A 20-year experiment on the effects of deer and hare on eastern hemlock regeneration

Eastern hemlock (Tsuga canadensis (L.) Carrière) is a shade-tolerant, slow-growing tree once common in forests across the Great Lakes region. It was heavily exploited in the late 19th and early 20th centuries and now experiences limited regeneration across much of its range. This failure to regenerate has been ascribed to poor seedbed conditions, insufficient canopy openings, warmer climate, and browsing by white-tailed deer (Odocoileus virginianus (Zimmermann, 1780)) or snowshoe hare (Lepus americanus Erxleben, 1777). To test whether deer or hare limit hemlock regeneration, we studied >2000 hemlock seedlings inside and adjacent to experimental deer exclosures at 59 sites randomly distributed across hemlock and hemlock-component stands in northern Wisconsin and the western Upper Peninsula of Michigan in the United States. We monitored local deer and hare abundance, seedling growth, and seedling survival for 20 years. Two First Nations reservations showed lower deer density and greater survival and growth of unprotected seedlings than three national forests of the United States. Cohorts of hemlock seedlings protected from deer survived at a rate four times higher than those exposed to deer (59.3% vs. 15.0%) and shared a combined height 5.2 times greater. Hare densities significantly affected seedling survival only within exclosures. This extensive, long-term study identifies deer as the primary factor constraining hemlock regeneration in the region.