Jaguar
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2008-07-11T03:07:58Z
Xian englund
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{{Taxobox
| name = Jaguar<ref name=MSW3>{{MSW3 Wozencraft | pages = 546–547}}</ref>
| status = NT
| status_system = iucn3.1
| status_ref = <ref name=iucn><span class="plainlinks">{{IUCN2006|assessors=Nowell, K., Breitenmoser, U., Breitenmoser, C. & Jackson |year=2002|id=15953|title=Panthera onca |downloaded=11 August 2006}} Database entry includes justification for why this species is near threatened.</span></ref>
| trend = down
| image = Jaguar head shot.jpg
| image2 = Jaguar sitting.jpg
| image2_caption = A jaguar at the [[Milwaukee County Zoological Gardens]]
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Felidae]]
| genus = ''[[Panthera]]''
| species = '''''P. onca'''''
| binomial = ''Panthera onca''
| binomial_authority = [[Carolus Linnaeus|Linnaeus]], 1758
| range_map = Jag_distribution.gif
| range_map_caption = Jaguar range
| range_map_width = 200px
}}
The '''jaguar''' (''Panthera onca'', {{PronEng|ˡdʒægjuɚ}} in [[British English]], or {{IPA|/ˡdʒægwɑr/}} in [[American English]]) is a [[New World]] [[mammal]] of the [[Felidae]] family and one of four "[[big cat]]s" in the ''[[Panthera]]'' [[genus]], along with the [[tiger]], [[lion]], and [[leopard]] of the [[Old World]]. The jaguar is the third-largest feline after the tiger and the lion, and on average the largest and most powerful feline in the [[Western Hemisphere]]. The jaguar's present range extends from [[Mexico]] (with occasional sightings in the southwestern [[United States]]) across much of [[Central America]] and south to [[Paraguay]] and northern [[Argentina]].
This spotted cat most closely resembles the leopard physically, although it is usually larger and of sturdier build and its behavioural and [[Habitat (ecology)|habitat]] characteristics are closer to those of the tiger. While dense [[rainforest]] is its preferred habitat, the jaguar will range across a variety of forested and open terrain. It is strongly associated with the presence of water and is notable, along with the tiger, as a feline that enjoys swimming. The jaguar is a largely [[solitary]], stalk-and-ambush [[predator]], and is opportunistic in prey selection. It is also an [[apex predator|apex]] and [[keystone predator]], playing an important role in stabilizing ecosystems and regulating the populations of prey species. The jaguar has developed an exceptionally powerful bite, even relative to the other big cats.<ref name=Bite>{{cite journal | author = Stephen Wroe, Colin McHenry, and Jeffrey Thomason | title = Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa | year = 2006 | journal = Proceedings of the Royal Society B | volume = Online | format = PDF | accessdate = 2006-08-07 | url = http://www.bio.usyd.edu.au/staff/research/swroe/Wroeetal2005Biteclub.pdf}}</ref> This allows it to pierce the shells of armoured reptiles and to employ an unusual killing method: it bites directly through the skull of prey between the ears to deliver a fatal blow to the [[brain]].<ref name=HAMDIG>{{cite web | url = http://www.ecology.info/ecology-jaguar-puma.htm | first = Paul | last = Hamdig | title = Sympatric Jaguar and Puma | publisher = Ecology Online Sweden | accessdate = August 30 | accessyear = 2006}}</ref>
The jaguar is a [[near threatened]] species and its numbers are declining. Threats include habitat loss and fragmentation. While international trade in jaguars or their parts is prohibited, the cat is still regularly killed by humans, particularly in conflicts with ranchers and farmers in South America. Although reduced, its range remains large; given its historical distribution, the jaguar has featured prominently in the mythology of numerous [[indigenous peoples of the Americas|indigenous American cultures]], including that of the [[Maya civilization|Maya]] and [[Aztec]].
==Etymology==
The etymology of the word ''jaguar'' is unclear. Some sources suggest a borrowing from the [[South American]] [[Tupi language]] to English via [[Portuguese language|Portuguese]],<ref name=etymology>{{cite web | url = http://www.etymonline.com/index.php?term=jaguar | title = "Jaguar" | work = Online Etymology Dictionary | publisher = Douglas Harper | accessdate = August 12 | accessdate = 2006-08-06}}</ref> while others attribute the term to the related [[Guaraní language]]s. In the Tupi language, the original and complete indigenous name for the species is ''jaguara'', which has been reported as a denotation for any carnivorous animal<ref name=etymology/><ref name=TakeOurWord>{{cite web | url = http://www.takeourword.com/TOW198/page2.html#jaguar | title = "Word to the Wise" | work = Take our word for it, issue 198, p. 2 | publisher = [[The Institute for Etymological Research and Education]] | accessdate = August 11 | accessyear = 2006}}</ref>—in the compound form ''jaguareté'', -''eté'' means "true".<ref name=TakeOurWord/> In the related Guaraní languages, ''yaguareté'' has been variously translated as "the real fierce beast",<ref>{{cite web | url = http://www.jaguares.com.ar/afiches/index.html | title = "Yaguareté - La Verdadera Fiera" | work = RED Yaguareté | accessdate = September 27 | accessyear = 2006|language=Spanish}}</ref> "dog-bodied",<ref>{{cite web | url = http://www.indigenas.bioetica.org/inves44-2.htm | title = Breve Vocabulario | publisher = Faculty of Law, University of Buenos Aires | accessdate = September 29 | accessyear = 2006|language=Spanish}}</ref> or "fierce dog".<ref>{{cite book|title=Nativas|author=Eduardo Acevedo Díaz|date=1890|chapter=Notas| url=http://es.wikisource.org/wiki/Nativa_:_Notas| accessdate = September 29 | accessdate = 2006-09-29|language=Spanish}}</ref> Early etymological reports were that ''jaguara'' means "a beast that kills its prey with one bound", and this claim persists in a number of sources. However, this has been challenged as incorrect.<ref name=TakeOurWord/> In many Central and South American countries, the cat is referred to as ''el tigre'' ("the tiger").
The first component of its own scientific designation, ''Panthera onca'', is often presumed to derive from Greek ''pan-'' ("all") and ''ther'' ("beast"), but this may be a [[folk etymology]]. Although it came into English through the classical languages, ''panthera'' is possibly of Sanskrit origin, from ''pundarikam'', the Sanskrit word for "tiger".<ref>{{cite web | url = http://www.etymonline.com/index.php?term=panther | title = "Panther" | work = Online Etymology Dictionary | publisher = Douglas Harper | accessdate = 2006-10-26}}</ref>
''Onca'' is said to denote "barb" or "hook", a reference to the animal's powerful claws, but the most correct etymology is simply that it is an adaptation of the current Portuguese name for the animal, ''onça'' (on-sa), with the [[cedilla]] dropped for typographical reasons.
==Taxonomy==
The jaguar, ''Panthera onca'', is the only extant New World member of the ''Panthera'' genus. DNA evidence shows that the lion, tiger, leopard, jaguar, [[snow leopard]], and [[clouded leopard]] share a common ancestor and that this group is between six and ten million years old;<ref name=Johnson2006>{{cite journal | author = Johnson, W.E., Eizirik, E., Pecon-Slattery, J., Murphy, W.J., Antunes, A., Teeling, E. & O'Brien, S.J. | year = 2006 | doi = 10.1126/science.1122277 | title = The Late Miocene radiation of modern Felidae: A genetic assessment. | journal = [[Science (journal)|Science]] | volume = 311 | pages = 73–77 | pmid = 16400146}}</ref> the fossil record points to the emergence of ''Panthera'' just two to 3.8 million years ago.<ref name=Johnson2006/><ref name="Turner1987">{{ cite journal | last = Turner | first = A. | authorlink = | coauthors = | year = 1987 | month = | title = New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora) | journal = Annals of the Transvaal Museum | volume = 34 | issue = | pages = 319–347 | issn = 0041-1752 | url = | accessdate = | quote = }}</ref> [[Phylogenetic]] studies generally have shown that the clouded leopard (''Neofelis nebulosa'') is [[Basal (phylogenetics)|basal]] to this group.<ref name=Johnson2006/><ref name=Yu>{{cite journal | author = Yu L & Zhang YP | year = 2005 | title = Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores | journal = Molecular Phylogenetics and Evolution | volume = 35 | issue = 2 | pages = 483–495 | doi = 10.1016/j.ympev.2005.01.017 <!--Retrieved from CrossRef by DOI bot-->}}</ref><ref name=Johnson1997>{{cite journal | author = Johnson WE & Obrien SJ | year = 1997 | title = Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes | journal = Journal of Molecular Evolution | volume = 44 | pages = S098 | doi = 10.1007/PL00000060 <!--Retrieved from CrossRef by DOI bot-->}}</ref><ref name=Janczewski>{{cite journal | author = Dianne N. Janczewski, William S. Modi, J. Claiborne Stephens, and Stephen J. O'Brien | year = 1996 | title = Molecular Evolution of Mitochondrial 12S RNA and Cytochrome b Sequences in the Pantherine Lineage of Felidae | journal = Molecular Biology and Evolution | volume = 12 | issue = 4 | pages = 690 | url = http://mbe.oxfordjournals.org/cgi/reprint/12/4/690 | accessdate = 2006-08-06}}</ref> The position of the remaining species varies between studies and is effectively unresolved. Many studies place the snow leopard within the genus ''Panthera''<ref name=Johnson2006/><ref name=Yu/><ref name=Janczewski/> but there is no consensus whether the scientific name of the snow leopard should remain ''Uncia uncia''<ref name=iucn/><ref>Felid Taxon Advisory Group: Alan H. Shoemaker (1996) [http://www.felidtag.org/pages/Reports%5CProtocol%5Ctaxon_legal.htm ''Taxonomic and Legal Status of the Felidae'']</ref> or be moved to ''Panthera uncia''.<ref name=Johnson2006/><ref name=Yu/><ref name=Johnson1997/><ref name=Janczewski/>
Based on morphological evidence, British [[zoologist]] [[Reginald Pocock]] concluded that the jaguar is most closely related to the leopard.<ref name=Janczewski/> However, DNA evidence is inconclusive and the position of the jaguar relative to the other species varies between studies.<ref name=Johnson2006/><ref name=Yu/><ref name=Johnson1997/><ref name=Janczewski/> Fossils of extinct ''Panthera'' species, such as the [[European jaguar]] (''Panthera gombaszoegensis'') and the [[American lion]] (''Panthera atrox''), show characteristics of both the lion and the jaguar.<ref name=Janczewski/> Analysis of jaguar [[mitochondrial DNA]] has dated the species lineage to between 280,000 and 510,000 years ago, later than suggested by fossil records.<ref name=Eizirik>{{cite journal | author = Eizirik E, Kim JH, Menotti-Raymond M, Crawshaw PG Jr, O'Brien SJ, Johnson WE. | year = 2001 | title = Phylogeography, population history and conservation genetics of jaguars (Panthera onca, Mammalia, Felidae) | journal = Molecular Ecology | volume = 10 | issue = 1 | pages = 65 | url = http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=11251788&dopt=Abstract | accessdate = 2006-08-07 | doi = 10.1046/j.1365-294X.2001.01144.x <!--Retrieved from CrossRef by DOI bot-->}}</ref>
===Geographical variation===
[[Image:Jaguar tier 2.jpg|230px|thumb|right|While numerous subspecies of the jaguar have been recognized, recent research suggests just three. Geographical barriers, such as the [[Amazon river]], limit gene flow within the species.]]
The last taxonomic delineation of the jaguar subspecies was performed by Pocock in 1939. Based on geographic origins and skull [[morphology (biology)|morphology]], he recognized eight subspecies. However, he did not have access to sufficient specimens to critically evaluate all subspecies, and he expressed doubt about the status of several. Later consideration of his work suggested only three subspecies should be recognized.<ref name=Seymore>{{cite journal | author = Seymore, K.L. | year = 1989 | title = Panthera onca | journal = Mammalian Species | volume = 340 | pages = 1–9 | doi = 10.2307/3504096}}</ref>
Recent studies have also failed to find evidence for well defined subspecies, and are no longer recognized.<ref>Ronald M. Nowak (1999) Walker's Mammals of the World, 6th edition. The Johns Hopkins University Press, Baltimore, Maryland ISBN 0-8018-5789-9</ref> Larson (1997) studied the morphological variation in the jaguar and showed that there is [[Cline (population genetics)|clinal]] north–south variation, but also that the differentiation within the supposed subspecies is larger than that between them and thus does not warrant subspecies subdivision.<ref name=Larson>{{cite journal | author = Larson, Shawn E. | year = 1997 | title = Taxonomic re-evaluation of the jaguar | journal = Zoo Biology | volume = 16 | issue = 2 | pages = 107 | url = http://www3.interscience.wiley.com/cgi-bin/abstract/51449/ABSTRACT?CRETRY=1&SRETRY=0 | accessdate = 2006-08-07 | doi = 10.1002/(SICI)1098-2361(1997)16:2<107::AID-ZOO2>3.0.CO;2-E}}</ref> A genetic study by Eizirik and coworkers in 2001 confirmed the absence of a clear geographical subspecies structure, although they found that major geographical barriers such as the [[Amazon River]] limited the exchange of genes between the different populations.<ref name=Eizirik/> A subsequent, more detailed, study confirmed the predicted population structure within the [[Colombia]]n jaguars.<ref name = Columbia>{{cite journal | author = Ruiz-Garcia M, Payan E, Murillo A & Alvarez D | year = 2006 | title = DNA microsatellite characterization of the jaguar (''Panthera onca'') in Colombia | journal = Genes & Genetic Systems | volume = 81 | issue = 2 | pages = 115–127 | doi = 10.1266/ggs.81.115 <!--Retrieved from CrossRef by DOI bot-->}}</ref>
Pocock's subspecies divisions are still regularly listed in general descriptions of the cat.<ref name=MANUALTAXONOMY>"[http://www.jaguarssp.org/Animal%20Mgmt/JAGUAR%20HUSBANDRY%20MANUAL.pdf ''Guidelines for Captive Management of Jaguars''], Taxonomy, pp. 5–7, [[Jaguar Species Survival Plan]]</ref> Seymore grouped these in three subspecies.<ref name=Seymore/>
# ''Panthera onca onca'': Venezuela, south and east to Rio Grande do Sul in Brazil, including
#* ''P. onca peruviana'': Coastal Peru – [[Peruvian Jaguar]]
# ''P. onca hernandesii'': Western Mexico, including – [[Mexican Jaguar]]
#* ''P. onca centralis'': Central America—El Salvador to Colombia – [[Central American Jaguar]]
#* ''P. onca arizonensis'': Eastern Arizona to Sonora, Mexico – [[Arizonan Jaguar]]
#* ''P. onca veraecrucis'': Southeastern Mexico to central Texas
#* ''P. onca goldmani'': [[Yucatan Peninsula]] to Belize and Guatemala – [[Goldman's Jaguar]]
# ''P. onca palustris'' or ''P. onca paraguensis'': Paraguay and northeastern Argentina (The description of the first is based on a fossil, the latter on living individuals)<ref name=Seymore/>
The canonical ''Mammal Species of the World'' continues to recognize nine subspecies, the eight subspecies above and additionally ''P. o. paraguensis''.<ref name=MSW3/>
==Biology and behavior==
===Physical characteristics===
[[Image:Jaguar at Edinburgh Zoo.jpg|right|thumb|220px|The head of the jaguar is robust and the jaw extremely powerful. Size tends to increase from the north to the south.]]
The jaguar is a compact and well-muscled animal. There are significant variations in size: weights are normally in the range of 56–96 kilograms (124–211 [[pound (mass)|lb]]). Larger males have been recorded as weighing 159 kilograms (350 lb)<ref name="Animal">{{cite book |last=Kindersley |first= Dorling |year=2001,2005 |title=Animal |location=New York City |publisher=DK Publishing |isbn=0-7894-7764-5}}</ref> (roughly matching a tigress or lioness), and smaller ones have extremely low weights of 36 kilograms (80 lb). Females are typically 10–20% smaller than males. The length of the cat varies from 1.62–1.83 meters (5.3–6 ft), and its tail may add a further 75 centimeters (30 in). It stands about 67–76 centimeters (27–30 in) tall at the shoulders.<ref name=WCS>{{cite web | url = http://savethejaguar.com/jag-index/jag-allabout/jag-aboutecology | title = "All about Jaguars: ECOLOGY" | publisher = [[Wildlife Conservation Society]]| accessdate = August 11 | accessyear = 2006 }}</ref>
Further variations in size have been observed across regions and habitats, with size tending to increase from the north to south. A study of the jaguar in the [[Chamela-Cuixmala Biosphere Reserve]] on the Mexican Pacific coast, showed ranges of just 30–50 kilograms (66–110 lb), about the size of the [[cougar]].<ref name = foodhabits>{{cite journal | author = Rodrigo Nuanaez, Brian Miller, and Fred Lindzey | year = 2000 | title = Food habits of jaguars and pumas in Jalisco, Mexico | journal = Journal of Zoology | volume = 252 | issue = 3 | pages = 373 | url = http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=58851 | accessdate = 2006-08-08}}</ref> By contrast, a study of the Jaguar in the [[Brazil]]ian [[Pantanal]] region found average weights of 100 kilograms (220 lb).<ref name=JSSP>{{cite web | url = http://jaguarssp.org/pdf/FACT%20SHEET%20with%20graphics.pdf | title = "Jaguar Fact Sheet" | publisher = [[Jaguar Species Survival Plan]]| accessdate = August 14 | accessyear = 2006}}</ref> Forest jaguars are frequently darker and considerably smaller than those found in open areas (the Pantanal is an open wetland basin), possibly due to the fewer large herbivorous prey in forest areas.<ref name=CAP> Nowell, K. and Jackson, P. (compilers and editors) 1996. [http://carnivoractionplans1.free.fr/wildcats.pdf ''Wild Cats. Status Survey and Conservation Action Plan''] (PDF). IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland. (see ''Panthera Onca,'' pp 118–122)</ref>
A short and stocky limb structure makes the jaguar adept at climbing, crawling and swimming.<ref name=WCS/> The head is robust and the jaw extremely powerful. It has been suggested that the jaguar has the strongest bite of all felids, and the second strongest of all mammals; this strength is an adaptation that allows the jaguar to pierce turtle shells.<ref name=HAMDIG/> A comparative study of bite force adjusted for body size ranked it as the top felid, alongside the clouded leopard and ahead of the lion and tiger.<ref>{{cite web | url = http://www.aptv.org/Schedule/showinfo.asp?ID=76679&Nola1=NAGS | title = "Search for the Jaguar" | work = National Geographic Specials | publisher = [[Alabama Public Television]] | accessdate = August 11 | accessyear = 2006 }}</ref> It has been reported that "an individual jaguar can drag a 360 kg (800 lb) bull 8 m (25 ft) in its jaws and pulverize the heaviest bones".<ref>{{cite web | last = McGrath | first = Susan | url = http://magazine.audubon.org/features0408/belize.html | title = "Top Cat" | publisher = [[National Audubon Society]] | date=August 2004 | accessdate = August 11 | accessyear = 2006 }}</ref> The jaguar hunts wild animals weighing up to 300 kilograms (660 lb) in dense jungle, and its short and sturdy physique is thus an adaptation to its prey and environment.
[[Image:Jaguar.jpg|thumb|left|220px|A melanistic jaguar at the [[Henry Doorly Zoo]]. Melanism is result of a dominant [[allele]] and remains relatively rare in jaguars.]]
The base coat of the jaguar is generally a tawny yellow, but can range to reddish-brown and black. The cat is covered in [[Rosette (zoology)|rosettes]] for camouflage in its jungle habitat. The spots vary over individual coats and between individual Jaguars: rosettes may include one or several dots, and the shape of the dots varies. The spots on the head and neck are generally solid, as are those on the tail, where they may merge to form a band. The underbelly, throat and outer surface of the legs and lower flanks are white.<ref name=WCS/>
A condition known as [[melanism]] occurs in the species. The melanistic form is less common than the spotted form (it occurs at about six percent of the population)<ref>{{cite web | first = Vladmir | last = Dinets | url = http://dinets.travel.ru/blackjaguar.htm | title = First documentation of melanism in the jaguar (Panthera onca) from northern Mexico | accessdate = September 29 | accessyear = 2006}}</ref> of jaguars in their South American range have been reported to possess it—and is the result of a dominant [[allele]].<ref name=Meyer>Meyer, John R. "[http://biological-diversity.info/Black_Jaguar.htm Black jaguars in Belize?: A survey of melanism in the jaguar, Panthera onca]", biological-diversity.info, Belize Explorer Group, 1994.</ref> Jaguars with melanism appear entirely black, although their spots are still visible on close examination. Melanistic Jaguars are informally known as [[black panther]]s, but do not form a separate species. Rare [[albino]] individuals, sometimes called [[white panther]]s, occur among jaguars, as with the other big cats.<ref name=CAP/>
The jaguar closely resembles the leopard, but is sturdier and heavier, and the two animals can be distinguished by their rosettes: the rosettes on a jaguar's coat are larger, fewer in number, usually darker, and have thicker lines and small spots in the middle that the leopard lacks. Jaguars also have rounder heads and shorter, stockier limbs compared to leopards.<ref name=akron>{{cite web | url = http://www.akronzoo.org/learn/jaguar.asp | title = "Jaguar (panthera onca)" | work = Our animals | publisher = [[Akron Zoo]] | accessdate = August 11 | accessyear = 2006 }}</ref>
===Reproduction and life cycle===
Jaguar females reach sexual maturity at about two years of age, and males at three or four.<!--the difference arises from competition for mates among males.''hide until sourced''--> The cat is believed to mate throughout the year in the wild, although births may increase when prey is plentiful.<ref name=MANUALREPRODUCTION>"Guidelines", Reproduction, pp. 28–38</ref> Research on captive male jaguars supports the year-round mating hypothesis, with no seasonal variation in semen traits and ejaculatory quality; low reproductive success has also been observed in captivity.<ref>{{cite journal | author = Ronaldo Gonçalves Morato, Marcelo Alcindo Barros de Vaz Guimaraes, Fernando Ferriera, Ieda Terezinha do Nascimento Verreschi, Renato Campanarut Barnabe | year = 1999 | title = Reproductive characteristics of captive male jaguars | journal = Brazilian Journal of Veterinary Research and Animal Science | volume = 36 | issue = 5 | url = http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1413-95961999000500008&lng=pt&nrm=iso&tlng=en | accessdate = 2006-08-08}}</ref> Female [[estrous]] is 6–17 days out of a full 37-day cycle, and females will advertise fertility with urinary scent marks and increased vocalization.<ref name=MANUALREPRODUCTION/> Both sexes will range more widely than usual during courtship.
Mating pairs separate after the act, and females provide all parenting. The gestation period lasts 93–105 days; females give birth to up to four cubs, and most commonly to two. The mother will not tolerate the presence of males after the birth of cubs, given a risk of infant [[cannibalism]]; this behaviour is also found in the tiger.<ref name=MANUALHIST&BEHAV>"Guidelines", Natural History & Behavior, pp. 8–16</ref>
The young are born blind, gaining sight after two weeks. Cubs are weaned at three months but remain in the birth den for six months before leaving to accompany their mother on hunts.<ref name=SWWL>{{cite web | url = http://web.archive.org/web/www.southwestwildlife.org/nwsletter/Spring06Newsletter.pdf | title = "Jaguars: Magnificence in the Southwest" | work = Newsletter, Spring 2006 | publisher = [[Southwest Wildlife Rehabilitation & Educational Foundation]] | accessdate = August 14 | accessyear = 2006}}</ref> They will continue in their mother's company for one to two years before leaving to establish a territory for themselves. Young males are at first nomadic, jostling with their older counterparts until they succeed in claiming a territory. Typical lifespan in the wild is estimated at around 12–15 years; in captivity, the jaguar lives up to 23 years, placing it among the longest-lived cats.<ref name=JSSP/>
===Social activity===
Like most cats, the jaguar is solitary outside mother-cub groups. Adults generally meet only to court and mate (though limited non-courting socialization has been observed anecdotally<ref name=MANUALHIST&BEHAV/>) and carve out large territories for themselves. Female territories, from 25 to 40 square kilometers in size, may overlap, but the animals generally avoid one another. Male ranges cover roughly twice as much area, varying in size with the availability of game and space, and do not overlap.<ref name=MANUALHIST&BEHAV/><ref>{{cite journal | author = George B. Schaller, Peter Gransden Crawshaw, Jr. | year = 1980 | title = Movement Patterns of Jaguar | journal = Biotropica | volume = 12 | issue = 3 | pages = 161 | url = http://links.jstor.org/sici?sici=0006-3606(198009)12%3A3%3C161%3AMPOJ%3E2.0.CO%3B2-H/ | accessdate = 2006-08-08 | doi = 10.2307/2387967}}</ref> Scrape marks, urine, and feces are used to mark territory.<ref name=Rabinowitz>{{cite journal | author = Rabinowitz, AR., Nottingham, BG Jr | year = 1986 | title = Ecology and behaviour of the Jaguar (Panthera onca) in Belize, Central America | journal = Journal of Zoology | volume = 210 | issue = 1 | pages = 149 | url = http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=1524401&q=jaguar+&uid=788304424&setcookie=yes | accessdate = 2006-08-11}} ''Overlapping male ranges are observed in this study in Belize. Note the overall size of ranges is about half of normal.''</ref>
Like the other big cats, the jaguar is capable of roaring (the male more powerfully) and does so to warn territorial and mating competitors away; intensive bouts of counter-calling between individuals have been observed in the wild.<ref name=Emmons87>{{cite journal | author = Emmons, Louise H. | year = 1987 | title = Comparative feeding ecology of felids in a neotropical rainforest | journal = Behavioral Ecology and Sociobiology | volume = 20 | issue = 4 | pages = 271 | url = http://www.springerlink.com/content/h715034593114546/ | accessdate = 2006-08-08 | doi = 10.1007/BF00292180 <!--Retrieved from CrossRef by DOI bot-->}}</ref> Their roar often resembles a repetitive cough, and they may also vocalize mews and grunts.<ref name=JSSP/> Mating fights between males occur, but are rare, and aggression avoidance behaviour has been observed in the wild.<ref name=Rabinowitz/> When it occurs, conflict is typically over territory: a male's range may encompass that of two or three females, and he will not tolerate intrusions by other adult males.<ref name=MANUALHIST&BEHAV/>
The jaguar is often described as [[nocturnal]], but is more specifically [[crepuscular]] (peak activity around dawn and dusk). Both sexes hunt, but males travel farther each day than females, befitting their larger territories. The jaguar may hunt during the day if game is available and is a relatively energetic feline, spending as much as 50–60% of its time active.<ref name=CAP/> The jaguar's elusive nature and the inaccessibility of much of its preferred habitat make it a difficult animal to sight, let alone study.
===Hunting and diet===
Like all cats, the jaguar is an obligate [[carnivore]], feeding only on meat. It is an opportunistic hunter and its diet encompasses 87 species.<ref name=CAP/> The jaguar prefers large prey and will take [[deer]], [[tapir]]s, [[peccary|peccaries]], [[dog]]s, [[fox]]es, and sometimes even [[anaconda]]s and [[caiman]]. However, the cat will eat any small species that can be caught, including [[frog]]s, [[mice]], [[birds]], [[fish]], [[sloth]]s, [[monkey]]s, [[turtle]]s, [[capybara]], and domestic livestock.<ref>{{cite web | url = http://www.kidsplanet.org/factsheets/jaguar.html | title = Jaguar -- Kids' Planet -- Defenders of Wildlife | accessdate = 2006-09-23 | author = Defenders of Wildlife}}</ref>
[[Image:Panthera onca at the Toronto Zoo 2.jpg|thumb|left|220px|The jaguar has developed an exceptionally powerful bite, even relative to the other big cats. It is an adaptation that allows it to pierce the shells of armoured reptiles.]]
While the jaguar employs the deep-throat bite-and-suffocation technique typical among ''Panthera'', it prefers a killing method unique amongst cats: it pierces directly through the [[temporal bone]]s of the skull between the ears of prey (especially the [[Capybara]]) with its canine teeth, piercing the brain. This may be an adaptation to "cracking open" turtle shells; following the late Pleistocene extinctions, armoured reptiles such as turtles would have formed an abundant prey base for the jaguar.<ref name=Emmons87/><ref name=CAP/> The skull bite is employed with mammals in particular; with reptiles such as caiman, the jaguar may leap on to the back of the prey and sever the [[cervical vertebrae]], immobilizing the target. While capable of cracking turtle shells, the jaguar may simply reach into the shell and scoop out the flesh.<ref name=MANUALHIST&BEHAV/> With prey such as dogs, a paw swipe to crush the skull may be sufficient.
The jaguar is a stalk-and-ambush rather than a chase predator. The cat will walk slowly down forest paths, listening for and stalking prey before rushing or ambushing. The jaguar attacks from cover and usually from a target's blind spot with a quick pounce; the species' ambushing abilities are considered nearly peerless in the animal kingdom by both indigenous people and field researchers, and are probably a product of its role as an apex predator in several different environments. The ambush may include leaping into water after prey, as a jaguar is quite capable of carrying a large kill while swimming; its strength is such that carcasses as large as a heifer can be hauled up a tree to avoid flood levels.<ref name=MANUALHIST&BEHAV/>
On killing prey, the jaguar will drag the carcass to a thicket or other secluded spot. It begins eating at the neck and chest, rather than the midsection. The heart and lungs are consumed, followed by the shoulders.<ref name=MANUALHIST&BEHAV/> The daily food requirement of a 34 kilogram animal, at the extreme low end of the species' weight range, has been estimated at 1.4 kilograms.<ref name=FEDERAL>{{cite web | url = http://www.epa.gov/fedrgstr/EPA-SPECIES/2006/July/Day-12/e10644.htm | title = Determination That Designation of Critical Habitat Is Not Prudent for the Jaguar | access = 2006-08-30 | date = 2006-07-12 | work = Federal Register Environmental Documents}}</ref> For captive animals in the 50–60 kilogram range, more than 2 kilograms of meat daily is recommended.<ref>"Guidelines", Hand-rearing, pp. 62–75 (see table 5)</ref> In the wild, consumption is naturally more erratic; wild cats expend considerable energy in the capture and kill of prey, and may consume up to 25 kilograms of meat at one feeding, followed by periods of famine.<ref name=MANUALNUTRITION>"Guidelines", Nutrition, pp. 55–61</ref>
==Ecology==
===Distribution and habitat===
The jaguar has been attested in the fossil record for two million years<ref name=MANUALTAXONOMY/> and it has been an American cat since crossing the [[Bering Land Bridge]] during the [[Pleistocene]]; the immediate ancestor of modern animals is ''Panthera onca augusta'', which was larger than the contemporary cat.<ref name = Columbia/> Its present range extends from Mexico, through Central America and into South America, including much of Amazonian Brazil.<ref name=CONSERVATION>{{cite journal | author = Eric W. Sanderson, Kent H. Redford, Cheryl-Lesley B. Chetkiewicz, Rodrigo A. Medellin, Alan R. Rabinowitz, John G. Robinson, and Andrew B. Taber | year = 2002 | title = Planning to Save a Species: the Jaguar as a Model | journal = Conservation Biology | volume = 16 | issue = 1 | pages = 58 | doi = 10.1046/j.1523-1739.2002.00352.x}} ''Detailed analysis of present range and terrain types provided here.''</ref> The countries included in this range are [[Argentina]], [[Belize]], [[Bolivia]], [[Brazil]], [[Colombia]], [[Costa Rica]] (particularly on the [[Osa Peninsula]]), [[Ecuador]], [[French Guiana]], [[Guatemala]], [[Guyana]], [[Honduras]], [[Mexico]], [[Nicaragua]], [[Panama]], [[Paraguay]], [[Peru]], [[Suriname]] and [[Venezuela]]. The jaguar is now extinct in El Salvador and [[Uruguay]].<ref name="iucn"/> It occurs in the 400 km² [[Cockscomb Basin Wildlife Sanctuary]] in Belize, the 5,300 km² [[Sian Ka'an]] [[Biosphere]] Reserve in Mexico, the approximately 15,000 km² [[Manú National Park]] in Peru, the approximately 26,000 km² [[Xingu National Park]] in Brazil, and numerous other reserves throughout its range.
The inclusion of the United States in the list is based on occasional sightings in the southwest, particularly in [[Arizona]], [[New Mexico]] and [[Texas]]. In the early 1900s, the jaguar's range extended as far north as Southern [[California]] and western Texas.<ref name=FEDERAL/> The jaguar is a protected species in the United States under the [[Endangered Species Act]], which has stopped the shooting of the animal for its pelt. In 2004, wildlife officials in Arizona photographed and documented jaguars in the southern part of the state. For any permanent population to thrive, protection from killing, an adequate prey base, and connectivity with Mexican populations are essential.<ref>{{cite web | url = http://www.gf.state.az.us/w_c/es/jaguar_management.shtml | title = Jaguar Management | publisher = Arizona Game & Fish, | accessdate = 2006-08-08}}</ref>
[[Image:Standing jaguar.jpg|220px|thumb|right|The jaguar can range across a variety of forested and open habitat, but is strongly associated with presence of water.]]
The historic range of the species included much of the southern half of the United States, and in the south extended much farther to cover most of the South American continent. In total, its northern range has receded 1000 kilometers southward and its southern range 2000 km northward. Ice Age fossils of the jaguar, dated between 40,000 and 11,500 years ago, have been discovered in the United States, including some at an important site as far north as [[Missouri]]. Fossil evidence shows jaguars of up to 190 kg (420 lb), much larger than the contemporary average for the animal.<ref name=MSI>{{cite web | url = http://www.museum.state.il.us/exhibits/larson/felis_onca.html | title = "Jaguars" | work = The Midwestern United States 16 000 years ago | publisher = [[Illinois State Museum]] | accessdate = August 20 | accessyear = 2006}}</ref>
The habitat of the cat includes the rain forests of [[South America|South]] and [[Central America]], open, seasonally flooded wetlands, and dry grassland terrain. Of these habitats, the jaguar much prefers dense forest;<ref name=CAP/> the cat has lost range most rapidly in regions of drier habitat, such as the Argentinian [[pampas]], the arid grasslands of Mexico, and the southwestern United States.<ref name="iucn"/> The cat will range across tropical, subtropical, and dry deciduous forests (including, historically, oak forests in the United States). The jaguar is strongly associated with water and it often prefers to live by rivers, swamps, and in dense rainforest with thick cover for stalking prey. Jaguars have been found at elevations as high as 3800 m, but they typically avoid mountain forest and are not found in the high plateau of central Mexico or in the Andes.<ref name=CAP/>
===Ecological role===
The adult jaguar is an apex predator, meaning that it exists at the top of its food chain and is not preyed on in the wild. The jaguar has also been termed a keystone species, as it is assumed, through controlling the population levels of prey such as herbivorous and granivorous mammals, apex felids maintain the structural integrity of forest systems.<ref name=foodhabits/><ref name=PHEONIX>{{cite web | url = http://www.phoenixzoo.org/learn/animals/animal_detail.aspx?FACT_SHEET_ID=100018 | title = Jaguar (Panthera Onca) | publisher = [[Pheonix Zoo]] | accessdate = August 30 | accessyear = 2006}}</ref> However, accurately determining what effect species like the jaguar have on ecosystems is difficult, because data must be compared from regions where the species is absent as well as its current habitats, while controlling for the effects of human activity. It is accepted that mid-sized prey species see population increases in the absence of the keystone predators and it has been hypothesized that this has cascading negative effects,<ref name=MONGA>{{cite web | url = http://rainforests.mongabay.com/02keystone.htm | title = Structure and Character: Keystone Species | work = mongabay.com | publisher = Rhett Butler | accessdate = August 30 | accessyear = 2006}}</ref> however, field work has shown this may be natural variability and that the population increases may not be sustained. Thus, the keystone predator hypothesis is not favoured by all scientists.<ref>{{cite journal | author = Wright, SJ; Gompper, ME; DeLeon, B | year = 1994 | title = Are large predators keystone species in Neotropical forests? The evidence from Barro Colorado Island | journal = Oikos | volume = 71 | issue = 2 | pages = 279 | url = http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=3657385&q=&uid=788032445&setcookie=yes | accessdate = 2006-08-08 | doi = 10.2307/3546277 <!--Retrieved from CrossRef by DOI bot-->}}</ref>
The jaguar also has an effect on other predators. The jaguar and the [[cougar]], the next largest feline of the Americas, are often [[sympatric]] (related species sharing overlapping territory) and have often been studied in conjunction. Where sympatric with the jaguar, the cougar is smaller than normal and is smaller than the local jaguars. The jaguar tends to take larger prey and the cougar smaller, reducing the latter's size.<ref>{{cite journal | author = J. Agustin Iriarte, William L. Franklin, Warren E. Johnson, and Kent H. Redford | year = 1990 | title = Biogeographic variation of food habits and body size of the America puma | journal = [[Oecologia]] | volume = 85 | issue = 2 | pages = 185 | url = http://www.springerlink.com/content/nvk62r701822qq17/ | accessdate = 2006-08-09 | doi = 10.1007/BF00319400}}</ref> This situation may be advantageous to the cougar. Its broader prey niche, including its ability to take smaller prey, may give it an advantage over the jaguar in human-altered landscapes;<ref name=foodhabits/> while both are classified as [[near threatened|near-threatened]] species, the cougar has a significantly larger current distribution.
==Conservation status==
Jaguar populations are currently declining. The animal is considered [[Near Threatened]] by the [[International Union for Conservation of Nature and Natural Resources]],<ref name=iucn/> meaning it may be threatened with extinction in the near future. The loss of parts of its range, including its virtual elimination from its historic northern areas and the increasing fragmentation of the remaining range, have contributed to this status. The 1960s saw particularly significant declines, with more than 15,000 jaguar skins brought out of the [[Brazilian Amazon]] yearly; the [[Convention on International Trade in Endangered Species]] (CITES) of 1973 brought about a sharp decline in the pelt trade.<ref>{{cite journal |last=Weber |first=William |coauthors=Rabinowitz, Alan |year=1996 |month=August |title=A Global Perspective on Large Carnivore Conservation |journal= Conservation Biology |volume=10 |issue=4 |pages=1046–1054 | url= http://www.blackwell-synergy.com/doi/abs/10.1046/j.1523-1739.1996.10041046.x?journalCode=cbi | doi = 10.1046/j.1523-1739.1996.10041046.x <!--Retrieved from URL by DOI bot-->}}</ref> Detailed work performed under the auspices of the [[Wildlife Conservation Society]] reveal that the animal has lost 37% of its historic range, with its status unknown in an additional 18%. More encouragingly, the probability of long-term survival was considered high in 70% of its remaining range, particularly in the Amazon basin and the adjoining [[Gran Chaco]] and [[Pantanal]].<ref name=CONSERVATION/>
The major risks to the jaguar include [[deforestation]] across its habitat, increasing competition for food with human beings,<ref name=iucn/> [[poaching]], [[hurricanes]] in Northern parts of its range, and the behaviour of ranchers who will often kill the cat where it preys on livestock. When adapted to the prey, the jaguars has been shown to take cattle as a large portion of its diet; while land clearance for grazing is a problem for the species, the jaguar population may have increased when cattle were first introduced to South America as the animals took advantage of the new prey base. This willingness to take livestock has induced ranch owners to hire full-time jaguar hunters, and the cat is often shot on sight.<ref name=JSSP/>
[[Image:Pantanal.jpg|thumb|right|220px|The [[Pantanal]], [[Brazil]], seen here in flood condition, is a critical jaguar range area.]]
The jaguar is regulated as an Appendix I species under CITES: all international trade in jaguars or their parts is prohibited. All hunting of jaguars is prohibited in [[Argentina]], [[Belize]], [[Colombia]], [[French Guiana]], [[Honduras]], [[Nicaragua]], [[Panama]], [[Paraguay]], [[Suriname]], the [[United States]] (where it is listed as endangered under the [[Endangered Species Act]]), [[Uruguay]] and [[Venezuela]]. Hunting of jaguars is restricted to "problem animals" in [[Brazil]], [[Costa Rica]], [[Guatemala]], [[Mexico]] and [[Peru]], while trophy hunting is still permitted in [[Bolivia]]. The species has no legal protection in [[Ecuador]] or [[Guyana]].<ref name=MANUALTAXONOMY/>
Current conservation efforts often focus on educating ranch owners and promoting [[ecotourism]].<ref name=WWF>{{cite web | url = http://www.panda.org/about_wwf/where_we_work/latin_america_and_caribbean/country/venezuela/index.cfm?uProjectID=VE0854 | title = Jaguar Refuge in the Llanos Ecoregion | publisher = [[World Wildlife Fund]] | accessdate = September 1 | accessyear = 2006}}</ref> The jaguar is generally defined as an "umbrella species"—a species whose home range and habitat requirements are sufficiently broad that, if protected, numerous other species of smaller range will also be protected.<ref>{{cite web | url = http://www.co.pima.az.us/cmo/sdcp/kids/gloss.html | title = Glossary | work = Sonoran Desert Conservation Plan: Kids | publisher = [[Pima County Government]] | accessdate = September 1 | accessyear = 2006}}</ref> Umbrella species serve as "mobile links" at the landscape scale, in the jaguar's case through predation. Conservation organizations may thus focus on providing viable, connected habitat for the jaguar, with the knowledge that other species will also benefit.<ref name=WWF/>
Given the inaccessibility of much of the species' range—particularly the central Amazon—estimating jaguar numbers is difficult. Researchers typically focus on particular bioregions, and thus species-wide analysis is scant. In 1991, 600–1,000 (the highest total) were estimated to be living in [[Belize]]. A year earlier, 125–180 jaguars were estimated to be living in Mexico's 4,000 square kilometer (2400 mi²) [[Calakmul Biosphere Reserve]], with another 350 in the state of [[Chiapas]]. The adjoining [[Maya Biosphere Reserve]] in Guatemala, with an area measuring 15,000 square kilometers (9,000 mi²), may have 465–550 animals.<ref name=GUIDELINESPOPULATION>"Guidelines", Protection and Population Status, p. 4.</ref> Work employing [[Global Positioning System|GPS]]-[[telemetry]] in 2003 and 2004 found densities of only six to seven jaguars per 100 square kilometers in the critical Pantanal region, compared with 10 to 11 using traditional methods; this suggests that widely used sampling methods may inflate the actual numbers of cats.<ref>{{cite journal | author = Marianne K. Soisalo, Sandra M.C. Cavalcanti. | year = 2006 | title = Estimating the density of a jaguar population in the Brazilian Pantanal using camera-traps and capture–recapture sampling in combination with GPS radio-telemetry | journal = Biological Conservation | volume = 129 | pages = 487 | url = http://www.procarnivoros.org.br/pdfs/Soisalo_e_Cavalcanti_2006.pdf#search=%22jaguar%20population%20numbers%22 | accessdate = 2006-08-08 | doi = 10.1016/j.biocon.2005.11.023}}</ref>
On [[7 January]], 2008 [[United States Fish and Wildlife Service]] Director [[H. Dale Hall]] approved an unprecedented decision by the [[George W. Bush Administration]] to abandon jaguar recovery as a federal goal under the Endangered Species Act. The decision is the first of its kind in the 34-year history of the Endangered Species Act. Some critics of the decision said that the jaguar is being sacrificed for the government's new border fence, which is to be built along many of the cat's typical crossings between the United States and Mexico.<ref name=sfchron>[http://www.sfgate.com/cgi-bin/article.cgi?f=/n/a/2008/01/17/national/w154058S41.DTL&type=health US Abandons Bid for Jaguar Recovery Plan]</ref>
==In mythology and culture==
===Mesoamerican culture===
:''Main article: [[Jaguars in Mesoamerican culture]]'' <ref>See [[Jaguars in Mesoamerican culture#References|main article references]] for bibliography on this section</ref>
[[Image:Jaguar warrior.jpg|thumb|150px|right|Jaguar warrior in Mayan culture]]
[[Image:MocheJaguarLarcoMuseum.jpg|thumb|right|150px| Moche Jaguar. 300 A.D. [[Larco Museum]] Lima, Peru]]
In Central and South America, the jaguar has long been a symbol of power and strength. The Chavín cult of the jaguar became accepted over most of what is today Peru by 900 BC. Concurrent with Chavin, the [[Olmec]], the progenitor culture of [[Pre-Columbian]] [[Mesoamerica]], developed a distinct "were-jaguar" motif of sculptures and figurines showing stylized jaguars or humans with jaguar characteristics. The [[Moche]] culture of Northern Peru used the jaguar as a symbol of power in many of their ceramics.<ref>Berrin, Katherine & Larco Museum. ''The Spirit of Ancient Peru:Treasures from the [[Larco Museum|Museo Arqueológico Rafael Larco Herrera]].'' New York: [[Thames and Hudson]], 1997.</ref> In the later [[Maya civilization]], the jaguar was believed to facilitate communication between the living and the dead and to protect the royal household. The Maya saw these powerful felines as their companions in the spiritual world, and kings were typically given a royal name incorporating the word jaguar. The Aztec civilization shared this image of the jaguar as the representative of the ruler and as a warrior. The [[Aztec]]s formed an elite warrior class known as the [[Jaguar warrior|Jaguar Knights]]. In [[Aztec mythology]], the jaguar was considered to be the [[totem]] animal of the powerful deity [[Tezcatlipoca]].
===Contemporary culture===
The jaguar is widely used as a symbol in contemporary culture. It is the national animal of [[Guyana]], and is featured in its [[coat of arms]].<ref>[http://www.rbcradio.com/knowguyana.html Guyana], RBC Radio</ref> It is widely used as a product name, most prominently for a [[Jaguar (car)|luxury car brand]]. The name has been adopted by sports franchises, including [[NFL]]'s [[Jacksonville Jaguars]] and the Mexican national soccer league team the [[Jaguares de Chiapas]]. The crest of [[Argentina|Argentina's]] [[Argentine Rugby Union|national federation]] in [[rugby union]] features a jaguar; however, because of a historic accident, the country's [[Argentina national rugby union team|national team]] is nicknamed ''Los Pumas''.
==Notes and references==
{{reflist|2}}
==External links==
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{{commons| Panthera onca}}
{{wikispecies| Panthera onca}}
* [http://www.biocrawler.com/videos/displayimage.php?album=topn&cat=0&pos=2 Short narrated video about the jaguar]
*[http://www.nature.org/animals/mammals/animals/jaguar.html The Nature Conservancy's Species Profile: Jaguar] – Learn more about the Jaguar
* [http://www.maniacworld.com/Jaguar-vs-Anaconda.html Jaguar preys on Anaconda]
*[http://www.arkive.org/species/GES/mammals/Panthera_onca/ Images of the jaguar ''(Panthera onca)''] from “ARKive images of life on Earth”
* The Jaguar in Argentina [http://www.redyaguarete.com.ar/ Red Yaguareté Argentina]
* [http://www.cordobes.com/30dias/noticia106.html History about the last Jaguars in Misiones Jungle, (Argentina) - Free video]
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[[Category:Mammals of South America]]
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