Wolves communicate using vocalizations, body postures, scent, touch, and taste. [1] The lunar phases have no effect on wolf vocalisation. Despite popular belief, wolves do not howl at the Moon. [2] Gray wolves howl to assemble the pack, usually before and after hunts, to pass on an alarm particularly at a den site, to locate each other during a storm or while crossing unfamiliar territory, and to communicate across great distances. [3] Other vocalisations include growls, barks and whines. [4] Wolves do not bark as loudly or continuously as dogs do but they bark a few times and then retreat from a perceived danger. [4] Aggressive or self-assertive wolves are characterized by their slow and deliberate movements, high body posture and raised hackles, while submissive ones carry their bodies low, sleeken their fur, and lower their ears and tail. [5] Raised leg urination is considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of all scent marks observed. [6]
The gray wolf's expressive behavior is more complex than that of the coyote and golden jackal, as necessitated by its group living and hunting habits. While less gregarious canids generally possess simple repertoires of visual signals, wolves have more varied signals that subtly inter grade in intensity. [7] [8] When neutral, the legs are not stiffened, the tail hangs down loosely, the face is smooth, the lips untensed, and the ears point in no particular direction. [9] Postural communication in wolves consists of a variety of facial expressions, tail positions and piloerection. [10] Aggressive, or self-assertive wolves are characterized by their slow and deliberate movements, high body posture and raised hackles, while submissive ones carry their bodies low, sleeken their fur and lower their ears and tail. [5] When a breeding male encounters a subordinate family member, it may stare at it, standing erect and still with the tail horizontal to its spine. [11] Two forms of submissive behavior are recognized: passive and active. Passive submission usually occurs as a reaction to the approach of a dominant animal, and consists of the submissive wolf lying partly on its back and allowing the dominant wolf to sniff its anogenital area. Active submission occurs often as a form of greeting, and involves the submissive wolf approaching another in a low posture, and licking the other wolf's face. [12] When wolves are together, they commonly indulge in behaviors such as nose pushing, jaw wrestling, cheek rubbing and facial licking. The mouthing of each other's muzzles is a friendly gesture, while clamping on the muzzle with bared teeth is a dominance display. [13]
Similar to humans, gray wolves have facial color patterns in which the gaze direction can be easily identified, although this is often not the case in other canid species. In 2014, a study compared the facial color pattern across 25 canid species. The results suggested that the facial color pattern of canid species is related to their gaze communication, and that especially gray wolves use the gaze signal in conspecific communication. [14]
Feature | Aggressive | Fearful |
---|---|---|
Eyes | Direct stare Open wide | Looking away Closed to slits |
Ears | Erect and forward | Flattened and turned down to side |
Lips | Horizontal contraction ("agonistic pucker") | Horizontal retraction ("submissive grin") |
Mouth | Opened | Closed |
Teeth | Canines bared | Canines covered |
Tongue | Retracted | Extended ("lick intention") |
Nose | Shortened (skin folded) | Lengthened (skin smoothed) |
Forehead | Contracted (bulging over eyes) | Stretched (smoothed) |
Head | Held high | Lowered |
Neck | Arched | Extended |
Hair | Erect (bristled) | Sleeked |
Body | Erect, tall | Crouched, low |
Tail | Held high Quivering | Tucked under body Wagging |
Gray wolves howl to assemble the pack (usually before and after hunts), to pass on an alarm (particularly at a den site), to locate each other during a storm or unfamiliar territory and to communicate across great distances. [3] Wolf howls can under certain conditions be heard over areas of up to 130 km2 (50 sq mi). [15] Wolf howls are generally indistinguishable from those of large dogs. [16] Male wolves give voice through an octave, passing to a deep bass with a stress on "O", while females produce a modulated nasal baritone with stress on "U". Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps. [17] Howling consists of a fundamental frequency that may lie between 150 and 780 Hz, and consists of up to 12 harmonically related overtones. The pitch usually remains constant or varies smoothly, and may change direction as many as four or five times. [18] Howls used for calling pack mates to a kill are long, smooth sounds similar to the beginning of the cry of a great horned owl. When pursuing prey, they emit a higher pitched howl, vibrating on two notes. When closing in on their prey, they emit a combination of a short bark and a howl. [16] When howling together, wolves harmonize rather than chorus on the same note, thus creating the illusion of there being more wolves than there actually are. [3] Lone wolves typically avoid howling in areas where other packs are present. [19] Wolves from different geographic locations may howl in different fashions: according to Erik Zimen, the howls of European wolves are much more protracted and melodious than those of North American wolves, whose howls are louder and have a stronger emphasis on the first syllable. [20]
Other vocalisations of wolves are usually divided into three categories: growls, barks and whines. [4] Barking has a fundamental frequency between 320–904 Hz, [18] and is usually emitted by startled wolves. Wolves do not bark as loudly or continuously as dogs do, but bark a few times and retreat from perceived danger. [4] Growling has a fundamental frequency of 380–450 Hz, [18] and is usually emitted during food challenges. Pups commonly growl when playing. One variation of the howl is accompanied by a high pitched whine, which precedes a lunging attack. [3] Whining is associated with situations of anxiety, curiosity, inquiry and intimacy such as greeting, feeding pups and playing. [4]
Olfaction is probably the wolf's most acute sense, and plays a fundamental role in communication. The wolf has a large number of apocrine sweat glands on the face, lips, back, and between the toes. The odor produced by these glands varies according to the individual wolf's microflora and diet, giving each a distinct "odor fingerprint". A combination of apocrine and eccrine sweat glands on the feet allows the wolf to deposit its scent whilst scratching the ground, which usually occurs after urine marking and defecation during the breeding season. The follicles present on the guard hairs from the wolf's back have clusters of apocrine and sebaceous glands at their bases. As the skin on the back is usually folded, this provides a microclimate for bacterial propagation around the glands. During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent. [21]
The precaudal scent glands may play a role in expressing aggression, as combative wolves raise the base of their tails whilst drooping the tip, thus positioning the scent glands at the highest point. [16] The wolf possesses a pair of anal sacs beneath the rectum, which contain both apocrine and sebaceous glands. The components of anal sac secretions vary according to season and gender, thus indicating that the secretions provide information related to gender and reproductive state. The secretions of the preputial glands may advertise hormonal condition or social position, as dominant wolves have been observed to stand over subordinates, apparently presenting the genital area for investigation, [21] which may include genital licking. [22]
During the breeding season, female wolves secrete substances from the vagina, which communicate the females' reproductive state, and can be detected by males from long distances. Urine marking is the best-studied means of olfactory communication in wolves. [24] Its exact function is debated, though most researchers agree that its primary purpose is to establish boundaries. Wolves urine mark more frequently and vigorously in unfamiliar areas, or areas of intrusion, where the scent of other wolves or canids is present. So-called raised leg urination (RLU) is more common in male wolves than in females, and may serve the purpose of maximizing the possibility of detection by conspecifics, as well as reflect the height of the marking wolf. Only dominant wolves typically use RLU, with subordinate males continuing to use the juvenile standing posture throughout adulthood. [21] RLU is considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of all scent marks observed. [25] Like other canids, wolves urinate on their food caches in order to identify them. [26] Their urine contains pyrazine analogs that act as kairomones, repelling their prey. [27]
The coyote, also known as the American jackal, prairie wolf, or brush wolf is a species of 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; however, the coyote is generally larger.
Canidae is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid. The family includes three subfamilies: the Caninae, and the extinct Borophaginae and Hesperocyoninae. The Caninae are known as canines, and include domestic dogs, wolves, coyotes, foxes, jackals and other species.
The wolf, also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of Canis lupus have been recognized, including the dog and dingo, though gray wolves, as popularly understood, only comprise naturally-occurring wild subspecies. The wolf is the largest extant member of the family Canidae, and is further distinguished from other Canis species by its less pointed ears and muzzle, as well as a shorter torso and a longer tail. The wolf is nonetheless related closely enough to smaller Canis species, such as the coyote and the golden jackal, to produce fertile hybrids with them. The wolf's fur is usually mottled white, brown, gray, and black, although subspecies in the arctic region may be nearly all white.
Canis is a genus of the Caninae which includes multiple extant species, such as wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.
In ethology, territory is the sociographical area that an animal consistently defends against conspecific competition using agonistic behaviors or real physical aggression. Animals that actively defend territories in this way are referred to as being territorial or displaying territorialism.
The eastern wolf, also known as the timber wolf, Algonquin wolf and eastern timber wolf, is a canine of debated taxonomy native to the Great Lakes region and southeastern Canada. It is considered to be either a unique subspecies of gray wolf or red wolf or a separate species from both. Many studies have found the eastern wolf to be the product of ancient and recent genetic admixture between the gray wolf and the coyote, while other studies have found some or all populations of the eastern wolf, as well as coyotes, originally separated from a common ancestor with the wolf over 1 million years ago and that these populations of the eastern wolf may be the same species as or a closely related species to the red wolf of the Southeastern United States. Regardless of its status, it is regarded as unique and therefore worthy of conservation with Canada citing the population in eastern Canada as being the eastern wolf population subject to protection.
Dog communication is the transfer of information between dogs, as well as between dogs and humans. Behaviors associated with dog communication are categorized into visual and vocal. Visual communication includes mouth shape and head position, licking and sniffing, ear and tail positioning, eye gaze, facial expression, and body posture. Dog vocalizations, or auditory communication, can include barks, growls, howls, whines and whimpers, screams, pants and sighs. Dogs also communicate via gustatory communication, utilizing scent and pheromones.
The golden jackal, also called the common jackal, is a wolf-like canid that is native to Eurasia. The golden jackal's coat varies in color from a pale creamy yellow in summer to a dark tawny beige in winter. It is smaller and has shorter legs, a shorter tail, a more elongated torso, a less-prominent forehead, and a narrower and more pointed muzzle than the Arabian wolf. It is listed as Least Concern on the IUCN Red List due to its widespread distribution and high density in areas with plenty of available food and optimum shelter.
A bark is a sound most often produced by dogs. Other animals that make this noise include, but are not limited to, wolves, coyotes, foxes, seals, frogs, and barking owls. "Bark" is also a verb that describes the sound of many canids.
Scent gland are exocrine glands found in most mammals. They produce semi-viscous secretions which contain pheromones and other semiochemical compounds. These odor-messengers indicate information such as status, territorial marking, mood, and sexual behaviour. The odor may be subliminal—not consciously detectable. Though it is not their primary function, the salivary glands may also function as scent glands in some animals.
The violet gland or supracaudal gland is a gland located on the upper surface of the tail of certain mammals, including European badgers and canids such as foxes, wolves, and the domestic dog, as well as the domestic cat. Like many other mammalian secretion glands, the violet gland consists of modified sweat glands and sebaceous glands.
Cats communicate for a variety of reasons, including to show happiness, express anger, solicit attention, and observe potential prey. Additionally, they collaborate, play, and share resources. When cats communicate with humans, they do so to get what they need or want, such as food, water, attention, or play. As such, cat communication methods have been significantly altered by domestication. Studies have shown that domestic cats tend to meow much more than feral cats. They rarely meow to communicate with fellow cats or other animals. Cats can socialize with each other and are known to form "social ladders," where a dominant cat is leading a few lesser cats. This is common in multi-cat households.
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Dog behavior is the internally coordinated responses of individuals or groups of domestic dogs to internal and external stimuli. It has been shaped by millennia of contact with humans and their lifestyles. As a result of this physical and social evolution, dogs have acquired the ability to understand and communicate with humans. Behavioral scientists have uncovered a wide range of social-cognitive abilities in domestic dogs.
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Erich Klinghammer was a wolf biologist best known for his contributions to the fields of ethology and behavioural ecology, particularly that of canids. He was the founder of Wolf Park in Indiana and a professor of animal behaviour at Purdue University.
The Kenai Peninsula wolf, also known as the Kenai Peninsula grey wolf, is an extinct subspecies of the gray wolf that lived on the Kenai Peninsula in southern Alaska.
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