Epistatic strength, modularity, and locus heterogeneity shape the number of local optima in fitness landscapes
Fitness landscapes provide a quantitative framework for understanding how natural selection shapes evolutionary trajectories. A central feature of these landscapes is their number of local optima, which determines whether fitness-increasing evolution can proceed towards a global optimum or become trapped on suboptimal peaks. Although multiple peaks are known to require reciprocal sign epistasis, the quantitative relationship between epistasis and number of peaks remains incompletely understood. Here, we show that for a broad class of unstructured fitness landscapes, i.e. isotropic Gaussian random fields, the expected number of local optima is determined by a single local measure of epistasis: the correlation of fitness effects. This provides a baseline prediction for the number of peaks in typical unstructured landscapes and links peak density directly to the amount of reciprocal sign epistasis. This baseline changes when epistatic interactions are structured. We show that clustering interactions within blocks of loci slightly increases the number of local optima. In contrast, strong heterogeneity between loci, where only a small subset of loci participate in epistatic interactions, causes the number of peaks to collapse. These results show that the number of local optima is governed not only by the overall strength of epistasis, but also by how epistatic interactions are distributed across the genotype space. Our framework therefore reconciles the central role of reciprocal sign epistasis with the observation that landscapes with similar amounts of epistasis can differ substantially in ruggedness, and provides a guide to the range of peak numbers expected in typical landscapes.