Territory (animal)

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For other uses, see Territory (disambiguation).
A male South China tiger scent-marking his territory

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.

Contents

Classic territories [edit]

Northern Elephant Seals (Mirounga angustirostris) fighting for territory and mates.

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.[1]

Invertebrates which show territoriality include some ants and bees, and the owl limpet.[2]

Scent marking [edit]

Wikimedia Commons has media related to: Scent marking
See also: Spraying (animal behavior), Pheromones, and Spoor (animal)

Scent marking (also known as spraying or territorial marking) is behavior used by animals to identify their territory.[3] 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.[4][5]

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.[citation needed] Prosimians and New World monkeys also use scent marking, including urine washing (self-anointing the body with urine), to communicate.[6][7] Many ungulates, for example the Blue Wildebeest, use scent marking from two glands, the preorbital gland and a scent gland in the hoof.[citation needed]

A female goat urinating

While the primary purpose of urination is the same across the animal kingdom, urination often serves a social purpose beyond the expulsion of waste material. In dogs and other animals, urination can mark territory or express submissiveness. In small rodents such as rats and mice, it marks familiar paths.

The urine and urination of animals of differing physiology or sex sometimes have different characteristics. For example, the urine of birds and reptiles is whitish, consisting of a pastelike suspension of uric acid crystals, and discharged with the feces of the animal via the cloaca, whereas mammals' urine is a yellowish colour, with mostly urea instead of uric acid, and is discharged via the urethra, separately from the feces. Some animals' (example: carnivores') urine possesses a strong odour, especially when it is used to mark territory or communicate in other ways.[clarify]

Stallions sometimes exhibit Flehmen response by smelling the urine of a mare in heat.[8] A stallion sometimes scent marks his urination spots in order to make clear his position as herd stallion.[9] A male horse's penis is protected by a sheath when it is not in use for urination.[10]

Ring-tailed lemurs have also been shown to mark using urine. Behaviorally, there is a difference between regular urination, where the tail is slightly raised and a stream of urine is produced, and the urine marking behavior, where the tail is held up in display and only a few drops of urine are used.[11][12] The urine-marking behavior is typically used by females to mark territory, and has been observed primarily at the edges of the troop's territory and in areas where other troops may frequent.[13] The urine marking behavior also is most frequent during the mating season, and may play a role in reproductive communication between groups.[11] Some other primate species use also urine for scent-marking. The white-headed capuchin sometimes engages in a practice known as "urine washing", in which the monkey rubs urine on its feet.[14] Urine washing, in which urine is rubbed on the hands and feet, is also used by the Panamanian night monkey.[15] In some cases, strepsirrhines may also anoint themselves with urine.[16]

Hyenas do not raise their legs as canids do when urinating, as urination serves no territorial function for them. Instead, hyenas mark their territories using their anal glands, a trait found also in viverrids and mustelids, but not canids and felids.[17] Unlike other female mammals, female spotted hyenas urinate, copulate, and give birth through an organ called the pseudo-penis.[18][19]

Canidae [edit]

A dog marking his territory

All canids (with the possible exception of dholes[20]) use urine (combined with preputial gland secretions) to mark their territories. Many species of canids, including hoary foxes,[21] cape foxes,[22] and golden jackals,[23] use a raised-leg posture when urinating.[24][25] The scent of their urine is usually strongest in the winter, before the mating season.[25]

Domestic dogs mark their territories by urinating on vertical surfaces (usually at nose level), sometimes marking over the urine of other dogs.[26] When one dog marks over another dog's urine, this is known as "counter-marking" or "overmarking".[27][28] Male dogs urine-mark more frequently than female dogs,[29] typically beginning after the onset of sexual maturity.[30] 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",[31][32][33][33][34] "shadow-urination",[35] or "pseudo-urination".[36] 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.[37] Marking behavior is present in both male dogs and female dogs, and are especially pronounced in male dogs that have not been neutered.[37]

Raised-leg urination is the most significant form of scent marking in wolves, and is most frequent around the breeding season.[38] Wolves urine-mark more frequently when they detect the scent of other wolves, or other canid species.[39] Leg-lifting is more common in male wolves than female wolves, although dominant females also use the raised-leg posture.[40] 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.[41]

A wolf marking its territory

Other types of urine-marking in wolves are FLU (flexed-leg urination), STU (standing urination), and SQU (squatting urination).[42] Breeding pairs of wolves will sometimes urinate on the same spot: this is known as "double-marking".[43][44][45][46][47][48] Double-marking is practiced by coyotes as well as wolves.,[49][50][51] and also by foxes.[52]

Coyotes mark their territories by urinating on bushes, trees, or rocks.[53] All male coyotes lift their legs when urinating.[54] However, females sometimes also raise their legs, and males sometimes squat.[55] Urine marking is also associated with pair bonding in coyotes[clarification needed][56] Coyotes sometimes urinate on their food, possibly to claim ownership over it.[57]

Red foxes use their urine to mark their territories.[58][59][60][61][62] 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.[63][64] Urine is also used to mark empty cache sites, as reminders not to waste time investigating them.[65][66][67] Red foxes use various postures[clarify] to urinate, depending on where they are leaving a scent mark.[63][68]

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.[69][70] When male bush dogs urinate, they create a spray instead of a stream.[71]

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.[72] 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.[73] (At the Rotterdam Zoo, this smell once set the police on a hunt for cannabis smokers.[73][74])

Felidae [edit]

Within the Felidae, male felids usually urinate backwards by curving the tip of the glans penis backward.[75][76] Urine marking by felids is also known as "spray-urinating"[77] or "spray-marking".[78] To identify their territories, male tigers mark trees by spraying urine[79][80] and anal gland secretions, as well as marking trails with scat. Males show a grimacing face, called the Flehmen response, when identifying a female's reproductive condition by sniffing their urine markings.

Male cheetah marking territory

Lions use urine to mark their territories. They often scrape the ground while urinating, and the urine often flows in short spurts, instead of flowing continuously. They often urinate on vegetation, or on tree trunks at least one meter high.[81] Male lions spray 1-20 jets of urine at an angle of 20-30 degrees upward, at a range of up to 4 meters behind them.[82]

Male cheetahs mark their territory by urinating on objects that stand out, such as trees, logs, or termite mounds. The whole coalition contributes to the scent. Males will attempt to kill any intruders, and fights result in serious injury or death.[83] When male cheetahs urine-mark their territories, they stand a meter away from a tree or rock surface with the tail raised, pointing the penis either horizontally backward or 60° upward.[84] The odor of cheetah urine (unlike that of other large felids) cannot be easily detected by humans.[85]

Black-footed cats use scent marking throughout their ranges, with males spraying urine up to 12 times an hour.[86]

Cervidae [edit]

Wikimedia Commons has media related to: Urinating deer
Video of a reindeer urinating

Bull elk often dig holes in the ground, in which they urinate and roll their body. The urine soaks into their hair and gives them a distinct smell which attracts cows.[87] One type of scent-marking behavior in elk is known as "thrash-urination."[clarify][88][89] Scent-marking by elk typically involves palpitation[clarify].[90][91][92] A male elk's urethra points upward so that urine is sprayed almost at a right angle to the penis.[91]

Unlike other deer species, chital do not spray urine on their bodies.[93] Instead, male chital mark their territory by dripping urine in scrapes, and then pawing them.[94]

Sambar stags will wallow and dig their antlers in urine soaked soil and then rub against tree trunks.[95][96] A stag will also mark himself by spraying urine directly in the face with a highly mobile penis, which is often erect during rutting activities.[95] Similar urine-spraying behavior is common in other deer species, and is known as automarking.[97]

Throughout the year white-tailed 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.[98] Bucks rub-urinate more frequently during the breeding season.[99]

References [edit]

White-tailed deer [edit]

See also: Urine spraying#Cervidae

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.[100] 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.[101]

Sign-post marking (scrapes and rubs) are a very obvious way that white-tailed deer communicate.[101] 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.


Defense [edit]

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).

Two territorial American Kestrels attacking a Red-tailed Hawk.

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 [102] 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.

Polyterritoriality [edit]

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.[103] 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 [edit]

In specific species [edit]

References [edit]

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Further reading [edit]

  • 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.
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