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PhylogenyandSpeciationofFelids

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Cladistics 16, 232–253 (2000)

doi:10.1006/clad.2000.0132, available online at http://www.idealibrary.com on

Phylogeny and Speciation of Felids

Michelle Y. Mattern and Deborah A. McLennan1

Centre for Comparative Biology and Biodiversity, Department of Zoology, University of Toronto,25 Harbord Street, Toronto, Ontario M5S 3G5, Canada

Accepted January 15, 2000

The phylogeny of the Felidae is reconstructed using atotal evidence approach combining sequences from 12SrRNA, 16S rRNA, NADH-5, and cytochrome b genes withmorphological and karyological characters. The 1504-character data set generated two equally parsimonioustrees (CI 5 0.413, 1795 steps) of which a strict consensusrevealed one polytomy in the placement of the bay catgroup. The tree supports several traditional groupingssuch as the genera Panthera and Lynx and the ocelotgroup of small South American felids, and it providesadditional resolution of relationships within and amongthe major felid lineages. Combining phylogenetic, distri-butional, and ecological data indicates that vicariant spe-ciation has played a relatively minor role in thediversification of the felids (approximately 26% ofevents), while sympatric speciation has been moreimportant than expected on theoretical grounds (approx-imately 51.8% of events), although postspeciation dis-persal may have blurred the boundaries betweensympatric, parapatric, and peripheral isolate modes. Anexamination of ecological changes on the felid tree showsrepeated patterns of resource partitioning in time (activ-

ity patterns), space (preferred habitat type), and food(as measured by body size) among closely related species.The rapid diversification of the cats thus appears to have

1To whom correspondence should be addressed. E-mail:mclennan@zoo.utoronto.ca.

232

been associated more with ecological than with geologi-cal separation. q 2000 The Willi Hennig Society

Key Words: biogeography; evolution; Felidae; speciation;molecular; morphology; phylogeny; total evidence.

What sort of philosophers are we, who know abso-lutely nothing of the origin and destiny of cats?

—Henry David Thoreau

Humans have had a long association with manymembers of the cat family. We hunt them for theirfur and medicinal properties, display them in zoos,associate them with magic and witchcraft, worshipthem as gods, domesticate them for pest control, andkeep them as pets. Although cats are an integral partof our history and our everyday lives, we know verylittle about most members of the Felidae and theirevolutionary relationships. Information is scatteredand generally biased toward the large, charismaticmembers of the genus Panthera, which includes thelion and tiger. Fascination with the larger cats has beencoupled with the relative neglect of the smaller, moresecretive felids. It is, after all, much easier logisticallyto track and observe lions than to struggle through the

jungles of Southeast Asia searching for the nocturnalleopard cat. These problems in data collection, com-bined with the observation that many felid species

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distributions with the hypothesized genealogical rela-

Phylogeny and Speciation of Felids

appear to display a frustrating combination of “primi-tive” and “specialized” characters, have plagued re-searchers interested in delineating the relationshipsamong cats ever since Linnaeus erected the genus Felisin 1758. For example, Pocock (1917, p. 329) describedthe results of early systematic attempts as “chaotic con-fusion,” a description that was echoed over half a cen-tury later with Collier and O’Brien’s (1985) assessmentthat felid relationships were still “widely disputed.”

Part of the early confusion stemmed from the lackof a robust methodology to reconstruct genealogicalrelationships from the wealth of data being amassed byresearchers (e.g., Schreber, 1774; Pocock, 1917; Wiegel,1961; Ewer, 1973; Radinsky, 1975; Hemmer, 1978, andextensive references therein; Wurster-Hill and Cen-terwall, 1982). Although the methodological problemwas solved with the publication of Hennig’s Phyloge-netic Systematics in 1950 (Hull, 1988), the first attemptto focus the phylogenetic beam upon the Felidae didnot appear until 36 years later (Herrington, 1986). Un-fortunately it is difficult to assess Herrington’s resultsbecause details of the analysis, including a data matrix,algorithms used, and tree statistics, were not includedin the discussion. In the 13 years following Her-rington’s study, the situation has improved past thepoint of chaotic confusion, but there are still glaringgaps in our understanding of felid relationships. Forexample, a strict consensus of tree topologies recon-structed using phylogenetic systematic methodology(Herrington, 1986; Salles, 1992; Johnson and O’Brien,1997) produces an almost complete polytomy among35 species (for a list of scientific and common namessee Appendix 1), with only three identifiable clades:(1) a polytomy comprising (Panthera tigris 1 Pan. onca1 Pan. pardus 1 Pan. leo); (2) (Lynx rufus (Lynx cana-densis 1 Lynx lynx)); and (3) (Leopardus pardalis 1 Leo-wiedii). A majority rule consensus of the same studiesgives five completely resolved groupings: (1) (Prionail-urus planiceps 1 (Pri. viverrinus 1 Pri. bengalensis)); (2)(Neofelis nebulosa (Pan. uncia (Pan. tigris (Pan. onca (Pan.pardus 1 Pan. leo))))); (3) (Pardofelis badia 1 Profelis tem-mincki); (4) (Puma concolor 1 Herpailurus yagouaroundi);and (5) Lynx as above. The monophyly of the domesticcat (Felis chaus 1 F. libyca 1 F. catus 1 F. silvestris 1 F.nigripes 1 F. margarita) and ocelot (Lynchailurus colocolo

1 Leo. geoffroyi 1 Oncifelis guigna 1 Leo. tigrinus 1 (Leo.pardalis 1 Leo. wiedii)) lineages is also supported, butthe resolution within those groups is minimal. Like the

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233

situation described for the strict consensus analysis,the relationships among the seven lineages, includingthe remaining seven felid species, are completely unre-solved, producing a large basal polytomy.

All members of the Felidae have at least one majorpopulation on Appendix I or II of CITES and on theIUCN Red List (Baillie and Groombridge, 1996) ofthreatened, endangered, and extinct species. In somecases, entire species have made the list (Lynx pardinus,N. nebulosa, On. guigna, Oreailurus jacobita, Pan. leo, Pan.pardus, Pan. uncia, Mayailurus iriomotensis, Ictailurusplaniceps). Collecting information about the cats is thusan urgent priority. Biologists have become increasinglyinterested in using phylogenetic trees as frameworksfor making conservation decisions (Brooks et al., 1992).The strength of those decisions, of course, is directlydependent upon the robustness of the phylogenetictree. Kluge (1989, 1998; see also Siddall, 1997) arguedthat the most robust trees could be reconstructed onlyby combining all available characters in one, total evi-dence analysis, ceasing the often endless debates aboutwhich types of data are more likely to contain “thetruth” about phylogeny (see also Flynn, 1996). Fortu-nately, as discussed above, researchers have been con-tributing valuable information about the evolutionarydiversification of the Felidae for over a century now.In fact, a few of those researchers have themselvesadvocated combining data from all sources to investi-gate the phylogeny of the enigmatic felids (Hemmer,1978; Leyhausen, 1979; Flynn, 1996). In this paper wewill heed that advice, combining all of the publisheddata, along with some new characters gathered fromthe literature, in order to reconstruct the most robustphylogenetic tree for the felids possible. We will thencombine data about general ecology and geographic

tionships to investigate patterns of speciation withinthe clade.

METHODS

Characters Used in the Phylogenetic Analysis

Both morphological evidence (Wozencraft, 1989;Wyss and Flynn, 1993) and molecular evidence (Dra-

goo and Honeycutt, 1997) place the Hyaenidae, Viverri-dae, and Herpestidae as successive sister-groups to the

234

Felidae. Crocuta crocuta (Hyaenidae) and Galidia elegans(Herpestidae) were thus chosen as outgroups to polar-ize characters in our analysis.

Karyology (characters 1–10; for descriptions see Ap-pendix 2). Details of chromosome number, morphol-ogy, and G-banding patterns for felid species and theoutgroups are from Wurster-Hill and Centerwall(1982), Dutrillaux and Couturier (1983), and Wurster-Hill et al. (1987).

Morphology (characters 11–68; for descriptions seeAppendix 2). The morphological data matrix con-structed by Salles (1992) was verified by M.Y.M. usingspecimens from the Royal Ontario Museum. One newcharacter was added to this matrix (68) based on re-ports that Pan. leo, Pan. onca, Pan. pardus, and Pan. tigrishave vocal folds with a thick pad of fibro-elastic tissue,whereas all other felids have sharp-edged vocal folds(Hast, 1986, 1989; Peters and Hast, 1994). Outgroupswere uninformative for this trait, so functional out-group analysis (Watrous and Wheeler, 1981) was usedto determine character polarity.

Molecular sequences (characters 69–1504). 16SrRNA (379 base pairs) and mitochondrial NADH-5(318 bp) sequences were available for 35 felid taxa andthe outgroups (Johnson and O’Brien, 1997). Mitochon-drial cytochrome b (363 bp) and 12S rRNA (376 bp)sequences were available for 20 felid taxa and the out-groups (Masuda et al., 1996). Sequences were alignedusing Malign, version 2.7 (Wheeler and Gladstein,1994), with transversions weighted 2:1 relative to tran-sitions and a gap creation penalty of 5. Several otherweights (1:1, 5:1, and 10:1) and gap creation penalties(2 and 10) were tested. They did not significantly im-pact the final alignment or the phylogenetic analysis.NADH-5 and 16S rRNA sequences are available fromGenBank (Accession Nos. AF006387–AF006460). Theentire aligned molecular data set is available fromEMBL (Alignment No. DS38847).

Phylogenetic Reconstruction

A total of 1504 characters (for a partial data matrix,see Appendix 3) were run unordered and unweightedin PAUP (version 3.1.1; Swofford, 1993) using the heu-ristic search algorithm (random addition sequence and

tree-bisection-reconnection branch-swapping). Thedata matrix was too large to analyze with the branch-and-bound algorithm, so eight separate heuristic

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Mattern and McLennan

searches (two of 1000 repetitions, three of 3000 repeti-tions, and three of 5000 repetitions) were run. A lackof information forced the exclusion of F. bieti, M. iriomo-tensis, Or. jacobita, and Lynx pardinus from the analysis.

Character Mapping

We collected data about habitat preferences, activitypatterns (nocturnal, diurnal, or both), and estimates ofbody size from the literature (Kitchener, 1991; Nowak,1991; Caro, 1994; Ortolani and Caro, 1996, and refer-ences therein; for descriptions see Appendix 1). Theseparticular traits are the only ones that have been re-corded for all or most of the felid species. None ofthese traits were included in the data matrix used toreconstruct the phylogeny. Characters were optimized(Farris, 1970; Maddison et al., 1984) by hand on the treeand the optimizations were verified using MacClade(version 3.04; Maddison and Maddison, 1992). Bodysize, as determined by weight, was used because thereis a general relationship between predator and preysize in carnivores (Gittleman, 1985). Changes in bodysize (morphology) are thus assumed to be associatedwith different prey preferences (ecology) in the felids(Kiltie, 1984, 1988). Three categories for size were used:small (1–10 kg), medium (11–40 kg), and large (.40kg). We recognize that these categories are somewhatarbitrary, but they are useful to a preliminary examina-tion of the relationship between felid body size evolu-tion and speciation. In order to conduct a more rigorousinvestigation, data about the range in size for malesand females across numerous populations within eachspecies are required, particularly for the less well stud-ied species like Par. badia, He. yagouaroundi, and Lynch-ailurus colocolo. Hyaenids fall into our large category,varying in weight from a maximum of 47.5 kg (Hy.brunnea), to 55 kg (Hy. hyeana), to 86 kg (Cr. crocuta;Nowak, 1991). We therefore used “large” as the out-group state for body size.

Modes of Speciation

Vicariant speciation (allopatric mode I) occurs whenan ancestral species is geographically separated intotwo or more relatively large and isolated populations,

with subsequent lineage divergence by the isolateddescendant populations (Rosen, 1978; Wiley, 1978,

by dispersal and secondary contact. Details of felid

less than the amount predicted under a random model

Phylogeny and Speciation of Felids

1981; Cracraft, 1982, 1983a, 1983b, 1986). Vicariant spe-ciation is generally assumed to be the null model forspeciation studies because the mechanism initiatingspeciation is independent of any particular biologicalsystem. Peripheral isolate speciation (allopatric modeII) occurs when a new species arises from a small,isolated population usually, but not always, on theperiphery of the larger ancestral population. Speciationcould happen rapidly (founder effect: Carson, 1975,1982; Barton, 1989) or it could be a relatively gradualprocess driven by adaptation to the new (peripheral)environment (Mayr, 1954, 1963). Because the peripheralpopulation is small and released somewhat from thehomeostatic constraints of large-scale gene flow, theperipheral descendant may exhibit more autapomor-phic traits than the central population (Hennig, 1966;Wiley, 1981), but this is not a requirement of the model(Lynch, 1989; Frey, 1993). Parapatric speciation (Endler,1977, 1982) occurs when two populations of an ances-tral species differentiate into descendant species de-spite the maintenance of limited gene flow and geo-graphical overlap during the process. Stochastic events(e.g., drift) and/or adaptive responses to local selectionpressures initiate the differentiation, while low vagilityamong members of the populations or a decrease inheterozygote/hybrid fitness leading to positive as-sortative mating promotes it. Sympatric speciation(Darwin, 1859; Maynard Smith, 1966; Bush, 1975a,1975b, 1982; Diehl and Bush, 1989; Kawecki, 1997) oc-curs when one or more new species arise without geo-graphical segregation of populations. Unlike the allo-patric models, which postulate that gene flow betweenpopulations is initially severed by factors extrinsic tothe biological system, sympatric speciation requiresthe involvement of biological processes intrinsic to thesystem; e.g., hybridization, ecological partitioning, ora change in mate recognition.

Demonstration of sympatric speciation is predicatedupon the assumption that the influence of geographicalseparation during speciation has not been obscuredby rampant dispersal of the descendant species. Thisassumption does not state that dispersal is unimport-ant, only that postspeciation dispersal does not over-whelm speciation patterns. We have reason to believethat this assumption has been violated to some extentwithin the Felidae. Many felids are large, highly mobile

creatures (e.g., a male mountain lion may occupy ahome range of up to 500 square miles) (Kitchener, 1991;

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235

Kobalenko, 1997). Given these dispersal abilities, canwe use sympatric overlap of sister-species today asevidence for sympatric speciation, or is it more realisticto postulate that speciation occurred in allopatry fol-lowed by secondary contact between the newly fledgedsister-pairs? This question is particularly pertinentwhen we are considering the widespread sympatrybetween ancestral sister-species buried deep in thephylogeny of the group because we would expect theobfuscating effects of dispersal to increase with increas-ing age of the speciation event (Brown and Gibson,1983; Chesser and Zink, 1994). Given this, we will takea conservative approach in this paper and err on theside of the null hypothesis. Sympatric, parapatric, and,to a lesser extent, peripheral isolate speciation all re-quire the evolutionary diversification of some specialecological or genetic characteristics that could, in them-selves, produce independent species; therefore, if twosister-species show any degree of overlap, but do notshow any ecological character change, we will tenta-tively assign that event to vicariant speciation followed

geographic distributions are given in Appendix 4.

RESULTS

The 1504-character data matrix yielded two equallyparsimonious trees, each with a length of 1795 steps.A strict consensus of the two trees (Fig. 1) revealedonly one point of incongruence: Par. marmorata is thesister-species to a clade comprising either (1) (domesticcat group (caracal group 1 leopard cat group)) or (2)(bay cat group (domestic cat group (caracal group 1

leopard cat group))). The final tree has a consistencyindex of 0.413, a retention index of 0.496, and a rescaledconsistency index of 0.205. Meier et al. (1991) developedthe homoplasy slope ratio (HSR), a measure to deter-mine whether the amount of homoplasy in a matrixdeviates significantly from random given the numberof taxa and characters. The HSR for this matrix is 0.16,well within the expected range of 0.03 to 0.46, indicat-ing that the amount of homoplasy, although large, is

for a data matrix of this size.Molecular data contributed 44, morphological data

also Table 2) and putative ages of each lineage (MYA, million years ago) based upon amount of sequence divergence (Johnson and O’Brien,On

group))), and (3) medium body size originated inde-

1997) are written across the top of the tree. L. cat, leopard cat clade.of genera).

contributed 8, and karyological data contributed 3 un-ambiguous (non-homoplasious) synapomorphies tothe overall tree topology (Table 1). x 2 analysis indicatesthat karyological and morphological characters con-tributed significantly more, and molecular characterssignificantly fewer, non-homoplasious synapomor-phies than expected based upon the total number ofeach character type used to reconstruct the tree (x 2 5

34.9, P . 0.01). More importantly, some nodes wereunambiguously supported by only one type of data(Table 1). The 12S rRNA and cytochrome b sequencessupport many of the basic groupings, but since theywere available for only 20 taxa, they do not affect theoverall topology of the tree when removed. These char-acters serve only to resolve a polytomy present in thedomestic cat lineage as well as to lend support to other

parts of the tree. The number and type of autapomor-phies for each species are listed in Table 2.

Acctran optimization of body weight (Fig. 2) onto

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ly species names are indicated on the tree (see Appendix 1 for names

the consensus tree indicates that large size is plesiomor-phic for the Felidae with six independent changes tomedium size, one reversal to large size, and three ori-gins of small body size. Deltran optimization gives asimilar picture, with the following exceptions: (1) theancestor of the (Puma 1 Lynx) clade is plesiomorphi-cally large, (2) there are two changes from large to smallin the ancestor of the ocelot clade and the ancestor ofthe clade comprising (the bay cat group, Par. marmorata(domestic cat group (caracal group 1 leopard cat

236 Mattern and McLennan

FIG. 1. Strict consensus tree of two equally parsimonious phylogenetic trees for the Felidae based upon 1504 characters (consistency index0.413; retention index 0.496; rescaled consistency index 0.205). Small numbers refer to nodes (Table 1). Names of monophyletic lineages (see

pendently in the ancestor of the caracal group and inPri. viverrinus.

DISCUSSION

Flynn (1996) made a passionate plea for Kluge’s(1989) total evidence approach. We concur with Flynn’s

two equally parsimonious trees.a Number in parenthesis indicates number of unambiguous (non-homoplasious) synapomorphies.b

Unambiguous synapomorphies from this type of character.

conclusion and believe that our total evidence analysishas produced the most robust hypothesis of phyloge-netic relationships among felid species to date. Thefinal tree supports all of the major monophyletic lin-eages (Panthera, ocelot, Lynx, puma, caracal, leopardcat, bay cat, and domestic cat) proposed by Johnsonand O’Brien (1997). Their molecular data, however,gave only poor resolution of the relationships amongthe eight lineages. As we had hoped, the resolution ofinterlineage relationships was improved in the totalevidence analysis, demonstrating that different datasets added resolution to different parts of the phyloge-netic tree. For example, although there were far moremolecular characters, unambiguous support for fivenodes was provided only by morphological characters,while support at two nodes was increased by the pres-ence of karyological synapomorphies (Table 1).

The distribution of characters on the tree confirmsearlier suspicions that cats are difficult to work with

in a systematic framework because they display a largenumber of autapomorphies and symplesiomorphies(Flynn, 1996; Johnson and O’Brien, 1997). The number

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of symplesiomorphic 1 autapomorphic characters out-weighs the number of synapomorphies 1.7:1 in thisdata set. This pattern may be typical of groups thathave undergone relatively rapid diversification earlyin their evolutionary history. Rapid diversification issupported by Johnson and O’Brien’s age estimates forthe eight lineages, which appear to have diverged fromone another within a range of about 5–8 million yearsago (MYA). Mapping the age estimates onto the con-sensus tree, however, indicates that the age estimatefor the puma clade is incorrect (Fig. 1). No phylogeneticanalysis has ever placed the puma lineage as the basalfelid group, a position it must hold if it is indeed theoldest lineage. Age estimates are generally based uponthe amount of sequence divergence, under the assump-tion of a constant divergence rate within a clade (seeexcellent discussion in Flynn, 1996). Given that thepuma lineage displays high levels of autapomorphies(Table 2) compared with its sister-group (Mann–

Phylogeny and Speciation of Felids 237

TABLE 1

Amount of Support for Each Node on the Felid Consensus Tree

Character type Character type

Node Total No.a Ka Mo N5 16s Cb 12s Node Total No.a Ka Mo N5 16s Cb 12s

1 55 (11) — 4 11 15b 16b 9b 11 24 (3) — — 5 3 10 6b

2 38 (8) — 6 4b 2 22b 4b 12 29 (1) — — 14b 3 10 23 22 (2) — 5 11b 6b — — 13 17 (1) — 1 6 5 3 2b

4 17 (1) — 4b 12 1 — — 14 14 (1) — 1 4 3 6b —5 5 (0) — — 3 2 — — 15 19 (2) 1 4b 8 6 — —6 7 (0) — — 6 1 — — 16 26 (2) 1 4 14b — 6b 17 22 (0) — 2 5 4 9 2 17 12 (1) — 4b 6 2 — —8 26 (1) 2 2b 8 1 10 3 18 18 (0) — 1 13 4 — —9 24 (4) 2b 1 7b 2 9 3b 19? 15 (0) — 4 3 2 1 5

10 17 (0) — 1 6 2 7 1 20? 6 (0) 1 1 4 — — —21 14 (3) 2b 1 11b — — — 28 3 (0) — 2 — 1 — —22 12 (0) 1 1 3 7 — — 29 2 (0) — — — — 2 —23 15 (0) 1 2 3 2 5 2 30 20 (1) — 2 6 1 10 1b

24 27 (5) — 3 2 3b 11b 8b 31 18 (2) — 7 3 8b — —25 25 (1) 1 2b 8 2 9 3 32 10 (0) — 1 1 1 7 —26 16 (0) — 1 5 3 6 1 33 11 (0) — 1 8 2 — —27 38 (2) — 1 15 6 14b 2b 34 21 (3) 1 0 14b 6 — —

Note. Boldface type indicates nodes with unambiguous support from only one type of data. Character types: Ka, karyological; Mo, morphologi-cal; N5, NADH-5; 16s, 16S rRNA; CB, cytochrome b; 12s, 12S rRNA. ? denotes that nodes are ambiguous. Support shown is from one of the

Whitney test: z 5 21.96, P , 0.05) and compared withthe levels across the remaining members of the Felidae(Mann–Whitney test: z 5 22.72, P , 0.007), it is not

Lep. serval 32 (0/2/30)Ca. caracal 30 (0/6/24)

Pro. aurata 22 (1/2/19)

surprising that an age estimate based upon quantityof change places the puma group as unusually oldwithin the felids. Because sister-groups are, by defini-

tion, the same age (Mayden, 1986), we would have topostulate that the Lynx group also originated at least

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Mattern and McLennan

8.2 MYA and that the two basal lineages on the tree,the ocelot and Panthera groups, are even older. So,either the ages of the basal members of the clade, andhence estimates of felid origins in general, are incorrector the age of the puma lineage is in error.

Our analysis supports Pocock’s (1917) original sub-family Pantherinae, with the inclusion of N. nebulosaas the basal member of the clade (see also Johnson andO’Brien, 1997) based upon 38 synapomorphies, 8 ofwhich are unambiguous molecular characters (Table1). The biogeography of this group is complicated, inpart because there have been so many extinctions bothin distant time and, more alarmingly, in recent times(Kitchener, 1991; Baillie and Groombridge, 1996).Matching the distributions (Appendix 4) with the phy-logenetic patterns hints at the following sequence ofspeciation events (Fig. 3): Node 2: sympatric speciationaccompanied by a body size increase in ancestor 1;Node 3: peripheral isolate speciation following move-ment of a subset of ancestor 1 into the high mountains,facilitated, perhaps, by the change in body size. It isimpossible to compare the number of autapomorphiesin Pan. uncia with the number in its ancestor/sister-group, because that ancestor has itself undergone asubstantial amount of speciation. Although the twogroups are the same age, the amount of speciation inthis larger clade, combined with the high degree ofadaptive change involving habitat shifts, body sizeevolution, and changes in activity pattern, will haveobscured the initial comparison between the peripheralisolate and its ancestor. Some time following this event,ancestor 1 itself underwent a shift from a diurnal to anocturnal activity pattern (ancestor 2: Acctran optimi-zation. Deltran shows two independent origins of diur-nal behavior associated with the speciation of N. nebu-losa and Pan. uncia); Node 4: sympatric speciationaccompanied by a second increase in body size (ances-tor 3); Node 5: sympatric speciation associated withthe movement by the ancestor of Pan. leo out of theforest into the savannah and a change from a nocturnalto a more generalized activity pattern; Node 6: Fossilevidence indicates that species with both tiger-like andjaguar-like features existed in northern China (Pan.palaeosinensis) and northern Eurasia (Pan. gomba-zoegensis) during the Pliocene (Hemmer, 1979). If the

238

TABLE 2

Number and Type (Karyological/Morphological/Molecular) ofAutapomorphies for Each Felid Species (Genera Names inAppendix 1)

Species Autapomorphies

Panthera groupN. nebulosa 25 (2/2/21)Pan. uncia 31 (0/7/24)Pan. pardus 17 (0/0/17)Pan. tigris 22 (0/2/20)Pan. onca 31 (0/2/29)Pan. leo 8 (0/1/7)

Ocelot groupLeo. wiedii 32 (0/1/31)Leo. pardalis 22 (0/0/22)On. guigna 22 (0/1/21)Leo. geoffroyi 17 (0/2/15)Leo. tigrina 8 (0/1/7)Leo. colocolo 29 (1/4/24)

Puma groupA. jubatus 61 (1/15/45)He. yagouaroundi 70 (1/5/64)Puma concolor 43 (0/3/40)

Lynx groupLynx rufus 13 (0/1/12)Lynx lynx 12 (0/2/10)Lynx canadensis 23 (0/1/22)

Bay cat groupPro. temmincki 29 (0/2/27)Par. badia 23 (0/2/21)

Marbled catPar. marmorata 28 (0/2/26)

Felis groupPri. rubiginosa 33 (1/2/30)Ot. manul 42 (0/8/34)Felis nigripes 55 (0/5/50)Felis chaus 35 (0/5/30)Felis silvestris 9 (0/1/8)Felis margarita 22 (0/4/18)Felis libyca 9 (0/2/7)Felis catus 6 (0/0/6)

Leopard cat groupPri. viverrina 21 (0/0/21)I. planiceps 49 (1/4/44)Pri. bengalensis 23 (0/5/18)

Caracal group

two extinct species are the sister-group of Pan. onca,then it is possible that the common ancestor of the

FIG. 2. Acctran optimization of body size (weight) onto one of the two equally parsimonious trees for the Felidae. The optimizations donot change on the other tree. White, small (1–10 kg), gray, medium (11–40 kg); black, large (.40 kg); hatched lines, indeterminate. Names

ca

of felid lineages are written across the top of the tree. L. cat, leopard

three was spread across northeastern Eurasia intoNorth America and was subdivided by the rupture ofthe land bridge between Asia and North America or byclimate changes associated with one of the Pleistoceneglaciation events (but see Klicka and Zink, 1997). It isalso possible that Pan. tigris, Pan. gombazoegensis, Pan.palaeosinensis, and Pan. onca may represent a series ofperipheral isolate speciation events along a dispersalroute across Asia to South America, something akin toisland hopping. If this hypothesis is correct, we wouldexpect to see some differences in ecology between Pan.tigris and Pan. onca. The ecological data are difficult toassess for these two species. Greene and Losos (1988)reported that Pan. onca tended to be associated withwater courses, even in xeric regions. Members of Pan.tigris are powerful swimmers; there is even evidencethat they mate in the water occasionally. So the at-traction to water, if anything, only provides further

support for the close relationship of these two cats.Given the ambiguities, Node 6 is assigned the conser-vative status of vicariant speciation.

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t clade.

The monophyly of the South American ocelot lineage(Johnson and O’Brien, 1997) is supported by 26 synapo-morphies, one of which, a reduction in the number ofchromosome pairs from 38 to 36 due to the tandemfusion of chromosomes F2 and F3 into C3, is unambigu-ous. Our analysis preserves Johnson and O’Brien’sthree sister-pairs (Leo. pardalis 1 Leo. wiedii, Leo. geoffroyi1 On. guigna, Leo. tigrinus 1 Lync. colocolo), but placesLeo. pardalis 1 Leo. wiedii as the sister-group to theremaining four species (see also two of four trees inMasuda et al., 1996). The following speciation modesare suggested (Fig. 4): Node 8: the distributions indi-cate parapatric speciation of ancestor 1 and ancestor 2,but there are no obvious functional changes associatedwith this event. As discussed previously, this node istentatively assigned the status of vicariant speciationfollowed by dispersal and secondary contact; Node 9:sympatric speciation accompanied by an increase inbody size (small to medium) in Leo. pardalis and a

Phylogeny and Speciation of Felids 239

change in ecology (from primarily terrestrial to arbo-real) in Leo. wiedii; Node 10: the distributions indicate

FIG. 3. Ecological and body size data optimized onto the phylogenetic tree for the Panthera group. Boldface type indicates apomorphic

of the fossil data (Van Valkenburgh et al., 1990) indi-

changes in the character.

sympatric speciation of ancestor 3 and ancestor 4, butagain there are no obvious functional changes associ-ated with this event (vicariant speciation with posts-peciation dispersal); Node 11: peripheral isolate specia-tion of On. guigna, presumably following themovement of a small population of ancestor 3 westacross the Andes into unoccupied, plesiomorphic habi-tat (classic waif dispersal; Frey, 1993). Although Leo.geoffroyi shows two apomorphic changes in ecology,there is no difference in the number of autapomorphies(Table 2) between the putative peripheral isolate andits sister-species (x2 5 0.64, NS); Node 12: parapatricspeciation associated with an ecological shift by Lync.colocolo out of the forests into grasslands and sur-rounding woods.

The puma and Lynx clades are placed as sister-groupson the basis of 14 synapomorphies, with one unambig-uous character provided by cytochrome b (see alsoneighbor-joining tree in Johnson and O’Brien, 1997).The puma clade is supported by 19 synapomorphies,2 of which, the greater development of an anteriorly

projecting flange on the head of the fibula (56) and theconnective structure of the tendon for the extensor

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digitorium longus (57), are unambiguous morphologi-cal characters (Table 1). The monophyly of the lynxesis supported by 12 synapomorphies, including one un-ambiguous morphological change, a reduction in thenumber of caudal vertebrae. The following speciationmodes are suggested (Fig. 5): Node 14: sympatric speci-ation associated with a change from strictly nocturnalto nocturnal/diurnal in ancestor 2 and a shift in bodysize in either ancestor 2 (medium to large: Acctran)or ancestor 1 (large to medium: Deltran); Node 17:sympatric speciation accompanied by a widening ofLynx rufus’ preferred habitat to include semidesertsand brushlands and a widening of ancestor 3’s habitatto include the equally arid (but colder) tundra; Node18: vicariant speciation on either side of the Atlantic.This speciation event was not accompanied by anyobvious ecological diversification; Node 15: sympatricspeciation accompanied by movement into more openenvironments and completely diurnal behavior in theancestor of the Acinonyx clade. Phylogenetic analysis

240 Mattern and McLennan

cates that the Acinonyx group speciated along a NorthAmerica (Miracinonyx inexpectatus and Mi. trumani)–

ic

FIG. 4. Ecological and body size data optimized onto the phylogenetin the character.

Europe (A. pardinensis)–Africa (A. jubatus) route (Mar-tin et al., 1977; Adams, 1979). This wave of speciationwas accompanied by the evolution of tibia–fibula fu-sion (A. jubatus), restricting climbing ability, and theenlargement of the nasal aperture (ancestor of Mi. inex-pectatus 1 Acinonyx), which presumably allowedgreater air intake for sprinting; Node 16: sympatricspeciation associated with a dramatic decrease in bodysize and partial movement off the ground into the trees,in He. yagouaroundi.

Grouping Par. badia with Pro. temmincki is supportedby 14 synapomorphies, 3 of which (including a distinc-tive banding pattern on chromosomes A1 and D1) areunambiguous (see also Pocock, 1932; Hemmer, 1978;Johnson and O’Brien, 1997). The production of thissister-pair represents another instance of sympatricspeciation (node 21 on Fig. 1) associated with an apo-morphic increase in body size from small to mediumin Pro. temmincki (Fig. 2). It is equally parsimonious toplace this sister-pair on either side of Par. marmorata.Too little is known about the ecology of Par. marmoratato propose any putative character change involved inits speciation.

The domestic cat clade of Johnson and O’Brien (1997)is expanded to include Otocolobus manul 1 Pri. rubigino-sus on the basis of 15 synapomorphies (Table 1), none

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tree for the ocelot group. Boldface type indicates apomorphic changes

of which is unambiguous. The relationships among F.margarita, F. silvestris, F. libyca, and F. catus are differentthan those postulated by Johnson and O’Brien, whoplaced F. margarita as the sister-group to the remainingthree species. The pairings are, however, congruentwith those postulated by Kratochvıl (1975), based onthe morphology of the os penis. The hypothesized se-quence of speciation is as follows (Fig. 6): Node 23:the distributions indicate sympatric speciation of an-cestors 1 and 2 in the Indian–central Asian area. Thereare no obvious functional changes associated with thisevent, so this node is tentatively assigned the statusof vicariant speciation followed by postspeciation dis-persal; Node 24: peripheral isolate speciation of Oc.manul following movement by a small population ofancestor 1 into the high steppes and deserts of theHimalayas. There is no difference in the number ofautapomorphies (Table 2) between the putative periph-eral isolate and its sister-species (x2 5 1.08, NS); Node25: peripheral isolate speciation of F. nigripes associatedwith no obvious ecological change. Felis nigripes is sep-arated from its sister-group by thousands of miles.It is possible that the original speciation event was

Phylogeny and Speciation of Felids 241

triggered by a widespread climate/habitat change innorthern Africa (e.g., the origin and expansion of the

FIG. 5. Ecological and body size data optimized onto the phylogenetic tree for the puma 1 Lynx group. Boldface type indicates apomorphic

changes in the character.

Sahara desert) followed by long-term habitat destruc-tion, fragmentation, and extinction of F. nigripes popu-lations. It is difficult to compare the number of autapo-morphies in F. nigripes with the number in its ancestor/sister-group because that ancestor has itself undergonea substantial amount of speciation; Node 26: sympatricspeciation associated with an increase in body size(from small to medium) and a change from a nocturnalto a diurnal activity pattern in F. chaus; Node 27: thedistributions indicate parapatric speciation of ancestor5 and ancestor 6 with the area of overlap in northernAfrica, but again there are no obvious functionalchanges associated with this event (vicariant speciationwith postspeciation dispersal); Node 28: parapatricspeciation associated with a movement into arid habi-tats, including sand dunes, by F. margarita; Node 29:artificial selection. The origin of F. catus has been de-bated for decades (Todd, 1978; Robinson, 1984; Serpell,1988; Daniels et al., 1998). Randi and Ragni (1991) com-bined F. silvestris, F. libyca, and F. catus into one poly-

typic species (F. silvestris) based upon allozyme data.Essop et al. (1997) concurred, reporting that the restric-tion map sites from mitochondrial DNA of F. libyca

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and F. catus are indistinguishable and the divergence(0.9%) between F. silvestris and the other two is notenough to designate them as separate species. Thisclassification can be supported by our analysis only ifF. margarita is also reduced to a subspecies of F. silvestris.

The most recently derived felid lineage is the leopardcat 1 the caracal clade. The sister-group status of thesetwo clades is supported by 20 synapomorphies, oneof which is unambiguous (from 12S rRNA). The leop-ard cat group is supported by 18 synapomorphies, 2 ofwhich are unambiguous (from 16S rRNA). The sister-group relationship of Pri. bengalensis and I. planiceps iscongruent with one of the trees published by Masudaet al. (1996) and by Salles (1992), but differs from thePri. viverrinus 1 Pri. bengalensis relationship postulatedby Johnson and O’Brien (1997). The caracal group issupported by 11 synapomorphies, none of which areunambiguous. It includes the generally accepted rela-tionship between Pro. aurata and Caracal caracal (Collierand O’Brien, 1985; Janczewski et al., 1995; O’Brien et al.,

242 Mattern and McLennan

1987; Johnson and O’Brien, 1997) and the less commonpostulate that Leptailurus serval is the sister-group tothat pair (but see Kratochvıl, 1982). The hypothesized

FIG. 6. Ecological and body size data optimized onto the phylogenetic tree for the domestic cat group. Boldface type indicates apomorphic

changes in the character.

sequence of speciation is as follows (Fig. 7): Node 30:vicariant speciation associated with the origin of swim-ming ability in ancestor 2 and possibly an increase inbody size (from small to medium: Deltran) in ancestor1; Node 33: sympatric speciation associated with theexpansion of ancestor 3’s preferred habitat to includedry forests as well as savannah, and the subsequentorigin of climbing ability in that ancestor; Node 34:sympatric speciation associated with the movement byPro. aurata into thicker rainforests and a change in itsactivity pattern from nocturnal to diurnal. Speciationin this group has involved a gradual movement fromsavannah (Lep. serval), to dry woods/savannah (Ca.caracal), to thicker forests (Pro. aurata); Node 31: sym-patric speciation associated with either a decrease inbody size (medium to small: Acctran) in ancestor 4 oran increase in body size (small to medium: Deltran) inPri. viverrinus; Node 32: sympatric speciation associ-ated with movement into thicker forest, coupled with

the origin of climbing ability in Pri. bengalensis, andthe widening of activity patterns to include diurnal aswell as nocturnal hunting, by I. planiceps. Interestingly,

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this clade shows the same trend toward a gradual inva-sion of the forest habitats as does its sister-group, in thiscase following a sequence from the vegetation aroundmangrove swamps and creeks (Pri. viverrinus) to riv-erbanks (I. planiceps), to forested areas (Pri. bengalensis).

Unlike the situation described for several other verte-brate clades (e.g., Lynch, 1989; Brooks and McLennan,1991, and references therein), vicariant speciationseems to have played a limited role in the Felidae. Fiveof seven putative instances of vicariance are highlysuspect and were accorded that status solely becausewe could not detect any ecological character changebetween sympatrically or parapatrically distributedsister-species with our limited data set. The majorityof these cases (4/5) are buried deep within the phyloge-netic tree, so it is not untoward to postulate that posts-peciation dispersal may have led to secondary contactand overlap in these cases (Brown and Gibson, 1983;

Phylogeny and Speciation of Felids 243

Chesser and Zink, 1994). Nevertheless, even with theseambiguous cases, vicariant speciation accounts for only25.9% of felid diversification. The studies depicted in

enet

FIG. 7. Ecological and body size data optimized onto the phylogapomorphic changes in the character.

Table 3 indicate that the relative importance of specia-

Note. Values represent the percentage of each mode as a function of t(N ). Ecological and body size character changes associated with a given

a Data taken from Lynch (1989).b Data taken from Chesser and Zink (1994).

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ic tree for the caracal 1 leopard cat group. Boldface type indicates

244 Mattern and McLennan

biological “fates and tendencies” (Brooks and Wiley,

1988), and these tendencies will play a role in determin-tion modes can vary widely among different groups

of vertebrates. In other words, clades have their own ing which types of speciation are more or less likely

TABLE 3

Postulated Frequencies of Speciation Modes in Different Vertebrate Groups, Including the Felidae

Frequency of speciation mode

Clade N Vicariant Peripheral isolates Parapatric Sympatric

Ranaa 19 89.4 5.3 Not examined 5.3Ceratophrysa 6 83.3 16.7 Not examined 0Eleutherodactylusa 7 71.4 14.3 Not examined 14.3Fundulusa 4 100 0 Not examined 0Heterandriaa 8 62.5 37.5 Not examined 0Xiphophorusa 13 61.5 23.1 Not examined 15.4Poephilaa 5 80.0 0 Not examined 20.0Passerinesb 30 66.7 3.3 Not examined 30.0Felidae 27 25.9 14.8 7.4 51.9Type of character change associated Body size 0 Body size 0 Body size 0 Body size 8

with particular speciation modes Habitat 1 Habitat 4 Habitat 2 Habitat 9in the Felidae Activity 0 Activity 1 Activity 0 Activity 6

he total number of events that could be assigned a speciation modespeciation mode are shown for the felids.

involvement, the fate of the cats may parallel the be-

Phylogeny and Speciation of Felids

within a group. We believe, therefore, that while it isimportant to begin with vicariant speciation as the nullhypothesis, it is equally important to allow informationfrom phylogeny, distributions, ecology, and genetics toidentify instances of peripheral isolate, parapatric, andsympatric speciation that refute the null hypothesis.Biology may be influenced by geology, but it is notcompletely subservient to it.

A cursory examination of ecological data shows re-peated patterns of resource partitioning (Schoener,1965, 1974) among closely related species. The parti-tioning of the environment by sympatric sister-speciesprovides more support for the hypothesis that adapta-tion via ecological character change may play a causalrole in the speciation process (see excellent discussionsin Losos, 1990, 1992; Bush, 1994; Schluter, 1996; Ka-wecki, 1997; Losos et al., 1997; Nagel and Schluter,1998, and references therein). The macroevolutionarypatterns plus distributional data on their own, how-ever, cannot tell us anything about the underlying pro-cesses (Brooks and McLennan, 1991; McLennan, 1996).For example, Toft (1985) concluded that resource parti-tioning in a wide variety of amphibians and reptilesresulted from a number of causes, including competi-tion, predation, and physiological constraints, actingalone or in concert. In order to understand how thesedifferent selective forces have influenced the evolutionof resource partitioning and the subsequent speciationof many felids, we need more detailed informationabout the physiology, ecology, and behavior of the fe-lids in general. We also need to demonstrate that thechanges in ecology are linked with the evolution ofreproductive isolation between the sister-pairs (May-

nard Smith, 1966; Rosenzweig, 1978; Wilson and Ture-lli, 1986; Diehl and Bush, 1989; Bush, 1994; Nagel and

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245

Schluter, 1998). Overall, though, this preliminary inves-tigation indicates that closely related cats are able tocoexist (Hutchinson, 1959; MacArthur and Levins,1967; Houle, 1997) because of changes in one of thethree traditional resource dimension categories (Pi-anka, 1975): displacement in habitat (space), displace-ment in time (stay in the same place, but hunt at differ-ent times), and displacement in energy source (huntin the same area at the same time, but focus on differentprey items).

Most of the character changes associated with specia-tion in the felids (Table 3) were autapomorphic (bodysize 5 4/8 (50%); habitat 5 13/16 (81.2%); activitypattern 5 5/7 (71.4%)). The patterns of ecological evo-lution within the Felidae (conservative general feedingmode (predator) coupled with widespread, autapo-morphic changes in prey preference, habitat prefer-ence, and activity pattern) parallels the general trendtoward a high percentage of symplesiomorphic plusautapomorphic characters uncovered for morphologi-cal, karyological, and molecular data. It is this couplingof phylogenetic stasis (conservatism) and adaptiveflexibility that makes the cats at once an evolutionarysuccess story and a systematic enigma. Unfortunately,many felid species are so endangered now that it isbecoming increasingly difficult to collect the detailedecological and behavioral data that are required to for-mulate robust hypotheses about felid origin and diver-sification. As biologists, we need to take an active roleboth in providing the scientific framework for conser-vation decisions and in the decision-making processitself (Greene and Losos, 1988; Brooks et al., 1992;O’Brien, 1994; Miththapala et al., 1996). Without that

havior of one of their most famous members, fadinglike a smile into the night.

246 Mattern and McLennan

APPENDIX 1

Scientific Names,a Common Names, and Ecological Characters of Felid Species

Scientific name Common name Abbreviation General habitat Within habitat Activity

Acinonyx jubatus Cheetah jubatus Brush/grassland Terrestrial DiurnalCaracal caracal Caracal caracal Forest/savannah Terrestrial/arboreal NocturnalFelis bieti Chinese desert cat Excluded — — —Felis catus Domestic cat catus Widespread Terrestrial NocturnalFelis chaus Jungle cat chaus Forest (open) Terrestrial DiurnalFelis libyca African wild cat libyca Widespread Terrestrial NocturnalFelis margarita Sand cat margarita Deserts Terrestrial NocturnalFelis nigripes Black-footed cat nigripes Forest (open) Terrestrial NocturnalFelis silvestris European wild cat silvestris Widespread Terrestrial NocturnalHerpailurus yagouaroundi Jaguaroundi yagouaroundi Lowland forests Terrestrial/arboreal BothIctailurus planiceps Flat-headed cat planiceps Riverbanks Terrestrial/swims BothLeopardus geoffroyi Geoffroy’s cat geoffroyi Open brush Terrestrial/swims NocturnalLeopardus pardalis Ocelot pardalis Forest (dense to open) Terrestrial NocturnalLeopardus tigrinus Little tiger cat, tigrina tigrinus Forest ? ?Leopardus wiedii Margay wiedii Forest Arboreal ?Leptailurus serval Serval serval Savannah, around streams Terrestrial NocturnalLynchailurus colocolo Pampas’ cat colocolo Forest/grassland Terrestrial NocturnalLynx canadensis Canadian lynx canadensis Widespread (dense Forest Terrestrial Nocturnal

to open tundra)Lynx lynx Eurasian lynx lynx Widespread as above Terrestrial NocturnalLynx pardinus Spanish lynx Excluded — — —Lynx rufus Bobcat rufus Widespread (forest to Terrestrial Nocturnal

Semidesert, open brush)Mayailurus iriomotensis Iriomote cat Excluded — — —Neofelis nebulosa Clouded leopard nebulosa Forests Terrestrial DiurnalOncifelis guigna Kodkod guigna Forest Terrestrial NocturnalOreailurus jacobita Andean mountain cat Excluded — — —Otocolobus manul Pallas’ cat manul High steppes & deserts Terrestrial NocturnalPanthera leo Lion leo Savannah Terrestrial BothPanthera onca Jaguar onca Widespread Terrestrial/swims NocturnalPanthera pardus Leopard pardus Widespread Terrestrial NocturnalPanthera tigris Tiger tigris Widespread Terrestrial/swims NocturnalPanthera uncia Snow leopard uncia High mountains Terrestrial DiurnalPardofelis badia Borean bay cat badia Forest ? ?Pardofelis marmorata Marbled cat marmorata Forests ? NocturnalPrionailurus bengalensis Leopard cat bengalensis Forests (high–low) Terrestrial/arboreal NocturnalPrionailurus rubiginosus Rusty-spotted cat rubiginosus Widespread (grasslands Terrestrial Nocturnal

to montane forests)Prionailurus viverrinus Fishing cat viverrinus Mangrove swamps Terrestrial/swims ?Profelis aurata African golden cat aurata Forests & mountains Terrestrial/arboreal DiurnalProfelis temmincki Asian golden cat, temmincki Forest Terrestrial ?

Temminckii’s cat

Puma concolor Cougar, mountain concolor Widespread Terrestrial Both

lion, puma

a Classification follows Ewer (1973).

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Phylogeny and Speciation of Felids

APPENDIX 2 DESCRIPTIONS OFKARYLOGICAL AND MORPHOLOGICALCHARACTER STATES

Karyology: Characters 1–10 (Wurster-Hilland Centerwall, 1982; Wurster-Hill et al.,1987; Dutrillaux and Couturier, 1983)

1. Distinctive banding pattern on chromosome A1: 0 5 absent;1 5 present

2. Pericentric inversion of chromosome B4: 0 5 absent; 1 5 present3. Distinctive banding pattern on chromosome D1: 0 5 absent;

1 5 present4. Distal placement of the major band in the short arm in contrast

to proximal placement on chromosome D2: 0 5 proximal; 1 5 distal5. Distinctive feature of chromosome D2: 0 5 absent; 1 5 present6. Conversion of chromosome F2 to E5: 0 5 F2; 1 5 E57. Presence of short arms on chromosome F2: 0 5 absent; 1 5

present8. Conversion of chromosome F3 to E4 to F1: 0 5 F3; 1 5 E4;

2 5 F19. Presence of short arms on chromosome F3: 0 5 absent; 1 5

present10. Fusion of chromosomes F2 and F3 into C3: 0 5 F2 and F3

unfused; 1 5 fused

Morphology: Characters 11–67 (Salles,1992); Character 68 (Peters and Hast,1994)

Cranial Features

11. Anterior part of the dentary: 0 5 intermediate curvature;1 5 reduced curvature; 2 5 marked curvature

Teeth

12. Upper fourth deciduous premolar, parastyle, and metastyleroots: 0 5 bifurcate; 1 5 partially fused; 2 5 totally fused

13. Upper third deciduous premolar, secondary parastyle cusp:0 5 well developed; 1 5 reduced

14. Lower third deciduous premolar, second posterior accessorycusp: 0 5 prominent, cylindrical cone; 1 5 reduced, flattened cone

15. Lower deciduous canine, lateral accessory cusp: 0 5 unicus-pid; 1 5 bicuspid

16. Upper third premolar, parastyle: 0 5 reduced or absent; 1 5

greatly enlarged17. Upper fourth premolar, protocone: 0 5 poorly or moderately

to well developed; 1 5 markedly reduced; 2 5 almost totally absent18. Upper third premolar, metastyle: 0 5 well developed; 1 5 re-

duced19. Upper third premolar, lingual ridge: 0 5 absent; 1 5 present20. Dorsoventral elongation of the upper canine length: 0 5 ab-

sent; 1 5 present

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247

21. Upper canine lingual ridge: 0 5 absent, 1 5 weakly developedridge, 2 5 well-developed ridge

22. Lower canine lingual cavity: 0 5 absent, 1 5 present23. Lower third premolar crown: 0 5 not elongated, 1 5 elongated24. Lower first molar, paraconid crest: 0 5 absent, 1 5 present25. Upper fourth and third premolars, relative positions on the

maxilla: 0 5 P3 and P4 aligned, 1 5 P3 projects laterally in relationto P4

Basicranial Morphology

26. Relative position of foramen rotundum to basicranial plane:0 5 on orbital wall; 1 5 at the basicraniums’s plane

27. External pterygoid fossa: 0 5 poorly to moderately developed;1 5 well developed

28. Palatine bones: 0 5 intermediate inflection; 1 5 reduced in-flection; 2 5 increased inflection

29. Occipital condyles: 0 5 no enlargement; 1 5 enlarged condyles

Auditory Region

30. Subarcuate fossa: 0 5 deep fossa; 1 5 poorly developed;2 5 absent

31. Internal auditory meatus, marginal surface: 0 5 distinctiveborder absent; 1 5 present

32. Longitudinal ridge of auditory meatus: 0 5 absent; 1 5 moder-ately developed ridge; 2 5 well-developed ridge

33. Malleus, processus muscularis: 0 5 thinner and pointed; 1 5

thicker and cylindrical34. Malleus, processus brevis: 0 5 not anteriorly reflected; 1 5

anteriorly reflected35. Incus, inferior head with malleus: 0 5 less prominent projec-

tion; 1 5 prominent projection36. Groove for stylomastoid foramen: 0 5 “small superior part

of the posterior wall of the anterior crus of the ectotympanic fusedwith the squamosal, which blends posteriorly with the mastoid pro-cess and displays an inferior crest quasi-enclosing the groove”;1 5 “lacks the circular aspect of the groove . . . [and] the mastoidprocess does not have an anteroventral projection over the stylomas-toid opening . . . a fissure separates the anterior crus of the ectotym-panic from the complex formed by the mastoid process and thesquamosal”; 2 5 “the posterior wall of the anterior crus of theectotympanic completely fused with the squamosal and forming acontinuous compact lateral surface with the mastoid process” (Salles,1992, p. 28)

Frontal Sinus

37. Frontal sinus, relative position on the skull: 0 5 frontal sinusrestricted to postorbital region; 1 5 centrally positioned; 2 5 anteri-orly positioned

38. Anterodorsal frontal sinus cavity: 0 5 not enlarged; 1 5 en-larged

39. Position of posterior part of the first caudal ethmoturbinatescroll: 0 5 posterior; 1 5 anterior

40. Frontal sinus, volume: 0 5 not enlarged; 1 5 enlarged

41. Frontal bone, outer surface depression: 0 5 absent; 1 5 present42. Frontal bone, lateral expansion: 0 5 none; 1 5 moderate expan-

sion; 2 5 extreme expansion

248

Rostrum43. Frontonasal region, dorsal profile: 0 5 less than 258; 1 5 greater

than 458

44. Frontonasal region, upper nasoturbinal chamber expansion:0 5 absent; 1 5 present

45. Frontonasal region, fossa along sagittal suture: 0 5 absent;1 5 present

46. Frontonasal region, salient ridge along upper orbital border:0 5 absent; 1 5 present

47. Rostral constriction: 0 5 absent; 1 5 moderate; 2 5 extensive48. Nasal curvature: 0 5 convex; 1 5 concave49. Narrow interorbital breadth: 0 5 absent; 1 5 present50. Reduction of the infraorbital foramen: 0 5 absent; 1 5 present51. Maxilla expansion over infraorbital foramen: 0 5 absent;

1 5 present52. Jugal and frontal postorbital processes: 0 5 not fused; 1 5

extended but not fused; 2 5 fused53. Jugal anterior process projected over the infraorbital foramen:

0 5 absent; 1 5 present54. Lower rim of the orbit flattened, enlarged and medially in-

flected: 0 5 absent; 1 5 present; 2 5 present with anterior projection

Other Morphological Features55. Ossification of the hyoid apparatus: 0 5 complete; 1 5 incom-

plete

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Mattern and McLennan

56. Anterior projecting flange on head of fibula: 0 5 absent orpoorly developed; 1 5 well developed

57. Tendon for extensor digitorium longus: 0 5 2; 1 5 tendonoriginates on the lateral epicondyle of the femur, passing througha notch on the head of the tibia and under a flange

58. Olecranon of the ulna: 0 5 lateral side larger; 1 5 medialside larger

59. Reduction of caudal vertebrae: 0 5 absent; 1 5 present60. Pinnae, shape of tip: 0 5 rounded; 1 5 pointed61. Pinnae, ear tufts: 0 5 absent; 1 5 moderately developed;

2 5 greatly developed62. Reduction of the rhinarium: 0 5 absent; 1 5 present63. Reduction of interdigital webbing of hind foot: 0 5 absent;

1 5 present64. Cutaneous lobe protecting retracted claw: 0 5 absent; 1 5

present65. Neck fur, direction of growth reversed: 0 5 grows backward;

1 5 grows forward66. Pupil shape, when contracted: 0 5 narrow slit; 1 5 round67. Spine patch on tongue 0 5 begins closer to tip; 1 5 begins

close to apex68. Thick pad on vocal folds: 0 5 absent; 1 5 present

2Plesiomorphic state for this character is not described in eitherSalles (1992) or Herrington (1986).

Phylogeny and Speciation of Felids 249

APPENDIX 3

Partial Data Matrix Used for Cladistic Analysis (Karyology & Morphology)

Characters

11111111112222222222333333333344444444445555555555666666666Taxa 12345678901234567890123456789012345678901234567890123456789012345678

crocuta 0??????0?0000000000000000000000000000000000000000000000000000000000?elegans 0????????0000000000000000000000000000000000000000000000000000000000?aurata 01010002000000?000000000000000000?0010100000001000001000000000011??0badia ??????????0?????00000000001001000?001010020000100001100000000001???0bengalensis 00010001000000001000000001000000001010100100002000011000000000010000canadensis 0101000000000??01000000000000211000010100110000000000000001120010??0caracal 00010002000000000000000000100210010010100010001000001000000120110??0catus 00000002000?0??00000000000100000010010100210000000001100000101010?00chaus 000000020000000000000000000000?00000101002000010000010000001110100?0colocolo 00000000010?00000010210001000000001010100100002000001000000000010??0concolor 00010001000100000000000000000200000010000100001000001001100000010110geoffroyi 00010000010?000001000000000000000?1010100000002000001000000000010??0guigna ??????????0000000000000000000100001010100100002000001000000000010??0jubatus 01000001001200012000000000020212011110110210000001000001110000100100leo 01010000000101010000100000000201100201000000000000000010000000001111libyca 000000020000000000000000001000000100101002100010000011000001110100?0lynx 010100000000000010000000000000100100101001100000000000000011200100?0manul 00000001000000001?0021011000000001101010021000010001?20001000?010??0margarita 0000000200000000110021000010000001001010021000010000?200000101010??0marmorata 010100000000000?00000000001001000?0010100210010000021000000000010??0nebulosa 01000000102101010001100000000201000210000000000000001000000000010000nigripes 00000002000?00?01100001000100000010020100210000100001200000101010??0onca 01010000000101010000000000000201110200000000000000000010000000001111pardalis 00021000010000000000000000000200001010100100002000001000000000011000pardus 01010000000101010000100000000201100200000000000000000010000000001111planiceps 01010001000000000000010001010100000010100000000010021000000000010??0rubiginosus 000000110000000001002100001000000?0010100200002000011000000000010??0rufus 010100000000000010000000000002100000101001100010000010000011200100?0serval 00010000000000001000000000000000010010100110001000001000000?000100?0silvestris 00000002000?0??01000000000100000010020100210000100001100000101010?00temmincki 101100000000000000000000001001000100101001000020000010000000000100?0tigrinus 00010000010000000000000000000100001010100100002000001000000000010??0tigris 01010000000101010000100000001201000200000000000000000010000000001111uncia 01010000000?????00001000000002010111101010010000001000100000000001?0viverrinus 00010001000?00000000000001000100001010100000001010011000000000010000

wiedii 00021000010000000000000001000200001010100100002000001000000000011??0yagouroundi 00010101000010100000000000000000001010100100102000001001100000010110

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250 Mattern and McLennan

APPENDIX 4

Geographic Distributions of Felid Species

Species Distribution

Panthera groupNeofelis nebulosa Southeast Asia, including Borneo and Himalayan foothillsPanthera uncia Himalayas, Tibetan plateau, mountains surrounding the western border of China into MongoliaPanthera pardus South of the Sahara desert and throughout southern AsiaPanthera tigris Southern and eastern AsiaPanthera onca Central and South America, from southern Mexico to the southern Pampas region, east of the Andes; middle

Pleistocene fossils from West Virginia (Van Valkenburgh et al., 1990) to Bolivia (Hemmer, 1979)Panthera leo Isolated populations from Asia Minor through central India, northern Africa south of the Sahara, including

Europe and the Middle East (through biblical and Roman times)Ocelot group

Leopardus pardalis Southern Texas through Mexico and Central America, most of South America, not west of the Andes orsouth into Argentina and Uruguay

Leopardus wiedii Overlaps completely with Leo. pardalis in the Amazon River BasinLeopardus geoffroyi East of the Andes in ArgentinaOncifelis guigna West of the Andes in Chile (range approximately 15% that of Leo. geoffroyi)Leopardus tigrinus Northern hills of South America and hilly region of southern BrazilLynchailurus colocolo Northern Pampas region of Argentina, overlaps with Leo. tigrinus in central Brazil and the east face of the

Andes in Peru, Bolivia, and northern ArgentinaPuma group

Acinonyx jubatus Central and southern Africa, small population in Iran, previously east to India; fossil members of group incentral North America (Miracinonyx inexpectatus and M. trumani: Martin et al., 1977; Adams, 1979; VanValkenburgh et al., 1990) and Eurasia (Acinonyx pardinensis; reviewed in Van Valkenburgh et al., 1990).

Herpailurus yagouaroundi Mexico through Central America to South America east of the AndesPuma concolor Western North America to southern Argentina with an isolated population in central Florida

Lynx groupLynx lynx Northern EurasiaLynx canadensis Northern North America, previously as far south as Nebraska (Nowak, 1991)Lynx rufus Southern North America, currently sympatric with Lynx canadensis only in the Rocky Mountains

Bay cat groupProfelis temmincki Southeast Asia including BorneoPardofelis badia Borneo

Marbled catPardofelis marmorata Malay Peninsula into Sumatra, Java, and Borneo

Felis groupPrionailurus rubiginosus South of the Himalayas on the Indian subcontinentOtocolobus manul Himalayan Mountains, ranging through the Hindu Kush and into the Tibetan plateauFelis nigripes Restricted to a small area in southern AfricaFelis chaus South east Asia, through India into Iran, south of the Caspian SeaFelis silvestris Northern China and southern Mongolia, western India through Europe along the southern half of AsiaFelis margarita Most of Algeria in Africa, eastern Egypt, Saudi Arabia, and east of the Caspian Sea; overlaps with F. silvestris

around the Caspian SeaFelis libyca All of Africa except the Sahara and the west central rainforest (Congo Basin) and along the coast of Saudi ArabiaFelis catus Cosmopolitan

Leopard cat groupPrionailurus viverrinus Peninsula of South east Asia (into eastern India), the Malay peninsula, and SumatraIctailurus planiceps Widely sympatric with Pri. viverrinus, Borneo and Malay peninsulaPrionailurus bengalensis Borneo and Malay peninsula of South east Asia extending north into eastern China and most of India (except

central region), all south of the HimalayasCaracal group

Leptailurus serval Central to southern AfricaCaracal caracal Africa, middle East, northern India, sympatric with Lep. serval over a large part of its rangeProfelis aurata Central Africa

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Phylogeny and Speciation of Felids

ACKNOWLEDGMENTS

We thank D. R. Brooks for helpful comments on the original draftof this paper and F. Marques for his assistance in using PAUP andMalign. Thanks are also due to two anonymous reviewers and J.Wenzel, whose detailed suggestions greatly improved the clarity

and flow of this paper. This research was funded by a NaturalSciences and Engineering Research Council of Canada grant to D.A.M.

REFERENCES

Adams, D. B. (1979). The cheetah: Native American. Science 205,1155–1157.

Baillie, J., and Groombridge, B. (1996). “1996 IUCN Red List ofThreatened Animals.” International Union for Conservation ofNature and Natural Resources, Gland, Switzerland.

Barton, N. H. (1989). Founder effect speciation. In “Speciation andIts Consequences” (D. Otte and J. A. Endler, Eds.), pp. 229–256.Sinauer, Sunderland, MA.

Brooks, D. R., and McLennan, D. A. (1991). “Phylogeny, Ecologyand Behavior.” Univ. of Chicago Press, Chicago.

Brooks, D. R., and Wiley, E. O. (1988). “Evolution as Entropy: Towarda Unified Theory of Biology,” 2nd ed. Univ. of Chicago Press,Chicago.

Brooks, D. R., Mayden, R. M., and McLennan, D. A. (1992). Phylogenyand biodiversity: Conserving our evolutionary legacy. Trends Ecol.Evol. 7, 55–59.

Brown, J. H., and Gibson, A. C. (1983). “Biogeography.” Mosby, St.Louis, MO.

Bush, G. L. (1975a). Modes of animal speciation. Annu. Rev. Ecol.Syst. 6, 339–364.

Bush, G. L. (1975b). Sympatric speciation in phytophagous parasiticinsects. In “Evolutionary Strategies of Parasitic Insects and Mites”(P. W. Price, Ed.), pp. 187–206. Plenum, New York.

Bush, G. L. (1982). What do we really know about speciation? In“Perspectives on Evolution” (P. Milkman, Ed.), pp. 119–128.Sinauer, Sunderland, MA.

Bush, G. L. (1994). Sympatric speciation in animals: New wine in oldbottles. Trends Ecol. Evol. 9, 285–288.

Caro, T. M. (1994). “Cheetahs of the Serengeti Plains: Group Livingin an Asocial Species.” Univ. of Chicago Press, Chicago.

Carson, H. L. (1975). The genetics of speciation at the diploid level.Am. Nat. 109, 83–92.

Carson, H. L. (1982). Speciation as a major reorganization of polygenicbalances. In “Mechanisms of Speciation” (C. Barigozzi, Ed.), pp.

411–433. A. R. Liss, New York.

Chesser, R. T., and Zink, R. M. (1994). Modes of speciation in birds:A test of Lynch’s method. Evolution 48, 490–497.

Copyright q 2000 by The Willi Hennig SocietyAll rights of reproduction in any form reserved

251

Collier, G. E., and O’Brien, S. J. (1985). A molecular phylogeny ofthe Felidae: Immunological distance. Evolution 39, 473–487.

Cracraft, J. (1982). Geographic differentiation, cladistics and vicariancebiogeography: Reconstructing the tempo and mode of evolution.Am. Zool. 22, 411–424.

Cracraft, J. (1983a). Species concepts and speciation analysis. In “Cur-rent Ornithology” (R. F. Johnston, Ed.), Vol. 1, pp. 159–187. Plenum,New York.

Cracraft, J. (1983b). Cladistic analysis and vicariance biogeography.Amer. Sci. 7, 273–281.

Cracraft, J. (1986). Origin and evolution of continental biotas: Specia-tion and historical congruence within the Australian avifauna. Evolu-tion 40, 977–996.

Daniels, M. J., Balharry, D., Hirst, D., Kitchener, A. C., and Aspinall,R. J. (1998). Morphological and pelage characteristics of wild catsliving in Scotland: Implications for defining the ‘wildcat’. J. Zool.244, 231–247.

Darwin, C. (1859). “The Origin of Species by Means of Natural Selec-tion,” 1st ed. Murray, London.

Diehl, S. R., and Bush, G. L. (1989). The role of habitat preferencein adaptation and speciation. In “Speciation and Its Consequences”(D. Otte and J. A. Endler, Eds.), pp. 345–365. Sinauer, Sunder-land, MA.

Dragoo, J. W., and Honeycutt, R. L. (1997). Systematics of mustelid-like carnivores. J. Mammol. 78, 426–443.

Dutrillaux, B., and Couturier, J. (1983). The ancestral karyotype ofCarnivora: Comparison with that of platyrrhine monkeys. Cytogenet.Cell Genet. 35, 200–208.

Endler, J. A. (1977). Geographic variation, speciation, and clines.Monographs in Population Biology 10. Princeton Univ. Press,Princeton, NJ.

Endler, J. A. (1982). Problems in distinguishing historical from ecologi-cal factors in biogeography. Am. Zool. 22, 441–452.

Essop, M. F., Mda, N., Flamand, J., and Harley, E. H. (1997). Mitochon-drial DNA comparisons between the African wild cat, Europeanwild cat and the domestic cat. S. Afr. J. Wildlife Res. 27, 71–72.

Ewer, R. F. (1973). “The Carnivores.” Cornell Univ. Press, Ithaca, NY.

Farris, J. S. (1970). Methods for computing Wagner trees. Syst. Zool.19, 83–92.

Flynn, J. G. (1996). Carnivoran phylogeny and rates of evolution:Morphological, taxic, and molecular. In “Carnivore Behavior, Ecol-ogy, and Evolution” (J.L. Gittleman, Ed.), Vol. 2, pp. 542–581.Cornell Univ. Press, Ithaca.

Frey, J. K. (1993). Modes of peripheral isolate formation and speciation.Syst. Biol. 42, 373–381.

Gittleman, J. L. (1985). Carnivore body size: Ecological and taxonomiccorrelation. Oecologia 67, 540–554.

Greene, H. W., and Losos, J. B. (1988). Systematics, natural history,and conservation. BioScience 38, 458–462.

Hast, M. H. (1986). The larynx of roaring and non-roaring cats. J.

Anat. 149, 221–222.

Hast, M. H. (1989). The larynx of roaring and non-roaring cats. J.Anat. 163, 117–121.

252

Hemmer, H. (1978). The evolutionary systematics of living Felidae:Present status and current problems. Carnivore 1, 71–79.

Hemmer, H. (1979). Fossil history of the living Felidae. Carnivore2, 58–61.

Hennig, W. (1966). “Phylogenetic Systematics.” Univ. of IllinoisPress, Urbana.

Herrington, S. J. (1986). “Phylogenetic Relationships of the Wild Catsof the World.” Ph.D. thesis, University of Kansas, Lawrence.

Houle, A. (1997). The role of phylogeny and behavioral competitionin the evolution of coexistence among primates. Can. J. Zool. 75,827–846.

Hull, D. L. (1988). “Science as a Process.” Univ. of Chicago Press,Chicago.

Hutchinson, G. E. (1959). Homage to Santa Rosalina, or why are thereso many kinds of animals? Am. Nat. 93, 145–159.

Janczewski, D. N., Modi, W. S., Stephens, J. C., and O’Brien, S.J. (1995). Molecular evolution of mitochondrial 12S rRNA andcytochrome b sequences in the pantherine lineage of Felidae. Mol.Biol. Evol. 12, 690–707.

Johnson, W. E., and O’Brien, S. J. (1997). Phylogenetic reconstructionof the Felidae using 16S rRNA and NADH-5 mitochondrial genes.J. Mol. Evol. 44, S98–S116.

Kawecki, T. J. (1997). Sympatric speciation via habitat specializationdriven by deleterious mutations. Evolution 51, 1751–1763.

Kiltie, R. A. (1984). Size ratios among sympatric neotropical cats.Oecologia 61, 411–416.

Kiltie, R. A. (1988). Interspecific size regularities in tropical felidassemblages. Oecologia 76, 97–105.

Kitchener, A. (1991). “The Natural History of the Wild Cats.” ComstockCornell, Ithaca, NY.

Klicka J., and Zink, R. M. (1997). The importance of recent ice agesin speciation: A failed paradigm. Science 277, 1666–1668.

Kluge, A. J. (1989). A concern for evidence and a phylogenetic hypoth-esis of relationships among Epicrates (Boidae, Serpentes). Syst. Zool.38, 7–25.

Kluge, A. J. (1998). Total evidence or taxonomic congruence: Cladisticsor consensus classification. Cladistics 14, 151–158.

Kobalenko, J. (1997). “Forest Cats of North America.” Firefly Books,Willowdale, Ontario.

Kratochvıl, J. (1975). Os penis of central European Felidae. Zool. Listy24, 289–296.

Kratochvıl, J. (1982). The karyotype and system of the family Felidae(Carnivora, Mammalia). Fol. Zool. 31, 289–304.

Leyhausen, P. (1979). “Cat Behaviour.” Garland STPM Press, NewYork.

Linnaeus (1758). “Systema Naturæ: Sistens Regna Tria Naturæ, inClasses et Ordines, Genera et Species Redacta, Tabulisque ÆneisIllustrata.” Impensis Godofr, Leipzig.

Losos, J. B. (1990). A phylogenetic analysis of character displacementin Caribbean Anolis lizards. Evolution 44, 558–569.

Losos, J. B. (1992). The evolution of convergent structure in CaribbeanAnolis communities. Syst. Biol. 41, 403–420.

Copyright q 2000 by The Willi Hennig SocietyAll rights of reproduction in any form reserved

Mattern and McLennan

Losos, J. B., Warheit, K. I., and Schoener, T. W. (1997). Adaptivedifferentiation following experimental island colonization in Anolislizards. Nature 387, 70–73.

Lynch, J. D. (1989). The gauge of speciation: On the frequency ofmodes of speciation. In “Speciation and Its Consequences” (D. Otteand J. A. Endler, Eds.), pp. 527–553. Sinauer, Sunderland, MA.

MacArthur, R. H., and Levins, R. (1967). The limiting similarity,convergence, and divergence of co-existing species. Am. Nat. 101,377–385.

Maddison, W. P., and Maddison, D. R. (1992). “MacClade: Analysisof Phylogeny and Character Evolution,” Version 3.04. Sinauer, Sun-derland, MA.

Maddison, W. P., Donoghue, M. J., and Maddison, D. R. (1984).Outgroup analysis and parsimony. Syst. Zool. 33, 83–103.

Martin, L. D., Gilbert, B. M., and Adams, D. B. (1977). A cheetah-like cat in the North American Pleistocene. Science 195, 981–982.

Masuda, R., Lopez, J. V., Pecon Slattery, J., Yuhki, N., and O’Brien,S. J. (1996). Molecular phylogeny of mitochondrial cytochrome-band 12S rRNA sequences in the Felidae: Ocelot and domestic catlineages. Mol. Phylogenet. Evol. 6, 351–365.

Mayden, R. M. (1986). Speciose and depauperate phylads and testsof punctuated and gradual evolution: Fact or artifact? Syst. Zool.35, 591–602.

Maynard Smith, J. (1966). Sympatric speciation. Am. Nat. 100,637–650.

Mayr, E. (1954). Change of genetic environment and evolution. In“Evolution as a Process” (J. Huxley, A. C. Hardy, and E. B. Ford,Eds.), pp. 157–180. Allen and Unwin, London.

Mayr, E. (1963). “Animal Species and Evolution.” Harvard Univ.Press, Cambridge.

McLennan, D. A. (1996). Integrating phylogenetic and experimentalanalyses: The volution of male and female nuptial coloration in thestickleback fishes (Gasterosteidae). Syst. Biol. 45, 261–277.

Meier, R., Kores, P., and Darwin, S. (1991). Homoplasy slope ratio:A better measurement of observed homoplasy in cladistic analyses.Syst. Zool. 40, 74–88.

Miththapala, S., Seidensticker, J., and O’Brien, S. J. (1996). Phylogeo-graphic subspecies recognition in leopards (Panthera pardus):Molecular genetic variation. Cons. Biol. 10, 1115–1132.

Nagel, L., and Schluter, D. (1998). Body size, natural selection, andspeciation in sticklebacks. Evolution 52, 209–218.

Nowak, R. M. (1991). “Walker’s Mammals of the World,” 5th ed.,Vol. 2. Johns Hopkins Univ. Press, Baltimore, MD.

O’Brien, S. J. (1994). A role for molecular genetics in biologicalconservation. Proc. Natl. Acad. Sci. USA 91, 5748–5755.

O’Brien, S. J., Collier, G. E., Benveniste, R. E., Nash, W. G., Newman,A. K., Simonson, J. M., Eichelberger, M. A., Seal, U. S., Janssen,D., Bush, M., and Wildt, D. E. (1987). Setting the molecular clockin Felidae: The great cats, Panthera. In “Tigers of the World: TheBiology, Biopolitics, Management and Conservation of an Endan-gered Species” (R. L. Tilson and U. S. Seal, Eds.), pp. 10–27. Noyes,

Park Ridge, NJ.

Ortolani, A., and Caro, T. M. (1996). The adaptive significance of

Phylogeny and Speciation of Felids

color patterns in carnivores: Phylogenetic tests of classic hypotheses.In “Carnivore Behavior, Ecology, and Evolution” (J.L. Gittleman,Ed.), Vol. 2, pp. 132–188. Cornell Univ. Press, Ithaca, NY.

Peters, G., and Hast, M. H. (1994). Hyoid structure, laryngeal anatomy,and vocalization in felids (Mammalia: Carnivora: Felidae). Z. Sauge-tierkunde 59, 87–104.

Pianka, E. R. (1975). Niche relations of desert lizards. In “Ecologyand Evolution of Communities” (M. L. Cody and J. M. Diamond,Eds.), pp. 292–314. Harvard Univ. Press, Cambridge.

Pocock, R. I. (1917). The classification of existing Felidae. Ann. Mag.Nat. Hist. 20, 329–350.

Pocock, R. I. (1932). The marbled cat (Pardofelis marmorata) andsome other Oriental species, with the definition of a new genus ofthe Felidae. Proc. Zool. Soc. London 132, 741–766.

Radinsky, L. (1975). Evolution of the felid brain. Brain Behav. Evol.11, 214–254.

Randi, E., and Ragni, B. (1991). Genetic variability and biochemicalsystematics of domestic and wild cat populations (Felis silvestris:Felidae). J. Mammal. 72, 79–88.

Robinson, R. (1984). Cat. In “Evolution of Domestic Animals” (I. L.Mason, Ed.), pp. 217–224. Longman, London.

Rosen, D. E. (1978). Vicariant patterns and historical explanation inbiogeography. Syst. Zool. 27, 159–188.

Rosenzweig, M. L. (1978). Competitive speciation. Biol. J. Linn. Soc.10, 274–289.

Salles, L. O. (1992). Felid phylogenetics: Extant taxa and skull mor-phology (Felidae, Aeluroidea). Am. Mus. Novit. 3047, 1–67.

Schoener, T. W. (1965). The Anolis lizards of Bimini: Resource parti-tioning in a complex fauna. Ecology 49, 704–726.

Schoener, T. W. (1974). Resource partitioning in ecological communi-ties. Science 185, 27–39.

Schluter, D. (1996). Ecological causes of adaptive radiation. Am. Nat.148, s40–s64.

Schreber, J. C. D. (1774). “Die Saugethiere in Abbildungen nach der

Natur mit Beschreibungen,” Vol. 3. Walther, Erlangen.

Serpell, J. A. (1988). The domestication and history of the cat. In “The

Copyright q 2000 by The Willi Hennig SocietyAll rights of reproduction in any form reserved

253

Domestic Cat: The Biology of Its Behaviour” (D.C. Turner, and P.Bateson, Eds.), pp. 151–158. Cambridge Univ. Press, Cambridge.

Siddall, M. (1997). Prior agreement: Arbitration or arbitrary? Syst.Biol. 46, 765–769.

Swofford, D. L. (1993). “PAUP: Phylogenetic Analysis Using Parsi-mony,” Version 3.1.1. Illinois Natural History Survey, Urbana.

Todd, N. B. (1978). An ecological, behavioral genetic model for thedomestication of the cat. Carnivore 1, 52–60.

Toft, C. A. (1985). Resource partitioning in amphibians and reptiles.Copeia 1985, 1–21.

Van Valkenburgh, B., Grady, F., and Kurten, B. (1990). The Plio-Pleistocene cheetah-like cat Miracinonyx inexpectus of NorthAmerica. J. Vert. Paleontol. 10, 434–454.

Watrous, L. E., and Wheeler, Q. D. (1981). The outgroup comparisonmethod of character analysis. Syst. Zool. 30, 1–11.

Weigel, I. (1961). Das Fellmuster der wildebenden Katzenarten und derHauskatze in vergleichender und stammesgeschichtlicher Hinsicht.Saugetierkunde. Mitt. 9, 1–120.

Wheeler, W., and Gladstein, D. (1994). “Malign,” Version 2.7. Ameri-can Museum of Natural History, New York.

Wiley, E. O. (1978). The evolutionary species concept revisited. Syst.Zool. 27, 17–26.

Wiley, E. O. (1981). “Phylogenetics: The Theory and Practice of Phylo-genetic Systematics.” Wiley–Interscience, New York.

Wilson, D. S., and Turelli, M. J. (1986). Stable underdominance andthe evolutionary invasion of empty niches. Am. Nat. 127, 835–850.

Wozencraft, W. C. (1989). The phylogeny of recent Carnivora. In“Carnivore Behavior, Ecology and Evolution” (J.G. Gittleman, Ed.),pp. 495–535. Cornell Univ. Press, Ithaca, NY.

Wurster-Hill, D. H., and Centerwall, W. R. (1982). The interrelation-ships of chromosome banding patterns in Procyonids, Viverrids andFelids. Cytogenet. Cell Genet. 34, 178–192.

Wurster-Hill, D. H., Doi, T., Izawa, M., and Ono, Y. (1987). Bandedchromosome study of the Iriomote cat. J. Heredity 78, 105–107.

Wyss, A., and Flynn, J. (1993). A phylogenetic analysis and definition

of the Carnivora. In “Mammal Phylogeny: Placentals” (M. Novacekand M. McKenna, Eds.), pp. 32–52. Springer-Verlag, New York.