Questions:Open-grown shade trees retained in pastures are legacies of agricultural history that influence recolonization dynamics and create structural heterogeneity in post-agricultural forests. This work addresses the long-term influence of pasture trees on forest structure and composition. I ask: Do signs of successional nucleation in microhabitat, structure and composition near pasture trees persist decades after canopy closure? If so, does variation primarily reflect seed deposition or environmental facilitation?Location: Southeastern Ohio, USA.Methods: Thirteen open-grown trees greater than 70 cm in diameter in secondgrowth forests 40-90 years old were selected as study sites. Trees, saplings, herbs, and environmental variables were sampled and compared at three positions around each tree: under the canopy, at the dripline, and in the forest matrix.
Results: Soil pH was lower and coarse woody debris most abundant under pasture tree canopies. Stems >10 cm in diameter at breast height were nearly absent under each focal tree's canopy and most abundant at its dripline. Composition under the canopy was limited to shade-tolerant trees, whereas the dripline was characterized by vertebrate-dispersed species. Ant-dispersed, adhesive, and unassisted herb species were most abundant under embedded pasture trees compared with the forest matrix. Discussion: Evidence of nucleation around open-grown trees persists for decades after canopy closure. Contrasts in structure and composition reflect spatially segregated interactions of competitive suppression and facilitation mediated by both dispersal and environment. Pasture trees contribute variation in microhabitat, composition, and structure to post-agricultural forests, and their retention should be encouraged. K E Y W O R D S agricultural legacies, community assembly, land-use history, nucleation, pasture trees, second-growth forest, succession | 455
Journal of Vegetation Science
HOLMES
| INTRODUC TI ONNearly all of the deciduous forests of eastern North America were cut by the start of the 20th century, and most of the cutover land was converted to European-style agriculture (Williams, 1992). Despite clearing, remnant trees were common in the agricultural landscape.Pasturing practiced by settlers often included retention of trees to provide shelter and food for livestock, a millennia-old practice that can be traced to the wood-pastures of Europe. Pasture trees were historically nut-bearing species such as Quercus, Fagus, and Castanea, or fleshy-fruited species such as Pyrus in silvopastoral systems that combine animal pasturing with fruit production (Harding and Rose, 1986;Hartel et al., 2013).The ecological contributions of pasture trees reflect both their place in the landscape matrix and their distinctive structure. Trees in open environments, such as pastures after clearing, spread laterally in response to light availability, resulting in large individuals with distinctive horizontal branching that persists after canopy closure (McEwan and McCarthy, 2008;Varga et al...