Canis latrans, Say, 1823

Don E. Wilson & Russell A. Mittermeier, 2009, Canidae, Handbook of the Mammals of the World – Volume 1 Carnivores, Barcelona: Lynx Edicions, pp. 352-446 : 415-416

3.

Coyote

Canis latrans

French: Coyote / German: Kojote / Spanish: Coyote

Other common names: Brush Wolf, Prairie Wolf

Taxonomy. Canis latrans Say, 1823 ,

Nebraska, USA.

The ancestral Coyote, C. lepophagus, is believed to have become widespread throughout North America by the late Pliocene. In the north-eastern USA, the eastern Coyote may be a subspecies with some introgression of wolf and dog genes. Nineteen subspecies have been recognized. However, the taxonomic validity of some subspecies is questionable.

Subspecies and Distribution.

C. l. latrans Say, 1823 — S Canada and USA (Great Plains region).

C. l. cagottis Hamilton-Smith, 1839 — SE Mexico.

C. l. clepticus Elliot, 1903 — Mexico (N Baja California) and USA (S California).

C. l. dickeyi Nelson, 1932 — Costa Rica, El Salvador, W Honduras, Nicaragua, and Panama.

C. l. frustror Woodhouse, 1850 — USA (Missouri, Kansas, parts of Oklahoma & E Texas).

C. l. goldmani Merriam, 1904 — Belize, Guatemala, and S Mexico.

C. l. hondurensis Goldman, 1936 — E Honduras.

C. l. impavidus J. A. Allen, 1903 - W Mexico.

C. l. incolatus Hall, 1934 - Alaska and NW Canada.

C. l. ¡jamesi Townsend. 1912 - Mexico (Tiburon l. Baia California).

c. [l.lestes lam Merriam, 1897 1897 - sw canada Canada and w USA USA Õnzenndunzain Region & NW).

v C. l. mearnsi Merriam, 1897 - NW Mexico and SW USA.

C. l. mıcrodon Merriam, 1897 — NE Mexico and S USA (S Texas).

C. l. ochropus Eschscholtz, 1829 - W USA (W coast).

C. l. peninsulae Merriam, 1897 — Mexico (S Baja California).

C. l. texensıs Bailey, 1905 - S USA (W Texas 8c New Mexico).

C. l. thamnos Jackson, 1949 - N-C Canada and E USA.

C. l. umpquensıs Jackson. 1949 - USA (NW coast).

C. l. vigilis Merriam. 1897 — SW Mexico.

Coyotes did not originally occur on the USA E coast or Florida. They (probably thamnos) have expanded into the area with the clearing of forests and been introduced to Florida and Georgia (subspecies unknown).

Descriptive notes. Head-body 74-94 cm for males and 74-94 cm for females,tail 29— 36- 3 cm for males and 26-34- 3 cm for females; weight 7-8-15- 8 kg for males and 7-7- 14- 5 kg for females. Slender appearance with a long, pointed nose, large pointed ears, slender legs with small feet, and a bushy tail. Size varies geographically, although adult males are heavier and larger than adult females. Coyotes range in color from pure gray to rufous; melanistic Coyotes are rare. Fur texture and color varies geographically: northern subspecies have long coarse hair. Coyotes in the desert tend to be fulvous in color, while Coyotes at higher latitudes are darker and grayer. The belly and throat are paler than the rest of the body and have a mantle of darker hair over the shoulders. The tip of the tail is usually black. Hairs are about 50-90 mm long, mane hairs tend to be 80-110 mm long. Pelage during summer is shorter than in winter. The dental formulais13/3,C1/1,PM 4/4, M 2/3 = 42.

Habitat. Coyotes utilize almost all available habitats throughout their range including prairie, forest, desert, mountain, and tropical ecosystems. Their ability to exploit human resources also allows them to occupy urban areas. Water availability may limit distribution in some desert environments.

Food and Feeding. Opportunistic, generalist predators that eat a wide variety of food items, ranging from fruit and insects to small mammals to large ungulates and live-stock, typically consuming items in relation to availability. Livestock and wild ungulates are often consumed as carrion, but predation on large ungulates (native and domestic) does occur. Predation on neonates of native ungulates can be high during fawning. Coyotes in suburban areas are adept at exploiting human-made food resources and will readily consume refuse, pet food or other human-related items. Studies of predatory behavior show that Coyote age, habitat, and wind and snow conditions all influence their ability to capture small mammals. Coyotes hunt small mammals alone, even when pack size is large. When preying on native ungulates, cooperation among pack members may facilitate the capture of prey, but is not essential. Environmental factors are important to the success of an attack on adult ungulates, as is the presence of the alpha Coyote pair, and younger animals generally do not participate. The number of Coyotes is not as important as which individuals are involved in the attack. The outcome is also affected by the ability of the quarry to escape into water, its defensive abilities, and its nutritional state. In areas with an ungulate prey base in winter, competition with other sympatric carnivores for a carcass may be intense and there can even be competition among members of the same pack.

Activity patterns. Coyotes may be active throughout the day, but they tend to be more active during the early morning and around sunset. Activity patterns change seasonally (e.g. during winter, when there is a change in the food base in some areas) or in response to human disturbance and persecution.

Movements, Home range and Social organization. The basic social unit is the bonded alpha pair, which may breed for up to 10-12 years. Associate animals often remain in the pack and possibly inherit the pack or displace members of the breeding pair and become alphas themselves. Associates participate in territorial maintenance and pup rearing, but not to the extent of the alpha pair. Other Coyotes exist outside of the resident packs as transient or nomadic individuals. Transients travel alone over larger areas and do not breed, but will move into territories when vacancies occur. One factor that may affect Coyote sociality is prey size or prey biomass. In populations where rodents are the major prey, Coyotes tend to be in pairs or trios. In populations where Elk and deer are available, large packs of up to ten individuals may form. Coyotes are territorial, with a dominance hierarchy within each resident pack. The dominance hierarchy influences access to food resources within the pack. In captivity, pups show early development of aggressive behavior and engage in dominance fights when 19-24 days old. The process of establishing hierarchy within litters appears to last up to 4-5 months. Territoriality regulates Coyote numbers as packs space themselves across the landscape in relation to available food and habitat. Home range size varies with energetic requirements, physiographic makeup, habitat, and food distribution. Home range size is influenced by social organization, with transients using larger areas, and residents occupying distinct territories. Only packs (2-10 animals) maintain and defend territories, both by direct confrontation and indirectly with scent marking and howling. Fidelity to the home range is high and may persist for many years. Shifts in territorial boundaries may occur in response to the loss of one or both of the alpha pair. Pups, yearlings, and non-breeding adults of lower social rank may disperse from the natal site into adjacent areas or farther afield. Dispersal seems to be voluntary, as social and nutritional pressures intensify during winter when food becomes limited. Juveniles usually disperse during autumn and early winter. Pre-dispersal forays may occur. Coyotes communicate using auditory, visual, olfactory, and tactile cues. Studies have identified different types of vocalizations, seasonal patterns, and the influence of social status on vocalization rates. Howling plays a role in territorial maintenance and pack spacing by advertising boundaries and signaling the presence of alpha animals that will confront intruders and defend the territory. Scent marking contributes to territory maintenance and is performed mostly by alpha individuals. Scent marking may also be an indicator of sexual condition, maturity, or synchrony.

Breeding. Both males and females show annual cyclic changes in reproductive anatomy and physiology. Females are seasonally monoestrous, showing one period of estrus per year between January and March, depending on latitude. Courtship behavior begins 2-3 months before copulation. Copulation ends with a copulatory tie lasting up to 25 minutes. The percentage of females breeding each year varies with local conditions and food supply. Usually, about 60-90% of adult females and up to 70% of female yearlings producelitters. Gestation lasts approximately 63 days. Litter size averages six (range 1-9) and may be affected by population density and food availability during the previous winter. In northern latitudes, litter size changes in response to population cycles in Snowshoe Hares (Lepus americanus). Litter size has been found to increase after cold, snowy winters, when more ungulate carcasses are available to ovulating females. Coyotes may den in brush-covered slopes, steep banks, under rock ledges, thickets, and hollow logs. Dens of other animals may also be used, and may have more than one entrance and interconnecting tunnels. The same den may be used from year to year. Denning and pup rearing are the focal point of Coyote families for several months, until the pups are large and mobile. Pups are born blind and helpless in the den. Birth weight is 240-275 g. Their eyes open at about 14 days and they emerge from the den at about three weeks. The young are cared for by the parents and other associates, usually siblings from a previous year. Pups are weaned at about 5-7 weeks of age and reach adult weight by about nine months.

Status and Conservation. CITES not listed. Classified as Least Concern on The [UCN Red List. Coyotes are abundant throughout their range, which may be expanding due to their ability to successfully live in human-modified landscapes. Elimination of Wolves may also have facilitated Coyote expansion. Density varies geographically with food and climate, and seasonally due to mortality, changes in pack structure, predator density and food abundance. Coyotes are considered a pest species in many regions. Control programs temporarily reduce numbers on a short-term basis, but Coyote populations are generally stable in most areas and free of threats throughout their range. Conservation measures have not been needed to maintain viable populations. Coyotes adapt to human environments and occupy most developed habitats, including urban and agricultural areas. Hybridization with dogs and Gray Wolves may be occurring in Some areas.

Bibliography. Andelt (1985, 1987), Bekoff (1978b), Bekoff & Diamond (1976), Bekoff & Gese (2003), Bekoff & Wells (1986), Bekoff et al. (1981), Camenzind (1978), Gese & Bekoff (2004), Gese & Grothe (1995), Gese et al. (1996a, 1996b, 1996¢), Gier (1968), Kitchen et al. (2000a, 2000b), Knowlton et al. (1999), Laundre & Keller (1984), O'Donoghue et al. (1997), Richens & Hugie (1974), Thurber & Peterson (1991), Todd & Keith (1983), Wayne & Lehman (1992), Young & Jackson (1951).

Figures

2. Red Wolf (Canis rufus), 3. Coyote (Canis latrans), 4. Ethiopian Wolf (Canis simensis), 5. Golden Jackal (Canis aureus), 6. Side-striped Jackal (Canis adustus), 7. Black-backed Jackal (Canis mesomelas)

Subspecies and Distribution. C. l. latrans Say, 1823 — S Canada and USA (Great Plains region). C. l. cagottis Hamilton-Smith, 1839 — SE Mexico. C. l. clepticus Elliot, 1903 — Mexico (N Baja California) and USA (S California). C. l. dickeyi Nelson, 1932 — Costa Rica , El Salvador , W Honduras , Nicaragua , and Panama . C. l. frustror Woodhouse, 1850 — USA (Missouri, Kansas, parts of Oklahoma & E Texas). C. l. goldmani Merriam, 1904 — Belize , Guatemala , and S Mexico . C. l. hondurensis Goldman, 1936 — E Honduras . C. l. impavidus J. A. Allen, 1903 - W Mexico. C. l. incolatus Hall, 1934 - Alaska and NW Canada. C. l. ¡jamesi Townsend. 1912 - Mexico (Tiburon l. Baia California). c. [l.lestes lam Merriam, 1897 1897 - sw canada Canada and w USA USA Õnzenndunzain Region & NW). v C. l. mearnsi Merriam, 1897 - NW Mexico and SW USA. C. l. mıcrodon Merriam, 1897 — NE Mexico and S USA (S Texas). C. l. ochropus Eschscholtz, 1829 - W USA (W coast). C. l. peninsulae Merriam, 1897 — Mexico (S Baja California). C. l. texensıs Bailey, 1905 - S USA (W Texas 8c New Mexico). C. l. thamnos Jackson, 1949 - N-C Canada and E USA. C. l. umpquensıs Jackson. 1949 - USA (NW coast). C. l. vigilis Merriam. 1897 — SW Mexico. Coyotes did not originally occur on the USA E coast or Florida. They (probably thamnos) have expanded into the area with the clearing of forests and been introduced to Florida and Georgia (subspecies unknown).