445:
92:
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977:) is a large (up to 8 cm in length) limpet. It lives in association with an approximately 1,000 cm^2 area of algal film in which its grazing marks can be seen, whereas the remainder of the rock surface is usually free of any visible film. These areas of algal film represent the territories of the Lottia; within them the animals do all their grazing. They keep their territories free of other organisms by shoving off any intruders: other Lottia, grazing limpets of the genus Acmaea, predatory snails, and sessile organisms such as anemones and barnacles.
371:
1048:), playbacks of neighbour and stranger songs at three periods of the breeding season show that neighbours are dear enemies in the middle of the season, when territories are stable, but not at the beginning of the breeding season, during settlement and pair formation, nor at the end, when bird density increases due to the presence of young birds becoming independent. Thus, this dear enemy territoriality relationship is not a fixed pattern but a flexible one likely to evolve with social and ecological circumstances.
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404:
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105:
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436:
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790:
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232:, a home range may be as small as 30 hectares (74 acres) in a good rural habitat, but as large as 300 hectares (740 acres) in a poor habitat. On average, a territory may be approximately 50 hectares (120 acres), with main setts normally at least 500 metres (1,600 ft) apart. In urban areas, territories can be as small as 5 hectares (12 acres), if they can obtain enough food from bird tables,
703:). Many animals have scent glands in their paws or deposit fur during tree-marking, so tree-marking may be a combination of both visual and olfactory advertising of the territory. The male ring-tailed lemur has a specialised adaptation to assist in leaving visual/olfactory territorial marks. On their inner forearm (antebrachial) is a scent gland which is covered by a
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territorial males patrolled an area around a tree and used the same territory for up to 49 days. It also appeared that they gave up territories to new males without violence. Males defend territories solely for mating, and no other resources such as fragrances, nests, nest construction materials, nectar, or pollen are found at these territories.
310:
707:. In a behaviour called "spur marking", they grasp the substrate, usually a small sapling, and drag the spur over it, cutting into the wood and spreading the gland's secretions. When on the ground, ring-tailed lemurs preferentially mark small saplings and when high in the trees, they usually mark small vertical branches.
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illusion of there being more wolves than there actually are. 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.
1058:
a non-social bee species, males have been observed to occasionally form aggregations of fragrance-rich territories, considered to be leks. These leks serve only a facultative purpose for this species, in which the more fragrance-rich sites there are, the greater the number of habitable territories.
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scratch their hind feet in the dung. By continuing to walk, they "transport" their own smell around the paths, thus establishing a scent-marked trail. Another method of visually marking their territory is wiping their horns on bushes or the ground and scraping with the feet, although this is likely
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bulls mark their territory with faeces and urine (olfactory marking). The dung is laid in well defined piles. There may be 20 to 30 of these piles to alert passing rhinoceroses that it is occupied territory. Other males may deposit dung over the piles of another and subsequently the sign-post grows
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Visual sign-posts may be a short-term or long-term mode of advertising a territory. Short-term communication includes the colouration or behaviour of the animal, which can only be communicated when the resident is present. Other animals may use more long-term visual signals such as faecal deposits,
339:
toward each other. This is a series of stylised postures, vocalisations, displays, etc. which function to solve the territory dispute without actual fighting as this could injure either or both animals. Ritualized aggression often ends by one of the animals fleeing (generally the intruder). If this
326:
Rather than retaining a territory simply by fighting, for some animals this can be a 3-stage process. Many animals create "sign-posts" to advertise their territory. Sometimes these sign-posts are on the boundary thereby demarcating the territory, or, may be scattered throughout the territory. These
656:
are noted for their highly aggressive territorial behaviour. They attack other males that stray into their territories, and have been observed attacking other small birds without apparent provocation. Such attacks sometimes lead to fatalities, accounting for up to 10% of adult robin deaths in some
870:. In this model of aggression, two contestants compete for a resource by persisting while constantly accumulating costs over the time that the contest lasts. Strategically, the game is an auction in which the prize goes to the player with the highest bid, and each player pays the loser's low bid.
829:
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, i.e. males defend territories only against other males and females only against other females. In this case, if the species is
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Many birds, particularly seabirds, nest in dense communities but are nonetheless territorial in defending their nesting site to within the distance they can reach while brooding. This is necessary to prevent attacks on their own chicks or nesting material from neighbours. Commonly the resulting
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An animal chooses its territory by deciding what part of its home range it will defend. In selecting a territory, the size and quality play crucial roles in determining an animal's habitat. Territory size generally tends to be no larger than the organism requires to survive, because defending a
188:
of the animals expressing the behaviour. Fitness in this biological sense relates to the ability of an animal to survive and raise young. The proximate functions of territory defense vary. For some animals, the reason for such protective behaviour is to acquire and protect food sources, nesting
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adult sons (and daughters) that are believed not to breed in their natal group and all group members participate in territorial defence. Males defend their territory using a nasty neighbour strategy. Group-living male breeders are nearly five times more aggressive towards their neighbours than
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is different from normal urination, which is done while squatting. This posture is exclusive to alpha wolves of either sex, although the alpha male does this most often. The alpha female usually urinates on a scent post that her breeding partner has just urinated on, although during the mating
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advertise their territories to other packs through a combination of scent marking and howling. Under certain conditions, wolf howls can be heard over areas of up to 130 km (50 sq mi). When howling together, wolves harmonize rather than chorus on the same note, thus creating the
220:
Reports of territory size can be confused by a lack of distinction between home range and the defended territory. The size and shape of a territory can vary according to its purpose, season, the amount and quality of resources it contains, or the geography. The size is usually a compromise of
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orchid bee. Males in this species of bee show alternative behaviours of territoriality and transiency. Transient male bees did not defend territories, but instead flew from one territory to the other. They also did not engage in physical contact with the territorial males. On the other hand,
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towards strangers, leading to the prediction that neighbours are the most important competitors for paternity. Using a molecular parentage analysis it has been shown that 28% of offspring are sired by neighbouring males and only 7% by strangers. In certain species of butterflies, such as the
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in which a territory-holder shows heightened aggression toward neighbouring territory-holders but unaffected aggression to unfamiliar animals or distant territory-holders. These contrasting strategies depend on which intruder (familiar or unfamiliar) poses the greatest threat to the resident
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hypothesis, claims that males have territories at distances sufficiently great that females are unable to discern already-mated males. The observation that males travelled long distances, ranging from 200m to 3.5 km, to find a second mate supports this argument. The debate about
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Behavioural ecologists have argued that food distribution determines whether a species is territorial or not, however, this may be too narrow a perspective. Several other type of resource may be defended including partners, potential mates, offspring, nests or lairs, display areas or
1037:) is dependent on the presence of females. Reduced aggression consistent with the dear enemy effect occurs between conspecific neighbours in the absence of females, but the presence of a female in a male's territory instigates comparably greater aggression between the neighbours.
661:) advertises its territory with urine scent marks. When it is urinating for marking purposes, it holds its extremely distinctive tail high in the air adding a visual component to the advertisement; when it is urinating for eliminative purposes, its tail is only slightly raised.
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superimposition of the short-range repulsion onto the long-range attraction characteristically leads to the well-known roughly hexagonal spacing of nests. One gets a similar hexagonal spacing resulting from the territorial behaviour of gardening limpets such as species of
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Many animals use vocalisations to advertise their territory. These are short-term signals transmitted only when the animal is present, but can travel long distances and over varied habitats. Examples of animals which use auditory signals include birds, frogs and canids.
1421:
801:. Such defense frequently involves a graded series of behaviours or displays that include threatening gestures (such as vocalizations, spreading of wings or gill covers, lifting and presentation of claws, head bobbing, tail and body beating) and finally, direct attack.
863:. This model pits a hawk strategy (always try to injure your opponent and only withdraw from the contest if an injury is received) against a dove strategy (always use a non-injurious display if the rival is another dove and always withdraw if the rival is a hawk).
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in which two neighbouring territorial animals become less aggressive toward one another once territorial borders are well-established and they are familiar to each other, but aggression toward unfamiliar animals remains unaffected. The converse of this is the
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913:. Territoriality emerges 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 excessive effort. Territoriality is often most strong towards
312:
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may be very territorial, especially in the breeding season during which they not only threaten or attack many kinds of intruders, but have stereotyped display behaviour to deter conspecifics sharing neighbouring nesting spots.
314:
599:, both sexes have glands that evolved for marking the nest. Males, although they have the gland, are unable to produce the marking substance. Females secrete it near the nest site entrance to establish their territory.
168:. The home ranges of different groups of animals often overlap, and in these overlap areas the groups tend to avoid each other rather than seeking to confront and expel each other. Within the home range there may be a
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larger territory incurs greater energy, time and risk of injury costs. For some animals, the territory size is not the most important aspect of territoriality, but rather the quality of the defended territory.
216:
Type F: Winter territory which typically includes foraging areas and roost sites. May be equivalent (in terms of location) to the Type A territory, or for a migratory species, may be on the wintering grounds.
2665:
Leiser, J.K. (2003). "When are neighbours 'dear enemies' and when are they not? The responses of territorial male variegated pupfish, Cyprinodon variegatus, to neighbours, strangers and heterospecifics".
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Territoriality is only shown by a minority of species. More commonly, an individual or a group of animals occupies an area that it habitually uses but does not necessarily defend; this is called its
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season, the female may first urinate on the ground. All other females in the pack, and also young wolves and low-ranking male wolves, urinate while squatting. Similar urination postures are used by
335:, auditory, or visual means, or a combination of these. If an intruder progresses further into the territory beyond the sign-posts and encounters the territory-holder, both animals may begin
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Animals use a range of behaviours to intimidate intruders and defend their territories, but without engaging in fights which are expensive in terms of energy and the risk of injury. This is
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Several species scratch or chew trees leaving a visual mark of their territory. This is sometimes combined with rubbing on the tree which may leave tufts of fur. These include the
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defend territories as pairs during the breeding season but as individuals during the winter. Resource availability may cause changes in territoriality. For example, some
2802:
Alatalo, Rauno V.; Carlson, Allan; Lundberg, Arne; Ulfstrand, Staffan (1981). "The
Conflict Between Male Polygamy and Female Monogamy: The Case of the Pied Flycatcher
1332:
Partecke, J.; von
Haeseler, A.; Wikelski, M. (2002). "Territory establishment in lekking marine iguanas, Amblyrhynchus cristatus: support for the hotshot mechanism".
1106:), researchers assert that males exhibit polyterritoriality to deceive females of the species into entering into polygynous relationships. This hypothesis, named the
2190:
1447:
Hurst, J.L.; Robertson, D.H.L.; Tolladay, U.; Beynon, R.J. (May 1998). "Proteins in urine scent marks of male house mice extend the longevity of olfactory signals".
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Since these territories are aggregated, females have a large selection of males with whom to potentially mate within the aggregation, giving females the power of
750:
931:
A western marsh harrier is mobbed by a northern lapwing. The marsh harrier, a male, had been quartering the ground in which lapwing and redshank were nesting.
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After leaving a urination mark, some animals scrape or dig the ground nearby, thereby leaving a visual advertisement of the territory. This includes domestic
64:
477:, is a behaviour used by animals to identify their territory. Most commonly, this is accomplished by depositing strong-smelling substances contained in the
627:
Ring-tailed lemurs hold their distinctive tails high in the air during territorial scent marking. They also engage in "stink fights" with intruding males.
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require an extensive protected area to guarantee their food supply. This territoriality only breaks down when there is a glut of food, for example when
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areas. The red breast of the bird (i.e. badge) is highly visible when it sings (vocal marking) at the boundary of its territory. The ring-tailed lemur (
652:
Some animals have prominent "badges" or visual displays to advertise their territory, often in combination with scent marking or auditory signals. Male
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rarely defend an area larger than the nest. Conversely, other insectivorous birds that occupy more constrained territories, such as the ground-nesting
2123:
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communicate to other animals that the territory is occupied and may also communicate additional information such as the sex, reproductive status or
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combined with the smell of the marking animal. The territorial male scrape-marks every 30 m (98 ft) or so around its territory boundary.
605:
use feces to mark their territory. They have evolved specialized intestinal anatomy to produce cubical feces to ensure the feces do not roll away.
2775:
Stern, David L. (1991-10-01). "Male
Territoriality and Alternative Male Behaviors in the Euglossine Bee, Eulaema meriana (Hymenoptera: Apidae)".
988:. They vigorously defend their gardens of particular species of algae, that extend for perhaps 1–2 cm around the periphery of their shells.
1212:
1536:
Ryon, Jenny; Fentress, J. C.; Harrington, F. H.; Bragdon, Susan (1986-03-01). "Scent rubbing in wolves (Canis lupus): the effect of novelty".
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polyterritoriality in this species may initiate research about the evolution and reasons for polyterritoriality in other unrelated species.
286:) forage on beaches and sandflats. When on beaches, they feed either in flocks or individual territories of 10 to 120 metres of shoreline.
2206:
Piñeiro, A.; Barja, I. (2002). "The plant physical features selected by wildcats as signal posts: an economic approach to fecal marking".
1027:, the male defends territories that receptive females are likely to fly through such as sunny hilltops and sunspots on a forest's floor.
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Food related territoriality is least likely with insectivorous birds, where the food supply is plentiful but unpredictably distributed.
590:. To do this, they perform a handstand to mark vertical surfaces, grasping the highest point with their feet while applying the scent.
51:
1014:) is group living with one single breeding male and up to 4 communally breeding females per group. Groups typically contain several
1921:
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or unmated pair, or a group. Territoriality is not always a fixed behavioural characteristic of a species. For example, red foxes (
1434:
2012:
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may defend feeding territories that are distant from their nest sites, and in some species that form leks, for example in the
717:) deposit their faecal marks on plants with high visual conspicuousness that enhances the visual effectiveness of the signal.
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or marks on the vegetation or ground. Visual marking of territory is often combined with other modes of animal communication.
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Burst, T.L. and Pelton, M.R., ( ). Black bear mark trees in the Smoky
Mountains. Int. Conf. Bear Res. and Manage., 5: 45-53
1977:
Gerling, Dan; Hermann, Henry R (1978). "Biology and mating behavior of
Xylocopa virginica L. (Hymenoptera, Anthophoridae)".
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Many species demonstrate polyterritoriality, referring to the act of claiming or defending more than one territory. In the
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Some species of penguin defend their nests from intruders trying to steal the pebbles from which the nest is constructed.
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can operate whereby the cost to the benefactor in helping defend the territory is less than the gains to the beneficiary.
1894:
2483:"A test for correlations among fitness-linked behavioural traits in the spider Agelenopsis aperta (Araneae, Agelenidae)"
2740:
Kimsey, Lynn Siri (1980). "The behaviour of male orchid bees (Apidae, Hymenoptera, Insecta) and the question of leks".
2620:
Davies, N.B. (1978). "Territorial defense in the speckled wood butterfly (Pararge aegeria): The resident always wins".
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Although most territories contain multiple (potential) resources, some territories are defended for only one purpose.
2866:
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562:, it lifts a hind leg and urinates on a scent post (usually an elevated position like a tree, rock, or bush). This
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Type C: A nesting territory which includes the nest plus a small area around it. Common in colonial waterbirds.
17:
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Type A: An 'all-purpose territory' in which all activities occur, e.g. courtship, mating, nesting and foraging
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does not happen, the territory may be defended by actual fighting, although this is generally a last resort.
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larger and larger. Such a dung heap can become up to five metres wide and one metre high. After defecating,
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245:
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to stabilize the odours and maintain them for longer. The animal sniffing the scent frequently displays a
2917:
2394:
Nursall, J.R. (1977). "Territoriality in Redlip blennies (Ophioblennius atlanticus-Pisces: Blenniidae)".
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often engages in fights over its territory and the most combative spiders have the largest territories.
2036:
1951:
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470:
31:
2530:"The nasty neighbour in the striped mouse (Rhabdomys pumilio) steals paternity and elicits aggression"
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1216:
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389:
2703:"When to be a dear enemy: flexible acoustic relationships of neighbouring skylarks, Alauda arvensis"
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586:) scent-mark both vertical and horizontal surfaces at the overlaps in their home ranges using their
180:
The ultimate function of animals inhabiting and defending a territory is to increase the individual
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reptile. Males start to establish small display territories two months ahead of the mating season.
244:) have highly variable territory sizes, ranging from less than 4,000 hectares (9,900 acres) in the
210:
Type D: A pairing and mating territory. The type of territory defended by males in lekking species.
2581:"The mating system of Vanessa kershawi: Males defend landmark territories as mate encounter sites"
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Fisher, J. (1954). "Evolution and bird sociality". In Huxley, J.; Hardy, A.; Ford, E. (eds.).
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1308:"Territory size in wintering Sanderlings: the effects of prey abundance and intruder density"
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Branch, G.M., Griffiths, C., Beckley, L.E., Branch, M.L.; Two Oceans. Pub. Struik, 2010.
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Some species of bees also exhibit territoriality to defend mating sites. For example, in
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have territories of only a few square centimetres in the immediate vicinity of the nest.
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Type B: A mating and nesting territory, not including most of the area used for foraging.
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The
Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates
1927:
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The
Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates
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1998:
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817:) either establish stable home ranges within particular areas or are itinerant with
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in many mammals. In many mammal species, scent marking is more frequent during the
498:
356:
355:"Urine spraying" redirects here. For spraying of urine as an animal repellent, see
264:
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that no other individual group uses, but, again, this is as a result of avoidance.
2718:
1090:) and the marine iguana, males defend the lek site which is used only for mating.
2194:
1867:
Johnston, Robert E.; Müller-Schwarze, Dietland; Sorenson, Peter W. (2012-12-06).
1841:
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834:, one male territory probably contains several female territories, while in some
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Territorial scent marking may involve behaviours specific to this activity. When
544:
506:
249:
229:
153:. Animals that actively defend territories in this way are referred to as being
2061:
1177:"Patterns of scent marking with urine and faeces amongst carnivore communities."
641:
The antebrachial scent gland and spur on the forearm of a male ring-tailed lemur
403:
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Marchlewska-Koj, Anna; Lepri, John J.; Müller-Schwarze, Dietland (2012-12-06).
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2062:"Urine marking and urination in Lemur catta: a comparison of design features"
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was created from a revision of this article dated 5 September 2019
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921:. This is because the conspecifics share exactly the same set of resources.
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status of the territory-holder. Sign-posts may communicate information by
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resource needs, defense costs, predation pressure and reproductive needs.
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Two domestic cats posturing during ritualized aggression over a territory
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1990:
1897:. wolfbehavior: All you need to know about Wolves. 2005. Archived from
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271:) territories are about 600 square metres (6,500 sq ft) and
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2347:"'Nasty neighbours' rather than 'dear enemies' in a social carnivore"
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Symposia of the
Zoological Society of London. Vol. 45. No. 107. 1980.
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to assist in detecting the mark. Scent marking is often performed by
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104:
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Wild
Mammals of North America: Biology, Management, and Conservation
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Gazelles and their relatives : a study in territorial behavior
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Habitats and territories; a study of the use of space by animals
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Animals may use several strategies to defend their territories.
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located on various areas of the body. Often, the scent contains
42:
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602:
568:
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131:
114:
1598:
Elbroch, Mark; Kresky, Michael; Evans, Jonah Wy (2012-02-11).
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A reassessment of the function of scent marking in territories
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may claim as much as 10 hectares (25 acres) of territory. For
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482:
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1803:
Mittermeier, R. A.; Rylands, A. B.; Konstant, W. R. (1999).
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The Spotted Hyena: A Study of Predation and Social Behaviour
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The time to develop territories varies between animals. The
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Among birds, territories have been classified as six types.
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Feldhamer, G.A., Thompson, B.C. and Chapman, J.A., (2003).
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512:
457:
2013:"Scientist figures out how wombats poop cubes | CBC Radio"
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Several types of resource in a territory may be defended.
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scent-mark by urinating on or rubbing against vegetation.
2527:
1923:
Mammalian Olfaction, Reproductive Processes, and Behavior
1815:(6th ed.). Johns Hopkins University Press. pp.
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681:
821:. Territories may vary with time (season). For example,
2572:
1655:
The Chemistry of Pheromones and Other Semiochemicals II
667:
have poor vision but may use visual marking. Dominant
2861:
Stroudsburg, Pa., Dowden, Hutchinson & Ross 398,
263:) have territories of 9,000 hectares (22,000 acres),
2528:
Schradin, C.; Schneider, C.; Lindholm, A.K. (2010).
2421:
Stimson J (1969). "Territoriality of the owl limpet
469:
Scent marking, also known as territorial marking or
1601:
Field Guide to Animal Tracks and Scat of California
1305:
2843:Walther, F. R., E. C. Mungall, G. A. Grau. (1983)
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2087:
1949:
1808:
1761:
1597:
1306:Myers, J.P.; Connors, P.G.; Pitelka, F.A. (1979).
1283:
866:Another strategy used in territory defence is the
2700:
1570:
535:also use scent marking, including urine washing (
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2847:Park Ridge, N.J. : Noyes Publications 239,
1678:
248:to over 100,000 hectares (250,000 acres) in the
2477:
2313:harvnb error: no target: CITEREFMacdonald1987 (
1945:
1943:
1840:L. David Mech; Luigi Boitani (1 October 2010).
1833:
1489:
1284:Ehrlich, P.R.; Dobkin, D.S.; Wheye, D. (1998).
938:Large solitary (or paired) carnivores, such as
2059:
1976:
1073:
1030:Territory defence in male variegated pupfish (
2165:
2085:
1759:
1679:Sunquist, Mel; Sunquist, Fiona (2017-05-15).
1564:
1191:"Ornithology: territoriality and coloniality"
2344:
2205:
1940:
1492:"Urinary Volatile Constituents of the Lion,
1063:. Similar behaviour is also observed in the
809:Territories may be held by an individual, a
343:
141:(or, occasionally, against animals of other
2777:Journal of the Kansas Entomological Society
2613:
2121:
2094:. University of California Press. pp.
2079:
1870:Advances in Chemical Signals in Vertebrates
1843:Wolves: Behavior, Ecology, and Conservation
1768:. University of California Press. pp.
890:In territory defence by groups of animals,
539:the body with urine), to communicate. Many
236:or artificial feeding in suburban gardens.
189:sites, mating areas, or to attract a mate.
2701:Briefer, E.; Rybak, F.; Aubin, T. (2008).
2334:. London: Allen and Unwin. pp. 71–83.
1958:. Wisconsin Primate Research Center (WPRC)
1732:
1574:Carnivore Behavior, Ecology, and Evolution
2633:
2596:
2555:
2545:
2370:
2308:
1873:. Springer Science & Business Media.
1753:
1726:
1658:. Springer Science & Business Media.
1577:. Springer Science & Business Media.
1400:. Springer Science & Business Media.
1370:Societies of Wolves and Free-ranging Dogs
1279:
1277:
547:, use scent marking from two glands, the
2928:, and does not reflect subsequent edits.
2911:
2290:The Wolf: His Place in the Natural World
2029:
1952:"Primate Factsheets: Ring-tailed lemur (
1805:"Primates of the world: An introduction"
1169:
926:
788:
778:
308:
304:
67:of all important aspects of the article.
2420:
2393:
873:Some animals use a strategy termed the
27:Area a wild animal consistently defends
14:
2945:
2739:
2664:
2619:
2585:Journal of Research on the Lepidoptera
2327:
2302:
1651:
1440:
1366:
1274:
1210:
63:Please consider expanding the lead to
2795:
2774:
1360:
1093:
899:
2110:urine OR urination OR scent marking.
1919:
1705:
1624:
1490:Andersen, K.F.; Vulpius, T. (1999).
1437:." Science 171.3970 (1971): 443-449.
36:
2579:Alcock, John; Gwynne Daryl (1988).
2345:Müller, C.A.; Manser, M.B. (2007).
2124:"Factfile:Greater one horned rhino"
1979:Behavioral Ecology and Sociobiology
1334:Behavioral Ecology and Sociobiology
130:is the sociographical area that an
24:
2898:
2837:
2481:; Hedrick, Ann V. (October 1993).
2408:10.1111/j.1469-7998.1977.tb04156.x
1604:. University of California Press.
1571:John L. Gittleman (9 March 2013).
1264:. University of California Press,
1213:"Squirrel behaviour and territory"
731:
25:
2974:
2879:
2272:J.M. Dent and Sons Limited. p.38
2060:Palagi, E.; Dapporto, L. (2006).
1397:Chemical Signals in Vertebrates 9
1237:
1188:
192:
149:or (less commonly) real physical
117:mark their territories with urine
2910:
1424:." Ethology 60.2 (1982): 89-118.
762:Problems playing this file? See
747:
634:
620:
493:or carrier proteins such as the
443:
434:
411:
402:
378:
369:
350:
103:
90:
41:
2768:
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2658:
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2414:
2387:
2338:
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2242:
2199:
2181:
2159:
2137:
2115:
2053:
2005:
1970:
1913:
1887:
1860:
1846:. University of Chicago Press.
1796:
1736:The Study of Prosimian Behavior
1699:
1685:. University of Chicago Press.
1672:
1645:
1618:
1591:
1529:
1483:
1427:
1414:
1387:
55:may be too short to adequately
2857:Stokes, A. W. (editor) (1974)
1733:G.A. Doyle (2 December 2012).
1373:. Cambridge University Press.
1367:Spotte, Stephen (2012-03-15).
1325:
1299:
1254:
1231:
1204:
1182:
842:, this situation is reversed.
593:In the Eastern carpenter bee,
65:provide an accessible overview
13:
1:
2754:10.1016/s0003-3472(80)80088-1
2719:10.1016/j.anbehav.2008.06.017
1811:Walker's Mammals of the World
1652:Schulz, Stefan (2005-01-07).
1163:
845:
422:Wolves mark their territories
134:consistently defends against
2875:New York, Basic Books 117 p.
2644:10.1016/0003-3472(78)90013-1
1920:Doty, Richard (2012-12-02).
1631:. Cornell University Press.
857:of fighting is known as the
7:
1950:Cawthon Lang, K.A. (2005).
1538:Canadian Journal of Zoology
1114:
1074:Single resource territories
950:are attracted to migrating
720:
322:scent-marking her territory
278:Territories can be linear.
213:Type E: Roosting territory.
175:
10:
2979:
2168:"Black Bear Marking Trees"
2037:"The RSPB-Robin:Territory"
1706:Ewer, R. F. (2013-12-11).
1157:The Territorial Imperative
917:, as shown in the case of
804:
782:
560:a wolf marks its territory
460:spray urine on scent posts
354:
32:Territory (disambiguation)
29:
2220:10.1007/s00114-012-0962-9
1807:. In Nowak, R. M. (ed.).
1346:10.1007/s00265-002-0469-z
991:The desert grass spider,
674:greater one-horned rhinos
611:
551:and a scent gland in the
344:Advertising the territory
2292:. Souvenir Press. p.73.
2145:"chemical communication"
1100:European pied flycatcher
2958:Biological interactions
2808:The American Naturalist
1515:10.1093/chemse/24.2.179
1435:Mammalian scent marking
1025:speckled wood butterfly
1021:Australian painted lady
588:anogenital scent glands
485:, or, from specialised
295:Amblyrhynchus cristatus
2906:
2886:Listen to this article
2871:Klopfer, P. H. (1969)
2680:10.1006/anbe.2003.2087
2547:10.1186/1742-9994-7-19
2499:10.1006/anbe.1993.1243
2363:10.1098/rspb.2006.0222
2331:Evolution As a Process
2166:Cabrera, K.A. (2013).
2086:Richard Estes (1991).
1760:Richard Estes (1991).
1682:Wild Cats of the World
1461:10.1006/anbe.1997.0650
1242:. www.badgerland.co.uk
1141:Dear enemy recognition
1131:Biological interaction
932:
883:nasty neighbour effect
794:
736:
495:major urinary proteins
323:
2963:Animals by adaptation
2905:
2268:Lopez, B.H., (1978).
1005:Mating opportunities:
930:
799:ritualized aggression
792:
785:Ritualized aggression
779:Ritualized aggression
735:
337:ritualized aggression
317:
305:Retaining a territory
2937:More spoken articles
2534:Frontiers in Zoology
2252:. JHU Press. p.496.
1625:Ewer, R. F. (1998).
1175:MacDonald, David W.
981:Nests and offspring:
838:species such as the
564:raised leg urination
30:For other uses, see
2288:Zimen, E., (1981).
2208:Naturwissenschaften
1709:Ethology of Mammals
1433:Ralls, Katherine. "
1260:Kruuk, H., (1972).
1080:European blackbirds
1054:Euglossa imperialis
1007:The striped mouse (
893:reciprocal altruism
697:American black bear
473:when this involves
147:agonistic behaviors
2907:
2804:Ficedula hypoleuca
2479:Riechert, Susan E.
2396:Journal of Zoology
2270:Of Wolves and Men.
2193:2010-12-03 at the
2069:Ann. Zool. Fennici
1991:10.1007/BF00294984
1104:Ficedula hypoleuca
1094:Polyterritoriality
1023:butterfly and the
994:Agelenopsis aperta
963:blacksmith lapwing
933:
900:Resources defended
887:territory-holder.
795:
737:
596:Xylocopa virginica
580:ring-tailed lemurs
543:, for example the
324:
2903:
2467:978-1-77007-772-0
2357:(1612): 959–965.
2105:978-0-520-08085-0
1933:978-0-323-15450-5
1880:978-1-4615-4733-4
1853:978-0-226-51698-1
1826:978-0-8018-6251-9
1779:978-0-520-08085-0
1746:978-0-323-14306-6
1719:978-1-4899-4656-0
1692:978-0-226-51823-7
1665:978-3-540-21308-6
1638:978-0-8014-8493-3
1611:978-0-520-25378-0
1584:978-1-4757-4716-4
1407:978-1-4615-0671-3
1380:978-1-107-37910-7
876:dear enemy effect
855:game theory model
752:
711:European wildcats
578:Males and female
533:New World monkeys
315:
269:Empidonax minimus
265:least flycatchers
261:Aquila chrysaetos
246:Ngorongoro Crater
186:inclusive fitness
82:
81:
16:(Redirected from
2970:
2927:
2925:
2914:
2913:
2904:
2894:
2892:
2887:
2832:
2831:
2799:
2793:
2792:
2772:
2766:
2765:
2742:Animal Behaviour
2737:
2731:
2730:
2713:(4): 1319–1325.
2707:Animal Behaviour
2698:
2692:
2691:
2668:Animal Behaviour
2662:
2656:
2655:
2637:
2622:Animal Behaviour
2617:
2611:
2610:
2600:
2598:10.5962/p.266708
2591:(1–4): 116–124.
2576:
2570:
2569:
2559:
2549:
2525:
2519:
2518:
2487:Animal Behaviour
2475:
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2154:
2152:
2141:
2135:
2134:
2132:
2130:
2122:von Houwald, F.
2119:
2113:
2112:
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2009:
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2002:
1974:
1968:
1967:
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1938:
1937:
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1911:
1910:
1908:
1906:
1901:on June 29, 2013
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1500:
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1481:
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1449:Animal Behaviour
1444:
1438:
1431:
1425:
1420:Gosling, L. M. "
1418:
1412:
1411:
1391:
1385:
1384:
1364:
1358:
1357:
1329:
1323:
1322:
1312:
1303:
1297:
1296:
1294:
1292:
1286:"Territoriality"
1281:
1272:
1258:
1252:
1251:
1249:
1247:
1235:
1229:
1228:
1226:
1224:
1219:on June 15, 2013
1215:. Archived from
1208:
1202:
1201:
1199:
1197:
1186:
1180:
1173:
1088:grazing antelope
868:war of attrition
754:
753:
734:
715:Felis silvestris
701:Ursus americanus
638:
624:
549:preorbital gland
499:flehmen response
447:
438:
415:
406:
382:
373:
357:Animal repellent
316:
230:European badgers
224:Some species of
107:
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77:
74:
68:
45:
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2929:
2923:
2921:
2918:This audio file
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2908:
2899:
2896:
2890:
2889:
2885:
2882:
2840:
2838:Further reading
2835:
2800:
2796:
2773:
2769:
2748:(4): 996–1004.
2738:
2734:
2699:
2695:
2663:
2659:
2635:10.1.1.513.4639
2618:
2614:
2577:
2573:
2526:
2522:
2476:
2472:
2460:
2456:
2441:10.2307/1933604
2423:Lottia gigantea
2419:
2415:
2392:
2388:
2351:Proc. R. Soc. B
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2322:
2312:
2307:
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2263:
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2214:(10): 801–809.
2204:
2200:
2195:Wayback Machine
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1895:"The Territory"
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1170:
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1126:Alpha (biology)
1117:
1096:
1076:
1066:Eulaema meriana
1046:Alauda arvensis
975:Lottia gigantea
942:and the bigger
902:
848:
840:northern jacana
823:European robins
807:
787:
781:
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693:Lynx canadensis
654:European robins
646:
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545:blue wildebeest
507:breeding season
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390:rub their heads
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242:Crocuta crocuta
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2880:External links
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2820:10.1086/283756
2814:(5): 738–753.
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2309:Macdonald 1987
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1791:urine marking.
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1628:The Carnivores
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860:hawk-dove game
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819:no fixed abode
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783:Main article:
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743:Wolves howling
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59:the key points
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18:Territoriality
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2041:RSPB website
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1494:Panthera leo
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2628:: 138–147.
1956:) Behavior"
1954:Lemur catta
1785:25 December
1211:Turpin, K.
1061:mate choice
1016:philopatric
836:polyandrous
689:Canada lynx
669:white rhino
659:Lemur catta
584:Lemur catta
318:Video of a
280:Sanderlings
155:territorial
139:competition
136:conspecific
2947:Categories
2933:Audio help
2924:2019-09-05
2258:0801874165
2075:: 280–284.
2046:2008-05-17
2022:2018-11-20
1321:: 551–561.
1270:0226455084
1164:References
1146:Home range
1121:Aggression
1084:Uganda kob
1035:variegatus
1033:Cyprinodon
971:owl limpet
853:The first
846:Strategies
832:polygynous
764:media help
695:) and the
665:Rhinoceros
529:Prosimians
491:pheromones
424:with urine
255:In birds,
234:food waste
166:home range
151:aggression
2859:Territory
2630:CiteSeerX
2607:127040681
2507:0003-3472
1558:0008-4301
1108:deception
1010:Rhabdomys
541:ungulates
475:urination
333:olfactory
329:dominance
226:squirrels
170:core area
128:territory
57:summarize
2953:Ethology
2935: ·
2828:85184225
2789:25085309
2762:53161684
2727:21703019
2688:53166315
2566:20573184
2515:53201859
2381:17251103
2228:22926138
2191:Archived
2173:June 28,
2151:June 28,
2129:June 25,
1999:33309938
1962:June 29,
1905:June 28,
1523:10321819
1291:June 24,
1246:June 23,
1223:June 22,
1196:June 22,
1151:Property
1115:See also
721:Auditory
521:leopards
519:such as
471:spraying
250:Kalahari
176:Function
145:) using
124:ethology
2922: (
2893:minutes
2652:7445027
2557:2908079
2449:1933604
2428:Ecology
2372:2141673
2236:2309455
1477:9879771
1469:9632512
1354:2132629
1042:Skylark
1040:In the
1012:pumilio
944:raptors
805:Defense
603:Wombats
569:coyotes
525:jaguars
388:Felids
320:tigress
299:lekking
297:) is a
182:fitness
143:species
115:Jackals
2865:
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1238:Anon.
1189:Anon.
959:Swifts
952:salmon
772:Wolves
612:Visual
517:felids
483:faeces
454:Tigers
132:animal
2824:S2CID
2785:JSTOR
2758:S2CID
2723:S2CID
2684:S2CID
2648:S2CID
2603:S2CID
2511:S2CID
2445:JSTOR
2398:. 2.
2232:S2CID
2065:(PDF)
1995:S2CID
1499:(PDF)
1473:S2CID
1350:S2CID
1311:(PDF)
940:bears
936:Food:
811:mated
513:Bears
479:urine
458:lions
273:gulls
2863:ISBN
2849:ISBN
2562:PMID
2503:ISSN
2463:ISBN
2377:PMID
2315:help
2294:ISBN
2274:ISBN
2254:ISBN
2224:PMID
2175:2013
2153:2013
2131:2013
2100:ISBN
1964:2013
1928:ISBN
1907:2013
1875:ISBN
1848:ISBN
1821:ISBN
1817:1–52
1787:2012
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1633:ISBN
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969:The
911:leks
705:spur
682:dogs
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