Type A territory, or for a migratory species, may be on the wintering grounds. 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 resource needs, defense costs, predation pressure and reproductive needs.
30 hectares in a good rural habitat, but as large as 300 hectares in a poor habitat. On average, a territory may be approximately 50 hectares, with main setts normally at least 500 metres apart. In urban areas, territories can be as small as 5 hectares, if they can obtain enough food from bird tables, food waste or artificial feeding in suburban gardens. 4,000 hectares in the Ngorongoro Crater to over 100,000 hectares in the Kalahari. 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.
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. 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 «ritualised aggression» toward each other.
This is a series of stylised postures, vocalisations, displays, etc. If this does not happen, the territory may be defended by actual fighting, although this is generally a last resort. Territorial scent marking may involve behaviours specific to this activity. 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.
To do this, they perform a handstand to mark vertical surfaces, grasping the highest point with their feet while applying the scent. Males, although they have the gland, are unable to produce the marking substance. Female secrete it near the nest site entrance to establish their territory. Ring-tailed lemurs hold their distinctive tails high in the air during territorial scent marking. They also engage in «stink fights» with intruding males. 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, 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. They attack other males that stray into their territories, and have been observed attacking other small birds without apparent provocation. 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 larger and larger.
Such a dung heap can become up to five metres wide and one metre high. 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. After leaving a urination mark, some animals scrape or dig the ground nearby, thereby leaving a visual advertisement of the territory. 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. 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. 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. 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. When howling together, wolves harmonize rather than chorus on 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.