Multi-mode fluctuating selection in host-parasite coevolution

Understanding fluctuating selection is important for our understanding of patterns of spatial and temporal diversity in nature. Host-parasite theory has classically assumed fluctuations either occur between highly specific genotypes (matching allele: MA) or from specialism to generalism (gene-for-gene: GFG). However, while MA can only generate one mode of fluctuating selection, we show that GFG can in fact produce both rapid 'within-range' fluctuations (among genotypes with identical levels of investment but which specialise on different subsets of the population) and slower cycling 'between ranges' (different levels of investment), emphasising that MA is a subset of GFG. Our findings closely match empirical observations, although sampling rates need to be high to detect these novel dynamics empirically. Within-range cycling is an overlooked process by which fluctuating selection can occur in nature, suggesting that fluctuating selection may be a more common and important process than previously thought in generating and maintaining diversity.