|This article may be expanded with text translated from the corresponding article in the Japanese Wikipedia. (February 2013)|
In ethology the term territory refers to any sociographical area that an animal of a particular species consistently defends against conspecifics (and, occasionally, animals of other species). Animals that defend territories in this way are referred to as territorial.
Classic territories 
Territorial animals defend areas that contain a nest, den or mating site, and sufficient food resources for themselves and their young. Defense rarely takes the form of overt fights: more usually there is a highly noticeable display, which may be visual (as in the red breast of the robin), auditory (as in much bird song, or the calls of gibbons) or olfactory, through the deposit of scent marks. Many territorial mammals use scent-marking to signal the boundaries of their territories; the marks may be deposited by urination, by defecation, or by rubbing parts of the bodies that bear specialised scent glands against the substrate. For example, dogs and other canids scent-mark by urination and defecation[further explanation needed], while cats scent-mark by rubbing their faces and flanks against objects, as well as by the notoriously persistently smelly spraying of urine by tomcats. Many prosimians use territorial marking; for example, the Red-bellied Lemur creates territories for groups of two to ten individuals in the rainforests of eastern Madagascar by scent marking: the male Diademed Sifaka also scent marks defended territories in some of these same rainforests. Humans build physical impediments and visual warnings to mark defended territories. The male Western fence lizard defends a territory by posturing and combat, but less intensely after the mating season.
Scent marking 
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Scent marking (also known as spraying or territorial marking) is behavior used by animals to identify their territory. Most commonly, this is accomplished by depositing strong-smelling substances, sometimes by urinating on prominent objects within the territory. Often the scent contains carrier proteins, such as the major urinary proteins, to stabilize the odours and maintain them for longer.
Not only does the marking communicate to others of the same species, but it is also noted by prey species and avoided. For example, felids such as leopards and jaguars mark by rubbing themselves against vegetation. Prosimians and New World monkeys also use scent marking, including urine washing (self-anointing the body with urine), to communicate. Many ungulates, for example the Blue Wildebeest, use scent marking from two glands, the preorbital gland and a scent gland in the hoof.
- Urine marking by canids [ edit ]
All canids (with the possible exception of dholes) use urine (combined with preputial gland secretions) to mark their territories. Many species of canids, including hoary foxes, cape foxes, and golden jackals, use a raised-leg posture when urinating. The scent of their urine is usually strongest in the winter, before the mating season.
Domestic dogs mark their territories by urinating on vertical surfaces (usually at nose level), sometimes marking over the urine of other dogs. When one dog marks over another dog's urine, this is known as "counter-marking" or "overmarking". Male dogs urine-mark more frequently than female dogs, typically beginning after the onset of sexual maturity. Male dogs, as well as wolves, sometimes lift a leg and attempt to urinate even when their bladders are empty - this is known as a "raised-leg display", "shadow-urination", or "pseudo-urination". They typically mark their territory due to the presence of new stimuli or social triggers in a dog's environment, as well as out of anxiety. Marking behavior is present in both male dogs and female dogs, and are especially pronounced in male dogs that have not been neutered.
Raised-leg urination is the most significant form of scent marking in wolves, and is most frequent around the breeding season. Wolves urine-mark more frequently when they detect the scent of other wolves, or other canid species. Leg-lifting is more common in male wolves than female wolves, although dominant females also use the raised-leg posture. Urine marking is the best-studied means of olfactory communication 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.
Other types of urine-marking in wolves are FLU (flexed-leg urination), STU (standing urination), and SQU (squatting urination). Breeding pairs of wolves will sometimes urinate on the same spot: this is known as "double-marking". Double-marking is practiced by coyotes as well as wolves., and also by foxes.
Coyotes mark their territories by urinating on bushes, trees, or rocks. All male coyotes lift their legs when urinating. However, females sometimes also raise their legs, and males sometimes squat. Urine marking is also associated with pair bonding in coyotes[clarification needed] Coyotes sometimes urinate on their food, possibly to claim ownership over it.
Red foxes use their urine to mark their territories. A male fox raises one hind leg and his urine is sprayed forward in front of him, whereas a female fox squats down so that the urine is sprayed in the ground between the hind legs. Urine is also used to mark empty cache sites, as reminders not to waste time investigating them. Red foxes use [clarify] to urinate, depending on where they are leaving a scent mark.
As in most other canids, male bush dogs lift their hind legs when urinating. However, female bush dogs use a kind of handstand posture, which is less common in other canids. When male bush dogs urinate, they create a spray instead of a stream.
Both male and female maned wolves use their urine to communicate, e.g. to mark their hunting paths, or the places where they have buried hunted prey. The urine has a very distinctive smell, which some people liken to hops or cannabis. The responsible substance is very likely a pyrazine, which occurs in both plants. (At the Rotterdam Zoo, this smell once set the police on a hunt for cannabis smokers.)
White-tailed deer 
White-tailed deer possess many glands that allow them to produce scents, some of which are so potent they can be detected by the human nose. Four major glands are the pre-orbital, forehead, tarsal, and metatarsal glands. It was originally thought that secretions from the preorbital glands (in front of the eye) were rubbed on tree branches; recent[when?] research suggests this is not so. It has been found that scent from the forehead or sudoriferous glands (found on the head, between the antlers and eyes) is used to deposit scent on branches that overhang "scrapes" (areas scraped by the deer's front hooves prior to rub-urination). The tarsal glands are found on the upper inside of the hock (middle joint) on each hind leg. Scent is deposited from these glands when deer walk through and rub against vegetation. These scrapes are used by bucks as a sort of "sign-post" by which bucks know which other bucks are in the area, and to let does know that a buck is regularly passing through the area—for breeding purposes. The scent from the metatarsal glands, found on the outside of each hind leg, between the ankle and hooves, may be used as an alarm scent. The scent from the Interdigital glands, which are located in between the hooves of each foot, emit a yellow waxy substance with an offensive odor. Deer can be seen stomping their hooves if they sense danger through sight, sound, or smell, this action leaves an excessive amount of odor for the purpose of warning other deer of possible danger.
Throughout the year deer will rub-urinate, a process during which a deer squats while urinating so that urine will run down the insides of the deer's legs, over the tarsal glands, and onto the hair covering these glands. Bucks rub-urinate more frequently during the breeding season. Secretions from the tarsal gland mix with the urine and bacteria to produce a strong smelling odor. During the breeding season does release hormones and pheromones that tell bucks that a doe is in heat and able to breed. Bucks also rub trees and shrubs with their antlers and head during the breeding season, possibly transferring scent from the forehead glands to the tree, leaving a scent other deer can detect.
Sign-post marking (scrapes and rubs) are a very obvious way that white-tailed deer communicate. Although bucks do most of the marking, does visit these locations often. To make a rub, a buck will use its antlers to strip the bark off of small diameter trees, helping to mark his territory and polish his antlers. To mark areas they regularly pass through bucks will make scrapes. Often occurring in patterns known as scrape lines, scrapes are areas where a buck has used its front hooves to expose bare earth. They often rub-urinate into these scrapes, which are often found under twigs that have been marked with scent from the forehead glands.
Territories may be held by an individual, a mated pair, or a group. Territoriality is not a fixed property of a species: for example, robins defend territories as pairs during the breeding season and as individuals during the winter, while some nectarivores defend territories only during the mornings (when plants are richest in nectar). In species that do not form pair bonds, male and female territories are often independent, in the sense that males defend territories only against other males, and females only against other females; in this case, if the species is polygynous, one male territory will probably contain several female territories, while in some polyandrous species such as the Northern Jacana, this situation is reversed.
Quite often territories that only yield a single resource are defended. For example, European Blackbirds may defend feeding territories that are distant from their nest sites, and in some species that form leks, for example the Uganda kob (a grazing antelope), males defend the lek site (which is used only for mating).
Territoriality is only shown by a minority of species. More commonly, an individual or a group of animals will have an area that it habitually uses but does not necessarily defend; this is called its home range. The home ranges of different groups often overlap, and in the overlap areas the groups will tend to avoid each other rather than seeking to expel each other. Within the home range there may be a core area that no other individual group uses, but again this is as a result of avoidance rather than defense.
Behavioural ecologists have argued that food distribution determines whether a species will be territorial or not. This however, though true as far as it goes, is too narrow a point of view. As mentioned above, there are several kinds of territoriality; for example, the defence of lek areas by kob has nothing to do with food. Many other examples of territorial defence, including fish, birds or even invertebrates, are related to competition for mates or safe lairs, rather than food. Territoriality will emerge where there is a focused resource that provides enough for the individual or group, within a boundary that is small enough to be defended without the expenditure of too much effort.
Many birds, particularly seabirds, though they nest in dense communities, are nonetheless territorial in that they defend their nesting site to within the distance that they can reach while brooding. This is necessary to prevent attacks on their own chicks or nesting material from neighbours. Commonly the resulting superimposition of the short-range repulsion onto the long-range attraction characteristically leads to the well-known roughly hexagonal spacing of nests. Interestingly, one gets a similar hexagonal spacing resulting from the territorial behaviour of gardening limpets such as species of Scutellastra  They vigorously defend their gardens of particular species of algae, that extend for perhaps 1–2 cm around the periphery of their shells.
Territoriality is least likely with insectivorous birds, where the food supply is plentiful but unpredictably distributed. Swifts rarely defend an area larger than the nest. Conversely, other insectivorous birds that occupy more constrained territories, such as the ground-nesting Blacksmith Lapwing may be very territorial, especially in the breeding season, where they not only threaten or attack many kinds of intruders, but have stereotyped display behaviour to deter conspecifics sharing neighbouring nesting spots.
Conversely, large solitary (or paired) carnivores, such as bears and the bigger raptors require an extensive protected area to guarantee their food supply. This territoriality will only break down when there is a glut of food, for example when Grizzly Bears are attracted to migrating salmon.
Many species demonstrate the behavior of polyterritoriality, referring to the act of claiming or defending more than one territory. In the European pied flycatcher (Ficedula hypoleuca), researchers assert that males exhibit polyterritoriality in order to deceive females of the species into entering into polygynous relationships. This hypothesis, named the deception hypothesis, claims that males have territories at far enough distances that females will not be able to discern already-mated males. The observation that males traveled long distances, ranging from 200m to 3.5km, to find a second mate seemed to be supportive of the argument. The debate about the polyterritoriality in this species may be able to spark research about the evolution of and reasons for polyterritoriality in other completely unrelated species.
See also 
In specific species 
- Dhole#Social and territorial behaviours
- * C. Michael Hogan (2008) "Western fence lizard (Sceloporus occidentalis)", Globaltwitcher, ed. Nicklas Stromberg 
- Stimson, J. (1969). "Territoriality of the owl limpet Lottia gigantea". Ecology 51 (1): 113–118. doi:10.2307/1933604. JSTOR 1933604.
- Hurst JL, Robertson DHL, Tolladay U, Beynon RJ (May 1998). "Proteins in urine scent marks of male house mice extend the longevity of olfactory signals". Anim Behav 55 (5): 1289–97. doi:10.1006/anbe.1997.0650. PMID 9632512.
- "V133.CHP:Corel VENTURA" (PDF). Retrieved 2012-11-20.
- Richard Estes (1991). The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates. University of California Press. pp. 464–. ISBN 978-0-520-08085-0. Retrieved 25 December 2012.
- Mittermeier, R. A.; Rylands, A. B.; Konstant, W. R. (1999). "Primates of the world: An introduction". In Nowak, R. M. Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 1–52. ISBN 978-0-8018-6251-9.
- Fox, M. W. (1984). The Whistling Hunters: Field Studies of the Indian Wild Dog (Cuon Alpinus). Steven Simpson Books. ISBN 0-9524390-6-9)
- Sillero-Zubiri, Claudio (2004-07-28). Canids: Foxes, Wolves, Jackals and Dogs: Status Survey and Conservation Action Plan. ISBN 9782831707860.
- Apps, Peter (1920-01-01). Wild Ways: Field Guide to the Behaviour of Southern African Mammals. ISBN 9781868724437.
- Grzimek, Bernhard (1972). Grzimek's Animal life encyclopedia.
- The Intelligence of Dogs: A Guide to the Thoughts, Emotions, and Inner Lives ... - Stanley Coren - Google Books. Books.google.com. Retrieved 2012-11-20.
- Lawrence Mark Elbroch, Michael Raymond Kresky, Jonah Wy Evans. Field Guide to Animal Tracks and Scat of California. Books.google.com. Retrieved 2013-02-10.
- Stanley Coren. The Intelligence of Dogs: A Guide to the Thoughts, Emotions, and Inner Lives .... Books.google.com. Retrieved 2013-02-10.
- Linda P. Case (2010). Canine and Feline Behavior and Training: A Complete Guide to Understanding Our Two Best Friends. Cengage Learning. pp. 46–. ISBN 978-1-4283-1053-7. Retrieved 22 March 2013.
- Tristram D. Wyatt (27 February 2003). Pheromones and Animal Behaviour: Communication by Smell and Taste. Cambridge University Press. pp. 99–. ISBN 978-0-521-48526-5. Retrieved 22 March 2013.
- James Serpell (21 September 1995). The Domestic Dog: Its Evolution, Behaviour and Interactions with People. Cambridge University Press. pp. 120–. ISBN 978-0-521-42537-7. Retrieved 21 March 2013.
- D. Caroline Coile; Margaret H. Bonham (2 September 2008). Why Do Dogs Like Balls?: More Than 200 Canine Quirks, Curiosities, and Conundrums Revealed. Sterling Publishing Company, Inc. ISBN 978-1-4027-5039-7. Retrieved 28 January 2013.
- "Mammalian Communication: A Behavioral Analysis of Meaning - Roger Peters - Google Books". Books.google.com. Retrieved 2013-04-29.
- "Wolf ethogram - Patricia A. Goodmann, Erich Klinghammer, Monty Sloan - Google Books". Books.google.com. Retrieved 2013-04-29.
- "Inside of a Dog: What Dogs See, Smell, and Know - Alexandra Horowitz - Google Books". Books.google.com. Retrieved 2013-04-29.
- Marc Bekoff (2006). Animal Passions And Beastly Virtues: Reflections On Redecorating Nature. Temple University Press. pp. 112–. ISBN 978-1-59213-349-9. Retrieved 21 March 2013.
- Michael W. Fox (1 July 2007). Dog Body, Dog Mind: Exploring Canine Consciousness and Total Well-Being. Globe Pequot. pp. 26–. ISBN 978-1-59921-661-4. Retrieved 23 February 2013.
- L. David Mech; Luigi Boitani (1 October 2010). Wolves: Behavior, Ecology, and Conservation. University of Chicago Press. pp. 85–. ISBN 978-0-226-51698-1. Retrieved 23 February 2013.
- "Urine Marking in Dogs: Why do Dogs Urinate to Mark Territory?". Pets.webmd.com. Retrieved 2012-11-20.
- The Wolves of Minnesota: Howl in the Heartland - L. David Mech - Google Books. Books.google.com. Retrieved 2012-11-20.
- Wolves: Behavior, Ecology, and Conservation - Google Books. Books.google.com. 2003-11-23. Retrieved 2012-11-20.
- Field Guide to Mammal Tracking in North America - James C. Halfpenny, Elizabeth Biesiot - Google Books. Books.google.com. Retrieved 2012-11-20.
- Mech & Boitani 2003, pp. 80–86
- Partha P. Majumder (1993). Human population genetics: a centennial tribute fo J.B.S. Haldane. Plenum Press. ISBN 978-0-306-44572-9. Retrieved 23 February 2013.
- Cambridge Scientific Abstracts, Inc (1986). Chemoreception Abstracts. Washington. Retrieved 22 March 2013.
- Meredith F. Small (1986). Zoo biology. ISBN 978-0-8451-3403-0. Retrieved 22 March 2013.
- Jane Williams (RCVS.) (16 June 2009). The complete textbook of animal health and welfare. Saunders/Elsevier. ISBN 978-0-7020-2944-8. Retrieved 22 March 2013.
- David W. Macdonald; Claudio Sillero-Zubiri (24 June 2004). The Biology and Conservation of Wild Canids. Oxford University Press. ISBN 978-0-19-152335-9. Retrieved 22 March 2013.
- Steven R. Lindsay (9 January 2008). Handbook of Applied Dog Behavior and Training, Procedures and Protocols. John Wiley & Sons. pp. 512–. ISBN 978-0-470-34413-2. Retrieved 22 March 2013.
- Durward Leon Allen (1979). Wolves of Minong: Isle Royal's Wild Community. University of Michigan Press. pp. 282–. ISBN 978-0-472-08237-7. Retrieved 21 March 2013.
- Animal Behaviour Abstracts. Cambridge Scientific Abstracts. 1998. Retrieved 22 March 2013.
- Ecology Abstracts. Cambridge Scientific Abstracts. 1998. Retrieved 22 March 2013.
- David Moskowitz (19 May 2010). Wildlife of the Pacific Northwest: Tracking and Identifying Mammals, Birds, Reptiles, Amphibians, and Invertebrates. Timber Press. pp. 221–. ISBN 978-0-88192-949-2. Retrieved 21 March 2013.
- Dietland Müller-Schwarze; Robert Milton Silverstein (1980). Chemical signals: vertebrates and aquatic invertebrates. Plenum Press. ISBN 978-0-306-40339-2. Retrieved 22 March 2013.
- Joanne Mattern (1 August 1999). The Coyote. Capstone. pp. 21–. ISBN 978-0-7368-8485-3. Retrieved 25 February 2013.
- Field Guide to Mammal Tracking in North America - James C. Halfpenny - Google Books. Books.google.com. Retrieved 2012-11-30.
- Hope Ryden (2005-05-30). God's Dog: A Celebration of the North American Coyote. Books.google.com. Retrieved 2013-02-10.
- Elbroch, Lawrence Mark; Kresky, Michael Raymond; Evans, Jonah Wy (2012-02-11). Field Guide to Animal Tracks and Scat of California. ISBN 9780520253780.
- Young & Jackson 1978, p. 98
- "Journal of Ethology, November 2012, John K. Fawcett, Jeanne M. Fawcett, Carl D. Soulsbury". Link.springer.com. 2013-01-01. Retrieved 2013-02-10.
- MacDonald, D. W. (2010). "Some Observations and Field Experiments on the Urine Marking Behaviour of the Red Fox, Vulpes vulpes L". Zeitschrift für Tierpsychologie 51: 1–0. doi:10.1111/j.1439-0310.1979.tb00667.x.
- Jorgenson, J. W.; Novotny, M.; Carmack, M.; Copland, G. B.; Wilson, S. R.; Katona, S.; Whitten, W. K. (1978). "Chemical Scent Constituents in the Urine of the Red Fox (Vulpes vulpes L.) During the Winter Season". Science 199 (4330): 796–798. doi:10.1126/science.199.4330.796. PMID 17836296.
- Whitten, W. K.; Wilson, M. C.; Wilson, S. R.; Jorgenson, J. W.; Novotny, M.; Carmack, M. (1980). "Induction of marking behavior in wild red foxes (Vulpes vulpes L.) by synthetic urinary constituents". Journal of Chemical Ecology 6: 49. doi:10.1007/BF00987526.
- Harrington, Fred H. (1981). "Urine-Marking and Caching Behavior in the Wolf". Behaviour 76 (3/4): 280–288. doi:10.1163/156853981X00112. JSTOR 4534102.
- Walters, Martin; Bang, Preben; Dahlstrøm, Preben (2001). Animal tracks and signs. Oxford: Oxford University Press. pp. 202–3. ISBN 0-19-850796-8.
- Lawrence Mark Elbroch, Michael Raymond Kresky, Jonah Wy Evans. Field Guide to Animal Tracks and Scat of California. Books.google.com. Retrieved 2013-02-10.
- Macdonald 1987, p. 125
- Henry, J. David (1977). "The Use of Urine Marking in the Scavenging Behavior of the Red Fox (Vulpes vulpes)". Behaviour 61 (1/2): 82–106. doi:10.1163/156853977X00496. JSTOR 4533812.
- Andersen, K. F.; Vulpius, T. (1999). "Urinary Volatile Constituents of the Lion, Panthera leo". Chemical Senses 24 (2): 179–189. doi:10.1093/chemse/24.2.179. PMID 10321819.
- Lawrence Mark Elbroch; Michael Raymond Kresky; Jonah Wy Evans (11 February 2012). Field Guide to Animal Tracks and Scat of California. University of California Press. pp. 189–. ISBN 978-0-520-25378-0. Retrieved 3 December 2012.
- Wolves and Other Dogs - Andrew Solway - Google Books. Books.google.com. Retrieved 2012-11-20.
- Canids: Foxes, Wolves, Jackals And Dogs: Status Survey And Conservation ... - Claudio Sillero-Zubiri - Google Books. Books.google.com. 2004-07-28. Retrieved 2012-11-30.
- Claudio Sillero-Zubiri (28 July 2004). Canids: Foxes, Wolves, Jackals And Dogs: Status Survey And Conservation Action Plan. IUCN. pp. 79–. ISBN 978-2-8317-0786-0. Retrieved 10 February 2013.
- Cristian Frers. "Un lobo de crin llamado Aguará Guazú". Retrieved 2007-04-23.
- Brian Switek (2011-03-10). "Maned Wolf Pee Demystified". Wired. Retrieved 2011-06-05.
- Süddeutsche Zeitung, 2006-09-02, p3
- Karen J. Alexy, Jonathan W. Gassett, David A. Osborn and Karl V. Miller, (2001). "White-Tailed Deer Rubs and Scrapes: Spatial, Temporal and Physical Characteristics and Social Role", Wildlife Society Bulletin, 29(3): 873-878.
- Terry L. Kile and R. Larry Marchinton, (1977). "White-Tailed Deer Rubs and Scrapes: Spatial, Temporal and Physical Characteristics and Social Role", American Midland Naturalist, 97(2): 257-266.
- Branch,G.M., Griffiths,C., Beckley,L.E., Branch,M.L.; Two Oceans. Pub. Struik, 2010. ISBN 978-1-77007-772-0
- Alatalo, Rauno V.; Carlson, Allan; Lundberg, Arne; Ulfstrand, Staffan (1981). "The Conflict Between Male Polygamy and Female Monogamy: The Case of the Pied Flycatcher Ficedula hypoleuca". The American Naturalist 117 (5): 738–753. doi:10.1086/283756.
Further reading 
- Walther, F. R., E. C. Mungall, G. A. Grau. (1983) Gazelles and their relatives : a study in territorial behavior Park Ridge, N.J. : Noyes Publications 239, ISBN 0-8155-0928-6
- Stokes, A. W. (editor) (1974) Territory Stroudsburg, Pa., Dowden, Hutchinson & Ross 398, ISBN 0-87933-113-5
- Klopfer, P. H. (1969) Habitats and territories; a study of the use of space by animals New York, Basic Books 117 p.