Evolutionary drivers of caching behaviour in corvids.

Caching has recurrently evolved across a range of animal taxa to withstand fluctuations in food availability and in the context of intraspecific competition. It is widespread in the corvid family, which exhibit considerable interspecific variation in their behavioural and morphological adaptations to caching. However, the evolutionary drivers responsible for this diversity have seldom been explored. The present study systematically reviews the literature on caching behaviour in corvids globally to determine (1) which food caching strategies species have adopted (specialist, generalist or non-cacher) and (2) whether ecological factors affect the occurrence of different strategies, namely (a) climate breadth, (b) trophic niche, (c) habitat breadth, (d) centroid latitude, (e) centroid longitude, (f) breeding system, and (g) body mass. In addition, the ancestral states of caching are reconstructed to assess the evolutionary trajectory of each strategy. Caching strategies were identified in 63 species from 16 genera (out of 128 corvid species and 22 genera). Ancestral state analysis suggested specialist caching as the ancestral state in corvids. Type of caching is associated with distance from equator and by average body mass, with generalist caching concentrated around the equatorial zone and among heavier corvids, while specialist caching occurring more commonly in smaller species found farther from the equator. Although specialist caching most likely was the ancestral state in corvids, both specialist and generalist caching evolved several times independently in the family of corvids. Our results show caching to be widespread in corvids and affected by body size and latitude but ecological factors such as topic niche and habitat breadth and breeding system, not to be strong drivers shaping caching behaviour.