The Roots of Multilevel Selection: Concepts of Biological Individuality in the Early Twentieth Century

As multilevel selection theory has gained greater acceptance over the past quarter-century, scientists and scholars have shown an increased interest in the theory's historical antecedents. Despite this interest, however, the early twentieth century remains largely unexplored. It is generally assumed that biologists thought "naively" about evolutionary dynamics during this era, and that their attempts to explain biological phenomena often lacked sophistication. Now that several recent works have called attention to the complex relationship between biological individuality and the levels of selection, we believe it will prove instructive to revisit these early-twentieth-century biologists and reassess their criteria for biological individuality. Doing so reveals that they constructed a multilevel explanation of evolution that anticipated modern interpretations in several important ways. Though it is certainly true that most of these early biologists failed to recognize natural selection's pervasive agency, it is no less true that one of them, termite expert Alfred Emerson, artfully united the multilevel theory of "emergent evolution" with natural selection in a way that differs but little from the theory of multilevel selection that many scientists and scholars now promote. After reviewing the historical record, we place these early-twentieth-century biologists in their proper historical context, and we compare their interpretation of evolution with modern interpretations.