Mating System: The coyote mating system is facultative monogamy. Monogamy means that only one male breeds with one female and this heterosexual pair remains together both in and out of the mating season, as is true for the coyote (Kleiman, 1977). The coyote mated pair remains together for multiple breeding seasons, generally until one of the individuals dies (Young, 1951a). If a mate is lost, it will be replaced by a new individual. Monogamy in mammals is very rare, applying to less than 3% of mammalian species. The coyote also meet other traits of monogamy. Species with long-term pair bonding often show little sexual dimorphism. In canids, the male is generally a little larger than the female, but the difference is not as significant as in most polygamous species. Furthermore, the coyote is monoestrus, which means that mating is restricted to a single short period each year. If the female does not conceive during this period, she will not have another chance until the next year. For the coyote and other canids, however, the female is receptive for several days during which mating may occur up to three times per day (Kleiman, 1977).

Image provided by Cody Forrester

There are two types of monogamy, obligate and facultative. Although obligate monogamy does not occurr in the coyote, it occurs when it would be impossible for a solitary female to rear a litter without the help of a male. Facultative monogamy means that the male's help is not absolutely necessary to raise the young. This form of monogamy may result when a species exists at low densities, so that males and females are so spread out that only a single member of the opposite sex is available (Kleiman, 1977). This may contribute to the coyote's practice of facultative monogamy because individuals tend to live in low densities. The male is not essential to the female after copulation has occurred, however, he does provide benefits such as food gathering and defense that decrease the female's required energy investment and helps increase the chances of pup survival. The male's opportunity for polygyny is limited in the coyote due to wide dispersal of females and because it would be virtually impossible for him to provide for more than one female during the denning season. As the males main roles are gathering food and defense, it would not be possible to defend multiple dens at a time.

 

Male/Female Pairing and Mating Behavior: If not already mated, coyotes are primarily involved in mate selection from December to February. Males seek out a mate and will travel well beyond their home range to find one. Scent-marking with urine appears to be very important in this process. Females increase the use of leg lift urination posture during this period, which may represent an increased scent-marking ability that may help them to find a mate. Soon after female urination increases, male urination rate increases as well, and they show additional interest in the female urine (Bekoff & Diamond, 1976).

Once couples are mated, den-selection behavior increases and generally takes place within the core areas of the pair's territory. The female does most of the den digging, which takes place from late December until early March. (Bekoff & Diamond, 1976). Interestingly, coyote return to the same denning sites annually, even when their pups have been killed by den hunters the year before, especially if the nest has not been damaged severely. Also, although females generally select the denning site, when a female of a pair is killed, the male may bring his new mate to the old denning site (Young, 1951b).

Early February marks a behavior shift in that the male demonstrates more interest in the female by sniffing and licking her vulva. It is not until the female finishes her den digging, however, that she is receptive to the male's actions of sniffing and licking the vulva. When the female is ready copulation occurs. A "copulatory tie", as is common for many canids, occurs at the end of copulation (Bekoff & Diamond, 1976). This means that during a copulation the male a female are locked together, due to their sexual anatomy, in a "tie" that may last up to 30 minutes (as cited in Kleiman, 1977). Copulatory behavior generally takes place sometime from late February to mid-March (Bekoff & Diamond, 1976).

Once successful copulation has occurred, the gestation period lasts approximately 60 days and the the average litter size is seven. Whelping season fluctuates with latitude in the United States (Bekoff & Diamond, 1976). A study by Hamlett (1938) indicated that onset in the northern states seems to be slightly earlier than in the southern states.

 

Parent/Offspring Relationship:

When the pups are born they are altricial for the first 2 months of life, which means they are fully dependent on parental care (Bekoff & Wells, 1980; Davis & Schmidly, 1997). They are born blind and helpless and in early life need to be fed, stimulated for elimination, kept warm and protected. In general, both adults help in raising the young.

While in the den, pups severely limit a mother's ability to acquire food and also increase her need for nutrition. As a result the male often helps to compensate by bringing food for the mother and the young (Laundré & Keller, 1981). Pups first emerge from the den around 3 weeks of age. Pups are weaned around 6-8 weeks of age, but remain at least semi-dependent until 4-5 months (Bekoff & Wells, 1986). As the pups are weaned, the mother and father regurgitate partially digested meat to feed them (as cited in Kleiman, 1977). If the mother dies before the pups are able to eat meat, they will die. If the pups are able to eat meat the father will generally care for them because he will be able to supply food. As previously discussed in the monogamy section, if the father dies, it would be possible for the mother to raise the young on her own (Young, 1951a).

Pup attendance remained relatively high even after weaning. This is probably to protect the young from predators, including other coyotes which may commit infanticide. The parent care roles become more equal after the pups are weaned. Parental travel from the den increases after weaning. This may be because a greater quantity of food is needed to feel the pups and the adults must travel further from the den to find enough food (Harrison & Gilbert, 1985). Once pups reach independence the parent/offspring bond begins to decrease.

A study by Althoff and Gibson examined the general proximity of related individuals throughout the day. After emerging from the den, related pups tended to stay near each other both during the day and at night, remaining within .5km of each other. Mated adults showed nearly five times the distance separation of the pups, averaging 1.85 km at night and 1.63 km during the day. Distances between pups and adults were slightly less than the distances between the mated pairs, but still substantial, with an average of 1.15 km during the day and 1.59 km at night. The pups tended to remain in the core of the family's territory, while the mothers spent most of their time near the border of the territory. The distance between adults and young for the majority of the data indicates that coyote mothers do not maintain close bonds with their pups after the denning period. In this study, beginning around November, some of the pups began dispersing (Althoff & Gipson,1981). Other studies have shown dispersal to begin earlier, however, with juveniles heading out as early as September and October (Harrison et al. 1991).

Photograph by David and Valerie Peters
(Pup in this photo is 3 months of age)

 

Continuation of a parental/offspring relationship depends on the dispersal of the juveniles. For individuals who disperse, this is generally the end of their relationship with their parent because the juvenile will eventually find its own territory or home range. Coyotes use directional travel when dispersing which helps them travel further from their natal territory, decreasing the risk of inbreeding and increasing the odds of reaching unclaimed territories, when all areas near the natal territory are occupied. Both sexes have equal dispersal rates. Because dispersal exposes juveniles to unfamiliar areas, it increases the risk for unexpected encounters with predators, hunters, vehicles, and traps (Harrison, 1992).

Not all juveniles disperse and those who remain with their parents cause a pack to form. Availability of large food items increases the likelihood of delayed dispersal and pack formation. When only small prey is available it is more advantageous to hunt alone, and therefore there is little need for a pack (Camenzind, 1978). Non-dispersing individuals may either remain directly with their parents and serve as alloparents for the next year's litter or they may have a less direct relationship. Juveniles that do not disperse and do not help with young rearing are known as "roamers" and typically hang on the periphery of the territory. These individuals have very little interaction with the other individuals in the pack (Camenzind, 1978). Unlike juveniles who associate directly with the pack, these individuals may actually hurt the pack by creating competition for available food resources but not providing any compensation to the mated pair, such as alloparenting (Messier & Barrette, 1982).

 

 

Cooperation in the coyote focuses on the relationship
between alloparent sub-adults and the mated pair.
It is a system of costs and benefits, with
neither group helping solely for the benefit of the other.