Patterns of exploitation of annually varying Pinus sylvestris cone crops by seed‐eaters of differing dispersal ability

Food consumption by animals depends on functional and numerical responses, and particularly the latter for food specialists. Some birds and mammals are conifer seed specialists and are likely to exploit the varying seed production in different ways according to differences in dispersal capabilities. This in turn may determine which species is more likely to affect the evolution of cone morphology. This study examined the inter‐annual pattern of foraging on Scots pine Pinus sylvestris cones by crossbills Loxia spp. (highly dispersive) and red squirrels Sciurus vulgaris (weakly dispersive) over 16 yr in three stands of ancient native pines at Abernethy Forest in Highland Scotland. There were synchronous annual variations in cone production across the stands, and an indication of a three‐year cycle. The number of cones taken by crossbills was correlated with cone production, indicating a numerical response by the birds, up to a certain limit of cone production. By contrast, the number of cones taken by red squirrels was not correlated with cone production. Rather, the percentage of cones taken by squirrels was high when cone production was low, and low when production was high. There was also a long‐term decline in the number of cones taken by squirrels, suggesting a decline in squirrel numbers. Although a high percentage of cones was removed from some cohorts on some trees, either by crossbills (maximum of 94.8%) or red squirrels (100%), the mean percentage of cones taken by crossbills from trees was small, ranging from 3.7 to 17.1% across all cone cohorts. For red squirrels, mean values ranged from 0.1 to 46.1% across all cohorts. However, given that crossbills can track changes in cone production by rapid numerical responses (i.e. through migration), and take a larger percentage of cones from cohorts of high production (10.7%), compared with red squirrels (3.6%), crossbills may be more important in driving the evolution of cone morphology because future trees are more likely to come from cohorts of high cone production.