14 15 16 17 18 KEYWORDS (4-6): Evolutionary stable strategies, masting, scatterhoarding, pilferage, seed caching, 19 synzoochory. 20 21 22 23 Number of words: 7127 24 ABSTRACT 25Many plant species worldwide are dispersed by scatterhoarding granivores: animals that hide seeds in 26 numerous, small caches for future consumption. Yet, the evolution of scatterhoarding is difficult to 27 explain because undefended caches are at high risk of pilferage. Previous models have attempted to 28 solve this problem by giving cache owners unrealistically large advantages in cache recovery, by kin 29 selection (but individuals that cache and those that pilfer are usually unrelated), or by introducing 30 reciprocal pilferage of "shared" seed resources. However, the role of environmental variability has 31 been so far overlooked in this context. One important form of such variability is masting, which is 32 displayed by many plant species dispersed by scatterhoarders. We use a mathematical model to 33 investigate the influence of masting on the evolution of scatter-hoarding. The model accounts for 34 periodically varying annual seed fall, caching and pilfering behavior, and the demography of 35 scatterhoarders. Masting, through its effects on population density, reduces cache pilferage and 36 lowers the reproductive cost of caching (i.e. the cost of caching for the future rather than using seeds 37 for current reproduction). These reductions promote the evolution of scatter-hoarding behavior 38 especially when interannual variation in seed fall and the period between masting events are high. 39the per capita advantage of owner versus pilferer, but under natural conditions there typically are 58 many potential pilferers per each cache owner. Thus, caches are more likely to be emptied by a 59 pilferer than by the owner even when per capita advantage of the latter is substantial. Second, most 60 empirical studies have been short-term, with cache recovery examined over several days at most. 61Long-term (e.g. overwinter) removal rates might be very similar for cache owners and pilferers 62 because cache locations are forgotten over time (Balda and Kamil 1992;Bednekoff et al. 1997; 63 Barkley and Jacobs 1998). Over the long-term, cache owners will increasingly rely on olfaction or 64 random search, just like naïve individuals do. As a consequence, the owners advantage appears to be