Feliformia is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, viverrids, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Caniformia (also Canoidea, "dog-like" carnivorans).[1]
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.[2][3] Indeed, molecular phylogenies suggest that all extant Feliformia are monophyletic.[4]
Systematic classifications dealing with both extant and extinct taxa vary more widely.[5][6] Some separate the feliforms (extant and extinct) as Aeluroidea (superfamily) and Feliformia (suborder).[6] Others include all feliforms (extant, extinct and "possible ancestors") into the Feliformia suborder.[5] Some studies suggest this inclusion of "possible ancestors" into Feliformia (or even Carnivora) may be spurious.[7] 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).
Characteristics
Feliformia skull showing double-chambered bullae
All extant feliforms share a common attribute: their auditory bullae (bony capsules enclosing the middle and inner ear).[8] 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.[9]
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 (snouts) 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 feeding. However, omnivorous feliforms also exist, particularly in the family Viverridae.
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). Most caniforms are terrestrial and have non-retractile claws.
Seven families are extant, with 12 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 semiarboreal ambush hunters. Target prey vary 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.
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).[10][11] Phylogenetic analysis supports this view and places all of the Malagasy carnivorans in the family Eupleridae.[12]
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 (1lb) and the largest species at up to 12kg (26lb). 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.
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 2kg (4.5lb) to the tiger (Panthera tigris) at 300kg (660lb). Diet ranges from large to small mammals, birds and insects (depending on species size).
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 80kg (176lb) and represent one of the most prolific large carnivorans on the planet. The aardwolf is much smaller and is a specialised feeder, eating mainly harvester termites.
Family Herpestidae (mongooses, kusimanses, and the meerkat) 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 1kg (2.2lb) to 5kg (11lb), 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.
Family Nandiniidae (the 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 5kg (11lb).
Family Prionodontidae (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.[13]
Family Viverridae (all but two civets, genets, oyans, and the binturong) has 30 living species. They all have long bodies, short legs with retractile claws, and usually long tails. In weight, the species range from 0.5–14kg (1.1–30.9lb). 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
Feliform evolutionary timeline
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.[7]
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.
Carnivora is an order of placental mammals that have specialized in primarily eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the fifth largest order of mammals, comprising at least 279 species.
Viverridae is a family of small to medium-sized, feliform mammals. The viverrids comprise 33 species placed in 14 genera. This family was named and first described by John Edward Gray in 1821. Viverrids occur all over Africa, southern Europe, and South and Southeast Asia, across the Wallace Line.
The Asiatic linsang (Prionodon) is a genus comprising two species native to Southeast Asia: the banded linsang and the spotted linsang. Prionodon is considered a sister taxon of the Felidae.
Fissipedia is a former suborder of placental mammals comprising the largely land-based families of the order Carnivora. In some former taxonomic classifications, Pinnipedia is treated as an suborder in order Carnivora along side Fissipedia and Creodonta,. rather than as a subgroup of "dog-like" carnivorans.
Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include dogs, bears, raccoons, and mustelids. The Pinnipedia are also assigned to this group. The center of diversification for the Caniformia is North America and northern Eurasia. Caniformia stands in contrast to the other suborder of Carnivora, the Feliformia, the center of diversification of which was in Africa and southern Asia.
Mammalia is a class of animal within the phylum Chordata. Mammal classification has been through several iterations since Carl Linnaeus initially defined the class. No classification system is universally accepted; McKenna & Bell (1997) and Wilson & Reader (2005) provide useful recent compendiums. Many earlier ideas from Linnaeus et al. have been completely abandoned by modern taxonomists, among these are the idea that bats are related to birds or that humans represent a group outside of other living things. Competing ideas about the relationships of mammal orders do persist and are currently in development. Most significantly in recent years, cladistic thinking has led to an effort to ensure that all taxonomic designations represent monophyletic groups. The field has also seen a recent surge in interest and modification due to the results of molecular phylogenetics.
The fossa is a slender, long-tailed, cat-like mammal that is endemic to Madagascar. It is a member of the carnivoran family Eupleridae.
The spotted linsang is a linsang, a tree-dwelling carnivorous mammal, native to much of Southeast Asia. It is widely, though usually sparsely, recorded, and listed as Least Concern on the IUCN Red List.
Eupleridae is a family of carnivorans endemic to Madagascar and comprising 10 known living species in seven genera, commonly known as euplerids, Malagasy mongooses or Malagasy carnivorans. The best known species is the fossa, in the subfamily Euplerinae. All species of Euplerinae were formerly classified as viverrids, while all species in the subfamily Galidiinae were classified as herpestids.
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 ringed tails.
Civet cat is an imprecise term that is used for a variety of cat-like creatures including:
A civet is a small, lean, mostly nocturnal mammal native to tropical Asia and Africa, especially the tropical forests. The term civet applies to over a dozen different species, mostly from the family Viverridae. Most of the species's diversity is found in southeast Asia. Civets do not form a monophyletic group, as they consist only of certain members of the Viverridae, Eupleridae, and Nandiniidae.
Viverroidea is a clade within Feliformia, containing both the family Viverridae, and the superfamily Herpestoidea.
The Viverrinae represent the largest subfamily of the Viverridae comprising three genera, which are subdivided into six species native to Africa and Southeast Asia. This subfamily was denominated and first described by John Edward Gray in 1864.
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.
The linsangs are four species of tree-dwelling carnivorous mammals. The name of these species originated in the Javanese language as linsang or wlinsang, and previously, was translated incorrectly in English dictionaries as "otter". The two African species belong to the family Viverridae and the two Asiatic species belong to the family Prionodontidae. Formerly, both linsang genera were placed in the subfamily Viverrinae, along with several other genera, but recent research suggests that their relationships may be somewhat different.
Asiavorator is an extinct genus of civet-like carnivoran belonging in the family Stenoplesictidae. It was endemic to Asia and lived during the Eocene and Oligocene epochs.
Gustafsonia is an extinct genus of carnivoran belonging to the family Amphicyonidae. The type species, Gustafsonia cognita, was described in 1986 by Eric Paul Gustafson, who originally interpreted it as a miacid and named it Miacis cognitus. It was subsequently considered to be the only species of the diverse genus Miacis that belonged to the crown-group Carnivora, within the Caniformia, and it was ultimately assigned to the family Amphicyonidae. The type specimen or holotype was discovered in Reeve's bonebed, western Texas, in the Chambers Tuff Formation in 1986. The University of Texas holds this specimen. It is the only confirmed fossil of this species.
Herpestoidea is a superfamily of mammalia carnivores which includes mongooses, Malagasy carnivorans and the hyenas.
Alagtsavbaatar is an extinct species of carnivorous cat-like carnivoran belonging to the infraorder Aeluroidea. It was endemic to Asia, with all known specimens being found in Mongolia, and lived during the late Eocene epoch. It is a monotypic genus, with the type and only known species being A. indigenus, and is named after the Alag Tsav locality where its remains were first discovered.
↑ 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
1 2 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. ISSN1477-2019. S2CID86755875.
↑ 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?
↑ 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
1 2 Egi, N.; Tsubamoto, T.; etal. (2016). "Taxonomic revisions on nimravids and small feliforms (Mammalia, Carnivora) from the Upper Eocene of Mongolia". Historical Biology. 28 (1–2): 105–119. doi:10.1080/08912963.2015.1012508. S2CID86239933.
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