Coyote Canis latrans clepticus Class: Mammalia Order: Carnivora Family: Canidae Dudek and Associates Species Accounts Status: Federal: None State: None Data Characterization The MSHCP data base includes 400 records for the coyote. Of the 400 records, 323 (81%) are precision code ?1" (an ?x" and ?y" coordinate that allows for good precision in the location), 45 (11%) are precision code ?2" (one ?x" or ?y" coordinate or equivalent), and the remaining 32 (8%) are precision codes ?3" or ?4" (relatively imprecise locations from general areas) or have no precision codes indicated (5 records). Most of the records are relatively recent, with 258 (64%) since 1990, 125 (31%) from the 1980s, and nine (2%) from the 1970s. The remaining six records include a 1947 observation and five records with no date noted in the data base. Data records for the coyote are distributed throughout the study area, with clusters on the Santa Rosa Plateau, Lake Skinner, Sage, Lake Mathews, Anza Valley, Banning/Beaumont, and Moreno Valley. Based on the recency of most of the data and the distribution of records throughout the study area, the data base appears to provide a fairly good representation of the population distribution of this species in the study area. Habitat and Habitat Associations Coyotes utilize all habitats types and often are found in urban areas adjacent to open land. Primary habitats include grasslands, short-grass prairies, semiarid sagebrush, and broken forests (Gier 1975). Within their geographic range, coyotes are limited by the absence of open areas (Gier 1975). Natal dens are associated with brush-covered slopes, thickets, hollow logs, rocky ledges, and burrows. For example, in eastern Maine, dens varied from shallow depressions to multichambered burrows extending 1-2 meters in depth (Harrison and Gilbert 1985). The coyote has been recorded within virtually all upland and riparian habitat and land cover types in the MSHCP study area. The majority of the 400 records are from chaparral (107 records or 27% of the total). Seventy-eight (19%) of the records are from scrub habitats (coastal scrub, Diegan coastal sage scrub, Riversidean alluvial fan scrub, and Riversidean sage scrub), 75 (19%) from annual and native grassland, 58 (15%) from field crops and grove/orchard, 58 (15%) from residential/urban/exotic, 14 (3%) from oak woodlands (coast live oak woodland, dense Engelmann oak woodland, oak woodland), five (1%) from riparian (riparian scrub, southern cottonwood-willow riparian), one (<1%) from lower montane coniferous forest, one (<1%) from peninsular juniper woodland, and one (<1%) from freshwater marsh. Biogeography The coyote's geographic range has expanded dramatically in the last 150 years and includes the contiguous United States, western Canada and eastern Alaska, north to Hudson Bay and south to Guatemala (Hall 1981). Marginal records for the subspecies C. l. clepticus are San Marcos, Julian, and Jacumba in San Diego County, and into Baja California, Mexico (Hall 1981). The type locality for the subspecies is from the San Pedro Martir Mountains in Baja California (Bekoff 1977). The range of the subspecies appears to include western Riverside County, but the boundary with the range of the subspecies to the north, C. l. ochropus, is not clearly defined (Hall 1981). Range The coyote occurs throughout the planning area. Key Populations in Planning Area All planning area subregions. Biology Genetics: The diploid number of chromosome of the coyote is 78 (Wayne 1998). Phylogenetically, the coyote is closely related to the gray wolf (Canis lupus) and Simien jackal (Canis simensis) based on allozyme genetic distance and chromosome morphology (Wayne 1998). Studies of coyote mitochondrial DNA (mtDNA) genotypes indicate little geographic variation and the same genotypes may be present at widely-spaced localities (Wayne 1998). Lehman and Wayne (1991) found 32 mtDNA genotypes in coyotes from most parts of their North American range. The genotypes were not strongly geographically segregated, indicating high gene flow between subpopulations. Genetic relationships among populations of coyotes also are confusing because interbreeding between dogs and coyotes, coyotes and gray wolves, and coyotes and red wolves (Canis rufus) is not uncommon. Brownlow (1996) indicates that the molecular genetic studies have determined that the red wolf is in fact a hybrid of the gray wolf and coyote. Consequently the currently described subspecies of coyote may not hold up under molecular genetic analyses. Diet: Coyotes are omnivores, but basically carnivores, and their diet strongly reflect the prey and other food items available (Andelt et al. 1987; Gier 1975). Gier (1975), for example, lists the following food items: bison, deer, elk, sheep, rabbits, rodents, birds, non-toad amphibians, lizards, most snakes, crustaceans, and insects, blackberries, blueberries, peaches, apples, pears, prickly-pear cactus apples, chapotes, persimmons, peanuts, watermelon, cantaloupe, and grasses. They also consume inedible but chewable items such as harness straps, rubber or leather shoe soles, scraps of automobile tires, and paper wrappings. Coyotes prefer fresh meat, but will scavenge carrion. Coyotes also can be a major predator on domestic animals and pets such as cattle, lambs, chickens, turkeys, ducks, cats and dogs. For example, Crooks (unpublished manuscript) found cat remains in 21% of coyote scats in urban fragments in southwestern San Diego County. In central and southern California, lagomorphs (rabbits and hares) and rodents are primary prey items (Cypher et al. 1996; Pierce et al. 2000; Weintraub 1986). Although coyotes are opportunistic omnivores, there is some evidence of differential use of food items by age class that probably reflects foraging experience. Cypher et al. (1996) found that pups (<1 year of age) in the southern Central Valley of California consumed more insects than did yearlings (1 year of age) and adults (> 1 year of age). Also, adults and yearlings differed in their secondary food selection (jackrabbits were the primary prey of both age groups); adults' secondary prey was mostly rodents while yearlings' secondary prey was livestock and rodents. Coyotes also exhibit seasonal selection of prey, probably in relation to availability. Smith (1990) found that coyotes' diet shifted from small mammals such as kangaroo rats (Dipodomys spp.), voles (Microtus sp.), and squirrels (Spermophilus sp.) in the spring to fruit (manzanita berries) in the fall. Coyotes generally hunt by coursing in open areas, where they approach, test, and pursue prey, but they also may sit and wait for smaller prey. Also, in some areas with cover, they use the cover to allow them to approach prey more closely before they pounce (Murray et al. 1995). Coyotes also may occasionally hunt in packs for larger or difficult prey (Gier 1975). Rathbun et al. (1980) documented a family group of a male, female, and two-year old male offspring attacking and killing a badger (Taxidea taxus). When taking large prey such as mule deer (Odocoileus hemionus), coyotes hunt in packs. However, there is no evidence that they select for deer in poor condition as do some other carnivores (Pierce et. al. 2000). Coyotes exhibit hunting associations with badgers in some parts of their range. Minta et al. (1992) observed coyotes and badgers hunting in association for Uinta ground squirrels (Spermophilus armatus) in northwestern Wyoming. When hunting with badgers in brushy vegetation, coyotes were more successful than hunting alone because badgers would flush squirrels to the surface. The badger, which hunts underground, benefitted as well because squirrels tended to enter and stay in burrows in the presence of coyotes, thus increasing the capture success of the badger. Coyotes can be sustained on about 400-600 g of meat per day, or about 250 kg per year (Gier 1975). Daily Activity: Coyotes may be active anytime of the day, but primarily are nocturnal and crepuscular (Nowak 1990). A coyote typically travels about 4 km (2.5 miles) during a night of hunting (Nowak 1990). Daily movements, however, also depend on reproductive activities. During the nursing season, males and females spend more time near the den. Daily movements increase during weaning and are much larger when the pups are weaned, probably because of the increased energetic needs of the pups and the requirement of larger prey (Harrison and Gilbert 1985). Human disturbances may modify the temporal and spatial pattern of coyote activities. For example, Gese et al. (1989) studied the effects of military training activity on coyote movements in Colorado and found that individual coyotes responded differently depending on the amount of cover in their range and the level of military activities. Coyotes with high cover and little military activity in their range contracted their ranges, while coyotes with little cover and moderate levels of military activity in their range expanded their ranges. Also, coyotes generally increased their level of diurnal activity in relation to training Survival: Coyotes live about ten years (Gier 1975), but mortality in the first year is high. About 10-15% of the pups die within a few days of birth from several causes, including parasitic infections (hookworm, roundworm), accidents, predation (hawks, owls, eagle), neighboring coyotes, the loss of the parents, and general physical weaknesses. By late summer, about 50% of the pups may have perished. Socio-Spatial Behavior: Coyotes typically establish consistent home ranges and exhibit fairly extensive intraspecific home range overlap (Gier 1975). Home ranges may be quite variable, with a range of 10-100 sq km (Laundré and Keller 1984). Coyotes apparently are only strongly territorial during the denning season when their pups are at risk of being killed by other coyotes (Gier 1975). As described above, coyotes may come together to hunt in packs comprised of a family unit or a temporary non-family "pack" of two to six individuals comprised of bachelor males, non-reproductive females, and near-mature young. Their spatial structure is related to the adequacy, type, and distribution of the food supply (e.g., a pack is required to take larger prey such as deer), denning territory, and intraspecific and interspecific competition for resources. Human disturbances may cause coyotes to alter their spatial behavior, such as the use of their home range and dens. As described above, Gese et al. (1989) found that coyotes shifted their ranges and centers of activity in response to military training activities. Harrison and Gilbert (1985) observed that coyotes moved their dens after human disturbance. On the other hand, range management practices in south Texas did not appear to have substantial effects on coyotes' establishment and use of home ranges (Bradley and Fagre 1988). In this study on an experimental ranch, roads and fencelines did not limit home range establishment, and, in fact, animals tended to use roads and fencelines for movement more than expected by chance. There was a slight avoidance of pastures with cattle. Coyotes are a highly social species and use a variety of techniques to mark their ranges, probably to communicate their presence and location, and to attract the opposite sex (Gier 1975). They primarily communicate through calls and scent marking with urine and feces; urine markings attract the opposite sex more than the same sex. Reproduction: Coyotes are monestrous, meaning that females only come into estrus once per year (Gier 1975) and lost litters are not compensated for in the same breeding season. Receptivity typically occurs in mid- to late-winter (January to March) and may last up to a month. About 90% of adult females are sexually active and 0-60% of "yearling" (9-10 months) are active during this period. Receptive females attract several male consorts and may mate with several males, which the female appears to select. Ovulation occurs about three to four days before the end of receptivity. Gestation is about 58 to 65 days. Litter sizes range from two to 12 pups, but some dens may contain more than one litter. Males and females establish a pairbond, whereupon they select a territory, prepare a den, hunt and sleep together during the pregnancy, and both provide care for the pups (Gier 1975). The male is the primary hunter during the nursing and weaning period and food is brought back to the den and regurgitated for the pups. The territory around the den is defended from predators and other coyotes. Pups are weaned by 8-10 weeks, at which time the den is abandoned (Harrison and Gilbert 1985). At this time, the pups concentrate their activity around rendezvous sites, and over time adult visits to the rendezvous sites become progressively less frequent. The family unit may remain intact until about November, but pups may wander long distances in November and December. Adult weights are achieved by the ninth month. Dispersal: Coyotes are highly mobile and capable of moving long distances. Young disperse in the fall and winter and may move 80-160 km (50-100 miles) from the parental range (Gier 1975). In Iowa, individuals traveled an average of 31 km (19 miles), but up to 323 km (200 miles) (Nowak 1990). Threats Rangewide, the coyote is not threatened with extinction and this species will persist in western Riverside County under any reserve scenario. However, its distribution in the MSHCP study area likely will be affected by the pattern of urbanization and fragmentation and isolation of patches of habitat that may be inaccessible or too small to attract or sustain a coyote population. As described above and reiterated below, the loss of coyotes may result in the decline of species richness in small habitat patches through the loss of the native fauna to mesopredators. Exacerbating the loss of coyotes in urban habitat patches is the increased risk of vehicle collisions and any predator control activities conducted by local agencies. Special Biological Considerations Although the coyote is not at great risk of extirpation from the MSHCP planning area, it appears to be a key species in maintaining species richness in smaller habitat fragments. Conceptually, habitat fragmentation may alter the composition and structure of animal communities by modifying several ecological processes, including predation. Predators may respond to landscape features that are affected by habitat fragmentation, such as proximity to habitat edges, size of habitat patches, and habitat diversity. For example, edges may act either as physical barriers or movement conduits, causing predators to move along them and thus encounter prey at the edge at a higher rate. In addition, predator densities (and prey vulnerability) may be higher in small habitat patches, particularly for some generalist predators whose patches of habitat are surrounded by matrices that offer human-derived foods (see Oehler and Litvaitis 1996). As demonstrated by the work by Crooks in San Diego County, coyotes can help maintain native fauna by controlling mesopredators such as raccoons, skunks, gray foxes, and feral and domestic house cats (Crooks and Soulé 1999; Crooks unpublished manuscript). Consequently the reserve system should allow coyotes access to as much of the reserve as possible. Literature Cited Andelt, W.F., J.G. Kie, F.R. Knowlton, and K. Cardwell. 1987. Variation in coyote diets associated with season and successional changes in vegetation. Journal of Wildlife Management 5:273-277. 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Mytton. 1989. Changes in coyote movements due to military activity. Journal of Wildlife Management 53:335-339. Gier, H.T. 1975. Ecology and behavior of the coyote (Canis latrans). In The Wild Canids, M.W. Fox (ed.), Van Nostrand Reinhold, New York, pp. 247-262. Hall, E.R. 1981. The Mammals of North America. John Wiley and Sons, New York, 2 Vol. 1181 pp. Harrison, D.J. and J.R. Gilbert. 1985. Denning ecology and movement of coyote (Canis latrans) in Maine during pup rearing. Journal of Mammalogy 66:712-719. Laundré, J.W. and B.L. Keller. 1984. Home-range size of coyotes: a critical review. Journal of Wildlife Management 48:127-139. Lehman, N. and R.K. Wayne. 1991. Analysis of coyote mitochondrial DNA genotype frequencies: estimation of the effective number of alleles. Genetics 128:405-416. Minta, S.C., K.A. Minta, and D.F. Lott. 1992. Hunting associations between badgers (Taxidea taxus) and coyotes (Canis latrans). Journal of Mammalogy 73:814-820. Murray, D.L., S. Boutin, M. O'Donoghue, and V.O. Nams. 1995. Hunting behaviour of a sympatric felid and canid in relation to vegetative cover. Animal Behaviour 50:1203- 1210. Nowak, R.M. 1991. Mammals of the World, Fifth Edition. The Johns Hopkins University Press, Baltimore. pp.1068-1070. Oehler, J.D. and J.A. Litvaitis. 1996. The role of spatial scale in understanding responses of medium-sized carnivores to forest fragmentation. Canadian Journal of Zoology 74:2070-2079. Pierce, B.M., V.C. Bleich, and R.T. Bowyer. 2000. Selection of mule deer by mountain lions and coyotes: effects of hunting style, body size, and reproductive status. Journal of Mammalogy 81:462-472. Rathbun, A.P., M.C. Wells, and M. Bekoff. 1980. Cooperative predation by coyotes on badgers. Journal of Mammalogy 61:375-376. Sargeant, A.B. and S.H. Allen. 1989. Observed interactions between coyotes and red foxes. Journal of Mammalogy 70:631-633. Smith, J.R. 1990. Coyotes diets associated with seasonal mule deer activities in California. 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