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Patterns of Plant Community Structure Within and Among Primary and Second-Growth Northern Hardwood Forest Stands

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Patterns of Plant Community Structure Within and Among Primary and Second-Growth Northern Hardwood Forest Stands Forest Ecology and Management 258 (2009) 2556–2568 Contents lists available at ScienceDirect Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco Patterns of plant community structure within and among primary and second-growth northern hardwood forest stands Julia I. Burton a,*, Eric K. Zenner b, Lee E. Frelich c, Meredith W. Cornett d a Department of Forest and Wildlife Ecology, University of Wisconsin – Madison, 1630 Linden Drive, Madison, WI 53706, United States b School of Forest Resources, The Pennsylvania State University, 305 Forest Resources Building, University Park, PA 16802, United States c Department of Forest Resources, University of Minnesota, 115 Green Hall, 1530 Cleveland Avenue North, St. Paul, MN 55108, United States d The Nature Conservancy in Minnesota, 394 Lake Avenue South, Duluth, MN 55802, United States ARTICLE INFO ABSTRACT Article history: Forest scientists advocate the use of natural disturbance-based forest management for restoring the Received 4 May 2009 characteristics of old-growth forests to younger second-growth northern hardwood stands. However, Received in revised form 4 August 2009 prescriptions rely upon studies that have (1) not spanned the full range of conditions and species Accepted 5 September 2009 assemblages, and (2) focused primarily on contrasting old-growth and mature second-growth stands at a single scale. To examine how the legacy of historical logging activities influences forest structure and Keywords: function, we compared and contrasted patterns of plant community structure within and among second- Forest structure growth and primary stands on the north shore of Lake Superior in Minnesota, USA — near the current Natural disturbance-based forest range limits of the dominant species, sugar maple (Acer saccharum). We expected second-growth stands management Old growth to be in younger developmental stages, and structurally less heterogeneous both within and among Primary forest stands. Furthermore, we expected those differences to be associated with patterns of plant community Second-growth forest composition and diversity. Understory vegetation Three of the four primary stands and one of the eight second-growth stands were in the old-growth stage of development. Yellow birch (Betula alleghaniensis) and conifers as a group (Thuja occidentalis, Picea glauca and Abies balsamea) were more abundant, and yellow birch was more variable, within primary stands than second-growth stands. The volume and heterogeneity of coarse woody debris in intermediate decay- and size classes was also greater within and among primary stands relative to second-growth stands. While mean subplot richness of overstory tree species was greater in primary stands, mean quadrat richness, and rates of species accumulation for forest herbs as well as total herbaceous cover, and graminoid cover were greater in second-growth stands. Furthermore, total basal area (BA), the BA of conifer species, the density of yellow birch trees, understory vegetation and light transmittance were more variable among second-growth stands. At the multivariate level, primary stands were distinguished from second-growth stands not by differences in stand structure, but by a greater abundance of yellow birch and conifer species in the canopy, which was also related to O-horizon depth and understory plant species composition and structure. Differences in community structure between primary and second-growth stands may have resulted from the original cutover as well as high- grade logging, which together may have disrupted the mechanisms that maintain populations of important co-dominant tree species and associated understory plant communities in northern hardwood stands. ß 2009 Elsevier B.V. All rights reserved. 1. Introduction natural disturbance, historical landscape and stand structures, and historical ranges of variation (Attiwill, 1994; Franklin et al., 2002; The potential degradation of forest ecosystem goods and Seymour et al., 2002; Drever et al., 2006). For northern hardwood services and loss of biodiversity due to extensive logging has forests, specific efforts include those that attempt to accelerate generated an impetus for developing silvicultural systems that succession in younger second-growth stands by imposing the sustain biodiversity and productivity by mimicking patterns of structures that characterize historical landscapes and old-growth stands (Lorimer and Frelich, 1994; Keeton, 2006). However, previous studies have focused primarily on average conditions * Corresponding author. Tel.: +1 608 265 6321; fax: +1 608 262 9922. or a single scale and have not included stands near the range limits E-mail address: [email protected] (J.I. Burton). of north-temperate tree species; thus, the legacy of historical 0378-1127/$ – see front matter ß 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2009.09.012 J.I. Burton et al. / Forest Ecology and Management 258 (2009) 2556–2568 2557 logging events on the composition, structure and function of light transmittance and understory plant communities, which are second-growth stands is not fully understood. Because interactions more fine-grained and patchy in old-growth stands (Crow et al., among structure, composition and variability at multiple scales 2002; Scheller and Mladenoff, 2002). Although Duffy and Meier may be important for maintaining the resilience of northern (1992) report devastating effects of logging on the richness and hardwood forest ecosystems, restoration strategies based solely on cover of forest herbs in the southern Appalachians, evidence from average conditions and successional processes may serve only to the Lake States suggests that timber harvesting may increase reinforce current differences between primary stands that were herbaceous species richness and cover due to the immigration of never logged and second-growth stands. weedy and non-native species (Metzger and Schultz, 1984; Most of the northern hardwood forests in the upper Great Lakes Scheller and Mladenoff, 2002). region regenerated after extensive logging, known as the cutover, Here we investigate differences in community structure and that occurred at the turn of the 20th century (Zon, 1925). Logging heterogeneity between primary forests with a history of natural at this time ranged from the selective removal of economically disturbance and regeneration, and second-growth forests that valuable tree species to heavy partial cuts and clear-cutting were logged during the cutover and subsequently high-graded in oftentimes followed by repeated uncontrolled slash fires (Stearns, northeastern Minnesota. We expected second-growth stands to be 1997). This series of events in particular reduced the area of in younger developmental stages, and structurally less hetero- unharvested, primary stands and stands in the old-growth stage of geneous both within and among stands. Furthermore, we expected development in Minnesota to 0.2% and 2% of the pre-Euro- those differences to be associated with patterns of diversity and American extent, respectively (Frelich, 1995). plant community composition and structure. By comparing and While the cutover greatly reduced the extent of old growth on contrasting primary and second-growth stands with respect to the landscape, it may have also initiated more persistent changes structure, species composition and diversity at multiple scales, we in community structure in regenerating second-growth stands. In can characterize the persistent effects of natural and anthropo- northern hardwoods of the upper Great Lakes region, the natural genic disturbance regimes to inform guidelines for managing and disturbance regime resulted in a shifting mosaic of stands in restoring structure and composition to second-growth stands. different stages of development dominated by old growth (Frelich and Lorimer, 1991). Within stands, the volume of coarse woody 2. Study area debris (CWD), snag basal area (BA), total area in gaps and average gap size increase in later stages of development (Tyrrell and Crow, Sugar maple-dominated northern hardwood forests are dis- 1994; Dahir and Lorimer, 1996). Studies from the Pacific North- tributed along the ridge tops on the north shore of Lake Superior west show that the range of structural variation and structural within the transition zone between north-temperate and south- heterogeneity indeed increases with stand age (Spies and Franklin, ern-boreal forests. Elevations range from 200 to 700 m and the 1988; Zenner, 2004). Furthermore, compared to conventional topography is gently rolling to steep. Lake Superior moderates the systems of forest harvesting, natural disturbances can also lead to climate, which is cold-temperate continental, with a mean more complex structures and greater residual heterogeneity growing season length of 104–168 frost-free days (base tempera- (Hanson and Lorimer, 2007). ture = 0 8C), mean annual temperature of 4.72 8C, and mean annual Heterogeneity of important community attributes can be both precipitation of 77.50 cm with 150.4 cm of snowfall (1971–2000, affected by disturbance, and influence the response of forests to Midwest Regional Climate Center, http://mcc.sws.uiuc.edu). disturbance (Fraterrigo and Rusak, 2008). In northern hardwood This study is part of the pre-treatment phase of a larger project forests, sugar maple trees can form dense layers of advance that examines methods of restoring the composition and structure regeneration that excludes light-seeded and mid-tolerant tree of
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