Coyotes
not within the same pack tend to avoid one another which helps to limit territorial
defense.
Even
when
boundaries
overlap
or were
continuous, the coyotes avoided being in the same areas. Scent marking, especially
with urine is thought to aide in this avoidance and with boundary marking.
Howling is also thought to be used to convey location. Although it is preferred
to avoid confrontation, as fighting creates high risks for the individual,
encounters do occur and must be taken seriously in order to indicate to
intruders that their presence will not be tolerated.
Photo from http://www.coyotehunter.freeservers.com/photo.htmlVocalizations: Coyotes have a wide ranges of calls used for auditory communication. These calls can be used over a variety of distances and appear to have varied significance. Vocalizations such as the woof, growl, bark, and yelp all relate to agonistic encounters. These calls indicate varying levels of threat intensity and meaning. Other calls such as the lone howl and the group howl are related to contact and are generally used during separation from the group and in response to distant howls (Lehner, 1978).
Coyote vocalizations generally take place between sunset and sunrise, but are quiet during the day (Canadian Wildlife, 1990). Howling behavior is most commonly associated with mated pairs and packs. Also, within a resident pack, alpha individuals rate of howling was three times greater than that of the beta and the pups, which were relatively equal. Pack size did not affect the rate of howling. The location of howling was most frequent at the periphery of the territory than in the center (Gese & Ruff, 1998). Non-resident members were not observed to initiate or participate in howling, indicating that this behavior is directly related to territoriality.
Vocalizations occur most often from November until April and again in June and July (Camanzind, 1978, Gese & Ruff, 1998). This period surrounds the breeding, gestation, and dispersal seasons and emphasizes the high need for territoriality during this period (Gese & Ruff, 1998).
Scent Marking: To some extent, territories are maintained by scent marking. Scent marking is most often through secreting on strong-smelling liquids from the anal glands or through urination (Lehner, 1978). Scent marking for territories provides a warning to the intruder that there is a risk in entering the area. The likelihood of trespassing is then at least semi-dependent on the non-residents perception of risk of injury if a physical encounter occurs (Messier & Barrette, 1982). In a study by Gese and Ruff (1997), alpha coyote were the primary individuals to perform scent-marking at territory boundaries. This demonstrates the alpha coyotes dominance and emphasizes the concept of marking as a warning because the alpha coyote is the main aggressor in territorial defense. Also, scent marking may indicate dominance, as the scent-marking rates were the highest during the breeding season, when there is an increased need to keep out intruders in order to protect pups from infanticide.Other uses for scent marking may include laying trails, signaling alarm, identifying individuals, recognizing group or species members, and attracting sexual partners (Allen et al., 1999). In a study on scent-marking and territoriality, Allen, Bekoff, and Crabtree (1999) observed that scent marking occurred most frequently on the periphery of territorial core areas . Scratching often occurs at the ground after elimination, and this may provide a visual marker in addition to the scent mark (Camanzind, 1978; Allen et al., 1999). In territorial markings, there was no scent-marking overlap for individuals from separate groups, indicating that scent marking is strongly associated with identifying and maintaining individuals' territories. Also, territorial coyote scent marked at a much higher rate than did transient coyotes and home range individuals who did not defend a home-range (Gese& Ruff, 1997).
Encounters and Aggressive Behaviors: Coyote encounters are fairly unlikely in comparison to other defense mechanisms such as vocalizations and scent-marking. Non-associated coyotes can generally avoid one another, even when intruding on another's territory because coyote generally live in fairly low density populations and on large areas. Also, because encounters are relatively infrequent, when an encounter occurs, the resident must display high levels of aggressiveness to assert that the boundaries are to be enforced because there will be a great risk to the individual if an encounter occurs (Messier & Barrette, 1982).
Agonistic encounters involve displays that show threat. Coyotes provide visual cues for others using their mouth, eyes, ears, shoulder hackles and tail. The coyote is primarily visually oriented. Facial fur coloring helps others see the expression being made by the coyote. The various faces have express meaning such as defensive aggression, low intensity aggression, neutral, alert, or submissive. The tail also has color markings that enhance tail movement and position.
In an encounter, the individuals size one another up and judge intent. Signs of threat include arching the back, raising the tail and wagging it stiffly with strong force (Camanzind, 1978). In agonistic encounters, the height of a tail seems to indicate a coyotes intent. For example, a raised tail indicates alertness or aggression , while a tucked tail indicates submission (Lehner, 1978). Coyote also display their canine teeth and retract their upper lip. Submissive behaviors include tucking the tail between the legs, dropping the head, and ears are commonly flattened (Camanzind, 1978).
Most agonistic encounters result in a chase in which the intruder was driven from the territory. Defense behaviors generally remain in the territory and the territory (Gese, 2001). In many cases the intruder is simply chased from there territory by a resident member (Camanzind, 1978).
Note: The below photographs are a series of three that demonstrate a pack running off a foreign member while they are protecting their kill-(kill not shown in picture).
Photographs by David and Valerie Peters
In this photo the coyote in the right hand corner is approaching a group. The individuals are probably sizing each other up. The two coyote in the center are displaying clear visual threat signals to the intruder.
Notice the submissive posture of the intruder. He is crouched low to the ground with his tail tucked, head low, and ears pressed against his head. The group follows him as they escort him away from their territory. The dominating individuals stand tall with backs slightly arched and their fur bristled.
This photograph demonstrates the chase, as the intruder makes a quick attempt to get away.
Although most encounters only result in a chase, physical contact sometimes occurs. Fight are aggressive. They usually do not end in death,however, but sometimes draw blood. In a study of territorial defense by Gese (2001), the alpha pair, as the dominant aggressors, participated in 87% of these actions. Pack members dominated over intruders in 75% of the interactions (Camanzind, 1978).
Overall, direct territorial behavior provide short term defense of the territory and protect against a specific intruder. Indirect territorial behavior, on the other hand, provides a longer lasting display of ownership. As mentioned, scent marking is a major component of indirect territorial behavior (Camanzind, 1978).
Social Spacing of Solitary Individuals
Solitary individuals utilize overlapping home ranges (Messier & Barrette, 1982; Laurence & Michael, 1985). These home ranges often partially overlap resident territories (Bekoff & Wells, 1980). The average home range size of solitary individuals was 30.1 square kilometers (Althoff, 1978). Single individuals need larger home ranges because they must rely on smaller prey year round, and this requires a greater area to obtain enough food (Bowen, 1982). Yearlings and juveniles also had very large home ranges, however, these were probably a result of dispersal activity (Althoff, 1978). Dispersing individuals may also pass through home ranges and territories as they travel, however these individuals do not use a specific home range and are nomadic in their land use until they find a new home range or territory (Bowen, 1982).
Mobility of non-resident males increases in late winter and this could be due to the approaching mating season as individuals set out to find a mate. Dispersing, although generally necessary in order to find a territory or mate, increases the coyote's risk for predation (Laurence & Michael, 1985). Unlike individuals in the mated pairs and packs who remain in one area and are familiar with the space, dispersing individuals do not have this familiarity.
Why this social spacing?
This social spacing is due to food distribution. Lone coyote cannot kill large mammals, so they are dependent of smaller prey such as rodents. Prey such as rodents are scattered and cannot be defended and a lone coyote cannot defend the larger food resources that are defendable by groups (Bekoff & Wells, 1980). Furthermore, without a mate, the coyote has no need for den sites as do groups containing mated pairs. Therefore, there would be no benefit to the lone coyote utilizing a territory.
