Facilitation by Pinus flexilis during succession: a hierarchy of mechanisms benefits other plant species.
Studies of facilitation have primarily been limited to single mechanisms, species, or environments. We examined interacting mechanisms governing the facilitative effects of Pinus flexilis on two later successional understory species, Pseudotsuga menziesii and Ribes cereum, in different microhabitats and seasons at the ecotone between the Rocky Mountain forests and Great Plains grasslands in Montana, USA. In field surveys, 69% of Pseudotsuga and 91% of Ribes were located beneath P. flexilis even though P. flexilis subcrowns accounted for a small proportion of available habitat. For three years, we monitored the survival of Pseudotsuga and Ribes seedlings experimentally planted beneath P. flexilis and in the open at a windward and a leeward site. Survival of both species was highest beneath P. flexilis at a site topographically protected from strong unidirectional winds (38% for Pseudotsuga and 63% for Ribes), and lowest at a windward site and in the open where tree crowns did not provide shelter from winds (2% and 6%, respectively). These results suggest that wind amelioration contributed to the facilitative effect of P. flexilis. However, even at the leeward site, where wind speed was low, survival of Pseudotsuga and Ribes was higher beneath P. flexilis, suggesting the importance of shade. To explore the relative importance of different mechanisms, we designed an experiment with six treatments: "shade," "shade + wind," "shade + drift," "wind," "drift," and a "control." After two years, we found shade to be of overwhelming importance for the survival of Pseudotsuga and Ribes. Without shade, no other treatments were significant, but once shade was provided, wind amelioration and snow pack accumulation increased survival of Pseudotsuga, suggesting that these different facilitative mechanisms functioned in a nested hierarchical manner: some mechanisms were important only when others were already functioning. Many studies have demonstrated multiple interacting mechanisms in the way that plants interact, but to our knowledge hierarchical interactive processes have not been previously documented. If the effects of positive or competitive mechanisms are often hierarchical, then studies of isolated mechanisms may not accurately assess their importance in nature.