The jaguar (Panthera onca) is a large catspecies and the only living member of the genusPanthera native to the Americas. With a body length of up to 1.85m (6ft 1in) and a weight of up to 158kg (348lb), it is the largest cat species in the Americas and the third largest in the world. Its distinctively marked coat features pale yellow to tan colored fur covered by spots that transition to rosettes on the sides, although a melanistic black coat appears in some individuals. The jaguar's 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 mammalianprey between the ears to deliver a fatal blow to the brain.
Reginald Innes Pocock placed the jaguar in the genusPanthera 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.
The lineage of the jaguar appears to have originated in Africa and spread to Eurasia 1.95–1.77 mya. The modern species may have descended from Panthera gombaszoegensis, which is thought to have entered the American continent via Beringia, the land bridge that once spanned the Bering Strait. Fossils of modern jaguars have been found in North America dating to over 850,000 years ago. Results of mitochondrial DNA analysis of 37 jaguars indicate that current populations evolved between 510,000 and 280,000 years ago in northern South America and subsequently recolonized North and Central America after the extinction of jaguars there during the Late Pleistocene.
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 68 to 75cm (26.8 to 29.5in) tall at the shoulders. Its size and weight vary considerably: weights are normally in the range of 56–96kg (123–212lb). Exceptionally big males have been recorded to weigh as much as 158kg (348lb). The smallest females weigh about 36kg (79lb). 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 1.12 to 1.85m (3ft 8in to 6ft 1in). The tail is 45 to 75cm (18 to 30in) 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 50kg (110lb), which is about the size of a female cougar (Puma concolor). Jaguars in Venezuela and Brazil are much larger, with average weights of about 95kg (209lb) in males and of about 56–78kg (123–172lb) in females.
The jaguar's coat ranges from pale yellow to tan or reddish-yellow, with a whitish underside and 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.
The jaguar closely resembles the leopard but 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 100kg (220lb) jaguar can bite with a force of 4.939kN (1,110lbf) with the canine teeth and 6.922kN (1,556lbf) at the carnassial notch.
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. Jaguars are good swimmers and play and hunt in the water, possibly more than tigers. They have been recorded moving between islands and the shore. Jaguars are also good at climbing trees but do so less often than cougars.
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 seed-eating mammals and thus maintains the structural integrity of forest systems. 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 22kg (49lb). The cougar's prey usually weighs between 2 and 22kg (4 and 49lb), 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 1 to 130kg (2.2 to 286.6lb); it prefers prey weighing 45–85kg (99–187lb), 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 (Dicotyles 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. One remote population in the Brazilian Pantanal is recorded to primarily feed on aquatic reptiles and fish. The jaguar also preys on livestock in cattleranching areas where wild prey is scarce. The daily food requirement of a captive jaguar weighing 34kg (75lb) was estimated at 1.4kg (3.1lb) 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). It employs 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.
The jaguar is generally 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 25–38km2 (9.7–14.7sqmi), 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 15.3km2 (5.9sqmi) in the Pantanal to 53.6km2 (20.7sqmi) in the Amazon to 233.5km2 (90.2sqmi) in the Atlantic Forest. Male jaguar home ranges vary from 25km2 (9.7sqmi) in the Pantanal to 180.3km2 (69.6sqmi) in the Amazon to 591.4km2 (228.3sqmi) in the Atlantic Forest and 807.4km2 (311.7sqmi) in the Cerrado. Studies employing GPStelemetry in 2003 and 2004 found densities of only six to seven jaguars per 100km2 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. In one wetland population with broken down territorial boundaries and a high population density, adults of the same sex have been observed fishing, traveling and playing together.
The jaguar roars or grunts for long-distance communication; intensive bouts of counter-calling between individuals have been observed in the wild. This vocalization is described as "hoarse" and contains five or six guttural notes.Chuffing is produced by individuals when greeting, during courting, or by a mother comforting her cubs. This sound is described as short, low intensity, non-threatening snorts, possibly intended to signal tranquility and passivity. Cubs have been recorded bleating, gurgling and mewing.
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. Infanticide may be combated by the female hiding her cubs and distracting the male with courtship behavior.
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.
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 since 2002, 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 83.759km2 (32.340sqmi), 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 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 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.
The jaguar is listed on CITES Appendix I, which means that all international commercial 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 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 566 to 67,598km2 (219 to 26,100sqmi); 51 JCUs were designated in 36 geographic regions including:
Optimal routes of travel between core jaguar population units were identified across its range in 2010 to implement 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. They cover an area of 2,600,000km2 (1,000,000sqmi) and range in length from 3 to 1,102km (1.9 to 684.8mi) in Mexico and Central America and from 489.14 to 1,607km (303.94 to 998.54mi) in South America. Cooperation with local landowners and municipal, state, or federal agencies is 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 14.25km (8.85mi) and a length of no more than 320km (200mi). The other corridors may hamper passage, as they are narrower and longer.
In August 2012, the United States Fish and Wildlife Service set aside 3,392.20km2 (838,232 acres) in Arizona and New Mexico for the protection of the jaguar. The Jaguar Recovery Plan was published in April 2019, in which Interstate 10 is considered to form the northern boundary of the Jaguar Recovery Unit in Arizona and New Mexico.
In Mexico, a national conservation strategy was developed from 2005 on and published in 2016. The Mexican jaguar population increased from an estimated 4,000 individuals in 2010 to about 4,800 individuals in 2018. This increase is seen as a positive effect of conservation measures that were implemented in cooperation with governmental and non-governmental institutions and landowners.
An evaluation of JCUs from Mexico to Argentina revealed that they overlap with high-quality habitats of about 1,500 mammals to varying degrees. Since co-occurring mammals benefit from the JCU approach, the jaguar has been called an umbrella species. Central American JCUs overlap with the habitat of 187 of 304 regional endemic amphibian and reptile species, of which 19 amphibians occur only in the jaguar range.
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.
Current conservation efforts often focus on educating ranch owners and promoting ecotourism. 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. A key concern in jaguar ecotourism is the considerable habitat space the species requires. 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 will be put into place that are large enough to support a growing jaguar population.
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 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 may have kept jaguars as pets.
Felidae is a family of mammals in the order Carnivora, colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid. The term "cat" refers both to felids in general and specifically to the domestic cat.
The tiger is the largest living cat species and a member of the genus Panthera. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily preys on ungulates such as deer and wild boar. It is territorial and generally a solitary but social predator, requiring large contiguous areas of habitat, which support its requirements for prey and rearing of its offspring. Tiger cubs stay with their mother for about two years, then become independent and leave their mother's home range to establish their own.
A black panther is the melanistic colour variant of the leopard and the jaguar. Black panthers of both species have excess black pigments, but their typical rosettes are also present. They have been documented mostly in tropical forests, with black leopards in Kenya, India, Sri Lanka, Nepal, Thailand, Peninsular Malaysia and Java, and black jaguars of the Americas in Mexico, Panama, Costa Rica and Paraguay. Melanism is caused by a recessive allele in the leopard, and by a dominant allele in the jaguar.
The leopard is one of the five extant species in the genus Panthera, a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia, and on the Indian subcontinent to Southeast and East Asia. It is listed as Vulnerable on the IUCN Red List because leopard populations are threatened by habitat loss and fragmentation, and are declining in large parts of the global range. The leopard is considered locally extinct in Hong Kong, Singapore, South Korea, Jordan, Morocco, Togo, the United Arab Emirates, Uzbekistan, Lebanon, Mauritania, Kuwait, Syria, Libya, Tunisia and most likely in North Korea, Gambia, Laos, Lesotho, Tajikistan, Vietnam and Israel. Contemporary records suggest that the leopard occurs in only 25% of its historical global range.
Panthera is a genus within the family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as comprising the tiger, lion, jaguar, and leopard on the basis of common cranial features. Results of genetic analysis indicate that the snow leopard also belongs to the genus Panthera, a classification that was accepted by IUCN Red List assessors in 2008.
The cougar is a large cat native to the Americas. Its range spans from the Canadian Yukon to the southern Andes in South America and is the most widespread of any large wild terrestrial mammal in the Western Hemisphere. It is an adaptable, generalist species, occurring in most American habitat types. Due to its wide range, it has many names, including puma, mountain lion, catamount and panther.
The term "big cat" is typically used to refer to any of the five living members of the genus Panthera, namely the tiger, lion, jaguar, leopard, and snow leopard, as well as the non-pantherine cheetah and cougar. However, only the first 4 of these species are able to roar.
The ocelot is a medium-sized spotted wild cat that reaches 40–50 cm (15.7–19.7 in) at the shoulders and weighs between 8 and 15.5 kg. It was first described by Carl Linnaeus in 1758. Two subspecies are recognized. It is native to the southwestern United States, Mexico, Central and South America, and to the Caribbean islands of Trinidad and Margarita. It prefers areas close to water sources with dense vegetation cover and high prey availability.
The jaguarundi is a wild cat native to the Americas. Its range extends from central Argentina in the south to northern Mexico, through Central and South America east of the Andes. The jaguarundi is a medium-sized cat of slender build. Its coloration is uniform with two color morphs, gray and red. It has an elongated body, with relatively short legs, a small, narrow head, small, round ears, a short snout, and a long tail, resembling mustelids in these respects. It is about twice as large as a domestic cat, reaching nearly 360 mm (14 in) at the shoulder, and weighs 3.5–7 kg (7.7–15.4 lb).
The American lion, also known as the "North American lion", or "American cave lion", is an extinct pantherine cat that lived in North America during the Pleistocene epoch and the early Holocene epoch, about 340,000 to 11,000 years ago. Its fossils have been excavated from Alaska to Mexico. Genetic analysis has shown that the American lion and the Late Pleistocene Eurasian cave lion are sister lineages. It was about 25% larger than the modern lion, making it one of the largest known felids.
Pantherinae is a subfamily within the family Felidae; it was named and first described by Reginald Innes Pocock in 1917 as only including the Panthera species. The Pantherinae genetically diverged from a common ancestor between 9.32 to 4.47 million years ago and 10.67 to 3.76 million years ago.
The North American cougar is a cougar subspecies in North America. It was once common in eastern North America, and is still prevalent in the western half of the continent. This subspecies includes populations in western Canada, the western United States, Florida, Mexico and Central America, and possibly South America northwest of the Andes Mountains. It is the biggest cat in North America, with North American jaguars being fairly small. It thus includes the extirpated Eastern cougar and extant Florida panther populations.
Panthera gombaszoegensis, also known as the European jaguar, is a Panthera species that lived from about 2.0 to 0.35 million years ago in Europe. The first fossils were excavated in 1938 in Gombasek, Slovakia.
The snow leopard, also known as the ounce, is a felid in the genus Panthera native to the mountain ranges of Central and South Asia. It is listed as Vulnerable on the IUCN Red List because the global population is estimated to number fewer than 10,000 mature individuals and is expected to decline about 10% by 2040. It is threatened by poaching and habitat destruction following infrastructural developments. It inhabits alpine and subalpine zones at elevations of 3,000–4,500 m (9,800–14,800 ft), ranging from eastern Afghanistan, the Himalayas and the Tibetan Plateau to southern Siberia, Mongolia and western China. In the northern part of its range, it also lives at lower elevations.
The southern brown howler is a monkey subspecies of brown howler native to southeastern Brazil and far northeastern Argentina (Misiones). Gregorin, 2006, considered the southern brown howler to be a separate species, Alouatta clamitans, but this has not been universally accepted.
Panthera onca mesembrina is an extinct subspecies of the jaguar that was endemic to southern South America during the Pleistocene epoch. Its fossils have been excavated primarily in Argentina and Chile, though few fossils are known. Genetic analysis in 2016 showed that P. onca mesembrina was in an extinct sister lineage to extant Panthera onca species based on genetic evidence, and is the largest known subspecies of jaguar.
Panthera onca augusta, commonly known as the Pleistocene jaguar or simply the giant jaguar, is an extinct subspecies of the jaguar that was endemic to North America during the Pleistocene epoch.
The North American jaguar is a jaguar population in North America, ranging from the southwestern United States to Central America. This population has declined over decades and almost eliminated by 1960.
The South American jaguar is a jaguar population in South America. Though a number of subspecies of jaguar have been proposed for South America, morphological and genetic research did not reveal any evidence for subspecific differentiation.
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↑ de Paula, R.; Campos Neto, M. F. & Morato, R. G. (2008). "First Official Record of Human Killed by Jaguar in Brazil". Cat News (49): 31–32.
↑ Zapata Ríos, G.; Araguillin, E.; Cevallos, J.; Moreno, F.; Ortega, A.; Rengel, J. & Valarezo, N. (2014). Plan de Acción para la Conservación del Jaguar en el Ecuador[Action Plan for the Conservation of the Jaguar in Ecuador](PDF) (in Spanish). Quito: Ministerio del Ambiente y Wildlife Conservation Society Ecuador. Archived(PDF) from the original on 23 September 2020. Retrieved 13 February 2021.
↑ Park, Yumi (2012). Mirrors of Clay: Reflections of Ancient Andean Life in Ceramics from the Sam Olden Collection. University Press of Mississippi. p.49. ISBN9781617037955.
↑ Ocampo López, J. (2007). Grandes culturas indígenas de América – Great indigenous cultures of the Americas (in Spanish). Bogotá, Colombia: Plaza & Janes Editores Colombia S.A. p.231. ISBN978-958-14-0368-4.