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March 2009
Mark Swanson
Click here to view the PDF version of Working Paper 10 AcknowledgementsThis report was made possible by a grant from the Olympic Natural Resources Center with financial support from the U.S.F.S. Pacific Northwest Experiment Station. An early version of the literature review was funded by the Washington State Department of Natural Resources. The findings in the report hopefully mirror the results of the many pioneering research studies referenced in the literature review. Any opinions, findings, conclusions or recommendations in this report are those of the authors and do not necessarily reflect the views of contributing entities.
AbstractIn this literature review we identified over 700 articles following the Washington Forest Landscape Management Project, (WFLMP), which provided one of the early attempts to understand the implications of innovative silviculture and landscape management on conservation values. We enumerate many of the alternative treatments and important elements starting with the WFLMP while noting the several research pathways that developed from that work. We do not attempt to highlight the impact of each referenced article but follow the several directions research has taken and rely on the emphasis provided through several publications and conferences that have attempted to provide a comprehensive summary and review of the literature. Keywords: Landscape management; biodiversity; habitat suitability; wildlife models; innovative silviculture; spatial forest planning; endangered species protection; harvest scheduling; optimization; meta-heuristic; spatial and temporal landscape patterns; spatial explicit wildlife model; multi-species approach; GIS.
Table of Contents
Background and IntroductionIn June 1988, with intensifying public debate about forest management and ecosystem protection, Brian Boyle, Commissioner of Public Lands, established the Commission on Old Growth Alternatives for Washington’s Forest Trust Lands. The Commission brought together 32 diverse “stakeholder group” representatives. Their mission was to develop consensus recommendations for the Washington Department of Natural Resources (DNR) on how to balance the goals of providing revenue for education, protecting the biological diversity of the forest environment, and supporting local timber-dependent communities. Commission recommendations included the establishment of the 264,000-acre Olympic Experimental State Forest (OESF) on the western portion of the Olympic Peninsula as a commercial forest within which there would be a special opportunity to research harvest and regeneration methods to enhance habitat characteristics and commodities production. To focus and coordinate the long-term research program for the OESF, the Commission recommended creation of the Olympic Natural Resources Center, or ONRC (Commission on Old Growth Alternatives 1989). These investments in experimental research capacity reflected a broader trend in the field of forest management, in which the desirability of maintaining the inherent complexity and array of functions of natural forest ecosystems is recognized. Prominent Washington forest scientists offered papers suggesting that modifications to commercial forest management could achieve ecological benefits and accelerate development of desired old forest habitat structures while generating forest products and economic returns (Franklin 1989, Oliver 1992). In 1993, the Washington Forest Landscape Management Project (WFLMP), funded by the U.S. Congress and facilitated by the DNR (Carey et al. 1996), created for the first time a broad multi-disciplinary research group with the objective of determining the advantages of simultaneously managing forests for multiple species of threatened, endangered, and sensitive wildlife over large landscapes and across different ownerships including economic considerations. It was hoped that innovative approaches to forest management could integrate species habitat enhancement with harvest of forest resources to maximize environmental benefits and reduce the costs of species conservation on society. Many ideas about the impacts of changing forest structures on species conservation were discussed and evaluated by the study team. The WFLMP provided the first interdisciplinary opportunity to evaluate the potential costs and benefits of intentionally managing forest stands to restore old forest functionality following harvests and other anthropogenic disturbance. The management treatment strategies developed by the WFLMP scientific team became known as biodiversity pathways, or ‘biopathways’, designed to intentionally produce old forest habitat conditions through active management at a much lower cost than no-action or passive alternatives. From a literature review perspective, the project considered 150 then-current publications such that projects after 1996 generally had the benefit of the integrative work provided by the project. The project produced many publications (e.g., Carey, Eliot et al. 1996, Carey, Thysell et al.1996, 1999; Lippke et al. 1996) and began a decade of additional scientific research into the effectiveness of biopathway treatment regimes as compared to no-harvest reserves and commercial or longer than commercial rotations. In this literature review we identified over 700 articles following the WFLMP with implications for innovative silviculture and landscape management. We enumerate many of the alternative treatments and important elements starting with the WFLMP while noting the several research pathways that developed from that work. We do not attempt to highlight the impact of each referenced article, as this approach would be too long for a useable synthesis. Rather, we follow the several directions silvicultural and forest landscape research has taken, and we rely on the emphasis provided through several publications and conferences that have attempted to provide a comprehensive summary and review of the literature. Biodiversity Management and Protection PathwaysThe distinguishing features of “biodiversity pathways” developed in the WFLMP included retention of biological legacies at harvest, plantation and natural regeneration for mixed species forests, pre-commercial thinning to quickly bypass the stage of competitive exclusion in dense young stands, thinning at variable densities in maturing stands, extending final harvest rotation lengths through multiple entry thinning, and creation of snags and downed woody debris. The range of options examined has been largely oriented toward thinning earlier or later while retaining less or more trees over time with consideration for retention of legacy trees, stumps and debris. Riparian buffer management incorporated similar thinning treatments with extra attention to stream bank stability, sufficient vegetative and overstory retention for shade, and recruitment of large woody debris (LWD). Direct measures of forest ecosystem health were largely related to the capacity to support vertebrate diversity, forest floor function for mammals, ecological productivity based on the abundance of arboreal rodents as a critical element of the food chain and production of large vertebrates. These functions/proxies were developed as indices to estimate the ecological impacts of alternative management approaches. The biodiversity pathways approach emphasizes changes in forest structure based on growth model simulations, which were analyzed statistically and spatially. The structural composition changes were shown to restore forest stand structure distributions more like those present before European settlement, albeit at a cost to landowners. The spatial patterns were less revealing since the size of treatment areas was dictated by prior treatments and stand boundaries and regulatory or operational limits on the treatment scale. While biodiversity pathways on the entire landscape produced what were considered to be better spatial distributions for late seral forests, there were few interior species that would benefit. The spotted owl was believed to be more negatively affected by aggregated cuttings than dispersed (Carey et al. 1992, Carey and Peeler 1995). Other studies found no effects from interior forest management alterations and further concluded that no species of concern were “associated” with (benefited from) “competitive exclusion” of dense closed-canopy structures (Hansen et al. 1993). The conclusion that no species benefited from competitive exclusion structure, with most species negatively affected, has provided strong motivation for biopathway thinning in forests with a legacy of commercial management. The dominant forest structure class associated with commercial rotations (or no-action alternatives in previously-harvested forests) following regeneration strategies is competitive exclusion. This forest condition can potentially last for much more than a century following full stocking regeneration if stem densities are not reduced by thinning treatments or disturbance. As a result, many studies have emphasized the need to reduce stem densities and avoid or reduce the prevalence of closed-canopy competitive exclusion as important to achievement of biodiversity goals and old forest habitat objectives (Oliver 1992, Poage 2000, Poage and Tappeiner 2002, Acker et al. 1998, Agee 1991, Carey et al. 1999, Hayes et al. 1997, Garman et al. 2003, Tappeiner et al. 1997, Bailey and Tappeiner 1998). Functional equivalents of the biopathways approach have been developed for forests in other regions, such as the broadleaf forests of New England. At a spatial scale of the individual stand, the use of rotated-sigmoid diameter distributions in combination with specific strategies to enhance structural richness (e.g., snag creation, crown release of vigorous trees, etc.) can further contribute to these objectives (Keeton 2006, Kenefic and Nyland 2000). Lindenmayer and McCarthy (2002) propose similar alterations to intensive forest management in southeastern Australia to conserve fauna dependent on old, hollow trees and intact forest landscapes. Application of these recommendations in landscapes will validate degrees of effectiveness over the long time scales necessary to conduct research in forested landscapes. Measuring the Biological and Economic Success of PathwaysDevelopment of quantifiable and statistically robust measures of ecosystem health with which to assess the comparative success of alternative management treatments has been a challenge for researchers. Early studies were constrained by limited empirical stand data and with limited statistical analysis enriched by expert opinion. Gehringer (2006) developed a non-parametric statistical target of old forest conditions by analyzing actual old and previously not harvested forest inventory data and determining the attributes that most reliably discriminate those stands from all others. Following this analysis Gehringer was able to create management targets to test for development of old forest characteristics within a confidence interval representative of an old forest range of variability for Western Washington. Gehringer found that trees per acre, diameter and height were the dominant metrics of interest that differentiated old forest characteristics. When simulated treatments and growth for forest stands are modeled forward in time then the point at which stand characteristics resemble those of the target older forests can be identified. Two important performance metrics for assessing comparative old forest treatment options emerge: which treatments create desired conditions soonest and which treatment regimes most successfully extend the duration of time in the old forest target. Other variables thought to be important to habitat such as coarse woody debris (CWD) were characterized by such large variance that they did not provide a discriminating function for old forests. However, CWD may be a discriminator of better habitat for some critical species (Bunnell et al. 1999), and can be used as a fine filter discriminator to identify selected species habitats within the broader class of old forests (Zobrist and Gehringer 2004). Other metrics include an index of old growth (Acker et al. 1998), which has been used to quantify the rate of structural development in unmanaged Douglas-fir stands (Larson et al. 2008). Stand-level diameter distributions are also useful for quantifying differences between structurally simple young stands and complex, multi-cohort late-successional stands (Zenner 2005), especially under chronic disturbance regimes (D’Amato et al. 2008). Simulated treatment activities such as thinning harvests can be used to forecast potential log yields and economic returns. Old forest structure targets and simulated economic returns provide a means for identifying least-cost pathways for achievement of biological forest targets. Similar methodologies for statistical discrimination between forest conditions have been explored by Lundquist (Lundquist & Beatty 1998, Lundquist & Lidner 2000). et al. (2005a) used the Gehringer procedure to determine the most economic pathways to reach old forest structure targets. The method was to inform forest management options within riparian zones in the PNW Douglas-fir zone as well as to aid development of legacy structures in the U.S. Southeast, a project supported by the National Commission on Science for Sustainable Forestry, NCSSF (Zobrist et al. 2005b). A similar approach was used to investigate management options for old forests and spotted owl habitat on the DNR Olympic Experimental State Forest (Lippke, et al. 2007). The performance metrics for a zone-specific target of old forests with their discriminating structural differences were used in all these studies to identify which treatments most successfully reach the structural target objective at the lowest economic costs. Habitat Suitability ModelsOther forest structure research efforts have targeted the development of various habitat suitability measures for individual species, which are logically more sensitive to detailed segmentation of forest structure classes. The U.S. Fish and Wildlife Service developed a habitat evaluation procedure (HEP) (USDI 1980) based on characterizing the forest in a number of discrete conditions in order to provide the inputs for habitat suitability models. These methods were automated for a study in Western Washington that used a stand-level simulation model to conduct habitat suitability tests, demonstrating repeatability in evaluation methods (Ceder et al. 2002a, Marzluff et al. 2002). Johnson and O’Neil (2001) developed a wildlife-habitat model that covers entire vertebrate communities in Washington and Oregon. This habitat model recognizes three factors: wildlife habitats = wildlife cover type(s) plus structural condition(s) plus habitat element(s). Wildlife cover types refer to vegetation classifications that were determined based on similarity of wildlife use. Structural conditions describe the forest structure at the stand level. There are a total of 26 structural conditions defined by tree size, percent canopy cover, and the number of canopy layers. Finer scale habitat features are habitat elements such as downed logs and snags. The resulting Johnson and O’Neil wildlife-habitat matrix associates species types with 26 habitat structural conditions further defined by three habitat qualities: closely-associated, generally-associated, and present. Although habitat elements are considered important components in habitat models, elements are too variable and species-specific to generalize in the Johnson and O’Neil’s wildlife-habitat model although potentially important at the site or forest type level. Habitat suitability measures are useful tools for analysis but currently lack sufficient data to be as statistically robust as the Gehringer approach. As more data is collected on species-specific habitats, statistical links between habitat suitability and forest structure class will logically improve. However, another problem with classification systems is that habitat definitions tend to be limited to “in or out” rather than probabilistic gradients and consequently can exhibit knife-edge properties relative to management simulations obscuring performance at the margin (University of Washington College of Forest Resources 2007). Models integrating a non-binary perspective on habitat utilization are more likely to offer insights into landscape and stand influence on vertebrates (Marzluff et al. 2004, Folliard et al. 2000). The former focus in landscape ecology on the patch-mosaic model of landscape structure is being enriched by an understanding that many phenomena occur as gradients in landscapes (McGarigal and Cushman 2005). This is reflected by the use of logistic regression models parameterized at multiple scales to assess the importance of landscape context (including the abundance of various structural classes of forest) in determining organism abundance or viability (e.g., White et al. 2005, Daw and DeStefano 2001). These models can then be used to create maps of probability gradients, assess actual landscapes for areas of high-quality habitat, or plan restoration treatments that will create appropriate combinations of landscape elements. Variable Densities and Variable RetentionResearch on the ecological impacts of biopathways silvicultural interventions is still in formative stages. Individual tree growth in variable density units tends to vary as a function of proximity to gaps and whether the tree is in the thinned matrix of the unit (Roberts and Harrington 2008). While variable densities were considered important in the early biodiversity pathway work (Carey et al. 1996), natural variation as well as variation induced from equipment impacts made it difficult to establish useful measures of density within otherwise uniform stand structures. Thinning clearly favors the development of shade-tolerant trees in the understory of stands in the competitive exclusion/biomass accumulation stages (Chan et al. 2006). Underplanting may have to be employed to accelerate this process, especially when seed sources for shade-tolerant conifers are distant (Beach and Halpern 2001). A meta-analysis of thinning impacts on understory vegetation (Wilson and Puettmann 2007) found that variable-density thinning increased spatial variability in understory communities, but effects on richness varied from case study to case study, often as a function of past site management. Concerns regarding potential for wind damage related to larger openings associated with variable-density thinning were allayed by Roberts et al. (2007), who found no difference between unthinned controls and treatment plots. Certain taxa may benefit from variable-density thinning, such as litter invertebrates (Schowalter et al. 2003). Many of these results await confirmation over longer time scales and application in other forest types, but the results of current research favor the concept that spatially variable thinning is an effective, ecologically-based method for enhancing complexity and diversity in young forest stands. Disturbance in forest ecosystems, by definition, kill or remove trees and other organisms (Pickett and White 1985). Most natural disturbance processes, however, tend to leave a variety of living or dead structures as biological legacies (Franklin et al. 2000). These legacies promote ecosystem recovery (e.g., Keeton 2000) and the persistence of a wide variety of organisms at multiple spatial scales. It has been proposed that retaining structures at harvest can capture some of the functional benefits of biological legacies within the context of the managed forest environment (Franklin et al. 1997, Mitchell and Beese 2002). Variable retention has been shown to be effective for the conservation of understory plant communities (Nelson and Halpern 2005), water quality (MacDonald et al. 2003), lichen communities (Esseen et al. 1996), and other values. Ectomycorrhizal fungi have been found to be more abundant in the vicinity of retained individual trees (Cline et al. 2005), with proximity to patches is relatively more important than the size of the patch itself (Jones et al. 2008). A meta-analysis of research on green tree retention (variable retention) demonstrated the utility of the approach for maintaining organisms from several important functional groups, such as dispersal-limited lichens, birds, and ectomycorrhizal fungi (Rosenvald and Lohmus 2008). Survival and growth of retained trees is a major concern when applying variable retention harvest strategies. Survival is often influenced by tree wounding during the harvest or proximity to skid trails (Thorpe et al. 2008), and wind firmness is a major consideration when selecting trees for retention. Healthy, uninjured trees (even in older age classes), however, typically respond well to the release provided by a variable retention harvest event (Latham and Tappeiner 2002, Bebber et al. 2004). Experience in wet sclerophyll forests of Tasmania demonstrates that variable retention may aid in regeneration of species-rich forest communities by reducing dispersal distances (Tabor et al. 2007), and that while windthrow may damage retained trees, old-growth individuals tend to be relatively windfirm (Neyland 2004). Additionally, the retention of undisturbed forest aggregates in Tasmanian forest has been shown to favor amphibians (Lauck et al. 2008) and beetles (Baker et al. 2007). Experimental On-the-Ground TreatmentsMost early work on innovative silviculture to promote conservation values relied on simulations as the historical data on past treatments was generally lacking. This prompted a series of experimental studies, collectively known as the Capitol Forest Study, to apply different treatments on the ground in sample plots to estimate cost and environmental benefit over time (Curtis et al. 2004). Located on public land in Washington State, this is an exemplary experimental effort aimed at clarifying the impacts of various silvicultural treatments on the structure and composition of Douglas-fir stands in the competitive exclusion stage. These studies, unlike many from the ecological literature, provide useful operational cost information; however, the impacts on ecological measures are only now beginning to have enough maturation time to be indicative of the future. Future monitoring will help to determine comparative environmental effectiveness. Silvicultural research to assess the efficacy of different timber harvest regimes in maintaining habitat values or ecosystem processes have shown the value of various modifications of traditional timber harvesting. The DEMO study (Aubry et al. 2004) has produced a number of studies on stand-level response to various forms of variable retention, some of which have been discussed in the section on variable retention. Silvicultural experimentation has not been limited to the creation of later-successional structural conditions, however. An excellent example may be found in the oak forests of the Puget Sound region in western Washington, where partial and full crown release prescriptions have been shown to be effective in restoring vigor to overgrown Oregon white oak (Devine and Harrington 2006). At Fort Lewis in Western Washington, Churchill (2005) showed that on dry-sites Douglas-fir can grow in the understory and maintain its release potential for at least 20 years under moderate overstory stocking levels (30-55% full stocking). His results indicate that by combining elements of shelterwood, group selection, and single tree selection systems, multi-cohort, structurally complex stands can be created and maintained in a shifting mosaic of patches while also producing significant wood volume over time. Historic stand conditions cannot be duplicated rapidly following either stand replacement fires or high volume timber harvests. Dry pine forests present a further set of restoration challenges, since these forest types have experienced exploitative timber extraction, fire suppression, and agricultural conversion. Everett et al. (2007) provide a unique analysis of history based on stand reconstructions over 150 years. They concluded that in pre-settlement forests landscapes varied greatly with many structural stand types reflecting different stages of recovery from multiple disturbances of varying intensity and frequency but had a well-represented tree-understory with very few dead and down fuels. Following euro settlement, decreased fire effects allowed existing understory to continue development, which increased stand density and the proportion of shade tolerant species. Under fire suppression, landscape diversity of forest structure has been lost with a decline in early succession stages and an increase in old forest structure (Everett et al. 2008). Stands are currently transitioning from high post fire suppression tree densities to less dense stands as insect and pathogens thin stands from above and below. This stand thinning process has increased amounts of dead and down wood, previously maintained at low levels by frequent surface fires. Although the dry forest landscape has always operated under a mixed fire regime, the proportion of landscape subject to high severity fires has increased over the last several decades reflecting unsustainable conditions (see also Hessburg et al. 2005). Both forest structure and disturbance regimes are evolving. Restored landscape and forest structures will need to be sustainable under disturbance regimes that differ significantly from euro settlement conditions. Restoration objectives that include thinning and prescribed fire to emulate the inherent disturbance regimes of the area will also need to reflect post settlement socio-economic expectations. Research in the forests of the Southeast has shown that it is possible to restore old-growth longleaf and loblolly stands through modified group selection and single-tree selection, the application of prescribed fire, and the introduction of spatial heterogeneity (Bragg et al. 2008, Brockway et al. 2002). Experimental silviculture in the Ouachita Mountains of Arkansas has focused on restoring shortleaf pine stands formerly maintained by fire (Guldin 2004). Research in ponderosa pine ecosystems in the Black Hills (Shepperd and Battaglia 2002), the interior Pacific Northwest (Youngblood et al. 2006), and the Southwest (Allen et al. 2002) indicate that restoration of coarse woody debris, retention of large-diameter pines, reintroduction of fire, and creation of spatial complexity (gaps, clusters of trees, and randomness at various spatial scales) are important elements that diverge from traditional tenets of silvicultural practice. Arno and Fiedler (2005) examine long research histories from a number of western forest types where carefully applied silvicultural treatments and the resumption of historic fire regimes lead to a host of ecological, economic, and social benefits. Temperate systems from around the world offer similar experiences. For example, silvicultural trials at Warra, Tasmania, in eucalyptus-dominated stands show benefits to creation of ecological complexity within stands and restoration of elements of natural disturbance regimes (Neyland 2004). Stand Variability Over Greater Spatial ScalesLindenmayer and Franklin (2002) focused on enhancing biodiversity with the perspective that the scale requirements can not be met by partitioning off some lands as reserves while other lands are used for intensive timber production. They characterized a checklist for achievement of forest biodiversity conservation across a landscape based upon a matrix of conditions needed. A number of cases studies were summarized to highlight the contribution of a variety of landscape characteristics. Important areas: The Lindenmayer and Franklin checklist begins by identifying important areas that need to be protected. Important areas include aquatic systems such as stream networks and wetlands, wildlife corridors, specialized habitats such as cave and thermal protection, biological hotspots, and remnants of late successional and disturbance refugia such as forest areas with no prior history of harvest. Culturally and socially important areas are identified for protection as well. These areas are mid-spatial scale and are to be given special consideration before making a forest management plan. Aquatic ecosystems: Protecting aquatic ecosystems are given special attention as uniquely important to biodiversity in a forested landscape (Naiman et al. 1993, Naiman et al. 2000, Brinson and Verhoeven 1999, Calhoun 1999). Sixty percent of 480 wildlife species were observed in riparian forest in Washington State (Raedeke 1988). Important roles of riparian forests are 1) light and water temperature control, 2) organic matter inputs, 3) bank protection, and 4) source of large woody debris. Selection of the appropriate widths and lengths for streamside corridors should be based on the objectives of the buffer and the spatial pattern of relevant influences (Lindenmayer and Franklin 2002). Recruitment of large woody debris is a good example. Relatively narrow buffers (e.g. 10 meters or less) have generally not been considered adequate to address the biological and physical interactions between riparian forests and streams. The Forest Ecosystem Management Team (FEMAT 1993) created one standard that uses a site-potential tree height to define protection areas (the more productive the site; the wider the buffer). Two tree heights for fish-bearing streams and one tree height for other small streams were used as standards for federal lands in the northwestern United States. However, these standards were arbitrary, based upon expert opinion, and not derived from effectiveness studies. Effectiveness studies (Cross 2002, Ice 2000, NCASI 2000) tend to show the vast majority of the LWD, shade and particulate matter available to streams are concentrated in the first ten meters. Microclimatic impacts, however, may not be attenuated with narrower buffers (Brosofske et al. 1997). The length of protection along the stream is also an issue since low order streams are more common and have been difficult to identify (University of Washington College of Forest Resources 2007, DP7). A final issue is related to management on migratory floodplains, since buffers established adjacent to the current channel position will be insufficient to accommodate fluvial shifts in the channel (Naiman et al. 2000). Lakes, ponds, and other wetlands are considered important elements within aquatic ecosystem that are afforded special protection when landscape management plans are developed. The Northwest Forest Plan (USDA Forest Service and USDI Bureau of Land Management 1994a, b) assigned two site-potential tree heights to protect forests around lakes and natural ponds and one site potential tree height for other wetlands. et al. 2004). In particular, low gradient pools for salmon rearing and the LWD important to their formation were considered of high importance. Only certain elevation profiles, streams banks and stream configurations likely to contribute LWD are critical in the formation of these pools. Greater protection from urban influences was thought to be more important than needed for rural areas. While these and other elements were considered of high priority and could potentially support more effective criteria for management, there does not appear to be a direct body of research quantifying such relative priorities as a part of management planning. Road networks: Because road networks can heavily influence sediments and organic materials supply to wetlands, road location and construction in order to minimize sediment is another important forest management consideration (Lindenmayer and Franklin 2002). While there exists a substantial body of literature on sediment management it is generally beyond the scope of this review. Skid roads in thinning units may contribute to spatial variability in stand structure, as well as sites for establishment of shade-tolerant trees (Nyland 2002). For example, bats may utilize roads and trails for travel between feeding (Hayes and Loeb 2007). Additionally, demographic processes such as post-harvest tree mortality (a source process for snag/downed wood) may be influenced by the proximity of skid trails (Thorpe et al. 2008). Geologic features: Specialized habitats such as cliffs, caves, thermal habitats, meadows, and vernal pools were identified as needing special attention. Calving sites for ungulates, high-quality spawning habitat for fish, foraging sites with rare but essential food resources, overwintering habitat areas were also to be considered for special attention (Lindenmayer and Franklin 2002). Harvesting and natural disturbances: Spatial and temporal arrangement of harvest units were identified as important in landscape-level forest management. The size of harvest units, levels of structural complexity retained within units, and time interval between rotations and temporal arrangement of harvest units and their management prescriptions were all identified as important considerations (Lindenmayer and Franklin 2002) and are covered in considerable depth under the spatial management sections of this review. While young stands do not develop features associated with old forest species such as the spotted owl, heavy thinnings at 50 and 80 years were considered more effective than not thinning (Andrews and Perkins 2005). In effect long rotations without thinnings are less effective both economically and for producing old forest-like conditions. Multiple treatment approaches may also help to mimic the variability of natural disturbance regimes. Lindenmayer and Franklin (2002) summarize stand-level management methods by including biodiversity pathways, dispersed retention, aggregated retention, variable retention harvest systems, variable-density thinning, and snag creation. Longer rotations have been linked to greater biological diversity (Lindenmayer and Franklin 2002, Oliver 1992, Carey et al. 1999). However, extended rotations result in losses of economic return that, without compensation, will limit acceptability. A range of rotation periods instead of a single rotation period has been proposed. Multiple rotation ages better mimic the variability in frequency of natural disturbance regimes (Seymour and Hunter 1999), and short rotations on certain sites may enhance certain structures that benefit biodiversity (e.g., aspen stands, Arno and Fiedler 2005). A useful frame of reference that has been used by researchers is natural disturbance history. Disturbance regimes have been developed to imitate landscape patterns caused by natural processes. Natural disturbance regimes are divided into two classes, 1) intense episodic disturbance regimes and 2) chronic disturbance regimes. Intense episodic disturbance regimes are defined by catastrophic stand-replacing disturbances such as forest fire and floods. However, the scale of size and time of episodic disturbance regimes are usually much larger than can be implemented on the ground. Therefore, only a few cases have employed intense episodic disturbance regimes (Cissel et al. 1998, 1999). Events associated with many chronic disturbance regimes occur at smaller spatial scales than those associated with episodic disturbance regimes, and are common in landscapes with frequent wind disturbances or insect/disease patches (Deal and Tappeiner 2002). Gap based timber harvesting or group selection may mimic the spatial effects of many chronic disturbance regimes (Lindenmayer and Franklin 2002). Silviculture based on chronic disturbance regimes is intended to create spatially complex landscape structures through the use of relatively small harvesting units. The current management plan for federal lands throughout the Sierra Nevada provides one example of a project emulating a chronic disturbance regime (USDA Forest Service 2001). In the New England landscape, agricultural clearing and subsequent abandonment of farmlands and imposition of patch clearcutting timber harvest regimes have resulted in substantial, measurable changes to forest landscape patch size distribution, stand composition, and structures such as woody debris (Seymour et al. 2002, Howard et al. 2005). Seymour et al. (2002) emphasize the shift from intermediate-scale clearcutting to a within-stand, gap-based cutting system to regenerate the species composition and structure of historic stands in addition to restoring the large, chronically disturbed landscape patches of historic New England. Such approaches are being investigated for other parts of the world, such as the true-fir/European beech forests of southeastern Europe (e.g., Nagel and Svoboda 2008) and the southern beech forests of southern Argentina and Chile (e.g., Martínez-Pastur et al. 2000). The many reports on thinning cited earlier are largely focused on achieving outcomes similar to the impact of disturbances (Bailey and Tappeiner 1998, Carey 1998, Tappeiner et al. 1997). Bailey and Tappeiner (1998) noted substantial structural differences by different thinning treatments contributing to diversity and complexity but that there were greater differences in response across sites than across stand types suggesting stand locations are very resilient in restoring prior growth and structural conditions after disturbances. The dispersed management model that spatially distributes harvest units across a landscape has been applied by many government agencies. The dispersed management model is appealing since it creates heterogeneity across the landscape. However, there are disadvantages as well, including negative impacts on biodiversity (Franklin and Forman, 1987), average patch size and interior habitat reduction, edge habitat increase, and increase in wind susceptibility (Sinton et al. 2000). Many of these issues are related to lack of congruence with the spatial characteristics of the natural disturbance regime. Lindenmayer and Franklin (2002) recommend diversifying landscape-level approaches by imitating natural disturbance regimes as much as possible in harvest planning. Harvest unit size and shape are considered important in landscape-level forest management. The size of interior habitat required for target species and edge/interior ratio can provide one index (Hof and Joyce 1992, 1993; Bevers and Hof 1999; Ducheyne et al. 2006; Wei and Hoganson 2006). Bayne and Hobson (1997) did not find fragmentation contributed to predation and that forest patch harvesting may not be a serious problem. In other studies (1998) they reported the impact of fragmentation to be largely species specific with some doing better and others worse with the impact of total habitat the more dominant predictor. They noted many species did better outside of isolated patches of dense older forests. The conclusion that a variety of stand types is needed to meet the needs of all species is frequently noted (Chambers and McComb 1999, Oliver 1992, Daw and DeStefano 2001). Spatial connections between different habitat types are also noted in the literature; for example, Mladenoff et al. (1993) noted that connectivity between riparian forest and upland old growth has been disrupted in managed vs. unmanaged forest landscapes by timber harvest regimes that spatially differ from the natural disturbance regime. The early work on biodiversity pathways concluded that while sequencing different treatments had some impact on creating diversity in structures across the landscape, regulations and guidelines limiting the size of harvests or other treatments were more important in determining ultimate outcomes (Carey et al. 1996). A meta-analysis on the impact of patch sizes (Bender and Contreras 1998) concluded that generalist species were sensitive to total habitat, not configuration; interior species experienced somewhat greater loss than others from fragmentation while edge species experience somewhat less loss; and migrating species less loss than resident species. McGarigal and McComb (1995) found that bird abundance for several common forest birds in western Oregon increased in relatively fragmented landscapes. Fragmentation would appear to be most important to just a few species for management planning. Boutin and Hebert (2002) believe that landscape structure (fragmentation) has been overemphasized for some landscapes and that threshold effects that can be better detected by spatial modeling comparisons to natural disturbance regimes will become more important. For some organisms and processes, fragmentation is a serious concern, but it must be carefully disentangled from co-occurring processes such as simple habitat loss in order to design effective conservation strategies (Fahrig 2002, Lindenmayer and Fishcher 2006). Consideration of disturbance processes operative at landscape scales is also critical to the success of forest management. The enhanced contiguity of forest stands vulnerable to fire, disease, and insect infestation in the inland Northwest is a result of prolonged fire suppression in a system once characterized by short fire return intervals (Hessburg et al. 2000, Hessburg et al. 2005). Recommended silvicultural treatments would focus on restoring the spatial interspersion of patch types, reducing density in pine and Douglas-fir stands, and conserving late-successional, complex forest at points in the landscape where the biophysical environment would have historically favored them (Camp et al. 1997). Many of the advances in conceptual and empirical understanding discussed here reflect a trend towards recognizing that management imperatives are derived from considerations at multiple spatial scales. Hummel and Barbour (2005) employ the term “landscape silviculture” to demonstrate coordination of silvicultural activities across multiple spatial scales to achieve objectives. This is an important trend that diverts focus from standard operational scales of focus such as the production area and harvest units. Effectiveness measuresWhile there would appear to be some support in the literature for the concepts of biodiversity enhancement as described above, the depth of research studies that include effectiveness is limited, long-term conclusions potentially premature, and many of the concepts would be difficult to implement technically as well as economically. Such approaches are natural outgrowths of efforts to preserve or restore naturally found conditions rather than to characterize effectiveness measures for integrated management. One difficulty with characterizing the effectiveness across such large scales is how to measure the benefits. Statistical measures across large scales reveal the substantial variation associated with habitat and old forest functionality. While this validates conceptually the idea that substantial variation in forest conditions across the landscape has been the historic precedent, it is still difficult to determine what treatment to apply where for what result. Spatial analysis has emerged over the last two decades as a means of quantifying more directly the impact of any given set of treatment strategies. A substantial body of the literature has attempted to assimilate the latest landscape-level forest management studies that focus on the effects of spatial and temporal forest stand management arrangements on ecological as well as economic outcomes. Spatial and Temporal Forest Stand Management Modeling
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