Toward Hierarchy: Trends and Tensions in Evolutionary Theory
Is the synthesis a complete and satisfactory evolutionary theory? It would be a trivial and derogatory exercise indeed to depict the synthesis in utterly simplistic terms and then turn around and conclude that it is incomplete and unsatisfactory. Though I have subjected the synthesis to a series of purifying distillations through the course of this book so far, I have in mind by no means solely Mayr’s (1980, p. 1) two-sentence summary of the synthesis as I pose the question of just how complete, workable, and satisfactory a theory of the evolutionary process it is. No serious student of evolutionary theory could ever claim that the modem synthesis is “just population genetics.” Many more phenomena are included than the statics and dynamics of genetic change in populations. What does seem to be true of the synthesis in general is that it focuses its concerns on a certain range of biological entities and attendant processes, and espouses attitudes and ontological positions on others that appear (to me) to exclude the latter from effective integration into the theory per se. Specifically, the synthesis focuses on genes; their replication, recombination, and mutation; and the fate of allelic variation within populations. But species—certainly not the sole province of population genetics—are very much a part of the synthesis, if equivocally so. If it is true that most evolutionary phenomena considered by the synthesis are construed, at least in principle, to be explicable in terms of the dynamics of selection and drift of allelic variation in populations, it is not because other sorts of phenomena, such as macroevolutionary trends, are alleged not to exist. The synthesis takes the (on the whole commendable) attitude of the Missourian who must be shown. We have a highly corroborated theory of the origin, maintenance, and modification of adaptations—through pure, narrowly defined natural selection. The burden of demonstration lies on anyone who would maintain either that some other process builds such (organismic) adaptation or that an additional process (or more than one) is also at work in evolution. Certainly the entire discussion on levels of selection, a discussion to which I return in this chapter, is structured in this general sort of way.