Phylogeny and evolution of cats (Felidae)

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CHAPTER 2 

Phylogeny and evolution of cats (Felidae) 

Lars Werdelin, Nobuyuki Yamaguchi, lWarren E. Johnson, and 

Stephen J. O’Brien 

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Introduction 

Cats, wild as well as domestic, fossil as well as living, 

are familiar to people around the world. The family 

Felidae has a worldwide distribution and has been 

associated with humans in various ways throughout 

history (Quammen 2004). Their functional morphology, 

ecology, and behaviour have been the subject 

of intense scrutiny by scientists for over 200 

years. The fossil record of cats is extensive and some 

of its members are among the most recognizable of 

extinct animals. Despite all this, the phylogeny and 

evolution of the family Felidae, and even the content 

of the family, have remained poorly understood. In 

this review, we will first present the current state of 

knowledge with regard to the interrelationships 

of living Felidae and the timing of the radiation of 

modern cats. We will also present the fossil record of 

Felidae in broad outline, focusing first on describing 

the different groups of species and their characteristics 

and then discussing the general patterns of cat 

evolution that we can deduce from current data. 

Provided with this overview, we will attempt to identify 

those areas most in need of further research in 

order to achieve the aim of a fuller understanding of 

felid evolution, especially that of the living felids and 

their ecological and functional relationship to the 

extinct sabre-toothed felids. 

In this discussion, we will synthesize the available 

data, distinguishing as far as possible monophyletic 

groups of taxa, suggesting the most likely interrelationships 

of the fossil lineages, but also pointing out 

that there are many problem areas that need to be 

resolved. This section should be viewed as a challenge 

to investigators to use old data or discover 

new data to corroborate or refute the scenarios proposed 

herein. We end the paper with a small section 

demonstrating some evolutionary patterns among 

extant Felidae, suggesting that there is much to be 

gained from the deeper analysis of the current phylogenetic 

information. 

Felid morphology is described and discussed elsewhere 

(Kitchener et al., Chapter 3, this volume) and 

will not be reiterated here except as needed. Teeth of 

the upper jaw are referred to in upper case letters 

(I, C, P, and M) and teeth of the lower jaw in lower 

case letters (i, c, p, and m), followed by the appropriate 

number in the sequence. Character mapping on cladograms 

was carried out with Mesquite, version 1.12 

(Maddison and Maddison 2004). Stratigraphic ages


60 Biology and Conversations of Wild Felids 

of taxa as given in the text and figures were obtained 

from either primary literature or (for North America) 

the Paleobiology Database (www.paleodb.org) and 

(for Eurasia) the NOW database (www.helsinki.fi/science/now/database.html). 

Phylogeny 

Many attempts have been made to investigate the 

interrelationships of Felidae. These have followed 

two broad approaches. Some, like Matthew (1910), 

Kretzoi (1929a, b) and Beaumont (1978) have 

incorporated both fossil and extant felids in their 

analyses, while others, such as Pocock (1917a), Herrington 

(1986), and Salles (1992) have focused exclusively 

on the living members of the family. A new era 

in felid phylogenetics was ushered in with the introduction 

of molecular evidence (Collier and O’Brien 

1985; O’Brien et al. 1985a; Johnson et al. 1996), 

while the first study to use a total evidence approach 

was that of Mattern and McLennan (2000). 

All of these approaches have had their problems. 

In the case of fossil studies, confounding factors have 

included the relatively poor fossil record, the problem 

of finding useful characters in fragmentary material 

and the convergence between Nimravidae and 

Felidae. Though previously included in the Felidae 

(Matthew 1910; Piveteau 1961), the former, Nimravidae, 

is now known to be diphyletic. Its Paleogene 

(65.5–23.0 million years ago [Ma]; Gradstein et al. 

2004) members form a basal clade within either Feliformia 

or Carnivora as a whole (Neff 1983; Hunt 

1987; Morlo et al. 2004), while its Neogene (23.0 

Ma—recent) members are placed in a separate family, 

Barbourofelidae, with affinities to Felidae (see 

below). Morphological studies of extant felids have 

been hampered by the very uniform morphology of 

the members of the family, making it difficult to find 

and polarize characters for phylogenetic analysis. 

Molecular studies, on the other hand, have been 

particularly hampered by the apparently short timespan 

during which the clades of modern felids 

evolved. Thus, clades of closely related taxa have 

been identified but the interrelationships of these 

clades have been difficult to pinpoint. 

Recently, two of us (Warren E. Johnson and Stephen 

J. O’Brien) published a phylogeny of Felidae 

based on a data set of 22,789 base pairs of DNA, 

including autosomal, Y-linked, X-linked, and mitochondrial 

gene segments (Johnson et al. 2006b). The 

results of this study, while not immutable, provide a 

firm basis for understanding the interrelationships 

and evolution of the extant Felidae. The results confirm 

some prior results, both molecular and morphological, 

while providing new insights and surprises. 

The study distinguishes eight clades of extant felids 

(Fig. 2.1). First of these to split off from the stem 

lineage is the Panthera lineage (genera Neofelis and 

Panthera) atc. 10.8 Ma (Fig. 2.1, node A). Most previous 

studies of felid phylogeny have placed Panthera 

as the crown group, but a few (Turner and Antón 

1997; Mattern and McLennan 2000) also have the 

Panthera lineage as basal to other cats. Within this 

35 

A 

Million years before present 

10 

5 

B 

C 

D 

E 

F 

G 

P. linsang 

N. nebulosa 

N. diardi 

P. tigris 

P. uncia 

P. pardus 

P. leo 

P. onca 

P. marmorata 

P. badia 

P. temmincki 

L. serval 

C. caracal 

C. aurata 

L. pardalis 

L. wiedii 

L. colocolo 

L. jacobita 

L. tigrinus 

L. geoffroyi 

L. guigna 

L. rufus 

L. canadensis 

L. pardinus 

L. lynx 

A. jubatus 

P. concolor 

P. yagouaroundi 

F. chaus 

F. nigripes 

F. silvestris 

F. margarita 

O. manul 

P. rubiginosus 

P. planiceps 

P. bengalensis 

P. viverrinus 

Figure 2.1 The phylogeny of the extant Felidae. Thick 

lines indicate the presence of a fossil record, thin lines 

indicate the absence of a fossil record. Node labels as in 

the main text. (Based on the work of Johnson et al. 2006b.)


Phylogeny and evolution of cats (Felidae) 61 

lineage, the clouded leopard, Neofelis, with the two 

species N. nebulosa and N. diardi (Buckley-Beason 

et al. 2006; Kitchener et al. 2006) is placed basally, 

as would be expected from its distinctive morphology 

implying a long separate evolutionary lineage 

(Christiansen 2006), with the rest of the pantherines 

radiating within the last 4 million years. 

The next clade to branch off, at c. 9.4 Ma (Fig. 2.1, 

node B), is the bay cat lineage (genus Pardofelis). This 

clade consists of the poorly known bay cat, Asian 

golden cat and marbled cat. The last mentioned species 

has been linked to the Panthera lineage (e.g. 

Herrington [1986]) and this is reflected in its position 

here, as basal member of the clade branching off 

closest to the Panthera lineage. 

The third lineage is the Caracal lineage, with two 

genera, Caracal and Leptailurus, incorporating three 

African species: caracal, African golden cat, and serval. 

This lineage branches off at c. 8.5 Ma (Fig. 2.1, 

node C), with the serval basal to the other two 

species. 

The next lineage is the ocelot lineage (genus Leopardus), 

including most of the South American small 

cats (Seymour 1999). This lineage branches off at 

c. 8.0 Ma (Fig. 2.1, node D). The beginning of lineage 

is thus independent of the formation of the land 

bridge between South and North America about 3 

Ma (Marshall et al. 1982). However, the radiation of 

the extant species within this lineage shows dates 

that are compatible with a single origin of the extant 

radiation from a North American ancestor, as previously 

proposed (Werdelin 1989). 

The fifth lineage comprises the genus Lynx, 

splitting off at c. 7.2 Ma (Fig. 2.1, node E). This 

lineage has also often been linked to Panthera (e.g. 

Collier and O’Brien [1985]; Salles [1992]), but the 

recent more robust study by Johnson et al. (2006b) 

indicates that the relationship is more distant than 

previously thought. Within the clade, L. rufus is basal 

as has generally been thought, but L. canadensis and 

L. lynx are not reconstructed as sister taxa, unlike in 

previous analyses (Werdelin 1981). 

The next lineage is the Puma lineage, including the 

genera Puma and Acinonyx which split off at c. 6.7 Ma 

(Fig. 2.1, node F). This lineage has previously been 

recognized in both morphological (Herrington 1986, 

Van Valkenburgh et al. 1990) and molecular (Johnson 

and O’Brien 1997) studies. It is worth noting that 

the puma and jaguarundi probably split before the 

Great American Biotic Interchange that followed the 

formation of the land bridge between South and 

North America (Marshall et al. 1982), and thus both 

are of North American origin. 

The seventh and eighth lineages are the small cats 

of the Old World—the leopard cat and domestic cat 

lineages. They split from each other at c. 6.2 Ma (Fig. 

2.1, node G). The former includes the genera Otocolobus 

and Prionailurus and the latter the genus Felis. 

The splits within the former are much deeper than 

within the latter, suggesting that the genus Felis may 

be oversplit. This is also the conclusion of Driscoll 

et al. (2007), who distinguish only four species in Felis: 

F. chaus, F. nigripes, F. margarita, andF. silvestris. The 

last mentioned species now also includes F. ornata, 

F. bieti,andF. lybica, making it one of the most widespread 

small cat species. 

Most of the nodes in this phylogeny are robustly 

supported (Johnson et al. 2006b). A few, however, are 

still unstable, showing either low support or incongruence 

between different analyses and data sets. 

These as yet incompletely resolved nodes: the relative 

positions of Panthera leo, P. pardus, and P. onca,as 

well as the relative positions of P. tigris and P. uncia 

within this clade, the position of L. jacobita, the 

position of O. manul, the position of F. nigripes, and 

the clade uniting Felis and Prionailurus/Otocolobus to 

the exclusion of Puma/Acinonyx. 

The most notable fact about this phylogeny of 

extant cats lies in the short time intervals between 

the splits of the eight lineages. The radiation of 

lineages along the entire stem of the felid clade occurs 

within the Late Miocene (over a period of c. 6.3 

Ma) and such a short space of time suggests the 

occurrence of some sort of functional or ecological 

release, but what that may be is at present unknown. 

We shall return to the fossil record of extant cats 

below. 

The fossil record 

According to available molecular data, the Felidae 

originated some time at or just after the end of the 

Eocene (Gaubert and Véron 2003). This accords well 

with the fossil record. The earliest forms placed in 

the felid lineage, Proailurus and possibly Stenogale



and Haplogale (Hunt 1998; Peigné 1999), occur after 

the ‘Grande Coupure’ marking the Eocene/Oligocene 

boundary (c. 33.9 Ma; Gradstein et al. 2004). 

In the Mammals Paleogene (MP) level system of 

Paleogene terrestrial mammal stratigraphy in Europe, 

this boundary is placed between MP 20 and MP 21 

(Schmidt-Kittler 1990). In the fissure fillings of the 

Quercy region, France, where most of our knowledge 

of early European carnivorans originates, feliforms 

are not known before MP 21 (Hunt 1998). Owing to 

the scarcity of their remains, modern excavations 

have yet to establish the first occurrence of the Felidae. 

What we know, however, suggests that some 

older known finds may be from the Early Oligocene, 

that is, before 28.4 Ma (Gradstein et al. 2004). Thus, 

the earliest felids appeared sometime between c. 35 

Ma (age of the sister group) and 28.5 Ma (minimum 

age of the earliest fossils). 

It is well established on morphological grounds, 

basicranial as well as dental, that Proailurus, known 

from the Quercy fissure fills, but also from excellent 

material from the Early Miocene site of Saint-Gérand-le-Puy, 

France, (Mammal Neogene, MN 2 in the 

Neogene mammal zonation of Europe; 22.8–20 Ma) 

is a felid. Despite this, the morphological path leading 

to the felid condition is not well delineated. Hunt 

(1998) discusses changes to the auditory bulla seen 

in a variety of early feliforms, including Haplogale 

and Stenogale, and leading to the bulla of Proailurus. 

However, the placement of Asiatic linsangs (genus 

Prionodon) as the sister group to Felidae on molecular 

grounds by Gaubert and Véron (2003), instead of 

with the Viverridae, in which they have traditionally 

been placed, adds complexity to the story. Hunt 

(2001) placed Prionodon in a clade with ‘true’ viverrids, 

for example Genetta, on the basis of basicranial 

anatomy (but without consideration of other features). 

What this conflict between separate data sets 

consisting of non-overlapping characters means for 

our understanding of the fossil record of the precursors 

of Felidae and for the origins of the family has 

yet to be established. 

Early felids 

As noted, the earliest well-established felid is Proailurus 

(Figs. 2.2, letter A; 2.3, and 2.4). Peigné (1999) 

provides a discussion of the evolution of this species 

and its relationship to other early putative felids. 

M 

illion 

Y 

ears 

B 

efore 

P 

resent 

5 

10 

Felinae 

(Fig.1) 

Nimravides 

H 

Amphimachairodus 

I 

Machairodus 

F 

G 

K 

J 

 

L 

Megantereon 

 

Metailurus 

M 

 

Vampyrictis 

Dinofelis 

Sansanosmilus 

Xenosmilus 

Smilodon 

Homotherium 

Dinobastis 

Paramachaerodus 

Barbourofelis 

15 

C 

Styriofelis 

D 

Hyperailurictis 

E 

Pseudaelurus 

Syrtosmilus 

Afrosmilus 

20 

25 

Stenogale 

B 

Proailurus 

Ginsburgsmilus 

N 

 

Prosansanosmilus 

30 

Haplogale 

 

 

A 

 

35 

Figure 2.2 Summary of the proposed evolutionary tree of Felidae discussed herein. Thick lines indicate the presence of a 

fossil record, thin lines indicate the absence of a fossil record. Labels as in the main text and Table 2.1. 

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Figure 2.3 The skull of Proailurus lemanensis, MNHN SG 

3509 (holotype) from Saint-Gérand-le-Puy, France, in ventral 

view. The anterior and posterior halves do not meet. (Photo 

courtesy of Stephane Peigné.) 

Proailurus (with three species, P. Lemanensis, P. bourbonnensis, 

and P. major) is a medium-sized cat about 

the size of a bobcat, L. rufus. Dentally, it differs from 

living cats in the (variable) presence of p1, p2, m2, 

and P1, as well as the presence of a small metaconid 

and talonid on m1. Overall the dentition is thus very 

similar to that of living felids, but includes some 

elements that have been fully reduced in the modern 

clade. Further, the auditory bulla of Proailurus 

has a ventral process of the petrosal promontorium 

(Hunt 1989, 1998). This process is lost in living felids. 

When it was lost in felid, evolution has yet to be 

established, but it serves to distinguish at least the 

modern clade from the basally situated Proailurus. 

The geologically youngest Proailurus is from 

Laugnac, France, biostratigraphically placed in MN 

2 billion (>20 Ma). In Proailurus we have (as far as it 

is known) an essentially modern felid except for a 

few minor details of the dentition, auditory bulla, 

and postcranium, which has shorter limbs than 

modern felids. Coupled with the molecular date for 

the divergence of Prionodon and Felidae, this suggests 

that there must have been a stem lineage of perhaps 

5 Ma in the Early Oligocene leading up to the full 

felid morphology. Haplogale and Stenogale are likely 

to be members of that lineage (Hunt 1998; Peigné 

1999), but the details of the process have not been 

worked out. 

Proailurus is not known with certainty outside Europe. 

Hunt (1998) reports the presence of Proailurus 

Figure 2.4 Artist’s reconstruction of Proailurus lemanensis, the first cat. (Illustration courtesy of Mauricio Antón.)



Table 2.1 The internal nodes of Fig. 2.2: content, place of origin, and age. 

Letter Content Continent 

Age, Ma 

(approximate) 

A Felidae sensu stricto Europe 27 

B Pseudaelurine radiation and later Eurasia 22 

felids 

C Felinae (radiation of extant felids Eurasia 14–13 

D Nimravides North America 14 

E Machairodontinae Europe 14–13 

F Amphimachairodus lineage Eurasia 10 

G Homotheriini Eurasia 6 

H Derived Homotheriini Africa 5 

I North American Homotheriini North America 4–3 

J Paramachaeroduslineage Eurasia 11–10 

K Paramachaerodus and derivates Eurasia 9 

L Smilodontini Eurasia, Africa, and North 5–4 

America 

M Metailurini Eurasia 9–8 

N Barbourofelidae Eurasia and Africa 32 

sp. from the Hsanda Gol Formation, Mongolia. However, 

Peigné (1999) concludes, in our opinion correctly, 

that this specimen is better assigned to the 

Barbourofelidae. On the other hand, Hunt (1998) 

also describes the skull of a Proailurus-grade felid 

from the Ginn Quarry, Nebraska (Late Hemingfordian, 

c. 17–16.5 Ma). According to Hunt the basicranial 

structure of the Ginn Quarry felid is more 

plesiomorphic than that of European Proailurus. 

This suggests that phylogenetic diversification in 

Felidae had begun already in the Early Miocene and 

that North American ‘Pseudaelurus’ (see below) may 

have evolved from a Proailurus-grade ancestor rather 

than from a migration of early Pseudaelurus into 

North America. If so, felids may have migrated 

into North America as early as the beginning of the 

Hemingfordian (c. 19 Ma), along with a number of 

other carnivoran taxa (Qiu 2003). 

The next felids to evolve belong to the Pseudaelurus 

complex (Fig. 2.2, letter B; Fig. 2.5). This is a group of 

species with representatives in Europe, Arabia, Asia, 

and North America. The interrelationships of the 

species included in Pseudaelurus and the relationship 

of this genus (or genera) to the radiations of the 

subfamilies Felinae (conical-toothed cats) and Machairodontinae 

(sabretooths) are a major challenge 

to felid palaeontology. Pseudaelurus is clearly a grade 

rather than a monophyletic clade, and this complex 

includes the ancestors of all subsequent felids. A 

number of generic names are available for parts of 

this complex, including Styriofelis, Hyperailurictis, 

Miopanthera, Schizailurus, and Pseudaelurus itself. We 

will consider the validity and applicability of these in 

the discussion below. A fuller knowledge of the interrelationships 

within this group would go a long way 

towards an understanding of the evolutionary patterns 

of the Felidae. 

Pseudaelurus is first recorded from Wintershof- 

West in Germany (MN 3, 20–18 Ma; Dehm 1950). 

Hence, it does not overlap stratigraphically with 

Proailurus in Europe. Several reviews of Pseudaelurus 

have been published in the past decades (Heizmann 

1973; Ginsburg 1983; Rothwell 2003) and we refer to 

them for a fuller discussion of evolutionary details. 

Four species of Pseudaelurus are known from Europe. 

In the order of increasing size they are: P. turnauensis 

(¼ P. transitorius), P. lorteti, P. romieviensis, and 

P. quadridentatus (type species of the genus). They 

range in size from a modern wild cat to a lynx or 

small puma. Differences between them, apart from



Figure 2.5 Artist’s reconstruction of Styriofelis lorteti, a member of the stem lineage leading to the extant Felidae, together 

with the flying squirrel Petaurista sp. (Illustration courtesy of Mauricio Antón.) 

AQ3 

size, are minute (Heizmann 1973). The first species to 

appear is the smallest, P. turnauensis (Dehm 1950). 

However, all three remaining species appear in MN 4 

(18–17 Ma). This indicates a rapid radiation of the 

Pseudaelurus grade, suggesting a monophyletic origin 

of at least European Pseudaelurus from a single species 

of Proailurus. P. lorteti and P. romieviensis become 

extinct at the end of the Middle Miocene (c. 11.6 

Ma), but P. quadridentatus and P. turnauensis survive 

into the Late Miocene (MN 9, c. 11.2–9.5 Ma). They 

thus overlap stratigraphically with the earliest documented 

Machairodontinae (Miomachairodus pseudailuroides 

from Turkey; Schmidt-Kittler 1976; Viranta 

and Werdelin 2003) (Fig. 2.2, letter E). 

Pseudaelurus is poorly known from Asia, possibly 

due to a relative dearth of Middle Miocene localities 

on the continent. Two Chinese species are known. 

Cao et al. (1990) described P. guangheensis from 

Gansu and Wang et al. (1998) describe P. cuspidatus 

from Xinjiang. In addition, Qiu and Gu (1996) describe 

material referred to P. lorteti. All this material is 

Middle Miocene in age. What the relationship is 

between the Chinese and European species has not 

been determined, nor has their relationship to the 

North American radiation of the genus. 

The fossil record of Pseudaelurus in North America 

was recently reviewed by Rothwell (2003). There are 

five valid species: P. validus (stratigraphic range 

c. 17.5–16.5 Ma), P. skinneri (c. 17.5–17.1 Ma), P. intrepidus 

(c. 17.1–13.3 Ma), P. stouti (c. 15.2–12.7 Ma), 

and P. marshi (c. 16.4–12.7 Ma). Thus, Pseudaelurus 

appears later in North America and goes extinct 

sooner there than in Europe. This, and the cladistic 

analysis of Rothwell (2003), in which the three younger 

species (P. intrepidus, P. stouti, and P. marshi) form 

a clade with the two older species (P. validus and 

P. skinneri) as outgroups, are consistent with a single 

origin for North American Pseudaelurus. 

Finally, a single record of P. turnauensis has been 

reported from Saudi Arabia (Thomas et al. 1982) in 

deposits now considered to be of MN 5 age (17.0–15.2 

Ma). Material from Africa previously referred to P. africanus 

(Andrews 1914) is now referred to Afrosmilus, a 

barbourofelid (see Morales et al. [2001] and see below).



The endemic North American genus Nimravides 

undoubtedly originated from one of the above-mentioned 

North American species of Pseudaelurus (Baskin 

1981; Beaumont 1990), probably P. intrepidus or 

P. marshi, which both have a prominent chin, also 

seen in Nimravides (Fig. 2.2, letter D). Nimravides 

differs from its putative ancestors only in relatively 

minor features: it has a more prominent chin, more 

elongated, serrated canines, a more reduced P4 protocone 

and more developed P4 ectoparastyle. These 

are all features pointing towards a sabre-toothed 

morphology, not dissimilar to that seen in M. pseudailuroides 

and Machairodus aphanistus (see below), 

but evolved in parallel. Four species of Nimravides 

are known. N. thinobates (c. 11.0–9.6 Ma), N. pedionomus 

(c. 12.0–11.5 Ma), N. hibbardi (c. 7.0–6.4 Ma), 

and N. galiani (c. 11.6–10.7 Ma). Near the end of the 

Miocene, Nimravides became extinct, apparently 

without leaving descendant lineages. A North American 

felid of uncertain affinities that may possibly 

belong here is Pratifelis martini from the Late Miocene 

(c. 7–6 Ma) of Kansas (Hibbard 1934). This species 

has a distinctively enlarged m1 talonid and does 

not fit comfortably into any of the larger felid 

lineages. 

Sabretooths 

The further evolution of Felidae beyond the Pseudaelurus 

grade begins with M. pseudailuroides (Fig. 2.2, 

letter E). This taxon, which is at present known only 

from Turkey (Schmidt-Kittler 1976; Viranta and Werdelin 

2003), has cheek teeth that are very similar to 

those of P. quadridentatus, but the upper canines are 

more flattened and have small crenulations on the 

mesial and distal faces that are not present in Pseudaelurus 

spp. (Schmidt-Kittler 1976, figs. 114a, 1c, 2, 

and 3, plate 5). In an important contribution, 

Schmidt-Kittler (1976) discusses the relationship between 

M. pseudailuroides and the Pseudaelurus-grade 

and how the morphological transition may have 

occurred. However, he does not pinpoint any specific 

relationships between taxa nor does he extend his 

discussion to conical-toothed cats. M. pseudailuroides 

is at present known only from MN 7/8 and MN 9 

(c. 12.5–9.5 Ma). The taxonomic status of the species 

and genus has been discussed several times. Beaumont 

(1978) made Miomachairodus a subgenus of 

Machairodus, and included Machairodus robinsoni 

from the early Late Miocene (c. MN 9) of Tunisia 

(Kurtén 1976) in the subgenus. On the other hand, 

Ginsburg et al. (1981) synonymized M. pseudailuroides 

with M. aphanistus, type species of the genus 

Machairodus. Morlo (1997) followed this, but suggested 

that M. robinsoni in that case be considered a 

separate genus. This discussion is far from settled, 

but at the very least shows that these forms grade 

into one another. Another early form about which 

there is taxonomic disagreement is M. alberdiae from 

MN 9 of Spain. Ginsburg (1999) considers this to be 

the most primitive Machairodus, but Morlo (1997) 

synonymizes it with M. aphanistus. 

M. aphanistus was described by Kaup (1833) and 

was the first Miocene felid to be named. Its craniodental 

morphology was recently reviewed in detail 

(Antón et al. 2004). These authors found that the 

functional morphology of the killing bite in M. aphanistus, 

and characters related to this behaviour, were 

considerably more primitive than in later machairodonts 

from the Eurasian Late Miocene. They concluded 

that Machairodus should be restricted in 

content to Vallesian (c. 11.2–9.0 Ma) forms, while 

Turolian (c. 9.0–5.3 Ma) forms should be referred to 

Amphimachairodus (Fig. 2.2, letter F). Morlo and Semenov 

(2004) objected to this procedure, arguing that 

the evolution from Machairodus to Amphimachairodus 

was gradual and mosaic and that the two could not be 

generically distinct. However, making the distinction 

is taxonomically useful and in line with a trend in 

recent years of trying to restrict the usage of Machairodus 

to something other than a waste-basket taxon for 

any or all Miocene sabretooths (Beaumont 1978; 

Ginsburg et al. 1981; Ginsburg 1999). 

Some time in the Vallesian Machairodus probably 

migrated to North America, where it gave rise to 

M. coloradensis (c. 9.0–5.3 Ma). This is a fairly 

generalized species, similar to M. aphanistus. It is 

possible, if unlikely, that it evolved from the North 

American Nimravides. This would require extensive 

parallelism with Machairodus. The possibility has 

been noted before, however, and the generic name 

Heterofelis (Cook 1922) is available for this taxon. 

The next stage in the evolution of the machairodont 

lineage is the genus Amphimachairodus (Fig. 2.2, 

letter G). This genus includes a number of closely



related species that morphologically lead up to the 

Plio-Pleistocene tribe Homotheriini (Fig. 2.2, letter 

H), which includes the genera Homotherium, Dinobastis, 

and Xenosmilus. Amphimachairodus includes 

the species A. giganteus (Eurasia; c. 9–5.3 Ma), A. 

kurteni (Kazakhstan; c. 7.1–5.3 Ma), A. kabir (Chad 

and Libya; c. 7–5.5 Ma), and possibly A. irtyschensis 

(Russia; c. 7.1–5.3 Ma), though the latter may be a 

synonym of A. giganteus. Closely related is also Lokotunjailurus 

emageritus (Werdelin 2003b; c. 7.4–5.5 

Ma), which lacks a number of the derived cranial 

features of Amphimachairodus, but is dentally the 

most derived of the group. A. giganteus is, as the 

name implies, characterized by very large size, extremely 

long upper canines and a derived mastoid 

region relative to that of Machairodus, implying modifications 

to the killing bite. The mastoid region is 

further evolved in M. kurteni and M. kabir, but has not 

yet reached the condition seen in Homotherium. Dentally, 

the upper incisor arcade is modified and the 

cheek dentition progressively simplified, with reduction 

of p3/P3, complete loss of the m1 talonid and 

nearly complete loss of the P4 protocone. The dentition 

of L. emageritus is very close to that of primitive 

Homotherium, but the skull and skeleton of the former 

preclude it from the direct ancestry of that species 

(Werdelin 2003b). L. emageritus has an extremely 

enlarged dew claw (absolutely and relative to the 

other claws) on the manus and this feature appears 

to be present also in Homotherium (Ballesio 1963). 

The evolution of Machairodus and Amphimachairodus 

is paralleled in the sabretooth group by the 

evolution of the genus Paramachaerodus (Fig. 2.2, 

letters J and K). At least two and possibly as many 

as four species of this genus are known:P. ogygius 

(c. 9–7 Ma), P. orientalis (c. 8–6 Ma), P. indicus (age 

uncertain) and P. maximiliani (c. 7–5.3 Ma) (Salesa et al. 

2003). The latter two may be synonymous, with each 

other and with P. orientalis. Paramachaerodus is much 

smaller than Machairodus and (especially) Amphimachairodus 

(Paramachaerodus is leopard, rather than 

lion-sized or larger in the case of Amphimachairodus). 

Clearly, this genus and its larger relatives were dividing 

up the prey-spectrum by size, though the details of 

this are not yet understood. New material from the 

early Late Miocene of Spain is doing much to clarify 

the taxonomic, functional, and ecological relationships 

between these Miocene sabretooths (Antón 

et al. 2004; Salesa et al.2005). 

A further lineage that is likely to at least in part 

belong among the sabretooths, despite lacking the 

typical craniodental attributes of this functional 

grade, is the tribe Metailurini (Fig. 2.6). This tribe as 

generally conceived includes the larger genus Dinofelis 

(Fig. 2.6a), with at least ten species (Werdelin 

and Lewis 2001), Metailurus (Fig. 2.6b), with at least 

four species, and Stenailurus, with one species 

(though the latter may be a synonym of Metailurus). 

Dinofelis is in many ways convergent on Panthera, 

but its evolution is not straightforward convergence. 

Instead, various species of Dinofelis are more or 

less pantherine-like, while the oldest and youngest 

species are the most sabretooth like. The Metailurini 

is essentially a waste-basket for taxa that show 

some sabretooth features but can not be placed 

in either the Machairodus or the Paramachaerodus 

lineages. It is not clear that Dinofelis and Metailurus 

are closely related, nor what their respective antecedents 

are. Nor is it clear, although it seems 

likely, that Metailurus is a member of the subfamily 

Figure 2.6 (a) Skull of Dinofelis petteri, KNM ER 2612 

(holotype), Tulu Bor member, Koobi Fora Formation, Kenya; in 

left lateral view. (b) Skull of Metailurus parvulus PIU M3835, 

Locality 108, Baode Province, China; in left lateral view.



Machairodontinae (sabretooth cats). Dinofelis, however, 

shares several traits with derived sabretooths 

and can confidently be placed in this subfamily 

(Werdelin and Lewis 2001). Both of these genera 

originate in the Miocene and survive into the Plio- 

Pleistocene; Metailurus is mainly a Miocene genus, 

while Dinofelis has its main radiation in the Pliocene. 

The Plio-Pleistocene sees the appearance of the 

two derived sabretooth tribes, Homotheriini and 

Smilodontini (Fig. 2.2, letters H and L; Fig. 2.7). The 

Homotheriini includes the genera Dinobastis (with at 

least one species, D. serus) and Xenosmilus (with one 

species, X. hodsonae) from North America and Homotherium 

(Fig. 2.7b) (with several species, including 

H. crenatidens and H. problematicum) from Eurasia 

and Africa. The relationships between these genera 

will be discussed below. The Smilodontini includes 

two genera: Megantereon (with at least five species, 

M. cultridens, M. whitei, M. hesperus, M. falconeri, and 

M. ekidoit) from Africa, Eurasia, and North America; 

and Smilodon (with three species, S. gracilis, S. fatalis 

(Fig. 2.7a), and S. populator) from North, Central, and 

South America. 

Differences between Homotheriini and Smilodontini 

are substantial, both craniodentally and postcranially. 

The Homotheriini have relatively short, 

mediolaterally narrow upper canines with large crenulations 

on the anterior and posterior edges; their 

postcranial skeleton shows some adaptations to a 

cursorial lifestyle (except in Xenosmilus), with long, 

slender limbs and forequarters that are massive but 

not hyperdeveloped. The cheek dentition of Homotheriini 

is dominated by very large carnassials, 

which especially in Homotherium become larger in 

later forms, with the p4 also usurped into the cutting 

blade. The Smilodontini have very long, broad upper 

canines with minute serrations (lost in Megantereon). 

Their skeleton is very robust and the forequarters 

extremely massive. The cheek dentition is reduced 

but the carnassials are not elongated to the extent 

seen in Homotheriini. 

Figure 2.7 (a) Skull (cast) of Smilodon fatalis from Rancho 

La Brea, California, United States; in left lateral view. (b) 

Skull (cast) of Homotherium sp., unknown locality, China; in 

left lateral view. 

Conical-toothed cats 

The conical-toothed cats, subfamily Felinae, comprise 

the common ancestor of all living cats and all 

of its descendants (Fig. 2.8). As the name implies, 

conical-toothed cats differ from sabretooths in having 

a more rounded canine cross-section. They are 

also united by a few other features, such as the relatively 

long lower canine. The interrelationships of 

the living members of this subfamily were discussed 

above. Their fossil history is much less well known 

than that of the sabre-toothed cats. This could be 

for three reasons: (1) They were predominantly 

adapted to environments in which fossilization is 

less likely than in the environments inhabited by 

sabre-toothed cats (i.e. the poor fossil record reflects 

a taphonomic bias; species that today occur in habitats 

in which fossilization potential can be considered 

fair [e.g. cheetahs and lynx], have a reasonably 

good fossil record, while species that today inhabit



Figure 2.8 Skulls of extant Felidae in left lateral view: (a) 

Lion, Panthera leo; (b) Eurasian lynx, Lynx lynx; (c) Domestic 

cat, Felis silvestris catus. 

tropical, wet forests [e.g. golden cats and clouded 

leopards] tend to have a very poor fossil record); (2) 

They were less common in the past than sabretoothed 

cats (i.e. the poor fossil record reflects a 

true pattern that is an outcome of a consideration 

of intra-familial competition between sabre-toothed 

and conical-toothed cats); (3) They are more similar 

to each other in hard-tissue morphology than sabretoothed 

cats (i.e. the poor fossil record reflects a bias 

in investigator perception; there are great similarities 

between all conical-toothed cats in, for example 

mandibular morphology, a region in which sabretoothed 

cats exhibit a number of diagnostic differences). 

All three of these possibilities may be true to 

some extent. Finally, the poor fossil record of conical-toothed 

cats may also reflect the interests of researchers. 

Sabretooth cats are large, spectacular and 

to some extent mysterious, at least as far as their 

feeding behaviour is concerned. Conical-toothed 

cats are often small, nondescript and closely similar 

to living forms that are comparatively well known 

ecologically and functionally. Hence, the former receive 

far more attention in the palaeontological literature 

than the latter. 

Only one researcher, Helmut Hemmer, has focused 

almost exclusively on the fossil record of conical-toothed 

cats, and it is thus from his work (e.g. 

Hemmer [1974, 1976]; Hemmer et al. [2001, 2004]) 

that most of the information on the fossil record of 

this group is to be gleaned. In the following section, 

the fossil record of conical-toothed cats will be outlined, 

following the scheme of eight major lineages 

as found in the molecular phylogeny (Fig. 2.1). 

Focus will be on the earliest members of each lineage 

and/or species. 

Some early conical-toothed cats cannot with confidence 

be included in any of the eight lineages. 

These include the first ‘Felis’, ‘F.’ attica, known from 

MN 11–MN 13 (c. 9.0–5.3 Ma) in western Eurasia. 

This species is a little larger than a wildcat. In morphology 

it is very similar to smaller species of Pseudaelurus, 

but it has a dentition that is reduced 

beyond the Pseudaelurus grade. It is noteworthy 

that the stratigraphic range of ‘F.’attica is younger 

than the estimated age of the base of the radiation 

of extant Felidae (Fig. 2.1), so that it may belong 

within that radiation rather than to the stem lineage. 

The same is true of ‘F.’ christoli, another primitive 

cat, known from MN 13–MN 14 (c. 7.1–4.2 Ma) 

of Spain and France. In addition, there are significant 

collections of Late Miocene small cats from China 

that remain undescribed. This material may answer 

some questions regarding the early evolution of 

extant cats. 

The clade with by far the best fossil record is the 

Panthera lineage. Despite this, it is also the clade with 

the longest ghost lineage (cladistically reconstructed 

lineage undocumented by fossils). According to molecular 

data ( Johnson et al., 2006b) this lineage split 

off from the Felidae stem lineage about 10.8 Ma. 

However, the oldest fossils unequivocally assigned 

to the lineage are no older than 3.8 Ma (Barry 1987; 

Werdelin and Dehghani, in press), leaving a ghost 

lineage that is nearly twice as long as the documented 

lineage. The earliest fossil Panthera from Laetoli 

belong to two species: a lion-sized one and a leopardsized 

one. They have been suggested to belong to the 

AQ4



extant species (Turner 1990), but in fact differ from 

them morphologically (Werdelin and Dehghani, in 

press). The molecular dates suggest that they may 

belong to the stem lineage of these species and 

there is nothing in the fossils that would suggest 

otherwise. 

The first definite lions are from Olduvai, Bed 1 

(



recorded outside Asia (but see Herrington [1987]; 

Groiss [1996]). 

The bay cat lineage is not known with certainty in 

the fossil record. The Caracal lineage is represented 

in the fossil record by specimens dating back c. 4 Ma. 

These specimens group into two distinct size classes, 

large and small. Given the molecular ages of these 

lineages, these may represent members of the caracal/golden 

cat stem lineage and serval stem lineage, 

respectively. Whether any or all of the fossils, which 

are known from a number of sites in eastern and 

southern Africa, are conspecific with the extant 

forms is not determinable on the basis of the available 

material, which consists mainly of isolated teeth 

and fragmentary jaws. An intriguing recent suggestion 

is that ‘Felis’ issiodorensis, a species generally 

referred to the genus Lynx (Werdelin 1981) should 

instead be referred to Caracal (Morales et al. 2003b). 

This conclusion is based on the observation that the 

metric analyses of Werdelin (1981) showed that specimens 

identified as belonging to L. issiodorensis were 

more similar to specimens of Caracal than to specimens 

of Lynx. This possibility deserves further study, 

but it is well to remember that it is just as likely that 

the similarities between Caracal and L. issiodorensis 

are shared ancestral characters. 

The fossil record of the ocelot lineage is relatively 

poor. This record has recently been reviewed by Seymour 

(1999) with updates by Prevosti (2006). The 

South American record of the group is limited, and 

with the exception of some remains of Leopardus colocolo 

from Argentina in sediments dating as far back 

as c. 0.5–1 Ma, and the enigmatic ‘Felis’vorohuensis of 

about the same age, all records are latest Pleistocene in 

age. North American fossils unequivocally referable to 

this lineage are also from the Late Pleistocene (Werdelin 

1985). The inferred age of the radiation of the 

extant taxa at c. 2.9 Ma (Fig. 2.1) is younger than 

previous estimates and compatible with a radiation 

from a single immigration event into South America 

(Werdelin 1989). However, this leaves a long ghost 

lineage back to the reconstructed age of the node 

leading to this group at c. 8.0 Ma. A number of 

North American taxa have been proposed at one 

time or another as members of this ghost lineage, 

including ‘F.’lacustris, ‘F.’rexroadensis, ‘F.’longignathus, 

and ‘F.’proterolyncis (e.g. Werdelin [1985]; Seymour 

[1999]). The first of these is likely to belong to the 

Puma lineage, but the relationships of the others are 

unclear. They may belong to the Lynx or ocelot 

lineages, or be on the backbone of the phylogeny 

between them. The earliest members of several of 

these taxa are Late Miocene (c. 7–6 Ma) in age. 

The short phylogenetic distance between the ocelot 

and Lynx lineages may explain why several taxa mentioned 

above could be assigned to either. The genus 

Lynx is well represented in the fossil record, both in 

Eurasia and North America (Werdelin 1981). In light of 

the above it is likely that the earliest fossil members of 

the lineage are Late Miocene in age. The earliest record 

of unequivocal Lynx in the fossil record has been considered 

to be L. issiodorensis from the Pliocene and 

Pleistocene of western Europe (but see the opinion of 

Morales et al. [2003a], as discussed above). This species 

is not, however, found on the African continent as 

previously suggested (Hendey 1974; Werdelin 1981). 

The only record of the genus on that continent is the 

Pleistocene L. thomasi from Morocco (Geraads 1980). 

The Puma lineage has a long, if uneven, fossil record. 

The oldest fossils unequivocally belonging to 

this lineage are specimens referred to Acinonyx sp. 

from Laetoli (c. 3.7–3.4 Ma) (Barry 1987; Werdelin 

and Dehghani, in press). These specimens are about 

the size of the modern species but differ slightly in 

morphology. The cheetah subsequently has a continuous 

though sparse fossil record in Africa. The genus 

Acinonyx has a long history in Eurasia. The ‘giant’ 

species A. pardinensis appeared in western Europe a 

little over 3 Ma. This form is also found in China (as 

A. pleistocaenicus) and India (as A. brachygnathus). It 

was about the size of a small lion, though considerably 

lighter. In most other respects it displayed typical 

characters of Acinonyx, though the skull does not 

show the extreme vaulting seen in A. jubatus. During 

the later Pliocene there is a marked size reduction 

in Eurasian cheetahs, leading Thenius (1953) to 

describe the younger form as a separate species, 

A. intermedius. However, some Pleistocene specimens 

are as large as the Pliocene ones and we agree with 

Viret (1954) and Kurtén (1968) that the difference 

probably does not warrant specific separation. The 

Eurasian cheetah became extinct in the early Middle 

Pleistocene. The North American ‘cheetah’, Miracinonyx, 

with two species, M. inexpectatus and M. studeri 

(Adams 1979; Van Valkenburgh et al. 1990), is 

not the sister taxon to Acinonyx (Barnett et al. 2005).



Instead, it apparently evolved its cheetah-like features 

independently, from puma-like ancestors. The 

oldest members of this lineage are c. 2.5 Ma. However, 

the oldest ‘F.’lacustris is somewhat older than this. 

An interesting specimen of about the same age is the 

Felis sp. of Gustafson (1978) from the Blancan of 

Oregon, which may also belong to this lineage. The 

presence of Puma in Europe has also been suggested, 

in the form of P. pardoides (Hemmer et al. 2004). The 

oldest of this material is of Pliocene age and may be 

the oldest material of Puma on record. The suggestion 

that Puma is present at Laetoli is hardly tenable, 

however (Werdelin and Dehghani, in press). The oldest 

fossil jaguarundi is less than 0.5 Ma. 

The leopard cat lineage is very poorly known in the 

fossil record. A few fossils probably pertaining to this 

lineage and possibly to Prionailurus bengalensis have 

been found in Middle Pleistocene sites in South-east 

Asia (Hemmer 1976). In addition, fossils tentatively 

referred to O. manul have been recorded from Kamyk, 

Poland (Kurtén 1968). These may be more than 1 Ma. 

The fossil record of the domestic cat lineage is not 

poor, but much of it is hidden beneath the general 

designation of Felis sp., since the species are all but 

indistinguishable on the basis of incomplete remains. 

The oldest ‘Felis sp.’ that definitely belongs to this 

lineage is from Kanapoi, Kenya, dated to >4Ma(Werdelin 

2003a). If the molecular dates are correct, this 

material belongs to a member of the stem lineage of 

Felis. Further specimens belonging to this lineage 

occur intermittently in the African fossil record. A 

species of some interest that may be the oldest member 

of the F. silvestris group is F. lunensis from Europe. 

This species goes back at least to the Early Pleistocene 

and possibly to the Late Pliocene. Specimens referable 

to F. chaus have been found in Holocene strata of Java 

(outside the modern range of the species) (Hemmer 

1976). No specimens definitely referable to F. nigripes 

or F. margarita have been found in the fossil record. 

Nimravidae, a group that itself has been the subject 

of much phylogenetic discussion. The nimravids were 

once known as ‘paleo-felids’ because of their felid-like 

craniodental morphology. They are known from the 

Late Eocene to Late Oligocene of North America and 

Europe and include genera such as Nimravus, Hoplophoneus,andEusmilus. 

Studies of basicranial morphology 

have, however, clearly shown that nimravids are 

not felids (Neff 1983; Hunt 1987). They are therefore 

placed in the family Nimravidae. In its original conception, 

Nimravidae also included the barbourofelids, 

Miocene sabretooths with representatives both in 

North America and Europe (Schultz et al. 1970). 

These, however, have a basicranial morphology, including 

an ossified bulla, that differs from those in 

both Nimravidae and Felidae. Therefore, Morales et al. 

(2001) proposed removing them from the Nimravidae 

and placing them as the subfamily Barbourofelinae 

within the Felidae. This proposal was amended by 

Morlo et al. (2004), who proposed raising Barbourofelinae 

to full family status as the Barbourofelidae, 

which is the path followed here. The Nimravidae are 

likely to be basal Carnivora, while the Barbourofelidae 

are either the sister-group to Felidae or the sistergroup 

to other Aeluroidea (Fig. 2.2, letter N). Because 

of their phylogenetic and ecomorphological closeness, 

to Felidae, their fossil record is outlined here. 

In Africa, the likely centre of origin of Barbourofelidae, 

the family is known from a number of genera 

(Morales et al. 2001; Morlo et al. 2004). Afrosmilus has 

two east African species, A. africanus (Fig. 2.10) and 

A. turkanae, both c. 18–17 Ma. Ginsburgsmilus, the 

most primitive member of the family, has a single 

Barbourofelidae 

Finally, we must touch upon the family (or subfamily) 

Barbourofelidae (Fig. 2.2, letter N), which consists of a 

number of derived sabre-toothed forms (though not 

all may be sabre-toothed—see below). Traditionally, 

they have been seen as Neogene members of the 

Figure 2.10 Left horizontal mandibular ramus of 

Afrosmilus turkanae, KNM MO 15929, Moruorot, Kenya, a 

barbourofelid. Note the well-developed metaconid at the 

posterior end of the tooth—a diagnostic difference between 

Barbourofelidae and Felidae. (Adapted from Morlo et al. 

2004.)



east African species, G. napakensis (c. 20.5–17 Ma). 

Syrtosmilus with one species, S. syrtensis (c. 19–15 

Ma), and Vampyrictis with one, V. vipera (c. 12.5–9.5 

Ma), are North African representatives of the family. 

In Europe, Barbourofelidae is known from several 

genera, Prosansanosmilus, with two species, P. peregrinus 

(MN 4, c. 18–17 Ma) and P. eggeri (MN 5, c. 17–15.2 

Ma), Sansanosmilus with two species, S. palmidens (Fig. 

2.11) (MN 5–MN 7/8, c. 17–11.2 Ma) and S. jourdani 

(MN 6–MN 9, c. 15.2–9.5 Ma), and Afrosmilus with 

one European species, A. hispanicus (MN 5, c. 17–15.2 

Ma). These show a temporal progression towards larger 

and more sabretooth forms, though they are generally 

less extreme in their adaptations than the North 

American Barbourofelis spp. Sansanosmilus is also 

known from the Middle Miocene of China, though 

it is less common there than in Europe. 

In North America, the Barbourofelidae consists of 

the single genus Barbourofelis, with five species, B. 

fricki (c. 10 Ma), B. loveorum (c. 11–9.8 Ma), B. morrisi 

(c. 11.5 Ma), B. osborni (c. 11.5 Ma), and B. whitfordi 

(c. 12–11.5 Ma). They are all extreme sabretooth ecomorphs, 

with long sabres, large mental flanges and 

short, stout limbs (where known). 

Finally, two species from southern Africa must be 

mentioned, Diamantofelis ferox (the size of a small 

puma) and Namafelis minor (lynx-sized) (Morales 

et al. 1998, 2003a). Both are from the late Early– 

Figure 2.11 Artist’s reconstruction of the head of 

Sansanosmilus palmidens, a barbourofelid. (Illustration 

courtesy of Mauricio Antón.) 

earliest Middle Miocene of Arrisdrift, Namibia (c. 

17–15.2 Ma). These species are not, as far as is 

known, sabre-toothed in morphology, as neither 

has a squared-off symphyseal region, but they do 

share other mandibular and dental features with 

species of Afrosmilus. D. ferox has a short and deep 

mandible, while that of N. minor is longer and more 

slender. The oldest true felid from Africa is a small 

specimen from Songhor, Kenya (c. 18–17 Ma), probably 

referable to Pseudaelurus sensu lato. Thus,since 

Felidaeisrareornon-existentinAfricaatthistime, 

whereas Barbourofelidae is known from a number 

of sites and regions, and given the morphological 

similarities between them, it should at least be considered 

whether the Arrisdrift species might be ‘conical-toothed’ 

barbourofelids. 

The Barbourofelidae was a relatively short-lived 

group (c. 20.5–9.5 Ma), within which the vast majority 

of species were specialized sabretooths. Their extinction 

in the early Late Miocene may be tied to the 

spread of sabre-toothed Felidae at this time. 

Discussion 

In this section, we will attempt to draw some conclusions 

from the review above. We advocate the use 

of generic names for fossils that maximizes the number 

of monophyletic taxa by splitting up at least 

those genus-level groups that are obviously para- or 

polyphyletic. In so doing we hope to create a more 

consistent framework for future studies. Unfortunately, 

many of the assertions made in the following 

discussion are at present untested, though we hope 

that it will be possible to test them in the future. We 

aim to erect a series of hypotheses to establish the 

basic level of understanding of felid evolution, that 

of interrelationships. When the interrelationships of 

fossil felids have been better established, a foundation 

for the understanding of the ecological, biogeographical, 

and functional patterns of felid 

evolution will have been laid, in much the same 

way as the current phylogeny of living felids provides 

such a foundation for study of their radiation. 

We will also point out areas where we know too 

little, which is especially true of the fossil record of 

the living felids, which at present has not much to 

contribute to an understanding of the modern



radiation. Johnson et al. (2006b) estimate that fossil 

representation in the modern cat radiation is about 

24%, leaving large areas unknown (cf. Fig. 2.1). Fig. 

2.2 provides a graphical summary of the discussion 

in the following section. 

Early cats 

The origins of the family Felidae are relatively uncontroversial, 

though that may merely be because the 

gap between the earliest unequivocal felids and their 

ancestors among Carnivoramorpha is relatively substantial. 

Thus, Proailurus is unquestionably a felid 

and Stenogale and Haplogale are likely to belong to 

this family as well. All of these genera are undoubtedly 

closer to crown group Felidae than is the extant 

sister taxon, Prionodon. Aspects of this early evolution 

are covered by Hunt (1998) and Peigné (1999) 

and require no further elaboration here. 

The subsequent radiation of Felidae in the Early– 

Middle Miocene is far more complex, however. The 

genus Pseudaelurus comprises 11 named species, 4 

from Europe and Arabia, 2 from China, and 5 from 

North America. Unanswered questions surrounding 

this radiation include: Does Pseudaelurus have a single 

origin What are the interrelationships of the 

European species to each other What is the relationship 

between Chinese and European species From 

which species did North American Pseudaelurus originate 

Which species of Pseudaelurus belong to which 

lineages of later, more derived felids 

Answers to some of these questions have been 

proposed in the past, whereas some have rarely 

been discussed, if at all. An example of the latter is 

the first question raised above: Does Pseudaelurus 

have a single origin This has tacitly been assumed 

in discussions of felid evolution in the past. However, 

the data in favour of this hypothesis are largely 

circumstantial. The presumed ancestor, Proailurus, 

has a limited geographic distribution and Pseudaelurus 

from Europe is older than Pseudaelurus on 

other continents, arguing for a single origin and 

subsequent dispersal. There is, on the other hand, 

no phylogenetic framework in which this has been 

demonstrated to be the most parsimonious hypothesis. 

It is certainly also possible that different species 

of Proailurus or related genera gave rise to different 

species of Pseudaelurus, rendering the latter polyphyletic. 

The Ginn Quarry felid discussed above (Hunt 

1998) makes such a scenario more plausible. At present, 

there does not seem to be any way to resolve 

this issue definitively and the monophyletic origin 

of Pseudaelurus is assumed here as a working 

hypothesis. 

On the other hand, Pseudaelurus is undoubtedly 

paraphyletic, with different species groups giving 

rise to different descendant taxa. The paraphyletic 

nature of Pseudaelurus has been recognized for a 

long time, though perhaps the first to do so explicitly 

was Kretzoi (1929b), and this was also implicitly 

acknowledged by Viret (1951) before being elaborated 

on by Beaumont (1964, 1978). Beaumont 

(1964), like Kretzoi before him, split Pseudaelurus 

into a number of genera at the bases of several 

subsequent radiations. Though Beaumont (1978) reduced 

these to subgenera, his figure 2 remains the 

fullest envisioning of felid evolution to this day. If we 

ignore the subgenera, he split Pseudaelurus into three 

genera: Pseudaelurus (Gervais [1850], type species 

P. quadridentatus), Schizailurus (Viret [1951], type 

species P. lorteti), and Hyperailurictis (Kretzoi [1929b], 

type species P. intrepidus). The first-mentioned includes 

only the type species, while the second 

includes P. turnauensis in addition to P. lorteti. The 

third includes all North American species of Pseudaelurus 

listed above. However, it should be noted that 

Schizailurus is an objective junior synonym of Miopanthera 

Kretzoi (1938) (based on the same type species), 

and this, in turn is a subjective junior synonym 

of Styriofelis Kretzoi (1929a) (type species Felis turnauensis). 

Thus, the latter name is the senior valid 

synonym and is used here. 

Beaumont (1978) places Pseudaelurus at the base of 

the radiation of sabre-toothed cats, Styriofelis at the 

base of the radiation of conical-toothed cats, and 

Hyperailurictis at the base of the North American 

radiation, as well as the radiation of ‘intermediate’ 

forms such as Metailurus, Stenailurus, and Dinofelis. 

The radiation of these three genera takes place at 

letter B in Fig. 2.2. The evidence for this scenario is 

not particularly strong, as it is based mainly on the 

somewhat more sabretooth like characteristics of 

P. quadridentatus as opposed to the clearly conicaltoothed 

features of S. lorteti and S. turnauensis. 

S. lorteti Although the generic separation between 

AQ5



these taxa has generally not been considered in 

reviews of Pseudaelurus, (Heizmann 1973; Ginsburg 

1983), the separation has been implicitly acknowledged 

by several other workers (e.g. Morlo [1997]). 

The status of the North American pseudaelurines 

(Hyperailurictis) as a distinct, generic-level clade is 

somewhat more secure, as these species are relatively 

derived, very similar to each other, and also very 

similar to their presumed descendant Nimravides. 

This scenario provides possible answers to several 

of the questions posed above, apart from the question 

of the relationship of Pseudaelurus to later, more 

derived felids. Among European ‘Pseudaelurus’, S. lorteti 

and S. turnauensis are closely related and more 

distant from P. quadridentatus (the status of P. romieviensis 

is unclear). The North American Hyperailurictis 

did not evolve from any of them, though it may be 

related to one or both of the Chinese species. It is 

more likely, however, that Hyperailurictis descended 

from a felid similar to the Ginn Quarry felid described 

by Hunt (1998). The wholly unanswered 

question is the relationship between European and 

Chinese pseudaelurines. 

A reasonable consensus, however, is that Styriofelis 

gave rise to the radiation of modern cats (Fig. 2.2, 

letter C). Aside from S. lorteti and S. turnauensis, no 

species definitely belonging to the stem lineage are 

known (but see Felis attica and see below). 

In North America there is little doubt that Hyperailurictis 

gave rise to Nimravides (Fig. 2.2, letter D). 

Whether the former is mono- or paraphyletic is not 

known at this time. If it is paraphyletic, further nomenclatural 

complications may arise, but these need 

not concern us here. Nimravides seems to have gone 

extinct without leaving descendants, though it is 

just possible that M. coloradensis evolved from this 

genus rather than being an immigrant from Eurasia. 

The close similarity between M. coloradensis and the 

Eurasian early Late Miocene M. aphanistus argues 

against this, however. 

The relationship between Hyperailurictis and Dinofelis, 

Metailurus and Stenailurus is far less well established 

and is not followed here. These genera are 

usually grouped together as the Metailurini, though 

the monophyly of this tribe has not been satisfactorily 

demonstrated. This is clearly an Old World 

group, with the evolution of Metailurus centred in 

Eurasia and that of Dinofelis in Africa. This presents 

some biogeographic problems with an origin from 

Hyperailurictis in North America as suggested by 

Beaumont (1964, 1978). It is very tempting instead 

to associate this group with the Chinese pseudaelurines, 

though these are so poorly known that this 

remains pure speculation at present. Here we will 

consider the Metailurini to belong to the sabretooth 

cats (but see below), and thus a part of the radiation 

at letter E of Fig. 2.2. 

Upper Miocene to Pleistocene cats 

Pseudaelurus, sensu stricto, gave rise to the radiation of 

sabretooth cats that first appeared in the late Middle 

Miocene of Eurasia (and possibly Africa) and spread 

across the world in the Late Miocene (Fig. 2.2, letter 

E). There has been much controversy surrounding 

sabretooths (subfamily Machairodontinae) and considerable 

confusion regarding taxonomy and the 

allocation of specimens ever since Cuvier (1824) 

placed the first sabretooth specimens in the genus 

Ursus. Numerous genera and species have been 

named over the years and the course of evolution 

of the group has been poorly understood. Part of the 

problem has been the focus on Smilodon, a late and 

highly derived sabretooth, as the exemplar species in 

discussions of the functional morphology and evolution 

of the group (e.g. Bohlin [1940]; Simpson 

[1941]; Miller [1969]; Akersten [1985]). However, 

considerable progress in understanding these issues 

has come in recent years with the study of the excellently 

preserved material from the carnivore trap site 

of Batallones-1 in the Cerro de Batallones, Spain (e.g. 

Antón et al. [2004]; Salesa et al. [2005]). These studies 

show that the functional morphology of sabretooths 

was not uniform across taxa, evolved over time, and 

is compatible with a gradual origin from Pseudaelurus-grade 

forms. 

Nevertheless, numerous questions regarding the 

systematics and evolution of sabretooth cats remain. 

Some of these are: What is the relationship of Dinofelis 

and Metailurus to Machairodontinae What are 

the evolutionary patterns within the paraphyletic 

Amphimachairodus group What is the relationship 

between Homotherium and Dinobastis How did the 

Smilodontini evolve and which taxa are their



ancestors What did sabretooths feed on and how 

How and why did sabretooths become extinct 

The relationship of Dinofelis and Metailurus to Machairodontinae 

(and to each other) has always been 

controversial. Some authors, (e.g. Beaumont [1978]; 

Werdelin; Lewis [2001]), have considered them to be 

members of the Machairodontinae with slight to 

moderate sabretooth adaptations, while others (Kretzoi 

1929b; Hendey 1974) have considered them to be 

conical-toothed cats with a tendency to develop sabretooth 

adaptations. The main feature they share 

with Machairodontinae is a reduced lower canine 

relative to the upper canine. Dinofelis further shares 

with Machairodontinae a deep groove or pit superomedial 

to the trochlear notch of the ulna (Werdelin 

and Lewis 2001). This feature seems not to be present 

in Metailurus (Roussiakis et al. 2006). Thus, it appears 

likely that Dinofelis belongs in the Machairodontinae, 

but the position of Metailurus is equivocal. This 

also, of course, makes the relationship between the 

two genera uncertain. Thus, the position of this 

group at letter J ofFig. 2.2 is problematic, as is the 

placement, even existence, of the node at letter M. 

Amphimachairodus is clearly paraphyletic, as Homotherium 

evolved from within this species group. This 

is reflected in the intermediate position of letter G 

(Fig. 2.2), between letter F where Amphimachairodus 

splits off from a similarly paraphyletic Machairodus, 

and letter H, at the base of the monophyletic Homotheriini. 

What is not clear is exactly which species 

gave rise to Homotheriini (Fig. 2.2, letter H). L. emageritus 

from Kenya has a more derived dentition than 

any species currently assigned to Amphimachairodus, 

but is too primitive in other respects and too derived 

in a few to be the ancestral taxon. Of the species of 

Amphimachairodus, A. kurteni seems the most derived 

dentally, but A. kabir (if the material from Sahabi 

belongs there; cf. Sardella and Werdelin [2007]) has 

the most derived mastoid region. Whichever of these 

(or some as yet unknown taxon) is ancestral to Homotherium, 

Amphimachairodus as presently conceived 

becomes paraphyletic. To resolve this issue, the detailed 

relationships of Amphimachairodus spp. need 

to be better understood. 

The relationship between Homotherium and Dinobastis 

(and Xenosmilus) is particularly interesting 

(Fig. 2.2, letter I). Traditionally, they are synonymized 

in the genus Homotherium (Turner and Antón 

1997). However, early North American homotheriines 

such as that from the Delmont Local Fauna, 

South Dakota (Martin and Harksen 1974) (c. 2.9–2.6 

Ma) differ considerably from contemporary forms in 

Eurasia (see, e.g., Ficcarelli [1979]), suggesting a long, 

separate evolution. In addition, Homotherium and 

Dinobastis differ in a number of aspects of their morphology. 

As an example, the upper canine of Dinobastis 

is smaller than that of Homotherium in 

specimens of approximately equal skull size (Werdelin 

and Sardella 2006, plate 1, fig. 1). This is an area 

that deserves further in-depth study. 

Regardless of which species in the Amphimachairodus 

group is closest to Homotherium, it is nearly 

universally acknowledged that there is, broadly conceived, 

an ancestor–descendant relationship between 

the two genera. However, the origins of the 

other major Plio-Pleistocene sabretooth lineage, the 

Smilodontini (Fig. 2.2, letter L), is much less clear. 

This group consists of the genera Megantereon and 

Smilodon, which share features such as reduced or 

absent serrations on the teeth and extremely long 

and relatively mediolaterally broad upper canines 

compared to Homotheriini (the latter probably a 

plesiomorphic feature). It is tempting to associate 

them with the other Miocene sabretooth lineage, 

Paramachaerodus (Turner and Antón 1997) (Fig. 2.2, 

letter K), but the morphological distance between 

that genus and Plio-Pleistocene Smilodontini is considerable 

and the hypothesized relationship is not 

based on any clear synapomorphies. Another question 

germane to this issue is the difference between 

Smilodontini and Homotheriini: Why is it there and 

what does it mean for the functional morphology 

and ecology of the respective groups One answer 

would be that the former were closed-habitat taxa 

and the latter open-habitat taxa, but can such a simplistic 

view be maintained Martin (1980) and Martin 

et al. (2000) discuss some of these questions, but 

more research needs to be done on the functional 

differences between Homotheriini and Smilodontini, 

and in particular on the latest Miocene species 

of Paramachaerodus (P. orientalis and P. maximiliani), 

to understand their ecology and feeding behaviour 

and whether these can be directly related to those of 

Smilodontini. 

The extinction of Homotheriini and Smilodontini 

occurs at different times on different continents. In



Africa, both Homotherium and Megantereon became 

extinct some time before 1.4 Ma (with Dinofelis lingering 

on another 500 ka). In Europe, Homotherium 

became extinct at c. 0.5 Ma (the recent record of a 

Late Pleistocene Homotherium from North Sea sediments 

(Reumer et al. 2003) needs to be corroborated 

by further material before its implications can be 

fully assessed) and Megantereon at c. 1 Ma. In North 

America, on the other hand, both tribes survive into 

the latest Pleistocene, with the last occurrence of 

Dinobastis from Friesenhahn Cave (Texas) at 

c. 11,000 BP and the last occurrence of Smilodon 

from Rancho La Brea (California) at c. 13,000 BP. 

We don’t fully understand why these dates differ so 

much between continents. The differences may reflect 

the different first appearance datums on each 

continent of advanced hominid competitors in sufficient 

numbers to affect the populations of sabretooths 

through direct or indirect competition for 

resources. Or they could be the result of major faunal 

changes on each continent brought about by human 

interference, climatic change or a combination of 

the two. In building and testing these scenarios, it 

is also important to consider the conical-toothed 

cats and their impact on their sabretooth competitors, 

for example, the relatively rapid range expansion 

of lions from Africa through Eurasia during the 

Middle–Late Pleistocene (Yamaguchi et al. 2004a), 

understanding of which has been hampered by the 

poor fossil record of the conical-toothed cats. 

Possibly no subject in mammal palaeontology has 

been more debated than that of sabretooth feeding 

adaptations. How did they use their canines What 

did they feed on What was their killing behaviour 

like Questions like these have been posed and answered 

numerous times since sabretooths were first 

discovered. (See Kitchener et al., Chapter 3, this volume) 

To answer these questions, it is important to 

realize that this ecomorphology is not restricted to 

felids and their carnivoran relatives among nimravids 

and barbourofelids. The package (with variations) 

is also present in some creodonts, an extinct 

order of mammals that lived from the Paleocene to 

the Miocene (genera Apataelurus and Machaeroides, 

Early–Middle Eocene of North America), marsupials 

(genus Thylacosmilus; Miocene–Early Pleistocene of 

South America) and in various groups of synapsid 

‘reptiles’ of the Late Palaeozoic, for example, Gorgonopsia 

(Kemp 2004). Despite this, it is among felids, 

nimravids, and barbourofelids that the adaptation 

appears to have been most successful. Recent work 

on early felid sabretooths (Salesa et al. 2003; Antón 

et al. 2004; Salesa et al. 2005) has begun to close the 

functional gap between sabre-toothed and conicaltoothed 

cats. This and other lines of evidence, such 

as the meandering evolutionary history of Dinofelis 

from more sabretooth to less sabretooth and back 

(Werdelin and Lewis 2001), suggest that the ecomorphology 

of the feeding apparatus in felids is more of 

a continuum than a dichotomy. The implications of 

this for understanding the ecology of sabretooths 

and competition between sabretooths and conicaltoothed 

cats are in need of detailed investigation. 

One possible implication of the feeding apparatus 

of sabre-toothed and conical-toothed cats being on a 

continuum is that there may have been more direct 

competition between the two groups than previously 

thought. Previous models tend to emphasize the 

difference, with sabretooths specializing in larger 

prey than similar-sized conical-toothed cats. However, 

more recent analyses suggest that perhaps the two 

groups focused on very similar prey. In Africa, sabretooths 

are fairly common fossils and conical-toothed 

cats rare until around the time when the number of 

fossils of sabretooths decreases (Werdelin and Lewis 

2005). This can be explained if sabretooths were 

dominant in the most commonly sampled habitats 

and competitively excluded conical-toothed cats. 

Support for such an idea can be found at Laetoli. 

This site (or at least the Laetolil Member, Upper 

Beds) is unique among eastern African sites in not 

being near a large body of standing water. It is also 

unique among sites in having a large number of 

fossils of conical-toothed cats and very few fossils of 

sabretooths. Further research on competition between 

sabretooths and conical-toothed cats is needed, 

as is research on the competitive structure of the 

carnivore guild as a whole. 

The single most important issue impeding an 

increased understanding of the evolution of conical-toothed 

cats is the extensive ghost lineage between 

the oldest fossil members of the Panthera 

lineage and the common ancestor of all Felinae. 

Two explanations for this gap in the fossil record 

immediately spring to mind: a poor fossil record in 

the earliest Pliocene and the possibility that the



Panthera lineage (and the Felinae as a whole) evolved 

in an environment that is not conducive to the process 

of fossilization. Both of these factors are undoubtedly 

in play, but it is hard to escape the 

impression that pantherines are present in the fossil 

record prior to 4 Ma, but that they are misidentified 

for as yet unknown reasons. To either identify these 

fossils or explain why they have not been found is 

the most pressing issue in felid palaeontology and 

evolution and without progress here it will not be 

possible to move towards a fuller reconciliation of 

the fossil record with the molecular evidence for felid 

evolution as presented by Johnson et al. (2006b). 

The position of Barbourofelidae is, of course, very 

uncertain, since there is no consensus at present on 

how closely related it is to the Felidae. Here, we have 

opted for the view that it split off from the stem 

lineage leading to Felidae after Prionodon but before 

the evolution of Proailurus (Fig. 2.2, letter N). This 

leaves an extensive barbourofelid stem lineage that is 

at present entirely unknown. 

Evolutionary patterns 

The availability of a phylogeny of extant Felidae 

makes it possible to consider evolutionary patterns 

within the family in the absence of fossils. Such 

studies have been attempted in the past (Ortolani 

and Caro 1996; Werdelin and Olsson 1997; Ortolani 

1999; Mattern and McLennan 2000), but given that 

the current phylogeny (Fig. 2.1) is fully resolved and, 

we believe, better corroborated than older hypotheses, 

this work is worth reconsidering. Further, since 

the current hypothesis is based on molecular data, it 

is possible to study morphological character evolution 

without the need to discuss possible circularity 

in the results. Some such uses of the phylogeny were 

presented by Johnson et al. (2006b) and O’Brien and 

Johnson (2007) (intercontinental migrations, ghost 

lineage analysis), and we will only briefly present two 

further examples of the sort of work that can and 

should be done on felid evolution using the phylogeny 

as a baseline. For other examples based on previously 

proposed phylogenetic hypotheses, see in 

particular Mattern and McLennan (2000). 

Werdelin and Olsson (1997) presented a phylogenetic 

study of coat patterns in Felidae using a selection 

of then-current phylogenetic hypotheses as the 

baseline. Their conclusion was that ‘most transformations 

of coat pattern originate from the flecked 

pattern, which we consider to be primitive for the 

Felidae as a whole’ (Werdelin and Olsson 1997, 

p. 399). The current phylogeny has some substantial 

differences from the phylogenies used in that study, 

so the question arises whether the conclusions hold 

up. Fig. 2.12 shows coat pattern mapped on the 

current phylogeny. The data are identical to those 

in Werdelin and Olsson (1997) except for P. tigris, 

which has been recoded from vertical stripes to rosettes, 

as we believe that what appear to be vertical 

stripes in the tiger’s coat in reality are enormously 

vertically elongated rosettes. This is indicated 

through examination of various coat pattern anomalies 

in tigers and can be more simply seen by holding 

up an image of a tiger pelt nearly parallel to one’s line 

of sight. One difference from the previous results is 

immediately obvious: under the current phylogeny, 

the primitive coat pattern for Felidae as a whole is 

large blotches. This coat pattern is present in only 

two genera:Neofelis, clouded leopards and Pardofelis, 

marbled cat. Both are basal within their clades, and 

these clades are basal within the family and hence 

the primitive condition is reconstructed as large 

blotches. Above the node leading to Pardofelis, however, 

flecks are primitive as they were in the previous 

study. If we consider the number and direction of the 

state changes in the cladogram (Fig. 2.13), we can 

also see that changes to and from flecks are still the 

dominant transformations, though not quite as 

dominant as previously thought. Thus, the new phylogeny 

corroborates the main thrust of the results of 

Werdelin and Olsson (1997), but also leads to some 

modifications of specific parts of their conclusions. 

In a second demonstration of possible phylogenetic 

reconstructions, we mapped habitat (as open or 

closed), activity pattern (diurnal or nocturnal), and 

pupil shape (slit-like or rounded in the contracted 

state) in all felids. The data are partly from Mattern 

and McLennan (2000) and partly original. Many species 

occur in both open and closed habitats and the 

mapping reflects this, not showing any clear phylogenetic 

associations of open- or closed-habitat specialists 

(Fig. 2.14), although the Panthera and domestic cat 

lineages are dominated by open-habitat taxa and 

have this habitat reconstructed as primitive for the



Rosettes 

Large blotches 

Uniform 

Flecks 

Small blotches 

Stripes 

Polymorphic 

Prionodon linsang 

Neofelis nebulosa 

Panthera tigris 

P. uncia 

P. pardus 

P. leo 

P. onca 

Pardofelis marmorata 

P. badia 

P. temmincki 

Leptailurus serval 

Caracal caracal 

C. aurata 

Leopardus pardalis 

L. wiedii 

L. colocolo 

L. jacobita 

L. tigrinus 


L. guigna 

Lynx rufus 

L. canadensis 

L. pardinus 

L. lynx 

Acinonyx jubatus 

Puma concolor 


Felis chaus 

F. nigripes 

F. silvestris 

F. margarita 

Otocolobus manul 

Prionailurus rubiginosus 

P. planiceps 


P. viverrinus 

Figure 2.12 Coat patterns (as labelled) mapped on the phylogeny of extant Felidae. 

respective clades. Likewise, there are no clear phylogenetic 

patterns underlying activity patterns in modern 

felids (mapping not shown). Rounded pupils, on the 

other hand, only occur in three clades, the Pantheralineage, 

where all species except the two Neofelis (A. 

Kitchener, personal communication) have rounded 

pupils, the Puma-lineage, where all three species have 

rounded pupils, and the leopard cat lineage, where the 

single species O. manul has rounded pupils. 

The question of the occurrence and causes behind 

slit-like or rounded pupils has been intermittently 

discussed in the literature without a consensus 

being reached (see Chapter 3 (this volume) herein 

for a discussion of some recent research). One suggestion 

that has been considered is that slit-like pupils 

allow the pupil to be more completely closed 

than rounded pupils (Walls 1942). This would suggest 

that the former would be more useful in the



1 

0,2 

1 

2–4 

1-2 

0–1 

0–1 

3 

0–1 

Figure 2.13 The coat pattern transformations implied by the mapping in Fig. 2.12. The majority of transformations 

involve flecks (center pattern). Thus, clockwise from bottom, there are three transformations from flecks to uniform, 

one transformation from large blotches to flecks, two to four transformations from flecks to stripes, zero or two (no 

reconstruction allows for one) transformations from striped to flecks, one to two transformations from flecks to small 

blotches, and zero to one transformations from small blotches to flecks. Remaining reconstructed transformations are 

zero to one transformation from large blotches to uniform, one transformation from large blotches to rosettes, and 

zero or one transformations from small blotches to stripes. No other transformations are allowed by the phylogeny of extant 

Felidae. 

brighter light of day, that is, slit-like pupils should be 

preferentially present in diurnal species. However, a 

comparison between the patterns does not corroborate 

this idea (not shown). There seems to be no 

correlation at all between pupil shape and activity 

pattern. However, if we compare habitat and pupil 

shape (Fig. 2.14), we find that with the exception of 

Puma yagouaroundi, which, if the fossil record of this 

clade is taken into account, must be considered secondarily 

adapted to closed habitats, rounded pupils 

never occur in closed-habitat specialist species. All 

the taxa with rounded pupils are either open-habitat 

species or occur in a variety of habitats. Further, all 

three nodes where there is a change from slit-like to 

rounded pupils are also nodes where there is a shift 

from closed to open habitat preference. What this 

means in functional terms is beyond the scope of 

this chapter, but the results point to a fruitful avenue 

of research. These very tentative results must be corroborated 

by more in-depth study and statistical testing. 

More generally, phylogenetically based studies 

such as the ones discussed above can direct future 

research and provide tests of functional hypotheses 

that could otherwise not be investigated due to a lack 

of independent data. The existence of a well-corroborated 

phylogeny such as that in Fig. 2.1 is a powerful 

tool for future research on felid evolution. 

Final words 

This chapter presents one possible scenario for the 

evolution and interrelationships of cats. Some of this 

work, such as that which has led to the phylogeny of 

Johnson et al. (2006b), is strongly corroborated by 

and based on considerable amounts of data. The 

fossil record of Felidae is uneven. Some groups, 

such as parts of the Machairodontinae have a fairly 

extensive fossil record, while others, such as the lineage 

leading to the extant radiation, are much more



Open habitats 

Closed habitats 

Prionodon linsang 

Neofelis nebulosa 

Panthera tigris 

P. uncia 

P. pardus 

P. leo 

P. onca 

Pardofelis marmorata 

P. badia 

P. temmincki 

Leptailurus serval 

Caracal caracal 

C. aurata 

Leopardus pardalis 

L. wiedii 

L. colocolo 

L. jacobita 

L. tigrinus 


L. guigna 

Lynx rufus 

L. canadensis 

L. pardinus 

L. lynx 

Acinonyx jubatus 

Puma concolor 


Felis chaus 

F. nigripes 

F. silvestris 

F. margarita 

Otocolobus manul 

Prionailurus rubiginosus 

P. planiceps 


P. viverrinus 

Slit-like pupils 

Round pupils 

Figure 2.14 Habitat type preference and pupil shape mapped on the phylogeny of extant Felidae. 

poorly documented. In no case, however, can the 

fossil record be said to be adequate, either in quantity 

or quality. Nor can the fossil record of Felidae be said 

to have been adequately studied. Some areas, such as 

the functional morphology of sabretooths, have 

been investigated over and over, while others, such 

as the stem lineage of modern cats, have been relatively 

neglected. Overall, the phylogeny and evolution 

of fossil Felidae have been neglected in favour of 

studies of their functional morphology and ecology. 

Given the limited resources available for this work, 

this is understandable, as the latter topics have proven 

more tractable and have yielded interesting and 

significant results. But if our understanding of the 

group is to progress, we must try to address such 

pressing issues as the fossil record of living cats, the 

origins of Smilodontini and the relationship of Barbourofelidae 

to Felidae. This will require extending 

the work of Johnson et al. (2006b) into the realm of 

fossils, by comparing the fossil record with the results 

obtained from the phylogeny of extant cats on 

aspects such as continental migration (O’Brien and 

Johnson 2007) to see if the timing of intercontinental 

migrations of fossil cat groups can be matched up 

with those postulated for the extant cats based on 

phylogeny and geology. 

It must be understood that developing a phylogeny, 

or even the simpler task of testing some aspect of 

the scenario developed herein, requires more than a 

superficial glance at the record and doing a phylogenetic 

analysis of the first few characters that come to 

mind. It will require developing new characters and 

looking at the fossil record in new ways. If the fossil 

record and phylogeny of extant Felidae can be better 

integrated, we can expect to develop a significantly 

better understanding of the evolution of this



fascinating group and its conditions for existing, 

thereby not only enhancing current knowledge, but 

also building a better platform for the conservation 

of the many endangered species of Felidae today. 

Acknowledgements 

We would like to thank David Macdonald for 

inviting us to contribute to this volume, as well as 

for his encouragement and a careful reading of the 

manuscript. We thank Mauricio Antón for letting us 

use his reconstructions of extinct felids, Stéphane 

Peigné for the photographs of Proailurus, Mikael Axelsson 

for photographs of felid skulls, Mats Wedin for 

help with MacClade, and Margaret Lewis and Susan 

Cheyne for reading and commenting on the manuscript. 

Lars Werdelis acknowledges support from the 

Swedish Research Council. We thank Andrew Loveridge, 

Blaire van Valkenburgh, Andrew Kitchener and 

an anonymous reviewer for their thorough readings 

that helped us clarify and correct parts of the text, 

making the whole far more accessible to a broad 

audience than it otherwise would have been. 

Author Queries: 

AQ1. Please provide the Caption for the introductory figure. 

AQ2. Please specify whether “” can be changed to “” in the Fig. 2.2. Also, can the citation “Felinae (Fig. 1)” 

be changed to “Felinae (Fig 2.1)” in the Fig. 2.2. 

AQ3. Please check the genus name in Styriofelis lorteti. 

AQ4. Please update Werdelin and Dehghani (in press). 

AQ5. Please check expand the genus name in “S. Lorteti and S. turnauensis”.

Phylogeny and evolution of cats (Felidae)

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