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Management of oak forests: striking a balance between timber production, biodiversity and cultural services

Management of oak forests: striking a balance between timber production, biodiversity and... INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT, 2016 VOL. 12, NOS. 1–2, 59–73 http://dx.doi.org/10.1080/21513732.2015.1120780 Special Issue: Synergies between biodiversity and timber management Management of oak forests: striking a balance between timber production, biodiversity and cultural services a a a,b a a a Magnus Löf , Jörg Brunet , Anna Filyushkina , Matts Lindbladh , Jens Peter Skovsgaard and Adam Felton a b Southern Swedish Forest Research Centre, Swedish University of Agricultural Sciences, Alnarp, Sweden; Department of Food and Resource Economics, University of Copenhagen, Frederiksberg, Denmark ABSTRACT ARTICLE HISTORY Received 23 March 2015 Identification of the ecosystem services provided by oak-dominated forests in southern Accepted 12 November 2015 Sweden is a prerequisite for ensuring their conservation and sustainable management. These forests seem well-suited for multiple-use forestry, but knowledge is limited regard- EDITED BY ing how to manage them for multiple uses. Management for the production of high-value Nicholas Brokaw timber species like oaks and management to conserve biodiversity, or for cultural services KEYWORDS can be in conflict with each other. This study evaluates the capacity of three contrasting Ecosystem services; oak management regimes to provide societies with economic revenue from timber production, forests; Sweden; synergies; habitats for biodiversity and cultural services, and the study analyses associated trade-offs trade-offs; multiple-use and synergies. The three regimes were: intensive oak timber production (A), combined forestry; silviculture management for both timber production and biodiversity (B) and biodiversity conservation without management intervention (C). We synthesized relevant scientific literature, gov- ernmental statistics and grey literature. Our assessments identified that Regime A provided the highest levels of economic returns and the lowest level of biodiversity. Regime C provided higher levels of habitat provision but at expense of wood production and cultural services. In contrast, Regime B provided a balanced delivery of timber production, biodiversity conservation and cultural services. We identified several stand-management options which provide comparatively synergistic outcomes in ecosystem services delivery. The use of these management options in combination with more traditional stand-man- agement approaches may be a more effective means of achieving sustainable forest goods and services. Despite this potential, current knowledge is limited 1. Introduction regarding how to manage oak forests for such multi- Forest ecosystems dominated by oak (Quercus spp.) ple values. are common throughout Eurasia and the Americas, In Sweden, oak forests are limited to the south, where they are valued for providing ecosystem ser- temperate region of the country, and have been vices (Johnson et al. 2009). Oak-dominated forests widespread in the region since the Holocene provide high value timber for industry, biomass for (Figure 1). Currently however, due to a variety of bio-energy production, key habitats for biodiversity biogeographical changes and anthropogenic and valued environments for recreation and other impacts, oak comprises only 2% of the standing cultural services. The recognized capacity of oak- volume (Swedish Forest Agency 2014), and is dominated forests to provide for multiple ecosystem much less abundant than in previous centuries services aligns well with growing societal expectations (Lindbladh & Foster 2010). Instead, and partially that production forests are managed for multiple as a result of conifer-dominated production for- products and services (Gustafsson et al. 2012; estry, thelandscapeofsouthernSwedenprimarily Schwenk et al. 2012). Referred to as multiple-use or consists of a mixture of conifer plantations and multi-functional forests, these forest lands are mana- agriculture (Löf et al. 2012). Most of the remaining ged in a way that recognizes the importance of bal- oak forests are managed or have been managed for ancing non-timber values with sustainable timber timber production by non-industrial private forest production (Thompson et al. 2011). Within their owners in a variety of ways. Given appropriate native range, oak-production forests appear well-sui- management, these stands can produce highly valu- ted to meet the challenge of multi-use forest manage- able timber and other wood products providing ment, and in the northern edges of their distribution, substantial economic returns (Werner et al. 2000). can also aid in the adaptation of forest management Given that a substantial percentage of oak wood to warmer and possibly more extreme climates (Bolte used in Sweden is imported, there appears to be et al. 2009; Felton et al. 2010a; Löf et al. 2012). CONTACT Magnus Löf magnus.lof@slu.se © 2016 Informa UK Limited, trading as Taylor & Francis Group 60 M. LÖF ET AL. 3 –1 25 m ha 3 –1 15 m ha 3 –1 5 m ha 100 km 3 –1 2.5 m ha 3 –1 1 m ha 3 –1 < 1 m ha Figure 1. Distribution of standing volume of oak in southern Sweden south of ‘Limes Norrlandicus’, the northern limit of the temperate forest zone. The data are derived from Swedish University of Agricultural Sciences and the Swedish National Forest Inventory (SLU-SNFI). untapped potential for the Swedish oak resource 2009). Irrespective of these conflicts, the protected (Nylinder et al. 2006). area consisting of oak-dominated forest lands is In southern Sweden and elsewhere, managed oak expected to increase (Götmark 2013). Management stands also provide a range of benefits in addition to for the production of high-value timber species like economic returns and wood products. For example, oaks and management to conserve biodiversity can be stands of oak are among the most preferred forest in conflict with each other. This paper examines the habitats for recreation, especially near urbanized scientific basis for positive synergy between managing areas (Norman et al. 2010). Furthermore, these for- both for high-value oaks and for biodiversity. ests are also considered important environments for There is thus a range of distinctive and potentially biodiversity conservation (Götmark 2013). With the competing or synergistic societal goals influencing vast majority of Sweden’s forest outside of protected the management of oak production stands, but there areas (Swedish Forest Agency 2014), production for- is little information available regarding the relative ests are necessarily enlisted as part of biodiversity capacity of different management regimes to simulta- conservation efforts. Because of the high biodiversity neously balance the habitat requirements of forest value of the oak forests which remain (Berg et al. biodiversity and delivery of associated ecosystem ser- 1994), these forest areas often receive disproportion- vices for societal wellbeing. By reviewing the available ate pressure from the state to be set aside exclusively literature, we attempt to fill some of the relevant for conservation, which often leads to conflict knowledge gaps by contrasting three distinctive man- between forest owners and the authorities (Götmark agement regimes for oak forests in southern Sweden, INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 61 two of which are currently applied while one may although these forests may produce highly valuable timber, wood production often only plays a minor become a valuable alternative in the future. Our objective is to evaluate the capacity of these alterna- role in the economy of each forest owner and in the tives to provide societies with timber (including eco- overall forestry economy due to relatively slow growth and the limited timber resource. The asso- nomic revenue), habitat for biodiversity, and cultural services, while analyzing associated trade-offs and ciated industry (saw-mills, furniture manufacturers, synergies. We hope our conclusions stimulate the etc.) is under-developed, and attempts to improve the timber resource are hampered by the small size of development of new management guidelines for oak-dominated lands in southern Sweden and else- stands, their scattered placement in the landscape and where that are more effective at fulfilling multiple associated harvesting inefficiencies (Werner et al. 2000). In contrast, the forest industry is heavily societal goals. We also hope that our study stimulates field research comparing delivery of ecosystem ser- dependent on the intensive, production-oriented vices by alternative forest management plans, includ- management of a limited number of highly produc- tive tree species, resulting in a forest landscape in ing our proposed new management regime. Such comparative field experiments would provide a plat- southern Sweden dominated by Norway spruce, and form for more formalized approaches to ecological to a lesser degree, Scots pine and birch (Swedish and socio-economic valuation of ecosystem functions Forest Agency 2014). Norway spruce is heavily uti- and services (De Groot et al. 2002; Fontana et al. lized due to its rapid growth, ease of establishment 2013). and management, and its lower degree of palatability to browsing ungulates (Bergquist et al. 2009). Policies relating to temperate broad-leaved forests 2. The southern Swedish context have primarily concentrated on protecting remaining stands from conversion to other land uses, primarily The two oak species found in southern Sweden, Norway spruce-dominated forestry. As a result, forest Quercus robur and Q. petraea, overlap considerably owners are not allowed to convert oak forest to con- in range, forest types and ecological characteristics ifer forest. To compensate forest owners for the asso- and we consequently do not differentiate between ciated silvicultural costs of managing such stands, the them in this article. The history of oak in this region regeneration and early stand management of these is defined by a gradual decline over recent millennia stands are subsidized (Löf et al. 2012). In addition, in the prevalence of oak-dominated forests throughout the production–forest matrix, the conser- (Lindbladh & Foster 2010). Today, oaks are distrib- vation of temperate broad-leaved forests is encour- uted south of Limes Norrlandicus, the northern limit aged by the Forestry Act, which requires timber and of the temperate forest zone (Figure 1). Oak stands environmental values to be given equal consideration. are mainly found on sites with favorable climate and These requirements and the widespread adoption of soil conditions, often in the transition between farm- third-party forest certification programs have resulted land and coniferous forest, but also on nutrient-poor in the use of green tree retention practices which and dry sites in coastal areas (Diekmann 1999). In the often prioritize retention of broadleaf tree species region, around 60.000 – 70.000 ha consists of oak (Simonsson et al. 2015). forests (>50% oak by basal area), but a considerable Additional policies of relevance to oak stand and additional proportion of oak occurs as scattered management include the Right of Public Access trees in pastures and as mixtures with other tree which allows people to freely visit and experience species in forests (Almgren et al. 1984; Swedish forest environments regardless of forest ownership Forest Agency 2014). Common mixtures are oak/ (Bergfors 1990). This right is considered essential Norway spruce (Picea abies) and oak/Scots pine for the outdoor recreation experiences of Swedes. (Pinus sylvestris) with or without other broadleaf As a result a typical Swede visits forests for recrea- species, such as birch (Betula spp.), aspen (Populus tion at least once every two weeks, and over 40% tremula), beech (Fagus sylvatica) and lime tree (Tilia of the Swedish population would prefer a shorter cordata), or mixtures of oak and other broadleaf distance to the forest from their homes species (Drössler et al. 2012). Many of these pure (Lindhagen & Hörnsten 2000). Oak trees and for- oak or oak-rich mixed forests derive from historical ests are associated with a range of aesthetic, sym- land uses such as woodland pasture, coppice wood- bolic, religious, recreational and historical values land or, more recently, abandoned fields and pasture (Garrido 2014) and are considered important in (Götmark 2013). The mean annual increment in pure 3 −1 −1 providing stress relief and other health benefits oak stands ranges between 3 and 6 m ha year on (Annerstedt et al. 2010). Broad-leaved forests are relatively good sites, with top heights from 20 to 26 m generally favored by the public; however, mixed after 100 years (Carbonnier 1975). forests are the most preferred (Nielsen et al. 2007; Most temperate broad-leaved forests are owned by Norman et al. 2010). small private forest owners (Löf et al. 2012), and 62 M. LÖF ET AL. attractive to forest owners in the future. In this 3. The three management regimes study, A serves as a reference condition for evaluating All three management regimes considered are either the production potential of B and C, and C provides a already applied or may be feasibly applied to the reference condition for evaluating the biodiversity remaining oak-dominated forest lands in southern potential of A and B. The production potential and Sweden that originate from afforestation or reforesta- economic return will be evaluated for the entire rota- tion of abandoned agricultural fields or grazing pas- tion period, whereas the biodiversity and cultural tures, or from oak-rich woodland pastures. To potential will be evaluated mainly at the end of the simplify comparisons, we assume that that the three rotation period, that is, at the age of 120 years or regimes all have the same starting point. Thus, when more. The fact that C stands will undergo natural the regeneration is established, any living or dead trees processes of growth, disturbances and decay pro- are removed but there is a sparse layer of shrubs. This cesses after this period is nevertheless acknowledged, approach is adopted not to strictly mimic realistic estab- something that can have important positive conse- lishment circumstances, but to minimize confounding quences for biodiversity. variables when making our comparisons. In all cases, Option A is targeted at the production of high- the dense (2000–5000 seedlings per hectare) natural or value oak timber over the whole area of the stand artificial (direct seeding or planting) regeneration of (Figure 2) and corresponds to contemporary silvicul- oaks is initially developed under open conditions. ture practices for oak in Sweden (Carbonnier 1975). Normally, other species of trees and shrubs also colo- In the young stand, management Regime A involves nize the area through natural regeneration so that an early pre-commercial thinnings to remove undesired oak-rich mixture is developed with a composition tree species and wolf oak trees and to promote the depending on the surrounding seed sources. growth of the remaining oaks. Trees that do not Two of the management regimes considered (A interfere with oaks of good stem quality are usually and C) are already typical for southern Sweden, not removed. With time and growth of the oaks the whereas the third (B) is hypothetical (see Figure 2). share of other tree species usually decreases so that, at Alternative A is targeted at the production of high- the end of the rotation, oak trees dominate the over- value oak timber and follows contemporary silvicul- storey. At the age of 30–40 years, when the stand is tural practices for oak management, mainly with the around 10–15 m tall and the oak stem density has objective of maximizing the economic return. been reduced to around 500 trees per ha, 60–100 Alternative B is also targeted at the production of potential future crop trees per hectare will be selected high-value oak timber, but combined with concerns and marked (Almgren et al. 1984). Subsequently, for biodiversity (Jensen & Skovsgaard 2009; Wilhelm regular thinnings will be conducted to promote the & Rieger 2013). Alternative C is targeted at the con- selected potential oak crop trees to ensure their opti- servation of biodiversity, without management inter- mal crown development and diameter growth of ventions (Götmark 2013). From a Swedish policy stems. If other tree species are not interfering with perspective, Option B could become increasingly the crowns of the oaks, they may be kept as an A B C Figure 2. Schematic illustration of the three management regimes in oak forests at the end of the rotation for which the crowns of crop trees are indicated with light green circles and areas without management intervention in dark green. The crowns of harvested crop trees occupy the majority of the stand in Regime A, parts of the stand in Regime B and are absent in stand C for which no trees are harvested. Each regime is also illustrated with photos. The A regime alternative that lacks an understory is illustrated. Photos: Magnus Löf, Lars Drössler and Jörg Brunet. INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 63 most likely remain small and close rapidly. Overall, understorey. This understorey may be dominated by the stand will likely be denser than in Option A, but broad-leaved tree species or Norway spruce and may help prevent the development of epicormic shoots on will contain gaps especially where the managed oaks grow. the oaks. At the age of 30–40 years and onwards, Option C is targeted at the conservation of biodi- epicormic shoots may be pruned at regular intervals to produce premium quality timber. After a rotation versity, and the whole stand is left for natural forest development without management interventions period of 120–150 years, around 50–70 of these oaks (Götmark 2013). Competition and lack of regular dominate the overstorey and remain in the final crop, which may or may not possess a well-developed management interventions within this dense mixture will disfavor oaks and other light-demanding species. understorey. By then, the oaks will be around 25 m Therefore, the proportion of crown cover provided by tall, have a crown diameter of 10–14 m, a target diameter at breast height of 60–70 cm and a clear oaks after 120–150 years will most likely be lower bole of 6–8 m. In this option, only small amounts of than in Option B. Similar to the unmanaged parts of Option B, some small gaps may temporarily be dead wood or dead trees are present during the rota- tion due to frequent management interventions that present, and these stands will be denser and more remove trees. From the management, some slash and dead wood will be present compared to Options A thinning residues will be left on site. and B. However, in the absence of major disturbances Option B (Figure 2) is targeted at the production or targeted intervention, dead wood accumulation is of high-value oak timber and biodiversity conserva- a slow process requiring extended time periods tion. To achieve this, the production aspect is limited (Vandekerkhove et al. 2009). to a subsection of the stand, with the rest of the stand left for natural development without management interventions (Jensen & Skovsgaard 2009; Wilhelm 4. Assessment of the management regimes & Rieger 2013). The production-dedicated areas of 4.1. Timber production the stand are established and maintained essentially in the same way as in Option A, but with fewer crop A larger variety of marketable wood (small – and trees (for example, 15–35 per ha) managed for tim- large diameter timber) will be produced and har- ber. Regular thinning will occur only to promote the vested per area in management Regime A compared timber crop trees. At the end of the rotation, after to Regime B. In Regime C, no marketable wood will 120–150 years, the managed oaks will occupy be harvested. approximately 20–70% of the overstorey. If other We evaluated the economic consequences of the tree species are not interfering with the crowns of three management regimes (Table 1) in terms of the timber oaks, they may be kept as an understorey. economic indicators such as cash flow (age at first The unmanaged mixed parts of the stand will gradu- positive cash flow and pay-back period of initial ally become denser and contain more dead wood due investment), net stumpage value (accumulated undis- to competition and self-thinning. Here, oaks and counted cash flow) and internal rate of return (the other light-demanding tree species may suffer from discount rate at which the net present value of the competition during stand development, and at the investment equals zero) (Klemperer 1996). The net end of the rotation more shade-tolerant tree species stumpage value of thinning revenues relative to the can be expected to dominate the overstorey in these total revenue (thinning and final harvest) was used as parts of the stand. Canopy gaps may occur due to a supplementary indicator. The scenarios presented disturbance from wind and tree diseases, but will here for production of high-quality oak timber all rely Table 1. Rough estimates of economic indicators for scenarios under Option A: oak managed for high-quality timber with a target diameter (dbh) of 70 cm and an expected rotation length of 120 years, with or without an admixture and with or without pruning. Legend: CF = cash flow positive from the indicated age-class, Pay-back = the number of years before complete pay- pos back of establishment costs, NSV = net stumpage value (NSV refers to thinning revenues), IRR = internal rate of return thin (IRR refers to IRR of the marginal investment of a specified management action). Missing values (–) indicate that no specific marg information was available. All estimates are approximations and based on analyses by Hermansen (1956), Holten (1980, 1986), Ståål (1986), Staun (1989), Jensen (1989, 1993), Madsen (1991), Lindén (2003), Skovsgaard (2004), Lomholt (2006) and Jørgensen (2013). Pos. CF Pay-back NSV NSV / IRR IRR thin marg Admixture Pruning (age) (years) k€/ha NSV % % total None No 50 90 50–60 .40–.50 2.5–3.0 – Conifer No 25 40 55–65 – 3.5 – None Yes 50 100 55–75 .25–.35 – 2.0–3.5 Note: The examples all include selection and marking of 50–100 potential future crop trees per ha. Estimates of NPV are updated to the price level of 2015. 64 M. LÖF ET AL. be taken as an indication of a stable balance on suitable site conditions and a continuously opti- between supply and demand in the market for oak mal management throughout the rotation. Deviation from these conditions may reduce the economic timber. The main consequence of high-pruning is the outcome. increased revenue that results from the improved The most relevant alternative silvicultural scenar- ios for timber production under Option A include the quality of final crop trees. High-pruning remains profitable at an internal rate of return of 2.0–3.5% possible admixture of other tree species (hardwoods, when considering only the added value on the oak conifers or both), the use of permanently marked potential future crop trees and high-pruning. A timber (Jensen 1989, 1993; Skovsgaard 2004). This is based on conservative estimates and corresponds to stand managed without an understorey and without an added income of 20% or more from pruning. high-pruning is considered the base scenario against which the economic outcome of other alternatives Finally, the practice of leaving a few retention trees under Regime A will be compared (Table 1). The at regeneration as prescribed by Swedish legislation is considered of no or only marginal importance for the admixture of one or more hardwood species (instead of a conifer) does not significantly change any of the economic outcome of management Option A. economic indicators and this option is therefore not Retention trees in oak stands will usually be selected considered in the comparison. among those of inferior stem quality or located at The cash-flow profile of the base scenario for stand margins and are consequently of little monetary Regime A is strongly skewed because of establish- value. ment and tending costs that are not compensated There is essentially no information available for by early thinning revenues. In contrast, the regarding the actual costs and incomes of manage- admixture of a fast-growing conifer (for example ment Regime B or the factors upon which they Norway spruce) provides early revenues from timber depend. Option B is currently being explored in harvesting, which provides a potentially positive cash some commercial forest stands in the states of flow after 25 years. After approximately 50 years, net Rheinland-Pfalz and Saarland in Germany (Wilhelm income from the oak takes over as the conifer trees & Rieger 2013). In addition, in Denmark three thin- are gradually removed from the stand. The pay-back ning experiments in young oak include treatments period with an admixture of Norway spruce may be similar to Option B (Jensen & Skovsgaard 2009). less than half of the base scenario. These experiments were installed in 2002–2003 and The net stumpage value of the investment in growing no results are as yet available. Nevertheless, we can oak for high-quality timber in heavily thinned stands expect that the reduction in income will be propor- −1 under Regime A is approximately 55,000 € ha tional to the number of final timber trees negatively or affected by Option B plus some additional costs due more, depending mainly on market conditions and to affected logistics during forest operations the quality of the timber at final harvest (Table 1). The (Wilhelm & Rieger 2013). In the event of a larger ratio of revenues from thinnings to that of the total impact comprising half or more of the potential final revenue from thinnings and final harvest depends heav- timber trees, the income reduction would probably be ily on the quality of timber and on prices for premium marginally larger than the actual share of negatively timber and lower classes of grading. This is indicated by the variation in NSV affected potential final timber trees. In addition, no /NSV (Table 1) thin total timber trees will be managed or extracted from man- The magnitude of the added revenue from an agement Regime C which is why this regime will admixture of conifers obviously depends on the growth and proportion of conifers in the stand and produce zero costs and incomes. on the final quality of the oak stems. The example in this review has been estimated to provide an addi- 4.2. Biodiversity tional 10% compared to a pure stand of oak (Madsen 1991). The main economic effect of the conifers, 4.2.1. Vascular plants however, is improved (positive) cash-flow earlier in By increasing light availability and providing dis- the rotation. Mixed stands (i.e., oak and spruce) are turbed ground for plant recruitment, canopy thinning also considered less sensitive to changes in economic of oak forest (Regime A) generally increases under- conditions than monocultures (Lindén 2003). storey plant species richness. This increase may occur Based on sources used in this analysis (see refer- at least up to a decade into the rotation (Brunet et al. ences in text for Table 1), the internal rate of return 1996, 1997). Also, partial cutting, corresponding to on the investment in the base scenario has Option B, will probably have a clear effect on the remained relatively constant at 2.5–3.0% since the herbaceous layer and will likewise increase plant spe- 1950s, indicating a low investment risk. cies richness during the first growing season after Considering changes in technologies, work salaries cutting (Götmark et al. 2005). Most typical forest and society at large during this period, this can also plants are tolerant of canopy thinning, or may even INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 65 species with rapid and vigorous sprouting such as increase in abundance or frequency with its occur- hazel and lime tree (Leonardsson & Götmark 2015). rence (Brunet et al. 1996; Götmark et al. 2005). The abundance of many other plant species (ruderal, 4.2.2. Lichens and bryophytes grassland and habitat generalists) also increases with Previous studies have found that the thinning of trees thinning intensity, which results in a short-term increase of total species richness with increasing and shrubs around large oaks (Options A and B) management intensity (Brunet et al. 1996, 1997). helped to maintain the species density of epiphytic The composition of the vegetation emerging after lichens and bryophytes on oak trunks, including sev- cutting also depends on previous land use history eral species of conservation concern, relative to and its effects on the seed bank, and on patches of unmanaged forest (Nordén et al. 2012). In a related remnant grassland vegetation in the forest (Milberg study by Paltto et al. (2008), the composition of 1995; Jonason et al. 2014). lichens and bryophytes on tree stumps shifted Conversely, the gradual loss of light-demanding towards species adapted to drier dead wood as a herbaceous species under closing canopies in unma- result of thinning, implying an increase of lichen naged forest (C) is usually not compensated by an richness and a trend of decreasing bryophyte rich- equivalent gain of shade tolerant plant species, which ness. Similar trends were also recorded on tree logs. results in an overall decrease of species richness (Von However, no increases or decreases in species of Oheimb & Brunet 2007). Likewise, when comparing conservation concern could be detected (Paltto et al. different types of oak-hazel woodland, Hansson 2008). Similar to vascular plants, thinning had a (2001) found that areas managed as wooded meadow positive effect on bryophytes associated with the for- or wood pastures maintained a significantly higher est floor (Götmark 2013). We conclude from these total number of vascular plant species than unma- studies that overall epiphyte diversity may be highest naged woodlands. Herb layer plants greatly vary in at the stand level in Option B, by providing the abundance along the light gradient (Tyler 1989), and highest diversity of micro-habitats in terms of tree we conclude that total vascular plant diversity at the species and micro-climate (Table 2). The proportion end of the rotation period is maximized in oak forests of lichens compared to bryophytes will probably with both open and closed canopy parts. Therefore, decrease with decreasing management intensity vascular plant diversity is probably greatest under from A to C. Regime B which provides the highest structural het- erogeneity due to partial understorey clearing 4.2.3. Birds (Table 2). Bird diversity and composition is strongly influ- Similarly to the herbaceous vegetation, the diver- enced by the structural heterogeneity of a forest sity of regenerating woody plant species in the her- stand, and in particular the understorey density baceous layer generally increases after canopy (Hinsley et al. 2009;Hewsonetal. 2011). In a thinning (Brunet et al. 1996). Concerning oak itself, study of understorey importance for bird commu- seedling density was found to be threefold higher in nities in the oak-rich urban woodlands of western partially cut oak stands (Option B) than in unma- Sweden, 90% removal of the understorey had a naged control plots (Option C) three years after cut- significant negative affect on total breeding bird ting, and survival and growth were positively related species densities, while 50% removal did not affect to canopy openness (Götmark 2007). Repeated densities compared to the unmanaged controls understorey cutting may lead to changes in the com- (Heyman 2010). This indicates that a moderate position of the woody understorey and may favor disturbance of the understorey consistent with B (and some A alternatives) may not affect the avi- fauna negatively. Another Swedish study showed Table 2. Relative value of the different management options that closed oak-hazel woodlands derived from for- in relation to their promotion of structures important for mer wooded grassland, corresponding to Option C, biodiversity. Maximum value for positive effects at stand supportedalowernumberofbirdspeciesthan age of ca 120 years is +++. woodlands of similar origin yet retaining some Species Stand characteristics Option A Option B Option C groups degree of disturbance and open areas, and thus Stand structural + +++ ++ Plants, most closely resembling Option B (Hansson 2001). heterogeneity Epiphytes, Even if Option B does not involve a creation of Birds Tree species diversity + +++ ++ Epiphytes, grazed and fully open sections of the forests, Birds, repeated disturbances from the thinning and man- Beetles Live trees with cavities + ++ +++ Birds, agement of the crop trees are probably beneficial to and dead branches Beetles some speciesaswellasfor theoveralldiversityof Dead wood + ++ +++ Beetles, the stand. To summarize, management Options C Saproxylic fungi and B would produce more heterogeneous 66 M. LÖF ET AL. structure, with a more developed understorey, com- average 14.3 m /ha coarse dead wood, which is pared to A. To distinguish between B and C in this twice as much as the mean value for all forests in regard is more difficult. Even though C represents a Sweden (Nordén et al. 2004). In the same study, fine less disturbed habitat, B would include more struc- dead wood made up another 12 m /ha. The stands tural heterogeneity due to the juxtaposition of two evaluated in that study are similar to our models for relatively distinct habitat types, including unma- Option C (and parts of B), namely abandoned wood- naged areas. Furthermore, the crop trees will prob- land pastures. These stand types can be expected to ably be taller and coarser as compared to most of accumulate more dead wood through self-thinning, the trees in the unmanaged sections of the stand, and thus have higher beetle diversity compared to the creating an extra structural dimension (Table 2). pure production Option A. However, if logging resi- Tree species composition is also an important dues are left after thinnings in A and B, this can determinant of forest bird diversity (Poulsen 2002). readily benefit many species (Jonsell et al. 2007). Many bird species are exclusively associated with Stand openness is also important for many oak- broad-leaved, coniferous or mixed forests, thereby associated beetles (Ranius & Jansson 2000; Widerberg causing an important divergence in the composition et al. 2012; Gough et al. 2014). In a study of retained of bird communities associated with either forest type oaks in nine spruce production forests in southern (Bibby et al. 1989; Roberge & Angelstam 2006). Sweden, oaks exposed to intermediate insolation har- Several studies have shown that the bird communities bored more species than shaded oaks (Widerberg within mixed stands of conifers and broadleaf trees 2013). Similar results were obtained in an experiment are composed of a mixture of bird species represent- in 22 oak-rich stands in southern Sweden where ing both the broadleaf- and coniferous-associated about 25% of the tree basal area was cut in 1 ha fauna (Donald et al. 1998; Hausner et al. 2002). plots and then compared with uncut plots (Franc & Hence, the mixed overstories of Options B and C Götmark 2008). Saproxylic beetle species richness will probably result in higher bird diversity than the increased by about 35% in the harvested plots, rela- base Option A, which is managed towards a pure oak tive to the reference plots. The results of both of these overstorey (Table 2). studies indicate that Option B could be more bene- Old trees are another important structural feature ficial for saproxylic species than the alternatives con- for birds (Nilsson et al. 2001; Poulsen 2002), in par- sidered (Table 2), due to the combination of dead ticular for cavity nesters (Carlson et al. 1998). A wood and favorable light conditions. In addition, the recent study from southern Sweden comparing oak larger tree diversity in Options C and B will most plantations of different ages with natural oak-rich probably be beneficial for saproxylic beetles com- forests shows that mid-age oak plantations with a pared to the potential monocultures of some A alter- woody understorey and managed according to natives (Jonsell et al. 1998; Felton et al. 2010b). Option A had a comparable bird diversity to that of Species richness of fungal fruiting bodies on fine older natural forests. However, the community com- woody debris (1–10 cm in diameter) was negatively position differed primarily due to a lack of many tree affected by thinning but remained unaffected on hole-nesting species (Felton et al., unpublished coarse dead wood (>10 cm) in a study of experimen- results). Neither of the options assessed here include tal partial cutting in southern Swedish oak-rich for- trees older than the stand itself, and few of the trees ests. Overall species composition did not change will develop large amounts of ‘old-tree structures’ significantly as a result of partial cutting, but red- during the 120 years considered. Nevertheless, a few listed species tended to decrease in thinned plots trees in the unmanaged stand (C) and unmanaged (Nordén et al. 2008). The available data are too scarce parts of the stand in B can be expected to develop to allow general conclusions. However, the assess- cavities during the time frame considered, and ment made here indicates that the diversity of increasingly as time progresses, as large branches saproxylic fungi may be highest under Option C, die from shading and eventually break off. where a moister microclimate is maintained and also where the most dead wood will accumulate due to self-thinning. As long as substantial parts of the 4.2.4. Saproxylic beetles and fungi stand remain unmanaged in Option B, negative Oak is the most important tree in northern Europe effects of tree harvest may remain negligible, while for saproxylic beetles. Over 500 different species, Option A probably will cause a considerable decrease many of them red-listed, use oak during part or all of saproxylic fungal diversity (Table 2). of their life cycle (Jonsell et al. 1998; Dahlberg & Stokland 2004). There is a large body of literature showing the importance of the type and quantity of 4.3. Cultural services dead wood for saproxylic species richness (e.g., Jonsson et al. 2005; Franc et al. 2007). Semi-natural The nonmaterial benefits people obtain from ecosys- oak-rich stands in southern Sweden contained on tems are often referred to as cultural services (MEA, INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 67 trees, if they are in good condition (Bostedt & 2005). These services include spiritual and religious Mattsson 1995; Tönnes et al. 2004). Regime A is the values, inspiration, aesthetic values, recreational values and cultural heritage values, and have implica- most intensively managed out of the three alterna- tives. Whereas public preferences regarding forest tions for social relations and one’s sense of place. management can often vary among studies and with Their importance for society and thus need for inte- gration into decision-making has been increasingly the characteristics of the sample population, people tend to favor managed forests containing few visible acknowledged (Bowler et al. 2010; Daniel et al. traces of human activities or large disturbances (Ribe 2012). The most studied cultural services in forests are recreational and aesthetic values. Recreation 1989; Mattsson & Li 1994; Gundersen & Frivold 2008). Therefore, in regard to anthropogenic distur- values increases with the share of temperate broad- bances, clear-cuttings are believed to have the stron- leaved trees, such as oak found within a stand (Norman et al. 2010; Johansson et al. 2014). gest and the longest negative impact on scenic beauty. Structural composition of the stand is, however, Management Regime C may be perceived as ‘messy’ by the general public (Table 3). The pre- often considered more important than tree species composition (Edwards et al. 2012). Overall, tree size, sence of large amounts of dead wood (from self- openness and visual penetration into the stand as well thinning) coupled with a high density of trees and as visual diversity of the stand are the most common thus low visual penetration and absence of forest predictors of public preferences (Ribe 1989; management results in negative attitudes and low Gundersen & Frivold 2008). recreation and aesthetic values of such environ- The development stage of the stand has also been ments (Hultman 1983). However, variation in pre- shown to be an important contributor to its recrea- ferences exist between people with different levels tional value. According to a Danish study (Jensen & of knowledge about forests and their functioning Koch 2000), this effect is more prominent in broad- (Gundersen & Frivold 2008;Erikssonetal. 2012). leaved stands than in conifers. Lower recreational and Potentially countering these aspects, Regime C also aesthetic values of young stands can be attributed to contains a variety of features often considered their relatively high density (Ribe 1989). The same favorable, including medium to large trees and effect is often seen in relation to the presence of a structural diversity. Fallen twigs and branches asso- dense understorey and is probably linked to accessi- ciated with older trees may diminish recreational bility and safety considerations (Jørgensen et al. and aesthetic values, but a number of more recent 2009). For forest management this suggests the studies suggest that since the 1990s natural forests potential positive effects of thinning young and med- have been considered more suitable for recreation ium-aged stands, especially when leaving the most than they were in previous decades (Lindhagen & vigorous and attractive trees on the site. However, Hörnsten 2000; Tahvanainen et al. 2001). Relatedly, high-intensity thinnings in young oak stands could there seems to be an increasing acceptance and also decrease the impression of accessibility due to appreciation of dead wood (Tyrväinen et al. 2003; the presence of high volumes of thinning slash on the Nielsen et al. 2007;Heyman 2012). This develop- site (Jensen & Skovsgaard 2009). ment has been attributed to increasing levels of Management Regime A is likely to be preferred knowledge among the general public about especially in the later stages of the stand develop- biodiversity. ment, providing big trees that are well distributed, Management Regime B probably provides many open environments and visual penetration (Table 3). desirable features for the general public such as However, from the point of visual diversity, the struc- medium and large trees, lower overall tree density ture of monocultures is much simpler than that in of some sections of the stand and structural diver- natural or mixed forests. Thus, the presence of an sity (Table 3). Semi-open stands increase access to understorey may be beneficial for recreational and more distant areas and openings and provide a aesthetic values. In addition, aesthetic and recrea- higher sense of security compared to dense stands tional values also increase with presence of retention (Kaplan & Kaplan 1989; Gundersen & Frivold 2008; Heyman et al. 2011). Additionally, management targeting only specific trees and leaving other Table 3. Relative recreational and aesthetic values in relation areas untouched contributes to a patchy and to stand characteristics for the three management regimes. visually diverse environment, thus potentially Maximum value for positive effects at stand age of ca. increasing recreational and aesthetic values of the 120 years is +++. forest (Ribe 1989;Lindgren 1995; Nielsen et al. Stand characteristics Option A Option B Option C Number of medium/large trees +++ ++ + 2007;Heyman 2012). Uneven-aged stands with a Openness +++ ++ + mixture of trees of different sizes are not only Visibility into the stand +++ ++ + preferred over monocultures, but this effect is Visual/structural diversity + +++ ++ Positive anthropogenic disturbance + +++ + maintained throughout the rotation (Hultman 68 M. LÖF ET AL. As expected, our evaluation of the free develop- 1983; Lindhagen & Hörnsten 2000). Overall, this ment regime (C) indicated substantially higher levels regime seems to avoid and/or mitigate some of the negatively perceived features of Regime A and of habitat provision for a range of taxonomic groups, which came at the expense of contributions to wood Regime C, by excluding large disturbances such as production and cultural services. The long-term tra- clear-cuttings in the end of the rotation, reducing stand density and providing higher levels of visual jectory of oak biodiversity values in these stands is however somewhat uncertain, as it depends on the diversity. Thus, Regime B is expected to be slightly extent of canopy disturbance and associated light morefavored by thepublicthanRegimeA,and much more favored than Regime C. conditions beneficial to the natural regeneration of oak. The availability of light is a primary determinant of oak seedling and sapling growth following canopy 5. Discussion disturbance (e.g. Jones 1959; Collet et al. 1997; Lüpke 2008; Parker & Dey 2008; Gardiner et al. 2009). For Any management decision that alters the tree species this reason the occurrence of oak has often been composition, structure or disturbance regimes of a linked to various natural and anthropogenic forest forest has the corresponding capacity to alter the disturbances, (e.g., livestock grazing, harvesting, fire range and balance of ecosystem services derived and windthrow) that provide increased insolation for from that forest. Previous studies have highlighted, oak regeneration and stump sprouting (Vera 2000; for example, that an unbalanced prioritization of Johnson et al. 2009; Brose et al. 2013). intensive timber production will reduce forest biodi- There is thus a potential synergistic interaction versity (Lindenmayer & Franklin 2002; Gossner et al. between oak management which provides conditions 2014), as well as the provisioning of regulatory and beneficial for oak regeneration and associated habi- cultural services (Bennett et al. 2009; Raudsepp- tats for biodiversity. If sufficient openness is main- Hearne et al. 2010). The potential for trade-offs is, tained, due to natural or anthropogenic disturbance, however, tempered by the existence of synergistic to ensure the continued persistence of an oak-domi- relationships among biodiversity and some ecosystem nated canopy, then a range of structural features of services derived from production forests (Duncker relevance to biodiversity in oak reserves can be et al. 2012). Our assessment lends support to the expected to increase as these stands mature beyond existence of both trade-offs and synergies among 120 years. This is due to the natural processes of the ecosystem service deliverables of oak-dominated growth; aging and senescence facilitating an increased stands, with some management alternatives providing abundance of structural features (e.g., large diameter outcomes far more balanced in terms of the overall dead wood, tree holes) often closely associated with delivery of ecosystem services. high biodiversity values (Berg et al. 1994; Remm & With respect to trade-offs, the most intensive oak- Lõhmus 2011). The importance of maintaining such production forest management regime (A) provided features within landscapes like southern Sweden, the highest levels of economic returns, slightly where land-use practices have largely reduced their decreased cultural values and the lowest biodiversity. occurrence, requires that oak reserves continue to be Notably however, the intensive production alternative a key component of protected broadleaf areas in assessed did not consist of only one silvicultural southern Sweden. This may require that in oak option, but included a range of management pre- reserves, in which the occurrence of natural distur- scriptions with different implications for both biodi- bance regimes is unlikely to maintain an oak-domi- versity and recreational values. Oak production nated forest system, active intervention takes place to alternatives considered in A ranged from monocul- ensure the continuation of oak-favorable successional tures to mixed-species forests, and also varied in dynamics (Götmark 2013). terms of the extent of understorey vegetation per- In contrast to the categorically distinct production mitted to develop. Thus the intensive production and biodiversity regimes (A & C), the ‘combined forest alternative (A) assessed varied in terms of tree goal’ management species diversity and structural heterogeneity allowed regime (B) unites both of these for, two aspects of direct relevance to evaluations of management priorities. Option B appears to provide the same potential value for biodiversity as C, the biodiversity and cultural value of a stand. As our assessments indicate, the biodiversity and cultural although over a smaller spatial extent, combined value of these stands could therefore be increased with approximately the same cultural services as A. Because of this, Option B provides forest owners with with minimal reductions to production and economic outcomes, by adopting silvicultural approaches that a high degree of flexibility with respect to the relative prioritization of different ecosystem services. Forest increase tree species composition and understorey owners can for example determine the extent to structural heterogeneity (Bostedt & Mattsson 1995; Lindenmayer & Franklin 2002; Tönnes et al. 2004; which different areas of a stand are allocated to man- agement practices more conducive to production, Brockerhoff et al. 2008). INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 69 ‘combined goal management regimes’ on timber pro- biodiversity and cultural values. Despite these cap- duction economy. abilities, this management alternative is so far pri- marily a hypothetical construct in southern Sweden. When evaluating the relative biodiversity and cul- tural value of any production forest alternative, a Our assessments suggest, however, that this manage- strong determinant of outcomes is the reference con- ment alternative may be a suitable means of balan- cing the provision of a more comprehensive suite of dition chosen. In this study, we contrasted intensely managed oak production forests (A) with alternatives the ecosystem services within a stand than the cur- possessing a substantial (B) or encompassing extent rently adopted management alternatives in this region. This balanced provision appears to be consis- (C) of total area allocated to unmanaged develop- ment. Whereas the intensive oak production manage- tent with increasing demands that forests are mana- ment alternative fared poorly with respect to ged for multiple goals (Gustafsson et al. 2012; Gamfeldt et al. 2013). An alternative approach biodiversity in this comparison, this outcome would would require a continuation of more categorically likely be reversed if comparison was made with the most common silvicultural alternative in the region: distinctive oak production stands and reserves that provide for the ecosystem requirements of societies at Norway spruce-dominated production forests. This landscape scales. The landscape scale balancing of difference only increases if the management regime ecosystem services does, however, raise complications allows for retention forestry and a biodiversity-rich (Duncker et al. 2012), especially in regions such as understorey. This is because even the most intensive southern Sweden dominated by a large number of oak production alternative is nevertheless consistent small-scale forest owners (McDermott et al. 2010; with regional biodiversity goals for the increased use Gustafsson et al. 2015). of broadleaf tree species, as advocated by the Swedish Despite the apparent capacity of alternative B to Forest Agency and forest certification programs provide suitable habitat for a substantial proportion (Gustafsson et al. 2010; Gustafsson & Perhans 2010; of a region’s flora and fauna, we nevertheless wish to Johansson et al. 2013). Likewise, broadleaf forests are emphasize that protected areas are an essential part of consistently preferred over spruce-dominated alter- any comprehensive framework for the protection of natives in terms of their recreational and aesthetic forest biodiversity. This is because the most demand- values (Norman et al. 2010). ing species often require relatively contiguous areas Whereas oak timber production values have been of complex and heterogeneous forest lands for which relatively stable since at least the 1950s, their recrea- disturbance regimes are allowed to approximate nat- tional and aesthetic values may be less stable. For ural patterns and processes (Hunter et al. 1988; example, demographic differences between various Kuuluvainen et al. 2012; Götmark 2013). Caution is categories of users exist, even though public prefer- thus warranted when evaluating the capacity of pro- ences exhibit high degree of consensus (Stamps duction forest to provide a balanced delivery of eco- 1999). Children find dense forests more fascinating, system services. In relation to biodiversity, extensive and mushroom and berry pickers favor more natural- reliance on forest management alternatives that strike looking environments (Ribe 1989; Eriksson et al. an optimal balance may nevertheless compromise the 2012). Forestry professionals and students tend to regional viability of populations only sustained in prefer medium- to high-density stands, whereas land- protected areas. scape architects favor low-density stands (Petucco In relation to timber production and associated et al. 2013). In addition, variation in preferences economic returns, the outcome of Regime B will exist between people with different levels of knowl- depend on the number of crop trees produced. edge about forests and their functioning (Gundersen Thus, Regime B will often produce less marketable & Frivold 2008; Eriksson et al. 2012). When respon- timber per hectare compared to A. Therefore, an dents in Denmark and southern Sweden were extensive reliance on only Regime B would probably informed about the importance of biodiversity, they reduce the economic returns from oak production in provided higher values for its conservation (Bakhtiari the region. The small size of stands, and their scat- 2014). Thus, values can change relatively rapidly tered placement in the landscape already increases through education, with corresponding influences harvesting inefficiencies and reduces available timber on preferences for forest use. for the industry. Thus, Regime A and Regime B are Management for the production of high-value tim- not interchangeable, but instead are complementary ber species like oaks and management for cultural management alternatives. Furthermore, if decisions services, or to conserve biodiversity can be in conflict were taken to manage the majority of forests simi- with each other. The results from our review have larly, the resultant homogeneity would likely decrease identified a management regime with combined their recreational and aesthetic values (Axelsson- management goals for the oak-dominated forests Lindgren & Sorte 1987). We would also like to which appears to provide a balanced delivery of tim- emphasize that little is known about the effects of ber production, biodiversity conservation and 70 M. LÖF ET AL. Berg Å, Ehnström B, Gustafsson L, Hallingbäck T, Jonsell cultural services. We would, however, like to stress M, Weslien J. 1994. Threatened plant, animal, and fun- that insufficient studies have been conducted to gus species in Swedish forests - Distribution and habitat determine the precise capacity of management associations. Conserv Biol. 8:718–731. Regime B to provide for production, biodiversity Bergfors U. 1990. Allemansrätten ur rättsekonomisk syn- and cultural values. Instead, our assessments had to punkt: historia, regelverk och dagsläge. – Arbetsrapport rely on information derived from management nr 115. Umeå: Inst. för skogsekonomi SLU. Bergquist J, Löf M, Örlander G. 2009. Effects of roe deer regimes which closely approximated this manage- browsing and site preparation on performance of ment alternative. To validate and expand upon our planted broadleaved and conifer seedlings when using findings, targeted field experiments are necessary to temporary fences. Scand J For Res. 24:308–317. determine the precise capacity of this management Bibby CJ, Aston N, Bellamy PE. 1989. Effects of broad- alternative to provide for ecosystem services of socie- leaved trees on birds of upland conifer plantations in tal interest. These findings would help stakeholders North Wales. Biol Conserv. 49:17–29. Bolte A, Ammer C, Löf M, Madsen P, Nabuurs G-J, Schall evaluate the feasibility of the three options in terms of P, Spathelf P, Rock J. 2009. Adaptive forest management implementation capacity and their respective desir- in central Europe: climate change impacts, strategies and ability in output of ecosystem services. This may be integrative concept. Scand J For Res. 24:473–482. achieved using a systematic framework such as multi- Bostedt G, Mattsson L. 1995. The value of forests for criteria decision analysis, which quantifies ecosystem tourism in Sweden. Ann Tour Res. 22:671–680. Bowler DE, Buyung-Ali LM, Knight TM, Pullin AS. 2010. services while highlighting the contribution of A systematic review of evidence for the added benefits to selected services. (Fontana et al. 2013). Such a process health of exposure to natural environments. BMC Public of stakeholder involvement would help test the accu- Health. 10:456. racy and comprehensiveness of ecosystem service Brockerhoff EG, Jactel H, Parrotta JA, Quine CP, Sayer J. projections, and in addition, may identify additional 2008. Plantation forests and biodiversity: oxymoron or silvicultural pathways for optimizing the delivery of opportunity? Biodiv Conserv. 17:925–951. Brose PH, Dey DC, Phillips RJ, Waldrop TA. 2013.A goods and services from oak forests at stand and meta-analysis of the fire-oak hypothesis: does prescribed landscape scales. burning promote oak reproduction in eastern North America? For Sci. 59:322–334. Brunet J, Falkengren-Grerup U, Rühling Å, Tyler G. 1997. Regional differences in floristic change in south Swedish Disclosure statement oak forests as related to soil chemistry and land use. J No potential conflict of interest was reported by the Veg Sci. 8:329–336. authors. Brunet J, Falkengren-Grerup U, Tyler G. 1996. Herb layer vegetation of south Swedish beech and oak forests - effects of management and soil acidity during one dec- Funding ade. For Ecol Manage. 88:259–272. Carbonnier C. 1975. Produktionen av kulturbestånd av ek i This research was funded by the Broadleaves for the Future södra Sverige. Stud For Suec. 125:1–89. program. Anna Filyushkina was funded by European Carlson A, Sandström U, Olsson K. 1998. Availability and Commission under Erasmus Mundus joint Doctoral use of natural tree holes by cavity nesting birds in a Programme ‘Forest and Nature for Society’ (FONASO), Swedish deciduous forest. Ardea. 86:109–119. and Adam Felton was partially funded by Future Forests, Collet C, Colin F, Bernier F. 1997. Height growth, shoot a research program supported by the Foundation for elongation and branch development of young Quercus Strategic Environmental Research (MISTRA). petraea grown under different levels of resource avail- ability. Ann Sci For. 54:65–81. Dahlberg A, Stokland JN. 2004. Vedlevande arters krav på substrat (Rapport nr. 7). Jönköping: Skogsstyrelsen. References Daniel TC, Muhar A, Arnberger A, Aznar O, Boyd JW, Chan KM, Costanza R, Elmqvist T, Flint CG, Gobster Almgren G, Ingelög T, Ehnström B, Mörtnäs A. 1984. PH, et al. 2012. Contributions of cultural services to the Ädellövskog – ekologi och skötsel. Jönköping: ecosystem services agenda. Proc Natl Acad Sci. Skogsstyrelsen. 109:8812–8819. Annerstedt M, Norman J, Boman M, Mattsson L, Grahn P. De Groot RS, Wilson MA, Boumans RMJ. 2002. A typology 2010. Finding stress relief in a forest. Ecol Bull. 53:33–42. for the classification, description and valuation of eco- Axelsson-Lindgren C, Sorte G. 1987. Public response to system functions, goods and services. Ecol Econ. 41:393– differences between visually distinguishable forest stands in a recreation area. Landsc Urban Plan. 14:211–217. Diekmann M. 1999. Southern deciduous forests. In: Rydin Bakhtiari F. 2014. Revealing components of the economic H, Snoeijs P, Diekmann M, editors. Swedish plant geo- value for environmental goods and services from forest: graphy. Acta Phytogeogr Suec 84:33–53. an application of stated preference methods for forest Donald PF, Fuller RJ, Evans AD, Gough SJ. 1998.Effects valuation and conservation [PhD dissertation]. of forest management and grazing on breeding bird Copenhagen: University of Copenhagen; 214 p. communities in plantations of broadleaved and con- Bennett EM, Peterson GD, Gordon LJ. 2009. iferous trees in western England. Biol Conserv. Understanding relationships among multiple ecosystem 85:183–197. services. Ecol Lett. 12:1394–1404. INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 71 Drössler L, Attocchi G, Jensen AM. 2012. Occurence and Gough LA, Birkemoe T, Sverdrup-Thygeson A. 2014. management of oak in southern Swedish forests. Reactive forest management can also be proactive for Forstarchiv. 83:163–169. wood-living beetles in hollow oak trees. Biol Conserv. Duncker PS, Barreiro SM, Hengeveld GM, Lind T, Mason 180:75–83. WL, Ambrozy S, Spiecker H. 2012. Classification of Gundersen VS, Frivold LH. 2008. Public preferences for forest management approaches: a new conceptual frame- forest structures: a review of quantitative surveys from work and its applicability to European forestry. Ecol Soc. Finland. Urban For Urban Green. 7:241–258. 17:51. Gustafsson L, Baker SC, Bauhus J, Beese WJ, Brodie A, Edwards DM, Jay M, Jensen FS, Lucas B, Marzano M, Kouki J, Lindenmayer DB, Lohmus A, Martinez PG, Montagné C, Peace A, Weiss G. 2012. Public preferences Messier C, et al. 2012. Retention forestry to maintain across Europe for different forest stand types as sites for multifunctional forests: a world perspective. Bioscience. recreation. Ecol Soc. 17:27. 62:633–645. Eriksson L, Nordlund AM, Olsson O, Westin K. 2012. Gustafsson L, Felton A, Felton AM, Brunet J, Caruso A, Recreation in different forest settings: a scene preference Hjälten J, Lindbladh M, Ranius T, Roberge J-M, Weslien study. Forests. 3:923–943. J. 2015. Natural versus national boundaries: the impor- Felton A, Ellingson L, Andersson E, Drössler L, Blennow K. tance of considering biogeographical patterns in forest 2010a. Adapting production forests in southern Sweden conservation policy. Conserv Lett. 8:50–57. to climate change – constraints and opportunities for Gustafsson L, Kouki J, Sverdrup-Thygeson A. 2010. Tree risk spreading. Int J Clim Change Strat Manage. 2:84–97. retention as a conservation measure in clear-cut forests Felton A, Lindbladh M, Brunet J, Fritz Ö. 2010b. Replacing of northern Europe: a review of ecological consequences. coniferous monocultures with mixed-species production Scand J For Res. 25:295–308. stands: an assessment of the potential benefits for forest Gustafsson L, Perhans K. 2010. Biodiversity conservation in biodiversity in northern Europe. For Ecol Manage. Swedish forests: ways forward for a 30-year-old multi- 260:939–947. scaled approach. Ambio. 39:546–554. Fontana V, Radtke A, Fedrigotti VB, Tappeiner U, Tasser Hansson L. 2001. Traditional management of forests: plant E, Zerbe S, Buchholz T. 2013. Comparing land-use alter- and bird community responses to alternative restoration natives: using the ecosystem services concept to define a of oak-hazel woodland in Sweden. Biodiv Conserv. multi-criteria decision analysis. Ecol Econ. 93:128–136. 10:1865–1873. Franc N, Götmark F. 2008. Openness in management: Hausner VH, Yoccoz NG, Strann KB, Ims RA. 2002. hands-off vs partial cutting in conservation forests, and Changes in bird communities by planting non-native the response of beetles. Biol Conserv. 141:2310–2321. spruce in coastal birch forests of northern Norway. Franc N, Götmark F, Okland B, Nordén B, Paltto H. 2007. Ecoscience. 9:470–481. Factors and scales potentially important for saproxylic Hermansen NK. 1956. Fra Bregentveds egeskove. beetles in temperate mixed oak forest. Biol Conserv. Økonomiske analyser. Dansk Skovforenings Tidsskrift. 135:86–98. 41:507–561. Gamfeldt L, Snäll T, Bagchi R, Jonsson M, Gustafsson L, Hewson CM, Austin GE, Gough SJ, Fuller RJ. 2011. Kjellander P, Ruiz-Jaen MC, Fröberg M, Stendahl J, Species-specific responses of woodland birds to stand- Philipson CD, et al. 2013. Higher levels of multiple level habitat characteristics: the dual importance of for- ecosystem services are found in forests with more tree est structure and floristics. For Ecol Manage. 261:1224– species. Nature Comm. 4:1340. 1240. Gardiner ES, Löf M, O´Brien JJ, Stanturf JA, Madsen P. Heyman E. 2010. Clearance of understorey in urban wood- 2009. Photosynthetic characteristics of Fagus sylvatica lands: assessing impact on bird abundance and diversity. and Quercus robur established for stand conversion For Ecol Manage. 260:125–131. from Picea abies. For Ecol Manage. 258:868–878. Heyman E. 2012. Analysing recreational values and man- Garrido P. 2014. Cultural Oak landscapes as green infra- agement effects in an urban forest with the visitor- structure for human well-being [Licentiate Thesis]. employed photography method. Urban For Urban Skinnskatteberg: Swedish University of Agricultural Green. 11:267–277. Sciences. 74 p. Heyman E, Gunnarsson B, Stenseke M, Henningsson S, Gossner M, Schall P, Ammer C, Ammer U, Engel K, Tim G. 2011. Openness as a key-variable for analysis of Schubert H, Simo U, Utschick H, Weisser WW. 2014. management trade-offs in urban woodlands. Urban For Forest management intensity measures as alternative to Urban Green. 10:281–293. stand properties for quantifying effects on biodiversity. Hinsley SA, Hill RA, Fuller RJ, Bellamy PE, Rothery P. Ecosphere. 5:113. 2009. Bird species distributions across woodland canopy Götmark F. 2007. Careful partial harvesting in conservation structure gradients. Comm Ecol. 10:99–110. stands and retention of large oaks favour oak regenera- Holten NE. 1980. En af Hauchs egekulturer samt egedyrkning tion. Biol Conserv. 140:349–358. og inflation. Dansk Skovforenings Tidsskrift. 65:296–340. Götmark F. 2009. Conflicts in conservation: woodland key Holten NE. 1986. Eichenwirtschaft - dargestellt an habitats, authorities and private forest owners in Beispielen aus dem dänischen Forstamt Bregentved. Sweden. Scand J For Res. 24:504–514. Forstarchiv. 57:221–227. Götmark F. 2013. Habitat management alternatives for Hultman SG. 1983. Public judgement of forest environ- conservation forests in the temperate zone: review, ments as recreation areas. 2. A national survey. Report synthesis, and implications. For Ecol Manage. 306:292– No. 28. Uppsala: Section of Environmental Forestry, 307. Swedish University of Agricultural Sciences. (in Götmark F, Paltto H, Nordén B, Götmark E. 2005. Swedish, with English summary). Evaluating partial cutting in broadleaved temperate for- Hunter ML, Jacobson GL, Webb T. 1988. Paleoecology and est under strong experimental control: short-term effects the coarse-filter approach to maintaining biological on herbaceous plants. For Ecol Manage. 214:124–141. diversity. Conserv Biol. 2:375–385. 72 M. LÖF ET AL. Jensen FS, Koch NE. 2000. Measuring forest preferences of Lindhagen A, Hörnsten L. 2000. Forest recreation in 1977 the population - a Danish approach. Schweiz Z Forstwes. and 1997 in Sweden: changes in public preferences and 151:11–16. behaviour. Forestry. 73:143–153. Jensen FS, Skovsgaard JP. 2009. Precommercial thinning of Löf M, Brunet J, Hickler T, Birkedal M, Jensen MA. 2012. pedunculate oak: recreational preferences of the popula- Restoring broadleaved forests in southern Sweden as cli- tion of Denmark for different thinning practices in mate changes. In: Stanturf JA, Madsen P, Lamb D, editors. young stands. Scand J For Res. 24:28–36. A goal-oriented approach to forest landscape restoration. Jensen JS. 1989. Økonomien i vanrisopkapning. Dansk Berlin: World Forests XVI, Springer; p. 373–391. Skovbrugs Tidsskrift. 74:113–124. Lomholt A. 2006. Egens økonomiske tomgangsproblem. Jensen JS. 1993. Vanris på dansk stilkeg. Forskningsserien. Skoven. 38:122–125. 4:1–105. Lüpke BV. 2008. Einfluss unterschiedlicher Hiebsformen Johansson M, Gyllin M, Witzell J, Küller M. 2014. Does auf die Naturverjüngung eines Traubeneichen-Buchen- biological quality matter? Direct and reflected appraisal Mischbestandes (In german). Influence of various cut- of biodiversity in temperate deciduous broad-leaf forest. ting types on natural regeneration of a sessile oak-beech Urban For Urban Green. 13:28–37. mixed stand. Forsarchiv. 79:4–14. Johansson T, Hjälten J, De Jong J, Von Stedingk H. 2013. Madsen EM. 1991. Kan granen lösa tomgångsproblemet? Environmental considerations from legislation and certi- Skog & Forskning. 2:36–41. fication in managed forest stands: a review of their impor- Mattsson L, Li C. 1994. How do different forest manage- tance for biodiversity. For Ecol Manage. 303:98–112. ment practices affect the non-timber value of forests?— Johnson PS, Shifley SR, Rogers R. 2009. The ecology and an economic analysis. J Environ Manage. 41:79–88. silviculture of oaks. 2nd ed. Wallingford: CABI McDermott CL, Cashore B, Kanowski P. 2010. Global Publishing. environmental forest policies: an international compar- Jonason D, Ibbe M, Milberg P, Tunér A, Westerberg L, ison. Oxon: Earthscan. Bergman KO. 2014. Vegetation in clear-cuts depends on Milberg P. 1995. Soil seed bank after eighteen years of succes- previous land use: a century-old grassland legacy. Ecol sion from grassland to forest. Oikos. 72:3–13. Evol. 4:4287–4295. [MEA] Millennium Ecosystem Assessment. 2005. Ecosystems Jones EW. 1959. Biological flora of the British Isles: and human well-being: biodiversity synthesis [Internet]. Quercus L. J Ecol. 47:169–222. World Resources Institute; [cited 2015 Feb 24]. Available Jonsell M, Hansson J, Wedmo L. 2007. Diversity of from: http://www.unep.org/maweb/documents/document. saproxylic beetle species in logging residues in Sweden 354.aspx.pdf - Comparisons between tree species and diameters. Biol Nielsen AB, Olsen SB, Lundhede T. 2007. An economic Conserv. 138:89–99. valuation of the recreational benefits associated with Jonsell M, Weslien J, Ehnström B. 1998. Substrate require- nature-based forest management practices. Landsc ments of red-listed saproxylic invertebrates in Sweden. Urban Plan. 80:63–71. Biodivers Conserv. 7:749–764. Nilsson SG, Hedin J, Niklasson M. 2001. Biodiversity and Jonsson BG, Kruys N, Ranius T. 2005. Ecology of species its assessment in boreal and nemoral forests. Scand J living on dead wood - lessons for dead wood manage- Forest Res Suppl. 3:10–26. ment. Silva Fenn. 39:289–309. Nordén B, Götmark F, Ryberg M, Paltto H, Allmér J. 2008. Jørgensen BB, Jensen FS, Skovsgaard JP. 2009. Partial cutting reduces species richness of fungi on Underplantning af eg: øger underplantning skovens woody debris. Can J For Res. 38:1807–1816. rekreative værdi? Skoven. 41:203–207. Nordén B, Götmark F, Tönnberg M, Ryberg M. 2004. Dead Jørgensen JH. 2013. Bregentveds egedyrkning ved en wood in semi-natural temperate broadleaved woodland: milepæl. Skoven. 45:121–125. contribution of coarse and fine dead wood, Kaplan R, Kaplan S. 1989. The experience of nature. attached dead wood and stumps. For Ecol Manage. Cambridge: Cambridge University Press. 194:235–248. Klemperer WD. 1996. Forest resource economics and Nordén B, Paltto H, Claesson C, Götmark F. 2012. Partial finance. New York (NY): McGraw-Hill. cutting can enhance epiphyte conservation in temperate Kuuluvainen T, Tahvonen O, Aakala T. 2012. Even-aged oak-rich forests. For Ecol Manage. 270:35–44. and uneven-aged forest management in Boreal Norman J, Ellingson L, Boman M, Mattsson L. 2010. The Fennoscandia: a review. Ambio. 41:720–737. value of forests for outdoor recreation in southern Leonardsson J, Götmark F. 2015. Differential survival and Sweden : are broadleaved trees important ? Ecol Bull. growth of stumps in 14 woody species after conservation 53:21–31. thinning in mixed oak-rich temperate forests. Eur J Nylinder M, Woxblom L, Fryk H. 2006. Ädellöv – virke Forest Res. 134:199–209. och förädling. Uppsala: Sveriges Lantbruksuniversitet, Lindbladh M, Foster DR. 2010. Dynamics of long-lived Institutionen för skogens produkter och marknader. foundation species: the history of Quercus in southern Paltto H, Nordén B, Götmark F. 2008. Partial cutting as a Scandinavia. J Ecol. 98:1330–1345. conservation alternative for oak (Quercus spp.) forest— Lindén M. 2003. Increment and yield in mixed stands with Response of bryophytes and lichens on dead wood. For Norway spruce in southern Sweden [Doctoral thesis]. Ecol Manage. 256:536–547. Swedish University of Agricultural Sciences. Silvestria 260. Parker WC, Dey DC. 2008. Influence of overstory density Lindenmayer DB, Franklin JF. 2002. Conserving forest on ecophysiology of red oak (Quercus rubra) and sugar biodiversity: a comprehensive multiscaled approach. maple (Acer saccharum) seedlings in central Ontario Washington: Island Press. shelterwoods. Tree Physiol. 28:797–804. Lindgren. 1995. Forest aesthetics. In: Hytönen M, editor. Petucco C, Skovsgaard JP, Jensen FS. 2013.Recreational Multiple-use forestry in the Nordic Countries. Metla: preferences depending on thinning practice in young Finnish Forest Research Institute, Helsinki Research even-aged stands of pedunculate oak (Quercus robur Centre; p. 279–293. L.): comparing the opinions of forest and landscape INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 73 experts and the general population of Denmark. Scan J perceptions–visual versus verbal information. Landsc For Res. 28:668–676. Urban Plan. 53:53–70. Poulsen BO. 2002. Avian richness and abundance in tem- Thompson ID, Okabe K, Tylianakis JM, Kumar P, perate Danish forests: tree variables important to birds Brockerhoff EG, Schellhorn NA, Parrotta JA, Nasi R. and their conservation. Biodiv Conserv. 11:1551–1566. 2011. Forest biodiversity and the delivery of ecosystem Ranius T, Jansson N. 2000. The influence of forest regrowth, goods and services: translating science into policy. original canopy cover and tree size on saproxylic beetles Bioscience. 61:972–981. associated with old oaks. Biol Conserv. 95:85–94. Tönnes S, Karjalainen E, Löfström I, Neuvonen M. 2004. Raudsepp-Hearne C, Peterson GD, Bennett E. 2010. Scenic impacts of retention trees in clear-cutting areas. Ecosystem service bundles for analyzing tradeoffs in Scand J For Res. 19:348–357. diverse landscapes. Proc Natl Acad Sci. 107:5242–5247. Tyler G. 1989. Interacting effects of soil acidity and Remm J, Lõhmus A. 2011. Tree cavities in forests–the broad canopy cover on the species composition of field-layer distribution pattern of a keystone structure for biodiver- vegetation in oak/hornbeam forests. For Ecol Manage. sity. For Ecol Manage. 262:579–585. 28:101–114. Ribe RG. 1989. The aesthetics of forestry: what has empirical Tyrväinen L, Silvennoinen H, Kolehmainen O. 2003. preference research taught us? Environ Manage. 13:55–74. Ecological and aesthetic values in urban forest manage- Roberge JM, Angelstam P. 2006. Indicator species among ment. Urban For Urban Green. 1:135–149. resident forest birds – a cross-regional evaluation in Vandekerkhove K, Keersmaeker LD, Menke N, Meyer P, northern Europe. Biol Conserv. 130:134–147. Verschelde P. 2009. When nature takes over from man: Schwenk WS, Donovan TM, Keeton WS, Nunery JS. 2012. dead wood accumulation in previously managed oak and Carbon storage, timber production, and biodiversity: beech woodlands in North-western and Central Europe. comparing ecosystem services with multi-criteria deci- For Ecol Manage. 258:425–435. sion analysis. Ecol Appl. 22:1612–1627. Vera FWM. 2000. Grazing ecology and forest history. UK: Simonsson P, Gustafsson L, Östlund L. 2015. Retention Cabi Publishing. forestry in Sweden: driving forces, debate and imple- Von Oheimb G, Brunet J. 2007. Dalby Söderskog revisited: mentation 1968-2003. Scand J For Res. 30:154–173. long-term vegetation changes in a south Swedish decid- Skovsgaard JP. 2004. Produktion af kvalitetstræ i eg: er uous forest. Acta Oecol. 31:229–242. eventyret bæredygtigt? Skoven. 36:532–535. Werner M, Rytter L, Stener L-G. 2000. Förbättrat Ståål E. 1986. Eken i skogen och landskapet. Växjö: Södra lövvedutnyttjande för vidareförädling. Redogörelse nr skogsägarna. 3. Uppsala: SkogForsk. Stamps AE. 1999. Demographic effects in environmental Widerberg KM. 2013. Oak as retention tree in commercial aesthetics: a meta-analysis. J Plan Lit. 14:155–175. spruce forests [Doctoral thesis No. 2013:66]. Sweden: Staun H. 1989. Intensiv egedyrkning. Dansk Skovbrugs Faculty of Forest Sciences, SLU. Tidsskrift. 74:87–112. Widerberg KM, Ranius T, Drobyshev I, Nilsson U, Swedish Forest Agency. 2014. Statistical yearbook of for- Lindbladh M. 2012. Increased openness around retained estry 2014. Jönköping: Official statistics of Sweden, oaks increases species richness of saproxylic beetles. Swedish Forest Agency. Biodiv Conserv. 21:3035–3059. Tahvanainen L, Tyrväinen L, Ihalainen M, Vuorela N, Wilhelm GJ, Rieger H. 2013. Naturnahe Waldwirtschaft Kolehmainen O. 2001. Forest management and public - die QD-Strategie. Stuttgart: Eugen Ulmer Verlag. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png International Journal of Biodiversity Science, Ecosystem Services & Management Taylor & Francis

Management of oak forests: striking a balance between timber production, biodiversity and cultural services

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INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT, 2016 VOL. 12, NOS. 1–2, 59–73 http://dx.doi.org/10.1080/21513732.2015.1120780 Special Issue: Synergies between biodiversity and timber management Management of oak forests: striking a balance between timber production, biodiversity and cultural services a a a,b a a a Magnus Löf , Jörg Brunet , Anna Filyushkina , Matts Lindbladh , Jens Peter Skovsgaard and Adam Felton a b Southern Swedish Forest Research Centre, Swedish University of Agricultural Sciences, Alnarp, Sweden; Department of Food and Resource Economics, University of Copenhagen, Frederiksberg, Denmark ABSTRACT ARTICLE HISTORY Received 23 March 2015 Identification of the ecosystem services provided by oak-dominated forests in southern Accepted 12 November 2015 Sweden is a prerequisite for ensuring their conservation and sustainable management. These forests seem well-suited for multiple-use forestry, but knowledge is limited regard- EDITED BY ing how to manage them for multiple uses. Management for the production of high-value Nicholas Brokaw timber species like oaks and management to conserve biodiversity, or for cultural services KEYWORDS can be in conflict with each other. This study evaluates the capacity of three contrasting Ecosystem services; oak management regimes to provide societies with economic revenue from timber production, forests; Sweden; synergies; habitats for biodiversity and cultural services, and the study analyses associated trade-offs trade-offs; multiple-use and synergies. The three regimes were: intensive oak timber production (A), combined forestry; silviculture management for both timber production and biodiversity (B) and biodiversity conservation without management intervention (C). We synthesized relevant scientific literature, gov- ernmental statistics and grey literature. Our assessments identified that Regime A provided the highest levels of economic returns and the lowest level of biodiversity. Regime C provided higher levels of habitat provision but at expense of wood production and cultural services. In contrast, Regime B provided a balanced delivery of timber production, biodiversity conservation and cultural services. We identified several stand-management options which provide comparatively synergistic outcomes in ecosystem services delivery. The use of these management options in combination with more traditional stand-man- agement approaches may be a more effective means of achieving sustainable forest goods and services. Despite this potential, current knowledge is limited 1. Introduction regarding how to manage oak forests for such multi- Forest ecosystems dominated by oak (Quercus spp.) ple values. are common throughout Eurasia and the Americas, In Sweden, oak forests are limited to the south, where they are valued for providing ecosystem ser- temperate region of the country, and have been vices (Johnson et al. 2009). Oak-dominated forests widespread in the region since the Holocene provide high value timber for industry, biomass for (Figure 1). Currently however, due to a variety of bio-energy production, key habitats for biodiversity biogeographical changes and anthropogenic and valued environments for recreation and other impacts, oak comprises only 2% of the standing cultural services. The recognized capacity of oak- volume (Swedish Forest Agency 2014), and is dominated forests to provide for multiple ecosystem much less abundant than in previous centuries services aligns well with growing societal expectations (Lindbladh & Foster 2010). Instead, and partially that production forests are managed for multiple as a result of conifer-dominated production for- products and services (Gustafsson et al. 2012; estry, thelandscapeofsouthernSwedenprimarily Schwenk et al. 2012). Referred to as multiple-use or consists of a mixture of conifer plantations and multi-functional forests, these forest lands are mana- agriculture (Löf et al. 2012). Most of the remaining ged in a way that recognizes the importance of bal- oak forests are managed or have been managed for ancing non-timber values with sustainable timber timber production by non-industrial private forest production (Thompson et al. 2011). Within their owners in a variety of ways. Given appropriate native range, oak-production forests appear well-sui- management, these stands can produce highly valu- ted to meet the challenge of multi-use forest manage- able timber and other wood products providing ment, and in the northern edges of their distribution, substantial economic returns (Werner et al. 2000). can also aid in the adaptation of forest management Given that a substantial percentage of oak wood to warmer and possibly more extreme climates (Bolte used in Sweden is imported, there appears to be et al. 2009; Felton et al. 2010a; Löf et al. 2012). CONTACT Magnus Löf magnus.lof@slu.se © 2016 Informa UK Limited, trading as Taylor & Francis Group 60 M. LÖF ET AL. 3 –1 25 m ha 3 –1 15 m ha 3 –1 5 m ha 100 km 3 –1 2.5 m ha 3 –1 1 m ha 3 –1 < 1 m ha Figure 1. Distribution of standing volume of oak in southern Sweden south of ‘Limes Norrlandicus’, the northern limit of the temperate forest zone. The data are derived from Swedish University of Agricultural Sciences and the Swedish National Forest Inventory (SLU-SNFI). untapped potential for the Swedish oak resource 2009). Irrespective of these conflicts, the protected (Nylinder et al. 2006). area consisting of oak-dominated forest lands is In southern Sweden and elsewhere, managed oak expected to increase (Götmark 2013). Management stands also provide a range of benefits in addition to for the production of high-value timber species like economic returns and wood products. For example, oaks and management to conserve biodiversity can be stands of oak are among the most preferred forest in conflict with each other. This paper examines the habitats for recreation, especially near urbanized scientific basis for positive synergy between managing areas (Norman et al. 2010). Furthermore, these for- both for high-value oaks and for biodiversity. ests are also considered important environments for There is thus a range of distinctive and potentially biodiversity conservation (Götmark 2013). With the competing or synergistic societal goals influencing vast majority of Sweden’s forest outside of protected the management of oak production stands, but there areas (Swedish Forest Agency 2014), production for- is little information available regarding the relative ests are necessarily enlisted as part of biodiversity capacity of different management regimes to simulta- conservation efforts. Because of the high biodiversity neously balance the habitat requirements of forest value of the oak forests which remain (Berg et al. biodiversity and delivery of associated ecosystem ser- 1994), these forest areas often receive disproportion- vices for societal wellbeing. By reviewing the available ate pressure from the state to be set aside exclusively literature, we attempt to fill some of the relevant for conservation, which often leads to conflict knowledge gaps by contrasting three distinctive man- between forest owners and the authorities (Götmark agement regimes for oak forests in southern Sweden, INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 61 two of which are currently applied while one may although these forests may produce highly valuable timber, wood production often only plays a minor become a valuable alternative in the future. Our objective is to evaluate the capacity of these alterna- role in the economy of each forest owner and in the tives to provide societies with timber (including eco- overall forestry economy due to relatively slow growth and the limited timber resource. The asso- nomic revenue), habitat for biodiversity, and cultural services, while analyzing associated trade-offs and ciated industry (saw-mills, furniture manufacturers, synergies. We hope our conclusions stimulate the etc.) is under-developed, and attempts to improve the timber resource are hampered by the small size of development of new management guidelines for oak-dominated lands in southern Sweden and else- stands, their scattered placement in the landscape and where that are more effective at fulfilling multiple associated harvesting inefficiencies (Werner et al. 2000). In contrast, the forest industry is heavily societal goals. We also hope that our study stimulates field research comparing delivery of ecosystem ser- dependent on the intensive, production-oriented vices by alternative forest management plans, includ- management of a limited number of highly produc- tive tree species, resulting in a forest landscape in ing our proposed new management regime. Such comparative field experiments would provide a plat- southern Sweden dominated by Norway spruce, and form for more formalized approaches to ecological to a lesser degree, Scots pine and birch (Swedish and socio-economic valuation of ecosystem functions Forest Agency 2014). Norway spruce is heavily uti- and services (De Groot et al. 2002; Fontana et al. lized due to its rapid growth, ease of establishment 2013). and management, and its lower degree of palatability to browsing ungulates (Bergquist et al. 2009). Policies relating to temperate broad-leaved forests 2. The southern Swedish context have primarily concentrated on protecting remaining stands from conversion to other land uses, primarily The two oak species found in southern Sweden, Norway spruce-dominated forestry. As a result, forest Quercus robur and Q. petraea, overlap considerably owners are not allowed to convert oak forest to con- in range, forest types and ecological characteristics ifer forest. To compensate forest owners for the asso- and we consequently do not differentiate between ciated silvicultural costs of managing such stands, the them in this article. The history of oak in this region regeneration and early stand management of these is defined by a gradual decline over recent millennia stands are subsidized (Löf et al. 2012). In addition, in the prevalence of oak-dominated forests throughout the production–forest matrix, the conser- (Lindbladh & Foster 2010). Today, oaks are distrib- vation of temperate broad-leaved forests is encour- uted south of Limes Norrlandicus, the northern limit aged by the Forestry Act, which requires timber and of the temperate forest zone (Figure 1). Oak stands environmental values to be given equal consideration. are mainly found on sites with favorable climate and These requirements and the widespread adoption of soil conditions, often in the transition between farm- third-party forest certification programs have resulted land and coniferous forest, but also on nutrient-poor in the use of green tree retention practices which and dry sites in coastal areas (Diekmann 1999). In the often prioritize retention of broadleaf tree species region, around 60.000 – 70.000 ha consists of oak (Simonsson et al. 2015). forests (>50% oak by basal area), but a considerable Additional policies of relevance to oak stand and additional proportion of oak occurs as scattered management include the Right of Public Access trees in pastures and as mixtures with other tree which allows people to freely visit and experience species in forests (Almgren et al. 1984; Swedish forest environments regardless of forest ownership Forest Agency 2014). Common mixtures are oak/ (Bergfors 1990). This right is considered essential Norway spruce (Picea abies) and oak/Scots pine for the outdoor recreation experiences of Swedes. (Pinus sylvestris) with or without other broadleaf As a result a typical Swede visits forests for recrea- species, such as birch (Betula spp.), aspen (Populus tion at least once every two weeks, and over 40% tremula), beech (Fagus sylvatica) and lime tree (Tilia of the Swedish population would prefer a shorter cordata), or mixtures of oak and other broadleaf distance to the forest from their homes species (Drössler et al. 2012). Many of these pure (Lindhagen & Hörnsten 2000). Oak trees and for- oak or oak-rich mixed forests derive from historical ests are associated with a range of aesthetic, sym- land uses such as woodland pasture, coppice wood- bolic, religious, recreational and historical values land or, more recently, abandoned fields and pasture (Garrido 2014) and are considered important in (Götmark 2013). The mean annual increment in pure 3 −1 −1 providing stress relief and other health benefits oak stands ranges between 3 and 6 m ha year on (Annerstedt et al. 2010). Broad-leaved forests are relatively good sites, with top heights from 20 to 26 m generally favored by the public; however, mixed after 100 years (Carbonnier 1975). forests are the most preferred (Nielsen et al. 2007; Most temperate broad-leaved forests are owned by Norman et al. 2010). small private forest owners (Löf et al. 2012), and 62 M. LÖF ET AL. attractive to forest owners in the future. In this 3. The three management regimes study, A serves as a reference condition for evaluating All three management regimes considered are either the production potential of B and C, and C provides a already applied or may be feasibly applied to the reference condition for evaluating the biodiversity remaining oak-dominated forest lands in southern potential of A and B. The production potential and Sweden that originate from afforestation or reforesta- economic return will be evaluated for the entire rota- tion of abandoned agricultural fields or grazing pas- tion period, whereas the biodiversity and cultural tures, or from oak-rich woodland pastures. To potential will be evaluated mainly at the end of the simplify comparisons, we assume that that the three rotation period, that is, at the age of 120 years or regimes all have the same starting point. Thus, when more. The fact that C stands will undergo natural the regeneration is established, any living or dead trees processes of growth, disturbances and decay pro- are removed but there is a sparse layer of shrubs. This cesses after this period is nevertheless acknowledged, approach is adopted not to strictly mimic realistic estab- something that can have important positive conse- lishment circumstances, but to minimize confounding quences for biodiversity. variables when making our comparisons. In all cases, Option A is targeted at the production of high- the dense (2000–5000 seedlings per hectare) natural or value oak timber over the whole area of the stand artificial (direct seeding or planting) regeneration of (Figure 2) and corresponds to contemporary silvicul- oaks is initially developed under open conditions. ture practices for oak in Sweden (Carbonnier 1975). Normally, other species of trees and shrubs also colo- In the young stand, management Regime A involves nize the area through natural regeneration so that an early pre-commercial thinnings to remove undesired oak-rich mixture is developed with a composition tree species and wolf oak trees and to promote the depending on the surrounding seed sources. growth of the remaining oaks. Trees that do not Two of the management regimes considered (A interfere with oaks of good stem quality are usually and C) are already typical for southern Sweden, not removed. With time and growth of the oaks the whereas the third (B) is hypothetical (see Figure 2). share of other tree species usually decreases so that, at Alternative A is targeted at the production of high- the end of the rotation, oak trees dominate the over- value oak timber and follows contemporary silvicul- storey. At the age of 30–40 years, when the stand is tural practices for oak management, mainly with the around 10–15 m tall and the oak stem density has objective of maximizing the economic return. been reduced to around 500 trees per ha, 60–100 Alternative B is also targeted at the production of potential future crop trees per hectare will be selected high-value oak timber, but combined with concerns and marked (Almgren et al. 1984). Subsequently, for biodiversity (Jensen & Skovsgaard 2009; Wilhelm regular thinnings will be conducted to promote the & Rieger 2013). Alternative C is targeted at the con- selected potential oak crop trees to ensure their opti- servation of biodiversity, without management inter- mal crown development and diameter growth of ventions (Götmark 2013). From a Swedish policy stems. If other tree species are not interfering with perspective, Option B could become increasingly the crowns of the oaks, they may be kept as an A B C Figure 2. Schematic illustration of the three management regimes in oak forests at the end of the rotation for which the crowns of crop trees are indicated with light green circles and areas without management intervention in dark green. The crowns of harvested crop trees occupy the majority of the stand in Regime A, parts of the stand in Regime B and are absent in stand C for which no trees are harvested. Each regime is also illustrated with photos. The A regime alternative that lacks an understory is illustrated. Photos: Magnus Löf, Lars Drössler and Jörg Brunet. INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 63 most likely remain small and close rapidly. Overall, understorey. This understorey may be dominated by the stand will likely be denser than in Option A, but broad-leaved tree species or Norway spruce and may help prevent the development of epicormic shoots on will contain gaps especially where the managed oaks grow. the oaks. At the age of 30–40 years and onwards, Option C is targeted at the conservation of biodi- epicormic shoots may be pruned at regular intervals to produce premium quality timber. After a rotation versity, and the whole stand is left for natural forest development without management interventions period of 120–150 years, around 50–70 of these oaks (Götmark 2013). Competition and lack of regular dominate the overstorey and remain in the final crop, which may or may not possess a well-developed management interventions within this dense mixture will disfavor oaks and other light-demanding species. understorey. By then, the oaks will be around 25 m Therefore, the proportion of crown cover provided by tall, have a crown diameter of 10–14 m, a target diameter at breast height of 60–70 cm and a clear oaks after 120–150 years will most likely be lower bole of 6–8 m. In this option, only small amounts of than in Option B. Similar to the unmanaged parts of Option B, some small gaps may temporarily be dead wood or dead trees are present during the rota- tion due to frequent management interventions that present, and these stands will be denser and more remove trees. From the management, some slash and dead wood will be present compared to Options A thinning residues will be left on site. and B. However, in the absence of major disturbances Option B (Figure 2) is targeted at the production or targeted intervention, dead wood accumulation is of high-value oak timber and biodiversity conserva- a slow process requiring extended time periods tion. To achieve this, the production aspect is limited (Vandekerkhove et al. 2009). to a subsection of the stand, with the rest of the stand left for natural development without management interventions (Jensen & Skovsgaard 2009; Wilhelm 4. Assessment of the management regimes & Rieger 2013). The production-dedicated areas of 4.1. Timber production the stand are established and maintained essentially in the same way as in Option A, but with fewer crop A larger variety of marketable wood (small – and trees (for example, 15–35 per ha) managed for tim- large diameter timber) will be produced and har- ber. Regular thinning will occur only to promote the vested per area in management Regime A compared timber crop trees. At the end of the rotation, after to Regime B. In Regime C, no marketable wood will 120–150 years, the managed oaks will occupy be harvested. approximately 20–70% of the overstorey. If other We evaluated the economic consequences of the tree species are not interfering with the crowns of three management regimes (Table 1) in terms of the timber oaks, they may be kept as an understorey. economic indicators such as cash flow (age at first The unmanaged mixed parts of the stand will gradu- positive cash flow and pay-back period of initial ally become denser and contain more dead wood due investment), net stumpage value (accumulated undis- to competition and self-thinning. Here, oaks and counted cash flow) and internal rate of return (the other light-demanding tree species may suffer from discount rate at which the net present value of the competition during stand development, and at the investment equals zero) (Klemperer 1996). The net end of the rotation more shade-tolerant tree species stumpage value of thinning revenues relative to the can be expected to dominate the overstorey in these total revenue (thinning and final harvest) was used as parts of the stand. Canopy gaps may occur due to a supplementary indicator. The scenarios presented disturbance from wind and tree diseases, but will here for production of high-quality oak timber all rely Table 1. Rough estimates of economic indicators for scenarios under Option A: oak managed for high-quality timber with a target diameter (dbh) of 70 cm and an expected rotation length of 120 years, with or without an admixture and with or without pruning. Legend: CF = cash flow positive from the indicated age-class, Pay-back = the number of years before complete pay- pos back of establishment costs, NSV = net stumpage value (NSV refers to thinning revenues), IRR = internal rate of return thin (IRR refers to IRR of the marginal investment of a specified management action). Missing values (–) indicate that no specific marg information was available. All estimates are approximations and based on analyses by Hermansen (1956), Holten (1980, 1986), Ståål (1986), Staun (1989), Jensen (1989, 1993), Madsen (1991), Lindén (2003), Skovsgaard (2004), Lomholt (2006) and Jørgensen (2013). Pos. CF Pay-back NSV NSV / IRR IRR thin marg Admixture Pruning (age) (years) k€/ha NSV % % total None No 50 90 50–60 .40–.50 2.5–3.0 – Conifer No 25 40 55–65 – 3.5 – None Yes 50 100 55–75 .25–.35 – 2.0–3.5 Note: The examples all include selection and marking of 50–100 potential future crop trees per ha. Estimates of NPV are updated to the price level of 2015. 64 M. LÖF ET AL. be taken as an indication of a stable balance on suitable site conditions and a continuously opti- between supply and demand in the market for oak mal management throughout the rotation. Deviation from these conditions may reduce the economic timber. The main consequence of high-pruning is the outcome. increased revenue that results from the improved The most relevant alternative silvicultural scenar- ios for timber production under Option A include the quality of final crop trees. High-pruning remains profitable at an internal rate of return of 2.0–3.5% possible admixture of other tree species (hardwoods, when considering only the added value on the oak conifers or both), the use of permanently marked potential future crop trees and high-pruning. A timber (Jensen 1989, 1993; Skovsgaard 2004). This is based on conservative estimates and corresponds to stand managed without an understorey and without an added income of 20% or more from pruning. high-pruning is considered the base scenario against which the economic outcome of other alternatives Finally, the practice of leaving a few retention trees under Regime A will be compared (Table 1). The at regeneration as prescribed by Swedish legislation is considered of no or only marginal importance for the admixture of one or more hardwood species (instead of a conifer) does not significantly change any of the economic outcome of management Option A. economic indicators and this option is therefore not Retention trees in oak stands will usually be selected considered in the comparison. among those of inferior stem quality or located at The cash-flow profile of the base scenario for stand margins and are consequently of little monetary Regime A is strongly skewed because of establish- value. ment and tending costs that are not compensated There is essentially no information available for by early thinning revenues. In contrast, the regarding the actual costs and incomes of manage- admixture of a fast-growing conifer (for example ment Regime B or the factors upon which they Norway spruce) provides early revenues from timber depend. Option B is currently being explored in harvesting, which provides a potentially positive cash some commercial forest stands in the states of flow after 25 years. After approximately 50 years, net Rheinland-Pfalz and Saarland in Germany (Wilhelm income from the oak takes over as the conifer trees & Rieger 2013). In addition, in Denmark three thin- are gradually removed from the stand. The pay-back ning experiments in young oak include treatments period with an admixture of Norway spruce may be similar to Option B (Jensen & Skovsgaard 2009). less than half of the base scenario. These experiments were installed in 2002–2003 and The net stumpage value of the investment in growing no results are as yet available. Nevertheless, we can oak for high-quality timber in heavily thinned stands expect that the reduction in income will be propor- −1 under Regime A is approximately 55,000 € ha tional to the number of final timber trees negatively or affected by Option B plus some additional costs due more, depending mainly on market conditions and to affected logistics during forest operations the quality of the timber at final harvest (Table 1). The (Wilhelm & Rieger 2013). In the event of a larger ratio of revenues from thinnings to that of the total impact comprising half or more of the potential final revenue from thinnings and final harvest depends heav- timber trees, the income reduction would probably be ily on the quality of timber and on prices for premium marginally larger than the actual share of negatively timber and lower classes of grading. This is indicated by the variation in NSV affected potential final timber trees. In addition, no /NSV (Table 1) thin total timber trees will be managed or extracted from man- The magnitude of the added revenue from an agement Regime C which is why this regime will admixture of conifers obviously depends on the growth and proportion of conifers in the stand and produce zero costs and incomes. on the final quality of the oak stems. The example in this review has been estimated to provide an addi- 4.2. Biodiversity tional 10% compared to a pure stand of oak (Madsen 1991). The main economic effect of the conifers, 4.2.1. Vascular plants however, is improved (positive) cash-flow earlier in By increasing light availability and providing dis- the rotation. Mixed stands (i.e., oak and spruce) are turbed ground for plant recruitment, canopy thinning also considered less sensitive to changes in economic of oak forest (Regime A) generally increases under- conditions than monocultures (Lindén 2003). storey plant species richness. This increase may occur Based on sources used in this analysis (see refer- at least up to a decade into the rotation (Brunet et al. ences in text for Table 1), the internal rate of return 1996, 1997). Also, partial cutting, corresponding to on the investment in the base scenario has Option B, will probably have a clear effect on the remained relatively constant at 2.5–3.0% since the herbaceous layer and will likewise increase plant spe- 1950s, indicating a low investment risk. cies richness during the first growing season after Considering changes in technologies, work salaries cutting (Götmark et al. 2005). Most typical forest and society at large during this period, this can also plants are tolerant of canopy thinning, or may even INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 65 species with rapid and vigorous sprouting such as increase in abundance or frequency with its occur- hazel and lime tree (Leonardsson & Götmark 2015). rence (Brunet et al. 1996; Götmark et al. 2005). The abundance of many other plant species (ruderal, 4.2.2. Lichens and bryophytes grassland and habitat generalists) also increases with Previous studies have found that the thinning of trees thinning intensity, which results in a short-term increase of total species richness with increasing and shrubs around large oaks (Options A and B) management intensity (Brunet et al. 1996, 1997). helped to maintain the species density of epiphytic The composition of the vegetation emerging after lichens and bryophytes on oak trunks, including sev- cutting also depends on previous land use history eral species of conservation concern, relative to and its effects on the seed bank, and on patches of unmanaged forest (Nordén et al. 2012). In a related remnant grassland vegetation in the forest (Milberg study by Paltto et al. (2008), the composition of 1995; Jonason et al. 2014). lichens and bryophytes on tree stumps shifted Conversely, the gradual loss of light-demanding towards species adapted to drier dead wood as a herbaceous species under closing canopies in unma- result of thinning, implying an increase of lichen naged forest (C) is usually not compensated by an richness and a trend of decreasing bryophyte rich- equivalent gain of shade tolerant plant species, which ness. Similar trends were also recorded on tree logs. results in an overall decrease of species richness (Von However, no increases or decreases in species of Oheimb & Brunet 2007). Likewise, when comparing conservation concern could be detected (Paltto et al. different types of oak-hazel woodland, Hansson 2008). Similar to vascular plants, thinning had a (2001) found that areas managed as wooded meadow positive effect on bryophytes associated with the for- or wood pastures maintained a significantly higher est floor (Götmark 2013). We conclude from these total number of vascular plant species than unma- studies that overall epiphyte diversity may be highest naged woodlands. Herb layer plants greatly vary in at the stand level in Option B, by providing the abundance along the light gradient (Tyler 1989), and highest diversity of micro-habitats in terms of tree we conclude that total vascular plant diversity at the species and micro-climate (Table 2). The proportion end of the rotation period is maximized in oak forests of lichens compared to bryophytes will probably with both open and closed canopy parts. Therefore, decrease with decreasing management intensity vascular plant diversity is probably greatest under from A to C. Regime B which provides the highest structural het- erogeneity due to partial understorey clearing 4.2.3. Birds (Table 2). Bird diversity and composition is strongly influ- Similarly to the herbaceous vegetation, the diver- enced by the structural heterogeneity of a forest sity of regenerating woody plant species in the her- stand, and in particular the understorey density baceous layer generally increases after canopy (Hinsley et al. 2009;Hewsonetal. 2011). In a thinning (Brunet et al. 1996). Concerning oak itself, study of understorey importance for bird commu- seedling density was found to be threefold higher in nities in the oak-rich urban woodlands of western partially cut oak stands (Option B) than in unma- Sweden, 90% removal of the understorey had a naged control plots (Option C) three years after cut- significant negative affect on total breeding bird ting, and survival and growth were positively related species densities, while 50% removal did not affect to canopy openness (Götmark 2007). Repeated densities compared to the unmanaged controls understorey cutting may lead to changes in the com- (Heyman 2010). This indicates that a moderate position of the woody understorey and may favor disturbance of the understorey consistent with B (and some A alternatives) may not affect the avi- fauna negatively. Another Swedish study showed Table 2. Relative value of the different management options that closed oak-hazel woodlands derived from for- in relation to their promotion of structures important for mer wooded grassland, corresponding to Option C, biodiversity. Maximum value for positive effects at stand supportedalowernumberofbirdspeciesthan age of ca 120 years is +++. woodlands of similar origin yet retaining some Species Stand characteristics Option A Option B Option C groups degree of disturbance and open areas, and thus Stand structural + +++ ++ Plants, most closely resembling Option B (Hansson 2001). heterogeneity Epiphytes, Even if Option B does not involve a creation of Birds Tree species diversity + +++ ++ Epiphytes, grazed and fully open sections of the forests, Birds, repeated disturbances from the thinning and man- Beetles Live trees with cavities + ++ +++ Birds, agement of the crop trees are probably beneficial to and dead branches Beetles some speciesaswellasfor theoveralldiversityof Dead wood + ++ +++ Beetles, the stand. To summarize, management Options C Saproxylic fungi and B would produce more heterogeneous 66 M. LÖF ET AL. structure, with a more developed understorey, com- average 14.3 m /ha coarse dead wood, which is pared to A. To distinguish between B and C in this twice as much as the mean value for all forests in regard is more difficult. Even though C represents a Sweden (Nordén et al. 2004). In the same study, fine less disturbed habitat, B would include more struc- dead wood made up another 12 m /ha. The stands tural heterogeneity due to the juxtaposition of two evaluated in that study are similar to our models for relatively distinct habitat types, including unma- Option C (and parts of B), namely abandoned wood- naged areas. Furthermore, the crop trees will prob- land pastures. These stand types can be expected to ably be taller and coarser as compared to most of accumulate more dead wood through self-thinning, the trees in the unmanaged sections of the stand, and thus have higher beetle diversity compared to the creating an extra structural dimension (Table 2). pure production Option A. However, if logging resi- Tree species composition is also an important dues are left after thinnings in A and B, this can determinant of forest bird diversity (Poulsen 2002). readily benefit many species (Jonsell et al. 2007). Many bird species are exclusively associated with Stand openness is also important for many oak- broad-leaved, coniferous or mixed forests, thereby associated beetles (Ranius & Jansson 2000; Widerberg causing an important divergence in the composition et al. 2012; Gough et al. 2014). In a study of retained of bird communities associated with either forest type oaks in nine spruce production forests in southern (Bibby et al. 1989; Roberge & Angelstam 2006). Sweden, oaks exposed to intermediate insolation har- Several studies have shown that the bird communities bored more species than shaded oaks (Widerberg within mixed stands of conifers and broadleaf trees 2013). Similar results were obtained in an experiment are composed of a mixture of bird species represent- in 22 oak-rich stands in southern Sweden where ing both the broadleaf- and coniferous-associated about 25% of the tree basal area was cut in 1 ha fauna (Donald et al. 1998; Hausner et al. 2002). plots and then compared with uncut plots (Franc & Hence, the mixed overstories of Options B and C Götmark 2008). Saproxylic beetle species richness will probably result in higher bird diversity than the increased by about 35% in the harvested plots, rela- base Option A, which is managed towards a pure oak tive to the reference plots. The results of both of these overstorey (Table 2). studies indicate that Option B could be more bene- Old trees are another important structural feature ficial for saproxylic species than the alternatives con- for birds (Nilsson et al. 2001; Poulsen 2002), in par- sidered (Table 2), due to the combination of dead ticular for cavity nesters (Carlson et al. 1998). A wood and favorable light conditions. In addition, the recent study from southern Sweden comparing oak larger tree diversity in Options C and B will most plantations of different ages with natural oak-rich probably be beneficial for saproxylic beetles com- forests shows that mid-age oak plantations with a pared to the potential monocultures of some A alter- woody understorey and managed according to natives (Jonsell et al. 1998; Felton et al. 2010b). Option A had a comparable bird diversity to that of Species richness of fungal fruiting bodies on fine older natural forests. However, the community com- woody debris (1–10 cm in diameter) was negatively position differed primarily due to a lack of many tree affected by thinning but remained unaffected on hole-nesting species (Felton et al., unpublished coarse dead wood (>10 cm) in a study of experimen- results). Neither of the options assessed here include tal partial cutting in southern Swedish oak-rich for- trees older than the stand itself, and few of the trees ests. Overall species composition did not change will develop large amounts of ‘old-tree structures’ significantly as a result of partial cutting, but red- during the 120 years considered. Nevertheless, a few listed species tended to decrease in thinned plots trees in the unmanaged stand (C) and unmanaged (Nordén et al. 2008). The available data are too scarce parts of the stand in B can be expected to develop to allow general conclusions. However, the assess- cavities during the time frame considered, and ment made here indicates that the diversity of increasingly as time progresses, as large branches saproxylic fungi may be highest under Option C, die from shading and eventually break off. where a moister microclimate is maintained and also where the most dead wood will accumulate due to self-thinning. As long as substantial parts of the 4.2.4. Saproxylic beetles and fungi stand remain unmanaged in Option B, negative Oak is the most important tree in northern Europe effects of tree harvest may remain negligible, while for saproxylic beetles. Over 500 different species, Option A probably will cause a considerable decrease many of them red-listed, use oak during part or all of saproxylic fungal diversity (Table 2). of their life cycle (Jonsell et al. 1998; Dahlberg & Stokland 2004). There is a large body of literature showing the importance of the type and quantity of 4.3. Cultural services dead wood for saproxylic species richness (e.g., Jonsson et al. 2005; Franc et al. 2007). Semi-natural The nonmaterial benefits people obtain from ecosys- oak-rich stands in southern Sweden contained on tems are often referred to as cultural services (MEA, INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 67 trees, if they are in good condition (Bostedt & 2005). These services include spiritual and religious Mattsson 1995; Tönnes et al. 2004). Regime A is the values, inspiration, aesthetic values, recreational values and cultural heritage values, and have implica- most intensively managed out of the three alterna- tives. Whereas public preferences regarding forest tions for social relations and one’s sense of place. management can often vary among studies and with Their importance for society and thus need for inte- gration into decision-making has been increasingly the characteristics of the sample population, people tend to favor managed forests containing few visible acknowledged (Bowler et al. 2010; Daniel et al. traces of human activities or large disturbances (Ribe 2012). The most studied cultural services in forests are recreational and aesthetic values. Recreation 1989; Mattsson & Li 1994; Gundersen & Frivold 2008). Therefore, in regard to anthropogenic distur- values increases with the share of temperate broad- bances, clear-cuttings are believed to have the stron- leaved trees, such as oak found within a stand (Norman et al. 2010; Johansson et al. 2014). gest and the longest negative impact on scenic beauty. Structural composition of the stand is, however, Management Regime C may be perceived as ‘messy’ by the general public (Table 3). The pre- often considered more important than tree species composition (Edwards et al. 2012). Overall, tree size, sence of large amounts of dead wood (from self- openness and visual penetration into the stand as well thinning) coupled with a high density of trees and as visual diversity of the stand are the most common thus low visual penetration and absence of forest predictors of public preferences (Ribe 1989; management results in negative attitudes and low Gundersen & Frivold 2008). recreation and aesthetic values of such environ- The development stage of the stand has also been ments (Hultman 1983). However, variation in pre- shown to be an important contributor to its recrea- ferences exist between people with different levels tional value. According to a Danish study (Jensen & of knowledge about forests and their functioning Koch 2000), this effect is more prominent in broad- (Gundersen & Frivold 2008;Erikssonetal. 2012). leaved stands than in conifers. Lower recreational and Potentially countering these aspects, Regime C also aesthetic values of young stands can be attributed to contains a variety of features often considered their relatively high density (Ribe 1989). The same favorable, including medium to large trees and effect is often seen in relation to the presence of a structural diversity. Fallen twigs and branches asso- dense understorey and is probably linked to accessi- ciated with older trees may diminish recreational bility and safety considerations (Jørgensen et al. and aesthetic values, but a number of more recent 2009). For forest management this suggests the studies suggest that since the 1990s natural forests potential positive effects of thinning young and med- have been considered more suitable for recreation ium-aged stands, especially when leaving the most than they were in previous decades (Lindhagen & vigorous and attractive trees on the site. However, Hörnsten 2000; Tahvanainen et al. 2001). Relatedly, high-intensity thinnings in young oak stands could there seems to be an increasing acceptance and also decrease the impression of accessibility due to appreciation of dead wood (Tyrväinen et al. 2003; the presence of high volumes of thinning slash on the Nielsen et al. 2007;Heyman 2012). This develop- site (Jensen & Skovsgaard 2009). ment has been attributed to increasing levels of Management Regime A is likely to be preferred knowledge among the general public about especially in the later stages of the stand develop- biodiversity. ment, providing big trees that are well distributed, Management Regime B probably provides many open environments and visual penetration (Table 3). desirable features for the general public such as However, from the point of visual diversity, the struc- medium and large trees, lower overall tree density ture of monocultures is much simpler than that in of some sections of the stand and structural diver- natural or mixed forests. Thus, the presence of an sity (Table 3). Semi-open stands increase access to understorey may be beneficial for recreational and more distant areas and openings and provide a aesthetic values. In addition, aesthetic and recrea- higher sense of security compared to dense stands tional values also increase with presence of retention (Kaplan & Kaplan 1989; Gundersen & Frivold 2008; Heyman et al. 2011). Additionally, management targeting only specific trees and leaving other Table 3. Relative recreational and aesthetic values in relation areas untouched contributes to a patchy and to stand characteristics for the three management regimes. visually diverse environment, thus potentially Maximum value for positive effects at stand age of ca. increasing recreational and aesthetic values of the 120 years is +++. forest (Ribe 1989;Lindgren 1995; Nielsen et al. Stand characteristics Option A Option B Option C Number of medium/large trees +++ ++ + 2007;Heyman 2012). Uneven-aged stands with a Openness +++ ++ + mixture of trees of different sizes are not only Visibility into the stand +++ ++ + preferred over monocultures, but this effect is Visual/structural diversity + +++ ++ Positive anthropogenic disturbance + +++ + maintained throughout the rotation (Hultman 68 M. LÖF ET AL. As expected, our evaluation of the free develop- 1983; Lindhagen & Hörnsten 2000). Overall, this ment regime (C) indicated substantially higher levels regime seems to avoid and/or mitigate some of the negatively perceived features of Regime A and of habitat provision for a range of taxonomic groups, which came at the expense of contributions to wood Regime C, by excluding large disturbances such as production and cultural services. The long-term tra- clear-cuttings in the end of the rotation, reducing stand density and providing higher levels of visual jectory of oak biodiversity values in these stands is however somewhat uncertain, as it depends on the diversity. Thus, Regime B is expected to be slightly extent of canopy disturbance and associated light morefavored by thepublicthanRegimeA,and much more favored than Regime C. conditions beneficial to the natural regeneration of oak. The availability of light is a primary determinant of oak seedling and sapling growth following canopy 5. Discussion disturbance (e.g. Jones 1959; Collet et al. 1997; Lüpke 2008; Parker & Dey 2008; Gardiner et al. 2009). For Any management decision that alters the tree species this reason the occurrence of oak has often been composition, structure or disturbance regimes of a linked to various natural and anthropogenic forest forest has the corresponding capacity to alter the disturbances, (e.g., livestock grazing, harvesting, fire range and balance of ecosystem services derived and windthrow) that provide increased insolation for from that forest. Previous studies have highlighted, oak regeneration and stump sprouting (Vera 2000; for example, that an unbalanced prioritization of Johnson et al. 2009; Brose et al. 2013). intensive timber production will reduce forest biodi- There is thus a potential synergistic interaction versity (Lindenmayer & Franklin 2002; Gossner et al. between oak management which provides conditions 2014), as well as the provisioning of regulatory and beneficial for oak regeneration and associated habi- cultural services (Bennett et al. 2009; Raudsepp- tats for biodiversity. If sufficient openness is main- Hearne et al. 2010). The potential for trade-offs is, tained, due to natural or anthropogenic disturbance, however, tempered by the existence of synergistic to ensure the continued persistence of an oak-domi- relationships among biodiversity and some ecosystem nated canopy, then a range of structural features of services derived from production forests (Duncker relevance to biodiversity in oak reserves can be et al. 2012). Our assessment lends support to the expected to increase as these stands mature beyond existence of both trade-offs and synergies among 120 years. This is due to the natural processes of the ecosystem service deliverables of oak-dominated growth; aging and senescence facilitating an increased stands, with some management alternatives providing abundance of structural features (e.g., large diameter outcomes far more balanced in terms of the overall dead wood, tree holes) often closely associated with delivery of ecosystem services. high biodiversity values (Berg et al. 1994; Remm & With respect to trade-offs, the most intensive oak- Lõhmus 2011). The importance of maintaining such production forest management regime (A) provided features within landscapes like southern Sweden, the highest levels of economic returns, slightly where land-use practices have largely reduced their decreased cultural values and the lowest biodiversity. occurrence, requires that oak reserves continue to be Notably however, the intensive production alternative a key component of protected broadleaf areas in assessed did not consist of only one silvicultural southern Sweden. This may require that in oak option, but included a range of management pre- reserves, in which the occurrence of natural distur- scriptions with different implications for both biodi- bance regimes is unlikely to maintain an oak-domi- versity and recreational values. Oak production nated forest system, active intervention takes place to alternatives considered in A ranged from monocul- ensure the continuation of oak-favorable successional tures to mixed-species forests, and also varied in dynamics (Götmark 2013). terms of the extent of understorey vegetation per- In contrast to the categorically distinct production mitted to develop. Thus the intensive production and biodiversity regimes (A & C), the ‘combined forest alternative (A) assessed varied in terms of tree goal’ management species diversity and structural heterogeneity allowed regime (B) unites both of these for, two aspects of direct relevance to evaluations of management priorities. Option B appears to provide the same potential value for biodiversity as C, the biodiversity and cultural value of a stand. As our assessments indicate, the biodiversity and cultural although over a smaller spatial extent, combined value of these stands could therefore be increased with approximately the same cultural services as A. Because of this, Option B provides forest owners with with minimal reductions to production and economic outcomes, by adopting silvicultural approaches that a high degree of flexibility with respect to the relative prioritization of different ecosystem services. Forest increase tree species composition and understorey owners can for example determine the extent to structural heterogeneity (Bostedt & Mattsson 1995; Lindenmayer & Franklin 2002; Tönnes et al. 2004; which different areas of a stand are allocated to man- agement practices more conducive to production, Brockerhoff et al. 2008). INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 69 ‘combined goal management regimes’ on timber pro- biodiversity and cultural values. Despite these cap- duction economy. abilities, this management alternative is so far pri- marily a hypothetical construct in southern Sweden. When evaluating the relative biodiversity and cul- tural value of any production forest alternative, a Our assessments suggest, however, that this manage- strong determinant of outcomes is the reference con- ment alternative may be a suitable means of balan- cing the provision of a more comprehensive suite of dition chosen. In this study, we contrasted intensely managed oak production forests (A) with alternatives the ecosystem services within a stand than the cur- possessing a substantial (B) or encompassing extent rently adopted management alternatives in this region. This balanced provision appears to be consis- (C) of total area allocated to unmanaged develop- ment. Whereas the intensive oak production manage- tent with increasing demands that forests are mana- ment alternative fared poorly with respect to ged for multiple goals (Gustafsson et al. 2012; Gamfeldt et al. 2013). An alternative approach biodiversity in this comparison, this outcome would would require a continuation of more categorically likely be reversed if comparison was made with the most common silvicultural alternative in the region: distinctive oak production stands and reserves that provide for the ecosystem requirements of societies at Norway spruce-dominated production forests. This landscape scales. The landscape scale balancing of difference only increases if the management regime ecosystem services does, however, raise complications allows for retention forestry and a biodiversity-rich (Duncker et al. 2012), especially in regions such as understorey. This is because even the most intensive southern Sweden dominated by a large number of oak production alternative is nevertheless consistent small-scale forest owners (McDermott et al. 2010; with regional biodiversity goals for the increased use Gustafsson et al. 2015). of broadleaf tree species, as advocated by the Swedish Despite the apparent capacity of alternative B to Forest Agency and forest certification programs provide suitable habitat for a substantial proportion (Gustafsson et al. 2010; Gustafsson & Perhans 2010; of a region’s flora and fauna, we nevertheless wish to Johansson et al. 2013). Likewise, broadleaf forests are emphasize that protected areas are an essential part of consistently preferred over spruce-dominated alter- any comprehensive framework for the protection of natives in terms of their recreational and aesthetic forest biodiversity. This is because the most demand- values (Norman et al. 2010). ing species often require relatively contiguous areas Whereas oak timber production values have been of complex and heterogeneous forest lands for which relatively stable since at least the 1950s, their recrea- disturbance regimes are allowed to approximate nat- tional and aesthetic values may be less stable. For ural patterns and processes (Hunter et al. 1988; example, demographic differences between various Kuuluvainen et al. 2012; Götmark 2013). Caution is categories of users exist, even though public prefer- thus warranted when evaluating the capacity of pro- ences exhibit high degree of consensus (Stamps duction forest to provide a balanced delivery of eco- 1999). Children find dense forests more fascinating, system services. In relation to biodiversity, extensive and mushroom and berry pickers favor more natural- reliance on forest management alternatives that strike looking environments (Ribe 1989; Eriksson et al. an optimal balance may nevertheless compromise the 2012). Forestry professionals and students tend to regional viability of populations only sustained in prefer medium- to high-density stands, whereas land- protected areas. scape architects favor low-density stands (Petucco In relation to timber production and associated et al. 2013). In addition, variation in preferences economic returns, the outcome of Regime B will exist between people with different levels of knowl- depend on the number of crop trees produced. edge about forests and their functioning (Gundersen Thus, Regime B will often produce less marketable & Frivold 2008; Eriksson et al. 2012). When respon- timber per hectare compared to A. Therefore, an dents in Denmark and southern Sweden were extensive reliance on only Regime B would probably informed about the importance of biodiversity, they reduce the economic returns from oak production in provided higher values for its conservation (Bakhtiari the region. The small size of stands, and their scat- 2014). Thus, values can change relatively rapidly tered placement in the landscape already increases through education, with corresponding influences harvesting inefficiencies and reduces available timber on preferences for forest use. for the industry. Thus, Regime A and Regime B are Management for the production of high-value tim- not interchangeable, but instead are complementary ber species like oaks and management for cultural management alternatives. Furthermore, if decisions services, or to conserve biodiversity can be in conflict were taken to manage the majority of forests simi- with each other. The results from our review have larly, the resultant homogeneity would likely decrease identified a management regime with combined their recreational and aesthetic values (Axelsson- management goals for the oak-dominated forests Lindgren & Sorte 1987). We would also like to which appears to provide a balanced delivery of tim- emphasize that little is known about the effects of ber production, biodiversity conservation and 70 M. LÖF ET AL. Berg Å, Ehnström B, Gustafsson L, Hallingbäck T, Jonsell cultural services. We would, however, like to stress M, Weslien J. 1994. Threatened plant, animal, and fun- that insufficient studies have been conducted to gus species in Swedish forests - Distribution and habitat determine the precise capacity of management associations. Conserv Biol. 8:718–731. Regime B to provide for production, biodiversity Bergfors U. 1990. Allemansrätten ur rättsekonomisk syn- and cultural values. Instead, our assessments had to punkt: historia, regelverk och dagsläge. – Arbetsrapport rely on information derived from management nr 115. Umeå: Inst. för skogsekonomi SLU. Bergquist J, Löf M, Örlander G. 2009. Effects of roe deer regimes which closely approximated this manage- browsing and site preparation on performance of ment alternative. To validate and expand upon our planted broadleaved and conifer seedlings when using findings, targeted field experiments are necessary to temporary fences. Scand J For Res. 24:308–317. determine the precise capacity of this management Bibby CJ, Aston N, Bellamy PE. 1989. Effects of broad- alternative to provide for ecosystem services of socie- leaved trees on birds of upland conifer plantations in tal interest. These findings would help stakeholders North Wales. Biol Conserv. 49:17–29. Bolte A, Ammer C, Löf M, Madsen P, Nabuurs G-J, Schall evaluate the feasibility of the three options in terms of P, Spathelf P, Rock J. 2009. Adaptive forest management implementation capacity and their respective desir- in central Europe: climate change impacts, strategies and ability in output of ecosystem services. This may be integrative concept. Scand J For Res. 24:473–482. achieved using a systematic framework such as multi- Bostedt G, Mattsson L. 1995. The value of forests for criteria decision analysis, which quantifies ecosystem tourism in Sweden. Ann Tour Res. 22:671–680. Bowler DE, Buyung-Ali LM, Knight TM, Pullin AS. 2010. services while highlighting the contribution of A systematic review of evidence for the added benefits to selected services. (Fontana et al. 2013). Such a process health of exposure to natural environments. BMC Public of stakeholder involvement would help test the accu- Health. 10:456. racy and comprehensiveness of ecosystem service Brockerhoff EG, Jactel H, Parrotta JA, Quine CP, Sayer J. projections, and in addition, may identify additional 2008. Plantation forests and biodiversity: oxymoron or silvicultural pathways for optimizing the delivery of opportunity? Biodiv Conserv. 17:925–951. Brose PH, Dey DC, Phillips RJ, Waldrop TA. 2013.A goods and services from oak forests at stand and meta-analysis of the fire-oak hypothesis: does prescribed landscape scales. burning promote oak reproduction in eastern North America? For Sci. 59:322–334. Brunet J, Falkengren-Grerup U, Rühling Å, Tyler G. 1997. Regional differences in floristic change in south Swedish Disclosure statement oak forests as related to soil chemistry and land use. J No potential conflict of interest was reported by the Veg Sci. 8:329–336. authors. Brunet J, Falkengren-Grerup U, Tyler G. 1996. Herb layer vegetation of south Swedish beech and oak forests - effects of management and soil acidity during one dec- Funding ade. For Ecol Manage. 88:259–272. Carbonnier C. 1975. Produktionen av kulturbestånd av ek i This research was funded by the Broadleaves for the Future södra Sverige. Stud For Suec. 125:1–89. program. Anna Filyushkina was funded by European Carlson A, Sandström U, Olsson K. 1998. Availability and Commission under Erasmus Mundus joint Doctoral use of natural tree holes by cavity nesting birds in a Programme ‘Forest and Nature for Society’ (FONASO), Swedish deciduous forest. Ardea. 86:109–119. and Adam Felton was partially funded by Future Forests, Collet C, Colin F, Bernier F. 1997. Height growth, shoot a research program supported by the Foundation for elongation and branch development of young Quercus Strategic Environmental Research (MISTRA). petraea grown under different levels of resource avail- ability. Ann Sci For. 54:65–81. Dahlberg A, Stokland JN. 2004. Vedlevande arters krav på substrat (Rapport nr. 7). Jönköping: Skogsstyrelsen. References Daniel TC, Muhar A, Arnberger A, Aznar O, Boyd JW, Chan KM, Costanza R, Elmqvist T, Flint CG, Gobster Almgren G, Ingelög T, Ehnström B, Mörtnäs A. 1984. PH, et al. 2012. Contributions of cultural services to the Ädellövskog – ekologi och skötsel. Jönköping: ecosystem services agenda. Proc Natl Acad Sci. Skogsstyrelsen. 109:8812–8819. Annerstedt M, Norman J, Boman M, Mattsson L, Grahn P. De Groot RS, Wilson MA, Boumans RMJ. 2002. A typology 2010. Finding stress relief in a forest. Ecol Bull. 53:33–42. for the classification, description and valuation of eco- Axelsson-Lindgren C, Sorte G. 1987. Public response to system functions, goods and services. Ecol Econ. 41:393– differences between visually distinguishable forest stands in a recreation area. Landsc Urban Plan. 14:211–217. Diekmann M. 1999. Southern deciduous forests. In: Rydin Bakhtiari F. 2014. Revealing components of the economic H, Snoeijs P, Diekmann M, editors. Swedish plant geo- value for environmental goods and services from forest: graphy. Acta Phytogeogr Suec 84:33–53. an application of stated preference methods for forest Donald PF, Fuller RJ, Evans AD, Gough SJ. 1998.Effects valuation and conservation [PhD dissertation]. of forest management and grazing on breeding bird Copenhagen: University of Copenhagen; 214 p. communities in plantations of broadleaved and con- Bennett EM, Peterson GD, Gordon LJ. 2009. iferous trees in western England. Biol Conserv. Understanding relationships among multiple ecosystem 85:183–197. services. Ecol Lett. 12:1394–1404. INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 71 Drössler L, Attocchi G, Jensen AM. 2012. Occurence and Gough LA, Birkemoe T, Sverdrup-Thygeson A. 2014. management of oak in southern Swedish forests. Reactive forest management can also be proactive for Forstarchiv. 83:163–169. wood-living beetles in hollow oak trees. Biol Conserv. Duncker PS, Barreiro SM, Hengeveld GM, Lind T, Mason 180:75–83. WL, Ambrozy S, Spiecker H. 2012. Classification of Gundersen VS, Frivold LH. 2008. Public preferences for forest management approaches: a new conceptual frame- forest structures: a review of quantitative surveys from work and its applicability to European forestry. Ecol Soc. Finland. Urban For Urban Green. 7:241–258. 17:51. Gustafsson L, Baker SC, Bauhus J, Beese WJ, Brodie A, Edwards DM, Jay M, Jensen FS, Lucas B, Marzano M, Kouki J, Lindenmayer DB, Lohmus A, Martinez PG, Montagné C, Peace A, Weiss G. 2012. Public preferences Messier C, et al. 2012. Retention forestry to maintain across Europe for different forest stand types as sites for multifunctional forests: a world perspective. Bioscience. recreation. Ecol Soc. 17:27. 62:633–645. Eriksson L, Nordlund AM, Olsson O, Westin K. 2012. Gustafsson L, Felton A, Felton AM, Brunet J, Caruso A, Recreation in different forest settings: a scene preference Hjälten J, Lindbladh M, Ranius T, Roberge J-M, Weslien study. Forests. 3:923–943. J. 2015. Natural versus national boundaries: the impor- Felton A, Ellingson L, Andersson E, Drössler L, Blennow K. tance of considering biogeographical patterns in forest 2010a. Adapting production forests in southern Sweden conservation policy. Conserv Lett. 8:50–57. to climate change – constraints and opportunities for Gustafsson L, Kouki J, Sverdrup-Thygeson A. 2010. Tree risk spreading. Int J Clim Change Strat Manage. 2:84–97. retention as a conservation measure in clear-cut forests Felton A, Lindbladh M, Brunet J, Fritz Ö. 2010b. Replacing of northern Europe: a review of ecological consequences. coniferous monocultures with mixed-species production Scand J For Res. 25:295–308. stands: an assessment of the potential benefits for forest Gustafsson L, Perhans K. 2010. Biodiversity conservation in biodiversity in northern Europe. For Ecol Manage. Swedish forests: ways forward for a 30-year-old multi- 260:939–947. scaled approach. Ambio. 39:546–554. Fontana V, Radtke A, Fedrigotti VB, Tappeiner U, Tasser Hansson L. 2001. Traditional management of forests: plant E, Zerbe S, Buchholz T. 2013. Comparing land-use alter- and bird community responses to alternative restoration natives: using the ecosystem services concept to define a of oak-hazel woodland in Sweden. Biodiv Conserv. multi-criteria decision analysis. Ecol Econ. 93:128–136. 10:1865–1873. Franc N, Götmark F. 2008. Openness in management: Hausner VH, Yoccoz NG, Strann KB, Ims RA. 2002. hands-off vs partial cutting in conservation forests, and Changes in bird communities by planting non-native the response of beetles. Biol Conserv. 141:2310–2321. spruce in coastal birch forests of northern Norway. Franc N, Götmark F, Okland B, Nordén B, Paltto H. 2007. Ecoscience. 9:470–481. Factors and scales potentially important for saproxylic Hermansen NK. 1956. Fra Bregentveds egeskove. beetles in temperate mixed oak forest. Biol Conserv. Økonomiske analyser. Dansk Skovforenings Tidsskrift. 135:86–98. 41:507–561. Gamfeldt L, Snäll T, Bagchi R, Jonsson M, Gustafsson L, Hewson CM, Austin GE, Gough SJ, Fuller RJ. 2011. Kjellander P, Ruiz-Jaen MC, Fröberg M, Stendahl J, Species-specific responses of woodland birds to stand- Philipson CD, et al. 2013. Higher levels of multiple level habitat characteristics: the dual importance of for- ecosystem services are found in forests with more tree est structure and floristics. For Ecol Manage. 261:1224– species. Nature Comm. 4:1340. 1240. Gardiner ES, Löf M, O´Brien JJ, Stanturf JA, Madsen P. Heyman E. 2010. Clearance of understorey in urban wood- 2009. Photosynthetic characteristics of Fagus sylvatica lands: assessing impact on bird abundance and diversity. and Quercus robur established for stand conversion For Ecol Manage. 260:125–131. from Picea abies. For Ecol Manage. 258:868–878. Heyman E. 2012. Analysing recreational values and man- Garrido P. 2014. Cultural Oak landscapes as green infra- agement effects in an urban forest with the visitor- structure for human well-being [Licentiate Thesis]. employed photography method. Urban For Urban Skinnskatteberg: Swedish University of Agricultural Green. 11:267–277. Sciences. 74 p. Heyman E, Gunnarsson B, Stenseke M, Henningsson S, Gossner M, Schall P, Ammer C, Ammer U, Engel K, Tim G. 2011. Openness as a key-variable for analysis of Schubert H, Simo U, Utschick H, Weisser WW. 2014. management trade-offs in urban woodlands. Urban For Forest management intensity measures as alternative to Urban Green. 10:281–293. stand properties for quantifying effects on biodiversity. Hinsley SA, Hill RA, Fuller RJ, Bellamy PE, Rothery P. Ecosphere. 5:113. 2009. Bird species distributions across woodland canopy Götmark F. 2007. Careful partial harvesting in conservation structure gradients. Comm Ecol. 10:99–110. stands and retention of large oaks favour oak regenera- Holten NE. 1980. En af Hauchs egekulturer samt egedyrkning tion. Biol Conserv. 140:349–358. og inflation. Dansk Skovforenings Tidsskrift. 65:296–340. Götmark F. 2009. Conflicts in conservation: woodland key Holten NE. 1986. Eichenwirtschaft - dargestellt an habitats, authorities and private forest owners in Beispielen aus dem dänischen Forstamt Bregentved. Sweden. Scand J For Res. 24:504–514. Forstarchiv. 57:221–227. Götmark F. 2013. Habitat management alternatives for Hultman SG. 1983. Public judgement of forest environ- conservation forests in the temperate zone: review, ments as recreation areas. 2. A national survey. Report synthesis, and implications. For Ecol Manage. 306:292– No. 28. Uppsala: Section of Environmental Forestry, 307. Swedish University of Agricultural Sciences. (in Götmark F, Paltto H, Nordén B, Götmark E. 2005. Swedish, with English summary). Evaluating partial cutting in broadleaved temperate for- Hunter ML, Jacobson GL, Webb T. 1988. Paleoecology and est under strong experimental control: short-term effects the coarse-filter approach to maintaining biological on herbaceous plants. For Ecol Manage. 214:124–141. diversity. Conserv Biol. 2:375–385. 72 M. LÖF ET AL. Jensen FS, Koch NE. 2000. Measuring forest preferences of Lindhagen A, Hörnsten L. 2000. Forest recreation in 1977 the population - a Danish approach. Schweiz Z Forstwes. and 1997 in Sweden: changes in public preferences and 151:11–16. behaviour. Forestry. 73:143–153. Jensen FS, Skovsgaard JP. 2009. Precommercial thinning of Löf M, Brunet J, Hickler T, Birkedal M, Jensen MA. 2012. pedunculate oak: recreational preferences of the popula- Restoring broadleaved forests in southern Sweden as cli- tion of Denmark for different thinning practices in mate changes. In: Stanturf JA, Madsen P, Lamb D, editors. young stands. Scand J For Res. 24:28–36. A goal-oriented approach to forest landscape restoration. Jensen JS. 1989. Økonomien i vanrisopkapning. Dansk Berlin: World Forests XVI, Springer; p. 373–391. Skovbrugs Tidsskrift. 74:113–124. Lomholt A. 2006. Egens økonomiske tomgangsproblem. Jensen JS. 1993. Vanris på dansk stilkeg. Forskningsserien. Skoven. 38:122–125. 4:1–105. Lüpke BV. 2008. Einfluss unterschiedlicher Hiebsformen Johansson M, Gyllin M, Witzell J, Küller M. 2014. Does auf die Naturverjüngung eines Traubeneichen-Buchen- biological quality matter? Direct and reflected appraisal Mischbestandes (In german). Influence of various cut- of biodiversity in temperate deciduous broad-leaf forest. ting types on natural regeneration of a sessile oak-beech Urban For Urban Green. 13:28–37. mixed stand. Forsarchiv. 79:4–14. Johansson T, Hjälten J, De Jong J, Von Stedingk H. 2013. Madsen EM. 1991. Kan granen lösa tomgångsproblemet? Environmental considerations from legislation and certi- Skog & Forskning. 2:36–41. fication in managed forest stands: a review of their impor- Mattsson L, Li C. 1994. How do different forest manage- tance for biodiversity. For Ecol Manage. 303:98–112. ment practices affect the non-timber value of forests?— Johnson PS, Shifley SR, Rogers R. 2009. The ecology and an economic analysis. J Environ Manage. 41:79–88. silviculture of oaks. 2nd ed. Wallingford: CABI McDermott CL, Cashore B, Kanowski P. 2010. Global Publishing. environmental forest policies: an international compar- Jonason D, Ibbe M, Milberg P, Tunér A, Westerberg L, ison. Oxon: Earthscan. Bergman KO. 2014. Vegetation in clear-cuts depends on Milberg P. 1995. Soil seed bank after eighteen years of succes- previous land use: a century-old grassland legacy. Ecol sion from grassland to forest. Oikos. 72:3–13. Evol. 4:4287–4295. [MEA] Millennium Ecosystem Assessment. 2005. Ecosystems Jones EW. 1959. Biological flora of the British Isles: and human well-being: biodiversity synthesis [Internet]. Quercus L. J Ecol. 47:169–222. World Resources Institute; [cited 2015 Feb 24]. Available Jonsell M, Hansson J, Wedmo L. 2007. Diversity of from: http://www.unep.org/maweb/documents/document. saproxylic beetle species in logging residues in Sweden 354.aspx.pdf - Comparisons between tree species and diameters. Biol Nielsen AB, Olsen SB, Lundhede T. 2007. An economic Conserv. 138:89–99. valuation of the recreational benefits associated with Jonsell M, Weslien J, Ehnström B. 1998. Substrate require- nature-based forest management practices. Landsc ments of red-listed saproxylic invertebrates in Sweden. Urban Plan. 80:63–71. Biodivers Conserv. 7:749–764. Nilsson SG, Hedin J, Niklasson M. 2001. Biodiversity and Jonsson BG, Kruys N, Ranius T. 2005. Ecology of species its assessment in boreal and nemoral forests. Scand J living on dead wood - lessons for dead wood manage- Forest Res Suppl. 3:10–26. ment. Silva Fenn. 39:289–309. Nordén B, Götmark F, Ryberg M, Paltto H, Allmér J. 2008. Jørgensen BB, Jensen FS, Skovsgaard JP. 2009. Partial cutting reduces species richness of fungi on Underplantning af eg: øger underplantning skovens woody debris. Can J For Res. 38:1807–1816. rekreative værdi? Skoven. 41:203–207. Nordén B, Götmark F, Tönnberg M, Ryberg M. 2004. Dead Jørgensen JH. 2013. Bregentveds egedyrkning ved en wood in semi-natural temperate broadleaved woodland: milepæl. Skoven. 45:121–125. contribution of coarse and fine dead wood, Kaplan R, Kaplan S. 1989. The experience of nature. attached dead wood and stumps. For Ecol Manage. Cambridge: Cambridge University Press. 194:235–248. Klemperer WD. 1996. Forest resource economics and Nordén B, Paltto H, Claesson C, Götmark F. 2012. Partial finance. New York (NY): McGraw-Hill. cutting can enhance epiphyte conservation in temperate Kuuluvainen T, Tahvonen O, Aakala T. 2012. Even-aged oak-rich forests. For Ecol Manage. 270:35–44. and uneven-aged forest management in Boreal Norman J, Ellingson L, Boman M, Mattsson L. 2010. The Fennoscandia: a review. Ambio. 41:720–737. value of forests for outdoor recreation in southern Leonardsson J, Götmark F. 2015. Differential survival and Sweden : are broadleaved trees important ? Ecol Bull. growth of stumps in 14 woody species after conservation 53:21–31. thinning in mixed oak-rich temperate forests. Eur J Nylinder M, Woxblom L, Fryk H. 2006. Ädellöv – virke Forest Res. 134:199–209. och förädling. Uppsala: Sveriges Lantbruksuniversitet, Lindbladh M, Foster DR. 2010. Dynamics of long-lived Institutionen för skogens produkter och marknader. foundation species: the history of Quercus in southern Paltto H, Nordén B, Götmark F. 2008. Partial cutting as a Scandinavia. J Ecol. 98:1330–1345. conservation alternative for oak (Quercus spp.) forest— Lindén M. 2003. Increment and yield in mixed stands with Response of bryophytes and lichens on dead wood. For Norway spruce in southern Sweden [Doctoral thesis]. Ecol Manage. 256:536–547. Swedish University of Agricultural Sciences. Silvestria 260. Parker WC, Dey DC. 2008. Influence of overstory density Lindenmayer DB, Franklin JF. 2002. Conserving forest on ecophysiology of red oak (Quercus rubra) and sugar biodiversity: a comprehensive multiscaled approach. maple (Acer saccharum) seedlings in central Ontario Washington: Island Press. shelterwoods. Tree Physiol. 28:797–804. Lindgren. 1995. Forest aesthetics. In: Hytönen M, editor. Petucco C, Skovsgaard JP, Jensen FS. 2013.Recreational Multiple-use forestry in the Nordic Countries. Metla: preferences depending on thinning practice in young Finnish Forest Research Institute, Helsinki Research even-aged stands of pedunculate oak (Quercus robur Centre; p. 279–293. L.): comparing the opinions of forest and landscape INTERNATIONAL JOURNAL OF BIODIVERSITY SCIENCE, ECOSYSTEM SERVICES & MANAGEMENT 73 experts and the general population of Denmark. Scan J perceptions–visual versus verbal information. Landsc For Res. 28:668–676. Urban Plan. 53:53–70. Poulsen BO. 2002. Avian richness and abundance in tem- Thompson ID, Okabe K, Tylianakis JM, Kumar P, perate Danish forests: tree variables important to birds Brockerhoff EG, Schellhorn NA, Parrotta JA, Nasi R. and their conservation. Biodiv Conserv. 11:1551–1566. 2011. Forest biodiversity and the delivery of ecosystem Ranius T, Jansson N. 2000. The influence of forest regrowth, goods and services: translating science into policy. original canopy cover and tree size on saproxylic beetles Bioscience. 61:972–981. associated with old oaks. Biol Conserv. 95:85–94. Tönnes S, Karjalainen E, Löfström I, Neuvonen M. 2004. Raudsepp-Hearne C, Peterson GD, Bennett E. 2010. Scenic impacts of retention trees in clear-cutting areas. Ecosystem service bundles for analyzing tradeoffs in Scand J For Res. 19:348–357. diverse landscapes. Proc Natl Acad Sci. 107:5242–5247. Tyler G. 1989. Interacting effects of soil acidity and Remm J, Lõhmus A. 2011. Tree cavities in forests–the broad canopy cover on the species composition of field-layer distribution pattern of a keystone structure for biodiver- vegetation in oak/hornbeam forests. For Ecol Manage. sity. For Ecol Manage. 262:579–585. 28:101–114. Ribe RG. 1989. The aesthetics of forestry: what has empirical Tyrväinen L, Silvennoinen H, Kolehmainen O. 2003. preference research taught us? Environ Manage. 13:55–74. Ecological and aesthetic values in urban forest manage- Roberge JM, Angelstam P. 2006. Indicator species among ment. Urban For Urban Green. 1:135–149. resident forest birds – a cross-regional evaluation in Vandekerkhove K, Keersmaeker LD, Menke N, Meyer P, northern Europe. Biol Conserv. 130:134–147. Verschelde P. 2009. When nature takes over from man: Schwenk WS, Donovan TM, Keeton WS, Nunery JS. 2012. dead wood accumulation in previously managed oak and Carbon storage, timber production, and biodiversity: beech woodlands in North-western and Central Europe. comparing ecosystem services with multi-criteria deci- For Ecol Manage. 258:425–435. sion analysis. Ecol Appl. 22:1612–1627. Vera FWM. 2000. Grazing ecology and forest history. UK: Simonsson P, Gustafsson L, Östlund L. 2015. Retention Cabi Publishing. forestry in Sweden: driving forces, debate and imple- Von Oheimb G, Brunet J. 2007. Dalby Söderskog revisited: mentation 1968-2003. Scand J For Res. 30:154–173. long-term vegetation changes in a south Swedish decid- Skovsgaard JP. 2004. Produktion af kvalitetstræ i eg: er uous forest. Acta Oecol. 31:229–242. eventyret bæredygtigt? Skoven. 36:532–535. Werner M, Rytter L, Stener L-G. 2000. Förbättrat Ståål E. 1986. Eken i skogen och landskapet. Växjö: Södra lövvedutnyttjande för vidareförädling. Redogörelse nr skogsägarna. 3. Uppsala: SkogForsk. Stamps AE. 1999. Demographic effects in environmental Widerberg KM. 2013. Oak as retention tree in commercial aesthetics: a meta-analysis. J Plan Lit. 14:155–175. spruce forests [Doctoral thesis No. 2013:66]. Sweden: Staun H. 1989. Intensiv egedyrkning. Dansk Skovbrugs Faculty of Forest Sciences, SLU. Tidsskrift. 74:87–112. Widerberg KM, Ranius T, Drobyshev I, Nilsson U, Swedish Forest Agency. 2014. Statistical yearbook of for- Lindbladh M. 2012. Increased openness around retained estry 2014. Jönköping: Official statistics of Sweden, oaks increases species richness of saproxylic beetles. Swedish Forest Agency. Biodiv Conserv. 21:3035–3059. Tahvanainen L, Tyrväinen L, Ihalainen M, Vuorela N, Wilhelm GJ, Rieger H. 2013. Naturnahe Waldwirtschaft Kolehmainen O. 2001. Forest management and public - die QD-Strategie. Stuttgart: Eugen Ulmer Verlag.

Journal

International Journal of Biodiversity Science, Ecosystem Services & Management – Taylor & Francis

Published: Jan 2, 2016

Keywords: Ecosystem services; oak forests; Sweden; synergies; trade-offs; multiple-use forestry; silviculture

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