<strong>Forest</strong> <strong>Resilience</strong>, <strong>Biodiversity</strong>, <strong>and</strong> <strong>Climate</strong> <strong>Change</strong>6. Conclusions <strong>and</strong> ecologicalprinciplesThe biodiversity in a forest is linked to <strong>and</strong> underpinsthe ecosystem’s productivity, resilience, <strong>and</strong> stabilityover time <strong>and</strong> space. <strong>Biodiversity</strong> increases the longtermresilience <strong>and</strong> resistance of forest ecosystemstates, increases their primary production, <strong>and</strong>enhances ecosystem stability at all scales. Whilenot all species play important functional roles inecosystems, many do, <strong>and</strong> we may not know orunderst<strong>and</strong> the role of a given species. Further,under changed environmental conditions, specieswith previously minimal functional responsibilitiesmay become highly functional. The persistenceof these functional groups within ecosystems isessential for ecosystem functioning <strong>and</strong> resilience.Capacity for resilience <strong>and</strong> ecosystem stability isrequired to maintain essential ecosystem goods<strong>and</strong> services over space <strong>and</strong> time. Loss of resiliencemay be caused by the loss of functional groups,environmental change such as climate change, oralteration of natural disturbance regimes (Folke etal. 2004). Loss of resilience results in a regime shift,often to a state of the ecosystem that is undesirable<strong>and</strong> irreversible. <strong>Resilience</strong> needs to be viewed asthe capacity of natural systems to self-repair basedon their biodiversity, hence the loss of biodiversitycould mean a reduction of that capacity. This review,together with those of Loreau et al. (2001), Hooper etal. (2005), <strong>and</strong> Drever et al. (2006), suggested strongsupport for the following concepts specific to forestecosystems <strong>and</strong> their resilience:1. <strong>Resilience</strong> is an emergent ecosystem propertyconferred at multiple scales by the biodiversity inthe forest system. More specifically, forest resilienceis related to genetic diversity, functional speciesdiversity, <strong>and</strong> ecosystem diversity (beta diversity)across a forest l<strong>and</strong>scape <strong>and</strong> over time (table 2).2. Most natural forests are highly resilient ecosystems,adapted to various kinds of perturbations <strong>and</strong>disturbance regimes; but if disturbance exceeds thecapacity of the forest to recover (forest degradationowing to human use, for example, which reducesfunctional components), the system will recover toa different state that may or may not also be highlyresilient, but which is unlikely to provide the formerlevel of goods <strong>and</strong> services.3. Complex forest ecosystems are generally moreproductive <strong>and</strong> provide more goods <strong>and</strong> servicesthan those with low species richness. Productiveforests dominated by mature trees are generallyhighly stable ecosystems.4. There is niche differentiation among some treespecies in a forest, as well as competition, leadingto complexity <strong>and</strong> variability within <strong>and</strong> amongforest ecosystems <strong>and</strong> their processes. Some of thisvariability is caused by small local changes in siteconditions across a forest.5. Redundancy of functional species is common incomplex forest ecosystems <strong>and</strong> is directly relatedto ecosystem resilience. Redundancy providesinsurance against changing environmentalconditions, <strong>and</strong> species with limited functions underone set of conditions may become driver speciesunder an altered set of conditions.6. Diverse forest systems are more stable (withindefined bounds) than less diverse systems <strong>and</strong> thisis partly related to a robust regional species pool <strong>and</strong>the beta diversity among ecosystems.7. Nevertheless, even high diversity is no guaranteefor ecosystem resilience once climate conditionsmove beyond those experienced by most of thecomponent species.8. Although a forest may change states in responseto disturbances, the flow of goods <strong>and</strong> services maynot necessarily be highly altered, suggesting thatthe ecosystem is ecologically resilient, even thoughthe forest community structure may have changed.Ecological resilience is unlikely, however, in a systemthat has low redundancy, such as degraded forests.9. There is a negative relationship between speciesdiversity, l<strong>and</strong>scape diversity, <strong>and</strong> the capacity of aforest system to be invaded, especially by pests <strong>and</strong>diseases.10. Not all forest ecosystems are equally resilientto disturbances, including climate change. Effectsof climate change will vary in forests depending onbiome, tree species composition, natural disturbanceregime, <strong>and</strong> moisture, temperature <strong>and</strong> edaphicresponses to climate change.11. <strong>Resilience</strong> is necessary to maintain desirableecosystem states under variable environmentalconditions.6.1 Ecological principles to foster forest ecosystemresilience <strong>and</strong> stability under climate change43
<strong>Forest</strong> <strong>Resilience</strong>, <strong>Biodiversity</strong>, <strong>and</strong> <strong>Climate</strong> <strong>Change</strong><strong>Forest</strong>s have a capacity to resist environmentalchange owing to their multiple species <strong>and</strong> complexmultiple processes. However, a reduction inbiodiversity in forest systems has clear implicationsfor the functioning of the system <strong>and</strong> the amountsof goods <strong>and</strong> services that these systems are able toproduce. While it is relatively simple to plant trees<strong>and</strong> produce a short-term wood crop, the lack ofdiversity at all levels (i.e., gene, species of flora <strong>and</strong>fauna, <strong>and</strong> l<strong>and</strong>scape) in these systems reducesresilience, degrades the provision of goods <strong>and</strong>services that the system can provide, <strong>and</strong> renders itvulnerable to catastrophic disturbance.Specifically, with respect to mitigating CO 2 emissionsfrom deforestation <strong>and</strong> degradation, maintaininglong-term stable forest ecosystems will be critical,as opposed to for example, rapidly growing simplelow diversity forests that have limited longevity,resistance, resilience or adaptive capacity. Further,the application of ecological sustainability principlesin the recovery of degraded forests to redevelop theirresilience <strong>and</strong> their former goods <strong>and</strong> services willprovide part of a long-term approach to mitigating<strong>and</strong> adapting to climate change (e.g., Lamb etal. 2005, Innes et al. 2009). Hence, maintainingresilience in forests, in time <strong>and</strong> space, is importantto maintain their function as an important “buffer”in the global carbon cycle by maximizing theirpotential to sequester <strong>and</strong> store carbon; along withthe ongoing capacity to provide the other goods <strong>and</strong>services that humans require. To this end, humanuse of forests will need to change in order to ensuretheir conservation, sustainable use, <strong>and</strong> restoration.In managed forests, it is imperative that biodiversity<strong>and</strong> ecosystem resilience be maintained. Theprinciples of sustainable forest management areto maintain ecosystem processes by matchingmanagement practices to natural processes (orexpected processes, modified under climate change)at multiple scales (e.g., Attiwill 1994, Perera et al.2004). Restoration of degraded forest l<strong>and</strong>scapescan take advantage of the linkage betweenbiodiversity <strong>and</strong> ecosystem resilience, by planting toenhance species richness <strong>and</strong> through the additionof functional species (e.g., N-fixing species) whereknown (see: Lamb et al. 2005, Brockerhoff et al.2008, for management recommendations). Variousoptions for policies <strong>and</strong> measures are availableto promote forest conservation <strong>and</strong> biodiversity,particularly at l<strong>and</strong>scape <strong>and</strong> regional scales, inaddition to conventional protected areas, includingpayments for l<strong>and</strong> stewardship <strong>and</strong> ecosystemservices (USDA 2007), connectivity conservationprogrammes (Crooks <strong>and</strong> Sanjayan 2006), <strong>and</strong>schemes built around recognition of Indigenous <strong>and</strong>traditional l<strong>and</strong>s (Australian Government 2007).The capacity to conserve, sustainably use <strong>and</strong> restoreforests rests on our underst<strong>and</strong>ing <strong>and</strong> interpretationof pattern <strong>and</strong> process at several scales, therecognition of thresholds, <strong>and</strong> the ability to translateknowledge into appropriate management actionsin an adaptive manner (Frelich <strong>and</strong> Reich 1998,Gauthier et al. 2008). Caring for forests in waysthat maintain their diversity <strong>and</strong> resilience is beingmade even more complex owing to climate change(e.g., Chapin et al. 2007, Kellomaki et al. 2008). Wesuggest the following as ecological principles thatcan be employed to maintain <strong>and</strong> enhance longtermforest resilience, especially under climatechange (e.g., Thompson et al. 2002, Fischer et al.2006, Millar et al. 2007, Innes et al. 2009):1. Maintain genetic diversity in forests throughpractices that do not select only certain trees forharvesting based on site, growth rate, or form, orpractices that depend only on certain genotypes(clones) for planting (see e.g., Schaberg et al. 2008) .2. Maintain st<strong>and</strong> <strong>and</strong> l<strong>and</strong>scape structuralcomplexity using natural forests as models <strong>and</strong>benchmarks.3. Maintain connectivity across forest l<strong>and</strong>scapesby reducing fragmentation, recovering lost habitats(forest types), <strong>and</strong> exp<strong>and</strong>ing protected areanetworks (see 8. below).4. Maintain functional diversity (<strong>and</strong> redundancy)<strong>and</strong> eliminate conversion of diverse natural foreststo monotypic or reduced species plantations.5. Reduce non-natural competition by controllinginvasive species <strong>and</strong> reduce reliance on non-nativetree crop species for plantation, afforestation, orreforestation projects.6. Reduce the possibility of negative outcomes byapportioning some areas of assisted regenerationwith trees from regional provenances <strong>and</strong> fromclimates of the same region that approximateexpected conditions in the future.7. Maintain biodiversity at all scales (st<strong>and</strong>,l<strong>and</strong>scape, bioregional) <strong>and</strong> of all elements (genetic,species, community) <strong>and</strong> by taking specificactions including protecting isolated or disjunctpopulations of organisms, populations at margins44
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