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71-611: 5–10, see text. See text The tits , chickadees , and titmice constitute the Paridae , a large family of small passerine birds which occur mainly in the Northern Hemisphere and Africa . Most were formerly classified in the genus Parus . Eurasian and African members of this family are referred to as "tits", while North American species are called either "chickadees" ( onomatopoeic , derived from their distinctive "chick-a dee dee dee" alarm call ) or "titmice". The name titmouse

142-438: A core area that no other individual group uses, but, again, this is as a result of avoidance. The ultimate function of animals inhabiting and defending a territory is to increase the individual fitness or inclusive fitness 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,

213-456: A subfamily Parinae . Alternatively, all tits—save the two monotypic genera discussed in the preceding section and possibly Cyanistes , but including Hume's ground tit—could be lumped in Parus . In any case, four major clades of "typical" tits can be recognized: the dark-capped chickadees and their relatives ( Poecile including Sittiparus ), the long-crested Baeolophus and Lophophanes species,

284-471: A branch or twig and leaves from all angles while hanging upside down to feed. In areas where numerous species of tit coexist, different species forage in different parts of the tree, their niche determined in no small way by their morphology; larger species forage on the ground, medium-sized species foraging on larger branches, and the smallest species on the ends of branches. Having obtained larger prey items or seeds, tits engage in hold-hammering, where they hold

355-399: 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 spur . In a behaviour called "spur marking", they grasp the substrate, usually a small sapling, and drag the spur over it, cutting into

426-560: A disjunct distribution, with one species occurring in Europe and the other in central Asia. The genus Baeolophus is endemic to North America. The genus Parus includes the great tit that ranges from Western Europe to Indonesia. Cyanistes has a European and Asian distribution (also into northern Africa), and the three remaining genera, Pseudopodoces , Sylviparus , and Melanochlora , are all restricted to Asia. Tits are active, noisy, and social birds. They are territorial during

497-616: A facultative purpose for this species, in which the more fragrance-rich sites there are, the greater the number of habitable territories. 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 mate choice . Similar behaviour is also observed in the Eulaema meriana 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

568-412: A nasty neighbour strategy. Group-living male breeders are nearly five times more aggressive towards their neighbours than 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

639-428: A perspective. Several other type of resource may be defended including partners, potential mates, offspring, nests or lairs, display areas or leks . 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 conspecifics , as shown in

710-402: A rival's territory. Quiet contact calls are made while feeding to facilitate cohesion with others in their social group. Other calls are used for signalling alarm—a well-known example being the "chic-a-dee-dee" of North American species in the genus Poecile , the call which gives them their local common name, the chickadee. The call also serves as a rallying call to summon others to mob and harass

781-440: A single pair cannot find enough food to rear even one nestling and in drought years breeding is likely to be futile. Many African tit species, along with Pseudopodoces , are cooperative breeders , and even pair-breeding parids are often highly social and maintain stable flocks throughout the nonbreeding season. Tits also have a variety of methods for attracting mates, primarily through their intricate, bouncing mating dance. Only

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852-588: A strategy termed the dear enemy effect 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 nasty neighbour effect 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

923-450: Is a lekking reptile. Males start to establish small display territories two months ahead of the mating season. 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 communicate to other animals that

994-421: Is a behaviour used by animals to identify their territory. Most commonly, this is accomplished by depositing strong-smelling substances contained in the urine , faeces , or, from specialised scent glands located on various areas of the body. Often, the scent contains pheromones or carrier proteins such as the major urinary proteins to stabilize the odours and maintain them for longer. The animal sniffing

1065-523: 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

1136-484: 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 does not happen, the territory may be defended by actual fighting, although this is generally a last resort. Scent marking, also known as territorial marking or spraying when this involves urination ,

1207-636: 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 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 coyotes and golden jackals . Males and female ring-tailed lemurs ( Lemur catta ) scent-mark both vertical and horizontal surfaces at

1278-516: Is from 9 to 21 cm (3.5 to 8.3 in). In weight, the family ranges from 5 to 49 g (0.18 to 1.73 oz); this contracts to 7 to 29 g (0.25 to 1.02 oz) when the three atypical genera are removed. The majority of the variation within the family is in plumage , and particularly colour. The bills of the tits are generally short, varying between stout and fine, depending on diet. The more insectivorous species have finer bills, whereas those that consume more seeds have stouter bills. It

1349-457: 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 larger and larger. Such a dung heap can become up to five metres wide and one metre high. After defecating, greater one-horned rhinos scratch their hind feet in the dung. By continuing to walk, they "transport" their own smell around

1420-554: 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 during which they not only threaten or attack many kinds of intruders, but have stereotyped display behaviour to deter conspecifics sharing neighbouring nesting spots. The owl limpet ( Lottia gigantea )

1491-492: Is not always a fixed behavioural characteristic of a species. For example, red foxes ( Vulpes vulpes ) either establish stable home ranges within particular areas or are itinerant with no fixed abode . Territories may vary with time (season). For example, European robins defend territories as pairs during the breeding season but as individuals during the winter. Resource availability may cause changes in territoriality. For example, some nectarivores defend territories only during

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1562-412: 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 home range . 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

1633-465: Is recorded from the 14th century, composed of the Old English name for the bird, mase ( Proto-Germanic *maison , Dutch mees , German Meise ), and tit, denoting something small. The former spelling, "titmose", was influenced by mouse in the 16th century. Emigrants to New Zealand presumably identified some of the superficially similar birds of the genus Petroica of the family Petroicidae,

1704-399: Is reversed. Animals may use several strategies to defend their territories. The first game theory model of fighting is known as the hawk-dove game . 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

1775-598: Is said that tits are evolving longer beaks to reach into bird feeders. The most aberrant bill of the family is possessed by Hume's ground tit of Tibet and the Himalayas, which is long and decurved. The tits are a widespread family of birds, occurring over most of Europe, Asia, North America, and Africa. The genus Poecile occurs from Europe through Asia into North America, as far south as southern Mexico . American species in this genus are known as chickadees. Some species in this genus have quite large natural distributions; one,

1846-477: The Australian painted lady butterfly and the speckled wood butterfly , the male defends territories that receptive females are likely to fly through such as sunny hilltops and sunspots on a forest's floor. Territory defence in male variegated pupfish ( Cyprinodon variegatus ) is dependent on the presence of females. Reduced aggression consistent with the dear enemy effect occurs between conspecific neighbours in

1917-808: The Ngorongoro Crater to over 100,000 hectares (250,000 acres) in the Kalahari . In birds, golden eagles ( Aquila chrysaetos ) have territories of 9,000 hectares (22,000 acres), least flycatchers ' ( Empidonax minimus ) territories are about 600 square metres (6,500 sq ft) and gulls have territories of only a few square centimetres in the immediate vicinity of the nest. Territories can be linear. Sanderlings ( Calidris alba ) forage on beaches and sandflats. When on beaches, they feed either in flocks or individual territories of 10 to 120 metres of shoreline. The time to develop territories varies between animals. The marine iguana ( Amblyrhynchus cristatus )

1988-552: The Uganda kob (a grazing antelope ) and the marine iguana, males defend the lek site which is used only for mating. Many species demonstrate 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 to deceive females of the species into entering into polygynous relationships. This hypothesis, named

2059-532: The grey-headed chickadee , is distributed from Scandinavia to Alaska and Canada. The majority of the tits in the genus Periparus are found in the southeastern portion of Asia. This includes two species endemic to the Philippines . The coal tit , also in this genus, is a much more widespread species, ranging from the British Isles and North Africa to Japan. The two crested tits of the genus Lophophanes have

2130-461: The penduline tits and long-tailed tits , but while the former are quite close to the tits and could conceivably be included in that family together with the stenostirid "warblers" , the long-tailed tits are not. Indeed, the yellow-browed tit and the sultan tit are possibly more distant to the tits than the penduline tits are. If the two current families are lumped into the Paridae, the tits would be

2201-458: The rufous-vented tit of the Himalayas to as many as 10 to 14 in the blue tit of Europe. In favourable conditions, this species had laid as many as 19 eggs, which is the largest clutch of any altricial bird. Most tits are multibrooded, a necessary strategy to cope with either the harsh winters in which they reside in the Holarctic or the extremely erratic conditions of tropical Africa, where typically

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2272-553: The Australian robins, as members of the tit family, giving them the title tomtit , although, in fact, they are not related. These birds are mainly small, stocky, woodland species with short, stout bills . Some have crests. They range in length from 10 to 22 cm (3.9 to 8.7 in). They are adaptable birds, with a mixed diet including seeds and insects. Many species live around human habitation and come readily to bird feeders for nuts or seed, and learn to take other foods. With

2343-501: The absence of females, but the presence of a female in a male's territory instigates comparably greater aggression between the neighbours. In the Skylark ( Alauda arvensis ), 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

2414-432: 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. Wolves 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

2485-490: The available data as distinct from Parus . Today, this arrangement is considered paraphyletic as indicated by mtDNA cytochrome b sequence analysis, and Parus is best restricted to the Parus major — Parus fasciiventer clade , and even the latter species' closest relatives might be considered a distinct genus. In the Sibley-Ahlquist taxonomy , the family Paridae is much enlarged to include related groups such as

2556-661: The available molecular data. Tits have settled North America twice, probably at some time during the Early-Mid Pliocene . The first were the ancestors of Baeolophus , with chickadees arriving somewhat later. Remizidae Cephalopyrus flammiceps Sylviparus modestus Melanochlora sultanea Pardaliparus Periparus Baeolophus Lophophanes Sittiparus Poecile Cyanistes Pseudopodoces Parus monticolus Parus major Machlolophus Melaniparus Family: PARIDAE Chickadees Their name reputedly comes from

2627-582: The bird (i.e. badge) is highly visible when it sings (vocal marking) at the boundary of its territory. The ring-tailed lemur ( Lemur catta ) 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. Rhinoceros have poor vision but may use visual marking. Dominant white rhino bulls mark their territory with faeces and urine (olfactory marking). The dung

2698-553: The birds are found from the East Coast to the West Coast, and from Canada to Mexico. Chickadees' preferred habitats include mixed deciduous or coniferous forests, parks, open woods, cottonwood groves, willow thickets, and disturbed areas. Mountain chickadees are food-caching birds. A single bird can hide as many as 80,000 individual seeds, which they retrieve during the winter. Their ability to do so depends on their spatial memory of

2769-495: The blue tit is typically polygynous; all other species are generally monogamous. Courtship feeding is typical of pair-breeding tits to deal with the cost of rearing their large broods. Recently, the large Parus group has been gradually split into several genera (as indicated below), initially by North American ornithological authorities and later elsewhere. Whereas in the mid-1990s, only Pseudopodoces , Baeolophus , Melanochlora , and Sylviparus were considered well-supported by

2840-434: 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 . Territorial scent marking may involve behaviours specific to this activity. When a wolf marks its territory , it lifts a hind leg and urinates on a scent post (usually an elevated position like a tree, rock, or bush). This raised leg urination

2911-435: The breeding season and often join mixed-species feeding flocks during the nonbreeding season. The tits are highly adaptable, and after the corvids (crows and jays) and parrots , amongst the most intelligent of all birds. Tits recognize the difference between species that are dangerous or harmless to them, by this they can protect each other or their families. These birds do this by mobbing or escaping, however they also avoid

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2982-488: The case of redlip blenny . This is because the conspecifics share exactly the same set of resources. Several types of resource in a territory may be defended. Food: Large solitary (or paired) carnivores, such as bears and the bigger raptors 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 grizzly bears are attracted to migrating salmon . Food related territoriality

3053-682: The city of Calgary , Alberta. The chickadee is also the state bird of Maine , but a species has never been specified. A proposed bill in 2019 would have named the black-capped chickadee as the official species for Maine, but was unanimously voted down in committee. The de facto species for Maine remains the black-capped. One holarctic species is referred to by a different name in each part of its range: grey-headed chickadee in North America and Siberian tit in Eurasia. Chickadees are native to North America, where they are very common. In North America,

3124-571: 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. Some species of bees also exhibit territoriality to defend mating sites. For example, in Euglossa imperialis , 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

3195-410: The exception of the three monotypic genera Sylviparus , Melanochlora , and Pseudopodoces , the tits are extremely similar in appearance, and have been described as "one of the most conservative avian families in terms of general morphology". The typical body length of adult members of the family is between 10 and 16 cm (3.9 and 6.3 in) in length; when the monotypic genera are added, this range

3266-532: The fact that their calls make a distinctive "chick-a-dee-dee-dee", though their normal call is actually "fee-bee," and the "chick-a-dee-dee-dee" call is an alarm call. The number of "dees" depends on the predator. The chickadee (specifically the black-capped chickadee Poecile atricapillus , formerly Parus atricapillus ) is the official bird for the US state of Massachusetts , the Canadian province of New Brunswick , and

3337-430: The genus Acmaea, predatory snails, and sessile organisms such as anemones and barnacles. Nests and offspring: 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 superimposition of

3408-821: The geography. The size is usually a compromise of resource needs, defense costs, predation pressure and reproductive needs. Some species of squirrels may claim as much as 10 hectares (25 acres) of territory. For European badgers , 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, food waste or artificial feeding in suburban gardens. Spotted hyenas ( Crocuta crocuta ) have highly variable territory sizes, ranging from less than 4,000 hectares (9,900 acres) in

3479-489: The greatest threat to the resident territory-holder. In territory defence by groups of animals, reciprocal altruism can operate whereby the cost to the benefactor in helping defend the territory is less than the gains to the beneficiary. 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

3550-579: The ground nearby, thereby leaving a visual advertisement of the territory. This includes domestic dogs . 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 Canada lynx ( Lynx canadensis ) and the American black bear ( Ursus americanus ). Many animals have scent glands in their paws or deposit fur during tree-marking, so tree-marking may be

3621-479: The item between their feet and hammer it with their bill until it opens. In this fashion, they can even open hazelnuts in around 20 minutes. A number of genera engage in food caching, hoarding supplies of food during the winter. Tits are cavity-nesting birds, typically using trees, although Pseudopodoces builds a nest on the ground. Most tree-nesting tits excavate their nests, and clutch sizes are generally large for altricial birds, ranging from usually two eggs in

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3692-759: The locations. Birds that live in harsher conditions, where their ability to remember the location of food is more important, have been found to have better memory abilities, a larger hippocampus , and more neurons than chickadees that live in milder climates where food sources are easier to find without relying on memory. Territory (animal) In ethology , territory is the sociographical area that an animal consistently defends against conspecific competition (or, occasionally, against animals of other species ) using agonistic behaviors or (less commonly) real physical aggression . Animals that actively defend territories in this way are referred to as being territorial or displaying territorialism . Territoriality

3763-426: 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 polygynous , one male territory probably contains several female territories, while in some polyandrous species such as the northern jacana , this situation

3834-566: The most combative spiders have the largest territories. Some species of penguin defend their nests from intruders trying to steal the pebbles from which the nest is constructed. Mating opportunities: The striped mouse ( Rhabdomys pumilio ) is group living with one single breeding male and up to 4 communally breeding females per group. Groups typically contain several philopatric 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

3905-418: The nest site entrance to establish their territory. Wombats use feces to mark their territory. They have evolved specialized intestinal anatomy to produce cubical feces to ensure the feces do not roll away. 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

3976-523: The nest when the predators are present in order to avoid their families to be seen. Fission–fusion society has been documented in a number of avian taxa including this one. In brief, that means flocks can split into smaller groups or individuals, and subsequently reunite. The tits make a variety of calls and songs . They are amongst the most vocal of all birds, calling continuously in most situations, so much so that they are only ever silent for specific reasons such as avoiding predators or when intruding on

4047-407: The organism requires to survive, because defending a 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. Behavioural ecologists have argued that food distribution determines whether a species is territorial or not, however, this may be too narrow

4118-715: The other. They also did not engage in physical contact with the territorial males. On the other hand, 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. Although most territories contain multiple (potential) resources, some territories are defended for only one purpose. European blackbirds may defend feeding territories that are distant from their nest sites, and in some species that form leks, for example in

4189-476: The overlaps in their home ranges using their anogenital scent glands . To do this, they perform a handstand to mark vertical surfaces, grasping the highest point with their feet while applying the scent. In the Eastern carpenter bee, Xylocopa virginica , 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

4260-407: 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 combined with the smell of the marking animal. The territorial male scrape-marks every 30 m (98 ft) or so around its territory boundary. After leaving a urination mark, some animals scrape or dig

4331-422: The predator. The number of "dee" syllables at the end of the call increases with the level of danger the predator poses. The tits are generalist insectivores that consume a wide range of small insects and other invertebrates, particularly small defoliating caterpillars. They also consume seeds and nuts, particularly in the winter. One characteristic method of foraging in the family is hanging, where they inspect

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4402-439: The reason for such protective behaviour is to acquire and protect food sources, nesting sites, mating areas, or to attract a mate. Among birds, territories have been classified as six types. 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

4473-724: The resident is present. Other animals may use more long-term visual signals such as faecal deposits, or marks on the vegetation or ground. Visual marking of territory is often combined with other modes of animal communication. Some animals have prominent "badges" or visual displays to advertise their territory, often in combination with scent marking or auditory signals. Male European robins 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 areas. The red breast of

4544-418: The risk of injury. This is ritualized aggression . 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. Territories may be held by an individual, a mated or unmated pair, or a group. Territoriality

4615-408: The rival is a hawk). Another strategy used in territory defence is the war of attrition . 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. Some animals use

4686-514: The same note, thus creating the 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. 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

4757-460: The scent frequently displays a flehmen response to assist in detecting the mark. Scent marking is often performed by scent rubbing in many mammals. In many mammal species, scent marking is more frequent during the breeding season . Bears and felids such as leopards and jaguars scent-mark by urinating on or rubbing against vegetation. Prosimians and New World monkeys also use scent marking, including urine washing ( self-anointing

4828-511: 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 Scutellastra . They vigorously defend their gardens of particular species of algae, that extend for perhaps 1–2 cm around the periphery of their shells. The desert grass spider, Agelenopsis aperta , often engages in fights over its territory and

4899-449: The territory is occupied and may also communicate additional information such as the sex, reproductive status or dominance status of the territory-holder. Sign-posts may communicate information by olfactory , 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 ritualized aggression toward each other. This

4970-400: The usually tufted, white-cheeked Periparus (including Pardaliparus ) with more subdued coloration and finally Parus sensu stricto (including Melaniparus and Machlolophus ). Still, the interrelationship of these, as well as the relationships of many species within the clades, are not well-resolved at all; analysis of morphology and biogeography probably gives a more robust picture than

5041-474: 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. European wildcats ( Felis silvestris ) deposit their faecal marks on plants with high visual conspicuousness that enhances the visual effectiveness of the signal. Many animals use vocalisations to advertise their territory. These are short-term signals transmitted only when

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