Patterns of tree replacement: canopy effects on understory pattern in hemlock - northern hardwood forests

Abstract

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability.

Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy.

Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.

Nomenclature follows Fernald (1950).

Acknowledgements: There are many whose help has contributed to this paper but Robert Whittaker played the primary advisory role throughout. The research described sprang from his ideas and his help was crucial in the performance of the research. I am also grateful to those who helped in the field work and the agencies and individuals who allowed access to field sites. I thank Peter Marks and Margaret Davis for their assistance and suggestions and Robert Peet and Steward Pickett for thorough and helpful review of the manuscript. The research was financed - in part - through a National Science Fellowship and by a Mclntyre-Stennis grant.