Modelling the influence of different harvesting methods on forest dynamics in the boreal mixedwoods of western Quebec, Canada
Introduction
Forest management emphasises an understanding of the effects of natural disturbance regimes on forest dynamics, which can be used as templates for forest management and help to reduce the gaps between managed and unmanaged forest landscapes (Grumbine, 1994, Bergeron and Harvey, 1997). The past several decades have witnessed a gradual evolution in forest ecology from static descriptions of forest community distributions along environmental gradients, such as climate and site characteristics (Whittaker, 1975), to the reconstruction of forest dynamics in relation to biotic and abiotic disturbances and ecosystem processes (Bergeron et al., 2002a). In the prolonged absence of large-scale disturbances such as wildfire, canopy gaps control stand dynamics (Kneeshaw and Bergeron, 1998). In this context, efforts have been devoted to developing forest gap models for various forest types where individual trees followed to simulate successional dynamics of forest ecosystems at different spatial scales (Larocque et al., 2011, Elzein et al., 2020). Such models could support forest management (Gauthier et al., 2009) to better identify and minimise the influence of silvicultural practices on managed forests, thereby maintaining forest dynamics that are comparable to what is observed in unmanaged stands (Seymour and Hunter, 1999). Gap models can assess the effects of different silvicultural treatments on stand dynamics over various temporal scales, whilst providing detailed information on the residual composition and structure of managed stands (Coates et al., 2003, Papaik et al., 2010). These models are highly flexible in terms of integrating conditions that are not easily or accurately obtained from empirical growth and yield tables. Implementing these models might also improve forest policy and planning (Landsberg, 2003, Taylor et al., 2009).
Mixedwoods are important sources of wood (Penner, 2008) and are the most diverse and productive forest types within the boreal biome of North America (Chen and Popadiouk, 2002). The dynamics of boreal mixedwoods are broadly regulated by natural disturbances, such as catastrophic wildfire (Payette, 1992, Bergeron, 2000) and insect outbreaks (Morin, 1994), together with anthropogenic disturbances, such as timber harvesting. Following a stand-replacing wildfire, the boreal mixedwoods of eastern Canada typically undergo replacement of shade-intolerant deciduous species by shade-tolerant conifer species (Bergeron, 2000). More specifically, stand development can be described as moving through three successional stages, from hardwood-dominated to mixedwoods, which culminate in conifer-dominated stands (Bergeron and Dubuc, 1989, Bergeron, 2000). The transitional development of the boreal mixedwoods is at the core of a three-cohort model. This tool was developed by Bergeron et al. (2002b), and guides forest management in emulating natural forest dynamics. The three-cohort model is an ecological concept that is based primarily upon stand composition and structure, which vary in both time and space depending upon factors, such as the availability of seed and regeneration sources, secondary disturbances, and the fire cycle (Bergeron, 2000, Bergeron et al., 2002b). For instance, stands experiencing frequent wildfires are dominated by species that are adapted to fire, such as trembling aspen (Populus tremuloides Michaux), paper birch (Betula papyrifera Marshall), black spruce (Picea mariana [Miller] BSP), and jack pine (Pinus banksiana Lambert),. (Bergeron, 2000, Kurkowski et al., 2008). Where longer fire return intervals prevail, abundance of shade-tolerant species increases, especially for balsam fir (Abies balsamea (L.) Miller), white spruce (Picea glauca [Moench] Voss), and eastern white cedar (Thuya occidentalis L.). Increasing proportions of fir and spruce leads to the increased probability of spruce budworm (Choristoneura fumiferana [Clemens]; SBW) outbreaks. Gap dynamics following canopy tree mortality modulate both species composition and structure of post-fire stands throughout stand development (de Römer et al., 2007, Kneeshaw et al., 2011). Species with different ecological requirements, growth rates, and modes of regeneration compete to fill gaps and recruit into the upper canopy (Taylor and Chen, 2011, Colford-Gilks et al., 2012). Nevertheless, species replacement dynamics within gaps is a complex process that depends upon various allogenic and autogenic factors, such as gap size, site conditions, disturbance regime, and stand composition and structure prior to disturbance (Bergeron, 2000, Colford-Gilks et al., 2012).
In recent decades, harvesting with variable tree retention and harvesting patterns (spatial configurations) has been proposed to reproduce the composition and structure of unmanaged stands of the boreal mixedwoods (Gauthier et al., 2009). Within the framework of forest ecosystem management (Grumbine, 1994, Christensen et al., 1996), clear-cuts and other even-aged harvesting systems are used to emulate stand-replacing wildfire, whereas selective and partial harvesting are used to emulate partial disturbances, such as insect outbreaks and gap dynamics (Bergeron et al., 2002b, Gauthier et al., 2009). Moreover, partial harvesting that retains greater residual forest structure is conducted to maintain stand attributes and ecosystem functions similar to those of mid- and late-successional stages by accelerating the replacement of shade-intolerant hardwoods with shade-tolerant conifers (Gauthier et al., 2009). Nevertheless, emulating stand dynamics by harvesting strongly depends upon the size and spatial distribution of gaps that are created as growing spaces for new tree cohorts (Harvey et al., 2002, Bose et al., 2015). For example, high-severity (dispersed or aggregated) harvesting with low tree retention creates large gaps and, accordingly, leads to greater variability in tree size classes through aspen recruitment (McCarthy, 2001), whilst promoting conifer sapling growth when these are present (Brais et al., 2013). Conversely, low-severity dispersed harvesting has been shown to maintain a greater number of structural attributes of even-aged stands (Bose et al., 2015). To consider not only the forest ecosystem management perspectives, but also timber supply and economic needs, the effects of harvesting practices on stand dynamics require long-term evaluation (Ruel et al., 2013). These effects could be considered positive if the growth and survival of residual trees and recruitment are successful over long periods of time, thereby ensuring the continuous supply of timber and ecosystem services (Thorpe and Thomas, 2007).
In this study, we used SORTIE-ND, which is a spatially explicit, individual-based stand dynamics model, to simulate 100 years successional dynamics of three stand types (deciduous, mixed deciduous, and mixed coniferous) in western Quebec, Canada. SORTIE-ND simulates small gaps and, thus, is a particularly suitable model for studying the dynamics of mixed stands related to partial disturbances (Coates et al., 2003). This model has been used to predict and assess the long-term effects of various silvicultural treatments, which could not be obtained by empirical studies (Vanderwel et al., 2011, Bose et al., 2015). Using the simulated outputs, (i) we evaluated the changes in stand dynamics, species growth and recruitment in the three stand types following silvicultural treatments with harvesting intensities ranging from partial harvesting (i.e., 30% and 60% basal area removal) to clear-cuts (93% basal area removal). We hypothesized that both pre-harvest stand composition and harvest intensity define post-harvest changes in managed stands, in which clear-cuts and high harvest intensities in deciduous and mixed deciduous stands would generally induce higher abundances of trembling aspen (H1). Conversely, we anticipated that low harvesting intensities, regardless of stand composition, would favour conifer recruitment to reproduce stand characteristics (in terms of composition and structure) similar to unharvested, natural stands (H2). For partial harvesting, (ii) we also assessed the contribution of tree removal patterns to changes in stand dynamics by comparing dispersed harvesting to aggregated (gap) harvesting with similar levels of basal area removal. We anticipated that for similar basal area removal rates, aggregated cuts would generate larger gaps that trigger recruitment of shade-intolerant species, whereas dispersed tree removal can successfully emulate natural gap dynamics that lead to gradual conifer dominance during stand succession (H3). The objective of the prevailing forest ecosystem management model that has been imposed the study region (Bergeron et al., 2002b, Harvey et al., 2002), is directed towards maintaining the composition and structure of the natural forest mosaic (Bergeron and Harvey, 1997, Brais et al., 2004). Therefore, (iii) we determined which harvesting treatment best emulates the natural dynamics of stands over 100 years of succession following harvesting.
Section snippets
Study area
The study was conducted at the Lake Duparquet Research and Teaching Forest (LDRTF; 79°19′W—79°30′W, 48°86′N—48°32′N), which is in the boreal mixedwoods of western Quebec, Canada. According to Environment Canada (https://climate.weather.gc.ca/climate_normals), the climate characterizing LDRTF is cold-continental, with mean annual precipitation of 985 mm and mean annual temperature of 1.0 °C. The research landscape lies within the balsam fir-white birch bioclimatic domain; a mixed composition of
Species dynamics in unharvested stands over 100 years
The changes in relative basal area of deciduous and coniferous species in unharvested stands (Cntrl) are presented in Table 3, for every 50 years and for the time-steps prior to and following SBW outbreaks over 100 years of simulation. As shown depending upon stand type (at time-step = 0), and SBW outbreaks (occurred at time-steps 28, 61, and 94) relative basal area of hardwoods and conifers changes over time.
Fig. 3 illustrates the natural successional dynamics of species in unharvested stands
Discussion
This study assessed the long-term effects of different harvesting methods, which differ in basal area removal intensities and harvesting patterns, on stand successional dynamics in the boreal mixedwoods. In support of our first hypothesis (H1), high-intensity harvesting and clear-cutting in aspen-dominated stands (deciduous and mixed deciduous) maintained or increased the hardwood component, similar to what has been observed following stand-replacing wildfires in the study region (Bergeron, 2000
Conclusions and management implications
Mixedwoods, which considered to be the most sustainable source of quality timber in the North American boreal forest (Chen and Popadiouk, 2002), are complex ecosystems that are compositionally and structurally influenced by numerous factors and processes (Bergeron, 2000, Chen and Popadiouk, 2002, Taylor and Chen, 2011). Maintenance of this complexity as well as biodiversity and other ecosystem services through partial harvesting has gained increasing interest from a forest management
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
We are grateful to Hubert Morin for his advice regarding adjustments to spruce budworm severity, to Claude-Michel Bouchard for his assistance in adjusting the harvesting scenarios, and to Philippe Marchand for his advice regarding statistical analyses. We also recognise the input of the following individuals, who contributed over the years to the development of the model for the study area: Albanie Leduc, Dave Coates, Danielle Charron, Sybille Haeussler, Christian Messier, Brian Harvey and Aron
Author contributions
All authors participated in study the design. K.M. and B.L. organised the required data and information for simulation scenarios. K.M. prepared parameter files for simulations, applied the SORTIE-ND model for predictions, finalised and analysed the constructed outputs. Y.B., B.L. and A.L. helped with the conceptualization and advised in the process of data simulation and analysis. K.M. prepared the original draft and B.L., A.L. and Y.B. reviewed and revised the original draft. All authors read
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