This articlehas beenaccepted for publicationundergone and full peer review buthas not been a 3 2 1 Authors Running title generalist and deadwood-dwelling specialist fungi on size fragment forest and edges anthropogenic transient of effects long-term Contrasting HandlingMartin Nuñez Editor: Article StandardType: Paper Date: 03-Nov-2016 Accepted Revised Date:12-Oct-2016 Received Date: 10-Jun-2016 This article is This protectedAll rights byreserved. copyright. 10.1111/1365-2664.12835 Version the and version this between differences throughcopyediting, the typesetting, pagination and proofreading process, which may leadto [email protected] Sweden. Sweden. Affiliations Corresponding Telephone:author. +46 18 672453, Fax: +46 18 67 3537, E-mail: AcceptedDepartment of Natural Sciences, Mid Sweden University, Sundsvall, SE-851 70, Sweden. SwedishSpecies Information Centre, SLU.SE-750 07. Uppsala, Sweden. Ecology Department, Swedish UniversityAgricultural of Sciences SE-750 (SLU). 07. Uppsala, Article : Alejandro Ruete : Alejandro : Long-term effects of transient edges edges transient of effects Long-term : 1a , Tord Snäll 2 , BengtJonsson Gunnar of Record. Please cite this article as doi: asdoi: article this cite Please Record. of 3 , Mari, Jönsson 2

fungi whilst commongeneralist species like around old-growth forest fragments lead to lower predicted occupancy of old-growth indicator 3. Small fragment size (1–3.14 ha) and intensifie ha). sizes (1–20 and fragment 100%) 50%, (0%, 25%, in forest adjacent clear-cutting of scenarios of logsgroup a by of old-growth forest indicatorfungiand common two fungi underdifferent We2. usedBayesian models fittedto empirical datayears topredict 40 ofoccupancy dynamics dwelling fungi inborealold-growth forestfragments. deadwood- of occupancy future the sizeon fragment and edges anthropogenic of impacts future the predict we study, In this edges. fragment along uses land human to exposure greater and area reduced with patches forest creating worldwide, fragmented increasingly Forests becoming are 1. Summary This article is This protectedAll rights byreserved. copyright. Acceptedthe increase may buffers) dynamic (i.e. time over effect edge the reduces that way a in cutting clear- adjacent non-simultaneous zones. Alternatively, buffer permanent with areas conservation of size the increasing on focus should Management effects. edge transient degrading occupancy of old-growth fungi, small fragmentsmanaged needbe to protect to species from for the long-term measures conservation area-based effective As landscapes. forest managed in structures) (i.e.species, quality last habitat the vestiges high of represent often fragments Synthesis5. and applications 11. target Aichi asthe such areas, protected for targets conservation towards for working when accounted be at should edge the occupancy fungal in changes These clear-cutting. adjacent increased from effects long-term negative the buffered size fragment increasing where management, and size fragment between atrade-off was There 4. Article . Preserve what is left – but buffer for change. Small forest Small forest change. –but for buffer left is what . Preserve Fomitopsis pinicola Fomitopsis d forestry 50–100%clear-cuttingd forest with of

increased. increased. (Butchart (Butchart target this fulfil to measuresareneeded conservation area-based additional 2010), (CBD services’ ecosystem and biodiversity for importance particular areasof terrestrial of ‘17% protect biodiversity (Rodrigues been has as protected areas aside land Setting Introduction areas; set asides scenarios; protected size; projection fragment effect; Key-words: conservation. for concern special of species protecting in set-asides of performance improve and area effective This article is This protectedAll rights byreserved. copyright. forest ecosystems (Harper (Harper ecosystems forest in tropical ortemperate as boreal ecosystems, influence in onvegetation persistent or extensive (Haddad Jonsson 2008; Ruete, Snäll& Jönsson 2016),these andeffects magnify the passage with time of forest remnants (Laurance & Yensen 1991;Magura, Tóthmérész& Molnár 2001; Berglund & are of 2000).most Degrading effects fragmentation typically influential in andsmall isolated growth forests currently represent<2% of European forests,excluding (ParviainenRussia old- For example, Fonseca 2000). & Williamson (Gascon, area current their of independent focused onpreserving vestiges last the ofsmall isolated and old-growth fragments forest Accepted impli biodiversity about raise concerns edges and the response of severalorganism groups (Harper Article et al. et et al. et boreal forests; common and indicator species; core:edge ratio; deadwood; edge edge deadwood; ratio; core:edge species; indicator and common forests; boreal 2015). A recent review suggests that anthropogenic edges do not have not asstrong, anthropogenic do that edges suggests review 2015). Arecent 2015). In regions with a long forest management history, the effort is often often is effort the history, management forest along with regions In 2015). et al. et al. 2004). Inproposedthe light of Aichibiodiversity target 11to 2015). Nevertheless, notable differences between forest structure structure forest between differences notable Nevertheless, 2015). cations of extensive creation of anthropogenic anthropogenic of creation extensive of cations the foundation of global efforts toconserveglobal efforts foundation of the et al.

2015)natural toand anthropogenic et al.

al. fore can sustain edges by affected areheavily that set-asides other and areas protected small if unclear is it example, For edges. This article is This protectedAll rights byreserved. copyright. 1990; Magura atthe found conditions specific to adapted et al. Ruete 2004; Kauserud & (Stokland expected be can edges clear-cut younger to proximity close abundance of old-growth forests speciesdue tochanges insubstrate andexposure quality in thandetrimental of the habitateffects loss an 2006; Laurance resources and microclimatic conditions (Ries &Sisk 2004; Ries responses) negative neutral and positive, (e.g., interior conditions (Aune, Jonsson & Moen 2005).Species showvarying responses toedges in determining role critical play a edge-effects where th a landscape in Therefore, 1991). Yensen & (Laurance conditions forest-interior effective of core-area the determine shape and size embedded are typically areas conservation Since time? over species sustain managed to effectively be they can how If so, 11? target in measures’ fragments (typically1-3about ha) should be included‘effective as area-based conservation is likelyis toincrease (Humphreys&Kitchener 1982;Magura si fragment decreasing with declines ratio core:edge & Whittaker 2014). Therefore, the “preserve what is left” policy leaves open the question if if question the open leaves left” policy is what “preserve the Therefore, Whittaker 2014). &

Accepted Article 2011). 2016). At the same time, edge-effects can result in an influx of generalist species ofgeneralist species or in an influx can result Atthesame time, edge-effects 2016).

Half-way towards theHalf-way towards 2020 a Aichi Targets, et al. et et al. 2001; Laurance 2011).For many sessile forest species, edge-influencemay even be more st-dwelling species over time(Gascon st-dwelling et al. 2011; Ruete 2011; interface between two environments (Duelli twoenvironmentsbetween (Duelli interface d isolation(Moen &2003). Jonsson Declines in dependinggradients on edge-to-interior of what proportion of the fragment retains forest- retains fragment ofthe proportion what in a matrix of productive forest, the fragment fragment the forest, productive of amatrix in e normsmall is remnantsof old-growth forest, ze, the ratio of generalist to specialist species generalist ofze, theratio key question is whether very small forest et al.

2016). Hence, as the fragment fragment as the Hence, 2016). et al. et et al. 2001; Matthews, 2001; Cottee-Jones 2004; Ewers & Didham et al. 2000; Timonen et al.

et are known to respond differently to old-growth forest amount and isolation (Nordén (Nordén isolation and amount forest old-growth to differently respond to are known ArtDatab (e.g.loss andhabitat isolation forest ecosystemsworldwide, with a substantialbut fractionspecies of red-listeddue toold-growth fungiDeadwood-dwelling are ubiquitous and functionally important organisms forest within would not jeopardizefuture biodiversity values. help identify of levels anthropogeniccreation edge aroundgrowth small old forest fragmentsthat characteristicsedge-effects of under differentmanagement scenarios.simulationsSuch could 2015).However, thereareno simulation studies investigating the future, possibly transient, importantincreasingly tools for conservationeffective identifying (Mouquet strategies are Forecast dynamics ecological of fragment on studies size. be time dependent transient and in mitigateto edge-effectsneedsto be based a onset varied of edgeconditions, whichshould also (Reese times rotation andforest sizes current stand subjected to imminent clear-cut (>years 90 and mature a years) or 20 (< clear-cut a recent to adjacent is fragment forest old-growth random any of perimeter the of 25% - 20 that is estimated it forest boreal Swedish In the case. the not often is this However, forests). caseof the in clear-cuts (i.e. edges hard by surrounded completely that are they assuming 1991), & Yensen (Laurance recommended strongly has been fragments around practice standard a as zones buffer of implementation the theory, edge-effect of protected areas succeed or fail to deliver conservation outcomes (Geldmann highlighted that remainsevidence limited concerning the management regimes under which species specialist (Aune willprevent small forest edge-effects areas fro This article is This protectedAll rights byreserved. copyright. Accepted Article et al. 2005; Rybicki &Hanski 2013).A recent systematicreview et al. anken 2015). Generalist and specialist wood fungi fungi wood specialist and Generalist 2015). anken m functioning as refuges for forest-interior for asrefuges m functioning 2003). Hence, management strategies aiming aiming strategies management Hence, 2003). ≤ 110 years),110given typical forest stand

et al. 2013). In light et al. et al. 2013),

whilst whilst increasing theoccupancy of common fungi (Ruete fungi indicator of occupancy the years)reduces 40 of timea window (within least temporally at that gradients edge-to-interior transient create edges anthropogenic that know do we study recent Komonen 2011;2012; CrockattAbrego & Salcedo Ruete 2014; is known but little their about responseedge-effects to &(Snäll Jonsson 2001;Junninen & This article is This protectedAll rights byreserved. copyright. common species, using simulations fitted to empirical data (Ruete data(Ruete to empirical fitted common using simulations species, years)(40 future occupancy dynamicsgroupa of logs by of old-growthindicator species and two deadwood-dwellingfungi under currentmanagement forest We policies. project the long-term sustain can conservation for sizesetaside in ha 20 to 1 fragments forest if determine to aim We perspective. the numberand occupancyfungi, logs of byat time scales relevant foresta from management surr edges anthropogenic of impact dynamic the for spatio-temporal roleedges of on deadwood-dwelling fungi allowed ustoparameterizemodels Acceptedprojectionsoccupancy offungal explicit Spatially species. focal the of levels occupancy future the may influence creation edge and size between intertwi as two creation edge and size fragment deadwood availability and occupancy of logs by deadwood-dwelling fungi may be influenced by clear- to is adjacent that perimeter fragment’s the of percentages four under fragment forest growth old of sizes four combining scenarios ( spruce by Norway dominated stands forest 31 old-growth of inventory data onfungal occurrence anddeadwood dynamics were obtained throughtransect-baseda re- Article cuts. cuts. allowedThis us to ned factors. We hypothesize that trade-offs trade-offs that We hypothesize factors. ned of logs allowed us to better understand the oundingforest fragmentsof different sizes on

et al. et 2016). These new insightsthe on et al. et al. quantify thefuture how 2016). However, through a 2016).The empirical Picea abies ). We ). ran 16 dwelling species (Timonen (Timonen dwelling species andconservationpotentialmitig values tohostand ma forest from aside and set identified been About 87 000 small (median size 1.4 ha) and isolated “woodland key habitats” (WKHs) have landscape. the of percent few a only representing and forests managed of matrix a in isolated small, few and occurring are typically remnants forest Old-growth amounts of low deadwood. modernforestry, and characterizedbyeven-aged monocultures conifers, of few oldtrees, and by influenced strongly is region the in landscape forest The Sweden. central in 1968) Jalas & Hämet-Ahti Ahti, (sensu regions boreal south and middle the in forests covers realm study The The study realm methods and Materials landscape. forest boreal fragmented and a managed measures within conservation small of effectiveness area-based long-term This article is This protectedAll rights byreserved. copyright. Accepted species Two edges. to relation in logs of dynamics the predict applied models the and logs forest edges and natural mires,(e.g.,edges The specieslakes). grow chiefly on Norway spruce clear-cut from distance to relation in fragments forest old-growth in 1) (Table species fungal Ruete in models The Models tosimulate logsfungal and occupancy dynamics modelthe below). ( spruce Norway by dominated fragments forest probability ofdeadwood-dwelling by logs fungiderived fromre-inventories ofboreal old-growth Article et al. et (2016) predict the occupancy probability of 12 deadwood-dwelling et al. 2011).We applied recently published models of occupancy nagement in Sweden because of their high Picea abies Picea ate the decline of nationally red-listed forest- red-listed ofnationally decline the ate

) (Ruete ) (Ruete et al. et 2016; further details of ( This article is This protectedAll rights byreserved. copyright. Sweden (see Table S1 in the Supporting Informatio Supporting the S1 in Table (see Sweden species in terms of deadwood substrate resource and forest type (Nordén (Nordén type forest and resource substrate deadwood of terms in species (Ottosson dynamics deadwood spruce on influence a large have and landscape forest managed the in occur in closed forests (Nordén forests (Nordén in closed occur (s tree species trees,a particular dead diameter 2005). They are generally highly fragmentation-sensitive and on specialized natural logs, large- deadwo of continuity aslong-term such values temporal biasfruiting in (e.g. Abrego high mycelia abundancetypically also have a high fruiting rate (Ovaskainen Molecula fruiting patterns. andtemporal presence fruit long-lived and large relatively with species species, as indicator to the species refer individual frequencies weregenerally tooto low deadwood-dwelling fungi (Halme theresingle no is deadwood-dwelling fungal species that alonewouldgood a be indicatorfor all accounting for the spatio-temporal effect of different edge types and edge ages (Ruete (Ruete ages edge and types edge different of effect spatio-temporal the for accounting models linear generalized hierarchical weused Bayesian and time, space over species occupancy and availability substrate forest old-growth on edges anthropogenic of impact the simulate To m 2016). The original models were parameterized using data from 1381 transect segments of 140 Fomitopsis pinicola 2

Accepted in central fragments forest old-growth boreal 31 from units) as sample to referred (hereafter Article et al. 2014).The remaining 10are indicator speciesfor highnature conservation and Trichaptum abietinum et al. et al. et 2013). We studied indicator species as a group because because group aas species indicator Westudied 2013). et al. 2009; Junninen & Komonen 2011) and, because their F. pinicola 2016). pruce or pine) and stage of decay, and mainly mainly and decay, of stage and pine) or pruce od highand 2000;naturalness (Nitare Niemelä ) are non-fragmentation-sensitive and generalist model them individually. In theanalysis, we bodies we minimized sampling bias in mycelia r studies have shown such that species with , and n). A first set of models estimated the effect effect the estimated models of set Afirst n). T. abietinum

. Focusing. our inventory on et al. et et al. 2013), common 2013)and low et al.

seemed to be stable. See Appendix Ruete S1 and Appendix See seemed to be stable. i.e.the fungal over populations a was decade, detected clearchange although no simulations occupanc log mean fragment-level inover time change the and variability among-fragment The species. common the for time-independent but abietinum ( affected positively were species common while edges, clear-cut young to proximity by affected negatively were species Indicator species. common and indicator Binomialdistribution. These modelscaptured contrasting responsesclear-cut to edges by basedon binary presence-absence dataand the number logs per of sampleunit, assuming a was occupancy) log (i.e. unit sample particular a in a log on occurring species common and indicator an of probability The sample unit. per logs of number the in variation for accounting cut vsnatural),andageedge onthe occupancylogs ofindicator by and common species, clear- (i.e. type edge edge, to distance of effect the estimated models of set Another class. decay affected by distance theto edge but by the initia wasnot a decade after logs of number the in change rate of the However, edges. clear-cut young close of fallen to trees thenumber in increment show models a transient These decades. between of edgeson the numberlogsof persample and the change unit innumber logs of per sample unit This article is This protectedAll rights byreserved. copyright. sizes1, 3.14, and 10 ha 20 (i.e. 56,100, and 178 min radius, 252 respectively).Selected stand of 1) (Fig. stands forest circular to approximation closest the covering units) sample surveyed Accepted simulationsSpatially wereexplicit run on m 140 Fragment sizes and management scenarios simulated model. Article ). Anthropogenicedge-effects were transient forindicator species (reduced overtime), y were retained in l number of logs, as well as their volume andvolume their as aswell logs, number of l 2 et al et hexagonal grid cells (i.e. the same area as the the same area (i.e. hexagonalgrid cells . (2016). for detailed descriptionsthe of

F. pinicola F. the models used for ) or unaffected ( ) orunaffected T. T. • • FourStåhl 2002). realistic scenarios of adjacent forest managementwere used: & Jonsson (Kruys, Sweden central in time decomposition log average years the is also Forty 0, 10, 20, 30 and 40 years), allowing sufficient time for forest re-growth in adjacentclear-cuts. killed (e.g. Jönsson are trees spruce Norway old-growth of majority the that clear-cutting surrounding of exposure initial the during is it that knowledge the on based assumptions reasonable are These field. the andappliescreation edge the after predicted models were used. The simulated numberper of logs excludes cell further loginputs than those distance-to-edge fitted the where 1) (Fig. m 100 of DEI maximum the within were centroids cell grid the unless interior, forest the for estimated dynamics the followed also dynamics distributions estimated thefor in forest Ruete interior quality ofoccupancy logs andlogs by fungi assumed in themodel fitting process (Ruete includingstands radius with equal themaximum to depthedgeof influence (DEIm) 100 = sizes represent potential old-growtha range of fore This article is This protectedAll rights byreserved. copyright.

data published by the Swedish National Forest Inventory (Reese from We inferred region. study the in sizes stand forest and times rotation forest current given estimatededge proportion of any randomold-growth forest fragment adjacentaclear-cut, to at clear-cut is forest 25% ofthesurrounding Current: interior. simulated asforest are stand forest the in cells all i.e. is clear-cut; forest surrounding the of 0% Reference: Acceptedforest stands inthe studyregion haverecently been clear-cut (< 20years) may or imminently Article et al. 2007). Simulations were run for five time steps at decadal intervals (i.e. intervals atdecadal time steps for were run five Simulations 2007). within grid cells followed the probability followedtheprobability gridcells within et al. rate of deadwooddecomposition observed inthe 2016). Initial conditions for the number andthe number for conditions Initial 2016). st stands foundthe in et al t = 0. This scenario represents = the This represents 0. scenario

. (2016). Subsequent inter-decadal et al. 2003) that 25%of all landscape, specifically specifically landscape, This article is This protectedAll rights byreserved. copyright. There were clear opposite spatial edge-to-in opposite clear wereThere cutting (Fig.Log2). numbers were highestfragments insmall and decreasedover time(Fig. 3a). on cells closerto cl occurred inputs deadwood Extreme S2). Table (see aclear-cut to was adjacent edge fragment’s the of 25% when edge the (Ruete distributions posterior the fullparameter from parameters of set particular forest stand sizes and forestry scenarios we ran 10 000 simulation replicates each using a Simulations3.1 were(RR Development run in CoreTeam 2014).For each combination of • • Infragments smallerthan 3.14 ha ( size andEdge forestfragment effects Results grid Fig. S1 hexagonal Information). cell; Supporting (i.e. in way,this we obtainedprobability a distribution forresponse the variable each for sampleunit

be clear-cut (>90 and and (>90 clear-cut be Accepted Article locations. particular in happen that can case extreme Extreme:100% of the surroundingforestis clear-cut at section of the stands’ perimeter. biomass (Strengbom in thefuture mo towards trends current given times where rotation situation hypothetical Intensive: 50%the ofsurrounding clear-cut forest is at perimeter. stands’ the ≤ 110 years). The edge was simulated as a single continuous section of of section continuous asingle as simulated was edge years). The 110 ≤ re intensive forestry and increased extraction offorest extraction and intensiveincreasedforestry re 100 mradius) more than60% the of area was influenced by et al. 2011). The edgewassingleas simulated a continuous teriorgradients between the occupancy log ear-cut edges within the first decade after clear- after decade the first edges within ear-cut are significantly shortened,possible which is

t t = 0. This scenario represents an represents scenario This 0. = = 0. This scenario represents an an represents scenario This 0. = et al. 2016).In was lower in larger fragments, and decreased marg decreased and fragments, was lowerin larger similarsmaller to fragmentsover time (Fig. 3c).mean The occupancy of logs by 4b). (Fig. fragments forest smaller in decreasing continue they fragments, forest largest the for future the in arestable levels While occupancy edges. anthropogenic to exposure 4).The exceptions were indicator species, for which occupancy of decrease logs will dueto scenarioafteryearsexposure 40 of both in large smalland fragments (notevarying Fig. scales on reference the to converge to tend 4c-d) (Fig. species by common logs of occupancy and 4a) (Fig. logs of number future The clear-cut. adjacent the on regrowth forest implicit the and effect edge- the of character transient the by driven were time) over scenario, reference the to compared differences showing 4 (Fig. scenario reference this from Differences 3). (Fig. conditions interior forest assuming model original the with estimated occupancy mean stand in change decadal referencescenario clear-cut (0% scenarioandforest only interior conditions)shows the projected logs by of occupancy mean The 3b). (Fig. future the in stable remain or marginally increase to projected was and fragments, large in was higher species indicator by logs of occupancy mean The 2). (Fig. edge) the m from 100 beyond forest (i.e. interior forest the and edge) the mfrom 100 within forest (i.e. habitat edge the between levels occupancy mean in difference a marginal was there but edge, the from distance opposite pattern was projected for for wasprojected pattern opposite the whilst edge, the from distance decreasing by affected negatively was species indicator probabilities by indicator speciesand by This article is This protectedAll rights byreserved. copyright. Accepted Article F. pinicola was initially lower in larger fragments, fragments, inlarger lower initially was F. pinicola F. pinicola . The species . The inally over time (Fig.3d) (Fig. 2). The occupancy probability of of probability occupancy 2). The (Fig.

but was projected to increase to levels toincrease projected but was T. abietinum was unaffected by by unaffected was . These in trends the T. abietinum

reference and current scenarios, but increased to increased but scenarios, current and reference stable over time(Fig. 3b).The meanoccupancy oflogs by proportionslower adjacent of clear-cutting, and was projected to marginally increase remain or Themean occupancy of logsindicator by species was highermanagement in scenarios with cutting (Fig.3a), butwas projected converge to the reference to scenarioafteryears 40 (Fig. 4a). clear- adjacent of proportion greater a to exposed fragments in higher was numbers log Mean Edge effectsand forest management scenarios This article is This protectedAll rights byreserved. copyright. Acceptedsize fragment the of independent and transient was clear-cutting adjacent after species common ha forest fragment (Fig.5b). Theincrease number in oflogsand inoccupancy logs of by 1 smallest the in lower were regimes management between differences the although clear-cut), reference scenario under the intensive and extreme management scenarios (i.e. 50 and 100% scenario,twoyears;10at timeyears). < (t 40 points We = t found largestdepartures from the the reference compared scenarios, to management and all fragment sizes between effects o stable future sustain to important were forest the adjacent in clear-cutting of the extent and size fragment both species, forindicator Especially Trade-offsbetween fragment forest sizemanagement and scenarios 4b). (Fig. scenario management of logs indicator by was species projected toremain lowerthan the referencelevels underany of exposure undermanagement all scenarios (Fig. 4c-d).contrast,In the future mean occupancy years 40 after scenario reference the to converge to tend species Article common by logs of occupancy ofthe Projections 3d). (Fig. marginally overtime clear-cutting, anddecreased adjacent less over time(Fig. 3c). The mean occupancy of logsby ccupancy Figureccupancy levels. the 5 illustrates trade-off levels similar to intens to levels similar T. abietinum

F. pinicola was lower in scenarios with in scenarioswith waslower was initially lower under lower wasinitially ive and extreme scenarios extreme scenarios and ive pinicola growth specialistdeadwood-dwelling fungiwhilst some commongeneralist specieslike anthropogenicaround edgesold-growth forest fragments will lead todeclining occupancyof old- in future that Weprojected forest. productive woodland keyhabitats, dependon bothfragment size andmanagement the of the surrounding as such strategies, conservation area-based of effectiveness the that showed results Our fungi. deadwood-dwelling by of logs forestdifferent fragment sizes and clear-cutting of effects edge dynamic the investigates that projection first the is study This losses. species when developing and implementing cost-effective management actionsto mitigate further responsesUnderstanding thefuture of speciesto Discussion scenarios. management forest the different of impact thefuture in regulating a variable major was size fragment forest the Hence, 5b-d). (Fig. clear-cuts to exposed perimeter entire its with ha) (20 fragments radius radius fragments (3.14ha)with a quarter of perimeter its exposedto clear-cuts asm- in252 groups speciesAn interestingthat all result was the twosmallest fragments withmore than 50% for future the in declining to stable almost from range to seems species indicator of occupancy the for observed effects edge negative the However, c-d). 5a, (Fig. regime management or This article is This protectedAll rights byreserved. copyright. Accepted Article may increase. Two important results were de were important results Two may increase.

tensified forestry with extensive creation of creation extensive with tensified forestry had similar future occupancy levels in 100 m- in occupancylevels future had similar management scenarios on of its perimeter facing clear-cuts (Fig. 5b). 5b). (Fig. clear-cuts facing perimeter its of fragmentation and edge creation isessential rived. First, as expected, larger forest forest larger asexpected, First, rived.

the future occupancy F. F. ha) with a substantial interior core area, inte area, interiorcore asubstantial ha) with 20 (up to fragments larger in even Second, in thefuture. toincrease but wasprojected fragments, ofthe species common levels Occupancy scenarios. management intensive and current under at least conditions), forest (interior scenario reference simulate and manage nature the best way we can given current knowledge (Urban (Urban knowledge current given can we way best the nature manage and simulate responses underclimate change its infanc isin fungal of forecast the whilst Therefore, islacking. knowledge distribution geographical northern some fungalgroups (e.g. Kauserud in for changes evidence is There in thefu influential middle-aged to mature productive forest. the cut, areforest while most likelyinterior, s clear- arenot that forests We adjacent that assume forest. productive older suboptimal adjacent S2; Laurance& Yensen 1991; Aune reduceratio, significantly core:edge the reducing th certainly will index shape Increased index). shape possible lowest the having a circle (i.e. shape stand optimal the only handle becausethey optimistic are simulations These cases. management forest particular results to our applying ofwhen tobeaware principles some precautionary are There species. indicator forest old-growth of persistence long-term the compromise would fragments in levels occupancy Relative the future. in todecline to projected continue were scenario, reference the to relative species, indicator of levels occupancy fragments ha 1 In smallest the future. the in species indicator of levels occupancy on impacts negative to resistant more were fragments This article is This protectedAll rights byreserved. copyright. Accepted Article ≥ 3.14 ha, however,were projected toremain ture, possibly also influencing future fungal responses at the forest edge. edge. forest the at responses fungal future influencing also possibly ture, fungal fruiting patterns over time with changing climate for for changing climate with over time patterns fruiting fungal et al. et al. for 2010), but deadwood

Climate change effects are likely to become more become to effects are likely change Climate 2005). We may also underestimate maythe We of impact alsounderestimate 2005). nsive clear-cutting (50-100%) around (50-100%)nsive clear-cutting the fragment ituation is that these forests are dominated by dominated are forests these isthat ituation y, we cannot wait for it to e area with forestinte with e area F. pinicola

stable the in future was instead lower in larger lower in wasinstead -dwelling species-dwelling with a develop in order to to order in develop rior conditions (Figure at levels below the at levelsbelow et al. 2016). Penttilä Penttilä 50 or moreyears (Gu, Heikkilä & Hanski 2002; Sverdrup-Thygeson & Lindenmayer 2003; (generally <10 ha) forest fragments (Berglund & Jonsson 2005, 2008), but more evident after c. small in decades few a in already place take to shown been have species indicator growth fungal offuture with oursimulations Inagreement This article is This protectedAll rights byreserved. copyright. Accepted be to are anetwork such within areas conservation if considered be to needs size fragment and way toensure viablepopulations. Our studyhigh best the be seemsto areas conservation forest of network functional a species these for and fungi, specialist old-growth many conserve to needed naturalness and volumes deadwood the reach stands tolevelsnatural-like in stands. Theyalso concluded that managed forests are unlikely to deadwood increased stocks and required qualities were to elevate fungal diversity of spruce natural Norway spruce forests(Peltoniemi, Penttilä Mäkipää & 2013).Theyprojected that fungi toproject the future polypore diversity based ondeadwood dynamicsmanagedin and management simulation study incorporated snap-shot occupancy data of deadwood-dwelling forest recent One research. further needs environments edge forest to responses fungal regulating in traits ecological different of role overall the However, 2014). Melo & (Ryvarden edge at the conditions microclimatic drying and by be thus may the and more affected sapwood in of mycelia Articlemore andlarger species theformer of result pinicola earlier. commonBetween the generalist thetwo species, potential of stronger buffering like species against clear-cut edges and microclimatic changes compared to to compared changes microclimatic edges and clear-cut against et al. et F. pinicola F. pinicola 2006; Paltto occur in the heartwood, whereas the mycelia of of mycelia the whereas heartwood, the in occur inedgeenvironments of small conservationareas has notbeen reported et al. 2006; Ranius long-lived fruit bodies and mycelia. Also, the et al. lights that transient anthropogenic edge effects effects edge anthropogenic transient that lights occupancy, declines in the occurrence ofold- occurrence in the declines occupancy, 2008). Increasing occupancy of common

T. abietinum T. abietinum concentrate concentrate could be a F. (Nordén (Nordén due tocommonly alteredforest landscapes dominatedman-made by deadwood substrate types Shriver,& Cutler Doak2012). The future increasedoccupancy common offungi likely is also environmental anddemographic stochasticity (Boyce that indicator areas incore suggests logs even Siitonen& Jonsson Nordén 2012; Stokland, 2011; Komonen & Junninen (e.g. fragmentation habitat and oflogging, effects characteristics, stand forest in changes to vulnerability greater specialists’ the by explained likely are fungi common and specialists interior forest between development future contrasting The altered microclimate conditions. Snäll,Ribeiro & Roberge Rydin 2003; 1988; (Söderström environment edge the in dynamics substrate altered the to linked are partly declines future These creation. clear-cut adjacent the of extent the on dependent be will declines dec experience to likely more also are fungi thisknowledge by showing thatsmaller fragments withlower occupancy levels of indicator Jonsson 2001; Siitonen, Lehtinen AbregoSiitonen& 2005; Salcedo& 2014). Our extends study & (Snäll fragments small in occupancies lower explaining variables the of one is effect edge that (but see Roberge 2004; Didham & Ewers 2012), especially for lesser studied organisms like sessile cryptogams Sisk& seeRies (but few relatively are future the in biodiversity local influence effects edge functional for the future conservation of fungi. Simulation studies that examine how area and This article is This protectedAll rights byreserved. copyright. Accepted Article et al. 2013; Komonen et al. et 2011; Fedrowitz,2011; Kuusinen & Snäll 2012).For wood fungi,knowdo we et al. et al. et 2014). These substrate types favor common species like species like common favor types substrate These 2014). et al. et 2013). A high temporal variability in the occupancy of occupancy in the variability temporal A high 2013). lining populations in the 2011; Ruete species are particularly vulnerable to future to future vulnerable are particularly species et al.

et al. 2006;Wiklund Ruete, Snäll&2012; 2016), butmayalso be linkedto future, and that such such that and future, F. at the landscape scale (e.g. Peltoniemi Peltoniemi (e.g. scaleat the landscape to wayapproachesmight be the best of mixture ecosystemsunderstandto and balance theand pros cons of differentmanagement scenarios.A optionsis highly relevantfor forest management policies, enablingmanagers forest of management and trade-offs such Identifying clear-cut. was 25% only where fragment forest ha obtainedainha 20 fragment forest where 100%ofthe surrounding forest was clear-cuta 3 in as was species by of indicator logs levels the occupancy Forsame future example, case. particular mayeffect) disfavor interior specialist species (Nordén pinicola This article is This protectedAll rights byreserved. copyright. Acceptedrequiredadjacent habitats towith specific biodiversityvalues”(Forest Stewardship Council zones any (b) buffer and values biodiversity specific with habitats (a)small patches demanding care- as demarcate shall “Managers states that certification forest for FSC standard Swedish The zones. buffer management optionsand calls forstronger national scale policies promoting larger set-asides and 2013), thegeneral situation is to Agency Forestry less (Swedish or ha 1.4 of fragments aresmall Sweden in remnants forest Articlema forachieving positive room more the fragment, thevariable regulating future impact of adja a was major size Forest fragment and actions. policies management forforest implications has clear edge-creation of anthropogenic size extent and fragment the between trade-off The changes for buffer but left, is what Preserve , whoseincreased establishment, fruiting,and sporedeposition (i.e. a so-called bulk wards the worst-case scenario for indicator species. This limits limits This species. indicator for scenario worst-case the wards et al. 2013). However, given that 50% of the old-growth cent forest management. The larger the forest improve theviability species of forest-dwelling nagement results and for trade-offs in each ineach trade-offs for and nagementresults et al.

2013). Administrative Board, respectively, for for their he respectively, Board, Administrative Wegrateful are Kellner to Olle and UrbanGunnarsson Gävleborg at and Dalarna County Acknowledgements conservation. for concern ofspecial species protecting in their performance improve and fragments forest small old-growth of area the core may increase time dynamic buffers) over (i.e. effect edge zones, non-simultaneousclear-cuts of surrounding productiveforest ina waythatreduces the buffer permanent to alternative Asan habitat. forest once-edge restored the recolonizing species Berglundand Jonsson Longer (2008). rotation times mayincrease chances the of indicator forest set-asides as previously suggested by Dettki and Esseen (2003), Siitonen et al. (2005), and longer rotati of importance highlight the results manyspecialized deadwood-dwellingfungi(Snäll & JonssonRuete 2001; Nonetheless, bufferzones between 25-100are mlikely relevantfor reducing edge effects for highly context dependentin relation toaspect,type, forest and the taxonomicgroup considered. provide any specific recommendations for buffer zone widths, likely because buffer zonesare FSC not does Swedish The effects. edge temporal degrading from fragments such in species protect to managed be to need forests surrounding 11, target Aichi the in measures area effective torelation the size. Ifeffective fragment ol small in change future for anticipated to buffer important equally but size, oftheir regardless landscape managed forest the in left that are qualityhigh fragments the topreserve It important habitats. is these around uncut left are forest productive of zones buffer that advocate we areas, conservation small of size, effective or capacity, conservation long-term the maintain to In order 2010). This article is This protectedAll rights byreserved. copyright. Accepted Article on times and tosmall areas adjacent protected lp in providing monitoring data, selecting the data,in providing selecting monitoringlp d-growthforest fragmentsareto be included as

et al. et 2016). Our 2016). Boyce, M.S., Haridas, C.V., Lee, C.T. & the NCEAS Stochastic Demography Working Group. (2006) (2006) Group. Working Demography Stochastic NCEAS the & C.T. Lee, C.V., Haridas, M.S., Boyce, Berglund, H. & Jonsson, B.G. (2008) Assessing the extinction vulnerability of wood-inhabiting fungal fungal wood-inhabiting of theextinction Assessing vulnerability (2008) & B.G. H. Jonsson, Berglund, and northern amongdebt in lichens fungi an extinction Verifying (2005) & B.G. H. Jonsson, Berglund, Aune, K., Jonsson, B.G. Jonsson, K., Aune, & woodland effectsedge and (2005) amongMoen, Isolation J. habitats key in ArtDatabanken. (2015) Ahti, T., Hämet-Ahti, L. & Jalas, J. (1968) Vegetation zones and their sections in northwesternEurope. sectionsVegetation their and in zones (1968) J. Jalas, & L. Hämet-Ahti, T., Ahti, Abrego, N. Salcedo,& I. (2014) Response of wood-inhabitingfungal community tofragmentation in a Abrego, N., Halme, P., Purhonen, J. & Ovaskainen, O. (2016) Fruit bodybased inventories in wood- References https://dx.doi.org/10.6084/m9.figshare.4129821.v1 (Ruete 2016). Figshare in available are available are files shape and simulations for script R accessibility Data FacultyNatural of ResourcesSLU. and Sciences, Agricultural the and (Skogssällskapet) Society Forest Swedish the by funded was study The manuscript. HåkanBerglund, Jenni Nordén andPanu Halme whosecomments helped improve us the studyand sites contactingWe landowners.two anonymous thank reviewers, Anders Dahlberg, This article is This protectedAll rights byreserved. copyright. Accepted Article Demographyanincreasingly in variableworld. 3039. species in fragmentednorthern Swedishboreal forests. Swedish boreal forests. Sweden: Is forestpolicy promoting fragmentation? ArtDatabanken, SLU, Uppsala, Sweden. Annales Botanici Fennici beech forest landscape. landscape. forest beech inhabiting fungi: Should we replicate spacein or time? Rödlistade Arter I Sverige 2015 = The 2015 Red List of Swedish Species Swedish of Red List 2015 The 2015 = Sverige I Arter Rödlistade Conservation Biology , Fungal Ecology 5 , 169–211. , 8 , 18–27. , 18–27. , 19 Trends inEcology Evolution& , 338–348. Biological Conservation Biological

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Forest Ecology andManagement Science . Cambridge University Press, , 353 , 144 , aad8466. , 2061–2067. , 174 , Common*: species. Else, indicator speciesfollowing Nitare (2000) Trichaptum laricinum Trichaptum abietinum Phellinus viticola nigrolimitatus Phellinus Phellinus ferrugineofuscus Phellinus chrysoloma mollis Leptoporus Fomitopsis rosea span body life Fruiting pinicola Fomitopsis murraii Cystostereum Rottype borealis Climacocystis Status Red-list Asterodon ferruginosus name Scientific (2015). Artdatabanken according to status Table 1 Tables This article is This protectedAll rights byreserved. copyright. Accepted Article . Key ecological characteristics of the stud the of characteristics ecological Key . NT Brown Perennial Brown NT LC White Perennial Perennial White LC N Bon Annual Brown NT C rw Perennial Brown * LC NT White Perennial Perennial White NT NT White Perennial Perennial White NT NT White Perennial Perennial White NT C ht Perennial White * LC LC White Annual Annual White LC NT White Annual Annual White NT NT White Perennial Perennial White NT NT White Perennial Perennial White NT iedindicator and common species. Red-list

square). The color scale of each cell (140 m (140 cell each of scale color The square). (pink aclear-cut to adjacent edge fragment forest the of is25% that conditions; management 2 Figure hexagons. green for applied were models and occupancyofindicator b)by logs species (note negative scale),c) 4 Figure c) species, indicator b) by logs of probability 3 Figure 140 m C: FI:interior; forest r: radius; fragments. forest 1 Figure This article is This protectedAll rights byreserved. copyright. d) species, species, posterior distribution of the numberofLogs)(No.logs occupancy or (Occ.) of logs indicator by fragments (25-100% clear-cutting) overlap. smallest for theforest the lines In 3d, the panels. scale between the Note varying clear-cut. surroundingforest clear-cut, andthe 100%extr is quarter of surrounding the forest clear-cut, 50%isthe intensive scenariowith half the of a with scenario current is the 25% clear-cut, adjacent no with scenario interior) (forest reference the is where 0% scenarios, management indicate Line colors sizes. fragment forest largest in cells of old-growth forest fragments over time. Projections are shown only for the smallest and

AcceptedTrichaptum abietinum Article 2 (out of scale).Edge modelshabitat were applied for pinkhexagons, andforest interior Fomitopsis pinicola Fomitopsis : Starting conditions predicted by models at t < 10 yearsStarting 10 predicted: conditions models t < at by current under forestry : Projections of the differences in future mean values for a) the number of logs per cell cell per logs of number a)the for values mean future in differences the of Projections : occupancy cell and logs per of a) number for the mean values of the future : Projections ex ofthespatiallyrepresentation Schematic : in cells of old-growth forest old-growthof forest incells , and , and Trichaptum abietinum 2 sample units) shows the median shows valueofthe units) sample Fomitopsis pinicola clear-cut. Each hexagonal cell is a study unit of unit is astudy cell Each hexagonal clear-cut. eme scenario with all of the surrounding forest forest surrounding the of all with eme scenario plicit simulationplicit set-up of edge-effects on fragments over time compared to the to compared time over fragments .

and d) Fomitopsis pinicola Trichaptum abietinum and and

d) and occupancyofindicator b)by logs species (note negative scale),c) 5 Figure fragments (25-100% clear-cutting) overlap. smallest for theforest the lines In 4d, the panels. scale between the Note varying clear-cut. surroundingforest clear-cut, andthe 100%extr is the of half with scenario intensive is the 50% clear-cut, forest surrounding the of aquarter with interior) (forest reference the is Line sizes. fragment forest largest and smallest the for only shown are Projections clear-cut). 0% (i.e. scenario reference the for levels projected This article is This protectedAll rights byreserved. copyright. surrounding forest clear-cut. Note the varying scale between the panels. panels. the between scale varying the Note clear-cut. forest surrounding half of the surrounding forest clear-cut,and red theis extreme scenario with all of the with scenario intensive yellowis the clear-cut, forest surrounding the of quarter a with scenario green is thecurrent where scenarios, management x-axis indicate ofcircles the and Colors scenario(0%clear-cut). Circle size representsforest fragmentsize (i.e. 3.14, 1, 10, 20ha). and circles) at

Accepted Trichaptum abietinum Article : Projections of the differences in future mean values for a) the number of logs per cell cell per logs of number a)the for values mean future in differences the of Projections : t = 40 years (empty circles) compared to the projected levels for the reference reference the for levels projected the to compared circles) (empty years 40 = in cells of old-growthof forest incells scenariowith noadjacent clear-cut, the 25% current is scenario colors indicate management scenarios, where 0% 0% where scenarios, management indicate colors eme scenario with all of the surrounding forest forest surrounding the of all with eme scenario fragments fragments at two points time

Fomitopsis pinicola t < 10 (filled < and and S2. Figure S1. Figure S2. Table S1. Table S1. Appendix Information Supporting This article is This protectedAll rights byreserved. copyright. Accepted 1 Figure Article

Affected area given combination of fragment sizes and proportion of edge of proportion and sizes fragment of combination given area Affected Characteristics of the forest stands surveyed Expected Variabilityamong sample units Effectsofand shape index depthedge of influencecorearea on affected Survey of the focal forest fragments forest focal the of Survey

This article is This protectedAll rights byreserved. copyright. Figure 2 Figure AcceptedOcc. T. abietinum Occ. F.Article pinicola Occ. Indicator spp. No. Logs

1 ha 3.14 ha

10 ha 20 ha 20 Log occ. Log occ. Log occ. No. 0 0.11 0 0.35 0 0.05 0 1.7

This article is This protectedAll rights byreserved. copyright. 3 Figure c p(Occupancy) of F. pinicola Number of logs per cell a 0.1 0.2 0.3 0.4 0.5 0.0 Accepted Article0.0 0.5 1.0 1.5 102030400 102030400

20 ha 1 ha Years Years 100% 50% 25% 0% d p(Occupancy) of T. abietinum p(Occupancy) of Indicator species b 0.5 0.0 0.1 0.2 0.3 0.4 0.0 0.1 0.2 0.3 0.4 0.5 102030400 102030400

Years Years

This article is This protectedAll rights byreserved. copyright. Figure 4 4 Figure Δ p(Occupancy) of F. pinicola c Δ Number of logs per cell a

Accepted0.0 0.1 0.2 0.3 Article0.4 0.5 0.0 0.2 0.4 0.6 0.8 1.0 102030400 102030400 20 ha 1 ha Years Years 50% 25% 100%

Δ p(Occupancy) of T. abietinum d Δ p(Occupancy) of Indicator species b -0.10 -0.08 -0.06 -0.04 -0.02 0.00 0.08 0.10 0.00 0.02 0.04 0.06 102030400 102030400

Years Years

This article is This protectedAll rights byreserved. copyright. Figure 5 Figure a AcceptedΔ p(Occupancy) of F. pinicola c ArticleΔ Number of logs per cell 0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.1 0.2 0.3 0.4 0.5 55 100 50 25 55 100 50 25

Scenario Scenario b Δ p(Occupancy) of T. abietinum d Δ p(Occupancy) of Indicator species -0.10 -0.08 -0.06 -0.04 -0.02 0.00 0.02 0.04 0.06 0.08 0.10 0.00 55 100 50 25 100 50 25 Scenario Scenario