Felidae () is the family of

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

s in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...

colloquially referred to as cats, and constitutes a

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

. A member of this family is also called a felid (). The term "cat" refers both to felids in general and specifically to the

domestic cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members o ...

('' Felis catus''). Felidae species exhibit the most diverse fur pattern of all terrestrial carnivores. Cats have retractile claws, slender muscular bodies and strong flexible forelimbs. Their teeth and facial muscles allow for a powerful bite. They are all

obligate carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...

, and most are solitary predators ambushing or stalking their prey. Wild cats occur in Africa, Europe, Asia and the

Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...

. Some wild cat species are adapted to forest habitats, some to arid environments, and a few also to wetlands and mountainous terrain. Their activity patterns range from

nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...

and

crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...

to diurnal, depending on their preferred prey species.

Reginald Innes Pocock Reginald Innes Pocock F.R.S. (4 March 1863 – 9 August 1947) was a British zoologist. Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edward ...

divided the

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

Felidae into three subfamilies: the Pantherinae, the Felinae and the Acinonychinae, differing from each other by the ossification of the hyoid apparatus and by the cutaneous sheaths which protect their claws. This concept has been revised following developments in molecular biology and techniques for analysis of morphological data. Today, the living Felidae are divided in two subfamilies: the Pantherinae and Felinae, with the Acinonychinae subsumed into the latter. Pantherinae includes five ''

Panthera ''Panthera'' is a genus within the family (biology), family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as co ...

'' and two ''

Neofelis ''Neofelis'' is a genus comprising two extant cat species in Southeast Asia: the clouded leopard (''Neofelis nebulosa'') of mainland Asia, and the Sunda clouded leopard (''Neofelis diardi'') of Sumatra and Borneo The scientific name ''Neofelis' ...

'' species, while Felinae includes the other 34 species in ten

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

. The first cats emerged during the

Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...

about , with the appearance of '' Proailurus'' and '' Pseudaelurus''. The latter species complex was ancestral to two main lines of felids: the cats in the extant subfamilies and a group of

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

cats of the subfamily

Machairodontinae Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...

, which include the saber-toothed cats such as the '' Smilodon''. The "false sabre-toothed cats", the

Barbourofelidae Barbourofelidae is an extinct family of carnivorans of the suborder Feliformia, sometimes known as false saber-toothed cats, that lived in North America, Eurasia and Africa during the Miocene epoch (16.9—9.0 million years ago) and existed for a ...

and Nimravidae, are not true cats, but are closely related. Together with the Felidae, Viverridae, hyaenas and mongooses, they constitute the Feliformia.

Characteristics

All members of the cat family have the following characteristics in common: * They are

digitigrade In terrestrial vertebrates, digitigrade () locomotion is walking or running on the toes (from the Latin ''digitus'', 'finger', and ''gradior'', 'walk'). A digitigrade animal is one that stands or walks with its toes (metatarsals) touching the groun ...

, have five toes on their forefeet and four on their hind feet. Their curved claws are protractile and attached to the terminal bones of the toe with

ligament A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...

s and tendons. The claws are guarded by cutaneous

sheath Sheath pronounced as , may refer to: * Scabbard, a sheath for holding a sword, knife, or other large blade, as well as guns, such as rifles. * The outer covering of a cable * Condom, a kind of contraception * Debye sheath, a layer of a plasma in ...

s, except in the '' Acinonyx''. * The plantar pads of both fore and hind feet form compact three-lobed cushions. * They actively protract the claws by contracting muscles in the toe, and they passively retract them. The dewclaws are expanded but do not protract. * They have lithe and flexible bodies with muscular limbs. * Their skull is foreshortened with a rounded profile and large orbits. * They have 30 teeth with a dental formula of . The upper third premolar and lower molar are adapted as carnassial teeth, suited to tearing and cutting flesh. The canine teeth are large, reaching exceptional size in the extinct saber-toothed species. The lower carnassial is smaller than the upper carnassial and has a crown with two compressed blade-like pointed cusps. * Their tongue is covered with horny

papillae Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Basilar papilla, a sensory organ of lizards, amphibians and fish * Dental papilla, in a developing tooth * Dermal papillae, part of ...

, which rasp meat from prey and aid in grooming. * Their nose projects slightly beyond the lower jaw. * Their eyes are relatively large, situated to provide binocular vision. Their night vision is especially good due to the presence of a '' tapetum lucidum'', which reflects light back inside the eyeball, and gives felid eyes their distinctive shine. As a result, the eyes of felids are about six times more light-sensitive than those of humans, and many species are at least partially

. The retina of felids also contains a relatively high proportion of

rod cell Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in per ...

s, adapted for distinguishing moving objects in conditions of dim light, which are complemented by the presence of

cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...

s for sensing colour during the day. * They have well-developed and highly sensitive

whiskers Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to touch, sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are c ...

above the eyes, on the cheeks, on the muzzle, but not below the chin. Whiskers help to navigate in the dark and to capture and hold prey. * Their external ears are large and especially sensitive to high-frequency sounds in the smaller cat species. This sensitivity allows them to locate small rodent prey. * The penis is subconical, facing backward when not erect. The

baculum The baculum (also penis bone, penile bone, or ''os penis'', ''os genitale'' or ''os priapi'') is a bone found in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the ...

is small or vestigial, and shorter than in the

Canidae Canidae (; from Latin, ''canis'', "dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within th ...

. * Felids have a vomeronasal organ in the roof of the mouth, allowing them to "taste" the air. The use of this organ is associated with the Flehmen response. * They cannot detect the

sweetness Sweetness is a Taste#Basic tastes, basic taste most commonly Perception, perceived when eating foods rich in sugars. Sweet tastes are generally regarded as pleasure, pleasurable. In addition to sugars like sucrose, many other chemical compounds ...

of sugar, as they lack the sweet- taste receptor. * They share a broadly similar set of vocalisations, but with some variation between species. In particular, the pitch of calls varies, with larger species producing deeper sounds; overall, the frequency of felid calls ranges between 50 and 10,000 hertz. The standard sounds made by all felids include meowing, spitting, hissing, snarling and

growling Growling is a low, guttural vocalization produced by animals as an aggressive warning but can also be found in other contexts such as playful behaviors or mating. Different animals will use growling in specific contexts as a form of communicati ...

. Meowing is the main contact sound, whereas the others signify an aggressive motivation. * They can purr during both phases of respiration, though pantherine cats seem to purr only during oestrus and

copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetrat ...

, and as cubs when suckling. Purring is generally a low pitch sound of less than 2

kHz The hertz (symbol: Hz) is the unit of frequency in the International System of Units (SI), equivalent to one event (or cycle) per second. The hertz is an SI derived unit whose expression in terms of SI base units is s−1, meaning that on ...

and mixed with other vocalization types during the expiratory phase. The ability to roar comes from an elongated and specially adapted larynx and hyoid apparatus. When air passes through the larynx on the way from the lungs, the cartilage walls of the larynx vibrate, producing sound. Only lions, leopards, tigers, and jaguars are truly able to roar, although the loudest mews of snow leopards have a similar, if less structured, sound. The colour, length and density of their fur is very diverse. Fur colour covers the gamut from white to black, and fur pattern from distinctive small spots, stripes to small blotches and rosettes. Most cat species are born with a spotted fur, except the

jaguarundi The jaguarundi (''Herpailurus yagouaroundi'') is a wild cat native to the Americas. Its range extends from central Argentina in the south to northern Mexico, through Central and South America east of the Andes. The jaguarundi is a medium-sized ...

(''Herpailurus yagouaroundi''),

Asian golden cat The Asian golden cat (''Catopuma temminckii'') is a medium-sized wild cat native to the northeastern Indian subcontinent, Southeast Asia and China. It has been listed as Near Threatened on the IUCN Red List since 2008, and is threatened by poach ...

(''Catopuma temminckii'') and caracal (''Caracal caracal''). The spotted fur of lion (''Panthera leo'') and

cougar The cougar (''Puma concolor'') is a large Felidae, cat native to the Americas. Its Species distribution, range spans from the Canadian Yukon to the southern Andes in South America and is the most widespread of any large wild terrestrial mamm ...

(''Puma concolor'') cubs change to a uniform fur during their ontogeny. Those living in cold environments have thick fur with long hair, like the snow leopard (''Panthera uncia'') and the Pallas's cat (''Otocolobus manul''). Those living in tropical and hot climate zones have short fur. Several species exhibit melanism with all-black individuals. In the great majority of cat species, the tail is between a third and a half of the body length, although with some exceptions, like the ''

Lynx A lynx is a type of wild cat. Lynx may also refer to: Astronomy * Lynx (constellation) * Lynx (Chinese astronomy) * Lynx X-ray Observatory, a NASA-funded mission concept for a next-generation X-ray space observatory Places Canada * Lynx, Ontar ...

'' species and margay (''Leopardus wiedii''). Cat species vary greatly in body and skull sizes, and weights: * The largest cat species is the tiger (''Panthera tigris''), with a head-to-body length of up to , a weight range of at least , and a skull length ranging from . Although the maximum skull length of a lion is slightly greater at , it is generally smaller in head-to-body length than the former. * The smallest cat species are the

rusty-spotted cat The rusty-spotted cat (''Prionailurus rubiginosus'') is one of the cat family's smallest members, of which historical records are known only from India and Sri Lanka. In 2012, it was also recorded in the western Terai of Nepal. Since 2016, the gl ...

(''Prionailurus rubiginosus'') and the black-footed cat (''Felis nigripes''). The former is in length and weighs . The latter has a head-to-body length of and a maximum recorded weight of . Most cat species have a

haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...

number of 18 or 19. Central and South American cats have a haploid number of 18, possibly due to the combination of two smaller chromosomes into a larger one. Most cat species are also induced ovulators, although the margay appears to be a spontaneous ovulator.

Evolution

The family Felidae is part of the Feliformia, a suborder that diverged probably about into several families. The Felidae and the Asiatic linsangs are considered a

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

, which split about . The earliest cats probably appeared about . '' Proailurus'' is the oldest known cat that occurred after the Eocene–Oligocene extinction event about ; fossil remains were excavated in France and Mongolia's

Hsanda Gol Formation The Hsanda Gol Formation is a geological formation located in the Tsagan Nor Basin of central Mongolia that dates to the Rupelian age of the Oligocene epoch. It formed in continental desert-like conditions and is notable for its fossil mammals, ...

. Fossil occurrences indicate that the Felidae arrived in North America around . This is about 20million years later than the Ursidae and the Nimravidae, and about 10 million years later than the

. In the Early Miocene about , '' Pseudaelurus'' lived in Africa. Its fossil jaws were also excavated in geological formations of Europe's Vallesian, Asia's

Middle Miocene The Middle Miocene is a sub-epoch of the Miocene Epoch made up of two stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0.05 Ma to 11.608 ± 0.005 Ma (million y ...

and North America's late Hemingfordian to late

Barstovian The Barstovian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 16,300,000 to 13,600,000 years BP, a period of . It is usua ...

epochs. In the Early or Middle Miocene, the sabre-toothed

evolved in Africa and migrated northwards in the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

. With their large upper canines, they were adapted to prey on large-bodied megaherbivores. ''

Miomachairodus ''Miomachairodus'' is an extinct genus of large saber-toothed cats of the subfamily Machairodontinae. It is known from Miocene-age fossils in China and Turkey and persisted until the Late Miocene (early Vallesian).Metailurus'' lived in Africa and Eurasia about . Several ''

Paramachaerodus ''Paramachaerodus'' is an extinct genus of saber-tooth cat of the subfamily Machairodontinae, which was endemic to Europe and Asia during the late Miocene from 15 to 9 Ma. ''Paramacheirodus'' is one of the oldest known true saber-toothed cat ...

'' skeletons were found in Spain. '' Homotherium'' appeared in Africa, Eurasia and North America around , and '' Megantereon'' about . '' Smilodon'' lived in North and South America from about . This subfamily became extinct in the

Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...

. Results of

mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...

analysis indicate that the living Felidae species descended from a common ancestor, which originated in Asia in the Late Miocene epoch. They migrated to Africa, Europe and the Americas in the course of at least 10 migration waves during the past ~11 million years. Low sea levels,

interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...

and glacial periods facilitated these migrations. ''

Panthera blytheae ''Panthera blytheae'' is an extinct species of the genus '' Panthera'' that is thought to have existed during the late Messinian to early Zanclean ages approximately 5.95–4.1 million years ago. It is currently the oldest known ''Panthera'' sp ...

'' is the oldest known pantherine cat dated to the late

Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the first ...

to early Zanclean ages about . A fossil skull was excavated in 2010 in Zanda County on the Tibetan Plateau. ''

Panthera palaeosinensis ''Panthera palaeosinensis'' was an early Pleistocene species from northern China. It is often incorrectly referenced as the ancestor of the tiger, ''Panthera tigris'', although it shares features with all living large cats. Recent studies place i ...

'' from North China probably dates to the Late Miocene or Early Pliocene. The skull of the holotype is similar to that of a lion or leopard. '' Panthera zdanskyi'' dates to the Gelasian about . Several fossil skulls and jawbones were excavated in northwestern China. '' Panthera gombaszoegensis'' is the earliest known pantherine cat that lived in Europe about . Living felids fall into eight

evolutionary lineages An evolutionary lineage is a temporal series of populations, organisms, cells, or genes connected by a continuous line of descent from ancestor to descendant.The Oxford English Dictionary defines biological lineage as "a sequence of species each of ...

or species

s. Genotyping of nuclear DNA of all 41 felid species revealed that

hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...

between species occurred in the course of evolution within the majority of the eight lineages. Modelling of felid coat pattern transformations revealed that nearly all patterns evolved from small spots.

Classification

Traditionally, five subfamilies have been distinguished within the Felidae based on phenotypical features: the Pantherinae, the Felinae, the Acinonychinae, and the extinct

and

Proailurinae ''Proailurus'' is an extinct felid genus that lived in Europe and Asia approximately 25 million years ago in the Late Oligocene and Miocene. Fossils have been found in Mongolia, Germany, and Spain. Etymology The generic name ''Proailurus'' come ...

Phylogeny

The following cladogram based on Piras et al. (2013) depicts the phylogeny of basal living and extinct groups. The phylogenetic relationships of living felids are shown in the following cladogram:

References

External links