Coyote

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Coyote info via Wikipedia:

Coyote
Temporal range: Middle Pleistocene – present (0.74–0.85 Ma)[1]:p131
2009-Coyote-Yosemite.jpg
Mountain coyote (C. l. lestes)
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Family: Canidae
Genus: Canis
Species: C. latrans
Binomial name
Canis latrans
Say, 1823
Cypron-Range Canis latrans.svg
Modern range of Canis latrans
Synonyms[3]

The coyote (Canis latrans); from Nahuatl About this sound pronunciation ) is a canine native to North America. It is smaller than its close relative, the gray wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological niche as the golden jackal does in Eurasia, though it is larger and more predatory, and is sometimes called the American jackal by zoologists.

The coyote is listed as least concern by the International Union for Conservation of Nature due to its wide distribution and abundance throughout North America, southwards through Mexico, and into Central America. The species is versatile, able to adapt to and expand into environments modified by humans. It is enlarging its range, with coyotes moving into urban areas in the Eastern U.S., and was sighted in eastern Panama (across the Panama Canal from their home range) for the first time in 2013.

As of 2005[update], 19 coyote subspecies are recognized. The average male weighs 8 to 20 kg (18 to 44 lb) and the average female 7 to 18 kg (15 to 40 lb). Their fur color is predominantly light gray and red or fulvous interspersed with black and white, though it varies somewhat with geography. It is highly flexible in social organization, living either in a family unit or in loosely knit packs of unrelated individuals. It has a varied diet consisting primarily of animal meat, including deer, rabbits, hares, rodents, birds, reptiles, amphibians, fish, and invertebrates, though it may also eat fruits and vegetables on occasion. Its characteristic vocalization is a howl made by solitary individuals. Man is the coyote's greatest threat, followed by cougars and gray wolves. In spite of this, coyotes sometimes mate with gray, eastern, or red wolves, producing "coywolf" hybrids. In the northeastern United States and eastern Canada, the eastern coyote (a larger subspecies, though still smaller than wolves) is the result of various historical and recent matings with various types of wolves. Genetic studies show that most North American wolves contain some level of coyote DNA.

The coyote is a prominent character in Native American folklore, mainly in the Southwestern United States and Mexico, usually depicted as a trickster that alternately assumes the form of an actual coyote or a man. As with other trickster figures, the coyote uses deception and humor to rebel against social conventions. The animal was especially respected in Mesoamerican cosmology as a symbol of military might. After the European colonization of the Americas, it was reviled in Anglo-American culture as a cowardly and untrustworthy animal. Unlike wolves (gray, eastern, or red), which have undergone an improvement of their public image, attitudes towards the coyote remain largely negative.

Description

Closeup of a mountain coyote's (C. l. lestes) head

Coyote males average 8 to 20 kg (18 to 44 lb) in weight, while females average 7 to 18 kg (15 to 40 lb), though size varies geographically. Northern subspecies, which average 18 kg (40 lb), tend to grow larger than the southern subspecies of Mexico, which average 11.5 kg (25 lb). Body length ranges on average from 1.0 to 1.35 m (3 ft 3 in to 4 ft 5 in), and tail length 40 cm (16 in), with females being shorter in both body length and height.[4] The largest coyote on record was a male killed near Afton, Wyoming, on November 19, 1937, which measured 1.5 m (4 ft 11 in) from nose to tail, and weighed 34 kg (75 lb).[5] Scent glands are located at the upper side of the base of the tail and are a bluish-black color.[6]

The color and texture of the coyote's fur varies somewhat geographically.[4] The hair's predominant color is light gray and red or fulvous, interspersed around the body with black and white. Coyotes living at high elevations tend to have more black and gray shades than their desert-dwelling counterparts, which are more fulvous or whitish-gray.[7] The coyote's fur consists of short, soft underfur and long, coarse guard hairs. The fur of northern subspecies is longer and denser than in southern forms, with the fur of some Mexican and Central American forms being almost hispid (bristly).[8] Generally, adult coyotes (including coywolf hybrids) have a sable coat color, dark neonatal coat color, bushy tail with an active supracaudal gland, and a white facial mask.[9]Albinism is extremely rare in coyotes; out of a total of 750,000 coyotes harvested by federal and cooperative hunters between March 22, 1938, and June 30, 1945, only two were albinos.[7]

The coyote is typically smaller than the gray wolf, but has longer ears and a relatively larger braincase,[4] as well as a thinner frame, face, and muzzle. The scent glands are smaller than the gray wolf's, but are the same color.[6] Its fur color variation is much less varied than that of a wolf.[10] The coyote also carries its tail downwards when running or walking, rather than horizontally as the wolf does.[11]

Coyote tracks can be distinguished from those of dogs by their more elongated, less rounded shape.[12][13] Unlike dogs, the upper canines of coyotes extends past the mental foramina.[4]

Taxonomy and evolution

History

At the time of the European colonization of the Americas, coyotes were largely confined to open plains and arid regions of the western half of the continent.[14] In early post-Columbian historical records, distinguishing between coyotes and wolves is often difficult. One record from 1750 in Kaskaskia, Illinois, written by a local priest, noted that the "wolves" encountered there were smaller and less daring than European wolves. Another account from the early 1800s in Edwards County mentioned wolves howling at night, though these were likely coyotes.[15] This species was encountered several times during the Lewis and Clark Expedition (1804–1806), though it was already well known to European traders on the upper Missouri. Lewis, writing on May 5, 1805, in northeastern Montana, described the coyote in these terms:[16]

Toltec pictograph of coyote.

the small woolf or burrowing dog of the prairies are the inhabitants almost invariably of the open plains; they usually ascociate in bands of ten or twelve sometimes more and burrow near some pass or place much frequented by game; not being able alone to take deer or goat they are rarely ever found alone but hunt in bands; they frequently watch and seize their prey near their burrows; in these burrows they raise their young and to them they also resort when pursued; when a person approaches them they frequently bark, their note being precisely that of the small dog. they are of an intermediate size between that of the fox and dog, very active fleet and delicately formed; the ears large erect and pointed the head long and pointed more like that of the fox; tale long; . . . the hair and fur also resembles the fox tho' is much coarser and inferior. they are of a pale redish brown colour. the eye of a deep sea green colour small and piercing. their tallons [claws] are reather longer than those of the ordinary wolf or that common to the atlantic states, none of which are to be found in this quarter, nor I believe above the river Plat.

The coyote was first scientifically described by Thomas Say, a naturalist, in September 1819 on the site of Lewis and Clark's Council Bluffs, fifteen miles up the Missouri River from the mouth of the Platte during a government-sponsored expedition with Major Stephen Long. He had the first edition of the Lewis and Clark journals in hand, which contained Biddle's edited version of Lewis's observations dated May 5, 1805. His account was published in 1823.[17][18]

Naming and etymology

The earliest written reference to the species comes from the naturalist Francisco Hernández's Plantas y Animales de la Nueva España (1651), where it is described as a "Spanish fox" or "jackal". The first published usage of the word "coyote" (which is a Spanish borrowing of its Nahuatl name coyōtl) comes from the historian Francisco Javier Clavijero's Historia de México in 1780.[19] The first time it was used in English occurred in William Bullock's Six months' residence and travels in Mexico (1824), where it is variously transcribed as cayjotte and cocyotie. The word's spelling was standardized as "coyote" by the 1880s.[16][20] Alternative English names for the coyote include "prairie wolf", "brush wolf", "cased wolf",[21][a] "little wolf"[22] and "American jackal".[23] Its binomial name Canis latrans translates to "barking dog", a reference to the many vocalizations they produce.[24]

Local and indigenous names for Canis latrans

Evolution

Phylogenetic tree of the extant wolf-like canids
Caninae 3.5 Ma
3.0
2.7
1.9
1.6
1.3
1.1



Dog Tibetan mastiff (white background).jpg



Gray wolf Dogs, jackals, wolves, and foxes (Plate I).jpg




Himalayan wolf Dogs, jackals, wolves, and foxes (Plate III).jpg




Coyote Dogs, jackals, wolves, and foxes (Plate IX).jpg




African golden wolf Dogs, jackals, wolves, and foxes (Plate XI).jpg




Ethiopian wolf Dogs, jackals, wolves, and foxes (Plate VI).jpg




Golden jackal Dogs, jackals, wolves, and foxes (Plate X).jpg




Dhole Dogs, jackals, wolves, and foxes (Plate XLI).jpg




African wild dog Dogs, jackals, wolves, and foxes (Plate XLIV).jpg




2.6

Side-striped jackal Dogs, jackals, wolves, and foxes (Plate XIII).jpg



Black-backed jackal Dogs, jackals, wolves, and foxes (Plate XII).jpg





Phylogenetic relationships between the extant wolf-like clade of canids based on nuclear DNA sequence data taken from the cell nucleus,[40][41] except for the Himalayan wolf, based on mitochondrial DNA sequences.[41][42] Timing in millions of years.[41]

Fossil record

Skeleton of Pleistocene coyote (C. l. orcutti)

Xiaoming Wang and Richard H. Tedford, one of the foremost authorities on carnivore evolution,[43] proposed that the genus Canis was the descendant of the coyote-like Eucyon davisi and its remains first appeared in the Miocene 6 million years ago (Mya) in the southwestern USA and Mexico. By the Pliocene (5 Mya), the larger Canis lepophagus[44] appeared in the same region and by the early Pleistocene (1 Mya) C. latrans (the coyote) was in existence. They proposed that the progression from Eucyon davisi to C lepophagus to the coyote was linear evolution.[45]:p58 Additionally, C. edwardii, C. latrans, and C.  aureus form together a small clade and because C. edwardii appeared earliest spanning the mid-Blancan (late Pliocene) to the close of the Irvingtonian (late Pleistocene), it is proposed as the direct ancestor of the coyote.[1]:p175,180 Johnston describes C. lepophagus as having a more slender skull and skeleton than the modern coyote.[46]:385 Ronald Nowak found that the early populations had small, delicate, narrowly proportioned skulls that resemble small coyotes and appear to be ancestral to C. latrans.[47]:p241

C. lepophagus was a similar in weight to modern coyotes, but had shorter limb bones that indicates a less cursorial lifestyle. The coyote represents a more primitive form of Canis than the gray wolf, as shown by its relatively small size and its comparatively narrow skull and jaws, which lack the grasping power necessary to hold the large prey in which wolves specialize. This is further corroborated by the coyote's sagittal crest, which is low or totally flattened, thus indicating a weaker bite than the wolf's. The coyote is not a specialized carnivore as the wolf is, as shown by the larger chewing surfaces on the molars, reflecting the species' relative dependence on vegetable matter. In these respects, the coyote resembles the fox-like progenitors of the genus more so than the wolf.[48]

The oldest fossils that fall within the range of the modern coyote date to 0.74–0.85 Ma (million years) in Hamilton Cave, West Virginia; 0.73 Ma in Irvington, California; 0.35–0.48 Ma in Porcupine Cave, Colorado and in Cumberland Cave, Pennsylvania.[1]:p136 Modern coyotes arose 1,000 years after the Quaternary extinction event.[49] Compared to their modern Holocene counterparts, Pleistocene coyotes (C. l. orcutti) were larger and more robust, likely in response to larger competitors and prey.[49] Pleistocene coyotes were likely more specialized carnivores than their descendants, as their teeth were more adapted to shearing meat, showing fewer grinding surfaces suited for processing vegetation.[50] Their reduction in size occurred within 1000 years of the Quaternary extinction event, when their large prey died out.[49] Furthermore, Pleistocene coyotes were unable to exploit the big-game hunting niche left vacant after the extinction of the dire wolf (C. dirus), as it was rapidly filled by gray wolves, which likely actively killed off the large coyotes, with natural selection favoring the modern gracile morph.[50]

DNA evidence

In 1993, a study proposed that the wolves of North America display skull traits more similar towards the coyote than those wolves from Eurasia.[51] In 2010, a study found that the coyote was a basal member of the clade that included the Tibetan wolf, the Dog, the Mongolian wolf and the Eurasian wolf, with the Tibetan wolf diverging early from wolves and domestic dogs.[52] In 2016, a whole-genome DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago. The study also indicated that all North America wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry, and that the red wolf and eastern wolf are highly admixed with different proportions of gray wolf and coyote ancestry. One test indicated a wolf/coyote divergence time of 51,000 years before present that matched other studies indicating that the extant wolf came into being around this time. Another test indicated that the red wolf diverged from the coyote between 55,000 and 117,000 years before present and the Great Lakes region wolf 32,000 years before present. Other tests and modelling showed various divergence ranges and the conclusion was a range of less than 6,000 and 117,000 years before present. The study found that coyote ancestry was highest in red wolves from the southeastern United States and lowest among the Great Lakes region wolves. The theory proposed was that this pattern matched the south to north disappearance of the wolf due to European colonization and its resulting loss of habitat. Bounties led to the extirpation of wolves initially in the southeast, and as the wolf population declined, wolf-coyote admixture increased. Later, this process occurred in the Great Lakes region with the influx of coyotes replacing wolves, followed by the expansion of coyotes and their hybrids across the wider region.[53][54] The proposed timing of the wolf/coyote divergence conflicts with the finding of a coyote-like specimen in strata dated to 1 Mya.[45]

Subspecies

As of 2005[update], 19 subspecies are recognized.[23][55] Geographic variation in coyotes is not great, though taken as a whole, the eastern subspecies (C. l. thamnos and C. l. frustor) are large, dark-colored animals, with a gradual paling in color and reduction in size westward and northward (C. l. texensis, C. l. latrans, C. l. lestes, and C. l. incolatus), a brightening of ochraceous tones–deep orange or brown–towards the Pacific coast (C. l. ochropus, C. l. umpquensis), a reduction in size in the Southwestern United States (C. l. microdon, C. l. mearnsi) and a general trend towards dark reddish colors and short muzzles in Mexican and Central American populations.[56]

Hybridization

Melanistic coyotes owe their color to a mutation that first arose in domestic dogs.[68]

Coyotes have occasionally mated with dogs, sometimes producing crosses colloquially known as "coydogs".[69] Such matings are rare in the wild, as the mating cycles of dogs and coyotes do not coincide, and coyotes are usually antagonistic towards dogs. Hybridization usually only occurs when coyotes are expanding into areas where conspecifics are few, and dogs are the only alternatives. Even then, pup survival rates are lower than normal, as dogs do not form pair bonds with coyotes, thus making the rearing of pups more difficult.[70] In captivity, F1 hybrids (first generation) tend to be more mischievous and less manageable as pups than dogs, and are less trustworthy on maturity than wolf-dog hybrids.[69] Hybrids vary in appearance, but generally retain the coyote's usual characteristics. F1 hybrids tend to be intermediate in form between dogs and coyotes, while F2 hybrids (second generation) are more varied. Both F1 and F2 hybrids resemble their coyote parents in terms of shyness and intrasexual aggression.[9][71] Hybrids are fertile and can be successfully bred through four generations.[69]Melanistic coyotes owe their black pelts to a mutation that first arose in domestic dogs.[68] A population of nonalbino white coyotes in Newfoundland owe their coloration to a melanocortin 1 receptor mutation inherited from Golden Retrievers.[72]

Coywolf hybrid conceived in captivity between a male gray wolf and a female coyote

Coyotes have hybridized with wolves to varying degrees, particularly in the Eastern United States and Canada. The so-called "eastern coyote" of northeastern North America probably originated in the aftermath of the extermination of gray and eastern wolves in the northeast, thus allowing coyotes to colonize former wolf ranges and mix with remnant wolf populations. This hybrid is smaller than either the gray or eastern wolf, and holds smaller territories, but is in turn larger and holds more extensive home ranges than the typical western coyote. As of 2010, the eastern coyote's genetic makeup is fairly uniform, with minimal influence from eastern wolves or western coyotes.[73] Adult eastern coyotes are larger than western coyotes, with female eastern coyotes weighing 21% more than male western coyotes.[73][74] Physical differences become more apparent by the age of 35 days, with eastern coyote pups having longer legs than their western counterparts. Differences in dental development also occurs, with tooth eruption being later, and in a different order in the eastern coyote.[75] Aside from its size, the eastern coyote is physically similar to the western coyote. The four color phases range from dark brown to blond or reddish blond, though the most common phase is gray-brown, with reddish legs, ears, and flanks.[76] No significant differences exist between eastern and western coyotes in aggression and fighting, though eastern coyotes tend to fight less, and are more playful. Unlike western coyote pups, in which fighting precedes play behavior, fighting among eastern coyote pups occurs after the onset of play.[75] Eastern coyotes tend to reach sexual maturity at two years of age, much later than in western coyotes.[73]

Eastern and red wolves are also products of varying degrees of wolf-coyote hybridization. The eastern wolf probably was a result of a wolf-coyote admixture, combined with extensive backcrossing with parent gray wolf populations. The red wolf may have originated during a time of declining wolf populations in the southeastern United States, forcing a wolf-coyote hybridization as well as backcrossing with local parent coyote populations to the extent that about 75–80% of the modern red wolf's genome is of coyote derivation.[53][77]

Behavior

Social and reproductive behaviors

Mearns' coyote (C. l. mearnsi) pups playing
A pack of coyotes in Yellowstone National Park

Like the golden jackal, the coyote is gregarious, but not as dependent on conspecifics as more social canid species like wolves are. This is likely because the coyote is not a specialized hunter of large prey as the latter species is.[78] The basic social unit of a coyote pack is a family containing a reproductive female. However, unrelated coyotes may join forces for companionship, or to bring down prey too large to attack singly. Such "nonfamily" packs are only temporary, and may consist of bachelor males, nonreproductive females and subadult young. Families are formed in midwinter, when females enter estrus.[22] Pair bonding can occur 2–3 months before actual copulation takes place.[79] The copulatory tie can last 5–45 minutes.[80] A female entering estrus attracts males by scent marking[81] and howling with increasing frequency.[23] A single female in heat can attract up to seven reproductive males, which can follow her for as long as a month. Although some squabbling may occur among the males, once the female has selected a mate and copulates, the rejected males do not intervene, and move on once they detect other estrous females.[22] Unlike the wolf, which has been known to practice both monogamous and bigamous matings,[82] the coyote is strictly monogamous, even in areas with high coyote densities and abundant food.[83] Females that fail to mate sometimes assist their sisters or mothers in raising their pups, or join their siblings until the next time they can mate. The newly mated pair then establishes a territory and either constructs their own den or cleans out abandoned badger, marmot, or skunk earths. During the pregnancy, the male frequently hunts alone and brings back food for the female. The female may line the den with dried grass or with fur pulled from her belly.[22] The gestation period is 63 days, with an average litter size of six, though the number fluctuates depending on coyote population density and the abundance of food.[23]

Coyote pups are born in dens, hollow trees, or under ledges, and weigh 200 to 500 g (0.44 to 1.10 lb) at birth. They are altricial, and are completely dependent on milk for their first 10 days. The incisors erupt at about 12 days, the canines at 16, and the second premolars at 21. Their eyes open after 10 days, by which point the pups become increasingly more mobile, walking by 20 days, and running at the age of six weeks. The parents begin supplementing the pup's diet with regurgitated solid food after 12–15 days. By the age of four to six weeks, when their milk teeth are fully functional, the pups are given small food items such as mice, rabbits, or pieces of ungulate carcasses, with lactation steadily decreasing after two months.[22] Unlike wolf pups, coyote pups begin seriously fighting (as opposed to play fighting) prior to engaging in play behavior. A common play behavior includes the coyote "hip-slam".[71] By three weeks of age, coyote pups bite each other with less inhibition than wolf pups. By the age of four to five weeks, pups have established dominance hierarchies, and are by then more likely to play rather than fight.[84] The male plays an active role in feeding, grooming, and guarding the pups, but abandons them if the female goes missing before the pups are completely weaned. The den is abandoned by June to July, and the pups follow their parents in patrolling their territory and hunting. Pups may leave their families in August, though can remain for much longer. The pups attain adult dimensions at eight months, and gain adult weight a month later.[22]

Territorial and sheltering behaviors

Individual feeding territories vary in size from 0.4 to 62 km2 (0.15 to 24 sq mi), with the general concentration of coyotes in a given area depending on food abundance, adequate denning sites, and competition with conspecifics and other predators. The coyote generally does not defend its territory outside of the denning season,[22] and is much less aggressive towards intruders than the wolf is, typically chasing and sparring with them, but rarely killing them.[85] Conflicts between coyotes can arise during times of food shortage.[22]

Like wolves, coyotes use a den (usually the deserted holes of other species) when gestating and rearing young, though they may occasionally give birth under sagebrushes in the open. Coyote dens can be located in canyons, washouts, coulees, banks, rock bluffs, or level ground. Some dens have been found under abandoned homestead shacks, grain bins, drainage pipes, railroad tracks, hollow logs, thickets, and thistles. The den is continuously dug and cleaned out by the female until the pups are born. Should the den be disturbed or infested with fleas, the pups are moved into another den. A coyote den can have several entrances and passages branching out from the main chamber.[86] A single den can be used year after year.[23]

Hunting and feeding behaviors

While the popular consensus is that olfaction is very important for hunting,[87] two studies that experimentally investigated the role of olfactory, auditory, and visual cues found that visual cues are the most important ones for hunting in red foxes[88] and coyotes.[89][90]

When hunting large prey, the coyote often works in pairs or small groups.[4] Success in killing large ungulates depends on factors such as snow depth and crust density. Younger animals usually avoid participating in such hunts, with the breeding pair typically doing most of the work.[23] Unlike the wolf, which attacks large prey from the rear, the coyote approaches from the front, lacerating its prey's head and throat. Like other canids, the coyote caches excess food.[91] Coyotes catch mouse-sized rodents by pouncing, whereas ground squirrels are chased. Although coyotes can live in large groups, small prey is typically caught singly.[23] Coyotes have been observed to kill porcupines in pairs, using their paws to flip the rodents on their backs, then attacking the soft underbelly. Only old and experienced coyotes can successfully prey on porcupines, with many predation attempts by young coyotes resulting in them being injured by their prey's quills.[92] Coyotes sometimes urinate on their food, possibly to claim ownership over it.[93]

Coyotes may occasionally form mutualistic hunting relationships with American badgers, assisting each other in digging up rodent prey.[94] The relationship between the two species may occasionally border on apparent "friendship", as some coyotes have been observed laying their heads on their badger companions or licking their faces without protest. The amicable interactions between coyotes and badgers were known to pre-Columbian civilizations, as shown on a Mexican jar dated to 1250–1300 CE depicting the relationship between the two.[95]

Communication

A coyote howling

Body language

Being both a gregarious and solitary animal, the variability of the coyote's visual and vocal repertoire is intermediate between that of the solitary foxes and the highly social wolf.[78] The aggressive behavior of the coyote bears more similarities to that of foxes than it does that of wolves and dogs. An aggressive coyote arches its back and lowers its tail.[96] Unlike dogs, which solicit playful behavior by performing a "play-bow" followed by a "play-leap", play in coyotes consists of a bow, followed by side-to-side head flexions and a series of "spins" and "dives". Although coyotes will sometimes bite their playmates' scruff as dogs do, they typically approach low, and make upward-directed bites.[97] Pups fight each other regardless of sex, while among adults, aggression is typically reserved for members of the same sex. Combatants approach each other waving their tails and snarling with their jaws open, though fights are typically silent. Males tend to fight in a vertical stance, while females fight on all four paws. Fights among females tend to be more serious than ones among males, as females seize their opponents' forelegs, throat, and shoulders.[96]

Vocalizations

The coyote has been described as "the most vocal of all [wild] North American mammals".[98][99] Its loudness and range of vocalizations was the cause for its binomial name Canis latrans, meaning "barking dog". At least 11 different vocalizations are known in adult coyotes. These sounds are divided into three categories: agonistic and alarm, greeting, and contact. Vocalizations of the first category include woofs, growls, huffs, barks, bark howls, yelps, and high-frequency whines. Woofs are used as low-intensity threats or alarms, and are usually heard near den sites, prompting the pups to immediately retreat into their burrows. Growls are used as threats at short distances, but have also been heard among pups playing and copulating males. Huffs are high-intensity threat vocalizations produced by rapid expiration of air. Barks can be classed as both long-distance threat vocalizations and as alarm calls. Bark howls may serve similar functions. Yelps are emitted as a sign of submission, while high-frequency whines are produced by dominant animals acknowledging the submission of subordinates. Greeting vocalizations include low-frequency whines, 'wow-oo-wows', and group yip howls. Low-frequency whines are emitted by submissive animals, and are usually accompanied by tail wagging and muzzle nibbling. The sound known as 'wow-oo-wow' has been described as a "greeting song". The group yip howl is emitted when two or more pack members reunite, and may be the final act of a complex greeting ceremony. Contact calls include lone howls and group howls, as well as the previously mentioned group yip howls. The lone howl is the most iconic sound of the coyote, and may serve the purpose of announcing the presence of a lone individual separated from its pack. Group howls are used as both substitute group yip howls and as responses to either lone howls, group howls, or group yip howls.[24]

Ecology

Habitat

Urban coyote in Bernal Heights, San Francisco

Prior to the near extermination of wolves and cougars, the coyote was most numerous in grasslands inhabited by bison, pronghorn, elk, and other deer, doing particularly well in short-grass areas with prairie dogs, though it was just as much at home in semiarid areas with sagebrush and jackrabbits or in deserts inhabited by cactus, kangaroo rats, and rattlesnakes. As long as it was not in direct competition with the wolf, the coyote ranged from the Sonoran Desert to the alpine regions of adjoining mountains or the plains and mountainous areas of Alberta. With the extermination of the wolf, the coyote's range expanded to encompass broken forests from the tropics of Guatemala and the northern slope of Alaska.[22]

Coyotes walk around 5–16 kilometres (3–10 mi) per day, often along trails such as logging roads and paths; they may use iced-over rivers as travel routes in winter. They are often crepuscular, being more active around evening and the beginning of the night than during the day. Like many canids, coyotes are competent swimmers, reported to be able to travel at least 0.8 kilometres (0.5 mi) across water.[100]

A Sonoran Desert coyote at the Sonora Desert Museum in Tucson Arizona

Diet

The coyote is roughly the North American equivalent to the Old World golden jackal.[101] Likewise, the coyote is highly versatile in its choice of food, but is primarily carnivorous, with 90% of its diet consisting of meat. Prey species include bison, deer, sheep, rabbits, rodents, birds, amphibians (except toads), lizards, snakes, fish, crustaceans, and insects. Coyotes may be picky over the prey they target, as animals such as shrews, moles, and brown rats do not occur in their diet in proportion to their numbers.[22] More unusual prey include fishers,[102] young black bear cubs,[103]harp seals[104] and rattlesnakes. Coyotes kill rattlesnakes mostly for food (but also to protect their pups at their dens) by teasing the snakes until they stretch out and then biting their heads and snapping and shaking the snakes.[105] In Death Valley, coyotes may consume great quantities of hawkmoth caterpillars or beetles in the spring flowering months.[106] Although coyotes prefer fresh meat, they will scavenge when the opportunity presents itself. Excluding the insects, fruit, and grass eaten, the coyote requires an estimated 600 g (1.3 lb) of food daily, or 250 kg (550 lb) annually.[22] The coyote readily cannibalizes the carcasses of conspecifics, with coyote fat having been successfully used by coyote hunters as a lure or poisoned bait.[6] The coyote's winter diet consists mainly of large ungulate carcasses, with very little plant matter. Rodent prey increases in importance during the spring, summer, and fall.[4]

The coyote feeds on a variety of different produce, including blackberries, blueberries, peaches, pears, apples, prickly pears, chapotes, persimmons, peanuts, watermelons, cantaloupes, and carrots. During the winter and early spring, the coyote eats large quantities of grass, such as green wheat blades. It sometimes eats unusual items such as cotton cake, soybean meal, domestic animal droppings, beans, and cultivated grain such as corn, wheat, and sorghum.[22]

Enemies and competitors

Comparative illustration of coyote and gray wolf
Mountain coyotes (C. l. lestes) cornering a juvenile cougar

In areas where the ranges of coyotes and gray wolves overlap, interference competition and predation by wolves has been hypothesized to limit local coyote densities. Coyote ranges expanded during the 19th and 20th centuries following the extirpation of wolves, while coyotes were driven to extinction on Isle Royale after wolves colonized the island in the 1940s. One study conducted in Yellowstone National Park, where both species coexist, concluded that the coyote population in the Lamar River Valley declined by 39% following the reintroduction of wolves in the 1990s, while coyote populations in wolf inhabited areas of the Grand Teton National Park are 33% lower than in areas where they are absent.[107][108] Wolves have been observed to not tolerate coyotes in their vicinity, though coyotes have been known to trail wolves to feed on their kills.[95]

Coyotes rarely kill healthy adult red foxes, and have been observed to feed or den alongside them, though they often kill foxes caught in traps. Coyotes may kill fox kits, but this is not a major source of mortality.[109] In southern California, coyotes frequently kill gray foxes, and these smaller canids tend to avoid areas with high coyote densities.[110]

Coyotes may compete with cougars in some areas. In the eastern Sierra Nevadas, coyotes compete with cougars over mule deer. Cougars usually outcompete coyotes, and may kill them occasionally, thus reducing coyote predation pressure on smaller carnivores such as foxes and bobcats.[111]

In some areas, coyotes share their ranges with bobcats. These two similarly sized species rarely physically confront one another, though bobcat populations tend to diminish in areas with high coyote densities.[112] However, several studies have demonstrated interference competition between coyotes and bobcats, and in all cases coyotes dominated the interaction.[113][114] Multiple researchers[115][116][117][114][118] reported instances of coyotes killing bobcats, whereas bobcats killing coyotes is more rare.[113] Coyotes attack bobcats using a bite-and-shake method similar to what is used on medium-sized prey. Coyotes (both single individuals and groups) have been known to occasionally kill bobcats – in most cases, the bobcats were relatively small specimens, such as adult females and juveniles.[114] However, coyote attacks (by an unknown number of coyotes) on adult male bobcats have occurred. In California, coyote and bobcat populations are not negatively correlated across different habitat types, but predation by coyotes is an important source of mortality in bobcats.[110] Biologist Stanley Paul Young noted that in his entire trapping career, he had never successfully saved a captured bobcat from being killed by coyotes, and wrote of two incidents wherein coyotes chased bobcats up trees.[95] Coyotes have been documented to directly kill Canadian lynx on occasion,[119][120][121] and compete with them for prey, especially snowshoe hares.[119] In some areas, including central Alberta, lynx are more abundant where coyotes are few, thus interactions with coyotes appears to influence lynx populations more than the availability of snowshoe hares.[122]

Range

Range of coyote subspecies as of 1978, (1) Mexican coyote, (2) San Pedro Martir coyote, (3) Salvador coyote, (4) southeastern coyote, (5) Belize coyote, (6) Honduras coyote, (7) Durango coyote, (8) northern coyote, (9) Tiburón Island coyote, (10) plains coyote, (11) mountain coyote, (12) Mearns' coyote, (13) Lower Rio Grande coyote, (14) California valley coyote, (15) peninsula coyote, (16) Texas plains coyote, (17) northeastern coyote, (18) northwest coast coyote, (19) Colima coyote, (20) eastern coyote[58]

Due to the coyote's wide range and abundance throughout North America, it is listed as least concern by the International Union for Conservation of Nature (IUCN).[2] The coyote's pre-Columbian range was limited to the Southwest and Plains regions of the United States and Canada, and northern and central Mexico. By the 19th century, the species expanded north and east, expanding further after 1900, coinciding with land conversion and the extirpation of wolves. By this time, its range encompassed all of the United States and Mexico, southward into Central America, and northward into most of Canada and Alaska. This expansion is ongoing, and the species now occupies the majority of areas between 8°N (Panama) and 70°N (northern Alaska).[2]

Although it was once widely believed that coyotes are recent immigrants to southern Mexico and Central America, aided in their expansion by deforestation, Pleistocene and Early Holocene records, as well as records from the pre-Columbian period and early European colonization show that the animal was present in the area long before modern times. Nevertheless, range expansion did occur south of Costa Rica during the late 1970s and northern Panama in the early 1980s, following the expansion of cattle-grazing lands into tropical rainforests. The coyote is predicted to appear in northern Belize in the near future, as the habitat there is favorable to the species.[123] Concerns have been raised of a possible expansion into South America through the Panamanian Isthmus, should the Darién Gap ever be closed by the Pan-American Highway.[124] This fear was partially confirmed in January 2013, when the species was recorded in eastern Panama's Chepo District, beyond the Panama Canal.[61]

Diseases and parasites

California valley coyote (C. l. ochropus) suffering from sarcoptic mange

Among large North American carnivores, the coyote probably carries the largest number of diseases and parasites, likely due to its wide range and varied diet.[125]Viral diseases known to infect coyotes include rabies, canine distemper, infectious canine hepatitis, four strains of equine encephalitis, and oral papillomatosis. By the late 1970s, serious rabies outbreaks in coyotes had ceased to be a problem for over 60 years, though sporadic cases every 1–5 years did occur. Distemper causes the deaths of many pups in the wild, though some specimens can survive infection. Tularemia, a bacterial disease, infects coyotes through their rodent and lagomorph prey, and can be deadly for pups.[126]

Coyotes can be infected by both demodectic and sarcoptic mange, the latter being the most common. Mite infestations are rare and incidental in coyotes, while tick infestations are more common, with seasonal peaks depending on locality (May–August in the Northwest, March–November in Arkansas). Coyotes are only rarely infested with lice, while fleas infest coyotes from puphood, though they may be more a source of irritation than serious illness. Pulex simulans is the most common species to infest coyotes, while Ctenocephalides canis tends to occur only in areas where coyotes and dogs (its primary host) inhabit the same area. Although coyotes are rarely host to flukes, they can nevertheless have serious effects on coyotes, particularly Nanophyetus salmincola, which can infect them with salmon poisoning disease, a disease with a 90% mortality rate. Trematode Metorchis conjunctus can also infect coyotes.[127]Tapeworms have been recorded to infest 60–95% of all coyotes examined. The most common species to infest coyotes is Taenia pisiformis and T. crassiceps, which uses cottontail rabbits as intermediate hosts. The largest species known in coyotes is T. hydatigena, which enters coyotes through infected ungulates, and can grow to lengths of 80 to 400 cm (31 to 157 in). Though once largely limited to wolves, Echinococcus granulosus has expanded to coyotes since the latter began colonizing former wolf ranges. The most frequent ascaroid roundworm in coyotes is Toxascaris leonina, which dwells in the coyote's small intestine and has no ill effects, except for causing the host to eat more frequently. Hookworms of the genus Ancylostoma infest coyotes throughout their range, being particularly prevalent in humid areas. In areas of high moisture, such as coastal Texas, coyotes can carry up to 250 hookworms each. The blood-drinking A. caninum is particularly dangerous, as it damages the coyote through blood loss and lung congestion. A 10-day-old pup can die from being host to as few as 25 A. caninum worms.[126]

Relationships with humans

In folklore and mythology

Spirit shield fashioned from coyote skull and crow feathers

The coyote features prominently as a trickster figure in the folktales of America's indigenous peoples, alternately assuming the form of an actual coyote or a man. As with other trickster figures, the coyote acts as a picaresque hero which rebels against social convention through deception and humor.[128] The coyote was likely given its trickster role in light of the actual animal's intelligence and adaptability; pre-Columbian American people observed its behavior, and their folkloric representations reflected its attributes.[129] After the European colonization of the Americas, it was reviled in Anglo-American culture as a cowardly and untrustworthy animal.[130] Unlike the gray wolf, which has undergone a radical improvement of its public image, cultural attitudes towards the coyote remain largely negative.[131]

The coyote plays a role in various mythologies and creation myths of Native American folklore. It is variously credited for having brought fire to humanity, releasing the bison into the world, and of having slain monsters by petrifying them. The Maidu creation myth has the coyote introducing work, suffering, and death to the world. Zuni folklore has the coyote bringing winter into the world by stealing light from the kachinas. Some tribes, such as the Chinook, Maidu, Paiute, Pawnee, Tohono O'odham, and Ute portray the coyote as the companion of the creator. In the Paiute creation myth, the coyote was created by the wolf as a companion, and the two created land by piling soil on the water-covered world. A Tohono O'odham flood myth has the coyote helping Montezuma survive a global deluge that destroys humanity. After the Great Mystery creates humanity, the coyote and Montezuma teach people how to live. The Crow creation myth portrays Old Man Coyote as the creator. In Navajo mythology, the coyote was present in the First World with First Man and First Woman, though a different version has it being created in the Fourth World. The Navajo coyote brings death into the world, explaining that without death, too many people would exist, thus no room to plant corn.[132]

Mural from Atetelco, Teotihuacán depicting coyote warriors.

Prior to the Spanish conquest of the Aztec Empire, the coyote played a significant role in Mesoamerican cosmology. The coyote symbolized military might in Classic era Teotihuacan, with warriors dressing up in coyote costumes to call upon its predatory power. The species continued to be linked to Central Mexican warrior cults in the centuries leading up to the post-Classic Aztec rule.[133] In Aztec mythology, Huehuecóyotl (meaning "old coyote"), the god of dance, music and carnality, is depicted in several codices as a man with a coyote's head.[134] He is sometimes depicted as a womanizer, responsible for bringing war into the world by seducing Xochiquetzal, the goddess of love.[135] Epigrapher David H. Kelley argued that the god Quetzalcoatl owed its origins to pre-Aztec Uto-Aztecan mythological depictions of the coyote, which is portrayed as mankind's "Elder Brother", a creator, seducer, trickster, and culture hero linked to the morning star.[136]

Attacks on humans

A sign discouraging people from feeding coyotes, which can lead to them habituating themselves to human presence, thus increasing the likelihood of attacks

Coyote attacks on humans are uncommon and rarely cause serious injuries, due to the relatively small size of the coyote, but have been increasingly frequent, especially in California. There have been two confirmed fatal attacks: one on a three-year-old named Kelly Keen in Glendale, California[137] and another on a nineteen-year-old named Taylor Mitchell in Nova Scotia, Canada.[138] In the 30 years leading up to March 2006, at least 160 attacks occurred in the United States, mostly in the Los Angeles County area.[139] Data from United States Department of Agriculture (USDA) Wildlife Services, the California Department of Fish and Game, and other sources show that while 41 attacks occurred during the period of 1988–1997, 48 attacks were verified from 1998 through 2003. The majority of these incidents occurred in Southern California near the suburban-wildland interface.[137]

In the absence of the harassment of coyotes practiced by rural people, urban coyotes are losing their fear of humans, which is further worsened by people intentionally or unintentionally feeding coyotes. In such situations, some coyotes have begun to act aggressively toward humans, chasing joggers and bicyclists, confronting people walking their dogs, and stalking small children.[137] Non-rabid coyotes in these areas sometimes target small children, mostly under the age of 10, though some adults have been bitten.[140]

Although media reports of such attacks generally identify the animals in question as simply "coyotes", research into the genetics of the eastern coyote indicates those involved in attacks in northeast North America, including Pennsylvania, New York, New England, and eastern Canada, may have actually been coywolves, hybrids of Canis latrans and C. lupus, not fully coyotes.[141]

Livestock and pet predation

Coyote confronting a dog

Coyotes are presently the most abundant livestock predators in western North America, causing the majority of sheep, goat, and cattle losses.[142] For example, according to the National Agricultural Statistics Service, coyotes were responsible for 60.5% of the 224,000 sheep deaths attributed to predation in 2004.[143] The total number of sheep deaths in 2004 comprised 2.22% of the total sheep and lamb population in the United States,[144] which, according to the National Agricultural Statistics Service USDA report, totaled 4.66 million and 7.80 million heads respectively as of July 1, 2005.[145] Because coyote populations are typically many times greater and more widely distributed than those of wolves, coyotes cause more overall predation losses. The United States government agents routinely shoot, poison, trap, and kill about 90,000 coyotes each year to protect livestock.[146] An Idaho census taken in 2005 showed that individual coyotes were 5% as likely to attack livestock as individual wolves.[147]

Livestock guardian dogs are commonly used to aggressively repel predators and have worked well in both fenced pasture and range operations.[148] A 1986 survey of sheep producers in the USA found that 82% reported the use of dogs represented an economic asset.[149]

Re-wilding cattle, which involves increasing the natural protective tendencies of cattle, is a method for controlling coyotes discussed by Temple Grandin of Colorado State University.[150] This method is gaining popularity among producers who allow their herds to calve on the range and whose cattle graze open pastures throughout the year.[151]

Coyotes typically bite the throat just behind the jaw and below the ear when attacking adult sheep or goats, with death commonly resulting from suffocation. Blood loss is usually a secondary cause of death. Calves and heavily fleeced sheep are killed by attacking the flanks or hindquarters, causing shock and blood loss. When attacking smaller prey, such as young lambs, the kill is made by biting the skull and spinal regions, causing massive tissue and bone damage. Small or young prey may be completely carried off, leaving only blood as evidence of a kill. Coyotes usually leave the hide and most of the skeleton of larger animals relatively intact, unless food is scarce, in which case they may leave only the largest bones. Scattered bits of wool, skin, and other parts are characteristic where coyotes feed extensively on larger carcasses.[142]

Coyote with a typical throat hold on domestic sheep

Tracks are an important factor in distinguishing coyote from dog predation. Coyote tracks tend to be more oval-shaped and compact than those of domestic dogs, and their claw marks are less prominent and the tracks tend to follow a straight line more closely than those of dogs. With the exception of sighthounds, most dogs of similar weight to coyotes have a slightly shorter stride.[142] Coyote kills can be distinguished from wolf kills by less damage to the underlying tissues in the former. Also, coyote scat tends to be smaller than wolf scat.[152][153]

Coyotes are often attracted to dog food and animals that are small enough to appear as prey. Items such as garbage, pet food, and sometimes feeding stations for birds and squirrels attract coyotes into backyards. About three to five pets attacked by coyotes are brought into the Animal Urgent Care hospital of South Orange County (California) each week, the majority of which are dogs, since cats typically do not survive the attacks.[154] Scat analysis collected near Claremont, California, revealed that coyotes relied heavily on pets as a food source in winter and spring.[137] At one location in Southern California, coyotes began relying on a colony of feral cats as a food source. Over time, the coyotes killed most of the cats, and then continued to eat the cat food placed daily at the colony site by people who were maintaining the cat colony.[137] Coyotes usually attack smaller-sized dogs, but they have been known to attack even large, powerful breeds such as the Rottweiler in exceptional cases.[155] Dogs larger than coyotes, such as greyhounds, are generally able to drive them off, and have been known to kill coyotes.[156] Smaller breeds are more likely to suffer injury or death.[140]

Uses

Prior to the mid-19th century, coyote fur was considered worthless. This changed with the diminution of beavers, and by 1860, the hunting of coyotes for their fur became a great source of income (75 cents to $1.50 per skin) for wolfers in the Great Plains. Coyote pelts were of significant economic importance during the early 1950s, ranging in price from $5 to $25 per pelt, depending on locality.[157] The coyote's fur is not durable enough to make rugs,[158] but can be used for coats and jackets, scarves, or muffs. The majority of pelts are used for making trimmings, such as coat collars and sleeves for women's clothing. Coyote fur is sometimes dyed black as imitation silver fox.[157]

Coyotes were occasionally eaten by trappers and mountain men during the western expansion. Coyotes sometimes featured in the feasts of the Plains Indians, and coyote pups were eaten by the indigenous people of San Gabriel, California. The taste of coyote meat has been likened to that of the wolf, and is more tender than pork when boiled. Coyote fat, when taken in the fall, has been used on occasion to grease leather or eaten as a spread.[159]

Tameability

Coyotes were probably semidomesticated by various pre-Columbian cultures. Some 19th-century writers wrote of coyotes being kept in native villages in the Great Plains. The coyote is easily tamed as a pup, but can become destructive as an adult.[160] Both full-blooded and hybrid coyotes can be playful and confiding with their owners, but are suspicious and shy of strangers,[69] though coyotes being tractable enough to be used for practical purposes like retrieving[161] and pointing have been recorded.[162] A tame coyote named "Butch", caught in the summer of 1945, had a short-lived career in cinema, appearing in Smoky and Ramrod before being shot while raiding a henhouse.[160]

Notes

  1. ^ The name "cased wolf" originates from the fact that the coyote's skin was historically cased like that of the muskrat, whereas the wolf's was spread out flat like the beaver's.[21]

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Bibliography

Further reading

Books
Video
Audiobook
  • Olson, Jack (May 2015). "The Last Coyote" (8 hours). Narrated by Gary MacFadden. Originally published as Slaughter The Animals, Poison The Earth, Simon & Schuster, Oct. 11, 1971.

External links

source: http://en.wikipedia.org/wiki/Coyote

Gray fox chase

Gray fox | Urocyon cinereoargenteus

Printed with acrylic spray paint on paper and touched up with water color and pencil. This print was designed to imagine the foxes as alive, with one chasing the other.

Gray fox info via Wikipedia:

Gray fox
Grey Fox (Urocyon cinereoargenteus).jpg
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Urocyon
Species: U. cinereoargenteus
Binomial name
Urocyon cinereoargenteus
(Schreber, 1775)
Leefgebied grijze vos.JPG
Gray fox range

The gray fox (Urocyon cinereoargenteus), or grey fox, is a carnivorous mammal of the family Canidae, widespread throughout northern and central America. This species and its only congener, the diminutive Channel Island fox (Urocyon littoralis), are the only living members of the genus Urocyon, which is considered to be the most basal of the living canids. Though it was once the most common fox in the eastern United States, and still is found there, human advancement and deforestation allowed the red fox to become more dominant. The Pacific States still have the gray fox as a dominant. It is the only American canid that can climb trees. Its specific epithet cinereoargenteus means "ashen silver".

Description

The gray fox is mainly distinguished from most other canids by its grizzled upper parts, black-tipped tail and strong neck, while the skull can be easily distinguished from all other North American canids by its widely separated temporal ridges that form a U-shape. There is little sexual dimorphism, save for the females being slightly smaller than males. The gray fox ranges from 76 to 112.5 cm (29.9 to 44.3 in) in total length. The tail measures 27.5 to 44.3 cm (10.8 to 17.4 in) of that length and its hind feet measure 100 to 150 mm (3.9 to 5.9 in). The gray fox typically weighs 3.6 to 7 kg (7.9 to 15.4 lb), though exceptionally can weigh as much as 9 kg (20 lb).[2][3][4] It is readily differentiated from the red fox by the lack of "black stockings" that stand out on the latter and the stripe of black hair that runs along the middle of the tail as well as individual guard hairs being banded with white, gray, and black.[5] The gray fox displays white on the ears, throat, chest, belly and hind legs. [5] In contrast to all Vulpes and related (Arctic and fennec) foxes, the gray fox has oval (instead of slit-like) pupils.[6]

The dental formula of the U. cinereoargenteus is 3.1.4.23.1.4.3 = 42. [5]

Origin and genetics

Gray fox kit at the Palo Alto Baylands in California

The gray fox appeared in North America during the mid-Pliocene epoch 3.6 million years ago (AEO) with the first fossil evidence found at the lower 111 Ranch site, Graham County, Arizona with contemporary mammals like the giant sloth, the elephant-like Cuvieronius, the large-headed llama, and the early small horses of Nannippus and Equus.[7] Genetic analyses of the fox-like canids confirmed that the gray fox is a distinct genus from the red foxes (Vulpes spp.). Genetically, the gray fox often clusters with two other ancient lineages, the east Asian raccoon dog (Nyctereutes procyonoides) and the African bat-eared fox (Otocyon megalotis).[8] Chromosome number is 2n=66.[9] Faunal remains at two northern California cave sites confirm the presence of the gray fox during the late Pleistocene.[10] Genetic analysis has shown that the gray fox migrated into the northeastern United States post-Pleistocene in association with the Medieval Climate Anomaly warming trend.[11] Recent mitochondrial genetic studies suggests divergence of North American eastern and western gray foxes in the Irvingtonian mid-Pleistocene into separate sister taxa.[12]

The gray fox's dwarf relative, the Channel Island fox, is likely descended from mainland gray foxes.[13] These foxes apparently were transported by humans to the islands and from island to island, and are descended from a minimum of 3–4 matrilineal founders.[12] The genus Urocyon is considered to be the most basal of the living canids.[14]

Distribution and habitat

The species occurs throughout most rocky, wooded, brushy regions of the southern half of North America from southern Canada to the northern part of South America (Venezuela and Colombia), excluding the mountains of northwestern United States.[15] It is the only canid whose natural range spans both North and South America.[16] In some areas, high populkation densities exist near brush-covered bluffs.[5]

Behavior

A yawning gray fox, northern Florida

The gray fox's ability to climb trees is shared only with the Asian raccoon dog among canids. Its strong, hooked claws allow it to scramble up trees to escape many predators, such as the domestic dog or the coyote,[17] or to reach tree-bound or arboreal food sources. It can climb branchless, vertical trunks to heights of 18 meters and jump from branch to branch.[18] It descends primarily by jumping from branch to branch, or by descending slowly backwards as a domestic cat would do. The gray fox is primarily nocturnal or crepuscular and makes its den in hollow trees, stumps or appropriated burrows during the day. Such gray fox tree dens may be located 30 ft above the ground.[6] Prior to European colonization of North America, the red fox was found primarily in boreal forest and the gray fox in deciduous forest, but now the red fox is dominant in most of the eastern United States since they are the more adaptable species to development and urbanization.[19] In areas where both red and gray foxes exist, the gray fox is dominant.[20]

Reproduction

Gray fox, showing black tail stripe, Sierra Nevada

The gray fox is assumed monogamous. The breeding season of the gray fox varies geographically; in Michigan, the gray fox mates in early March, in Alabama, breeding peaks occur in February. The gestation period lasts approximately 53 days. Litter size ranges from 1 to 7, with a mean of 3.8 young per female. Kits begin to hunt with their parents at the age of 3 months. By the time that they are four months old, the kits will have developed their permanent dentition and can now easily forage on their own. The family group remains together until the autumn, when the young reach sexual maturity, then they disperse.[citation needed]

Dens are used at any time during the year but mostly during whelping season. Dens are built in brushy or wooded regions and are less obvious than the dens of the red fox. Logs, trees, rocks, burrows, or abandoned dwellings serve as suitable den sites.[5]

Diet

A gray fox at night
Adult male and female gray fox

The gray fox is an omnivorous, solitary hunter. It frequently preys on the eastern cottontail (Sylvilagus floridanus) in the eastern U.S., though it will readily catch voles, shrews, and birds. In California, the gray fox primarily eats rodents, followed by lagomorphs, e.g. jackrabbit, brush rabbit, etc.[17] In some parts of the Western United States (such as in the Zion National Park in Utah), the gray fox is primarily insectivorous and herbivorous.[20] Fruit is an important component of the diet of the gray fox and they seek whatever fruits are readily available, generally eating more vegetable matter than does the red fox (Vulpes vulpes).[2]

Subspecies

Gray fox skull

There are 16 subspecies recognized for the gray fox.[9]

  • Urocyon cinereoargenteus borealis (New England)
  • Urocyon cinereoargenteus californicus (southern California)
  • Urocyon cinereoargenteus cinereoargenteus (eastern United States)
  • Urocyon cinereoargenteus costaricensis (Costa Rica)
  • Urocyon cinereoargenteus floridanus (Gulf states)
  • Urocyon cinereoargenteus fraterculus (Yucatán)
  • Urocyon cinereoargenteus furvus (Panama)
  • Urocyon cinereoargenteus guatemalae (southernmost Mexico south to Nicaragua)
  • Urocyon cinereoargenteus madrensis (southern Sonora, south-west Chihuahua, and north-west Durango)
  • Urocyon cinereoargenteus nigrirostris (south-west Mexico)
  • Urocyon cinereoargenteus ocythous (Central Plains states)
  • Urocyon cinereoargenteus orinomus (southern Mexico, Isthmus of Tehuantepec)
  • Urocyon cinereoargenteus peninsularis (Baja California)
  • Urocyon cinereoargenteus scottii (south-western United States and northern Mexico)
  • Urocyon cinereoargenteus townsendi (northern California and Oregon)
  • Urocyon cinereoargenteus venezuelae (Colombia and Venezuela)

Parasites

Parasites of gray fox include trematode Metorchis conjunctus.[21]

See also

References

  1. ^ Cypher; et al. (2008). "Urocyon cinereoargenteus". IUCN Red List of Threatened Species. Version 2008. International Union for Conservation of Nature. Retrieved 6 May 2008. 
  2. ^ a b "Urocyon cinereoargenteus". Animal Diversity Web. Retrieved 2007-08-19. 
  3. ^ Boitani, Luigi (1984) Simon & Schuster's Guide to Mammals. Simon & Schuster/Touchstone Books, ISBN 978-0-671-42805-1
  4. ^ Common Gray Fox (Urocyon cinereoargenteus). Nsrl.ttu.edu. Retrieved on 2013-01-26.
  5. ^ a b c d e Fritzell, Haroldson, Erik, Kurt (November 1982). "Urocyon cinereoargenteus". Mammalian Species: 1–8. 
  6. ^ a b Alderton, p. 122.
  7. ^ Paleobiology database, Collection 19656, Graham County, Arizona. Authority by the Dr. John Alroy, 18 February 1993.
  8. ^ Geffen, E.; Mercure, A.; Girman, D. J.; MacDonald, D. W.; Wayne, R. K. (Sep 1992). "Phylogenetic relationships of the fox-like canids: mitochondrial DNA restriction fragment, site and cytochrome b sequence analyses". Journal of Zoology, London. 228: 27–39. doi:10.1111/j.1469-7998.1992.tb04430.x. 
  9. ^ a b Fritzell, Erik K.; Haroldson, Kurt J. (1982). "Urocyon cinereoargenteus" (PDF). Mammalian Species. 189: 1–8. doi:10.2307/3503957. Retrieved 2011-11-27. 
  10. ^ Graham RW; Lundelius Jr. EL. FAUNMAP II: New data for North America with a temporal extension for the Blancan, Irvingtonian and early Rancholabrean (Report). FAUNMAP II Database, version 1.0; 2010. Retrieved December 13, 2015. 
  11. ^ Bozarth, Christine A.; Lance, Stacey L.; Civitello, David J.; Glenn, Julie L.; Maldonado, Jesus E. (2011). "Phylogeography of the gray fox (Urocyon cinereoargenteus) in the eastern United States" (PDF). Journal of Mammalogy. 92 (2): 283–294. doi:10.1644/10-MAMM-A-141.1. Retrieved 2011-11-27. 
  12. ^ a b Natalie S. Goddard; Mark J. Statham; Benjamin N. Sacks (August 19, 2015). "Mitochondrial Analysis of the Most Basal Canid Reveals Deep Divergence between Eastern and Western North American Gray Foxes (Urocyon spp.) and Ancient Roots in Pleistocene California". PLOS ONE. 10: e0136329. doi:10.1371/journal.pone.0136329. Retrieved 2015-12-13. 
  13. ^ Fuller, T.K.; Cypher, B. L. (2004). C. Sillero-Zubiri; M. Hoffman; D. W. Macdonald, eds. Gray fox Urocyon cinereoargenteus. pp. 92–97 in Canids: foxes, wolves, jackals, and dogs. Status survey and conservation action plan (PDF). Cambridge, United Kingdom: IUCN Publications. Retrieved 2011-11-27. 
  14. ^ Wayne, R. K.; Geffen, E; Girman, D. J.; Koepfli, K. P.; Lau, L. M.; Marshall, C. R. (1997). "Molecular Systematics of the Canidae". Systematic Biology. 46 (4): 622–653. doi:10.1093/sysbio/46.4.622. PMID 11975336. 
  15. ^ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 582. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  16. ^ Kurten B, Anderson E (1980). Pleistocene mammals of North America. New York: Columbia University. ISBN 978-0231037334. 
  17. ^ a b Fedriani, J. M.; Fuller, T. K.; Sauvajot, R. M.; York, E. C. (2000). "Competition and intraguild predation among three sympatric carnivores". Oecologia. 125 (2): 258–270. doi:10.1007/s004420000448. PMID 24595837. 
  18. ^ Sillero-Zubiri, Claudio; Hoffman, Michael; and MacDonald David W. (2004) Canids: Foxes, Wolves, Jackals, and Dogs: Status Survey and Conservation Action Plan. Gland, Switzerland and Cambridge, UK: IUCN. p. 95
  19. ^ Goddard-Taylor, Gayle (Winter 2005–2006). "The Silver Ghost: The life and times of the gray fox". Sanctuary: the Journal of the Massachusetts Audubon Society. Massachusetts Audubon Society. 45 (2): 13–15. 
  20. ^ a b Alderton, p. 124.
  21. ^ Mills J. H., Hirth R. S. (1968). "Lesions Caused by the Hepatic Trematode, Metorchis conjunctus, Cobbold, 1860: A Comparative Study in Carnivora". Journal of Small Animal Practice. 9 (1): 1–6. doi:10.1111/j.1748-5827.1968.tb04678.x. 

Bibliography

  • Alderton, David (1998). Foxes, Wolves, Lions, and Wild Dogs of the World. London: Blandford ISBN 081605715X

External links

source: https://en.wikipedia.org/wiki/Gray_fox

Sika deer skull

86W_Cervus_nippon_vers1_30x24

Sika Deer | Cervus nippon

Sika Deer Info via Wikipedia:

Sika deer
Cervus nippon hortulorum qtl5.jpg
Male (stag)
Male sika breeding calls, UK
Juni 2012 Alte Fasanerie Sikahirsch-Kuh.JPG
Female (hind)
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Cervidae
Subfamily: Cervinae
Genus: Cervus
Species: C. nippon
Binomial name
Cervus nippon
Temminck, 1838
Subspecies

See text

The sika deer (Cervus nippon) also known as the spotted deer or the Japanese deer, is a species of deer native to much of East Asia, and introduced to various other parts of the world. Previously found from northern Vietnam in the south to the Russian Far East in the north,[1] it is now uncommon in these areas, excluding Japan, where the species is overabundant.[2]

Etymology

Its name comes from shika (鹿), the Japanese word for "deer". In Japan the species is known as the nihonjika (ニホンジカ(日本鹿), lit. "Japan deer").

Taxonomy

The sika deer is a member of the genus Cervus, a group of deer also known as the "true deer".[citation needed] Formerly, sika were grouped together in this genus with nine other species. Now, only the sika and red deer remain, the latter being divided into three separate species: European red deer, central Asian red deer and American elk (though this remains controversial).[3]

Recent DNA evidence indicates these deer are not as closely related as previously thought, resulting in the creation of new species and genera. The genera Rucervus, Rusa, and Przewalskium are where most of the former Cervus species now belong. The ancestor of all Cervus species probably originated in central Asia and resembled sika deer.[4] All Cervus species can crossbreed and produce hybrids in areas where they coexist (for example, introduced sika hybridize with native red deer in the Scottish Highlands, where this is a serious threat to the gene pool of the red deer population).

Subspecies

Serious genetic pollution has occurred in many populations, especially in China. Therefore, the status of many subspecies remains unclear.[1] The status of C. n. hortulorum is particularly uncertain and might in fact be of mixed origin, hence it is not listed here.

Description

A Sika deer in Shiretoko Peninsula, Hokkaido, Japan.

The sika deer is one of the few deer species that does not lose its spots upon reaching maturity. Spot patterns vary with region. The mainland subspecies have larger and more obvious spots, in contrast to the Taiwanese and Japanese subspecies, whose spots are nearly invisible. Many introduced populations are from Japan, so they also lack significant spots.

The color of the fur ranges from mahogany to black, and white individuals are also known. During winter, the coat becomes darker and shaggier and the spots less prominent, and a mane forms on the back of the males' necks.[6] They are medium-sized herbivores, though they show notable size variation across their several subspecies and considerable sexual dimorphism, with males invariably much larger than females. They can vary from 50 to 110 cm (20 to 43 in) tall at the shoulder and from 95 to 180 cm (37 to 71 in) in head-and-body length. The tail measures about 7.5–13 cm (3.0–5.1 in) long.

The largest subspecies is the Manchurian sika deer (C. n. mantchuricus), in which males commonly weigh about 68–109 kg (150–240 lb) and females weigh 45–50 kg (99–110 lb), with large stags scaling up to 160 kg (350 lb). On the other end of the size spectrum, in the Japanese sika deer (C. n. nippon), males weigh 40–70 kg (88–154 lb) and females weigh 30–40 kg (66–88 lb).[7][8] All sikas are compact and dainty-legged, with short, trim, wedge-shaped heads and a boisterous disposition. When alarmed, they will often display a distinctive flared rump, much like the American elk.

Sika stags have stout, upright antlers with an extra buttress up from the brow tine and a very thick wall. A forward-facing intermediate tine breaks the line to the top, which is usually forked. Occasionally, sika antlers develop some palmation (flat areas). Females carry a pair of distinctive black bumps on the forehead. Antlers can range from 28 to 45 centimetres (11 to 18 in) to more than 80 centimetres (30 in), depending on the subspecies. Stags also have distinctive manes during their mating period (rut).

Behavior

The sika deer can be active throughout the day, though in areas with heavy human disturbance, they tend to be nocturnal. Seasonal migration is known to occur in mountainous areas, such as Japan, with winter ranges being up to 700 metres (2,300 ft) lower in elevation than summer ranges.[6]

Lifestyles vary between individuals, with some occurring alone while others are found in single-sex groups. Large herds will gather in autumn and winter. The sika deer is a highly vocal species, with over 10 individual sounds, ranging from soft whistles to loud screams.

A male young Sika deer in Nara.
A male Sika Deer calling, recorded at Wareham, Dorset, England, October 1964.

Sika males are territorial and keep harems of females during their mating period (rut), which peaks from early September through October, but may last well into the winter months. Territory size varies with habitat type and size of the buck; strong, prime bucks may hold up to 2 hectares (5 acres). Territories are marked with a series of shallow pits or "scrapes", into which the males urinate and from which emanates a strong, musky odor. Fights between rival males are sometimes fierce and long, and may even be fatal.

A fawn of sika deer at the Wildpark Alte Fasanerie in Klein-Auheim

The gestation period lasts for 7 months. Hinds give birth to a single fawn, weighing 4.5 to 7 kilograms (9.9 to 15.4 lb), which is nursed for up to 10 months. The fawn becomes independent 10 to 12 months after birth, and attains sexual maturity at 16 to 18 months in both sexes.[9]

The sika deer may interbreed with the red deer; hybrid descendants may have adaptive advantages over purebred relatives.[9]

The average lifespan is 15 to 18 years in captivity, although one case is recorded as living 25 years and 5 months.[9]

In Nara Prefecture, Japan, the deer are also known as "bowing deer", as they bow their heads before being fed special shika senbei (鹿せんべい, called "deer cookies"). However, deer bow heads to signal that they are about to headbutt. Therefore, when a human 'bows' to a deer, the deer will assume the same stance and may charge and injure the human. Deer headbutt both for play and to assert dominance, as do goats. Sika deer are found throughout the city of Nara and its many parks and temples like Tōdai-ji, as they are considered to be the messengers of the Shinto gods.[citation needed]

Habitat

Sika deer are found in the temperate and subtropical forests of eastern Asia, preferring areas with dense understory, and where snowfall does not exceed 10–20 cm (3.9–7.9 in). They tend to forage in patchy clearings of forests. Introduced populations are found in areas with similar habitats to their native ranges, including Western and Central Europe, Eastern United States, and New Zealand.

Population

The sika deer inhabits temperate and subtropical woodlands, which often occupy areas suitable for farming and other human exploitation. Its range encompasses some of the most densely populated areas in the world, where forests were cleared hundreds of years ago. Their population status varies significantly in different countries. Although the species as a whole is thriving, it is endangered and extinct in many areas.

Japan has by far the largest native sika population in the world. Though the exact population is uncertain, it is likely to be in the hundred thousand range and is still increasing,[citation needed] mainly due to recent conservation efforts and the extinction of its main predator, the wolf (Canis lupus hodophilax), over a century ago. Without its main enemy, the population of sika exploded and it is now overpopulated in many areas, posing a threat to both forests and farmlands. Efforts are now being made to control its population instead of conserving it. None of its subspecies is endangered except the Kerama deer (C. n. keramae) in the tiny Kerama Islands.[2] In 2015, Japanese Ministry of the Environment estimated the population at 3,080,000 in Japan, including Hokkaido.[10]

China used to have the largest population of sika, but thousands of years of hunting and habitat loss have reduced the population to less than 1,000. Of the five subspecies in China, the North China sika deer (C. n. mandarinus) is believed to be extinct in the wild since the 1930s; the Shanxi sika deer (C. n. grassianus) has not been seen in the wild since the 1980s and is also believed to be extinct in the wild. The status of Manchurian sika deer in China is unclear, though it is believed to be extinct, as well, and the sightings there are actually feral populations.

The South China sika deer (C. n. kopschi) and Sichuan sika deer (C. n. sichuanicus) are the only remaining subspecies in the wild. The former exists in fragmented populations of around 300 in southeast China, while the latter is found in a single population of over 400. The feral population is likely to be much higher than the wild, though most of them are descended from domesticated sikas of mixed subspecies. All of the subspecies are present in captivity, but a lack of suitable habitats and government efforts prevent their reintroduction.

The Formosan sika deer (C. n. taioanus) has been extinct for almost two decades before individuals from zoos were introduced to Kenting National Park; the population now numbers 200. Reintroduction programs are also under way in Vietnam, where the Vietnamese sika deer (C. n. pseudaxis) is extinct or nearly so.

Russia has a relatively large and stable population of 9,000 individuals of the Manchurian subspecies, but this is limited to a small area in Primorsky Krai. Small populations might exist in North Korea, but the political situation makes investigation impossible. The species is extinct in South Korea, with no plans for reintroduction.

Introduced populations

A Sika deer outside of a store on the island of Miyajima.

Sika deer have been introduced into a number of other countries, including Estonia, Latvia, Lithuania, Austria, Belgium, Denmark, France, Germany, Ireland, Netherlands, Norway, Switzerland, Russia, Romania, New Zealand, Australia, the Philippines (Jolo Island), Poland, Sweden, Finland, Canada, the United Kingdom, and the United States (Maryland, Oklahoma, Nebraska, Pennsylvania, Wisconsin, Virginia, Indiana, Michigan, Minnesota, Maine, Wyoming, Washington, and Kansas).[11] In many cases, they were originally introduced as ornamental animals in parklands, but have established themselves in the wild. On Spieden Island in the San Juan Islands of Washington, they were introduced as a game animal.

In the UK and Ireland, several distinct feral populations now exist.[12] Some of these are in isolated areas, for example on the island of Lundy, but others are contiguous with populations of the native red deer. Since the two species sometimes hybridize, there is a serious conservation concern.[13] In research which rated the negative impact of introduced mammals in Europe, the sika deer was found to be among the most damaging to the environment and economy, along with the brown rat and muskrat.[14]

In the 1900s, King Edward VII presented a pair of sika deer to John, the second Baron Montagu of Beaulieu. This pair escaped into Sowley Wood and were the basis of the sika to be found in the New Forest today.[citation needed] They were so prolific, culling had to be introduced in the 1930s to control their numbers.[15]

Hunting

Tsukioka Yoshitoshi Ukiyo-e depicting the Minamoto no Tsunemoto hunting a sika with a yumi.

Across its original range and in many areas to which it has been introduced, the sika is regarded as a particularly prized and elusive sportsman's quarry. In Britain, Ireland, and mainland Europe, sika display very different survival strategies and escape tactics from the indigenous deer. They have a marked tendency to use concealment in circumstances when red deer, for example, would flee, and have been seen to squat and lie belly-flat when danger threatens.

In the British Isles, sika are widely regarded as a serious threat to new and established woodlands, and public and private forestry bodies adopt policies of rigorous year-round culling.[16]

The main predator of sika deer include tiger and wolves[9], leopard, brown bear. Lynx and golden eagle targeted fawn.

Velvet antler

A tame deer wandering the streets of Miyajima, Japan.

Velvet antler (dried immature antlers) is a popular ingredient in traditional Chinese medicine, and sika in China were domesticated long ago for the antler trade, along with several other species. In Taiwan, both Formosan sika deer and Formosan sambar deer (Cervus unicolor swinhoei) have been farmed for velvet antlers. Japan is the only country in eastern Asia where sika deer were not farmed for velvet antlers.

Other deer raised for the antler trade were Thorold's deer (Cervus albirostris), central Asian red deer (Cervus affinis) and American elk (Cervus canadensis).

References

  1. ^ a b c Harris, R.B. (2008). "Cervus nippon". IUCN Red List of Threatened Species. Version 2008. International Union for Conservation of Nature. Retrieved 5 April 2009.  Database entry includes a brief justification of why this species is of least concern.
  2. ^ a b Kaji, Koichi; Takashi Saitoh; Hiroyuki Uno; Hiroyuki Matsuda; Kohji Yamamura. "Adaptive management of sika deer populations in Hokkaido, Japan: theory and practice" (PDF). Retrieved 19 January 2011. 
  3. ^ Ludt, Christian J.; Wolf Schroeder; Oswald Rottmann; Ralph Kuehn. "Mitochondrial DNA phylogeography of red deer (Cervus elaphus)" (PDF). Molecular Phylogenetics and Evolution 31 (2004) 1064–1083. Elsevier. Archived from the original (PDF) on 27 September 2004. Retrieved 6 October 2006. 
  4. ^ Geist, Valerius (1998). Deer of the World: Their Evolution, Behavior, and Ecology. Mechanicsburg, Pa: Stackpole Books. ISBN 0-8117-0496-3. 
  5. ^ "ITIS Standard Report Page: Cervus nippon soloensis". Retrieved 14 February 2016. 
  6. ^ a b Landesman, N. "Sika deer, Japanese deer". Ultimate Ungulate. [full citation needed]
  7. ^ Sika Deer. Bds.org.uk. Retrieved on 2012-08-23.
  8. ^ Nowak, R. M. 1991. Walker's Mammals of the World. Fifth Edition. Volume Two. Johns Hopkins University Press, Baltimore.
  9. ^ a b c d Landesman, N. "Cervus nippon". University of Michigan Museum of Zoology. Animal Diversity Web. 
  10. ^ "環境省_(お知らせ)改正鳥獣法に基づく指定管理鳥獣捕獲等事業の推進に向けたニホンジカ及びイノシシの生息状況等緊急調査事業の結果について". Retrieved 14 February 2016. 
  11. ^ "Sika Deer - North America Introduced - Big Game Hunting Records - Safari Club International Online Record Book". Retrieved 14 February 2016. 
  12. ^ https://docs.google.com/viewer?a=v&pid=sites&srcid=ZGVmYXVsdGRvbWFpbnxndWlkZXRvc3BoYWdudW18Z3g6NjA0ZDIzYmYwNWRmZDAwYw
  13. ^ "Cross-breeding 'threat' to deer". BBC. 22 January 2009. 
  14. ^ Rats top invasive mammals table. BBC. 7 May 2010.
  15. ^ "British Mammals: Sika Deer". BBC. 15 June 2007. Retrieved 8 October 2009. 
  16. ^ http://www.nonnativespecies.org/downloadDocument.cfm?id=355
  • "Cervus nippon". Integrated Taxonomic Information System. Retrieved 10 February 2006. 
  • Igota, H., Sakagura, M., Uno, H., Kaji, K., Maneko, M., Akamatsu, R., & Maekawa, (in press). Seasonal patterns of female sika deer in eastern Hokkaidō, Japan. Ecological Research, 19.

Further reading

O'Brien, D.J., Rooney, S.M. and Hayden, T.J. 2009. A differential vulnerability to hunting between the sexes in Sika-type calves. I. Nat. J. 30: 7- 9.

External links

source: http://en.wikipedia.org/wiki/Sika_deer

Domesticated Cow

90W_Bos_primagenius_vers1_24x18

Domesticated Cow Skull | Bos primigenius

Domesticated Cow info via Wikipedia:

Cattle
CH cow 2 cropped.jpg
A Swiss Braunvieh cow wearing a cowbell
Domesticated
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Bovidae
Subfamily: Bovinae
Genus: Bos
Species: B. taurus
Binomial name
Bos taurus
Linnaeus, 1758
Bovine range-2013-14-02.png
Bovine range
Synonyms
  • Bos primigenius
  • Bos indicus

Cattle—colloquially cows[note 1]—are the most common type of large domesticated ungulates. They are a prominent modern member of the subfamily Bovinae, are the most widespread species of the genus Bos, and are most commonly classified collectively as Bos taurus.

Cattle are commonly raised as livestock for meat (beef and veal), as dairy animals for milk and other dairy products, and as draft animals (oxen or bullocks that pull carts, plows and other implements). Other products include leather and dung for manure or fuel. In some regions, such as parts of India, cattle have significant religious meaning.

Around 10,500 years ago, cattle were domesticated from as few as 80 progenitors in southeast Turkey.[1] According to an estimate from 2011, there are 1.4 billion cattle in the world.[2] In 2009, cattle became one of the first livestock animals to have a fully mapped genome.[3] Some consider cattle the oldest form of wealth, and cattle raiding consequently one of the earliest forms of theft.

Taxonomy

A Holstein Fresian cow, a typical member of the Bos taurus taurus sub-species
Żubroń, a wisent and cattle hybrid

Cattle were originally identified as three separate species: Bos taurus, the European or "taurine" cattle (including similar types from Africa and Asia); Bos indicus, the zebu; and the extinct Bos primigenius, the aurochs. The aurochs is ancestral to both zebu and taurine cattle.[4] These have been reclassified as one species, Bos taurus, with three subspecies: Bos taurus primigenius, Bos taurus indicus, and Bos taurus taurus.[5][6]

Complicating the matter is the ability of cattle to interbreed with other closely related species. Hybrid individuals and even breeds exist, not only between taurine cattle and zebu (such as the sanga cattle, Bos taurus africanus), but also between one or both of these and some other members of the genus Bos – yaks (the dzo or yattle[7]), banteng, and gaur. Hybrids such as the beefalo breed can even occur between taurine cattle and either species of bison, leading some authors to consider them part of the genus Bos, as well.[8] The hybrid origin of some types may not be obvious – for example, genetic testing of the Dwarf Lulu breed, the only taurine-type cattle in Nepal, found them to be a mix of taurine cattle, zebu, and yak.[9] However, cattle cannot be successfully hybridized with more distantly related bovines such as water buffalo or African buffalo.

The aurochs originally ranged throughout Europe, North Africa, and much of Asia. In historical times, its range became restricted to Europe, and the last known individual died in Mazovia, Poland, in about 1627.[10] Breeders have attempted to recreate cattle of similar appearance to aurochs by crossing traditional types of domesticated cattle, creating the Heck cattle breed.

Etymology

Cattle did not originate as the term for bovine animals. It was borrowed from Anglo-Norman catel, itself from medieval Latin capitale 'principal sum of money, capital', itself derived in turn from Latin caput 'head'. Cattle originally meant movable personal property, especially livestock of any kind, as opposed to real property (the land, which also included wild or small free-roaming animals such as chickens — they were sold as part of the land).[11] The word is a variant of chattel (a unit of personal property) and closely related to capital in the economic sense.[12] The term replaced earlier Old English feoh 'cattle, property', which survives today as fee (cf. German: Vieh, Dutch: vee, Gothic: faihu).

The word "cow" came via Anglo-Saxon (plural ), from Common Indo-European gʷōus (genitive gʷowés) = "a bovine animal", compare Persian error: {{lang}}: unrecognized language code: per (help), Sanskrit go-, Welsh buwch.[13] The plural became ki or kie in Middle English, and an additional plural ending was often added, giving kine, kien, but also kies, kuin and others. This is the origin of the now archaic English plural, "kine". The Scots language singular is coo or cou, and the plural is "kye".

In older English sources such as the King James Version of the Bible, "cattle" refers to livestock, as opposed to "deer" which refers to wildlife. "Wild cattle" may refer to feral cattle or to undomesticated species of the genus Bos. Today, when used without any other qualifier, the modern meaning of "cattle" is usually restricted to domesticated bovines.[14]

Terminology

An Ongole bull
A Hereford bull

In general, the same words are used in different parts of the world, but with minor differences in the definitions. The terminology described here contrasts the differences in definition between the United Kingdom and other British-influenced parts of the world such as Canada, Australia, New Zealand, Ireland and the United States.[15]

  • An "intact" (i.e., not castrated) adult male is called a bull. A wild, young, unmarked bull is known as a micky in Australia.[16] An unbranded bovine of either sex is called a maverick in the US and Canada.
  • An adult female that has had a calf (or two, depending on regional usage) is a cow.
  • A young female before she has had a calf of her own[17] and is under three years of age is called a heifer (/ˈhɛfər/ HEF-ər).[18] A young female that has had only one calf is occasionally called a first-calf heifer.
  • Young cattle of both sexes are called calves until they are weaned, then weaners until they are a year old in some areas; in other areas, particularly with male beef cattle, they may be known as feeder calves or simply feeders. After that, they are referred to as yearlings or stirks[19] if between one and two years of age.[20]
  • A castrated male is called a steer in the United States; older steers are often called bullocks in other parts of the world,[21] but in North America this term refers to a young bull. Piker bullocks are micky bulls (uncastrated young male bulls) that were caught, castrated and then later lost.[16] In Australia, the term Japanese ox is used for grain-fed steers in the weight range of 500 to 650 kg that are destined for the Japanese meat trade.[22] In North America, draft cattle under four years old are called working steers. Improper or late castration on a bull results in it becoming a coarse steer known as a stag in Australia, Canada and New Zealand.[23] In some countries, an incompletely castrated male is known also as a rig.
  • A castrated male (occasionally a female or in some areas a bull) kept for draft purposes is called an ox (plural oxen); ox may also be used to refer to some carcass products from any adult cattle, such as ox-hide, ox-blood, oxtail, or ox-liver.[18]
  • A springer is a cow or heifer close to calving.[24]
  • In all cattle species, a female twin of a bull usually becomes an infertile partial intersex, and is called a freemartin.
  • Neat (horned oxen, from which neatsfoot oil is derived), beef (young ox) and beefing (young animal fit for slaughtering) are obsolete terms, although poll, pollard and polled cattle are still terms in use for naturally hornless animals, or in some areas also for those that have been disbudded or dehorned.
  • Cattle raised for human consumption are called beef cattle. Within the American beef cattle industry, the older term beef (plural beeves) is still used to refer to an animal of either sex. Some Australian, Canadian, New Zealand and British people use the term beast.[25]
  • Cattle bred specifically for milk production are called milking or dairy cattle;[15] a cow kept to provide milk for one family may be called a house cow or milker. A fresh cow is a dairy term for a cow or first-calf heifer who has recently given birth, or "freshened."
  • The adjective applying to cattle in general is usually bovine. The terms bull, cow and calf are also used by extension to denote the sex or age of other large animals, including whales, hippopotamuses, camels, elk and elephants.

Singular terminology issue

Cattle can only be used in the plural and not in the singular: it is a plurale tantum.[26] Thus one may refer to "three cattle" or "some cattle", but not "one cattle". No universally used singular form in modern English of cattle exists, other than the sex- and age-specific terms such as cow, bull, steer and heifer. Historically, "ox" was not a sex-specific term for adult cattle, but generally this is now used only for draft cattle, especially adult castrated males. The term is also incorporated into the names of other species, such as the musk ox and "grunting ox" (yak), and is used in some areas to describe certain cattle products such as ox-hide and oxtail.[27]

A Brahman calf

Cow is in general use as a singular for the collective cattle, despite the objections by those who insist it to be a female-specific term. Although the phrase "that cow is a bull" is absurd from a lexicographic standpoint, the word cow is easy to use when a singular is needed and the sex is unknown or irrelevant – when "there is a cow in the road", for example. Further, any herd of fully mature cattle in or near a pasture is statistically likely to consist mostly of cows, so the term is probably accurate even in the restrictive sense. Other than the few bulls needed for breeding, the vast majority of male cattle are castrated as calves and slaughtered for meat before the age of three years. Thus, in a pastured herd, any calves or herd bulls usually are clearly distinguishable from the cows due to distinctively different sizes and clear anatomical differences. Merriam-Webster, a US dictionary, recognizes the sex-nonspecific use of cow as an alternate definition,[28] whereas Collins and the OED, UK dictionaries, do not.

Colloquially, more general nonspecific terms may denote cattle when a singular form is needed. Australian, New Zealand and British farmers use the term beast or cattle beast. Bovine is also used in Britain. The term critter is common in the western United States and Canada, particularly when referring to young cattle.[29] In some areas of the American South (particularly the Appalachian region), where both dairy and beef cattle are present, an individual animal was once called a "beef critter", though that term is becoming archaic.

Other terminology

Cattle raised for human consumption are called beef cattle. Within the beef cattle industry in parts of the United States, the term beef (plural beeves) is still used in its archaic sense to refer to an animal of either sex. Cows of certain breeds that are kept for the milk they give are called dairy cows or milking cows (formerly milch cows). Most young male offspring of dairy cows are sold for veal, and may be referred to as veal calves.

The term dogies is used to describe orphaned calves in the context of ranch work in the American West, as in "Keep them dogies moving".[30] In some places, a cow kept to provide milk for one family is called a "house cow". Other obsolete terms for cattle include "neat" (this use survives in "neatsfoot oil", extracted from the feet and legs of cattle), and "beefing" (young animal fit for slaughter).

An onomatopoeic term for one of the most common sounds made by cattle is moo (also called lowing). There are a number of other sounds made by cattle, including calves bawling, and bulls bellowing. Bawling is most common for cows after weaning of a calf. The bullroarer makes a sound similar to a bull's territorial call.[31]

Characteristics

Anatomy

Bones are mounted on a black board
Displayed skeleton of a domestic cow

Cattle are large quadrupedal ungulate mammals with cloven hooves. Most breeds have horns, which can be as large as the Texas Longhorn or small like a scur. Careful genetic selection has allowed polled (hornless) cattle to become widespread.

Anatomy model of a cow

Digestive system

Cattle are ruminants, meaning their digestive system is highly specialized to allow the use of poorly digestible plants as food. Cattle have one stomach with four compartments, the rumen, reticulum, omasum, and abomasum, with the rumen being the largest compartment. The reticulum, the smallest compartment, is known as the "honeycomb". Cattle sometimes consume metal objects which are deposited in the reticulum and irritation from the metal objects causes hardware disease. The omasum's main function is to absorb water and nutrients from the digestible feed. The omasum is known as the "many plies". The abomasum is like the human stomach; this is why it is known as the "true stomach".

Cattle are known for regurgitating and re-chewing their food, known as cud chewing, like most ruminants. While the animal is feeding, the food is swallowed without being chewed and goes into the rumen for storage until the animal can find a quiet place to continue the digestion process. The food is regurgitated, a mouthful at a time, back up to the mouth, where the food, now called the cud, is chewed by the molars, grinding down the coarse vegetation to small particles. The cud is then swallowed again and further digested by specialized microorganisms in the rumen. These microbes are primarily responsible for decomposing cellulose and other carbohydrates into volatile fatty acids cattle use as their primary metabolic fuel. The microbes inside the rumen also synthesize amino acids from non-protein nitrogenous sources, such as urea and ammonia. As these microbes reproduce in the rumen, older generations die and their cells continue on through the digestive tract. These cells are then partially digested in the small intestines, allowing cattle to gain a high-quality protein source. These features allow cattle to thrive on grasses and other tough vegetation.

Gestation and size

The gestation period for a cow is about nine months long. A newborn calf's size can vary among breeds, but a typical calf weighs between 25 to 45 kg (55 to 99 lb). Adult size and weight vary significantly among breeds and sex. Steers are generally killed before reaching 750 kg (1,650 lb). Breeding stock may be allowed a longer lifespan, occasionally living as long as 25 years. The oldest recorded cow, Big Bertha, died at the age of 48 in 1993.

Reproduction

Reproductive system of a bovine female.
Ox testis.

On farms it is very common to use artificial insemination (AI)), a medically assisted reproduction technique consisting of the artificial deposition of semen in the female's genital tract.[32] It is used in cases where the spermatozoa can not reach the fallopian tubes or simply by choice of the owner of the animal. It consists of transferring, to the uterine cavity, spermatozoa previously collected and processed, with the selection of morphologically more normal and mobile spermatozoa.

A cow's udder contains two pairs of mammary glands, (commonly referred to as teats) creating four "quarters".[33] The front ones are referred to as fore quarters and the rear ones rear quarters.[34]

Bulls become fertile at about seven months of age. Their fertility is closely related to the size of their testicles, and one simple test of fertility is to measure the circumference of the scrotum: a young bull is likely to be fertile once this reaches 28 centimetres (11 in); that of a fully adult bull may be over 40 centimetres (16 in).[35][36]

Bulls have a fibro-elastic penis. Given the small amount of erectile tissue, there is little enlargement after erection. The penis is quite rigid when non-erect, and becomes even more rigid during erection. Protrusion is not affected much by erection, but more by relaxation of the retractor penis muscle and straightening of the sigmoid flexure.[37][38][39]

Induced ovulation can be manipulated to produce farming benefits. For example, to synchronise ovulation of the cattle to benefit dairy farming.

Weight

The world record for the heaviest bull was 1,740 kg (3,840 lb), a Chianina named Donetto, when he was exhibited at the Arezzo show in 1955.[40] The heaviest steer was eight-year-old ‘Old Ben’, a Shorthorn/Hereford cross weighing in at 2,140 kg (4,720 lb) in 1910.[41]

The weight of adult cattle varies, depending on the breed. Smaller kinds, such as Dexter and Jersey adults, range between 272 to 454 kg (600 to 1,000 lb). Large Continental breeds, such as Charolais, Marchigiana, Belgian Blue and Chianina, adults range from 635 to 1,134 kg (1,400 to 2,500 lb). British breeds, such as Hereford, Angus, and Shorthorn, mature between 454 to 907 kg (1,000 to 2,000 lb), occasionally higher, particularly with Angus and Hereford.[42]

Bulls will be a bit larger than cows of the same breed by a few hundred kilograms. Chianina bulls can weigh up to 1,500 kg (3,300 lb); British bulls, such as Angus and Hereford, can weigh as little as 907 kg (2,000 lb) to as much as 1,361 kg (3,000 lb).[citation needed]

It is difficult to generalize or average out the weight of all cattle because different kinds have different averages of weights. However, according to some sources, the average weight of all cattle is 753 kg (1,660 lb). Finishing steers in the feedlot average about 640 kg (1,410 lb); cows about 725 kg (1,600 lb), and bulls about 1,090 kg (2,400 lb).[43]

In the United States, the average weight of beef cattle has steadily increased, especially since the 1970s, requiring the building of new slaughterhouses able to handle larger carcasses. New packing plants in the 1980s stimulated a large increase in cattle weights.[44] Before 1790 beef cattle averaged only 160 kg (350 lb) net; and thereafter weights climbed steadily.[45][46]

Cognition

In laboratory studies, young cattle are able to memorize the locations of several food sources and retain this memory for at least 8 hours, although this declined after 12 hours.[47] Fifteen-month-old heifers learn more quickly than adult cows which have had either one or two calvings, but their longer-term memory is less stable.[48] Mature cattle perform well in spatial learning tasks and have a good long-term memory in these tests. Cattle tested in a radial arm maze are able to remember the locations of high-quality food for at least 30 days. Although they initially learn to avoid low-quality food, this memory diminishes over the same duration.[49] Under less artificial testing conditions, young cattle showed they were able to remember the location of feed for at least 48 days.[50] Cattle can make an association between a visual stimulus and food within 1 day – memory of this association can be retained for 1 year, despite a slight decay.[51]

Calves are capable of discrimination learning[52] and adult cattle compare favourably with small mammals in their learning ability in the Closed-field Test.[53]

They are also able to discriminate between familiar individuals, and among humans. Cattle can tell the difference between familiar and unfamiliar animals of the same species (conspecifics). Studies show they behave less aggressively toward familiar individuals when they are forming a new group.[54] Calves can also discriminate between humans based on previous experience, as shown by approaching those who handled them positively and avoiding those who handled them aversively.[55] Although cattle can discriminate between humans by their faces alone, they also use other cues such as the color of clothes when these are available.[56]

In audio play-back studies, calves prefer their own mother's vocalizations compared to the vocalizations of an unfamiliar mother.[57]

In laboratory studies using images, cattle can discriminate between images of the heads of cattle and other animal species.[58] They are also able to distinguish between familiar and unfamiliar conspecifics. Furthermore, they are able to categorize images as familiar and unfamiliar individuals.[54]

When mixed with other individuals, cloned calves from the same donor form subgroups, indicating that kin discrimination occurs and may be a basis of grouping behaviour. It has also been shown using images of cattle that both artificially inseminated and cloned calves have similar cognitive capacities of kin and non-kin discrimination.[59]

Cattle can recognize familiar individuals. Visual individual recognition is a more complex mental process than visual discrimination. It requires the recollection of the learned idiosyncratic identity of an individual that has been previously encountered and the formation of a mental representation.[60] By using 2-dimensional images of the heads of one cow (face, profiles, ¾ views), all the tested heifers showed individual recognition of familiar and unfamiliar individuals from their own breed. Furthermore, almost all the heifers recognized unknown individuals from different breeds, although this was achieved with greater difficulty. Individual recognition was most difficult when the visual features of the breed being tested were quite different from the breed in the image, for example, the breed being tested had no spots whereas the image was of a spotted breed.[61]

Cattle use visual/brain lateralisation in their visual scanning of novel and familiar stimuli.[62] Domestic cattle prefer to view novel stimuli with the left eye, i.e. using the right brain hemisphere (similar to horses, Australian magpies, chicks, toads and fish) but use the right eye, i.e. using the left hemisphere, for viewing familiar stimuli.[63]

Temperament and emotions

Ear postures of cows are studied as indicators of their emotional state and overall animal welfare.[64]

In cattle, temperament can affect production traits such as carcass and meat quality or milk yield as well as affecting the animal's overall health and reproduction. Cattle temperament is defined as "the consistent behavioral and physiological difference observed between individuals in response to a stressor or environmental challenge and is used to describe the relatively stable difference in the behavioral predisposition of an animal, which can be related to psychobiological mechanisms".[65] Generally, cattle temperament is assumed to be multidimensional. Five underlying categories of temperament traits have been proposed:[66]

  • shyness-boldness
  • exploration-avoidance
  • activity
  • aggressiveness
  • sociability

In a study on Holstein–Friesian heifers learning to press a panel to open a gate for access to a food reward, the researchers also recorded the heart rate and behavior of the heifers when moving along the race towards the food. When the heifers made clear improvements in learning, they had higher heart rates and tended to move more vigorously along the race. The researchers concluded this was an indication that cattle may react emotionally to their own learning improvement.[67]

Negative emotional states are associated with a bias toward negative (pessimistic) responses towards ambiguous cues in judgement tasks – as encapsulated in the question of "is the glass half empty or half full?". After separation from their mothers, Holstein calves showed such a cognitive bias indicative of low mood.[68] A similar study showed that after hot-iron disbudding (dehorning), calves had a similar negative bias indicating that post-operative pain following this routine procedure results in a negative change in emotional state.[69]

In studies of visual discrimination, the position of the ears has been used as an indicator of emotional state.[54] When cattle are stressed, this can be recognised by other cattle as it is communicated by alarm substances in the urine.[70]

Cattle are very gregarious and even short-term isolation is considered to cause severe psychological stress. When Aubrac and Fresian heifers are isolated, they increase their vocalizations and experience increased heart rate and plasma cortisol concentrations. These physiological changes are greater in Aubracs. When visual contact is re-instated, vocalisations rapidly decline, regardless of the familiarity of the returning cattle, however, heart rate decreases are greater if the returning cattle are familiar to the previously-isolated individual.[71] Mirrors have been used to reduce stress in isolated cattle.[72]

Senses

Cattle use all of the five widely recognized sensory modalities. These can assist in some complex behavioural patterns, for example, in grazing behaviour. Cattle eat mixed diets, but when given the opportunity, show a partial preference of approximately 70% clover and 30% grass. This preference has a diurnal pattern, with a stronger preference for clover in the morning, and the proportion of grass increasing towards the evening.[73]

Vision

Vision is the dominant sense in cattle and they obtain almost 50% of their information visually. [74]

Cattle are a prey animal and to assist predator detection, their eyes are located on the sides of their head rather than the front. This gives them a wide field of view of 330° but limits binocular vision (and therefore stereopsis) to 30° to 50° compared to 140° in humans.[54][75] This means they have a blind spot directly behind them. Cattle have good visual acuity (1/20)[54] but compared to humans, the visual accommodation of cattle is poor.[74]

Cattle have two kinds of color receptors in the cone cells of their retinas. This means that cattle are dichromatic, as are most other non-primate land mammals.[76][77] There are two to three rods per cone in the fovea centralis but five to six near the optic papilla.[75] Cattle can distinguish long wavelength colors (yellow, orange and red) much better than the shorter wavelengths (blue, grey and green). Calves are able to discriminate between long (red) and short (blue) or medium (green) wavelengths, but have limited ability to discriminate between the short and medium. They also approach handlers more quickly under red light.[78] Whilst having good color sensitivity, it is not as good as humans or sheep.[54]

A common misconception about cattle (particularly bulls) is that they are enraged by the color red (something provocative is often said to be "like a red flag to a bull"). This is a myth. In bullfighting, it is the movement of the red flag or cape that irritates the bull and incites it to charge.[79]

Taste

Cattle have a well-developed sense of taste and can distinguish the four primary tastes (sweet, salty, bitter and sour). They possess around 20,000 taste buds. The strength of taste perception depends on the individual's current food requirements. They avoid bitter-tasting foods (potentially toxic) and have a marked preference for sweet (high calorific value) and salty foods (electrolyte balance). Their sensitivity to sour-tasting foods helps them to maintain optimal ruminal pH.[74]

Plants have low levels of sodium and cattle have developed the capacity of seeking salt by taste and smell. If cattle become depleted of sodium salts, they show increased locomotion directed to searching for these. To assist in their search, the olfactory and gustatory receptors able to detect minute amounts of sodium salts increase their sensitivity as biochemical disruption develops with sodium salt depletion.[80][81]

Audition

Cattle hearing ranges from 23 Hz to 35 kHz. Their frequency of best sensitivity is 8 kHz and they have a lowest threshold of −21 db (re 20 μN/m−2), which means their hearing is more acute than horses (lowest threshold of 7 db).[82] Sound localization acuity thresholds are an average of 30°. This means that cattle are less able to localise sounds compared to goats (18°), dogs (8°) and humans (0.8°).[83] Because cattle have a broad foveal fields of view covering almost the entire horizon, they may not need very accurate locus information from their auditory systems to direct their gaze to a sound source.

Vocalisations are an important mode of communication amongst cattle and can provide information on the age, sex, dominance status and reproductive status of the caller. Calves can recognize their mothers using vocal and vocal behaviour may play a role by indicating estrus and competitive display by bulls.[84]

Olfaction and gustation

Several senses are used in social relationships between cattle

Cattle have a range of odiferous glands over their body including interdigital, infraorbital, inguinal and sebaceous glands, indicating that olfaction probably plays a large role in their social life. Both the primary olfactory system using the olfactory bulbs, and the secondary olfactory system using the vomeronasal organ are used.[85] This latter olfactory system is used in the flehmen response. There is evidence that when cattle are stressed, this can be recognised by other cattle and this is communicated by alarm substances in the urine.[70] The odour of dog faeces induces behavioural changes prior to cattle feeding, whereas the odours of urine from either stressed or non-stressed conspecifics and blood have no effect.[86]

In the laboratory, cattle can be trained to recognise conspecific individuals using olfaction only.[85]

In general, cattle use their sense of smell to "expand" on information detected by other sensory modalities. However, in the case of social and reproductive behaviours, olfaction is a key source of information.[74]

Touch

Cattle have tactile sensations detected mainly by mechanoreceptors, thermoreceptors and nociceptors in the skin and muzzle. These are used most frequently when cattle explore their environment.[74]

Magnetoreception

There is conflicting evidence for magnetoreception in cattle. One study reported that resting and grazing cattle tend to align their body axes in the geomagnetic North-South (N-S) direction.[87] In a follow-up study, cattle exposed to various magnetic fields directly beneath or in the vicinity of power lines trending in various magnetic directions exhibited distinct patterns of alignment.[88] However, in 2011, a group of Czech researchers reported their failed attempt to replicate the finding using Google Earth images.[89]

Behavior

Under natural conditions, calves stay with their mother until weaning at 8 to 11 months. Heifer and bull calves are equally attached to their mothers in the first few months of life.[90] Cattle are considered to be "hider" type animals, but in the artificial environment of small calving pens, close proximity between cow and calf is maintained by the mother at the first three calvings but this changes to being mediated by the calf after these. Primiparous dams show a higher incidence of abnormal maternal behavior.[91]

Video of a calf suckling

Beef-calves reared on the range suckle an average of 5.0 times every 24 hours with an average total time of 46 min spent suckling. There is a diurnal rhythm in suckling activity with peaks between 05:00–07:00, 10:00–13:00 and 17:00–21:00.[92]

Studies on the natural weaning of zebu cattle (Bos indicus) have shown that the cow weans her calves over a 2-week period, but after that, she continues to show strong affiliatory behavior with her offspring and preferentially chooses them for grooming and as grazing partners for at least 4–5 years.[93]

Reproductive behavior

Semi-wild Highland cattle heifers first give birth at 2 or 3 years of age and the timing of birth is synchronized with increases in natural food quality. Average calving interval is 391 days, and calving mortality within the first year of life is 5%.[94]

Dominance and leadership

One study showed that over a 4-year period, dominance relationships within a herd of semi-wild highland cattle were very firm. There were few overt aggressive conflicts and the majority of disputes were settled by agonistic (non-aggressive, competitive) behaviors that involved no physical contact between opponents (e.g. threatening and spontaneous withdrawing). Such agonistic behavior reduces the risk of injury. Dominance status depended on age and sex, with older animals generally being dominant to young ones and males dominant to females. Young bulls gained superior dominance status over adult cows when they reached about 2 years of age.[94]

As with many animal dominance hierarchies, dominance-associated aggressiveness does not correlate with rank position, but is closely related to rank distance between individuals.[94]

Dominance is maintained in several ways. Cattle often engage in mock fights where they test each other's strength in a non-aggressive way. Licking is primarily performed by subordinates and received by dominant animals. Mounting is a playful behavior shown by calves of both sexes and by bulls and sometimes by cows in estrus,[95] however, this is not a dominance related behavior as has been found in other species.[94]

The horns of cattle are "honest signals" used in mate selection. Furthermore, horned cattle attempt to keep greater distances between themselves and have fewer physical interactions than hornless cattle. This leads to more stable social relationships.[96]

In calves, the frequency of agonistic behavior decreases as space allowance increases, but this does not occur for changes in group size. However, in adult cattle, the number of agonistic encounters increases as the group size increases.[97]

Grazing behavior

When grazing, cattle vary several aspects of their bite, i.e. tongue and jaw movements, depending on characteristics of the plant they are eating. Bite area decreases with the density of the plants but increases with their height. Bite area is determined by the sweep of the tongue; in one study observing 750-kilogram (1,650 lb) steers, bite area reached a maximum of approximately 170 cm2 (30 sq in). Bite depth increases with the height of the plants. By adjusting their behavior, cattle obtain heavier bites in swards that are tall and sparse compared with short, dense swards of equal mass/area.[98] Cattle adjust other aspects of their grazing behavior in relation to the available food; foraging velocity decreases and intake rate increases in areas of abundant palatable forage.[99]

Cattle avoid grazing areas contaminated by the faeces of other cattle more strongly than they avoid areas contaminated by sheep,[100] but they do not avoid pasture contaminated by rabbit faeces.[101]

Genetics

In the 24 April 2009, edition of the journal Science, a team of researchers led by the National Institutes of Health and the US Department of Agriculture reported having mapped the bovine genome.[102] The scientists found cattle have about 22,000 genes, and 80% of their genes are shared with humans, and they share about 1000 genes with dogs and rodents, but are not found in humans. Using this bovine "HapMap", researchers can track the differences between the breeds that affect the quality of meat and milk yields.[103]

Behavioral traits of cattle can be as heritable as some production traits, and often, the two can be related.[104] The heritability of fear varies markedly in cattle from low (0.1) to high (0.53); such high variation is also found in pigs and sheep, probably due to differences in the methods used.[105] The heritability of temperament (response to isolation during handling) has been calculated as 0.36 and 0.46 for habituation to handling.[106] Rangeland assessments show that the heritability of aggressiveness in cattle is around 0.36.[107]

Quantitative trait loci (QTLs) have been found for a range of production and behavioral characteristics for both dairy and beef cattle.[108]

Domestication and husbandry

Texas Longhorns are a US breed

Cattle occupy a unique role in human history, having been domesticated since at least the early neolithic age.

Archeozoological and genetic data indicate that cattle were first domesticated from wild aurochs (Bos primigenius) approximately 10,500 years ago. There were two major areas of domestication: one in the area that is now Turkey, giving rise to the taurine line, and a second in the area that is now Pakistan, resulting in the indicine line.[109] Modern mitochondrial DNA variation indicates the taurine line may have arisen from as few as 80 aurochs tamed in the upper reaches of Mesopotamia near the villages of Çayönü Tepesi in southeastern Turkey and Dja'de el-Mughara in northern Iraq.[1]

Although European cattle are largely descended from the taurine lineage, gene flow from African cattle (partially of indicine origin) contributed substantial genomic components to both southern European cattle breeds and their New World descendants.[109] A study on 134 breeds showed that modern taurine cattle originated from Africa, Asia, North and South America, Australia, and Europe.[110] Some researchers have suggested that African taurine cattle are derived from a third independent domestication from North African aurochsen.[109]

Usage as money

As early as 9000 BC both grain and cattle were used as money or as barter (Davies) (the first grain remains found, considered to be evidence of pre-agricultural practice date to 17,000 BC).[111][112][113] Some evidence also exists to suggest that other animals, such as camels and goats, may have been used as currency in some parts of the world.[114] One of the advantages of using cattle as currency is that it allows the seller to set a fixed price. It even created the standard pricing. For example, two chickens were traded for one cow as cows were deemed to be more valuable than chickens.[112]

Modern husbandry

This Hereford is being inspected for ticks; cattle are often restrained or confined in cattle crushes (squeeze chutes) when given medical attention.
This young bovine has a nose ring to prevent it from suckling, which is usually to assist in weaning.

Cattle are often raised by allowing herds to graze on the grasses of large tracts of rangeland. Raising cattle in this manner allows the use of land that might be unsuitable for growing crops. The most common interactions with cattle involve daily feeding, cleaning and milking. Many routine husbandry practices involve ear tagging, dehorning, loading, medical operations, vaccinations and hoof care, as well as training for agricultural shows and preparations. Also, some cultural differences occur in working with cattle; the cattle husbandry of Fulani men rests on behavioural techniques, whereas in Europe, cattle are controlled primarily by physical means, such as fences.[115] Breeders use cattle husbandry to reduce M. bovis infection susceptibility by selective breeding and maintaining herd health to avoid concurrent disease.[116]

Cattle are farmed for beef, veal, dairy, and leather, and they are less commonly used for conservation grazing, simply to maintain grassland for wildlife – for example, in Epping Forest, England. They are often used in some of the most wild places for livestock. Depending on the breed, cattle can survive on hill grazing, heaths, marshes, moors and semidesert. Modern cattle are more commercial than older breeds and, having become more specialized, are less versatile. For this reason, many smaller farmers still favor old breeds, such as the Jersey dairy breed. In Portugal, Spain, southern France and some Latin American countries, bulls are used in the activity of bullfighting; Jallikattu in India is a bull taming sport radically different from European bullfighting, humans are unarmed and bulls are not killed. In many other countries bullfighting is illegal. Other activities such as bull riding are seen as part of a rodeo, especially in North America. Bull-leaping, a central ritual in Bronze Age Minoan culture (see Sacred Bull), still exists in southwestern France. In modern times, cattle are also entered into agricultural competitions. These competitions can involve live cattle or cattle carcases in hoof and hook events.

In terms of food intake by humans, consumption of cattle is less efficient than of grain or vegetables with regard to land use, and hence cattle grazing consumes more area than such other agricultural production when raised on grains.[117] Nonetheless, cattle and other forms of domesticated animals can sometimes help to use plant resources in areas not easily amenable to other forms of agriculture.

Sleep

The average sleep time of a domestic cow is about 4 hours a day.[118] Cattle do have a stay apparatus,[119] but do not sleep standing up,[120] they lie down to sleep deeply.[121] In spite of the urban legend, cows cannot be tipped over by people pushing on them.[122]

Economy

Holstein cattle are the primary dairy breed, bred for high milk production.

The meat of adult cattle is known as beef, and that of calves is veal. Other animal parts are also used as food products, including blood, liver, kidney, heart and oxtail. Cattle also produce milk, and dairy cattle are specifically bred to produce the large quantities of milk processed and sold for human consumption. Cattle today are the basis of a multibillion-dollar industry worldwide. The international trade in beef for 2000 was over $30 billion and represented only 23% of world beef production.[123] The production of milk, which is also made into cheese, butter, yogurt, and other dairy products, is comparable in economic size to beef production, and provides an important part of the food supply for many of the world's people. Cattle hides, used for leather to make shoes, couches and clothing, are another widespread product. Cattle remain broadly used as draft animals in many developing countries, such as India. Cattle are also used in some sporting games, including rodeo and bullfighting.

Cattle meat production

Cattle meat production (kt)
Country 2008 2009 2010 2011
Argentina 3132 3378 2630 2497
Australia 2132 2124 2630 2420
Brazil 9024 9395 9115 9030
China 5841 6060 6244 6182
Germany 1199 1190 1205 1170
Japan 520 517 515 500
USA 12163 11891 12046 11988

Source: Helgi Library,[124] World Bank, FAOSTAT

About half the world's meat comes from cattle.[125]

Dairy

Dairy farming and the milking of cattle was once performed largely by hand, but is now usually replaced by machine

Certain breeds of cattle, such as the Holstein-Friesian, are used to produce milk,[126][127] which can be processed into dairy products such as milk, cheese or yogurt. Dairy cattle are usually kept on specialized dairy farms designed for milk production. Most cows are milked twice per day, with milk processed at a dairy, which may be onsite at the farm or the milk may be shipped to a dairy plant for eventual sale of a dairy product.[128] For dairy cattle to continue producing milk, they must give birth to one calf per year. If the calf is male, it generally is slaughtered at a young age to produce veal.[129] They will continue to produce milk until three weeks before birth.[127] Over the last fifty years, dairy farming has become more intensive to increase the yield of milk produced by each cow. The Holstein-Friesian is the breed of dairy cow most common in the UK, Europe and the United States. It has been bred selectively to produce the highest yields of milk of any cow. Around 22 litres per day is average in the UK.[126][127]

Hides

Most cattle are not kept solely for hides, which are usually a by-product of beef production. Hides are most commonly used for leather which can be made into a variety of product including shoes. In 2012 India was the world's largest producer of cattle hides.[130]

Feral cattle

Feral cattle are defined as being 'cattle that are not domesticated or cultivated'.[131] Populations of feral cattle are known to come from and exist in: Australia, United States of America, Colombia, Argentina, Spain, France and many islands, including New Guinea, Hawaii, Galapagos, Juan Fernández Islands, Hispaniola (Dominican Republic and Haiti), Tristan da Cunha and Île Amsterdam,[132] two islands of Kuchinoshima[133] and Kazura Island next to Naru Island in Japan.[134][135]Chillingham cattle is sometimes regarded as a feral breed.[136]Aleutian wild cattles can be found on Aleutian Islands.[137] The "Kinmen cattle" which is dominantly found on Kinmen Island, Taiwan is mostly domesticated while smaller portion of the population is believed to live in the wild due to accidental releases.[138]

Other notable examples include cattles vicinity to Hong Kong (in the Shing Mun Country Park,[139] among Sai Kung District[140] and Lantau Island[141] and on Grass Island[142]), and semi-feral animals in Yangmingshan, Taiwan.[143]

Environmental impact

Cattle in dry landscape north of Alice Springs, Australia (CSIRO)

Gut flora in cattle include methanogens that produce methane as a byproduct of enteric fermentation, which cattle belch out. The same volume of atmospheric methane has a higher global warming potential than atmospheric carbon dioxide.[144][145] Methane belching from cattle can be reduced with genetic selection, immunization, rumen defaunation, diet modification and grazing management, among others.[146][147][148]

A report from the Food and Agriculture Organization (FAO) states that the livestock sector is "responsible for 18% of greenhouse gas emissions".[149] The IPCC estimates that Cattle and other livestock emit about 80 to 93 Megatonnes of methane per year,[150] accounting for an estimated 37% of anthropogenic methane emissions,[149] and additional methane is produced by anaerobic fermentation of manure in manure lagoons and other manure storage structures.[151] The net change in atmospheric methane content was recently about 1 Megatonne per year,[152] and in some recent years there has been no increase in atmospheric methane content.[153] While cattle fed forage actually produce more methane than grain-fed cattle, the increase may be offset by the increased carbon recapture of pastures, which recapture three times the CO2 of cropland used for grain.[154]

One of the cited changes suggested to reduce greenhouse gas emissions is intensification of the livestock industry, since intensification leads to less land for a given level of production. This assertion is supported by studies of the US beef production system, suggesting practices prevailing in 2007 involved 8.6% less fossil fuel use, 16.3% less greenhouse gas emissions, 12.1% less water use, and 33.0% less land use, per unit mass of beef produced, than those used in 1977.[155] The analysis took into account not only practices in feedlots, but also feed production (with less feed needed in more intensive production systems), forage-based cow-calf operations and back-grounding before cattle enter a feedlot (with more beef produced per head of cattle from those sources, in more intensive systems), and beef from animals derived from the dairy industry.

The number of American cattle kept in confined feedlot conditions fluctuates. From 1 January 2002 through 1 January 2012, there was no significant overall upward or downward trend in the number of US cattle on feed for slaughter, which averaged about 14.046 million head over that period.[156][157] Previously, the number had increased; it was 12.453 million in 1985.[158] Cattle on feed (for slaughter) numbered about 14.121 million on 1 January 2012, i.e. about 15.5% of the estimated inventory of 90.8 million US cattle (including calves) on that date. Of the 14.121 million, US cattle on feed (for slaughter) in operations with 1000 head or more were estimated to number 11.9 million.[157] Cattle feedlots in this size category correspond to the regulatory definition of "large" concentrated animal feeding operations (CAFOs) for cattle other than mature dairy cows or veal calves.[159] Significant numbers of dairy, as well as beef cattle, are confined in CAFOs, defined as "new and existing operations which stable or confine and feed or maintain for a total of 45 days or more in any 12-month period more than the number of animals specified"[160] where "[c]rops, vegetation, forage growth, or post-harvest residues are not sustained in the normal growing season over any portion of the lot or facility."[161] They may be designated as small, medium and large. Such designation of cattle CAFOs is according to cattle type (mature dairy cows, veal calves or other) and cattle numbers, but medium CAFOs are so designated only if they meet certain discharge criteria, and small CAFOs are designated only on a case-by-case basis.[162]

A CAFO that discharges pollutants is required to obtain a permit, which requires a plan to manage nutrient runoff, manure, chemicals, contaminants, and other wastewater pursuant to the US Clean Water Act.[163] The regulations involving CAFO permitting have been extensively litigated.[164] Commonly, CAFO wastewater and manure nutrients are applied to land at agronomic rates for use by forages or crops, and it is often assumed that various constituents of wastewater and manure, e.g. organic contaminants and pathogens, will be retained, inactivated or degraded on the land with application at such rates; however, additional evidence is needed to test reliability of such assumptions .[165] Concerns raised by opponents of CAFOs have included risks of contaminated water due to feedlot runoff,[166] soil erosion, human and animal exposure to toxic chemicals, development of antibiotic resistant bacteria and an increase in E. coli contamination.[167] While research suggests some of these impacts can be mitigated by developing wastewater treatment systems[166] and planting cover crops in larger setback zones,[168] the Union of Concerned Scientists released a report in 2008 concluding that CAFOs are generally unsustainable and externalize costs.[154]

An estimated 935,000 cattle operations were operating in the USA in 2010.[169] In 2001, the US Environmental Protection Agency (EPA) tallied 5,990 cattle CAFOs then regulated, consisting of beef (2,200), dairy (3,150), heifer (620) and veal operations (20).[170] Since that time, the EPA has established CAFOs as an enforcement priority. EPA enforcement highlights for fiscal year 2010 indicated enforcement actions against 12 cattle CAFOs for violations that included failures to obtain a permit, failures to meet the terms of a permit, and discharges of contaminated water.[171]

Cattle grazing in a high-elevation environment at the Big Pasture Plateau, Slovenia

Another concern is manure, which if not well-managed, can lead to adverse environmental consequences. However, manure also is a valuable source of nutrients and organic matter when used as a fertilizer.[172] Manure was used as a fertilizer on about 15.8 million acres of US cropland in 2006, with manure from cattle accounting for nearly 70% of manure applications to soybeans and about 80% or more of manure applications to corn, wheat, barley, oats and sorghum.[173] Substitution of manure for synthetic fertilizers in crop production can be environmentally significant, as between 43 and 88 megajoules of fossil fuel energy would be used per kg of nitrogen in manufacture of synthetic nitrogenous fertilizers.[174]

Grazing by cattle at low intensities can create a favourable environment for native herbs and forbs; in many world regions, though, cattle are reducing biodiversity due to overgrazing.[175] A survey of refuge managers on 123 National Wildlife Refuges in the US tallied 86 species of wildlife considered positively affected and 82 considered negatively affected by refuge cattle grazing or haying.[176] Proper management of pastures, notably managed intensive rotational grazing and grazing at low intensities can lead to less use of fossil fuel energy, increased recapture of carbon dioxide, fewer ammonia emissions into the atmosphere, reduced soil erosion, better air quality, and less water pollution.[154]

Health

The veterinary discipline dealing with cattle and cattle diseases (bovine veterinary) is called buiatrics.[177] Veterinarians and professionals working on cattle health issues are pooled in the World Association for Buiatrics, founded in 1960.[178] National associations and affiliates also exist.[179]

Cattle diseases were in the center of attention in the 1980s and 1990s when the Bovine spongiform encephalopathy (BSE), also known as mad cow disease, was of concern. Cattle might catch and develop various other diseases, like blackleg, bluetongue, foot rot too.[180][181][182]

In most states, as cattle health is not only a veterinarian issue, but also a public health issue, public health and food safety standards and farming regulations directly affect the daily work of farmers who keep cattle.[183] However, said rules change frequently and are often debated. For instance, in the U.K., it was proposed in 2011 that milk from tuberculosis-infected cattle should be allowed to enter the food chain.[184] Internal food safety regulations might affect a country's trade policy as well. For example, the United States has just reviewed its beef import rules according to the "mad cow standards"; while Mexico forbids the entry of cattle who are older than 30 months.[185]

Cow urine is commonly used in India for internal medical purposes.[186][187] It is distilled and then consumed by patients seeking treatment for a wide variety of illnesses.[188] At present, no conclusive medical evidence shows this has any effect.[189] However, an Indian medicine containing cow urine has already obtained U.S. patents.[190]

Digital dermatitis is caused by the bacteria from the genus Treponema. It differs from foot rot and can appear under unsanitary conditions such as poor hygiene or inadequate hoof trimming, among other causes. It primarily affects dairy cattle and has been known to lower the quantity of milk produced, however the milk quality remains unaffected. Cattle are also susceptible to ringworm caused by the fungus, Trichophyton verrucosum, a contagious skin disease which may be transferred to humans exposed to infected cows.[191]

Mycobacterium vaccae is a non pathogenic, possibly even beneficial bacteria, that is seen naturally in soil;[192] that was first isolated from cow dung.[193]

Effect of high stocking density

Stocking density refers to the number of animals within a specified area. When stocking density reaches high levels, the behavioural needs of the animals may not be met. This can negatively influence health, welfare and production performance.[194]

The effect of overstocking in cows can have a negative effect on milk production and reproduction rates which are two very important traits for dairy farmers. Overcrowding of cows in barns has been found to reduced feeding, resting and rumination.[194] Although they consume the same amount of dry matter within the span of a day, they consume the food at a much more rapid rate, and this behaviour in cows can lead to further complications.[195] The feeding behaviour of cows during their post-milking period is very important as it has been proven that the longer animals can eat after milking, the longer they will be standing up and therefore causing less contamination to the teat ends.[196] This is necessary to reduce the risk of mastitis as infection has been shown to increase the chances of embryonic loss.[197] Sufficient rest is important for dairy cows because it is during this period that their resting blood flow increases up to 50%, this is directly proportionate to milk production.[196] Each additional hour of rest can be seen to translate to 2 to 3.5 more pounds of milk per cow daily. Stocking densities of anything over 120% have been shown to decrease the amount of time cows spend lying down.[198]

Cortisol is an important stress hormone; its plasma concentrations increase greatly when subjected to high levels of stress.[199] Increased concentration levels of cortisol have been associated with significant increases in gonadotrophin levels and lowered progestin levels. Reduction of stress is important in the reproductive state of cows as an increase in gonadotrophin and lowered progesterone levels may impinge on the ovulatory and lutenization process and to reduce the chances of successful implantation.[200] A high cortisol level will also stimulate the degradation of fats and proteins which may make it difficult for the animal to sustain its pregnancy if implanted successfully.[199]

Oxen

Draft Zebus in Mumbai, Maharashtra, India

Oxen (singular ox) are cattle trained as draft animals. Often they are adult, castrated males of larger breeds, although females and bulls are also used in some areas. Usually, an ox is over four years old due to the need for training and to allow it to grow to full size. Oxen are used for plowing, transport, hauling cargo, grain-grinding by trampling or by powering machines, irrigation by powering pumps, and wagon drawing. Oxen were commonly used to skid logs in forests, and sometimes still are, in low-impact, select-cut logging. Oxen are most often used in teams of two, paired, for light work such as carting, with additional pairs added when more power is required, sometimes up to a total of 20 or more.

Oxen used in Plowing

An ox is a mature bovine which has learned to respond appropriately to a teamster's signals. These signals are given by verbal commands or by noise (whip cracks). Verbal commands vary according to dialect and local tradition. In one tradition in North America, the commands are:[citation needed]

  • "Back up": go backwards
  • "Gee": turn right
  • "Get up": walk forward
  • "Haw": turn left
  • "Whoa": stop
Riding an ox in Hova, Sweden

Oxen can pull harder and longer than horses. Though not as fast as horses, they are less prone to injury because they are more sure-footed.

Many oxen are used worldwide, especially in developing countries. About 11.3 million draft oxen are used in sub-Saharan Africa.[201] In India, the number of draft cattle in 1998 was estimated at 65.7 million head.[202] About half the world's crop production is thought to depend on land preparation (such as plowing) made possible by animal traction.[203]

The "Ure-Ox" (Aurochs) by Edward Topsell, 1658

Religion, traditions and folklore

Hindu tradition

Cattle are venerated within the Hindu religion of India. In the Vedic period they were a symbol of plenty [204]:130 and were frequently slaughtered. In later times they gradually acquired their present status. According to the Mahabharata they are to be treated with the same respect 'as one's mother'.[205] In the middle of the first millennium, the consumption of beef began to be disfavoured by lawgivers.[204]:144 Although there has never been any cow-goddesses or temples dedicated to them,[204]:146 cows appear in numerous stories from the Vedas and Puranas. The deity Krishna was brought up in a family of cowherders, and given the name Govinda (protector of the cows). Also, Shiva is traditionally said to ride on the back of a bull named Nandi.

Milk and milk products were used in Vedic rituals.[204]:130 In the postvedic period products of the cow – milk, curd, ghee, but also cow dung and urine (gomutra), or the combination of these five (panchagavya) – began to assume an increasingly important role in ritual purification and expiation.[204]:130–1

Veneration of the cow has become a symbol of the identity of Hindus as a community,[204]:20 especially since the end of the 19th century. Slaughter of cows (including oxen, bulls and calves) is forbidden by law in several states of the Indian Union. McDonald's outlets in India do not serve any beef burgers. In Maharaja Ranjit Singh's empire of the early 19th century, the killing of a cow was punishable by death.[206]

Other traditions

Legend of the founding of Durham Cathedral is that monks carrying the body of Saint Cuthbert were led to the location by a milk maid who had lost her dun cow, which was found resting on the spot.
An idealized depiction of girl cow herders in 19th-century Norway by Knud Bergslien.
  • The Evangelist St. Luke is depicted as an ox in Christian art.
  • In Judaism, as described in Numbers 19:2, the ashes of a sacrificed unblemished red heifer that has never been yoked can be used for ritual purification of people who came into contact with a corpse.
  • The ox is one of the 12-year cycle of animals which appear in the Chinese zodiac related to the Chinese calendar. See: Ox (Zodiac).
  • The constellation Taurus represents a bull.
  • An apocryphal story has it that a cow started the Great Chicago Fire by kicking over a kerosene lamp. Michael Ahern, the reporter who created the cow story, admitted in 1893 that he had fabricated it for more colorful copy.
  • On 18 February 1930, Elm Farm Ollie became the first cow to fly in an airplane and also the first cow to be milked in an airplane.
  • The first known law requiring branding in North America was enacted on 5 February 1644, by Connecticut. It said that all cattle and pigs had to have a registered brand or earmark by 1 May 1644.[207]
  • The akabeko (赤べこ, red cow) is a traditional toy from the Aizu region of Japan that is thought to ward off illness.[208]
  • The case of Sherwood v. Walker—involving a supposedly barren heifer that was actually pregnant—-first enunciated the concept of mutual mistake as a means of destroying the meeting of the minds in contract law.[citation needed]
  • The Fulani of West Africa are the world's largest nomadic cattle-herders.
  • The Maasai tribe of East Africa traditionally believe their god Engai entitled them to divine rights to the ownership of all cattle on earth.[209]

In heraldry

Cattle are typically represented in heraldry by the bull.

Population

For 2013, the FAO estimated global cattle numbers at 1.47 billion.[210] Regionally, the FAO estimate for 2013 includes: Asia 495 million; South America 348 million; Africa 305 million; Europe 122 million; North America 102 million; Central America 46 million; Oceania 42 million; and Caribbean 9 million. The following table shows the cattle population in 2009.[211]

As of 2003[update], Africa had about 231 million head of cattle, raised in both traditional and non-traditional systems, but often an "integral" part of the culture and way of life.[212]

Cattle population
Region 2009 2013 [213]
 India 285,000,000 (By 2003)[214] 194,655,285
 Brazil 187,087,000 186,646,205
 China 139,721,000 102,668,900
 USA 96,669,000 96,956,461
 European Union 87,650,000
 Argentina 51,062,000 52,509,049
 Pakistan 38,300,000 26,007,848
 Australia 29,202,000 27,249,291
 Mexico 26,489,000 31,222,196
 Bangladesh 22,976,000 22,844,190
 Russia 18,370,000 28,685,315
 South Africa 14,187,000 13,526,296
 Canada 13,945,000 13,287,866
Others 49,756,000

Gallery

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