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Extending microevolutionary theory to a macroevolutionary theory of complex adaptations

Over the last two decades of the 20th century, the macroevolutionary theory was transformed. Held at the outset of that period, the 1980 Chicago colloquium “Macroevolution: Pattern and Process” became notorious for its heated debates about punctuated equilibria and the adequacy of “Neo-Darwinism.” The debates were so intense that the proceedings were never published (Futuyma 2015). Punctuated equilibria may have dominated the debate but advocates of that theory were not the only workers who found Neo-Darwinism inadequate to explain macroevolution. What seemed missing, especially to morphologists, were the processes, both developmental and evolutionary, that could explain complex adaptations. Even those willing to accept, on principle, that gradual allelic substitution is a model for evolutionary change and that adaptation is due to natural selection, might still wonder whether that was enough to explain complex adaptations. What seemed missing was articulated by Gould in his promise of a new theory (1980, p. 129): “... the new theory will restore to biology a concept of organism. In an exceedingly curious and unconscious bit of irony, strict selectionism... debased what had been a mainstay of biology–the organism as an integrated entity exerting constraint over its history”. That view of organisms as integrated was indeed a mainstay of morphology; as Weidenreich (1941, p.323) put it, “It is an entirely inconceivable idea that each morphological unit should have developed independently from all others and from the body as a whole, as an isolated and incidental variation, subsequently tested as to its feasibility for the organism...” Defending the theory, Charlesworth, Slatkin, and Lande responded that “...the concept of organism, including constraints of history, development and architecture, which Gould (1980)