Natural history

Migration

Many artiodactyls undertake seasonal migrations between their breeding grounds and feeding areas or between different feeding areas. They can then take advantage of the seasonal changes in different areas. This means that larger populations, and hence a larger biomass (i.e., the total weight of all individuals in an area), can be supported than if all passed their lives in one area. The North American mule deer (Odocoileus hemionus) comes from its summer pastures at high altitudes as the first snow falls and returns at the end of winter, several weeks after the snow has melted.

Social behaviour

Although the popular image of artiodactyls is one of great herds numbering thousands of individuals, some species are solitary, and many others form only small family groups. The maternal family unit, in fact, is the most cohesive one, providing the basis for herd formation. Most artiodactyls are more or less social, and grazing forms may be found in especially large aggregations. It appears that the practice of aggregating gives protection, favouring those members of the species that are the most active contributors to the gene pool (thus the most available to natural selection), since the individuals most frequently taken by predators are old, solitary males, males maintaining territories, and animals of either sex separated from the herd.

Social facilitation (the instigation of collective behaviour) takes place in herds. After one animal flees, all of the others flee, and the predator may thus not catch any. Social facilitation may also promote a restricted season for births; this helps survival of the young by denying these easy-prey individuals to predators through much of the year, and keeps the predator population lower than if young were available throughout the year. Another advantage of herding is that the older generation in a herd can guide migrations to water, feeding areas, or mating grounds.

Females and young are usually in herds separate from those of the younger males, but territorial (the older, proven) males may accompany the females. There are some variations of this behaviour. In the Eurasian roe deer (Capreolus capreolus), for example, the basic unit includes the doe, her litter of two, and often the young of the previous year. During the rutting (mating) season males associate with females in heat but do not gather harems. The female herds of red deer (Cervus elephas) are separate from the males except in the breeding season, when the stag will defend his female herd against other males. Among cattle and related species, the males associate with the females and young, but the bulls are ranked below a so-called master bull, each defending its place within the rank order. Female hippopotamuses and their young form a group in water and have a favourite resting and basking sandbank. The males have their resting places around this area. Each male’s rank in the social hierarchy determines how close to the females he may be.

There can be some flexibility of social organization within a species. During the rutting season the male Rocky Mountain goat (Oreamnos americanus) makes little effort to herd females within a fixed area if there is little snow, but he does drive off other males. When there is much snow, he neither fights other males nor defends individual females.

Forest-dwelling artiodactyls often live singly, as does the okapi (Okapia johnstoni) of central Africa; individuals meet only for mating. Female moose (Alces alces) with calves are intolerant of their own young of the previous year and of adults, so even small herds do not form.

The territory of an animal is an area from which the possessor attempts to exclude other individuals of the same species (and occasionally other species). An animal in an area lacking its own scent is more timid and ready to flee. Among solitary artiodactyls the territory holder defends an area sufficient to meet his needs for food and shelter. Among social artiodactyls the territorial system is interwoven with breeding activities, and territories are normally defended only by certain males. Other males are driven off, and a percentage of males are prevented from mating.

The most simple territorial organization among artiodactyls is that of the common wild pig (Sus scrofa), which lives within a home range including resting, feeding, drinking, and wallowing places. There is little sign of territorial defense, and the herd (called the sounder) may move to a new area. At the other extreme, male Uganda kob antelopes (Kobus kob) hold territories, for breeding only, that are as small as 15 to 30 metres (50 to 100 feet) in diameter. There are 30 to 40 territories on the breeding ground of a herd, and groups of females and young move about the territories despite the efforts of individual males to detain them. The semi-arid Serengeti plains of northern Tanzania contain nomadic aggregations of blue wildebeest (Connochaetes taurinus), males of which defend temporary territories only while an aggregation remains stationary.

In territorial defense an aggressive encounter between males is generally preceded by visual signalling of intentions. Chital deer (Cervus axis), for example, have several sorts of threatening displays. When sharp, potentially lethal horns appeared in early ruminants, intimidating displays rather than combats would doubtless have been favoured. Horns or antlers eventually functioned to maintain head contact during struggles rather than to bruise, slash, or gore. This stylized fighting, in which the competing males interlock horns or antlers and try to “outwrestle” each other, minimizes the danger of killing an opponent of the same species (conspecific). It evolved in two ways: further development of the wrestling, found in stags and some of the antelopes, and ramming, as in sheep. In sheep the horns are the sole organs of display. They increase in size throughout life and parallel the dominance order of the males, so that unnecessary fighting is minimized. Ramming may have intermediate forms; goats, for example, butt with a sideways hooking motion. In the fighting of hornless artiodactyls, such as pigs, the combatants may be badly mauled or even killed. The fighting behaviour of camels retains primitive elements of biting, kicking, and neck wrestling.