Jaguar

In 1758, Carl Linnaeus described the jaguar in his work Systema Naturae and gave it the scientific name Felis onca.[6] In the 19th and 20th centuries, several jaguar type specimens formed the basis for descriptions of subspecies.[7] In 1939, Reginald Innes Pocock recognized eight subspecies based on geographic origins and skull morphology of these specimens.[8] 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.[9] By 2005, nine subspecies were considered to be valid taxa.[7]

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.[8] Results of morphological and genetic research indicate a clinal north–south variation between populations, but no evidence for subspecific differentiation.[17][18] DNA analysis of 84 jaguar samples from South America revealed that the gene flow between jaguar populations in Colombia was high in the past.[19]

Since 2017, the jaguar is therefore considered to be a monotypic taxon.[20]

Fossil skull of a jaguar dating to the Pleistocene

The evolutionary radiation of the Felidae began in the late Miocene between 14.45 to 8.38 million years ago and 16.76 to 6.46 million years ago in Asia.[21][22] The clouded leopard (Neofelis nebulosa) is the first cat that genetically diverged from the common ancestor of the Felidae.[23][24][25] The genetic lineage of the Panthera is estimated to have diverged between 9.32 to 4.47 million years ago[21] and 11.75 to 0.97 million years ago.[22]

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.[17] Fossil remains of the extinct jaguar subspecies Panthera onca augusta and Panthera onca mesembrina excavated in North and South America date to the Pleistocene and are up to 130,000 years old.[26]

A black jaguar

Melanistic jaguars are informally known as black panthers. The black morph is less common than the spotted one.[39] Melanism in the jaguar is caused by deletions in the melanocortin 1 receptor gene and inherited through a dominant allele.[40]

Melanistic jaguars had been documented in Central and South America, in the southern portion of their range. In 2004, the first black jaguar was recorded in the northern portion of their range, in the Sierra Madre Occidental area of Mexico.[41] Black jaguars were also recorded in Costa Rica's Alberto Manuel Brenes Biological Reserve and in the mountains of the Cordillera de Talamanca.[42][43]

Albino jaguars, sometimes called white panthers, are extremely rare.[29]

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.[35][48][49]

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 undergo population increases in the absence of the keystone predators, which has been hypothesized to have cascading negative effects.[50] 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.[51]

The jaguar also coexists with other predators. The jaguar and the cougar, which is the next-largest cat in South America, but is usually larger than the jaguar in Central or North America,[35] are often sympatric (related species sharing overlapping territory) and have often been studied in conjunction. In northern Mexico, the jaguar and cougar were found to almost always live in the same areas with neither species having areas where it was the only one of the two living.[52] A study in central Mexico found that the favorite prey of both species was white-tailed deer, with it making up 54% and 66% of jaguar and cougar's prey by biomass, respectively.[35] Therefore, in Mexico and Central America, where cougars are usually equal to or slightly larger than the jaguar in size, neither would be considered to dominant over the other.[52]

In South America, however, where the jaguar is larger than the cougar, the jaguar tends to take larger prey, usually over 22 kg (49 lb) and the cougar smaller prey, usually between 2 and 22 kg (4 and 49 lb), which reduces the cougar's average size.[53] 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;[35] while both are classified as near-threatened species, the cougar has a significantly larger current distribution. Depending on the availability of prey, the cougar and jaguar may even share it.[52]

The jaguar has an exceptionally powerful bite that allows it to pierce the shells of armored prey.

Illustration of a jaguar killing a tapir, the largest native land animal in its range

Like all cats, the jaguar is an obligate carnivore, feeding only on meat. It is an opportunistic hunter, and its diet encompasses at least 87 species.[29] An analysis of 53 studies documenting the diet of the jaguar across its distribution revealed that its prey ranges in weight from 1–130 kg (2.2–286.6 lb); it prefers prey weighing 45–85 kg (99–187 lb), with capybara (Hydrochoerus hydrochaeris), giant anteater (Myrmecophaga tridactyla), collared peccary (Pecari tajacu), nine-banded armadillo (Dasypus novemcinctus), wild boar (Sus scrofa) and white-nosed coati (Nasua narica) being the most preferred species. Where present, it also preys on Odocoileus deer, spectacled caiman (Caiman crocodilus), South American tapir (Tapirus terrestris), crab-eating fox (Cerdocyon thous), tapeti (Sylvilagus brasiliensis) and agouti species.[27] Some jaguars also prey on livestock such as horses, cattle, and llamas.[54] In the Arizona mountains, a jaguar killed and fed on an American black bear (Ursus americanus).[55]

The jaguar has an exceptionally powerful bite, even relative to the other big cats.[36] Its bite force allows it to pierce the shells of armored reptiles and turtles[56] and to employ an unusual killing method with mammals: it bites directly through the skull of prey between the ears to deliver a fatal bite to the brain.[57] Although the jaguar bites into the throat of South American tapir (Tapirus terrestris) and other large prey until the victim suffocates, 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.[58] This may 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.[56] It has been reported that an individual jaguar can drag an 360 kg (800 lb) bull 8 m (25 ft) in its jaws and pulverize the heaviest bones.[59]

The activity patterns of the jaguar have been found to coincide with the activity of their main prey species in their biomes.[60] Camera trap studies have shown that jaguars primarily have a crepuscular–nocturnal activity pattern in all the biomes that they are found in; however jaguars have been recorded to have considerable diurnal activity in thickly forested regions of the Amazon Rainforest and the Pantanal, as well as purely nocturnal activity in other regions such as the Atlantic forest.[61]

The jaguar uses a stalk-and-ambush strategy when hunting, rather than chasing prey. 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.[62] 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, rather than the midsection. The heart and lungs are consumed, followed by the shoulders.[62]

The daily food requirement of a 34 kg (75 lb) animal, at the extreme low end of the species' weight range, has been estimated at 1.4 kg (3 lb).[63] For captive animals in the 50–60 kg (110–130 lb) range, more than 2 kg (4 lb) of meat daily are recommended.[64] In the wild, consumption is naturally more erratic; wild cats expend considerable energy in the capture and kill of prey, and they may consume up to 25 kg (55 lb) of meat at one feeding, followed by periods of famine.[65] Though carnivorous, there is evidence that wild jaguars consume the roots of Banisteriopsis caapi.[66]

Four month old cub at Salzburg Zoo

Jaguar females reach sexual maturity at about two years of age, and males at three or four. The cat probably mates throughout the year in the wild, with births increasing when prey is plentiful.[67] 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.[68] Generation length of the jaguar is 9.8 years.[69]

Female estrus lasts 6–17 days out of a full 37-day cycle, and they advertise fertility with urinary scent marks and increased vocalization.[67] Females range more widely than usual during courtship. Breeding pairs separate after mating, and females provide all parenting. After a gestation period of 93–105 days, they give birth to two cubs, sometimes up to four. After birth, females do not tolerate the presence of the father; there is a risk of infanticide.[62]

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.[70] They stay in their mother's company for one to two years before establishing 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.[71]

Like most cats, the jaguar is solitary outside mother–cub groups. Adults generally meet only to court and mate (though limited noncourting socialization has been observed anecdotally[62]) and carve out large territories for themselves. Female territories, which range from 25 to 40 km² 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. The territory of a male can contain those of several females.[62][72] The jaguar uses scrape marks, urine, and feces to mark its territory.[73][74]

Like the other big cats except the snow leopard, the jaguar is capable of roaring[75][76] and does so to warn territorial and mating competitors away; intensive bouts of counter-calling between individuals have been observed in the wild.[56] Their roar often resembles a repetitive cough, and they may also vocalize mews and grunts.[71] Mating fights between males occur, but are rare, and aggression avoidance behavior has been observed in the wild.[73] 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.[62]

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 percent of its time active.[29] The jaguar's elusive nature and the inaccessibility of much of its preferred habitat make it a difficult animal to sight, let alone study.

The evolutionary food sources of jaguars did not include large primates, so they do not normally see humans as prey. [77] Experts have cited them as 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.[78] However, such behavior appears to be more frequent whenever humans enter jaguar habitat, and decrease access to prey.[79] } When the Spanish conquistadors arrived in the Americas, they feared jaguars. However, at this time, the jaguar's chief prey was the capybara in South America, and the peccary further north. Charles Darwin reported that the Indigenous peoples of the Americas stated that their people did not need to fear the jaguar, as long as capybaras were abundant.[80] Captive jaguars sometimes attack zookeepers.[81]

Jaguar conservation is complicated because of the species' large range spanning 18 countries with different policies and regulations. Specific areas of high importance for jaguar conservation, so-called "Jaguar Conservation Units" (JCU) were determined in 2000. These are large areas inhabited by at least 50 jaguars. Each unit was assessed and evaluated on the basis of size, connectivity, habitat quality for both jaguar and prey, and jaguar population status. That way, 51 Jaguar Conservation Units were determined in 36 geographic regions as priority areas for jaguar conservation including:[45]

• the Sierra Madre of Mexico
• the Selva Maya tropical forests extending over Mexico, Belize, and Guatemala
• the Chocó-Darién moist forests from Honduras, Panama to Colombia
• Sierra de Tamaulipas
• Venezuelan Llanos
• northern Cerrado and Amazon basin in Brazil
• Misiones Province in Argentina

Recent studies underlined that to maintain the robust exchange across the jaguar gene pool necessary for maintaining the species, it is important that jaguar habitats are interconnected. The Paseo del Jaguar project (Path of the Jaguar), has been established to connect several jaguar hotspots.[86] In 1986, the Cockscomb Basin Wildlife Sanctuary was established in Belize as the world's first protected area for jaguar conservation.[87]

Given the inaccessibility of much of the species' range, particularly the central Amazon, estimating jaguar numbers is difficult. Researchers typically focus on particular bioregions, 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-km² (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 km² (9,000 mi²), may have 465–550 animals.[88] Work employing GPS telemetry in 2003 and 2004 found densities of only six to seven jaguars per 100 km² in the critical Pantanal region, compared with 10 to 11 using traditional methods; this suggests the widely used sampling methods may inflate the actual numbers of cats.[89]

Jaguars are occasionally sighted in Arizona and New Mexico.[90][91][92] In August 2012, the USFWS proposed setting aside 3,392.20 km² (838,232 acres) in Arizona and New Mexico for the protection of the jaguar.[93]

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.[94]

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.[89][95] Current conservation efforts often focus on educating ranch owners and promoting ecotourism.[96] 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.[97] 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.[96]

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.[98]

Jaguar warrior in the Aztec culture

Statuette of Karajà in the Museum of Toulouse

Moche jaguar figurine dating to 300 CE, at the Larco Museum in Lima, Peru

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.[100][101][102] In the religion of the Muisca, who inhabited the cool Altiplano Cundiboyacense in the Colombian Andes, the jaguar was considered a sacred animal and during their religious rituals the people dressed in jaguar skins.[103] The skins were traded with the lowland peoples of the tropical Orinoquía Region.[104] The name of zipa Nemequene was derived from the Muysccubun words nymy and quyne, meaning "force of the jaguar".[105][106]

In Mesoamerica, the Olmec—an early and influential culture of the Gulf Coast of Mexico roughly contemporaneous with the Chavín—developed a distinct "Olmec were-jaguar" motif of sculptures and figurines showing stylised jaguars or humans with jaguar characteristics. 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 a number of 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. 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.[62][107] Remains of jaguar bones were discovered in a burial site in Guatemala which indicates that Mayans kept jaguars as pets.[108]

The jaguar and its name are widely used as a symbol in contemporary culture. It is the national animal of Guyana, and is featured in its Coat of arms of Guyana.[109] The flag of the Department of Amazonas features a black jaguar silhouette pouncing towards a hunter.[110] The jaguar also appears in banknotes of the Brazilian real. The jaguar is also a common fixture in the mythology of several native cultures in South America.[111]

The crest of the Argentine Rugby Union features a jaguar; however, the Argentina national rugby union team is nicknamed Los Pumas.[112] In the spirit of the ancient Mayan culture, the 1968 Summer Olympics in Mexico City adopted a red jaguar as the first official Olympic mascot.[113]

• Baker, W. K. Jr.; et al. Law, Christopher (ed.). Guidelines for Captive Management of Jaguars (PDF). Jaguar Species Survival Plan. American Zoo and Aquarium Association. Archived from the original (PDF) on 13 January 2012. Retrieved 11 November 2011.