Patchy invasion and the origin of a hemlock-hardwoods forest mosaic

The record of forest invasion by eastern hemlock (Tsuga canadensis) during the course of Holocene migration provides useful information about invasion processes in temperate forest, a system that has been invaded by few exotic species. We used fossil pollen preserved in small forest hollows, which record forest composition on the scale of 1-3 ha, to study hemlock invasion of forests in the Sylvania Wilderness in the upper peninsula of Michigan, where there is now a mosaic of 3-30 ha stands dominated either by hemlock or by sugar maple (Acer saccharum) and basswood (Tilia americana). Fossil pollen was interpreted by comparison with 66 surface samples from small hollows in Michigan and Wisconsin, using three different statistical methods: pollen ratios, a dissimilarity index, and canonical variates analysis (CVA) ordination. We found that four hemlock stands along a 10-km transect across Sylvania originated as patches of white pine (Pinus strobus) forest that were invaded by hemlock ~3000 yr ago (calibrated ¹⁴C dating) when hemlock expanded its range in northern Michigan. Invasion occurred at about the same time (within 800 yr) at all sites but was not associated with disturbance at any site. Over the next several thousand years hemlock coexisted with white pine, but eventually, at a time that differs from site to site, hemlock became dominant, and white pine disappeared from all but one of the four stands. These changes were apparently driven by climate changes over the last 4000 yr that caused the water table to rise (Brugam and Johnson 1997). The history of four nearby maple stands is more variable and less well understood. Unlike the hemlock stands, three of the four maple patches were not dominated by pine at the time of hemlock invasion, but instead had abundant oak (Quercus) and/or maple. Two of these stands were not invaded by hemlock, and the third, if invaded at all, was invaded for a few centuries by low densities of hemlock trees. Thus invasion by hemlock was sensitive to the species composition of the resident forest. Sugar maple and basswood increased in these stands, and by 2000 to 800 yr ago, depending on site, they resembled modern maple stands. The fourth patch was invaded by hemlock, but it was converted to a maple stand 1000-500 yr ago. A wood layer in the sediment is evidence that a catastrophic windstorm may have been responsible.