Behavioral Ecology Population Dynamics and Community Structure

The ways in which organisms make decisions about habitats, food, and associates have many important implications for the structure and functioning of ecological systems. We will describe two of those implications here. First, animals with complex social organization often achieve remarkably high abundances. Second, the ways in which animals select habitats and food, combined with their interactions with individuals of other species, may influence the range of habitat and foods a species uses in nature.

Social animals may achieve great abundances

The abundances achieved by some social animals are impressive. For example, up to 94 percent of the individuals and 86 percent of the biomass of arthropods in the canopies (tree-

Large Australian Termites
53.13 Termite Mounds Are Large and Complex These immense Australian termite mounds are constructed over many years by millions of worker termites. Elaborate nests or burrows, which are very costly to construct and maintain, characterize nearly all eusocial animals.

tops) of tropical rainforests are social ants. Termites (also social insects) are the primary consumers of plant tissues in the savannas of Africa. They live in and build large mounds, within which many other species of animals live. Termites may extract nutrients from the soil at depths as great as 80 meters. In parts of Australia, termite density may reach 1,000 colonies per hectare (Figure 53.13).

Ants and termites have achieved these remarkable abundances in part because their social organization allows them to exploit the services of other organisms in harvesting vital resources. The most abundant and highly productive ants and termites actively cultivate fungi that break down difficult-to-digest plant tissues, including wood. Some ants tend aphids and other insects that tap phloem fluids, protecting the phloem-suckers from predators. Because phloem is rich in carbohydrates but poor in proteins, phloem-suckers ingest more carbohydrates than they can use. They eject the excess in the form of sugar-rich anal drops (see Figure 36.13), which the ants eat. Because the ants can easily obtain enough carbohydrates in this manner, they need to get only proteins in other ways, such as by eating other insects. Moreover, with their high, sugar-based metabolic rates, the ants can expend the energy needed to drive other predatory insects away from their food sources. In this way, ants strongly influence the community of insects in tropical forest canopies.

Social living also enables organisms to find and use temporally and spatially patchy foods. The wildebeest, which travels in large herds, is the most abundant large mammal in Africa (see Figure 53.2). More than a million individuals are found in the herd that migrates between the Masai Mara in Kenya and the Serengeti in Tanzania to feed on the rapidly growing grass that follows seasonal rains in each area.

Even more striking is the abundance achieved by our own species (Figure 53.14). Social living enabled members of human groups to specialize in different activities. Among the benefits of specialization were domestication of plants and animals and cultivation of land. These innovations enabled our ancestors to increase the resources at their disposal dramatically. Those increases, in turn, stimulated rapid population growth up to the limit determined by the agricultural productivity that was possible with human- and animal-powered tools. Agricultural machines and artificial fertilizers, made possible by the tapping of fossil fuels, greatly increased agricultural productivity and removed that earlier limit. In addition, the development of modern medicine reduced the mortality rate in human populations. Medicine and better hygiene have also allowed people to live in large numbers in areas where diseases formerly kept numbers very low. However, these successes have been accompanied by many problems, some of which we will discuss in subsequent chapters.

Interspecific interactions influence animal distributions

As we have seen, animals assess habitat quality and settle preferentially in better places. They also select the food items that give them the best return for the time and energy they expend in getting them. The optimality modeling approach used to develop and test hypotheses about how such choices are made has yielded an important general "rule of thumb" of behavioral ecology: As much as possible, organisms concentrate on doing what they do best and avoid doing what they do poorly.

However, interspecific interactions may prevent animals from living in those environments in which they would do best. Individuals of a behaviorally dominant species may be able to exclude individuals of a subordinate species from its preferred foraging areas. How such behavioral dominance influences use of foraging areas can be illustrated by observing hummingbird behavior.

Hummingbirds extract nectar from flowers and often defend patches of flowers from other hummingbirds. In an ex

Densities Cities

53.14 Social Organization Allows Humans to Live at High Densities Human cities such as Benidorm on Spain's Costa Blanca are examples of how social organization allows our species to achieve and sustain extreme population densities.

periment done in southeastern Arizona, investigators created artificial "flower patches" by setting up an array of feeders. Some feeders contained artificial nectar that was rich in sucrose; others contained a more dilute solution that was a poorer source of sucrose. Hummingbirds quickly learned which were the high-quality feeders because the rich ones had blue bee guards; the poor ones had yellow bee guards.

Males of three hummingbird species visited the feeders. Interactions were strongest between two of them: Male blue-throated hummingbirds, which weigh about 8.3 g on average, behaviorally dominated the smaller male black-chinned hummingbirds, which weighed only about 3.2 g. When no male blue-throats were present, black-chinned males fed almost exclusively at the rich feeders; but when male blue-throats were present, black-chinned males fed at poor feeders as often as they fed at rich feeders. Even though the nectar at the poor feeders was more dilute, the black-chins achieved about the same rate of energy from them as from the rich feeders because they were able to feed longer at the poor feeders without being chased away by the larger blue-throat males.

These kinds of observations show us that the ways in which animals choose what to eat, where to seek food, and with whom to associate influence the sizes and distributions of populations of many species and how they interact in nature. These aspects of populations will form the focus of the next chapter, on population ecology.

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