Impacts of salvage logging on biodiversity: A meta-analysis

Abstract

Logging to “salvage” economic returns from forests affected by natural disturbances has become increasingly prevalent globally. Despite potential negative effects on biodiversity, salvage logging is often conducted, even in areas otherwise excluded from logging and reserved for nature conservation, inter alia because strategic priorities for post‐disturbance management are widely lacking.
A review of the existing literature revealed that most studies investigating the effects of salvage logging on biodiversity have been conducted less than 5 years following natural disturbances, and focused on non‐saproxylic organisms.
A meta‐analysis across 24 species groups revealed that salvage logging significantly decreases numbers of species of eight taxonomic groups. Richness of dead wood dependent taxa (i.e. saproxylic organisms) decreased more strongly than richness of non‐saproxylic taxa. In contrast, taxonomic groups typically associated with open habitats increased in the number of species after salvage logging.
By analysing 134 original species abundance matrices, we demonstrate that salvage logging significantly alters community composition in 7 of 17 species groups, particularly affecting saproxylic assemblages.
Synthesis and applications . Our results suggest that salvage logging is not consistent with the management objectives of protected areas. Substantial changes, such as the retention of dead wood in naturally disturbed forests, are needed to support biodiversity. Future research should investigate the amount and spatio‐temporal distribution of retained dead wood needed to maintain all components of biodiversity.

Full text

J Appl Ecol. 2018;55:279–289. wileyonlinelibrary.com/journal/jpe 
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279
© 2017 The Authors. Journal of Applied Ecology
© 2017 British Ecological Society
Received:16January2017
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Accepted:18May2017
DOI: 10.1111/1365-2664.12945
REVIEW
Impacts of salvage logging on biodiversity: A meta- analysis
Simon Thorn1 | Claus Bässler2 | Roland Brandl3 | Philip J. Burton4 | Rebecca
Cahall5 | John L. Campbell5 | Jorge Castro6 | Chang-Yong Choi7 | Tyler Cobb8 |
Daniel C. Donato9 | Ewa Durska10 | Joseph B. Fontaine11 | Sylvie Gauthier12 |
Christian Hebert12 | Torsten Hothorn13 | Richard L. Hutto14 | Eun-Jae Lee15 |
Alexandro B. Leverkus16 | David B. Lindenmayer17 | Martin K. Obrist18 |
Josep Rost19,20 | Sebastian Seibold2,21 | Rupert Seidl22 | Dominik Thom22 |
Kaysandra Waldron23 | Beat Wermelinger24 | Maria-Barbara Winter25 |
Michal Zmihorski26 | Jörg Müller1,2
1FieldStationFabrikschleichach,DepartmentofAnimalEcologyandTropicalBiology(ZoologyIII),Julius-Maximilians-UniversityWürzburg,Rauhenebrach,Germany
2BavarianForestNationalPark,Grafenau,Germany
3DepartmentofEcology,AnimalEcology,FacultyofBiology,Philipps-UniversitätMarburg,Marburg,Germany
4UniversityofNorthernBritishColumbia,Terrace,BC,Canada
5DepartmentofForestEcosystemsandSociety,OregonStateUniversity,Corvallis,OR,USA
6DepartmentofEcology,UniversityofGranada,Granada,Spain
7DepartmentofForestSciences,SeoulNationalUniversity,Seoul,Korea
8RoyalAlbertaMuseum,Edmonton,AB,Canada
9SchoolofEnvironmental&ForestSciences,UniversityofWashington,Seattle,WA,USA
10DepartmentofEcologyandBiodiversity,MuseumandInstituteofZoology,PolishAcademyofSciences,Warsaw,Poland
11SchoolofVeterinaryandLifeSciences,MurdochUniversity,Murdoch,WA,Australia
12NaturalResourcesCanada,CanadianForestService,LaurentianForestryCentre,Quebec,QC,Canada
13DivisionofBiostatistics,UniversityofZürich,Zürich,Switzerland
14DivisionofBiologicalSciences,UniversityofMontana,Missoula,MT,USA
15UrbanPlanningResearchGroup,DaejeonSejongResearchInstitute,Daejeon,Korea
16EcologyUnit,DepartmentofLifeSciences,UniversityofAlcalá,AlcaládeHenares,Madrid,Spain
17FennerSchoolofEnvironmentandSociety,TheAustralianNationalUniversity,Canberra,ACT,Australia
18WSLSwissFederalInstituteforForest,SnowandLandscapeResearch,BiodiversityandConservationBiology,Birmensdorf,Switzerland
19DepartmentofEnvironmentalSciencesandFoodIndustries,UniversityofVic-CentralUniversityofCatalonia,Catalonia,Vic.,Spain
20DepartmentofEnvironmentalSciences,UniversityofGirona,Girona,Spain
21DepartmentofEcologyandEcosystemManagement,ChairforTerrestrialEcology,TechnischeUniversitätMünchen,Freising,Germany
22InstituteofSilviculture,DepartmentofForest-andSoilSciences,UniversityofNaturalResourcesandLifeSciencesVienna,Vienna,Austria
23DepartmentofWoodandForestSciences,LavalUniversity,Québec,QC,Canada
24WSLSwissFederalInstituteforForest,SnowandLandscapeResearch,ForestDynamics–ForestEntomology,Birmensdorf,Switzerland
25ForestResearchInstituteofBaden-Württemberg(FVA),Freiburg,Germany
26DepartmentofEcology,SwedishUniversityofAgriculturalSciences,Uppsala,Sweden

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1 | INTRODUCTION
The frequency and extent of stand-replacing natural disturbances,
such as wildfires, windstorms and insect outbreaks, has increased
considerably during recent decades, particularly in the Northern
Hemisphere(Kurz etal.,2008;Seidl, Schelhaas,Rammer,&Verkerk,
2014).Naturaldisturbancescanenhancethestructuralheterogeneity
offorests,createhabitatsforspecies-richassemblagesofhighconser-
vationvalueandincreasethelong-termresilienceofforeststofuture
stressors(Swansonetal.,2011).However,societaldemandfortimber
and/orpestreductioncompelsforestmanagersto“salvage”timberby
logging before it deteriorates,a common practice even in locations
otherwise exempt from conventional green-tree harvesting, such
as national parks orwilderness areas (Figure1) (Chylarecki & Selva,
2016;Thorn etal., 2014).Suchsalvage loggingreducesthe amount
of dead wood, alters successional trajectories, affects biodiversity,
and can influence restoration costs and subsequent fire hazards
(Lindenmayer,Burton, & Franklin, 2008; Waldron,Ruel, & Gauthier,
2013).Consequently,conflictsoftenemergebetweennaturalresource
managers,policy-makersandconservationistsonhowtohandlenatu-
rallydisturbedforests(González&Veblen,2007;Lindenmayer,Thorn,
& Banks, 2017; Lindenmayer etal., 2004; Schmiegelow, Stepnisky,
Stambaugh,& Koivula,2006).This hasresultedinintensepublicde-
bates(Lindenmayeretal.,2017;Nikiforuk,2011;Stokstad,2006).
Differentnaturaldisturbance regimesleavedistincttypesofbio-
logicaland/orstructural legacies(Franklinetal.,2000).Forinstance,
forestskilledbywildfireorinsectoutbreaksarecharacterizedbylarge
numbersofsnags,whilewindstormscreateuprootedtrees(Swanson
etal., 2011). Salvage logging typicallyremoves or alters these lega-
cies.Theresponsesofsaproxylicandnon-saproxylicspeciesgroupsto
salvageloggingthusdependontheirrelationto(deadwood)legacies
affectedbysalvagelogging(Lindenmayeretal.,2008).Consequently,
differenttaxonomicgroupsindifferenttypesofnaturaldisturbances
may respond differently to salvage logging (Zmihorski & Durska,
2011).Numerousstudieshavefocusedontheeffectsofsalvagelog-
gingafternaturaldisturbancesonspeciesrichnessandthecommunity
compositionofvarioustaxasuchasvascularplants(Blair,McBurney,
Blanchard, Banks, & Lindenmayer, 2016; Macdonald, 2007; Stuart,
Correspondence
SimonThorn
Email:simon@thornonline.de
Funding information
GermanEnvironmentalFoundation;Austrian
Science Fund
HandlingEditor:MatthewStruebig
Abstract
1. Loggingto“salvage”economicreturnsfromforestsaffectedbynaturaldisturbances
hasbecomeincreasingly prevalentglobally.Despite potentialnegativeeffects on
biodiversity,salvageloggingisoftenconducted,eveninareasotherwiseexcluded
from logging and reserved for nature conservation, inter alia because
strategicprioritiesforpost-disturbancemanagementarewidelylacking.
2. Areviewoftheexistingliteraturerevealedthatmoststudiesinvestigatingtheef-
fects of salvage logging on biodiversity have been conducted less than 5years
followingnaturaldisturbances,andfocusedonnon-saproxylicorganisms.
3. Ameta-analysisacross24speciesgroupsrevealedthatsalvageloggingsignificantly
decreasesnumbers ofspecies ofeighttaxonomicgroups.Richnessofdeadwood
dependenttaxa(i.e.saproxylicorganisms)decreasedmorestronglythanrichnessof
non-saproxylictaxa.Incontrast, taxonomicgroupstypically associatedwithopen
habitatsincreasedinthenumberofspeciesaftersalvagelogging.
4. Byanalysing134originalspeciesabundancematrices,wedemonstratethatsalvage
logging significantly alters community composition in 7 of 17 species groups,
particularlyaffectingsaproxylicassemblages.
5. Synthesis and applications.Ourresultssuggestthatsalvageloggingisnotconsistent
withthe managementobjectivesofprotectedareas. Substantialchanges,suchas
theretentionofdeadwoodin naturallydisturbedforests, areneededto support
biodiversity.Futureresearchshouldinvestigatetheamountandspatio-temporal
distribution of retained dead wood needed to maintain all components of
biodiversity.
KEYWORDS
barkbeetle,climatechange,deadwood,disturbedforest,fire,naturaldisturbance,post-
disturbancelogging,salvagelogging,saproxylictaxa,windstorm

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Grifantini,Fox,&Fox,1993),carabids(Cobb,Langor,&Spence,2007;
Koivula & Spence, 2006; Phillips, Cobb,Spence, & Brigham, 2006),
birds(Castro,Moreno-Rueda,&Hódar,2010;Choi,Lee,Nam,Lee,&
Lim,2014;Nappi&Drapeau, 2009; Saab, Russell,&Dudley, 2009;
Thorn,Werner,etal.,2016; Zmihorski,2010),and saproxylicorgan-
isms(i.e.thosedependingon dead wood during somepartoftheir
life cycles; Cobb etal., 2011; Norvez, Hébert, Bélanger, Hebert, &
Belanger,2013).
Twomain effects of salvage logging on biodiversityarise recur-
rentlyfrom the existing bodyofliterature. First,salvageloggingre-
duces the richness of taxonomicgroups or abundance of particular
speciesthatdependondeadwood.Forinstance,salvageloggingde-
creasednestingdensity ofcavity-nesting-birdsthat usuallybreedin
fire-killedtrees(Hutto&Gallo,2006).Similarly,post-stormloggingde-
creasedthetotalnumberofsaproxylicbeetlespeciesandthenumber
ofthreatenedspecies(Thornetal.,2014).Second,studiesthatinves-
tigateasetofdifferenttaxonomicgroupshavedemonstratedthatsal-
vageloggingcanalterthecommunitycompositionofbothsaproxylic
andnon-saproxylicorganisms,whiletheeffectsontheoverallnumber
ofspeciescanbesmall(Thorn,Bässler,Bernhardt-Römermann,etal.,
2016).Forinstance, post-stormsalvagelogginginMinnesota greatly
diminishedbirdcommunities,whilefewerdifferencesinthetreecover
weredetected(Lain,Haney,Burris,&Burton,2008).However,previ-
ousattemptstosummarizeknowledgeontheeffectsofsalvagelog-
gingonbiodiversityhavefocusedmainlyonsalvageloggingofburned
forests(Lindenmayer&Noss,2006;Lindenmayeretal.,2008;McIver
&Starr,2000;Thorn,Bässler,Svoboda,&Müller,2016),andaquan-
titativeassessment ofsalvagelogging impactsonbiodiversityisstill
lacking,particularlyacrossdifferenttaxonomicgroupsandinresponse
todifferenttypesofdisturbances(Figure1).
FIGURE1 Salvagelogging(SL)iscommonlyappliedafterwildfires,windstormsorinsectoutbreaks,andleadstochangesinhabitatsand
communitycompositionsinvariousforestecosystemsaroundtheworld(ashighlightedbythestudiesillustratedinpanels(a–l).Studylocations
(colouredcircles)representstudysitesthatcontributeddatatoourmeta-analysis.Photographsbyauthors.[Colourfigurecanbeviewedat
wileyonlinelibrary.com]

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Here,wereviewedthescientificliteratureandcompiledexisting
data to quantify the effectsof salvage logging after wildfire,wind-
storms and insect outbreaks on (1) species numbers via a meta-
analysisof238individualcomparisonsofsalvaged/unsalvagedareas;
and (2) community composition, based on a subset of134 original
speciesabundancematrices.Wealsotestedthehypothesisthat the
impactsofsalvagelogging aremorepronouncedfor saproxylicspe-
ciesgroups thanfor non-saproxylicgroupsregardingthenumber of
speciesandcommunitycompositionwithindifferenttypesofnatural
disturbances.
2 | MATERIALS AND METHODS
2.1 | Literature search
We followed guidelines for systematic literature reviews (Pullin &
Stewart,2006) tocompile comparisonsofspeciesrichnessbetween
salvagedandunsalvagedfire-, wind- or insect-affectedforests.We
screenedtheelectronicdatabasesWebofScience,ScopusandGoogle
Scholaron15 February 2016 byusingthe simplified search strings
[salvageloggingORpost$disturbance*ORsalvaging]and[forest$OR
vegetationORdisturbanceORecosystem].Fromthis bodyoflitera-
ture(>2,000articles),weretainedonlyfield-basedstudiesafterhav-
ingscreenedthetitleandabstract.Modellingstudieswereexcluded.
Wealsoaddedrelevantpapersfromreferencelistsinpublishedstud-
ies. We restricted studies to those providing comparisons between
completely salvage logged plots and completely unsalvaged control
plotsaccordingtotheinformationgivenintherespectivestudies.This
meansthatonsalvageloggedplots,morethan75%ofthetreeswere
affectedbynatural disturbanceandthen completelysalvage logged
withoutfurthertreatment suchastree plantingorlegacy retention.
Lowerintensitiesofnaturaldisturbanceshavebeenrarelytargetedby
scientificstudies.Salvageloggingoperationsthusresembledconven-
tionalclear-cutting.Unsalvaged control plots hadtobeaffected by
thesamenaturaldisturbanceeventbutwithoutanyhumaninterven-
tion.Salvageloggedplotshadtobeofsimilarsize,surveyedwiththe
samefieldmethodsduringthesamestudyperiodandwiththesame
samplingeffortasunsalvagedcontrolplots.
Toexaminewhetherpseudo-replication(i.e.allplotsnestedwithin
onearea)mightbiasthe resultsofour meta-analysis(Ramage etal.,
2013), we carefullyselected the studies according to their designs,
andweusedstatisticsthataccountforpseudo-replication(seebelow).
Thespatialarrangementofplotsinallstudieswascheckedbasedon
methoddescriptionsand/ororiginalgeographiccoordinates.Wecon-
tactedauthorstoprovidedataortoclarifytheirstudydesignswhere
necessary(seeDatasourcessection).Studieswithouttrue replicates
(e.g.allsalvagedplotsnestedandseparatedfromunsalvagedcontrol
plots) were excluded from the analysis to ensure valid effect sizes
(Halme etal., 2010). Studies using the same set offield plots and/
orthesame studyarea(e.g.Samcheok Forest,Korea)wereidentified
andnestedinallsubsequentstatisticalanalysestocontrolforpseudo-
replicationwithinstudyareas.Wealsoexcludedstudiesthatsampled
forests undergoing multiple types of disturbances. Salvage logging
hadtobe conducted immediately(<12months)afternaturaldistur-
bancetookplace.Meannumberofspeciesandstandarddeviationval-
uespersamplingunitwereextractedfrompublishedtextandtables,
or from figures using PLOT DIGITIZER 2.6.2. (www.plotdigitizer.
sourceforge.net). Last, we compiled data on covariates by extract-
ing information on the disturbance type and the time since distur-
bance, and the time since subsequent salvage logging. In addition,
we compiled original species abundance matrices that underpinned
the published papers, which allowed us to explore the effects of
salvageloggingoncommunitycomposition.
2.2 | Meta- analysis
Allanalyseswereconductedinr3.3.1(www.r-project.org).Priortosta-
tisticalanalysis,specieswereassignedtooneofthefollowingtaxonomic
groupsandtoassociationwithdeadwood(i.e.saproxylic/non-saproxylic)
basedonthedescriptioninthearticles.Thesewhere:amphibians,ants,
bats, bees and wasps, birds, carabids, epigeal lichens, epigeal mosses,
epigealspiders,epixyliclichens,epixylicmosses,harvestmen,hoverflies,
landsnails,nocturnalmoths,non-saproxylicbeetles(excludingcarabids),
reptiles,rodents,saproxylic beetles,scuttleflies, springtails, truebugs,
vascular plants and wood-inhabiting fungi. For the analysis compar-
ingresponses ofsaproxylicand non-saproxylicspecies groups,wede-
finedsaproxylicbeetles,wood-inhabitingfungi,andepixyliclichensand
mossesassaproxylicandallotherspeciesgroupsasnon-saproxylic.
Forcomparing numbersofspecies betweensalvagedand unsal-
vagednaturallydisturbedplotsdescribedin thepublished literature,
weusedHedges’d,whichaccountsfordifferencesinsamplingeffort
across studies and for small sample sizes (Hedges & Olkin, 1985).
PositivevaluesofHedges’dindicatehighernumbersofspeciesinsal-
vageloggedplots,whereasnegativevaluesindicatealossinnumbers
ofspeciesattributedtosalvagelogging(i.e.highernumbersofspecies
inunsalvagednaturallydisturbedplots).Meanabsoluteeffectsizesof
d=0.2indicateasmalleffect,d=0.5amoderateeffect,andd = 0.8 a
largeeffect(Koricheva,Gurevitch,&Mengersen,2013).
We used multi-level linear mixed-effectsmodels, provided by
the r function “rma.mv” in the “metafor” package (Viechtbauer,
2010),totest theeffectoftaxonomicgroup asacategorical pre-
dictor and year since disturbance as a numerical covariate on
Hedges’dastheresponsevariable.Hedges’dvalueswereweighted
bythecorrespondingsamplingvariancewithinthestatisticalmodel.
Furthermore,thestudysitewasincludedasarandomeffectinthe
model (i.e. moderator term) to control for unmeasured site spec-
ificities and repeated measurements (pseudo-replication) within
onestudysite.Thismeansthatmultipledatapointsperstudywere
possibleifstudiesexaminedmultipletaxonomicgroupsorifstudies
lastedformorethan1year.Wesubtracted theinterceptfromthe
effectsizes (byincluding “−1”in themodelformula)to evaluateif
observedHedges’ddifferedsignificantlyfrom zero(fordetails and
modelformulaseeTableS1).
Toevaluate theeffectsof salvagelogging on saproxylicvs.non-
saproxylic groups, we fitted a second model with Hedges’d as re-
sponsevariable.Weagainincludedtheyearafternaturaldisturbance

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andsubsequentlogging as a numericalpredictorvariable andstudy
siteas wellastaxonomicgroupasrandomfactors. Furthermore,we
addedtheinteractionofdeadwooddependence(i.e.saproxylic/non-
saproxylic)withnaturaldisturbancetypeaspredictorstotestwhether
theeffectofsalvageloggingonthe numberofspecies insaproxylic
andnon-saproxylicgroups differedwithin different typesofnatural
disturbances. Weimplemented a simultaneous inference procedure
tocomparesaproxylicandnon-saproxylicspeciesgroupswithineach
disturbancetype(Hothorn, Bretz,& Westfall,2008). Thisprocedure
allowedus totest ifresponsesofsaproxylicandnon-saproxylictaxa
varyamong fire-,wind-and insect-disturbedforests(fordetails and
model formula see TableS2). Last, we conducted funnel plots by
meansofthefunction“funnel”fromthe“metafor”packagetoassess
publicationbias(Korichevaetal.,2013;FigureS1).
2.3 | Analysis of community composition
Based on the reviewed literature, we compiled original species
abundance matrices to quantify changes in community composi-
tioninducedby salvage logging.Quantifyingchangesin community
compositionamong largeheterogeneousdatasetsischallenging and
requires statistical methods able to deal with issues such as unbal-
ancedsamplingeffortandwhichgenerateastandardizedeffect size
thatis comparableamong differentspecies groupsand surveytech-
niques.Thus,weusedpermutationalmultivariateanalysisofvariance
usingdistance matrices(Legendre&Anderson,1999),performedby
meansofthefunction“adonis”inthepackage“vegan”(Oksanenetal.,
2016).ThisanalysisprovidesapseudoF-value,basedon999permu-
tations,thatquantifies thedeviancefrom thenull-hypothesis,while
simultaneouslyaccounting forimbalanced studydesigns (McArdle&
Anderson,2001).Consequently,largevaluesofFcorrespondtolarge
changesin communitycompositioninducedbysalvagelogging.This
F-value represents the standardized difference between communi-
tiesinsalvageloggedandunsalvagednaturallydisturbedplotswithin
onespeciesabundancematrix(e.g. differences in bird communities
6yearsafterwildfire andsalvagelogging inOregon).We rigorously
restrictedthisanalysistothoseabundancematricesthatyieldedvalid
pseudoF-values overthe courseof permutations;that is,those ma-
triceswhichgeneratedlessthan99realpermutationswereexcluded.
Theserestrictions resultedin atotal numberof 134matrices, which
suppliedF-valuesfortheanalysisoutlinedbelow.
Totestifsalvageloggingchangedcommunitycompositionindif-
ferenttaxonomicgroups,wemodelledpseudoF-valuesinlinearmixed
modelsprovidedbythefunction“lmer”inthe“lme4”packageassum-
ingaGaussianerrordistribution(Bolkeretal.,2009).Weincludedthe
taxonomicgroupasacategoricalpredictorand theyearsincedistur-
banceas a numericalcovariate. Furthermore,weincludedthe study
siteasarandomeffecttocontrolforpossibledifferencesamongstudy
sitesand repeatedmeasurementswithin onestudysite.Weomitted
theinterceptfromthemodelformulatodetermineifF-valuesdiffered
significantlyfromzero.Thus,significant changesin communitycom-
positionofataxonomicgroupduetosalvageloggingwereindicated
by F-valuessignificantlylargerthanzero(fordetailsandmodelformula
seeTableS3).
AsfortheanalysisofHedges’d,asecondmodelwasfittedtotest
whether the effects of salvage logging on community composition
differed between saproxylic and non-saproxylic species groups in
differenttypesofdisturbances.Therefore,weincludedtheyearafter
disturbance and the interaction of saproxylic/non-saproxylic with
disturbancetypeaspredictors.Taxonomicgroupandstudysitewere
includedasrandomfactorsinthismodel.Weimplementedasimulta-
neousinferenceproceduretocomparesaproxylicandnon-saproxylic
species groups within each disturbance type (for details and model
formulaseeTableS4).
3 | RESULTS
Our meta-analysis showed that the effects of salvage logging have
been studied primarily for birds, vascular plants and carabids, par-
ticularly in burned forests. Studies were conducted primarily in
NorthAmerica andEurope, butlackingintropicalregions(Figure1).
Furthermore,therewasaclearlackofstudiesinvestigatingsaproxylic
taxa.Ofthe238compileddatapoints,170coveredaperiodof5years
or less after disturbance, with studies addressing the long-term ef-
fectsofsalvageloggingbeingrare(Figure2).Onlyonestudy(Hutto&
Gallo,2006)wasavailablethatprovideddataontheeffectsofsalvage
loggingformorethan20yearsafterdisturbances(Figure2).
Halfofthe individual comparisonsproducedvalues ofHedges’d
lowerthanzero,indicatinghighernumbers ofspeciesinnon-salvage
loggedareasthan salvageloggedareas (Figure3).Wefoundsignifi-
cantly lowerspecies numbers of epigeal and epixylic mosses, birds,
wood-inhabiting fungi, saproxylic beetles, springtails and epixylic
aswellas epigeallichensin salvage loggedareas comparedtonon-
salvagelogged areas(Figure3a).Incontrast,thenumbers ofspecies
of land snails, epigeal spiders and carabids were higher in salvage
logged areas than in unsalvaged areas (Figure3a). Thirteen of the
24 taxonomic groups, includingvascular plants, exhibited no signif-
icantresponsein numbers ofspeciestosalvage logging (Figure3a).
The numbers of species of saproxylic taxa significantly decreased
compared to non-saproxylic taxa in storm-affected and burned for-
ests(Figure4a).Thenegativeeffectofsalvageloggingonnumberof
speciesincreasedwithtimeelapsedsincedisturbanceandsubsequent
salvagelogging,althoughlong-termdataonsalvageloggingarescarce.
Salvageloggingwas associatedwithsignificant changes incom-
munitycompositionin 7 of 17taxonomicgroups(Figure3b).These
seven groups were epigeal spiders, carabids, vascular plants, birds,
wood-inhabiting fungi, saproxylic beetles and epixylic lichens
(Figure3b). Time elapsed since disturbance had no effect on the
strength of logging-induced changes to community composition
(TableS3).Furthermore,logging-inducedchangesincommunitycom-
positionwerestrongerforsaproxylictaxathanfornon-saproxylictaxa
in storm-disturbed forests. However, data availabilitywas scarce in
insect-affectedforestandlackinginburnedforests(Figure4b).

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4 | DISCUSSION
Our study revealed that salvage logging can result in significant
changes in species numbers and/or in altered community composi-
tion.Negativeeffects were particularlystrongfor taxa thatdepend
ondeadwood. Incontrast,thenumbers ofspeciesof taxa thatare
commonly characterized by species-rich communities in open habi-
tats, such as carabids and epigeal spiders, responded positively to
salvage logging. Despite positive effects of salvage logging on taxa
associated with open habitats, strong negative effects on saprox-
ylicgroupscall forsubstantialchanges inhowdisturbed forestsare
routinelymanaged.
Naturallydisturbedforestsarecharacterizedbylargevolumesof
deadwoodwithhighstructuraldiversity(Swansonetal.,2011).Incon-
trast,salvageloggingtypicallyreducestheamountandheterogeneity
of deadwood by removing tree trunks (Keyser, Smith, & Shepperd,
2009;Priewasser,Brang,Bachofen,Bugmann,&Wohlgemuth,2013).
Notsurprisingly, salvageloggingreduced thenumbersof speciesof
saproxylicgroups (Figures3and4). However,not onlyadecreasing
deadwoodamountbutlikewisealogging-inducedshiftindeadwood
qualitymay haveadditionalimpacts onsaproxylictaxa. Salvagelog-
gingnotonlyreducestheamountoflargetreetrunksbutalsoalters
characteristicconditions,such as decaystages or diameterdistribu-
tions,oftheremainingdeadwood(Waldronetal.,2013).Forinstance,
branchescutduringpost-stormloggingremainonthegroundbutare
overgrownbygroundvegetation.Theresultingshiftin microclimatic
conditionsthenmodifiesresourcequality,leadingtoalossofsaprox-
ylicbeetles thatdependon sun-exposed, drybranches (Thornetal.,
2014).
It is important to note that losses of saproxylic species can be
presentalso within taxonomicgroups thatdisplayedno responsein
theiroverallspeciesnumbers(Figure3a).Forinstance,birds(themost
studiedvertebrategroup)wereslightlynegativelyaffectedbysalvage
logging(Figure3a),despitefewspeciesbeing directlydependent on
dead wood. Nevertheless, several forest-dwelling bird species de-
pend on snags, cavities or natural regenerationin post-disturbance
forest stands. The removal of such legacies bysalvage logging can
causealossofassociated bird speciesandconsequentlyan overall
lowernumber ofbirdspecies in loggedareas(Hutto &Gallo,2006;
Werner,Müller,Heurich,&Thorn,2015).Althoughtheoverallnumber
ofbirdspeciesdecreasedlessstronglythan,forinstance,thenumber
ofsaproxylic beetlespecies(Figure3a),birdspecies thatdependon
post-disturbancehabitatcharacteristicsareoftenofhighconservation
interest.Forinstance,salvageloggingafterhighseveritywildfirescan
leadtolowersite occupanciesofNorthernSpottedOwls(Strix occi-
dentalis caurina)onlogged than onunloggedsites in Oregon(Clark,
Anthony,&Andrews,2013).
Ourstudyrevealedthatsalvageloggingcausedsignificantchanges
incommunitycomposition forsevenspeciesgroups(Figure3b),with
saproxylic species groups being affected most strongly (Figure4b).
Suchalterationsin community compositionmightreflect the estab-
lishmentofopen-habitatspeciesand/orasimultaneouslossofforest
specialists.Forinstance, salvagelogging canincreasetheabundance
ofopen-habitatcarabidbeetles(Koivula&Spence,2006)orpromote
the establishment of non-forest vegetation (Stuartetal., 1993; Van
Nieuwstadt,Sheil, &Kartawinata,2001).Hence,speciesgroupsthat
are commonly characterized by species-rich communities in open
habitats,suchascarabidsorepigealspiders,candisplayanoverallin-
creaseinnumbersofspeciesinresponsetosalvagelogging(Figure3a).
Likewise, salvagelogging can cause an increase in herb- and grass-
feedingmothspeciesbutadecreaseinsaproxylicanddetritus-feeding
mothspecies (Thornetal., 2015).Such contrastingresponseswithin
andbetweenspeciesgroupscan masktheoverallimpactofsalvage
logging on biodiversity in coarse-scale analyses (i.e. Thom & Seidl,
2016).Numerousspecies ofhighconservationinterest, suchasthe
Red-cockadedWoodpecker(Leuconotopicus borealis),dependondead
woodin burnedforests(Conner,Rudolph,& Walters,2001).The re-
sultsofourstudythereforeindicatethat thebiodiversityofsaprox-
ylictaxacouldbeenhanced bya modifiedmanagementofnaturally
disturbedforests.Incontrast,populationsofspeciesassociatedwith
open habitats, such as the Sharp-tailed Grouse (Tympanuchus pha-
sianellus)inNorthAmerica,maypersistorevenincreaseinthelarger
remaining area subject to unmodified management, that is, salvage
logging(Radeloff,Mladenoff,&Boyce,2000).
Thetwomajor incentivesfor salvageloggingare toreduceeco-
nomic losses caused by a naturaldisturbance and to omit mass re-
productionand spreadofinsectpests thatdevelop intrees killedor
weakenedbya preceding naturaldisturbance.Forinstance,salvage
logging of storm-felled Norway spruce (Picea abies) decreased new
infestationsofnearbytreesbytheEuropeansprucebarkbeetle (Ips
FIGURE2 Distributionofstudies
investigatingtheeffectsofsalvagelogging
onbiodiversityafterwildfire,windstorms
andinsectoutbreaksaccordingtotheyears
afterdisturbance.[Colourfigurecanbe
viewedatwileyonlinelibrary.com]

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typographus) at a landscape scale (Stadelmann, Bugmann, Meier,
Wermelinger,&Bigler,2013).Salvageloggingisthereforethepredom-
inantresponsetonaturaldisturbancesinwoodproductionforests,but
pestcontrolis regularlyusedtojustify salvage logginginprotected
areas.Forinstance,theBiałowieżaForestNationalParkontheborder
betweenPolandandBelarus,whichis thelast primevallowland for-
estinEurope,iscurrentlyobligedtosalvageloggingofareasaffected
by I. typographusonattempttoavoidfurtherinfestations(Chylarecki
&Selva,2016). Suchan approachtodisturbed forestsneglects that
regionalfactors,suchassummerdrought,canpromoteoutbreaksof
I. typographus more strongly than local standvariables (Seidl etal.,
2015).Furthermore,salvageloggedtimberis usuallyofsubstantially
lowereconomicvaluethannormallyharvestedtimberduetoarapid
colonization by wood-inhabiting fungi and to the fact that distur-
bancesaffect forestsofanyage,sothat generalizedsalvagelogging
operationsnecessarilyincludeyoungerstands thatotherwisewould
notbeharvested(Leverkus,Puerta-Pinero,Guzmán-Álvarez,Navarro,
& Castro, 2012). Our results demonstrate that salvage logging has
strongandnegative effects on manytaxonomic groups, particularly
thoseassociatedwith deadwood,and thatitisthus notconsistent
with biodiversity conservation goals.Along with questionable eco-
nomicoutputs andpestreducingeffects,weargue thatsalvagelog-
gingshouldbeexcludedfromprotectedareassuchasnationalparks.
The incidence of stand-replacing natural disturbances remains
spatially and temporally unpredictable (Berry etal., 2015), creating
inherentuncertaintyaboutappropriatemanagementofnaturallydis-
turbedforests.Hence,managementplansneedtobejointlydeveloped
with(andconfirmedby)stakeholders,scientistsandnaturalresource
managers before the nextdisturban ceoccurs (Lindenmayer, Likens,
& Franklin, 2010). Such management plans could,for instance, en-
compassan aprioriidentificationofsalvageloggingexclusionzones
basedonecologicaldata(e.g.Nappietal.,2011).Forestmanagersalso
may targetthe preservation of structural key attributes in naturally
disturbed forests, including snags or tipped uproot plates of wind-
throwntrees(Hutto,2006).Retentionoftreesduringgreen-treehar-
vestshas becomeanincreasingly commontool aroundtheglobe to
helpconserveforestbiodiversity(Fedrowitzetal.,2014;Gustafsson
etal.,2012;Mori&Kitagawa,2014).Toobtainsomeeconomicreturn
whileretainingdeadwood-dependenttaxa,werecommendasimple
expansionofthe green-treeretentionapproach toincludenaturally
disturbedforests.Retentionapproachesinnaturallydisturbedforests
couldbeexpectedtobelesscostlythaningreen-treeharvestdueto
theloweropportunitycostofnotharvestingdisturbance-killedtrees.
Approximately70%ofthestudieswecompiledspannedlessthan
5years; studies addressing the long-term effects of salvage logging
arerare(Figure2). However,deadwood, and particularlysnags, are
long-lastingkeybiologicallegacies,andtheirlosscanhavelong-lasting
effectson biodiversity(Hutto,2006). Hence,future researchshould
target the long-term effects of salvage logging after natural distur-
bances.Therearealsotaxonomicbiasesinexistingstudiesinvestigating
FIGURE3 (a)Estimatedresponse
ofHedges’dbasedon238individual
comparisonsofspeciesnumbersin
salvageloggedandunsalvagedforests
affectedbynaturaldisturbances.Higher
speciesnumbersinsalvageloggedareas
correspondtopositiveHedges’d,whereas
negativevaluesindicatelowerspecies
numbersinsalvageloggedareas.(b)Pseudo
F-valuesofpermutationalmultivariate
analysisofvariancebasedon134individual
speciesabundancematrices.Largerpseudo
F-valuescorrespondtolargerchangesin
communitycompositioninducedbysalvage
logging.Asterisksindicatesignificant
responses(seeTablesS1andS2for
statisticaldetails).Forillustrativepurposes,
greydots(andthegreylinejoiningthemfor
emphasis)representthemeaneffectsizein
eachtaxonomicgroup.[Colourfigurecan
beviewedatwileyonlinelibrary.com]

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theeffectsofsalvageloggingafternaturaldisturbances.Inparticular,
saproxylicgroupssuchaswood-inhabitingfungihavebeenunderrep-
resentedin empirical studiesdespitetheir high diversityand impor-
tance for ecosystem functioning. Future research should therefore
targetparticularlysaproxylicspeciesgroups.Incontrast,othergroups
havebeenrelativelywellstudiedinonedisturbancetype(e.g.birdsin
burnedforests),but lessinothers,and studies wereconducted pri-
marilyin NorthAmerica, Europeand Asia,butlacking intropicalre-
gions(Figure1).However,differenttypesofnatural disturbancesin
differentpartsoftheworldcanactatverydifferentspatialscalesand
mayrequiredifferentretentionapproaches(Kulakowskietal.,2016).
Furthermore,coniferousforestsoftheNorthernHemisphere—incon-
trasttotropicalforests—arenaturallypronetolarge-scalenaturaldis-
turbances(Lindenmayeretal.,2008),whereasdisturbancesintropical
forestsmostlyhave anthropogeniccausesassociatedwithlong-term
land-use change (e.g. fire to open space for livestock grazing and
agriculture; Peres,Barlow, & Laurance, 2006). Nevertheless, natural
disturbancessuchaswindstormsaffecttropicalforestsaswellastem-
perateforests,andsalvageloggingeffectson tropicalforestsshould
betargetedinfutureresearch(e.g.Lawton&Putz,1988).
In conclusion, these data from a wide range of studies demon-
strate that salvage logging has a range of effects on species num-
bers and community composition ofvarious taxonomic groups,with
important negative consequences for several groups, especially
saproxylic ones. Whilecurrent policies for enhancing biodiversity and
ecosystemservices,suchasgreen-treeretention(e.g.,Gustafssonetal.,
2012), focus mainly on forestssubjected to traditional logging opera-
tions,suchpolicies are largelyabsent fromnaturallydisturbed forests.
Wethereforecallforanexpansionofthegreen-treeretentionapproach
toincludenaturallydisturbedforestsbyleavingsubstantialamountsof
deadwoodonsitetoreducetheimpactofsalvageloggingonbiodiversity.
ACKNOWLEDGEMENTS
Wethanknumerouscontributorsforclarifyingtheirstudiesandthree
anonymousreviewersfortheircommentsonanearlierversionofthis
manuscript.S.T.andS.S.werefundedbytheGermanEnvironmental
Foundation. R.S. and D.T. acknowledge support from the Austrian
ScienceFund(FWF,STARTgrantY895-B25).J.C.acknowledgessup-
portfromgrantP12-RNM-2705andA.L.fromSpanishMINECO(FJCI-
2015-23687).D.B.L.wassupportedbyanARCLaureateFellowship.
AUTHORS' CONTRIBUTIONS
S.T.andJ.M.initiatedthestudy.S.T.analysedandinterpretedthedata
andwrotethefirstdraftofthepaper.TheauthorsnamedfromS.T.to
J.M.arelistedalphabetically,astheycontributedequallyingathering
fielddata,providingcorrectionstosubsequentmanuscriptdraftsand
discussingideas.
DATA ACCESSIBILITY
All data are from previously published articles, see “Data sources”.
Datafromthesearticlescan bemadeavailable uponreasonable re-
questtooriginaldataowners.Alistofdatasourcesusedinthestudy
isprovidedintheDataSourcessection.
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reference.[Colourfigurecanbeviewedatwileyonlinelibrary.com]

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How to cite this article:ThornS,BässlerC,BrandlR,etal.
Impactsofsalvageloggingonbiodiversity:Ameta-analysis.
J Appl Ecol. 2018;55:279–289. https://doi.org/10.1111/1365-
2664.12945