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Some of Howard's theoretical ideas were far ahead of their time. He had data on wasps that showed how behavior, including learning, could affect evolution, and he discussed the general importance of this especially in his 1966 book on sand wasps (see also Evans, 2002). Howard's ideas on how behavior can take the lead in evolution are now being cited more frequently than before, because finally evolutionary biologists are seeing how the condition sensitivity of organisms can supplement the genetic study of evolution. Howard saw that connection long ago, and because of it he wrote important critiques of overly gene-centered thinking, such as some analyses involving kin selection (e.g., see Evans, 1977). -- MARY JANE WEST-EBERHARD
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