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Abstract:
Although conclusive evidence is lacking for its establishment, the thesis that complexity adds stability to communities is probably accepted by the majority of ecologists. I believe this attitude found its origins in the indisputable fact that there are latitudinal and altitudinal changes in community complexity. As one progresses northward or southward from the equator, or higher in altitude in most parts of the world, one cannot help but notice that communities tend to become simpler, that is, there are fewer species per community. At the same time, these communities appear to become less stable. But perhaps this change in stability is in appearance only; they appear to be less stable because of the relatively greater number of individuals comprising each species population in temperate areas. Each population, because of its greater numbers, is therefore conspicuous, and changes in these numbers are noticed. We are particularly aware of such changes because populations in these areas of the world have been comparatively well studied. Many of the most studied populations include species of economic importance where changes in population numbers are vital to agricultural or forestry practices. Equatorial populations, on the other hand, contain smaller numbers of individuals of each species because of the greater number of species present. Number changes are simply not as noticeable because the population itself is not as obvious among the other populations. It may be that when (if ever) we have as much data on equatorial populations as we have on those of temperate climates, we will find fluctuations of equal relative magnitude (but not of equal numbers, of course). If, on the other hand, we really do find a correlation between complexity and stability, the suggestion by May (12) that stability permits complexity may be well worth investigating. Because of its organization and physical setting, the Rhopalomyia community I have studied might be expected to have considerable stability. In fact, however, it does not. Each of the populations in the community fluctuates greatly and irregularly in both percentages and numbers, and these populations apparently become locally extinct occasionally, because they sometimes cannot be found even in extensive collections. After studying several of the more important parasitoid species, it is evident to me that there is little or nothing about their interactions that might induce greater community stability. Each species seems to have evolved into the community with no higher purpose than simply to usurp what it can from some other member, and it does this by concentrating its energies on better competitive mechanisms rather than higher reproductive capacities. There are never empty niches to be filled by organisms having the "correct specifications" because new niches are created out of parts of older, broader niches which were occupied by other, more r-selected organisms. Thus, perhaps we have read too much into community organization. Perhaps the "ifiling of niches" is essentially nothing more than the haphazard result of competitive jostling among species; and that as communities develop, they are not necessarily programmed for such things as greater stability or better energy utilizationthe species merely become more closely packed.