Jaguar

The jaguar (Panthera onca) is a large felid species and the only living member of the genus Panthera native to the Americas. Its distinctively marked coat features pale yellow to tan colored fur covered by spots that transition to rosettes on the sides. With a body length of up to , it is the largest cat species in the Americas and the third largest in the world. Its powerful bite allows it to pierce the carapaces of turtles and tortoises, and to employ an unusual killing method: it bites directly through the skull of mammalian prey between the ears to deliver a fatal blow to the brain.
The jaguar most likely entered the Americas from Eurasia during the Early Pleistocene via the land bridge that once spanned the Bering Strait. Jaguar fossils excavated in the Americas date back to 130,000 years BP. Today, the jaguar's range extends from extreme southern Arizona in the United States across Mexico and much of Central America, the Amazon rainforest and south to Paraguay and northern Argentina. It inhabits a variety of forested and open terrains, but its preferred habitat is tropical and subtropical moist broadleaf forest, wetlands and wooded regions. It is adept at swimming and is largely a solitary, opportunistic, stalk-and-ambush apex predator. As a keystone species, it plays an important role in stabilizing ecosystems and regulating prey populations.
The jaguar is threatened by habitat loss, habitat fragmentation, poaching for trade with its body parts and killings in human–wildlife conflict situations, particularly with ranchers in Central and South America. It has been listed as Near Threatened on the IUCN Red List since 2002. The wild population is thought to have declined since the late 1990s. Priority areas for jaguar conservation comprise 51 Jaguar Conservation Units (JCUs), defined as large areas inhabited by at least 50 breeding jaguars. The JCUs are located in 36 geographic regions ranging from Mexico to Argentina.
The jaguar has featured prominently in the mythology of numerous indigenous peoples of the Americas, including those of the Maya and Aztec civilizations.
==Etymology==
The word jaguar is derived from , a word in one of the indigenous languages of Brazil for a wild spotted cat that is larger than a wolf. Onca is derived from the Portuguese name for a spotted cat in Brazil that is larger than a lynx.
Indigenous peoples in Guyana call it .
The word panther is derived from classical Latin , itself from the ancient Greek ().
== Classification ==
===Taxonomy===
In 1758, Carl Linnaeus described the jaguar in his work Systema Naturae and gave it the scientific name Felis onca.
In the 19th and 20th centuries, several jaguar type specimens formed the basis for descriptions of subspecies.
In 1939, Reginald Innes Pocock recognized eight subspecies based on the geographic origins and skull morphology of these specimens.
Pocock did not have access to sufficient zoological specimens to critically evaluate their subspecific status, but expressed doubt about the status of several. Later consideration of his work suggested only three subspecies should be recognized. The description of P. o. palustris was based on a fossil skull.
By 2005, nine subspecies were considered to be valid taxa.
Reginald Innes Pocock placed the jaguar in the genus Panthera and observed that it shares several morphological features with the leopard (P. pardus). He, therefore, concluded that they are most closely related to each other. Results of morphological and genetic research indicate a clinal north–south variation between populations, but no evidence for subspecific differentiation.
DNA analysis of 84 jaguar samples from South America revealed that the gene flow between jaguar populations in Colombia was high in the past.
Since 2017, the jaguar is considered to be a monotypic taxon.
===Evolution===
thumb|right|Fossil skull of a jaguar dating to the Pleistocene
The evolutionary radiation of the Felidae began in the late Miocene between and in Asia.
The clouded leopard (Neofelis nebulosa) is the first cat that genetically diverged from the common ancestor of the Felidae.
The genetic lineage of the Panthera is estimated to have diverged between and .
The jaguar is thought to have entered the American continent in the early Pleistocene via Beringia, the land bridge that once spanned the Bering Strait. Results of mitochondrial DNA analysis of 37 jaguars indicate that the species evolved between 510,000 and 280,000 years ago.
The oldest fossil remains of the extinct jaguar subspecies Panthera onca augusta and Panthera onca mesembrina excavated in North and South America date to the Last Interglacial about 130,000 years before present.
==Description==
The jaguar is a compact and well-muscled animal. It is the largest cat native to the Americas and the third largest in the world, exceeded in size only by the tiger and the lion. It stands tall at the shoulders.
Its size and weight vary considerably: weights are normally in the range of . Exceptionally big males have been recorded to weigh as much as .
The smallest females weigh about .
It is sexually dimorphic, with females typically being 10–20% smaller than males. The length, from the nose to the base of the tail, varies from . The tail is long and the shortest of any big cat.
Its muscular legs are shorter than the legs of other Panthera species with similar body weight.
Further variations in size have been observed across regions and habitats, with size tending to increase from north to south. Jaguars in the Chamela-Cuixmala Biosphere Reserve on the Pacific coast of central Mexico weighed around , about the size of a female cougar (Puma concolor).
Jaguars in Venezuela and Brazil are much larger, with average weights of about in males and of about in females.
The jaguar's coat ranges from pale yellow to tan or reddish-yellow, while the ventral areas are whitish, and is covered in black spots. The spots and their shapes vary: on the sides they become rosettes which may include one or several dots. The spots on the head and neck are generally solid, as are those on the tail where they may merge to form bands near the end and create a black tip. They are elongated on the middle of the back, often connecting to create a median stripe, and blotchy on the belly. These patterns serve as camouflage in areas with dense vegetation and patchy shadows.
Jaguars living in forests are often darker and considerably smaller than those living in open areas, possibly due to the smaller numbers of large, herbivorous prey in forest areas.
While the jaguar closely resembles the leopard, it is generally more robust, with stockier limbs and a more square head. The rosettes on a jaguar's coat are larger, darker, fewer in number and have thicker lines, with a small spot in the middle.
It has powerful jaws with the third-highest bite force of all felids, after the tiger and the lion.
It has an average bite force at the canine tip of 887.0 Newton and a bite force quotient at the canine tip of 118.6.
A jaguar can bite with a force of with the canine teeth and at the carnassial notch.
===Color variation===
Melanistic jaguars are also known as black panthers. The black morph is less common than the spotted one.
Black jaguars have been documented in Central and South America. Melanism in the jaguar is caused by deletions in the melanocortin 1 receptor gene and inherited through a dominant allele.
In 2004, a camera trap in the Sierra Madre Occidental mountains photographed the first documented black jaguar in Northern Mexico. Black jaguars were also photographed in Costa Rica's Alberto Manuel Brenes Biological Reserve, in the mountains of the Cordillera de Talamanca, in Barbilla National Park and in eastern Panama.
==Distribution and habitat==
At present, the jaguar's range extends from Mexico through Central America to South America, including much of Amazonian Brazil. The countries included in its range are Argentina, Belize, Bolivia, Colombia, Costa Rica, particularly on the Osa Peninsula, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Nicaragua, Panama, Paraguay, Peru, Suriname, the United States and Venezuela. It is considered to be locally extinct in El Salvador and Uruguay.
In 1999, its historic range at the turn of the 20th century was estimated at , stretching from the southern United States through Central America to southern Argentina. By the turn of the 21st century, its global range had decreased to about , most of it in the southern United States, northern Mexico, northern Brazil, and southern Argentina.
The jaguar prefers dense forest and typically inhabits dry deciduous forests, tropical and subtropical moist broadleaf forests, rainforests and cloud forests in Central and South America; open, seasonally flooded wetlands, dry grassland and historically also oak forests in the United States. It has been recorded at elevations up to but avoids montane forests. It favors riverine habitat and swamps with dense vegetation cover. Results of a study in the Mayan forests of Mexico and Guatemala showed that 11 GPS-collared jaguars preferred undisturbed dense habitat away from roads; females avoided even areas with low levels of human activity, whereas males appeared less disturbed by human population density.
Sightings of jaguars as far north as the North Platte River in Colorado were recorded in the 19th century.
In 1919, sightings of jaguars were reported to have occurred in the Monterey, California region.
Jaguars have been occasionally sighted in Arizona, New Mexico and Texas.
Between 2012 and 2015, a male vagrant jaguar was recorded in 23 locations in the Santa Rita Mountains.
==Behavior and ecology==
The jaguar is mostly active at night and during twilight.
However, jaguars living in densely forested regions of the Amazon Rainforest and the Pantanal are largely active by day, whereas jaguars in the Atlantic Forest are primarily active by night.
The activity pattern of the jaguar coincides with the activity of its main prey species.
It is an expert climber and swimmer. In seasonally flooded forests, it lives partially in trees and preys on arboreal wildlife.
===Ecological role===
The adult jaguar is an apex predator, meaning it is at the top of the food chain and is not preyed upon in the wild. The jaguar has also been termed a keystone species, as it is assumed that it controls the population levels of prey such as herbivorous and granivorous mammals, and thus maintains the structural integrity of forest systems.
However, accurately determining what effect species like the jaguar have on ecosystems is difficult, because data must be compared from its current habitats and regions where it is absent, and while controlling for the effects of human activity. It is accepted that mid-sized prey species undergo population increases in the absence of the keystone predators, which has been hypothesized to have cascading negative effects.
However, field work has shown this may be natural variability, and the population increases may not be sustained. Thus, the keystone predator hypothesis is not accepted by all scientists.
The jaguar is sympatric with the cougar (Puma concolor). In central Mexico, both prey on white-tailed deer (Odocoileus virginianus), which makes up 54% and 66% of jaguar and cougar's prey, respectively. In northern Mexico, the jaguar and the cougar share the same habitat, and their diet overlaps dependent on prey availability. Jaguars seemed to prefer deer and calves. In Mexico and Central America, neither of the two cats are considered to be the dominant predator.
In South America, the jaguar is larger than the cougar and tends to take larger prey, usually over . The cougar's prey usually weighs between , which is thought to be the reason for its smaller size.
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.
===Hunting and diet===
The jaguar is an obligate carnivore and depends solely on flesh for its nutrient requirements. An analysis of 53 studies documenting the diet of the jaguar revealed that its prey ranges in weight from ; it prefers prey weighing , with capybara (Hydrochoerus hydrochaeris) and giant anteater (Myrmecophaga tridactyla) being significantly preferred. When available, it also preys on marsh deer (Blastocerus dichotomus), southern tamandua (Tamandua tetradactyla), collared peccary (Pecari tajacu) and black agouti (Dasyprocta fuliginosa). In floodplains, jaguars opportunistically take reptiles such as turtles and caimans. Consumption of reptiles appears to be more frequent in jaguars than in other big cats.
The jaguar also preys on livestock in cattle ranching areas where wild prey is scarce.
The daily food requirement of a captive jaguar weighing was estimated at of meat.
The jaguar's bite force allows it to pierce the carapaces of the yellow-spotted Amazon river turtle (Podocnemis unifilis) and the yellow-footed tortoise (Chelonoidis denticulatus)
and to employ an unusual killing method: it bites mammalian prey directly through the skull between the ears to deliver a fatal bite to the brain. It kills capybara by piercing its canine teeth through the temporal bones of its skull, breaking its zygomatic arch and mandible and penetrating its brain, often through the ears.
It has been hypothesized to be an adaptation to "cracking open" turtle shells; armored reptiles may have formed an abundant prey base for the jaguar following the late Pleistocene extinctions. However, this is disputed, as even in areas where jaguars prey on reptiles, they are taken relatively infrequently in comparison to their abundance and mammals still dominate the cat's diet.
Between October 2001 and April 2004, 10 jaguars were monitored in the southern Pantanal. In the dry season from April to September, they killed prey at intervals ranging from one to seven days; and ranging from one to 16 days in the wet season from October to March.
The jaguar uses a stalk-and-ambush strategy when hunting rather than chasing prey. The cat will slowly walk 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. After killing prey, the jaguar will drag the carcass to a thicket or other secluded spot. It begins eating at the neck and chest. The heart and lungs are consumed, followed by the shoulders.Baker, Natural History and Behavior, pp. 8–16.
===Social activity===
The jaguar is solitary except for females with cubs. In 1977, groups consisting of a male, female and cubs, and two females with two males were sighted several times in a study area in the Paraguay River valley. A radio-collared female moved in a home range of , which partly overlapped with another female. The home range of the male in this study area overlapped with several females.
The jaguar uses scrape marks, urine, and feces to mark its territory.
The size of home ranges depends on the level of deforestation and human population density. The home ranges of females vary from in the Pantanal to in the Amazon to in the Atlantic Forest. Male jaguar home ranges vary from in the Pantanal to in the Amazon to in the Atlantic Forest and in the Cerrado.
Studies employing GPS telemetry in 2003 and 2004 found densities of only six to seven jaguars per 100 km2 in the Pantanal region, compared with 10 to 11 using traditional methods; this suggests the widely used sampling methods may inflate the actual numbers of individuals in a sampling area.
Fights between males occur but are rare, and avoidance behavior has been observed in the wild. The jaguar roars or grunts to warn territorial and mating competitors; intensive bouts of counter-calling between individuals have been observed in the wild. This vocalization contains five or six guttural notes. The female jaguar prustens when approached by the male. This sound is described as a short, low intensity, non-threatening vocalization, possibly intended to signal tranquility and passivity.
It is often used between two cats as a greeting, during courting, or by a mother comforting her cubs. Cubs have been recorded bleating, gurgling and mewing.
===Reproduction and life cycle===
In captivity, the female jaguar reaches sexual maturity at the age of about 2.5 years. Estrus lasts 7–15 days with an estrus cycle of 41.8 to 52.6 days. During estrus, she exhibits increased restlessness with rolling and prolonged vocalizations.
She is an induced ovulator but can also ovulate spontaneously.
Gestation lasts 91 to 111 days.
The male is sexually mature at the age of three to four years.
His mean ejaculate volume is 8.6±1.3 ml.
Generation length of the jaguar is 9.8 years.
In the Pantanal, breeding pairs were observed to stay together for up to five days. Females had one to two cubs.
The young are born with closed eyes but open them after two weeks. Cubs are weaned at the age of three months but remain in the birth den for six months before leaving to accompany their mother on hunts.
Jaguars remain with their mothers for up to two years. They appear to rarely live beyond 11 years, but captive individuals may live 22 years.
In 2001, a male jaguar killed and partially consumed two cubs in Emas National Park. DNA paternity testing of blood samples revealed that the male was the father of the cubs. Two more cases of infanticide were documented in the northern Pantanal in 2013.
===Attacks on humans===
The Spanish conquistadors feared the jaguar. According to Charles Darwin, the indigenous peoples of South America stated that people did not need to fear the jaguar as long as capybaras were abundant.
The first official record of a jaguar killing a human in Brazil dates to June 2008.
Two children were attacked by jaguars in Guyana.
The jaguar is the least likely of all big cats to kill and eat humans, and the majority of attacks come when it has been cornered or wounded.
==Threats==
The jaguar is threatened by loss and fragmentation of habitat, illegal killing in retaliation for livestock depredation and for illegal trade in jaguar body parts. It is listed as Near Threatened on the IUCN Red List, as the jaguar population has probably declined by 20–25% since the mid-1990s. Deforestation is a major threat to the jaguar across its range. Habitat loss was most rapid in drier regions such as the Argentine pampas, the arid grasslands of Mexico and the southwestern United States.
In 2002, it was estimated that the range of the jaguar had declined to about 46% of its range in the early 20th century. In 2018, it was estimated that its range had declined by 55% in the last century. The only remaining stronghold is the Amazon rainforest, a region that is rapidly being fragmented by deforestation.
Between 2000 and 2012, forest loss in the jaguar range amounted to , with fragmentation increasing in particular in corridors between Jaguar Conservation Units (JCUs).
By 2014, direct linkages between two JCUs in Bolivia were lost, and two JCUs in northern Argentina became completely isolated due to deforestation.
In Mexico, the jaguar is primarily threatened by poaching. Its habitat is fragmented in northern Mexico, in the Gulf of Mexico and in the Yucatán Peninsula, caused by changes in land use, construction of roads and tourism infrastructure.
In Panama, 220 of 230 jaguars were killed in retaliation for predation on livestock between 1998 and 2014.
In Venezuela, the jaguar was extirpated in about 26% of its range in the country since 1940, mostly in dry savannas and unproductive scrubland in the northeastern region of Anzoátegui.
In Ecuador, the jaguar is threatened by a reduced prey availability in areas where the expansion of the road network facilitated access of human hunters to forests.
In the Alto Paraná Atlantic forests, at least 117 jaguars were killed in Iguaçu National Park and the adjacent Misiones Province between 1995 and 2008.
Some Afro-Colombians in the Colombian Chocó Department hunt jaguars for consumption and sale of meat.
Between 2008 and 2012, at least 15 jaguars were killed by livestock farmers in central Belize.
The international trade of jaguar skins boomed between the end of the Second World War and the early 1970s.
Significant declines occurred in the 1960s, as more than 15,000 jaguars were yearly killed for their skins in the Brazilian Amazon alone; the trade in jaguar skins decreased since 1973 when the Convention on International Trade in Endangered Species was enacted.
Interview surveys with 533 people in the northwestern Bolivian Amazon revealed that local people killed jaguars out of fear, in retaliation, and for trade.
Between August 2016 and August 2019, jaguar skins and body parts were seen for sale in tourist markets in the Peruvian cities of Lima, Iquitos and Pucallpa.
Human-wildlife conflict, opportunistic hunting and hunting for trade in domestic markets are key drivers for killing jaguars in Belize and Guatemala.
Seizure reports indicate that at least 857 jaguars were involved in trade between 2012 and 2018, including 482 individuals in Bolivia alone; 31 jaguars were seized in China.
Between 2014 and early 2019, 760 jaguar fangs were seized that originated in Bolivia and were destined for China. Undercover investigations revealed that the smuggling of jaguar body parts is run by Chinese residents in Bolivia.
==Conservation==
The jaguar is listed on CITES Appendix I, which means that all international trade in jaguars or their body parts is prohibited. Hunting jaguars is prohibited in Argentina, Brazil, Colombia, French Guiana, Honduras, Nicaragua, Panama, Paraguay, Suriname, the United States, and Venezuela. Hunting jaguars is restricted in Guatemala and Peru. In Ecuador, hunting jaguars is prohibited, and it is classified as threatened with extinction.
In Guyana, it is protected as an endangered species, and hunting it is illegal.
In 1986, the Cockscomb Basin Wildlife Sanctuary was established in Belize as the world's first protected area for jaguar conservation.
===Jaguar Conservation Units===
In 1999, field scientists from 18 jaguar range countries determined the most important areas for long-term jaguar conservation based on the status of jaguar population units, stability of prey base and quality of habitat. These areas, called "Jaguar Conservation Units" (JCUs), are large enough for at least 50 breeding individuals and range in size from ; 51 JCUs were designated in 36 geographic regions including:
* the Sierra Madre Occidental and Sierra de Tamaulipas in Mexico
* the Selva Maya tropical forests extending over Mexico, Belize and Guatemala
* the Chocó–Darién moist forests from Honduras and Panama to Colombia
* Venezuelan Llanos
* northern Cerrado and Amazon basin in Brazil
* Tropical Andes in Bolivia, Peru and Argentina
* Misiones Province in Argentina
Optimal routes of travel between core jaguar population units were identified across its range in 2010 with the aim of implementing wildlife corridors that connect JCUs. These corridors represent areas with the shortest distance between jaguar breeding populations, require the least possible energy input of dispersing individuals and pose a low mortality risk. These corridors cover an area of and range in length from in Mexico and Central America and from in South America.
Cooperation with local landowners and municipal, state, or federal agencies are essential to maintain connected populations and prevent fragmentation in both JCUs and corridors.
Seven of 13 corridors in Mexico are functioning with a width of at least and a length of no more than . The other corridors may hamper passage, as they are narrower and longer.
In August 2012, the United States Fish and Wildlife Service set aside in Arizona and New Mexico for the protection of the jaguar.
===Approaches===
In setting up protected reserves, efforts generally also have to be focused on the surrounding areas, as jaguars are unlikely to confine themselves to the bounds of a reservation, especially if the population is increasing in size. Human attitudes in the areas surrounding reserves and laws and regulations to prevent poaching are essential to make conservation areas effective.
To estimate population sizes within specific areas and to keep track of individual jaguars, camera trapping and wildlife tracking telemetry are widely used, and feces may be sought out with the help of detector dogs to study jaguar health and diet.
Current conservation efforts often focus on educating ranch owners and promoting ecotourism.
The jaguar is generally defined as an umbrella species – its home range and habitat requirements are sufficiently broad that, if protected, numerous other species of smaller range will also be protected.
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 other species will also benefit.
Ecotourism setups are being used to generate public interest in charismatic animals such as the jaguar while at the same time generating revenue that can be used in conservation efforts. Audits done in Africa have shown that ecotourism has helped in African cat conservation. As with large African cats, a key concern in jaguar ecotourism is the considerable habitat space the species requires. So if ecotourism is used to aid in jaguar conservation, some considerations need to be made as to how existing ecosystems will be kept intact or how new ecosystems that are large enough to support a growing jaguar population will be put into place.
==In culture and mythology==
In pre-Columbian Central and South America, the jaguar was a symbol of power and strength. In the Andes, a jaguar cult disseminated by the early Chavín culture became accepted over most of today's Peru by 900 BC. The later Moche culture of northern Peru used the jaguar as a symbol of power in many of their ceramics.
In the Muisca religion in Altiplano Cundiboyacense, the jaguar was considered a sacred animal, and people dressed in jaguar skins during religious rituals.
The skins were traded with peoples in the nearby Orinoquía Region.
The name of the Muisca ruler Nemequene was derived from the Chibcha words nymy and quyne, meaning "force of the jaguar".
Sculptures with "Olmec were-jaguar" motifs were found on the Yucatán Peninsula in Veracruz and Tabasco; they show stylized jaguars with half-human faces.
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 several Maya rulers bore names that incorporated the Mayan word for jaguar (b'alam in many of the Mayan languages). Balam (Jaguar) remains a common Maya surname, and it is also the name of Chilam Balam, a legendary author to whom are attributed 17th and 18th-centuries Maya miscellanies preserving much important knowledge. Remains of jaguar bones were discovered in a burial site in Guatemala, which indicates that Mayans kept jaguars as pets.
The Aztec civilization shared this image of the jaguar as the representative of the ruler and as a warrior. The Aztecs formed an elite warrior class known as the Jaguar warrior. In Aztec mythology, the jaguar was considered to be the totem animal of the powerful deity Tezcatlipoca.
A conch shell gorget depicting a jaguar was found in a burial mound in Benton County, Missouri. The gorget shows evenly-engraved lines and measures .
Rock drawings made by the Hopi, Anasazi and Pueblo all over the desert and chaparral regions of the American Southwest show an explicitly spotted cat, presumably a jaguar, as it is drawn much larger than an ocelot.
The jaguar is also used as a symbol in contemporary culture. It is the national animal of Guyana and is featured in its coat of arms.
The flag of the Department of Amazonas features a black jaguar silhouette leaping towards a hunter.
The crest of the Argentine Rugby Union features a jaguar.
==See also==
* List of largest cats
== References ==
===Bibliography===
* 
==External links==
* 
* 
* People and Jaguars a Guide for Coexistence
* Felidae Conservation Fund
* 
 
Category:Apex predators
Category:Big cats
Category:ESA endangered species
Category:Fauna of the Amazon
Category:Fauna of the Cerrado
Category:Fauna of the Pantanal
Category:Fauna of the Southwestern United States
Category:Felids of Central America
Category:Felids of North America
Category:Felids of South America
Category:Mammals described in 1758
Category:Near threatened animals
Category:Near threatened biota of South America
Category:Near threatened biota of North America
onca
Category:Taxa named by Carl Linnaeus