 Perhaps the most important of all of the criticisms of “Pop Sociobiology” was that it was excessively adaptationist; similar sorts of criticisms are leveled at modern descendants of sociobiology, such as behavioral ecology. Behavioral ecologists, including HBEs, are really methodological adaptationists (Godfrey Smith, 2001), in that they assume that natural selection is optimizing the trait they are studying as a heuristic, in order to establish other things about that trait; as such, no strong commitment about the power of natural selection is necessarily required on their part (as discussed earlier). This did lead many behavioral ecologists to regard Gould and Lewontin’s (1979) criticisms as a cautionary tale, rather than as properly directed at them—see, for example, Gould and Lewontin’s objection that adaptationist hypotheses were untestable, which the behavioral ecologists could easily rebut (Mayr, 1983). Similarly, Parker and Maynard Smith (1990) responded to Gould and Lewontin’s objection that adaptationists do not properly account for phylogenetic and developmental constraints; Parker and Maynard Smith argued that they are taken into account—even where developmental constraints and tradeoffs are not explicitly represented in optimality models, they are often built into the strategy set or are otherwise implicit in those models. However, it is reasonable to point out that Parker and Maynard Smith’s paper did not fully respond to Gould and Lewontin’s objection: Gould and Lewontin recognized that adaptationist models appeal to constraints; their worry was that such constraints are appealed to ad hoc to fill in gaps in models, and are not subjected to any external test.