Abstract

Spatial variation in cone serotiny in Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) across Yellowstone National Park influences initial pine recruitment after stand‐replacing fire with tremendous population, community, and ecosystem consequences. A previous study showed that much of the spatial variation in serotiny results from the balance of selection arising from high frequencies of fire favoring serotiny countered by opposing selection exerted by American red squirrels (Tamiasciurus hudsonicus) as seed predators. This earlier study, however, assumed stable local red squirrel densities over multiple generations of pines. Here, we examine environmental properties that might contribute to long‐term stability in the densities of red squirrels among sites. We found that the amount of clay in the soil, an indicator of plant and fungal growth—the latter an important food resource for red squirrels—and the coefficient of variation (CV) in diameter at breast height (DBH) of forest trees together account for a substantial amount of variation in red squirrel density. Soil development occurs over very long time scales, and thus, intersite variation in the amount of clay is unlikely to shift across pine generations. However, CV of DBH and squirrel density increase with stand age, which acts to amplify selection against serotiny with increasing interfire interval. Regardless, much of the variation in the CV of DBH is accounted for by soil bulk density, mean annual temperature, and surface curvature, which are unlikely to vary in their relative differences among sites over time. Consequently, these soil and abiotic attributes could contribute to consistent spatial patterns of red squirrel densities from one pine generation to the next, resulting in consistent local and spatial variation in selection exerted by red squirrels against serotiny.

Highlights

  • When we think of species having large and disproportionate impacts on communities, apex predators that drive trophic cascades come to mind (Estes et al, 2011)

  • The other variable in the models accounting for red squirrel density, namely variation in tree size at a site (CV of diameter at breast height (DBH)), increases with stand age, presumably because of gaps formed from tree mortality and subsequent infilling; most trees in young stands will have germinated soon after a fire, so DBH should be relatively homogeneous initially

  • A model that incorporated abiotic factors unlikely to vary with stand age, and are likely to contribute to consistent spatial differences among locations, accounted for a large proportion of the variation in DBH (Table 2C)

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Summary

Introduction

When we think of species having large and disproportionate impacts on communities (keystone species), apex predators that drive trophic cascades come to mind (Estes et al, 2011). The huge difference between having kelp forests and their diverse community of fishes, sea lions, and eagles, versus largely kelp-­less barrens arises from contemporary ecological processes, otters directly eating urchins and indirectly facilitating the increase in kelp. Such cascading effects are thought to be widespread both on land and in water where you have. Strongly interacting predators and herbivores potentially have strong evolutionary effects on their prey (Benkman, 2013; Benkman, Siepielski, & Parchman, 2008; Steinberg, Estes, & Winter, 1995) When such prey dominates a landscape (e.g., foundation species), the ecological consequences of these evolutionary effects could be profound. The evolutionary effect is the result of differential seed predation by American red squirrels (Fig. 1A; Tamiasciurus hudsonicus; Talluto & Benkman, 2014)

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