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Wolves communicate using vocalizations, body postures, scent, touch, and taste. The
lunar phase Concerning the lunar month of ~29.53 days as viewed from Earth, the lunar phase or Moon phase is the shape of the Moon's directly sunlit portion, which can be expressed quantitatively using areas or angles, or described qualitatively using the t ...
s have no effect on wolf vocalisation. Despite popular belief, wolves do not howl at the Moon. Gray wolves
howl Howl most often refers to: *Howling, an animal vocalization in many canine species *Howl (poem), a 1956 poem by Allen Ginsberg Howl may also refer to: Film * ''The Howl'', a 1970 Italian film * ''Howl'' (2010 film), a 2010 American arthouse b ...
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. Other vocalisations include growls,
barks Barks may refer to: * Carl Barks (1901 – 2000) was an American cartoonist, author, and painter * Coleman Barks (b. 1937), an American poet, and former literature faculty at the University of Georgia * Samantha Barks (b. 1990), a Manx actress an ...
and whines. Wolves do not bark as loudly or continuously as dogs do but they bark a few times and then retreat from a perceived danger. Aggressive or self-assertive wolves are characterized by their slow and deliberate movements, high body
posture Posture or posturing may refer to: Medicine * Human position ** Abnormal posturing, in neurotrauma ** Spinal posture ** List of human positions * Posturography Posturography is the technique used to quantify postural control in upright stance in ...
and raised
hackles Hackles are the erectile plumage or hair in the neck area of some birds and mammals. In birds, the hackle is the group of feathers found along the back and side of the neck. The hackles of some types of chicken, particularly roosters, are long, ...
, while submissive ones carry their bodies low, sleeken their fur, and lower their ears and tail. 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.


Visual

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. 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. Postural communication in wolves consists of a variety of facial expressions, tail positions and
piloerection Goose bumps, goosebumps or goose-pimples are the bumps on a person's skin at the base of body hairs which may involuntarily develop when a person is tickled, cold or experiencing strong emotions such as fear, euphoria or sexual arousal. The f ...
. Aggressive, or self-assertive wolves are characterized by their slow and deliberate movements, high body
posture Posture or posturing may refer to: Medicine * Human position ** Abnormal posturing, in neurotrauma ** Spinal posture ** List of human positions * Posturography Posturography is the technique used to quantify postural control in upright stance in ...
and raised
hackles Hackles are the erectile plumage or hair in the neck area of some birds and mammals. In birds, the hackle is the group of feathers found along the back and side of the neck. The hackles of some types of chicken, particularly roosters, are long, ...
, while submissive ones carry their bodies low, sleeken their fur and lower their ears and tail. 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. 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. 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. 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.


Auditory

Gray wolves
howl Howl most often refers to: *Howling, an animal vocalization in many canine species *Howl (poem), a 1956 poem by Allen Ginsberg Howl may also refer to: Film * ''The Howl'', a 1970 Italian film * ''Howl'' (2010 film), a 2010 American arthouse b ...
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. Wolf howls can under certain conditions be heard over areas of up to . Wolf howls are generally indistinguishable from those of large dogs. Male wolves give voice through an
octave In music, an octave ( la, octavus: eighth) or perfect octave (sometimes called the diapason) is the interval between one musical pitch and another with double its frequency. The octave relationship is a natural phenomenon that has been refer ...
, passing to a deep bass with a stress on "''O''", while females produce a modulated nasal
baritone A baritone is a type of classical male singing voice whose vocal range lies between the bass and the tenor voice-types. The term originates from the Greek (), meaning "heavy sounding". Composers typically write music for this voice in the r ...
with stress on "''U''". Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps. 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. 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 The great horned owl (''Bubo virginianus''), also known as the tiger owl (originally derived from early naturalists' description as the "winged tiger" or "tiger of the air"), or the hoot owl, is a large owl native to the Americas. It is an extrem ...
. 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. 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. Lone wolves typically avoid howling in areas where other packs are present. Wolves from different geographic locations may howl in different fashions: 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. The two are however
mutually intelligible In linguistics, mutual intelligibility is a relationship between languages or dialects in which speakers of different but related varieties can readily understand each other without prior familiarity or special effort. It is sometimes used as an ...
, as North American wolves have been recorded to respond to European-style howls made by biologists. Other vocalisations of wolves are usually divided into three categories: growls,
barks Barks may refer to: * Carl Barks (1901 – 2000) was an American cartoonist, author, and painter * Coleman Barks (b. 1937), an American poet, and former literature faculty at the University of Georgia * Samantha Barks (b. 1990), a Manx actress an ...
and whines. Barking has a fundamental frequency between 320–904 Hz, 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. Growling has a fundamental frequency of 380–450 Hz, 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. Whining is associated with situations of anxiety, curiosity, inquiry and intimacy such as greeting, feeding pups and playing.


Olfactory

Olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
is probably the wolf's most acute sense, and plays a fundamental role in communication. The wolf has a large number of
apocrine sweat gland An apocrine sweat gland (; from Greek ''apo'' 'away' and ''krinein'' 'to separate') is composed of a coiled secretory portion located at the junction of the dermis and subcutaneous fat, from which a straight portion inserts and secretes into the ...
s on the face, lips, back, and between the toes. The odor produced by these glands varies according to the individual wolf's
microflora Microbiota are the range of microorganisms that may be commensal, symbiotic, or pathogenic found in and on all multicellular organisms, including plants. Microbiota include bacteria, archaea, protists, fungi, and viruses, and have been found to ...
and diet, giving each a distinct "odor fingerprint". A combination of apocrine and
eccrine sweat gland Eccrine sweat glands (; from Greek ''ekkrinein'' 'secrete'; sometimes called merocrine glands) are the major sweat glands of the human body, found in virtually all skin, with the highest density in palm and soles, then on the head, but much less ...
s 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 Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
. The follicles present on the guard hairs from the wolf's back have clusters of apocrine and
sebaceous gland A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest number ...
s at their bases. As the skin on the back is usually folded, this provides a
microclimate A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squ ...
for bacterial propagation around the glands. During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent. The
precaudal scent gland A Rhodesian Ridgeback (sex unknown) with "stud tail": the violet gland lost hair and appears as a dark dimple The violet gland or supracaudal gland is a gland located on the upper surface of the tail of certain mammals, including European badge ...
s may play a role in expressing aggression, as combative wolves raise the base of their tails whilst drooping the tip, thus positioning the
scent gland 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 sexu ...
s at the highest point. The wolf possesses a pair of
anal sac Anal may refer to: Related to the anus *Related to the anus of animals: ** Anal fin, in fish anatomy ** Anal vein, in insect anatomy ** Anal scale, in reptile anatomy *Related to the human anus: ** Anal sex, a type of sexual activity involvin ...
s 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 gland Preputial glands are exocrine glands that are located in the folds of skin front of the genitals of some mammals. They occur in several species, including mice, ferrets, rhinoceroses, and even-toed ungulates and produce pheromones. The glands pla ...
s may advertise hormonal condition or social position, as dominant wolves have been observed to stand over subordinates, apparently presenting the genital area for investigation, which may include genital licking. 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 Animal communication is the transfer of information from one or a group of animals (sender or senders) to one or more other animals (receiver or receivers) that affects the current or future behavior of the receivers. Information may be sent int ...
in wolves. 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. 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. Their urine contains
pyrazine Pyrazine is a heterocyclic aromatic organic compound with the chemical formula C4H4N2. It is a symmetrical molecule with point group D2h. Pyrazine is less basic than pyridine, pyridazine and pyrimidine. It is a ''"deliquescent crystal or wax-lik ...
analogs that act as
kairomone A kairomone (a coinage using the Greek καιρός ''opportune moment'', paralleling pheromone"kairomone, n.". OED Online. September 2012. Oxford University Press. http://www.oed.com/view/Entry/241005?redirectedFrom=kairomone (accessed 3 Octobe ...
s, repelling their prey.Osada, Kazumi, et al.
Pyrazine analogues are active components of wolf urine that induce avoidance and freezing behaviours in mice
" PLoS One 8.4 (2013): e61753.


See also

*
Dog communication 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, li ...


Bibliography

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References

{{Lupus Animal communication Wolves