Carnivora
{{short description|Order of mammals}}
{{About||information on all meat-eating organisms|Carnivores|the alternative medicine "Carnivorans"|Venus flytrap#In alternative medicine}}
{{Use British English|date=March 2025}}
{{Use dmy dates|date=March 2025}}
{{Automatic taxobox
|name = Carnivorans
|fossil_range = {{fossil_range|51.88|0|earliest=66.0}}Early Eocene – Holocene{{Cite journal |last=Foley |first=N. M. |last2=Mason |first2=V. C. |last3=Harris |first3=A. J. |last4=Bredemeyer |first4=K. R. |last5=Damas |first5=J. |last6=Lewin |first6=H. A. |last7=Eizirik |first7=E. |last8=Gatesy |first8=J. |last9=Karlsson |first9=E. K. |last10=Lindblad-Toh |first10=K. |last11=Zoonomia Consortium |last12=Springer |first12=M. S. |last13=Murphy |first13=W. J. |year=2023 |title=A genomic timescale for placental mammal evolution |journal=Science |volume=380 |issue=6643 |at=eabl8189 |doi=10.1126/science.abl8189 |pmc=10233747 |pmid=37104581}}
|image =
File:Carnivora_portraits.jpg|300px
rect 580 485 0 0 Cheetah
rect 581 487 1190 971 Wolf
rect 0 486 577 972 Spotted hyena
rect 582 0 1191 485 Brown bear
rect 0 975 577 1387 Binturong
rect 582 974 1190 1388 Raccoon
rect 0 1392 577 1805 Indian grey mongoose
rect 581 1391 1188 1807 American mink
rect 0 1811 577 2231 Fossa
rect 581 1810 1190 2231 Walrus
|image_upright = 1.15
|image_caption = Various carnivorans, with feliforms to the left, and caniforms to the right
|display_parents = 2
|taxon = Carnivora
|authority = Bowdich, 1821Bowditch, T. E. 1821. An analysis of the natural classifications of Mammalia for the use of students and travelers J. Smith Paris. 115. (refer pages 24, 33)
|subdivision_ranks = Suborders
|subdivision =
|synonyms =
{{collapsible list |bullets=true |title=list of synonyms:
|Caniformes {{small|(Zagorodniuk, 2008)}}Zagorodniuk, I. (2008) [https://www.academia.edu/27585773/%D0%9D%D0%B0%D1%83%D0%BA%D0%BE%D0%B2%D1%96_%D0%BD%D0%B0%D0%B7%D0%B2%D0%B8_%D1%80%D1%8F%D0%B4%D1%96%D0%B2_%D1%81%D1%81%D0%B0%D0%B2%D1%86%D1%96%D0%B2_%D0%B2%D1%96%D0%B4_%D0%BE%D0%BF%D0%B8%D1%81%D0%BE%D0%B2%D0%B8%D1%85_%D0%B4%D0%BE_%D1%83%D0%BD%D1%96%D1%84%D1%96%D0%BA%D0%BE%D0%B2%D0%B0%D0%BD%D0%B8%D1%85_Scientific_names_of_mammalian_orders_from_descriptive_to_uniform_ "Scientific names of mammal orders: from descriptive to uniform"] Visnyk of Lviv University, Biology series, Is. 48. P. 33–43Zagorodniuk, I. (2014) [http://terioshkola.org.ua/library/pts12-divers/pts12-11-zag-mamdivers.pdf "Changes In Taxonomic Diversity Of Ukrainian Mammals For The Last Three Centuries: Extinct, Phantom And Alien Species"]. Proceedings of the Theriological School, Vol. 12: 3–16
|Carnaria {{small|(Haeckel, 1866)}}Haeckel, Ernst (1866.) [https://www.biodiversitylibrary.org/item/52177#page/11/mode/1up "Generelle Morphologie der Organismen."] Berlin: Georg Reimer.
|Carnassidentia {{small|(Wortman, 1901)}}J. L. Wortman (1901.) [https://www.biodiversitylibrary.org/page/40243865#page/157/mode/1up "Studies of Eocene Mammalia in the Marsh Collection, Peabody Museum."] The American Journal of Science, series 4 12:193-206
|Carnivoramorphia {{small|(Kalandadze & Rautian, 1992)}}Kalandadze, N. N. and S. A. Rautian (1992.) "Systema mlekopitayushchikh i istorygeskaya zoogeographei [The system of mammals and historical zoogeography]." Sbornik Trudov Zoologicheskogo Muzeya Moskovskogo Goschdarstvennoro Universiteta 29:44–152.
|Carnivores {{small|(Cuvier, 1817)}}Georges Cuvier, Pierre André Latreille (1817.) [https://archive.org/details/bub_gb_H9sTAAAAQAAJ/page/n3/mode/2up "Le Règne Animal Distribué d'après son Organisation, pour Servir de Base à l'Histoire Naturelle des Animaux et d'Introduction à l'Anatomie Comparée"] Déterville libraire, Imprimerie de A. Belin, Paris, 4 Volumes
|Carnivori {{small|(Vieq d'Azyr, 1792)}}Félix Vicq-Dazyr (1792.) [https://books.google.com/books?id=8E5wN2Ake3kC&pg=PP9 "Encyclopédie Méthodique, Vol. 2: Système Anatomique, Quadrupèdes"] Panckoucke
|Carnivorida {{small|(Pearse, 1936)}}Arthur Sperry Pearse, (1936) [https://archive.org/details/zoologicalnamesl1936pear/page/24/mode/2up?q=Creodontina "Zoological names. A list of phyla, classes, and orders, prepared for section F, American Association for the Advancement of Science"] American Association for the Advancement of ScienceG. G. Simpson (1952) [https://www.jstor.org/stable/2411849?seq=1 "For and Against Uniform Endings in Zoological Nomenclature"] in "Systematic Zoology Vol. 1, No. 1 (Spring, 1952)", pp. 20–23, Published By: Taylor & Francis, Ltd.
|Carnivoriformes {{small|(Kinman, 1994)}}Kenneth E. Kinman (1994.) "The Kinman System: Toward a Stable Cladisto-Eclectic Classification of Organisms: Living and Extinct, 48 Phyla, 269 Classes, 1,719 Orders", Hays, Kan. (P. O. Box 1377, Hays 67601), 88 pages
|Carnivoripedida {{small|(Vyalov, 1966)}}O. S. Vyalov (1966.) "Sledy Zhiznedeyatel'nosti Organizmov i ikh Paleontologicheskoye Znacheniye [Traces of Vital Activity of Organisms and their Paleontological Significance]" Naukova Dumka, Kyiv, 1-219
|Cynofeliformia {{small|(Ginsburg, 1982)}}
|Cynofeloidea {{small|(Hough, 1953)}}Hough, J. R. (1953.) [https://pubs.usgs.gov/pp/0243g/report.pdf "Auditory region in North American fossil Felidae: Its significance in phylogeny."] United States Geological SurveyProfessional Papers, 243-G,95–115.
|Cynosia {{small|(Rafinesque, 1815)}}{{Cite journal |last=Rafinesque |first=Constantine Samuel |date=1815 |title=Analyse de la Nature ou tableau de l'univers et des corps organisés |volume=1815 |url=https://www.biodiversitylibrary.org/item/188066#page/11/mode/1up |publisher=Palermo, Aux dépens de l'auteur, 223 pp.}}
|Digitigrada {{small|(Illiger, 1811)}}Johann Karl Wilhelm Illiger (1811.) [https://www.biodiversitylibrary.org/item/188747#page/7/mode/1up "Prodromus Systematis Mammalium et Avium."] Berlin: Sumptibus C. Salfeld, xviii, 301 pages
|Digitigradae {{small|(Gray, 1821)}}{{cite journal |last=Gray |first=J. E. |year=1821 |title=On the natural arrangement of vertebrose animals |url=https://archive.org/stream/londonmedicalre08unkngoog#page/n314/mode/1up |journal=London Medical Repository |volume=15 |issue=1 |pages=296–310}}
|Eucarnivora {{small|(Mekayev, 2002)}}Mekayev, Y. A. (2002.) "The faunagenesis and classification of mammals." Petrov’s Academy of Sciences and Arts, St. Petersburg, 1–895.
|Ferae {{small|(Linnaeus, 1758)}}
|Fissipeda {{small|(Blumenbach, 1791)}}Johann Friedrich Blumenbach (1791.) [https://www.deutschestextarchiv.de/book/view/blumenbach_naturgeschichte_1791?p=5 "Handbuch der Naturgeschichte. Vierte auflage."] Göttingen, Johann Christian Dieterich, xii+704+[33] pp., 3 pls.
|Neocarnivora {{small|(Radinsky, 1977)}}Leonard Radinsky (1977.) [https://www.cambridge.org/core/journals/paleobiology/article/abs/brains-of-early-carnivores/AE507D4080D40AA8254D1306CC349824 "Brains of early carnivores."] Paleobiology, Volume 3, Issue 4, pp. 333 – 349
|Plantigrada {{small|(Illiger, 1811)}}
}}
|range_map = Carnivora_Distribution.png
|range_map_caption = The extant distribution and density of Carnivora species.
}}
Carnivora ({{IPAc-en|k|ɑr|ˈ|n|ɪ|v|ər|ə}} {{respell|kar|NIH|vər|ə}}) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals,{{cite web |url= https://www.mammaldiversity.org/taxa.html |title= Higher Taxonomy |author= |date= |website= ASM Mammal Diversity Database |publisher= American Society of Mammalogy |access-date= 22 August 2024}} comprising at least 279 species. Carnivorans are found on every major landmass and in a variety of habitats, ranging from the cold polar regions of Earth to the hyper-arid region of the Sahara Desert and the open seas. Carnivorans exhibit a wide array of body plans, varying greatly in size and shape.
Carnivora are divided into two suborders, the Feliformia, containing the true felids and several {{nowrap|"cat-like"}} animals; and the Caniformia, containing the true canids and many {{nowrap|"dog-like"}} animals. The feliforms include the Felidae, Viverridae, hyena, and mongoose families, the majority of which live only in the Old World; cats are the only exception, occurring in the Old World and the New World, entering the Americas via the Bering land bridge. The caniforms include the Caninae, Procyonidae, bears, mustelids, skunks, and pinnipeds that occur worldwide with immense diversity in their morphology, diet, and behavior.
Etymology
The word Carnivora is derived from Latin carō (stem carn-) 'flesh' and vorāre 'to devour'.{{cite web |title=Carnivora {{!}} Etymology of the name Carnivora by etymonline |url=https://www.etymonline.com/word/Carnivora |website=www.etymonline.com |access-date=13 February 2025 |language=en}}
Phylogeny
The oldest known carnivoran line mammals (Carnivoramorpha) appeared in North America 6 million years after the Cretaceous–Paleogene extinction event.{{cite book|last1=Hunter|first1=L.|title=Field Guide to Carnivores of the World.|publisher=Bloomsbury Wildlife|year=2018|isbn=978-1472950796|pages=1–271}} These early ancestors of carnivorans would have resembled small weasel or genet-like mammals, occupying a nocturnal shift on the forest floor or in the trees, as other groups of mammals like the mesonychians and later the creodonts were occupying the megafaunal faunivorous niche. However, following the extinction of mesonychians and the oxyaenid creodonts at the end of the Eocene, carnivorans quickly moved into this niche, with forms like the nimravids being the dominant large-bodied ambush predators during the Oligocene alongside the hyaenodont creodonts (which similarly produced larger, more open-country forms at the start of the Oligocene). By the time Miocene epoch appeared, most if not all of the major lineages and families of carnivorans had diversified and become the most dominant group of large terrestrial predators in Eurasia and North America, with various lineages being successful in megafaunal faunivorous niches at different intervals during the Miocene and later epochs.
Systematics
=Evolution=
Image:Tapocyon robustus.jpg robustus, a species of miacid]]
The order Carnivora belongs to a group of mammals known as Laurasiatheria, which also includes other groups such as bats and ungulates.{{Cite journal | last1 = Waddell | first1 = Peter J.| last2 = Okada | first2 = Norihiro| last3 = Hasegawa | first3 = Masami| doi = 10.1093/sysbio/48.1.1 | title = Towards Resolving the Interordinal Relationships of Placental Mammals | journal = Systematic Biology | volume = 48 | issue = 1 | pages = 1–5 | year = 1999 | pmid = 12078634| doi-access = free }}{{cite journal |last1=Tsagkogeorga |first1=G |last2=Parker |first2=J |last3=Stupka |first3=E |last4=Cotton |first4=J.A. |last5=Rossiter |first5=S.J. |year=2013 |title=Phylogenomic analyses elucidate the evolutionary relationships of bats |journal=Current Biology |volume=23 |issue=22 |pages=2262–2267 |doi=10.1016/j.cub.2013.09.014 |pmid=24184098|doi-access=free |bibcode=2013CBio...23.2262T }} Within this group the carnivorans are placed in the clade Ferae. Ferae includes the closest extant relative of carnivorans, the pangolins, as well as several extinct groups of mostly Paleogene carnivorous placentals such as the creodonts, the arctocyonians, and mesonychians.{{cite journal|last1=Halliday|first1=Thomas J. D.|last2=Upchurch|first2=Paul|last3=Goswami|first3=Anjali|title=Resolving the relationships of Paleocene placental mammals|journal=Biological Reviews|volume=92|issue=1|year=2015|pages=521–550|issn=1464-7931|doi=10.1111/brv.12242|pmid=28075073|url=http://discovery.ucl.ac.uk/1473028/1/Halliday_et_al-Biological_Reviews.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://discovery.ucl.ac.uk/1473028/1/Halliday_et_al-Biological_Reviews.pdf |archive-date=2022-10-09 |url-status=live|pmc=6849585}} The creodonts were originally thought of as the sister taxon to the carnivorans, perhaps even ancestral to, based on the presence of the carnassial teeth,{{cite book | last = McKenna | first = M. C. | date = 1975 | chapter = Toward a phylogenetic classification of the Mammalia | pages = 21–46 | editor1-first = W. P. | editor1-last = Luckett | editor2-first = F. S. | editor2-last = Szalay | title = Phylogeny of the Primates | publisher = Plenum | location = New York }} but the nature of the carnassial teeth is different between the two groups. In carnivorans, the carnassials are positioned near the front of the molar row, while in the creodonts, they are positioned near the back of the molar row,{{cite book|first1=George A.|last1=Feldhamer|first2=Lee C.|last2=Drickamer|first3=Stephen H.|last3=Vessey|first4=Joseph F.|last4=Merritt|first5=Carey|last5=Krajewski|title=Mammalogy: Adaptation, Diversity, Ecology|location=Baltimore|publisher=Johns Hopkins University Press|year=2015|page=[https://archive.org/details/mammalogyadaptat03edunse/page/356 356]|isbn=978-0801886959|url=https://archive.org/details/mammalogyadaptat03edunse/page/356}} and this suggests a separate evolutionary history and an order-level distinction.{{cite book|first1=Alan|last1=Turner|first2=Mauricio|last2=Antón|title=Evolving Eden: An Illustrated Guide to the Evolution of the African Large-Mammal Fauna|location=New York|publisher=Columbia University Press|year=2004|page=77|isbn=978-0-231-11944-3}} In addition, phylogenetic analysis suggests that creodonts are more closely related to pangolins while mesonychians might be the sister group to carnivorans and their stem-relatives.
The closest stem-carnivorans are the miacoids. The miacoids include the families Viverravidae and Miacidae, and together the Carnivora and Miacoidea form the stem-clade Carnivoramorpha. The miacoids were small, genet-like carnivoramorphs that occupy a variety of niches such as terrestrial and arboreal habitats. Studies have shown that while viverravids are a monophyletic basal group, the miacids are paraphyletic with respect to Carnivora (as shown in the phylogeny below).Bryant, H.N., and M. Wolson (2004) [https://web.archive.org/web/20080706090739/http://www.ohiou.edu/phylocode/IPNM.pdf "Phylogenetic Nomenclature of Carnivoran Mammals."] First International Phylogenetic Nomenclature Meeting. Paris, Museum National d’Histoire Naturelle{{cite book |author1=John J. Flynn |author2=John A. Finarelli |author3=Michelle Spaulding |year=2010 |chapter=Phylogeny of the Carnivora and Carnivoramorpha, and the use of the fossil record to enhance understanding of evolutionary transformations |editor1=Anjali Goswami |editor2=Anthony Friscia |title=Carnivoran evolution. New views on phylogeny, form and function |url=https://archive.org/details/carnivoranevolut00gosw |url-access=limited |publisher=Cambridge University Press |pages=[https://archive.org/details/carnivoranevolut00gosw/page/n40 25]–63 |isbn=9781139193436 |doi=10.1017/CBO9781139193436.003}}
Carnivoramorpha as a whole first appeared in the Paleocene of North America about 60 million years ago.{{cite journal | author = Polly, David, Gina D. Wesley-Hunt, Ronald E. Heinrich, Graham Davis and Peter Houde | year = 2006 | title = Earliest known carnivoran auditory bulla and support for a recent origin of crown-clade carnivora (Eutheria, Mammalia) | journal = Palaeontology | volume = 49 | issue = 5 | pages = 1019–1027 | doi = 10.1111/j.1475-4983.2006.00586.x | doi-access = free }} Crown carnivorans first appeared around 42 million years ago in the Middle Eocene.{{cite journal |author=Heinrich, R.E. |author2=Strait, S.G. |author3=Houde, P. |date=January 2008 |title=Earliest Eocene Miacidae (Mammalia: Carnivora) from northwestern Wyoming |journal=Journal of Paleontology |volume=82 |issue=1 |pages=154–162 |doi=10.1666/05-118.1|bibcode=2008JPal...82..154H |s2cid=35030667 }} Their molecular phylogeny shows the extant Carnivora are a monophyletic group, the crown group of the Carnivoramorpha.{{cite journal |last1=Eizirik |first1=E. |last2=Murphy |first2=W.J. |last3=Koepfli |first3=K.P. |last4=Johnson |first4=W.E. |last5=Dragoo |first5=J.W. |last6=O'Brien |first6=S.J. |date=July 2010 |title=Pattern and timing of the diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences |journal=Molecular Phylogenetics and Evolution |volume=56 |issue= 1|pages=49–63 |doi=10.1016/j.ympev.2010.01.033 |pmid=20138220|pmc=7034395 |bibcode=2010MolPE..56...49E }} From there carnivorans have split into two clades based on the composition of the bony structures that surround the middle ear of the skull, the cat-like feliforms and the dog-like caniforms.{{cite book | last1 = Wang | first1 = X. | last2 = Tedford | first2 = R. H. | year = 2008 | title = Dogs: Their Fossil Relatives and Evolutionary History| publisher = Columbia University Press |location = New York | pages = 1–232 |isbn = 978-0-231-13529-0}} 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.{{cite book |author=R. F. Ewer |title=The Carnivores |url=https://books.google.com/books?id=IETMd3-lSlkC |year=1973 |publisher=Cornell University Press |isbn=0-8014-8493-6}} Initially, the early representatives of carnivorans were small as the creodonts (specifically, the oxyaenids) and mesonychians dominated the apex predator niches during the Eocene, but in the Oligocene, carnivorans became a dominant group of apex predators with the nimravids, and by the Miocene most of the extant carnivoran families have diversified and become the primary terrestrial predators in the Northern Hemisphere.
=Classification of the extant carnivorans=
{{see also|List of carnivorans}}
In 1758, the Swedish botanist Carl Linnaeus placed all carnivorans known at the time into the group Ferae (not to be confused with the modern concept of Ferae which also includes pangolins) in the tenth edition of his book Systema Naturae. He recognized six genera: Canis (canids and hyaenids), Phoca (pinnipeds), Felis (felids), Viverra (viverrids, herpestids, and mephitids), Mustela (non-badger mustelids), Ursus (ursids, large species of mustelids, and procyonids).{{cite book | author = Linnaeus, C. | year = 1758 | title = Sistema naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonimis locis. Tomus I | publisher = Impensis direct. Laurentii Salvii, Holmia | pages = 20–32}} It was not until 1821 that the English writer and traveler Thomas Edward Bowdich gave the group its modern and accepted name.Bowditch, T. E. 1821. An analysis of the natural classifications of Mammalia for the use of students and travelers J. Smith Paris. 115. (refer pages 24, 33)
Initially, the modern concept of Carnivora was divided into two suborders: the terrestrial Fissipedia and the marine Pinnipedia.{{cite journal | last1 = Simpson | first1 = G.G. | year = 1945 | title = The principles of classification and a classification of mammals | journal = Bulletin of the AMNH | volume = 85 | pages = 1–350| hdl = 2246/1104 }} Below is the classification of how the extant families were related to each other after American paleontologist George Gaylord Simpson in 1945:
- Order Carnivora Bowdich, 1821
- Suborder Fissipedia Blumenbach, 1791
- Superfamily Canoidea G. Fischer de Waldheim, 1817
- Family Canidae G. Fischer de Waldheim, 1817 – dogs
- Family Ursidae G. Fischer de Waldheim, 1817 – bears
- Family Procyonidae Bonaparte, 1850 – raccoons, ringtails and coatis (also included red pandas as subfamily Ailurinae, which is now treated as a family)
- Family Mustelidae G. Fischer de Waldheim, 1817 – badgers, otters, weasels and skunks (as subfamily Mephitinae, now treated as family)
- Superfamily Feloidea G. Fischer de Waldheim, 1817
- Family Viverridae J. E. Gray, 1821 – civets and allies, including mongooses (now family Herpestidae), African palm civets (now family Nandiniidae) and Asiatic linsangs (now family Prionodontidae)
- Family Hyaenidae J. E. Gray, 1821 – hyenas
- Family Felidae G. Fischer de Waldheim, 1817 – cats
- Suborder Pinnipedia Iliger, 1811
- Family Otariidae J. E. Gray, 1825 – eared seals
- Family Odobenidae J. A. Allen, 1880 – walruses
- Family Phocidae J. E. Gray, 1821 – earless seals
Since then, however, the methods in which mammalogists use to assess the phylogenetic relationships among the carnivoran families has been improved with using more complicated and intensive incorporation of genetics, morphology and the fossil record. Research into Carnivora phylogeny since 1945 has found Fissipedia to be paraphyletic in respect to Pinnipedia,{{Cite journal
|author1=Arnason, U. |author2=Gullberg, A. |author3=Janke, A. |author4=Kullberg, M. |title=Mitogenomic analyses of caniform relationships |doi=10.1016/j.ympev.2007.06.019 |journal=Molecular Phylogenetics and Evolution |volume=45 |issue=3 |pages=863–74 |year=2007 |pmid=17919938|bibcode=2007MolPE..45..863A }} with pinnipeds being either more closely related to bears or to weasels.{{cite journal |author1=Lento, G. M. |author2=Hickson, R. E. |author3=Chambers, G. K. |author4=Penny, D. |year=1995 |title=Use of spectral analysis to test hypotheses on the origin of pinnipeds |journal=Molecular Biology and Evolution |volume=12 |issue=1 |pages=28–52 |pmid=7877495 |doi=10.1093/oxfordjournals.molbev.a040189|doi-access= }}{{cite journal|author1=Hunt, R. M. Jr. |author2=Barnes, L. G. |year=1994|title=Basicranial evidence for ursid affinity of the oldest pinnipeds|journal=Proceedings of the San Diego Society of Natural History|volume=29|pages=57–67|url=https://archive.org/details/cbarchive_36692_basicranialevidenceforursidaff1990|format=PDF}}{{Cite journal |author1=Higdon, J. W. |author2=Bininda-Emonds, O. R. |author3=Beck, R. M. |author4=Ferguson, S. H. |title=Phylogeny and divergence of the pinnipeds (Carnivora: Mammalia) assessed using a multigene dataset |doi=10.1186/1471-2148-7-216 |journal=BMC Evolutionary Biology |volume=7 |page=216 |year=2007 |issue=1 |pmid=17996107 |pmc=2245807 |doi-access=free |bibcode=2007BMCEE...7..216H }}{{Cite journal
|author1=Sato, J. J. |author2=Wolsan, M. |author3=Suzuki, H. |author4=Hosoda, T. |author5=Yamaguchi, Y. |author6=Hiyama, K. |author7=Kobayashi, M. |author8=Minami, S. |doi=10.2108/zsj.23.125 |title=Evidence from nuclear DNA sequences sheds light on the phylogenetic relationships of Pinnipedia: Single origin with affinity to Musteloidea |journal=Zoological Science |volume=23 |issue=2 |pages=125–46 |year=2006 |pmid=16603806|hdl=2115/13508 |s2cid=25795496 |hdl-access=free }}{{Cite journal |author1=Flynn, J. J. |author2=Finarelli, J. A. |author3=Zehr, S. |author4=Hsu, J. |author5=Nedbal, M. A. |title=Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships |doi=10.1080/10635150590923326 |journal=Systematic Biology |volume=54 |issue=2 |pages=317–37 |year=2005 |pmid=16012099|doi-access=free }} The small carnivoran families Viverridae,{{cite journal | last1 = Gaubert | first1 = P. | last2 = Veron | first2 = G. | year = 2003 | title = Exhaustive sample set among Viverridae reveals the sister-group of felids: the linsangs as a case of extreme morphological convergence within Feliformia | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 270 | issue = 1532| pages = 2523–2530 | doi = 10.1098/rspb.2003.2521 | pmid = 14667345 | pmc = 1691530 }} Procyonidae, and Mustelidae have been found to be polyphyletic:
- Mongooses and a handful of Malagasy endemic species are found to be in a clade with hyenas, with the Malagasy species being in their own family Eupleridae.Anne D. Yoder and John J. Flynn 2003: [https://web.archive.org/web/20070612131729/http://www.biology.duke.edu/yoderlab/pdfs/ipYoderFlynnNHMad.pdf Origin of Malagasy Carnivora]Yoder, A., M. Burns, S. Zehr, T. Delefosse, G. Veron, S. Goodman, J. Flynn. 2003: [https://web.archive.org/web/20070612131729/http://www.biology.duke.edu/yoderlab/pdfs/ipYoderFlynnNHMad.pdf Single origin of Malagasy Carnivora from an African ancestor – Letters to Nature]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?
- The African palm civet is a basal cat-like carnivoran.{{Cite journal |title=Pattern and timing of diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences |first1=E. |last1=Eizirik |first2=W. J. |last2=Murphy |first3=K. P. |last3=Koepfli |first4=W. E. |last4=Johnson |first5=J. W. |last5=Dragoo |first6 = R. K. |last6=Wayne |first7=S. J. |last7=O'Brien |journal=Molecular Phylogenetics and Evolution |date=2010 |pages=49–63 |volume=56 |issue=1 |doi=10.1016/j.ympev.2010.01.033 |pmid=20138220|pmc=7034395 |bibcode=2010MolPE..56...49E }}
- The linsang is more closely related to cats.{{cite book | last1 = Gaubert | first1 = P. | year = 2009 | chapter = Family Prionodontidae (Linsangs) | editor1-last = Wilson | editor1-first = D.E.| editor2-last = Mittermeier | editor2-first = R.A. | title = Handbook of the Mammals of the World – Volume 1 | publisher = Lynx Ediciones | location=Barcelona | pages =170–173 | isbn = 978-84-96553-49-1}}
- Pandas are not procyonids nor are they a natural grouping.{{cite journal |author=Salesa, M. |author2=M. Antón |author3=S. Peigné |author4=J. Morales |year=2006 |title=Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas |journal=Proceedings of the National Academy of Sciences |volume=103 |pages=379–382 |doi=10.1073/pnas.0504899102 |pmid=16387860 |issue=2 |pmc=1326154|bibcode=2006PNAS..103..379S |doi-access=free }} The giant panda is a true bear{{cite journal |last1=Yu |first1=Li |last2=Li |first2=Yi-Wei |last3=Ryder |first3=Oliver A. |last4=Zhang |first4=Ya-Ping |title=Analysis of complete mitochondrial genome sequences increases phylogenetic resolution of bears (Ursidae), a mammalian family that experienced rapid speciation| journal=BMC Evolutionary Biology |date=2007 |volume=7 |issue=198 |pages=198 |doi=10.1186/1471-2148-7-198 |pmid=17956639 |pmc=2151078 |doi-access=free |bibcode=2007BMCEE...7..198Y }}{{Cite journal | last=Krause | first=J. | author2=Unger, T. | author3=Noçon, A. | author4=Malaspinas, A. | author5=Kolokotronis, S. | author6=Stiller, M. | author7=Soibelzon, L. | author8=Spriggs, H. | author9=Dear, P. H. | author10=Briggs, A. W. | author11=Bray, S. C. E. | author12=O'Brien, S. J. | author13=Rabeder, G. | author14=Matheus, P. | author15=Cooper, A. | author16=Slatkin, M. | author17=Pääbo, S. | author18=Hofreiter, M. | title=Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary | journal=BMC Evolutionary Biology | volume=8 | issue=220 | page=220 | year=2008 | pmid=18662376 | pmc=2518930| doi=10.1186/1471-2148-8-220 | doi-access=free | bibcode=2008BMCEE...8..220K }} while the red panda is a distinct family.{{Cite journal |last1=Mehta |first1=R. S. |last2=Slater|first2=G. J.|last3=Law|first3=C. J.|date=2018 |title=Lineage Diversity and Size Disparity in Musteloidea: Testing Patterns of Adaptive Radiation Using Molecular and Fossil-Based Methods |journal=Systematic Biology |volume=67 |issue=1 |pages=127–144 |doi=10.1093/sysbio/syx047 |pmid=28472434|issn=1063-5157|doi-access=free }}
- Skunks and stink badgers are placed in their own family, and are the sister group to a clade containing Ailuridae, Procyonidae and Mustelidae sensu stricto.{{cite journal |vauthors=Koepfli KP, Deere KA, Slater GJ, etal |title=Multigene phylogeny of the Mustelidae: Resolving relationships, tempo and biogeographic history of a mammalian adaptive radiation |journal=BMC Biol. |volume=6 |pages=4–5 |year=2008 |pmid=18275614 |doi=10.1186/1741-7007-6-10 |pmc=2276185 |doi-access=free }}
Below is a table chart of the extant carnivoran families and number of extant species recognized by various authors of the first (2009{{cite book |title=Handbook of the Mammals of the World – Volume 1 |publisher=Lynx Ediciones |year=2009 |isbn=978-84-96553-49-1 |editor1-last=Wilson |editor1-first=D.E. |location=Barcelona |pages=1–728 |editor2-last=Mittermeier |editor2-first=R.A.}}) and fourth (2014{{cite book |title=Handbook of the Mammals of the World – Volume 4 |publisher=Lynx Ediciones |year=2014 |isbn=978-84-96553-93-4 |editor1-last=Wilson |editor1-first=D.E. |location=Barcelona |pages=1–614 |editor2-last=Mittermeier |editor2-first=R.A.}}) volumes of the Handbook of the Mammals of the World:
class="wikitable"
|colspan="100%" align="center" bgcolor="#c2c2a9"|Carnivora Bowdich, 1821 | |||||
colspan="100%" align="center" bgcolor="#d9d9c1"|Feliformia Kretzoi, 1945 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Nandinioidea Pocock, 1929 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
---|---|---|---|---|---|
Nandiniidae Pocock, 1929
|1 |Nandinia binotata (J. E. Gray, 1830) | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Feloidea G. Fischer de Waldheim, 1817 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Felidae G. Fischer de Waldheim, 1817
|Cats (including domestic cats, tigers, leopards, jaguars, lions, cheetahs, ocelots, etc.) |Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands) |37 |Felis catus Linnaeus, 1758 | |||||
Prionodontidae Horsfield, 1822
|2 |Prionodon linsang (Hardwicke, 1821) | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Viverroidea J. E. Gray, 1821 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Viverridae J. E. Gray, 1821
|Southern Europe, Indomalayan realm, and Africa (introduced to Madagascar) |34 |Viverra zibetha Linnaeus, 1758 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Herpestoidea Bonaparte, 1845 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Hyaenidae J. E. Gray, 1821
|Hyenas |Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent |4 |Hyaena hyaena (Linnaeus, 1758) | |||||
Herpestidae Bonaparte, 1845
|Mongooses |Iberian Peninsula, Africa, the Middle East, the Caucasus, Central Asia, and the Indomalayan realm |34 |Herpestes ichneumon (Linnaeus, 1758) | |||||
Eupleridae Chenu, 1850
|Malagasy mongooses and civets |8 |Eupleres goudotii Doyère, 1835 | |||||
colspan="100%" align="center" bgcolor="#d9d9c1"|Caniformia Kretzoi, 1945 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Canoidea G. Fischer de Waldheim, 1817 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Canidae G. Fischer de Waldheim, 1817
|Dogs (including domestic dogs, wolves, foxes, dingoes, jackals, coyotes, etc.) |Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands) |35 |Canis familiaris Linnaeus, 1758 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Ursoidea G. Fischer de Waldheim, 1817 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Ursidae G. Fischer de Waldheim, 1817
|Bears |8 |Ursus arctos Linnaeus, 1758 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Phocoidea J. E. Gray, 1821 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Odobenidae J. A. Allen, 1880
|Walruses |The North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere |1 |Odobenus rosmarus (Linnaeus, 1758) | |||||
Otariidae J. E. Gray, 1825
|Subpolar, temperate, and equatorial waters throughout the Pacific and Southern Oceans and the southern Indian and Atlantic Oceans |15 |Otaria flavescens (Shaw, 1800) | |||||
Phocidae J. E. Gray, 1821
|The sea and Lake Baikal |18 |Phoca vitulina Linnaeus, 1758 | |||||
colspan="100%" align="center" bgcolor="#ebebd2"|Musteloidea G. Fischer de Waldheim, 1817 | |||||
Family | English name | Distribution | Number of extant species | Type taxon | Image figure |
Mephitidae Bonaparte, 1845
|Skunks and stink badgers |Americas, western Philippines, and Indonesia and Malaysia |12 |Mephitis mephitis (Schreber, 1776) | |||||
Ailuridae J. E. Gray, 1843
|Eastern Himalayas and southwestern China |1 |Ailurus fulgens F. Cuvier, 1825 | |||||
Procyonidae J. E. Gray, 1825
|Raccoons, olingos, ringtails, coatis, cacomistles, and kinkajous |Americas (introduced to Europe, the Caucasus, and Japan) |12 |Procyon lotor (Linnaeus, 1758) | |||||
Mustelidae G. Fischer de Waldheim, 1817
|Weasels, otters, wolverines, polecats, badgers, martens, and grisons |Americas, Africa, and Eurasia (introduced to Australasia and several islands) |57 |Mustela erminea Linnaeus, 1758 |
Anatomy
=Skull=
Image:Em - Cryptoprocta ferox - 2.jpg (Cryptoprocta ferox). Note the large and conical canine and carnassial teeth common in feliforms.]]
The canine teeth are usually large, conical, thick and stress resistant. All of the terrestrial species of carnivorans have three incisors on each side of each jaw (the exception is the sea otter (Enhydra lutris) which only has two lower incisor teeth). The third molar has been lost. The carnassial pair is made up of the fourth upper premolar and the first lower molar teeth. Like most mammals, the dentition is heterodont, though in some species, such as the aardwolf (Proteles cristata), the teeth have been greatly reduced and the cheek teeth are specialised for eating insects. In pinnipeds, the teeth are homodont as they have evolved to grasp or catch fish, and the cheek teeth are often lost. In bears and raccoons, the carnassial pair is secondarily reduced. The skulls are heavily built with a strong zygomatic arch. Often a sagittal crest is present, sometimes more evident in sexually dimorphic species such as sea lions and fur seals, though it has also been greatly reduced in some small carnivorans.{{cite book | last1 = Vaughan | first1 = T. A. | last2 = Ryan | first2 = J. M. | last3 = Czaplewski | first3 = N. J. | year = 2013 | title = Mammalogy | publisher = Jones & Bartlett Learning | location = Burlington, Massachusetts | pages = 1–750 |isbn = 9781284032093}} The braincase is enlarged with the frontoparietal bone at the front. In most species, the eyes are at the front of the face. In caniforms, the rostrum is usually long with many teeth, while in feliforms it is shorter with fewer teeth. The carnassial teeth of feliforms are generally more sectional than those of caniforms.
The turbinates are large and complex in comparison to other mammals, providing a large surface area for olfactory receptors.
=Postcranial region=
Image:Canis mesomelas vs. Arctocephalus pusillus.jpg (Lupulella mesomelas) trying to predate on a brown fur seal (Arctocephalus pusillus) pup. These two species illustrate the diversity in bodyplan seen among carnivorans, especially between pinnipeds and their terrestrial relatives.]]
Aside from an accumulation of characteristics in the dental and cranial features, not much of their overall anatomy unites carnivorans as a group.{{cite book | last1 = Nowak | first1 = R. M. | year = 2005 | title = Walker's Carnivores of the World | url = https://archive.org/details/walkerscarnivore00nowa | url-access = limited | publisher = Johns Hopkins University Press | location= Baltimore, Maryland | pages = [https://archive.org/details/walkerscarnivore00nowa/page/n11 1]–328 |isbn = 0801880335}} All species of carnivorans are quadrupedal and most have five digits on the front feet and four digits on the back feet. In terrestrial carnivorans, the feet have soft pads. The feet can either be digitigrade as seen in cats, hyenas, and dogs or plantigrade as seen in bears, skunks, raccoons, weasels, civets, and mongooses. In pinnipeds, the limbs have been modified into flippers.File:Panthera tigris11.jpg, have pads on their feet.]]
Unlike cetaceans and sirenians, which have fully functional tails to help them swim, pinnipeds use their limbs underwater to swim. Earless seals use their back flippers; sea lions and fur seals use their front flippers, and the walrus uses all of its limbs. As a result, pinnipeds have significantly shorter tails than other carnivorans.
Aside from the pinnipeds, dogs, bears, hyenas, and cats all have distinct and recognizable appearances. Dogs are usually cursorial mammals and are gracile in appearance, often relying on their teeth to hold prey; bears are much larger and rely on their physical strength to forage for food. Compared to dogs and bears, cats have longer and stronger forelimbs armed with retractable claws to hold on to prey. Hyenas are dog-like feliforms that have sloping backs due to their front legs being longer than their hind legs. The raccoon family and red panda are small, bear-like carnivorans with long tails. The other small carnivoran families Nandiniidae, Prionodontidae, Viverridae, Herpestidae, Eupleridae, Mephitidae, and Mustelidae have through convergent evolution maintained the small, ancestral appearance of the miacoids, though there is some variation seen such as the robust and stout physicality of badgers and the wolverine (Gulo gulo).
Most carnivoran species have a well-defined breeding season.{{Cite book |last=Knobil |first=Ernst |url=https://books.google.com/books?id=6j-mOAcNnPUC&dq=carnivora&pg=PA776 |title=Knobil and Neill's Physiology of Reproduction |date=2006 |publisher=Gulf Professional Publishing |isbn=978-0-12-515401-7 |language=en}} Male carnivorans will usually have bacula, which are absent in hyenas and binturongs.{{Cite journal|last1=Schultz|first1=Nicholas G.|last2=Lough-Stevens|first2=Michael|last3=Abreu|first3=Eric|last4=Orr|first4=Teri|last5=Dean|first5=Matthew D.|date=2016-06-01|title=The Baculum was Gained and Lost Multiple Times during Mammalian Evolution|url= |journal=Integrative and Comparative Biology|volume=56|issue=4|language=en|pages=644–56|doi=10.1093/icb/icw034|issn=1540-7063|pmid=27252214|pmc=6080509}}
The length and density of the fur can vary depending on the environment that the species inhabits. In warm climate species, the fur is often short in length and lighter. In cold climate species, the fur is can be either dense or long, often with an oily substance that helps to retain heat. The pelage coloration differs between species, often including black, white, orange, yellow, red, and many shades of grey and brown. Some are striped, spotted, blotched, banded, or otherwise boldly patterned. There may be a correlation between habitat and color pattern; for example, spotted or banded species tend to be found in heavily forested environments. Some species like the grey wolf are polymorphic with different individual having different coat colors. The arctic fox (Vulpes lagopus) and the stoat (Mustela erminea) have fur that changes from white and dense in the winter to brown and sparse in the summer. In pinnipeds and polar bears, a thick insulating layer of blubber helps maintain their body temperature.
Sexual dimorphism
{{excerpt|Sexual dimorphism in Carnivorans}}
Relationship with humans
Carnivorans are arguably the group of mammals of most interest to humans. The dog is noteworthy for not only being the first species of carnivoran to be domesticated, but also the first species of any taxon. In the last 10,000 to 12,000 years, humans have selectively bred dogs for a variety of different tasks and today there are well over 400 breeds. The cat is another domesticated carnivoran and it is today considered one of the most successful species on the planet, due to their close proximity to humans and the popularity of cats as pets. Many other species are popular, and they are often charismatic megafauna. Many civilizations have incorporated a species of carnivoran into their culture: a prominent example is the lion, viewed as a symbol of power and royalty in many societies. Yet many species such as wolves and the big cats have been broadly hunted, resulting in extirpation in some areas. Habitat loss and human encroachment as well as climate change have been the primary cause of many species going into decline. Four species of carnivorans have gone extinct since the 1600s: Falkland Island wolf (Dusicyon australis) in 1876; the sea mink (Neogale macrodon) in 1894; the Japanese sea lion (Zalophus japonicus) in 1951 and the Caribbean monk seal (Neomonachus tropicalis) in 1952. Some species such as the red fox (Vulpes vulpes) and stoat (Mustela erminea) have been introduced to Australasia and have caused many native species to become endangered or even extinct.{{cite web|title=100 of the World's Worst Invasive Species|url=http://www.issg.org/database/species/ecology.asp?si=98&fr=1&sts=&lang=EN|publisher=Invasive Species Specialist Group}}
See also
References
{{Reflist}}
External links
{{Wikispecies|Carnivora}}
{{Wikibooks|Dichotomous Key|Carnivora}}
{{EB1911 poster|Carnivora}}
- [http://brainmaps.org/index.php?p=datasets-species#carnivore High-Resolution Images of Carnivore Brains]{{cbignore|bot=medic}}
{{Mammals}}
{{Carnivora}}
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{{Authority control}}