Proc. Nat. Acad. Sci. USA Vol. 69, No. 11, pp. 3151-3155, November 1972 Competition, Competitive Repulsion, and Coexistence (faunal equilibriums/exclusion principle/niche/ecologic range) P. J. DARLINGTON, JR. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 Contributed by P. J. Darlington, Jr., August 14, 1972 ABSTRACT This manuscript is concerned with con- it related to competition? It can be argued that fox and rabbit cepts rather than abstruse details or mathematics. Dis- compete for the energy the rabbit or for the space cussed are: competition; extended competition, proposed represents for competition in the strict sense, extended and modified the rabbit occupies; if there were fewer foxes there would be by all related interactions including predation, parasitism, more rabbits. Predation can have additional, indirect effects. disease, and even cooperation, all of which can be "weapons If a predator depends on one species of prey, the prey may of competition"; competitive repulsion, proposed for survive indefinitely, with only limited population fluctuations. the sum of forces that determine spacings, including But if the predator utilizes two prey species, one may be re- ecologic spacings, of individuals and populations; Darwin (biotic) equilibriums; competitive extinction, Gause's duced to extinction. In this case, one species of prey becomes principle, limited and limiting resources, and single- extinct as a result of the presence of the other. Is this com- resource competition; de facto coexistence of competing petition? Disease and parasitism can have similar effects; species, exemplified by green plants competing for sun- species that tolerate them can eliminate or those that light; niche competition; the two concepts of competitive repel exclusion; devision of resources and of their utilizers; do not. AMayr (ref. 1, p. 75), quoting Haldane, calls disease a cause and effect in real situations; and niches, niche "weapon of competition." A recent note in Nature (2) sum- overlap, and coexistence. Stressed is the complexity of the marizes an apparent example: tsetse-fly-carried trypano- real world, and the confusion that can and does arise from somes, tolerated by native African big game but fatal to many modeling it too simply. incoming herbivores (e.g., to cattle brought by man) may This manuscript is not concerned with abstruse details or protect the African fauna against immigrants from other mathematics, but is a reassessment of some general ecological continents. Is this competition? and evolutionary concepts. These concepts, including those of Attempts to answer these questions and to fix a precise competition, repulsion, biotic equilibrium, exclusion, coexis- biological definition of competition do not seem to clarify the tence, and the niche, are under intense study now by both subject. Recognition of different categories of competition naturalists and mathematical biologists, but are not con- seems more likely to do so. Of the many categories that might sistently defined and not yet adequately understood. In order be distinguished, I shall now propose and discuss three: ex- to facilitate understanding among all interested persons, I tended competition, single-resource competition, and niche think it is important to define and discuss these concepts in competition. terms consistent as possible with ordinary usage. Extended competition and competitive repulsion THEORY The term extended competition is now proposed to include competition, in the strict sense, extended and modified by Definitions and categories of competition any and all related processes and interactions, including preda- In ordinary usage "competition" means a contending of rivals tion, parasitism, disease, and even cooperation-cooperation for almost anything, either directly by almost any means is a "weapon of competition" if success of the cooperators ("all is fair in love and war") or indirectly (as for resources or reduces the success of competing forms. This totality of inter- markets in commerce). Biologists usually define "competi- actions is, of course, Darwin's "struggle for existence." It can tion" more narrowly, as including only actions and inter- be considered as "competition for places in the world" and as actions (or exploitations and interferences, or "active de- including "any interaction among (plants and) animals, no mand") of individuals or species seeking the same "resources." matter how complex and indirect it may be, that is or may be This definition is intended to be precise but in fact is not, for disadvantageous to any of them" (ref. 3, I. 23). it leaves open the question of what "resources" are. Food and A useful, evocative term derived from "extended compe- (for green plants) light clearly are resources that are com- tition" is competitive repulsion (see physicists' "repulsion" peted for. Many biologists would say that competition occurs among atoms), now proposed for the sum of all forces that also for such things as mates and territories. But are mates determine the spacings (including ecologic spacings), and and territories resources? Is a multidimensional "niche" a thus the numbers, both of individuals in populations and of resource? Is space as such a resource? If so, are places in a populations everywhere. [I coined this term, then found that biota or simply places on the earth resources? E. 0. Wilson (ref. 4, p. 195) had already used "repulsion" ill Biologists usually exclude predation from their definition a similar but more restricted way, and it may well have been of competition, and this raises further questions. When a fox used by other biologists.] Competitive repulsions determine kills and eats a rabbit, is this competition, and if not, how is biotic equilibriums. The existence of biotic equilibriums is one 3151 Downloaded by guest on September 25, 2021 3152 Zoology: Darlington Proc. Nat. Acad. Sci. USA 69 (1972) of the fundamental facts of evolution, biogeography, and Gause said (not in exactly these words) that extinction of one ecology. It is not a new discovery of mathematicians, but has of two competing populations will follow when a resource been recognized since Darwin (ref. 5, p. 109), who said, competed for is completely utilized (he said "completely seized"); and in Gause's model limitation of the resource was . .as from the high geometrical powers of increase of all organic beings, each area is already fully stocked with inhabi- the only factor limiting growth of the populations. The prin- tants, it follows that as each selected and favored form increases ciple that Gause did in fact expound may reasonably be called in number (of individuals), so will the less favored forms de- Gause's principle and may be restated thus: two populations crease. .. But we may go further than this; for as new forms (species) cannot long coexist if they compete for a vital re- are continually and slowly being produced, unless we believe source limitation of which is the direct and only factor limiting that the number of (species) goes on perpetually and almost both populations. As thus restated, the principle is, I think, indefinitely increasing, numbers must inevitably become extinct. valid without exception. This is the first statement of the relation between additions The relatively simple competition involved in this case may and subtractions of species that produces biotic equilibriums. be called single-resource competition. It may occur among the Because Darwin did state this relation, even though impre- most diverse organisms. For example, grass-attacking viruses, cisely, and in order to emphasize the continuity of this and grasshoppers, geese, and cattle may compete for grass, many other concepts from the pre-mathematical era of biology and any one of them may cause competitive extinction of any to the present, I propose that the equilibriums that determine other. numbers of individuals and of species in "each area" be called Darwin equilibriums. Limited and limiting resources Evidences of such equilibriums in time and space, and their Resource competition does not always lead to extinction. Two relation to competition, have been summarized elsewhere or more species that compete for an essential resource often (ref. 3, p. 553), thus: seem to coexist, and obvious ways can be suggested in which Throughout the recorded history of (animals), whenever the they may do so. For example, two species that depend on the record is good enough, the world as a whole and each main same resource may coexist if their populations are limited by part of it has been inhabited by . fauna(s) that (have) been separate density-dependent factors (say, each by its own virus reasonably constant in size and adaptive structure. Neither disease) so that the two together do not fully utilize the re- the world nor any part of it has been overfull of animals in one source. A case like this is analyzed mathematically and graph- epoch and empty in the next, and no great ecological roles have ically by Wilson and Bossert (ref. 8, chap. 3, p. 161, Fig. been long unfilled . Existing faunas show the same balance. 18, case 4). A more complex case (competition for light by Every continent has a fauna reasonably proportionate to its diverse plants) is considered below. In these cases, limitation area and climate, and each main fauna has a reasonable pro- of the resources no longer directly and simply limits the popu- portion of herbivores, carnivores, etc. This cannot be due to chance. Something holds the size and composition of faunas lations, and the cases do not invalidate Gause's principle. within limits in spite of continual changes and successions in Cases like these emphasize the need to distinguish limited separate phylogenetic groups. Only (extended) competition from limiting, and directly and simply from indirectly and com- can do this, and to do it competition (competitive repulsion) plexly limiting resources. The world is a limited area; all re- must be a fundamental, ever-present force (maintaining faunal sources are limited; and all normal populations will increase to equilibriums).