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Invisible Trade-offs: Van Noordwijk and de Jong and Life-History Evolution

When I started my PhD in 2000, I was a part of a cohort of graduate students delighted with the prospect that evolutionary biology could be used to predict and explain patterns of traits observed in nature. We were surrounded by a wealth of wonderful theory at varying levels of abstraction from reality, and our goal was to apply this theory to data from the specific systems that we studied. Making predictions about traits in evolutionary biology invariably involves discussions of trade-offs. Everything eventually comes back to survival and fertility, and if there is nothing to hold back survival (or fertility), you can predict nothing more interesting than that survival (or fertility) should bemaximized. Consequently, almost every graduate student in my cohort was looking for a trade-off. Not one of us could find one. It was frustrating; we thought Darwinian demons were unlikely. We knew that, somewhere, allocation decisions must have a cost; but in systems ranging from birds to plants, none of us had any success in quantifying anything that looked like one fitness component going up while another came down. In my system, plants that grew better also survived better and often produced more seeds. And then one of us stumbled across van Noordwijk and de Jong’s 1986 paper in The American Naturalist. There are many reasons to treasure this paper—for example, it contains my favorite definition of life-history theory: “an elaborate answer to the simple question of why havingmore offspring is not always selected for” (p. 137)—but it also