Territory (animal)
Encyclopedia
In ethology
Ethology
Ethology is the scientific study of animal behavior, and a sub-topic of zoology....

 the term territory refers to any sociographical area that an animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

 of a particular species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 consistently defends against conspecifics
Conspecificity
Conspecificity is a concept in biology. Two or more individual organisms, populations, or taxa are conspecific if they belong to the same species....

 (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
Nest
A nest is a place of refuge to hold an animal's eggs or provide a place to live or raise offspring. They are usually made of some organic material such as twigs, grass, and leaves; or may simply be a depression in the ground, or a hole in a tree, rock or building...

, 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
Bird song
Bird vocalization includes both bird calls and bird songs. In non-technical use, bird songs are the bird sounds that are melodious to the human ear. In ornithology and birding, songs are distinguished by function from calls.-Definition:The distinction between songs and calls is based upon...

, or the calls of gibbon
Gibbon
Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...

s) 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
Urination
Urination, also known as micturition, voiding, peeing, weeing, pissing, and more rarely, emiction, is the ejection of urine from the urinary bladder through the urethra to the outside of the body. In healthy humans the process of urination is under voluntary control...

, by defecation
Defecation
Defecation is the final act of digestion by which organisms eliminate solid, semisolid or liquid waste material from the digestive tract via the anus. Waves of muscular contraction known as peristalsis in the walls of the colon move fecal matter through the digestive tract towards the rectum...

, or by rubbing parts of the bodies that bear specialised scent gland
Scent gland
Scent glands are found in the genital area of most mammals and in various other parts of the body, such as the underarms of humans and the preorbital glands of deer and muskox. They produce a semi-viscous fluid which contains pheromones. These odor-messengers indicate information such as status,...

s against the substrate. For example, dog
Dog
The domestic dog is a domesticated form of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The dog may have been the first animal to be domesticated, and has been the most widely kept working, hunting, and companion animal in...

s and other canids scent-mark by urination and defecation, 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 prosimian
Prosimian
Prosimians are a grouping of mammals defined as being primates, but not monkeys or apes. They include, among others, lemurs, bushbabies, and tarsiers. They are considered to have characteristics that are more primitive than those of monkeys and apes. Prosimians are the only primates native to...

s use territorial marking; for example, the Red-bellied Lemur
Red-bellied Lemur
The Red-bellied Lemur is a medium sized prosimian with a luxuriant chestnut brown coat. This lemur is endemic to eastern Madagascan rainforests and is distinguished by patches of white skin below the eyes, giving rise to a "teardrop" effect, particularly conspicuous in the male.The species, first...

 creates territories for groups of two to ten individuals in the rainforest
Rainforest
Rainforests are forests characterized by high rainfall, with definitions based on a minimum normal annual rainfall of 1750-2000 mm...

s of eastern Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...

 by scent marking: the male Diademed Sifaka
Diademed Sifaka
The diademed sifaka , or diademed simpona, is an endangered species of sifaka, one of the lemurs endemic to certain rainforests in eastern Madagascar. This species is one of the world's largest living lemurs, with a total adult length of approximately 105 centimetres , half of which is tail...

 also scent marks defended territories in some of these same rainforests. Humans build physical impediments
Fence
A fence is a freestanding structure designed to restrict or prevent movement across a boundary. It is generally distinguished from a wall by the lightness of its construction: a wall is usually restricted to such barriers made from solid brick or concrete, blocking vision as well as passage .Fences...

 and visual warnings to mark defended territories. The male Western fence lizard
Western fence lizard
The western fence lizard is a common lizard of California and the surrounding area. Because the ventral abdomen of an adult is characteristically blue, it is also known as the blue-belly...

 defends a territory by posturing and combat, but less intensely after the mating season.

Invertebrates which show territoriality include some ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s and bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s, and the owl limpet 

Spraying

Spraying (also known as territorial marking) is behavior
Behavior
Behavior or behaviour refers to the actions and mannerisms made by organisms, systems, or artificial entities in conjunction with its environment, which includes the other systems or organisms around as well as the physical environment...

 used by animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s to identify their territory. Most commonly, this is scent marking, accomplished by depositing strong-smelling chemicals
Chemical compound
A chemical compound is a pure chemical substance consisting of two or more different chemical elements that can be separated into simpler substances by chemical reactions. Chemical compounds have a unique and defined chemical structure; they consist of a fixed ratio of atoms that are held together...

 such as urine
Urine
Urine is a typically sterile liquid by-product of the body that is secreted by the kidneys through a process called urination and excreted through the urethra. Cellular metabolism generates numerous by-products, many rich in nitrogen, that require elimination from the bloodstream...

 at prominent locations within the territory. Often the scent contains carrier proteins, such as the major urinary proteins
Major urinary proteins
Major urinary proteins , also known as α2u-globulins, are a subfamily of proteins found in abundance in the urine and other secretions of many animals. Mups provide a small range of identifying information about the donor animal, when detected by the vomeronasal organ of the receiving animal. They...

, to stabilize the odors and maintain them for longer.

Not only does the marking communicate to others of the same species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

, but it is also noted by prey species and avoided. For example felids such as leopard
Leopard
The leopard , Panthera pardus, is a member of the Felidae family and the smallest of the four "big cats" in the genus Panthera, the other three being the tiger, lion, and jaguar. The leopard was once distributed across eastern and southern Asia and Africa, from Siberia to South Africa, but its...

s and jaguar
Jaguar
The jaguar is a big cat, a feline in the Panthera genus, and is the only Panthera species found in the Americas. The jaguar is the third-largest feline after the tiger and the lion, and the largest in the Western Hemisphere. The jaguar's present range extends from Southern United States and Mexico...

s mark by rubbing themselves against vegetation. Some prosimian
Prosimian
Prosimians are a grouping of mammals defined as being primates, but not monkeys or apes. They include, among others, lemurs, bushbabies, and tarsiers. They are considered to have characteristics that are more primitive than those of monkeys and apes. Prosimians are the only primates native to...

s, such as the Red-bellied Lemur
Red-bellied Lemur
The Red-bellied Lemur is a medium sized prosimian with a luxuriant chestnut brown coat. This lemur is endemic to eastern Madagascan rainforests and is distinguished by patches of white skin below the eyes, giving rise to a "teardrop" effect, particularly conspicuous in the male.The species, first...

, also use scent marking to establish a territory. Many ungulate
Ungulate
Ungulates are several groups of mammals, most of which use the tips of their toes, usually hoofed, to sustain their whole body weight while moving. They make up several orders of mammals, of which six to eight survive...

s, for example the Blue Wildebeest
Blue Wildebeest
The Blue Wildebeest , also called the Common Wildebeest, is a large antelope and one of two species of wildebeest. It grows to 115–145 cm shoulder height and attains a body mass of 168–274 kg. They range the open plains, bushveld and dry woodlands of Southern and East Africa, living for...

, use scent marking from two glands, the preorbital gland and a scent gland in the hoof
Hoof
A hoof , plural hooves or hoofs , is the tip of a toe of an ungulate mammal, strengthened by a thick horny covering. The hoof consists of a hard or rubbery sole, and a hard wall formed by a thick nail rolled around the tip of the toe. The weight of the animal is normally borne by both the sole...

.

Defense

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
European Robin
The European Robin , most commonly known in Anglophone Europe simply as the Robin, is a small insectivorous passerine bird that was formerly classed as a member of the thrush family , but is now considered to be an Old World flycatcher...

 defend territories as pairs during the breeding season and as individuals during the winter, while some nectarivore
Nectarivore
In zoology, nectarivore is an animal which eats the sugar-rich nectar produced by flowering plants. Most nectarivores are insects or birds, but there are also nectarivorous mammals, notably several species of bats in the Southwestern United States and Mexico, as well as the Australian Honey Possum...

s 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
Polygyny
Polygyny is a form of marriage in which a man has two or more wives at the same time. In countries where the practice is illegal, the man is referred to as a bigamist or a polygamist...

, one male territory will probably contain several female territories, while in some polyandrous
Polyandry
Polyandry refers to a form of marriage in which a woman has two or more husbands at the same time. The form of polyandry in which a woman is married to two or more brothers is known as "fraternal polyandry", and it is believed by many anthropologists to be the most frequently encountered...

 species such as the Northern Jacana
Northern Jacana
The Northern Jacana or Northern Jaçana is a wader which is a resident breeder from coastal Mexico to western Panama, and on Cuba, Jamaica and Hispaniola. It sometimes breeds in Texas, USA, and has also been recorded on several occasions as a vagrant in Arizona...

, 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
Kob
The Kob is an antelope found across Sub-Saharan Africa from Senegal to South Sudan. Found along the Northern Savanna, often seen in Murchison Falls and Queen Elizabeth National Park, Uganda; Garamba and Virunga National Park, Democratic Republic of the Congo, as well as grassy floodplains of...

 (a grazing antelope
Grazing antelope
A grazing antelope is any of the species of antelope that make up the subfamily Hippotraginae in the family Bovidae, which also includes sheep, goats, and cattle.* FAMILY BOVIDAE...

), 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
Home range
Home range is the area where an animal lives and travels in. It is closely related to, but not identical with, the concept of "territory".The concept that can be traced back to a publication in 1943 by W. H. Burt, who constructed maps delineating the spatial extent or outside boundary of an...

. 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
Scutellastra
Scutellastra is a genus of sea snails with gills, typical true limpets, marine gastropod mollusks in the family Patellidae, the true limpets.-Species:Species within the genus Scutellastra include:* Scutellastra aphanes...

  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. Swift
Swift
The swifts are a family, Apodidae, of highly aerial birds. They are superficially similar to swallows, but are actually not closely related to passerine species at all; swifts are in the separate order Apodiformes, which they share with hummingbirds...

s rarely defend an area larger than the nest. Conversely, other insectivorous birds that occupy more constrained territories, such as the ground-nesting Blacksmith Lapwing
Blacksmith Lapwing
The Blacksmith Lapwing or Blacksmith Plover occurs commonly from Kenya through central Tanzania to southern and southwestern Africa...

 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 bear
Bear
Bears are mammals of the family Ursidae. Bears are classified as caniforms, or doglike carnivorans, with the pinnipeds being their closest living relatives. Although there are only eight living species of bear, they are widespread, appearing in a wide variety of habitats throughout the Northern...

s and the bigger raptors
Bird of prey
Birds of prey are birds that hunt for food primarily on the wing, using their keen senses, especially vision. They are defined as birds that primarily hunt vertebrates, including other birds. Their talons and beaks tend to be relatively large, powerful and adapted for tearing and/or piercing flesh....

 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 Bear
Grizzly Bear
The grizzly bear , also known as the silvertip bear, the grizzly, or the North American brown bear, is a subspecies of brown bear that generally lives in the uplands of western North America...

s are attracted to migrating salmon
Salmon
Salmon is the common name for several species of fish in the family Salmonidae. Several other fish in the same family are called trout; the difference is often said to be that salmon migrate and trout are resident, but this distinction does not strictly hold true...

.

See also

  • Aggression
    Aggression
    In psychology, as well as other social and behavioral sciences, aggression refers to behavior between members of the same species that is intended to cause humiliation, pain, or harm. Ferguson and Beaver defined aggressive behavior as "Behavior which is intended to increase the social dominance of...

  • Alpha (biology)
    Alpha (biology)
    In social animals, the alpha is the individual in the community with the highest rank. Where one male and one female fulfill this role, they are referred to as the alpha pair...

  • Biological interaction
    Biological interaction
    Biological interactions are the effects organisms in a community have on one another. In the natural world no organism exists in absolute isolation, and thus every organism must interact with the environment and other organisms...

  • Dear enemy recognition
    Dear enemy recognition
    In ethology, dear enemy recognition is a situation in which a territorial animal responds more strongly to strangers than to its neighbors from adjacent territories. This phenomenon may be generally advantageous to an animal because it minimizes time and energy spent on territorial defense, and...


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.
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