The Red Queen

Abstract

Maynard Smith (1976a) has given a useful alternative derivation of the prediction, from the Red Queen's Hypothesis, of the constancy of extinction. It seems appropriate, however, to note that all his conclusions are stated or implied also in my first paper on the subject (Van Valen 1973, pp. 17-19). We therefore do not seem to be in any important disagreement, as he implies. Even his lag load (Maynard Smith 1976b) is the genetical equivalent of my ecologically based environmental load (Van Valen 1973). I believe that we developed this concept independently. The Red Queen's Hypothesis, that any gain in fitness by one unit of evolution is balanced by losses in fitness by others, is most easily justified by equating realized fitness with control of trophic energy. (This is only a first approximation; I have justified the deeply but clandestinely radical approach of energetics and have given more accurate expressions elsewhere [Van Valen 1976].) More than a very small average rate of change in energy availability over geological time would give absurdly low or high values in the Paleozoic or even later. And geochemical evidence (Van Valen 1976) suggests even less change than this argument requires. Even on ecological time scales the usual approximate constancy of available energy is an important underlying part of the ancient notion of the balance of nature. The Red Queen's domain is less than universal, but first-order processes deserve first attention. Maynard Smith and I do disagree on one minor point. He says (1976a, p. 329), "The difficulty [with the zero-sum assumption for resources] is that not all evolutionary changes in a species increase its share of available resources. For example, adaptation of a prey species to escape predation need not increase its numbers and in general will not do so unless predation is a density-limiting factor." But individuals that escape predators continue to control energy, while those that succumb lose all their control. Predator-resistant phenotypes thereby come to control more energy. Selection is on the individual level here, not the group level, so it is not surprising that realized group fitness is not increased. Prospective group fitness may well be increased, but that is a very different matter (Van Valen 1976). The general approach of the Red Queen was used by Fisher, Darwin, and Lyell (Van Valen 1973). Darwin's use is even clearer in his recently published book on natural selection (Darwin 1975) than in the Origin. In fact, his analogy of wedges comes from his earliest thoughts on evolution after reading Malthus and is recorded in his third notebook, of 1838 (Darwin 1967, p. 163). Darwin's thoughts were presumably inspired by Lyell's equilibrial outlook. Everything changes, as Lyell (1832) knew from the fossil record, but everything is the same. On Lyell's use of group selection in 1832 in a shifting balance between selective extinction and presumably random creation, see Van Valen (1975). Darwin, but not Lyell, saw that such a process gives an average increase in competitive fitness, but Darwin remained true to Lyell's denial of overall upward progress of life through geologic time. Lyell's mechanism was the same as Darwin's, on a different level, and was also based on competition: the antievolutionist Lyell