- Open Access
Forest restoration, biodiversity and ecosystem functioning
BMC Ecology volume 11, Article number: 29 (2011)
Globally, forests cover nearly one third of the land area and they contain over 80% of terrestrial biodiversity. Both the extent and quality of forest habitat continue to decrease and the associated loss of biodiversity jeopardizes forest ecosystem functioning and the ability of forests to provide ecosystem services. In the light of the increasing population pressure, it is of major importance not only to conserve, but also to restore forest ecosystems.
Ecological restoration has recently started to adopt insights from the biodiversity-ecosystem functioning (BEF) perspective. Central is the focus on restoring the relation between biodiversity and ecosystem functioning. Here we provide an overview of important considerations related to forest restoration that can be inferred from this BEF-perspective.
Restoring multiple forest functions requires multiple species. It is highly unlikely that species-poor plantations, which may be optimal for above-ground biomass production, will outperform species diverse assemblages for a combination of functions, including overall carbon storage and control over water and nutrient flows. Restoring stable forest functions also requires multiple species. In particular in the light of global climatic change scenarios, which predict more frequent extreme disturbances and climatic events, it is important to incorporate insights from the relation between biodiversity and stability of ecosystem functioning into forest restoration projects. Rather than focussing on species per se, focussing on functional diversity of tree species assemblages seems appropriate when selecting tree species for restoration. Finally, also plant genetic diversity and above - below-ground linkages should be considered during the restoration process, as these likely have prominent but until now poorly understood effects at the level of the ecosystem.
The BEF-approach provides a useful framework to evaluate forest restoration in an ecosystem functioning context, but it also highlights that much remains to be understood, especially regarding the relation between forest functioning on the one side and genetic diversity and above-ground-below-ground species associations on the other. The strong emphasis of the BEF-approach on functional rather than taxonomic diversity may also be the beginning of a paradigm shift in restoration ecology, increasing the tolerance towards allochthonous species.
Globally, forests cover nearly one third of the land area and contain over 80% of terrestrial biodiversity . The income of more than 1.6 billion people depends on forests and sustainable management of forests can contribute to sustainable development, poverty eradication and the achievement of internationally agreed development goals [1, 2]. Despite increasing efforts for sustainable forest management and forest conservation , the extent of forest habitat, in particular in the tropics, continues to decrease, mainly by forest conversion to agriculture and land uses related to urban population growth [4, 5]. Between 1980 and 2000 more than half of the new agricultural land across the tropics was obtained by clearing intact forests [6, 7]. Also, many disturbed and secondary forests, which are increasingly important habitat for many forest species [8, 9], are eventually cleared for agricultural purposes.
In the remaining forests and forest fragments, decreasing habitat patch sizes result in increased deleterious edge effects  and decreasing plant and animal population sizes , which, in turn, may lower population viability and genetic variation [12, 13]. The negative effects of forest fragmentation and isolation are expected to be exacerbated by other anthropogenic threats such as fire [14, 15], in particular in the light of global climatic change [16, 17]. Parallel to forest loss and forest fragmentation, cryptic deforestation [18, 19] - the selective logging and internal degradation of forests - alters forest structure and plant communities, jeopardizing biodiversity, regeneration capacity and vitality of forests . The simultaneous reduction of both forest quantity and quality is expected to lead to massive extinction of many species inhabiting forest habitats . For a wide range of taxa, including trees and lianas, birds, fruit-feeding butterflies, leaf-litter amphibians, large mammals, epigeic arachnids, lizards, dung beetles and bats, biodiversity has been shown to decline significantly over a forest degradation gradient, from primary over secondary to plantation forest .
Loss of forest biodiversity may seriously jeopardize the functioning of forest ecosystems (i.e. the activities, processes or properties of forests, such as decomposition of organic matter, soil nutrient cycling and water retention), and consequently the ability of forest to provide ecosystem services . Ecosystem services have been defined as the benefits that people obtain from ecosystems  and have been categorized into four broad categories. These include provisioning services such as food, water, timber, and fiber; regulating services that affect climate (e.g. though carbon sequestration), pollination, biological pest control, floods, disease, wastes, and water quality; cultural services that provide recreational, aesthetic, and spiritual benefits; and supporting services such as soil formation, photosynthesis, and nutrient cycling [25–27].
Clearly, the role of forests as sanctuaries of biodiversity and as providers of ecosystem services cannot be overestimated. In the light of the increasing human population, however, conserving the remaining forests and their biodiversity, functions and services of forests is unlikely to be sufficient . To meet the increasing demands for ecosystem services provided by forests - in particular the many provisioning services of forests as many people heavily rely on forests for livelihoods and products such as timber, medicines, thatch, fiber and meat  - large-scale (passive or active) forest restoration is probably the only solution that will be effective in the long term [28–30]. Establishing short-rotation single- or multiple-species plantations on degraded soils, restoration plantings in secondary forests or assisted regeneration in selectively logged forest are a few examples of the wide spectrum of forest restoration approaches . They all have in common that they consist of management interventions that aim at recovering ecosystems that have been degraded, damaged or destroyed by human activities [29, 32]. Ecological restoration is therefore an important practice that may increase levels of biodiversity in human-altered ecosystems  and may mitigate the impact of climate change . To this end, restoration ecology has adopted insights from both community and ecosystem ecology, and more recently, from the integrated biodiversity-ecosystem functioning (BEF) perspective [35–37]. The main aim of this article is to discuss how forest restoration may benefit from insights originating from the emerging BEF framework.
Traditional approaches to ecological restoration
The community approach
A biological community is a group of organisms that interact and share an environment. Within a community, organisms may compete for the same resources (competition), profit from the presence of other organisms (facilitation) [38, 39] or use other organisms as a food source (trophic interaction) . In stable communities, these interactions lead to predictable, directional changes in community structure known as ecological succession. Succession is an important guiding principle in the community approach to ecological restoration . The restoring forest is a dynamic ecosystem, with changing species composition and forest structure, but interventions and management steer the forest towards a desired climax or pre-disturbance community structure. These interventions are usually designed to accelerate natural succession or to bypass intermediate successional phases. Basically, the community approach is focussing on restoring forest biodiversity per se. The many studies that apply facilitation as a restoration tool of woody communities  are typical examples of the community approach to forest restoration. Planting late-successional tree species (protégé species) under early-successional shrubs (nurse species) has been shown to be an effective means of restoring forests under high abiotic stress [42, 43] (Figure 1).
Insights from alternative stable states theory have also been useful to guide restoration practices that focus on community structure . In severely degraded systems, alternative stable states may make efforts to restore pre-disturbance communities difficult, if not impossible . In such cases, a single intervention may not suffice to induce forest regrowth: succession fails and the community is blocked in a low diversity/low biomass state. Exclusion of grazing animals may be an effective means for woodland restoration in degraded drylands, but only when soil moisture conditions also improve. Wet pulses caused by climatic oscillations such as the El Niño Southern Oscillation may provide such necessary additional impulse to induce a regime shift that leads to forest restoration . Similarly, planting and sowing of late successional tree species (an intervention to overcome seed limitation) has been found effective for the restoration of highly complex forest on bauxite mined sites, but only after careful site preparation and topsoil handling or replacement (interventions to overcome survival limitation caused by soil compaction, decreased soil porosity and infiltration capacity, and the loss of soil biota) [47, 48].
The ecosystem approach
Restoration of species richness and community structure over time implies increasing ecosystem complexity and functionality . In the ecosystem approach, restoration of ecosystem functions such as primary production, energy flows and nutrient cycles, is the guiding principle on which restoration efforts are based . Basically, this approach aims at restoring suitable abiotic conditions that allow (passive) recolonization of species. The ecosystem perspective typically starts from a landscape point of view, building on spatial heterogeneity and broad spatial scales . The connections or barriers between neighbouring ecosystems have an effect on the resource balances and set limits on the communities that can be restored . Reforestation of degraded sites with trees that alter the physical and chemical characteristics of the soil and that affect the biochemical cycles through litter fall or root activity presents a typical example of the ecosystem approach to forest restoration [51, 52].
The biodiversity - ecosystem function approach to ecological forest restoration
The study of the relation between biodiversity and ecosystem functioning is a rapidly growing field (see the volume edited by Naeem et al.  for an exhaustive state of the art). The traditional view that has dominated ecology until the 1990's started from the idea that species distribution patterns resulted directly from the abiotic and biotic (species interactions) components determining the environment. In the early 1990's, however, this view was challenged, when one started to realize that species diversity also affects the abiotic environment, and even the functioning of ecosystems . The functioning of an ecosystem incorporates processes such as decomposition of organic matter, fixation of carbon, nutrient and water cycling and degradation of toxic compounds. Meta-analyses of the results of mainly small-scale biodiversity experiments have shown that, on average, ecosystem functions increase with increasing species number [e.g. ]. The success of the idea that biodiversity affects ecosystem properties and functions - some have called it a paradigm shift in ecology  - can be explained by the fact that it offers a comprehensive framework to evaluate the consequences of biodiversity loss caused by human activities, and at the same time provides a powerful incentive for biodiversity conservation and ecological restoration [37, 57].
Naeem  was the first to propose that restoration ecology may benefit from insights from the BEF framework, and this idea has been further elaborated by Wright et al. . Here we build on these ideas and put them in a forest restoration context. In contrast to more traditional approaches, restoration based on the BEF perspective strongly focuses on restoring the relationship between biodiversity and ecosystem functioning . In what follows we list some important considerations regarding forest restoration that can be inferred from the BEF framework. We are aware that foresters have already adopted the BEF framework in setting up large experiments where the effects of tree species richness on ecosystem functions are evaluated [e.g., [58, 59]]. Nevertheless, we believe that forest restoration efforts may benefit from such an overview, in particular since ecosystem functioning and functional (bio)diversity has received very little attention in a forest restoration context so far (Figure 2).
Restoring multiple forest functions requires multiple species
One of the major functions of forest ecosystems is carbon fixation , which is directly related to the ecosystem services carbon sequestration and the provision of fire and construction wood. There is evidence that tree diversity has a positive effect on ecosystem production (see Thompson et al.  for an overview). Based on the largest data set ever analysed in this context to date (12.000 permanent forest plots in eastern Canada), Paquette & Messier  reported that, after controlling for environmental and climate differences between plots, tree productivity was positively related to stand biodiversity. These results confirm earlier work in 5000 permanent plots in Mediterranean forests across Catalonia (NE Spain) . In a reforestation context, Piotto et al.  found that mixed plantations in Costa Rica performed better than monocultures for all growth variables considered, including height, diameter at breast height, volume, and above-ground biomass. Also in natural stands of tropical forest with high environmental and spatial variation, positive effects of tree species diversity on tree carbon storage were found . Positive effects of tree diversity on above-ground productivity are certainly not an universal pattern, however [61, 66], and above-ground biomass production and soil carbon fixation may also respond differently to tree diversity in plantation forests . This corroborates the result of a meta-analysis of BEF experiments where it was found that high biodiversity treatments do not always outperform the best performing monoculture . In a forest restoration context, where fast growing tree species with strong global timber markets are readily available, this may suggest that monocultures are an option. However, evidence is accumulating that focussing on one single ecosystem function often overlooks an important aspect of biodiversity: the possibility of one species to contribute to different ecosystem functions at the same time . Because different species often influence different ecosystem functions, focussing on one function in isolation will strongly underestimate the biodiversity required for maintaining an ecosystem with multiple functions, at multiple times and places in a changing environment . Although the evidence only comes from grasslands and aquatic environments so far, it convincingly shows that species redundancy is unlikely to occur when several ecosystem functions and services are considered in combination [68–71].
Therefore, it is highly unlikely that species poor plantations will outperform species diverse tree assemblages for a combination of forest ecosystem functions , including above-ground biomass production, disease resistance, carbon fixation, nectar provision, erosion control, water capitation, N2-fixation and fruit production. It is therefore of special importance that reforestation efforts clearly define the ecosystem services and functions that the restored forest is intended to deliver. Also, it is important to realize that ecosystem functions of restoring forests may change over time because of changes in tree sizes, forest structure and relative importance of functional groups, even if there are no changes in tree species composition . Finally, it should be noticed that although there is already some knowledge on the effects of tree diversity on forest productivity, it is not known how understory shrub diversity, and even herbaceous species, affect forest productivity or other ecosystem functions. This may, for example, happen through these species' impacts on litter decomposition, on water capture and on the diversity of soil biota .
Restoring stable forest functions requires multiple species
The hypothesis that larger species diversity leads to higher stability of ecosystem functioning has been a point of debate for half a century, and it has re-emerged within the BEF framework [53, 74, 75]. The main ideas behind the biodiversity vs. ecosystem stability concept are functional response diversity and functional compensation [61, 76]. This occurs when positive changes in the level of functioning of one species (a species becoming functionally dominant) are associated with negative changes in the functioning of other species. This compensation drives the stabilization of ecosystem properties such as biomass production . Basically, the stability of the functioning of an ecosystem can be measured in three ways: i) the long term variability of an ecosystem property through time in relation to background environmental variation (variance); ii) the impact (resistance); and iii) the recovery (resilience) of ecosystem properties to discrete disturbances [61, 78]. As it is expected that these discrete and extreme disturbances such as extreme climate events and pest and disease outbreaks will become more frequent under the predicted climate change , it is very important to incorporate insights from the relation between biodiversity and stability of ecosystem functioning into forest restoration projects. It is crucial to realize that, just as the degree of species redundancy decreases when multiple ecosystem functions are considered (see earlier), there is currently strong experimental evidence that in changing environments, more species are required to guarantee ecosystem functioning than in constant environments [e.g., [69, 80]].
Evidence for the latter comes from studies that have related forest tree diversity with measures of stability of forest ecosystem functioning. Lloret et al.  used satellite imagery to estimate the impact of the extreme 2003 summer drought on canopy greenness of different forests types in Spain, by quantifying the NDVI (normalized difference vegetation index). NDVI correlates with ecosystem CO2 fluxes. These authors reported a positive relation between woody species diversity and resistance of canopy greenness against drought in forests on dry locations, whereas no such relation was discovered in more moist forests. Similarly, DeClerck et al.  related stability in stand productivity across 64 years with conifer diversity in the Sierra Nevada, USA. They found a significant relation between species richness and resilience of stand productivity after recurrent severe droughts. Resistance to drought was, however, not related to species diversity. These studies partly support positive biodiversity effects on stability of biomass production, but they also show that patterns may be complex, vary across ecosystem types, and depend on the measures that are used to quantify stability. In any case, temporal stability of ecosystem functioning is an important consideration for projects aiming at forest restoration, especially under the current global change scenario. Again, it is not known whether understory shrubs and herbaceous species contribute to the stability of forest ecosystem functioning.
Focus on functional diversity rather than on taxonomic diversity
Whereas general biodiversity measures are based on taxonomy in the first place (species presence or absence), functional diversity measures relate to what organisms effectively do in an ecosystem, quantify the distribution of traits in a community or measure the relative magnitude of species similarities and differences. How to best measure functional diversity is a much debated question, but Cadotte et al.  summarize five useful multivariate functional diversity measures. Some authors have suggested that functional diversity measures are particularly suitable or even better to predict the interactions between biodiversity and ecosystem processes [83–85]. Using a tree diversity index based on among-species variation in seed mass, wood density and maximum height Paquet and Messier  showed that this measure outperformed a taxonomically based diversity index in explaining tree productivity. Bunker et al.  demonstrated that removing certain functional groups from a tropical forest had more important effects on the above-ground carbon pool than randomly removing species. Vila et al. , on the contrary, reported that functional group richness performed worse than tree species richness, but this was likely due to a rather rudimentary functional group delineation. Thus, when selecting tree species for forest restoration, these findings suggest focusing on functional groups based on relevant plant traits. While these traits are readily available for species from temperate regions by now, the establishment of plant trait databases for tropical tree species and the centralisation of all available data in a general database are important works in progress [87, 88]. Maximizing functional diversity can be achieved by quantifying the functional diversity of the species mix used for restoration. This can be done by delineating emergent or functional groups (assemblages of species performing similar functional roles) [e.g. [61, 89]], or by using more complex, continuous or non-grouping measures of functional diversity . The selection of relevant plant traits remains, however, crucial with respect to the forest ecosystem functions to be restored. Scherer-Lorenzen et al.  provide a comprehensive list of species traits that can be used to quantify functional diversity of tree mixtures used for reforestation of European temperate forests. Selected traits included nominal (e.g. leaf type, crown architecture), ordinal (e.g. adult light requirements, height growth vigour) and scale variables (e.g. leaf N concentration, litter C:N ratio). A better mechanistic understanding of how species traits and their interactions affect ecosystem functioning is also important, however, to be able to proactively analyse different reforestation scenarios and their impact on forest functioning. In this context, it is important to realize that relationships between functional traits and ecosystem functions such as carbon storage in natural populations are not always transferable to tree plantations and vice versa .
Effects of genetic diversity extend up to the ecosystem level
Whereas conservation biologists have acknowledged the negative fitness consequences of reduced genetic diversity for decades, forest restoration projects may still incorporate very few genotypes . There is evidence, however, that monoclonal populations are more vulnerable to pathogens than genetically diverse assemblages [e.g. [92, 93]]. The point that we want to make here, however, is that the effects of stand genetic diversity can be expected to extend far beyond the fitness of the individual trees or stands. It is only recently that it has become clear that variation in population genetic diversity or in genotype composition can have far-reaching ecological effects. The ecological consequences of genetic diversity (coined 'community genetics') have been demonstrated at different levels of organization, from the population over the community to the ecosystem [94–96]. For example, plant genotypic diversity and genotype identity have been shown to affect biomass production and community invasibility, and also the invertebrate diversity of the higher trophic levels [97, 98]. It was also shown that litter decomposition and nutrient release differed between different Populus genotypes, indicating that selection of tree genotypes may have profound and long lasting effects on ecosystem functioning of restored forests [99, 100]. Although a discipline as community genetics is in its infancy, there is already some evidence to suggest that there are extended consequences of plant genetic variation, up to the level of the ecosystem properties . The selection of specific genotypes, and the genotypic diversity of tree assemblages, may therefore have major implications for the functioning and the resilience of forests .
Synchronize above- and below-ground biodiversity
The above-ground biodiversity of forests also comprises fauna with important ecosystem services that include pollination, pest control and seed dispersal. The ecosystem services of birds, for instance, have been well documented  and in the light of forest restoration birds have been shown essential for dispersing tree seeds into restoring areas and overcoming seed dispersal and germination limitation [102, 103]. Far less is known about the role of below-ground biota and the linkages between trees and these biota. The study of soil microbial community structure and functioning has traditionally received little attention in ecology. But as with above-ground biodiversity, there is evidence that below-ground diversity has a significant impact on ecosystem functioning. In a series of simplified tropical forests, Lovelock and Ewel  found significant positive relationships between the diversity of arbuscular mycorrhizal fungi (AMF) and ecosystem net primary productivity, and between AM fungal community evenness and ecosystem phosphorus-use efficiency. The quick development and availability of molecular tools such as t-RFLP and next-generation sequencing to quantify microbial diversity [e.g. [105, 106]], together with the strong focus of the BEF approach on the functionality of ecosystems, has resulted in an increased interest in the role of microbial soil community diversity in driving processes such as organic matter decomposition and plant nutrient uptake. Because ecological restoration is usually occurring on highly disturbed or degraded sites, it is important that above-ground-below-ground species linkages are permanently considered during the restoration process , and more specific, that there is a synchronisation between above- and below-ground species associations . Clearly, the crucial question is whether below-ground microbial community simply follows the introduced tree and shrub species, or whether some kind of inoculation is required [see e.g. ]. Among the relevant soil micro-organisms, arbuscular mycorrhizal fungi (AMF) and ectomycorrhizal fungi (ECMF) can be expected to play a major role during restoration of degraded sites. Many tree and shrub species associate with AMF and ECMF, which provide nutrients in exchange for plant carbohydrates. Recent evidence has shown that at least ECMF are dispersal limited, and are less abundant on isolated trees . This finding may urge for some kind of active inoculation of degraded restoration sites. How to successfully apply soil microorganisms in particular restoration projects, however, is an almost empty research field. Whereas fundamental insights in the role of AMF in structuring grassland communities are growing [but see ], it remains largely unknown how these fungi contribute to successful restoration and the few available reports on the effects of large scale inoculations in grasslands did result in contradictory conclusions (White et al. vs. Smith et al. ). Also, the inoculation of tree roots with mycorrhiza has received some attention in forest restoration projects, but the results are not straightforward [e.g., [113, 114]]. This leads to the conclusion that at present much remains to be understood about how below-ground microbial diversity contributes to successful restoration of forest functions. Newly available molecular tools to quantify microbial diversity combined with detailed measurements of forest functioning are likely to increase our insights in how to apply below-ground biodiversity for restoration purposes.
Restored forests are often novel ecosystems
While restored forests may deliver similar ecosystem services and conserve levels of biodiversity comparable to the pre-disturbance vegetation, restored forests rarely match the composition and structure of the original forest cover . Large changes in ecosystems will usually result in novel systems, comprising different species, interactions and functions [116, 117]. In this context, it is important to realize that both the recent tendency towards accepting perennial, global change driven changes to the environment and the increasing application of the BEF framework to ecological restoration may facilitate the acceptance of using non-native species in forest restoration. While many ecologists still consider autochthony of species a prerequisite for their use in ecological restoration [see e.g. ], a focus on species' functions rather than on species' origins is already advocated by others  as being a "more dynamic and pragmatic approach to the conservation and management of species". In this sense, the BEF approach may be at the source of a paradigm shift in restoration ecology .
The BEF approach provides a useful framework to evaluate forest restoration in an ecosystem functioning context. It highlights different aspects of forest restoration that do not always receive sufficient attention in the more traditional approaches to restoration. At the same time the BEF framework confronts us with huge knowledge gaps still present in restoration science. The mechanistic understanding of how plant functional traits and their mutual interactions affect ecosystem functioning, understanding the role of genetic diversity in ecosystem functioning, and acquiring insights in the interactions between below-ground biodiversity and forest functioning and restoration success, are the most urgent research needs.
Raf Aerts is a forest engineer and tropical field ecologist and is a post-doctoral research fellow at the Division Forest, Nature and Landscape of the University of Leuven (K.U.Leuven). His research focuses on (tropical) forest conservation and forest restoration. He applies principles of community ecology and ecological genetics to trees, birds, epiphytic orchids and wild arabica coffee. Olivier Honnay is associate professor of plant ecology and plant conservation biology at the Biology Department of the University of Leuven.
arbuscular mycorrhizal fungi
biodiversity - ecosystem functioning
normalized difference vegetation index
United Nations International Year of Forests, 2011. [http://www.un.org/en/events/iyof2011/]
United Nations Resolution adopted by the General Assembly 61/193 International Year of Forests, 2011. [http://www.un.org/en/events/iyof2011/resolution.shtml]
Butchart SHM, Walpole M, Collen B, van Strien A, Scharlemann JPW, Almond REA, Baillie JEM, Bomhard B, Brown C, Bruno J, Carpenter KE, Carr GM, Chanson J, Chenery AM, Csirke J, Davidson NC, Dentener F, Foster M, Galli A, Galloway JN, Genovesi P, Gregory RD, Hockings M, Kapos V, Lamarque JF, Leverington F, Loh J, McGeoch MA, McRae L, Minasyan A: Global biodiversity: indicators of recent declines. Science. 2010, 328 (5982): 1164-1168.
Ahrends A, Burgess ND, Milledge SAH, Bulling MT, Fisher B, Smart JCR, Clarke GP, Mhoro BE, Lewis SL: Predictable waves of sequential forest degradation and biodiversity loss spreading from an African city. Proc Natl Acad Sci USA. 2010, 107 (33): 14556-14561.
DeFries RS, Rudel T, Uriarte M, Hansen M: Deforestation driven by urban population growth and agricultural trade in the twenty-first century. Nature Geosci. 2010, 3 (3): 178-181.
Gibbs HK, Ruesch AS, Achard F, Clayton MK, Holmgren P, Ramankutty N, Foley JA: Tropical forests were the primary sources of new agricultural land in the 1980s and 1990s. Proc Natl Acad Sci USA. 2010, 107 (38): 16732-16737.
Lambin EF, Meyfroidt P: Global land use change, economic globalization, and the looming land scarcity. Proc Natl Acad Sci USA. 2011, 108 (9): 3465-3472.
Farwig N, Sajita N, Bohning-Gaese K: Conservation value of forest plantations for bird communities in western Kenya. For Ecol Manage. 2008, 255 (11): 3885-3892.
Edwards DP, Larsen TH, Docherty TDS, Ansell FA, Hsu WW, Derhe MA, Hamer KC, Wilcove DS: Degraded lands worth protecting: the biological importance of Southeast Asia's repeatedly logged forests. Proc Roy Soc B Biol Sci. 2011, 278 (1702): 82-90.
Laurance WF, Nascimento HEM, Laurance SG, Andrade A, Ewers RM, Harms KE, Luizao RCC, Ribeiro JE: Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis. PLoS ONE. 2007, 2 (10):
Laurance WF, Laurance SG, Hilbert DW: Long-term dynamics of a fragmented rainforest mammal assemblage. Conserv Biol. 2008, 22 (5): 1154-1164.
Honnay O, Jacquemyn H, Bossuyt B, Hermy M: Forest fragmentation effects on patch occupancy and population viability of herbaceous plant species. New Phyt. 2005, 166 (3): 723-736.
Aguilar R, Quesada M, Ashworth L, Herrerias-Diego Y, Lobo J: Genetic consequences of habitat fragmentation in plant populations: susceptible signals in plant traits and methodological approaches. Molec Ecol. 2008, 17 (24): 5177-5188.
Siegert F, Ruecker G, Hinrichs A, Hoffmann AA: Increased damage from fires in logged forests during droughts caused by El Niño. Nature. 2001, 414 (6862): 437-440.
Laurance WF, Camargo JLC, Luizao RCC, Laurance SG, Pimm SL, Bruna EM, Stouffer PC, Williamson GB, Benitez-Malvido J, Vasconcelos HL, Van Houtan KS, Zartman CE, Boyle SA, Didham RK, Andrade A, Lovejoy TE: The fate of Amazonian forest fragments: a 32-year investigation. Biol Conserv. 2011, 144 (1): 56-67.
Malhi Y, Aragao L, Galbraith D, Huntingford C, Fisher R, Zelazowski P, Sitch S, McSweeney C, Meir P: Exploring the likelihood and mechanism of a climate-change-induced dieback of the Amazon rainforest. Proc Natl Acad Sci USA. 2009, 106 (49): 20610-20615.
Aragao L, Shimabukuro YE: The incidence of fire in Amazonian forests with implications for REDD. Science. 2010, 328 (5983): 1275-1278.
Nepstad DC, Verissimo A, Alencar A, Nobre C, Lima E, Lefebvre P, Schlesinger P, Potter C, Moutinho P, Mendoza E, Cochrane M, Brooks V: Large-scale impoverishment of Amazonian forests by logging and fire. Nature. 1999, 398 (6727): 505-508.
Puyravaud JP, Davidar P, Laurance WF: Cryptic loss of India's native forests. Science. 2010, 329 (5987): 32-32.
Aerts R, Hundera K, Berecha G, Gijbels P, Baeten M, Van Mechelen M, Hermy M, Muys B, Honnay O: Semi-forest coffee cultivation and the conservation of Ethiopian Afromontane rainforest fragments. For Ecol Manage. 2011, 261: 1034-1041.
Wright SJ, Muller-Landau HC: The future of tropical forest species. Biotropica. 2006, 38: 287-301.
Barlow J, Gardner TA, Araujo IS, Ávila-Pires TC, Bonaldo AB, Costa JE, Esposito MC, Ferreira LV, Hawes J, Hernandez MIM: Quantifying the biodiversity value of tropical primary, secondary, and plantation forests. Proc Natl Acad Sci USA. 2007, 104 (47): 18555-18560.
Duffy JE: Why biodiversity is important to the functioning of real-world ecosystems. Front Ecol Environ. 2009, 7 (8): 437-444.
Millenium Ecosystem Assessment. Ecosystems and Human Well-being: Synthesis. 2005, Washington, DC.: Island Press
Costanza R, dArge R, deGroot R, Farber S, Grasso M, Hannon B, Limburg K, Naeem S, Oneill RV, Paruelo J, Raskin RG, Sutton P, vandenBelt M: The value of the world's ecosystem services and natural capital. Nature. 1997, 387 (6630): 253-260.
Ricketts TH, Daily GC, Ehrlich PR, Michener CD: Economic value of tropical forest to coffee production. Proc Natl Acad Sci USA. 2004, 101 (34): 12579-12582.
Kremen C: Managing ecosystem services: what do we need to know about their ecology?. Ecol Lett. 2005, 8: 468-479.
Lamb D, Erskine PD, Parrotta JA: Restoration of degraded tropical forest landscapes. Science. 2005, 310 (5754): 1628-1632.
Rey Benayas JM, Newton AC, Diaz A, Bullock JM: Enhancement of biodiversity and ecosystem services by ecological restoration: a meta-analysis. Science. 2009, 325 (5944): 1121-1124.
Holl KD, Aide TM: When and where to actively restore ecosystems?. For Ecol Manage. 2011, 261 (10): 1558-1563.
Lamb D: Regreening the bare hills: tropical forest restoration in the Asia-Pacific region. 2010, World Forests vol. 8: Springer Verlag
Birch JC, Newton AC, Aquino CA, Cantarello E, Echeverria C, Kitzberger T, Schiappacasse I, Garavito NT: Cost-effectiveness of dryland forest restoration evaluated by spatial analysis of ecosystem services. Proc Natl Acad Sci USA. 2010, 107 (50): 21925-21930.
Brudvig LA: The restoration of biodiversity: where has research been and where does it need to go?. Am J Bot. 2011, 98 (3): 549-558.
Harris JA: Soil microbial communities and restoration ecology: facilitators or followers?. Science. 2009, 325 (5940): 573-574.
Naeem S: Biodiversity and ecosystem functioning in restored ecosystems: extracting principles for a synthetic perspective. Foundations of Restoration Ecology. Edited by: Falk DA, Palmer MA, Zedler JB. 2006, Washington, D.C.: Society for Ecological Restoration International, Island press, 210-237.
Wright JP, Symstad AJ, Bullock JM, Engelhardt KAM, Jackson LE, Bernhardt ES: Restoring biodiversity and ecosystem function: will an integrated approach improve results?. Biodiversity, ecosystem functioning, and human wellbeing. Edited by: Naeem S, Bunker DE, Hector A, Loreau M, Perrings C. 2009, Oxford: Oxford University Press, 167-177.
DeClerck FAJ, Chazdon R, Holl KD, Milder JC, Finegan B, Martinez-Salinas A, Imbach P, Canet L, Ramos Z: Biodiversity conservation in human-modified landscapes of Mesoamerica: Past, present and future. Biol Conserv. 2010, 143 (10): 2301-2313.
Callaway RM, Walker LR: Competition and facilitation: a synthetic approach to interactions in plant communities. Ecology. 1997, 78 (7): 1958-1965.
Cardinale BJ, Palmer MA, Collins SL: Species diversity enhances ecosystem functioning through interspecific facilitation. Nature. 2002, 415 (6870): 426-429.
Palmer MA, Ambrose RF, Poff NL: Ecological theory and community restoration ecology. Restor Ecol. 1997, 5 (4): 291-300.
Gómez-Aparicio L: The role of plant interactions in the restoration of degraded ecosystems: a meta-analysis across life-forms and ecosystems. J Ecol. 2009, 97 (6): 1202-1214.
Gómez-Aparicio L, Zamora R, Gómez JM, Hódar JA, Castro J, Baraza E: Applying plant facilitation to forest restoration: a meta-analysis of the use of shrubs as nurse plants. Ecol Applic. 2004, 14 (4): 1128-1138.
Aerts R, Negussie A, Maes W, November E, Hermy M, Muys B: Restoration of dry Afromontane forest using pioneer shrubs as nurse plants for Olea europaea ssp. cuspidata. Restor Ecol. 2007, 15 (1): 129-138.
Holmgren M, Scheffer M: El Niño as a window of opportunity for the restoration of degraded arid ecosystems. Ecosystems. 2001, 4 (2): 151-159.
Fukami T, Lee WG: Alternative stable states, trait dispersion and ecological restoration. Oikos. 2006, 113 (2): 353-356.
Holmgren M, Scheffer M, Ezcurra E, Gutiérrez JR, Mohren GMJ: El Niño effects on the dynamics of terrestrial ecosystems. Trends Ecol Evol. 2001, 16 (2): 89-94.
Parrotta JA, Knowles OH: Restoration of tropical moist forests on bauxite-mined lands in the Brazilian Amazon. Restor Ecol. 1999, 7 (2): 103-116.
Parrotta JA, Knowles OH: Restoring tropical forests on lands mined for bauxite: Examples from the Brazilian Amazon. Ecol Engin. 2001, 17 (2-3): 219-239.
Bell SS, Fonseca MS, Motten LB: Linking restoration and landscape ecology. Restor Ecol. 1997, 5 (4): 318-323.
Ehrenfeld JG, Toth LA: Restoration ecology and the ecosystem perspective. Restor Ecol. 1997, 5 (4): 307-317.
Russell AE, Cambardella CA, Ewel JJ, Parkin TB: Species, rotation, and life-form diversity effects on soil carbon in experimental tropical ecosystems. Ecol Applic. 2004, 14 (1): 47-60.
Paul M, Catterall CP, Pollard PC, Kanowski J: Recovery of soil properties and functions in different rainforest restoration pathways. For Ecol Manage. 2010, 259 (10): 2083-2092.
Naeem S, Bunker DE, Hector A, Loreau M, Perrings C, (eds.): Biodiversity, ecosystem functioning and human wellbeing. An ecological and economic perspective. 2009, Oxford: Oxford University Press
Schulze E-D, Mooney HA, (eds.): Biodiversity and ecosystem functioning. 1993, New York: Springer Verlag
Cardinale BJ, Wright JP, Cadotte MW, Carroll IT, Hector A, Srivastava DS, Loreau M, Weis JJ: Impacts of plant diversity on biomass production increase through time because of species complementarity. Proc Natl Acad Sci USA. 2007, 104 (46): 18123-18128.
Naeem S: Ecosystem consequences of biodiversity loss: The evolution of a paradigm. Ecology. 2002, 83 (6): 1537-1552.
Ruiz-Jaen MC, Potvin C: Can we predict carbon stocks in tropical ecosystems from tree diversity? Comparing species and functional diversity in a plantation and a natural forest. New Phyt. 2011, 189 (4): 978-987.
Scherer-Lorenzen M, Schulze ED, Don A, Schumacher J, Weller E: Exploring the functional significance of forest diversity: A new long-term experiment with temperate tree species (BIOTREE). Persp Plant Ecol Evol System. 2007, 9 (2): 53-70.
Pan Y, Birdsey RA, Fang J, Houghton R, Kauppi PE, Kurz WA, Phillips OL, Shvidenko A, Lewis SL, Canadell JG, Ciais P, Jackson RB, Pacala SW, McGuire AD, Piao S, Rautiainen A, Sitch S, Hayes D: A large and persistent carbon sink in the world's forests. Science. 2011, 333 (6045): 988-993.
Thompson ID, Mackey B, McNulty S, Mosseler A: Forest resilience, biodiversity and climate change. Montreal: Secretariat on the Convention of Biological Diversity, Technical Series No. 43. 2009
Paquette A, Messier C: The role of plantations in managing the world's forests in the Anthropocene. Front Ecol Environ. 2010, 8 (1): 27-34.
Vila M, Vayreda J, Comas L, Ibanez JJ, Mata T, Obon B: Species richness and wood production: a positive association in Mediterranean forests. Ecol Lett. 2007, 10 (3): 241-250.
Piotto D, Craven D, Montagnini F, Alice F: Silvicultural and economic aspects of pure and mixed native tree species plantations on degraded pasturelands in humid Costa Rica. New For. 2010, 39 (3): 369-385.
Ruiz-Jaen MC, Potvin C: Tree diversity explains variation in ecosystem function in a neotropical forest in Panama. Biotropica. 2010, 42 (6): 638-646.
Nadrowski K, Wirth C, Scherer-Lorenzen M: Is forest diversity driving ecosystem function and service?. Curr Opin Environ Sust. 2010, 2 (1-2): 75-79.
Potvin C, Mancilla L, Buchmann N, Monteza J, Moore T, Murphy M, Oelmann Y, Scherer-Lorenzen M, Turner BL, Wilcke W: An ecosystem approach to biodiversity effects: Carbon pools in a tropical tree plantation. For Ecol Manage. 2011, 261 (10): 1614-1624.
Gamfeldt L, Hillebrand H, Jonsson PR: Multiple functions increase the importance of biodiversity for overall ecosystem functioning. Ecology. 2008, 89 (5): 1223-1231.
Isbell F, Calcagno V, Hector A, Connolly J, Harpole WS, Reich PB, Scherer-Lorenzen M, Schmid B, Tilman D, van Ruijven J, Weigelt A, Wilsey BJ, Zavaleta ES, Loreau M: High plant diversity is needed to maintain ecosystem services. Nature. 2011, 477 (7363): 199-202.
Hector A, Bagchi R: Biodiversity and ecosystem multifunctionality. Nature. 2007, 448 (7150): 188-190.
De Deyn GB, Shiel RS, Ostle NJ, McNamara NP, Oakley S, Young I, Freeman C, Fenner N, Quirk H, Bardgett RD: Additional carbon sequestration benefits of grassland diversity restoration. J Appl Ecol. 2011, 48 (3): 600-608.
Chazdon RL, Finegan B, Capers RS, Salgado-Negret B, Casanoves F, Boukili V, Norden N: Composition and dynamics of functional groups of trees during tropical forest succession in northeastern Costa Rica. Biotropica. 2010, 42 (1): 31-40.
Eisenhauer N, Yee K, Johnson EA, Maraun M, Parkinson D, Straube D, Scheu S: Positive relationship between herbaceous layer diversity and the performance of soil biota in a temperate forest. Soil Biol Biochem. 2011, 43 (2): 462-465.
Elton CS: The ecology of invasions by animals and plants. 1958, London: Methuen
May RM: Will a large complex system be stable?. Nature. 1972, 238 (5364): 413-414.
Laliberte E, Wells JA, DeClerck F, Metcalfe DJ, Catterall CP, Queiroz C, Aubin I, Bonser SP, Ding Y, Fraterrigo JM, McNamara S, Morgan JW, Merlos DS, Vesk PA, Mayfield MM: Land-use intensification reduces functional redundancy and response diversity in plant communities. Ecol Lett. 2010, 13 (1): 76-86.
Loreau M, Naeem S, Inchausti P, Bengtsson J, Grime JP, Hector A, Hooper DU, Huston MA, Raffaelli D, Schmid B, Tilman D, Wardle DA: Ecology - biodiversity and ecosystem functioning: Current knowledge and future challenges. Science. 2001, 294 (5543): 804-808.
Griffin JN, O'Gorman EJ, Emmerson MC, Jenkins SR, Klein AM, Loreau M, Symstad AJ: Biodiversity and the stability of ecosystem functioning. Biodiversity, ecosystem functioning and human wellbeing. Edited by: Naeem S, Bunker DE, Hector A, Loreau M, Perrings C. 2009, Oxford: Oxford University Press, 78-94.
Pachauri RK, Reisinger A, (eds.): Climate change 2007 synthesis report of the fourth assessment report of the Intergovernmental Panel on Climate Change. 2007, Geneva: IPCC
Hector A, Joshi J, Scherer-Lorenzen M, Schmid B, Spehn EM, Wacker L, Weilenmann M, Bazeley-White E, Beierkuhnlein C, Caldeira MC, Dimitrakopoulos PG, Finn JA, Huss-Danell K, Jumpponen A, Leadley PW, Loreau M, Mulder CPH, Nesshoover C, Palmborg C, Read DJ, Siamantziouras ASD, Terry AC, Troumbis AY: Biodiversity and ecosystem functioning: reconciling the results of experimental and observational studies. Funct Ecol. 2007, 21 (5): 998-1002.
Lloret F, Lobo A, Estevan H, Maisongrande P, Vayreda J, Terradas J: Woody plant richness and NDVI response to drought events in Catalonian (northeastern Spain) forests. Ecology. 2007, 88 (9): 2270-2279.
DeClerck FAJ, Barbour MG, Sawyer JO: Species richness and stand stability in conifer forests of the Sierra Nevada. Ecology. 2006, 87 (11): 2787-2799.
Cadotte MW, Carscadden K, Mirotchnick N: Beyond species: functional diversity and the maintenance of ecological processes and services. J Appl Ecol. 2011
Mouchet MA, Villeger S, Mason NWH, Mouillot D: Functional diversity measures: an overview of their redundancy and their ability to discriminate community assembly rules. Funct Ecol. 2010, 24 (4): 867-876.
Mouillot D, Villeger S, Scherer-Lorenzen M, Mason NWH: Functional structure of biological communities predicts ecosystem multifunctionality. PLoS ONE. 2011, 6 (3):
Bunker DE, DeClerck F, Bradford JC, Colwell RK, Perfecto I, Phillips OL, Sankaran M, Naeem S: Species loss and aboveground carbon storage in a tropical forest. Science. 2005, 310 (5750): 1029-1031.
Naeem S, Bunker DE: TraitNet: furthering biodiversity research through the curation, discovery and sharing of species trait data. Biodiversity, ecosystem functioning and human wellbeing An ecological and economic perspective. Edited by: Naeem S, Bunker DE, Hector A, Loreau M, Perrings C. 2009, Oxford: Oxford University Press, 281-289.
Poorter L, McDonald I, Alarcon A, Fichtler E, Licona JC, Pena-Claros M, Sterck F, Villegas Z, Sass-Klaassen U: The importance of wood traits and hydraulic conductance for the performance and life history strategies of 42 rainforest tree species. New Phyt. 2010, 185 (2): 481-492.
Herault B, Honnay O: The relative importance of local, regional and historical factors determining the distribution of plants in fragmented riverine forests: an emergent group approach. J Biogeogr. 2005, 32 (12): 2069-2081.
Petchey OL, O'Gorman EJ, Flynn DFB: A functional guide to functional diversity measures. Biodiversity, ecosystem functioning, and human wellbeing An ecological and economic perspective. Edited by: Naeem S, Bunker DE, Hector A, Loreau M, Perrings C. 2009, Oxford: Oxford University Press, 49-60.
Bettinger P, Clutter M, Siry J, Kane M, Pait J: Broad implications of southern United States pine clonal forestry on planning and management of forests. Intern For Rev. 2009, 11 (3): 331-345.
Zhu YY, Chen HR, Fan JH, Wang YY, Li Y, Chen JB, Fan JX, Yang SS, Hu LP, Leung H, Mew TW, Teng PS, Wang ZH, Mundt CC: Genetic diversity and disease control in rice. Nature. 2000, 406 (6797): 718-722.
Burdon RD: Genetic diversity and disease resistance: some considerations for research, breeding, and deployment. Can J For Res. 2001, 31 (4): 596-606.
Antonovics J: Towards community genetics. Plant resistance to herbivores and pathogens: ecology, evolution and genetics. Edited by: Frit RS, Simms EL. 1992, Chicago: University of Chicago Press, 426-450.
Hughes AR, Inouye BD, Johnson MTJ, Underwood N, Vellend M: Ecological consequences of genetic diversity. Ecol Lett. 2008, 11 (6): 609-623.
Bailey JK, Schweitzer JA, Ubeda F, Koricheva J, LeRoy CJ, Madritch MD, Rehill BJ, Bangert RK, Fischer DG, Allan GJ, Whitham TG: From genes to ecosystems: a synthesis of the effects of plant genetic factors across levels of organization. Phil Trans Roy Soc B Biol Sci. 2009, 364 (1523): 1607-1616.
Crutsinger GM, Collins MD, Fordyce JA, Gompert Z, Nice CC, Sanders NJ: Plant genotypic diversity predicts community structure and governs an ecosystem process. Science. 2006, 313 (5789): 966-968.
Vellend M, Drummond EBM, Tomimatsu H: Effects of genotype identity and diversity on the invasiveness and invasibility of plant populations. Oecologia. 2010, 162 (2): 371-381.
Schweitzer JA, Bailey JK, Hart SC, Whitham TG: Nonadditive effects of mixing cottonwood genotypes on litter decomposition and nutrient dynamics. Ecology. 2005, 86 (10): 2834-2840.
Madritch M, Donaldson JR, Lindroth RL: Genetic identity of Populus tremuloides litter influences decomposition and nutrient release in a mixed forest stand. Ecosystems. 2006, 9 (4): 528-537.
Sekercioglu CH, Daily GC, Ehrlich PR: Ecosystem consequences of bird declines. Proc Natl Acad Sci USA. 2004, 101 (52): 18042-18047.
Sekercioglu CH: Increasing awareness of avian ecological function. Trends Ecol Evol. 2006, 21 (8): 464-471.
Aerts R, Lerouge F, November E, Lens L, Hermy M, Muys B: Land rehabilitation and the conservation of birds in a degraded Afromontane landscape in northern Ethiopia. Biodiv Conserv. 2008, 17 (1): 53-69.
Lovelock CE, Ewel JJ: Links between tree species, symbiotic fungal diversity and ecosystem functioning in simplified tropical ecosystems. New Phyt. 2005, 167 (1): 219-228.
Verbruggen E, Roling WFM, Gamper HA, Kowalchuk GA, Verhoef HA, van der Heijden MGA: Positive effects of organic farming on below-ground mutualists: large-scale comparison of mycorrhizal fungal communities in agricultural soils. New Phyt. 2010, 186 (4): 968-979.
Dumbrell AJ, Ashton PD, Aziz N, Feng G, Nelson M, Dytham C, Fitter AH, Helgason T: Distinct seasonal assemblages of arbuscular mycorrhizal fungi revealed by massively parallel pyrosequencing. New Phyt. 2011, 190 (3): 794-804.
Kardol P, Wardle DA: How understanding aboveground-belowground linkages can assist restoration ecology. Trends Ecol Evol. 2010, 25 (11): 670-679.
Allen EB, Allen ME, Egerton-Warburton L, Corkidi L, Gomez-Pompa A: Impacts of early- and late-seral mycorrhizae during restoration in seasonal tropical forest, Mexico. Ecol Applic. 2003, 13 (6): 1701-1717.
Peay KG, Garbelotto M, Bruns TD: Evidence of dispersal limitation in soil microorganisms: Isolation reduces species richness on mycorrhizal tree islands. Ecology. 2010, 91 (12): 3631-3640.
Klironomos J, Zobel M, Tibbett M, Stock WD, Rillig MC, Parrent JL, Moora M, Koch AM, Facelli JM, Facelli E, Dickie IA, Bever JD: Forces that structure plant communities: quantifying the importance of the mycorrhizal symbiosis. New Phyt. 2011, 189 (2): 366-370.
White JA, Tallaksen J, Charvat I: The effects of arbuscular mycorrhizal fungal inoculation at a roadside prairie restoration site. Mycologia. 2008, 100 (1): 6-11.
Smith MR, Charvat I, Jacobson RL: Arbuscular mycorrhizae promote establishment of prairie species in a tallgrass prairie restoration. Can J Bot. 1998, 76 (11): 1947-1954.
Caravaca F, Barea JM, Figueroa D, Roldan A: Assessing the effectiveness of mycorrhizal inoculation and soil compost addition for enhancing reafforestation with Olea europaea subsp. sylvestris through changes in soil biological and physical parameters. Appl Soil Ecol. 2002, 20 (2): 107-118.
Caravaca F, Barea JM, Palenzuela J, Figueroa D, Alguacil MM, Roldan A: Establishment of shrub species in a degraded semiarid site after inoculation with native or allochthonous arbuscular mycorrhizal fungi. Appl Soil Ecol. 2003, 22 (2): 103-111.
Chazdon RL: Beyond deforestation: restoring forests and ecosystem services on degraded lands. Science. 2008, 320 (5882): 1458-1460.
Hobbs RJ, Higgs E, Harris JA: Novel ecosystems: implications for conservation and restoration. Trends Ecol Evol. 2009, 24 (11): 599-605.
Lugo AE: The emerging era of novel tropical forests. Biotropica. 2009, 41 (5): 589-591.
Hall JS, Ashton MS, Garen EJ, Jose S: The ecology and ecosystem services of native trees: Implications for reforestation and land restoration in Mesoamerica. For Ecol Manage. 2011, 261 (10): 1553-1557.
Davis MA, Chew MK, Hobbs RJ, Lugo AE, Ewel JJ, Vermeij GJ, Brown JH, Rosenzweig ML, Gardener MR, Carroll SP: Don't judge species on their origins. Nature. 2011, 474 (7350): 153-154.
Aerts R, Honnay O: Seeds of change for restoration ecology. Science. 2011, 333 (6039): 156-
This review was written on the occasion of the United Nations International Year of Forests 2011. RA holds a post-doctoral fellowship of the Research Foundation - Flanders (FWO). The support of Wim Aertsen and the constructive comments by Hans Jacquemyn, Robin Chazdon and two anonymous reviewers are gratefully acknowledged.
The authors declare that they have no competing interests.
RA and OH contributed equally to the design of the review. The authors drafted the manuscript together. Both authors read and approved the final manuscript.
Raf Aerts and Olivier Honnay contributed equally to this work.
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Aerts, R., Honnay, O. Forest restoration, biodiversity and ecosystem functioning. BMC Ecol 11, 29 (2011). https://0-doi-org.brum.beds.ac.uk/10.1186/1472-6785-11-29
- Ecosystem Service
- Arbuscular Mycorrhizal Fungus
- Ecosystem Function
- Ecosystem Functioning
- Forest Restoration