Two
kinds of organisms that are each using a resource that is in short supply cm
with one another. Interspecific competition is often greatest between organisms
obtain their food in similar ways; thus green plants compete mainly with other
green plants, herbivores, with other herbivores, and carnivores with carnivores.
In addi competition is more acute between closely similar kinds of organisms
than bet1 ones that are less similar. Interspecific competition is to be distinguished
from i petition between individuals of a single species, which is called intrasperific
coi tition. More than 50 years ago, the Soviet ecologist G.F. Gause formulated
what is called the principle of competitive exclusion. This principle states
that if two species are competing with one another for the same limited resource,
then one of the species will be able to use that resource more efficiently than
the other, and the former will therefore eventually eliminate the latter locally.
In many experiments, the results of growing individuals of two species together
in the laboratory have not always been easily predictable. For example, when
Thomas Park and his colleagues at the University of Chicago grew two species
of flour beetle, Tribolium, together in the same container of flour, one always
became extinct. The species that survived, however, varied. Tribolium castaneum
usually won under relatively hot and damp conditions, whereas Tribolium confusum
won under cooler, drier conditions. Subsequent experiments with these species
demonstrated that a genetic component was also involved in the unpredictability
of the outcome. Some strains of one species would win over some-but not all-strains
of the other, under a given set of conditions.
Similarly, John Harper and his
colleagues at the University College of North Wales grew two species of clover
together, Trifolium repens (white clover) and Trifo-lium fragiferum (strawberry
clover). Each was sown at two densities, 36 and 64 plants per square foot (930
square centimeters), using all the possible combinations. Al-ihough white clover
initially formed a dense canopy of leaves, the slower-growing strawberry clover,
whose leaf stalks are longer, eventually produced enough leaves lhat overtopped
the lower while clover and overcame it. Strawberry clover did this by mmpeting
more effectively for light; the outcome was the same regardless of the ini-lial
densities at which the plants were sown.
Among plants generally, competition between root systems for the nutrients that
ihe plants require and which they obtain from soil is of central importance.
The roots of one species not only may deplete essential minerals and thus outcompete
another, but may also excrete toxic substances that depress the growth of the
other species. Such interactions are poorly understood, but experimental studies
are beginning to resolve their complexity. Competition: Examples from Nature
Examples that demonstrate the principle of competitive exclusion can be found
in nature and studied, both by observation and by experiment. Such examples
are directly related to the structure of the communities in which they occur,
and they will be discussed more extensively. However, we include a few examples
here, as we consider the contribution of competitive interactions to the structure
of natural communities.
The competitive interactions between two species of barnacles that grew together
on the same rocks along the coast of Scotland have been investigated by J.H.
Connell of the University of California, Santa Barbara- Barnacles are marine
animals (crustaceans) that have free-swimming larvae that settle down, cement
themselves to rocks, and then remain permanently attached to that point. Of
the two species Connell studied, Chthamalus stellatus lives in shallower water,
where it is often exposed to air by tidal action, and Balanus balanoides occurs
lower down, where it is rarely exposed to the atmosphere. In this deeper zone,
Balanus could always outcompete Chthamalus by crowding it off the rocks, undercutting
it, and replacing it even where it had begun to grow. When Connell removed Balanus
from the area, however, Chlhamalus was easily able to occupy the deeper zone,
indicating that no physiological or other general obstacles prevented it from
becoming established there. In contrast, Balanus cannot survive in the shallow-water
habitats where Chthamalus normally occurs. It evidently does not have the special
physiological and morphological adaptations that allow Chthamalus to occupy
this zone.
Viewing the same situation in evolutionary terms, one might suppose that Chthamalus
evolved later than Balanus or that it reached the Scottish coast later than
its rival. Unable to compete with Balanus in its main area of distribution,
it nevertheless could survive in the relatively less favorable shallow-water
habitats where Balanus did not occur. Conversely, it is possible that Balanus
evolved more recently or reached the area more recently than Chlhamalus and
possibly eliminated Chthamalus from most of its former range. In doing so it
would have restricted Chthamalus to the less favorable sites where it was subject
to periodic drying.
In another set of field observations, the late Princeton ecologist Robert Mac-Arthur
studied five species of warblers, which are small, insect-eating birds that
coexist part of the year in the forests of the northeastern United States and
adjacent Canada. Although they all appeared to be competing for the same resources,
MacArchur found that each species actually spent most of its time feeding in
different parts of the trees, and so each ate different subsets of insects in
those trees. Some of the species of warblers fed on insects near the ends of
the branches, whereas others regularly penetrated well into the foliage; some
stayed high on the trees, others fed on the lower branches; and these patterns
were recombined in different ways characteristic of each of the warbler species.
As a result of these different feeding habits, each species of warbler actually
occupied a different niche; in other words, they had different ways of using
the resources of the environment and thus were not really in direct competition
with one another for limited resources.
Species of barnacles, warblers, and many other kinds of organisms coexist because
the niches of the species involved differ.

