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68-609: Oxyaenidae ("sharp hyenas ") is a family of extinct carnivorous placental mammals. Traditionally classified in order Creodonta , this group is now classified in its own order Oxyaenodonta ("sharp tooth hyenas") within clade Pan-Carnivora in mirorder Ferae . The group contains four subfamilies comprising fourteen genera. Oxyaenids were the first to appear during the late Paleocene in North America , while smaller radiations of oxyaenids in Europe and Asia occurred during

136-438: 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,

204-581: A change in climate, along with the arrival of canids into Eurasia. Of the dog-like hyena lineage, only the insectivorous aardwolf survived, while the bone-crushing hyenas (including the extant spotted, brown and striped hyenas) became the undisputed top scavengers of Eurasia and Africa. The descendants of Plioviverrops reached their peak 15 million years ago, with more than 30 species having been identified. Unlike most modern hyena species, which are specialised bone-crushers, these dog-like hyenas were nimble-bodied, wolfish animals; one species among them

272-399: 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

340-621: 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

408-412: 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

476-428: 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

544-881: A short, broad skull, deep jaws, and teeth designed for crushing rather than shearing, as in the hyaenodonts or modern cats. Oxyaenids were specialized carnivores that preyed on other terrestrial vertebrates, eggs and insects. They were capable of climbing trees, which is suggested by fossil evidence of their paws. Cladogram according to Gunnel in 1991: Tytthaena parrisi Tytthaena lichna Palaeonictis occidentalis Palaeonictis peloria Dipsalodon matthewi Dipsalodon churchillorum Dipsalidictis krausei Dipsalidictis transiens Dipsalidictis platypus Dipsalidictis aequidens Oxyaena gulo Oxyaena intermedia Oxyaena forcipata Hyena Hyenas or hyaenas ( / h aɪ ˈ iː n ə z / hi- EE -nəz ; from Ancient Greek ὕαινα , hýaina ) are feliform carnivoran mammals belonging to

612-588: 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

680-450: 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

748-421: 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

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816-527: 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

884-484: 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 ,

952-636: 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

1020-457: 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

1088-554: 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 )

1156-546: Is likely that its unrivaled ability to digest the terpene excretions from soldier termites is a modification of the strong digestive system its ancestors used to consume fetid carrion. The striped hyena may have evolved from Hyaenictitherium namaquensis of Pliocene Africa . Striped hyena fossils are common in Africa, with records going back as far as the Villafranchian . As fossil striped hyenas are absent from

1224-492: 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

1292-412: 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

1360-399: 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

1428-483: 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

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1496-615: The Eocene . The name of order Oxyaenodonta comes from Ancient Greek ὀξύς (oxús)  'sharp', name of hyena genus Hyaena and from Ancient Greek ὀδούς (odoús)  'tooth'. The name of family Oxyaenidae comes from Ancient Greek ὀξύς (oxús)  'sharp', name of hyena genus Hyaena and taxonomic suffix " -idae ". They were superficially cat-like mammals that walked on flat feet , in contrast to modern cats , which walk and run on their toes . Anatomically, characteristic features include

1564-1193: The Furninha Cave in Portugal and the Genista Caves in Gibraltar . The European form was similar in appearance to modern populations, but was larger, being comparable in size to the brown hyena . The spotted hyena ( Crocuta crocuta ) diverged from the striped and brown hyena 10 million years ago. Its direct ancestor was the Indian Crocuta sivalensis , which lived during the Villafranchian. Ancestral spotted hyenas probably developed social behaviours in response to increased pressure from rivals on carcasses, thus forcing them to operate in teams. Spotted hyenas evolved sharp carnassials behind their crushing premolars, therefore they did not need to wait for their prey to die, and thus became pack hunters as well as scavengers. They began forming increasingly larger territories , necessitated by

1632-809: The Mediterranean region, it is likely that the species is a relatively late invader to Eurasia, having likely spread outside Africa only after the extinction of spotted hyenas in Asia at the end of the Ice Age . The striped hyena occurred for some time in Europe during the Pleistocene, having been particularly widespread in France and Germany . It also occurred in Montmaurin , Hollabrunn in Austria ,

1700-577: The Mid-Pleistocene transition . The four extant species are the striped hyena ( Hyaena hyaena ), the brown hyena ( Parahyaena brunnea ), the spotted hyena ( Crocuta crocuta ), and the aardwolf ( Proteles cristata ). The aardwolf can trace its lineage directly back to Plioviverrops 15 million years ago, and is the only survivor of the dog-like hyena lineage. Its success is partly attributed to its insectivorous diet, for which it faced no competition from canids crossing from North America. It

1768-821: 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 )

1836-561: 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

1904-893: The family Hyaenidae ( / h aɪ ˈ ɛ n ɪ d iː / ). With just four extant species (each in its own genus ), it is the fifth-smallest family in the order Carnivora and one of the smallest in the class Mammalia . Despite their low diversity, hyenas are unique and vital components of most African ecosystems. Although phylogenetically closer to felines and viverrids , hyenas are behaviourally and morphologically similar to canids in several elements due to convergent evolution : both hyenas and canines are non- arboreal , cursorial hunters that catch prey with their teeth rather than claws. Both eat food quickly and may store it, and their calloused feet with large, blunt, nonretractable claws are adapted for running and making sharp turns. However, hyenas' grooming, scent marking , defecation habits, mating and parental behavior are consistent with

1972-459: The middle ear and dentition. The lineage of Plioviverrops prospered, and gave rise to descendants with longer legs and more pointed jaws, a direction similar to that taken by canids in North America . Hyenas then diversified into two distinct types: lightly built dog-like hyenas and robust bone-crushing hyenas. Although the dog-like hyenas thrived 15 million years ago (with one taxon having colonised North America), they became extinct after

2040-609: The World for extant genera. The percrocutids are, in contrast to McKenna and Bell's classification, not included as a subfamily into the Hyaenidae, but as the separate family Percrocutidae (though they are generally grouped as sister-taxa to hyenas ). Furthermore, the living brown hyena and its closest extinct relatives are not included in the genus Pachycrocuta , but in the genus Parahyaena . However, some research has suggested Parahyaena may be synonymous with Pachycrocuta , making

2108-506: 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

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2176-421: The ancestral bone-crushing hyenas coincided with the decline of the similarly built family Percrocutidae . The bone-crushing hyenas survived the changes in climate and the arrival of canids, which wiped out the dog-like hyenas, though they never crossed into North America, as their niche there had already been taken by the dog subfamily Borophaginae . By 5 million years ago, the bone-crushing hyenas had become

2244-432: 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

2312-438: The behavior of other feliforms. Hyenas feature prominently in the folklore and mythology of human cultures that live alongside them. Hyenas are commonly viewed as frightening and worthy of contempt. In some cultures, hyenas are thought to influence people's spirits, rob graves, and steal livestock and children. Other cultures associate them with witchcraft, using their body parts in traditional medicine . Hyenas originated in

2380-582: 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

2448-434: 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

2516-419: The broader context of late-Quaternary extinctions , as the late Pleistocene and early Holocene saw the disappearance of many primarily large mammals from Europe and the world. Expansion or duplication of the olfatory receptor gene family has been found in all 4 extant species, which would have led to the evolution of the more specialised feeding habits of hyenas. Expansion in immune-related gene families

2584-1301: The brown hyena the only extant member of this genus. The following cladogram illustrates the phylogenetic relationships between extant and extinct hyaenids based on the morphological analysis by Werdelin & Solounias (1991), as updated by Turner et al. (2008). Protictitherium crassum "Protictitherium" cingulatum "Protictitherium" intermedium "Protictitherium" llopisi "Protictitherium" punicum " Protictitherium" gaillardi "Protictitherium" sumegense "Protictitherium" csakvarense Plioviverrops gervaisi Plioviverrops orbignyi Plioviverrops guerini Plioviverrops faventinus Plioviverrops gaudryi Tungurictis spocki Thalassictis robusta "Thalassictis" certa "Thalassictis" montadai "Thalassictis" proava "Thalassictis" sarmatica "Thalassictis" spelaea Tongxinictis primordialis Proteles cristatus (aardwolf) [REDACTED] Proteles amplidentus Ictitherium viverrinum Ictitherium ebu Ictitherium tauricum Ictitherium ibericum Ictitherium kurteni Ictitherium intuberculatum Ictitherium pannonicum Miohyaenotherium bessarabicum Hyaenotherium wongii Hyaenictitherium hyaenoides "Hyaenictitherium" pilgrimi Scent marking Territoriality

2652-488: 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

2720-418: The dominant scavengers of Eurasia, primarily feeding on large herbivore carcasses felled by sabre-toothed cats . One genus, Pachycrocuta , was a 110 kg (240 lb) mega-scavenger that could splinter the bones of elephants . Starting in the early Middle Pleistocene Pachycrocuta was replaced by the smaller Crocuta and Hyena , which corresponds to a general faunal change, perhaps in connection to

2788-410: The end of the last glacial period and a subsequent displacement of open grassland by closed forests, which favoured wolves and humans instead. However, analyses have shown that climate change alone is insufficient to explain the spotted hyena's disappearance from Europe, suggesting that other factors – such as human pressure – must have played a role. This suggests that the events must be seen within

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2856-576: 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

2924-482: The evolution of the feeding of termites Trinervitermes in this species. Mutations and variants in genes related to craniofacial shape were also found ( GARS , GMPR , STIP1 , SMO and PAPSS2 ). Another gene is related to protective epidermis function ( DSC1 ). The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997) and Wozencraft (2005) in Wilson and Reeders Mammal Species of

2992-480: The fact that their prey was often migratory, and long chases in a small territory would have caused them to encroach into another clan's turf. Spotted hyenas spread from their original homeland during the Middle Pleistocene , and quickly colonised a very wide area from Europe, to southern Africa and China . The eventual disappearance of the spotted hyena from Europe has traditionally been attributed to

3060-430: 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

3128-821: 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

3196-489: 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

3264-588: 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

3332-402: The jungles of Miocene Eurasia 22 million years ago, when most early feliform species were still largely arboreal . The first ancestral hyenas were likely similar to the modern African civet ; one of the earliest hyena species described, Plioviverrops , was a lithe, civet-like animal that inhabited Eurasia 20–22 million years ago, and is identifiable as a hyaenid by the structure of

3400-550: The land bridge into North America, being the only hyena to do so. Chasmaporthetes managed to survive for some time in North America by deviating from the endurance-running and bone-crushing niches monopolized by canids, and developing into a cheetah -like sprinter. Most of the dog-like hyenas had died off by 1.5 million years ago. By 10–14 million years ago, the hyena family had split into two distinct groups: dog-like hyenas and bone-crushing hyenas. The arrival of

3468-426: 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

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3536-566: 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

3604-418: 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

3672-407: 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

3740-715: 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

3808-481: 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

3876-407: 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

3944-439: 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

4012-724: 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

4080-418: 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

4148-408: 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

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4216-514: 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

4284-460: 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

4352-511: 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

4420-449: 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

4488-474: 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

4556-521: Was Ictitherium viverrinum , which was similar to a jackal . The dog-like hyenas were numerous; in some Miocene fossil sites, the remains of Ictitherium and other dog-like hyenas outnumber those of all other carnivores combined. The decline of the dog-like hyenas began 5–7 million years ago during a period of climate change, exacerbated by canids crossing the Bering land bridge to Eurasia. One species, Chasmaporthetes ossifragus , managed to cross

4624-584: Was also found in the spotted hyena, striped hyena and brown hyena, which would have led to the evolution of the scavenging in these species. Mutations and variants were also found in digestion-related genes ( ASH1L , PTPN5 , PKP3 , AQP10 ). One of these digestion-related genes has variants also related to enhanced bone mineralisation ( PTPN5 ), while other have also a role in inflammatory skin responses ( PKP3 ). In aardwolves, expansion of genes related to toxin response were found ( Lipocalin and UDP Glucuronosyltransferase gene families), which would have led to

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