Caught between birds and squirrels, limber pines go both ways

Responding to natural selection often means compromising between different selective forces. A brief paper published online early at Evolution documents one such case – limber pine trees' compromise between protecting their seeds from squirrels, and making them accessible to the birds that disperse them. Pulled between these conflicting selective sources, some limber pine populations grow cones in a wider variety of shapes [$a].

Cones of the limber pine must balance protection against squirrels with accessibility for seed-dispersing Clark's nutcrackers. Photos by Fool-On-The-Hill and almiyi.

Seeds of the limber pine are cache-dispersed by Clark's nutcrackers. That is, the birds collect pine seeds to cache as a winter food source, but they collect many more than they need, and forget lots of them, and forgotten seeds are often able to sprout. However, squirrels also like pine seeds, and they harvest them before the birds start caching seeds. These two seed-harvesters generate conflicting selection on limber pine cones [$a]. Nutcrackers go for cones with lots of seeds protected by thinner scales; but so do squirrels.

However, pine-nut-eating squirrels are not present everywhere limber pines grow. The new study's authors, Siepielski and Benkman, take advantage of this quirk of distributions to perform a natural experiment, comparing pines that only need to satisfy their seed dispersers with pines that also need to defend against seed predators. Surveying cone shapes in populations of each class, they found that limber pine populations facing conflicting selection were bimodal, with trees mainly growing either squirrel-defended short, thick-scaled cones, or nutcracker-friendly longer, thin-scaled cones. Populations growing in regions without squirrels produced only nutcracker-friendly cones.

This apparently simple pattern conceals more complicated dynamics – in fact, as the authors disclose in the Discussion section, many other limber pine populations are solely composed of trees producing squirrel-defended cones. This is because, when pines establish in areas with large squirrel populations, nutcrackers may never colonize the area, or may visit less frequently and disperse fewer seeds. Without nutcracker dispersal, seeds are mainly dispersed after the cones fall, by rodent species that (unlike squirrels) forage on the ground. This makes squirrel defense the only selective priority. Populations displaying both cone types probably only arise in unique conditions, the authors say, where squirrels are present but not at high density.

References

Siepielski, A., & Benkman, C. (2007). Convergent patterns in the selection mosaic for two North American bird-dispersed pines. Ecological Monographs, 77 (2), 203-20 DOI: 10.1890/06-0929

Siepielski, A., & Benkman, C. (2009). Conflicting selection from an antagonist and a mutualist enhances phenotypic variation in a plant. Evolution DOI: 10.1111/j.1558-5646.2009.00867.x