Habitat heterogeneity and life-history traits influence presettlement distributions of early-successional tree species in a late-successional, hemlock-hardwood landscape

In landscapes dominated by late-successional plant communities, early-successional species may lead a tenuous existence, persisting only as fugitives or relying on refuges in marginal habitats to provide a persistent seed source. The objective of this study was to relate fine-scale distributions of early-successional tree species in hemlock-hardwood forests of northern Wisconsin, USA to potential landscape persistence strategies. A special emphasis was placed on eastern white pine (Pinus strobus), a restoration priority in the region. Witness tree data from nineteenth century US Public Land Survey records (encompassing 40,610?km² and 106,790 trees) were used with modern environmental data to relate species distributions to habitat characteristics. Early-successional tree species had strong positive associations with marginal habitats such as inclusions of sandy soil and margins of lakes, wetlands, and rivers. Marginal habitats occupied ~44 % of the landscape, which may help account for the abundance of early-successional species in our study area relative to other hemlock-hardwood forests. Populations of early-successional species in marginal habitats could also have provided important seed sources for the upland mesic landscape matrix, as >70 % of the landscape was within 200?m of these habitats. The degree to which early-successional species were limited to marginal habitats largely followed predictions based on species life-history characteristics, except that white pine was more common than expected in upland mesic habitats. These findings illustrate the potential importance of landscape heterogeneity for persistence of early-successional species in late-successional forest landscapes and provide baseline information on habitat associations and landscape dynamics that will be useful in restoration efforts.