Vegetation patterns in heterogeneous landscapes: The importance of history and environment

Throughout the eastern United States, plant species distributions and community patterns have developed in response to heterogeneous environmental conditions and a wide range of historical factors, including complex histories of natural and anthropogenic disturbance. Despite increased recognition of the importance of disturbance in determining forest composition and structure, few studies have assessed the relative influence of current environment and historical factors on modern vegetation, in part because detailed knowledge of prior disturbance is often lacking. In the present study, we investigate modern and historical factors that control vegetation patterns at Harvard Forest in central Massachusetts, USA. Similar to the forested uplands throughout the northeastern United States, the site is physiographically heterogeneous and has a long and complex history of natural and anthropogenic disturbance. However, data on forest composition and disturbance history collected over the past > 90 years allow us to evaluate the importance of historical factors rigorously, which is rarely possible on other sites. Soil analyses and historical sources document four categories of historical land use on areas that are all forested today: cultivated fields, improved pastures/mowings, unimproved pastures, and continuously forested woodlots. Ordination and logistic regressions indicate that although species have responded individualistically to a wide range of environmental and disturbance factors, many species are influenced by three factors: soil drainage, land use history, and C:N ratios. Few species vary in accordance with ionic gradients, damage from the 1938 hurricane, or a 1957 fire. Contrary to our expectation that the effects of disturbance will diminish over time, historical land use predicts 1992 vegetation composition better than 1937 composition, perhaps because historical woodlots have become increasingly differentiated from post-agricultural stands through the 20th century. Interpretations of modern vegetation must consider the importance of historical factors in addition to current environmental conditions. However, because disturbances such as land use practices and wind damage are complex, it is often difficult to detect disturbance effects using multivariate approaches, even when the broad history of disturbance is known.