Costs and Benefits of Social Life

Many think that the reason so many animals live with others of their species is that social creatures are higher up the evolutionary scale and so are better adapted and leave more offspring than do animals that live solitary lives. However, in each and every species, generation after generation, relatively social and relatively solitary types compete unconsciously with one another in ways that determine who leaves more offspring on average. In some species, the more social individuals have won out, but in a large majority, it is the solitary types that have consistently left more surviving descendants on average.

But how can living alone ever be superior to living together? Under some conditions, a cost-benefit comparison favors solitary life over a more social existence. For example, among most social species, animals have to expend time and energy competing for social status. Those that do not occupy the top positions regularly have to signal their submissive state to their superiors if they are to be permitted to remain in the group. This can take up a major share of a social subordinate's life. In fact, even in small social groups there are both subtle competition and not-so-subtle competition.

Social groups also offer opportunities for reproductive interference. Breeding males that live in close association with more attractive rivals may lose their mates to these individuals. In addition, sociality has two other potential disadvantages. The first is heightened competition for food, which occurs in animals as different as colonial fieldfares (a kind of songbird) and groups of lions, whose females are often pushed from their food by hungry males. The second is increased vulnerability to parasites and disease, which plague social species of all sorts. While it is true that some social animals have evolved special responses designed to combat parasites and disease, those responses can only reduce, but cannot totally eliminate, the damage caused by those threats, and the responses may even carry their own costs. Thus, honeybees warm their hives in response to an infestation by a fungal pathogen, which apparently helps kill the heat-sensitive fungus, but at the price of time and energy expended by the heat-producing workers.

If social living carries a heightened risk of infection, then the larger the group, the greater the risk. This prediction holds for cliff swallows, which pack their nests side by side in colonies composed of anywhere from a handful of birds to several thousand pairs. The more swallows nesting together, the greater the chance that at least one bird will be infested with swallow bugs, which can then readily spread from one nest to another.

The parasites and fungi that make life miserable for swallows and other social creatures demonstrate that if sociality is to evolve, the assorted costs of living together must be outweighed by compensatory benefits. Cliff swallows may join others to take advantage of the improved foraging that comes from following companions to good feeding sites, while other animals, such as male emperor penguins, save thermal energy by huddling shoulder to shoulder during the brutal Antarctic winter. Still others, such as lionesses, join forces to fend off enemies of their own species.

The most widespread fitness benefit for social animals, however, probably is improved protection against predators. Many studies have shown that animals in groups gain by reducing the individual risk of being captured, or by spotting danger sooner, or by attacking their enemies in groups. Males in nesting colonies of the bluegill sunfish cooperate in driving egg-eating bullhead catfish away from their nests at the bottom of a freshwater lake. While bluegills have adopted social behavior to avoid predation, closely related species that nest alone have evolved means to protect themselves while nesting alone. Thus, the solitary pumpkinseed sunfish, a member of the same genus as the bluegill, has powerful biting jaws and so can repel egg-eating enemies on its own, whereas bluegills have small, delicate mouths good only for inhaling small, soft-bodied insect larvae. Pumpkinseed sunfish are in no way inferior to or less well adapted than bluegills because they are solitary; they simply gain less through social living, which makes solitary nesting the adaptive tactic for them.