Biotic and abiotic constraints on shifts in temperate savanna ecotones at lower treeline

In contrast to documented increases in woody plant dominance of former savannas and grasslands of North America, oak (Quercus L.) savannas that form lower treelines in the southwestern United States and northwestern Mexico have been relatively stable over the past millennium. This research identified potential biotic and abiotic constraints on seedling recruitment of Quercus emoryi Torr. (Emory oak) within the context of potential shifts in lower treeline. Field surveys were used to describe seedling distribution at and below lower treeline, and to determine the potential for acorn dispersal from lower treeline into adjacent grassland. Field and greenhouse experiments designed to determine constraints on seedling establishment included reciprocal soil transfers, nutrient amendment studies, provision of artificial shade, and manipulation of seasonal precipitation inputs. Results indicate that rates of Q. emoryi recruitment within grasslands below treeline are relatively low, and are constrained by low rates of seed dispersal coupled with a low probability of seedling emergence. Seedling recruitment rates were directly correlated with quantity of summer precipitation, but were independent of winter precipitation. Results of this and complementary research suggest that lower treeline in southern Arizona is stabilized by self-enhancing feedback mechanisms of overstory shade, seed dispersal, and seedling establishment coupled with strong abiotic constraints beyond the current ecotone. The observed shift in treeline in the last millennium was less likely the result of slow, spatial progression of autogenic safe sites than the result of episodic and infrequent allogenic processes that simulated, or negated the importance of, conspecific, biogenic safe sites. Increases in summer precipitation are one such process that would facilitate (historic or potential future) downslope shifts in lower treeline. This interpretation is consistent with observations that downslope shifts in lower treeline which occurred 700-1700 ybp coincided with a period of particularly high summer precipitation in the region (i.e., the "Medieval Warm" period, 645-1295 ybp)