Feliformia

Feliformia (also Feloidea) is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, civets, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Caniformia ("dog-like" carnivorans).

The separation of the Carnivora into the broad groups of feliforms and caniforms is widely accepted, as is the definition of Feliformia and Caniformia as suborders (sometimes superfamilies). The classification of feliforms as part of the Feliformia suborder or under separate groupings continues to evolve.

Systematic classifications dealing with only extant taxa include all feliforms into the Feliformia suborder, though variations exist in the definition and grouping of families and genera.[1][2] Indeed, molecular phylogenies suggest that all extant Feliformia are monophyletic.[3]

The extant families as reflected in the taxa chart at right and the discussions in this article reflect the most contemporary and well-supported views (as at the time of writing this article).

Systematic classifications dealing with both extant and extinct taxa vary more widely.[4][5] Some separate the feliforms (extant and extinct) as: Aeluroidea (superfamily) and Feliformia (suborder).[5] Others include all feliforms (extant, extinct and "possible ancestors") into the Feliformia suborder.[4] Some studies suggest this inclusion of "possible ancestors" into Feliformia (or even Carnivora) may be spurious.[6] The extinct (†) families as reflected in the taxa chart are the least problematic in terms of their relationship with extant feliforms (with the most problematic being Nimravidae).

Feliforms
Temporal range: Eocene-Holocene
Feliform portraits
Several extant feliform families: Eupleridae, Felidae, Hyaenidae, Herpestidae and Viverridae.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Kretzoi, 1945
Families

Characteristics

Feliformia Skull v01-2
Feliformia skull showing double-chambered bullae

All extant feliforms share a common attribute: their auditory bullae (bony capsules enclosing the middle and inner ear).[7] This is a key diagnostic in classifying species as feliform versus caniform. In feliforms, the auditory bullae are double-chambered, composed of two bones joined by a septum. Caniforms have single-chambered or partially divided auditory bullae, composed of a single bone. This feature, however, is problematic for the classification of the extinct Nimravidae as feliforms. Nimravid fossils show ossified bullae with no septum, or no trace at all of the entire bulla. It is assumed that they had a cartilaginous housing of the ear mechanism.[8]

The specific characteristics of extant feliform bullae suggest a common ancestor, though one has not been identified in the fossil records. There are other characteristics that differentiate feliforms from caniforms and probably existed in their stem taxa. But, due to speciation, these do not apply unambiguously to all extant species.

Feliforms tend to have shorter rostrums than caniforms, fewer teeth, and more specialized carnassials. Feliforms tend to be more carnivorous and are generally ambush hunters. Caniforms tend more toward omnivorous and opportunity-based feeders.

Many feliforms have retractile or semi-retractile claws and many are arboreal or semi-arboreal. Feliforms also tend to be more digitigrade (walking on toes). In contrast, most caniforms are terrestrial, have non-retractile claws and tend to be plantigrade.

Extant families

Bodleian Libraries, Handbill of Merchant's Hall, 1739, announcing A lion, lionesses, tigers, etc.
A 1739 advert by Charles Benjamin Incledon featuring feliforms: the Mesopotamian lion from the vicinity of Bassorah, Cape lion, tiger from the East Indies, panther from Buenos Aires, Hyaena hyaena from West Africa, and leopard from Turkey, besides a "Man tyger" from Africa.

There are seven extant families, twelve subfamilies, 56 genera and 114 species in the Feliformia suborder. They range natively across all continents except Australia and Antarctica. Most species are arboreal or semi-arboreal ambush hunters. Target prey varies based on the species size and available food sources (with the larger species feeding mainly on small mammals and the smallest species feeding on insects or invertebrates).

An overview of each family is provided here. For detailed taxa and descriptions of the species in each family, follow the links to other articles and external references.

Fossa
Fossa (Cryptoprocta ferox)

Family Eupleridae (the "Malagasy carnivorans") includes fossa, falanouc, Malagasy civet and Malagasy mongooses, all of which are restricted to the island of Madagascar. The eight species in the family exhibit significant variations in form. These differences initially led to the species in this family sharing common names with, and being placed in the different families of, apparently more similar species on the mainland (e.g. civets and mongoose). However, phylogenetic analysis of DNA provides strong evidence that all Malagasy carnivorans evolved from a single common ancestor that was a herpestid (Yoder et al. 2003).[9][10] Phylogenetic analysis supports this view and places all of the Malagasy carnivorans in the family Eupleridae.[11]

The differences in form make it difficult to concisely summarise the species in this family. The range in size is as diverse as the range in form, with smaller species at less than 500 g (1 lb) and the largest species at up to 12 kg (26 lb). Some have retractile or semi-retractile claws (the fossa and the Malagasy civet) and others do not (the falanouc and Malagasy mongooses). They all tend to have slender bodies and pointed rostra (except the fossa, which has a blunt snout). Diet varies with size and form of the species and, like their mainland counterparts, ranges from small mammals, insects and invertebrates through to crustaceans and molluscs.

Cheetah4
Cheetah (Acinonyx jubatus)

Family Felidae (cats) are the most widespread of the "cat-like" carnivorans. There are 41 extant species, and all but a few have retractile claws. This family is represented on all continents except Australia (where domestic cats have been introduced) and the Antarctic. The species vary in size from the tiny black-footed cat (Felis nigripes) at only 2 kg (4.5 lb) to the tiger (Panthera tigris) at 300 kg (660 lb). Diet ranges from large to small mammals, birds and insects (depending on species size.)

Crocuta crocuta sideview
Spotted hyena (Crocuta crocuta)

Family Hyaenidae (hyenas and aardwolf) has four extant species and two subspecies. All show features of convergent evolution with canids, including non-retractile claws, long muzzles, and adaptations to running for long distances. They are extant in the Middle East, India and Africa. Hyenas are large, powerful animals, up to 80 kg (176 lb) and represent one of the most prolific large carnivorans on the planet. The aardwolf is much smaller at 27 kg (60 lb) and is a specialised feeder, eating mainly harvester termites.

Suricata
Meerkat (Suricata suricatta)

Family Herpestidae (the mongooses, kusimanses, meerkat, etc.) has 32 species. Previously, these were placed in the family Viverridae. However, Wilson and Reeder (1993) established the herpestids as morphologically and genetically distinct from viverrids. They are extant in Africa, Middle East and Asia. All have non-retractile claws. They are smaller as a family, ranging from 1 kg (2.2 lb) to 5 kg (11 lb), and typically have long, slender bodies and short legs. Diet varies based on species size and available food sources, ranging from small mammals, birds to reptiles, insects and crabs. Some species are omnivorous, including fruits and tubers in their diet.

Binturong in Overloon
Binturong (Arctictis binturong)

Family Nandiniidae (African palm civet) has only one species (Nandinia binotata), extant across sub-Saharan Africa. They have retractile claws and are slender-bodied, arboreal omnivores (with fruit making up much of their diet). They are relatively small with the larger males weighing up to 5 kg (11 lb).

Family Prionodontidae (the Asiatic linsangs) has two extant species in one genus. They live in Southern-East Asia. All are arboreal hypercarnivorans. They are the closest living relatives of the family Felidae.[12]

Family Viverridae has 30 living species including binturong, large Indian civet, genets and oyans. They all have long bodies, short legs with retractile claws, and usually long tails. In weight, the species range from 0.5–14 kg (1.1–30.9 lb). Some occur in Southern Europe, but most in Africa and Asia. Their diet ranges from fruit and plants to insects, crustaceans and molluscs, and small mammals.

Evolution

In the Middle Palaeocene (60 million years ago), Miacoidea appears. Miacoids were a group of paraphyletic taxa believed to be basal to Carnivora. They had Carnivora-like carnassials but lacked fully ossified auditory bullae. Miacids were small arboreal carnivorans and, based on their size (roughly that of mongooses), they probably fed on insects, small mammals and birds.

The miacoids are divided into two groups: the miacids, with a full complement of molars, and the viverravines with a reduced number of molars and more specialized carnassials. These dental differences resemble the difference between Caniforms (with more teeth) and Feliforms (with fewer teeth) but this may not mean evolutionary lineages. It was thought that Viverravidae was basal to the Feliforms. However, some studies suggest this is not the case.[6]

In the Middle Eocene (about 42 mya), the miacids started to branch into two distinct groups of the order Carnivora: the Feliforms and Caniforms. The miacid precursors to the extant Feliforms remained forest-dwelling, arboreal or semi-arboreal ambush hunters, while the Caniform precursors were more mobile, opportunistic hunters. While it is clear that the first Feliforms appeared at this time, there is no clear common ancestor of the Feliform families in the fossil records. As forest dwellers, the early Feliforms were subject to more rapid decomposition in the absence of sedimentary materials, resulting in large gaps in the fossil records.

For more discussion on feliform evolution and the divergence from the caniforms, together with additional external references on this subject, see the articles on Carnivora, Miacoidea and Carnivoramorpha.

Phylogenetic tree

   Feliformia   

NimravidaeHoplophoneus.jpg

Stenoplesictidae

PercrocutidaeDinocrocuta gigantea.jpg

Nandiniidae Nandinia binotata, Manchester Museum.jpg

Prionodontidae

BarbourofelidaeEusmilus285wide.jpg

Felidae Amur or Siberian tiger (Panthera tigris altaica), Tierpark Hagenbeck, Hamburg, Germany - 20070514.jpg

Viverridae AfricanCivet.jpg

Hyaenidae Crocuta crocuta.jpg

Herpestidae Mongoose.jpg

Eupleridae Fossa.jpg

References

  1. ^ Taxonomic references - extant species (1): Supporting descriptive information and pictures: Diversity Web (online) – Feliformia
  2. ^ Taxonomic references - extant species (2): Integrated Taxonomic Information System (ITIS)
  3. ^ Eizirik, E., W.J. Murphy, K.P. Koepfli, W.E. Johnson, J.W. Dragoo, R.K.Wayne, en S.J. O’Brien, 2010. Pattern and timing of the diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences. Molecular Phylogenetics and Evolution 56: 49-63. doi:10.1016/j.ympev.2010.01.033
  4. ^ a b Fossil record data (with taxonomic references) extant and extinct species: The Paleaobiology Database
  5. ^ a b Supporting taxonomic references extant and extinct species: Systema Naturae 2000 / Classification - Suborder Feliformia
  6. ^ a b Wesley‐Hunt, Gina D.; Flynn, John J. (2005). "Phylogeny of the carnivora: Basal relationships among the carnivoramorphans, and assessment of the position of 'miacoidea' relative to carnivora". Journal of Systematic Palaeontology. 3 (1): 1–28. doi:10.1017/S1477201904001518. ISSN 1477-2019.
  7. ^ R. F. Ewer (1973). The Carnivores. Cornell University Press. ISBN 0-8014-8493-6.
  8. ^ Turner, Alan (1997). The Big Cats and their Fossil Relatives: an illustrated guide. New York: Columbia University Press. p. 234. ISBN 0-231-10228-3.
  9. ^ Anne D. Yoder and John J. Flynn 2003: Origin of Malagasy Carnivora
  10. ^ Yoder, A., M. Burns, S. Zehr, T. Delefosse, G. Veron, S. Goodman, J. Flynn. 2003: Single origin of Malagasy Carnivora from an African ancestor – Letters to Nature
  11. ^ Philippe Gaubert, W. Chris Wozencraft, Pedro Cordeiro-Estrela and Géraldine Veron. 2005 - Mosaics of Convergences and Noise in Morphological Phylogenies: What's in a Viverrid-Like Carnivoran?
  12. ^ Gaubert, P., & Veron, G. (2003). "Exhaustive sample set among Viverridae reveals the sister-group of felids: the linsangs as a case of extreme morphological convergence within Feliformia". Proceedings of the Royal Society, Series B, 270 270 (1532): 2523–30. doi:10.1098/rspb.2003.2521
Barbourofelidae

Barbourofelidae is an extinct family of carnivorans of the suborder Feliformia that lived in North America, Eurasia and Africa during the Miocene epoch (16.9—9.0 Ma) and existed for about 7.9 million years.

Caniformia

Caniformia, or Canoidea (literally "dog-like"), is a suborder within the order Carnivora. They typically possess a long snout and nonretractile claws (in contrast to the cat-like carnivorans, the Feliformia). The Pinnipedia (seals, walruses and sea lions) are also assigned to this group. The center of diversification for Caniformia is North America and northern Eurasia. This contrasts with the feliforms, the center of diversification of which was in Africa and southern Asia.

Caraval

The caraval (also called a cara-serval) is the cross between a male caracal and a female serval. They have a spotted pattern similar to the Serval, but on a darker background. These are bred for the pet market.

A servical is the cross between a male serval and a female caracal. A litter of servicals occurred by accident when the two animals were kept in the same enclosure at a Los Angeles zoo. The hybrids were given to an animal shelter. The only photos show them as tawny kittens.

These hybrids can theoretically be backcrossed to their parent species in various ways:

Ser-caraval (¾ serval, ¼ caracal) - a cross between a male serval and a female caraval.

Car-servical (¾ caracal, ¼ serval) - a cross between a male caracal and a servical.

Ser-servicals (¾ serval, ¼ caracal) - a male serval and a female servical.

Car-caravals (¾ caracal, ¼ serval) - a male caracal and female caraval.Currently, only ser-caravals have been documented. All of these hybrids will more closely resemble the species that contributed the greater fraction of their genes.

Carnivora

Carnivora (; from Latin carō (stem carn-) "flesh" and vorāre "to devour") is a diverse scrotiferan order that includes over 280 species of placental mammals. Its members are formally referred to as carnivorans, whereas the word "carnivore" (often popularly applied to members of this group) can refer to any meat-eating organism. Carnivorans are the most diverse in size of any mammalian order, ranging from the least weasel (Mustela nivalis), at as little as 25 g (0.88 oz) and 11 cm (4.3 in), to the polar bear (Ursus maritimus), which can weigh up to 1,000 kg (2,200 lb), to the southern elephant seal (Mirounga leonina), whose adult males weigh up to 5,000 kg (11,000 lb) and measure up to 6.7 m (22 ft) in length.

Carnivorans have teeth and claws adapted for catching and eating other animals. Many hunt in packs and are social animals, giving them an advantage over larger prey. Some carnivorans, such as cats and pinnipeds, depend entirely on meat for their nutrition. Others, such as raccoons and bears, are more omnivorous, depending on the habitat. The giant panda is largely a herbivore, but also feeds on fish, eggs and insects. The polar bear subsists mainly on seals.

Carnivorans are split into two suborders: Feliformia ("catlike") and Caniformia ("doglike").

Carnivoramorpha

Carnivoramorpha are a clade of mammals that includes the modern order Carnivora.

Eupleridae

Eupleridae is a family of carnivorans endemic to Madagascar and comprising 10 known living species in seven genera, commonly known as euplerids, or Malagasy mongooses. The best known species is the fossa (Cryptoprocta ferox), in the subfamily Euplerinae. All species of Euplerinae were formerly classified as viverrids, while all species in the subfamily Galidiinae were classified as herpestids.

Recent molecular studies indicate that the 10 living species of Madagascar carnivorans evolved from one ancestor that is thought to have rafted over from mainland Africa 18-24 million years ago. This makes Malagasy carnivorans a clade. They are closely allied with the true herpestid mongooses, their closest living relatives. The fossa and the Malagasy civet (Fossa fossana) are each evolutionarily quite distinct from each other and from the rest of the clade.

All Eupleridae are considered threatened species due to habitat destruction, as well as predation and competition from non-native species.

Galidiinae

Galidiinae is a subfamily of carnivorans that is restricted to Madagascar and includes six species classified into four genera. Together with the three other species of indigenous Malagasy carnivorans, including the fossa, they are currently classified in the family Eupleridae within the suborder Feliformia. Galidiinae are the smallest of the Malagasy carnivorans, generally weighing about 600 to 900 g. They are agile, short-legged animals with long, bushy tails.In some of these characters, they resemble the mongooses (family Herpestidae) of continental Africa and southern Eurasia, with which they were classified until 2006, and accordingly they are said to be "mongoose-like" or even described as "Malagasy mongooses".

Haile Quarry site

The Haile Quarry or Haile sites are an Early Miocene and Pleistocene assemblage of vertebrate fossils located in the Haile quarries, Alachua County, northern Florida. The assemblage was discovered during phosphate mining, which began in the late 1940s. Haile sites are found in the Alachua Formation. Two sites within the Ocala Limestone yielded Upper Eocene Valvatida (sea stars) and mollusks.

University of Florida and Florida Museum of Natural History paleontologists numbered the Haile fossil sites with Arabic and Roman numbers and letters in order to define locations more distinctly. UF scientists used Roman numbering and the FLMNH scientists used Arabic.

Inglis quarry

The Inglis quarry or Inglis quarry sites 1A and 1C are assemblages of vertebrate fossils dating from the Pleistocene ~1.8 Mya—300,000 years ago, located in the phosphate quarries near the town of Inglis, Citrus County, northern Florida.

Inglis sites FCi-1, FCi-2, Inglis Formation, Florida Geological Survey C-11, Inglis Member, Moodys Branch Formation, and Dunellon Phosphate Company pit no. 5 are composed of a variety of bivalves, echinoderms, gastropods, crustaceans (mud shrimp), crinoids dating from the Eocene to Early Oligocene of ~48—33.9 Mya.

Linsang

Linsangs is an originally Javanese name applied to four species of tree-dwelling carnivorous mammals. The two African species belong to the family Viverridae, the two Asiatic species belong to the family Prionodontidae.

Both linsang genera (the African Poiana and the Asian Prionodon) were formerly placed in the subfamily Viverrinae (of Viverridae), along with several other genera, but recent research suggests that their actual relationships may be somewhat different. The linsangs are remarkable for their morphological resemblance to cats, family Felidae, which is greater than in the other viverrids. As the relationship between linsangs and cats was thought to be rather distant (the two groups belonging to different families within the superfamily Feliformia), this was considered an example of convergent evolution. However, DNA analysis indicates that while the African linsangs (Poiana) are true viverrids closely related to the genets, the Asiatic linsangs (Prionodon) are not and may instead be the closest living relatives of the family Felidae. The similarities between Asiatic linsangs and cats are thus more likely to be due to common ancestry, while the similarities between the two genera of linsangs must be convergent.

The name linsang is from Javanese linsang or wlinsang, which used to be wrongly translated as "otter" in English dictionaries. Linsangs are nocturnal, generally solitary tree dwellers. They are carnivorous, eating squirrels and other rodents, small birds, lizards and insects. Typical size is a little over 30 cm (1 foot), with a tail that more than doubles that length. Bodies are long, with short legs, giving a low appearance. Both species have yellowish bodies with black markings (stripes, blotches and spots), though the distribution and nature of the markings varies between the two species.

The species of African linsangs are:

Poiana leightoni - Leighton's linsang

Poiana richardsonii - African linsangThe species of Asiatic linsangs are:

Prionodon linsang - banded linsang

Prionodon pardicolor - spotted linsang

Mongoose

Mongoose is the popular English name for 29 of the 34 species in the 14 genera of the family Herpestidae, which are small feliform carnivorans native to southern Eurasia and mainland Africa. The other five species (all African) in the family are the four kusimanses in the genus Crossarchus, and the species Suricata suricatta, commonly called meerkat in English.

Six species in the family Eupleridae are endemic to the island of Madagascar. These are called "mongoose" and were originally classified as a genus within the family Herpestidae, but genetic evidence has since shown that they are more closely related to other Madagascar carnivorans in the family Eupleridae; they have been classified in the subfamily Galidiinae within Eupleridae since 2006.

Herpestidae is placed within the suborder Feliformia, together with the cat, hyena, and Viverridae families.

Nimravidae

Nimravidae is an extinct family of mammalian carnivores, sometimes known as false saber-toothed cats, whose fossils are found in North America, and Eurasia. Not considered to belong to the true cats (family Felidae), the nimravids are generally considered closely related and classified as a distinct family in the suborder Feliformia. Fossils have been dated from the Middle Eocene through the Late Miocene epochs (Bartonian through Tortonian stages, 40.4—7.2 mya), spanning about 33.2 million years.The barbourofelids were once previously classified as a subfamily of Nimravidae, in 2004, but were reassigned to their own distinct family Barbourofelidae.

Paradoxurus

Paradoxurus is a genus within the viverrid family that was denominated and first described by Frédéric Cuvier in 1822. As of 2005, this genus was defined as comprising three species native to Southeast Asia:

the Asian palm civet (P. hermaphroditus)

the golden palm civet (P. zeylonensis)

the brown palm civet (P. jerdoni)In 2009, it was proposed to also include the golden wet-zone palm civet (P. aureus), the Sri Lankan brown palm civet (P. montanus) and the golden dry-zone palm civet (P. stenocephalus), which are endemic to Sri Lanka.

Poiana (genus)

The African linsangs also known as oyans are two species classified in the mammalian subfamily Viverrinae, in the family Viverridae. There is one genus, Poiana.

Both linsang genera (Poiana and the Asian Prionodon) were formerly placed in the subfamily Viverrinae (of Viverridae), along with several other genera, but recent research suggests that their actual relationships may be somewhat different. The linsangs are remarkable for their morphological resemblance to cats, family Felidae, which is greater than in the other viverrids. As the relationship between linsangs and cats was thought to be rather distant (the two groups belonging to different families within the superfamily Feliformia), this was considered an example of convergent evolution. However, DNA analysis indicates that while the African linsangs (Poiana) are true viverrids closely related to the genets, the Asiatic linsangs (Prionodon) are not and may instead be the closest living relatives of the family Felidae. The similarities between Asiatic linsangs and cats are thus more likely to be due to common ancestry, while the similarities between the two genera of linsangs must be convergent.

The name linsang is from Javanese linsang or wlinsang, which used to be wrongly translated as "otter" in English dictionaries. Linsangs are nocturnal, generally solitary tree dwellers. They are carnivorous, eating squirrels and other rodents, small birds, lizards and insects. Typical size is a little over 30 cm (1 foot), with a tail that more than doubles that length. Bodies are long, with short legs, giving a low appearance. Both species have yellowish bodies with black markings (stripes, blotches and spots), though the distribution and nature of the markings varies between the two species.

The species of African linsangs are:

Poiana leightoni - West African oyan

Poiana richardsonii - Central African oyan

Prosansanosmilus

Prosansanosmilus is an extinct genus of mammalian carnivores of the suborder Feliformia, family Barbourofelidae, which lived in Europe during the Miocene epoch (16.9—16.0 mya), existing for approximately 0.9 million years.

Quercygale

Quercygale is an extinct genus of Miacidae, primitive carnivores that lived during the Eocene. The genus contains four species: Q. angustidens, Q. hastingsae, Q. helvetica, and Q. smithi. Phylogenetic analysis of the basicranial morphology of miacid carnivoramorphans suggests Quercygale is the most advanced miacid and sister to crown group Carnivora, predating the split between Feliformia and Caniformia., although another recent study places them as a stem group within Feliformia.

Viverridae

Viverridae is a family of small to medium-sized mammals, the viverrids (), comprising 15 genera, which are subdivided into 38 species. This family was named and first described by John Edward Gray in 1821. Members of this family are commonly called civets or genets. Viverrids are found in South and Southeast Asia, across the Wallace Line, all over Africa, and into southern Europe. Their occurrence in Sulawesi and in some of the adjoining islands shows them to be ancient inhabitants of the Old World tropics.

Viverrinae

The Viverrinae represent the largest subfamily within the Viverridae comprising five genera, which are subdivided into 22 species native to Africa and Southeast Asia. This subfamily was denominated and first described by John Edward Gray in 1864.

Viverroidea

Viverroidea is an infraorder of feliformia, containing both the family Viverridae, and the superfamily Herpestoidea.

Extant Carnivora species

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