Abstract Ecosystem services (ES) are the benefits that people receive from ecosystems. Understanding the impact of forest management on their supply can inform policy and practice for meeting societal demand. The objectives of this paper are to identify and review the effect of management intensity on priority ES supply and identify synergies and trade-offs among ES of different management approaches. We review seven priority ES identified from UK land and forestry strategy and policy documents: timber/biomass production, carbon storage, biodiversity, health and recreation, water supply and quality and flood protection. We present a synthesis of the impact of management on relative ES supply. It shows that low intensity management is unsuitable for high biomass production, yet provides high or moderately high levels of other services. Higher intensity management impacts negatively on biodiversity, health and recreation and water supply services. Combined objective forestry provides high or moderately high levels for all services except biomass. We recommend that a diversity of management approaches is needed to maintain multiple ecosystem service provision. The ES framework offers opportunities to forest management by revealing areas of conflict or co-production and potential trade-offs that may arise from adjusting management intensity. Introduction Forests provide a broad range of environmental, economic and social benefits. Framing these benefits as ecosystem services (ES) is a way of linking ecosystem functions to human wellbeing and the value of nature to society (Turner and Daily, 2007; Braat and De Groot, 2012). The Millennium Ecosystem Assessment chapter on forest and woodland systems discusses the diversity of forest services for all forest types that includes fibre, fuel and non-timber forest products, carbon sequestration, biodiversity, soil and water protection and socio-cultural values (Shvidenko et al., 2005). Recognizing their value, and maintaining or increasing their supply is an important objective for forest managers to meet society’s demands now and into the future. Recent research has shown that a focus on ES provision as part of the wider ecosystem approach (CBD SBSTTA, 2000) is emerging as the new paradigm for forestry, which overlaps with existing sustainable forest management approaches (Kline et al., 2013; Quine et al., 2013; Winkel, 2014; Raum and Potter, 2015). ES have been embedded in the Convention on Biological Diversity’s Strategic Plan for Biodiversity and Aichi Biodiversity Targets in 2010 (Nayar, 2010), further establishing their relevance for natural resource managers. The ES that a forest provides will vary according to a range of variables. Research has examined the effects of species composition (Gamfeldt et al., 2013) including monocultures (Mason and Connolly, 2014), silvicultural treatments (Biber et al., 2015), stand age (Luyssaert et al., 2008) and harvesting techniques (Walmsley et al., 2009; Clarke et al., 2015) on ES provision. Natural disturbances will also impact on the ability of forests to provide goods and services (Boyd et al., 2013; Thom and Seidl, 2016). Lindner et al. (2010) discuss the potential climate change impacts of increasing temperature and atmospheric CO2 levels which are favourable to tree growth in northern and western Europe, but are offset by increasing disturbance and drought risks in southern and eastern regions for wood production. A focus on ES can support integrated, sustainable and equitable natural resource management (Potschin and Haines-Young, 2011; Seppelt et al., 2011) by delivering evidence to natural resource managers on the impacts of their decisions for meeting societal demands and raise awareness of trade-offs among ES. Making trade-offs explicit is widely regarded as a ‘core function’ of ES research (Carpenter et al. 2009; Daily et al. 2009). Forest managers are required to maintain and enhance the sustainability of a range of environmental, social and economic benefits while faced with future climate, socio-economic, political and environmental uncertainty and the decisions they make will have implications on the suite of ES that forests provide. For example Turner et al. (2011) demonstrate the positive effect of reducing logging intensity following the North West Forest Plan for species conservation on carbon sequestration. This has also been explored for several case studies including the UK (Ray et al., 2015), Germany (Frank et al., 2015) and Sweden (Zanchi et al., 2014), where alternative management strategies result in differing ES provision. Duncker et al. (2012b) quantified the effects of management for virtual forest landscapes in Europe which showed the conflicts between managing for biomass production, water quality and biodiversity. Yet the implementation of ES knowledge at the operational level remains a challenge (De Groot et al., 2010; Braat and De Groot, 2012). Institutional barriers such as conventional economic growth objectives, and cultural entrenchment in traditional forest management (Kelly, 2014) restrict the operationalization of new knowledge from ES in decision-making (Cowell and Lennon, 2014; Jordan and Russel, 2014; Waylen and Young, 2014). To transition from science to policy and practice, the ES framework requires greater comprehension and clear methods, together with support from the research community. Several initiatives, including the Intergovernmental Panel on Biodiversity and Ecosystem Services (IPBES; http://www.ipbes.net accessed 8 September 2017), The Economics of Ecosystems and Biodiversity (TEEB; http://www.teebweb.org accessed 8 September 2017), the Ecosystem Services Community Scotland (ESCOM; http://escom.scot accessed 8 September 2017) and the Belgium Ecosystem Services community of practice (BEES; http://www.beescommunity.be/en/ accessed 8 September 2017), aim to address this by improving dialogue among research, policy and practice. The objectives of this paper are (1) to identify and review the effect of management intensity on priority forest ES supply and (2) identify synergies and trade-offs among ES of different management approaches. As foresters are required to consider a broad set of services in their management decision-making, and to improve ES knowledge in decision-making, we have undertaken a literature review to evaluate the effects of management intensity for Britain’s temperate deciduous, conifer and mixed forests. We also identify decision support tools which have been developed to provide evidence of ES for decision-makers. We then synthesize the current understanding of management practice impacts on ES provision in a matrix to inform practitioners of how their decisions affect ES provision and the trade-offs among ES that may result. These trade-offs are internationally relevant as temperate broadleaf/mixed forests and temperate conifer forests are important forest biomes (Olson et al., 2001). Method Forest policy is the articulation of a society’s values for its forests and how those values should be realized (McDermott et al., 2010). In the UK, forest policy has been devolved to the governments of Scotland, Wales and Northern Ireland, while English policy is retained by the UK Government. Taking policy as an indicator of national scale ES demand, we identified the priority ES for UK forests based on the frequency with which they are referred to within the national forest and land use policy documents of England (Department for Environment Food and Rural Affairs, 2013), Wales (Welsh Assembly Government, 2009), Scotland (Scottish Executive, 2006; The Scottish Government, 2016) and Northern Ireland (Forest Service NI 2006, 2009). These were cross-referenced with a sample of English language forest policy documents for other European countries (Germany, Switzerland, Netherlands and Ireland) to identify trans-national themes (Bundesministerium fur Ernahrung Landwirtschaft und Verbraucherschutz, 2011; Federal Office for the Environment, 2013; Department of Agriculture Food and the Marine, 2014; Ministry of Economic Affairs, 2014). We conducted a literature search for each ES, selecting papers that focus on the impact of management for temperate forests internationally. Papers were identified from a Web of Science literature search using the search criteria ‘forest management’ and each ecosystem service identified in the policy documents. Papers were selected based on their relevance to temperate conifer and broadleaved forests. The impact on ES supply was evaluated on a scale from high to low according to the type of management described. We have used the ‘Forest Management Approach’ (FMA) typology as described by Duncker et al. (2012a) that classifies management into five FMA classes of increasing management intensity (Table 1): (1) unmanaged forest reserve, (2) close-to-nature forestry, (3) combined objective forestry, (4) intensive even-aged forestry, and (5) short rotation forestry. Table 1 Description of Forest Management Approaches (FMAs), arranged in increasing management intensity, following Duncker et al. (2012a). FMA Description Management intensity 1 Unmanaged forest reserve Interventions restricted to recreation provision (e.g. car parks and trails) and protection from browsing 2 Close-to-nature Interventions mimic natural processes, e.g. long rotation lengths, harvesting by single stem and group selection. 3 Combined objective Limited interventions to deliver timber and other ecosystem services in an integrative (not zoned) approach. Longer rotations at or above maximum mean annual increment. 4 Intensive even-aged Interventions follow production goals: single-aged monocultures or small per cent admixture species, typically harvested by clearcut. Rotation lengths at or below maximum mean annual increment. 5 Short rotation forestry Intensive management for maximum biomass production, single-aged monocultures grown on short rotations (~20 years), whole tree harvesting to maximize volume FMA Description Management intensity 1 Unmanaged forest reserve Interventions restricted to recreation provision (e.g. car parks and trails) and protection from browsing 2 Close-to-nature Interventions mimic natural processes, e.g. long rotation lengths, harvesting by single stem and group selection. 3 Combined objective Limited interventions to deliver timber and other ecosystem services in an integrative (not zoned) approach. Longer rotations at or above maximum mean annual increment. 4 Intensive even-aged Interventions follow production goals: single-aged monocultures or small per cent admixture species, typically harvested by clearcut. Rotation lengths at or below maximum mean annual increment. 5 Short rotation forestry Intensive management for maximum biomass production, single-aged monocultures grown on short rotations (~20 years), whole tree harvesting to maximize volume The area of woodland in the UK is 3.16 million hectares, ~13 per cent of the land area. This is one of the lowest percentages of forest cover in Europe, where the average forest area is 46 per cent (including the Russian Federation). The forests of Wales and Northern Ireland have approximately equal areas of broadleaved and conifer forests, whereas England’s forests are predominantly broadleaved (76 per cent) and Scotland’s forests are predominantly conifers (76 per cent) (Forestry Commission, 2016). Britain’s conifer forests are dominated by Sitka spruce (Picea sitchensis), which accounts for approximately half the total standing coniferous volume and stocked area. Oak (Quercus spp), beech (Fagus sylvatica) and ash (Fraxinus excelsior) are the principle broadleaved species by standing volume (Forestry Commission, 2011, 2013). The dominance of conifers in upland areas is a result of twentieth century afforestation to stimulate a UK based timber processing sector. Sitka spruce plantations are predominantly managed as FMA 4 intensive even-aged stands on short (up to 50 year) rotations (Mason and Perks, 2011). However current forest and land use policies are directing management towards alternative strategies, to increase resilience to the threats of climate change, pests and diseases, and to deliver a greater range of benefits. The potential forest management map for British forests (Hengeveld et al., 2012) suggests large areas are suitable for FMA 3 in Scotland with areas potentially suitable for FMA 5, while England has more forest suited to FMA 1 and 2. Wales and Northern Ireland have a mixture of FMA 2, 3 and 4. Although it is not widespread in the UK at present, FMA 5 (short rotation forestry) is a potential future management option to deliver bioenergy to contribute towards achieving climate mitigation targets (Moffat et al., 2010). The most frequently cited ES for the consulted policy documents in the UK are climate mitigation (carbon storage), timber and woodfuel supply, biodiversity, water quality, recreation, soil protection and flood protection (Table 2). Air quality is another frequently cited regulating service; this is more directly attributed to urban trees and therefore has not been included in this review. Comparison with other forestry policy and strategy documents for Germany, Switzerland, Ireland and Netherlands showed strong agreement across all ES apart from flood protection. Table 2 A set of priority ecosystem services Britain’s forests, identified from national forest and land use policy and strategy documents. 1Department for Environment Food and Rural Affairs (2013). 2Welsh Assembly Government (2009). 3Scottish Executive (2006), The Scottish Government (2016). 4Forest Service NI (2006, 2009). 5Bundesministerium fur Ernahrung Landwirtschaft und Verbraucherschutz (2011). 6Federal Office for the Environment (2013). 7Department of Agriculture Food and the Marine (2014). 8Ministry of Economic Affairs (2014). 9Canadian Council of Forest Ministers (2008). Impact of management on the supply of priority ecosystem services Fibre: timber and biomass production Historically, intensive management for timber production (FMA 4) has been characterized by single species conifer plantations harvested under patch clearcutting on short (<50 year) rotations (Mason and Perks, 2011), with rotation lengths intended to maximize site productivity (Burger, 2009). However, countries in central Europe are increasingly moving to close-to-nature continuous cover systems (FMA 2 and 3) in regions where endemic wind damage is less severe (Troen and Petersen, 1989). A shift to close-to-nature forestry has been shown to increase variation in log size, which may impact on timber processing sector without significantly affecting timber quality, while edge effects created under shelterwood systems may negatively affect timber quality (Macdonald et al., 2010). Computer simulations of future productivity have shown that overall yield is likely to fall with a shift to lower intensity management systems (FMA 1–2) (Duncker et al., 2012a; Ray et al., 2015). Removal of harvest residue, stump and thinnings under FMA 5 for biomass production increases production volume; however, the reduction in site nutrition can affect second rotation growth (Walmsley et al., 2009; Wall, 2012). Recent increases in domestic demand for woodfuel in developed countries are expected to continue, in part driven by bioenergy policies for reducing greenhouse gas emissions (Söderberg and Eckerberg, 2013). Thus, increasing the area of woodland under management and increasing woodland cover are two approaches identified by policymakers to stimulate supply in response to increasing demand, and to offset the decreased production that results from policies encouraging combined objective and close-to-nature forestry. For instance, broadleaves can deliver additional wood products (Bentrup et al., 2009; Deal et al., 2013). European forests have the capacity to increase harvesting intensity: currently ~40 per cent of the annual increment of Europe’s forests is harvested (FOREST EUROPE UNECE and FAO, 2011). However designations for other ES that limit management intensity to FMA 1 and 2 have been shown to restrict wood harvesting: Verkerk et al. (2014) calculated that 10 per cent of the annual theoretical potential wood supply is unavailable due to protection status in Europe’s forests. Climate mitigation: carbon storage Forest carbon (C) is stored above ground within the trees, ground vegetation and litter, and below ground in the roots and soil (Lal, 2005). The relationship between stand age and carbon sequestration is dependent on the time over which carbon is sequestered and the size of the tree (Harrison et al., 2014). Forest carbon stocks fluctuate over the lifecycle of a stand (Lippke et al., 2011), peaking during the full vigour growth phase and falling post-harvest through removal of wood material, reduction in litter and soil carbon released by disturbance (Morison et al., 2012). Increasing site productivity results in higher levels of on-site carbon accumulation (Jandl et al., 2007; Morison et al., 2012), including use of improved forest stock (Mason and Perks, 2011). Yet rotation lengths at or below the mean maximum annual increment for biomass and timber production (FMA 3, 4 and 5) result in lower long-term forest carbon stocks. This is due to the relationship between tree size and duration of growth during the full vigour phase. Conversely forests containing deadwood and trees that are grown into mature and old growth phases (FMA 1 and 2) have been shown to store more carbon (Luyssaert et al., 2008; D’Amato et al., 2011; Fortin et al., 2012; Harrison et al., 2014). Empirical modelling has shown that Low Impact Silvicultural Systems such as Continuous Cover Forestry (FMA 2) that contain greater age diversity can provide greater long-term carbon stocks compared with single age stands (Duncker et al., 2012b; Ray et al., 2015). However, lower carbon stocks have been recorded in stands with lower basal area intended to promote natural regeneration (Mason and Perks, 2011). Transforming single-aged stands to increase structural diversity also increases the risk of wind disturbance with consequences for the loss of standing carbon stocks (Mason and Perks, 2011). In addition to controlling stand age structure, forest management influences carbon stocks directly, through thinning and harvesting and indirectly through influencing growth conditions, for example by fertilization (Nabuurs et al., 2008). Losses of soil carbon through harvesting are soil-, species- and practice-dependent (Clarke et al., 2015), for example whole tree harvesting reduces soil carbon (Johnson and Curtis, 2001). A number of studies have shown the link between reduced harvest levels and greater on-site carbon storage (Bradford et al., 2013; Man et al., 2013), and alternative residue management and site preparation can mitigate losses (Jandl et al., 2007; Nave et al., 2010). In addition, forest management strategies can reduce carbon losses by lowering the risk of tree mortality due to natural disturbances such as storms, drought and fire (Galik and Jackson, 2009; McKinley et al., 2011; Nave et al., 2011; Law and Waring, 2015). Carbon storage benefits are also provided as C pools existing in harvested wood products, or though fossil fuel substitution, either through supply of bioenergy or construction materials (Nabuurs et al., 2008; Lippke et al., 2011). The additional long-term carbon storage benefits of these products depends on the lifecycle of wood products and consumer uptake (Markewitz, 2006; Suttie et al., 2009). Shifting to shorter rotations (FMA 5) will increase the supply of biomass for bioenergy, though this may potentially compete with the supply of traditional wood products (Malmsheimer et al., 2011). Carbon has become a monetized commodity, with voluntary carbon offset markets trading the additional sequestration benefits resulting from afforestation/reforestation and improved forest management (IFM) schemes in private ownership. Carbon accounting tools have been developed to support these markets to calculate the carbon in harvested wood products (PRoduct EStimation Tool Online, PRESTO; http://www.nrs.fs.fed.us/pubs/47240 accessed 8 September 2017) and future carbon from afforestation projects (UK Woodland Carbon Code; http://www.forestry.gov.uk/carboncode accessed 8 September 2017). However, fluctuations in carbon unit price linked to policy signals (Forest Trends, 2015) together with price and land value uncertainty affect financial returns and land owner decision making (Reeson et al., 2015). Biodiversity Forests provide a range of habitats, micro-habitats and niches for woodland species, delivering a critical underpinning role for other ES (Harrison et al., 2014) including forest productivity (Ishii et al., 2004; Thompson et al., 2011). Biodiversity within the forest tends to be greater in stands that are structurally diverse in terms of their age, species, patch edge, understory and deadwood component (Gilliam, 2007; Moning et al., 2009; Lassauce et al., 2012a; Bereczki et al., 2014; Gao et al., 2014; Larrieu et al., 2014; Humphrey et al., 2015). Patch size and quality are important for maintaining populations in fragmented landscapes (van Halder et al., 2015; Humphrey et al., 2015). Management intensity has been shown to influence species richness and abundance, with species dependent on the continuity of forest cover, deadwood and large trees negatively affected by more intensive management (FMA 4 and 5) (e.g. Halpern and Spies, 1995; Paillet et al., 2010; Summerville, 2013), such as long-term losses in species diversity and abundance. Research conducted on avian populations and intensive forest management found a negative impact caused by greater herbicide use (Betts et al., 2013). Forest management that mimics natural disturbances (FMA 2 and 3) delivers greater biodiversity benefits through diversifying species and age classes of even-aged stands (Lindenmayer et al., 2006; Ares et al., 2010), though this may be at the expense of reduced timber production (Brockerhoff et al., 2008; Deal et al., 2013). Yet non-native plantation forests (FMA 4) also deliver biodiversity benefits, by buffering native forest remnants and enhancing landscape connectivity where native woodland is scarce, in addition to providing habitat in the landscape (Humphrey et al., 2000; Brockerhoff et al., 2008; Quine and Humphrey, 2010; Irwin et al., 2014; Procter et al., 2015). Species mixtures also enhance biodiversity in plantation forests (Oxbrough et al., 2016). There is evidence of fluctuations in the biodiversity benefit of single species plantations over time: young eucalyptus plantations have been shown to provide habitat for shrubland species (Calviño-Cancela et al., 2012), whereas old-growth spruce stands provide habitat for native fungi (Humphrey et al., 2000). Clearcut harvesting (FMA 3, 4 and 5) results in short-term loss of biodiversity prior to recolonization from nearby mature stands; this can be mitigated by adjusting clearcut practice to retain mature trees (Deal et al., 2013; Baker et al., 2015). Shelterwood systems can enhance habitat for some species (Goodale et al., 2009; Summerville, 2013), but not others (Newell and Rodewald, 2012; Nascimbene et al., 2013). However removal of harvest residues negatively affects biodiversity by reducing the availability of deadwood habitats (Lassauce et al., 2012b). At the landscape scale, management actions to improve functional conditions for biodiversity can be achieved by enhancing the condition and/or size of existing forest patches (e.g. afforestation buffering existing woodlands) and reducing isolation between patches (Honnay et al., 2002; Bailey, 2007; Humphrey et al., 2015). Financial mechanisms that provide incentives to manage for biodiversity benefits have been criticized amid concern of making public goods commodities for financial gain (Muradian et al., 2013; Schröter et al., 2014). Nevertheless, working forest conservation easements in the USA have been shown to increase ES supply from private forests (Sedjo, 2007). A conservation easement is a legal agreement between a land owner and trust or governmental body that limits the uses of the land to protect specified conservation values or public ES benefits by compensating for the loss of revenue from reduced timber harvesting. Health and recreation Recreation within natural environments provides health benefits through physical activity, social interaction and mental restoration (Bowler et al., 2010; Hartig et al., 2011; O’Brien and Morris, 2014; Boncinelli et al., 2015; Bratman et al., 2015). Recent evidence has revealed the human health benefit from the exposure to weak concentrations of volatile organic chemicals and other compounds released by trees that stimulate the body and its immune system (Moore, 2015). Naturally, access is key to the realization of these health and well-being benefits (O’Brien and Morris, 2014). Woodlands that are located close to population centres have higher visitor numbers, as the time and cost of travel to sites influences frequency of visits (Cho et al., 2014). Consequently, there has been a programme of targeted planting and management of woodlands close to population centres delivered through the ‘Woodlands In and Around Towns’ initiative in Scotland, the results of which are still being researched (Silveirinha de Oliveira et al., 2013), but for which initial results showed positive impacts for local communities (Ward Thompson et al., 2010). There are synergies with timber production where forest roads provide access for recreation (Harshaw and Sheppard, 2013), particularly consumptive activities (Hunt et al., 2010). Social research methods have been used to understand how management affects recreational value (Boxall and Macnab, 2000; Horne et al., 2005; Christie et al., 2007; Tyrväinen et al., 2013; Schmidt et al., 2016). Willingness to pay (WTP) elicited from choice experiments are influenced by a range of variables that includes forest type, location and survey method (Barrio and Loureiro, 2010), and results highlight the heterogeneity in forest management preferences within and between user groups (Christie et al., 2007; Berninger et al., 2010; Hunt et al., 2010). In general, however, studies have shown public preference for more diverse, open forest structures that appear more accessible and lower impact silvicultural systems associated with FMA 2 and 3 than single species, single-aged stands managed on patch clearcut systems FMA 4 and 5 (Scarpa et al., 2000; Christie et al., 2007; Carvalho-Ribeiro and Lovett, 2011; Edwards et al., 2012; Petucco et al., 2013). Choice experiments of forest management practices have revealed preferences for close-to-nature management, including contoured forest edges, species diversity, vertical layering and older trees (Hanley et al., 1998; Horne et al., 2005; Christie et al., 2007; Juutinen et al., 2014; Giergiczny et al., 2015). Water resources: supply and quality The impact of intensively managed plantations (FMA 4 and 5) on water supply through interception losses and use is difficult to quantify due to a wide range of factors, such as stand age, soil type and condition and wider landscape factors, which influence water yield and confound experiment results (Sahin and Hall, 1996; Stednick, 1996; Marc and Robinson, 2007). Species choice has been shown to affect water supply, in particular the higher interception losses of conifers compared to broadleaved species (Keenan and Van Dijk, 2010), and the higher water demand from fast growing species such as eucalyptus (Nisbet et al., 2011; Ellison et al., 2012). In their overview paper, van Dijk and Keenan (2007) set out the current understanding of the effects of planted forests on water, focusing on the impact on water resources and other water-related issues in agricultural landscapes. Forest water use tends to be higher than for non-irrigated agricultural crops, leading to reduced annual flows from catchments, and fast growing plantation species have been found to cause major reductions in catchment flows (Calder, 2007). Water draining from forests is generally of high quality (Kauffman and Belden, 2010), and forests are used to protect water quality around the world; in fact, many of the world’s largest cities rely on water draining from forest protected areas (Dudley and Stolton, 2003). However, forest management practices may have a detrimental impact on water quality by increasing diffuse pollution, soil disturbance resulting from cultivation, drainage, road construction and harvesting operations can increase turbidity and sedimentation (Brown and Binkley, 1994). There is also a risk of nutrient runoff, particularly in fertilized areas (Nisbet, 2001). Diffuse pollution from fertilization applications (Holland et al., 2015) and pollutant scavenging by forest canopies (Nisbet et al., 2011) can result in further acidification of surface waters, particularly in areas with sensitive geology and mature conifer forests (Nisbet, 2001). Best management practices, such as the use of riparian buffers of native tree species along watercourses and species selection for site conditions have been shown to reduce these impacts (Lowrance et al., 1984; Nisbet, 2001; Broadmeadow and Nisbet, 2004; McBroom et al., 2008; Kuglerová et al., 2014). In practice, less intensive management approaches (FMA 1, 2 and 3) such as stand restructuring with broadleaved species, reduced pesticide use and low-impact silvicultural systems have been the principal approaches for achieving good quality drinking water. Research from other areas demonstrates the links between management and benefit provision. For example, forested catchments have reduced water treatment costs in the USA (Postel and Thompson, 2005) and France (Fiquepron et al., 2013) relative to other land uses. Tools exist to support management decision-making (Zhang and Barten, 2009) and identification of catchments (Weidner and Todd, 2011). Ownership is a major route to delivery, with forests that are either state-owned or owned by water utility companies delivering drinking water for towns and cities across Europe and North America (Turner and Daily, 2007; Neary et al., 2009; Richards et al., 2012; Fiquepron et al., 2013; Blanchard et al., 2015), though not always successfully (Herbert, 2007). Many provide additional recreation benefits, though in some cases access to forests is restricted (Dudley and Stolton, 2003). Incentives for private owners to plant new woodlands or reduce management intensity (reducing felling coupes and the use of fertilizers and pesticides) for drinking water provision have been used in a number of regions (International Union for Conservation of Nature, 2009). Hazard regulation: flood protection Forests can provide flood protection benefits at the local scale through the higher infiltration capacity of soils under trees (Marshall et al., 2014), and the hydraulic roughness of floodplain and riparian woodlands which slows peak flows and enhances storage (Sakals et al., 2006; Nisbet and Thomas, 2008). The higher water use of conifer forests, and particularly the higher interception loss, provides some scope for flood reduction in poorly drained soils (Robinson et al., 2003; Nisbet, 2005; Keenan and Van Dijk, 2010), although the effect reduces with increasing storm size (Nisbet et al., 2011). Riparian woodlands are a natural source of large woody debris vital for the formation of dams and pools which increase upstream storage (Linstead and Gurnell, 1998). However this debris can contribute to flooding by blocking bridges and culverts in flood prone areas (Broadmeadow and Nisbet, 2004). Brash from clearcut sites can make a similar contribution to improving storage upstream during flood peaks (Robinson et al., 2003), while at other times it can fill dams and block upstream fish migration (Broadmeadow and Nisbet, 2004). However, as for water supply and quality, the impact of forests on flood protection at the catchment scale is difficult to quantify, due to the contributions of different land use and land cover across a catchment. Studies have shown that the ability of forests and woodlands to attenuate peak flows in streams and rivers occurs only for smaller flood events in small catchments (Robinson et al., 2003; O’Connell et al., 2004). However a recent review indicates greater benefits of up to 19 per cent reduction in peak flows from natural flood management (Dixon et al., 2016). Forest management practices may also deliver disbenefits (Calder, 2007). For example, forest roads, drainage channels created to drain wet soils for conifer plantations and clearcut harvesting (FMA 4 and 5) have been shown to increase peak flows (Robinson et al., 2003; Neary et al., 2009). Active management of riparian buffer zones in flood prone watersheds (FMA 2 and 3) and around plantations reduces the risks associated with large woody debris from unmanaged stands which can block channels and contribute to flooding or affect other users including migrating fish (Broadmeadow and Nisbet, 2004; Keenan and Van Dijk, 2010). Riparian planting has been undertaken as part of an integrated approach to flood alleviation in the UK, where flood alleviation is a policy priority (Nisbet et al., 2015). Synergies and trade-offs Based on the literature we constructed an impact matrix (Table 3) to capture how management intensity affects ecosystem services supply. We have then used this matrix to plot the relative supply of each ecosystem service with increasing management intensity (Figure 1). The matrix and figure demonstrate some of the likely trade-offs and synergies among ES for a particular management approach, as well as the potential outcomes in ES supply from changing (either increasing or reducing) management intensity. Where forest stands are managed as intensive even-aged plantations or short rotation forestry with single species for timber and biomass production (FMA 4 and 5), they also deliver local- to global-scale benefits through carbon sequestration, increased water supply, enhanced slope stability and provision of recreation. However interventions such as harvesting, road construction and site preparation cause disturbances which can reduce slope stability, affect water quality, release carbon, impact on biodiversity and restrict recreational use (Jandl et al., 2007; Duncker et al., 2012b). There is also the potential impact of fast grown species on increased water use within catchments (Başkent et al., 2010; Chisholm, 2010; Dymond et al., 2012). Clearcut harvesting disturbs or damages habitats (Deal et al., 2013; Baker et al., 2015), while monocultures of non-native species have lower biodiversity value (Halpern and Spies, 1995). Table 3 Impact matrix of the effects of management intensity on the supply of priority ecosystem services, using the Forest Management Approaches classification system (Duncker et al., 2012a). Figure 1 View largeDownload slide Impact of forest management intensity on the relative supply of priority ecosystem services, using the Forest Management Approaches classification system (Duncker et al., 2012a). Carbon is ‘in-forest’ carbon stocks and does not account for carbon stored in harvested wood products and the substitution of fossil fuels. Estimated change in relative supply based on the average rate of C accumulation in Sitka spruce, values not available for FMA 5 (Morison et al., 2012). Figure 1 View largeDownload slide Impact of forest management intensity on the relative supply of priority ecosystem services, using the Forest Management Approaches classification system (Duncker et al., 2012a). Carbon is ‘in-forest’ carbon stocks and does not account for carbon stored in harvested wood products and the substitution of fossil fuels. Estimated change in relative supply based on the average rate of C accumulation in Sitka spruce, values not available for FMA 5 (Morison et al., 2012). Conversely, where the management intensity of existing forests is lower (FMA 2 and 3) to achieve other objectives such as biodiversity, drinking water or natural hazard protection, there is a shift in the suite of ES supplied. Forests managed for local and regional drinking water supply and natural hazard protection areas are necessarily located close to the demand for the service. These forests provide favourable conditions for biodiversity and recreation, though in some cases access to forests is restricted (Dudley and Stolton, 2003). The continuity of forest cover on these sites maintains carbon sequestration benefits over the long term. For example, in North America, reducing the intensity of timber production to deliver biodiversity protection for rare species in the North West Forest Plan, USA, increased carbon sequestration, though at the expense of timber and woodfuel supply and regional employment (Eichman et al., 2010; Turner et al., 2011). The relationship between carbon and timber production is widely discussed in the literature. There are synergies from tree growth accumulating carbon which remains a store for the lifespan of harvested wood products, as well as from fossil fuel substitution in energy and construction (Lippke et al., 2011; Malmsheimer et al., 2011; McKinley et al., 2011). Increasing site productivity results in higher levels of on-site carbon accumulation. However trade-offs occur as a result of forest operations reducing in situ carbon pools (Jandl et al., 2007; Seidl et al., 2007; Nave et al., 2010; Nunery and Keeton, 2010; McKinley et al., 2011; Vanhala et al., 2013), while reducing harvesting intensity to increase carbon pools impacts on timber production (Seidl et al., 2007; Schwenk et al., 2012). Bellassen and Luyssaert (2014) propose ‘win-win’ management strategies to increase both timber production and forest carbon stocks by protecting trees from herbivory and replacing low productivity forests. However, site selection for such a strategy should consider the impacts on other ES, such as species movement across the landscape as a result of fencing, or the other benefits that these low productivity forests are delivering. Discussion Identifying, mapping and quantifying ES supply and demand provides an interesting opportunity to inform forest management by highlighting the trade-offs between services from different management actions, and the possible consequences of adjusting management intensity on the provision of different ES. Mapping approaches can identify hotspots of ES supply and demand to aid in delivering targeted forest management and operations (Gimona and Van Der Horst, 2007; Gonzalez-Redin et al., 2016), by revealing areas of conflict or areas of co-production of two or more ES. Spatial prioritization of management actions can assist in mitigating ES trade-offs. Two alternative spatial approaches are segmentation and integration (Simončič et al., 2015). Where trade-offs occur between timber and biomass production and other priority services, spatial prioritization using a segmentation approach through protective designations protects highly valued social benefits such as biodiversity conservation, drinking water provision and natural hazard protection by limiting the intensity of forest management (Dudley and Stolton, 2003; Simončič et al., 2013). In contrast, where the synergies outweigh the benefits, for example where protective designations are not required as the ES values are lower, an integrative approach for multiple benefits is used. For example where recreation negatively impacts on biodiversity and habitat quality, such as trails affecting ground nesting birds, forest recreation could be zoned to maximize disturbance-free habitat (Thompson, 2015). The ability of ES concepts to influence decision-making will be dependent on the new insights they offer compared to the current ‘business as usual’ approach (Bagstad et al., 2013). Empirical and knowledge based models have been integrated with scenarios to compare the provision of ES under alternative climate change and forest management scenarios (Fürst et al., 2013; Petr et al., 2014; Frank et al., 2015; Ray et al., 2015) to test their robustness and resilience. This approach is particularly suitable for forest management due to the long time scales of forest planning. Scale is an important dimension to consider, since the supply of ES takes place at a range of spatial scales and time scales. Spatially, ecosystem services are supplied to beneficiaries at scales that vary from local to global, so that natural resource planning and management decisions which are generally made at the local level have impacts on the benefits received locally, regionally, nationally and internationally (Hein et al., 2006). For example, landscape scale analysis may be more appropriate for species when considering biodiversity, depending on species requirements and dispersal ability. Carbon storage is likely to be more useful when scaled up to regional or national scale, and should include the lifecycle of harvested wood products and fossil fuel substitution effects. Research incorporating multi-scale modelling can fill these knowledge gaps (Seidl et al., 2013; Seppelt et al., 2013). The impact matrix of management on ecosystem services used to synthesize this review (Table 3) does not include the impacts of afforestation and reforestation, which is widely discussed and identified in policy as a method to increase ES supply. However changing land use creates trade-offs that depend on the existing land use that is replaced, such as lowered food production (McKinley et al., 2011; Whitehead, 2011), as well as the success of new planting schemes (Thomas et al., 2015). Certain land uses that have high policy priority, including prime agricultural land and peatlands, have been ring-fenced and protected from large-scale afforestation projects in Scotland on this basis (Sing et al., 2013). A cross-sectoral analysis can explore the ES gains and losses for alternative land use scenarios. Indeed, if ES are to deliver landscape-scale benefits there needs to be a shift away from single sector governance (Quine et al., 2013). Currently, there is a mismatch between ecosystem processes and existing governance structures and decision-making processes (Primmer and Furman, 2012). Changes to environmental policies and governance structures will be required which cut across traditional single sector approaches to natural resource management (Carvalho-Ribeiro et al., 2010; Everard et al., 2014). Such changes are being explored, such as the nature-based solutions approach in the Netherlands (Ministry of Economic Affairs, 2014) and the Scottish Land Use Strategy (Scottish Government, 2011). While we have identified national priority ES for forest management in Table 1, we also recognize that there are likely to be different ES that are relevant at regional and local scales, particularly cultural services. For example forested catchments that deliver drinking water regionally also deliver recreation provision locally (e.g. Blanchard et al., 2015). Where this is the case, it will be important that the additional ES that local beneficiaries require or desire do not become marginalized in decision-making processes. While sustainable forest management recognizes the social, economic and environmental dimensions of forests, ES places a greater emphasis on beneficiaries and their temporal and spatial characteristics (Burkhard et al., 2012; Bagstad et al., 2014). This is an important strength given the long time scales over which forests grow. Greater understanding about beneficiaries gives context to the impact of trade-offs in ES supply that result from a particular management approach, or change in land use or management intensity (De Groot et al., 2010; Hauck et al., 2013), particularly where they may be remote from the forest as the ecosystem service providing area (García-Nieto et al., 2013; Palomo et al., 2013). Knowledge about the temporal scale at which benefits are demanded and supplied can improve understanding about the ES outcomes from different management approaches (Duncker et al., 2012b) or land use change such as afforestation (Gimona and Van Der Horst, 2007). Conclusion We have shown that ES are integrated in forest policy and our analysis has identified a consistent set of priority ES for which forest managers will be required to provide evidence of implementation and impact of forest policy. Temperate forests deliver a wide range of ES that are affected by management intensity. This paper has shown that low intensity management or no management is unsuitable for high biomass production, yet provide high or moderately high levels of other services. Higher intensity management provides the greatest biomass provision but impacts negatively on biodiversity, health and recreation and water supply services. Combined objective forestry provides high or moderately high levels for all priority ES except biomass. Maintaining the supply of ES at the forest scale will require a diversity of management approaches that build resilience in forests in the face of socio-economic and climate change uncertainty. Understanding how ES are affected by forest management can be useful in informing decision-making processes, in particular demonstrating trade-offs across ES and synergies in co-production of ES for particular management approaches. Funding This work was supported by Forestry Commission GB. M.J.M. and J.S.P. would like to acknowledge support from the European Commission Seventh Framework Programme under Grant (Agreement No. FP7-ENV-2012-308393-2) Operational Potential of Ecosystem Research Applications (OPERAs). Conflict of interest statement None declared. 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