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Forty-Two Years of Succession Following Strip Clearcutting in a Northern Hardwoods Forest in Northwestern Massachusetts Taber D

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Forty-Two Years of Succession Following Strip Clearcutting in a Northern Hardwoods Forest in Northwestern Massachusetts Taber D Forest Ecology and Management 182 (2003) 285–301 Forty-two years of succession following strip clearcutting in a northern hardwoods forest in northwestern Massachusetts Taber D. Allison*, Henry W. Art, Frank E. Cunningham1, Rebecca Teed2 Center for Environmental Studies, Williams College, Williamstown, MA 01267, USA Received 20 July 2002; received in revised form 26 August 2002; accepted 19 January 2003 Abstract We investigated the effects of strip width, slope position, and soil scarification in a split–split plot design on the regeneration of northern hardwoods in northwestern Massachusetts. Whole plots of 20 and 40 m in width were cut in 1954 in a second growth forest dominated by Betula papyrifera. Slope position and soil scarification were the split and split–split plot treatments, respectively. We measured height for all tree species present in randomly located 4 m2 plots beginning in 1955 and at irregular intervals over the following 42-year period. We measured all trees in the cut strips in 1996. Prunus pensylvanica was the dominant species initially, but had nearly disappeared from the cut strips by 1996. Soil scarification significantly increased initial establishment of B. papyrifera, but density and basal area of this species did not differ by soil treatment in 1996. Tree composition in cut strips was weakly correlated with soil moisture, soil scarification, and initial tree density immediately following cutting, but high spatial variation in species composition and low replication made it difficult to detect any significant correlations among the distribution and abundance of different species and selected environmental variables. The canopy of the cut strips is even-aged; establishment of most canopy trees occurred within 5 years following cutting. A comparison of successional trends in adjacent uncut strips with the trends in the cut strips indicates that cutting has altered the sequence of successional changes in forest composition increasing the abundance of some species that were of low importance prior to cutting. In 1996, Acer rubrum and A. saccharum are replacing B. papyrifera in the canopy of the uncut strips. The canopy of the cut strips consists of a diverse and spatially varying mixture of intermediate hardwoods including Quercus rubra, Fraxinus americana, Betula lenta, Acer rubrum, B. papyrifera, and an understory of late successional hardwoods. # 2003 Elsevier Science B.V. All rights reserved. Keywords: Clearcutting; Forest succession; Betula papyrifera; Tree age; Tree height; Prunus pensylvanica 1. Introduction The study of forest succession is one of the oldest * Corresponding author. Present address: Massachusetts Audu- and most interesting topics in forest ecology (e.g. bon Society, 208 South Great Road, Lincoln, MA 01773, USA. Cowles, 1899; Thoreau, 1906). Much of what we have E-mail address: [email protected] (T.D. Allison). 1 Deceased. learned about the pattern of forest succession and 2 University of Maryland, Unit 29216, APO, AE 09102, Maryland, the responses of species to disturbance, especially USA. in the northeastern United States, comes from studies 0378-1127/$ – see front matter # 2003 Elsevier Science B.V. All rights reserved. doi:10.1016/S0378-1127(03)00066-5 286 T.D. Allison et al. / Forest Ecology and Management 182 (2003) 285–301 of forest regeneration following clearcutting (e.g. In addition to examining the effects of the pre- Bormann and Likens, 1979; Marquis, 1967; Leak, scribed treatments on paper birch regeneration we 1991; McClure and Lee, 1993). Studies of logged were interested in the effect of these treatments and forests often provide the longest sequence of forest other non-manipulated factors in the physical envir- change on individual sites as many of these studies onment on forest composition 42 years later. Speci- date from the 1930s and 1940s. Clearcutting studies fically, to what extent could the current composition often are established with controlled, replicated (1996) of the cut forest be predicted from the compo- experimental designs that allow the statistical analysis sition during the establishment phase immediately of forest response under different environmental con- following cutting? It was difficult to determine a ditions. This avoids the problems of pseudoreplication priori, all possible initial conditions that might be or the uncontrolled and confounded comparisons of important. Some of the initial conditions in this study stands compiled along a chronosequence. Data usually were determined by the specific treatments employed are collected on permanent plots established immedi- by the USFS, but disentangling the importance of ately after cutting, and, occasionally, forest composi- other non-controlled factors would readily have tion is measured prior to cutting. degenerated into a fishing expedition with no statis- In this paper we report on the initial response of tical power. To avoid this we asked specific questions strips cut in 1954 in the Hopkins Memorial Forest including whether variation in the abundance of paper (HMF) in northwestern Massachusetts and the subse- birch in 1996 could be predicted from the abundance quent changes in species’ composition and dominance of the initial establishment of paper birch? We also over a 42-year period. Cunningham initiated the clear- asked whether soil moisture or initial density of pin cutting experiment under the auspices of the United cherry (Prunus pensylvanica L.) following cutting States Forest Service (USFS), which owned and man- affected initial stand composition and forest composi- aged the HMF from 1935 to 1968. The experiment’s tion 42 years later. primary objective was to examine the influence of strip width and seedbed scarification on the natural regen- eration and early development of paper birch (Betula 2. Methods papyrifera Marsh.) (Cunningham, unpublished manu- script). Clearcutting in strips leaves uncut intervening 2.1. Description of study area strips as a source of seed and shade. Exposure of mineral soil was thought to enhance establishment of light- The study area is located in the HMF now owned by seeded tree species, such as paper birch, and had sub- Williams College, Williamstown, Massachusetts. The sequently been shown to enhance initial establishment HMF has topography typical of the western New of paper and yellow birch (Hutnik and Cunningham, England Highlands. The study area is situated below 1961; Marquis et al., 1964; Marquis, 1965). Because of the Taconic Crest at an elevation ranging from 380 to the topography of the study area, the effect of slope 450 m above sea level. The aspect of the study site is position on paper birch regeneration was also examined. northeast; the slope varies from 0 to 47% and averages Data on abundance of tree species after cutting were 25%. The bedrock of the site is primarily schistose and collected by the USFS from permanent 4 m2 plots in phyllitic rock characteristic of the Taconic Range, and 1955, 1957, 1958, and 1962 at which time the project the soils are primarily thin, stony acid loams (Art and was temporarily abandoned. Allison and Art relocated Dethier, 1986; Scanu, 1988). the plots in 1974 and made several measurements on Just prior to cutting, the study site was an even-aged species composition in 1974, 1975, 1989 (measured stand dominated by paper birch (50% density and by Teed), and 1996. The length of observation of this basal area of trees over 11.7 cm diameter at breast experiment allowed us to examine not only the initial height (DBH)) and red maple (Acer rubrum L.) (25% effect of treatment on paper birch regeneration, and of the basal area and 28% of the density). Sugar maple other tree species, but also whether or not the effect of (A. saccharum Marsh.) was the next most impor- this treatment persisted over the 42-year period of tant species at 9% of the basal area and 11% of the measurement. density. The remainder of the stand consisted of small T.D. Allison et al. / Forest Ecology and Management 182 (2003) 285–301 287 amounts of black birch (Betula lenta L.), yellow birch the HMF. No logging or agricultural activity took (B. alleghaniensis Britton), red oak (Quercus rubra L.), place in the study area after this time (Rosenburg, white ash (Fraxinus americana L.), American beech personal communication). Tree cores removed by (Fagus grandifolia L.), aspen (Populus spp.), Eastern the USFS in 1936, primarily from paper birch and hophornbeam (Ostrya virginiana K. Koch.), and red maple growing in nearby stands, showed tree ages striped maple (A. pensylvanicum L.). at breast height ranging between 40 and 83 years; Massachusetts tax records indicate that the study mean and median ages were 54 and 50 years, res- area was used for pasture and cultivation until the pectively. Permanent plot studies elsewhere in mid-1860s. The study area was forested in 1901 the HMF have shown that paper birch dominates when Amos Lawrence Hopkins purchased the early to mid-successional stands on lands that had land and incorporated it into the property that became been used for cultivation immediately prior to their Fig. 1. Field layout of experimental design showing orientation of cut and uncut strips and 16 split–split plots (experimental units). 288 T.D. Allison et al. / Forest Ecology and Management 182 (2003) 285–301 abandonment from agriculture (Art, 1974; Art and amount of scarification. Plots were then assigned to Dethier, 1986). classes based on percent scarification—the scale ran- ged from 0 to 100% at 25% intervals. 2.2. Experimental design 2.3. Data collection In 1952, Cunningham and colleagues surveyed a por- tion of the selected stand, measuring 200 and 240 m. Cunningham and USFS colleagues counted and This area was subdivided as follows: four cut strips were measured the height of all tree stems in the ninety- separated from each other by intervening uncut strips six 4 m2 plots in the fall of 1955, 1957, 1958, and 1962; each measuring 20 m in width (see Fig. 1).
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