Sampling bias exaggerates a textbook example of a trophic cascade

Understanding trophic cascades in terrestrial wildlife communities is a major challenge because these systems are difficult to sample properly. We show how a tradition of nonrandom sampling has confounded this understanding in a textbook system (Yellowstone National Park) where carnivore [Canis lupus (wolf)] recovery is associated with a trophic cascade involving changes in herbivore [Cervus canadensis (elk)] behavior and density that promote plant regeneration. Long-term data indicate a practice of sampling only the tallest young plants overestimated regeneration of overstory aspen (Populus tremuloides) by a factor of 3-8 compared to random sampling because it favored plants taller than the preferred browsing height of elk and overlooked non-regenerating aspen stands. Random sampling described a trophic cascade, but it was weaker than the one that nonrandom sampling described. Our findings highlight the critical importance of basic sampling principles (e.g., randomization) for achieving an accurate understanding of trophic cascades in terrestrial wildlife systems.