See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/266755142 Phylogeny and evolution of cats (Felidae) Chapter · January 2010 CITATIONS READS 52 35,030 4 authors, including: Lars Werdelin Nobuyuki Yamaguchi Swedish Museum of Natural History Universiti Malaysia Terengganu 168 PUBLICATIONS 4,315 CITATIONS 106 PUBLICATIONS 2,208 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Sytematics and evolution of Erinaceidae View project Wildlife diseases View project All content following this page was uploaded by Nobuyuki Yamaguchi on 13 October 2014. The user has requested enhancement of the downloaded file. OUP CORRECTED PROOF – FINAL, 1/5/2010, SPi CHAPTER 2 Phylogeny and evolution of cats (Felidae) Lars Werdelin, Nobuyuki Yamaguchi, Warren E. Johnson, and Stephen J. O’Brien Artist’s reconstruction of the sabre-toothed cat Megantereon cultridens stalking its prey. (Illustration courtesy of Mauricio Anto´n.) Introduction felid evolution, especially that of the living felids and their ecological and functional relationship to the Cats, wild as well as domestic, fossil as well as living, extinct sabre-toothed felids. are familiar to people around the world. The family In this discussion, we will synthesize the available Felidae has a worldwide distribution and has been data, distinguishing as far as possible monophyletic associated with humans in various ways throughout groups of taxa, suggesting the most likely interrela- history (Quammen 2004). Their functional mor- tionships of the fossil lineages, but also pointing out phology, ecology, and behaviour have been the sub- that there are many problem areas that need to be ject of intense scrutiny by scientists for over 200 resolved. This section should be viewed as a chal- years. The fossil record of cats is extensive and some lenge to investigators to use old data or discover of its members are among the most recognizable of new data to corroborate or refute the scenarios pro- extinct animals. Despite all this, the phylogeny and posed herein. We end the paper with a small section evolution of the family Felidae, and even the content demonstrating some evolutionary patterns among of the family, have remained poorly understood. In extant Felidae, suggesting that there is much to be this review, we will first present the current state of gained from the deeper analysis of the current phy- knowledge with regard to the interrelationships logenetic information. of living Felidae and the timing of the radiation of Felid morphology is described and discussed else- modern cats. We will also present the fossil record of where (Kitchener et al., Chapter 3, this volume) and Felidae in broad outline, focusing first on describing will not be reiterated here except as needed. Teeth of the different groups of species and their characteris- the upper jaw are referred to in upper-case letters tics, and then discussing the general patterns of cat (I, C, P, and M) and teeth of the lower jaw in lower- evolution that we can deduce from current data. case letters (i, c, p, and m), followed by the appropriate Provided with this overview, we will attempt to iden- number in the sequence. Character mapping on tify those areas most in need of further research in cladograms was carried out with Mesquite, version order to achieve the aim of a fuller understanding of 1.12 (Maddison and Maddison 2004). Stratigraphic OUP CORRECTED PROOF – FINAL, 1/5/2010, SPi 60 Biology and Conservation of Wild Felids ages of taxa as given in the text and figures were based on a data set of 22,789 base pairs of DNA, obtained from either primary literature or (for North including autosomal, Y-linked, X-linked, and mito- America) the Paleobiology Database (www.paleodb. chondrial gene segments (Johnson et al. 2006b). The org) and (for Eurasia) the NOW database (www.hel- results of this study, while not immutable, provide a sinki.fi/science/now/database.html). firm basis for understanding the interrelationships and evolution of the extant Felidae. The results con- firm some prior results, both molecular and morpho- Phylogeny logical, while providing new insights and surprises. The study distinguishes eight clades of extant fe- Many attempts have been made to investigate the lids (Fig. 2.1). The first of these to split off from the interrelationships of Felidae. These have followed stem lineage is the Panthera lineage (genera Neofelis two broad approaches. Some, like Matthew (1910), and Panthera)atc. 10.8 Ma (Fig. 2.1, node A). Most Kretzoi (1929a, b) and Beaumont (1978) have previous studies of felid phylogeny have placed incorporated both fossil and extant felids in their Panthera as the crown group, but a few (Turner and analyses, while others, such as Pocock (1917a), Her- Anto´n 1997; Mattern and McLennan 2000) also have rington (1986), and Salles (1992) have focused exclu- the Panthera lineage as basal to other cats. Within this sively on the living members of the family. A new era in felid phylogenetics was ushered in with the intro- duction of molecular evidence (Collier and O’Brien Million years before present 35 10 5 1985; O’Brien et al. 1985a; Johnson et al. 1996), while the first study to use a total evidence approach P. linsang was that of Mattern and McLennan (2000). N. nebulosa All of these approaches have had their problems. N. diardi P. tigris In the case of fossil studies, confounding factors have P. uncia included the relatively poor fossil record, the prob- P. pardus P. leo lem of finding useful characters in fragmentary ma- P. onca A P. marmorata terial and the convergence between Nimravidae and P. badia Felidae. Though previously included in the Felidae P. temmincki L. serval (Matthew 1910; Piveteau 1961), the former, Nimra- C. caracal vidae, is now known to be diphyletic. Its Paleogene B C. aurata L. pardalis (65.5–23.0 million years ago [Ma]; Gradstein et al. L. wiedii L. colocolo 2004) members form a basal clade within either Feli- C L. jacobita formia or Carnivora as a whole (Neff 1983; Hunt L. tigrinus L. geoffroyi 1987; Morlo et al. 2004), while its Neogene (23.0 L. guigna D L. rufus Ma—recent) members are placed in a separate family, L. canadensis Barbourofelidae, with affinities to Felidae (see L. pardinus L. lynx below). Morphological studies of extant felids have A. jubatus E been hampered by the very uniform morphology of P. concolor P. yagouaroundi the members of the family, making it difficult to find F. chaus F F. nigripes and polarize characters for phylogenetic analysis. F. silvestris F. margarita Molecular studies, on the other hand, have been G O. manul particularly hampered by the apparently short time- P. rubiginosus span during which the clades of modern felids P. planiceps P. bengalensis evolved. Thus, clades of closely related taxa have P. viverrinus been identified but the interrelationships of these Figure 2.1 The phylogeny of the extant Felidae. Thick clades have been difficult to pinpoint. lines indicate the presence of a fossil record, thin lines Recently, two of us (Warren E. Johnson and Ste- indicate the absence of a fossil record. Node labels as in phen J. O’Brien) published a phylogeny of Felidae the main text. (Based on the work of Johnson et al. 2006b.) OUP CORRECTED PROOF – FINAL, 1/5/2010, SPi Phylogeny and evolution of cats (Felidae) 61 lineage, the clouded leopard, Neofelis, with the two (Johnson and O’Brien 1997) studies. It is worth not- species N. nebulosa and N. diardi (Buckley-Beason ing that the puma and jaguarundi probably split be- et al. 2006; Kitchener et al. 2006) is placed basally, fore the Great American Biotic Interchange that as would be expected from its distinctive morphol- followed the formation of the land bridge between ogy implying a long separate evolutionary lineage South and North America (Marshall et al. 1982), and (Christiansen 2006), with the rest of the pantherines thus both are of North American origin. radiating within the last 4 million years. The seventh and eighth lineages are the small cats The next clade to branch off, at c. 9.4 Ma (Fig. 2.1, of the Old World—the leopard cat and domestic cat node B), is the bay cat lineage (genus Pardofelis). This lineages. They split from each other at c. 6.2 Ma (Fig. clade consists of the poorly known bay cat (P. badia), 2.1, node G). The former includes the genera Otoco- Asian golden cat (P. temminckii), and marbled cat lobus and Prionailurus and the latter the genus Felis. (P. marmorata). The last mentioned species has been The splits within the former are much deeper than linked to the Panthera lineage (e.g. Herrington 1986) within the latter, suggesting that the genus Felis may and this is reflected in its position here, as basal be oversplit. This is also the conclusion of Driscoll member of the clade branching off closest to the et al. (2007), who distinguish only four species in Felis: Panthera lineage. F. chaus, F. nigripes, F. margarita,andF. silvestris.The The third lineage is the Caracal lineage, with two last mentioned species now also includes F. ornata, genera, Caracal and Leptailurus, incorporating three F. bieti,andF. lybica, making it one of the most wide- African species: caracal (Caracal), African golden cat spread small cat species. (C. aurata), and serval (S. leptail urus serval). This Most of the nodes in this phylogeny are robustly lineage branches off at c. 8.5 Ma (Fig. 2.1, node C), supported (Johnson et al. 2006b). A few, however, are with the serval basal to the other two species. still unstable, showing either low support or incon- The next lineage is the ocelot lineage (genus Leo- gruence between different analyses and data sets.