632 THE STRUCTURE OF EVOLUTIONARY THEORY
genes in future generations. I shall use the language of convenience. Unless
otherwise stated, 'altruistic behavior' and 'selfish behavior' will mean behavior
directed by one animal body towards another [p. 50]... We shall continue to treat
the individual as a selfish machine, programmed to do whatever is best for his
genes as a whole. This is the language of convenience [p. 71]."
In a later book (1982, p. 4) Dawkins admits that proxy—that is, must select
gene replicators via organisms as causal actors: "The most important kind of
replicator is the 'gene' or small genetic fragment. Replicators are not, of course,
selected directly, but by proxy; they are judged by their phenotypic effects."
This argument, I think, has truly become an inadaptive meme, destined for
eventual extinction, but propagated wherever gene selectionism survives, whether
in technical literature or popular presentation. A major popular book on this topic
holds (Cronin, 1991, p. 289): "If organisms are not replicators, what are they? The
answer is that they are vehicles of replicators... Groups, too, are vehicles, but far
less distinct, less unified ... In this weak sense, then, 'group selection' could occur...
[but it] would in no way undermine the status of genes as the only units of
replicator selection. This does not mean that higher-level entities are unimportant
in evolution. They are important, but in a different way: as vehicles."
Bookkeeping and causality: the fundamental error of
gene selectionism
The error and the incoherence of gene selectionism, as documented above, can be
summarized in a single statement illustrating the fruitful, "Pareto-like" character of
the central fallacy: proponents of gene selectionism have confused bookkeeping
with causality. This error achieves its Pareto status of substantial utility because
changes recorded at the genetic level do play a fundamental part in characterizing
evolution, and records of these changes (bookkeeping) do maintain an important
role in evolutionary theory. But the error remains: bookkeeping* is not causality;
natural selection is a causal process, and units or agents of selection must be
defined as overt actors in the mechanism, not merely as preferred items for
tabulating results.
No one has ever stated the issue more accurately or succinctly than George
Williams himself (1992, p. 13), thus increasing my puzzlement at his failure to
recognize how his own formulation invalidates the gene selectionism that still wins
his lip service: "For natural selection to occur and be a factor in evolution,
replicators must manifest themselves in interactors, the concrete realities that
confront a biologist. The truth and usefulness of a biological theory
*Working through the logic and problems of this vexatious issue has been pursued
as a collective enterprise among many biologists and philosophers for more than 20
years. I have used the terminology of bookkeeping and causality for some time (Gould,
1994), and have developed or sharpened some of the arguments. But I do not think that I
devised the labels. I believe that I first picked up the terminology of bookkeeping from
arguments presented by the University of Chicago philosopher Bill Wimsatt. Many
authors have used this fruitful distinction for some time.