In classical evolutionary theory, traits evolve because they facilitate organismal survival and/or reproduction. We discuss a different type of evolutionary mechanism that relies upon differential dispersal. Traits that enhance rates of dispersal inevitably accumulate at expanding range edges, and assortative mating between fast-dispersing individuals at the invasion front results in an evolutionary increase in dispersal rates in successive generations. This cumulative process (which we dub "spatial sorting") generates novel phenotypes that are adept at rapid dispersal, irrespective of how the underlying genes affect an organism's survival or its reproductive success. Although the concept is not original with us, its revolutionary implications for evolutionary theory have been overlooked. A range of biological phenomena (e.g., acceleration of invasion fronts, insular flightlessness, preadaptation) may have evolved via spatial sorting as well as (or rather than) by natural selection, and this evolutionary mechanism warrants further study.colonization | evolution | spatial disequilibrium | nonadaptive evolution I n 1859, Charles Darwin proposed a mechanism to explain the process by which organisms become well matched to local conditions. That mechanism was natural selection (1). At its heart lies the concept of differential lifetime reproductive success (LRS). Significant extensions to the paradigm since Darwin's work-such as multiple levels of selection (2, 3)-all rely upon the basic principle that, through time, some genes leave more copies of themselves than do others (4). Here we describe an additional mechanism whereby traits evolve because genes are differentially successful through space rather than time. This idea is not new; the process was described long ago (5), has been explored in several spatially explicit models of nonequilibrial populations (6-8), and is widely recognized by researchers who work with range-edge dynamics (9). The basis of the idea is that on expanding range edges evolutionary change can arise from differential dispersal rates (spatial sorting) as well as from differential survival or reproductive success. Spatial sorting and classical natural selection both require heritable variation, and both result in deterministic shifts in phenotypic attributes, but the two evolutionary processes rely on fundamentally different mechanisms (spatial filtering versus temporal filtering). Mainstream biology has failed to recognize that evolutionary change can be caused by spatial sorting as well as by conventional natural selection.
Spatial SortingImagine a species expanding its range into hitherto unoccupied territory and with a genetic basis to variation among individuals in dispersal rates (6-8). For example, continuously distributed variation may occur in dispersal-relevant morphological traits [e.g., seed shape (5), flight musculature and wing size (7, 10), leg length (11), foot size (12)], behavior [movement patterns (13)], and physiology [locomotor endurance (14)]. Alleles that confer the hi...