one sense rivals.â Wallace, in return, was consistently deferential. In 1864, he wrote to Darwin: âAs to the theory of Natural Selection itself, I shall always maintain it to be actually yours and yours only. You had worked it out in details I had never thought of, years before I had a ray of light on the subject, and my paper would never have convinced anybody or been noticed as more than an ingenious speculation, whereas your book has revolutionized the study of Natural History, and carried away captive the best men of the present age.â
This genuine affection and mutual support masked a serious disagreement on what may be the fundamental question in evolutionary theoryâboth then and today. How exclusive is natural selection as an agent of evolutionary change? Must all features of organisms be viewed as adaptations? Yet Wallaceâs role as Darwinâs subordinate alter ego is so firmly fixed in popular accounts that few students of evolution are even aware that they ever differed on theoretical questions. Moreover, in the one specific area where their public disagreement is a matter of recordâthe origin of human intellectâmany writers have told the story backwards because they failed to locate this debate in the context of a more general disagreement on the power of natural selection.
All subtle ideas can be trivialized, even vulgarized, by portrayal in uncompromising and absolute terms. Marx felt compelled to deny that he was a marxist, while Einstein contended with the serious misstatement that he meant to say âall is relative.â Darwin lived to see his name appropriated for an extreme view that he never heldâfor âDarwinismâ has often been defined, both in his day and in our own, as the belief that virtually all evolutionary change is the product of natural selection. In fact Darwin often complained, with uncharacteristic bitterness, about this misappropriation of his name. He wrote in the last edition of the Origin (1872): âAs my conclusions have lately been much misrepresented, and it has been stated that I attribute the modification of species exclusively to natural selection, I may be permitted to remark that in the first edition of this work, and subsequently, I placed in a most conspicuous positionânamely, at the close of the Introductionâthe following words: âI am convinced that natural selection has been the main but not the exclusive means of modification.â This has been of no avail. Great is the power of steady misrepresentation.â
However, England did house a small group of strict selectionistsââDarwiniansâ in the misappropriated senseâand Alfred Russel Wallace was their leader. These biologists did attribute all evolutionary change to natural selection. They viewed each bit of morphology, each function of an organ, each behavior as an adaptation, a product of selection leading to a âbetterâ organism. They held a deep belief in natureâs ârightness,â in the exquisite fit of all creatures to their environments. In a curious sense, they almost reintroduced the creationist notion of natural harmony by substituting an omnipotent force of natural selection for a benevolent deity. Darwin, on the other hand, was a consistent pluralist gazing upon a messier universe. He saw much fit and harmony, for he believed that natural selection holds pride of place among evolutionary forces. But other processes work as well, and organisms display an array of features that are not adaptations and do not promote survival directly. Darwin emphasized two principles leading to nonadaptive change: (1) organisms are integrated systems and adaptive change in one part can lead to nonadaptive modifications of other features (âcorrelations of growthâ in Darwinâs phrase); (2) an organ built under the influence of selection for a specific role may be able, as a consequence of its structure, to