome generalists or being in a high-latitude climate warmer than previously hypothesized but did not experience the boost in speciation rates associated with a warming climate as predicated under habitat theory. In fact, an increase in overall feloid diversity isn’t seen for two million years after the event which does follow a rapid warm excursion immediately after the initial cold excursion of the Eo-Oligocene transition (Zachos et al., 2001). The Oligocene saw rapid diversification of nimravid species (Barrett, 2021), but also in those of stenoplesictids and palaeogalids (Hunt, 1998a; Peigné, 1999; Peigné and De Bonis, 1999; Hunt, 2001a; Welsh, 2021; Famoso and Orcutt, 2022). Ecologically, save the living African Palm Civet (Nandinia), all analyzed stenoplesictids are hypothesized to be hypercarnivorous feliforms, though not derived to the degree seen in contemporary nimravids, nor in later hyaenids or felids. This suggests that the global faunal turnover of the Eo-Oligocene transition may have provided a large opportunity (contingent event) for Feliform diversification, as supported by the nimravid- specific RBL analysis of this chapter, though not yet assessed at the clade level for stenoplesictids nor palaeogalids. The end of the Oligocene saw rapid global warming that was sustained into the early Miocene. Much like the Grande Coupure before it, another mass immigration event occurred, the African- Eurasian interchange. This event saw the immigration of African endemic species out of Arica, such as proboscideans, while Eurasian taxa including carnivorans and ungulates into Africa. However, there is a bias in what species immigrated and subsequently diversified. For example, feliforms were the dominant carnivorans to immigrate into Africa, while caniforms are only represented by two species (Pickford and Morales, 1994). For African immigrants, 55 proboscideans, hyracoids and aardvarks were the only groups to find substantial success when leaving Africa, likely beginning in the latest Oligocene (Lucas and Bendukidze, 1997; Antoine et al., 2003; Sen, 2013). Habitat Theory would predict large-scale immigration of tropical biome specialists (i.e., most feliforms) to the newly accessible landmass with the same fundamental habitat within which they currently reside, especially during a period of global warming (Figure 5). It would also predict the exclusion of boreal, or Palearctic in this case, species from Africa, possibly related to differential reproductive strategies as influenced by seasonality and day length changes. At high latitudes, temperature and day length fluctuations are the greatest signals of seasonality, while at low latitudes these same fluctuations are minor and humidity changes dominate, lending the tropics to wet/dry seasonality (Pickford and Morales, 1994). Nimravids were perhaps the first carnivorans that arrived in Africa, presumably from Asia, given the lack of known European nimravid material of this age (Friscia et al., 2020; Barrett et al., 2021; Werdelin, 2021). However, a diversity of stenoplesictid, and stem feloid taxa also immigrated to Africa around the same time (Pickford and Morales, 1994; Morales and Pickford, 2021). This period (~25-20 Ma) correlates with an increase in speciation rate in the skyline plot of Figure 1; the divergence of hyaenids from herpestid/euplerid ancestors, the origin of felids, the origin and diversification of lophocyonids, and initial diversification of the Barbourofelini. In short, a great diversification of feliform taxa occurred resulting in the origin of most living families. By 16 Ma, the earth would enter the Middle Miocene Climatic Optimum (Figure 5), the last excursion into a warming planet before a steady decline into the cooler and more familiar northern and southern icesheet driven system of the present (Zachos et al., 2001). With the rise in global temperatures, one would expect a correlated rise in feliform diversity under Habitat 56 theory. In fact, the phylogeny of Chapter II (Figure 2), recovers this as the MRCA for the extant radiation of viverrids and herpestids + euplerids, and the divergence of machairodontine felids from felines + pantherines. These values are further supported by the analysis of Slater and Friscia (2019) on a more inclusive taxon set of extant species. Globally, there is also a large increase in Mammalian diversity during the middle Miocene, including that of feliform taxa (Alroy, 1992; Pickford and Morales, 1994; Morales and Pickford, 2011; Grohé et al., 2020; Wang et al., 2020; Morales and Pickford, 2021). The present analysis found felid dietary evolution was best supported by an early burst model (Table 6); this was also the case for RBL evolution of viverrids and euplerids in Slater and Friscia (2019), inclusive of almost all living species. The latter finding for euplerids is perhaps no surprise as they are one of the most exemplary clades for contingency, with a highly supported sweepstakes dispersal to Madagascar from the African mainland (Yoder et al., 2003). Thus, it is no additional surprise that nearly all variables analyzed by Slater and Friscia (2019) found an early burst model as the best supported for euplerid evolution. The remaining feliform clades, herpestids and hyaenids, show patterns of ecological evolution better explained by other processes. Herpestids have the most recent origin among the feliform clades, likely in Africa with a single Asian immigration event (Wesley-Hunt et al., 2010; Zhou et al., 2017). Slater and Friscia (2019) found both RBL and body mass evolution in herpestids was best supported by an OU model of evolution, suggesting there is an optimal value the clade was either constrained to, or pulled towards, and not an adaptive radiation. Furthermore, there appears to be no obvious correlation in global temperature at the extant herpestid origin (Zachos et al., 2001). Similarly, hyaenids seem drawn towards optimal body mass and dental toolkits, though the present analysis finds these values are correlated to a given dietary category. This is 57 the same result that was found by Slater (2015) for canid evolution, though a durophagous category was not included in that study. The analogy between canid and hyaenid evolution has been made before, where hyaenids are the Old World ecological analogs to the New World canids (Hunt, 1996; Van Valkenburgh, 2007; Tseng and Wang, 2011), and thus the results of this work offer a new line of inference on convergent evolution between these clades. Both hyaenids and canids found remarkably similar optimal body mass (save mesocarnivory) per dietary category (Table 10). However, the inferred phylogenetic half-life for these optima differed by two orders of magnitude for hyaenids and canids, 1.96 Ma to 936 Ma respectively. For hyaenids this suggests an incredibly strong pull towards optimal values within a given category, but for canids, such weak attraction was better interpreted as a slow, sustained trend in increased body mass across all three dietary categories, an assessment supported by the next best supported model of Slater (2015), trended random walk. Relative blade length of m1 was not assessed by Slater (2015), but relative lower grinding area (RLGA) was, being another proxy for dental toolkit, and by extension diet. Higher values for this proxy indicate a greater grinding proportion (compared to slicing), and unsurprisingly optimal values from hypo- to hypercarnivory show a progressive decline in RLGA. Hyaenids show a similar pattern in reducing the grinding component with increased hypercarnivory. However, values for hypo- and mesocarnivory in hyaenids seemed backwards, such that hypocarnivory had greater optimal RBL values than mesocarnivory. This may be explained by the trigonid not necessarily being dedicated to a slicing function amongst carnivorans and instead being modified to a grinding or puncturing (in the case of arthropod food items) component like the talonid behind it. Thus, retaining a large trigonid, as in Plioviverrops, would be functionally advantageous. Closer inspection of the total grinding versus slicing component of the dental toolkit in hyaenids and 58 canids shows distinct differences. Specifically, there is a substantially greater grinding component retained in hypercarnivorous canids than hyaenids, even though the pull towards each clade’s optimal dietary value was comparably strong, t1/2= 1.99 Ma for hyaenids and 3.85 Ma for canids. This distinction may result from phylogenetic constraint unique to each clade, especially in the context of dental morphology and diet (Hopkins et al., 2021). This constraint has been hypothesized for durophagy in the two clades where the retention of posterior molars may have resulted in the shift of bone-crushing teeth/cusps in borophagine canids to the anterior P4 and lower m1 compared to that of non-carnassial premolars in hyaenids whom otherwise lose the posterior molars (Tseng and Wang, 2011). Table 10. Median optimal body mass (in kg) values from model fits of 500 random trees from the stable Bayesian posterior of the phylogenetic analysis for hyaenids of the present analysis and that of canids in Slater (2015). Hypocarnivore Mesocarnivore Hypercarnivore Hyaenidae 3.19 16.11 25.46 Canidae 3.19 4.71 20.70 A core concept of adaptive radiations has been an initial rapid diversification and phenotypic evolution that “...fit the divergent environments they exploit” (Schluter, 2000). I subsequently assessed this evolutionary pattern via early burst models of decelerating rates of phenotypic evolution. The underlying process behind these patterns is primarily explained by the work of Simpson (1944, 1953) where high rates of evolution are required to move through nonadaptive realms of the adaptive landscape to new stable realms. This view that evolution precedes through 59 ‘quantum’ (Simpson, 1944), ‘punctuated’ (Eldredge and Gould, 1972) or only minimally stable states has existed since at least the work of Francis Galton (1869), who analogized evolution proceeding as a polyhedron flipping from one stable facet to another. Within each adaptive zone, Simpson suggested the existence of adaptive subzones giving opportunities for fine-scale ecological diversification. For example, felids and canids occupy their own unique subzones which can be grouped into a more inclusive ‘fissiped’ subzone as opposed to one of pinnipeds, and even more inclusively, all of the aforementioned can be grouped into carnivores as opposed to herbivores. Slater (2015) suggests that canids can be subdivided into canid hypo-, meso- and hypercarnivore subzones within which have their own optimal state. OU models can be viewed as evolutionary constraint, which in this context could be interpreted as a constraint in the dental morphology required to occupy a given dietary niche or subzone. The lack of an early burst signal for canids was suggested by Slater to be a result of early burst or adaptive radiations only existing at more inclusive clades. In fact, Slater and Friscia (2019) found that among extant Carnivora there was a preponderance of early burst dynamics, but primarily in only diet/dental related traits. At the family level this falls apart into far greater nuance of Brownian motion or OU models as best supported, a hierarchical distinction previously suggested in theoretical literature (Osborn, 1902; Simpson, 1944, 1953; Gould, 2002; Humphreys and Barraclough, 2014). The presented results suggest that feliform ecological evolution may best be explained as a pull towards optimal dental toolkits, and to a lesser extent body mass, per dietary category. However, these optima are filtered through contingent events, likely correlated to global climatic and/or tectonic dynamics. Nimravids and felids explosively entered a nimravid/cat-like zone that ecologically may have prevented these clades from escaping (i.e., evolutionary ratchet, Van Valkenburgh et al., 2004) 60 related to phylogenetic constraints inherent to the clade. These constraints, much like tooth count in canids versus hyaenids, may have been contingent in themselves, such that the path to hypercarnivory from a hypo- or mesocarnivorous ancestor has several routes, e.g., cat/nimravid versus dog/hyaenid, and that one path may be more restrictive than the other. Canid and hyaenid hypercarnivores utilize more of the carcass than felids based on observation of living taxa (Van Valkenburgh, 1996) and dental microwear and enamel structure in fossil forms (Tseng, 2012; Desantis et al., 2015). The consumption of bone or otherwise resilient vertebrate material in addition to soft flesh may have facilitated the retention or development of robust molars in canids and bone-cracking premolars in hyaenids. Thus, the difference in what parts of vertebrate prey are consumed may be a more important distinction than how much vertebrate prey is consumed and the resulting evolutionary patterns. For canids, this could provide an escape to a more omnivorous diet and the ability to specialize in a given dietary subzone. For felids and nimravids, explosive entry into soft flesh hypercarnivory may have been irreversible, a potential generalization for mammals (Brocklehurst, 2019). Even with findings that mammal cusp number and placement are under the control of relatively limited genetic pathways allowing potentially a diverse array of dental complexity from early crown developmental (Jernvall and Jung, 2000; Kangas et al., 2004; Kavanagh et al., 2007; Harjunmaa et al., 2014), the loss of talon(ids) is seemingly inflexible to recovery (Solé and Ladevèze, 2017). Rapid felid and nimravid hypercarnivorous shifts seem to be tied to global climatic/tectonic dynamics, in turn associated with faunal turnover. These turnover events may provide contingent opportunities for soft-flesh specialist hypercarnivores to quickly exploit new resources by rapid dental toolkit specialization, but ones that constrain descendant taxa to remain within this ecospace. 61 The ecological evolution of hyaenids and canids saw exploration of multiple subzones, one of which was hypercarnivory. This path was still rapid given the estimated phylogenetic half-life values of the present study and that of Slater (2015), but it does not seem correlated with the type of global events that habitat theory would predict. Hyaenids may occupy a common canid-like subzone of the Neogene where optima from a hypo- or mesocarnivorous ancestor mediated the pull towards each dietary category, compared to a singular pull to extreme hypercarnivory. A possible explanation for this may lie in the postcranial bauplän of felids/nimravids versus canids and hyaenids. Early members of both the Canidae and Hyaneidae exhibit incipient cursorial morphology, including reduced ability to retract the distal phalanges and supinate/pronate the forelimb (Hunt Jr. and Solounias, 1991; Wang, 1993; Figueirido et al., 2015). These changes likely enhanced efficiency in covering large distances (Janis and Wilhelm, 1993), but also reduced the ability to grapple with prey, as do modern felids. Becoming increasingly face- orientated in predation, canids and hyaenids may have relied more upon the integration and optimization of body size and dental tool kit than do soft-flesh specialists and thus set them down a distinct ecospace trajectory. A third situation is seemingly possible in the case of small feliforms, where body mass and dental toolkit evolve via random walk. This third subzone of small carnivorans and associated dietary subzones, achieves broad dietary ecologies, possibly related to their small size (Pineda- Munoz et al., 2016). Evolutionary patterns of small feliforms may correlate to the morphological refuge discussed for canids and hyaenids, in that a hypocarnivorous (and specifically insectivorous) ancestor may better facilitate evolution into varied dietary categories at small size. This bias in dietary shifts across mammalian evolution has been recovered in the work of Price et al. (Price et al., 2012) and Reuter (2021), where far greater dietary transitions occur from diets 62 that feature invertivory than any other. This malleability in ecology is likely facilitated by being small, where metabolic and structural burdens have minimal impact compared to mammals of larger size (Carbone et al., 1999, 2007). The current study suggests that the hypothesis of early burst evolution being only observable at higher taxonomic levels, while stabilizing selection or Brownian motion dominates lower levels is overly simplistic. Instead, there appears to be a story of contingent opportunity mediated by global tectonic and climatic factors, but these too are filtered through phylogenetic baggage accrued through entrance into different ecological zones. Soft-flesh specialists rapidly appear at times of massive faunal turnover, from lineages with plesiomorphic carnivoramorphan postcranial morphology. However, the descendants of these lineages are constrained to remain within this ecospace with the loss of dental hardware lending itself to different diets. Climatic/tectonic events also facilitate the diversification and appearance of other feliform clades, but the evolutionary patterns expressed differ from those of felids and nimravids. Hyaenids experienced vast ecological diversity optimized for body mass and dental toolkit. This ecospace trajectory may have been necessitated by the early postcranial modifications to cursoriality in the clade and correlated face-centric predation. With the loss of this ancestral postcranial morphology, both hyaenids and canids may have been excluded from soft-flesh specialization, even if the opportunity existed, such as the “cat-gap” in the early Miocene of North America. However, retention of more generalized dentition combined with cursorial adaptations may channel body size and dental tool kit optimization per dietary category. Finally, small feliforms experience apparent random evolution in both body size and dentition. These clades’ appearance and diversification can also be correlated with global tectonic and climatic events, but given the absolute size of their members, are free to drift across a range of ecological 63 options, being freed of the constraints of large size. However, once opportunities do present themselves the above ecospace trajectories come into play with growing body size and postcranial adaptations. For example, the fossa (Cryptoprocta ferox) acquired familiar felid ecomorphology in Madagascar after a sweepstakes dispersal of a herpestid-like ancestor from Africa. Further analysis within a paleoecological framework may find more similarities than differences in subzone occupation of additional Carnivoran clades. The caniform groups of Mustelidae, Mephitidae, Ailuridae and Procyonidae may occupy the small ecospace of sister group small feliforms, while certain subfamilies of ursids and amphicyonids may fall within that of canids and hyaenids. Regardless, contingent opportunity from global events seems to drive the evolution of feliform carnivorans, but the path they take is mediated by ecological pathways previously taken and consequent phylogenetic baggage. 64 Chapter IV Evolution of feliform cranial shape Introduction The processes that govern the evolution of shape amongst the pageant of life, are central questions in biology (Dobzhansky, 1951; Hutchinson, 1959; Gould, 2002). Studies suggest that there is a variable degree of connection between morphology and ecology (Marroig and Cheverud, 2005; Hunt, 2007; Harmon et al., 2010; Santana and Cheung, 2016; Slater and Friscia, 2019). For example, mammals have a strong association between dental and gnathic morphology and their diet (Evans et al., 2007; Boyer, 2008; Christensen, 2014; Pineda-Munoz et al., 2016; Grossnickle, 2020; Wang et al., 2021). Taxa that consume large amounts of vertebrate material possess reduced dentary lengths and enhanced shearing surfaces to maximize power and efficiency in processing animal tissue, while folivorous mammals lengthen the dentary and increase tooth occlusal area and complexity to aid in grinding of tough floral material. However, how this form and function correlation extends to overall cranial shape has remained less clear (Figueirido et al., 2011; Tseng, 2013; Tseng and Flynn, 2018). While some authors have found a connection between diet and cranial shape (Sacco and Van Valkenburgh, 2004; Wroe and Milne, 2007), others recover only a predictive connection for bite force (Christiansen, 2008b; Maestri et al., 2016; Law et al., 2018b), which when tied to diet, only discriminates between hypercarnivores and generalists in terrestrial carnivorans (Radinsky, 1981b; Rovinsky et al., 2021). For the latter studies, the lack of predictive power for a broad swath of diet (0-70% vertebrate material = hypo-mesocarnivores) may relate to constraint in phylogeny, which, by limiting 65 possible morphologies, creates stochastic variation that disrupts expected mapping of form to function (Raup and Gould, 1974). However, ecomorphological categories have been assessed in the fossil and living record (Martin, 1989; Werdelin, 1996; Van Valkenburgh, 2007; Turner et al., 2008; Coca-Ortega and Pérez-Claros, 2019; Barrett, 2021) by the iterative appearance of certain morphologies over geologic time, and presumably comparable ecologies to which they belong. This observation can be extended to the conclusion that a lineage that entered many dietary ecologies should be more morphologically disparate than one that did not, and by some metrics, more successful (Guillerme et al., 2020). Of the carnivoran clades, low amounts of cranial-dental variation have been historically presented in literature for “cat-like” carnivores such as felids and nimravids (Radinsky, 1981a, 1982; Van Valkenburgh, 1991). Foreshortened faces and reduced tooth counts optimize forces in slicing meat at the scissor-like carnassials and prey seizing with the canines. This reduced raw material for selection is argued to relate to increased specialization and narrow morphological disparity amongst hypercarnivores with concomitant increased extinction risk (Holliday and Steppan, 2004; Van Valkenburgh et al., 2004; Van Valkenburgh, 2007). Specifically, loss of posterior grinding molars in soft-flesh specialist hypercarnivores may have put them on an evolutionary road of no return (evolutionary ratchet), resulting in an ecomorphological constraint that did not allow them to evolve additional ecologies. However, recent 3D Geometric Morphometric (3DGM) analyses suggest that cranial disparity for hypercarnivores may be greater than that of generalists among living carnivorans, particularly amongst feliforms (Michaud et al., 2018, 2020). This disparity may in turn be an underestimate, for living carnivores do not occupy the entire range of morphologies, and presumably ecologies, of extinct lineages. Sabertooth and bone-cracking carnivores were once species-rich components of past ecosystems, and it has been suggested that their cranial 66 morphology may have operated under different rules than their extant relatives in terms of disparity and cranial modularity (Holliday and Steppan, 2004; Goswami et al., 2015). Even with the seeming inability to recover molar crushing basins (Solé and Ladevèze, 2017), there may be another reason for aberrant disparity patterns in the crania of soft-flesh hypercarnivores (i.e. felids and nimravids). Recent literature has shown that morphology is often integrated from developmental genetic sequences (Gehring, 1996; Shubin et al., 1997; Tomarev et al., 1997) to gross anatomy (Goswami, 2006; Goswami and Polly, 2010a; Goswami et al., 2014, 2015) in such ways that structures can covary, generating morphology that changes in sync. Do felids and nimravids present limited evolutionary potential resulting from overly integrated cranial development compared to other clades, or does developmental integration provide a fruitful channel to more extreme morphologies (Goswami et al., 2015)? In this study, I use 3DGM analyses to assess feliform cranial allometry and morphology within and among 73 species spanning approximately 34 million years of evolution. Specifically, I ask the following questions: (1) Are there distinct cranial morphologies for each ecological category? (2) Does disparity differ among these categories? (3) What is the source of disparity in more morphologically diverse clades? (3a) Does morphological disparity increase in clades that have had more time to evolve, or (3b) Does greater variance in size allow a line of least evolutionary resistance for morphological variation via allometry? (3c) If allometry is important, is there a common allometric trajectory amongst feliforms? (4) Does integration (an indicator of developmental constraints), enable or limit potential in the evolution of feliform cranial shape? 67 Methods Materials The morphological dataset is composed of 157 cranial specimens, representing 73 extinct and extant feliform species (Appendix I). When possible, I sampled one male and female individual of a species to approximate the range of sexual dimorphism for that species, while I chose species to sample the cranial disparity of a given clade, including an “average” shape and more outlying morphologies as well. I then sorted these species into ten monophyletic clades as recovered in Chapter II: Felinae/Pantherinae (FP), Machairodontinae, Felidae, Stenoplesictidae, Nimravidae, Viverridae, Hyaenidae, Herpestidae, Eupleridae, and Prionodontidae. I digitized 69 three-dimensional (3D) landmarks (Figure 6, Table 11) using a Microscribe G2LX digitizer (Immersion Corp., San Jose, CA). During data collection I stabilized individual crania with clay supports and digitized landmarks in two sets, one in dorsal view and the other in ventral view. I used a set of four reference landmarks in both views to facilitate alignment of the dorsal and ventral datasets. The two sets were then stitched together using the ‘unifyLandmarks’ function of the ‘StereoMorph’ package (Olsen and Westneat, 2015) in R v. 4.1.2 (R Core Team, 2020). I considered alignments sufficient when alignment error was less than or equal to the manufacturer stated accuracy of the Microscribe G2LX digitizer (0.3mm). I supplemented the Microscribe- generated data set with a few rare taxa/specimens landmarked from published CT scans or photogrammetry-generated 3D models. These latter specimens were digitized in Landmark Editor v. 3.0 (Institute for Data Analysis and Visualization, University of California, Davis). 68 Figure 6. Position of the landmarks taken on the cranium for 3DGM analyses (69 landmarks) to quantify variation in shape of feliform crania. A, dorsal view. B, ventral view. C, right lateral view. Landmarks are subset into two modules, Neural Crest (NC, blue) and Mesoderm (MD, red), based on tissue origin of cranial elements. The skull used for this representation is a leopard (Panthera pardus, Felidae). 69 Table 11. Definition of anatomical landmarks used in the three-dimensional geometric morphometric analyses. Landmark Definition 1 Anterior border of canine at alveolus-left 2 Anterior border of canine at alveolus-right 3 Anterior border of P3 at alveolus-left 4 Anterior border of P3 at alveolus-right 5 Anterior midline suture of premaxilla 6 Anteromedial point of mastoid process-left 7 Anteromedial point of mastoid process-right 8 Anterolateral border of P4 alveolus-left 9 Anterolateral border of P4 alveolus-right 10 Anteromesial border of I3 at alveolus-left 11 Anteromesial border of I3 at alveolus-right 12 Basioccipital basisphenoid bulla suture-left 13 Basioccipital basisphenoid bulla suture-right 14 Basion 15 Basisphenoid basioccipital midline suture 16 Basisphenoid presphenoid suture-left 17 Basisphenoid presphenoid suture-right 18 Bulla anterior medial extreme-left 19 Bulla anterior medial extreme-right 20 Bulla posterior lateral extreme-left 21 Bulla posterior lateral extreme-right 22 Frontal parietal alisphenoid suture-left 23 Frontal parietal alisphenoid suture-right 24 Glenoid fossa lateral extreme-left 25 Glenoid fossa lateral extreme-right 70 Table 12 continued. Landmark Definition 26 Jugal maxilla ventral suture-left 27 Jugal maxilla ventral suture-right 28 Jugal maxilla lacrimal suture-left 29 Jugal maxilla lacrimal suture-right 30 Jugal squamosal ventral suture-left 31 Jugal squamosal ventral suture-right 32 Lacrimal frontal maxilla suture-left 33 Lacrimal frontal maxilla suture-right 34 Mastoid process ventral tip-left 35 Mastoid process ventral tip-right 36 Maxilla-palatine midline suture 37 Maxilla frontal nasal suture-left 38 Maxilla frontal nasal suture-right 39 Medial border of P4 at widest point of alveolus-left 40 Medial border of P4 at widest point of alveolus-right 41 Medial M1 alveolus-left 42 Medial M1 alveolus-right 43 Nasal-anterior midline suture 44 Nasal-premaxilla suture anterior-left 45 Nasal-premaxilla suture anterior-right 46 Nasals frontal midline suture 47 Occipital condyle lateral extreme-left 48 Occipital condyle lateral extreme-right 49 Parietal-Frontal midline suture 50 Parietal-squamosal-alisphenoid suture-left 51 Parietal-squamosal-alisphenoid suture-right 52 Parietals occipital midline suture 53 Paroccipital process tip-left 71 Table 13 continued. Landmark Definition 54 Paroccipital process tip-right 55 Posterior border of canine at alveolus-left 56 Posterior border of canine at alveolus-right 57 Posterior border of palatine at midline 58 Posterior of carnassial at alveolus-left 59 Posterior of carnassial at alveolus-right 60 Posterolateral border alveolus M1-left 61 Posterolateral border alveolus M1-right 62 Posterodorsal tip of occiput at midline 63 Posterolateral border of I3 at alveolus-left 64 Posterolateral border of I3 at alveolus-right 65 Postorbital process base (jugal)-right 66 Postorbital process base (jugal)-left 67 Postorbital process tip (frontal)-left 68 Postorbital process tip (frontal)-right 69 Premaxilla maxilla ventral midline suture Because the crania of extinct taxa were often incompletely preserved, I imputed missing landmarks to maximize the size of the dataset. Two additional fossil specimens, TMM 933-3444 (Homotherium serum) and CB 07 (Eusmilus sicarius) were retrodeformed to correct for postdepositional shape changes using the ‘symmetrize’ function of the ‘Morpho’ R package (Schlager, 2017). I first performed imputation via reflection across the plane of symmetry using the ‘mirrorfill’ function in the ‘paleomorph’ package (Lucas and Goswami, 2017). These mirrored models I then used to impute absent landmarks by the ‘estimate.missing’ function in the ‘geomorph’ package (Adams et al., 2021) with the thin-plate spline method. Imputation 72 proceeded in three iterations to maximize accuracy, given the ‘estimate missing’ function interpolates landmarks using a reference specimen within the analyzed dataset. Thus, felids and nimravids were grouped for imputation, with a second group composed of durophagous hyaenids, and a final group containing the remaining taxa (i.e., viverroids). This final group included the hyaenid genera: Ictitherium, Hyaenotherium and Tungurictis. While paraphyletic, these hyaenids and other viverroid families exhibit similar dolichocephalic cranial shapes. The digitized landmarks for all crania I then subjected to Generalized Procrustes Analysis (GPA) superimposition which translates the specimens to a common origin, scales each specimen to unit centroid size, and rotates each specimen to minimize landmark distances using a least-squares calculation. The resulting Procrustes coordinates I projected into tangent space with a principal components analysis (PCA) using the ‘gm.prcomp’ geomorph function to identify the major axes of cranial shape among all the crania in the analysis. I identified morphological changes associated with the major PC axes by calculating a series of indices of individual crania and then examined their Pearson’s correlation to said axes. Disparity and Integration I calculated disparity as the Procrustes variance (trace of the group covariance matrix, divided by group n) of each clade of interest using the ‘morphol.disparity’ function in geomorph v. 4.0.1 (Adams et al., 2021; Baken et al., 2021) with 1000 randomized residual permutations (Collyer et al., 2015). I performed this calculation while assessing the impact of allometry and clade age on disparity, because certain feliform clades span orders of magnitude in body mass while others do not. Thus, strong allometric trends may provide a line of least evolutionary resistance towards generating disparity in clades with large size variance. Similarly, morphological disparity for a clade may simply be the result of having more time to accrue within relatively ancient lineages 73 compared to recently diverging ones. I further calculated disparity per dietary category (Table 12) as determined in Chapter III, while adding extant invertebrate specialists as those with ≥70% invertebrate material in their diet as determined by the Elton Trait dataset (Wilman et al., 2014), checked against the recent carnivoran ecological analysis of Hopkins et al. (2021). I compared ecological cranial disparity by using the ‘morphol.disparity’ function and model coords ~ log(Csize) with 1000 randomized residual permutations. Randomization of residuals allows assessment of model effects and summary statistics, especially important for the high dimensionality of trait data versus sample size of 3DGM analyses (Adams, 2014a; Adams and Collyer, 2015). The Bonferroni correction accounted for multiple comparisons (Bonferroni, 1936). Table 14. Definitions of dietary categories for analysis of cranial shape. Dietary Category Definition Invertivore ≥70% invertebrate material Hypocarnivore ≤40% vertebrate material Mesocarnivore 50-69% vertebrate material Hypercarnivore ≥70% vertebrate material Durophagous Hypercarnivore with bone-cracking morphology Sabertooth Hypercarnivore with sabertooth morphology I calculated cranial integration two ways. The first measure used was eigenvalue dispersion (equations 7&8, Goswami and Polly, 2010b), equal to the standard deviation of eigenvalues from the correlation matrix of the PCA. With high values of eigenvalue dispersion, variance will be concentrated in the first few eigenvectors resulting from high covariance between landmarks. 74 The second measure calculated the integration of the mesoderm (MD) and neural crest (NC) developmental modules. I determined module assignment for each landmark by the tissue origin of the bone on which a given landmark was located (Figure 6). The origins of cranial bones, as neural crest or mesoderm derived, were assigned following Ferguson and Atit (2019) for the face and vault elements, and McBratney-Owen et al. (2008) for the cranial base. The squamosal is derived from both Neural crest and Mesoderm tissues (Ferguson and Atit, 2019), thus landmarks 30-31 were treated as NC, while 24-25 were assigned to MD. Landmark 49 was located on the sutural boundary between the NC and MD modules and was assigned to the MD module, as the coronal suture is of mesoderm origin (Mishina and Snider, 2014). I assessed integration of these two developmental modules with the ‘integration.test’ function geomorph. This is a two-block partial least squares analysis (PLS, Rohlf and Corti, 2000), also referred to as a singular warps analysis (Bookstein et al., 2003). The z-scores were used for clade comparisons while significance was derived from 1000 random permutations of individuals in one partition to those in the other. I tested the relationship between morphological disparity and integration by performing two sets of regressions using the base R function lm(). I regressed morphological disparity against both eigenvalue dispersion values and against between‐module integration (z‐scores) of the MD and NC regions. Allometric Variation I assessed the impact of allometry on clade evolution in two ways. The first looked at per-clade influence of allometry on shape variation by using the geomorph function ‘procD.lm’ and the equation (Procrustes coordinates) ~ log(centroid size)+RCS. Relative canine size (RCS) was 75 included in particular to assess its role in predicting cranial shape in sabertooth carnivorans (Slater and Van Valkenburgh, 2008). Significance was evaluated with Goodall’s (1991) F-test with 1,000 permutations for this and the subsequent analyses. For the second set of analyses, variation in evolutionary (among-clades) allometries was tested using an ANCOVA model where Procrustes shape coordinates were regressed against log centroid size and clade membership with the model coords ~ log(Csize) * clade. A post hoc test using the package RRPP (ver. 1.1.2; Collyer and Adams, 2018, 2019) function pairwise evaluated whether the evolutionary allometric slopes of clades (n =10) significantly differ from one another. Multiple comparisons were accounted for by reducing α to .01. The model was visualized by plotting the regression scores of shape on size versus log centroid size (Drake and Klingenberg, 2008). I also compared these results with a phylogenetic ANCOVA (pANCOVA) using geomorph’s procD.pgls (Adams, 2014a), which executes the ANCOVA model in a phylogenetic framework. This pANCOVA used mean averaged crania from all 73 species and a pruned version of the phylogenetic tree of Chapter 2 (Figure 7) that only included taxa sampled for this analysis. Phylogenetic Signal and Evolutionary Rates I calculated the phylogenetic signal of the cranial landmark dataset with a multivariate K-value (Kmult) (Adams, 2014b) using the ‘physignal’ function in geomorph. Kmult uses 1000 random permutations to assess how well the morphometric data fit a phylogenetic tree. A Kmult value of zero corresponds to absence of phylogenetic signal, whereas a Kmult value greater than one indicates a strong phylogenetic signal, implying that morphological traits are conserved within the phylogeny. 76 Figure 7. Pruned MCC phylogeny of Chapter II to include only those feliforms analyzed in the present cranial morphology chapter. I calculated the morphological rates of evolution for each clade of interest by using the geomorph function ‘compare.evol.rates’ on the same pruned tree. I performed 1000 simulations of Brownian motion trait evolution which were summarized with the net evolutionary rate of each clade obtained from the sum of squared distances between taxa and the origin of their clade (O’Meara et al., 2006; Adams, 2014c). This procedure was also performed on mean natural log centroid sizes for species crania to estimate rates of cranial size evolution. I compared the clade- wise rates of size and shape evolution and morphological disparity with ordinary least squares regression analysis in the base R function lm(). I further compared rates of evolution per dietary category making pairwise comparisons (over 1000 iterations) of shape evolution rates between all dietary categories. The Bonferroni correction accounted for multiple comparisons (Bonferroni, 1936). 77 Dietary Impact I tested the effect of diet on feliform cranial shape by first calculating the degree of carnivory (proportion of vertebrate material in the diet) for extant taxa, taken from the Elton Traits (Wilman et al., 2014) database and checked against Hopkins et al. (2021). I then regressed this value against relative blade length (RBL) of the lower first molar (taken from the dataset in Appendix H) and estimated the degree of carnivory for extinct taxa using the base lm() and predict() functions in R. Degree of carnivory for all taxa was then regressed against mean cranial shape for feliforms using the geomorph procD.pgls function within a phylogenetic framework. Results PC1 (Figure 8) accounts for 44.68% of feliform shape variation and describes morphological changes associated with sabertooth morphology, with progressively more derived sabertooth taxa towards the negative side of the axis. This is echoed by the correlation of relative canine size (distance from landmark 1 to 55 divided by centroid size) to the PC1 axis, (r = -0.87, p = 2.2x10- 16). PC2 (14.57% of variation) describes changes from brachycephalic to dolichocephalic morphology, reflected in the correlation of relative palate length (distance from landmark 5 to 57 divided by centroid size) to the PC2 axis, (r = 0.70, p = 2.2x10-16). Every other PC axis describes less than 10% of remaining cranial shape variation. Within the PC 1 and 2 morphospace, all viverroids save hyaenids overlap each other with positive scores on the PC1 axis and positive to slightly negatives scores on the PC2. Hyaenids occupy an area near the origin, but primarily extend into positive PC2 morphospace. Felines and pantherines extend the furthest into negative PC2, with the largest members also having the most negative PC1 scores. The aforementioned 78 taxa generally occupy an upper right to lower left orientation in the PC1 and 2 morphospace while extending perpendicular to them lie the entirety of the sabertooth machairodontine and nimravid taxa. This is a similar pattern to that recovered in the 2D GM analysis of Slater and Van Valkenburgh (2008). Figure 8. PCA of all cranial feliform specimens. Convex hulls surround clades of interest while individual taxa are indicated by a surface scan of their place in morphospace. Below and to the left side of PC1 and 2 are wire frame models of the change in landmarks at the extreme of each principal component from a right lateral and dorsal perspective. 79 Procrustes ANOVA Results Clade membership, log centroid size, and relative canine size are significant predictors of cranial shape, though the influence on each clade of the latter two variables differs, Table 13 and Table 14. Felines and pantherines had by far the greatest allometric signal (49%) while nimravids had the least (9%) and almost all other clades (save herpestids) had an intermediate signal between 20 and 26%. Relative canine size was minimally explanatory for cranial variation in most clades (3-7%) except for the euplerids (47%) and sabertooth machairodontines (19%) and nimravids (24%). Table 15. Allometry and effect of relative canine size (RCS) on cranial shape as determined by ANOVA of shape (Procrustes coordinates) ~ log(centroid size)+RCS for each clade. Disparity was calculated by Procrustes variance adjusted for allometry and clade age. Statistically significant (p <0.05) values are in bold. Eigenvalue dispersion does not include a significance assessment. ln Centroid RCS Disparity Eigenvalue Developmental Size dispersion module integration Eupleridae 0.204 0.466 0.012 0.408 1.909 Felinae/Pantherinae 0.487 0.035 0.013 0.167 4.708 Herpestidae 0.119 0.070 0.008 0.236 3.522 Hyaenidae 0.246 0.043 0.016 0.218 3.013 Machairodontinae 0.259 0.185 0.020 0.250 3.763 Nimravidae 0.090 0.238 0.017 0.236 3.411 Viverridae 0.256 0.071 0.011 0.229 2.459 80 Table 16. Evolutionary allometry ANCOVA. Evolutionary allometry (among clades) uses the mean shapes and mean centroid sizes of the 73 species, which were then grouped into ten clades (including the one monospecific lineage). Df = degrees of freedom, SS = sum of squares, MS = mean square, R2 = coefficient of determination, F= F statistic, Z= effect sizes, Pr(>F) = significance value. Df SS MS R2 F Z Pr(>F) log(Csize) 1 0.871 0.871 0.271 120.64 5.158 <0.001 clade 9 1.159 0.129 0.361 17.842 12.253 <0.001 log(Csize):clade 9 0.193 0.021 0.06 2.968 5.48 <0.001 Residuals 137 0.989 0.007 0.308 Total 156 3.21 Levels of Morphological Disparity and Integration Age of lineage had a significant correlation with allometrically adjusted disparity (Figure 9). Adjusting Procrustes variance for both allometry and lineage age found machairodontines with the highest level of morphological cranial disparity (Table 13), followed by nimravids and hyaenids. Felines/pantherines, euplerids and viverrids possessed moderate levels of disparity, while herpestids had the least overall. Disparity per dietary category (Table 15) recovered sabertooth taxa having the greatest amount of size-adjusted variance. This is followed by durophagous taxa in second; mesocarnivores, hypercarnivores and invertivores with comparable intermediate values, and hypocarnivores with the least. However, Bonferroni-corrected significance values only recover sabertooth and hypocarnivores as significantly different in disparity. 81 Table 17. Cranial disparity (Procrustes variance adjusted for size) per dietary category. Upper triangle is absolute difference between dietary categories while lower triangle contains p-values. Hypocarn. Mesocarn. Hypercarn. Durophag. Sabertooth Invertivor. Hypocarn. - 0.0062 0.0058 0.0086 0.0136 0.0044 Mesocarn. 1 - 3.00E-04 0.0024 0.0074 0.0018 Hypercarn. 1 1 - 0.0028 0.0077 0.0014 Durophag. 1 1 1 - 0.0049 0.0042 Sabertooth 0.015 1 0.105 1 - 0.0091 Invertivor. 1 1 1 1 0.495 - Integration, as determined by both eigenvalue dispersion and developmental module integration, has no significant correlation to allometric and lineage age adjusted disparity (Figure 10 and Figure 11). Eigenvalue dispersion was similar for all clades save euplerids, which possessed a value almost twice as great as other feliforms. However, felines/pantherines had the greatest between-module integration value. There was also a greater spread of this integration metric for other clades, with the sabertooth clades, hyaenids, and herpestids having moderate values, and viverrids and euplerids the least overall. Phylogenetic Signal and Evolutionary Rates The phylogenetic signal of feliform cranial shape is K = 0.59 (p < 0.001). This is substantially greater than that observed across extant Carnivora in Michaud et al. (K = 0.38: 2018), though less than that of extant feliforms in Michaud et al. (K= 0.78: 2020), implying that feliforms have a more conserved ancestral cranial morphology than carnivorans overall, and that fossil taxa may have a disproportionate impact on this calculation. Clade-wise rates of size evolution (Figure 12) 82 Figure 9. Relationship between feliform lineage age and disparity (Procrustes variance). Disparity has been adjusted for allometry. 83 Figure 10. Relationship between feliform cranial integration (Eigenvalue dispersion) and disparity (Procrustes variance). Disparity has been adjusted for allometry and age of lineage. 84 Figure 11. Relationship between feliform cranial integration (between developmental module) and disparity (Procrustes variance). Disparity has been adjusted for allometry and age of lineage. poorly predicted rates of morphological shape evolution (R2= -0.08, p< 0.52). Of note, machairodontines and nimravids appear as outliers, having the greatest rates of shape evolution, but even with their removal the fit is non-significant (R2= 0.46, p< 0.08). For the rates of size and shape evolution, only that of shape evolution significantly predicted cranial disparity (Figure 12). Pairwise analysis of shape evolution rates amongst dietary categories revealed that crania of sabertooth carnivores evolved 2.2-5.1 times faster than those of non-sabertooths (Table 16; p < 85 0.02). A similar result was found for durophagous and hypercarnivorous carnivores relative to hypocarnivorous ones, with a value 1.9 times (p < 0.02) faster for the former, and 1.74 times (p < 0.02) faster for the latter. All other pairwise comparisons were nonsignificant. Allometric Variation Feliform cranial morphology exhibits substantial evolutionary (among-clades) allometry (R2 = 0.27, p<0.001). This is less than that of clade membership (R2 = 0.36, p<0.001) indicating strength in phylogenetic signal over a common allometric trajectory (Table 14). The ANCOVA of the interaction between clade membership and log centroid size further shows very little support for distinct allometric trends per clade (R2 = 0.06, p<0.001). However, the pANCOVA (Table 17) of mean shapes against size returned contradictory results with size being the only significant variable accounting for 7% of variation. All other variables were insignificant within this phylogenetic framework. Figure 13B shows the similarity in evolutionary trajectories for each clade, which when taken with the post hoc test for homogeneity of slopes found that, out of 55 pairwise comparisons, only two had significant differences in slopes (Table 18). The significantly different comparisons are the felines/pantherines to the hyaenids (p<0.002) and nimravids as compared to the machairodontines (p<0.004). Dietary Impact The regression of RBL on degree of carnivory returned a significant correlation (R2= 0.55, p< 2.2e-16). However, the phylogenetic ANOVA of cranial shape on diet was not significant R2= 0.02, p<0.12). Figure 14 shows the relationship between these variables, where from 0 to 70% vertebrate material in the diet, feliforms possess a narrow range of cranial shapes, while at ≥ 80% feliforms occupy a very diverse range. This pattern is reflected in the cranial disparity 86 Figure 12. Relationship between feliform size and shape evolutionary rates, and morphological disparity (Procrustes variance). Disparity has been adjusted for allometry and age of lineage. 87 Table 18. Pairwise cranial shape evolution rates between dietary ecologies in upper triangle, with p-value significance in lower triangle. Hypocarn. Mesocarn. Hypercarn. Durophag. Sabertooth Invertivor. Hypocarn. - 1.29 1.74 2.36 5.07 1.81 Mesocarn. 1 - 1.35 1.82 3.92 1.4 Hypercarn. 0.015 1 - 1.35 2.91 1.04 Durophag. 0.015 0.06 0.944 - 2.15 1.3 Sabertooth 0.015 0.015 0.015 0.015 - 2.8 Invertivor. 0.03 1 1 1 0.015 - (Procrustes variance) of species with ≥80% vertebrate material in their diet being greater than that of those ≤ 70%, 0.014 versus 0.009. Furthermore, phylogenetic signal was greater for those species with ≤ 70% vertebrate material in their diet (K= 0.6587, p< 0.001), while less for those with ≥80% (K= 0.4003, p< 0.001). Discussion Ecological cranial shape and disparity The PCA of cranial morphospace (Figure 8) and phylogenetic regression of degree of carnivory on cranial shape (Figure 14) found broad connections to ecology, but not the degree reported in some past analyses (Wroe and Milne, 2007; Goswami et al., 2011). There is a distinct region of morphospace occupied by sabertooth feliforms, but almost the entirety of hypo-mesocarnivores, i.e., herpestids, viverrids, euplerids, stenoplesictids, prionodontids overlap each other. What remains are two other distinct regions, occupied by hyaenids and felids respectively (with some overlap from sabertooth clades) indicating that phylogeny might have substantial input on morphospace occupation compared to dietary category in most cases among feliforms. 88 Figure 13. Log centroid size versus the regression scores of shape and size for each specimen (A). Idealized allometric trajectories predicted values for each clade, highlighting similarities in evolutionary allometric slopes (B). 89 Table 19. Evolutionary allometry pANCOVA. Csize = centroid size, Df = degrees of freedom, SS = sum of squares, MS = mean square, R2 = coefficient of determination, F= F statistic, Z= effect sizes, Pr(>F) = significance value. Df SS MS Rsq F Z Pr(>F) log(Csize) 1 0.006 0.006 0.073 5.216 3.506 <0.001 clade 9 0.006 0.001 0.07 0.553 -2.666 <0.994 log(Csize):clade 8 0.009 0.001 0.102 0.91 -0.132 <0.55 Residuals 54 0.064 0.001 0.756 Total 72 0.085 This lack of convergence of skull shape on diet or ecology is reflected in the regression of degree of carnivory, where there exists a narrow range of cranial shapes for the vast majority of the dietary spectrum, and only within the realm of hypercarnivory do cranial shapes occupy new and more variable zones. This result is more aligned with recent studies that were unable to find significant correlation between cranial shape and diet among carnivorans (Law et al., 2018b; Rovinsky et al., 2021). The present results support a decoupling of cranial shape and diet until hypercarnivory is reached. The vast majority of diets are served by a narrow cranial shape, or “one-to-many” ecological mapping. This dietary regime does coincide with changes in dental morphology, implying dentition is far more labile relative to a species’ environment than what its skull looks like. Crania within this range are thus little altered (or subject to stabilizing selection) from ancestral morphology as indicated by the stronger phylogenetic signal (K=0.66), with observed variation likely the outcome of phylogenetic stochastic processes (Raup and Gould, 1974). 90 Table 20. Pair-wise homogeneity of slope test summary of p values for distinct evolutionary allometric trajectories. Significantly different comparisons are in bold. Fel Pan = Felinae + Pantherinae, Machair. = Machairodontinae, Prionodont. = Prionodontidae, Stenoples = Stenoplesictidae Eupleridae Stem FelPan Herpestidae Hyaenidae Machair. Nimravidae Prionodont. Stenoples. Viverridae Felid Eupleridae 1 0.94 0.745 0.383 0.982 0.97 0.926 0.95 0.473 0.257 Stem Felid 0.94 1 0.353 0.681 0.347 0.986 0.669 0.942 0.936 0.605 FelPan 0.745 0.353 1 0.311 0.002 1 0.515 0.9 0.854 0.663 Herpestidae 0.383 0.681 0.311 1 0.976 0.944 0.138 0.936 0.703 0.964 Hyaenidae 0.982 0.347 0.002 0.976 1 0.822 0.09 0.928 0.411 0.659 Machair. 0.97 0.986 1 0.944 0.822 1 0.004 0.948 0.998 0.99 Nimravidae 0.926 0.669 0.515 0.138 0.09 0.004 1 0.91 0.341 0.499 Prionodont. 0.95 0.942 0.9 0.936 0.928 0.948 0.91 1 0.95 0.936 Stenoples. 0.473 0.936 0.854 0.703 0.411 0.998 0.341 0.95 1 0.313 Viverridae 0.257 0.605 0.663 0.964 0.659 0.99 0.499 0.936 0.313 1 91 Figure 14. Relationship between degree of carnivory and feliform cranial shape (PGLS). Mean cranial shape (in right lateral and dorsal views) of species with ≤ 70% vertebrate material in their diet below and ≥80% above. Presented K values indicate associated phylogenetic signal for the cranial shapes of those species. Hypercarnivores, on the other hand, occupy a diverse range of cranial shapes that likely are needed to subdue and consume a proportionally diverse set of prey or animal materials. This can be viewed as “many-to-one” mapping of morphology to ecology, with the “one” truly referring to the narrow band of almost exclusively vertebrate material in the diet. Within this zone there 92 are differential demands upon the feliform skull that are not met by the ancestral shape, and thus no hypercarnivore taxon analyzed occupied that zone. Other studies have shown the demands of bite force and its effect on cranial shape when it comes to hypercarnivory (Law et al., 2018b), durophagy (Tseng and Wang, 2011; Figueirido et al., 2013; Tseng, 2013), or sabertooth morphology (Figueirido et al., 2011; Christiansen, 2012). Taken together, these demands require different optimization leading to distinct cranial shapes, such as the dissipation of compressive force in durophagous hyaenids, the required increase in the gape of sabertooth feliforms, and overall cranial rigidity and strength in bite force for hypercarnivores in subduing/consuming their prey. Surprisingly, only one of the analyzed pair-wise comparisons of ecological category and cranial disparity found significance: sabertooth forms versus hypocarnivores. While each ecological category has a distinct cranial morphology, the range of variance within each category is similar. This feature of sabertooths was also observed in the osteometric analysis of Holliday and Steppan (2004) where increasing specialization did not correlate with morphological diversity except in the most specialized sabertooth taxa. The uniqueness of sabertooth taxa may relate to niche partitioning of scimitar- and dirk-tooth forms occupying additional morphospace (Lautenschlager et al., 2020). This greater disparity is reflected in extremely high rates of morphological evolution (Table 16), greater than all other ecological categories (2.2-5.1 times faster), suggesting high selective pressures for these cranial shapes. High rates were also observed for durophagous and hypercarnivorous carnivores relative to hypocarnivorous ones, but the associated disparity of these categories was not significantly different. The disparity of non- sabertooth hypercarnivores compared to more generalized ecomorphs has been contradictory. Holliday and Steppan (2004) recovered hypercarnivores with less disparity than non- 93 hypercarnivorous sister groups, while Michaud et al. (2020), found greater cranial disparity among hypercarnivorous feliforms than generalists (all other taxa) in their analysis. However, both may be accurate in describing certain patterns through respective methodological approach. Holliday and Steppan (2004)’s data primarily looked at dentognathic measurements, such as specific tooth lengths and widths and dentary output lever lengths. As previously stated, hypercarnivory showed trends in tooth loss and tooth row length reduction to accommodate higher bite forces, while dentition was often simplified in overall topography to facilitate slicing or bone-crushing mechanics. With the loss of posterior molars, overall variance would necessarily go down (no variance for these structures), leading to less overall disparity than taxa with broader diets. Michaud et al. (2020) and the present study suggest that from a cranial shape point of view, hypercarnivores are just as capable of occupying a range of shapes as other dietary categories with many pathways/shapes lending themselves to a hypercarnivorous ecology (many- to-one mapping). Conversely, dentognathic restrictions on a diet high in vertebrate material are limiting, especially in regard to evolvability of non-hypercarnivorous ecologies and broader diets. Clade disparity and impact of lineage age As questioned at the start of this study, the age of a lineage may play a substantial role in explaining disparity among clades, as an ancient lineage may have simply accrued substantial disparity by random walk dynamics over geologic time, disparity that younger clades may also achieve if provided with enough time. Indeed, there is a correlation of cranial disparity and age of lineage, where nimravids, as the oldest lineage, (34.3 Ma), have the greatest disparity. Similarly, herpestids have the least and are the youngest lineage (9.52 Ma), while all others are intermediate. Thus, adjusting for both allometry and age of lineage, where do we find the 94 greatest variance in cranial shapes amongst feliforms? In other words, on a more level playing field of size and time to evolve numerous and varied shapes, do certain clades still possess exceptional cranial disparity? Yes, with this adjustment machairodontines become the most diverse clade of cranial shapes, followed by similar values for nimravids and hyaenids. Felines/pantherines, euplerids and viverrids form a clump of similar values in third, while herpestids still remain the least disparate clade. In short, felids possess the greatest cranial disparity amongst feliforms. Setting aside sabertooth morphs for a moment, this great level of disparity is still observed in felines/pantherines, having comparable values to ecologically and taxonomically diverse euplerids and vivverids. Though perhaps not apparent in the outward appearance of many felids, derived cranial shapes have been the subject of several analyses, such as the unique skull of the cheetah (Acinonyx jubatus) that possesses resistance to compressive forces more analogous to a durophage and unlike the expectation for a cursorial specialist (Chamoli and Wroe, 2011; Tseng, 2013). The incipient sabertooth morphology of the clouded leopard (Christiansen, 2006, 2008c) also adds to the diverse morphological range of this clade, as does the primarily piscivorous flat-headed cat (Prionailurus planiceps), with a cranial shape consistent with its name. These relatively high cranial disparity values for felids were also observed by Michaud et al. (2018) and Figueiridoe et al. (2011) in their analyses, in spite of a different method of assessment and range of taxa. Perhaps unsurprisingly, relative canine size had substantial impact on cranial shape for machairodontines (~19%) and nimravids (~24%), the clades containing sabertooth feliforms. However, euplerids saw the greatest impact of this variable (~47%), a result which may be correlated with the taxa sampled. Only four species of euplerid were included in this study, representing extremes and apparent average of cranial shape for the family. Most euplerids 95 appear similar to their African herpestid relatives (e.g., Salanoia), while others are ecomorphologically divergent towards invertebrate diets, such as Eupleres, or hypercarnivory in Cryptoprocta. This high cranial variance was found to be the greatest among living feliforms by Michaud et al. (2018), but with the narrow sample of this study, the hypercarnivorous outlier of Cryptoprocta may be driving an overrepresented relative canine size signal. Greater taxonomic inclusion may reduce the RCS signal for the clade and/or inform on the extent of modification that occurs in the euplerid skull to accommodate hypercarnivorous canines. For the sabertooth clades, hypertrophied canines no doubt contribute to the greater range of cranial shapes not seen in other hypercarnivores. The required reorganization of the cranium to wield such hardware has been argued as a possible escape of soft-flesh specialists into new realms of disparity, when retreating back into omnivory or a more general diet is no longer possible (Holliday and Steppan, 2004). Nimravids’ high disparity may be summarized across several ecomorphs that would later appear in machairodontines and maybe even felines (Barrett, 2021). The “cat-like” cranial morphology may represent the only way to exist as a soft flesh specialist unless sabertooth modifications are undertaken. However, once that evolutionary path is opened, differential niches in scimitar- or dirk-tooth morphology may allow renewed variation to exist as large canines (and associated behavioral adaptations) are incorporated into the hypercarnivore mold. What follows are high rates of morphological evolution, with rapid selection for these features, generating a relationship between rates of morphological evolution and disparity (Figure 12), not seen in other clades (Slater, 2013; Jones et al., 2015; Michaud et al., 2018; Simons et al., 2020). 96 Allometry The relatively large predictive influence of size on cranial shape (27%) suggests that allometry is indeed important within Feliformia (for some clades such as felines and pantherines extremely so) such that there may be a common allometric trajectory for this clade. Furthermore, if size variation represents a line of least evolutionary resistance for morphological evolution, it would be expected that (1) rates of size evolution would be correlated with rates of cranial shape evolution, and (2) rates of size evolution would be predictive of cranial disparity. Taken at face value these connections do not exist within the results of the present study (Figure 12). However, when comparing size and shape rates of evolution, machairodontines and nimravids exist as outliers from what appears to be an otherwise linear relationship. Removal of these clades still results in a non-significant relationship (R2= 0.46, p< 0.08), but the reduced power of fewer observations may obfuscate a real correlation. As discussed throughout, sabertooth taxa exhibit anomalously high rates of evolution, and nimravids are recovered with the lowest levels of allometry of any feliform clade assessed, a non-statistically significant 9%. Part of this may relate to the existence of highly derived (e.g. dirk-tooth) nimravid taxa at either end of the size spectrum (Barrett, 2021), demonstrating that size is not required to generate these extreme morphologies. These low allometric values are not seen in machairodontines, but the even higher rates of morphological evolution within this clade fits into a narrative of high selective pressure and rapid evolution for this ecology. Even with these deviations in morphological rates of evolution for sabertooth clades, support for some level of common allometry amongst feliforms was found in this study (Table 14, Figure 13). Save sabertooth taxa and the vermivore specialist Eupleres, most taxa present a linear relationship in shape with increasing size. Comparisons of idealized allometric trajectories 97 further found only two significantly different results, that of machairodontines and nimraivds; and hyaenids with felines/pantherines. Machairodontines and nimravids evolved similar cranial shapes but did so under apparently different allometric trajectories. Implying that size, once again, has a differential impact on cranial evolution for these clades. The other clades that differ significantly, hyaenids and non-sabertooth felids, perhaps demonstrate the uniqueness in size- correlated changes amongst soft-flesh specialists with reduced tooth counts and brachiocephalic crania compared to those of incipient to fully developed durophagy and comparatively dolichocephalic crania. Regardless, the deviants are all clades with large size ranges, and all contain hypercarnivores, with hyaneids the only of these clades containing taxa that are not solely hypercarnivorous. The lack of significance between other clades may thus relate to a lack of selective pressure to pull them off this common allometric trajectory of utility of mapping a narrow range of generalized cranial shapes onto a broad swath of dietary possibilities. The lack of significant differences in allometric trajectories between hypercarnivorous clades and more generalized ones may then simply be a result of lack of overlap in size ranges to robustly infer these differences, or small members of hypercarnivous clades are not that different from large members of generalist clades, as in hyaenids. A variable connection in macroevolutionary size to shape is observed in several clades. For example, Simons et al. (2020) found no correlation in catarrhine primate morphological evolutionary rates (size or shape) and disparity, but did find a correlation for size and shape evolutionary rates, unlike the present study. However, their additional finding that most of the variation in within-clade allometries is best explained by a shared allometry model indicates that allometry is still an important factor in catarrhine cranial evolution. This hypothesis is supported by numerous other primate studies (Collard and O’Higgins, 2001; Singleton, 2002; Mitteroecker 98 et al., 2004; Schaefer et al., 2004), which found that allometric trends are apparent in interspecific cranial shape differences, but that species do not merely share allometric scaling trajectories. Instead, details of primate evolution are likely constrained by the necessity of maintaining the numerous functional roles of the cranium throughout an organism’s life span preventing size from being the sole force in the production of morphological disparity (Simons et al., 2020). Comparatively, Marcy et al. (2020) found strong support for conserved allometric slopes across Australian murids, save for carnivore and folivore specialists. The static (within-species) allometric trajectories were self-similar to evolutionary (among-species) Australian murid allometry. This pattern is more similar to that seen in non-mammalian vertebrate clades. Bird cranial evolution seems to possess strong allometric patterns; for example, the shape of bird of prey beaks is controlled almost entirely by size (~80%, Bright et al., 2016). In another example of this strong allometry in birds, most nightbird (Strisores) cranial shapes display changes associated with heterochrony (Navalón et al., 2021). Similarly, encephalization, or growth of the brain causes changes in the topology of its bony base inside the cranium leading to predictable effects on the shape of the whole skull (Marugán-Lobón et al., 2022). Overall, mammals are thought to have a high degree of evolvability in cranial allometry (Tsuboi et al., 2018), such that specific or generic allometric slopes can be far more variable than seen at higher taxonomic levels. Taken together with the present study, it appears that feliforms share relatively similar allometric trajectories until ecological selection on some aspect of hypercarnivory pulls them off into new realms of shape space. Certain clades, such as hyaenids, appear to deviate less from generalized small feliform taxa, while soft-flesh specialists like felids 99 and nimravids possess the greatest shift in allometric intercepts and possibly slopes to accommodate their derived ecology. Cranial integration The idea that variation, or the raw material for evolution, may not be isotropic in nature, but instead has limitations that facilitate or constrain subsequent evolution, has been a prominent hypothesis within evolutionary theory for well over a century (Galton, 1869; Gould, 2002). Strong correlations between traits may limit or outright prevent certain traits from existing, while on the other hand, such correlations may funnel or fruitfully channel variation in extreme directions that increase the rate and/or magnitude of observed variation (Goswami et al., 2014). This idea of constraint may then be a fertile hypothesis for why certain feliform clades never achieved a given level of disparity in their evolutionary history, representing a “smoking gun” of underlying processes worthy of investigation. For the present study, magnitude of cranial integration was employed as a proxy of developmental constraint, one that has the additional benefit of being collected from the fossil record. Amongst carnivorans, there have been contradictory results and arguments for the effect of integration on the generation of cranial disparity. For example, increasing integration has been used to explain increased disparity across the latest Pleistocene in Smilodon and dire wolves (Goswami et al., 2015), while that same integration may also be responsible for differential morphospace occupation of machairodontines compared to nimravids (Goswami et al., 2011). Conversely, in assessment of cranial module integration and disparity for living terrestrial carnivorans and primates, over half of comparisons found no significant correlation between these variables, but when significant results were found the effect of integration on disparity is in limitation of morphological variability (Goswami and Polly, 2010a). A similarly complicated 100 result was found for pinnipeds, wherein ecology may be driving integration values (Randau et al., 2019), while no correlation was found in levels of integration and disparity generated when comparing domesticated mammals with their wild ancestors (Wilson et al., 2021). Like these latter studies, the predictive power of integration on disparity of this analysis returned insignificant results (Figure 10 and Figure 11). This was still the case when values of raw clade disparity were used for eigenvalue dispersion (R2= -0.16, p<0.69), while even poorer results for between-module integration (R2= -0.20, p<0.90). These results suggest that no connection exists between how tightly integrated (developmentally or otherwise) a feliform’s cranium is and the range of morphological variation it can achieve. Instead, crania remain generally similar to each other, fulfilling a wide variety of dietary and ecological roles until the constraint of hypercarnivory comes into play. At this time, necessary considerations for subduing or processing non-compliant vertebrate prey overrides the ancestral cranial shape in feliforms generating new morphologies. Some of these shapes appear to fall on a common allometric trajectory of smaller clades (i.e., hyaenids), suggesting that the path to generating diverse prey material hypercarnivory and durophagy has a solution different than that of soft-flesh specialized hypercarnivores. Felids and nimravids occupy their own allometric and morphospace realms reflecting a presumed similar ecology and path into their varied cranial shapes, perhaps ending in sabertooth morphology. Regardless, feliform cranial shape seems to remain the same until it isn’t. There is no underlying restriction (as assessed via integration) that prevents them from achieving disparate cranial forms; instead, a high dietary/ecological bar must be cleared before selection pressure directs cranial evolution into new realms. 101 Chapter V Conclusion Feliforms of small size exist within a realm of opportunity. This ecospace allows the exploitation of diverse resources including invertebrate-dominated diets, folivory, frugivory, hypocarnivory and mesocarnivory. A narrow range of cranial shapes accomplishes these diverse diets, while the dental toolkit is unencumbered to optimize for the challenges in processing each food material (Figure 15). This pattern is seen in the random walk small feliform body size and dental toolkit take as they evolve, the labile nature of convergent evolution of living small feliform species (Gaubert et al., 2005), and the challenges in resolving phylogenetic relationships amongst fossil taxa of this ecospace. However, there is a tipping point where extreme diet, and to a lesser extent body mass, take over, to shape the morphology and ecology of feliform evolution. Hypercarnivory, as observed at ≥80% vertebrate material in the diet, forces selection for more robust cranial morphology to subdue and process prey items. This selection necessitates vacating the ancestral feliform cranial mold and occupying a distinct, broader area of shape space. However, what path a feliform takes into this realm depends upon its style of hypercarnivory and distinct ecospace. Soft-flesh specialists such as felids, nimravids, Cryptoprocta and Proailurus rapidly acquire their familiar cranial and dentognathic condition, with loss of posterior molars, enhancement of the shearing surfaces of the carnassials, robust canines and brachiocephalic crania. With the loss of grinding dentition and the associated capacity to process plant material, there appears to be no return to former ecospaces. This is not all bad, for soft-flesh hypercarnivory is tenable across multiple orders of magnitude in body size; one can be minute as in the black-footed cat (Felis nigripes), less than 2.5 kg (Renard et al., 2015), or enormous as in 102 the extinct North American Lion (Panthera leo atrox), up to possibly 457 kg (Wheeler and Jefferson, 2009). Furthermore, allometry may offer a line of least evolutionary resistance for these hypercarnivores, for access to new prey items may only require the cranial (and likely post- cranial) changes associated with allometric scaling (Meachen-Samuels and Van Valkenburgh, 2009; Slater and Van Valkenburgh, 2009), scaling that differs compared to non-soft-flesh- specialists. Though, a more extreme realm exists beyond these familiar cat-like shapes from this ecospace, one of sabertooth morphology. Becoming sabertooth opens new doors into cranial disparity, variance not seen within any other ecomorphology. An obvious source for this increased variance is the reorganization of the cranium to facilitate wielding such hardware, but two new ecologies appear to open up as well, that of scimitar- and dirk-tooth morphs. Scimitar- tooth carnivores fall closer to the common allometric line of other feliforms (Fig. 13A), requiring fewer modifications in terms of gape management than dirk-tooth morphs. Studies of machairodontines find differentiation in their respective diets via isotopic analysis, which is also reflected in their respective baupläne argued to favor pursuit or ambush predation (Feranec, 2004; DeSantis et al., 2021). Regardless, body mass doesn’t appear to be that important as a sabertooth carnivore. Some of the smallest nimravids were also the most derived dirk-tooth morphs, such as Eusmilus cerebralis, but the inferred largest feliform of this entire study, Barbourofelis fricki, was possibly the most derived dirk-tooth placental mammal to have ever existed (Barrett, 2021). This range of extremes is not as complemented in equal extremes of body mass in machairodontines, but bobcat sized members are known to have also existed (de Bonis et al., 2018), though unable to be included in this study. 103 Figure 15. Hypothesis of terrestrial carnivoran evolution with regards to ecology and cranial morphology. Silhouettes reference relative body size possible within each ecospace (small to large), while food icons an approximation of dietary spectrum. Silhouettes taken from Phylopic.org. The other feliform path into hypercarnivory is that occupied by hyaenids. Unlike soft-flesh specialists, hyaenids facilitate their cranial shape by use of the common allometric trajectory of most feliforms, those occupying the small-bodied ecospace. Hyaenids are notable in their ecological exploitation of nearly all dietary categories examined in this study, though with two exceptions, soft-flesh specialization as in living felids, and sabertooth morphology. Why didn’t hyaenids evolve these ecomorphologies? As stated above, reduction or loss of posterior molars is associated with felid ecomorphology. This reduction is seen in Hyaena, and to a greater degree in Crocuta, where M1s are typically not present. However, all hypercarnivorous hyaenids 104 maintain robust premolar dentition, something not seen in felids nor nimravids. Indeed, the upper P3 and lower p3-4 are used for durophagy in many hyaenids, but not all, such as Chasmaporthetes (Coca-Ortega and Pérez-Claros, 2019). So again, why the difference? It is informative that canids and hyaenids returned similar results in the mode of their ecological evolution. Both clades found the best fit for an optimal body mass and dental toolkit per dietary category, and canids also did not converge on soft-flesh specialization in the same way as felids and nimravids. Amongst canids, hypercarnivory is a delayed acquisition of ecospace evolution (Slater, 2015). It forms a terminus of ecomorphology across all three subfamilies. Compare this to the paleoecology analyses of felids and nimravids, which saw greatest support for early burst dynamics directly into hypercarnivory. A clearer picture of the evolutionary trajectory for hyaenids will require a more comprehensive phylogeny, that better samples the ‘ictitheres’ and the more hypo- to mesocarnivorous lineages. However, current literature suggests that the evolution of hypercarnivory in hyaenids is similar to that of canids with delayed acquisition (Werdelin and Solounias, 1991; Turner et al., 2008). Hyaenids may be unable to access soft-flesh specialization, as were canids, from insurmountable phylogenetic inertia. The fossil record suggests that both lineages developed cursorial morphology, early on losing the ability to supinate and pronate the forelimbs, and to retract the distal phalanges (Hunt Jr. and Solounias, 1991; Wang, 1993; Figueirido et al., 2015; Gracia, 2015). Generally speaking, the felid postcranial condition is plesiomorphic among Carnivoramorpha (Spaulding and Flynn, 2009, 2012; Solé et al., 2014, 2016) and likely associated with arboreal adaptations to the dense canopy forests of the Eocene (Tomiya et al., 2021). Cursoriality favors face-orientated predation in canid-like hypercarnivores, compared to ambush and grappling strategies found in felids, and presumably nimravids (Ewer, 1973; Andersson, 2005). The cranium of soft-flesh specialists 105 maximizes gape capability while balancing bite force (even in non-sabertooths) to allow quick dispatch of prey, a benefit in solitary predators (Figueirido et al., 2011; Meloro et al., 2015). Thus, the early trajectory of hyaenids and canids into cursoriality with increasing body size may not have allowed them to switch tracks to the ecospace shared by felids and nimravids. Instead, the hyaenid/canid postcranial situation may have predisposed these lineages to communal hypercarnivory, necessitating a different strategy in capturing large prey (Van Valkenburgh and Koepfli, 1993; Van Valkenburgh et al., 2003). This hyaenid/canid ecospace and associated evolutionary trajectory may also be shared by the amphicyonid lineages, Daphoeninae and Temnocyoninae, and the ursid hemicyonines (Hunt, 1996, 1998b, 2001b). All experienced similar patterns of cursorial adaptations and dolichocephalic cranial shapes with broad dental tool kits capable of omnivory. Furthermore, all of these lineages produced members with adaptations to durophagy via hypercarnivory, save perhaps hemicyonines. Future paleoecological and cranial morphology analyses could assess the validity of the comparison, but sufficient phylogenies are also in need. Interestingly, a fourth ecospace category may exist, encompassing extant ursines. Turning to the fossil record it appears that the Amphicyoninae lineage once occupied this ecospace, featuring large size, relative plantigrade posture, but a range of hypo-hypercarnivorous diets. Much like the above comparisons to other caniforms, future study may find a unique hypercarnivorous end member for this ecospace, one possibly occupied by taxa such as Ysengrinia and Magericyon (Viranta, 1996; Peigné et al., 2008; Soledad Domingo et al., 2013). Furthermore, unlike the previous ecospaces, both ursids and amphicyonids seemed capable of switching from canid/hyaenid ecospace to the ursine space during their evolutionary history and thus offer a 106 unique area of inquiry of why these clades where able to make the shift, while hyaenids and canids were not. The above discussion offers potential pathways in carnivoran evolution, but not the process- based push that may get them moving or shifting to new zones. For that, contingency in natural history may be the best explanation. Small feliforms, and maybe even caniforms such as mustelids, procyonids, ailurids, mephitids, seem labile in their ability to shift rapidly in diet and associated dental toolkits. However, climatic and tectonic sourced opportunity may be required to facilitate rapid radiations into soft-flesh specialized hypercarnivory or the wide ecological occupation of canids and hyaenids. Nimravids gained purchase in North America, through circum-Pacific immigration from Asia, into an increasingly mosaic forested landscape after the Mid-Eocene climatic optimum. This was also shortly after the extinction of sabertooth machaeroidines (Zack et al., 2022), facilitating a potential contingent opportunity for nimravids. The Oligo-Miocene transition saw both global warming and the African-Eurasian interchange. This period correlates with the divergence of hyaenids from their common ancestors with herpestids and euplerids the origin of felids, the origin and diversification of lophocyonids, and initial diversification of the barbourofelins. In short, an immense diversification, in both taxonomic and ecological diversity of feliforms. The most recent example of such contingent global opportunities may be the Middle Miocene Climatic Optimum, which recovers this time as the appearance of the most recent common ancestor for the extant radiation of viverrids and herpestids + euplerids, and the divergence of machairodontine felids from felines + pantherines. The success of euplerids is itself a phenomenal example of contingent evolution with the sweepstakes dispersal of a herpestid-like ancestor rafting to Madagascar and diversifying into the 107 ecologically broad family known today. So, viewed in one light, feliforms have been capable of invading numerous ecologies around the globe; they just required the opportunity to do so. Considering the original question: Is there evolutionary constraint in feliforms? Apparently yes, but not in the way typically framed. Herpestids, viverrids, euplerids, prionodontids, etc. are constrained in staying small to exploit the diverse range of ecologies they occupy, small size that frees them from metabolic and functional challenges of greater mass. Additionally, they can do all of this with relatively little change in cranial morphology. However, once a feliform enters one of the other ecospace pathways outlined above, new constraints come into play, such as retreating from soft-flesh specialist hypercarnivory into a broader dietary category or switching tracks from the canid/hyaenid ecospace into the aforementioned felid/nimravid space. These constraints fruitfully channel the extreme endmembers of cranial evolution and ecology that otherwise are inaccessible to other groups, i.e., sabertooth and durophagous morphology. An important distinction of this work is the lack of support for an inherent developmental or cranial disparity constraint on the variation observed for supposedly stymied specialists, such as felids. Quite the opposite in fact; felids possess the greatest range of cranial variety of any feliform. Though narrow in their vertebrate dominated diet, they seem to have explored a diverse set of options in cranial shape, options that do not exist for more generalist feliforms, or ones that maybe could not exist without the constraints that have affected their evolutionary trajectory. In the end, it seems Galton was correct. Only certain channels or facets seem viable for the route of evolution. This is expressed in the irreversible loss of the cusps and teeth in hypercarnivores, or the “re-development” of grappling forelimbs with retractile claws in lineages with cursorial postcrania. These kinds of limitations shape the evolutionary landscape, generating channels down which variation can proceed. This in turn is seen as convergence in the fossil record, and 108 for this study, the smoking gun of channelized evolution among terrestrial mammalian carnivores. Indeed, the above discussion points out these patterns among carnivorans, but broader taxonomic inclusion finds similar ecospaces for Sparassodonts in South America (Prevosti et al., 2012; Forasiepi and Sánchez-Villagra, 2014), marsupials in Australia (Rovinsky et al., 2021) and hyaenodont and oxyaenid “creodonts” (Borths et al., 2016; Zack, 2019b), the latter of which also developed sabertooth morphology within the machaeroidines (Zack et al., 2022). These broad repeated patterns suggest that the interface of terrestrial mammalian carnivores with Cenozoic ecosystems has been consistent. That within this framework only certain trajectories are possible to be a successful carnivore. These channels do provide a limit on what ecology and morphology is feasible, but they also facilitate the fruitful possibilities of extreme carnivory, unique for each channel. 109 Appendix A Genbank Accession numbers for molecular data in phylogenetic analysis Nuclear Mitogenome IRBP CH TTR- MT- MT- MT- MT- MT- MT- MT- MT- MT- MT- MT- MT- MT- RN intron ATP6 ATP8 CO1 CO2 CO3 CYB ND1 ND2 ND3 ND4 ND4L ND5 ND6 A-1 1 Acinonyx XM_0 DQ0 DQ0 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 AF34 jubatus 270624 8182 8251 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 4830. 97 6 7 1 1 1 1 1 1 1 1 1 1 1 1 1 Arctictis DQ683 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 KX44 binturong 125 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 9332. 1 1 1 1 1 1 1 1 1 1 1 1 1 Atilax AY7 JF444 AF52 AY75 paludinosus 5060 116.1 2324. 0647. 4 1 1 Bdeogale AY9 AY95 AY97 nigripes 5064 0655. 4022. 3 1 1 Canis latrans KT447 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 DQ48 969 0511 0511 0511 0511 0511 0511 0511 0511 0511 0511 0511 0511 0511 Caracal caracal DQ0 DQ0 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 8183 8252 2272 2272 2272 2272 2272 2272 2272 2272 2272 2272 2272 2272 2272 1 1 Catopuma AY525 DQ0 DQ0 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 KP27 temminckii 034 8184 8253 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 1500. 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 Civettictis AY170 GU9 AY1 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 civetta 078 3102 7002 33378 33378 33378 33378 33378 33378 33378 33378 33378 33378 33378 33378 33378 4 3 Crocuta crocuta AY170 AY9 DQ0 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 JF894 087 2870 8254 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 377.1 0 0 Crocuta crocuta MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 MN3 spelaea 20462 20462 20462 20462 20462 20462 20462 20462 20462 20462 20462 20462 20462 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 110 Crossarchus AY170 AF03 JF444 AY17 AY17 obscurus 071 9726 131.1 0101. 0041. 1 1 Cryptoprocta AY170 AY9 AY1 MG4 MG4 ferox 066 2870 7001 52178 52299 5 8 .1 .1 Cynogale DQ68 DQ68 bennettii 3992. 3983 1 Eupleres KY30 MG4 goudotii 5488. 52300 1 .1 Felis lybica DQ0 DQ0 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 ornata 8181 8250 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 2275. 4 7 1 1 1 1 1 1 1 1 1 1 1 1 1 Fossa fossana AY170 GU9 AY1 AF51 AY17 067 3102 7001 1062. 0037. 2 9 1 1 Galerella AY9 AF52 AY97 sanguinea 5064 2331. 4027. 6 1 1 Genetta DQ267 AY2 KJ19 AF51 DQ68 maculata 562 3262 2810. 1055. 3986. 7 1 1 1 Helogale DQ0 AY7 AF52 AY75 parvula 8185 5060 2333. 0649. 2 5 1 1 Hemigalus AY170 AY1 AY17 AY17 derbyanus 082 7002 0109. 0052. 7 1 1 Herpestes AY170 AY1 DQ51 AY17 edwardsii 080 7002 9053 0050 5 Homotherium MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 latidens 1701 1701 1701 1701 1701 1701 1701 1701 1701 1701 1701 1701 Homotherium MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 serum 1703 1703 1703 1703 1703 1703 1703 1703 1703 1703 1703 1703 Hyaena hyaena DQ267 AY9 DQ2 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 570 2870 6755 20669 20669 20669 20669 20669 20669 20669 20669 20669 20669 20669 20669 20669 2 2 111 Ichneumia GU9 AY9 MN3 AF52 AY97 albicauda 3102 5065 26093 2341. 4034. 0 1 .1 1 1 Leopardus AB109 DQ0 DQ0 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 pardalis 335 8183 8252 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 2284. 4 4 1 1 1 1 1 1 1 1 1 1 1 1 1 Lynx canadensis DQ205 AY9 DQ0 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 KP20 910 2869 8252 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 2281. 5 0 1 1 1 1 1 1 1 1 1 1 1 1 1 Miracinonyx DQ09 DQ09 DQ09 trumani 7168. 7175. 7170. 1 1 1 Mungos mungo AY170 AY9 AY1 AF52 AY17 065 2869 7001 2347. 0035. 8 7 1 1 Nandinia AY170 AY9 AF03 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 binotata 083 2870 9729 24567 24567 24567 24567 24567 24567 24567 24567 24567 24567 24567 24567 24567 4 Neofelis AY525 DQ0 DQ0 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 DQ25 nebulosa 032 8184 8253 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 7669. 9 9 1 1 1 1 1 1 1 1 1 1 1 1 1 Otocolobus AY525 DQ0 DQ0 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 MH9 manul 039 8181 8251 78908 78908 78908 78908 78908 78908 78908 78908 78908 78908 78908 78908 78908 9 1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 Paguma larvata AY525 AY5 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 KT19 intrudens 040 2505 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 1130. 5 1 1 1 1 1 1 1 1 1 1 1 1 1 Panthera atrox DQ89 9945. 1 Panthera leo AY525 AY9 AF03 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 036 2869 9725 28302 28302 28302 28302 28302 28302 28302 28302 28302 28302 28302 28302 28302 4 Panthera leo KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 KX25 spelaea 8452 8452 8452 8452 8452 8452 8452 8452 8452 8452 8452 8452 8452 Panthera pardus AY525 DQ0 DQ0 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 KP00 041 8184 8253 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 1507. 5 5 1 1 1 1 1 1 1 1 1 1 1 1 1 Paradoxurus AY170 AY9 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 MG2 hermaphroditus 086 2869 00264 00264 00264 00264 00264 00264 00264 00264 00264 00264 00264 00264 00264 112 laotum 7 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 Poiana DQ267 AY2 AF51 richardsonii 559 3262 1049. 0 1 Prionailurus DQ0 DQ0 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 KY68 planiceps 8182 8251 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 2729. 3 5 1 1 1 1 1 1 1 1 1 1 1 1 1 Prionodon JN414 DQ0 DQ2 MT55 MT55 MT55 MT55 MT55 MT55 MT55 MT55 MT55 MT55 linsang 782 8185 6755 9410. 9410. 9410. 9410. 9410. 9410. 9410. 9410. 9410. 9410. 0 1 1 1 1 1 1 1 1 1 1 1 Profelis aurata DQ0 DQ0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 8183 8252 28299 28299 28299 28299 28299 28299 28299 28299 28299 28299 28299 28299 28299 2 2 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 Proteles AY9 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 MH6 cristatus 2870 62445 62445 62445 62445 62445 62445 62445 62445 62445 62445 62445 62445 62445 3 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 Puma concolor DQ205 DQ0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 911 8182 16470 16470 16470 16470 16470 16470 16470 16470 16470 16470 16470 16470 16470 4 Rhynchogale GU9 AY9 AF52 AY97 melleri 3102 5065 2344. 4036. 1 3 1 1 Salanoia AY7 AY18 MG4 concolor 5060 7007. 52301 7 1 .1 Smilodon MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 MF87 populator 1700 1700 1700 1700 1700 1700 1700 1700 1700 1700 1700 1700 1700 Suricata AY170 DQ0 DQ0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 NC_0 suricata 084 8185 8254 45900 45900 45900 45900 45900 45900 45900 45900 45900 45900 45900 45900 45900 3 2 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 .1 Taxidea taxus AB285 AF4 AY7 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 HM1 jacksoni 379 9814 5059 06330 06330 06330 06330 06330 06330 06330 06330 06330 06330 06330 06330 06330 8 8 . 1 .1 .1 .1 .1 .1 . 1 .1 . 1 . 1 . 1 . 1 .1 Viverra AY170 AF03 AF51 AY17 tangalunga 085 9731 1045. 0055. tangalunga 1 1 Viverricula DQ267 AY2 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 KX89 indica schlegelii 568 3261 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 1745. 8 1 1 1 1 1 1 1 1 1 1 1 1 1 113 Vulpes vulpes GQ214 AY9 AF03 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 KP34 fulva 077 2869 9733 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 2452. 3 1 1 1 1 1 1 1 1 1 1 1 1 1 114 Appendix B Highest supported partition and evolutionary model scheme of molecular data as determined via PartitionFinder 2 Settings alignment : ./infile.phy branchlengths : linked models : JC, K80, TRNEF, SYM, HKY, TRN, GTR, HKY+X, TRN+X, GTR+X, JC+G, K80+G, TRNEF+G, SYM+G, HKY+G, TRN+G, GTR+G, HKY+G+X, TRN+G+X, GTR+G+X, JC+I, K80+I, TRNEF+I, SYM+I, HKY+I, TRN+I, GTR+I, HKY+I+X, TRN+I+X, GTR+I+X, JC+I+G, K80+I+G, TRNEF+I+G, SYM+I+G, HKY+I+G, TRN+I+G, GTR+I+G, HKY+I+G+X, TRN+I+G+X, GTR+I+G+X model_selection : bic search : greedy Best partitioning scheme Scheme Name : step_10 Scheme lnL : -85150.29748535156 Scheme BIC : 171846.019025 Number of params : 169 Number of sites : 9363 Number of subsets : 10 Subset | Best Model | # sites | subset id | Partition names 1 | HKY+I+G+X | 1678 | 1810c769b8f916a03fd2a6c2d9373381 | ND4L_1stpos, COX3, ATP6, COX2, COX1 115 2 | GTR+I+G+X | 3990 | 08696fe9fcedeb6bee35f068c3f81ae3 | ND4, CYB, ND1, ATP8, ND5, ND3 3 | SYM+G | 1521 | c9715b8586452f904d516ab02113b915 | TTR, CHRNA_1 4 | TRN+G+X | 421 | 87e5cb9ee47837231e1b3b3e8777f356 | IRBP_1stpos 5 | TRNEF+I | 421 | 8201d351e9239325f36dd35b4c61c9af | IRBP_2ndpos 6 | HKY+G+X | 421 | 785a339441a44dfd0e936358a6abde9b | IRBP_3rdpos 7 | HKY+I+G+X | 396 | c61fec9e72928c8aa7170ad04a8e6142 | ND2 8 | HKY+I+G+X | 98 | eeb82570db336fc244e80a4f45ebdce5 | ND4L_2ndpos 9 | HKY+I+G+X | 98 | 321271c4f6bceb00245ec5d843fdc285 | ND4L_3rdpos 10 | HKY+G+X | 319 | eaeee0a798be6d486926234462330d84 | ND6 Scheme Description in PartitionFinder format Scheme_step_10 = (ND4L_1stpos, COX3, ATP6, COX2, COX1) (ND4, CYB, ND1, ATP8, ND5, ND3) (TTR, CHRNA_1) (IRBP_1stpos) (IRBP_2ndpos) (IRBP_3rdpos) (ND2) (ND4L_2ndpos) (ND4L_3rdpos) (ND6); Nexus formatted character sets begin sets; charset Subset1 = 5160-5453\3 1827-2164 1-125 1388-1826 710-1387; charset Subset2 = 5454-6268 2165-2846 4110-4644 126-321 6269-7911 5041-5159; charset Subset3 = 8231-9363 322-709; charset Subset4 = 2847-4109\3; charset Subset5 = 2848-4109\3; charset Subset6 = 2849-4109\3; charset Subset7 = 4645-5040; charset Subset8 = 5161-5453\3; charset Subset9 = 5162-5453\3; charset Subset10 = 7912-8230; 116 charpartition PartitionFinder = Group1:Subset1, Group2:Subset2, Group3:Subset3, Group4:Subset4, Group5:Subset5, Group6:Subset6, Group7:Subset7, Group8:Subset8, Group9:Subset9, Group10:Subset10; end; 117 Appendix C Source Material for Morphological Character Scoring and Continuous Variables American Museum of Natural History, New York City, New York, USA (AMNH) American Museum of Natural History – Frick Collection, New York City, New York, USA (F:AM) American Museum of Natural History, Comparative Anatomy Collection, New York City, New York, USA (AMNH C.A.) Field Museum of Natural History, Chicago, Illinois, USA (FMNH) Smithsonian National Museum of Natural History, Washington D.C., USA (USNM) John Day Fossil Beds National Monument, Kimberly, Oregon, USA (JODA) Grand Canyon National Park, Grand Canyon, Arizona, USA (GRCA) University of Nebraska Lincoln State Museum, Lincoln, Nebraska, USA (UNSM) Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA (UF) Texas Memorial Museum, University of Texas at Austin, Austin, Texas, USA (TMM) Museum of Evolution of Uppsala University, Uppsala, Sweden (M) University of California Museum of Paleontology, Berkeley, California, USA (UCMP) Yale Peabody Museum, New Haven, Connecticut, USA (YPM) 118 Yale Peabody Museum, Princeton University Collection, New Haven, Connecticut, USA (YPM PU) University of Kansas, Natural History Museum, Lawrence, Kansas, USA (KUVP) Raymond M. Alf Museum of Paleontology, Claremont, California, USA (RAM) Museum of Geology, South Dakota School of Mines and Technology, Rapid City, South Dakota, USA (SDSM) University of Oregon, Museum of Natural and Cultural History-Modern Collection, Eugene, Oregon, USA (B) Los Angeles County Museum, George C. Page Museum (Hancock Collection), Los Angeles, California, USA (LACMHC) Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain (MNCN) Museum National d'Histoire Naturelle, Paris, France (MNHN) Museum für Naturkunde, Berlin, Germany (MFN MB. Ma.) Beijing Natural History Museum, Beijing, China (BNHM) Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing, China (IVPP) Collection du Quercy, Faculté des Sciences de Marseille, France (FSM PQ) University of Florence, Florence, Italy (IGF) University of Washington Burke Museum, Seattle, Washington, USA (UWBM) 119 Kenya National Museum, Nairobi, Kenya (KNM) Uganda Museum, Kampala, Uganda (UM) Taxa Specimens Literature Tungurictis spocki AMNH 26600, 26610, 99146 (Hunt Jr. and Solounias, 1991; Wang, 2004) Ictitherium viverrinum AMNH 20696, China 51-L437; F:AM (Coca-Ortega and Pérez-Claros, 144906, 144905 2019) Hyaenotherium wongii AMNH 20555, 22878, 99082, 99088, (Coca-Ortega and Pérez-Claros, 99084, 99091, 99081, 99093, 99080, 2019) 99095, 20554, 20586 Hyaenictitherium hyaenoides AMNH 144883; F:AM 144892, (Coca-Ortega and Pérez-Claros, 144889 2019) Lycyaena chaeretis F:AM 145002, 144899, 144902 - Chasmaporthetes lunensis AMNH 99789, 99786, 99781, 26955, (Baryshnikov and Averianov, 101261, 113847; F:AM 99787, 99783; 1993a; Antón et al., 2006; Coca- MNCN 67100 Ortega and Pérez-Claros, 2019) Chasmaporthetes ossifragus UF 18088, 18089, 27366 (Berta, 1981) Palinhyaena reperta F:AM 144897 - Adcrocuta exima China -41L-339; AMNH 22880, (Baryshnikov and Averianov, 140301, 140298, 140299, 140300; 1993a; Kovachev, 2012; Coca- F:AM 144903 Ortega and Pérez-Claros, 2019) Pachycrocuta brevirostris AMNH 27757, 27756 (Werdelin and Solounias, 1991; Baryshnikov and Averianov, 1993a; Mutter et al., 2001; Palmqvist et al., 2011; Liu et al., 2021) Tongxinictis primordialis - (Qiu et al., 1988b) Allohyaena kidici - (Howell and Petter, 1985; Werdelin and Kurten, 1999; Coca-Ortega and Pérez-Claros, 2019) Dinocrocuta gigantea - (Qiu et al., 1988a; Koufos, 1995; Zhang, 2005; Tseng and Binder, 120 2010; Xiong, 2019) Percrocuta carnifex - (Howell and Petter, 1985; Morales and Pickford, 2006; Ghaffar et al., 2019) Percrocuta algeriensis - (Howell and Petter, 1985) Percrocuta tobieni - (Howell and Petter, 1985; Werdelin, 2019) Proteles cristatus FMNH 211365; AMNH 24219 (Baryshnikov and Averianov, 1993a) Crocuta crocuta FMNH 34582, 18855 (Baryshnikov and Averianov, 1993a) Hyaena hyaena FMNH 101982, 47416 (Baryshnikov and Averianov, 1993a) Plioviverrops orbignyi MFN MB. Ma. 29580 (de Beaumont, 1969; Koufos, 2006, 2009, 2012) Crocuta crocuta spelaea MFN MB. Ma. 43, 44381, 29593, (Anyonge, 1993; Baryshnikov and 29940, 30359, 30360, 29945, 30357, Averianov, 1993a; Coca-Ortega and 29730, 29744, 29735, 29733, 29745, Pérez-Claros, 2019) 51190.9, 92 Protictitherium crassum - (Chen and Schmidt-Kittler, 1983; Gracia, 2015) Belbus beaumonti - (de Beaumont, 1968; Werdelin and Solounias, 1991; Koufos, 2011) Proailurus lemanensis AMNH 101931 (cast of holotype (Schmidt-Kittler, 1976) MNHN S.G. 3509) Pseudaelurus validus AMNH 62128, 62167; F:AM 61847 (Hunt, 1998; Rothwell, 2001, 2003) Promegantereon ogygia - (Salesa, 2002; Salesa et al., 2005, 2010a, 2010b; Christiansen, 2013; Siliceo et al., 2014) Metailurus major AMNH 131854 (cast of IVPP 5679), (Kovatchev, 2001) AMNH 26379, China-57-L548, Bx45L399; M 3841 Nimravides pedionomus F:AM 61855, 61852, 25205, 25206, - 62156-B, 62156-C, 62154, 62153-A, 62155, 62174, 62158, 62157-B 121 Machairodus catocopis F:AM 104044; Hig. 315-2306, KAN- (Antón et al., 2013) 30-18, KAN-85 125 Megantereon cultridens AMNH 105446, 113842 (cast of IGF (Vekua, 1995; Antón and Werdelin, 831), 113848 (cast of IGF 827), 1998; Qiu et al., 2004; Christiansen 101471, 105087 and Adolfssen, 2007; Palmqvist et al., 2007; Christiansen, 2013) Amphimachairodus giganteus F:AM 50478, 50476 (Koufos, 2016) Amphimachairodus coloradoensis F:AM 104726/104727, 104725, (Antón et al., 2013) 69263; #1745 Bx 50 DKLA, GY-147- 4995-H, #4142 Bx 117, #1066 Bx 34, #901 Bx 29, #3711 Bx 101, #368 Bx 16, #3492 Bx 99, RED 175-3262 Dinofelis diastemata F:AM 50445, 50446 (Werdelin and Lewis, 2001) Homotherium serum F:AM 142497, 128069, 116841; TMM (Meade, 1961; Collins, 1981; 933-3444 Christiansen, 2013; Antón et al., 2014) Homotherium latidens AMNH 104641; F:AM 50461, 50462, (Anyonge, 1993; Sardella and 50469 Iurino, 2012; Antón et al., 2014) Homotherium ischyrus AMNH 95297, F:AM 131893, (Hearst et al., 2011) Chan.35-856, Chan-42-929 Smilodon fatalis AMNH 14349; LACMHC 2002-R- (Matthew, 1910; Merriam and 289, R10864, R10688, A-3708, K-65, Stock, 1932; Christiansen and K-876, K-1541, K-2281, Q-1609, Q- Harris, 2005) 2965, 36960, 37415, 37873, 38504, 38731, 39221, U-3585, R-423, R- 6739, 40416, K-3309, K-4507, K- 4671, J-6083, T-5113, T-3068, Q- 5192, Q-3903, 46699, K-2768, N- 1219, N-1737, 133528, 60175 Smilodon populator MFN MB. Ma. 48108 (cast of MNHN (Kurtén and Werdelin, 1990; F.BRD21) Christiansen and Harris, 2005; Rincón, 2006; de Castro and Langer, 2008; Christiansen, 2013; Wallace and Hulbert, 2013; McDonald and Werdelin, 2018) Xenosmilus hodsonae UF 60000, 19400, 22907, 45340, (Martin et al., 2011; Christiansen, 45428, 223829, 223830, 223831, 2013) 223832, 244471, 274280, 312700 Caracal caracal FMNH 135042, 57220 - Lynx canadensis FMNH 129341 (Leche, 1915) 122 Felis silvestris FMNH 97861 (Leche, 1915) Octolobus manul MNHN 2010-646, 2009-251 - Prionailurus planiceps MNHN CG 1873-228 - Catopuma temminckii MNHN 1941-293 - Profelis aurata MNHN 1932-5233 - Leopardus pardalis FMNH 88887, 68895, 93174 (Schmidt-Kittler, 1976; Christiansen, 2008) Acinonyx jubatus AMNH C.A. 145071; FMNH 34589, (Leche, 1915; Van Valkenburgh et 29635 al., 1990) Miracinonyx trumani GRCA 21734 (Orr, 1969; Adams, 1979; Van Valkenburgh et al., 1990; Wang and Martin, 1993) Puma concolor FMNH 15532 (Van Valkenburgh et al., 1990; Christiansen, 2008) Neofelis nebulosa FMNH 75831 - Panthera pardus MNHN C.G. 1962-2884, MNHN (Leche, 1915; Christiansen, 2008) A7927, MNHN 1996-521 Panthera leo FMNH 31121; B 8707 (Leche, 1915; Hunt, 1987; Salles, 1992; Hsieh and Takemura, 1994; Mattern and McLennan, 2000; Christiansen, 2008; Lueders et al., 2012; De Schepper, 2016) Panthera leo spelaea MFN MB. Ma. 0948, 1956, 2246, (Anyonge, 1993) 1953, 1906, 30090, 30118, 30112a, 30114a, b; MFN MB. Ma. 14322, 14321, 14317, 14320, 30124, 3642, 3643, 1964 Panthera atrox AMNH 14397; LACMHC 572, 575, (Merriam and Stock, 1932; Wheeler 597, 1564, 2903-R-6, 14621, X-6716, and Jefferson, 2009) 14661, 14844, 14926, 14696, 14905, X-5036, R6740, 16311, 2911-R-1, 2907-R-4, 14722, 2908-R-7, 15297, 15460, 15379, 15536, 15582, X-7067, X-5552, 2937-2, X-5642, X-6274, X- 6405, X-7279, 16043, X-9759 Barbourofelis morrisi AMNH 79999; F:AM 61876, 61900, - 80000, 61895, 25204, 69359, 61893, 123 61976, 61898, 61970, 125665, 61882, Bx-5-Bx-69 #25138-B Oriensmilus liupanensis AMNH 144755 (cast of an (Wang et al., 2020) uncatalogued BNHM specimen) Sansanosmilus palmidens - (Filhol, 1890; Ginsburg, 1961; Morlo et al., 2004; Peigné, 2012) Barbourofelis fricki AMNH 108193 (cast of UNSM (Schultz et al., 1970; Joeckel and 76000); F:AM 61986, 61994, 61997, Stavas, 1996) 99258, 68234, 2672, 61984, 61983, 125670, 116854, 61991; UWBM 72291 Barbourofelis loveorum UF 24447, 36855, 36871, 37000, (Baskin, 1981; Neff, 1983; Hunt, 36867, 23796, 24432, 36867, 25081, 1987; Bryant, 1988a, 1988b, 1991) 25103, 25156, 25191, 25228, 25249, 25267, 25283, 25294, 25302, 27258, 37939, 25054, 25034, 36800, 25013, 24429; AMNH 125125 Albanosmilus whitfordi AMNH 14308; F:AM 61856, 61858, - 61861, 61864, 61849, 61680, 61885 Eusmilus cerebralis AMNH 6941; F:AM 98189, 69377; - UCMP 123180, 123181; JODA 7047 Hoplophoneus oharrai SDSM 2417 - Hoplophoneus primaevus F:AM 62007, AMNH 5338, 82440, (Bryant, 1988a; Barrett, 2016) 38980; USNM 18184; YPM PU 10741 Hoplophoneus occidentalis F:AM 102387, 62025, 62022; AMNH - 1407, 655; KUVP 2874, 2561; RAM 10356 Nimravus brachyops UCMP 1681, 2556, 76111; AMNH (Welsh et al., 2015; Barrett, 2016) 6930, 6993, 6940, 6933; F:AM 62020, 62151; JODA 1312 Nimravus intermedius AMNH 137130 (cast of FSM PQ 327), (Schmidt-Kittler, 1976; Peigné and 105390 (cast of CM 2587); MFN MB. De Bonis, 1999; Peigné, 2003; Egi Ma. 29987, 29988, 29986, 29991, et al., 2016) 29993, 29994, 29995; YPM PU 11569 Pogonodon platycopis AMNH 6938, 6953 (Barrett, 2016) Pogonodon davisi UCMP 789; AMNH 102156; F:AM (Barrett, 2016) 62026, 62042, 62024, 62018; JODA 5841; YPM 10520; YPM PU 11430 124 Dinictis felina AMNH 6937, 8777, 38805; YPM PU (Barrett, 2016) 12551, 13625, 12577, 11431 Nanosmilus kurteni UNSM 25505 - Ginsburgsmilus napakensis KNM-SO-5670; UM-P67-13 (Morales et al., 2001, 2008; Morlo et al., 2004; Morales and Pickford, 2018) Prosansanosmilus eggeri - (Morlo et al., 2004) Prosansanosmilus peregrinus - (Heizmann et al., 1980; Morlo et al., 2004; Morlo, 2006) Afrosmilus hispanicus - (Belinchón and Morales, 1989; Azanza et al., 1993; Morales et al., 2001) Afrosmilus turkanae KNM-MO-15929, KNM-RU-15984, (Morales et al., 2001; Werdelin and KNM-RU-15986, KNM-SO15973 Peigné, 2010; Morales and Pickford, 2018) Afrosmilus africanus - (Andrews, 1914; Savage, 1965; Morales et al., 2001) Albanosmilus jourdani - (Robles et al., 2013) Maofelis cantonensis - (Averianov et al., 2016) Quercylurus major - (Ginsburg, 1979; Peigné, 2003) Dinailurictis bonali MFN MB. Ma. 29985 (Ginsburg, 1979; Peigné, 2003) Eofelis edwardsii - (Ginsburg, 1979; Peigné, 2000; de Bonis et al., 2019) Dinaelurus crassus - (Eaton, 1922) MA-PHQ 348 - (Peigné, 2001) Eusmilus dakotensis YPM PU 11079; UNSM 1068 (Hatcher, 1895; Morea, 1975; Bryant, 1996) Eusmilus sicarius YPM PU 12953 A (cast of YPM PU - 12953) Eusmilus adelos USNM 12820, 18214 - Eusmilus bidentatus MFN MB. Ma. 30154, 30151 (Piveteau, 1931; Ringeade and Michel, 1994; Peigné and Brunet, 2001; Joeckel et al., 2002) Eusmilus villebramarensis - (Peigné and Brunet, 2001; Peigné, 125 2003) Prionodon linsang FMNH 88606 (Leche, 1915) Nandinia binotata AMNH 2409 C.A.; FMNH 149361, (Wesley-Hunt and Flynn, 2005; 55758 Spaulding and Flynn, 2012; Wible and Spaulding, 2013) Hemigalus derbyanus AMNH C.A. 9; FMNH 68717 (Leche, 1915) Arctictis binturong AMNH C.A. 1182; FMNH 53747 (Leche, 1915) Civettictis civetta FMNH 108174 (Leche, 1915) Viverra tangalunga FMNH 85116 (Leche, 1915) Genetta maculata FMNH 73044, 85987 - Paradoxurus hermaphroditus FMNH 338000, 140476 (Leche, 1915) Paguma larvata MNHN CG 1988-163, 1962-2062, - 1913-577 Cynogale bennettii MNHN CG 1962-170; MNHN A-2094 (Leche, 1915) Poiana richardsonii MNHN CG 1976-389 - Viverricula indica MNHN CG 1962-2113 (Leche, 1915) Atilax paludinosus MNHN CG 1995-426, 1877-153; - MNHN 1871-121 Herpestes edwardsii FMNH 83097 (Spaulding and Flynn, 2012) Ichneumia albicauda FMNH 73024, 157991 (Leche, 1915) Bdeogale nigripes FMNH 167685 - Mungos mungo FMNH 149365 - Suricata suricata FMNH 38348, 180674 - Crossarchus obscurus FMNH 54410 - Rhynchogale melleri MNHN CG 1962-992 - Galerella sanguinea MNHN CG 2000-1054, 2001-2188 - Helogale parvula MNHN CG1962-1056, 1987-176 - Fossa fossana FMNH 85196 - Cryptoprocta ferox FMNH 33950, 5655 (Carlsson, 1911) 126 Eupleres goudotii FMNH 30492, MNHN CG 1962-2105 - Salanoia concolor MNHN CG 1962-2111 - Kanuites lewisae - (Dehghani and Werdelin, 2008; Werdelin, 2019) Kichechia zamanae - (Savage, 1965; Schmidt-Kittler, 1987; Morales and Pickford, 2011; Adrian et al., 2018) Herpestides antiquus - (Chen and Schmidt-Kittler, 1983; Hunt, 1991; Morlo, 1996; Wolsan and Morlo, 1997; Wesley-Hunt and Flynn, 2005; Spaulding and Flynn, 2012) Euboictis aliverensis - (Schmidt-Kittler, 1983; Fejfar and Schmidt-Kittler, 1984) Sivanasua viverroides - (Fejfar and Schmidt-Kittler, 1984; Fejfar et al., 1997; Ginsburg and Morales, 1999) Izmirictis cani - (Morales et al., 2019) Palaeoprionodon lamandini - (Hunt, 1998, 2001; Wesley-Hunt and Flynn, 2005; Hans-Volker et al., 2007; Spaulding and Flynn, 2012) Stenogale julieni - (Hunt, 1998; Wesley-Hunt and Flynn, 2005; Spaulding and Flynn, 2012; Wang et al., 2019) Tapocyon robustus - (Wesley and Flynn, 2003; Wesley- Hunt and Flynn, 2005; Spaulding and Flynn, 2012) Procynodictis vulpiceps AMNH 2514 (Wesley-Hunt and Flynn, 2005; Spaulding and Flynn, 2012) Hesperocyon gregarius AMNH 50276 (Bryant, 1992; Wang, 1993, 1994; Wesley-Hunt and Flynn, 2005; Spaulding and Flynn, 2012) Canis latrans FMNH 135222 (Baryshnikov and Averianov, 1993b) Vulpes vulpes FMNH 64610 (Baryshnikov and Averianov, 1993b) 127 Taxidea taxus FMNH 47747 (Leche, 1915) References Adams, D. B. 1979. The cheetah:Native American. Science 205:1155–1158. Adrian, B., L. Werdelin, and A. Grossman. 2018. New Miocene Carnivora (Mammalia) from Moruorot and Kalodirr, Kenya. Palaeontologia Electronica 21. Andrews, C. W. 1914. On the Lower Miocene vertebrates from British East Africa, collected by Dr. Felix Oswald. Quarterly Journal of the Geological Society of London 70:163–186. Antón, A. M., M. J. Salesa, and G. Siliceo. 2013. Machairodont Adaptations and Affinities of the Holarctic Late Miocene Homotherin Machairodus (Mammalia, Carnivora, Felidae): The Case of Machairodus Catocopis Cope, 1887. Journal of Vertebrate Paleontology 33:1202– 1213. Antón, M., and L. Werdelin. 1998. Too well restored? The case of the Megantereon skull from Seneze. Lethaia Seminar 31:158–160. Antón, M., A. Turner, M. J. Salesa, and J. Morales. 2006. A complete skull of Chasmaporthetes lunensis (Carnivora, Hyaenidae) from the Spanish Pliocene site of la Puebla de Valverde (Teruel). Estudios Geologicos 62:375–388. Antón, M., M. J. Salesa, A. Galobart, and Z. J. Tseng. 2014. The Plio-Pleistocene scimitar- toothed felid genus Homotherium Fabrini, 1890 (Machairodontinae, Homotherini): Diversity, palaeogeography and taxonomic implications. Quaternary Science Reviews 96:259–268. Anyonge, W. 1993. Body mass in large extant and extinct carnivores. Journal of Zoology 231:339–350. 128 Averianov, A., E. Obraztsova, I. Danilov, P. Skutschas, and J. Jin. 2016. First nimravid skull from Asia. Scientific Reports 6:1–8. Azanza, B., E. Cerdeno, L. Ginsburg, J. Van Der Made, J. Morales, and P. Tassy. 1993. Les grands mammifères du Miocène inférieur d’Artesilla, bassin de Calatayud-Teruel (province de Saragosse, Espagne). Bulletin Du Museum National d’Histoire Naturelle Section C Sciences de La Terre Paleontologie Geologie Mineralogie 15:105–153. Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658. Baryshnikov, G. F., and A. O. Averianov. 1993a. Deciduous teeth of carnivorous mammals (Order Carnivora) Part V. Families Protelidae and Hyaenidae. Russian Academy of Sciences Proceedings of the Zoological Institute St. Petersburg 263:46–84. Baryshnikov, G. F., and A. O. Averianov. 1993b. Deciduous teeth of carnivorous mammals (Order Carnivora) Part IV. Families Amphicyonidae and Canidae. Russian Academy of Sciences Proceedings of the Zoological Institute St. Petersburg 249:158–196. 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. de Beaumont, G. 1968. Une intéressante mandibule de Hyaenidae (Carnivora) du Pontien de Samos. Archives Des Sciences 21:21–26. de Beaumont, G. 1969. Brèves remarques sur Plioviverrops Kretzoi (Carnivora). Bulletin de La Société Vaudoise Des Sciences Naturelles 70:1–7. Belinchón, M., and J. Morales. 1989. Los carnívoros del Mioceno Inferior de Buñol (Valencia, España). Revista Española de Paleontología 4:3–8. Berta, A. 1981. The Plio-Pleistocene hyaena Chasmaporthetes ossifragus from Florida. Journal of Vertebrate Paleontology 1:341–356. 129 de Bonis, L., A. Gardin, and C. Blondel. 2019. Carnivora form the early Oligocene of the “Phosphorites du Quercy” in southwestern France. Geodiversitas 41:601–621. Bryant, H. N. 1988a. The anatomy, phylogenetic relationships and systematics of the Nimravidae (Mammalia: Carnivora). University of Toronto, 414 pp. Bryant, H. N. 1988b. Delayed Eruption of the Deciduous Upper Canine in the Sabertoothed Carnivore Barbourofelis lovei (Carnivora, Nimravidae). Journal of Vertebrate Paleontology 8:298–306. Bryant, H. N. 1991. Phylogenetic relationships and systematics of the Nimravidae (Carnivora). Journal of Mammalogy 72:56–78. Bryant, H. N. 1992. The Carnivora of the Lac Pelletier Lower Fauna (Eocene: Duchesnean), Cypress Hills Formation, Saskatchewan. Journal of Paleontology 66:847–855. 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. Carlsson, A. 1911. Über Cryptoprocta ferox. Zoologische Jahrbücher Abteilung Für Systematik 30:419–470. de Castro, M. C., and M. C. Langer. 2008. New postcranial remains of Smilodon populator Lund, 1842 from southeastern Brazil. Revista Brasileira de Paleontologia 11:199–206. Chen, G. F., and N. Schmidt-Kittler. 1983. The deciduous dentition of Percrocuta Kretzoi and the diphyletic origin of the hyaenas (Carnivora, Mammalia). Paläontologische Zeitschrift 57:159–169. Christiansen, P. 2008. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics 24:977–992. 130 Christiansen, P. 2013. Phylogeny of the sabertoothed felids (Carnivora: Felidae: Machairodontinae). Cladistics 29:543–559. Christiansen, P., and J. M. Harris. 2005. Body size of Smilodon (Mammalia: Felidae). Journal of Morphology 266:369–384. Christiansen, P., and J. S. Adolfssen. 2007. Osteology and ecology of Megantereon cultridens SE311 (Mammalia; Felidae; Machairodontinae), a sabrecat from the Late Pliocene – Early Pleistocene of Senéze, France. Zoological Journal of the Linnean Society 151:833–884. Coca-Ortega, C., and J. A. Pérez-Claros. 2019. Characterizing ecomorphological patterns in hyenids: A multivariate approach using postcanine dentition. PeerJ 6:e6238. Collins, R. L. 1981. Scimitar cats, Homotherium serum Cope from Gassaway Fissure, Cannon County, Tennessee and the North American distribution of Homotherium. Journal of the Tennessee Academy of Science 56:15–19. Dehghani, R., and L. Werdelin. 2008. A new small carnivoran from the Middle Miocene of Fort Ternan, Kenya. Neues Jahrbuch Fur Geologie Und Palaontologie - Abhandlungen 248:233– 244. Eaton, G. F. 1922. John Day Felidae in the Marsh Collection. American Journal of Science 4:425–452. 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. Fejfar, O., and N. Schmidt-Kittler. 1984. Sivanasua und Euboictis n. gen. - zwei pflanzenfressende Schleichkatzenvorläufer (Viverridae, Carnivora, Mammalia) im europäischen Untermiozän. Mainzer Geowissenschaftliche Mitteilungen 13:49–72. 131 Fejfar, O., N. Schmidt-Kittler, and M. Rummel. 1997. Sivanasua viverroides (Schlosser, 1916) from the Lower Miocene fissure filling Rothenstein 1/13/. Münchner Geowissenschaftliche Abhandlungen Reihe A Geologie Und Paläontologie 34:93–110. Filhol, H. 1890. Études sur les Mammifères Fossiles de Sansan. Bibliothèque Des Hautes Études Section Des Sciences Naturelles 37:1–319. Ghaffar, A., M. Akhtar, M. A. Khan, and M. A. Babar. 2019. Comments on percrocuta carnifex (carnivora, percrocutidae) based on new fossil material from the nagri formation (middle siwaliks) of hasnot, pakistan. Geologica Acta 17:1–9. 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. Ginsburg, L. 1979. Révision taxonomique des Nimravini (Carnivora Felidae) de l’Oligocène des Phosphorites du Quercy. Bulletin Du Muséum National d’Histoire Naturelle Paris 4e série 1:35–49. Ginsburg, L., and J. Morales. 1999. Le genre sivanasua (Lophocyoninae, Hyaenodontidae, Creodonta, Mammalia) dans le miocene de france. Estudios Geológicos 55:173–180. Gracia, S. F. 2015. Estudio de Protictitherium crassum del Cerro de los Batallones (Torrejón de Velasco, Madrid): aportación a la filogenia y evolución de la familia hyaenidae. Universidad Complutense de Madrid, 365 pp. Hans-Volker, K., E. Gröning, and C. Brauckmann. 2007. Comment on a fossil civet skull from the Lower Oligocene of the Weisselster Basin (Saxonia, Germany). Studia Geologica Salmanticensia 43:215–225. Hatcher, J. B. 1895. Discovery, in the Oligocene of South Dakota, of Eusmilus, a genus of sabre- toothed cats new to North America. The American Naturalist 29:1091–1093. 132 Hearst, J. M., L. D. Martin, J. P. Babiarz, and V. L. Naples. 2011. Osteology and myology of Homotherium ischyrus from Idaho; pp. 123–186 in V. L. Naples, L. D. Martin, and J. P. Babiarz (eds.), The other saber-tooths: Scimitar-tooth cats of the Western Hemisphere. The John Hopkins University Press, Baltimore. Heizmann, E. P. J., L. Ginsburg, and C. Bulot. 1980. Prosansanosmilus peregrinus, ein neuer machairodontider Felide aus dem Miocän Deutschlands und Frankreichs. Stuttgarter Beiträge Zur Naturkunde B 58:1–27. Howell, F. C., and G. Petter. 1985. Comparative observations on some middle and upper Miocene hyaenids. Genera: Percrocuta Kretzoi, Allohyaena Kretzoi, Adcrocuta Kretzoi (Mammalia, Carnivora, Hyaenidae). Geobios 18:419–476. Hsieh, H.-M., and A. Takemura. 1994. The Rete Mirabile of the Maxillary Artery in the Lion (Panthera leo). Okajimas Folia Anatomica Japonica 71:1–12. Hunt Jr., R. M., and N. Solounias. 1991. Evolution of the aeluroid Carnivora: hyaenid affinities of the Miocene carnivoran Tungurictis spocki from Inner Mongolia. American Museum Novitates 1–25. Hunt, R. M. 1987. Evolution of the aeluroid Carnivora: significance of auditory structure in the nimravid cat Dinictis. American Museum Novitates 2886:1–74. Hunt, R. M. 1991. Evolution of the aeluroid Carnivora: viverrid affinities of the Miocene carnivoran Herpestides. American Museum Novitates 3023:1–34. 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. 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. 133 Joeckel, R. M., and J. M. Stavas. 1996. New Insights into the Cranial Anatomy of Barbourofelis fricki (Mammalia, Carnivora). Journal of Vertebrate Paleontology 16:585–591. 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. Koufos, G. D. 1995. The late Miocene percrocutas of Macedonia (Greece). Palaeovertebrata 24:68–84. Koufos, G. D. 2006. The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece: 4. Carnivora. Palaeontographica, Abteilung A: Palaozoologie - Stratigraphie 276:39–74. Koufos, G. D. 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 5. Carnivora. Beiträge Zur Paläontologie 31:57–105. Koufos, G. D. 2011. The Miocene carnivore assemblage of Greece. Estudios Geologicos 67:291–320. Koufos, G. D. 2012. New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley, Macedonia, Greece. Comptes Rendus Palevol 11:49–64. Koufos, G. D. 2016. Palaeontology of the upper Miocene vertebrate localities of Nikiti (Chalkidiki Peninsula, Macedonia, Greece). Carnivora. Geobios 49:53–67. Kovachev, D. 2012. A complete skeleton of Adcrocuta eximia (Roth and Wagner, 1854) from the Upper Maeotian (Turolian) of Hadzhidimovo, SW Bulgaria. Geologica Balcanica 41:77–95. Kovatchev, D. 2001. Description d’un squelette complet de Metailurus (Felidae, Carnivora, Mammalia) du Miocène supérieur de Bulgarie. Geologica Balcanica 31:71–88. 134 Kurtén, B., and L. Werdelin. 1990. Relationships between North and South American Smilodon. Journal of Vertebrate Paleontology 10:158–169. Leche, W. 1915. Zur Frage nach der stammesgeschichtlichen Bedeutung des Milchgebisses bei den Säugetieren. 11. Viverridae, Hyaenidae, Telidae, Mustelidae, Creodonta. Zoologische Jahrbücher. Abteilung Für Systematik, Geographie Und Biologie Der Tiere 38:275–370. Liu, J., J. Liu, H. Zhang, J. Wagner, Q. Jiangzuo, Y. Song, S. Liu, Y. Wang, and C. Jin. 2021. The giant short-faced hyena Pachycrocuta brevirostris (Mammalia, Carnivora, Hyaenidae) from Northeast Asia: A reinterpretation of subspecies differentiation and intercontinental dispersal. Quaternary International. Lueders, I., I. Luther, G. Scheepers, and G. van der Horst. 2012. Improved semen collection method for wild felids: Urethral catheterization yields high sperm quality in African lions (Panthera leo). Theriogenology 78:696–701. Martin, L. D., J. P. Barbiarz, and V. L. Naples. 2011. The osteology of a cookie-cutter cat, Xenosmilus hodsonae; pp. 42–97 in V. L. Naples, L. D. Martin, and J. P. Babiarz (eds.), The other saber-tooths: scimitar-tooth cats of the Western Hemisphere. Johns Hopkins University Press, Baltimore. Mattern, M. Y., and D. A. McLennan. 2000. Phylogeny and Speciation of Felids. Cladistics 16:232–253. Matthew, W. D. 1910. The phylogeny of the Felidae. Bulletin of the American Museum of Natural History 28:289–316. McDonald, H. G., and L. Werdelin. 2018. The Sabertooth Cat, Smilodon populator (Carnivora, Felidae), from Cueva del Milodón, Chile; pp. 53–75 in L. Werdelin, H. G. McDonald, and C. A. Shaw (eds.), Smilodon: The Iconic Sabertooth. Johns Hopkins University Press, Baltimore. Meade, G. E. 1961. The Saber-toothed Cat Dinobastis serus. Bulletin of the Texas Memorial Museum 2:23–60. 135 Merriam, J. C., and C. Stock. 1932. The Felidae of Rancho La Brea. The Carnegie Institution, Washington, 231 pp. Morales, J., and M. Pickford. 2006. A large Percrocutid Carnivore from the Late Miocene (ca. 10-9 Ma) of Nakali, Kenya. Annales de Paleontologie 92:359–366. Morales, J., and M. Pickford. 2011. Un nouveau carnivore paradoxuriné du Miocène supérieur des Siwaliks (Inde) et une révision des viverridés bunodontes d’Afrique. Geobios 44:271– 277. 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. Morales, J., M. Pickford, and M. J. Salesa. 2008. Creodonta and Carnivora from the Early Miocene of the Northern Sperrgebiet, Namibia. Memoirs of the Geological Survey of Namibia 20:291–310. 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. Morales, J., S. Mayda, A. Valenciano, D. DeMiguel, and T. Kaya. 2019. A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey. Journal of Systematic Palaeontology 17:1127–1138. Morea, F. M. 1975. On the species of Hoplophoneus and Eusmilus (Carnivora, Felidae). South Dakota School of Mines and Technology, 74 pp. Morlo, M. 1996. Carnivoren aus dem Unter-Miozän des Mainzer Beckens - 2. Mustelida, Pinnipedia, Feliformia, Palaeogale. Senckenbergiana Lethaea 76:193–249. 136 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. 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. Mutter, R. J., L. R. Berger, and P. Schmid. 2001. New evidence of the giant hyaena, Pachycrocuta brevirostris (Carnivora, Hyaenidae), from the Gladysvale cave deposit (Plio- Pleistocene, John Nash Nature Reserve, Gauteng, South Africa). Palaeontologia Africana 113:103–113. Neff, N. A. 1983. The basicranial anatomy of the Nimravidae (Mammalia: Carnivora) : character analyses and phylogenetic inferences. City University of New York, 642 pp. Orr, P. C. 1969. Felis trumani a new radiocarbon dated cat skull from Crypt Cave, Nevada. Bulletin of Santa Barbara Museum of Natural History Department of Geology 2:1–8. 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. Palmqvist, P., B. Martínez-Navarro, J. A. Pérez-Claros, V. Torregrosa, B. Figueirido, J. M. Jiménez-Arenas, M. Patrocinio Espigares, S. Ros-Montoya, and M. De Renzi. 2011. The giant hyena Pachycrocuta brevirostris: Modelling the bone-cracking behavior of an extinct carnivore. Quaternary International 243:61–79. Peigné, S. 2000. A new species of Eofelis (Carnivora: Nimravidae) from the Phosphorites of Quercy, France. Comptes Rendus de l’Académie Des Sciences, Paris 330:653–658. Peigné, S. 2001. A primitive nimravine skull from the Quercy fissures, France: implications for the origin and evolution of Nimravidae (Carnivora). Zoological Journal of the Linnean Society 132:401–410. 137 Peigné, S. 2003. Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32:199–229. Peigné, S. 2012. Les Carnivora de Sansan; pp. 559–660 in S. E. N. S. (ed.), Mammifères de Sansan. Muséum national d’Histoire naturelle, Paris. Peigné, S., and L. De Bonis. 1999. Le premier crane de Nimravus (Mammalia, Carnivora) d’Eurasie et ses relations avec N. brachyops d’Amerique du Nord. Revue De Paleobiolgie 18:57–67. Peigné, S., and M. Brunet. 2001. Une Nouvelle Espèce Du Genre Eusmilus (Carnivora: Nimravidae) De L’Oligocène (MP 22) D’Europe. Geobios 34:657–672. Piveteau, J. 1931. Les Chats des Phosphorites du Quercy. Annales de Paléontologie 20:107–163. Qiu, Z.-X., J. Y. Xie, and D. F. Yan. 1988a. Discovery of the skull of Dinocrocuta gigantea. Vertebrata PalAsiatica 26:128–138. Qiu, Z., J. Ye, and J. Cao. 1988b. A new species of Percrocuta from Tongxin, Ningxia. Vertebrata Palasiatica 26:116–127. Qiu, Z. -x., T. Deng, and B. -y. Wang. 2004. Early Pleistocene mammalian fauna from Longdan, Dongxiang, Gansu, China. Palaeontologia Sinica, New Series C 191:1–198. Rincón, A. D. 2006. A first record of the Pleistocene saber-toothed cat Smilodon populator Lund, 1842 (Carnivora: Felidae: Machairodontinae) from Venezuela. Ameghiniana 43:499– 501. Ringeade, M., and P. Michel. 1994. A propos de l’Eusmilus (Eusmilus bidentatus ringeadei Ringeade et Michel, 1994) de Soumailles, lieu-dit de la commune de Pardaillan, Canton de Duras (Lot et Garonne, France) : étude descriptive. Paléo 6:5–37. 138 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. Rothwell, T. P. 2001. Phylogenetic systematics of North American Pseudaelurus (Carnivora: Felidae). Columbia University, 358 pp. Rothwell, T. P. 2003. Phylogenetic systematics of North American Pseudaelurus (Carnivora, Felidae). American Museum Novitates 3403. Salesa, M. J. 2002. Estudio anatómico, biomecánico, paleoecológico y filogenético de Paramachairodus ogygia (Kaup, 1832) Pilgrim, 1913 (Felidae, Machairodontinae) del yacimiento vallesiense (Mioceno superior) de Batallones-1 (Torrejón de Velasco, Madrid). Universidad Complutense de Madrid, Madrid, 370 pp. 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. 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. 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. Salles, L. O. 1992. Felid Phylogenetics: Extant Taxa and Skull Morphology (Felidae, Aeluroidea). American Museum Novitates 3047:67. 139 Sardella, R., and D. A. Iurino. 2012. The latest Early Pleistocene sabertoothed cat Homotherium (Felidae, Mammalia) from Monte Peglia (Umbria, central Italy). Bollettino Della Societa Paleontologica Italiana 51:15–22. Savage, R. J. G. 1965. Fossil Mammal of Africa: The Miocene Carnivora of East Africa. Bulletin of the British Museum (Natural History) 10:239–316. De Schepper, M. 2016. A comparative study in morphological defects of semen from African Lions (Panthera leo) and Caracal (Caracal caracal): collected by urethral catheterization and electro-ejaculation. Ghent University, 31 pp. Schmidt-Kittler, N. 1976. Raubtiere aus dem Jungtertiär Kleinasiens. Palaeontographica Abteilung A 155:1–131. Schmidt-Kittler, N. 1983. The Mammals from the Lower Miocene of Aliveri (Island of Evia, Greece). III. On a new species of Sivanasua Pilgrim, 1931 (Feliformia, Carnivora) and the phylogenetic position of this genus. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, C 86:301–318. Schmidt-Kittler, N. 1987. The Carnivora (Fissipeda) from the lower Miocene of East Africa. Palaeontographica A197:85–126. 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. Siliceo, G., M. J. Salesa, M. Antón, M. F. G. Monescillo, and J. Morales. 2014. Promegantereon ogygia (Felidae, Machairodontinae, Smilodontini) from the Vallesian (late Miocene, MN 10) of Spain: morphological and functional differences in two noncontemporary populations. Journal of Vertebrate Paleontology 34:407–418. Spaulding, M., and J. J. Flynn. 2012. Phylogeny of the Carnivoramorpha: The impact of postcranial characters. Journal of Systematic Palaeontology 10:653–677. 140 Tseng, Z. J., and W. J. Binder. 2010. Mandibular biomechanics of Crocuta crocuta, Canis lupus, and the late Miocene Dinocrocuta gigantea (Carnivora, Mammalia). Zoological Journal of the Linnean Society 158:683–696. Van Valkenburgh, B., F. Grady, and B. Kurtén. 1990. The Plio-Pleistocene cheetah-like cat Miracinonyx inexpectatus of North America. Journal of Vertebrate Paleontology 10:434– 454. Vekua, A. 1995. Die Wirbeltierfauna des Villafranchium von Dmanisi und ihre biostratigraphische Bedeutung. Jahrbuch Des Romisch-Germanischen Zentralmuseums Mainz 42:77–180. Wallace, S. C., and R. C. Hulbert. 2013. A New Machairodont from the Palmetto Fauna (Early Pliocene) of Florida, with Comments on the Origin of the Smilodontini (Mammalia, Carnivora, Felidae). PLoS ONE 8:e56173. Wang, X. 1993. Transformation from plantigrady to digitigrady: functional morphology of locomotion in Hesperocyon (Canidae: Carnivora). American Museum Novitates 1–23. Wang, X. 1994. Phylogenetic systematics of the Hesperocyoninae (Carnivora, Canidae). Bulletin of the American Museum of Natural History 221. Wang, X. 2004. New materials of Tungurictis (Hyaenidae, Carnivora) from Tunggur Formation, Nei Mongol. Vertebrata PalAsiatica 42:145–153. Wang, X., and L. D. Martin. 1993. Natural Trap Cave. National Geographic Research & Exploration 9:422–435. 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. 141 Wang, X., Q. Li, L. Sun, C. Deng, Q. Li, and W. Dusu. 2019. Biostratigraphy, magnetostratigraphy, and geochronology of lower Miocene Aoerban strata in Central Inner Mongolia. Palaeogeography, Palaeoclimatology, Palaeoecology 518:187–205. Welsh, E., C. A. Boyd, K. Spearing, and P. Z. Barrett. 2015. Stratigraphic and taxonomic revision of a North American False Saber-toothed Cat Cub. Proceedings of the South Dakota Academy of Science 94:141–153. Werdelin, L. 2019. Middle miocene carnivora and hyaenodonta from fort Ternan, Western Kenya. Geodiversitas 41:267–283. Werdelin, L., and N. Solounias. 1991. The Hyaenidae : taxonomy, systematics and evolution. Fossils and Strata 30:104. Werdelin, L., and B. Kurten. 1999. Allohyaena (Mammalia: Carnivora): giant hyaenid from the Late Miocene of Hungary. Zoological Journal of the Linnean Society 126:319–334. Werdelin, L., and M. E. Lewis. 2001. A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132:147–258. Werdelin, L., and S. Peigné. 2010. Carnivora; pp. 603–657 in L. Werdelin and W. J. Sanders (eds.), Cenozoic Mammals of Africa. University of California Press, Berkeley, Los Angeles, London. 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. Wesley, G. D., and J. J. Flynn. 2003. a Revision of Tapocyon (Carnivoramorpha), Including Analysis of the First Cranial Specimens and Identification of a New Species. Journal of Paleontology 77:769–783. 142 Wheeler, H. T., and G. T. Jefferson. 2009. Panthera atrox: body proportions, size, sexual dimorphism, and behavior of the cursorial lion of the North American plains. Museum of Northern Arizona Bulletin 65:423–444. Wible, J. R., and M. Spaulding. 2013. On the cranial osteology of the African palm civet, Nandinia Binotata (Gray, 1830) (Mammalia, Carnivora, Feliformia). Annals of Carnegie Museum 82:1–114. Wolsan, M., and M. Morlo. 1997. The status of “Plesictis” croizeti, “Plesictis” gracilis and “Lutra” minor: synonyms of the early Miocene viverrid Herpestides antiquus (Mammalia, Carnivora). Bulletin of the Natural History Museum London (Geol) 53:1–9. Xiong, W.-Y. 2019. Basicranial morphology of Late Miocene Dinocrocuta gigantea (Carnivora: Hyaenidae) from Fugu, Shaanxi. Vertebrata Palasiatica 57:274–307. Zhang, Z. 2005. New materials of Dinocrocuta (Percrocutidae, Carnivora) from Lantian, Shaanxi Province, China, and remarks on Chinese Late Miocene biochronology. Geobios 38:685– 689. 143 Appendix D Morphological Character List 1. Lacrimal facial process: (0), broad rostral flange; (1), small, present on face; (2), not present on face; (3), orbital flange reduced to area around lacrimal foramen (Wesley-Hunt and Flynn, 2005: character 1). 2. Ventral exposure of premaxilla; posterior extent of premaxilla, lateral to palatal foramen: (0), lateral to canine; (1), anterior to canine (Wesley-Hunt and Flynn, 2005: character 2). 3. Shape of infraorbital foramen: (0), elongate; (1), round (Wesley-Hunt and Flynn, 2005: character 3). 4. Length of palate – position of the posterior edge of palatine midline relative to tooth row: (0) posterior to upper tooth row; (1) anterior or equal to upper tooth row (Wesley-Hunt and Flynn, 2005: character 5). 5. Palatine canal primary anterior opening: (0), opening through palatine; (1), at maxilla–palatine suture; (2) opening through maxilla (Wesley-Hunt and Flynn, 2005: character 6). 6. Relative length of frontal and parietal at midline: (0) parietal greater than frontal; (1) parietal equal or subequal to frontal; (2) frontal midline much longer than parietal (Wesley-Hunt and Flynn, 2005: character 7). 144 7. Postorbital process: (0) prominent; (1) small, reduced (Wesley-Hunt and Flynn, 2005: character 8). 8. Paroccipital process size: (0) well-developed; (1) reduced (Wesley-Hunt and Flynn, 2005: character 9). 9. Paroccipital process shape: (0) simple process; (1) laterally flattened, thin, but is distinct process; (2) cupped around bulla, process not distinct; (3) absent (Wesley-Hunt and Flynn, 2005: character 10). 10. Placement of postglenoid foramen: (0) medially placed; (1) more lateral, external, very near edge of skull (Wesley-Hunt and Flynn, 2005: character 11). 11. Postglenoid foramen: (0) present; (1) greatly reduced, or missing (Wesley-Hunt and Flynn, 2005: character 12). 12. Shape of mastoid process: (0) forming a distinct process, extending out farther than paroccipital process, or subequal; (1) blunt, rounded, does not protrude significantly, more a swelling of the mastoid; (2) thin plate, no distinct process; (3) large tabular process, anteriorly sloping, obscuring much, if not all, of the auditory bulla; (4) large tabular process, extremely anteriorly inclined, closing or nearly enclosing the external auditory meatus by its junction with the glenoid pedicle, medially encroaching and affixed to the ventral floor of the bulla in the 145 vicinity of the ectotympanic (modified from Wesley-Hunt and Flynn, 2005: character 13 to incorporate states of Christiansen (2013): character 35). 13. Direction of mastoid process extension: (0) lateral – ventral; (1) ventral; (2) lateral; (3) none, or only swelling (Wesley-Hunt and Flynn, 2005: character 14). 14. Condyloid (hypoglossal) foramen position relative to postlacerate foramen: (0) distant; (1) close (less than the diameter of the hypoglossal foramen away); (2) conjoined with posterior lacerate foramen (Wesley-Hunt and Flynn, 2005: character 15). 15. Condyloid (hypoglossal) foramen position relative to groove between the occipital condyle and the paroccipital process: (0) inline or within groove; (1) anterior to groove (Wesley-Hunt and Flynn, 2005: character 16). 16. Posterior lacerate foramen: (0), present as a vacuity between the promontorium and the basioccipital; (1), present as an individual foramen (Wesley-Hunt and Flynn, 2005: character 17). 17. Fenestra cochleae (rotunda) position relative to mastoid tubercle: (0) posterior to mastoid tubercle; (1) anterior, subequal to mastoid tubercle (Wesley-Hunt and Flynn, 2005: character 18). 146 18. Relative distance between the foramen ovale and the alisphenoid canal: (0) separated by at least the diameter of the alisphenoid canal; (1) separated only by a thin wall; (2) no alisphenoid canal present (Wesley-Hunt and Flynn, 2005: character 19). 19. Ossification of tegmen tympani: (0) facial nerve exposed ventrally; (1) facial nerve partially embedded within tegmen tympani and floored in anteromedial segment; (2) facial nerve beneath a bony sheath that defines the fossa for tensor tympani muscle (Wesley-Hunt and Flynn, 2005: character 20). 20. Composition of mastoid tubercle: (0) mastoid tubercle formed by petrosal; (1) mastoid tubercle formed by squamosal (Wesley-Hunt and Flynn, 2005: character 22). 21. Anterior loop of internal carotid artery: (0) lack of an anterior loop of the internal carotid artery; (1) presence of the loop – excavation in basisphenoid; (2) presence of loop – but extrabullar (Wesley-Hunt and Flynn, 2005: character 23). 22. Suprameatal fossa (fossa on squamosal anterior to mastoid: (0) absent; (1) small; (2) large, well developed (Wesley-Hunt and Flynn, 2005: character 24). 23. Position of internal carotid artery: (0) internal carotid artery laterally positioned, transpromontorial, runs close to margin of fenestra cochlea, presence of a promontory artery, groove for stapedial artery may or may not be present; (1) internal carotid artery transpromontorial but medially positioned, course far from fenestra cochlea; (2) internal carotid 147 artery medial, extrabullar, inside a bony canal formed by the caudal entotympanic (Wesley-Hunt and Flynn, 2005: character 25). 24. Apron shelf on promontorium posterior to fenestra cochleae for entotympanic attachment: (0) absent; (1) blunt – surface present posterior to fenestra cochleae, but no extensive attachment possible; (2) extended, large area for attachment, may roof posterior bullar chamber (Wesley- Hunt and Flynn, 2005: character 26). 25. Ventral process of promontorium: (0) absent; (1) present, medially positioned on promontorium (Wesley-Hunt and Flynn, 2005: character 27). 26. Shape of the promontorium, anterior extension: (0) elongate, apron extension tapers to a point anteriorly, almond like in appearance; (1) elongate, rounded anteriorly; (2) blunt, quickly truncating; (3) elongate, apron is broad, flat extension, not almond-shaped and not blunt (Wesley-Hunt and Flynn, 2005: character 28). 27. Facet on promontorium indicative of ectotympanic contact: 0, absent; 1, present (Wesley- Hunt and Flynn, 2005: character 29). 28. Inferior petrosal sinus: (0) inferior petrosal sinus small; (1) inferior petrosal sinus greatly enlarged (Wesley-Hunt and Flynn, 2005: character 31). 148 29. A deep, well developed fossa or pit on the squamosal/alisphenoid recording the contact with the anterior crus or anterior face of the ectotympanic: (0) absent, may have slight/shallow indentation; (1) present, well developed, or bulla present and fully ossified (Wesley-Hunt and Flynn, 2005: character 32). 30. Shelf between mastoid process and paroccipital process: (0) laterally wide, curved trough with smooth surface; (1) laterally wide, could have flat surface, rugose or bulbous, no smoothed out trough; (2) very thin, outside edge could be raised; (3) no shelf present (Wesley-Hunt and Flynn, 2005: character 33). 31. Extent of flange on basiocciptal lateral edge bordering auditory region: (0) absent; (1) small, nascent; (2) well developed when compared to basal ‘miacids’ (Wesley-Hunt and Flynn, 2005: character 34). 32. Evidence on basisphenoid and basioccipital for marked medial inflation of the entotympanic: (0) absent; (1) present – inflation of entotympanic pushing medially onto and over the basioccipital (Wesley-Hunt and Flynn, 2005: character 35). 33. Evidence of marked posterior inflation of the entotympanic; entotympanic attached during life to paroccipital process or to extensive area posterior to the petrosal: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 36). 149 34. Epitympanic wing of the petrosal forms ventral floor to the anterior medial corner of the fossa for the tensor tympani muscle: (0) absent; (1) present, but relatively flat and horizontal; (2) ventral floor present, but not horizontal, instead it forms a delicate ‘tube’, the bony floor is not an extension of the petrosal (Wesley-Hunt and Flynn, 2005: character 38). 35. Placement of middle lacerate foramen: (0) foramen a vacuity – not defined anteriorly nor posteriorly, positioned directly anterior to petrosal; (1) foramen anteriorly defined, posteriorly bordered by petrosal – positioned equal or posterior to basisphenoid/basioccipital suture; (2) foramen defined anteriorly, petrosal may be undefined posterior border, foramen positioned in basisphenoid (or edge of alisphenoid) just anterior to basisphenoid/basioccipital suture; (3) foramen defined anteriorly and posteriorly completely bordered by basisphenoid, foramen positioned far anterior to basisphenoid/basioccipital suture; (4) lack of a foramen (Wesley-Hunt and Flynn, 2005: character 40). 36. M1, a defined cingulum continuous around the lingual face of the protocone: (0) absent; (1) complete cingulum present; (2) anterior segment of cingulae absent or smaller than posterior cingulae (Wesley-Hunt and Flynn, 2005: character 41). 37. M1, protocone height relative to paracone: (0) protocone shorter than paracone; (1) protocone equal or subequal to height of paracone; (2) protocone absent or lacks a cusp associated with the root (modified from Wesley-Hunt and Flynn, 2005: character 42). 150 38. M1, parastyle projects farther labially than metastyle: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 44) 39. M1, parastyle direction: (0) buccally with anterior direction; (1) buccally (Wesley-Hunt and Flynn, 2005: character 45). 40. M1, size of posterior lingual cingular shelf at base of protocone: (0) posterior lingual cingular shelf equal or subequal to anterior cingulum; (1) posterior lingual cingular shelf more pronounced, larger than anterior cingulum (Wesley-Hunt and Flynn, 2005: character 47). 41. M1, relative height of paracone and metacone: (0) paracone equals metacone in height; (1) paracone greater than metacone; (2) metacone absent (modified from Wesley-Hunt and Flynn, 2005: character 48). 42. M1, relative height of paraconule and metaconule: (0) paraconule greater than metaconule; (1) paraconule equal or subequal to metaconule; (2) both absent (Wesley-Hunt and Flynn, 2005: character 49). 43. M1, presence of hypocone: (0) absent; (1) present (distinct cusp); (2) present, formed by swelling of entire cingulum ridge (Wesley-Hunt and Flynn, 2005: character 50). 44. M1, width of parastylar shelf: (0) lack of a shelf; (1) broad; (2) narrow, consisting mainly of ridge (Wesley-Hunt and Flynn, 2005: character 51). 151 45. Presence of M3: (0) present; (1) absent (Wesley-Hunt and Flynn, 2005: character 53). 46. P4 protocone: (0) large, well-developed; (1) reduced or absent (Wesley-Hunt and Flynn, 2005: character 56). 47. Posterior accessory cusps on P3: (0) one cusp present; (1) two cusps present; (2) absent (Wesley-Hunt and Flynn, 2005: character 58). 48. Palatine, relative size: (0) midline length of palatine less than midline length of maxilla; (1) midline length greater than midline length of maxilla (Wesley-Hunt and Flynn, 2005: character 60). 49. Turbinal bones: (0) simple development of maxilloturbinals in nasal cavity; (1) maxilloturbinals large and branching, excluding nasoturbinals from narial opening (Wesley-Hunt and Flynn, 2005: character 62). 50. Jugal: (0) jugal reaches lacrimal, or is separated from it by only thin sliver of maxilla; (1) jugal widely separated from lacrimal, maxilla broadly laps posteriorly over anterior orbital rim (Wesley-Hunt and Flynn, 2005: character 64). 51. Anterior extent of palatine in orbit: (0) broadly contacts lacrimal; (1) fails to contact lacrimal (Wesley-Hunt and Flynn, 2005: character 65). 152 52. Postorbital constriction: (0) just anterior of frontoparietal suture, near posterior margin of frontal; (1) braincase expanded, with frontals making much greater contribution; fronto–parietal suture located more anteriorly in frontal (Wesley-Hunt and Flynn, 2005: character 66). 53. Posterior entrance of carotid artery into auditory capsule: (0) posterior entry, artery not enclosed in osseous tube; (1) posterior entry, artery enclosed in tube; (2) anterior entry, artery not enclosed in tube; (3) anterior entry, artery not enclosed in tube (Wesley-Hunt and Flynn, 2005: character 67). 54. Entotympanic: (0) fails to ossify, or is only weakly attached to auditory capsule; (1) ossified at least partially, and firmly fused to the skull (Wesley-Hunt and Flynn, 2005: character 68). 55. Ectotympanic contributes to external auditory meatal tube: (0) no; (1) yes (Wesley-Hunt and Flynn, 2005: character 69). 56. Ectotympanic septum: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 70). 57. Entotympanic septum: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 71). 58. Fenestra cochleae: (0) approximately equal in size to fenestra ovalis, cochlear fossula not developed; (1) at least three times the area of oval window, cochlear fossula well developed (Wesley-Hunt and Flynn, 2005: character 72). 153 59. Major a2 arterial shunt: (0) small; (1) large, intracranial rete (Wesley-Hunt and Flynn, 2005: character 75). 60. Major a4 arterial shunt: (0) present; (1) present, intracranial rete (Wesley-Hunt and Flynn, 2005: character 76). 61. Major anastomosis x: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 77). 62. P1: (0) present: (1) absent (Wesley-Hunt and Flynn, 2005: character 79). 63. P3 lingual cusp: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 80). 64. P4 hypocone: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 83). 65. p1: (0) present; (1) absent (Wesley-Hunt and Flynn, 2005: character 84). 66. Lower molars: (0); subequal in size; (1); m1 much larger than m2–3 and progressive decrease in size from m1–3 (Wesley-Hunt and Flynn, 2005: character 86). 67. M2 hypocone: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 87). 68. m3: (0) present; (1) absent (Wesley-Hunt and Flynn, 2005: character 88). 154 69. Baculum: (0) small and simple or absent; (1) long, stylized (Wesley-Hunt and Flynn, 2005: character 89). 70. Scapula, postscapular fossa: (0) absent; (1) present (Wesley-Hunt and Flynn, 2005: character 90). 71. Tail: (0) long; (1) reduced (Wesley-Hunt and Flynn, 2005: character 91). 72. Femur, third trocanter: (0) present; (1) absent (Wesley-Hunt and Flynn, 2005: character 94). 73. Cowper’s (bulbourethral) gland: (0) present; (1) absent (Wesley-Hunt and Flynn, 2005: character 95). 74. Prostate gland: (0) small/vestigial; (1) large, ampulla bilobed (Wesley-Hunt and Flynn, 2005: character 96). 75. Anal glands: (0) simple; (1) enlarged and having enlarged anal sac (Wesley-Hunt and Flynn, 2005: character 98). 76. Scapula – supraglenoid tubercle morphology: (0) expands out over the glenoid fossa (Didymictis); (1) blunt, does not extend over the glenoid fossa (Vulpavus) (Spaulding and Flynn, 2012: character 100). 155 77. Scapula – acromion process angle: Either a short or a long process can be dorsally directed at the acromion process apex, or remain in the same plane as the scapular spine: (0) process angles dorsally (Vulpavus); (1) process remains in same plane as the scapular spine (Canis) (Spaulding and Flynn, 2012: character 101). 78. Scapula – acromion process length: The acromion process can terminate far distally to the glenoid fossa or terminate at or before the border. A long acromion can be used as a proxy for the presence of clavicles in extinct taxa. (0) process extends past glenoid fossa (Vulpavus); 1– process terminates before or at glenoid fossa (Gulo) (Spaulding and Flynn, 2012: character 102). 79. Scapula – coracoid process: (0) present, as a large clearly projecting process (Felis); (1) extremely small or absent (Canis) (Spaulding and Flynn, 2012: character 103). 80. Scapula – scapular spine morphology: Coded while viewing the spine from a ventral view of the scapula. (0) spine forms one continuous, smooth, downward curve (Öodectes); (1) spine has a dip or embayment in the curve (Herpestes) (Spaulding and Flynn, 2012: character 104). 81. Scapula – metacromion process: (0) present and well developed (Herpestes); (1) weak or absent (Otariidae) (Spaulding and Flynn, 2012: character 105). 82. Humerus – indentation on the anterior surface of the capitulum: (0) present (Hyeanodon); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 106). 156 83. Humerus – olecranon fossa shape: (0) shallow and round (Vulpavus); (1) deep and slot-like (Miacis uintensis); (2) perforated (Didymictis) (Spaulding and Flynn, 2012: character 107). 84. Humerus – medial edge of posterior trochlea: (0) vertical (Vulpavus); (1) slanted (Öodectes) (Spaulding and Flynn, 2012: character 108). 85. Humerus – delto-pectoral crest: (0) present (Vulpavus); (1) absent (Didymictis) (Spaulding and Flynn, 2012: character 109). 86. Humerus – medial epicondyle: (0) ends with a well rounded head (Vulpavus); (1) poorly defined, appearing more like a blunt tubercle (Didymictis) (Spaulding and Flynn, 2012: character 110). 87. Humerus – medial epicondyle orientation (new): (0) lacks curvature (is straight) (Öodectes); (1) curves posteriorly (Didymictis) (Spaulding and Flynn, 2012: character 111). 88. Humerus – greater tuberosity height: (0) extends past head (Didymictis); (1) flush with head in height (Vulpavus) (Spaulding and Flynn, 2012: character 112). 89. Humerus – posterior trochlea: (0) not bound by high ridges (Öodectes); (1) bound by high ridges (Canis) (Spaulding and Flynn, 2012: character 113). 157 90. Humerus – brachial flange: (0) present and large, extending out from the body of the bone as a flat surface (Vulpavus); (1) small, nothing but a small raised line of bone (Didymictis) (Spaulding and Flynn, 2012: character 114). 91. Humerus – lesser tuberosity with a crest or ridge of bone leading from this feature down the shaft: (0) present (Didymictis); (1) absent (Canis) (Spaulding and Flynn, 2012: character 115). 92. Humerus – trochlea extent: (0) extends distally past capitulum when viewed anteriorly (Didymictis); (1) two articular surfaces are more inline (Vulpavus) (Spaulding and Flynn, 2012: character 116). 93. Humerus – epicondylar foramen: (0) present and round (Hyaenodon); (1) present and elongated (Vulpavus) (Spaulding and Flynn, 2012: character 117). 94. Humerus – distal L-shaped ridge of bone on capitulum in distal view: (0) present (Vulpavus) (1) absent (Felis) (Spaulding and Flynn, 2012: character 118). 95. Humerus – orientation of bone on the lateral distal margin: (0) faces laterally (Hyaenodon); (1) faces posteriorly (Vulpavus) (Spaulding and Flynn, 2012: character 119). 96. Humerus – ulnar collateral ligament insertion site size: (0) very large, forming a distinct circular pit (Didymictis); (1) small, forming only a shallow depression (Vulpavus) (Spaulding and Flynn, 2012: character 120). 158 97. Humerus – greater tuberosity angle: (0) greater tuberosity angled away from head; smooth arch is not formed with lesser tuberosity (Didymictis); (1) greater tuberosity more flush with head (Vulpavus) (Spaulding and Flynn, 2012: character 121). 98. Humerus – prominence of bicipital groove: (0) groove is very noticeable and deep (Nandinia); (1) groove is very subtle if noticeable at all (Civetticits) (Spaulding and Flynn, 2012: character 122). 99. Humerus – capitulum shape: (0) flat for the whole length with a uniform distal margin (Thinocyon); (1) rounded (Vulpavus) (Spaulding and Flynn, 2012: character 123). 100. Humerus - tricipital line: (0) large and distinctive (Didymictis); (1) reduced (Nandinia) (Spaulding and Flynn, 2012: character 124). 101. Ulna – semi-lunar notch distal border morphology: (0) W-shaped distal border; an indentation occurs between the articular surface with the radius and the rest of the facet (Öodectes); (1) indentation lacking (Canis) (Spaulding and Flynn, 2012: character 125). 102. Ulna – semi-lunar notch proximal border extent: (0) proximal ridge extends far from shaft surface (Hyeanodon); (1) proximal ridge flush with shaft surface (Öodectes) (Spaulding and Flynn, 2012: character 126). 159 103. Ulna – m. bracialis insertion site: (0) present (Vulpavus); (1) absent (Ailurus) (Spaulding and Flynn, 2012: character 127). 104. Ulna – m. bracialis insertion site position: (0) on anterior surface of ulna (Vulpavus); (1) on the medial margin of the shaft (Öodectes) (Spaulding and Flynn, 2012: character 128). 105. Ulna – radial notch curvature: (0) absent (Vulpavus); (1) present (Ailurus) (Spaulding and Flynn, 2012: character 129). 106. Ulna – deep tendon groove on proximal end of ulna: (0) present (Hyaenodon); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 130). 107. Ulna – deep tendon groove on proximal end of ulna (Heinrich and Rose, 1997): (0) present (Hyaenodon); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 131). 108. Ulna – olecranon process shape: (0) mediolaterally robust, square in shape (Vulpavus); (1) mediolaterally compressed (Felis) (Spaulding and Flynn, 2012: character 132). 109. Ulna – anconeal process extent: (0) projecting from the shaft, shelf like (Didymictis); (1) flush with shaft (Vulpavus) (Spaulding and Flynn, 2012: character 133). 110. Ulna – proximal border of semi-lunar notch, position: (0) lateral to the shaft (Hyaenodon); (1) centered with respect to the shaft (Öodectes) (Spaulding and Flynn, 2012: character 134). 160 111. Ulna – anteriormedial protuberuance development (Taylor, 1974): (0) well developed (Nandinia); (1) small (Canis) (Spaulding and Flynn, 2012: character 135). 112. Ulna – groove on lateral side of shaft: (0) present (Hyaenodon); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 136). 113. Ulna shape – sigmoidal; the ulna is used as a proxy for the curved nature of all forelimb bones noted by Matthew (1909): (0) sigmoidal (Vulpavus); (1) straight (Canis) (Spaulding and Flynn, 2012: character 137). 114. Ulna – radial notch orientation (Heinrich and Rose, 1997): (0) faces anteriorly (Didymictis); (1) faces more laterally (Vulpavus) (Spaulding and Flynn, 2012: character 138). 115. Radius – radial head shape: (0) round (Vulpavus); (1) oval (Didymictis) (Spaulding and Flynn, 2012: character 139). 116. Radius – large scaphoid articulation surface: (0) present (Didymictis); (1) surface small (Vulpavus) (Spaulding and Flynn, 2012: character 141). 117. Radius – capitular eminence of radial head development; (0) small (Vulpavus); (1) large, disrupts radial rim (Canis) (Spaulding and Flynn, 2012: character 142). 161 118. Radius – distal articulation surface shape: (0) with a deep sulcus (Erinaceus); (1) smooth, lacks a deep sulcus (Vulpavus) (Spaulding and Flynn, 2012: character 143). 119. Carpus – cuneiform shape in proximal view: (0) triangular (Nandinia); (1) rectangular (Vulpavus) (Spaulding and Flynn, 2012: character 144). 120. Carpus – proximal surface of scapholunar, curvature: (0) fully convex (Nandinia); (1) convex and concave (Herpestes) (Spaulding and Flynn, 2012: character 145). 121. Carpus – uniciform width (new): (0) as wide as trapezoid and magnum combined (Canis); (1) small, only as wide as magnum alone (Nandinia) (Spaulding and Flynn, 2012: character 146). 122. Carpus – trapezium position (Yalden, 1970): (0) on the posterior surface of the trapezoid (Canis); (1) on the radial surface of the trapezoid (Felis) (Spaulding and Flynn, 2012: character 147). 123. Carpus – metacarpal III overlaps IV proximally (Bryant, 1991): (0) absent (Canis); (1) present (Vulpavus) (Spaulding and Flynn, 2012: character 148). 124. Carpus – distal surface of scapholunar shape (Yalden, 1970): (0) relatively smooth/even (Canis); (1) a sharp projection exists between the magnum and trapezoid (Ursus) (Spaulding and Flynn, 2012: character 149). 162 125. Carpus – cuneiform articulation with ulna, position: (0) on medial (radial) margin (Felis); (1) on distal surface of ulna (Ursus) (Spaulding and Flynn, 2012: character 150). 126. Carpus –metacarpal II strongly overlaps III proximally: (0) overlap small or absent (Canis); (1) overlap present and substantial (Felis) (Spaulding and Flynn, 2012: character 151). 127. Carpus – metacarpal length: (0) metacarpal lengths are equivalent to sum of phalanges (Vulpavus); 1– phalanges are longer than metacarpals (Otariidae) (Spaulding and Flynn, 2012: character 152). 128. Carpus – medial phalanx distal articular surface symmetry: (0) symmetrical (Vulpavus); (1) asymmetrical (Felis) (Spaulding and Flynn, 2012: character 153). 129. Carpus – proximal phalanges compressed dorsoventrally: (0) absent (Vulpavus); (1) present (Hoplophoneus) (Spaulding and Flynn, 2012: character 154). 130. Carpus – lateral excavation of the medial phalanx; (0) absent (Vulpavus); (1) present (Felis) (Spaulding and Flynn, 2012: character 155). 131. Femur – lesser trochanter orientation: (0) projects posteriorly (Didymictis); (1) projects medially (Vulpavus) (Spaulding and Flynn, 2012: character 156). 163 132. Femur – intertrochanteric crest extent: (0) extends to lesser trochanter (Didymictis); (1) becomes flush with shaft before reaching the lesser trochanter (Vulpavus) (Spaulding and Flynn, 2012: character 157). 133. Femur – position of lesser trochanter relative to the third trochanter: (0) third trochanter is lower (Vulpavus); (1) third trochanter is roughly at the same level (Didymictis) (Spaulding and Flynn, 2012: character 158). 134. Femur – supracondylar tuberosities: (0) absent (Vulpavus); (1) presence of two or more raised tuberosities just proximal to the condyles of the femur on the posterior shaft (Canis) (Spaulding and Flynn, 2012: character 160). 135. Femur – height of greater trochanter relative to head of femur: (0) greater trochanter is higher or sub-equal (Vulpavus); (1) greater trochanter is lower than head of femur (Ursus) (Spaulding and Flynn, 2012: character 161). 136. Femur – medial condyle morphology: (0) proximal edge of the articular surface of condyles are flush with the shaft, due to the lack of development of a condylar neck (Vulpavus); (1) proximal edge of the articular surface of condyles are not flush with the shaft, due to the development of a condylar neck (Canis) (Spaulding and Flynn, 2012: character 162). 164 137. Femur – lesser trochanter development: (0) strong, prominently projecting feature (Vulpavus); (1) small, barely more than a small bump or ridge on the shaft (Ursus) (Spaulding and Flynn, 2012: character 163). 138. Femur – greater trochanter broadest surface, orientation: (0) faces laterally (Vulpavus); (1) faces posteriorly (Canis) (Spaulding and Flynn, 2012: character 164). 139. Femur – patellar groove shape: (0) deep and narrow (Didymictis); (1) wide and flat (Vulpavus) (Spaulding and Flynn, 2012: character 166). 140. Femur – position of greater trochanter relative to shaft: (0) greater trochanter over distal midline (Erinaceus); (1) greater trochanter lateral to midline (Didymictis) (Spaulding and Flynn, 2012: character 167). 141. Tibia – deep groove on the posterior surface of the tibia (Matthew, 1909): (0) present (Vulpavus); (1) absent (Canis) (Spaulding and Flynn, 2012: character 169). 142. Tibia – posterior bone spur on distal tibia: (0) presence of a prominent bone spur on the posterior margin (Vulpavus); (1) spur absent (Nandinia) (Spaulding and Flynn, 2012: character 170). 143. Tibia – contact of inner borders of condyles: (0) in contact (Vulpavus); (1) separate (Canis) (Spaulding and Flynn, 2012: character 171). 165 144. Tibia – internal (medial) malleolus morphology: (0) forms a clear and distinct well-defined point (Vulpavus); (1) indistinct, forming a general extension of the medial surface of the tibia (Civettictis) (Spaulding and Flynn, 2012: character 172). 145. Pes – dorsal surface of astragalus, shape (Matthew, 1909): (0) smooth (Vulpavus); (1) grooved (Canis) (Spaulding and Flynn, 2012: character 174). 146. Pes – astragalus – height of fibular facet: (0) height subequal to length, resulting in a roughly square shaped facet (Vulpavus); (1) height much less than length, resulting in a more crescent shaped feature (Canis) (Spaulding and Flynn, 2012: character 175). 147. Pes – astragalar head shape when viewed dorsally: (0) rounded (Vulpavus); (1) flattened (Canis) (Spaulding and Flynn, 2012: character 176). 148. Pes – astragalus fossa, on the lateral edge, posterior ventral quadrant: (0) present (Didymictis); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 178). 149. Pes – astragalus lateral margin, anterior ventral edge expansion: (0) lateral expansion present (Vulpavus); (1) expansion absent, creating a much smoother margin (Procyon) (Spaulding and Flynn, 2012: character 179). 166 150. Pes – astragalar head medial articulation surface extent: (0) extends backwards a great distance, past the border of the sustentacular facet (Hyaenodon); (1) short, does not extend past the start of the neck (Vulpavus) (Spaulding and Flynn, 2012: character 180). 151. Pes – astragalus sustentacular facet shape: (0) has clear edges, is somewhat convex, and does not extend into gully (Vulpavus); (1) is more flat than convex and extends into gully (Canis) (Spaulding and Flynn, 2012: character 181). 152. Pes – astragalar foramen; (0) present and dorsally positioned (Vulpavus); (1) present and posteriorly positioned (Hyaenodon); (2) absent (Canis) (Spaulding and Flynn, 2012: character 182). 153. Pes – astragalus, posterior ridge for ligament attachments, presence: (0) present, high (Vulpavus); (1) low or absent (Felis) (Spaulding and Flynn, 2012: character 183). 154. Pes – astragalus, orientation of posterior ridge for ligament attachments: (0) orientated at an oblique angle relative to the long axis of the astragalar trochlea (Vulpavus); (1) orientated in line with the trochlea (Canis) (Spaulding and Flynn, 2012: character 184). 155. Pes – astragalus-tibia articulation surface, extent: (0) covers entire posterior dorsal surface (Canis); (1) restricted, fails to cover entire posterior dorsal surface, leaving a gap in the lateral posterior quadrant (Vulpavus) (Spaulding and Flynn, 2012: character 185). 167 156. Pes – astragalus, cotylar fossa presence: (0) present (Didymictis); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 187). 157. Pes – Calcaneus, sustentaculum position (Heinrich and Rose, 1997): (0) far from anterior border, roughly in the middle of the calcaneus (Didymictis); (1) close to anterior border (Vulpavus) (Spaulding and Flynn, 2012: character 188). 158. Pes – Calcaneus, peroneal tubercle development: (0) weakly developed, little more than a ridge (Didymictis); (1) well developed (Vulpavus) (Spaulding and Flynn, 2012: character 189). 159. Pes – astragalus neck development: (0) absent (lacks a clearly defined neck of astragalus) (Erinaceus); (1) present (Vulpavus) (Spaulding and Flynn, 2012: character 190). 160. Pes – calcaneus, cuboid facet shape: (0) irregularly shaped (Didymictis); (1) round (Vulpavus) (Spaulding and Flynn, 2012: character 191). 161. Pes – calcaneus, cuboid facet orientation: (0) angled dorsally (Vulpavus); (1) in planes other than dorsal (Didymictis) (Spaulding and Flynn, 2012: character 192). 162. Pes – calcaneus, fibular facet (Flynn and Galiano, 1982): (0) present (Didymictis); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 193). 168 163. Pes – calcaneus –dorsal facet morphology: (0) smooth (Vulpavus); (1) clearly defined, sharp corner (rather than a smooth curve) (Canis) (Spaulding and Flynn, 2012: character 194). 164. Pes – cuboid, contact with metatarsal V (Bryant, 1991): (0) small (less than 40%) articular surface for metatarsal V (Dinictis); (1) large, articular surface for metatarsal V at least 40% of distal surface (Canis) (Spaulding and Flynn, 2012: character 195). 165. Pes – cuboid shape: (0) relatively rectangular (Didymictis); (1) wider proximally than distally (Vulpavus) (Spaulding and Flynn, 2012: character 196). 166. Pes – cuboid, tubercle for the long plantar ligament, position (Heinrich and Rose, 1997): (0) distal to the ectocuneiform articulation facet (Didymictis); (1) even with the ectocuneiform articulation facet (Vulpavus) (Spaulding and Flynn, 2012: character 197). 167. Pes – cuboid/astragalus contact (Matthew, 1909): (0) present (Thinocyon); (1) absent (Vulpavus) (Spaulding and Flynn, 2012: character 198). 168. Pes – navicular height: (0) height less than width (Vulpavus); (1) height roughly equal to or greater than width (Canis) (Spaulding and Flynn, 2012: character 199). 169. Pelvis – ilium, anterior expansion: (0) not expanded (Felis); (1) expanded dorsoventrally (Canis) Primitively for Carnivoramorpha the anterior ilium is not dorsoventrally expanded; 169 however in Canis, Ursus, Hyaena, and Civettictis a clear expansion is seen (Spaulding and Flynn, 2012: character 200). 170. Pelvis – ischial spine position: (0) located just posterior of the border of the acetabulum (Didymictis); (1) far posterior from acetabulum (Vulpavus); (2) spine absent (Erinaceus) (Spaulding and Flynn, 2012: character 201). 171. Pelvis – dorsal margin of acetabulum, position: (0) even with or above dorsal surface of pelvis (Vulpavus); (1) ventral to dorsal surface of pelvis (Canis) (Spaulding and Flynn, 2012: character 202). 172. Pelvis – angle at pubic symphysis: (0) ‘U-like’ (Canis); (1) ‘V-like’ (Felis) (Spaulding and Flynn, 2012: character 203). 173. Pelvis – ilium, anterior region morphology: (0) broad and flat (Canis); (1) divided by a distinct ridge running antero-posteriorly (Vulpavus) (Spaulding and Flynn, 2012: character 204). 175. Pelvis – ilium, ventral surface morphology: (0) broad and flat (Thinocyon); (1) narrow (Vulpavus) (Spaulding and Flynn, 2012: character 205). 176. Atlas – alar foramen: (0) absent, only a notch is present (Canis); (1) present, confluent with lateral vertebral foramen (Herpestes); (2) present, separate from lateral vertebral foramen (Procyon) (Spaulding and Flynn, 2012: character 207). 170 177. Atlas – transverse processes, orientation: (0) projects at right angle from the body (Canis); (1) extends posteriorly at an acute angle to the body (Nandinia) (Spaulding and Flynn, 2012: character 208). 178. Atlas – ventral arch length (Gaubert et al., 2005): (0) equal to or shorter than dens of axis (Canis); (1) longer than dens of axis (Civettictis) (Spaulding and Flynn, 2012: character 209). 179. Axis – body length: (0) roughly as long as it is wide (Vulpavus); (1) elongated relative to its width (Felis) (Spaulding and Flynn, 2012: character 210). 180. Cervical vertebrae – keel: (0) large ventral keel present (Nandinia); (1) ventral keel absent (Dinictis) (Spaulding and Flynn, 2012: character 211). 181. Cervical vertebrae – spinous processes, size: (0) large (Canis); (1) small (Felis) (Spaulding and Flynn, 2012: character 212). 182. Cervical vertebrae – dorsal lateral margin shape: (0) concave (Öodectes); (1) convex (Felis) (Spaulding and Flynn, 2012: character 213). 183. Sacrum – size compared to pelvis: (0) small, does not reach border of acetabulum (Canis); (1) large, reaches border of acetabulum (Felis) (Spaulding and Flynn, 2012: character 215). 171 184. Sacrum – neural spines: (0) clear and distinct from one another (Nandinia); (1) fused to one another, appearing as one long keel (Canis) (Spaulding and Flynn, 2012: character 216). 185. Alisphenoid, foramen ovale (position): (0) anterior to glenoid fossa; (1) medial to glenoid fossa. Comparisons made to the posterior-most edge of the glenoid fossa, not the process. Serves for the passage of the trigeminal nerve (V), internal maxillary artery and the medial meningeal artery (modified from Gaubert et al., 2005: character 71). 186. Basisphenoid, medial lacerate foramen: (0) covered by bullae; (1) anteriomedial to bullae (e.g., Herpestes). [Serves for the passage of the ascending pharyngeal artery.] (Gaubert et al., 2005: character 79). 187. Ectotympanic/entotympanic division: (0) not externally visible; (1) externally visible with infolding at junction, without clear suture; (2) externally visible, with clear suture. [In the viverrids, the point of contact between the two elements is marked by an inbending of the plates of bone. In the other feliforms, although one can detect the point of contact, it is because of structural differences.] (Modified from Gaubert et al., 2005: character 85). 188. Entotympanic, ventral floor ossification: (0) ossified; (1) unossified (i.e. Nandinia) (modified from Gaubert et al., 2005: character 102). 172 189. Vagina processus hyoideus: (0) confluent with stylomastoid foramen (giving the appearance of one foramen); (1) slightly separate from stylomastoid foramen (Gaubert et al., 2005: character 129). 190. i1- i3 projection: (0) extends upward; (1) procumbent (less than 45 degrees) (Gaubert et al., 2005: character 151). 191. C1, flanges: (0) posterior flange present; (1) absent; (2) anterior and posterior flanges present (Gaubert et al., 2005: character 156). 192. P3 number or roots: (0) two roots; (1) one root; (2) three roots (Gaubert et al., 2005: character 164). 193. P3, posterolingual cingulum: (0) present, reduced; (1) present, broad; (2) vestigial/absent (Gaubert et al., 2005: character 165). 194. p3, talonid: (0) absent; (1) present (Gaubert et al., 2005: character 169). 195. P4, parastyle: (0) absent/vestigial; (1) present, small (less than size of protocone); (2) present, large (near equal to protocone); (3) present, larger than protocone (Gaubert et al., 2005: character 172). 173 196. P4 posterolingual cingulum: (0) present; (1) vestigial; (2) absent (Gaubert et al., 2005: character 177). 197. p4 talonid: (0) absent/vestigial; (1) present (Gaubert et al., 2005: character 178). 198. p4 posterior accessory cusps: (0) 2 cusps, linear; (1) 1 cusp; (2) no cusps; (3) 2 or more cusps, nonlinear (Gaubert et al., 2005: character 179). 199. P4-M1 carnassial embrasure pit: (0) present; (1) absent (Gaubert et al., 2005: character 181). 200. M1 roots: (0) three roots; (1) two roots; (2) one root (Gaubert et al., 2005: character 183). 201. m1, trigonid: (0) present, considerably higher than talonid; (1) present, near equal in height to talonid; (2) absent (modified from Gaubert et al., 2005: character 187). 202. m1, trigonid proportion: (0) length less than talonid; (1) length near equal length of talonid; (2) trigonid length is 60% to 73% of total length; (3) 77-87%; (4) 88% and higher; (5) no talonid (modified from Gaubert et al., 2005: character 188; Barrett, 2016: character 30). 203. m1 metaconid: (0) large; (1) considerably smaller than paraconid; (2) absent (Gaubert et al., 2005: character 189). 174 204. m1, protoconid : (0) present; (1) absent (Gaubert et al., 2005: character 190). 205. m1 entoconid: (0) present; (1) absent (Gaubert et al., 2005: character 191). 206. m1 hypoconid: (0) present; (1) absent (Gaubert et al., 2005: character 192). 207. m1, hypoconulid: (0) absent; (1) present (Gaubert et al., 2005: character 193). 208. m1, small cusp between hypoconid and protoconid: (0) absent; (1) present (Gaubert et al., 2005: character 194). 209. M2, paracone and metacone: (0) metacone and paracone present, paracone larger; (1) metacone absent; (2) tooth without distinguishable cusps; ? = tooth absent (Gaubert et al., 2005: character 195). 210. M2, roots: (0) three roots; (1) two roots; (2) one root; (3) tooth absent (modified from Gaubert et al., 2005: character 198). 211. m2, cusps: (0) high trigonid with extremely elongate talonid due to large hypoconulid, tooth elongate oval outline; (1) talonid not elongate, tooth oval outline, no enlarged hypoconulid; (2) low trigonid, extremely elongate with low, well developed talonid cusps; (3) low trigonid, well developed talonid cusps; (4) tooth with cusps but no clear trigonid or talonid; (5) tooth without clearly defined cusps; (6) absent (modified from Gaubert et al., 2005: character 202). 175 212. Buccal cingulum, upper molars: (0) small/not developed; (1) enlarged (Gaubert et al., 2005: character 206). 213. Mandible, mandibular symphysis: (0) strong (well fused); (1) weak (not well fused) (Gaubert et al., 2005: character 212). 214. Mandible, medial shelf of angular process: (0) present, small ridge; (1) present, wide shelf; (2) absent (Gaubert et al., 2005: character 214). 215. p4 anterior accessory cusp: (0) absent; (1) smaller than posterior; (2) about equal to posterior; (3) larger than posterior (Gaubert et al., 2005: character 221). 216. Volume contribution of the feloid auditory bulla: (0) caudal entotympanic envelopes ectotympanic chamber (e.g. Felidae); (1) both chambers meet over the petrosal and display an external ‘infolding’ at their juncture with a septum oblique to the long axis of the skull, such that the caudal entotympanic penetrates the anterointernal (rostromedial) corner of the auditory region (e.g. viverrids); (2) both chambers meet over the petrosal and display an external ‘infolding’ at their juncture with a septum orthogonal to the long axis of the skull (e.g. herpestids); (3) ectotympanic chamber dominates with caudal entotympanic chamber restricted to posterior third of bulla with the division separating anterior and posterior chambers rostral of the mastoid and paroccipital processes (e.g. Ictitheriines); (4) ectotympanic chamber dominates bulla, partially or totally overlaying caudal entotympanic chamber via ventral overgrowth, 176 recumbent septum concealing posterior chamber within mastoid and paroccipital process (e.g. Crocuta, Hyaena): (5) anterior chamber formed by rostral entotympanic or pseudoseptum. 217. Mastoid invasion by bulla: (0) absent; (1) present. 218. P4 nimravid “parastyle”: (0) absent; (1) present (Barrett et al., 2021: character 222). 219. P2: (0) double-rooted; (1) single-rooted; (2) absent (Barrett et al., 2021: character 224). 220. p3 posterior cusps: (0) no cusps; (1) one cusp (posterior cingular cusp); (2) two cusps (posterior cingular cusp and posterior accessory cusp) (Barrett et al., 2021: character 229). 221. Basicranial foramina: (0) petrobasilar venous sinus and sigmoid sinus merge intracranially and exit the posterior lacerate (jugular) foramen; (1) veins merge extracranially, passing through the petrobasilar and posterior lacerate foramina respectively (Barrett et al., 2021: character 231). 222. P4 protocone: (0) medial or posterior to paracone; (1) anterior to paracone or paracone- parastyle juncture; (2) anterior face of protocone approximately level with anterior face of parastyle; (3) protocone extending anteriorly of parastyle (modified from Werdelin and Solounias, 1991: character 9). 223. Shape of dentary tooth row: (0), tooth row straight; (l), tooth row curved, or convex (Werdelin and Solounias, 1991: character 10). 177 224. P4 metacone/metastyle blade length to paracone length ratio: (0): 0.54 or less; (1): 0.6-0.99; (2): 1-1.3; (3): 1.4 and greater (modified from Werdelin and Solounias, 1991: character 11). 225. Placement of camassials in tooth row: (0) carnassials in line with tooth row, i.e. at an angle relative to the sagittal plane); (l) carnassials parallel to sagittal plane (Werdelin and Solounias, 1991: character 14). 226. Placement of anterior accessory cusp of p4: (0) free of main cusp; (l) appressed to main cusp (Werdelin and Solounias, 1991: character 14). 227. Shape of anterior face of p3: (0) concave/straight; (l) convex (Werdelin and Solounias, 1991: character 15). 228. Position of infraorbital foramen: (0) positioned anterior to middle of P3; (1) positioned above middle to posterior of P3; (2) positioned above anterior edge of P4; (3) positioned above mid-posterior portion of P4 (modified from Werdelin and Solounias, 1991: character 22). 229. Position of anterior margin of orbit: (0) above or behind metastylar blade of P4; (1) above the anterior end of P4; (2) above P3 (modified from Werdelin and Solounias, 1991: character 23). 230. Suture between premaxillary and frontal on snout: (0) absent; (l) present (Werdelin and Solounias, 1991: character 24). 178 231. Size of inferior oblique muscle fossa at maxillary-lacrimal suture postero-dorsal to infra- orbital foramen: (0) small; (l) large; (2) absent (modified from Werdelin and Solounias, 1991: character 25). 232. Inferior oblique muscle Fossa II at juncture between maxillary, lacrimal, and Frontal: (0) absent; (l) present (Werdelin and Solounias, 1991: character 26). 233. Sphenoid foramen and postpalatine foramen position: (0) well separated, distinct foramina; (l) foramina located close together in a single depression (Werdelin and Solounias, 1991: character 27). 234. The contribution of the maxilla to the antero-internal rim of the zygomatic arch: (0) small to none; (l) maxillary makes up a substantial portion of the antero-dorsal margin of the zygomatic arch (Werdelin and Solounias, 1991: character 28). 235. Shape of basioccipital in ventral view: (0) flat; (l) lateral ridges and central groove (Werdelin and Solounias, 1991: character 32). 236. Position of premaxillary-maxillary suture on palate: (0) near the middle of the incisive fossa; (l) at the postero-lateral margin of the incisive fossa (Werdelin and Solounias, 1991: character 33). 179 237. Shape of jugal-maxillary suture in external view: (0) strongly sinusoid; (1) angled downwards posteriorly in a straight or weakly sinusoid manner; (2) horizontally straight; (3) vertical (modified from Werdelin and Solounias, 1991: character 36). 238. Position of external auditory meatus: (0) far forwards of nuchal crest; (l) level with nuchal crest (Werdelin and Solounias, 1991: character 43). 239. Shape of nuchal crest: (0) antero-posteriorly inclined; (1) nearly vertical or vertical (Werdelin and Solounias, 1991: character 44). 240. Size of metacarpal 1: (0) metacarpal 1 has a long, slender diaphysis resembling the other four metacarpals; (1) metacarpal 1 has a blunt and rectangular shape (e.g. felids); (2) vestigial (modified from Werdelin and Solounias, 1991: character 45). 241. Angle of scapular spine in posterior view: (0) angled; (1) straight (Werdelin and Solounias, 1991: character 47). 242. Relative width of p3 (ranked in order by ratio of Wp3 to Lp3): (0) 0.2-0.39; (1) 0.4-0.6; (2) 0.61-0.66; (3) 0.69 and greater (Tseng and Wang, 2007: character 12). 243. Relative width of p4 (ranked in order by ratio of Wp4 to Lp4): (0) 0.38 and less; (1) 0.40- 0.54; (2) 0.55-0.68; (3) 0.72 and greater (Tseng and Wang, 2007: character 13). 180 244. Relative width of P3 (ranked in order by ratio of WP3 to LP3): (0) 0.38 and less; (1) 0.4- 0.65; (2) 0.66-0.79; (3) 0.81 and above (Tseng and Wang, 2007: character 14). 245. Labiolingual compression of the C1: (0) very slightly compressed (length/breadth index smaller than 1.5); (1) markedly compressed (index between 1.5 and 1.8); (2) very compressed (index 1.81-3.0); (3) extremely compressed 4.0 and greater (Robles et al., 2013: character 3). 245. C1 vertical grooves: (0) absent; (1) present (Robles et al., 2013: character 4). 247. P3 anterior cingulum cusp: (0) absent; (1) present (Robles et al., 2013: character 7). 248. P4 preparastyle: (0) absent; (1) present (Robles et al., 2013: character 10). 249. p2: (0) present; (1) absent (Robles et al., 2013: character 16). 250. Orbit: (0) open; (1) closed (Robles et al., 2013: character 23). 251. Position of M1. Polarity: (0) distal to P4; (1) medial to P4 (Robles et al., 2013: character 27). 252. c1: (0) larger than i3; (1) same size (Robles et al., 2013: character 31). 181 253. Ratio P4 length /p4 length: (0) 0.93 and less; (1) 1.0-1.87; (2) 1.95-2.1; (3) 2.2 and higher (modified from Robles et al., 2013: character 32). 254. Masseteric fossa on the lateral surface of the maxilla and jugal: (0) shallow or absent; (1) deep with distinct dorsal margin (Barrett, 2016: character 3). 255. Fossa on the medial face of the zygomatic arch, below the postorbital process: (0) no fossa; (1) presence of a marked fossa (Peigné, 2003: character 12). 256. Zygomata shape in dorsal view: (0) broadly circular; (1) triangular (Barrett, 2016: character 5). 257. Shape of the glenoid socket of the craniomandibular joint: (0) anterior lip is missing; (1) posterior lip of the glenoid socket projects more ventrally than anterior lip; (2) anterior lip and posterior lip project equally ventrally (Barrett, 2016: character 10). 258. Anterior mandible position: (0) in line with the tooth row, mandibular border of cheek tooth row is in the same plane as the mandibular border of the incisors and canines; (1) elevated above tooth row; (2) cheek teeth and anterior teeth brought again into same plane by elevation of cheek teeth on pedestal (Barrett, 2016: character 13). 259. Development and orientation of the coronoid process of the dentary: (0) posteriorly 182 orientated posterior border without surpassing the level of the mandibular condyle; (1) surpassing the level of the mandibular condyle; (2) vertically orientated posterior border; (3) anteriorly orientated posterior border (Barrett, 2016: character 20). 260. Size of the genial flange of the dentary in adult taxa. Measured as the height of the genial flange from the anterior portion of the postcanine diastema to the ventral apex of the genial flange/length of dentary from the posterior articular surface to the most anterior aspect: (0) no flange, the ventral rim of the chin is regularly curved; (1) no flange, but the ventral rim of the chin is distinctly angulate; (2) short flange, between 22 and 31% of the total length of mandible; (3) deep flange, 32-50% of the total length of the mandible; (4) extremely deep flange, 54% or more of the total mandibular length (Barrett, 2016: character 15). 261. Mental fossa. Fossa on the ventral-internal face of the chin: (0) no fossa; (1) fossa present and marked (Barrett, 2016: character 16). 262. Incisors: (0) spatulate; (1) mostly spatulate, I3 caniform; (2) all caniform (Barrett, 2016: character 17). 263. C1 length. Mesial-distal length of C1 measured at the dentine/enamel boundary: (0) less than that of P4; (1) greater than that of P4 (Barrett, 2016: character 18). 264. Serration density of permanent upper canines per millimeter. Measured over an average of 5 mm: (0) none; (1) 0.7-2.7; (2) 2.8-5.0; (3) ≥ 5.5 (modified from Barrett, 2016: character 19). 183 265. Size of P3 vs. size of P4. Measured as a ratio of crown height (base of cingulum to apex of tooth) on adult minimally worn teeth: (0) 0.69-1.1; (1) 1.2 and greater; (2) 0.48-0.69; (3) 0.45 and lower (modified from Barrett, 2016: character 20). 266. p3 crown height compared to p4: (0) 0.6-0.98; (1) 1.0 and greater; (2) less than 0.58; (3) p3 absent (modified from Barrett, 2016: character 26). 267. Cheek tooth serrations. Serrations on adult minimally worn cheek teeth: (0) absent; (1) present (Barrett, 2016: character 31). 268. Articulation between the calcaneum and navicular : (0) absent; (1) present (Barrett, 2016: character 33). 269. Anterior palatine canal opening location: (0) level with P3; (1) between P2 and P3; (2) P4; (3) M1; (4) P2 or anterior (modified from Bryant, 1996: character 4). 270. Position of nasals relative to maxilla-frontal suture: (0) nasals extend beyond the maxillofrontal suture; (1) nasals shortened, the posterior border lies across or anterior to the maxillofrontal suture (Peigné, 2003: character 9). 271. P4, orientation of the protocone: (0) anterolingually projected; (1) lingually projected (modified from Peigné, 2003: character 23). 184 272: Processus muscularis on malleus: (0) short, cone-shaped and with tapering apex; (1) distinctly enlarged and cylindrical; (2) absent (modified from Christiansen, 2008: character 2). 273: Lambdoidal-squamous sutures: (0) converging along lamboid crest; (1) separate (modified from Christiansen, 2008: character 12). 274. Dorsal part of frontoparietal (coronal) suture: (0) smooth, lacking a parietal process; (1) suture irregular with a distinct, often very large, medio-dorsally directed parietal process (Christiansen, 2008: character 13). 275. Dorsal Jugal-squamosal suture in zygomatic arch: (0) abuts the postorbital process; (1) does not abut the postorbital process (modified from Christiansen, 2008: character 19). 276. Paroccipital process: (0) ventrally directed; (1) posteriorly directed (modified from Christiansen, 2008: character 22). 277. Angle of long axis of bulla to long axis of skull: (0) Less than or equal to 19°; (1) 20°-27°; (2) 28°-37° (modified from Christiansen, 2008: character 29). 278: Entotympanic process: (0) narrow and raised; (1) wide and flattened; (2) absent (modified from Christiansen, 2008: character 30). 185 279. Upper incisor arcade: (0) strongly parabolic; (1) slightly parabolic such that the anterior edge of I3 does not contact a straight edge held across the anterior of the arcade; (2) straight (modified from Christiansen, 2013: character 1). 280. P4 anterior cingulum: (0) indistinct; (1) distinct and often forming a ridge; (2) forming a distinct ectoparastyle (Christiansen, 2013: character 11). 281. Mediolateral width of P3 across the metacone relative to width across the paracone: (0) 56 and less; (1) 58- 68; (2) narrow (~70–93%); (3) wide (~95–130%) (modified from Christiansen, 2013: character 17). 282. Relative height of the m1 major cusps, measured from tip of cusp to base of cingulum: (0) 61% and lower; (1) height of paraconid low compared with height of protoconid (65–93%); (2) height of paraconid tall compared with height of protoconid (95–130%). (modified from Christiansen, 2013: character 24). 283. Relative length of p4: (0) 7% and lower; (1) small relative to mandible length (~7.5–9%); (2) large relative to mandible length (~10–15%) (modified from Christiansen, 2013: character 26). 284. Snout area elevation compared with braincase: (0) snout area low, line from dorsal narial aperture to juncture of occipital/sagittal crest lies below frontal postorbital process; (1) snout area elevated, line passes above frontal postorbital process (modified from Christiansen, 2013: character 34). 186 285. Size of the jugal postorbital process: (0) small, rounded, often almost absent; (1) tall, triangular and tapering (Christiansen, 2013: character 39). 286. Relative width across the upper incisor arcade compared to cranial basilar length (CBL): (0) 11% and less; (1) narrow (12–15% of CBL); (2) wide (16–19% of CBL); (3) extremely wide (21–25% of CBL) (modified from Christiansen, 2013: character 42). 287. Palatal region relative width compared to CBL: (0) 14% and lower; (1) 16-22%; (2) ~23– 25%; (3) palatal region across center of P3 relatively wide (~27–34% of CBL); (4) palatal region across center of P3 extremely wide (~36–43% of CBL) (modified from Christiansen, 2013: character 43). 288. Mandibular fossa termination: (0) posterior to the carnassial; (1) terminating below talonid or before carnassial notch: (2) well anterior of the posterior edge of the carnassial, frequently terminating around carnassial saddle or even at the M1/P4 junction (modified from Christiansen, 2013: character 49). 289. Height of horizontal dentary ramus (below m1) relative to ramus length (articular to symphysis between i1s): (0) 14 and less; (1) slender (~15–17%); (2) robust (~18–25%) (modified from Christiansen, 2013: character 50). 187 290. Lower canines, buccal vertical groove/ridge: (0) present; (1) absent (modified from Salesa et al., 2012: character 22). 291. C1 lingual ridge: (0) absent; (1) present (Salles, 1992: character 11). 292. Relative position of p3 and p4: (0) aligned; (1) not aligned (Salesa et al., 2010: character 24). 293. Palatal ridge/groove: (0) no visibly obvious groove or ridge (but may be felt by touch); (1) visibly obvious but shallow groove with ridges; (2) deep groove with distinctive ridges; (3) extreme medial ridges displacing medial palate ventrally from the main plane of the palate (Sakamoto and Ruta, 2012: character 39). 294. Stylomastoid groove: (0) groove originating from the stylomastoid foramen; (1) groove originating from the tympanohyal pit (Sakamoto and Ruta, 2012: character 41). 295. Metatarsal 1: (0) functional metatarsal, articulating with a phalanx; (1) vestigial metatarsal 1 (Rothwell, 2003: character 20). 296. Orientation of premaxillary-maxillary suture relative to upper tooth row: (0) 50 and less; (1) 52-70°; (2) moderately steep, 70-80°; (3) very steep, greater than 80° (modified from Van Valkenburgh et al., 1990: character 3). 188 297. Shape of sphenopalatine foramen: (0) oval with long axis oriented anteroposteriorly; (1) round/oval with long axis oriented dorsoventrally (Van Valkenburgh et al., 1990: character 12). 298. Teres major fossa: (0) absent; (1) small, triangular; (2) long (modified from Van Valkenburgh et al., 1990: character 20). 299. Radius, orientation of distal ulnar facet: (0) horizontal; (1) vertical (Van Valkenburgh et al., 1990: character 23). 300. Tibia, cnemial crest in lateral view: (0) straight; (1) concave (Van Valkenburgh et al., 1990: character 27). 301. Fibula, shape of proximal end: (0) without anterior projection; (1) expanded anteriorly with bony process (Van Valkenburgh et al., 1990: character 29). 302. Patella, ridge separating articular facets: (0) absent; (1) present (Van Valkenburgh et al., 1990: character 30). 303. Scapholunar, articular surface for trapezoid: (0) convex and concave; (1) concave only (Van Valkenburgh et al., 1990: character 37). 304. Pisiform, shape of cuneiform facet: (0) oval; (1) triangular (Van Valkenburgh et al., 1990: character 41). 189 305. Astragalus and calcaneum articulation: (0) astragalus reaches distal end of calcaneum; (1) astragalus does not reach distal end of calcaneum (Van Valkenburgh et al., 1990: character 42). 306. Seventh lumbar vertebra, shape of transverse processes: (0) similar to preceding vertebrae; (1) shorter and angled anteriorly (Van Valkenburgh et al., 1990: character 43). 307. m1 anterolingual cingulum: (0) absent; (1) present. 308. Buccal cingulum, lower molars: (0) absent; (1) present. 309. P2 parastyle: (0) absent or vestigial; (1) present-cuspidate (Morales et al., 2019: character 2). 310. P2 metastyle (posterior cusp): (0) absent or vestigial; (1) strong (Morales et al., 2019: character 3). 311. P3 metastyle (blade on P3): (0) absent or vestigial; (1) strong (Morales et al., 2019: character 8). 312. P3 mesostyle: (0) absent or weak; (1) strong (Morales et al., 2019: character 12). 313. P3 Paracone-Metastyle disposition: (0) aligned with the antero-posterior axis; (1) inclined as in the P4 (Morales et al., 2019: character 14). 190 314. P4 mesostyle: (0) absent or weak; (1) strong (Morales et al., 2019: character 23). 315. P4 alignment of protocone with paracone and metastylar blade: (0) absent; (1) present (modified from Morales et al., 2019: character 16). 316. M1 paracone-metacone shape: (0) coniform; (1) crestiform (Morales et al., 2019: character 26). 317. M1 protocone shape: (0) coniform with cristas; (1) crestiform with undifferentiated cristas (Morales et al., 2019: character 27). 318. p2 molarization: (0) metaconid absent; (1) metaconid present (Morales et al., 2019: character 38). 319. p3 molarization: (0) no metaconid; (1) with metaconid (Morales et al., 2019: character 40). 320. Masseteric fossa on maxilla below orbit: (0) absent; (1) present. 321. Auriform process of basioccipital: (0) absent; (1) present, large wing-like projection. 322. Frontal sinus, caudal extension: (0) terminates antero-dorsally to the brain cavity; (1) terminates dorsally above brain cavity, >50% of its length; (2) expands caudally over the complete length of the brain cavity. 191 323. dp4 metaconid alignment: (0) non-linear, triangular alignment with paraconid and protoconid, separated from talonid by transverse deep valley; (1) linear alignment with paraconid and protoconid; (2) no metaconid. 324. Petrobasilar foramen: (0) open vacuity; (1) closed foramen in basioccipital (Barrett, 2021: character 223). 325. Crenulations on the anterior face of the paracone of P4 and occlusal face of the protocone: (0) absent; (1) present (Barrett, 2021: character 225). References Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658. Barrett, P. Z. 2021. The largest hoplophonine and a complex new hypothesis of nimravid evolution. Scientific Reports 11:1–9. Barrett, P. Z., S. S. B. Hopkins, and S. A. Price. 2021. How many sabretooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae. Journal of Vertebrate Paleontology e1923523. Bryant, H. N. 1991. Phylogenetic relationships and systematics of the Nimravidae (Carnivora). Journal of Mammalogy 72:56–78. 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. Christiansen, P. 2008. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics 24:977–992. 192 Christiansen, P. 2013. Phylogeny of the sabertoothed felids (Carnivora: Felidae: Machairodontinae). Cladistics 29:543–559. 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. Gaubert, P., W. C. Wozencraft, P. Cordeiro-Estrela, and G. Veron. 2005. Mosaics of Convergences and Noise in Morphological Phylogenies: What’s in a Viverrid-Like Carnivoran? Systematic Biology 56:865–894. Heinrich, R. E., and K. D. Rose. 1997. Postcranial morphology and locomotor behaviour of two early Eocene miacoidcarnivorans, Vulpavus and Didymictis. Palaeontolgy 40:279–305. Matthew, W. D. 1909. The Carnivora and the Insectivora of the Bridger Basin, middle Eocene. Memoirs of the American Museum of Natural History 9:291–576. Morales, J., S. Mayda, A. Valenciano, D. DeMiguel, and T. Kaya. 2019. A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey. Journal of Systematic Palaeontology 17:1127–1138. Peigné, S. 2003. Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32:199–229. 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. Rothwell, T. P. 2003. Phylogenetic systematics of North American Pseudaelurus (Carnivora, Felidae). American Museum Novitates 3403. Sakamoto, M., and M. Ruta. 2012. Convergence and Divergence in the Evolution of Cat Skulls: Temporal and Spatial Patterns of Morphological Diversity. PLoS ONE 7:e39752. Salesa, M. J., M. Antón, J. Morales, and S. Peigné. 2012. Systematics and phylogeny of the small felines (Carnivora, Felidae) from the Late Miocene of Europe: a new species of Felinae from the Vallesian of Batallones (MN 10, Madrid, Spain). Journal of Systematic Palaeontology 10:87–102. Salesa, M. J., M. Antón, A. Turner, L. Alcalá, P. Montoya, and J. Morales. 2010. Systematic revision of the late Miocene sabre-toothed felid Paramachaerodus in Spain. Palaeontolgy 53:1369–1391. 193 Salles, L. O. 1992. Felid Phylogenetics: Extant Taxa and Skull Morphology (Felidae, Aeluroidea). American Museum Novitates 3047:67. Spaulding, M., and J. J. Flynn. 2012. Phylogeny of the Carnivoramorpha: The impact of postcranial characters. Journal of Systematic Palaeontology 10:653–677. Taylor, M. E. 1974. The functional anatomy of the forelimb of some African Viverridae (Carnivora). Journal of Morphology 143:307–336. Tseng, Z. J., and X. Wang. 2007. The first record of the late Miocene Hyaenictitherium hyaenoides Zdansky (Carnivora: Hyaenidae) in Inner Mongolia and an evaluation of the genus. Journal of Vertebrate Paleontology 27:699–708. Van Valkenburgh, B., F. Grady, and B. Kurtén. 1990. The Plio-Pleistocene cheetah-like cat Miracinonyx inexpectatus of North America. Journal of Vertebrate Paleontology 10:434– 454. Werdelin, L., and N. Solounias. 1991. The Hyaenidae : taxonomy, systematics and evolution. Fossils and Strata 30:104. 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. Yalden, D. W. 1970. The functional morphology of the carpal bones in carnivores. Acta Anatomica 77:481–500. 194 Appendix E Nexus formatted morphological character matrix #NEXUS [written Fri Oct 01 13:40:37 PDT 2021 by Mesquite version 3.61 (build 927) at LAPTOP- CRLI5NPQ/128.223.221.117] BEGIN TAXA; TITLE Taxa; DIMENSIONS NTAX=129; TAXLABELS Tungurictis_spocki Ictitherium_viverrinum Hyaenotherium_wongii Hyaenictitherium_hyaenoides Lycyaena_chaeretis Chasmaporthetes_lunensis Chasmaporthetes_ossifragus Palinhyaena_reperta Adcrocuta_eximia Pachycrocuta_brevirostris Crocuta_crocuta Crocuta_crocuta_spelaea Hyaena_hyaena Proteles_cristata Plioviverrops_orbignyi Protictitherium_crassum Belbus_beaumonti Tongxinictis_primordialis Allohyaena_kadici Dinocrocuta_gigantea Percrocuta_carnifex Percrocuta_algeriensis Percrocuta_tobieni Proailurus_lemanensis Pseudaelurus_validus Metailurus_major Nimravides_pedionomus Machairodus_catocopis Megantereon_cultridens Amphimachairodus_giganteus Amphimachairodus_coloradoensis Dinofelis_diastemata Homotherium_ischyrus Homotherium_serum Homotherium_latidens Promegantereon_ogygia Smilodon_fatalis Smilodon_populator Xenosmilus_hodsonae Neofelis_nebulosa Panthera_pardus Panthera_leo Panthera_leo_spelaea Panthera_atrox Caracal_caracal Lynx_canadensis Felis_lybica Leopardus_pardalis Acinonyx_jubatus Miracinonyx_trumani Puma_concolor Otocolobus_manul Prionailurus_planiceps Catopuma_temminckii Profelis_aurata Prionodon_linsang Arctictis_binturong Civettictis_civetta Paradoxurus_hermaphroditus Hemigalus_derbyanus Viverra_tangalunga Genetta_maculata Paguma_larvata Cynogale_bennettii Poiana_richardsonii Viverricula_indica Bdeogale_nigripes Mungos_mungo Suricata_suricata Herpestes_edwardsii Ichneumia_albicauda Crossarchus_obscurus Atilax_paludinosus Rhynchogale_melleri Galerella_sanguinea Helogale_parvula Fossa_fossana Cryptoprocta_ferox Eupleres_goudotii Salanoia_concolor Kichechia_zamanae Kanuites_lewisae Izmirictis_cani Euboictis_aliverensis Sivanasua_viverroides Nandinia_binotata Palaeoprionodon_lamandini Stenogale_julieni Herpestides_antiquus Ginsburgsmilus_napakensis Prosansanosmilus_eggeri Prosansanosmilus_peregrinus Afrosmilus_hispanicus Afrosmilus_turkanae Afrosmilus_africanus Albanosmilus_jourdani Maofelis_cantonensis Eofelis_edwardsii Dinailurictis_bonali 195 Quercylurus_major Dinaelurus_crassus Hoplophoneus_oharrai 'MA-PHQ 348' Hoplophoneus_dakotensis Hoplophoneus_sicarius Hoplophoneus_adelos Hoplophoneus_bidentatus Hoplophoneus_villebramarensis Barbourofelis_morrisi Oriensmilus_liupanensis Sansanosmilus_palmidens Barbourofelis_fricki Barbourofelis_loveorum Albanosmilus_whitfordi Hoplophoneus_cerebralis Hoplophoneus_primaevus Hoplophoneus_occidentalis Nimravus_brachyops Pogonodon_platycopis Pogonodon_davisi Dinictis_felina Nanosmilus_kurteni Nimravus_intermedius Tapocyon_robustus Procynodictis_vulpiceps Hesperocyon_gregarius Canis_latrans_frustror Vulpes_vulpes Taxidea_taxus ; END; BEGIN CHARACTERS; TITLE Character_Matrix; DIMENSIONS NCHAR=325; FORMAT DATATYPE = STANDARD RESPECTCASE GAP = - MISSING = ? SYMBOLS = " 0 1 2 3 4 5 6"; MATRIX Tungurictis_spocki 201020000?113210102??0??2?0?1?211020011?12011001???1?1?110???0000101????????????? ????????????????????????????????????????????????????????????1?01100110211010111011?1 111????????????????1100???010100010021000100031??0410000311000111???001?00??111?? ?0001?10000?0???0???00000??00122103??10010???001?2????????0?1{0 1}000000001??000?- 0 Ictitherium_viverrinum 2010200020110210?0???0??????12211??101101202100{0 1}?0?{0 1}21111????0000101???????????????????????????????????????????????????????????????????? ????????????????????????????????????????????????10000010102013000210001000311223100 2021{1 2}000{1 2}11001000?00??111101000101?0000000000000?101?10?122102220?111100000?1?????????? 1100000000100002?-0 Hyaenotherium_wongii 201020002?113210002??01??20?122110?1011012021000?0?0211110???0000101???1???????? ?02111?011102111011110011?0111?1101001?1???0???00001?00100011011111001021101001 196 0011????0????????????????1000002010201000021000{0 1}0004{0 1}1223100202110002100?0000000??111100000101001000000000000101?10?1121022210111 11000??1???0??1?0?1100000000100002?-0 Hyaenictitherium_hyaenoides 200020002?103210?????0??????12211??1011012021001?0?121111????0000101????????????? ??????????????????????????????????????????????????????????????????????????????????????? ????????????????1000002010200000021000100141122310020211001211???0?0?00??11100{0 1}00010100000?00000000?101?1011221022201111210000?1??????????1100000000100002?-0 Lycyaena_chaeretis 200020?02?113210?1???0??????12211??0011?12021001?0?021111????000{0 1}1?1?????????????????????????????????????????????????????????????????????????????????? ??????????????????????????????????100000101020000003{1 2}00010?3{5 6}112{2 3}410010111001110??0?00000??11100100010100100000000000?401??0?022102220?1212100 00?21?????????1100000000100002?-0 Chasmaporthetes_lunensis 201020102?113210?2????1?2???12211??0011?{0 1}20{1 2}10{0 1}{0 1}?0?021111????{0 1}0011?1??????????????????????????????????????????????????????????????????????????????? ?????????????????????????????????????10?0?020003000000{3 4}2011{0 1}0?3601234000202{0 1}2001{1 2}1{0 1}??0?01000??11{1 2}00{0 1}0001010?100000100000?401?100022101220023121000??1{0 1}?????????110000?0001??0022-0 Chasmaporthetes_ossifragus ??10????????????????????????????????????????110??????????????000?1?1???1?????????????? ?0????????00?0???????????????0?1????0??01???10??0?00??0?11??????????????????????????? ????????????????????00030000204201100?36?121??001?2120011?????????????111000000101 0???000??00?00???1???????10122????22?00??11??10??????1?0000000??001???-0 Palinhyaena_reperta 30102?002?113210?2???0??????12211??0011?12021001?11121111????01001?1????????????? ??????0????????01?????????????????????????????????????????????????????????????????????? ????????????????100000221020000003100010?151123410010111011111001101010??2220010 0010100100000000000?401?1111122012201121210000?21?????????1100000000100001?-0 Adcrocuta_eximia 300022002?100210?2???0??2???12211??0011?12021100?1?021111????00011?1??11????11?0? 0101100111021011110?0???1???1??1???????1011?110?0???????????1?0110001121101001?1? ?11?10????0???????????10000020203?000003200{0 1}00?36012341001021200111000110?110{1 2}?32{1 2}000000101001000000000000401???1122201220?{2 3}3121000011?0??????0?11000000001000022-0 197 Pachycrocuta_brevirostris 300021{0 1}02??13??????????????????????0011?12021000?1???1111????00011?1????????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ?????????000?0202030010003200010?360122410010211100121???1?110{0 1}??3220010001010?100?00000000?400???10??2112?00??0?1000??11?????????11000000001 00??20-0 Crocuta_crocuta 301021002?113210022?00122?001221103?????????10010{0 1}11211110?1?00011?10111???01110102011?011110101011110011101111110100111001100 110001?00100011111110001121101001111111110111001110010101010001020103001020420 1100?36010141001021211002000011110021222000000?01001011000001100401?1010221032 201131210101120010000100000000000??000022-0 Crocuta_crocuta_spelaea 3000220020013210122?00121?001221103?????2?0?10010101211110???00011?1?1?1???0011? ?0211100111101010110100111011111111011?1???0??????00?01100011111110010121101001 1111?????111?010100111010100000201030010104201100?36?1014100102121100100001111 00?132200100010100100000?00110?400?100022102?201230210101?21210??1?0?110000000? ?00002{0 2}-0 Hyaena_hyaena 300021002?113210022?0012220012211030011?020110000{0 1}1121111011000011?1000111101110102011?0111101010110001?1101111110100111000100 110000?00100011111110000121101001111110110111001111010101{0 1}100010201020010103100100?360101410010211110021011110110202210010001010011110 00000000401?0{0 1}1020112220112021010112011000010?110000000?1000020-0 Proteles_cristata 211022012?113210122?00121200122110202???2?0011200001211110???1001??10001???0111 0002011?0110101010111101?11011111101001?11?01?0110000?001000111111100001211010 010111111101110011110111000200011?12002??1{1 2}25211100?36002?2102?0?0?0??30100100111020??200001000?001010000101?0040??00101 210??001130110?0?11{0 1}010000?0?00??00000??000001-? Plioviverrops_orbignyi 2?1022{0 1}120113210?0???01?1???12211??0011?1?011001?00121011????0000101??????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????110000?{0 2}101013101200000000111013000003110?00210??00?010??12100100010100000000000000? 400??0002210311001100?010??10?????????0100000000100000?-0 Protictitherium_crassum 20002?002?11300000???012?200122110?0010112011001?0?0210110???0000101??01???0110 0012011000110200?011010??11011110101011????????011?00?00100010?101?10?012110110 1?1111111001110111101???10110010?010101310001000100031101300020312001111???100 198 1100111110000010100001200000000?000???0002012120011111010110?0?00???0?110000000 01000000-0 Belbus_beaumonti ????????????????????????????????????????????????????????????????01?1??????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ???????????????0??????00??03100000??5?101????2??1??11??????1???????11?????0??0?????1 000????00??????????????22????121?0??????????????11?????????00????-? Tongxinictis_primordialis ???022112001?210?0??0?122?0?1?211?3?0???????1001???021011????0001?????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????????1100???0?02?010??????????3??1?3310010312001021????00100??2210010001? 10000??0?100000??00??0??22102??0012????00???????????????0000000??00100?-0 Allohyaena_kadici ????????????????????????????????????????????100??????????????10011?1?????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????????????2102000??03101100?35???3??001?212?00??????????????221??110???1??? ???00???000???1?????????22?????0???0??????????????110000000??00???0-0 Dinocrocuta_gigantea 200022102?100110?02?00122?0011211040210??2011100?0?0211110???01011?1???????????? ??????????????????????????????????????????????????????????????????????????????????????? ?????????????????000010002032000103200000?360123410010112101010????00001??222001 0001010?101000100000?401??011321022200221211001?1??????????11000000001001121-0 Percrocuta_carnifex ?????????????????????????????????????1??220?110????????????????011?1?????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????????????{0 2}0030000104200000?36???3??001?112?0???????????????222??100?1?1??????00??0??0???1? ???????0?2?????1???0??????????????11??00000??00???1-0 Percrocuta_algeriensis ???0???????????????????????????????0010?22011101?????????????00011?1????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ???????????????????02030010103200000?36???1??001?212101??????????????323?01000101? ???1000?00?10???1???????102??????0???1??????????????1100000000100???1-0 Percrocuta_tobieni ???????????????????????????????????001??220111????????????????0011?1?????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????????????02?3001?103201000?36?1?1??000?2?2?10??????????????22???100??01??? ?1000????00???0????????0??2????021?0???????????????10000000??00???1-0 199 Proailurus_lemanensis ?00?1??01001311000210012221012211030011?1201101??1??300??1???00001?1????????????? ??????????????????????????????????????????????????????????????????????????????????????? ????????????????10?10?202011000004101100?35012200002020200011????001200??1110000 0000111010200?00000?2???1??12?200121?01120000??2??????????00000000000000001-0 Pseudaelurus_validus 201010002?100110102??0122201122110?0011?12011011?000310110???00011?1??01???0??0? ?110111110102000001011011100010110100111111?01211111?001{0 1}0010?10110001001101011111000?100???????????????101000?02011000103201100?360122 000{0 1}201020011102000011001011110000000101011000000?001000?0101202{0 1}3220?02120?0{0 1}1010?000??00?0000000000000000?-0 Metailurus_major 201010001?10121012???01?0???122110?0211?22011011?00031011????10011?1??01???001?1 ?10?110101?02?????1??0????0??1??1?10?????????1????10??010001???01?0??1??????0?1?1?? ???100?110??0001???10100000202031000104201100?360123000220102101210200001100?11 11101010101001101101000000001???0010103120011111001111?0?0????0?00??0000000?000 0?-0 Nimravides_pedionomus ?00?1?????1?????1?2??012?20?1????0?0211?22011011??????0110???10011?1???1?????????11 01111011020010010100111010101111001????????????00?00100110110110001011101011111 0???????????10001?????1??0?0200032000104201100?360123??012?10{1 2}00121????0?10????11110101?10101?1011010100010?0???????203{1 2}2????1{1 2}100???2??000???0?000000000???00???-0 Machairodus_catocopis ?00012001?121210?22??012?20?12211???????????10?1?0?131011????10011?1?001???001000 110111101102001001010011101010111100111111101011101?00100010110110001011101001 111000110011101100010101010001020??3200010{3 4}201100?360123000220102001200???0111001111?20101010101110010101?011000???01201 1?22002?121001?11?00000000?0{0 1}??00000???0000?-0 Megantereon_cultridens 201010010012121012???012??0?10211???????????1111?0?131011????10011?1???1???010?00 110011001112??1011011011?01110110100101111101211100?0000001?1101?001112110?11? 01100??100111011010101110100010200032000105201100?360121010210111101210???00?1 00???112000100?1001{1 2}12301002000001??011201{0 1}32201231210031?1??????????01??00000??00000?-0 Amphimachairodus_giganteus 200012001?121210?????012????12211??0211?220111{0 1}1?00031011????10011?1???1?????????????????????????????????????????????????????????? 00??0??0010??0????????????01?1110?????????????????????100010201032000104201100?360 200 123010110111101200200001210??110201010{0 1}0100111020201201?000??001200232210221{0 1}10121?11?????????00??00000???0000?-0 Amphimachairodus_coloradoensis ?0?????01??????????????????????????0211?2201111??????????????10011?1???1????????????? ???????????????100111???101111001?1??10?1111110?001001101101100010211010010110?? ??0?????????????????????0201030000105201100?360123??01??10310?2??????????????11201 010101???110202012011??1???????023121???12100??????000?000?010000000????0?0?-0 Dinofelis_diastemata 201011001?101210?1???01?????12211??0211?11011111?0?131011????10011?1??01???0???0? 1111110011021010111100111001100101011?11?11?1011111??0?10110?011?0?000211010010 110?0110??????????01???0100000200032000105101100?360121000210102001110200?01100 1?1111010101010?1110101012000000??000301032200{1 2}3121001?1{1 2}1??0??0?0?00??00000???0000?-0 Homotherium_ischyrus 101111011?121210112??012??0110211???211?22011101?0?0310110???10011?1??11???01000 0110100001102101011010??11011101111001?1??11?1???101?0?10?1?1111110010021101000 {0 1}11000?10000?0?111?10101?100010201032000205201100?3601230102001?3001200???01?2 001121120001010100101??02013011011?1001200022?1033???0??111??100000?001??00000? ??0000?-0 Homotherium_serum 100011001?121210?2???0?2000010211???????????1101?0?0310110???10011?1??11???001?00 1101110011021010010100111??111111100??1{0 1}?11?111?010?00?0011?1011?0?????110??00????10?101?1?0111111010?010001021?032010 205201100?360122000210112001300000?1?2001?31220?01010200101{0 2}2{0 1}20??011201??001000032{1 2}1033121002110000?0???0000?????00???0000?-0 Homotherium_latidens 2{0 1}10{1 2}{1 2}001?121210?2???0??????10211??0211?22011101?00131011????10011?1?????????????????? ??????????????????????????????????????????????????????????????????????????????????????? ???????????100010201032000205201100?360123010210112101300200010100??31120101010 {1 3}00111{0 2}2{0 1}201321?011?1001{1 2}000{2 3}?{1 2}10{0 3}31{1 2}10021?{0 1}0?????????01??00000???0000?-0 Promegantereon_ogygia 200011101?101210?2???0??0?00122110?0011?22011011?0?03?011????1001101??01???01000 001011?001?02??1?01?110111010101101011?11?11?121?100?00?001101101100{0 1}{0 1}0{0 2}1101001111001110011?0110001?1010100000202032000104101100?3601020002201020011 10??0?01100111111000{0 1}0101101110101002000201?000010{1 2}13220011111010111?00000001?00??000000?00000?-0 201 Smilodon_fatalis 2000{0 1}0{0 1}10?131210122?021?020010211030211?220111{0 1}100{0 1}0310110???10011?10111???0010011111101011020010110100111001101111001111111010 11101?001000110101100001{0 1}01011010110101100011011010110010100010202032000105{1 2}01100?3601220102?011200?2101{0 1}001110111?11200010101001212212012{2 3}{0 1}1211000010222322002{3 4}1{1 2}10?3112{0 1}00000000000??0000000?00001-0 Smilodon_populator 2?0??1?10??31210?????0??????11211??021??2201111??00031011????10011?1??11?????????? 10?1???1??2??1??1?100111001101111011???????1?1?111?00?1000??1011???01011011010110 ?????021?01????01?????00010202032000105201100?36010201020011210?210??00??101??31 1201010101001213212112{2 3}01?11??011??1222201241210131?20?????????00??0000000?0000?-0 Xenosmilus_hodsonae 2000{1 2}0111?12121012???0?20???1{2 3}211???2???2?0?11?1?0?031011????10011?1?111???001001110110001102101011011011110 1101110000?1??11?11?1?10?0010010011011000012110110011100??10??????????00?0??1000 1020??31010?05201100?36?1210002?000200?300????1?11011?1320?010?1100111220201?31 1211??000300012210331210?2??1?000000?0?00??0000000??000?-0 Neofelis_nebulosa 201010101?101210122?00120?0011211030211?220111110101310110???10011?10101???0110 001111111010021010110111?1100010110100110111101211101?011000111101100110011010 110110011100111011000111110100000202032000204101100?36012300012010200102000000 111011111000010101100110000000000201000012120322011211011001210000?00000000000 00??00000?-0 Panthera_pardus 201011002?101210122?00120?00122110?0211?220110110001310110???10011?10101???0110 001{1 2}011100100210100111101110111011{0 1}1001?1??11?1110100?001100111101101101211011011110101100011011000111010100000 20203201010{4 5}201100?360123000010101000210000001000111110010101010001100010000002011100021 202220112110101011110000000000000000000?00001-0 Panthera_leo 200010102?101210122?00120?0011211030211?22011111000031011011?10011?10001?1?011 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??????????????????????????????????????????????????????????????????????????????????????? ?????????????????10210??2?01?000103101100?35?1?110001020100002?????0?100??113??00 000?1??0?0?00?0?000?0?0?11?002?011?1???0?1?00??1??????????0000000000100000?-0 Herpestides_antiquus 201020002011311000200112121012211022111012011?0??001210110???0100101??0???????? ??????????????????????????????????????????????????????????????????????????????????????? ??????????????????111000?0001000100210000001311?1100020311001121????00200??11100 00000010001000??00000?000??0?102103110???011?01??1??????????11000000001000000-0 Ginsburgsmilus_napakensis ?--?????????- ??????????????????????0011?1202101??????????????1001101?????????????-????-???????- ??????????-?????????????????1????1???????????-???????????????- ?????????????????????????????????20-0010000?????????36- 209 ??3??102?2010000?????????????11111000?0?1????1?11?02000???0????????0-??-???0-?0--???- ?????0????-0000000?00?????-0 Prosansanosmilus_eggeri ?--?????????- 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12251121001210010?????11?01??211211111{0 1}121?11233120?221???1??0011200-22-1341- 10--??1-?????????0-??00000???0000?-0 210 Maofelis_cantonensis 1--12??01??0- 010?0?????10????01????0011?12021001?0?0?????????000?????????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????1??10?22-002??01?????????3?- ????000?11?10??020????0?100????20000?00??0?00????1010???0?0?1?010?10-??-001?-?0-- ??2-??????????-??0000000??000?00 Eofelis_edwardsii ?--?????????- ???????????????????????011??202100??????????????{0 1}00{0 1}1?1?????????????-????- ???????-??????????-??????????????????????????????????-???????????????- ?????????????????????????????????22-00?010103201100?3{5 6}- 121??001?1010001?????????????11110000?001????101010200????0???????10-12-???1-10-- ???-?????????0-0000000?000?????1 Dinailurictis_bonali ?--12???????- ??????????????????????0011?22021001??????????????00?1?1?????????????-01??-???????- 0??1??????-?????????????????????????1????????-???????????????- ?????????????????????????????????22-001010103201100?36- 121??0?1?1010001??2?0????????11110000?0?1????1?10?02?0??0?0????????0-1?-???0-?0-- ???-?????????0-??00000?0?00?0??1 Quercylurus_major ?--?2???????- ???????????????????????????2?0?100??????????????100?????????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ??????????????????????????????????2-001??01?????????3?- ?????00??1?10??20??????????????1??00??0?????????????0?0?0?0????????0-??-????-??--???- ??????????-0000000?????????1 Dinaelurus_crassus ?--122101000- 010?0??????????10211??0211?22021001????01000????100?????????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????10?1??20-002??11?????????3?- ???5002?11?10??020????00?00????10000?00??1111????0000?0?000???012210-??-013?-?0-- ??2-??????????-??00000????000?10 Hoplophoneus_oharrai 1--120000002- 01010???0010?0110211????11?22021101?00001000????10011?1??00???0?1?00-0100- 0001020-0101110000-11010011?0?1?0???1????1?1100100011-110????????????- ???????????010?10111???????10?100?0-0??010004101100?36- 12050?01110?00?110?00?00100?1?1?20?0100010011123021??2??100??0?11200-?1-?231-10-- ??1-??0??1???0-0000000?0?0000?00 'MA-PHQ 348' ?--1??01?000- 010?00???0201011021110?????????10??????01?00????00??????????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????10?1??22-?????1????????????- 211 ????0?0?11??0??0??????0??00????200???00????0??????0??????00?????2?1?-??-?12?-?0--??2- ??????????-????????????00?0? Hoplophoneus_dakotensis 2--1?0010?12- ????????0??????10211??0211?2202110??0?0?1000????10011?1?????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ????????????????????????????0?1?020-002010104201100?36- 12150?2?110?00?200???0??200???11200?100110012223021?231?101?1001??00-11-0??1-?0-- ??1-?????????0-??00000????0?0?00 Hoplophoneus_sicarius 2--122010?12- 11011???00?0??110211??0211?2202110??00001000????10011?1?????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????10?10020-002010104201100?36- 1215012?110100?110200011111???103000100110012234021?231?101?10011?00-11-0131-10- -??1-?????????0-??00000????000?00 Hoplophoneus_adelos 2--122010012- 10010???00?0?01?0211??0210?22021100?00001000????100?1?1???????????0?-0?0?-?00?02?- ???0?1?010-???0?0011?1???????????????????????-???????????????- ?????????????????10?01100??10?1??20-002??01?????????36-?2?501{1 2}?11?10??110??0011201?1?01?000?00?10012????2132?1?111?10110?00-??-0231-?0--??1- ??????????-00000000???000?00 Hoplophoneus_bidentatus 2--122010?12- 210112?100202011021101?21??1?0?1100?001010000???10011?1???0???000000-0?01- 000??2?-????1?0???-0???000??00?????????21?11???0?0011-???????????????- 0??????????0?0?????0?????1010?10020-00?010004{1 2}01100?36-1215012?10{0 1}200?110200?10?0111?{0 1}1{1 2}00010011???22340203331?101?10?12210-12-?230-10-- ?02-????????00-??00000????000?00 Hoplophoneus_villebramarensis 2--12?010?12- 000?0??1001020110211?10211?22021101?0??01000????10011?1?????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????10?1??20-002010004{1 2}01100?36- 1225012?100200?110????10201???102000100?1???213?0?1?33??1?1????11200-?2-?230-?0-- ???-?????????0-??00000????000?00 Barbourofelis_morrisi 3--120010?12-{0 2}00002??1020?0010211??0211?220?1111?1?0110??0???10011?1???0?????????-0111- 0011021-1011010011-101101111001??1????1????11000?0001-10?????????????- 1?0110??????00?1???????????10001020-002000105201100?36-1225112?001{2 3}10?21000001?101??31121111111310122331202321?001?10012200-22-1241-?0--1?1- ?00??????0-??00000???00?0?-0 212 Oriensmilus_liupanensis 3--120000?12-00010???0020001102110?0211?2202111{0 1}?0?111000????10011?1?????????????-????-???????-??????????- ??????????????????????????????????-???????????????-???????????????????????????10?0?020- 002000105201100?36-123501210{0 1}12101210???00?100??111211110{0 1}0100101221202?21?00{0 1}?10012200-22-0??0-10--???-?????????0-??00000????000?-0 Sansanosmilus_palmidens 2--120010012- 210102?00020001102110?0211?22021111?1?0110??????10011?1???0???0?????-0111- 001102?-1011?11011-1111001110?1????1??1?11110100?00?1-1101?0000011101- 11?110???10?00?1????0?????010001020-002000105201100?36- ??3511210112101210????001011?11121111000110121?3120222?0000?10012200-2?-0240-?0-- 1?2-?0?????0?0-??00000???0000?-0 Barbourofelis_fricki 3--122013012- 210012??0020001102110??2???2???1101????110??0???10011?1???0?????????-0111-0011011- 1011010010-1011?1111001?1??11??????1100110000-100????????????- 0?0110?????100?11111???????10001021-002000205?0110??36- 12351121001?101200???011101??31331111111310122341202?2??011?10011200-?2-1240-10- -1?1-?00??0???0-??00000???0001?-0 Barbourofelis_loveorum 3--122013?12- 200102??0020001102110??2???1?0?1100?1?0110??????10011?1?0?0???001000-0111-0011{0 1}11-10110100{0 1}1-101101111001?1111101211111000000?1-1101100?0011101- 11011000?100???1??1????????1000?021-002000205201100?36- 12351120001310120?????1??011?111?111111131?11233120232100?1??0?12200-22-1241-10-- ?1?-000??0?0?0-??00000???0000?-0 Albanosmilus_whitfordi 3--12?013?12- 200?12??002??0?102110?0211?220?1111?0??110??????10011?1???0?????????-0111-0011011- 10110?????-????????1?0?????????????1100?00000-???????????????- 1?0110?????????????????????10000020-002000105201100?36- 123511220012101210???011100??2112111{0 1}?1121012{1 2}231202321?0?1?1?012200-22- 1241-10--??1-?????????0-??00000???0000?-0 Hoplophoneus_cerebralis 2--1220100{0 1}2- 210112?0001020110211030211?22021100?0?1010000???10011?1???0???0??00?-0101- 0001011-1101111??0-1??????11001????????????110000000?-?1010?00?00110?- 1?1110??????00?1?100011101000?10020-00201010{4 5}201100?36- 1215012?100110?110200{0 1}10201???11201010012001223402033310111?10111210-11- ?1?1-10--??2-??0??????0-??000000???000?00 Hoplophoneus_primaevus 2--121010002- 010102??0010201102110?0211?22021101?0?0010000???10011?1?100???000000-0100- 0001020-1101111000-110100111011?00?111121111100000010-100100001021101- 111110001100101111{0 1}1?11101000?1?020-00201010{3 4}101100?36-121501{0 213 1}111020002102000111001111110{0 1}010011001112{2 3}02022210100?10011200-12-0231- 10--?01-100000?000-00000000??0000?00 Hoplophoneus_occidentalis 2--120010012- 010112?1001020110211000211?220211?1?000010000???10011?1??00???0??0??-0100- 0001020-1101111000-11010111100110011?1121111100000011-110100000021101- 1111100011001011111101010??10?1?020-???01010{3 4}101100?36- 122501211102000200000?1110011111200?10011001112302022210100?1?01020?-11-0230- 10--??1-?00?0?000{0 1}-??000000??0000?00 Nimravus_brachyops 2--122101000-{0 1}0010210{0 1}02000110211000211?22021001?000010000???{0 1}0011?1??00?????????-0101-0011021- 1101011010-111100111001?11?110121111100010001-110110000011101- 01011000110010111100011101?10?11020-00201010{3 4}201100?35-12350{0 1}0111020001102?00001001?111100000001110110110010101000011012210-12-0120-10-- ?12-?00??0?0?0-00000000?00000110 Pogonodon_platycopis 2--12?101000- 000102??002010110211??0011?12021001???101000????10011?1???0?????????-????-???????- ??????????-????????????????????????1100010001-???????????????- ???????????????????????????00?11020-002010003201100?36- 121500011101001200??0000100??12110000000100101120211001?000?1??10?10-11-?220-10- -??1-?????????0-00000000000000?10 Pogonodon_davisi 2--122101000- 00000???002010110211?00011?12021000?0?1010000???10011?1???0???0??0??-0101- ?001020-0101111??0-1100?0111?0?????????????111???00??-???110000010100- 110110?01?0010?11????1?????10?1?020-002010003201100?36-121500{0 1 2}11101000200???001100??121200000001001011{2 3}01020011100?1?010?10-1{1 2}-0220- 10--??1-????0??0?0-00000000000000?10 Dinictis_felina 2--122101000-0{0 1}0102?0002010110211000011?12021001?001010000???10011?1?100???000000-0101- 0001020-0101111000-110100111011?10?110121111100010001-100100000010101- 110110001?0?10?111{0 1}0011101000?1102{0 2}-002010003101100?3{4 5}-12{1 2}50001110100011020000110011{0 1}1{0 1}10000000100{0 1}100201020010000?10012210- 12-0{1 2}21-10--10{2 3}-100?00?000-00000000000000?10 Nanosmilus_kurteni 2--122?00002- 000102?1001020110211000211?22021101?0?0010000???10011?1?????????????-????-???????- ??????????-??????????????????????????????????-???????????????- ???????????????????????????00?1?020-002010104201100?36- 121501111102000210????01?00??111?00010011???1123020?221?101??0?112?0-12-?121-1?-- ??1-?????????0-00000000??0000?00 214 Nimravus_intermedius 2--1?{1 2}?01000-0001?2??0020001102110?0211?{1 2}2021001?0??010000???00001?1?????????????-{0 1}101-?01102{0 1}-1??10?????- ????????1?1?????1??1??1???????????-???????????????- 0?0110?????????????????????10?11?20-002010104201100?35- ??3500011101000110????00100??1111000000011101101100101010?0?11012210-12-?1?0-10-- ??1-?????????0-00000000?00000110 Tapocyon_robustus 10101?0000010010112010010100000010?201101102000????0000??0???0000100??0????0100? ?1100010000020010100010111100101?11111??0?1??1?1?11?0001001100000?0??1?100111?1 ??1?0??10????????????????1??1?0?2?0000000021010000131103?000201010001100?000??00? ?11110000000111001000000000?200????1??201110100010000??1??????????01000000101000 00?-0 Procynodictis_vulpiceps 1?1??0000001111011??11000100101000010111110100{0 1}??00?000??0???00001?0???????0110?01111111010020010?001101110101?1111111???????? ?0010101010001000011010112101110111?001110?1????110010???????????0200003?0020000 00??10?20??002?1010?10?????????????111000000??1????0?0??00000???0????????021??????? ?00??????????????0000000000100?0??-0 Hesperocyon_gregarius 10111?110001211000201121030011200112011111201010?001010010???0000100??00???1?1? ?012011?0000020000000110100010101111011????????000000?1010101011011111101101011 1111??111?0101111???0000000000?00020000000021000000030121?00020101011110????011 00??1111000000010?000000000200?200????112112120011011001??1??????????00000000001 000000-0 Canis_latrans_frustror 1001111000010111012?2{0 1}2103001120111101111120110010010100100010000110100100001110102011?01110010101 10{0 1}0011101111110101111000000100000?10111011110111101120101001011110?11101001000 01000011000102000000000021000000030120?000201010?11001010000100111010000000101 0000000002000200?1001200021100110010001110010000?0000000000000000000-0 Vulpes_vulpes 2001110000010111012?2021030011201111011110201000100101001000100001101001000011 10102011?011100101011000011101111110101111100000100001?10111011110111101121101 001011110?1110100100001001011000102020000000021000100030120?000201010?11002010 0001001010{0 1}00000001110000000000000200?{0 1}00120103110001001000112001100000000000000000000000-0 Taxidea_taxus 2000111000000001022?0221030011201111111110211001100121100000110111?11001001001 110001110100102000011010000110001011101111010000200001?00110011110100001120001 111111111?10010100?????10110200010200010001002000011?340020?100001010?121020100 215 100001112000000001000102001000000301??0102210021012102000000101?000??000000000 ?000000000-0 ; END; 216 Appendix F Custom Evolutionary Models (Rate Matrices) Beast 2.6.3 does not include a method to order characters from the BEAUti graphic user interface, thus only non-ordered characters are available for an analysis that does not manually edit the xml file. Below are schematic depictions of the rate matrices used in this analysis, with diagonal elements removed. The xml versions of these models are visible in the provided xml code. 3-state Characters Ordered Feliformia_trim_2_3_ordc_ratematrix Characters: 5, 6, 14, 18, 19, 22, 24, 31, 37, 41, 42, 83, 185, 196, 200, 203, 219, 220, 229, 257, 258, 262, 279, 280, 282, 283, 288, 289, 322, 323 Ordered, multipath Feliformia_trim_2_3m_ordc_ratematrix Characters: 47, 127, 192, 193, 231 217 4-state Characters Ordered Feliformia_trim_2_4c_ord_ratematrix Characters: 1, 195, 210, 215, 222, 224, 228, 242, 243, 244, 245, 253, 264, 277, 281, 286, 293, 296 Ordered, multipath 1, irreversible Feliformia_trim_2_4mi_1ord_ratematrix Characters: 266 Ordered, multipath 2 Feliformia_trim_2_4m_2ord_ratematrix Characters: 198 218 Ordered, multipath 3 Feliformia_trim_2_4m_3ord_ratematrix Characters: 12, 259, 265 Ordered, multipath 4 Feliformia_trim_2_4m_4ordered_ratematrix Characters: 53 5-state Characters Ordered Feliformia_trim_2_5c_ord_ratematrix 219 Characters: 260, 287 Ordered, multipath Feliformia_trim_2_5m_ordered_ratematrix Characters: 35 Ordered, multipath 2 Feliformia_trim_2_5m_2ordered_ratematrix Characters: 269 6-state Characters Ordered 220 Feliformia_trim_2_6_ordered_ratematrix Characters: 202 Ordered, multipath Feliformia_trim_2_6m_ordered_ratematrix Characters: 216 221 7-state Characters Ordered Feliformia_trim_2_7_ordered_ratematrix Characters: 211 222 Appendix G Stratigraphic Ranges and Tip Date Priors Extant Taxa Due to the preponderance of molecular data, compared to morphological data, for extant taxa in this analysis, tip dates were fixed to zero for these taxa (n=45). All remaining taxa were given uniform distributions of equal probability between their FAD and LAD, unless otherwise stated. Caniformia Hesperocyon gregarious. Locality: Numerous across North America, see Wang (1994). Absolute age estimate: 41.4-29.75 Ma. A relatively ubiquitous taxon of the Chadronian through Whitneyan, the earliest records extend into possibly the earliest Duchesnean (Wang, 1994). These earliest specimens come from the Lac Pelletier Lower Fauna of Saskatchewan, but precise age constraints for this fauna is still debated (Bryant, 1992). Thus, the age range chosen for this taxon was the entirety of the Duchesnean through Whitneyan (as defined by Kelly et al., 2012; Ogg et al., 2016) pending more precise dating. Procynodictis vulpiceps. Locality: Wyoming, USA. Absolute age estimate: 45.15-41.4 Ma. The stratigraphic range of this taxon appears to encompass the entire Uintan (Robinson et al., 2004), though the latest occurrence is in some doubt (Spaulding et al., 2010; Solé et al., 2016). For this study the entire Uintan NALMA was used (as defined by Kelly et al., 2012; Ogg et al., 2016). Feliformia 223 Tapocyon robustus. Locality: California, Montana, Wyoming, Utah, USA. Stratigraphy: All Uintan. Absolute age estimate: 46.5 - 39.7 Ma (Robinson et al., 2004). Tapocyon has typically been placed as the immediate sister taxon to crown Carnivora based on parsimony analysis of morphological data (Wesley-Hunt and Flynn, 2005). However, Barrett et al (2021) recovered it as a stem feliform, as did Tomiya and Tseng (2016). Feloidea Stem Feloids Proailurus lemanensis. Localites: Quercy Fissures, Laugnac, Paulhiac, Bassin de St-Gérand, France. Absolute age estimate: 27.82-20.44 Ma (Slater and Friscia, 2019). Stenogale julieni Locality: Quercy Fissures, France. Absolute age estimate: 27.82 - 23.03 Ma (Slater and Friscia, 2019). Palaeoprionodon lamandini Locality: Quercy Fissures, France. Absolute age estimate: 27.82- 23.03 Ma (Slater and Friscia, 2019). Herpestides antiquus Locality: France, Germany. Absolute age estimate: 23.03-20.44 Ma. Following the revision of Wolsan and Morlo (1997), this taxon is known from the Aquitanian/Agenian (Hunt, 1991). The absolute dates of the former were taken from Ogg et al. (2016). 224 Felidae Pseudaelurus validus Localities: Various in New Mexico, Nebraska, USA (see Rothwell, 2001, 2003) Absolute age estimate: 17.0-14.5 Ma (Slater and Friscia, 2019). Promegantereon ogygia Localities: Eppelsheim, Germany; Batallones-1 and Crevillente-2, Spain. Absolute age estimate: 11.1-7.75 Ma (Slater and Friscia, 2019). Metailurus major Locality: Bulgaria, Greece, China. Absolute age estimate: 7.6-5.7 Ma. The referred AMNH material is from the Mahui Formation, 6.7-5.8 Ma (Qiu et al., 2013). The oldest dated specimen of this taxon is from the MN11/12 boundary of Bulgaria (Kovatchev, 2001). Roussiakis (2001) lists further European occurrences from MN12-13. However, the taxonomic revision of Metailurus by Spassov and Geraads (2015) casts doubt on the diagnosis of many of these specimens. Confidently referred material comes from Pikermi, Greece (7.37-7.11 Ma; Roussiakis, 2001; Böhme et al., 2017), the type skull from Loc. 30, Tay-Chia-Kou (5.7 Ma; Kaakinen et al., 2013), and intermediate aged dates from Greece and China (Spassov and Geraads, 2015). Thus, the absolute date of this taxon follows the material that was examined for this study and referable to Metailurus major by Spassov and Geraads (2015). Absolute age dates given for MN zones follow Hilgen et al. (2012). Dinofelis diastemata Locality: France, China. Absolute age estimate: 5.3-1.0 Ma. Referred specimens F:AM 50445, 50446 differ in osteometrics from published values of Dinofelis diastemata, namely in smaller values of absolute C1 and P3 size (length, width) (Werdelin and Lewis, 2001). Furthermore, they are substantially smaller than that of Asian D. cristata, though a 225 geographically more-likely taxon, than the French D. diastemata (Werdelin and Lewis, 2001). Another possibility is the same species to which the Georgian Melaani skull belongs (Gabunia and Vekua, 1998; Werdelin and Lewis, 2001). The Melaani skull also differs from the above named species by having smaller osteometrics, but more compressed canines, and is stated by Werdelin and Lewis (2001) to be most-similar in dimensions to South African D. barlowi. However, barring a more in-depth taxonomic review of additional Eurasian Dinofelis specimens, I chose to diagnose F:AM 50445, 50446 to D. diastemata based upon similar length of the P4, unreduced P3 and shape of the sagittal crest. Dinofelis diastemata is attributed to MN 14 in Europe (Werdelin and Lewis, 2001). The referred Chinese material has an attributed date of Ruscinian-Villafranchian, thus overlapping absolute age dates are given above based on Rook and Martínez-Navarro (2010) and Hilgen et al. (2012). Nimravides pedionomus Locality: Nebraska, USA. Absolute age estimate: 13.55-9.95 Ma. Referred material to this taxon has been collected from the Nenzel quarry, Valentine Formation, Crookston Bridge Member; Xmas quarry, channel fills within the Ash Hollow Formation, Cap Rock Member and UCMP locality V-3327 (Antón et al., 2013). The Crookston Bridge Member lies above the Hurlbut Ash, dated at 13.55 ±0.09 Ma (Tedford et al., 2004), while the Xmas quarry was viewed as stratigraphically correlative with the Machairodus quarry by Skinner and Johnson (1984), which is capped by a dated ash at 9.95 ±0.8 Ma. UCMP locality V-3327 is where the holotype of Nimravides pedionomus comes, also Cap Rock Member of the Ash Hollow Formation, and is thought to be intermediate in age of the previously listed sites (Tedford et al., 2004). 226 Machairodus catocopis Locality: Kansas, USA. Absolute age estimate: 7.5-6.6 Ma. This taxon is well known from the Jack Swayze Quarry and Sebastian Ranch fossil localities (Antón et al., 2013). Both of these localities are viewed to be Hh2 in age (Martin, 1998). Absolute dates for this Hemphillian subunit follows Hilgen et al. (2012). Amphimachairodus coloradoensis Locality: USA. Absolute age estimate: 6.6-4.9 Ma. This taxon is known from various late Hemphillian sites in the American west (Antón et al., 2013), while material assigned to this taxon from Arroyo Tepalcates, Mexico has since been removed to a new species (Ruiz-Ramoni et al., 2020). Hh3-Hh4 were chosen as the time range for this taxon given imprecise dates for many of these localities. Absolute dates for the Hemphillian subunits follow Hilgen et al. (2012). Amphimachairodus giganteus Locality: Various, Europe, Asia. Absolute age estimate: 8.7-5.3 Ma (Slater and Friscia, 2019). Xenosmilus hodsonae Locality: Florida, USA. Absolute age estimate: 2.5-1.0 Ma. Specimens from University of Florida localities span the latest Blancan to early Irvingtonian. The Santa Fe River locality is likely the oldest at approximately 2.5 Ma, while the youngest locality is likely Haile 21A, site of the holotype, inferred to be slightly younger than the dated Leisy Shell Pit at 1.1-1.55 Ma (Bell et al., 2004). Homotherium ischyrus Locality: North America. Absolute age estimate: 4.9-2.0 Ma. Following the findings of Antón et al. (2014), I limited the biostratigraphic age of this species to the 227 Blancan, compared to the later, morphologically diverse Homotherium latidens (Paijmans et al., 2017). Dates follow Hilgen et al. (2012). Homotherium latidens Locality: Europe. Absolute age estimate: 0.033049-0.031674 Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, Natuurmuseum Rotterdam catalogue number 02-011, GenBank: MF871701. Six total radiocarbon dates have been collected on this specimen (Reumer et al., 2003). The last two of these dates were considered most reliable, with inferred contamination in the first four. Of these two dates I chose the date obtained from the tooth (28,100 ± 220 14C age yr. BP), as it is the one primarily reported by Reumer et al. (2003) for the age of this specimen. This date in two sigma calibrated years (using CalPal Rev 8.1.0) is the range reported above. A normal distribution spanning this range (2 sigma confidence interval, mean=0.03236, sigma = 0.0004) was used as the tip prior in the phylogenetic analysis. Homotherium serum Locality: North America. Absolute age estimate: > 0.0565 Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, Canadian Museum of Nature, Ottawa, CMN46442; GenBank: MF871703. The provided radiocarbon date (>56,500 14C age yr. BP; Paijmans et al., 2017) is beyond the 45–50 ka resolution limit of radiocarbon dating and thus provides a minimum age of the specimen. I chose an exponential distribution (mean 0.0511, 95% quantile 0.2 Ma, offset 0.0565) such that the 95% quantile extended to the start of the Rancholabrean, the first occurrence of this species in North America (Slater and Friscia, 2019). 228 Megantereon cultridens Locality: Various, Africa, Eurasia, ?North America. Absolute age estimate: 3.5-0.4 Ma (Slater and Friscia, 2019). Smilodon fatalis Localities: North America (USA, Mexico) and possibly western South America. Absolute age estimate: 0.6-0.0095 Ma (Slater and Friscia, 2019). Smilodon populator Localities: Various, Venezuela, Ecuador, Bolivia, Argentina, Brazil. Absolute age estimate: 0.013372-0.013235 Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, an uncatalogued left tibia from the Kruimel collection, Naturalis, Leiden, GenBank: MF871700 (11,335 ± 30 14C age yr. BP; Paijmans et al., 2017). This date in two sigma calibrated years (using CalPal Rev 8.1.0) is the range reported above. A normal distribution spanning this range (2 sigma confidence interval, mean=0.0133035, sigma = 0.00004) was used as the tip prior in the phylogenetic analysis. Panthera leo spelaea Localities: Various, Europe, Asia, North America (Barnett et al., 2009). Absolute age estimate: > 0.061Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, hair bolus, sample F-2678/70, GenBank: KX258452 (Barnett et al., 2016). However, there is a discrepancy in the radiocarbon dates for the hair sample and that of associated skeletal material (Kirillova et al., 2015). This discrepancy the authors attributed to contamination of the hair sample, while not of the skeletal material. Thus, I chose to use the radiocarbon date for the skeletal material, but given its extent beyond the resolution limit of radiocarbon dates I applied an exponential distribution (mean 0.61, 95% quantile 1.89 Ma, offset 229 0.061) such that the 95% quantile extended to inferred split between Panthera leo and Panthera leo spelaea (Barnett et al., 2016). Panthera atrox Localites: Various, North America. Absolute age estimate: 0.013401-0.013226 Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, Metatarsal II, Royal Alberta Museum EDM P89, GenBank: DQ899945.1 (11,355 ± 55 14C age yr. BP; Barnett et al., 2009). This date in two sigma calibrated years (using CalPal Rev 8.1.0) is the range reported above. A normal distribution spanning this range (2 sigma confidence interval, mean = 0.0133135, sigma = 0.00004) was used as the tip prior in the phylogenetic analysis. Miracinonyx trumani Localites: Various, North America. Absolute age estimate: 0.024037-0.023781 Ma. The date for this species is tied to the radiocarbon date of the specimen sampled for DNA, University of Kansas, KS28 NTC43093, GenBank: DQ097168.1, DQ097175.1, DQ097170.1 (19,765 ± 80 14C age yr. BP; Barnett et al., 2005). This date in two sigma calibrated years (using CalPal Rev 8.1.0) is the range reported above. A normal distribution spanning this range (2 sigma confidence interval, mean=0.023909, sigma = 0.0001) was used as the tip prior in the phylogenetic analysis. Lophocyonidae Izmirictis cani Localites: Sabuncubeli, Turkey. Absolute age estimate: 19.5-17.2 Ma. This taxon comes from the lower part of the Soma Formation of western Turkey (Morales et al., 2019). No absolute dates are available for this locality, but biostratigraphy suggests an early MN 230 3 fauna (Morales et al., 2019). However, barring more-detailed temporal placement, I chose to use the entire MN 3 zone for this phylogeny, following Hilgen et al. (2012). Euboictis aliverensis Localites: Aliveri, Greece. Absolute age estimate: 18.01-17.5 Ma. Hoek Ostende et al. (2015) place the only known fossil locality of this taxon at the above absolute age. Sivanasua viverroides Localites: France, Germany. Absolute age estimate: 17.2-11.2 Ma. This taxon is known from MN4-MN7/8 (Morales et al., 2019), with absolute ages following Hilgen et al. (2012). Herpestidae Kichechia zamanae Localites: Rusinga, Kenya. Absolute age estimate: 20.0-17.0 Ma. A recent revision has split this taxon between at least two species, with Kichechia zamanae now only explicitly known from Rusinga and Songhor (Adrian et al., 2018). Additional eastern African localities report this taxon (e.g. Kichechia sp. from Napak; Werdelin and Peigné, 2010), but barring additional taxonomic study at these sites I chose to only use the temporal age associated with Rusinga and Songhor (Werdelin and Peigné, 2010; Adrian et al., 2018). Kanuites lewisae Localites: Fort Ternan, Kenya. Absolute age estimate: 14.0-13.4 Ma. All material attributable to this taxon is known from Fort Ternan (Werdelin, 2019). The age of this locality is well known from whole-rock K/Ar and single-crystal 40Ar/39Ar at 13.7 ± 0.3 Ma (Pickford et al., 2006). This date in two sigma years is the range reported above. A normal 231 distribution spanning this range (2 sigma confidence interval, mean=13.7, sigma = 0.2) was used as the tip prior in the phylogenetic analysis. Hyaenidae Tongxinictis primordialis Locality: Tongxin, China. Absolute age estimate: 13.8-12.5 Ma. As described by Qiu et al. (1988), this taxon is known from the earliest Tungurian, roughly equivalent to MN6. The absolute dates follow Agustí et al. (2001). Allohyaena kadici Locality: Hungary, Germany. Absolute age estimate: 8.7-7.5 Ma. The described temporal age (MN 11) and geographic range of this taxon follow Turner et al. (2008), while the absolute dates follow Agustí et al. (2001). Plioviverrops orbignyi Locality Greece, Spain: Absolute age estimate: 9.7-6.8 Ma. The described temporal age (MN 10-12) and geographic range of this taxon follow Turner et al. (2008), while the absolute dates follow Agustí et al. (2001). Protictitherium crassum Locality: France, Germany, Greece, Spain, Turkey. Absolute age estimate: 17.1-5.1 Ma. Following the in-depth study of this species (particularly at Cerro de los Batallones, Spain) by Gracia (2015), the temporal range now extends from the end of MN 4 to MN 13. 232 Tungurictis spocki Locality: Tung Gur, China. Absolute age estimate: 12.75-11.1 Ma (Slater and Friscia, 2019). Ictitherium viverrinum Locality: Various Europe, Asia. Absolute age estimate: 11.5-4.9 Ma. The described temporal age and geographic range of this taxon follow Turner et al. (2008), while the absolute dates follow Agustí et al. (2001). Hyaenictitherium wongii Locality: Germany, Greece, Ukraine, Turkey, Iran, Kazakhstan, China. Absolute age estimate: 11.5-4.2 Ma. This cosmopolitan species ranges from MN 9-14 (Tseng and Wang, 2007). However, there is debate as to the generic assignment of this and other Hyaenictitherium taxa (see Semenov, 2008). Here, I followed the taxonomy of Tseng and Wang (2007) and Turner et al. (2008). Hyaenictitherium hyaenoides Locality: China, Ukraine, Iran, Turkey, Kazakhstan, Mongolia. Absolute age estimate: 7.5-4.2 Ma. Temporal (MN 11-14) and geographic ranges follow Tseng and Wang (2007). Lycyaena chaeretis Locality: Greece, Spain, Ukraine. Absolute age estimate: 8.7-6.8 Ma. The described temporal age (MN 11-12) and geographic range of this taxon follow Turner et al. (2008), while the absolute dates follow Agustí et al. (2001). 233 Palinhyaena reperta Locality: China. Absolute age estimate: 7.5-6.8 Ma. The described temporal age (MN 12) and geographic range of this taxon follow Werdelin and Solounias (1991), while the absolute dates follow Agustí et al. (2001). Belbus beaumonti Locality: Greece, Turkey. Absolute age estimate: 8.7-6.8 Ma. The described temporal age (MN 11-12) and geographic range of this taxon follow Turner et al. (2008), while the absolute dates follow Agustí et al. (2001). Adcrocuta eximia Locality: Various China, Kyrgyzstan, Europe. Absolute age estimate: 8.7-4.9 Ma (Slater and Friscia, 2019; Miller et al., 2020). Pachycrocuta brevirostris Locality: Various South and East Africa, China, Europe and South Asia. Absolute age estimate: 3.0 (Asia, Africa) - 0.5 (Europe) Ma (Slater and Friscia, 2019). Chasmaporthetes lunensis Locality: Various China, Europe, Asia. Absolute age estimate: 5.332- 2.0 Ma (Slater and Friscia, 2019). Chasmaporthetes ossifragus Locality: Various; USA, Mexico, China? Absolute age estimate: 5.332-0.85 Ma. Werdelin and Solounias (1991) recognized most North American hyaenid material as C. ossifragus, expect for the Florida specimens, which were distinguished by a longer m1 relative to the p4. However, Tseng et al. (2013) found this value to be on a continuum of other Chasmaporthetes specimens, and thus cast doubt on the validity of this specific character. 234 Thus, until more clear specific diagnostic criteria for a purported Florida species of Chasmaporthete is determined, I chose to include this material into C. ossifragus. Chasmaporthetes is a Blancan index taxon and thus was used as the first appearance datum (Bell et al., 2004; Tseng et al., 2013). The latest described specimens appear to come from the Mexican El Golfo Fauna, thought to be Irvingtonian in age, but with no associated radiometric or paleomagnetic dates (Bell et al., 2004). For this study an Irvingtonian 1 latest occurrence was used, given the preponderance of Blancan dates for this taxon, and lack of other known Irvingtonian occurrences. Crocuta crocuta spelaea Localites: Various, Europe, Asia. Absolute age estimate: 0.021889- 0.126 Ma. The date for this species is tied to the age of the specimen sampled for DNA, petrous bone, GenBank: MN320462.1 (Westbury et al., 2020). Unfortunately, there is no connection between GenBank accession numbers and the radiocarbon dates of the three German cave hyena specimens in Westbury et al. (2020). Thus, I chose to use a uniform prior to encompass the dates of all three German cave hyena specimens to be assured of sampling the most likely date. The youngest specimen, Ccsp043, has a reported date of 18,030±50 14C age yr. BP, while the oldest specimen, Ccsp040, as “Late Pleistocene” (Westbury et al., 2020). The Late Pleistocecne is an informal unit, but has been defined by the Subcomission on Quaternary Stratigraphy (http://quaternary.stratigraphy.org/definitions/pleistocene-subdivision/ accessed on 12/22/2020) at the base of the Eemian, or 0.126 Ma. The youngest date in two sigma calibrated years (using CalPal Rev 8.1.0) returns 0.021889-0.022191 Ma. A uniform distribution spanning the minimum 2 sigma confidence interval to the base of the Late Pleistocene was used as the tip prior in the phylogenetic analysis. 235 Dinocrocuta gigantea Localites: Bulgaria, Mongolia, China. Absolute age estimate: 11.5-8.0 Ma. The precise geologic age of this taxon has long been unclear (Zhang, 2005). However, European remains attributed to this taxon are thought to be Vallesian in age (Spassov and Koufos, 2002), with a slightly later appearance at approximately Chron 5n.2n in China (Zhang, 2005). The youngest dated record is at approximately 8.0 Ma from the Red Clays of Fugu, China (Zhang, 2005). Percrocuta carnifex Localites: Pakistan. Absolute age estimate: 14.2-9.5 Ma. Recent work has reviewed the occurrence of this taxon in the Middle Siwalik (Ghaffar et al., 2019). Percrocuta carnifex is known from the Chinji and Nagaro Formations, with referrals to the younger Dhok Pathan Formation as unlikely (Ghaffar et al., 2019). Percrocuta algeriensis Localites: Algeria. Absolute age estimate: 12.18-7.0 Ma. Howell and Petter (1985) give localities at Bou Hanifia (12.18-9.7 Ma.; Bernor et al., 1980) and Menacer. The latter locality is viewed as Middle Turolian by Howell and Petter (1985), and approximately 7 Ma by Benefit et al. (2008). Percrocuta tobieni Localites: Kenya, Tunisia. Absolute age estimate: 13.8-11.0 Ma. Recent finds of this taxon at Fort Ternan extend the temporal range of this taxon to 13.8 Ma (Werdelin, 2019), while the youngest occurrences span the Ngorora Formation (12.5-11.0 Ma; Morales and Pickford, 2005; Werdelin, 2019). 236 Nimravidae Oriensmilus liupanensis. Locality: Tongxin area of Ningxia Hui Autonomous Region, northern China (Wang et al., 2020). Absolute age estimate: 16.0-15.0 Ma (Qiu et al., 2013). The holotype and additionally referred material is from an informal middle member of the Zhang’enbao Formation, from which emanates the Ma’erzhuizi Gou Fauna (Wang et al., 2016), that the authors included within the Dingjia’ergou Fauna. The latter fauna is the date to which I applied the age estimate of this taxon, as did Wang et al. (2020). Sansanosmilus palmidens. Locality: France. Absolute age estimate: 16.0-13.0 Ma. The majority of S. palmidens material comes from southern France at the locality of Sansan, MN 6 (Peigné, 2012). However, additional fragmentary material has been described from Savigné-sur- Lathan (MN5) of western France (Ginsburg, 2001). Barbourofelis fricki Locality: Kansas, Nebraska, Texas, Nevada, USA. Absolute age estimate: 9.0–7.0 Ma (Tseng et al., 2010). Barbourofelis loveorum Locality: Florida, USA. Absolute age estimate: 9.5–8.0 Ma (Tseng et al., 2010). Barbourofelis morrisi Locality: Nevada, South Dakota, Texas, USA. Absolute age estimate: 12.0–9.5 Ma (Tseng et al., 2010). 237 Albanosmilus whitfordi Locality: California, Colorado, Nebraska, Texas, Florida, USA. Absolute age estimate: 12.0–7.0 Ma (Tseng et al., 2010). Albanosmilus jourdani Locality: Spain, France, Turkey. Absolute age estimate:11.9-9.7 Ma (Robles et al., 2013). Hoplophoneus cerebralis. Locality: South Dakota, California, Oregon, Wyoming, USA. Absolute age estimate: 34.7-29.586 Ma. The earliest specimen referable to this taxon is an edentulous dentary from the Crazy Johnson Member of the Chadron Formation (Barrett, 2016). The latest well-documented occurrence is from below the “Blue Basin Tuff” of the Turtle Cove Member of the John Day Formation (29.586 Ma; Famoso et al., 2015). Hoplophoneus primaevus. Locality: Nebraska, South Dakota, Wyoming, Colorado, Oregon, USA. Absolute age estimate: 35.7-30.58 Ma. Following the taxonomy of Barrett (2016), the oldest occurrence of this taxon is from Flagstaff Rim Wyoming (Bryant, 1996; Prothero and Emry, 2004). The youngest specimens attributed to this taxon are found just below the “Upper Whitney Ash” in Nebraska and the basal Leptauchenia beds of the Poleslide Member of the Brule in South Dakota (Bryant, 1996; Prothero and Emry, 2004). Hoplophoneus occidentalis. Locality: Nebraska, North Dakota, South Dakota, Wyoming, USA. Absolute age estimate: 33.4-31.4 Ma. The earliest stratigraphically constrained specimens of this taxon come from the “lower nodules” of the Scenic Member of the Brule Formation, correlated to 33.4-33.1 Ma (Bryant, 1996; Prothero and Emry, 2004). The latest occurring 238 specimens emanate from the Protoceros channel sandstones of the Poleslide Member of the Brule Formation (Bryant, 1996; Barrett, 2016). Nimravus brachyops. Locality: Nebraska, South Dakota, California, Oregon, Wyoming (USA); Saskatchewan (Canada). Absolute age estimate: 31.4- 27.14 Ma. This is a geographically wide- ranging taxon in North America with its first occurrence in the Protoceros channel sandstones of the Poleslide Member of the Brule Formation, South Dakota (Bryant, 1996; Prothero and Emry, 2004). The last reported occurrence of this taxon is from the K1 unit of the Turtle Cove Member of the John Day Formation (Famoso et al., 2015). The K1 unit is capped by the “Biotite Tuff” and forms the LAD for this study. Pogonodon platycopis. Locality: South Dakota, Nebraska, Oregon, USA. Absolute age estimate: 32.0-29.586 Ma. Numerous specimens of Whitneyan age are referred to this taxon, with none being clearly older (Bryant, 1996; Barrett, 2016). The youngest age comes from Unit E of the Turtle Cove Member of the John Day Formation, below the “Blue Basin Tuff” (Famoso et al., 2015). Pogonodon davisi. Locality: Wyoming, Nebraska, South Dakota, Oregon, USA. Absolute age estimate: 32.0-25.9 Ma. Like P. platycopis, the earliest record of this taxon is Whitneyan in age (Bryant, 1996; Barrett, 2016). The youngest specimen (JODA 5841) attributable to this taxon comes from the K2 unit of the Turtle Cove Member of the John Day Formation. This specimen was referred to as a possible new species of Pogonodon by Bryant and Fremd (1998) and Albright et al. (2008), but is included in this taxon following the diagnosis of Barrett (2016). 239 Dinictis felina. Locality: South Dakota, North Dakota, Nebraska, Oregon, Montana, Wyoming (USA); Saskatchewan (Canada). Absolute age estimate: 35.7- 29.586 Ma. The oldest record known is at Flagstaff Rim, at just over 35.5 Ma, though several slightly older occurrences are possible from fragmentary remains and provisional dates (Bryant, 1996). For example, Dinictis sp. has been collected from the Medicine Poles local fauna of southwestern North Dakota, thought to be Ch2 in age (Boyd and Webster, 2018). This latter occurrence is used as the FAD of this taxon. The LAD includes the occurrence of Dinictis cyclops (= D. felina; Barrett, 2016) below the Blue Basin Tuff (29.586 Ma; Famoso et al., 2015). Nanosmilus kurteni. Locality: Nebraska, USA. Absolute age estimate: 33.89-32.0 Ma. The holotype and only known specimen is from the Orella Member of the Brule Formation (Martin, 1992). Absolute radiometrically dated boundaries for the Orella Member are not known, but bracketed to 34.6-31.5 Ma (Zanazzi et al., 2009) based on stratigraphically and geographically, nearest dated ash layers. However, dates for the Orellan NALMA are better known and what is used for the range applied to this taxon (Prothero and Emry, 2004; Zanazzi et al., 2009). Nimravus intermedius. Locality: France, Germany, Mongolia. Absolute age estimate: 35.37- 27.24 Ma. Numerous fragmentary remains are known from the old Quercy collections, as well as the new stratigraphically documented ones (MP 22 to 25; Peigné, 2003; Peigné et al., 2014). Additional French material comes from Villebramar, MP 22 (Peigné, 2003). Of similar age in Southern Germany are attributed dentition at Möhren 13 (MP 22) and Liptingen (MP 22 or MP 23; Peigné, 2003). Mongolian Nimravus material, of Ergilian age, was separated into its own 240 species, Nimravus mongoliensis, for much of the 20th century (Gromova, 1959; Dashzeveg, 1996), but was synonymized to N. intermedius by Peigné (2003). Most recently this material was re-established as a valid species by Egi et al. (2016) based upon depth of the horizontal ramus and relative size of p1 and p2. However, the variability in anterior premolars for Nimravus has been well documented in the literature (Toohey, 1959; Peigné, 2003; Barrett, 2016). Thus, until more complete material of this purported species is described I chose to follow the assement of Peigné (2003) in its synonymy. The Thai, ‘Nimravus’ material described by Peigné et al. (2000) lacks the apomorphy of p4 morphology for this genus, and may be more closely related to Maofelis for which no lower dentognathic remains are known. Ginsburgsmilus napakensis. Locality: Langental, Fiskus, Grillental-6, Sperrgebiet, Namibia; Songhor, Kenya; Napak, Uganda. Absolute age estimate: 20.0-19.0 Ma. An updated occurrence list can be found in Morales and Pickford (2018), while Werdelin (2010) describes all of the above sites to be correlative to 20-19 Ma. Prosansanosmilus eggeri. Locality: Bavaria, Germany. Absolute age estimate: 16.7-15.97 Ma. Thus far, the only known specimens of this taxon come from Sandelzhausen (MN 5) of Bavaria (Morlo et al., 2004). Sandelzhausen is constrained in age to about 16 Ma based on magnetochronological events and biostratigraphy; see Moser et al. (2009) for a detailed overview. Prosansanosmilus peregrinus. Locality: Germany, France. Absolute age estimate: 18.0-16.0 Ma. In their original description, Heizmann et al. (1980) named three localities for the 241 occurrence of this taxon: Langenau 1 of Germany; and Bézian and Artenay of France. To this list, Petersbuch 2 of Germany was added by Morlo (2006). Petersbuch 2 and Artenay are MN 4a in age while Langenau 1 and Bézian are MN 4b. The age of MN 4 is skewed slightly older in the depositional basins of Switzerland and Southern Germany (see Reichenbacher et al., 2013) than the European “standard” (compare to Agustí et al., 2001). Furthermore, the exact age of the above fossil localities within their respective subzones is unequally known, save with Langenau 1 at slightly less than 17.2 Ma (Reichenbacher et al., 2013). Thus conservatively, the base of the South German MN 4a was used as the FAD of this taxon, while the LAD the “standard” top of MN 4. Afrosmilus africanus. Locality: Locherangan, Rusinga, and Karungu (Kenya); Buñol, Spain. Absolute age estimate: 17.8-16.0 Ma. A. africanus is known from the African sites of Locherangan, Rusinga, and Karungu (Kenya), as well as the Spanish locality of Buñol. The Kenyan locality Locherangan (c. 17.5 Ma; Anyonge, 1991) is located west of Lake Turkana, while Rusinga (c. 17.8 Ma; Werdelin, 2010) and Karungu (c. 17.7-17.5 Ma; Drake et al., 1988), though likely contemporaneous with Rusinga (Werdelin, 2010) or slightly older (Morales and Pickford, 2018) based on biostratigraphy, are localities along the northwestern shore of Lake Victoria. The Spanish locality of Buñol (MN4, 16.6-16.0 Ma; Agustí et al., 2001) is located in the east of Spain in the province of Valencia. Afrosmilus turkanae. Locality: Moruorot Hill and Rusinga, Kenya. Absolute age estimate: 17.8-16.8 Ma. A. turkanae is known from the African localities of Moruorot Hill and Rusinga (Kenya). Moruorot Hill (c. 17.5-16.8 Ma; Boschetto et al., 1992) is located west of Lake 242 Turkana, while Rusinga is the same locality (and chronology) as mentioned above for A. africanus. Afrosmilus hispanicus. Locality: Artesilla, Spain. Absolute age estimate: 16.7-16.0 Ma. A. hispanicus is the only non-African originating species of Afrosmilus, known only from the Spanish locality of Artesilla (c. 16.7 Ma; Azanza et al., 1993; Morales et al., 2001), in the northeast of the country in the province of Saragossa. A. hispanicus is typically correlated to the MN 4 zone (Morlo, 2006; Morales and Pickford, 2018), thus the younger date of 16.0 Ma is applied to its range, being the boundary between MN 4 and 5 (Agustí et al., 2001). Maofelis cantonensis. Locality: Maoming Basin, Guangdong Province, China. Absolute age estimate: 41.03-33.9 Ma. The holotype, and only known specimen, is from the Youganwo Formation, middle-upper Eocene (Averianov et al., 2016). The vertebrate fauna is only known from the upper portion of the formation containing dark oil shales and alternating mudstones (Averianov et al., 2016). The precise stratigraphic horizon of the holotype is unknown, but based on magnetostratigraphy and biostratigraphy it has been correlated to the Bartonian through Priabonian (Li et al., 2016; Averianov et al., 2019). Eofelis edwardsii. Locality: France, Mongolia. Absolute age estimate: 35.37-30.83 Ma. The vast majority of material referred to Eofelis edwardsii comes from the old collections of the phosphorites of Quercy with no precise age (Peigné, 2000). However, material questionably referred to this taxon has been described from Villebramar, France, MP 22 (Peigné, 2000). Egi et al. (2016) described Eofelis sp. from the Ergilin Dzo Formation, Mongolia, which based on their 243 measurements most closely resembles Eofelis edwardsii compared to the larger E. giganteus (Peigné, 2000). The Mongolian specimens are dated to the Ergilian ALMA (Egi et al., 2016) and thus forms the oldest occurrence of this taxon while MP 22 the youngest for this study. Dinailurictis bonali. Locality: France, Spain. Absolute age estimate: 32.63-27.24 Ma Most material attributed to this taxon comes from the Quercy fissures, dated MP 22 to 25, and Villebramar, MP 22 (Peigné, 2003; Peigné et al., 2014). The Spanish material, canine fragments, comes from Carrascosa del Campo, approximately MP 25 (Peigné, 2003). Thus, the range of this taxon was chosen as MP 22-25, with absolute dates for these zones following Ogg et al. (2016). Quercylurus major. Locality: France, Spain. Absolute age estimate: 28.82-27.24 Ma The French material, including the holotype, come from the old collections of Quercy which have no precise age (Peigné, 2003). The Spanish material from Carrascosa del Campo has an estimated age of MP 25 and thus the only reliably dated material (Peigné, 2003). Absolute dates follow Ogg et al. (2016). Dinaelurus crassus. Locality: Oregon, USA. Absolute age estimate: 29.5-28.5 Ma. The precise stratigraphic context for the holotype, and only known specimen of this taxon, were unknown when Eaton (1922) initially described it. However, a provisional Early Arikareean (Upper John Day Formation) was attributed to the specimen based on preservation and associated matrix (Eaton, 1922; Bryant, 1996). Albright et al. (2008) suggest the origin to be the E unit of the Turtle Cove Member of the John Day Formation. This assessment is given without evidence, but is one of most fossiliferous units of the Turtle Cove Member. The associated absolute dates for 244 the E unit were used for this study pending additional justification for the provenience of this taxon. Hoplophoneus oharrai. Locality: South Dakota, USA. Absolute age estimate: 35.7-34.7 Ma. The type and only specimen comes from the Crazy Johnson Member of the Chadron Formation in southwest South Dakota (Barrett, 2016). Absolute dates are not available for the Crazy Johnson member, though it contains a middle Chadronian fauna (Ch3: 34.7-35.7 Ma: Janis et al., 2008) which is used for its range here. MA-PHQ 348. Locality: Phosphorites of Quercy, south-western France. Absolute age estimate: 47.4-23.03 Ma. This plesiomorphic nimravid specimen comes from the old collections of Quercy and thus has little to no stratigraphic context. The Phosphorites have a total extent from the base of MP 11 to the early Miocene (Sigé et al., 1991; Legendre et al., 1992). Given the poor understanding of this specimen’s stratigraphic context a broad prior from the base of MP 11 to the end of the Oligocene was chosen. Eusmilus dakotensis. Locality: South Dakota, USA. Absolute age estimate: 30.5-29.75 Ma. Three referable specimens come from the Brule Fm., Poleslide Member of South Dakota (Barrett, 2016). Bryant (1996) limits the first appearance to the latter Whitneyan, during Chron C12n or C11r. This leads to the FAD as stated with the LAD being the end of the Whitneyan. Eusmilus sicarius. Locality: South Dakota, USA. Absolute age estimate: 33.4-33.1 Ma. Following the results of this study, the only known specimen belonging to this species is the 245 holotype, which comes from the lower nodule zone of the Scenic Member of the Brule Formation. This stratum is roughly correlative to Chron C13n (Prothero and Emry, 2004). Eusmilus adelos. Locality: Niobrara County, Wyoming, USA. Absolute age estimate: 33.7-32.0 Ma. The holotype specimen (Barrett, 2021) comes from the Northwest corner of Seaman Hills, Niobrara County, Wyoming. The additional referred specimen was collected about one mile North of Whitman, Wyoming. Both of these specimens come from the Seaman Hills which spans the entirety of the Orellan in typical outcrops (Prothero and Whittlesey, 1998; Prothero and Emry, 2004). Unfortunately, with no greater stratigraphic or geographic resolution, the entirety of the Orellan was used as the occurrence for this taxon. Eusmilus bidentatus. Locality: France, Germany. Absolute age estimate: 33.77-32.63 Ma. This taxon is well known in the Phosphorites of Quercy, but also Soumailles (Ringeade and Michel, 1994; Peigné and Brunet, 2001). Additional material is known from southern Germany, but all of it (along with the French material) has been correlated to MP 21 (Peigné and Brunet, 2001). Eusmilus villebramarensis. Locality: France, Switzerland, Germany. Absolute age estimate: 32.63-30.83 Ma. Most material for this taxon comes from France in the old collections of Quercy, but also Villebramar (Peigné and Brunet, 2001). However, occurrences in Switzerland and Germany are also known, which with the Villebramar material also has a MP 22 date (Peigné and Brunet, 2001). 246 References Adrian, B., L. Werdelin, and A. Grossman. 2018. New Miocene Carnivora (Mammalia) from Moruorot and Kalodirr, Kenya. Palaeontologia Electronica 21. Agustí, J., L. Cabrera, M. Garcés, W. Krijgsman, O. Oms, and J. M. Parés. 2001. A calibrated mammal scale for the Neogene of Western Europe. State of the art. Earth Science Reviews 52:247–260. 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. Antón, A. M., M. J. Salesa, and G. Siliceo. 2013. Machairodont Adaptations and Affinities of the Holarctic Late Miocene Homotherin Machairodus (Mammalia, Carnivora, Felidae): The Case of Machairodus Catocopis Cope, 1887. Journal of Vertebrate Paleontology 33:1202– 1213. Antón, M., M. J. Salesa, A. Galobart, and Z. J. Tseng. 2014. The Plio-Pleistocene scimitar- toothed felid genus Homotherium Fabrini, 1890 (Machairodontinae, Homotherini): Diversity, palaeogeography and taxonomic implications. Quaternary Science Reviews 96:259–268. Anyonge, W. 1991. Fauna from a New Lower Miocene Locality West of Lake Turkana, Kenya. Journal of Vertebrate Paleontology 11:378–390. Averianov, A., E. Obraztsova, I. Danilov, and J. Jin. 2019. Anthracotheriid artiodactyl Anthracokeryx and an upper Eocene age for the Youganwo Formation of southern China. Historical Biology 31:1115–1122. Averianov, A., E. Obraztsova, I. Danilov, P. Skutschas, and J. Jin. 2016. First nimravid skull from Asia. Scientific Reports 6:1–8. 247 Azanza, B., E. Cerdeno, L. Ginsburg, J. Van Der Made, J. Morales, and P. Tassy. 1993. Les grands mammifères du Miocène inférieur d’Artesilla, bassin de Calatayud-Teruel (province de Saragosse, Espagne). Bulletin Du Museum National d’Histoire Naturelle Section C Sciences de La Terre Paleontologie Geologie Mineralogie 15:105–153. Barnett, R., M. L. Z. Mendoza, A. E. R. Soares, S. Y. W. Ho, G. Zazula, N. Yamaguchi, B. Shapiro, I. V. Kirillova, G. Larson, and M. T. P. Gilbert. 2016. Mitogenomics of the extinct cave lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the panthera cats. Open Quaternary 2:1–11. Barnett, R., B. Shapiro, I. Barnes, S. Y. Ho, J. Burger, N. Yamaguchi, T. F. Higham, H. T. Wheeler, W. Rosendahl, A. V Sher, M. Sotnikova, T. Kuznetsova, G. F. Baryshnikov, L. D. Martin, C. R. Harington, J. A. Burns, and A. Cooper. 2009. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity. Molecular Ecology 18:1668–1677. 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. Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658. Barrett, P. Z. 2021. The largest hoplophonine and a complex new hypothesis of nimravid evolution. Scientific Reports 11:1–9. Barrett, P. Z., S. S. B. Hopkins, and S. A. Price. 2021. How many sabretooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae. Journal of Vertebrate Paleontology e1923523. Bell, C. J., E. L. Lundelius Jr., A. D. Barnosky, R. W. Graham, E. H. Lindsay, D. R. Ruez Jr., H. A. Semken Jr., S. D. Webb, and R. J. Zakrzewski. 2004. The Blancan, Irvingtonian, and Rancholabrean Mammal Ages; pp. 232–314 in M. O. Woodburne (ed.), Late Cretaceous and Cenozoic Mammals of North American. Columbia University Press, New York. Benefit, B. R., M. McCrossin, N. T. Boaz, and P. Pavlakis. 2008. New Fossil Cercopithecoids from the Late Miocene of As Sahabi, Libya. Garyounis Scientific Bulletin 5:265–282. 248 Bernor, R. L., M. O. Woodburne, and J. A. Van Couvering. 1980. A contribution to the chronology of some Old World miocene faunas based on hipparionine horses. Geobios 13:705–739. Böhme, M., N. Spassov, M. Ebner, D. Geraads, L. Hristova, U. Kirscher, S. Kötter, U. Linnemann, J. Prieto, S. Roussiakis, G. Theodorou, G. Uhlig, and M. Winklhofer. 2017. Messinian age and savannah environment of the possible hominin Graecopithecus from Europe. PLoS ONE 12:e0177347. Boschetto, H. B., F. H. Brown, and I. M. McDougall. 1992. Stratigraphy of the Lothidok Range, northern Kenya, and K/Ar ages of its Miocene primates. Journal of Human Evolution 22:47–71. Boyd, C. A., and J. R. Webster. 2018. Depositional History of the Chadron Formation in North Dakota. North Dakota Geological Survey Report of Investigation 120:107. Bryant, H. N. 1992. The Carnivora of the Lac Pelletier Lower Fauna (Eocene: Duchesnean), Cypress Hills Formation, Saskatchewan. Journal of Paleontology 66:847–855. 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. Bryant, H. N., and T. J. Fremd. 1998. Revised Biostratigraphy of the Nimravidae (Carnivora) from the John Day Basin of Oregon. Journal of Vertebrate Paleontology 18:30A. Dashzeveg, D. 1996. Some Carnivorous Mammals from the Paleogene of the Eastern Gobi Desert , Mongolia , and the Application of Oligocene Carnivores to Stratigraphic Correlation. American Museum Novitates 3179:1–14. Drake, R. E., J. A. Van Couvering, M. H. Pickford, G. H. Curtis, and J. A. Harris. 1988. New chronology for the Early Miocene mammalian faunas of Kisingiri, Western Kenya. Journal of the Geological Society, London 145:479–491. Eaton, G. F. 1922. John Day Felidae in the Marsh Collection. American Journal of Science 4:425–452. 249 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. Famoso, N. A., J. X. Samuels, S. S. B. Hopkins, M. M. Emery, and E. B. Davis. 2015. Updated Biostratigraphy of the Turtle Cove Member (John Day Formation) in the John Day Basin, Oregon; pp. 6 in Western association of vertebrate Paleontology annual Meeting: Program with abstracts. PaleoBios 32(1). vol. 32. Gabunia, L., and A. Vekua. 1998. The find of Dinofelis in the Pliocene of Georgia. Bulletin of the Georgian Academy of Sciences 157:335–338. Ghaffar, A., M. Akhtar, M. A. Khan, and M. A. Babar. 2019. Comments on percrocuta carnifex (carnivora, percrocutidae) based on new fossil material from the nagri formation (middle siwaliks) of hasnot, pakistan. Geologica Acta 17:1–9. Ginsburg, L. 2001. Les faunes de mammifères terrestres du Miocène moyen des Faluns du bassin de Savigné-sur-Lathan (France). Geodiversitas 23:381–394. Gracia, S. F. 2015. Estudio de Protictitherium crassum del Cerro de los Batallones (Torrejón de Velasco, Madrid): aportación a la filogenia y evolución de la familia hyaenidae. Universidad Complutense de Madrid, 365 pp. Gromova, V. 1959. Premiere decoverte d’un chat primitif au Paleogene d’Asie Centrale. Vertebrata PalAsiatica 3:59–72. Heizmann, E. P. J., L. Ginsburg, and C. Bulot. 1980. Prosansanosmilus peregrinus, ein neuer machairodontider Felide aus dem Miocän Deutschlands und Frankreichs. Stuttgarter Beiträge Zur Naturkunde B 58:1–27. Hilgen, F. J., L. J. Lourens, J. A. Van Dam, A. G. Beu, A. F. Boyes, R. A. Cooper, W. Krijgsman, J. G. Ogg, W. E. Piller, and D. S. Wilson. 2012. The Neogene Period; pp. 923– 978 in F. M. Gradstein, J. G. Ogg, M. D. Schmitz, and G. M. Ogg (eds.), The geologic time scale. Elsevier, Amsterdam. 250 Hoek Ostende, L. W. van den, S. Mayda, A. Oliver, A. Madern, V. Hernández-Ballarín, and P. Peláez-Campomanes. 2015. Aliveri revisited, a biogeographical appraisal of the early Miocene mammals from the eastern Mediterranean. Palaeobiodiversity and Palaeoenvironments 95:271–284. Howell, F. C., and G. Petter. 1985. Comparative observations on some middle and upper Miocene hyaenids. Genera: Percrocuta Kretzoi, Allohyaena Kretzoi, Adcrocuta Kretzoi (Mammalia, Carnivora, Hyaenidae). Geobios 18:419–476. Hunt, R. M. 1991. Evolution of the aeluroid Carnivora: viverrid affinities of the Miocene carnivoran Herpestides. American Museum Novitates 3023:1–34. Janis, C. M., G. F. Gunnell, and M. D. Uhen. 2008. Evolution of Tertiary Mammals of North America. Volume 2: Small Mammals, Xenarthrans, and Marine Mammals. Cambridge University Press, Cambridge, 795 pp. Kaakinen, A., B. H. Passey, Z.-Q. Zhang, L.-P. Liu, L. J. Pesonen, and M. Fortelius. 2013. Stratigraphy and Paleoecology of the Classical Dragon Bone Localities of Baode County, Shanxi Province; pp. 203–217 in Fossil mammals of Asia: Neogene biostratigraphy and chronology. Columbia University Press, New York. Kelly, T. S., P. C. Murphey, and S. L. Walsh. 2012. New Records of Small Mammals From the Middle Eocene Duchesne River Formation, Utah, and Their Implications for the Uintan- Duchesnean North American Land Mammal Age Transition. Paludicola 8:208–251. Kirillova, I. V., A. V. Tiunov, V. A. Levchenko, O. F. Chernova, V. G. Yudin, F. Bertuch, and F. K. Shidlovskiy. 2015. On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia). Quaternary Science Reviews 117:135–151. Kovatchev, D. 2001. Description d’un squelette complet de Metailurus (Felidae, Carnivora, Mammalia) du Miocène supérieur de Bulgarie. Geologica Balcanica 31:71–88. Legendre, S., B. Marandat, B. Sigé, J. Crochet, M. Godinot, J. L. Hartenberger, J. Sudre, and M. Vianey-Liaud. 1992. La faune de mammifères de Vielase (phosphorites du Quercy, Sud de la France): Preuve paléontologique d’une karstification du Quercy dès l’Eocène inférieur Par Montpellier ; Toulouse Débutées en 1965 , l ’ exploration et l ’ exploitation paléontologique no. Neues Jahrbuch Für Geologie Und Paläontologie 7:414–428. 251 Li, Y. X., W. J. Jiao, Z. H. Liu, J. H. Jin, D. H. Wang, Y. X. He, and C. Quan. 2016. Terrestrial responses of low-latitude Asia to the Eocene-Oligocene climate transition revealed by integrated chronostratigraphy. Climate of the Past 12:255–272. Martin, L. D. 1992. A new miniature saber-tooth nimravid from the Oligocene of Nebraska. Annales Zoologici Fennici 28:341–348. Martin, L. D. 1998. 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. Miller, S. A., P. Z. Barrett, W. N. F. McLaughlin, and S. S. B. Hopkins. 2020. Endemism and migration in the Kochkor Basin? Identification and description of Adcrocuta eximia (Mammalia: Carnivora: Hyaenidae) and C.F. Paramachaerodus (Mammalia: Carnivora: Felidae) fossils at the Miocene locality of Ortok, Kyrgyzstan. Palaeontologia Electronica 23:a45. Morales, J., and M. Pickford. 2005. Carnivores from the middle miocene ngorora formation (13- 12 MA), Kenya. Estudios Geologicos 61:271–284. 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. 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. Morales, J., S. Mayda, A. Valenciano, D. DeMiguel, and T. Kaya. 2019. A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey. Journal of Systematic Palaeontology 17:1127–1138. 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. 252 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. Moser, M., G. E. Rössner, U. B. Göhlich, M. Böhme, and V. Fahlbusch. 2009. The fossil lagerstätte Sandelzhausen (Miocene; southern Germany): History of investigation, geology, fauna, and age. Palaontologische Zeitschrift 83:7–23. Ogg, J. G., G. Ogg, and F. M. Gradstein. 2016. A Concise Geologic Time Scale. Elsevier, 240 pp. 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. Peigné, S. 2000. A new species of Eofelis (Carnivora: Nimravidae) from the Phosphorites of Quercy, France. Comptes Rendus de l’Académie Des Sciences, Paris 330:653–658. Peigné, S. 2003. Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32:199–229. Peigné, S. 2012. Les Carnivora de Sansan; pp. 559–660 in S. E. N. S. (ed.), Mammifères de Sansan. Muséum national d’Histoire naturelle, Paris. Peigné, S., and M. Brunet. 2001. Une Nouvelle Espèce Du Genre Eusmilus (Carnivora: Nimravidae) De L’Oligocène (MP 22) D’Europe. Geobios 34:657–672. Peigné, S., M. Vianey-Liaud, T. Pélissié, and B. Sigé. 2014. Valbro: A new site of vertebrates from the early Oligocene (MP22) of France (Quercy). I – Geological context; Mammalia: Rodentia, Hyaenodontida, Carnivora. Annales de Paleontologie 100:1–45. Peigné, S., Y. Chaimanee, J.-J. Jaeger, V. Suteethorn, and S. Ducrocq. 2000. Eocene nimravid carnivorans from Thailand. Journal of Vertebrate Paleontology 20:157–163. 253 Pickford, M., Y. Sawada, R. Tayama, Y. ko Matsuda, T. Itaya, H. Hyodo, and B. Senut. 2006. Refinement of the age of the Middle Miocene Fort Ternan Beds, Western Kenya, and its implications for Old World biochronology. Comptes Rendus - Geoscience 338:545–555. Prothero, D. R., and K. E. Whittlesey. 1998. Magnetic stratigraphy and biostratigraphy of the Orellan and Whitneyan land-mammal “ages” in the White River Group. Geological Society of America Special Paper 325:39–61. Prothero, D. R., and R. J. Emry. 2004. The Chadronian, Orellan, and Whitneyan North American land mammal ages; pp. 156–168 in M. O. Woodburne (ed.), Late Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology. Columbia University Press, New York City. Qiu, Z.-X., Z.-D. Qiu, T. Deng, C.-K. Li, Z.-Q. Zhang, B.-Y. Wang, and X. Wang. 2013. Neogene land mammal stages/ages of China – toward the goal to establish an Asian land mammal stage/age scheme; pp. 29–90 in X. Wang, L. J. Flynn, and M. Fortelius (eds.), Fossil mammals of Asia: Neogene biostratigraphy and chronology. Columbia University Press, New York. Qiu, Z., J. Ye, and J. Cao. 1988. A new species of Percrocuta from Tongxin, Ningxia. Vertebrata Palasiatica 26:116–127. Reichenbacher, B., W. Krijgsman, Y. Lataster, M. Pippèrr, C. G. C. Van Baak, G. Doppler, L. Chang, D. Kälin, J. Jost, D. Jung, J. Prieto, H. A. A. Bohme, M. Böhme, J. Garnish, U. Kirscher, and V. Bachtadse. 2013. A new magnetostratigraphic framework for the Lower Miocene A new magnetostratigraphic framework for the Lower Miocene ( Burdigalian / Ottnangian , Karpatian ) in the North Alpine Foreland Basin. Swiss Journal of Geosciences 106:309–334. Reumer, J. W. F., L. Rook, K. Van Der Borg, K. Post, D. Mol, and J. De Vos. 2003. Late pleistocene survival of the saber-toothed cat Homotherium in Northwestern Europe. Journal of Vertebrate Paleontology 23:260–262. Ringeade, M., and P. Michel. 1994. A propos de l’Eusmilus (Eusmilus bidentatus ringeadei Ringeade et Michel, 1994) de Soumailles, lieu-dit de la commune de Pardaillan, Canton de Duras (Lot et Garonne, France) : étude descriptive. Paléo 6:5–37. 254 Robinson, P., G. F. Gunnell, S. L. Walsh, W. C. Clyde, J. E. Storer, R. K. Stucky, D. J. Froehlich, I. Ferrusquia-Villafranca, M. C. McKenna, and M. Woodburne. 2004. Wasatchian through duchesnean biochronology.; pp. 106–155 in M. Woodburne (ed.), Late Cretaceous and Cenozoic mammals of North America: Biostratigraphy and Geochronology. Columbia University Press, New York. 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. Rook, L., and B. Martínez-Navarro. 2010. Villafranchian: The long story of a Plio-Pleistocene European large mammal biochronologic unit. Quaternary International 219:134–144. Rothwell, T. P. 2001. Phylogenetic systematics of North American Pseudaelurus (Carnivora: Felidae). Columbia University, 358 pp. Rothwell, T. P. 2003. Phylogenetic systematics of North American Pseudaelurus (Carnivora, Felidae). American Museum Novitates 3403. Roussiakis, S. J. 2001. Metailurus major Zdansky, 1924 (Carnivora, Mammalia) from the classical locality of Pikermi (Attica, Greece). Annales de Paleontologie 87:119–132. Ruiz-Ramoni, D., A. D. Rincón, and M. Montellano-Ballesteros. 2020. Taxonomic revision of a Machairodontinae (Felidae) from the Late Hemphillian of México. Historical Biology 32:1312–1319. Semenov, Y. 2008. Taxonomical reappraisal of “ictitheres” (Mammalia, Carnivora) from the Late Miocene of Kenya. Comptes Rendus Palevol 7:529–539. Sigé, B., J. P. Aguilar, J. G. Astruc, and B. Marandat. 1991. Extension au Miocène inférieur des remplissages phosphatés du Quercy. La faune de Vertébrés de Crémat (Lot, France). Geobios 24:497–502. 255 Skinner, M. F., F. W. Johnson, and C. Frick. 1984. Tertiary stratigraphy and the Frick Collection of fossil vertebrates from north-central Nebraska. Bulletin of the American Museum of Natural History 178:368. Slater, G. J., and A. R. Friscia. 2019. Hierarchy in adaptive radiation: A case study using the Carnivora (Mammalia). Evolution 73:524–539. 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. Spassov, N., and G. D. Koufos. 2002. The first appearance of Dinocrocuta gigantea and Macbairodus aphanistus (Mammalia: Carnivora) in the Miocene of Bulgaria. Mitteilungen Der Bayerischen Staatssammlung Fur Palaontologie Und Historische Geologie 42:83–101. Spassov, N., and D. Geraads. 2015. A New Felid from the Late Miocene of the Balkans and the Contents of the Genus Metailurus Zdansky, 1924 (Carnivora, Felidae). Journal of Mammalian Evolution 22:45–56. Spaulding, M., J. J. Flynn, and R. K. Stucky. 2010. A new basal carnivoramorphan (Mammalia) from the “Bridger B” (Black’s fork member, bridger formation, bridgerian nalma, middle eocene) of wyoming, USA. Palaeontology 53:815–832. Tedford, R. H., L. B. I. Albright, A. D. Barnosky, I. Ferrusquia−Villafranca, R. M. J. Hunt, J. E. Storer, C. C. I. Swisher, M. R. Voorhies, S. D. Webb, and D. . Whistler. 2004. Mammalian biochronology of the Arikareean through Hemphillian interval (late Oligocene through early Pliocene epochs); pp. 169–231 in M.O. Woodburne (ed.), Late Cretaceous and Cenozoic Mammals of North America. Columbia University Press, New York. 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. Toohey, L. 1959. The Species of Nimravus (Carnivora, Felidae). Princeton University, New York, 130 pp. 256 Tseng, Z. J., and X. Wang. 2007. The first record of the late Miocene Hyaenictitherium hyaenoides Zdansky (Carnivora: Hyaenidae) in Inner Mongolia and an evaluation of the genus. Journal of Vertebrate Paleontology 27:699–708. 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. Tseng, Z. J., Q. Li, and X. Wang. 2013. A new cursorial hyena from Tibet, and analysis of biostratigraphy, paleozoogeography, and dental morphology of Chasmaporthetes (Mammalia, Carnivora). Journal of Vertebrate Paleontology 33:1457–1471. Turner, A., M. Antón, and L. Werdelin. 2008. Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe. Geobios 41:677–687. Wang, S.-Q., L.-Y. Zong, Q. Yang, B.-Y. Sun, Y. Li, Q.-Q. Shi, X.-W. Yang, J. Ye, and W.-Y. Wu. 2016. Biostratigraphic subdividing of the Neogene Dingjia’ergou mammalian fauna, Tongxin County, Ningxia Province, and its background for the uplift of the Tibetan Plateau36. Quaternary Sciences 36:789–809. Wang, X. 1994. Phylogenetic systematics of the Hesperocyoninae (Carnivora, Canidae). Bulletin of the American Museum of Natural History 221. 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. Werdelin, L. 2010. Chronology of Neogene Mammal Localities; pp. 27–43 in L. Werdelin and W. J. Sanders (eds.), Cenozoic Mammals of Africa. University of California Press, Berkeley, Los Angeles, London. Werdelin, L. 2019. Middle miocene carnivora and hyaenodonta from fort Ternan, Western Kenya. Geodiversitas 41:267–283. 257 Werdelin, L., and N. Solounias. 1991. The Hyaenidae : taxonomy, systematics and evolution. Fossils and Strata 30:104. Werdelin, L., and M. E. Lewis. 2001. A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132:147–258. Werdelin, L., and S. Peigné. 2010. Carnivora; pp. 603–657 in L. Werdelin and W. J. Sanders (eds.), Cenozoic Mammals of Africa. University of California Press, Berkeley, Los Angeles, London. 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. Westbury, M. V., S. Hartmann, A. Barlow, M. Preick, B. Ridush, D. Nagel, T. Rathgeber, R. Ziegler, G. Baryshnikov, G. Sheng, A. Ludwig, I. Wiesel, L. Dalen, F. Bibi, L. Werdelin, R. Heller, and M. Hofreiter. 2020. Hyena paleogenomes reveal a complex evolutionary history of cross-continental gene flow between spotted and cave hyena. Science Advances 6:1–11. Wolsan, M., and M. Morlo. 1997. The status of “Plesictis” croizeti, “Plesictis” gracilis and “Lutra” minor: synonyms of the early Miocene viverrid Herpestides antiquus (Mammalia, Carnivora). Bulletin of the Natural History Museum London (Geol) 53:1–9. Zanazzi, A., M. J. Kohn, and D. O. Terry. 2009. Biostratigraphy and paleoclimatology of the Eocene-Oligocene boundary section at Toadstool Park (northwestern Nebraska). Geological Society of America Special Paper 452:197–214. Zhang, Z. 2005. New materials of Dinocrocuta (Percrocutidae, Carnivora) from Lantian, Shaanxi Province, China, and remarks on Chinese Late Miocene biochronology. Geobios 38:685– 689. 258 Appendix H Dietary Classification and Ecological Data Tables Table 1. Extant taxa and associated data used for CVA training set. Dentognathic metrics are averages per species taken from Slater and Friscia (2019). C1 = compression of the upper C1, P4S = ratio of width to length of fourth lower premolar, RBL = relative blade length of the trigonid of m1, M1BS = blade length of the m1 relative to total dentary length. Degree of carnivory is calculated by percent of diet belonging to the Diet-Vend (vertebrate endotherm), Diet-Vect (vertebrate ectotherm), Diet-Vfish (vertebrate fish), Diet-Vunk (vertebrate unknown), Diet-Scav (scavenge) categories of the Elton Traits dataset (Wilman et al., 2014). Diet reflects the following categories: 1 = hypocarnivore, 2 = mesocarnivore, 3 = hypercarnivore. C1 P4S RBL M1BS degree diet clade of carnivor y Acinonyx jubatus 0.77205 0.40163 1 0.15139 100 3 Felidae 4 2 4 Ailurus fulgens 0.66926 0.63478 0.62610 0.09532 10 1 Ailuridae 3 6 6 5 Arctictis binturong 0.61020 0.72164 0.5939 0.05107 50 2 Viverridae 8 4 7 Arctogalidia 0.72781 0.64677 0.59326 0.05687 60 2 Viverridae trivirgata 8 4 7 9 259 Atelocynus 0.66531 0.47592 0.64459 0.08397 80 3 Canidae microtis 4 5 2 Atilax paludinosus 0.72967 0.68234 0.72531 0.09809 60 2 Herpestidae 1 5 2 1 Bassaricyon alleni 0.80870 0.79115 0.45601 0.04745 10 1 Procyonidae 7 7 1 2 Bassaricyon gabbii 0.78306 0.85210 0.49278 0.05059 10 1 Procyonidae 9 2 8 6 Bassariscus 0.70332 0.50174 0.61185 0.08803 10 1 Procyonidae astutus 3 7 3 3 Bassariscus 0.69778 0.60897 0.61452 0.07765 10 1 Procyonidae sumichrasti 4 8 7 1 Bdeogale 0.72887 0.58803 0.56413 0.05421 20 1 Herpestidae crassicauda 5 4 3 3 Bdeogale nigripes 0.68366 0.62357 0.47748 0.04761 40 1 Herpestidae 8 4 9 Canis adustus 0.63840 0.45317 0.62803 0.08493 50 2 Canidae 5 8 6 9 Canis aureus 0.61618 0.45807 0.68265 0.10987 80 3 Canidae 1 9 7 4 Canis latrans 0.57667 0.44749 0.69641 0.10958 100 3 Canidae 6 6 1 3 Canis lupus 0.62145 0.52157 0.70792 0.11052 100 3 Canidae 260 5 4 7 8 Canis mesomelas 0.61704 0.44972 0.65814 0.10824 60 2 Canidae 5 4 2 2 Canis simensis 0.63720 0.45429 0.68465 0.08964 100 3 Canidae 7 3 2 7 Caracal caracal 0.75985 0.46037 1 0.14873 100 3 Felidae 6 9 7 Caracal serval 0.74340 0.42630 1 0.14057 100 3 Felidae 4 7 1 Cerdocyon thous 0.65387 0.46994 0.62643 0.08952 50 2 Canidae 5 1 6 Chrotogale 0.59932 0.39607 0.42447 0.03764 10 1 Viverridae owstoni 3 5 3 1 Chrysocyon 0.62425 0.50251 0.64114 0.08739 70 3 Canidae brachyurus 8 5 3 Civettictis civetta 0.80786 0.59631 0.61508 0.08220 50 2 Viverridae 7 7 7 Crocuta crocuta 0.72886 0.55983 0.91677 0.14806 100 3 Hyaenidae 4 2 7 3 Crossarchus 0.64787 0.59487 0.54375 0.05782 20 1 Herpestidae alexandri 1 1 8 Crossarchus 1.11303 0.56667 0.48447 0.05396 20 1 Herpestidae obscurus 6 2 2 4 261 Crossarchus 0.59279 0.54115 0.51644 0.05680 50 2 Herpestidae platycephalus 8 2 1 9 Cryptoprocta ferox 0.75258 0.56267 0.80478 0.11767 90 3 Eupleridae 3 3 9 8 Cuon alpinus 0.60538 0.49566 0.72617 0.12377 100 3 Canidae 1 1 9 6 Cynictis penicillata 0.68673 0.51814 0.70852 0.08666 10 1 Herpestidae 3 7 2 5 Cynogale bennettii 0.70279 0.45827 0.48085 0.05338 50 2 Viverridae 7 8 6 Diplogale hosei 0.68627 0.52584 0.47788 0.04921 40 1 Viverridae 5 7 6 8 Dusicyon australis 0.63463 0.48011 0.71256 0.10422 100 3 Canidae 6 4 6 1 Eupleres goudotii 0.69980 0.35062 0.58566 0.05052 10 1 Eupleridae 4 7 6 4 Felis chaus 0.78590 0.38494 1 0.14050 100 3 Felidae 9 4 8 Felis margarita 0.78431 0.43255 1 0.14258 100 3 Felidae 4 1 5 Felis nigripes 0.72576 0.46701 1 0.14099 100 3 Felidae 8 4 1 Felis silvestris 0.77786 0.43040 1 0.14629 90 3 Felidae 262 3 4 Fossa fossana 0.65510 0.45010 0.59251 0.07058 20 1 Eupleridae 7 7 7 4 Galerella 0.68484 0.48016 0.63871 0.08980 20 1 Herpestidae pulverulenta 5 1 9 Galerella 0.66263 0.48112 0.74556 0.10447 20 1 Herpestidae sanguinea 6 6 7 8 Galidia elegans 0.69114 0.50299 0.65632 0.10911 80 3 Eupleridae 9 2 6 Galidictis fasciata 0.67052 0.57047 0.62718 0.08372 70 3 Eupleridae 5 9 2 9 Genetta angolensis 0.74320 0.44367 0.73872 0.09010 90 3 Viverridae 9 1 3 5 Genetta genetta 0.71593 0.41667 0.74356 0.0892 90 3 Viverridae 9 1 Genetta maculata 0.73610 0.41833 0.72120 0.08754 90 3 Viverridae 3 4 6 7 Genetta pardina 0.68675 0.36535 0.67536 0.08288 70 3 Viverridae 3 1 4 Genetta servalina 0.70336 0.39411 0.73857 0.08590 90 3 Viverridae 7 9 9 Genetta thierryi 0.74921 0.37385 0.65037 0.07870 90 3 Viverridae 8 9 8 6 263 Genetta tigrina 0.77994 0.40295 0.69505 0.08791 90 3 Viverridae 5 2 7 Helogale hirtula 0.62981 0.64546 0.82431 0.09499 20 1 Herpestidae 8 4 6 6 Helogale parvula 0.60952 0.61413 0.78429 0.08741 20 1 Herpestidae 8 8 8 5 Hemigalus 0.65491 0.48340 0.43740 0.04683 0 1 Viverridae derbyanus 8 4 7 9 Herpestes 0.71376 0.56041 0.61636 0.08512 60 2 Herpestidae brachyurus 3 9 2 2 Herpestes 0.72307 0.50759 0.63488 0.09126 70 3 Herpestidae edwardsii 2 8 9 6 Herpestes fuscus 0.69979 0.49810 0.61004 0.07851 60 2 Herpestidae 1 8 7 8 Herpestes 0.69149 0.49819 0.72273 0.09655 60 2 Herpestidae ichneumon 5 5 8 7 Herpestes 0.70009 0.49491 0.62057 0.08465 60 2 Herpestidae javanicus 6 1 9 2 Herpestes naso 0.71421 0.52738 0.62267 0.08236 60 2 Herpestidae 4 2 2 Herpestes urva 0.70438 0.56240 0.64142 0.09024 60 2 Herpestidae 2 7 8 1 Herpestes 0.65284 0.54782 0.58815 0.06880 60 2 Herpestidae 264 vitticollis 9 7 1 3 Hyaena hyaena 0.68306 0.55372 0.78850 0.10393 80 3 Hyaenidae 4 1 2 4 Ichneumia 0.72193 0.50991 0.64076 0.07146 20 1 Herpestidae albicauda 7 9 4 3 Leopardus 0.69735 0.50534 1 0.15216 100 3 Felidae colocolo 7 3 Leopardus 0.71350 0.44556 1 0.15004 100 3 Felidae geoffroyi 2 1 6 Leopardus guigna 0.79109 0.45977 1 0.12585 100 3 Felidae 7 3 Leopardus 0.75554 0.44825 1 0.13775 100 3 Felidae pardalis 9 4 Leopardus tigrinus 0.74622 0.40528 1 0.13745 100 3 Felidae 8 3 Leopardus wiedii 0.71439 0.43518 1 0.14158 80 3 Felidae 6 2 6 Liberiictis kuhni 0.62619 0.47969 0.52670 0.04932 20 1 Herpestidae 1 1 4 7 Lycaon pictus 0.64928 0.49252 0.71066 0.11969 100 3 Canidae 8 3 1 8 Lynx canadensis 0.78898 0.44634 1 0.15941 100 3 Felidae 1 2 2 265 Lynx lynx 0.79083 0.50001 1 0.14758 100 3 Felidae 4 2 3 Lynx pardinus 0.81483 0.48392 1 0.15210 100 3 Felidae 8 3 Lynx rufus 0.80711 0.49118 1 0.14546 100 3 Felidae 5 2 9 Mungos mungo 0.65398 0.65529 0.75450 0.07647 20 1 Herpestidae 3 9 2 Mungotictis 0.73336 0.54262 0.60088 0.08917 20 1 Eupleridae decemlineata 1 6 4 3 Nandinia binotata 0.69825 0.65049 0.66409 0.06867 10 1 Nandiniidae 2 6 7 4 Nasua narica 0.56090 0.64758 0.54355 0.05286 20 1 Procyonidae 6 3 9 4 Nasua nasua 0.52934 0.57890 0.50048 0.04732 20 1 Procyonidae 1 2 8 Nasuella olivacea 0.43911 0.56308 0.45791 0.03555 20 1 Procyonidae 6 4 4 Neofelis nebulosa 0.73510 0.46239 1 0.13438 100 3 Felidae 8 9 2 Nyctereutes 0.67459 0.44269 0.62048 0.09377 40 1 Canidae procyonoides 7 3 2 Otocolobus manul 0.76252 0.46755 1 0.16595 100 3 Felidae 266 7 8 1 Paguma larvata 0.67564 0.75779 0.56497 0.06633 60 2 Viverridae 3 4 7 1 Panthera uncia 0.82155 0.48066 1 0.14734 100 3 Felidae 9 1 1 Panthera leo 0.69909 0.47720 1 0.12219 100 3 Felidae 2 8 2 Panthera onca 0.80370 0.45631 1 0.12654 100 3 Felidae 3 4 Panthera pardus 0.77458 0.49165 1 0.13171 100 3 Felidae 6 5 Panthera tigris 0.74551 0.47559 1 0.12687 100 3 Felidae 9 3 5 Paracynictis 0.65567 0.50331 0.68850 0.07914 40 1 Herpestidae selousi 9 2 1 7 Paradoxurus 0.71923 0.6925 0.59609 0.07465 40 1 Viverridae hermaphroditus 6 2 3 Paradoxurus 0.69575 0.65321 0.60528 0.07331 40 1 Viverridae zeylonensis 4 2 Parahyaena 0.76328 0.60451 0.82032 0.11484 80 3 Hyaenidae brunnea 7 8 5 1 Pardofelis badia 0.76913 0.45521 1 0.15492 100 3 Felidae 5 2 1 267 Pardofelis 0.67560 0.47841 1 0.16031 100 3 Felidae marmorata 7 1 3 Pardofelis 0.77640 0.41433 1 0.14784 100 3 Felidae temminckii 4 2 Potos flavus 0.70417 0.72558 0.44098 0.04392 10 1 Procyonidae 8 6 2 Prionailurus 0.75377 0.43890 1 0.13446 100 3 Felidae bengalensis 2 9 Prionailurus 0.75193 0.34981 1 0.12842 100 3 Felidae planiceps 2 5 5 Prionailurus 0.77443 0.44923 1 0.14432 100 3 Felidae rubiginosus 3 6 9 Prionailurus 0.76886 0.46159 1 0.13566 80 3 Felidae viverrinus 3 8 1 Prionodon linsang 0.6281 0.35400 0.73947 0.10292 70 3 Prionodontidae 2 1 5 Prionodon 0.70536 0.39311 0.77266 0.10606 70 3 Prionodontidae pardicolor 5 9 1 3 Procyon 0.81627 0.75597 0.56880 0.07588 30 1 Procyonidae cancrivorus 9 4 5 5 Procyon lotor 0.73123 0.67924 0.57708 0.07438 20 1 Procyonidae 4 3 5 4 Pseudalopex 0.66431 0.40767 0.67306 0.09345 80 3 Canidae 268 culpaeus 6 5 5 Pseudalopex 0.60407 0.44854 0.62969 0.09501 60 2 Canidae fulvipes 2 8 3 1 Pseudalopex 0.66291 0.39103 0.64165 0.10007 60 2 Canidae griseus 3 9 3 1 Pseudalopex 0.63880 0.40692 0.63459 0.09649 50 2 Canidae gymnocercus 5 2 9 1 Pseudalopex 0.65726 0.47037 0.62018 0.08393 60 2 Canidae sechurae 2 3 2 3 Pseudalopex 0.71278 0.50764 0.63281 0.08457 60 2 Canidae vetulus 8 5 3 3 Puma concolor 0.82354 0.48746 1 0.13520 100 3 Felidae 3 8 3 Puma yaguarondi 0.77382 0.47649 1 0.14714 80 3 Felidae 5 4 5 Rhynchogale 0.87628 0.58857 0.53478 0.05118 30 1 Herpestidae melleri 4 5 4 Salanoia concolor 0.66319 0.51300 0.57352 0.08124 30 1 Eupleridae 6 6 5 Speothos venaticus 0.70441 0.52094 0.74695 0.10685 100 3 Canidae 9 3 1 Suricata suricatta 0.70286 0.63987 0.70184 0.08038 20 1 Herpestidae 3 9 2 5 269 Urocyon 0.65778 0.47745 0.63150 0.08830 30 1 Canidae cinereoargenteus 2 9 9 7 Urocyon littoralis 0.65270 0.46811 0.62808 0.09530 30 1 Canidae 1 1 6 4 Viverra megaspila 0.66866 0.42413 0.60889 0.08615 70 3 Viverridae 3 8 2 4 Viverra 0.67364 0.45058 0.59888 0.08093 70 3 Viverridae tangalunga 8 1 6 3 Viverra zibetha 0.70795 0.49263 0.56774 0.08149 80 3 Viverridae 7 6 2 Viverricula indica 0.78034 0.41796 0.57179 0.07149 60 2 Viverridae 9 1 7 8 Vulpes chama 0.68879 0.45126 0.62393 0.08838 70 3 Canidae 9 8 1 2 Vulpes ferrilata 0.69365 0.45923 0.69462 0.10508 100 3 Canidae 1 6 2 5 Vulpes lagopus 0.63524 0.44493 0.70144 0.10541 90 3 Canidae 4 9 8 Vulpes rueppelli 0.63766 0.42088 0.65720 0.09894 60 2 Canidae 9 3 5 2 Vulpes velox 0.66293 0.37019 0.67880 0.09655 90 3 Canidae 4 7 1 Vulpes vulpes 0.64426 0.40331 0.67798 0.09955 70 3 Canidae 270 5 8 3 3 Vulpes zerda 0.67714 0.39484 0.60908 0.09014 50 2 Canidae 8 7 8 Table 2. Extinct taxa with associated averages of ecomorphological metrics. Species C1 P4S RBL M1BS Adcrocuta eximia 0.747188 0.588566 0.835985 0.112602 Afrosmilus africanus 0.406736 0.489209 NA NA Afrosmilus hispanicus 0.513333 NA 0.983271 NA Afrosmilus turkanae NA NA 0.969421 NA Albanosmilus jourdani 0.373684 0.433526 1 0.167953 Albanosmilus whitfordi 0.386364 0.444976 1 0.171829 Allohyaena kadici NA 0.571292 0.830248 NA Amphimachairodus 0.396489 0.461898 0.935223 0.113028 giganteus Barbourofelis fricki 0.157042 0.497861 1 0.133913 Barbourofelis loveorum NA 0.427078 1 0.176374 Barbourofelis morrisi 0.355795 0.447192 1 0.167111 Belbus beaumonti NA 0.525 0.75261 0.118533 Chasmaporthetes lunensis 0.750759 0.492451 0.83533 0.10658 Chasmaporthetes 0.897561 0.399386 0.899157 0.126683 ossifragus 271 Crocuta crocuta spelaea 0.724907 0.618027 0.910248 0.134365 Dinaelurus crassus 0.709677 NA NA NA Dinailurictis bonali 0.563907 0.428571 0.812039 NA Dinictis felina 0.551178 0.455047 0.826374 0.120949 Dinocrocuta gigantea 0.842105 0.56474 0.860564 0.120973 Dinofelis diastemata 0.650263 0.470874 0.955863 0.14701 Eofelis edwardsii 0.666667 0.443872 0.820345 0.143968 Euboictis aliverensis NA 0.772059 NA NA Ginsburgsmilus 0.583333 0.59434 1 NA napakensis Herpestides antiquus 0.767521 0.458333 0.678161 0.073861 Homotherium ischyrus 0.366998 0.44666 0.966745 NA Homotherium latidens 0.416956 0.458315 1 0.1394 Homotherium serum 0.460004 0.432468 1 0.14193 Hoplophoneus adelos NA 0.33574 NA NA Hoplophoneus bidentatus 0.487805 0.424895 0.914269 0.145238 Hoplophoneus cerebralis 0.477814 0.500697 0.926823 0.151966 Hoplophoneus dakotensis 0.443933 0.53125 0.929063 0.123486 Hoplophoneus occidentalis 0.440957 0.491782 0.950486 0.125868 Hoplophoneus oharrai 0.465559 0.44174 0.907312 0.122276 Hoplophoneus primaevus 0.537219 0.49914 0.906273 0.147058 Hoplophoneus sicarius 0.240255 0.420154 0.955703 0.11689 Hoplophoneus 0.327681 0.436975 0.921053 0.146346 272 villebramarensis Ictitherium viverrinum 0.636882 0.510467 0.673976 0.089825 Izmirictis cani NA 0.514256 0.617248 NA Kanuites lewisae 0.729167 NA 0.606173 NA Kichechia zamanae 0.682927 0.619048 0.63871 NA Lycyaena chaeretis 0.756263 0.523179 0.769853 0.107619 Machairodus catocopis 0.415963 0.431696 0.904503 0.126069 Machairodus 0.363719 0.401679 0.958988 0.131272 coloradoensis Maofelis cantonensis 0.76506 NA NA NA MA-PHQ 348 0.725038 NA NA NA Megantereon cultridens 0.497118 0.462991 0.974003 0.134235 Metailurus major 0.57248 0.434636 0.904655 0.098354 Miracinonyx trumani 0.816667 0.48125 0.95892 0.125692 Nanosmilus kurteni 0.404018 0.474055 0.881467 0.155248 Nimravides pedionomus 0.572538 0.457513 0.938835 0.141835 Nimravus brachyops 0.56505 0.445368 0.830725 0.143222 Nimravus intermedius 0.610619 0.416926 0.848941 NA Oriensmilus liupanensis 0.445055 0.405316 1 0.191014 Pachycrocuta brevirostris 0.704545 0.651515 0.840678 NA Palaeoprionodon 0.658436 NA 0.783824 0.085664 lamandini Palhyaena reperta 0.74669 0.633072 0.778208 0.114786 273 Panthera atrox 0.733222 0.491752 1 0.115471 Panthera leo spelaea 0.719254 0.477838 1 0.116486 Percrocuta algeriensis NA 0.57193 0.831716 NA Percrocuta carnifex NA 0.573709 0.89735 NA Percrocuta tobieni NA 0.574713 0.838384 0.119752 Plioviverrops orbignyi 0.669246 0.472991 0.674444 0.076506 Pogonodon davisi 0.569334 0.599482 0.840437 0.119177 Pogonodon platycopis 0.556972 0.617188 0.838545 0.114899 Proailurus lemanensis 0.671711 0.458333 0.887452 0.122739 Promegantereon ogygia 0.59299 0.447216 0.930521 0.128398 Prosansanosmilus eggeri NA 0.522124 0.96875 NA Prosansanosmilus 0.467199 0.375 0.95241 0.114234 peregrinus Protictitherium crassum 0.591837 0.468531 0.419863 0.059764 Pseudaelurus validus 0.704031 0.491611 0.858555 0.119611 Quercylurus major NA NA NA NA Sansanosmilus palmidens 0.460606 0.412776 1 0.180789 Sivanasua viverroides 0.655518 0.680128 0.693564 NA Smilodon fatalis 0.4905 0.483591 1 0.136957 Smilodon populator 0.463153 0.451665 1 0.136113 Stenogale julieni NA 0.471074 0.831325 NA Tapocyon robustus 0.598083 0.47106 0.695688 0.065691 Thalassictis hyaenoides 0.741935 0.502005 0.737882 0.101481 274 Thalassictis wongii 0.601575 0.484962 0.727808 0.102206 Tongxinictis primordialis 0.683761 0.552486 NA NA Tungurictis spocki 0.454198 0.463883 0.707547 NA Xenosmilus hodsonae 0.525 0.510823 1 0.160784 Table 3. Dietary classification of Percrocuta species using the canonical variates analysis of Coca-Ortega and Pérez-Claros (2019). Diet category 3 = Fully developed bone cracker. Die Civet- cursorial_ Full_b Jackal/wolf Mongoose- Trans_b t like meat one -like like one Percrocuta 3 3.62E- 3.34E-09 0.9244 1.23E-05 2.82E-06 0.07552 carnifex 07 57 8 Percrocuta 3 7.68E- 0.045511 0.9544 1.96E-14 1.79E-30 1.08E- algeriensis 24 89 08 Percrocuta 3 0.0007 1.88E-10 0.9933 0.001611 8.33E-11 0.00426 tobieni 97 24 8 275 Table 4. Dietary classification of extinct non-durophagous and questionable sabertooth taxa based on dentognathic variables in canonical variates analysis. Diet: 1 = hypocarnivorous; 2 = mesocarnivorous; 3 = hypercarnivorous. diet P.hypo P.meso P.hyper P.saber Belbus beaumonti 3 0.070803 0.244877 0.531351 0.152969 Chasmaporthetes lunensis 3 0.058428 0.065953 0.87559 2.93E-05 Chasmaporthetes ossifragus 3 0.001783 0.008813 0.989404 3.78E-08 Dinaelurus crassus 3 0.312037 0.187957 0.497663 0.002343 Dinofelis diastemata 3 0.001254 0.012282 0.915439 0.071025 Euboictis aliverensis 1 0.922125 0.059729 0.010637 0.007509 Herpestides antiquus 1 0.426162 0.221477 0.352361 2.29E-07 Ictitherium viverrinum 2 0.340978 0.404937 0.253603 0.000481 Izmirictis cani 1 0.477878 0.406266 0.105721 0.010136 Kanuites lewisae 1 0.580569 0.326893 0.092534 4.08E-06 Kichechia zamanae 1 0.72625 0.238773 0.03492 5.75E-05 Lycyaena chaeretis 3 0.132535 0.193906 0.673541 1.86E-05 Maofelis cantonensis 3 0.276446 0.155494 0.567772 0.000289 MA-PHQ 348 3 0.302467 0.178764 0.517454 0.001314 Metailurus major 3 0.022531 0.01939 0.869325 0.088753 Miracinonyx trumani 3 0.004811 0.007994 0.987188 6.45E-06 Palaeoprionodon lamandini 3 0.122438 0.145578 0.72617 0.005814 Panthera atrox 3 0.008033 0.005211 0.986459 0.000297 Panthera leo spelaea 3 0.006048 0.00484 0.988571 0.000542 Plioviverrops orbignyi 1 0.434927 0.281416 0.283625 3.23E-05 Proailurus lemanensis 3 0.008773 0.030235 0.957263 0.003729 Promegantereon ogygia 3 0.002953 0.013493 0.760499 0.223055 Protictitherium crassum 2 0.392143 0.599815 0.008037 4.23E-06 Pseudaelurus validus 3 0.022985 0.056354 0.919908 0.000754 Sivanasua viverroides 1 0.795004 0.171522 0.032811 0.000662 276 Stenogale julieni 3 0.02149 0.050873 0.675502 0.252136 Tapocyon robustus 1 0.588692 0.180363 0.230443 0.000503 Thalassictis hyaenoides 3 0.162663 0.244622 0.5927 1.56E-05 Thalassictis wongii 3 0.133929 0.324479 0.529699 0.011893 Tungurictis spocki 3 0.155341 0.280326 0.513479 0.050853 Table 5. Ecological data used in Chapter III analyses. Diet refers to dietary category, 1 = hypocarnivorous; 2 = mesocarnivorous; 3 = hypercarnivorous; 4 = durophagous; 5 = sabertooth; RBL = relative blade length of the lower m1; g = body mass in grams. diet RBL log_cuberoot_mass g Acinonyx_jubatus 3 1 3.610417 50577 Adcrocuta_eximia 4 0.835985 3.616675 51535.48 Afrosmilus_africanus 5 NA NA NA Afrosmilus_hispanicus 5 0.983271 3.480055 34206.27 Afrosmilus_turkanae 5 0.969421 3.187158 14206.77 Albanosmilus_jourdani 5 1 3.85289 104680.8 Albanosmilus_whitfordi 5 1 3.8472 102908.8 Allohyaena_kadici 4 0.830248 3.588347 47336.72 Amphimachairodus_coloradoensis 5 0.935223 4.121573 234384.7 Amphimachairodus_giganteus 5 0.958988 4.150483 255620.7 Arctictis_binturong 2 0.5939 3.157543 12999 Atilax_paludinosus 2 0.725312 2.729563 3600 Barbourofelis_fricki 5 1 4.233938 328342.7 Barbourofelis_loveorum 5 1 4.179069 278509.7 Barbourofelis_morrisi 5 1 3.977234 152009.9 Bdeogale_nigripes 1 0.477484 2.624025 2623 Belbus_beaumonti 3 0.75261 3.461368 32341.43 Caracal_caracal 3 1 3.129886 11964 277 Catopuma_temminckii 3 1 2.984116 7726 Chasmaporthetes_lunensis 3 0.83533 3.454764 31706.94 Chasmaporthetes_ossifragus 3 0.899157 3.540839 41048.8 Civettictis_civetta 2 0.615087 3.132964 12075 Crocuta_crocuta 4 0.916777 3.685577 63369 Crocuta_crocuta_spelaea 4 0.910248 3.686889 63618.99 Crossarchus_obscurus 1 0.484472 2.41355 1395 Cryptoprocta_ferox 3 0.804789 3.053016 9500 Cynogale_bennettii 2 0.480858 2.784578 4246 Dinaelurus_crassus 3 NA 3.521503 38735.41 Dinailurictis_bonali 5 0.812039 3.954974 142190.5 Dinictis_felina 5 0.826374 3.589415 47488.67 Dinocrocuta_gigantea 4 0.860564 4.067907 199530 Dinofelis_diastemata 3 0.955863 3.814638 93331.72 Eofelis_edwardsii 5 0.820345 3.194692 14531.53 Euboictis_aliverensis 1 NA NA NA Eupleres_goudotii 1 0.585666 2.641357 2763 Felis_lybica 3 1 2.809308 4573 Fossa_fossana 1 0.592517 2.508187 1853 Galerella_sanguinea 1 0.745567 2.099036 543 Genetta_maculata 2 0.721206 2.525195 1950 Ginsburgsmilus_napakensis 5 1 3.260098 17681.83 Helogale_parvula 1 0.442353 1.879452 281 Hemigalus_derbyanus 1 0.437407 2.380151 1262 Herpestes_edwardsii 3 0.634889 2.391064 1304 Herpestides_antiquus 1 0.678161 2.836014 4954.46 Homotherium_ischyrus 5 1 4.102459 221322.4 Homotherium_latidens 5 1 4.141736 249000 Homotherium_serum 5 1 4.050205 189210.5 Hoplophoneus_adelos 5 NA 3.87149 110687.8 278 Hoplophoneus_bidentatus 5 0.914269 3.513513 37817.95 Hoplophoneus_cerebralis 5 0.926823 3.286854 19159.63 Hoplophoneus_dakotensis 5 0.929063 3.85158 104270.1 Hoplophoneus_occidentalis 5 0.950486 3.800259 89391.28 Hoplophoneus_oharrai 5 0.907312 3.668141 60139.55 Hoplophoneus_primaevus 5 0.906273 3.59462 48235.91 Hoplophoneus_sicarius 5 0.955703 3.634071 54296.46 Hoplophoneus_villebramarensis 5 0.921053 3.645912 56259.84 Hyaena_hyaena 4 0.788502 3.488367 35070 Hyaenictitherium_hyaenoides 3 0.737882 3.39525 26522.56 Hyaenotherium_wongii 3 0.727808 3.317363 20996.01 Ichneumia_albicauda 1 0.491228 2.732146 3628 Ictitherium_viverrinum 2 0.673976 3.245832 16941.06 Izmirictis_cani 1 0.617248 2.937566 6719.02 Kanuites_lewisae 1 0.606173 2.793518 4361.422 Kichechia_zamanae 1 0.63871 2.634546 2707.111 Leopardus_pardalis 3 1 3.127537 11880 Lycyaena_chaeretis 3 0.769853 3.426575 29135.83 Lynx_canadensis 3 1 3.059341 9682 MA_PHQ_348 3 NA 2.894135 5898.212 Machairodus_catocopis 5 0.904503 4.120902 233913.7 Maofelis_cantonensis 3 NA 3.576047 45621.75 Megantereon_cultridens 5 0.974003 3.688356 63899.53 Metailurus_major 3 0.904655 3.581296 46345.93 Miracinonyx_trumani 3 0.95892 3.535222 40362.89 Mungos_mungo 1 0.516129 2.379622 1260 Nandinia_binotata 1 0.664097 2.560366 2167 Nanosmilus_kurteni 5 0.881467 3.410991 27805.08 Neofelis_nebulosa 3 1 3.204044 14945 Nimravides_pedionomus 5 0.938835 3.877899 112836.6 279 Nimravus_brachyops 5 0.830725 3.896134 119181.3 Nimravus_intermedius 5 0.848941 3.759695 79148.88 Oriensmilus_liupanensis 5 0.936422 3.616048 51438.58 Otocolobus_manul 3 1 2.06081 6324.129 Pachycrocuta_brevirostris 4 0.840678 3.869412 110000 Paguma_larvata 1 0.564977 2.78879 4300 Palaeoprionodon_lamandini 3 0.783824 2.68948 3192.12 Palinhyaena_reperta 4 0.778208 3.203014 14898.91 Panthera_atrox 3 1 4.088114 212000 Panthera_leo 3 1 3.991429 158623 Panthera_leo_spelaea 3 1 4.141736 249000 Panthera_pardus 3 1 3.622214 52399 Paradoxurus_hermaphroditus 1 0.596092 2.690302 3200 Percrocuta_algeriensis 4 0.831716 3.627596 53251.82 Percrocuta_carnifex 4 0.89735 3.492805 35540.07 Percrocuta_tobieni 4 0.838384 3.407761 27536.88 Plioviverrops_orbignyi 1 0.674444 2.699183 3286.407 Pogonodon_davisi 5 0.840437 3.84183 101264.3 Pogonodon_platycopis 5 0.838545 3.905489 122573.8 Poiana_richardsonii 1 0.779359 2.115212 570 Prionailurus_planiceps 3 1 2.723301 3533 Prionodon_linsang 3 0.739471 2.175986 684 Proailurus_lemanensis 3 0.887452 3.271761 18311.48 Profelis_aurata 3 1 3.110174 11277 Promegantereon_ogygia 3 0.930521 3.585968 47000 Prosansanosmilus_eggeri 5 0.96875 3.198023 14677.45 Prosansanosmilus_peregrinus 5 0.95241 3.508625 37267.45 Proteles_cristata 1 NA 3.001474 8139 Protictitherium_crassum 2 0.419863 2.93761 6719.9 Pseudaelurus_validus 3 0.858555 3.483199 34530.5 280 Puma_concolor 3 1 3.631962 53954 Quercylurus_major 5 NA NA NA Rhynchogale_melleri 1 0.534785 2.57141 2240 Salanoia_concolor 1 0.730315 2.188891 711 Sansanosmilus_palmidens 5 1 3.593764 48112.23 Sivanasua_viverroides 1 0.693564 2.94273 6823.919 Smilodon_fatalis 5 1 4.098942 219000 Smilodon_populator 5 1 4.197112 294000 Stenogale_julieni 3 0.831325 2.67532 3059.356 Suricata_suricata 1 0.701842 2.197225 729 Tapocyon_robustus 1 0.695688 3.030226 8872.208 Tongxinictis_primordialis 4 NA 3.125784 11817.68 Tungurictis_spocki 3 0.707547 2.798452 4426.461 Viverra_tangalunga 1 0.598886 2.96744 7349 Viverricula_indica 1 0.571797 2.659551 2918 Xenosmilus_hodsonae 5 1 4.201001 297450.9 Table 6. Standard error of the mean in continuous variables listed in Table 3. RBL log_cuberoot_mass Acinonyx_jubatus 0 0.0345 Adcrocuta_eximia 0.0345 0.0345 Afrosmilus_africanus NA NA Afrosmilus_hispanicus 0.0345 0.0345 Afrosmilus_turkanae 0.0345 0.0345 Albanosmilus_jourdani 0 0.0345 Albanosmilus_whitfordi 0 0.0345 Allohyaena_kadici 0.0345 0.0345 Amphimachairodus_coloradoensis 0.0345 0.0345 Amphimachairodus_giganteus 0.0345 0.0345 281 Arctictis_binturong 0.013861 0.0345 Atilax_paludinosus 0.009469 0.0345 Barbourofelis_fricki 0 0.0345 Barbourofelis_loveorum 0 0.0345 Barbourofelis_morrisi 0 0.0345 Bdeogale_nigripes 0.025931 0.0345 Belbus_beaumonti 0.0345 0.0345 Caracal_caracal 0 0.0345 Catopuma_temminckii 0 0.0345 Chasmaporthetes_lunensis 0.0345 0.0345 Chasmaporthetes_ossifragus 0.0345 0.0345 Civettictis_civetta 0.010171 0.0345 Crocuta_crocuta 0.011597 0.0345 Crocuta_crocuta_spelaea 0.0345 0.0345 Crossarchus_obscurus 0.012224 0.0345 Cryptoprocta_ferox 0.011597 0.0345 Cynogale_bennettii 0.012966 0.0345 Dinaelurus_crassus NA 0.0345 Dinailurictis_bonali 0.0345 0.0345 Dinictis_felina 0.004529 0.0345 Dinocrocuta_gigantea 0.005321 0.0345 Dinofelis_diastemata 0.0345 0.0345 Eofelis_edwardsii 0.0345 0.0345 Euboictis_aliverensis NA NA Eupleres_goudotii 0.013861 0.0345 Felis_lybica 0 0.0345 Fossa_fossana 0.010586 0.0345 Galerella_sanguinea 0.009469 0.0345 Genetta_maculata 0.007486 0.0345 Ginsburgsmilus_napakensis 0 0.0345 282 Helogale_parvula 0.014971 0.0345 Hemigalus_derbyanus 0.010171 0.0345 Herpestes_edwardsii 0.025931 0.0345 Herpestides_antiquus 0.0345 0.0345 Homotherium_ischyrus 0 0.0345 Homotherium_latidens 0 0.0345 Homotherium_serum 0 0.0345 Hoplophoneus_adelos NA 0.0345 Hoplophoneus_bidentatus 0.0345 0.0345 Hoplophoneus_cerebralis 0.013624 0.0345 Hoplophoneus_dakotensis 0.0345 0.0345 Hoplophoneus_occidentalis 0.028924 0.0345 Hoplophoneus_oharrai 0.0345 0.0345 Hoplophoneus_primaevus 0.00555 0.0345 Hoplophoneus_sicarius 0.0345 0.0345 Hoplophoneus_villebramarensis 0.0345 0.0345 Hyaena_hyaena 0.011597 0.0345 Hyaenictitherium_hyaenoides 0.0345 0.0345 Hyaenotherium_wongii 0.0345 0.0345 Ichneumia_albicauda 0.009469 0.0345 Ictitherium_viverrinum 0.0345 0.0345 Izmirictis_cani 0.0345 0.0345 Kanuites_lewisae 0.0345 0.0345 Kichechia_zamanae 0.0345 0.0345 Leopardus_pardalis 0 0.0345 Lycyaena_chaeretis 0.0345 0.0345 Lynx_canadensis 0 0.0345 MA_PHQ_348 NA 0.0345 Machairodus_catocopis 0.0345 0.0345 Maofelis_cantonensis NA 0.0345 283 Megantereon_cultridens 0.0345 0.0345 Metailurus_major 0.0345 0.0345 Miracinonyx_trumani 0.0345 0.0345 Mungos_mungo 0.009469 0.0345 Nandinia_binotata 0.010586 0.0345 Nanosmilus_kurteni 0.0345 0.0345 Neofelis_nebulosa 0 0.0345 Nimravides_pedionomus 0.0345 0.0345 Nimravus_brachyops 0.008036 0.0345 Nimravus_intermedius 0.0345 0.0345 Oriensmilus_liupanensis 0.0345 0.0345 Otocolobus_manul 0 0.0345 Pachycrocuta_brevirostris 0.0345 0.0345 Paguma_larvata 0.010171 0.0345 Palaeoprionodon_lamandini 0.0345 0.0345 Palinhyaena_reperta 0.0345 0.0345 Panthera_atrox 0 0.0345 Panthera_leo 0 0.0345 Panthera_leo_spelaea 0 0.0345 Panthera_pardus 0 0.0345 Paradoxurus_hermaphroditus 0.014971 0.0345 Percrocuta_algeriensis 0.0345 0.0345 Percrocuta_carnifex 0.008929 0.0345 Percrocuta_tobieni 0.0345 0.0345 Plioviverrops_orbignyi 0.0345 0.0345 Pogonodon_davisi 0.011505 0.0345 Pogonodon_platycopis 0.0345 0.0345 Poiana_richardsonii 0.0345 0.0345 Prionailurus_planiceps 0 0.0345 Prionodon_linsang 0.014971 0.0345 284 Proailurus_lemanensis 0.0345 0.0345 Profelis_aurata 0 0.0345 Promegantereon_ogygia 0.0345 0.0345 Prosansanosmilus_eggeri 0.0345 0.0345 Prosansanosmilus_peregrinus 0.0345 0.0345 Proteles_cristata NA 0.0345 Protictitherium_crassum 0.0345 0.0345 Pseudaelurus_validus 0.0345 0.0345 Puma_concolor 0 0.0345 Quercylurus_major NA NA Rhynchogale_melleri 0.018336 0.0345 Salanoia_concolor 0.012224 0.0345 Sansanosmilus_palmidens 0 0.0345 Sivanasua_viverroides 0.0345 0.0345 Smilodon_fatalis 0 0.0345 Smilodon_populator 0 0.0345 Stenogale_julieni 0.0345 0.0345 Suricata_suricata 0.012966 0.0345 Tapocyon_robustus 0.0345 0.0345 Tongxinictis_primordialis NA 0.0345 Tungurictis_spocki 0.0345 0.0345 Viverra_tangalunga 0.0164 0.0345 Viverricula_indica 0.0164 0.0345 Xenosmilus_hodsonae 0 0.0345 285 References Coca-Ortega, C., and J. A. Pérez-Claros. 2019. Characterizing ecomorphological patterns in hyenids: A multivariate approach using postcanine dentition. PeerJ 6:e6238. Slater, G. J., and A. R. Friscia. 2019. Hierarchy in adaptive radiation: A case study using the Carnivora (Mammalia). Evolution 73:524–539. Wilman, H., B. J., S. J., de L. R. C., R. M., and J. W. 2014. EltonTraits 1.0 : Species-level foraging attributes of the world’s birds and mammals. Ecology 95:2027. 286 Appendix I Digitized Specimen List - Subsample Sample of crania used in the three-dimensional geometric morphometric analyses of Chapter IV. Unless otherwise specified, specimens were digitized via Microscribe with the methodology described in Chapter IV. Specimen number institutional codes are the same as in Appendix C, save CB = Bone Clones, Inc. 9200 Eton Ave. Chatsworth, CA 91311 USA. Clade Genus Species Specimen # Male Female Unknown Source (n) (n) sex (n) Eupleridae Cryptoprocta ferox FMNH 161793, 2 33950 Eupleridae Eupleres goudotii FMNH 30492; 2 MNHN CG 1962-2105 Eupleridae Fossa fossana FMNH 156648, 1 1 85196 Eupleridae Salanoia concolor MNHN CG 1 1 1866-233, 1962- 2111 Felidae Pseudaelurus validus FAM 61834, 3 61847; FM 287 61835 Felinae Acinonyx jubatus FM 45071; 1 2 FMNH 29635, 34589 Felinae Caracal caracal FMNH 135042, 1 1 32945 Felinae Catopuma temminckii MNHN 1939- 1 1 2152, 1941-293 Felinae Felis lybica FMNH 104579, 1 1 97861 Felinae Leopardus pardalis FMNH 85503, 2 1 88887, 93174 Felinae Lynx canadensis FMNH 138821, 1 1 145822 Felinae Otocolobus manul MNHN CG 1 1 2009-251, 2010- 646 Felinae Prionailurus planiceps MNHN CG 1 1873-228 Felinae Profelis aurata MNHN CG 1 1 1917-8, 1939- 687 288 Felinae Puma concolor FMNH 28334, 1 1 51875 Herpestidae Atilax paludinosus MNHN CG 1 1 1950-266, 1995- 426 Herpestidae Bdeogale nigripes FMNH 8383, 1 1 85974 Herpestidae Crossarchus obscurus FMNH 4374, 1 1 54410 Herpestidae Galerella sanguinea MNHN CG 2 2000-1054, 2000-1072 Herpestidae Helogale parvula MNHN CG 1 1 1969-72, 1987- 176 Herpestidae Herpestes edwardsii FMNH 83094, 1 1 97856 Herpestidae Ichneumia albicauda FMNH 73024, 1 1 85970 Herpestidae Mungos mungo FMNH 149365, 1 1 177229 Herpestidae Rhynchogale melleri MNHN CG 2 289 1962-1577, 1962-992 Herpestidae Suricata suricata FMNH 38348, 2 38349 Hyaenidae Adcrocuta eximia AMNH 22880; 5 China 32-L-255, 41L-339, C-L-6, 80-L613 Hyaenidae Chasmaporthetes lunensis MNCN 67100 1 CT scan (Tseng et al., 2011) Hyaenidae Crocuta crocuta FMNH 34582, 1 1 98952 Hyaenidae Crocuta crocuta spelaea MFN MB. Ma 2 44381, 49139 Hyaenidae Dinocrocuta gigantea IVPP V15649 1 CT scan (Tseng, 2009) Hyaenidae Hyaena hyaena FMNH 101982, 1 1 27008 Hyaenidae Hyaenotherium wongii Bx. 71-L665; 5 China (G) L-49, 42-l331, 89-L 746, 95-L789 290 Hyaenidae Ictitherium viverrinum China 51-L437; 2 FAM 19665 Hyaenidae Palinhyaena reperta FAM 129667, 2 144897 Hyaenidae Tungurictis spocki AMNH 26600 1 Machairodontinae Amphimachairodus giganteus FAM 50476 1 Machairodontinae Dinofelis diastemata FAM 50446 1 Machairodontinae Homotherium ischyrus AMNH 95297 1 Machairodontinae Homotherium latidens AMNH 104641 2 (cast); FAM 50462 Machairodontinae Homotherium serum TMM 933-3444 1 CT scan, DigiMorph Machairodontinae Megantereon cultridens AMNH 105446 1 (cast) Machairodontinae Metailurus major AMNH 131854 1 (cast) Machairodontinae Smilodon fatalis LACMHC 2001- 10 42, 2001-44, 2001-50, 2001- 58, 2001-59, 2001-63, 2001- 82, 2001-83; 291 LACMHC R- 10688, R-10864 Nimravidae Barbourofelis fricki AMNH 108193 1 (cast) Nimravidae Barbourofelis morrisi AMNH 61870, 2 79999 Nimravidae Dinictis felina AMNH 38805, 3 6937, 8777 Nimravidae Eusmilus cerebralis AMNH 6941; 2 Microscribe and JODA 7047 photogrammetry Nimravidae Eusmilus dakotensis CB 15 1 Photogrammetry, courtesy of D. Tamagnini Nimravidae Eusmilus sicarius CB 07 1 Photogrammetry, courtesy of D. Tamagnini Nimravidae Hoplophoneus occidentalis RAM 10356 1 CT scan, courtesy of A. Farke Nimravidae Hoplophoneus primaevus AMNH 38981, 4 38982, 9764; FAM 125662 292 Nimravidae Nanosmilus kurteni AMNH 1 140559.001 (cast) Nimravidae Nimravus brachyops AMNH 6930, 2 6933 Nimravidae Pogonodon davisi AMNH 102156 1 (cast) Nimravidae Pogonodon platycopis AMNH 6938 1 Pantherinae Neofelis nebulosa FMNH 75830, 1 1 75831 Pantherinae Panthera atrox AMNH 14397; 7 LACMHC 2900- 10, 2900-7, 2900-8, 2900-9; UCMP 14001, 20049 Pantherinae Panthera leo FMNH 20757, 3 23970 (cast), 35739 Pantherinae Panthera leo spelaea MFN MB. Ma. 3 48115.1, 50947, 50948 293 Pantherinae Panthera pardus MNHN 1962- 1 1 2884; MNHN CG 1996-521 Prionodontidae Prionodon linsang FMNH 8371, 1 1 88606 Stenoplesictidae Proailurus lemanensis AMNH 101931 1 (cast) Stenoplesictidae Nandinia binotata AMNH 2409 1 1 1 C.A; FMNH 55758, 73804 Viverridae Arctictis binturong AMNH C.A. 1 1 1 1182; FMNH 53744, 53747 Viverridae Civettictis Civetta FMNH 108174, 1 1 27278 Viverridae Cynogale bennettii MNHN A-2094; 2 MNHN CG 1962-170 Viverridae Genetta maculata FMNH 17525, 1 1 85984 Viverridae Hemigalus derbyanus AMNH 9 C.A; 1 1 1 FMNH 33465, 294 68717 Viverridae Paguma larvata MNHN CG 1 1 1962-1588, 1988-163 Viverridae Paradoxurus hermaphroditus FMNH 338000, 1 1 39345 Viverridae Poiana richardsonii MNHN CG 1 1976-389 Viverridae Viverra tangalunga FMNH 68704, 1 1 85116 Viverridae Viverricula indica MNHN CG 1 1932-3552 295 References Tseng, Z. J. 2009. Cranial function in a late Miocene Dinocrocuta gigantea (Mammalia: Carnivora) revealed by comparative finite element analysis. Biological Journal of the Linnean Society 96:51–67. Tseng, Z. J., M. Antón, and M. J. Salesa. 2011. The evolution of the bone-cracking model in carnivorans: cranial functional morphology of the Plio-Pleistocene cursorial hyaenid Chasmaporthetes lunensis (Mammalia: Carnivora). Paleobiology 37:140–1. 296 References Adams, D. C. 2014a. A method for assessing phylogenetic least squares models for shape and other high-dimensional multivariate data. Evolution 68:2675–2688. Adams, D. C. 2014b. A generalized K statistic for estimating phylogenetic signal from shape and other high-dimensional multivariate data. Systematic Biology 63:685–697. Adams, D. C. 2014c. Quantifying and comparing phylogenetic evolutionary rates for shape and other high-dimensional phenotypic data. Systematic Biology 63:166–177. Adams, D. C., and M. L. Collyer. 2015. Permutation tests for phylogenetic comparative analyses of high-dimensional shape data: What you shuffle matters. Evolution 69:823–829. Adams, D. C., M. L. Collyer, A. Kaliontzopoulou, and E. K. Baken. 2021. Geomorph: Software for geometric morphometric analyses. R package version 4.0. Https://Cran.r- Project.Org/Package=geomorph. Alroy, J. 1992. Conjunction among taxonomic distributions and the Miocene mammalian biochronology of the Great Plains. Paleobiology 18:326–343. Andersson, K. 2005. Were there pack-hunting canids in the Tertiary, and how can we know? Paleobiology 31:56–72. 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. Antón, A. M., M. J. Salesa, and G. Siliceo. 2013. Machairodont Adaptations and Affinities of the Holarctic Late Miocene Homotherin Machairodus (Mammalia, Carnivora, Felidae): The Case of Machairodus Catocopis Cope, 1887. Journal of Vertebrate Paleontology 33:1202– 1213. 297 Antón, M., M. J. Salesa, A. Galobart, and Z. J. Tseng. 2014. The Plio-Pleistocene scimitar- toothed felid genus Homotherium Fabrini, 1890 (Machairodontinae, Homotherini): Diversity, palaeogeography and taxonomic implications. Quaternary Science Reviews 96:259–268. Averianov, A., E. Obraztsova, I. Danilov, P. Skutschas, and J. Jin. 2016. First nimravid skull from Asia. Scientific Reports 6:1–8. Averyanov, A. O., and G. F. Baryshnikov. 1996. Origin and systematic position of aardwolf proteles cristatus (mammalia, carnivora). Zoologicheskii Zhurnal 75:1256–1258. Baken, E. K., M. L. Collyer, A. Kaliontzopoulou, and D. C. Adams. 2021. geomorph v4.0 and gmShiny: Enhanced analytics and a new graphical interface for a comprehensive morphometric experience. Methods in Ecology and Evolution 12:2355–2363. Bapst, D. W. 2012. Paleotree: an R Package for Paleontological and Phylogenetic Analyses of Evolution. Methods in Ecology and Evolution 3:803–807. Barnett, R., M. L. Z. Mendoza, A. E. R. Soares, S. Y. W. Ho, G. Zazula, N. Yamaguchi, B. Shapiro, I. V. Kirillova, G. Larson, and M. T. P. Gilbert. 2016. Mitogenomics of the extinct cave lion, Panthera spelaea (Goldfuss, 1810), resolve its position within the panthera cats. Open Quaternary 2:1–11. Barnett, R., B. Shapiro, I. Barnes, S. Y. Ho, J. Burger, N. Yamaguchi, T. F. Higham, H. T. Wheeler, W. Rosendahl, A. V Sher, M. Sotnikova, T. Kuznetsova, G. F. Baryshnikov, L. D. Martin, C. R. Harington, J. A. Burns, and A. Cooper. 2009. Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity. Molecular Ecology 18:1668–1677. 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 30:1–8. 298 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. Barrett, P. Z. 2016. Taxonomic and systematic revisions to the North American Nimravidae (Mammalia, Carnivora). PeerJ 4:e1658. Barrett, P. Z. 2021. The largest hoplophonine and a complex new hypothesis of nimravid evolution. Scientific Reports 11:1–9. Barrett, P. Z., S. S. B. Hopkins, and S. A. Price. 2021. How many sabretooths? Reevaluating the number of carnivoran sabretooth lineages with total-evidence Bayesian techniques and a novel origin of the Miocene Nimravidae. Journal of Vertebrate Paleontology e1923523. Baryshnikov, G. F., and A. O. Averianov. 1993. Deciduous teeth of carnivorous mammals (Order Carnivora) Part V. Families Protelidae and Hyaenidae. Russian Academy of Sciences Proceedings of the Zoological Institute St. Petersburg 263:46–84. Beaulieu, J. M., D. C. Jhwueng, C. Boettiger, and B. C. O’Meara. 2012. Modeling stabilizing selection: Expanding the Ornstein-Uhlenbeck model of adaptive evolution. Evolution 66:2369–2383. Bell, M. A., and G. T. Lloyd. 2014. Strap: Stratigraphic Tree Analysis for Palaeontology. R package version 1.4. https://CRAN.R-project.org/package=strap. Bergsten, J. 2005. A review of long-branch attraction. Cladistics 21:163–193. Blomberg, S. P., T. Garland, and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717–745. Bobe, R., A. K. Behrensmeyer, and R. E. Chapman. 2002. Faunal change, environmental variability and late Pliocene hominin evolution. Journal of Human Evolution 42:475–497. 299 Böhme, M., M. Aiglstorfer, P. O. Antoine, E. Appel, P. Havlik, G. Métais, L. T. Phuc, S. Schneider, F. Setzer, R. Tappert, D. N. Tran, D. Uhl, and J. Prieto. 2013. Na Duong (northern Vietnam) - An exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana Reihe A: Mitteilungen Der Bayerischen Staatssammlung Fur Palaontologie Und Geologie 53:121–167. Bonferroni, C. E. 1936. Teoria statistica delle classi e calcolo delle probabilità. Pubblicazioni Del R Istituto Superiore Di Scienze Economiche e Commerciali Di Firenze 8:1–62. de Bonis, L., S. Peigné, H. T. Mackaye, A. Likius, P. Vignaud, and M. Brunet. 2018. New sabre toothed felidae (Carnivora, mammalia) in the hominid-bearing sites of toros menalla (late miocene, Chad). Geodiversitas 40:69–86. Bookstein, F. L., P. Gunz, P. Mittercker, H. Prossinger, K. Schæfer, and H. Seidler. 2003. Cranial integration in Homo: Singular warps analysis of the midsagittal plane in ontogeny and evolution. Journal of Human Evolution 44:167–187. Borths, M. R., P. A. Holroyd, and E. R. Seiffert. 2016. Hyainailourine and teratodontine cranial material from the late Eocene of Egypt and the application of parsimony and Bayesian methods to the phylogeny and biogeography of Hyaenodonta (Placentalia, Mammalia). PeerJ 4:e2639. Bouckaert, R., J. Heled, D. Kühnert, T. Vaughan, C. H. Wu, D. Xie, M. A. Suchard, A. Rambaut, and A. J. Drummond. 2014. BEAST 2: A Software Platform for Bayesian Evolutionary Analysis. PLoS Computational Biology 10:e1003537. Boyer, D. M. 2008. Relief index of second mandibular molars is a correlate of diet among prosimian primates and other euarchontan mammals. Journal of Human Evolution 55:1118– 1137. Bright, J. A., J. Marugán-Lobón, S. N. Cobb, and E. J. Rayfield. 2016. The shapes of bird beaks are highly controlled by nondietary factors. Proceedings of the National Academy of Sciences of the United States of America 113:5352–5357. Brocklehurst, N. 2019. Morphological evolution in therocephalians breaks the hypercarnivore ratchet. Proceedings of the Royal Society B: Biological Sciences 286. 300 Browne, I., and R. E. Reynolds. 2015. Nimravides marshi (Felidae): an early scimitar-cat from the upper Barstovian section of the Mud Hills, Mojave Desert, California. Mojave Miocene 136–144. 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. Cantalapiedra, J. L., R. G. FitzJohn, T. S. Kuhn, M. H. Fernández, D. DeMiguel, B. Azanza, J. Morales, and A. Mooers. 2013. Dietary innovations spurred the diversification of ruminants during the Caenozoic. Proceedings of the Royal Society B: Biological Sciences 281. Carbone, C., A. Teacher, and J. M. Rowcliffe. 2007. The costs of carnivory. PLoS Biology 5:0363–0368. Carbone, C., G. M. Mace, S. C. Roberts, and D. W. Macdonald. 1999. Energetic constraints on the diet of terrestrial carnivores. Nature 402:286–288. Chamoli, U., and S. Wroe. 2011. Allometry in the distribution of material properties and geometry of the felid skull: Why larger species may need to change and how they may achieve it. Journal of Theoretical Biology 283:217–226. Chen, G. F., and N. Schmidt-Kittler. 1983. The deciduous dentition of Percrocuta Kretzoi and the diphyletic origin of the hyaenas (Carnivora, Mammalia). Paläontologische Zeitschrift 57:159–169. Chow, M. M. 1958. A record of the earliest sabre-toothed cats from the Eocene of Lushih, Honan. Science Record 2:347–349. Christensen, H. B. 2014. Similar associations of tooth microwear and morphology indicate similar diet across marsupial and placental mammals. PLoS ONE 9:e102789. Christiansen, P. 2006. Sabertooth characters in the clouded leopard (Neofelis nebulosa Griffiths 1821). Journal of Morphology 267:1186–1198. 301 Christiansen, P. 2008a. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics 24:977–992. Christiansen, P. 2008b. Evolution of skull and mandible shape in cats (Carnivora: Felidae). PLoS ONE 3:e2807. Christiansen, P. 2008c. Evolutionary convergence of primitive sabertooth craniomandibular morphology: the clouded leopard (Neofelis nebulosa) and Paramachairodus ogygia compared. Journal of Mammalian Evolution 15:155–179. Christiansen, P. 2012. The making of a monster: Postnatal ontogenetic changes in craniomandibular shape in the great sabercat Smilodon. PLoS ONE 7:19–21. Christiansen, P. 2013. Phylogeny of the sabertoothed felids (Carnivora: Felidae: Machairodontinae). Cladistics 29:543–559. Christiansen, P., and J. M. Harris. 2005. Body size of Smilodon (Mammalia: Felidae). Journal of Morphology 266:369–384. Clavel, J., G. Escarguel, and G. Merceron. 2015. mvMORPH: An R package for fitting multivariate evolutionary models to morphometric data. Methods in Ecology and Evolution 6:1311–1319. Coca-Ortega, C., and J. A. Pérez-Claros. 2019. Characterizing ecomorphological patterns in hyenids: A multivariate approach using postcanine dentition. PeerJ 6:e6238. Collard, M., and P. O’Higgins. 2001. Ontogeny and homoplasy in the papionin monkey face. Evolution and Development 3:322–331. Collyer, M. L., and D. C. Adams. 2018. RRPP: An R package for fitting linear models to high- dimensional data using residual randomization. Methods in Ecology and Evolution 9:1772– 1779. 302 Collyer, M. L., and D. C. Adams. 2019. RRPP: linear model evaluation with randomized residuals in a permutation procedure. R package version 1.1.2. Available at https://cran.r- project.org/packagepRRPP. Collyer, M. L., D. J. Sekora, and D. C. Adams. 2015. A method for analysis of phenotypic change for phenotypes described by high-dimensional data. Heredity 115:357–365. Costa, E., M. Garcés, A. Sáez, L. Cabrera, and M. López-blanco. 2011. The age of the “Grande Coupure” mammal turnover : New constraints from the Eocene – Oligocene record of the Eastern Ebro Basin ( NE Spain ). Palaeogeography, Palaeoclimatology, Palaeoecology 301:97–107. Dashzeveg, D. 1996. Some Carnivorous Mammals from the Paleogene of the Eastern Gobi Desert , Mongolia , and the Application of Oligocene Carnivores to Stratigraphic Correlation. American Museum Novitates 3179:1–14. Desantis, L. R. G., B. W. Schubert, E. Schmitt-linville, P. S. Ungar, S. L. Donohue, and R. J. Haupt. 2015. Dental microwear textures of carnivorans from the La Brea tar pits, California, and potential extinction implications. Natural History Museum of Los Angeles County, Science Series 42:37–52. DeSantis, L. R. G., R. S. Feranec, M. Antón, and E. L. Lundelius. 2021. Dietary ecology of the scimitar-toothed cat Homotherium serum. Current Biology 31:2674-2681.e3. Ding, S., J. Zhen, Y. Zhang, and Y. Tong. 1977. The age and characteristic of the Liuniu and the Dongjun faunas, Bose Basin of Guangxi. Vertebrata Palasiatica 15:35–45. Dobzhansky, T. 1951. Genetics and the Origin of Species, 3rd ed. Columbia University Press, New York, 364 pp. Drake, A. G., and C. P. Klingenberg. 2008. The pace of morphological change: Historical transformation of skull shape in St Bernard dogs. Proceedings of the Royal Society B: Biological Sciences 275:71–76. 303 Ducrocq, S., Y. Chaimanee, V. Suteethorn, and J. J. Jaeger. 1995. Mammalian faunas and the ages of the continental Tertiary fossiliferous localities from Thailand. Journal of Southeast Asian Earth Sciences 12:65–78. Eimer, G. H. T. 1890. Organic Evolution as the Result of the Inheritance of Acquired Characters According to the Laws of Organic Growth. Macmillan, London, UK, 435 pp. Eldredge, N., and S. J. Gould. 1972. Punctuated equilibria: an alternative to phyletic gradualism; pp. 82–115 in T. J. M. Schopf (ed.), Moddels in Paleobiology. Freeman, Cooper and Co., San Francisco. Evans, A. R., G. P. Wilson, M. Fortelius, and J. Jernvall. 2007. High-level similarity of dentitions in carnivorans and rodents. Nature 445:78–81. Evans, R. A., and S. Pineda-Munoz. 2018. Inferring mammal dietary ecology from dental morphology; pp. 37–51 in D. A. Croft, D. F. Su, and S. W. Simpson (eds.), Methods in Paleoecology: Reconstructing Cenozoic terrestrial environments and ecological communities, 1st ed. Springer. Ewer, R. F. 1973. The Carnivores. Weidenfeld and Nicolson, London, England, 494 pp. Faith, J. T., and A. K. Behrensmeyer. 2013. Climate change and faunal turnover: testing the mechanics of the turnover-pulse hypothesis with South African fossil data. Paleobiology 39:609–627. Famoso, N. A., and J. D. Orcutt. 2022. First occurrences of Palaeogale von Meyer, 1846 in the Pacific Northwest, United States. Geodiversitas 41:427–436. 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. Feranec, R. S. 2004. Isotopic evidence of saber-tooth development, growth rate, and diet from the adult canine of Smilodon fatalis from Rancho La Brea. Palaeogeography, Palaeoclimatology, Palaeoecology 206:303–310. 304 Ferguson, J. W., and R. P. Atit. 2019. A tale of two cities: The genetic mechanisms governing calvarial bone development. Genesis 57:e23248. Figueirido, B., Z. J. Tseng, and A. Martín-Serra. 2013. Skull shape evolution in durophagous carnivorans. Evolution 67:1975–1993. Figueirido, B., A. Martín-Serra, Z. J. Tseng, and C. M. Janis. 2015. Habitat changes and changing predatory habits in North American fossil canids. Nature Communications 6. Figueirido, B., N. MacLeod, J. Krieger, M. De Renzi, J. A. Pérez-Claros, and P. Palmqvist. 2011. Constraint and adaptation in the evolution of carnivoran skull shape. Paleobiology 37:490– 518. 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. Forasiepi, A. M., and M. R. Sánchez-Villagra. 2014. Heterochrony, dental ontogenetic diversity, and the circumvention of constraints in marsupial mammals and extinct relatives. Paleobiology 40:222–237. Friscia, A. R., B. Van Valkenburgh, and A. R. Biknevicius. 2007. An ecomorphological analysis of extant small carnivorans. Journal of Zoology 272:82–100. Friscia, A. R., M. Macharwas, S. Muteti, F. Ndiritu, and D. Tab Rasmussen. 2020. A Transitional Mammalian Carnivore Community from the Paleogene–Neogene Boundary in Northern Kenya. Journal of Vertebrate Paleontology 40. Frost, S. R. 2007. African Pliocene and Pleistocene cercopithecid evolution and global climate change; pp. 51–76 in R. Bobe, Z. Alemseged, and A. K. Behrensmeyer (eds.), Hominin environments in the East African Pliocene: an assessment of the faunal evidence. Springer, Dordrecht. Galton, F. 1869. Hereditary Genius: An Inquiry into Its Laws and Consequences. Macmillan, London, 390 pp. 305 Gaubert, P., and G. Veron. 2003. Exhaustive sample set among Viverridae reveals the sister- group of felids: The linsangs as a case of extreme morphological convergence within Feliformia. Proceedings of the Royal Society B: Biological Sciences 270:2523–2530. Gaubert, P., W. C. Wozencraft, P. Cordeiro-Estrela, and G. Veron. 2005. Mosaics of Convergences and Noise in Morphological Phylogenies: What’s in a Viverrid-Like Carnivoran? Systematic Biology 56:865–894. 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. Gehring, W. J. 1996. The master control gene for morphogenesis and evolution of the eye. Genes Cells 1:11–15. Goldschmidt, R. B. 1940. The Material Basis of Evolution. Yale University Press, New Haven, CT, 438 pp. Goloboff, P. A., and S. A. Catalano. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32:221–238. Goodall, C. 1991. Procrustes Methods in the Statistical Analysis of Shape. Journal of the Royal Statistical Society: Series B (Methodological) 53:285–321. Goswami, A. 2006. Morphological Integration in the Carnivoran Skull. Evolution 60:169–183. Goswami, A., and P. D. Polly. 2010a. The Influence of Modularity on Cranial Morphological Disparity in Carnivora and Primates (Mammalia). PLoS ONE 5:e9517. Goswami, A., and P. D. Polly. 2010b. Methods for Studying Morphological Integration and Modularity. The Paleontological Society Papers 16:213–243. Goswami, A., N. Milne, and S. Wroe. 2011. Biting through constraints: cranial morphology, disparity and convergence across living and fossil carnivorous mammals. Proceedings of the Royal Society B 278:1831–1839. 306 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. Goswami, A., W. J. Binder, J. Meachen, and F. R. O’Keefe. 2015. The fossil record of phenotypic integration and modularity: A deep-time perspective on developmental and evolutionary dynamics. Proceedings of the National Academy of Sciences of the United States of America 112:4891–4896. Gould, S. J. 1989. Wonderful Life: The Burgess Shale and the Nature of History. W. W. Norton & Company, New York City, 352 pp. Gould, S. J. 2002. The Structure of Evolutionary Theory. Belknap Press of Harvard University Press, Cambridge, Mass., 1433 pp. Gracia, S. F. 2015. Estudio de Protictitherium crassum del Cerro de los Batallones (Torrejón de Velasco, Madrid): aportación a la filogenia y evolución de la familia hyaenidae. Universidad Complutense de Madrid, 365 pp. Grohé, C., L. De Bonis, Y. Chaimanee, O. Chavasseau, M. Rugbumrung, C. Yamee, K. Suraprasit, C. Gibert, J. Surault, C. Blondel, and J. J. Jaeger. 2020. The Late Middle Miocene Mae Moh Basin of Northern Thailand: The Richest Neogene Assemblage of Carnivora from Southeast Asia and a Paleobiogeographic Analysis of Miocene Asian Carnivorans. American Museum Novitates 2020:1–57. Grossnickle, D. M. 2020. Feeding ecology has a stronger evolutionary influence on functional morphology than on body mass in mammals. Evolution 74:610–628. Guillerme, T., N. Cooper, S. L. Brusatte, K. E. Davis, A. L. Jackson, S. Gerber, A. Goswami, K. Healy, M. J. Hopkins, M. E. H. Jones, G. T. Lloyd, J. E. O’Reilly, A. Pate, M. N. Puttick, E. J. Rayfield, E. E. Saupe, E. Sherratt, G. J. Slater, V. Weisbecker, G. H. Thomas, and P. C. J. Donoghue. 2020. Disparities in the analysis of morphological disparity: Analysis of morphological disparity. Biology Letters 16. Haeckel, E. 1866. Generelle Morphologie Der Organismen: Allgemeine Grundzüge Der Organischen Formen-Wissenschaft, Mechanisch Begründet Durch Die von Charles Darwin Reformirte Decendenz-Theorie. Georg Reimer, Berlin, 574 pp. 307 Hans-Volker, K., E. Gröning, and C. Brauckmann. 2007. Comment on a fossil civet skull from the Lower Oligocene of the Weisselster Basin (Saxonia, Germany). Studia Geologica Salmanticensia 43:215–225. Hansen, T. F. 1997. Stabilizing selection and the comparative analysis of adaptation. Evolution 51:1341–1351. Hanson, C. B. 1996. Stratigraphy and vertebrate faunas of the Bridgerian-Duchesnean Clarno Formation, north-central Oregon; pp. 206–239 in D. R. Prothero and R. J. Emry (eds.), The Terrestrial Eocene-Oligocene Transition in North America. Cambridge University Press. Harjunmaa, E., K. Seidel, T. Häkkinen, E. Renvoisé, I. J. Corfe, A. Kallonen, Z. Q. Zhang, A. R. Evans, M. L. Mikkola, I. Salazar-Ciudad, O. D. Klein, and J. Jernvall. 2014. Replaying evolutionary transitions from the dental fossil record. Nature 512:44–48. Harmon, L. J., J. B. Losos, T. Jonathan Davies, R. G. Gillespie, J. L. Gittleman, W. Bryan Jennings, K. H. Kozak, M. A. McPeek, F. Moreno-Roark, T. J. Near, A. Purvis, R. E. Ricklefs, D. Schluter, J. A. Schulte, O. Seehausen, B. L. Sidlauskas, O. Torres-Carvajal, J. T. Weir, and A. T. Mooers. 2010. Early bursts of body size and shape evolution are rare in comparative data. Evolution 64:2385–2396. Hassanin, A., G. Veron, A. Ropiquet, B. J. van Vuuren, A. Lécu, S. M. Goodman, J. Haider, and T. T. Nguyen. 2021. Evolutionary history of Carnivora (Mammalia, Laurasiatheria) inferred from mitochondrial genomes. PLoS ONE 16:1–28. Holliday, J. A., and S. J. Steppan. 2004. Evolution of hypercarnivory: the effect of specialization on morphological and taxonomic diversity. Paleobiology 30:108–128. Hopkins, S. S. B., S. A. Price, and A. J. Chiono. 2021. Influence of phylogeny on the estimation of diet from dental morphology in the Carnivora. Paleobiology 1–16. Humphreys, A. M., and T. G. Barraclough. 2014. The evolutionary reality of higher taxa in mammals. Proceedings of the Royal Society B: Biological Sciences 281. 308 Hunt, G. 2007. The relative importance of directional change, random walks, and stasis in the evolution of fossil lineages. Proceedings of the National Academy of Sciences of the United States of America 104:18404–18408. Hunt Jr., R. M., and N. Solounias. 1991. Evolution of the aeluroid Carnivora: hyaenid affinities of the Miocene carnivoran Tungurictis spocki from Inner Mongolia. American Museum Novitates 1–25. Hunt, R. M. 1987. Evolution of the aeluroid Carnivora: significance of auditory structure in the nimravid cat Dinictis. American Museum Novitates 2886:1–74. 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. Hunt, R. M. 1991. Evolution of the aeluroid Carnivora: viverrid affinities of the Miocene carnivoran Herpestides. American Museum Novitates 3023:1–34. Hunt, R. M. 1996. Biogeography of the order Carnivora; pp. 485–541 in J. L. Gittleman (ed.), Carnivore Behavior, Ecology, and Evolution. Cornell University Press, New York. Hunt, R. M. 1998a. Evolution of the aeluroid Carnivora. Diversity of the earliest aeluroids from Eurasia (Quercy, Hsanda-Gol) and the origin of felids. American Museum Novitates 3252. Hunt, R. M. 1998b. Amphicyonidae; pp. 196–227 in C. M. Janis, K. Scott, and L. L. Jacobs (eds.), Evolution of Tertiary Mammals of North America, Volume 1: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals. Cambridge University Press, Cambridge. Hunt, R. M. 2001a. 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. Hunt, R. M. 2001b. Small Oligocene amphicyonids from North America (Paradaphoenus, Mammalia, Carnivora). American Museum Novitates 3331:1–20. 309 Hunter, J. P., and J. Jernvall. 1995. The hypocone as a key innovation in mammalian evolution. Proceedings of the National Academy of Sciences of the United States of America 92:10718–10722. Hutchinson, G. E. 1959. Homage to Santa Rosalia or why are there so many kinds of animals? American Naturalist 93:145–159. Hyatt, A. 1897. Cycle in the life of the individual (ontogeny) and in the evolution of its own group (phylogeny). Proceedings of the American Academy of Arts and Sciences 32:209– 224. Janis, C. M., and P. B. Wilhelm. 1993. Were there mammalian pursuit predators in the Tertiary? Dances with wolf avatars. Journal of Mammalian Evolution 1:103–125. Jernvall, J., and H. Jung. 2000. Genotype, phenotype, and developmental biology of molar tooth characters. American Journal of Physical Anthropology 113:171–190. Jones, K. E., J. Bielby, M. Cardillo, S. A. Fritz, J. O’Dell, C. D. L. Orme, K. Safi, W. Sechrest, E. H. Boakes, C. Carbone, C. Connolly, M. J. Cutts, J. K. Foster, R. Grenyer, M. Habib, C. A. Plaster, S. A. Price, E. A. Rigby, J. Rist, A. Teacher, O. R. P. Bininda-Emonds, J. L. Gittleman, G. M. Mace, and A. Purvis. 2009. PanTHERIA: a species-level database of life history, ecology, and geography of extant and recently extinct mammals. Ecology 90:2648– 2648. Jones, K. E., J. B. Smaers, and A. Goswami. 2015. Impact of the terrestrial-aquatic transition on disparity and rates of evolution in the carnivoran skull. BMC Evolutionary Biology 15. Kangas, A. T., A. R. Evans, I. Thesleff, and J. Jernvall. 2004. Nonindependence of mammalian dental characters. Nature 432:211–214. 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. Kavanagh, K. D., A. R. Evans, and J. Jernvall. 2007. Predicting evolutionary patterns of mammalian teeth from development. Nature 449:427–432. 310 Koepfli, K. P., S. M. Jenks, E. Eizirik, T. Zahirpour, B. Van Valkenburgh, and R. K. Wayne. 2006. Molecular systematics of the Hyaenidae: Relationships of a relictual lineage resolved by a molecular supermatrix. Molecular Phylogenetics and Evolution 38:603–620. Koufos, G. D., L. de Bonis, and S. Sen. 1995. Lophocyon paraskevaidisi, a new viverrid (Carnivora, Mammalia) from the middle Miocene of Chios island, Greece. Geobios 28:511– 523. Kretzoi, N. 1929. Materialien zur phylogenetischen Klassifikation der Aeluroïdeen . Congrès International de Zoologie 10 8:1293–1355. Lanfear, R., P. B. Frandsen, A. M. Wright, T. Senfeld, and B. Calcott. 2016. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses . Molecular Biology and Evolution 34:772–773. Lautenschlager, S., B. Figueirido, D. D. Cashmore, E. M. Bendel, and T. L. Stubbs. 2020. Morphological convergence obscures functional diversity in sabre-toothed carnivores: Sabre-tooth functional morphology. Proceedings of the Royal Society B: Biological Sciences 287. Law, C. J. 2021. Ecological drivers of carnivoran body shape evolution. American Naturalist 198:406–420. Law, C. J., G. J. Slater, and R. S. Mehta. 2018a. Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods. Systematic Biology 67:127–144. Law, C. J., E. Duran, N. Hung, E. Richards, I. Santillan, and R. S. Mehta. 2018b. Effects of diet on cranial morphology and biting ability in musteloid mammals. Journal of Evolutionary Biology 31:1918–1931. 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. 311 Li, Y., and N. Spassov. 2017. A new species of Paramachaerodus (Mammalia, Carnivora, Felidae) from the late Miocene of China and Bulgaria, and revision of Promegantereon Kretzoi, 1938 and Paramachaerodus Pilgrim, 1913. Paläontologische Zeitschrift 91:409– 426. 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. Liu, J., J. Liu, H. Zhang, J. Wagner, Q. Jiangzuo, Y. Song, S. Liu, Y. Wang, and C. Jin. 2021. The giant short-faced hyena Pachycrocuta brevirostris (Mammalia, Carnivora, Hyaenidae) from Northeast Asia: A reinterpretation of subspecies differentiation and intercontinental dispersal. Quaternary International. 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. Lucas, T., and A. Goswami. 2017. paleomorph: Geometric Morphometric Tools for Paleobiology. R package version 0.1.4. Https://CRAN.R- Project.Org/Package=paleomorph. Maestri, R., B. D. Patterson, R. Fornel, L. R. Monteiro, and T. R. O. de Freitas. 2016. Diet, bite force and skull morphology in the generalist rodent morphotype. Journal of Evolutionary Biology 29:2191–2204. Marcy, A. E., T. Guillerme, E. Sherratt, K. C. Rowe, M. J. Phillips, and V. Weisbecker. 2020. Australian rodents reveal conserved cranial evolutionary allometry across 10 million years of murid evolution. American Naturalist 196:755–768. Marroig, G., and J. M. Cheverud. 2005. Size as a line of least evolutionary resistance: Diet and adaptive morphological radiation in New World monkeys. Evolution 59:1128–1142. Martin, L. D. 1989. Fossil History of the Terrestrial Carnivora; pp. 536–568 in J. L. Gittleman (ed.), Carnivore Behavior, Ecology, and Evolution. Cornell University Press, Ithaca, New York. 312 Marugán-Lobón, J., S. M. Nebreda, G. Navalón, and R. B. J. Benson. 2022. Beyond the beak: Brain size and allometry in avian craniofacial evolution. Journal of Anatomy 240:197–209. Matthew, W. D. 1926. The evolution of the horse: a record and its interpretation. The Quarterly Review of Biology 1:139–185. McBratney-Owen, B., S. Iseki, S. D. Bamforth, B. R. Olsen, and G. M. Morriss-Kay. 2008. Development and Tissue Origins of the Mammalian Cranial Base. Developmental Biology 322:121–132. McHorse, B. K., A. A. Biewener, and S. E. Pierce. 2019. The Evolution of a Single Toe in Horses: Causes, Consequences, and the Way Forward. Integrative and Comparative Biology 59:638–655. McNab, B. K. 1986. The Influence of Food Habits on the Energetics of Eutherian Mammals. Ecological Monographs 56:1–19. Meachen-Samuels, J., and B. Van Valkenburgh. 2009. Craniodental indicators of prey size preference in the Felidae. Biological Journal of the Linnean Society 96:784–799. Meloro, C., M. Clauss, and P. Raia. 2015. Ecomorphology of Carnivora challenges convergent evolution. Organisms Diversity and Evolution 15:711–720. Michaud, M., G. Veron, and A. C. Fabre. 2020. Phenotypic integration in feliform carnivores: Covariation patterns and disparity in hypercarnivores versus generalists. Evolution 74:2681–2702. Michaud, M., G. Veron, S. Peigné, A. Blin, and A. C. Fabre. 2018. Are phenotypic disparity and rate of morphological evolution correlated with ecological diversity in Carnivora? Biological Journal of the Linnean Society 124:294–307. 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. 313 Mishina, Y., and T. N. Snider. 2014. Neural crest cell signaling pathways critical to cranial bone development and pathology. Experimental Cell Research 325:138–147. Mitteroecker, P., P. Gunz, M. Bernhard, K. Schaefer, and F. L. Bookstein. 2004. Comparison of cranial ontogenetic trajectories among great apes and humans. Journal of Human Evolution 46:679–698. Morales, J., and M. Pickford. 2011. A new paradoxurine carnivore from the Late Miocene Siwaliks of India and a review of the bunodont viverrids of Africa. Geobios 44:271–277. Morales, J., and M. Pickford. 2021. Taxonomic revision of the genus Leptoplesictis (Viverridae, Mammalia) with description of new fossils from Grillental VI (Namibia) and Moratilla 2 (Spain). Communications of the Geological Survey of Namibia 23:161–176. Morales, J., S. Mayda, A. Valenciano, D. DeMiguel, and T. Kaya. 2019. A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey. Journal of Systematic Palaeontology 17:1127–1138. Navalón, G., S. M. Nebreda, J. A. Bright, M. Fabbri, R. B. J. Benson, B. A. Bhullar, J. Marugán- Lobón, and E. J. Rayfield. 2021. Craniofacial development illuminates the evolution of nightbirds (Strisores). Proceedings of the Royal Society B: Biological Sciences 288. 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. O’Brien, C. L., M. Huber, E. Thomas, M. Pagani, J. R. Super, L. E. Elder, and P. M. Hull. 2020. The enigma of Oligocene climate and global surface temperature evolution. Proceedings of the National Academy of Sciences of the United States of America 117:25302–25309. O’Meara, B. C., C. Ané, M. J. Sanderson, and P. C. Wainwright. 2006. Testing for Different Rates of Continuous Trait Evolution Using Likelihood. Evolution 60:922. 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. 314 Olsen, A. M., and M. W. Westneat. 2015. StereoMorph: An R package for the collection of 3D landmarks and curves using a stereo camera set-up. Methods in Ecology and Evolution 6:351–356. Olson, E. C., and R. L. Miller. 1958. Morphological Integration. University of Chicago Press, Chicago, Il., 376 pp. Osborn, H. F. 1902. The Law of Adaptive Radiation. The American Naturalist 36:353–363. 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. Palmqvist, P., B. Martínez-Navarro, J. A. Pérez-Claros, V. Torregrosa, B. Figueirido, J. M. Jiménez-Arenas, M. Patrocinio Espigares, S. Ros-Montoya, and M. De Renzi. 2011. The giant hyena Pachycrocuta brevirostris: Modelling the bone-cracking behavior of an extinct carnivore. Quaternary International 243:61–79. 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. Peigné, S. 1999. Proailurus, l’ un des plus anciens Felidae (Carnivora) d’Eurasie: systématique et évolution. Bulletin de La Société d’histoire Naturelle de Toulouse 135:125–134. Peigné, S. 2003. Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. Zoologica Scripta 32:199–229. Peigné, S., and L. De Bonis. 1999. The genus Stenoplesictis Filhol (Mammalia, Carnivora) from the Oligocene deposits of the Phosphorites of Quercy, France. Journal of Vertebrate Paleontology 19:566–575. 315 Peigné, S., M. J. Salesa, M. Antón, and J. Morales. 2008. A new amphicyonine (Carnivora: Amphicyonidae) from the upper miocene of batallones-1 Madrid, Spain. Palaeontology 51:943–965. Peigné, S., Y. Chaimanee, J.-J. Jaeger, V. Suteethorn, and S. Ducrocq. 2000. Eocene nimravid carnivorans from Thailand. Journal of Vertebrate Paleontology 20:157–163. Pennell, M. W., J. M. Eastman, G. J. Slater, J. W. Brown, J. C. Uyeda, R. G. Fitzjohn, M. E. Alfaro, and L. J. Harmon. 2014. Geiger v2.0: An expanded suite of methods for fitting macroevolutionary models to phylogenetic trees. Bioinformatics 30:2216–2218. Pickford, M., and J. Morales. 1994. Biostratigraphy and palaeobiogeography of East Africa and the Iberian peninsula. Palaeogeography, Palaeoclimatology, Palaeoecology 112:297–322. Pineda-Munoz, S., A. R. Evans, and J. Alroy. 2016. The relationship between diet and body mass in terrestrial mammals. Paleobiology 42:659–669. 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. Pires, M. M., D. Silvestro, and T. B. Quental. 2017. Interactions within and between clades shaped the diversification of terrestrial carnivores. Evolution 71:1855–1864. Plessis, L. du. 2016. BDSKY Tools. R package version 0.0.1.0. Https://Github.Com/Laduplessis/Bdskytools/. Prevosti, F. J., A. M. Forasiepi, M. D. Ercoli, and G. F. Turazzini. 2012. Paleoecology of the mammalian carnivores (Metatheria, Sparassodonta) of the Santa Cruz Formation (late Early Miocene); pp. 173–193 in S. F. Vizcaíno, R. F. Kay, and M. S. Bargo (eds.), Early Miocene Paleobiology in Patagonia. Cambridge University Press, Cambridge, UK. Price, S. A., S. S. B. Hopkins, K. K. Smith, and V. L. Roth. 2012. Tempo of trophic evolution and its impact on mammalian diversification. Proceedings of the National Academy of Sciences of the United States of America 109:7008–7012. 316 Pyron, R. A. 2011. Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia. Systematic Biology 60:466–481. R Core Team. 2020. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/. Radinsky, L. B. 1981a. Evolution of skull shape in carnivores: 1. Representative modern carnivores. Biological Journal of the Linnean Society 15:369–388. Radinsky, L. B. 1981b. Evolution of the skull shape in carnivores 2. Additional modern carnivores. Biological Journal of the Linnean Society 16:337–355. Radinsky, L. B. 1982. Evolution of Skull Shape in Carnivores. 3. The Origin and Early Radiation of the Modern Carnivore Families. Paleobiology 8:177–195. 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 syy032. Randau, M., D. Sanfelice, and A. Goswami. 2019. Shifts in cranial integration associated with ecological specialization in pinnipeds (Mammalia, Carnivora). Royal Society Open Science 6. Raup, D. M. 1966. Geometric Analysis of Shell Coiling: General Problems. Journal of Paleontology 40:1178–1190. Raup, D. M., and S. J. ay Gould. 1974. Stochastic Simulation and Evolution of Morphology- Towards a Nomothetic Paleontology. Systematic Zoology 23:305–322. Renard, A., M. Lavoie, J. A. Pitt, and S. Larivière. 2015. Felis nigripes (Carnivora: Felidae). Mammalian Species 47:78–83. Reuter, D. M. 2021. Trophic structure evolution in Oregon Oligo-Miocene terrestrial communities. University of Oregon, 163 pp. 317 Revell, L. J. 2012. phytools: An R package for phylogenetic comparative biology (and other things). Methods Ecol. Evol. 3 217–223. 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. 2013a. 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. Robles, J. M., J. Madurell-Malapeira, J. Abella, C. Rotgers, R. Carmona, S. Almécija, J. Balaguer, and D. M. Alba. 2013b. New Pseudaelurus and Styriofelis remains (Carnivora: Felidae) from the Middle Miocene of Abocador de Can Mata (Vallès-Penedès Basin). Comptes Rendus - Palevol 12:101–113. Rohlf, F. J., and M. Corti. 2000. Use of two-block partial least-squares to study covariation in shape. Systematic Biology 49:740–753. 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. Rothwell, T. P. 2001. A Partial Skeleton of Pseudaelurus (Carnivora: Felidae) from the Nambé Member of the Tesuque Formation, Española Basin, New Mexico. American Museum Novitates 3342:1–31. Rothwell, T. P. 2003. Phylogenetic systematics of North American Pseudaelurus (Carnivora, Felidae). American Museum Novitates 3403. Rovinsky, D. S., A. R. Evans, and J. W. Adams. 2021. Functional ecological convergence between the thylacine and small prey-focused canids. BMC Ecology and Evolution 21. Sacco, T., and B. Van Valkenburgh. 2004. Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). Journal of Zoology 263:41–54. Sakamoto, M., and M. Ruta. 2012. Convergence and Divergence in the Evolution of Cat Skulls: Temporal and Spatial Patterns of Morphological Diversity. PLoS ONE 7:e39752. 318 Salesa, M. J., M. Antón, J. Morales, and S. Peigné. 2012. Systematics and phylogeny of the small felines (Carnivora, Felidae) from the Late Miocene of Europe: a new species of Felinae from the Vallesian of Batallones (MN 10, Madrid, Spain). Journal of Systematic Palaeontology 10:87–102. Salesa, M. J., M. Antón, A. Turner, L. Alcalá, P. Montoya, and J. Morales. 2010. Systematic revision of the late Miocene sabre-toothed felid Paramachaerodus in Spain. Palaeontolgy 53:1369–1391. Salles, L. O. 1992. Felid Phylogenetics: Extant Taxa and Skull Morphology (Felidae, Aeluroidea). American Museum Novitates 3047:67. Sanderson, M. J., and M. J. Donoghue. 1994. Shifts in Diversification Rate with the Origin of Angiosperms. Science 264:1590–1593. Santana, S. E., and E. Cheung. 2016. Go big or go fish: Morphological specializations in carnivorous bats. Proceedings of the Royal Society B: Biological Sciences 283:20160615. Schaefer, K., P. Mitteroecker, P. Gunz, M. Bernhard, and F. L. Bookstein. 2004. Craniofacial sexual dimorphism patterns and allometry among extant hominids. Annals of Anatomy 186:471–478. Schaeffer, B. 1948. The Origin of a Mammalian Ordinal Character. Evolution 2:164–175. Schlager, S. 2017. Morpho and Rvcg – Shape Analysis in R; pp. 217–256 in Z. G, L. S, and S. G (eds.), Statistical Shape and Deformation Analysis. Academic Press. Schluter, D. 2000. The Ecology of Adaptive Radiation. Oxford University Press, Oxford, 296 pp. Schmidt-Kittler, N. 1976. Raubtiere aus dem Jungtertiär Kleinasiens. Palaeontographica Abteilung A 155:1–131. Semenov, Y. 1989. Ictitheres and Morphologically Similar Hyaenas from the Neogene of the USSR. Naukova dumka, Kiev, 180 pp. 319 Semenov, Y. 2008. Taxonomical reappraisal of “ictitheres” (Mammalia, Carnivora) from the Late Miocene of Kenya. Comptes Rendus Palevol 7:529–539. Sen, S. 2013. Dispersal of African mammals in Eurasia during the Cenozoic: Ways and whys. Geobios 46:159–172. Shubin, N., C. Tabin, and S. Carroll. 1997. Fossils, genes and the evolution of animal limbs. Nature 388:639–648. Simons, E. A., S. R. Frost, K. Harvati, K. McNulty, and M. Singleton. 2020. Comparing Rates of Linage Diversification with Rates of Size and Shape Evolution in Catarrhine Crania. Evolutionary Biology 47:152–163. Simpson, G. G. 1944. Tempo and Mode in Evolution. Columbia University Press, New York, 237 pp. Simpson, G. G. 1953. Major Features of Evolution. Columbia University Press, New York, 434 pp. Singleton, M. 2002. Patterns of cranial shape variation in the Papionini (Primates: Cercopithecinae). Journal of Human Evolution 42:547–578. Slater, G. J. 2013. Phylogenetic evidence for a shift in the mode of mammalian body size evolution at the Cretaceous-Paleogene boundary. Methods in Ecology and Evolution 4:734– 744. Slater, G. J. 2015. Iterative adaptive radiations of fossil canids show no evidence for diversity- dependent trait evolution. Proceedings of the National Academy of Sciences of the United States of America 112:4897–4902. Slater, G. J., and B. Van Valkenburgh. 2008. Long in the tooth: evolution of sabertooth cat cranial shape. Paleobiology 34:403–419. Slater, G. J., and B. Van Valkenburgh. 2009. Allometry and performance: The evolution of skull form and function in felids. Journal of Evolutionary Biology 22:2278–2287. 320 Slater, G. J., and A. R. Friscia. 2019. Hierarchy in adaptive radiation: A case study using the Carnivora (Mammalia). Evolution 73:524–539. Solé, F. 2014. New carnivoraforms from the early Eocene of Europe and their bearing on the evolution of the Carnivoraformes. Palaeontology 57:963–978. Solé, F., and S. Ladevèze. 2017. Evolution of the hypercarnivorous dentition in mammals (Metatheria, Eutheria) and its bearing on the development of tribosphenic molars. Evolution and Development 19:56–68. 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. 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. Soledad Domingo, M., L. Domingo, C. Badgley, O. Sanisidro, and J. Morales. 2013. Resource partitioning among top predators in a Miocene food web. Proceedings of the Royal Society B: Biological Sciences 280. Spassov, N., and D. Geraads. 2015. A New Felid from the Late Miocene of the Balkans and the Contents of the Genus Metailurus Zdansky, 1924 (Carnivora, Felidae). Journal of Mammalian Evolution 22:45–56. Spaulding, M., and J. J. Flynn. 2009. Anatomy of the postcranial skeleton of “mlacis” uintensis (Mammalia: Carnivoramorpha). Journal of Vertebrate Paleontology 29:1212–1223. Spaulding, M., and J. J. Flynn. 2012. Phylogeny of the Carnivoramorpha: The impact of postcranial characters. Journal of Systematic Palaeontology 10:653–677. Stadler, T., D. Kühnert, 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. 321 Tomarev, S. I., P. Callaerts, L. Kos, R. Zinovieva, G. Haider, W. Gehring, and J. Piatigorsky. 1997. Squid Pax-6 and eye development. Proceedings of the National Academy of Sciences of the United States of America 94:2421–2426. 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. Tomiya, S., S. P. Zack, M. Spaulding, and J. J. Flynn. 2021. Carnivorous mammals from the middle Eocene Washakie Formation, Wyoming, USA, and their diversity trajectory in a post-warming world. Journal of Paleontology 95:1–115. 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. Tseng, J. Z., and J. J. Flynn. 2018. Structure-function covariation with nonfeeding ecological variables influences evolution of feeding specialization in carnivora. Science Advances 4. Tseng, Z., and X. Wang. 2011. Do convergent ecomorphs evolve through convergent morphological pathways? Cranial shape evolution in fossil hyaenids and borophagine canids (Carnivora, Mammalia). Paleobiology 37:470–489. Tseng, Z. J. 2009. Cranial function in a late Miocene Dinocrocuta gigantea (Mammalia: Carnivora) revealed by comparative finite element analysis. Biological Journal of the Linnean Society 96:51–67. Tseng, Z. J. 2012. Connecting Hunter-Schreger Band microstructure to enamel microwear features: New insights from durophagous carnivores. Acta Palaeontologica Polonica 57:473–484. Tseng, Z. J. 2013. Testing Adaptive Hypotheses of Convergence with Functional Landscapes: A Case Study of Bone-Cracking Hypercarnivores. PLoS ONE 8:e65305. 322 Tseng, Z. J., and X. Wang. 2007. The first record of the late Miocene Hyaenictitherium hyaenoides Zdansky (Carnivora: Hyaenidae) in Inner Mongolia and an evaluation of the genus. Journal of Vertebrate Paleontology 27:699–708. Tseng, Z. J., and W. J. Binder. 2010. Mandibular biomechanics of Crocuta crocuta, Canis lupus, and the late Miocene Dinocrocuta gigantea (Carnivora, Mammalia). Zoological Journal of the Linnean Society 158:683–696. Tsuboi, M., W. van der Bijl, B. T. Kopperud, J. Erritzøe, K. L. Voje, A. Kotrschal, K. E. Yopak, S. P. Collin, A. N. Iwaniuk, and N. Kolm. 2018. Breakdown of brain–body allometry and the encephalization of birds and mammals. Nature Ecology and Evolution 2:1492–1500. Turner, A., M. Antón, and L. Werdelin. 2008. Taxonomy and evolutionary patterns in the fossil Hyaenidae of Europe. Geobios 41:677–687. Uyeda, J. C., and L. J. Harmon. 2014. A novel Bayesian method for inferring and interpreting the dynamics of adaptive landscapes from phylogenetic comparative data. Systematic Biology 63:902–918. Vaidya, G., D. J. Lohman, and R. Meier. 2011. SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics 27:171–180. Van Valen, L. 1971. Adaptive zones and the orders of mammals. Evolution 25:420–428. Van Valkenburgh, B. 1990. Skeletal and dental predictors of body mass in carnivores; pp. 181– 205 in J. Damuth and B. J. MacFadden (eds.), Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press. Van Valkenburgh, B. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): Evolutionary interactions among sympatric predators. Paleobiology 17:340– 362. Van Valkenburgh, B. 1996. Feeding behavior in free-ranging, large African carnivores. Journal of Mammalogy 77:240–254. 323 Van Valkenburgh, B. 2007. Déjà vu: the evolution of feeding morphologies in the Carnivora. Integrative and Comparative Biology 47:147–163. Van Valkenburgh, B., and K.-P. Koepfli. 1993. Cranial and dental adaptations to predation in canids. Symposium Zoological Society of London 65:15–37. Van Valkenburgh, B., F. Grady, and B. Kurtén. 1990. The Plio-Pleistocene cheetah-like cat Miracinonyx inexpectatus of North America. Journal of Vertebrate Paleontology 10:434– 454. Van Valkenburgh, B., T. Sacco, and X. Wang. 2003. Pack hunting in Miocene borophagine dogs: Evidence from craniodental morphology and body size. Bulletin of the American Museum of Natural History 279:147–162. Van Valkenburgh, B., X. Wang, and J. Damuth. 2004. Cope’s rule, hypercarnivory, and extinction in North American canids. Science 306:101–104. Venables, W. N., and B. D. Ripley. 2002. Modern Applied Statistics with S, 4th Ed. Springer, New York, 495 pp. Vermeij, G. J. 1973. Adaptation, versatility, and evolution. Systematic Zoology 22:466–477. Viranta, S. 1996. European Miocene Amphicyonidae-taxonomy, systematics and ecology. Zoologica Fennica 204:1–61. Vrba, E. S. . 1992. Mammals as a Key to Evolutionary Theory. Journal of Mammalogy 73:1–28. Wang, B., M. Zelditch, and C. Badgley. 2021. Geometric morphometrics of mandibles for dietary differentiation of Bovidae (Mammalia: Artiodactyla). Current Zoology 1–13. Wang, J., and Z.-Q. Zhang. 2015. Phylogenetic analysis on Palaeogale (Palaeogalidae , Carnivora) based on specimens from Oligocene strata of Saint-Jacques , Nei Mongol. Vertebrata PalAsiatica 53:310–334. 324 Wang, X. 1993. Transformation from plantigrady to digitigrady: functional morphology of locomotion in Hesperocyon (Canidae: Carnivora). American Museum Novitates 1–23. Wang, X., Z. J. Tseng, W. Y. Wu, J. Ye, J. Meng, and S. Bi. 2020. A new species of Tungurictis Colbert, 1939 (Carnivora, hyaenidae) from the middle miocene of Junggar basin, Northwestern China and the early divergence of basal hyaenids in East Asia. Geodiversitas 42:29–45. Wang, Y., J. Meng, X. Ni, and C. Li. 2007. Major events of Paleogene mammal radiation in China. Geological Journal 42:415–430. Welsh, E. 2021. A new species of an enigmatic carnivore Palaeogale (Feliformia: Palaeogalidae) from Badlands National Park, South Dakota. Proceedings of the South Dakota Academy of Science 100:107–120. Werdelin, L. 1996. Carnivoran ecomorphology: a phylogenetic perspective; pp. 582–624 in J. L. Gittleman (ed.), Carnivore Behavior, Ecology, and Evolution. Cornell University Press, Ithaca, New York. Werdelin, L. 2019. Middle miocene carnivora and hyaenodonta from fort Ternan, Western Kenya. Geodiversitas 41:267–283. Werdelin, L. 2021. African Barbourofelinae (Mammalia, Nimravidae): a critical review. Historical Biology 00:1–9. Werdelin, L., and N. Solounias. 1991. The Hyaenidae : taxonomy, systematics and evolution. Fossils and Strata 30:104. Werdelin, L., and M. E. Lewis. 2001. A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132:147–258. Werdelin, L., and M. E. Lewis. 2005. Plio-Pleistocene Carnivora of eastern Africa: Species richness and turnover patterns. Zoological Journal of the Linnean Society 144:121–144. 325 Werdelin, L., and T. Flink. 2018. The Phylogenetic Context of Smilodon; pp. 14–29 in L. Werdelin, H. G. Mcdonald, and C. A. Shaw (eds.), Smilodon: The Iconic Sabertooth. Johns Hopkins University Press, Baltimore. 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. 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. Wesley-Hunt, G. D., R. Dehghani, and L. Werdelin. 2010. Comparative ecomorphology and biogeography of Herpestidae and Viverridae (Carnivora) in Africa and Asia; pp. 246–268 in A. Goswami and A. R. Friscia (eds.), Carnivoran Evolution: New Views on Phylogeny, Form, and Function. Cambridge University Press. Westbury, M. V., S. Hartmann, A. Barlow, M. Preick, B. Ridush, D. Nagel, T. Rathgeber, R. Ziegler, G. Baryshnikov, G. Sheng, A. Ludwig, I. Wiesel, L. Dalen, F. Bibi, L. Werdelin, R. Heller, and M. Hofreiter. 2020. Hyena paleogenomes reveal a complex evolutionary history of cross-continental gene flow between spotted and cave hyena. Science Advances 6:1–11. Wheeler, H. T., and G. T. Jefferson. 2009. Panthera atrox: body proportions, size, sexual dimorphism, and behavior of the cursorial lion of the North American plains. Museum of Northern Arizona Bulletin 65:423–444. Wible, J. R., and M. Spaulding. 2013. On the cranial osteology of the African palm civet, Nandinia Binotata (Gray, 1830) (Mammalia, Carnivora, Feliformia). Annals of Carnegie Museum 82:1–114. Wilman, H., B. J., S. J., de L. R. C., R. M., and J. W. 2014. EltonTraits 1.0 : Species-level foraging attributes of the world’s birds and mammals. Ecology 95:2027. Wilson, L. A. B., A. Balcarcel, M. Geiger, L. Heck, and M. R. Sánchez-Villagra. 2021. Modularity patterns in mammalian domestication: Assessing developmental hypotheses for diversification. Evolution Letters 5:385–396. 326 Wroe, S., and N. Milne. 2007. Convergence and remarkably consistent constraint in the evolution of carnivore skull shape. Evolution 61:1251–1260. Xiong, W.-Y. 2019. Basicranial morphology of Late Miocene Dinocrocuta gigantea (Carnivora: Hyaenidae) from Fugu, Shaanxi. Vertebrata Palasiatica 57:274–307. Yoder, A. D., M. M. Burns, S. Zehr, T. Delefosse, G. Veron, S. M. Goodman, and J. J. Flynn. 2003. Single origin of Malagasy Carnivora from an African ancestor. Nature 421:734–737. Zachos, J., H. Pagani, L. Sloan, E. Thomas, and K. Billups. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292:686–693. Zack, S. P. 2019a. The first North American propterodon (Hyaenodonta: Hyaenodontidae), a new species from the late Uintan of Utah. PeerJ 7:e8136. Zack, S. P. 2019b. A skeleton of a Uintan machaeroidine ‘creodont’ and the phylogeny of carnivorous eutherian mammals. Journal of Systematic Palaeontology 17:653–689. Zack, S. P., A. W. Poust, and H. Wagner. 2022. Diegoaelurus, a new machaeroidine (Oxyaenidae) from the Santiago Formation (late Uintan) of southern California and the relationships of Machaeroidinae, the oldest group of sabertooth mammals. PeerJ 10:e13032. Zheng, J. [郑家坚], Y. [汤英俊] Tang, R. [翟人杰] Zhai, S. [丁素因] Ding, and X. [黄学诗] Huang. 1978. Early Tertiary strata of Lunan Basin, Yunnan [云南路南盆地的早笫三纪地层]. Professional Papers on Stratigraphy and Paleontology, Beijing [地层古生物论文集] 7:22-29 [in Chinese]. 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. 327