“…The neutral diversity and adaptation in spatially expanding populations has been studied in computer simulations (Edmonds et al, 2004; Klopfstein et al, 2006; Kuhr et al, 2011; Kuhr and Stark, 2015; Lavrentovich and Nelson, 2014; Otwinowski and Krug, 2014), in the field (Ramachandran et al, 2005; White et al, 2013; Louppe et al, 2017), and in microbial colonies (Hallatschek et al, 2007; Fusco et al, 2016; Gralka et al, 2016b; Korolev et al, 2011), which can serve as a useful model system because short generation times and ease of handling allow for quantitative investigations of the evolutionary dynamics of range expansions. In microbial colonies, nutrient gradients and mechanical effects limit the number of proliferating individuals to a small region close to the colony perimeter called the growth layer (Grant et al, 2014; Gralka et al, 2016b; Warren et al, 2019). For mutations occurring inside the growth layer, most mutant offspring are concentrated in a relatively small number of enormously successful lineages that manage to remain at the front and 'surf’ on the expanding population wave (Excoffier and Ray, 2008).…”