Papers

The family of cats—delineation of the

feline basic type1
Barnabas Pendragon and Niko Winkler
Based on previously reported hybridizations, cats have long been considered to belong to a single basic type.2
However, there has been discussion concerning whether great cats and small cats might represent independent
sister clades. Recent DNA sequencing data confirm that such distinctions are not fundamental in nature and
that all cats share a common genetic ancestry. More recently described hybridizations between great cats and
small cats, along with various other studies described in the present article, further support the hypothesis that
all cats belong to a single clearly delineated basic type. The Nimravidae (paleosabers), the Machairodontinae
(neosabers) and the genus Panthera, each underwent a prominent radiation during the tertiary period. All three
taxa represent cat-like placental carnivores, and they may all have arisen from the same basic type.

Cats and their taxonomic position with numerous, horny papillae that are directed backwards;
digitigrade extremities with five toes on the forefoot and
T he cat family is placed within the order Carnivora,
which comprises nine extant families (or ten if
mongooses are considered a separate family; Herpestidae).
four on the hindfoot; claws that are sharp, strongly curved,
and usually highly retractile, protected by a fleshy sheath
(except in the genus Acinonyx, the cheetahs).4
The carnivores are grouped into two suborders: the cat-
like carnivores, or Feliformia, including the Felidae (cats),
One or two family histories?
the Hyaenidae (hyenas), the Viveridae (civets), and the
Herpestidae (mongooses); and the dog-like carnivores, The earliest cat-like carnivore family, the Nimravidae
or Caniformia, including the Canidae (dogs), the Ursidae (paleosabers), includes two lineages that suddenly appear
(bears), the Procyonidae (raccoons), and the Mustelidae in the late Eocene jungles of North America. One lineage,
(weasels), as well as two marine families, the Otariidae represented by Hoplophoneus, was saber-toothed; the other
(sea lions) and the Phocidae (seals). The present role of lineage, represented by Dinictis, was rather like a modern
carnivores in nature is regulatory, keeping in check the serval. So convincingly cat-like are these fossils that
numbers of herbivores. They are assumed to indirectly help originally they were called paleofelids and designated the
maintain healthy populations of herbivores by selectively first cats.5 The more recent cat family, the Felidae, include the
devouring non-healthy and phenotypically disadvantaged modern cats, Felinae; and the neosabers, Machairodontinae,
animals. The carnivores share a relatively homogeneous e.g. Smilodon.6 The Nimravidae and the Felidae display
phenotype. Many are capable of running quickly, possess only modest skeletal differences. Nevertheless, they are
conspicuous canine teeth often used for catching and killing placed into two distinct families. The most prominent
prey, and display the carnivore-typical carnassial teeth, skeletal difference is that nimravid fossils lack a bony wall
which include the last premolars of the upper jaw and the (septum) in their middle-ear chambers (auditory bullae) or
first molars of the lower jaw. Instead of having a grinding sometimes the whole chamber, implying these structures
surface, these teeth have a flattened, razor-like crown used were cartilaginous.7 The nimravid, Barbourofelis, which
for slicing through muscle tissue when devouring prey. In lived contemporaneously with the Felidae, had ossified
the omnivorous carnivores, such as bears, true carnassial auditory bullae, but there is no evidence of a bony septum.8
teeth do not develop. Such morphogenetic differences require modest selective
The 38 species of extant cat have a very characteristic change. Indeed even in extant cats the auditory bullae are
phenotype readily distinguished from other species of first cartilaginous and only later ossify. As versatile as
animals, even by laymen. Recently, the clouded leopard ossification of the auditory bullae has proven to be in helping
(Neofelis nebulosa) was separated into two species, which, define extant cat species, calling an extinct animal a non-cat
if acknowledged, brings the total number of species to 39.3 that is otherwise clearly a cat, simply for want of evidence
They possess a lithe, muscular, compact and deep-chested of such ossification, seems excessive.
body. Technical diagnostics include: pointed, elongate The Nimravidae-Felidae ‘two-family’ hypothesis is
canine teeth; large carnassials, strongly shearing in function; supported less by ossifications and more by phylogenetic
the dental formula 3/3, 1/1, 2-3/2, 1/1; ossified auditory considerations; specifically, that the Felidae arose from
bullae, inflated in appearance and divided by a bilaminate Proailurus during the Miocene in the Old World. 8
septum (except Leopardus jacobita, the Andean mountain Proailurus, the animal currently nominated the first ‘true’
cat, which has a double-chambered bulla);4 a tongue covered cat, was short, only 15 cm in length. It had very many

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civet-like features and was probably plantigrade (walking on

the soles of its feet) and not digitigrade (walking on its toes)
like cats today.9 Perhaps the need for a link to a postulated
‘common-civet ancestor’ is the primary motivation for
trying to classify Proailurus among the cats. It had been
previously classified among the civets (Viverridae) and
there is much to support this classification still; plantigrade
gait, flatter skull, extra teeth, the bones of Proailurus are
very similar to the living viverrid, Cryptoprocta, the fossa
of Madagascar.2,5,7 The next most recent cat, Pseudaelurus,
had a clearly cat-like skull, based on its general morphology
and dentition. It roamed Europe and North American during
the Miocene. Several different forms of Pseudaelurus
existed and these are believed to have formed the basis for
the later radiation and diversification of the neofelid cats
and the sabretooth tigers.9 Proailurus, from the Miocene,
is the reason why New World paleosabers from the Eocene Figure 1. The anatomy of the feline hyoid apparatus and its
‘complete’ ossification. The hyoid apparatus joins the larynx (La)
cannot be cats. As a result the origins of the paleosabers to the middle ear capsules (M = auditory bullae) via two parallel
have had to be placed elsewhere; e.g. within the dog-like chains of seven ‘bones’. The fifth bone links the two chains, forming
carnivores10 or even outside the carnivores11. It is easy to the central basihyoid. The first and seventh members of the chains
are always cartilaginous and are not displayed in the figure. Tr =
see how the two-family hypothesis could be specious, based trachea. (After Dahm in Peters and Hast, ref. 13.)
on preconceived notions.
Differences in ossification of skeletal features are not were replaced by woodland savannas. 15 These more
necessarily diagnostic of family status. A case in point is the open forest conditions resulted in adaptive radiations in
early attempt to separate small cats and big cats into separate herbivore groups including oreodonts, anchitheres and small
subfamilies based on ossifications in their hyoid apparatus rhinocerotoids (Janis 1998). Since these were the principle
and their ability to roar.7 Sir Richard Owen12 was the first prey of the Nimravidae, adaptive radiation in the herbivores
to explore hyoid anatomy and felid vocalization. The hyoid would account for the adaptive radiation observed in the
apparatus consists of two chains of seven bones, joined paleosabers during that period. Similarly, at the Miocene/
at the fifth (basihyoid). These stretch from the auditory Pliocene transition, grassland savannas were replaced by
bullae to the larynx, support the voice box, and confer prairies.15 Adaptive radiations of cursorial herbivore groups,
characteristic vocalizations (see figure 1). In small cats (the including camelids, antilocaprines, and equines was then
felines) the third bones (epihyoids) are ossified and short, accompanied by adaptive radiation of their predators, the
and they cannot roar. In the larger big cats (the pantherines) Felidae.16 Today both the paleosabers (Nimravidae) and
the third epihyoids are cartilaginous and extended, and all neosabers (Machairodontinae) are completely extinct.
but one of these cats can roar. In snow leopards (P. uncia; Only in the extant clouded leopard (Neofelis nebulosa) can
figure 2) the epihyoids are cartilaginous; however, they still a vestige of the saber tooth phenotype still be observed;
cannot roar because they lack additional pads in their vocal
although both the upper and the lower canines are
folds.13,14 Ossification of the seven hyoid bones in felids is
pronounced, this is a unique occurrence in the cat family.
never complete. Both the first (tympanohyoid) and seventh
Why these animals became extinct is something of a
(chondrohyoid) ‘bones’ are cartilaginous. These attach to
mystery. These large carnivores would have required large
the auditory bullae and cartilaginous larynx, respectively.13
prey; saber teeth are inefficient for capturing and killing
Bone ossifications are of less taxonomic importance than
small prey. Biome change may have led to an imbalance
genetic and hybridization data, which support the conclusion
in predator–prey relations, resulting both in the loss of
that small cats and big cats belong in the same subfamily.
sustainable populations of large prey and in the extinction
It is submitted that lack of ossification of the auditory
of these remarkable cats.
bullae in the Nimravidae is similarly of questionable
diagnostic use for the family’s phylogenetic position
Pelage pattern—a common origin
and that the Nimravidae represent at least a sister clade
of the Felidae, if not actual members of the same basic The development of melanistic coat colour patterns
type. Their distributions reflected the dominant biomes. within the cat family is not entirely understood, but some
During the course of the Tertiary period, gradual cooling interesting conclusions can be drawn. Weigel17 believed
and drying caused major shifts in these biomes. During that all of the extant melanistic coat colors and patterns
the Eocene/Oligocene transition, subtropical rainforests arose from a single original type of relatively large dark

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Phylogenetic Code Sci. Name English Name Lineage

Fca: Felis catus Domestic cat

Fsi: F. Silvestris European Wildcat
Fli: F. Libyca African Wildcat
Fbi: F. bieti Chinese Desert cat 8. Domestic cat line
Fma: F. margarita Desert Cat
Fni: F. nigripes Black-footed cat
Fch: F. chaus Jungle Cat

Oma: Otocolobus manul Pallas cat

Pru: Prionailurus rubiginosus Rusty spotted cat
Pbe: P. bengalensis Asian leopard cat 7. Leopard cat line
Pvi: P. viverrinus Fishing cat
Ppi: P. planiceps Flat-headed cat

Pco: Puma concolor Puma

Pya: P. yagouaroundi Jaguarundi 6. Puma line
Aju: Acinonyx jubatus Cheetah

Lyp: Lynx pardinus Iberian lynx

Lly: L. lynx Eurasian lynx
5. Lynx line
Lca: L. canadensis Canadian lynx
Lru: L. rufus Bobcat

Lpa: Leopardus pardalis Ocelot

Lwi: L. wiedii Margay
Lja: L. Jacobita Andean mountain cat
Lco: L. colocolo Pampas cat 4. Ocelot line
Lge: L. geoffroyl Geoffroys’s
Lgu: L. guigna Kodkod
Lti: L. tigrinus Tigrina

Cca: Caracal caracal Caracal

Cau: C. aurata African golden cat 3. Caracal line
Cse: C. serval Serval

Pba: Pardofelis badia Bay cat

Pte: P. temminckii Asian golden cat 2. Bay cat line
Cat ancestor Pma: P. marmorata Marbled cat

Ple: Panthera leo Lion

Pon: P. onca Jaguar
Ppa: P. pardus Leopard
1. Panthera line
Pti: P. tigris Tiger
Pun: P. uncia Snow leopard
Nne: Neofelis nebulosa Clouded leopard

Figure 3a. Phylogentic tree and distribution (see figure 3b, opposite page) of the extant Felidae. The terminal nomenclature refers to
the eight major cat lineages (see text for further details). Distribution codes from coded map are given preceding scientific names. (After
Johnson et al., ref. 23.)

spot. Werdelin and Olsson18 replaced Weigel’s proposal can be traced back to subtle variations of the basic pattern.
with evidence indicating that the original cat family These can adequately explain the changes observed during
coat pattern was actually small spots or ‘flecks’. This is both the development of individuals and the radiation of
highlighted by changes observed in pelage patterns of cat lineages. In other words, mechanisms of change such
jaguars and leopards during their development. Juveniles as recombination and natural selection appear sufficient to
possess a simple spot pattern but adults display a range of explain the variation in melanistic spot patterns observed
complex rosettes. A recent theoretical study by Liu and within the cat family. Other patterns like the stripes of the
colleagues,19 using mathematical models, confirmed that in tiger were not investigated in the study, but the same kind of
both the jaguar and leopard a single mathematical function mathematical modeling has been used to simulate formation
(based on a Turing-Model) could simulate both the simple of stripes in other animals. Whole coat melanism in the
fleck-pattern of juveniles and the complex rosette-pattern domestic cat, jaguar and jaguarundi has been elucidated at
of the adults. It appears that differences in pelage patterns the molecular level.20 These pelage pattern studies seem to

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and the black-footed cat (F. nigripes)

within the Domestic Cat Lineage, and
the hierarchy of the Panthera Lineage
are all questioned. Presence of a large
non-functional mtDNA translocation in
the nuclear genome of all extant members
of Panthera supports a late Pliocene
radiation of the genus,21 and a recent
cladistic study has thrown much light
onto the detailed relationships between
extant and extinct Pantherinae.22 In the
cat phylogeny of Johnson et al.,23 21 of 36
speciation branch points took less than 1
million years each, and seven speciation
events that usher in the eight major cat
lineages took on average just 600,000
Figure 3b. Distribution of the extant Felidae. years to complete. Brief and spectacular
radiation events appear to be the norm
reinforce the general observation that cats form a unique in most vertebrate phylogenies. It is the reason why large
and well-defined group of animals. numbers of genes must be employed in phylogenetic studies.
During the radiation of the cat family, significant migrations
Radiation of modern cats in the Miocene took place across continents and distribution zones. These
The phylogenetic history of extant cats is complex and are displayed in figure 3b.23
has been controversial for a long time, primarily because of
The taming of the cat
the extremely rapid speciation events that took place during
the Tertiary period. Recently, based on DNA sequencing Felis catus, the domestic cat, is believed to have
studies, relations between the various lineages and species arisen from the African wildcat, F. lybica, and not from
could be clarified. DNA analyses of a spectrum of genes, the European wildcat, F. silvetris. Although taxonomists
along with modern statistical techniques, have revealed that sometimes lump these two wild species together, a number
the extant cat family can be subdivided into eight lineages of phenotypic differences are evident. F. lybica is easier to
(figure 3a): tame; it is far less shy of man. In appearance, F. lybica is
1. The Panthera Lineage (lion, jaguar, leopard, somewhat larger and stockier than F. silvestris. Felis lybica
tiger, snow leopard, and cloudy leopard) also has a black mark on the sole of its feet, continuing
2. The Bay Cat Lineage (bay cat, Asian golden between the toes. The domestic cat also displays a number
cat, and marbled cat) of differences from F. lybica, its ancestral species. Domestic
3. The Caracal line (caracal, African golden cat, cats tend to be smaller. They have a smaller brain and longer
serval) digestive tract. Domestic cats are also tamer than their
wildcat cousins. At least two features contribute to this.
4. The Ocelot Lineage (ocelot, margay, Andean
First, their appearance, including their behavior, is more
mountain cat, pampas cat, Geoffroy’s cat,
akin to the juvenile animal. Second, and perhaps related
kodkod, and tigrina)
to this, the relative size of their adrenal glands is smaller.
5. The Lynx Lineage (Iberian lynx, Eurasian lynx, The adrenal glands produce epinephrine (adrenaline), and
Canadian lynx, and bobcat) when animals are stressed this causes the fight-or-flight
6. The Puma line (puma, jaguarundi, and cheetah) behavior so familiar in wild animals. Because domestic
7. The Leopard Cat Lineage (Pallas’ cat, rusty cats have a smaller adrenal gland, less epinephrine is
spotted cat, Asian leopard cat, fishing cat, and produced and the animals are much quieter than wild
flat-headed cat) cats.24 These studies parallel observations by Trut et al.25 in
8. The Domestic Cat Lineage (domestic cat, studies of fox domestication. Together the studies suggest
European wildcat, African wildcat, Chinese that the domesticated phenotype lies dormant within the
desert cat, desert cat, black-footed cat, and genetic potential of many wild species. A recent extensive
jungle cat). study of mitochondrial DNA sequences from hundreds of
In spite of the excellent general consensus, a few domestic and wild cats across Europe, Asia, and Africa
phylogenetic relations are still uncertain. The placing of (including the domestic cat (F. catus), the European
the Andean mountain cat (Leopardus jacobita) within the wildcat (F. silvestris), the African wildcat (F. libyca) and
Ocelot Lineage, the placing of the jungle cat (Felis chaus) the Chinese desert cat (F. bieti), using the desert (sand) cat

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(F. margarita) as an outgroup, has clarified with great

certainty the various ancestral relationships. The domestic Nne Pun
cat is clearly descended from the African wildcat (F. libyca Ple Pti
Pba
or F. silvestris libyca), not the European wildcat (F. silvestris Ppa Pon
Pma Pte
or F. s. silvestris). In addition, the domestic cat, the European Fli
Fma Fsi
wildcat, the African wildcat, and also the Chinese desert
Fbi Cse
cat all appear to belong to a single species. Cementing this Fca Cau
idea is the fact that mitochondrial DNA from domestic cats Fni Cca
Fch
is routinely found in all three wild cat populations. Natural
hybridizations between them are frequent events.26
Lwi Lti
Pru
Hybridizations within the cat family Oma Pvi Lge Lco
Cats are beloved zoo and house animals, so there Ppl Pbe Lpa Lja
are many reports of hybridizations either occurring Lgu
spontaneously or deliberately undertaken (figure 4).4,27 Lru
Pco
Seven of the eight major cat lineages reported by Johnson Lca Lly
Aju Pya
et al.23 are linked by hybridizations. Only the Bay Cat Lyp
Lineage has not been linked by hybridization to another
lineage. Phenotypically, however, it is closely related to Figure 4. Hybridization network of the extant cat family. Cat
the Caracal lineage. Both the lineages contain golden lineages are depicted as circles, the great cat lineage (Panthera
cats. The Bay Cat Lineage includes the Asian golden cat, Lineage) is depicted in the uppermost circle. Each cat species is
indicated by a three-letter code obtained from the initial letter of
and the Caracal Lineage includes the African golden cat. the genus name and the first two letters of the species name, as
Moreover, in the Johnson et al. study they are placed as listed in figure 3a. Hybridization events are indicated by connecting
sister groups in the phylogenetic DNA-sequence tree. All lines. Hybrids with three pivotal species, often between lineages,
fourteen interlineage hybrids can be traced to one of three are indicated by single lines: F. catus (Fca; dotted lines), L. rufus
(Lru; dashed lines), P. concolor (Pco; continuous lines); all other
critical species: the domestic cat (F. catus), the bobcat (L. hybrids, within lineages, are indicated by double lines. All species
rufus), and the puma (P. concolor). Besides interlineage known to form interlineage hybrids are underlined. Additional
hybrids, numerous intralineage hybrids have also been information and many photographs of hybrids are available at
reported. Alderton4 describes a cross between a lion and ‘Hybrid and Mutant Big Cats, Mammals & Birds’, www.messybeast.
com/genetics/hybrid-cats.htm.
the hybrid of a jaguar and a leopard. The bobcat hybridizes
with all the lynxes (see figure 4). There is clear evidence × L. pardalis (11–16 kg) × L. wiedii (3–9 kg) × F. catus
that hybridizations occur between the major cat lineages (3–7 kg) × F. nigripes (1.5–2.5 kg). Intralineage hybrids
and that hybridizations occur within the lineages; therefore, of the lynx always involve the bobcat (L. rufus) and are
the hypothesis—all extant felids belong to a single basic referred to as Blynx or Lynx cats. Five cat hybrids are bred
type—has been reasonably proven. commercially as pets: F. chaus × F. silvestris (Euro-chaus),
Hybrids between cats are often given unique names. L. rufus × F. chaus (Jungle lynx), L. rufus × F. catus (desert
Many of these names are contracted fusions of their common lynx), F. catus × P. bengalensis (Bengal cat), and Bengal
names; the first part indicating the male parent and the cat × F. chaus (Jungle bob).
second part, the female parent. These include: P. leo × P.
tigris (Liger or Tigon), P. leo × P. pardus (Liard or Leopon), Feline-specific viruses
P. tigris × P. pardus (Tigard), P. onca × P. pardus (Jagulep or
Lepjag), P. onca × P. leo (Jaglion), P. concolor × P. pardus Viruses can only reproduce with the help of host cells.
(Pumapard), C. serval × C. caracal (servical or caraval), Viruses first attach to specific surface molecules on the
L. wiedii × L. pardalis (marlot). Female hybrids of lions host cell; then they infect the cell. Some viruses tend not
and tigers are fertile and reproduce. Their offspring also to be host-specific and are able to infect different spe-
enjoy a clear nomenclature. Li-ligers, li-tigons, ti-ligers, cies. A typical example of this is the influenza virus. It
and ti-tigons are all readily bred from female ligers and infects different families of birds, as well as pigs, other
tigons, and the female offspring of these crosses are also animals and, of course, man. The viruses attach to surface
fertile. P. concolor × L. pardalis (puma × ocelot) bridges the molecules common to warm-blooded animals in general.
gap between larger and smaller cats.28 Hybridization data Other viruses, however, are far more host-specific. Feline
connects the largest cats, P. tigris and the massive Liger (400+ Immunodeficiency Virus (FIV), the feline equivalent of
kg), to the smallest cat, F. nigripes, via seven documented HIV, has been shown to infect 30 of 37 felid species. At
hybrid steps: P. tigris (110–320 kg) × P. leo (120–250 least four FIV strains are species-specific. However, it also
kg) × P. pardus (30–85 kg) × P. concolor (35–100 kg) infects Crocuta crocuta (the spotted hyena) a member of

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the felids’ taxonomically closest family. Nevertheless, the P. leo (lion) was 36.4%, but the difference between P. onca
studies confirm that FIV transfer, even between cat species, (jaguar) and Cr. crocuta (spotted hyena) was only 35.9%.
is an infrequent event.29 The deadly viral disease, Feline Treatment of such values calls for best-judgment decisions.
Infectious Peritonitis (FIP), is specific to cats. Its causative Using a synopsis of all the values, the authors hypothesized
agent, Feline Corona Virus (FCoV), is very similar to the that the family of cats represent a single holobaramin.
human respiratory virus responsible for SARS and to coro- Over the years the two taxonomic concepts used to
naviruses from other animals. The mutation rate of the virus delineate a related group of species, the basic type and
is high (three mutations per virus genome per generation). the baramin, have been converging. Even in their 1998
Since the disease was first clinically described, in 1963, no article, Robinson & Cavanaugh33 used hybridization data
other natural host, besides members of the cat family, has as an indicator of inclusion in a holobaramin. They wrote,
been reported. There was a single report that FCoV could “The potential for interspecific hybridization provides an
induce infectious peritonitis in ferrets,30 and dogs can be important data set for elucidating monobaramins.”36 Further,
infected under laboratory conditions.31 Otherwise, FCoV Wood et al. wrote in their article about the HybriDatabase:
is an infallible indicator of the felid basic type, from house “Within-group reproductive viability and outgroup repro-
cats to lions, with cheetahs being especially susceptible.32 ductive isolation have been hypothesized to be important
Currently, host specificity of viruses is not considered di- characteristics of the holobaramin or basic type.”37 The more
agnostic of a basic type. Nevertheless, these are additional the hybridization criterion is given primary significance,
pieces of evidence supporting the unique character of the the closer the application of the holobaramin concept ap-
feline basic type because, essentially, cats alone are suscep- proaches the basic type definition. In turn, basic type re-
tible, with rare exceptions. search requires supplementary methods, such as statistical
tools, to place species that cannot be crossed, as is clearly
Basic-type or holobaramin? the case with fossils. In summary, it can be stated that the
family of cats represents both a basic type and a holobaramin
In 1998 an article about the family of cats was printed
according to external characteristics, hybridization data,
in the journal Creation Research Society Quarterly.33
and genetic evidence.
The authors placed the family of cats within a so-called
‘holobaramin’. A holobaramin, as originally defined, is ‘a Conclusion
complete set of organisms’ that are genetically related to
each other through common descent.34 Because common Ability to hybridize is the most important criteria
descent is harder to assess the further back in time one for including species within a common basic type.38 This
goes, eventually becoming empirically impossible to criterion cannot be used directly on fossil forms. However,
validate, Wood et al.35 proposed an alternative definition, because it indicates the extent of the morphogenetic
“a group of known organisms that share continuity (i.e. potential of a basic type, hybridization is an indirect
each member is continuous with at least one other member) indicator of fossil inclusion. Therefore, reports of extant cat
and are bounded by discontinuity”. To delimit baramin (a hybrids, fossil skeletal evidence, and various other features,
composite term derived from the Hebrew: bara = created, including molecular sequencing data, pelage patterns,
and min = kind), a modification of numerical taxonomy is and unique virus sensitivities, all seem to suggest that the
used. In their article, Robinson & Cavanaugh33 examined as family Felidae represents a single clearly delineated basic
many characteristics from cats and closely related animals as type. It is reasonable to assume that all felids arose from a
possible. Characteristics included information from ecology, single founder species and that they have passed through
morphology, and genetic evidence, such as size, weight, one or more adaptive radiations, exploiting their inherent
proportion of bone lengths to each other, food, chromosome morphogenetic potential to produce all of the known extant
numbers, etc., to name a few. In total the authors compared and extinct species of cat.1
287 criteria from cats, Crocuta crocuta (the spotted hyena),
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Barny Pendragon holds a B.Sc. (hons.) (1980) and an M.Sc.
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(1982) from the faculty of science at Manchester University,
21. Kim, J.-H., Antunes, A., Luo, S.-J., Menninger, J., Nash, W.G., O’Brien, England, a Ph.D. (1987) from the faculty of science at the
S.J., and Johnson, W.E., Evolutionary analysis of a large mtDNA
Justus Liebig University, Giessen, Germany, and a D.Sc.
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species, Gene 366:292–302, 2006.
(1998) from the faculty of medicine at Zurich University,
Switzerland. He has published almost 100 scientific papers
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and is a referee for ten international science journals. He has
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taught and held consultancy positions throughout Europe,
USA and Asia.
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24. Lumpkin, S., Small Cats Facts on File, New York, 1993.
Niko Winkler obtained a graduate degree in biology and
his Ph.D. in Microbiology. His dissertation research involved
25. Trut, L.N., Plyusnina, I.Z. and Oskina, I.N., An experiment on fox
studying microbes at the genetic and molecular levels. He
domestication and debatable issues of evolution of the dog, Russian J.
Genet. 40:644–655, 2004.
has been actively involved in the origins debate for many
years and has made various active contributions to basic-
26. Driscoll, C.A., Menotti-Raymond, M., Roca, A., Hupe, K., Johnson,
type research.
W.E., Geffen, E., Harley, E.H., Delibes, M., Pontier, D., Kitchener, A.C.,
Yamaguchi, N., O’Brien, S.J. and Macdonald, D., The Near Eastern origin
of cat domestication, Science 317:519–523, 2007.

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