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  • Raccoon

    The raccoon (/rəˈkuːn/ or US/ræˈkuːn/ Procyon lotor), also spelled racoon[3] and sometimes called the common raccoon or northern raccoon to distinguish it from the other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of 40 to 70 cm (16 to 28 in), and a body weight of 5 to 26 kg (11 to 57 lb). Its grayish coat mostly consists of dense underfur, which insulates it against cold weather. The animal’s most distinctive features include its extremely dexterous front paws, its facial mask, and its ringed tail, which are common themes in the mythologies of the Indigenous peoples of the Americas surrounding the species. The raccoon is noted for its intelligence, and studies show that it is able to remember the solution to tasks for at least three years. It is usually nocturnal and omnivorous, eating about 40% invertebrates, 33% plants, and 27% vertebrates.

    The original habitats of the raccoon are deciduous and mixed forests, but due to their adaptability, they have extended their range to mountainous areas, coastal marshes, and urban areas, where some homeowners consider them to be pests. As a result of escapes and deliberate introductions in the mid-20th century, raccoons are now also distributed across central Europe, the Caucasus, and Japan.

    In Europe, the raccoon has been included on the list of Invasive Alien Species of Union Concern since 2016.[4] This implies that this species cannot be imported, bred, transported, commercialized, or intentionally released into the environment in the whole of the European Union.[5]

    Though previously thought to be generally solitary, there is now evidence that raccoons engage in sex-specific social behavior. Related females often share a common area, while unrelated males live together in groups of up to four raccoons in order to maintain their positions against foreign males during the mating season and against other potential invaders. Home range sizes vary anywhere from 3 ha (7.4 acres) for females in cities, to 5,000 ha (12,000 acres) for males in prairies. After a gestation period of about 65 days, two to five young known as “kits” are born in spring. The kits are subsequently raised by their mother until dispersal in late fall. Although captive raccoons have been known to live over 20 years, their life expectancy in the wild is only 1.8 to 3.1 years. In many areas, hunting and vehicular injury are the two most common causes of death.

    Etymology

    The mask of a raccoon is often interrupted by a brown-black streak that extends from forehead to nose.[6]

    Names for the species include the common raccoon,[7] North American raccoon,[8] and northern raccoon.[9] In various North American native languages, the reference to the animal’s manual dexterity, or use of its hands is the source for the names.[10] The word raccoon was adopted into English from the native Powhatan term meaning ‘animal that scratches with its hands’, as used in the Colony of Virginia. It was recorded on John Smith‘s list of Powhatan words as aroughcun, and on that of William Strachey as arathkone.[11] It has also been identified as a reflex of a Proto-Algonquian root *ahrah-koon-em, meaning ‘[the] one who rubs, scrubs and scratches with its hands’.[12] The word is sometimes spelled as racoon.[13]

    In Spanish, the raccoon is called mapache, derived from the Nahuatl mapachtli of the Aztecs, meaning ‘[the] one who takes everything in its hands’.[14]

    Its Latin name, procyon lotor, literally means ‘before-dog washer’.[15] The genus Procyon was named by Gottlieb Conrad Christian Storr.[10] The animal’s observed habit of “washing” or “dousing” (see below) is the source of its name in other languages.[16][17] For example, the French “raton laveur” means “washing rat”.

    The colloquial abbreviation coon is used in words like coonskin for fur clothing and in phrases like old coon as a self-designation of trappers.[18][19] In the 1830s, the United States Whig Party used the raccoon as an emblem, causing them to be pejoratively known as “coons” by their political opponents, who saw them as too sympathetic to African-Americans. Soon after that the term became an ethnic slur,[20] especially in use between 1880 and 1920 (see coon song), and the term is still considered offensive.[21] Dogs bred to hunt raccoons are called coonhound and coon dog.[22] Due to having a habit of eating human garbage in urban environments, raccoons are also colloquially known as trash pandas.[23]

    Taxonomy

    Skins of P. lotor and P. cancrivorus

    Skulls of P. lotor and P. cancrivorus

    In the first decades after its discovery by the members of the expedition of Christopher Columbus, who were the first Europeans to leave a written record about the species, taxonomists thought the raccoon was related to many different species, including dogscatsbadgers and particularly bears.[24] Carl Linnaeus, the father of modern taxonomy, placed the raccoon in the genus Ursus, first as Ursus cauda elongata (‘long-tailed bear’) in the second edition of his Systema Naturae (1740), then as Ursus Lotor (‘washer bear’) in the tenth edition (1758–59).[25][26] In 1780, Gottlieb Conrad Christian Storr placed the raccoon in its own genus Procyon, which can be translated as either ‘before the dog’ or ‘doglike’.[27][28] It is also possible that Storr had its nocturnal lifestyle in mind and chose the star Procyon as eponym for the species.[29][30]

    Evolution

    Based on fossil evidence from Russia and Bulgaria, the first known members of the family Procyonidae lived in Europe in the late Oligocene about 25 million years ago.[31] Similar tooth and skull structures suggest procyonids and weasels share a common ancestor, but molecular analysis indicates a closer relationship between raccoons and bears.[32] After the then-existing species crossed the Bering Strait at least six million years later in the early Miocene, the center of its distribution was probably in Central America.[33] Coatis (Nasua and Nasuella) and raccoons (Procyon) have been considered to share common descent from a species in the genus Paranasua present between 5.2 and 6.0 million years ago.[34] This assumption, based on morphological comparisons of fossils, conflicts with a 2006 genetic analysis which indicates raccoons are more closely related to ringtails.[35] Unlike other procyonids, such as the crab-eating raccoon (Procyon cancrivorus), the ancestors of the common raccoon left tropical and subtropical areas and migrated farther north about 2.5 million years ago, in a migration that has been confirmed by the discovery of fossils in the Great Plains dating back to the middle of the Pliocene.[36][34] Its most recent ancestor was likely Procyon rexroadensis, a large Blancan raccoon from the Rexroad Formation characterized by its narrow back teeth and large lower jaw.[37]

    Subspecies

    A Torch Key raccoon (P. l. incautus) in Cudjoe Key, Florida. Subspecies inhabiting the Florida Keys are characterized by their small size and very pale fur.
    Female raccoon of the Vancouver Island subspecies at Sidney, British Columbia, with characteristic dark fur

    As of 2005, Mammal Species of the World recognizes 22 subspecies of raccoons.[38] Four of these subspecies living only on small Central American and Caribbean islands were often regarded as distinct species after their discovery. These are the Bahamian raccoon and Guadeloupe raccoon, which are very similar to each other; the Tres Marias raccoon, which is larger than average and has an angular skull; and the extinct Barbados raccoon. Studies of their morphological and genetic traits in 1999, 2003 and 2005 led all these island raccoons to be listed as subspecies of the common raccoon in Mammal Species of the World’s third edition. A fifth island raccoon population, the Cozumel raccoon, which weighs only 3 to 4 kg (6.6 to 8.8 lb) and has notably small teeth, is still regarded as a separate species.[39][40][41][42]

    The four smallest raccoon subspecies, with a typical weight of 1.8 to 2.7 kg (4.0 to 6.0 lb), live along the southern coast of Florida and on the adjacent islands; an example is the Ten Thousand Islands raccoon (Procyon lotor marinus).[43] Most of the other 15 subspecies differ only slightly from each other in coat color, size and other physical characteristics.[44][45] The two most widespread subspecies are the eastern raccoon (Procyon lotor lotor) and the Upper Mississippi Valley raccoon (Procyon lotor hirtus). Both share a comparatively dark coat with long hairs, but the Upper Mississippi Valley raccoon is larger than the eastern raccoon. The eastern raccoon occurs in all U.S. states and Canadian provinces to the north of South Carolina and Tennessee. The adjacent range of the Upper Mississippi Valley raccoon covers all U.S. states and Canadian provinces to the north of LouisianaTexas, and New Mexico.[46]

    The taxonomic identity of feral raccoons inhabiting Central Europe, Causasia and Japan is unknown, as the founding populations consisted of uncategorized specimens from zoos and fur farms.[47]

    SubspeciesImageTrinomial authorityDescriptionRangeSynonyms
    Eastern raccoon
    P. l. lotor
    Nominate subspecies
    Linnaeus, 1758A small and dark subspecies with long, soft fur.[48]Nova Scotia, southern New Brunswick, southern Quebec, and southern Ontario south through the eastern United States to North Carolina, and from the Atlantic coast west to Lake Michigan, Indiana, southern Illinois, western Kentucky, and probably eastern Tennessee.annulatus (G. Fischer, 1814)
    brachyurus (Wiegmann, 1837)
    fusca (Burmeister, 1850)
    gularis (C. E. H. Smith, 1848)
    melanus (J. E. Gray, 1864)
    obscurus (Wiegmann, 1837)
    rufescens (de Beaux, 1910)
    vulgaris (Tiedemann, 1808)
    Key Vaca raccoon
    P. l. auspicatus
    Nelson, 1930A very small and pale-furred subspecies.[49]Key Vaca and doubtless closely adjoining keys of the Key Vaca Group, a central section of the main chain off the southern coast of Florida.
    Florida raccoon
    P. l. elucus
    Bangs, 1898Generally a medium-sized and dark-colored subspecies with a prominent rusty rufous nuchal patch.[50]Peninsular Florida, except southwestern part inhabited by P. l. marinus, north to extreme southern Georgia; grading into P. l. varius in northwest Florida.
    Snake River Valley raccoon
    P. l. excelsus
    Nelson and Goldman, 1930A very large and pale subspecies.[51]Snake River drainage in southeastern Washington, eastern Oregon, and southern Idaho, the Humboldt River Valley, Nev., and river valleys of northeastern California.
    Texas raccoon
    P. l. fuscipes
    Mearns, 1914A large, dark grayish subspecies.[52]Texas, except extreme northern and western parts, southern Arkansas, Louisiana, except delta region of Mississippi, and south into northeastern Mexico, including Coahuila and Nuevo León, to southern Tamaulipas.
    † Barbados raccoon
    P. l. gloveralleni
    Nelson and Goldman, 1930A small, dark-furred subspecies with a lightly built skull.[53]Known only from the Island of Barbados.solutus (Nelson and Goldman, 1931)
    Baja California raccoon
    P. l. grinnelli
    Nelson and Goldman, 1930A large, pale-furred subspecies with high and broad skull.[54]Southern Baja California from the Cape region north at least to San Ignacio.
    Mexican raccoon
    P. l. hernandezii
    Wagler, 1831A large and dark grayish subspecies with a flattish skull and heavy dentition.[55]Mexican mainland (primarily central, western, and southern Mexico), extending southward through Guatemala, Belize, El Salvador, Honduras, Nicaragua, Costa Rica, and northern Panama.[56][42]crassidens (Hollister, 1914)
    dickeyi (Nelson and Goldman, 1931)
    mexicana (Baird, 1858)
    shufeldti (Nelson and Goldman, 1931)
    Upper Mississippi Valley raccoon
    P. l. hirtus
    Nelson and Goldman, 1930A large and dark-furred subspecies, whose pelage is usually suffused with ochraceous buff.[57]Upper Mississippi and Missouri River drainage areas from the eastern slopes of the Rocky Mountains east to Lake Michigan, and from southern Manitoba and probably southwestern Ontario and southeastern Alberta south to southern Oklahoma and Arkansas.
    Torch Key raccoon
    P. l. incautus
    Nelson, 1930A small subspecies with very pale fur (the palest of the Florida raccoons).[58]Big Pine Key Group, near southwestern end of chain of Florida Keys.
    Matecumbe Key raccoon
    P. l. inesperatus
    Nelson, 1930Similar to P. l. elucus, but smaller and grayer and with a flatter skull.[59]Key Largo Group, embracing fringing keys along the southeast coast of Florida, from Virginia Key south to Lower Matecumbe Key.
    Tres Marias raccoon
    P. l. insularis
    Merriam, 1898A large, massive-skulled subspecies with short and coarse fur.[60]Tres Marías Islands, off west coast of Nayarit, Mexico.vicinus (Nelson and Goldman, 1931)
    Saint Simon Island raccoon
    P. l. litoreus
    Nelson and Goldman, 1930Similar to P. l. elucus, being of medium size and having dark fur.[61]Coastal strip and islands of Georgia.
    Ten Thousand Islands raccoon
    P. l. marinus
    Nelson, 1930A very small subspecies with heavy dentition.[62]Keys of the Ten Thousand Islands Group, and adjoining mainland of southwestern Florida from Cape Sable north through the Everglades to Lake Okeechobee.maritimus (Dozier, 1948)
    Bahamian raccoon
    P. l. maynardi
    Bangs, 1898A small and slightly dark subspecies with a lightly built skull and dentition.[63]Known only from New Providence Island, Bahamas.flavidus (de Beaux, 1910)
    minor (Miller, 1911)
    varius (Nelson and Goldman, 1930)
    Mississippi Delta raccoon
    P. l. megalodous
    Lowery, 1943A medium-sized subspecies, with a massive skull and pale yellow fur suffused above with black.[64]Coast region of southern Louisiana from St. Bernard Parish west to Cameron Parish.
    Guadeloupe raccoonP. l. minorMiller, 1911A small subspecies with a delicate skull, a dark gray coat, and a slight ochre tint on the neck and shoulders.[65]Guadeloupe in the Lesser Antilles (both islands: Grande-Terre and Basse-Terre).
    Pacific Northwest raccoon
    P. l. pacificus
    Merriam, 1899A dark-furred subspecies with a relatively broad, flat skull.[66]Southwestern British Columbia, except Vancouver Island, northern, central, and western Washington, western Oregon, and extreme northwestern California.proteus (Brass, 1911)
    Colorado Desert raccoon
    P. l. pallidus
    Merriam, 1900One of the palest subspecies, around the same size as P. l. mexicanus.[67]Colorado and Gila River Valleys and adjoining territory from the delta north to northeastern Utah, and east to western Colorado and northwestern New Mexico.ochraceus (Mearns, 1914)
    California raccoon
    P. l. psora
    Gray, 1842A large and moderately dark subspecies with a broad, rather flat skull.[68]California, except extreme northwest coastal strip, the northeastern corner and southeastern desert region, ranging south through northwestern Baja California to San Quentin; extreme west central Nevada.californicus (Means, 1914)
    Isthmian raccoon
    P. l. pumilus
    Miller, 1911Similar to P. l. crassidens in color, but has a shorter, broader and flatter skull.[69]Panama and the Canal Zone from Porto Bello west to Boqueron, Chiriqui, though the limits of its range are unknown.
    † Short-faced raccoon
    P. l. simus
    Gidley, 1906Pleistocene subspecies similar to P. l. excelsus, but with a deeper lower jaw and a more robust dentition.[70][71]California.
    Vancouver Island raccoon
    P. l. vancouverensis
    Nelson and Goldman, 1930A dark-furred subspecies, similar to P. l. pacificus but smaller.[72]Known only from Vancouver Island.

    Description

    Physical characteristics

    Lower side of front paw with visible vibrissae on the tips of the digits

    Skeleton

    Skull with dentition: 2/2 molars, 4/4 premolars, 1/1 canines, 3/3 incisors

    Baculum or penis bone

    Female genitourinary system

    Male genitourinary system

    Head to hindquarters, raccoons measure between 40 and 70 cm (16 and 28 in), not including the bushy tail which can measure between 20 and 40 cm (7.9 and 15.7 in), but is usually not much longer than 25 cm (9.8 in).[73][74][75] The shoulder height is between 23 and 30 cm (9.1 and 11.8 in).[76] The body weight of an adult raccoon varies considerably with habitat, making the raccoon one of the most variably sized mammals. It can range from 2 to 26 kg (4.4 to 57.3 lb), but is usually between 5 and 12 kg (11 and 26 lb). The smallest specimens live in southern Florida, while those near the northern limits of the raccoon’s range tend to be the largest (see Bergmann’s rule).[77] Males are usually 15 to 20% heavier than females.[78] At the beginning of winter, a raccoon can weigh twice as much as in spring because of fat storage.[79][80][81] The largest recorded wild raccoon weighed 28.4 kg (63 lb) and measured 140 cm (55 in) in total length, by far the largest size recorded for a procyonid.[82][83]

    The most characteristic physical feature of the raccoon is the area of black fur around the eyes, which contrasts sharply with the surrounding white face coloring. This is reminiscent of a “bandit’s mask” and has thus enhanced the animal’s reputation for mischief.[84][85] The slightly rounded ears are also bordered by white fur. Raccoons are assumed to recognize the facial expression and posture of other members of their species more quickly because of the conspicuous facial coloration and the alternating light and dark rings on the tail.[86][87][88] The dark mask may also reduce glare and thus enhance night vision.[87][88] On other parts of the body, the long and stiff guard hairs, which shed moisture, are usually colored in shades of gray and, to a lesser extent, brown.[89] Raccoons with a very dark coat are more common in the German population because individuals with such coloring were among those initially released to the wild.[90] The dense underfur, which accounts for almost 90% of the coat, insulates against cold weather and is composed of 2 to 3 cm (0.79 to 1.18 in) long hairs.[89]

    The raccoon, whose method of locomotion is usually considered to be plantigrade, can stand on its hind legs to examine objects with its front paws.[91][92] As raccoons have short legs compared to their compact torso, they are usually not able either to run quickly or jump great distances.[93][94] Their top speed over short distances is 16 to 24 km/h (9.9 to 14.9 mph).[95][96] Raccoons can swim with an average speed of about 5 km/h (3.1 mph) and can stay in the water for several hours.[97][94] For climbing down a tree headfirst—an unusual ability for a mammal of its size—a raccoon rotates its hind feet so they are pointing backwards.[98][94] Raccoons have a dual cooling system to regulate their temperature; that is, they are able to both sweat and pant for heat dissipation.[99][100]

    Raccoon skulls have a short and wide facial region and a voluminous braincase. The facial length of the skull is less than the cranial, and their nasal bones are short and quite broad. The auditory bullae are inflated in form, and the sagittal crest is weakly developed.[101] The dentition—40 teeth with the dental formula:3.1.4.23.1.4.2—is adapted to their omnivorous diet: the carnassials are not as sharp and pointed as those of a full-time carnivore, but the molars are not as wide as those of a herbivore.[102] The penis bone of males is about 10 cm (3.9 in) long and strongly bent at the front end,[103][104] and its shape can be used to distinguish juvenile males from mature males.[105][106][107] Seven of the thirteen identified vocal calls are used in communication between the mother and her kits, one of these being the birdlike twittering of newborns.[108][109][100]

    Senses

    The most important sense for the raccoon is its sense of touch.[110][111][112] The “hyper sensitive”[111] front paws are protected by a thin horny layer that becomes pliable when wet.[113][114] The five digits of the paws have no webbing between them, which is unusual for a carnivoran.[115] Almost two-thirds of the area responsible for sensory perception in the raccoon’s cerebral cortex is specialized for the interpretation of tactile impulses, more than in any other studied animal.[116] They are able to identify objects before touching them with vibrissae located above their sharp, nonretractable claws.[91][112] The raccoon’s paws lack an opposable thumb; thus, it does not have the agility of the hands of primates.[112][114] There is no observed negative effect on tactile perception when a raccoon stands in water below 10 °C (50 °F) for hours.[117]

    Raccoons are thought to be color blind or at least poorly able to distinguish color, though their eyes are well-adapted for sensing green light.[118][119][120] Although their accommodation of 11 dioptre is comparable to that of humans and they see well in twilight because of the tapetum lucidum behind the retinavisual perception is of subordinate importance to raccoons because of their poor long-distance vision.[121][122][123] In addition to being useful for orientation in the dark, their sense of smell is important for intraspecific communication. Glandular secretions (usually from their anal glands), urine and feces are used for marking.[124][125][126] With their broad auditory range, they can perceive tones up to 50–85 kHz as well as quiet noises, like those produced by earthworms underground.[127][128]

    Intelligence

    Zoologist Clinton Hart Merriam described raccoons as “clever beasts”, and that “in certain directions their cunning surpasses that of the fox”. The animal’s intelligence gave rise to the epithet “sly coon”.[129] Only a few studies have been undertaken to determine the mental abilities of raccoons, most of them based on the animal’s sense of touch. In a study by the ethologist H. B. Davis in 1908, raccoons were able to open 11 of 13 complex locks in fewer than 10 tries and had no problems repeating the action when the locks were rearranged or turned upside down. Davis concluded that they understood the abstract principles of the locking mechanisms and their learning speed was equivalent to that of rhesus macaques.[130]

    Studies in 1963, 1973, 1975 and 1992 concentrated on raccoon memory showed that they can remember the solutions to tasks for at least three years.[131] In a study by B. Pohl in 1992, raccoons were able to instantly differentiate between identical and different symbols three years after the short initial learning phase.[131] Stanislas Dehaene reports in his book The Number Sense that raccoons can distinguish boxes containing two or four grapes from those containing three.[132] In research by Suzana Herculano-Houzel and other neuroscientists, raccoons have been found to be comparable to primates in density of neurons in the cerebral cortex, which they have proposed to be a neuroanatomical indicator of intelligence.[133][134]

    Behavior

    Social behavior

    Eastern raccoons (P. l. lotor) in a tree: The raccoon’s social structure is grouped into what Ulf Hohmann calls a “three-class society”.
    California raccoon (P. l. psora) climbing a tree in Lower Klamath National Wildlife Refuge

    Baby raccoon chatter

    Duration: 1 second.0:01


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    Studies in the 1990s by the ethologists Stanley D. Gehrt and Ulf Hohmann suggest that raccoons engage in sex-specific social behaviors and are not typically solitary, as was previously thought.[135][136] Related females often live in a so-called “fission-fusion society“; that is, they share a common area and occasionally meet at feeding or resting grounds.[137][138] Unrelated males often form loose male social groups to maintain their position against foreign males during the mating season—or against other potential invaders.[139] Such a group does not usually consist of more than four individuals.[140][141] Since some males show aggressive behavior towards unrelated kits, mothers will isolate themselves from other raccoons until their kits are big enough to defend themselves.[142]

    With respect to these three different modes of life prevalent among raccoons, Hohmann called their social structure a “three-class society”.[143] Samuel I. Zeveloff, professor of zoology at Weber State University and author of the book Raccoons: A Natural History, is more cautious in his interpretation and concludes at least the females are solitary most of the time and, according to Erik K. Fritzell’s study in North Dakota in 1978, males in areas with low population densities are solitary as well.[144]

    The shape and size of a raccoon’s home range varies depending on age, sex, and habitat, with adults claiming areas more than twice as large as juveniles.[145] While the size of home ranges in the habitat of North Dakota’s prairies lie between 7 and 50 km2 (3 and 20 sq mi) for males and between 2 and 16 km2 (1 and 6 sq mi) for females, the average size in a marsh at Lake Erie was 0.5 km2 (0.19 sq mi).[146] Irrespective of whether the home ranges of adjacent groups overlap, they are most likely not actively defended outside the mating season if food supplies are sufficient.[147] Odor marks on prominent spots are assumed to establish home ranges and identify individuals.[126] Urine and feces left at shared raccoon latrines may provide additional information about feeding grounds, since raccoons were observed to meet there later for collective eating, sleeping and playing.[148]

    Concerning the general behavior patterns of raccoons, Gehrt points out that “typically you’ll find 10 to 15 percent that will do the opposite” of what is expected.[149]

    Diet

    Though usually nocturnal, the raccoon is sometimes active in daylight to take advantage of available food sources.[150][151] Its diet consists of about 40% invertebrates, 33% plant material and 27% vertebrates.[152] Since its diet consists of such a variety of different foods, Zeveloff argues the raccoon “may well be one of the world’s most omnivorous animals”.[153] While its diet in spring and early summer consists mostly of insects, worms, and other animals already available early in the year, it prefers fruits and nuts, such as acorns and walnuts, which emerge in late summer and autumn, and represent a rich calorie source for building up fat needed for winter.[154][155]

    Contrary to popular belief, raccoons only occasionally eat active or large prey, such as birds and mammals. They prefer prey that is easier to catch, specifically crayfishinsects,[156] fishamphibians and bird eggs.[157] Raccoons are virulent predators of eggs and hatchlings in both birds and reptile nests, to such a degree that, for threatened prey species, raccoons may need to be removed from the area or nests may need to be relocated to mitigate the effect of their predations (i.e. in the case of some globally threatened turtles).[158][159][160][161][162] When food is plentiful, raccoons can develop strong individual preferences for specific foods.[80] In the northern parts of their range, raccoons go into a winter rest, reducing their activity drastically as long as a permanent snow cover makes searching for food difficult.[163]

    Dousing

    Captive raccoons often douse their food before eating.

    One aspect of raccoon behavior is so well known that it gives the animal part of its scientific name, Procyon lotorlotor is Latin for ‘washer’. In the wild, raccoons often dabble for underwater food near the shore-line. They then often pick up the food item with their front paws to examine it and rub the item, sometimes to remove unwanted parts. This gives the appearance of the raccoon “washing” the food. The tactile sensitivity of raccoons’ paws is increased if this rubbing action is performed underwater, since the water softens the hard layer covering the paws.[111][164] However, the behavior observed in captive raccoons in which they carry their food to water to “wash” or douse it before eating has not been observed in the wild.[165][166] Naturalist Georges-Louis Leclerc, Comte de Buffon, believed that raccoons do not have adequate saliva production to moisten food thereby necessitating dousing, but this hypothesis is now considered to be incorrect.[164][165][167][168] Captive raccoons douse their food more frequently when a watering hole with a layout similar to a stream is not farther away than 3 m (10 ft).[168] The widely accepted theory is that dousing in captive raccoons is a fixed action pattern from the dabbling behavior performed when foraging at shores for aquatic foods.[164][168][169][170] This is supported by the observation that aquatic foods are doused more frequently. Cleaning dirty food does not seem to be a reason for “washing”.[168]

    Reproduction

    Raccoons usually mate in a period triggered by increasing daylight between late January and mid-March.[171][172][173] However, there are large regional differences which are not completely explicable by solar conditions. For example, while raccoons in southern states typically mate later than average, the mating season in Manitoba also peaks later than usual in March and extends until June.[173] During the mating season, males restlessly roam their home ranges in search of females in an attempt to court them during the three- to four-day period when conception is possible. These encounters will often occur at central meeting places.[174][175][176] Copulation, including foreplay, can last over an hour and is repeated over several nights.[177] The weaker members of a male social group also are assumed to get the opportunity to mate, since the stronger ones cannot mate with all available females.[178] In a study in southern Texas during the mating seasons from 1990 to 1992, about one third of all females mated with more than one male.[179] If a female does not become pregnant or if she loses her kits early, she will sometimes become fertile again 80 to 140 days later.[180][181][182]

    An eastern raccoon (P. l. lotor) kit

    After usually 63 to 65 days of gestation (although anywhere from 54 to 70 days is possible), a litter of typically two to five young is born.[183][184] The average litter size varies widely with habitat, ranging from 2.5 in Alabama to 4.8 in North Dakota.[185][186] Larger litters are more common in areas with a high mortality rate, due, for example, to hunting or severe winters.[187][186] While male yearlings usually reach their sexual maturity only after the main mating season, female yearlings can compensate for high mortality rates and may be responsible for about 50% of all young born in a year.[188][189][190] Males have no part in raising young.[140][191][192] The kits (also called “cubs”) are blind and deaf at birth, but their mask is already visible against their light fur.[193][194] The birth weight of the roughly 10 cm (4 in)-long kits is between 60 and 75 g (2.1 and 2.6 oz).[194] Their ear canals open after around 18 to 23 days, a few days before their eyes open for the first time.[195] Once the kits weigh about 1 kg (2 lb), they begin to explore outside the den, consuming solid food for the first time after six to nine weeks.[196][197] After this point, their mother suckles them with decreasing frequency; they are usually weaned by 16 weeks.[198] In the fall, after their mother has shown them dens and feeding grounds, the juvenile group splits up.[199][200] While many females will stay close to the home range of their mother, males can sometimes move more than 20 km (12 mi) away.[201][202] This is considered an instinctive behavior, preventing inbreeding.[203][204] However, mother and offspring may share a den during the first winter in cold areas.[200]

    Life expectancy

    Young Florida raccoon (P. l. elucus) crossing a road

    Captive raccoons have been known to live for more than 20 years.[84] However, the species’ life expectancy in the wild is only 1.8 to 3.1 years, depending on the local conditions such as traffic volume, hunting, and weather severity.[205] It is not unusual for only half of the young born in one year to survive a full year.[188][206] After this point, the annual mortality rate drops to between 10% and 30%.[188] Young raccoons are vulnerable to losing their mother and to starvation, particularly in long and cold winters.[207] The most frequent natural cause of death in the North American raccoon population is distemper, which can reach epidemic proportions and kill most of a local raccoon population.[208] In areas with heavy vehicular traffic and extensive hunting, these factors can account for up to 90% of all deaths of adult raccoons.[209] The most important natural predators of the raccoon are bobcatscoyotes, and great horned owls, the latter mainly preying on young raccoons but capable of killing adults in some cases.[210][211][212][213][214][215] In Florida, they have been reported to fall victim to larger carnivores like American black bear and cougars and these species may also be a threat on occasion in other areas.[216][217][218] Where still present, gray wolves may still occasionally take raccoons as a supplemental prey item.[219][220] Also in the southeast, they are among the favored prey for adult American alligators.[221][222] On occasion, both bald and golden eagles will prey on raccoons.[223][224] In the tropics, raccoons are known to fall prey to smaller eagles such as ornate hawk-eagles and black hawk-eagles, although it is not clear whether adults or merely juvenile raccoons are taken by these.[225][226] In rare cases of overlap, they may fall victim from carnivores ranging from species averaging smaller than themselves such as fishers to those as large and formidable as jaguars in Mexico.[227][228] In their introduced range in the former Soviet Union, their main predators are wolveslynxes and Eurasian eagle-owls.[229] However, predation is not a significant cause of death, especially because larger predators have been exterminated in many areas inhabited by raccoons.[230]

    Range

    Mississippi Delta raccoon (P. l. megaloudus) searching for food on a lake shore

    Habitat

    Although they have thrived in sparsely wooded areas in the last decades, raccoons depend on vertical structures to climb when they feel threatened.[231][232] Therefore, they avoid open terrain and areas with high concentrations of beech trees, as beech bark is too smooth to climb.[233] Tree hollows in old oaks or other trees and rock crevices are preferred by raccoons as sleeping, winter and litter dens. If such dens are unavailable or accessing them is inconvenient, raccoons use burrows dug by other mammals, dense undergrowth or tree crotches.[234][235] In a study in the Solling range of hills in Germany, more than 60% of all sleeping places were used only once, but those used at least ten times accounted for about 70% of all uses.[236] Since amphibians, crustaceans, and other animals around the shore of lakes and rivers are an important part of the raccoon’s diet, lowland deciduous or mixed forests abundant with water and marshes sustain the highest population densities.[237][238] While population densities range from 0.5 to 3.2 animals per square kilometer (1.3 to 8.3 animals per square mile) in prairies and do not usually exceed 6 animals per square kilometer (15.5 animals per square mile) in upland hardwood forests, more than 20 raccoons per square kilometer (51.8 animals per square mile) can live in lowland forests and marshes.[237][239]

    Distribution in North America

    An albino Florida raccoon (P. l. elucus) in Virginia Key, Florida

    Raccoons are common throughout North America from Canada to Panama, where the subspecies Procyon lotor pumilus coexists with the crab-eating raccoon (Procyon cancrivorus).[240][241] The population on Hispaniola was exterminated as early as 1513 by Spanish colonists who hunted them for their meat.[242] Raccoons were also exterminated in Cuba and Jamaica, where the last sightings were reported in 1687.[243] The Barbados raccoon became extinct relatively recently, in 1964. When they were still considered separate species, the Bahamas raccoon, Guadeloupe raccoon and Tres Marias raccoon were classified as endangered by the IUCN in 1996.[244]

    A Raccoon sleeping on a tree in High ParkToronto

    There is archeological evidence that in pre-Columbian times raccoons were numerous only along rivers and in the woodlands of the Southeastern United States.[245] As raccoons were not mentioned in earlier reports of pioneers exploring the central and north-central parts of the United States,[246] their initial spread may have begun a few decades before the 20th century. Since the 1950s, raccoons have expanded their range from Vancouver Island—formerly the northernmost limit of their range—far into the northern portions of the four south-central Canadian provinces.[247] New habitats which have recently been occupied by raccoons (aside from urban areas) include mountain ranges, such as the Western Rocky Mountains, prairies and coastal marshes.[248] After a population explosion starting in the 1940s, the estimated number of raccoons in North America in the late 1980s was 15 to 20 times higher than in the 1930s, when raccoons were comparatively rare.[249] Urbanization, the expansion of agriculture, deliberate introductions, and the extermination of natural predators of the raccoon have probably caused this increase in abundance and distribution.[250]

    Distribution outside North America

    Distribution in Germany: Raccoons killed or found dead by hunters in the hunting years 2000–2001, 2001–2002 and 2002–2003 in the administrative districts of Germany

    As a result of escapes and deliberate introductions in the mid-20th century, the raccoon is now distributed in several European and Asian countries. Sightings have occurred in all the countries bordering Germany, which hosts the largest population outside of North America.[251] Another stable population exists in northern France, where several pet raccoons were released by members of the U.S. Air Force near the Laon-Couvron Air Base in 1966.[252] Furthermore, raccoons have been known to be in the area around Madrid since the early 1970s. In 2013, the city authorized “the capture and death of any specimen”.[253] It is also present in Italy, with one self-sustaining population in Lombardy.[254]

    About 1,240 animals were released in nine regions of the former Soviet Union between 1936 and 1958 for the purpose of establishing a population to be hunted for their fur. Two of these introductions were successful – one in the south of Belarus between 1954 and 1958, and another in Azerbaijan between 1941 and 1957. With a seasonal harvest of between 1,000~1,500 animals, in 1974 the estimated size of the population distributed in the Caucasus region was around 20,000 animals and the density was four animals per square kilometer (10 animals per square mile).[255]

    Distribution in Japan

    In Japan, up to 1,500 raccoons were imported as pets each year after the success of the anime series Rascal the Raccoon (1977). In 2004, the descendants of discarded or escaped animals lived in 42 of 47 prefectures.[256][257][258] The range of raccoons in the wild in Japan grew from 17 prefectures in 2000 to all 47 prefectures in 2008.[259] It is estimated that raccoons cause thirty million yen (~$275,000) of agricultural damage on Hokkaido alone.[260]

    Distribution in Germany

    In Germany – where the raccoon is called the Waschbär (literally, ‘wash-bear’ or ‘washing bear’) due to its habit of “dousing” food in water – two pairs of pet raccoons were released into the German countryside at the Edersee reservoir in the north of Hesse in April 1934 by a forester upon request of their owner, a poultry farmer.[261] He released them two weeks before receiving permission from the Prussian hunting office to “enrich the fauna“.[262] Several prior attempts to introduce raccoons in Germany had been unsuccessful.[263][264] A second population was established in eastern Germany in 1945 when 25 raccoons escaped from a fur farm at Wolfshagen (today district of Altlandsberg), east of Berlin, after an air strike. The two populations are parasitologically distinguishable: 70% of the raccoons of the Hessian population are infected with the roundworm Baylisascaris procyonis, but none of the Brandenburgian population is known to have the parasite.[265] In the Hessian region, there were an estimated 285 raccoons in 1956, which increased to over 20,000 in 1970; in 2008 there were between 200,000 and 400,000 raccoons in the whole of Germany.[208][263] By 2012 it was estimated that Germany now had more than a million raccoons.[266]

    The raccoon was once a protected species in Germany, but has been declared a game animal in 14 of the 16 German states since 1954.[267] Hunters and environmentalists argue the raccoon spreads uncontrollably, threatens protected bird species, and supersedes indigenous competitors.[90] This view is opposed by the zoologist Frank-Uwe Michler, who finds no evidence that a high population density of raccoons leads to negative effects on the biodiversity of an area.[90] Hohmann holds that extensive hunting cannot be justified by the absence of natural predators, because predation is not a significant cause of death in the North American raccoon population.[268]

    Graphs are unavailable due to technical issues. Updates on reimplementing the Graph extension, which will be known as the Chart extension, can be found on Phabricator and on MediaWiki.org.

    The raccoon is extensively hunted in Germany as it is seen as an invasive species and pest.[269][270] In the 1990s, only about 400 raccoons were hunted yearly. This increased dramatically over the next quarter-century: during the 2015–2016 hunting season, 128,100 raccoons were hunted, 60 percent of them in the state of Hesse.[271]

    Distribution in the former Soviet Union

    Experiments in acclimatising raccoons into the Soviet Union began in 1936, and were repeated a further 25 times until 1962. Overall, 1,222 individuals were released, 64 of which came from zoos and fur farms (38 of them having been imports from western Europe). The remainder originated from a population previously established in Transcaucasia. The range of Soviet raccoons was never single or continuous, as they were often introduced to different locations far from each other. All introductions into the Russian Far East failed; melanistic raccoons were released on Petrov Island near Vladivostok and some areas of southern Primorsky Krai, but died. In Central Asia, raccoons were released in Kyrgyzstan‘s Jalal-Abad Province, though they were later recorded as “practically absent” there in January 1963. A large and stable raccoon population (yielding 1,000~1,500 catches a year) was established in Azerbaijan after an introduction to the area in 1937. Raccoons apparently survived an introduction near Terek, along the Sulak River into the Dagestani lowlands. Attempts to settle raccoons on the Kuban River‘s left tributary and Kabardino-Balkaria were unsuccessful. A successful acclimatization occurred in Belarus, where three introductions (consisting of 52, 37, and 38 individuals in 1954 and 1958) took place. By January 1963, 700 individuals were recorded in the country.[272]

    Urban raccoons

    On the roof of a house in Albertshausen, Germany

    Due to its adaptability, the raccoon has been able to use urban areas as a habitat. The first sightings were recorded in a suburb of Cincinnati in the 1920s. Since the 1950s, raccoons have been present in metropolitan areas like Washington, DCChicagoToronto, and New York City.[273][274] Since the 1960s, Kassel has hosted Europe’s first and densest population in a large urban area, with about 50 to 150 animals per square kilometer (130 to 390 animals per square mile), a figure comparable to those of urban habitats in North America.[273][275] Home range sizes of urban raccoons are only 3 to 40 hectares (7.5 to 100 acres) for females and 8 to 80 hectares (20 to 200 acres) for males.[276] In small towns and suburbs, many raccoons sleep in a nearby forest after foraging in the settlement area.[273][277] Fruit and insects in gardens and leftovers in municipal waste are easily available food sources.[278] Furthermore, a large number of additional sleeping areas exist in these areas, such as hollows in old garden trees, cottages, garages, abandoned houses, and attics. The percentage of urban raccoons sleeping in abandoned or occupied houses varies from 15% in Washington, DC (1991) to 43% in Kassel (2003).[277][275]

    Health

    Raccoon roundworm Baylisascaris procyonis larvae

    Raccoons can carry rabies, a lethal disease caused by the neurotropic rabies virus carried in the saliva and transmitted by bites. Its spread began in Florida and Georgia in the 1950s and was facilitated by the introduction of infected individuals to Virginia and North Dakota in the late 1970s.[279] Of the 6,940 documented rabies cases reported in the United States in 2006, 2,615 (37.7%) were in raccoons.[280] The U.S. Department of Agriculture, as well as local authorities in several U.S. states and Canadian provinces, has developed oral vaccination programs to fight the spread of the disease in endangered populations.[281][282][283] Only one human fatality has been reported after transmission of the rabies virus strain commonly known as “raccoon rabies”.[284] Among the main symptoms for rabies in raccoons are a generally sickly appearance, impaired mobility, abnormal vocalization, and aggressiveness.[285] There may be no visible signs at all, however, and most individuals do not show the aggressive behavior seen in infected canids; rabid raccoons will often retire to their dens instead.[90][265][285] Organizations like the U.S. Forest Service encourage people to stay away from animals with unusual behavior or appearance, and to notify the proper authorities, such as an animal control officer from the local health department.[286][287] Since healthy animals, especially nursing mothers, will occasionally forage during the day, daylight activity is not a reliable indicator of illness in raccoons.[150][151]

    Unlike rabies and at least a dozen other pathogens carried by raccoons, distemper, an epizootic virus, does not affect humans.[288][289] This disease is the most frequent natural cause of death in the North American raccoon population and affects individuals of all age groups.[208] For example, 94 of 145 raccoons died during an outbreak in Clifton, Ohio, in 1968.[290] It may occur along with a following inflammation of the brain (encephalitis), causing the animal to display rabies-like symptoms.[279] In Germany, the first eight cases of distemper were reported in 2007.[208]

    Some of the most important bacterial diseases which affect raccoons are leptospirosislisteriosistetanus, and tularemia. Although internal parasites weaken their immune systems, well-fed individuals can carry a great many roundworms in their digestive tracts without showing symptoms.[291][289] The larvae of the roundworm Baylisascaris procyonis, which can be contained in the feces and seldom causes a severe illness in humans, can be ingested when cleaning raccoon latrines without wearing breathing protection.[292]

    While not endemic, the worm Trichinella does infect raccoons,[293] and undercooked raccoon meat has caused trichinosis in humans.[294]

    Trematode Metorchis conjunctus can also infect raccoons.[295]

    Relationship with humans

    Conflicts

    skunk and a California raccoon (P. l. psora) share cat food morsels in a Hollywood, California backyard

    Raccoons have become notorious in urban areas for consuming food waste. They possess impressive problem-solving abilities and can break into all but the most secure food waste bins, which has earned them the derisive nickname trash panda.[23] The presence of raccoons in close proximity to humans may be undesirable, as raccoon droppings (like most wild animals) contain parasites and other disease vectors. Raccoon roundworm is of particular concern to public health. It can be contracted in humans by accidental ingestion or inhalation of the eggs, which are present in the feces of infected raccoons. While usually harmless to the host, it causes progressive neurological damage in humans, and is eventually fatal if untreated. It is found in about 60% of adult raccoons. The general presence of raccoons in an area is not typically of concern, but nests or droppings found within or near structures should be destroyed. Roundworm eggs are very robust and bleach alone is insufficient; burning or treatment with hot solutions of sodium hydroxide is required. The keeping of raccoons as pets is illegal in some jurisdictions due to these risks.[296][297]

    The increasing number of raccoons in urban areas has resulted in diverse reactions in humans, ranging from outrage at their presence to deliberate feeding.[298] Some wildlife experts and most public authorities caution against feeding wild animals because they might become increasingly obtrusive and dependent on humans as a food source.[299] Other experts challenge such arguments and give advice on feeding raccoons and other wildlife in their books.[300][301] Raccoons without a fear of humans are a concern to those who attribute this trait to rabies, but scientists point out this behavior is much more likely to be a behavioral adjustment to living in habitats with regular contact to humans for many generations.[265][302] Raccoons usually do not prey on domestic cats and dogs, but isolated cases of killings have been reported.[303] Attacks on pets may also target their owners.[304]

    A Florida raccoon (P. l. elucus) in the Everglades approaches a group of humans, hoping to be fed

    While overturned waste containers and raided fruit trees are just a nuisance to homeowners, it can cost several thousand dollars to repair damage caused by the use of attic space as dens.[305] Relocating or killing raccoons without a permit is forbidden in many urban areas on grounds of animal welfare. These methods usually only solve problems with particularly wild or aggressive individuals, since adequate dens are either known to several raccoons or will quickly be rediscovered.[187][287][306] Loud noises, flashing lights, and unpleasant odors have proven particularly effective in driving away a mother and her kits before they would normally leave the nesting place (when the kits are about eight weeks old).[287][307] Typically, though, only precautionary measures to restrict access to food waste and den sites are effective in the long term.[287][308][309]

    Among all fruits and crops cultivated in agricultural areas, sweet corn in its milk stage is particularly popular among raccoons.[310][311] In a two-year study by Purdue University researchers, published in 2004, raccoons were responsible for 87% of the damage to corn plants.[312] Like other predators, raccoons searching for food can break into poultry houses to feed on chickens, ducks, their eggs, or food.[152][287][313]

    Mythology, arts, and entertainment

    See also: List of fictional raccoons

    Stylized raccoon skin as depicted on the Raccoon Priests gorget found at Spiro Mounds

    In the mythology of the Indigenous peoples of the Americas, the raccoon is the subject of folk tales.[314] Stories such as “How raccoons catch so many crayfish” from the Tuscarora centered on its skills at foraging.[315] In other tales, the raccoon played the role of the trickster which outsmarts other animals, like coyotes and wolves.[316] Among others, the Dakota believe the raccoon has natural spirit powers, since its mask resembles the facial paintings, two-fingered swashes of black and white, used during rituals to connect to spirit beings.[317] The Aztecs linked supernatural abilities especially to females, whose commitment to their young was associated with the role of wise women in their society.[318]

    The raccoon also appears in Native American art across a wide geographic range. Petroglyphs with engraved raccoon tracks were found in Lewis Canyon, Texas;[319] at the Crow Hollow petroglyph site in Grayson County, Kentucky;[320] and in river drainages near Tularosa, the San Francisco River of New Mexico and Arizona.[321] The meaning and significance of the Raccoon Priests Gorget, which features a stylized carving of a raccoon and was found at the Spiro Mounds, Oklahoma, remains unknown.[322][323]

    Hunting and fur trade

    Automobile coat made out of raccoon fur (1906, U.S.)
    Pen with climbing facilities, hiding places, and a watering hole (lower-left-side)

    The fur of raccoons is used for clothing, especially for coats and coonskin caps. At present, it is the material used for the inaccurately named “sealskin” cap worn by the Royal Fusiliers of Great Britain.[324] Sporrans made of raccoon pelt and hide have sometimes been used as part of traditional Scottish highland men’s apparel since the 18th century, especially in North America. Such sporrans may or may not be of the “full-mask” type.[325] Historically, Native American tribes not only used the fur for winter clothing, but also used the tails for ornament.[326] The famous Sioux leader Spotted Tail took his name from a raccoon skin hat with the tail attached he acquired from a fur trader. Since the late 18th century, various types of scent hounds, called coonhounds, which are able to tree animals have been bred in the United States.[327] In the 19th century, when coonskins occasionally even served as means of payment, several thousand raccoons were killed each year in the United States.[328][329] This number rose quickly when automobile coats became popular after the turn of the 20th century. In the 1920s, wearing a raccoon coat was regarded as status symbol among college students.[330] Attempts to breed raccoons in fur farms in the 1920s and 1930s in North America and Europe turned out not to be profitable, and farming was abandoned after prices for long-haired pelts dropped in the 1940s.[331][332] Although raccoons had become rare in the 1930s, at least 388,000 were killed during the hunting season of 1934–1935.[330][333]

    After persistent population increases began in the 1940s, the seasonal coon hunting harvest reached about one million animals in 1946–1947 and two million in 1962–1963.[334] The broadcast of three television episodes about the frontiersman Davy Crockett and the film Davy Crockett, King of the Wild Frontier in 1954 and 1955 led to a high demand for coonskin caps in the United States, although it is unlikely either Crockett or the actor who played him, Fess Parker, actually wore a cap made from raccoon fur.[335] The seasonal hunt reached an all-time high with 5.2 million animals in 1976–1977 and ranged between 3.2 and 4.7 million for most of the 1980s. In 1982, the average pelt price was $20.[336] As of 1987, the raccoon was identified as the most important wild furbearer in North America in terms of revenue.[337] In the first half of the 1990s, the seasonal hunt dropped to 0.9 from 1.9 million due to decreasing pelt prices.[338]

    Food

    While primarily hunted for their fur, raccoons were also a source of food for Native Americans and early American settlers.[339][340] According to Ernest Thompson Seton, young specimens killed without a fight are palatable, whereas old raccoons caught after a lengthy battle are inedible.[341] Raccoon meat was extensively eaten during the early years of California, where it was sold in the San Francisco market for $1–3 apiece.[342] American slaves occasionally ate raccoon at Christmas, but it was not necessarily a dish of the poor or rural. The first edition of The Joy of Cooking, released in 1931, contained a recipe for preparing raccoon, and US President Calvin Coolidge‘s pet raccoon Rebecca was originally sent to be served at the White House Thanksgiving Dinner.[343][344][345]

    Although the idea of eating raccoons may seem repulsive to most mainstream consumers, who see them as endearing, cute, or vermin, several thousand raccoons are still eaten each year in the United States, primarily in the Southern United States.[346][347][348][349] Some people tout the taste of the meat.[350]

    Other uses

    In addition to the fur and meat, the raccoon baculum (penis bone) have had numerous traditional uses in the Southern United States and beyond. Indigenous people used the bones as a pipe cleaning tool.[351] The bones were used by moonshine distillers to guide the flow of whiskey from the drip tube to the bottle.[352] With their tips filed down, the bones were used as toothpicks under the moniker “coon rods”.[a][353] In hoodoo, the folk magic of the American South, the baculum is sometimes worn as an amulet for love or luck.[354] The bones also have decorative uses[355] (e.g. on the trademark hat of stock car racer Richard Petty or as earrings by actresses Sarah Jessica Parker[354] and Vanessa Williams[354]).

    Pet raccoons

    Raccoons are sometimes kept as pets, which is discouraged by many experts because the raccoon is not a domesticated species. Raccoons may act unpredictably and aggressively and it is extremely difficult to teach them to obey commands.[356][357] In places where keeping raccoons as pets is not forbidden, such as in Wisconsin and other U.S. states, an exotic pet permit may be required.[358][359] One notable pet raccoon was Rebecca, kept by US president Calvin Coolidge.[360]

    Their propensity for unruly behavior exceeds that of captive skunks, and they are even less trustworthy when allowed to roam freely. Because of their intelligence and nimble forelimbs, even inexperienced raccoons are easily capable of unscrewing jars, uncorking bottles and opening door latches, with more experienced specimens having been recorded to open door knobs.[129] Sexually mature raccoons often show aggressive natural behaviors such as biting during the mating season.[356][361] Neutering them at around five or six months of age decreases the chances of aggressive behavior developing.[362] Raccoons can become obese and suffer from other disorders due to poor diet and lack of exercise.[363] When fed with cat food over a long time period, raccoons can develop gout.[364] With respect to the research results regarding their social behavior, it is now required by law in Austria and Germany to keep at least two individuals to prevent loneliness.[365][366] Raccoons are usually kept in a pen (indoor or outdoor), also a legal requirement in Austria and Germany, rather than in the apartment where their natural curiosity may result in damage to property.[365][366][356][367][368]

    When orphaned, it is possible for kits to be rehabilitated and reintroduced to the wild. However, it is uncertain whether they readapt well to life in the wild.[369] Feeding unweaned kits with cow’s milk rather than a kitten replacement milk or a similar product can be dangerous to their health.[356][370]

    Local and indigenous names

    Linguistic group or areaLocal / Indigenous name
    Canadian FrenchRaton laveur[371]
    ChocoTouaru[372]
    CocopahNYmaṣ[373]
    Creeᐱᓯᐢᑫᐢ (pisiskês)
    ᐅᒋᑯᒼᓯᐢ (ocikomsis)[374]
    Cree (Swampy and Woods)kimociškw-[375]
    CreekWot•ko[376]
    Lakotawičhítegleǧawičhá[377]
    LenapeNahënëm[378]
    Louisiana FrenchChaoui[379]
    MayanCulu[372]
    MiskitoSuksuk[372]
    NahuatlMapachitli[14]
    NavajoTábąąh mą’ii[380]
    Nez PerceK’ayk’áyoc[381]
    Oglala LakotaWee’-cha[382]
    OjibweEsiban[383]
    Omaha–Poncamiká[384]
    PawneeIcat[385]
    SpanishMapache[372]
    Mapachín[372]
    Tejón[372]
    Gato manglatero[372]
    WintuQari•lit[386]
    Yankton DakotaWay-atch-a

  • Cougar

    The cougar (Puma concolor) (/ˈkuːɡər/KOO-gər), sometimes called the mountain lioncatamountpuma, or panther is a large small cat native to the Americas. It inhabits NorthCentral and South America, making it the most widely distributed wild, terrestrial mammal in the Western Hemisphere, and one of the most widespread in the world. Its range spans the YukonBritish Columbia and Alberta provinces of Canada, the Rocky Mountains and areas in the western United States. Further south, its range extends through Mexico to the Amazon Rainforest and the southern Andes Mountains in Patagonia. It is an adaptable generalist species, occurring in most American habitat types. It prefers habitats with dense underbrush and rocky areas for stalking but also lives in open areas.

    The cougar is largely solitary. Its activity pattern varies from diurnality and cathemerality to crepuscularity and nocturnality between protected and non-protected areas, and is apparently correlated with the presence of other predators, prey species, livestock and humans. It is an ambush predator that pursues a wide variety of prey. Ungulates, particularly deer, are its primary prey, but it also hunts rodents. It is territorial and lives at low population densities. Individual home ranges depend on terrain, vegetation and abundance of prey. While large, it is not always the dominant apex predator in its range, yielding prey to other predators. It is reclusive and mostly avoids people. Fatal attacks on humans are rare but increased in North America as more people entered cougar habitat and built farms.

    The cougar is listed as Least Concern on the IUCN Red List. Intensive hunting following European colonization of the Americas and ongoing human development into cougar habitat has caused populations to decline in most parts of its historical range. In particular, the eastern cougar population is considered to be mostly locally extinct in eastern North America since the early 20th century, with the exception of the isolated Florida panther subpopulation.

    Naming and etymology

    Look up cougar in Wiktionary, the free dictionary.

    The cougar holds the Guinness record for the animal with the greatest number of names, with over 40 in English alone.[3] The word cougar is borrowed from the Portuguese çuçuarana, via French; it was originally derived from the Tupi language. A current form in Brazil is suçuarana.[4] In the 17th century, Georg Marcgrave named it cuguacu ara. Marcgrave’s rendering was reproduced in 1648 by his associate Willem PisoCuguacu ara was then adopted by John Ray in 1693.[5] In 1774, Georges-Louis Leclerc, Comte de Buffon converted cuguacu ara to cuguar, which was later modified to “cougar” in English.[6][7]

    The name puma is the common name used in Latin America and most parts of Europe and is occasionally used in the United States.[8][9][10][11] The first use of puma in English dates to 1777, introduced from Spanish from the Quechua language.[12]

    In the western United States and Canada, the name mountain lion is commonly used, first seen in writing in 1858.[13] The name catamount, a shortening of name “cat of the mountain”, has also been in English use for the cougar and other wild cats since at least 1664.[14] “Panther” is often used synonymously with cougarpuma or mountain lion.[15] The name painter is also sometimes used instead of panther, mostly in the southern United States.[16]

    Taxonomy and evolution

    Felis concolor was the scientific name proposed by Carl Linnaeus in 1771 for a cat with a long tail from Brazil.[17] It was placed in the genus Puma by William Jardine in 1834.[18] This genus is part of the Felinae.[2] The cougar is most closely related to the jaguarundi and the cheetah.[19][20]

    Subspecies

    South American cougar at Torres del Paine National Park, in the Chilean part of Patagonia

    Following Linnaeus’s first scientific description of the cougar, 32 cougar zoological specimens were described and proposed as subspecies until the late 1980s. Genetic analysis of cougar mitochondrial DNA indicates that many of these are too similar to be recognized as distinct at a molecular level but that only six phylogeographic groups exist. The Florida panther samples showed a low microsatellite variation, possibly due to inbreeding.[20] Following this research, the authors of Mammal Species of the World recognized the following six subspecies in 2005:[2]

    • P. c. concolor (Linnaeus, 1771) includes the synonyms bangsiincarumosgoodisoasoarannasussuaranasoderstromiisuçuaçuara, and wavula
    • P. c. puma (Molina, 1782) includes the synonyms araucanusconcolorpatagonicapearsoni, and puma (Trouessart, 1904)
    • P. c. couguar (Kerr, 1792) includes arundivagaaztecusbrownicalifornicafloridanahippolestesimprocerakaibabensismayensismissoulensisolympusoregonensisschorgeristanleyanavancouverensis, and youngi
    • P. c. costaricensis (Merriam, 1901)
    • P. c. anthonyi (Nelson and Goldman, 1931) includes acrocodiaborbensiscapricornensisconcolorgreeni, and nigra
    • P. c. cabrerae Pocock, 1940 includes hudsonii and puma proposed by Marcelli in 1922

    In 2006, the Florida panther was still referred to as a distinct subspecies P. c. coryi in research works.[21]

    As of 2017, the Cat Classification Taskforce of the Cat Specialist Group recognizes only two subspecies as valid:[22]

    • P. c. concolor in South America, possibly excluding the region northwest of the Andes
    • P. c. couguar in North and Central America and possibly northwestern South America

    Evolution

    Lynx lineageLynxPuma lineageAcinonyxCheetah A. jubatus PumaCougar HerpailurusJaguarundi H. yagouaroundi Domestic cat lineageFelisLeopard cat lineageOtocolobusPrionailurus
    The Puma lineage of the family Felidae, depicted along with closely related genera[23]

    The family Felidae is believed to have originated in Asia about 11 million years ago (Mya). Taxonomic research on felids remains partial, and much of what is known about their evolutionary history is based on mitochondrial DNA analysis.[19] Significant confidence intervals exist with suggested dates. In the latest genomic study of the Felidae, the common ancestor of today’s LeopardusLynxPumaPrionailurus, and Felis lineages migrated across the Bering land bridge into the Americas 8.0 to 8.5 million years ago. The lineages subsequently diverged in that order.[19] North American felids then invaded South America 2–4 Mya as part of the Great American Interchange, following the formation of the Isthmus of Panama.[20]

    The cheetah lineage is suggested by some studies to have diverged from the Puma lineage in the Americas and migrated back to Asia and Africa,[19][20] while other research suggests the cheetah diverged in the Old World itself.[24] A high level of genetic similarity has been found among North American cougar populations, suggesting they are all fairly recent descendants of a small ancestral group. Culver et al. propose the original North American cougar population was extirpated during the Pleistocene extinctions some 10,000 years ago, when other large mammals, such as Smilodon, also disappeared. North America was then repopulated by South American cougars.[20]

    coprolite identified as from a cougar was excavated in Argentina’s Catamarca Province and dated to 17,002–16,573 years old. It contained Toxascaris leonina eggs. This finding indicates that the cougar and the parasite have existed in South America since at least the Late Pleistocene.[25] The oldest fossil record of a cougar (Puma concolor) in South America (Argentina) is a partial skull from the late Calabrian (Ensenadan) age.[26]

    Characteristics

    Cougar skull and jawbone

    Front paw print of a cougar

    The head of the cougar is round, and the ears are erect. Its powerful forequarters, neck, and jaw serve to grasp and hold large prey. It has four retractile claws on its hind paws and five on its forepaws, of which one is a dewclaw. The larger front feet and claws are adaptations for clutching prey.[27]

    Cougars are slender and agile members of the Felidae. They are the fourth largest cat species worldwide;[28] adults stand about 60 to 90 cm (24 to 35 in) tall at the shoulders.[29] Adult males are around 2.4 m (7 ft 10 in) long from nose to tail tip, and females average 2.05 m (6 ft 9 in), with overall ranges between 1.50 to 2.75 m (4 ft 11 in to 9 ft 0 in) nose to tail suggested for the species in general.[30][31] Of this length, the tail typically accounts for 63 to 95 cm (25 to 37 in).[32] Males generally weigh 53 to 72 kg (117 to 159 lb). Females typically weigh between 34 and 48 kg (75 and 106 lb).[32][33] Cougar size is smallest close to the equator and larger towards the poles.[34] The largest recorded cougar, shot in 1901, weighed 105.2 kg (232 lb); claims of 125.2 kg (276 lb) and 118 kg (260 lb) have been reported, though they were probably exaggerated.[35] Male cougars in North America average 62 kg (137 lb), while the average female in the same region averages about 42 kg (93 lb).[36] On average, adult male cougars in British Columbia weigh 56.7 kg (125 lb) and adult females 45.4 kg (100 lb), though several male cougars in British Columbia weighed between 86.4 and 95.5 kg (190 and 211 lb).[37]

    Depending on the locality, cougars can be smaller or bigger than jaguars but are less muscular and not as powerfully built, so on average, their weight is less. Whereas the size of cougars tends to increase as much as distance from the equator increases,[34] which crosses the northern portion of South America, jaguars are generally smaller north of the Amazon River in South America and larger south of it. For example, while South American jaguars are comparatively large, and may exceed 90 kg (200 lb),[38] North American jaguars in Mexico’s Chamela-Cuixmala Biosphere Reserve weigh approximately 50 kg (110 lb), about the same as female cougars.[39]

    Cougar coloring is plain (hence the Latin concolor [“one color”] in the scientific name) but can vary greatly across individuals and even siblings. The coat is typically tawny, but it ranges from silvery-grey to reddish with lighter patches on the underbody, including the jaws, chin, and throat. Infants are spotted and born with blue eyes and rings on their tails;[33] juveniles are pale, and dark spots remain on their flanks.[31] A leucistic individual was seen in Serra dos Órgãos National Park in Rio de Janeiro in 2013 when it was recorded by a camera trap, indicating that pure white individuals do exist within the species, though they are extremely rare.[40][41]

    The cougar has large paws and proportionally the largest hind legs in the Felidae,[33] allowing for great leaping and powerful short sprints. It can leap from the ground up to 5.5 m (18 ft) high into a tree.[42]

    Distribution and habitat

    camera trap image of a cougar in Saguaro National ParkArizona

    The cougar has the most extensive range of any wild land animal in the Americas, spanning 110 degrees of latitude from the Yukon in Canada to the southern Andes in Chile.[1] The species was extirpated from eastern North America, aside from Florida, but they may be recolonizing their former range and isolated populations have been documented east of their contemporary ranges in both the Midwestern US and Canada.[43]

    The cougar lives in all forest types, lowland and mountainous deserts, and in open areas with little vegetation up to an elevation of 5,800 m (19,000 ft).[1] In the Santa Ana Mountains, it prefers steep canyons, escarpments, rim rocks and dense brush.[44] In Mexico, it was recorded in the Sierra de San Carlos.[45] In the Yucatán Peninsula, it inhabits secondary and semi-deciduous forests in El Eden Ecological Reserve.[46] In El Salvador, it was recorded in the lower montane forest in Montecristo National Park and in a river basin in the Morazán Department above 700 m (2,300 ft) in 2019.[47] In Colombia, it was recorded in a palm oil plantation close to a riparian forest in the Llanos Basin, and close to water bodies in the Magdalena River Valley.[48][49] In the human-modified landscape of central Argentina, it inhabits bushland with abundant vegetation cover and prey species.[50]

    Behavior and ecology

    Cougars are an important keystone species in Western Hemisphere ecosystems, linking numerous species at many trophic levels. In a comprehensive literature review of more than 160 studies on cougar ecology, ecological interactions with 485 other species in cougar-inhabited ecosystems have been shown to involve different areas of interaction, ranging from the use of other species as food sources and prey, fear effects on potential prey, effects from carcass remains left behind, to competitive effects on other predator species in shared habitat. The most common research topic in the literature used here was the cougar’s diet and its prey’s regulation.[51]

    Hunting and diet

    Camera trap image of cougar in the Santa Susana Mountains northwest of Los Angeles
    Cougar with deer kill

    The cougar is a generalist hypercarnivore. It prefers large mammals such as mule deerwhite-tailed deerelkmoosemountain goat and bighorn sheep. It opportunistically takes smaller prey such as rodentslagomorphs, smaller carnivores, birds, and even domestic animals, including pets.[52] The mean weight of cougar vertebrate prey increases with its body weight and is lower in areas closer to the equator. A survey of North America research found 68% of prey items were ungulates, especially deer. Only the Florida panther showed variation, often preferring feral hogs and armadillos.[34] Cougars have been known to prey on introduced gemsbok populations in New Mexico. One individual cougar was recorded as hunting 29 gemsbok, which made up 58% of its recorded kills. Most gemsbok kills were neonates, but some adults were also known to have been taken.[53] Elsewhere in the southwestern United States, they have been recorded to also prey on feral horses in the Great Basin,[54] as well as feral donkeys in the Sonoran and Mojave Deserts.[55]

    Investigations at Yellowstone National Park showed that elk and mule deer were the cougar’s primary prey; the prey base is shared with the park’s wolves, with which the cougar competes for resources.[56] A study on winter kills from November to April in Alberta showed that ungulates accounted for greater than 99% of the cougar diet. Learned, individual prey recognition was observed, as some cougars rarely killed bighorn sheep, while others relied heavily on the species.[57]

    In the Central and South American cougar range area, the ratio of deer in the diet declines. Small to mid-sized mammals, including large rodents such as the capybara, are preferred. Ungulates accounted for only 35% of prey items in one survey, about half that of North America. Competition with the larger jaguar in South America has been suggested for the decline in the size of prey items.[34] In Central or North America, the cougar and jaguar share the same prey, depending on its abundance.[58] Other listed prey species of the cougar include miceporcupinesAmerican beaversraccoonsharesguanacoespeccariesvicuñasrheas and wild turkeys.[59] Birds and small reptiles are sometimes preyed upon in the south, but this is rarely recorded in North America.[34] Magellanic penguins (Spheniscus magellanicus) constitute the majority of prey items in cougar diet in Patagonia‘s Bosques Petrificados de Jaramillo National Park and Monte León National Park.[60]

    Although capable of sprinting, the cougar is typically an ambush predator. It stalks through brush and trees, across ledges, or other covered spots, before delivering a powerful leap onto the back of its prey and a suffocating neck bite. The cougar can break the neck of some of its smaller prey with a strong bite and momentum bearing the animal to the ground.[27] Kills are generally estimated around one large ungulate every two weeks. The period shrinks for females raising young, and may be as short as one kill every three days when cubs are nearly mature around 15 months.[33] The cat drags a kill to a preferred spot, covers it with brush, and returns to feed over a period of days. The cougar is generally reported to not be a scavenger, but deer carcasses left exposed for study were scavenged by cougars in California, suggesting more opportunistic behavior.[61]

    Interactions with other predators

    Juvenile cougar in conflict with coyotes at National Elk Refuge, using a buck-and-rail fence for refuge

    Aside from humans, no species preys upon mature cougars in the wild, although conflicts with other predators or scavengers occur. Of the large predators in Yellowstone National Park – the grizzly and black bearsgray wolf and cougar – the massive grizzly bear appears dominant, often (though not always) able to drive a gray wolf pack, black bear or cougar off their kills. One study found that grizzlies and American black bears visited 24% of cougar kills in Yellowstone and Glacier National Parks, usurping 10% of carcasses. Bears gained up to 113%, and cougars lost up to 26% of their daily energy requirements from these encounters.[62] In Colorado and California, black bears were found to visit 48% and 77% of kills, respectively. In general, cougars are subordinate to black bears when it comes to killing, and when bears are most active, the cats take prey more frequently and spend less time feeding on each kill. Unlike several subordinate predators from other ecosystems, cougars do not appear to exploit spatial or temporal refuges to avoid competitors.[63][64]

    The gray wolf and the cougar compete more directly for prey, mostly in winter. Packs of wolves can steal cougars’ kills, and there are some documented cases of cougars being killed by them. One report describes a large pack of seven to 11 wolves killing a female cougar and her kittens,[65] while in nearby Sun Valley, Idaho, a 2-year-old male cougar was found dead, apparently killed by a wolf pack.[66] Conversely, one-to-one confrontations tend to be dominated by the cat, and there are various documented accounts where wolves have been ambushed and killed,[67][68][69][70] including adult male specimens.[71] Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities, and disrupting the feline’s behavior. Preliminary research in Yellowstone, for instance, has shown displacement of the cougar by wolves.[72] One researcher in Oregon noted: “When there is a pack around, cougars are not comfortable around their kills or raising kittens […] A lot of times a big cougar will kill a wolf, but the pack phenomenon changes the table.”[73] Both species are capable of killing mid-sized predators, such as bobcatsCanada lynxeswolverines and coyotes, and tend to suppress their numbers.[56] Although cougars can kill coyotes, the latter have been documented attempting to prey on cougar cubs.[74]

    The cougar and jaguar share overlapping territory in the southern portion of its range.[75] The jaguar tends to take the larger prey where ranges overlap, reducing both the cougar’s potential size and the likelihood of direct competition between the two cats.[34] Cougars appear better than jaguars at exploiting a broader prey niche and smaller prey.[76]

    Social spacing and interactions

    The cougar is a mostly solitary animal. Only mothers and kittens live in groups, with adults meeting rarely. While generally loners, cougars will reciprocally share kills and seem to organize themselves into small communities defined by the territories of dominant males. Cats within these areas socialize more frequently with each other than with outsiders.[77]

    In the vicinity of a cattle ranch in northern Mexico, cougars exhibited nocturnal activity that overlapped foremost with the activity of calves.[78] In a nature reserve in central Mexico, the activity of cougars was crepuscular and nocturnal, overlapping largely with the activity of the nine-banded armadillo (Dasypus novemcinctus).[79] Cougars in the montane Abra-Tanchipa Biosphere Reserve in southeastern Mexico displayed a cathemeral activity pattern.[80] Data from 12 years of camera trapping in the Pacific slope and Talamanca Cordillera of Costa Rica showed cougars as cathemeral.[81] Both cougars and jaguars in the Cockscomb Basin of Belize were nocturnal but avoided each other.[82] In a protected cloud forest in the central Andes of Colombia, cougars were active from late afternoon to shortly before sunrise and sometimes during noon and early afternoon.[83] In protected areas of the MadidiTambopata Landscape in Bolivia and Peru, cougars were active throughout the day but with a tendency to nocturnal activity that overlapped with the activity of main prey species.[84] During an 8-year-long study in a modified landscape in southeastern Brazil, male cougars were primarily nocturnal, but females were active at night and day.[85] Cougars were diurnal in the Brazilian Pantanal, but crepuscular and nocturnal in protected areas in the CerradoCaatinga and ecotone biomes.[86] Cougars in the Atlantic Forest were active throughout the day but displayed peak activity during early mornings in protected areas and crepuscular and nocturnal activity in less protected areas.[87] In central Argentina, cougars were active day and night in protected areas but were active immediately after sunset and before sunrise outside protected areas.[88] Cougars displayed a foremost crepuscular and nocturnal activity pattern in a ranching area in southern Argentina.[50]

    Home range sizes and overall cougar abundance depend on terrain, vegetation, and prey abundance.[89] Research suggests a lower limit of 25 km2 (9.7 sq mi) and upper limit of 1,300 km2 (500 sq mi) of home range for males.[90] Large male home ranges of 150 to 1,000 km2 (58 to 386 sq mi) with female ranges half that size.[91] One female adjacent to the San Andres Mountains was found with a big range of 215 km2 (83 sq mi), necessitated by poor prey abundance.[92] Research has shown cougar abundances from 0.5 animals to as many as seven per 100 km2 (39 sq mi).[93]

    Male home ranges include or overlap with females but, at least where studied, not with those of other males. The home ranges of females overlap slightly. Males create scrapes composed of leaves and duff with their hind feet, and mark them with urine and sometimes feces.[94] When males encounter each other, they vocalize and may engage in violent conflict if neither backs down.[95]

    Cougars communicate with various vocalizations. Aggressive sounds include growls, spits, snarls, and hisses. During the mating season, estrus females produce caterwauls or yowls to attract mates, and males respond with similar vocals. Mothers and offspring keep in contact with whistles, chirps, and mews.[52][96]

    Reproduction and life cycle

    North American cougar cub in the Santa Monica Mountains

    Cubs

    Females reach sexual maturity at the age of 18 months to three years and are in estrus for about eight days of a 23-day cycle; the gestation period is approximately 91 days.[33] Both adult males and females may mate with multiple partners, and a female’s litter can have multiple paternities.[52] Copulation is brief but frequent. Chronic stress can result in low reproductive rates in captivity as well as in the field.[97]

    Gestation is 82–103 days long.[52] Only females are involved in parenting. Litter size is between one and six cubs, typically two. Caves and other alcoves that offer protection are used as litter dens. Born blind, cubs are completely dependent on their mother at first and begin to be weaned at around three months of age. As they grow, they go out on forays with their mother, first visiting kill sites and, after six months, beginning to hunt small prey on their own.[90][52] Kitten survival rates are just over one per litter.[33]

    Juveniles remain with their mothers for one to two years.[52] When a female reaches estrous again, her offspring must disperse or the male will kill them. Males tend to disperse further than females.[98] One study has shown a high mortality rate among cougars that travel farthest from their maternal range, often due to conflicts with other cougars.[90] In a study area in New Mexico, males dispersed farther than females, traversed large expanses of non-cougar habitat and were probably most responsible for nuclear gene flow between habitat patches.[99]

    Life expectancy in the wild is reported at 8 to 13 years and probably averages 8 to 10; a female of at least 18 years was reported killed by hunters on Vancouver Island.[33] Cougars may live as long as 20 years in captivity. Causes of death in the wild include disability and disease, competition with other cougars, starvation, accidents, and, where allowed, hunting. The feline immunodeficiency virus is well-adapted to the cougar.[100]

    Conservation

    Two cougar kittens at White Oak Conservation

    The cougar has been listed as Least Concern on the IUCN Red List since 2008. However, it is also listed on CITES Appendix II.[1] Hunting it is prohibited in California, Costa RicaHondurasNicaraguaGuatemalaPanamaVenezuela, Colombia, French GuianaSurinameBolivia, Brazil, Chile, ParaguayUruguay and most of Argentina. Hunting is regulated in Canada, Mexico, Peru, and the United States.[33] Establishing wildlife corridors and protecting sufficient range areas are critical for the sustainability of cougar populations. Research simulations showed that it faces a low extinction risk in areas larger than 2,200 km2 (850 sq mi). Between one and four new individuals entering a population per decade markedly increases persistence, thus highlighting the importance of habitat corridors.[101]

    The Florida panther population is afforded protection under the Endangered Species Act.[102][103] The Texas Mountain Lion Conservation Project was launched in 2009 and aimed at raising local people’s awareness of the status and ecological role of the cougar and mitigating conflict between landowners and cougars.[104]

    The cougar is threatened by habitat losshabitat fragmentation, and depletion of its prey base due to poaching. Hunting is legal in the western United States. In Florida, heavy traffic causes frequent accidents involving cougars. Highways are a major barrier to the dispersal of cougars.[1] The cougar populations in California are becoming fragmented with the increase in human population and infrastructure growth in the state.[105]

    Human–wildlife conflict in proximity of 5 km2 (1.9 sq mi) of cougar habitat is pronounced in areas with a median human density of 32.48 inhabitants/km2 (84.1 inhabitants/sq mi) and a median livestock population density of 5.3 heads/km2 (14 heads/sq mi). Conflict is generally lower in areas more than 16.1 km (10.0 mi) away from roads and 27.8 km (17.3 mi) away from settlements.[106]

    Relationships with humans

    Attacks on humans

    In North America

    See also: List of fatal cougar attacks in North America

    Mountain lion warning sign in California, U.S.

    Due to the expanding human population, cougar ranges increasingly overlap with areas inhabited by humans.[107] Attacks on humans are very rare, as cougar prey recognition is a learned behavior and they do not generally recognize humans as prey.[108] In a 10-year study in New Mexico of wild cougars who were not habituated to humans, the animals did not exhibit threatening behavior to researchers who approached closely (median distance=18.5 m; 61 feet) except in 6% of cases; 1416 of those were females with cubs.[109] Attacks on people, livestock, and pets may occur when a puma habituates to humans or is in a condition of severe starvation. Attacks are most frequent during late spring and summer when juvenile cougars leave their mothers and search for new territory.[110]

    Between 1890 and 1990 in North America, there were 53 reported, confirmed attacks on humans, resulting in 48 nonfatal injuries and 10 deaths of humans (the total is greater than 53 because some attacks had more than one victim).[111] By 2004, the count had climbed to 88 attacks and 20 deaths.[112]

    Within North America, the distribution of attacks is not uniform. The heavily populated state of California saw a dozen attacks from 1986 to 2004 (after just three from 1890 to 1985), including three fatalities.[113] In March 2024, two brothers in California were attacked by a male cougar, with one being fatally wounded; it was the state’s first fatal attack in 20 years.[114][115] Washington state was the site of a fatal attack in 2018, its first since 1924.[116] Lightly populated New Mexico reported an attack in 2008, the first there since 1974.[117]

    As with many predators, a cougar may attack if cornered, if a fleeing human stimulates their instinct to chase, or if a person “plays dead“. Standing still may cause the cougar to consider a person easy prey.[118] Exaggerating the threat to the animal through intense eye contact, loud shouting, and any other action to appear larger and more menacing, may make the animal retreat. Fighting back with sticks and rocks, or even bare hands, is often effective in persuading an attacking cougar to disengage.[108][110]

    When cougars do attack, they usually employ their characteristic neck bite, attempting to position their teeth between the vertebrae and into the spinal cord. Neck, head, and spinal injuries are common and sometimes fatal.[108] Children are at greatest risk of attack and least likely to survive an encounter. Detailed research into attacks before 1991 showed that 64% of all victims – and almost all fatalities – were children. The same study showed the highest proportion of attacks to have occurred in British Columbia, particularly on Vancouver Island, where cougar populations are especially dense.[111] Preceding attacks on humans, cougars display aberrant behavior, such as activity during daylight hours, a lack of fear of humans, and stalking humans. There have sometimes been incidents of pet cougars mauling people.[119][120]

    Research on new wildlife collars may reduce human-animal conflicts by predicting when and where predatory animals hunt. This may save the lives of humans, pets, and livestock, as well as the lives of these large predatory mammals that are important to the balance of ecosystems.[121]

    In South America

    Cougars in the southern cone of South America are reputed to be extremely reluctant to attack people; in legend, they defended people against jaguars.[122] The nineteenth-century naturalists Félix de Azara[123] and William Henry Hudson[124] thought that attacks on people, even children or sleeping adults, did not happen. Hudson, citing anecdotal evidence from hunters, claimed that pumas were positively inhibited from attacking people, even in self-defense. Attacks on humans, although exceedingly rare, have occurred.[125][126]

    An early, authenticated, non-fatal case occurred near Lake Viedma, Patagonia, in 1877 when a female mauled the Argentine scientist Francisco P. Moreno; Moreno afterward showed the scars to Theodore Roosevelt. In this instance, however, Moreno had been wearing a guanaco-hide poncho round his neck and head as protection against the cold;[127] in Patagonia the guanaco is the puma’s chief prey animal.[128] Another authenticated case occurred in 1997 in Iguazú National Park in northeastern Argentina, when the 20-month-old son of a ranger was killed by a female puma. Forensic analysis found specimens of the child’s hair and clothing fibers in the animal’s stomach. The coatí is the puma’s chief prey in this area. Despite prohibitory signs, coatis are hand-fed by tourists in the park, causing unnatural approximation between cougars and humans. This particular puma had been raised in captivity and released into the wild.[129] On March 13, 2012, Erica Cruz, a 23-year-old shepherdess was found dead in a mountainous area near Rosario de Lerma, Salta Province, in northwestern Argentina.[130] Claw incisions, which severed a jugular vein, indicated that the attacker was a felid; differential diagnosis ruled out other possible perpetrators.[b] There were no bite marks on the victim, who had been herding goats.[131] In 2019 in Córdoba Province, Argentina an elderly man was badly injured by a cougar after he attempted to defend his dog from it, while in neighboring Chile a 28-year-old woman was attacked and killed in Corral, in Los Ríos Region, on October 20, 2020.[132]

    Fatal attacks by other carnivores, such as feral dogs, can be misattributed to cougars without appropriate forensic knowledge.[133]

    Predation on domestic animals

    The Cougar Hunt, a 1920s silent film created by the United States Department of Agriculture which explains the procedures to successfully hunt livestock-threatening cougars

    During the early years of ranching, cougars were considered on par with wolves in destructiveness. According to figures in Texas in 1990, 86 calves (0.0006% of Texas’s 13.4 million cattle and calves), 253 mohair goats, 302 mohair kids, 445 sheep (0.02% of Texas’s 2 million sheep and lambs) and 562 lambs (0.04% of Texas’s 1.2 million lambs) were confirmed to have been killed by cougars that year.[134][135] In Nevada in 1992, cougars were confirmed to have killed nine calves, one horse, four foals, five goats, 318 sheep, and 400 lambs. In both reports, sheep were the most frequently attacked. Some instances of surplus killing have resulted in the deaths of 20 sheep in one attack.[136] A cougar’s killing bite is applied to the back of the neck, head, or throat and the cat inflicts puncture marks with its claws usually seen on the sides and underside of the prey, sometimes also shredding the prey as it holds on. Coyotes also typically bite the throat, but the work of a cougar is generally clean, while bites inflicted by coyotes and dogs leave ragged edges. The size of the tooth puncture marks also helps distinguish kills made by cougars from those made by smaller predators.[137]

    Remedial hunting appears to have the paradoxical effect of increased livestock predation and complaints of human-cougar conflicts. In a 2013 study, the most important predictor of cougar problems was the remedial hunting of cougars the previous year. Each additional cougar on the landscape increased predation and human-cougar complaints by 5%, but each animal killed during the previous year increased complaints by 50%. The effect had a dose-response relationship with very heavy (100% removal of adult cougars) remedial hunting, leading to a 150–340% increase in livestock and human conflicts.[138] This effect is attributed to the removal of older cougars that have learned to avoid people and their replacement by younger males that react differently to humans. Remedial hunting enables younger males to enter the former territories of the older animals.[139][140] Predation by cougars on dogs “is widespread, but occurs at low frequencies”.[141]

    In mythology

    The grace and power of the cougar have been widely admired in the cultures of the indigenous peoples of the Americas. The Inca city of Cusco is reported to have been designed in the shape of a cougar, and the animal also gave its name to both Inca regions and people. The Moche people often represented the cougar in their ceramics.[142] The sky and thunder god of the Inca, Viracocha, has been associated with the animal.[143]

    In North America, mythological descriptions of the cougar have appeared in the stories of the Hocąk language (“Ho-Chunk” or “Winnebago”) of Wisconsin and Illinois[144] and the Cheyenne, among others. To the Apache and Walapai of the Southwestern United States, the wail of the cougar was a harbinger of death.[145] The Algonquins and Ojibwe believe that the cougar lived in the underworld and was wicked, whereas it was a sacred animal among the Cherokee.[