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Arthropod communities in fungal fruitbodies are weakly structured by climate and biogeography across European beech forests

  • Nationalpark Bayerischer Wald

Abstract and Figures

Aim The tinder fungus Fomes fomentarius is a pivotal wood decomposer in European beech Fagus sylvatica forests. The fungus, however, has regionally declined due to centuries of logging. To unravel biogeographical drivers of arthropod communities associated with this fungus, we investigated how space, climate and habitat amount structure alpha and beta diversity of arthropod communities in fruitbodies of F. fomentarius. Location Temperate zone of Europe. Taxon Arthropods. Methods We reared arthropods from fruitbodies sampled from 61 sites throughout the range of European beech and identified 13 orders taxonomically or by metabarcoding. We estimated the total number of species occurring in fruitbodies of F. fomentarius in European beech forests using the Chao2 estimator and determined the relative importance of space, climate and habitat amount by hierarchical partitioning for alpha diversity and generalized dissimilarity models for beta diversity. A subset of fungi samples was sequenced for identification of the fungus’ genetic structure. Results The total number of arthropod species occurring in fruitbodies of F. fomentarius across European beech forests was estimated to be 600. Alpha diversity increased with increasing fruitbody biomass; it decreased with increasing longitude, temperature and latitude. Beta diversity was mainly composed by turnover. Patterns of beta diversity were only weakly linked to space and the overall explanatory power was low. We could distinguish two genotypes of F. fomentarius, which showed no spatial structuring. Main conclusion Fomes fomentarius hosts a large number of arthropods in European beech forests. The low biogeographical and climatic structure of the communities suggests that fruitbodies represent a habitat that offers similar conditions across large gradients of climate and space, but are characterized by high local variability in community composition and colonized by species with high dispersal ability. For European beech forests, retention of trees with F. fomentarius and promoting its recolonization where it had declined seems a promising conservation strategy.
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Diversity and Distributions. 2019;25:783–796.  
  Revised:18O ctober2018 
  Accepted:11November2 018
Arthropod communities in fungal fruitbodies are weakly
structured by climate and biogeography across European beech
Nicolas Friess1| Jörg C. Müller2,3 | Pablo Aramendi4| Claus Bässler2|
Martin Brändle1| Christophe Bouget5| Antoine Brin6| Heinz Bussler7|
Kostadin B. Georgiev2,3| Radosław Gil8| Martin M. Gossner9|
Jacob Heilmann‐Clausen10| Gunnar Isacsson11| Anton Krištín12| Thibault Lachat13,14|
Laurent Larrieu15,16| Elodie Magnanou17,18 | Alexander Maringer19| Ulrich Mergner20|
Martin Mikoláš21,22| Lars Opgenoorth1| Jürgen Schmidl23| Miroslav Svoboda21|
Simon Thorn3| Kris Vandekerkhove24| Al Vrezec25| Thomas Wagner26|
Maria‐Barbara Winter27| Livia Zapponi28| Roland Brandl1| Sebastian Seibold29
11SwedishForestA gency,Hässleholm,Sweden
12InstituteofForestEcolog ySAS,Zvolen,Slovakia
15INRA ,UMR1201DYNAFOR ,ChemindeBordeRouge,UniversityofToulouse,CastanetTolosanCedex,France
16CRPFOC ,Tolosane,France
18Réser veNaturelleNationaledelaForêtdelaMassane,Argelès‐sur‐Mer,France
23Ecologygroup,DevelopmentalBiolog y,DepartmentBiology,UniversityofErlangen‐Nuremberg,Erlangen,Germany
©2019TheAuthors.Diversity and DistributionsPublishedbyJohnWiley&SonsLtd.
   FRIESS Et al.
26Depar tmentofBiolog y,UniversityofKoblenz‐Landau,Koblenz,Germany
27ForestResearchInstituteofBaden‐Würt temberg(FVA),Freiburg,Germany
29TerrestrialEcolog yResearchGroup,DepartmentofEcologyandEcosystemManagement,TechnischeUniversitätMünchen,Freising,Germany
ResearchGroup,Depar tmentofEcology
Funding information
Aim:ThetinderfungusFomes fomentariusisapivotalwooddecomposerinEuropean
beechFagus sylvaticaforests.Thefungus,however,hasregionallydeclinedduetocen
atedwiththisfungus,weinvestigatedhowspace,climateandhabitatamountstruc ture
alphaandbetadiversityofarthropodcommunitiesinfruitbodiesofF. fomentarius.
Tax o n:Arthropods.
coding.WeestimatedthetotalnumberofspeciesoccurringinfruitbodiesofF. fom en
tarius in European beech forests using the Chao2 estimator and determined the
Results:ThetotalnumberofarthropodspeciesoccurringinfruitbodiesofF. fomentarius
latitude. Beta diversitywas mainly composed by turnover.Patterns of beta diversity
distinguishtwogenotypesofF. fomentarius,whichshowednospatialstructuring.
Main conclusion: Fomes fomentariushost s a l a r g e n u m b er o f a r th ro p o ds i n Eu r o p e a n
suggeststhatfruitbodiesrepresentahabitatthatoffers similarconditionsacross
European beech forests, retention of trees withF. fomentariusandpromotingits
deadwood,Fagus sylvatica,Fomes fomentarius,insects,invertebrates,restoration,saproxylic,
Most parts of the temperate zone of Europe—from the Iberian
Peninsula to the Black Sea and from southern Italy to southern
Sweden—are naturallycoveredby forestsdominated by European
beechFagus sylvatica (Fig ure 1). These forests, however,have de‐
clined over recent centuries due to deforestation until around
sylvestris,Picea abies)plantations(Dirkx,1998;Schelhaas,Nabuurs,&
been reinforced by large‐scale clear‐cutting of old‐growth beech
forestsin regionsthat,until recently, were rather unaffected (e.g.,
al.,2017).Sincethe distributionofEuropean beechisrestricted to
thetemperatezoneofEurope,theEUhas acknowledgeditsglobal
FRIESS E t al.
UNESCO World Heritage “Ancient and PrimevalBeech Forests of
theCarpathians andOther RegionsofEurope”(http://whc.unesco.
driversof biodiversity inbeechforests remainslimited,hampering
systematic conservation planning, given prevalent area conflicts
reflecting different underlying mechanisms. European beech was
one of the last tree species to recolonize central and northern
ciation and is still expandingits range towards the north andeast
(Magri, 2008). Understorey plant diversity in European beech for‐
climate tolerance,Europeanbeech covers a widerange of climatic
conditions (Figure 1; Brunet, Fritz, & Richnau, 2010), which might
arthropodsassociatedtothesetrees (Brändle&Brandl,2001)may
These natural drivers ofcommunity structure in beech forests
interact with anthropogenic factors. Forest clearing and forest
manageme nt have been more intens e in western than in e astern
and conse quently fragme ntation of these fo rests from eas t–west
(Abrego,Bässler,Christensen, & Heilmann‐Clausen, 2015;Kaplan,
Krumhardt, & Zimmermann,2009;Larsson,2001). Manyspecialist
Onsmallerspatialscales,speciescommunities canbe affected
bytheregionalclimate acting as environmental filterasshown for
gradients of anthropogenic pressure can influence communitiesin
FIGURE 1 Mapofthe61samplingsitesofthisstudy.ThegreenareadepictsthepredictedcurrentdistributionofEuropeanbeech
Fagus sylvatica(Brusetal..,2011).ThenumbersinthemapcorrespondtothestudysiteIDinsupportinginformationAppendixS2and
withfruitbodiesofFomes fomentarius. PhotographbyThomasStephan.Rightinset:Meanannualtemperatureandannualprecipitationof
   FRIESS Et al.
beech forests butalsothe amountofavailablehabitat at localand
tivityofhabitat patches (Abrego et al., 2015; Nordén et al.,2018;
Fun giareth emainbiot icagent sofwooddecom posit ionan dt heir
astheycontain higherconcentrationsofnutrientsstoredin amore
ac ces sibl efor mtha ninundec ayed woo d(Fi lipi ak,S obczyk, &Weine r,
tatformany fungicolousarthropod species(Schigel,2012).Studies
host spe cies, but low amon g sites across host sp ecies (Komonen,
ThetinderfungusFomes fomentariusisoneofthemaindecom
posersofwoodinmanybeechforestsinEurope.However,F. fomen
tarius has a much l arger range tha n European bee ch covering the
temperate and boreal zones of Europe, Asia and Nor th America.
estsonBetula,Populus, Alnus orotherhardwoodtrees(Matthewman
itcan efficiently break down lignocellulose andcontributes tothe
death ofweakenedliving trees, thus promoting naturalforest dy‐
habitatformanyarthropodspecies (Schigel,2012).Their commu
Reibnitz, 1999; Thunes &Willassen, 1997).Thus, in order to cap‐
ture the w hole local com munity occur ring in F. fomentarius differ
Treescolonized by the fungus have been suggested as afocal
habitat for biodiversity conservation in beech forests (Larrieu
et al., 2018; Mül ler, 2005). Howe ver, due to centuri es of logging
and direct persecution for phytosanitary reasons, populations of
this fungus have declined orbecamelocally extinctin many areas
(Vandekerkhove etal., 2011;Zytynskaet al.,2018). Toguide con‐
servation planning andstrategies inEuropeanbeechforests, such
asthe selectionof areastobe setaside forconservation(Bouget,
Parmain,&G ilg ,2014)orforacti verestoratio nbyde adwoodenrich
ment(Dörfler,Gossner,Müller,&Weisser,2017), it isnecessaryto
In this study, we reared arthropods from fruitbody samples of
F. fomentarius across the whole distributional range of European
podsinfruitbodiesofF. fomentariusandtodise nt angletheeffect sof
weexpected (a) decreasing alphadiversity and increasingnested
creasingalphadiversity and increasing nestednessfromeast‐west
due to the an thropogenic l and use histor y, (c)i ncreasing tur nover
2.1 | Collection of Fomes fomentarius fruitbodies
We collected fruitbodies from 61 beech‐dominated forest sites
across the distributional range of F. sylvatica (Figure 1) between
distributionofF. sylvatica,aswellasthefullrangeofclimaticcondi‐
tions withinthis area(Figure1).Wewerenot abletoincludesites
from someparts of thedistributionalrange,for example southern
England,whereF. fomentarius isalmostabsentforhistoricalreasons
(Abreg o, Christensen, B ässler, & Ainswort h, 2017). Sites were lo‐
cated inunmanaged (36) and managed forests (25); both manage‐
Forarthropodrearing,wecollected10fruitbodiesofF. fomentar
iuspersitefoll owi nga stand ardizedpro toc ol.A ssem blagesi nhab iting
position.Therefore,wesampledfruitbodiesatdif ferentsuccessional
frui tbodies).Thelatterwer ee itherdrywhens tillatta chedtowoo d(3
didnotrepresent the local availabilityoffruitbodies as transporta
In additi on, we collect ed samples of li ving fruitb odies to anal‐
ysethe geneticstructure withinthe population of F. fomentarius in
2.2 | Arthropod rearing
“sample”) w ere put into a cardbo ard box (25cm×25cm×50cm)
in an unheated well‐ventilated storage room with a seasonal
FRIESS E t al.
temper ature regime. A tra nsparent collec ting jar was att ached to
podsinside theboxeswerecollectedbyhand. Rearingwascarried
2.3 | Arthropod identification and classification
Reared ar thropod specimens were stored in ethanol and beetles
were determined to species level by taxonomists. The remaining
fauna was identified by metabarcoding using next‐generation se
quencin g carried out by Adv anced Identific ation Methods G mbH
S1).Arthropodsequenceswerematched against the publicly avail
able DNA barcode library within the Barcode of Life (BOLD; Ratnasingham & Hebert, 2007). Laboratory
ples,includingspeciesthatusehollowfruitbodiesas shelterorde‐
However, since this i ncludes specie s that do not intera ct directly
with the fruitbody, we additionally analysed the data excluding
these species.Basedon literature,we classified species orgenera
thatare knownto feeddirectly on thefungaltissue or exclusively
2.4 | Environmental predictor variables
Coordinatesofeachsitewererecordedin the fieldusinghandheld
WorldClim database (Hijmans, Cameron, Parra, Jones, & Jarvis,
thesites.Thefirstprincipal componentsrepresented a gradient in
ality withhigh values forsites displaying hightemperature orpre‐
Too btain a proxy f or landscap e‐scale habi tat amount an d an‐
thropogenicpressure,we calculated the proportionofforestcover
surroundingthe sitesforradiifrom 100 to 5,000m(100‐msteps).
Forest coverwithina radiusof700maround siteshadthehighest
alphaand betadiversity,werecordedthetotal dryweightoffruit‐
2.5 | Statistical analyses
AllstatisticalanalyseswerecarriedoutusingR version3.3.2(RCore
Tea m, 2016) .T he main analyses i ncluded beetl es identified t axo‐
werethus restricted to the52sites for which metabarcoding data
Toestimate the overall species pool, we calculated the Chao2
estimator, as implemented in the vegan package version 2.4–3
(Oksan en et al., 2018). The Chao2 es timate is a functi on of spe‐
bound es timation for sp ecies richnes s based on inciden ces under
(Chao, 1987). Cal culations were b ased on data for a ll species and
predator a nd parasitoid ) on the 52 sites. In ad dition, we use d the
rarefaction–extrapolation framework based onsp eciesincidences
0(species richness),1 (the exponential of Shannon's entropy) and
2(the inverse of Simpson's concentration)to analyse thediversity
ofrare andcommonspecies within one framework. We used 999
replicated bootstraps to calculateconfidence intervalsaround the
species‐accumulationcurves using the iNEXT package (Hsieh, Ma,
Toestimate therelative importance of thepredictor variables, we
package version 1.0–4 (Walsh & Mac Nally,2013)—based ongen‐
aquasipoissonerrordistributionanda log‐linkfunctionin orderto
data.Ple asenotethatalternativelych oosingmod elsincludin ga no b
serva tion‐level ran dom effect o rm odels with a neg ative‐binomia l
alphadiversity as the dependentvariableandspace(latitude,lon
precipitation,precipitation seasonality)andhabitatamount(forest
   FRIESS Et al.
cover,sample size)aspredictor variable sets. Allcalculations were
all 52 sites using presence–absence information. The community
compositionofall speciesandfungi specialistswasvisualized using
to the resulting ordination as implemented in the envfit function
using the vegan package. In ad dition, we per formed an a nalysis of
similarityin ordertotest for groupdifferences in communitycom
position a mong managed and un managed sites, as wel l as among
biogeographicalregionsagainusingvegan (seeSupportinginforma
beta diversity in its turnover and nestedness components based
onthe Sørensenindexfamilyas implementedinbetapart (Baselga,
Toestimatethe relativeimportanceofthepredictor variables (lati
tude,longitude, meantemperature,temperatureseasonality,mean
for beta diversity, we calculated generalized dissimilarity models
arately. GDMs allow theanalysis of spatial patternsof community
compositionacrosslargeregions under consideration of nonlinear
AllGDMswerecalculatedusingthedefaultofthree I‐splines.The
calculatedcoefficientfor each of the threeI‐splinesrepresentsthe
thefirstI‐splineindicateahigh rateofchange alongthe first third
containingallpredictorsetsandamodel fromwhichthispredictor
Dataforbeetlesincludingabundances wereavailable for all61
ualsonalpha diversity andtheeffectof space,climate and habitat
ties. He re, we used Bray–Cur tis dissimil arities and de composed it
(i.e., individuals of some species ata site are substituted by equal
numbersof individualsat anothersite)and dissimilarityintroduced
In total, we identified 216 arthropod species emerging from fruit‐
bodiesof F. fomentariusfrom52sites.Speciesbelongedto13or
ders, with highest species richness found in Diptera ( n=72) and
The majority oftaxa (n=179) couldbe assigned to species bythe
taxonomist or by alignment of operationaltaxonomic units (OTUs;
see Supporting information Appendix S1)with existing databases.
bers ofthe Cecidomyiidae (Diptera), for whichbarcodes were not
ers, 68 speciesaspredatorsand17species as parasitoids.Genetic
analysisofF. fomentariussamplesrevealedtwogenotypesthatwere
previouslyidentified aspossiblesympatric crypticspecies(termed
Chao2 est imators indic ated an overall s pecies pool of 5 87(S E
=103) for all specie s, 249(S E =181) for fungi spe cialists, 40 2 (SE
=104) for consumer s, 163 (SE =43) fo r predators an d 42(S E =24)
for parasitoids associated with F. fomentarius in European beech
forest s. The observed effec tive number of typical species (q=1)
was87,whilethe observedeffective number of dominant species
(q=2)was 44 (Supporting informationAppendix S3, Figure S3.3).
Manyofthedominantspecieswereconsumers,such as beetles of
the family Ciidae, the Tenebrionidae Bolitophagus reticulatus, the
micro‐mothScardia boletellaandCecidomyiidaesp.3(Figure3).The
most frequent par asitoids were the hyme nopterans Astichus spp.
tobe “primeval forest relicts” (Eckeltet al., ), namely Bolitophagus
interruptus, Bolitochara lucida, Teredus cylindricus and Philothermus
evanescens, whi ch were each found at on e site (Slovenia, France ,
FIGURE 2 Piechartoftheproportionofspeciesfromdifferent
arthropodordersrearedfromfruitbodiesofFomes fomentariusfrom
FRIESS E t al.
was16(SE=6) with thelowest number (six species) foundin the
German Wetterauandthe highestnumber(36 species)locatedin
26% for the fung i specialists (F igure 4). The explain ed deviance
decreased from consumers(22%) topredators (16%)and parasit
(Table 1).According to hierarchical part itioning, habitat amount,
thatisforestcoverandsamplesize,explainedmostoft hedeviance
inourmodels(Fig ure4).Alp hadiver sit yofa llspecies,fungisp ecial
is t s ,cons u mersa n dpred a torsi ncrea s e dwit h incre a sing s amples ize
(Table1,Figure5a)andthatofconsumersalsoincreasedwith in
thatoffungispecialistsalsodecreasedwithlatitude.Alpha diver
Ordinationofthecommunitycompositionofallspeciesas well
sition acrossour study sites(Supporting informationAppendixS3:
Figure S3 .2). Except for a si gnificant ef fect of samp le size on the
communitycompositionofallspecies(r2=0.13,p < 0.05), environ‐
bynestedness with increasinglongitudinal distance between sites
sityoffungispecialists andconsumerspecies (Supportinginforma‐
(Suppor ting inform ation Append ix S3, Table S3.6 and Table S3 .7).
Our a n a l ysesfo r b e e t l esfroma l l61 sitesi n c l u d e dabundan c e data
for123 species (Supporting information AppendixS5).Here,alpha
increased with fungal sample size as the range in samplesize was
consider ably higher acr oss all 61 sites (Figure 5b) th an across the
subset of 52 sites (F igure 5a). Beetl e community co mposition was
FIGURE 3 Rank‐incidenceplotofall216arthropodspecies
rearedfromfruitbodiesofFomes fomentariusfrom52beech‐
FIGURE 4 Relativecontributionofpredictorsetsinexplaineddevianceofalphaandbetadiversityanditscomponentsturnoverand
Fungi specialists
All species
Alpha diversity
0510 15
Beta diversity
0510 15
0510 15
Explained deviance
in alpha-diversity (%)
Deviance explained exclusively
by sets of variables (%)
   FRIESS Et al.
affec ted by dissimilarit y in sample size and longitude. Here, bee
tle commu nities showed i ncreased rat es of turnover and balanc ed
changes of abundances withlongitudeand increased ratesofnest
beetle species explainedupto59%of the deviance in alpha diver
sity,34% inSørensen dissimilarity and 19%inBray–Curtisdissimi
Var iable slinkedtoha bit atamountconsis tentl yexplainedmos toft he
Overall,our results indicatethatfruitbodiesof F. fomentariusform
an important micro‐habitat in European beech forests, host inga
rich fauna (estimat ed ~600 art hropod species). Howe ver, t he ar
thropod communities included about 30 dominantspecies which
occurredatmost sitesacrossEurope andcan beconsideredtypi
cal for fr uitbodies of F. fomentarius. Moreover, there was a lar ge
number of species thatuse F. fomentarius fruitbodiesoccasionally.
The latter groupincludes fungicolousspecies usingawiderrange
offungal hosts (e.g.,Bolitophagus interruptus,Coleoptera,which is
more common on Ischnoderma spp.),speciesthatfeedonwhite‐rot‐
ten wood (e.g ., Corymbia scutellata, Coleoptera) or fu ngal mycelia
and spec ies that use cavi ties inside fru itbodies sim ply for shelter
(e.g. , Amaurobius fenestralis, Aranaea)or that benefit from arthro
podprey(e.g.,Plegaderus dissectus, Coleoptera).Alpha diversity in
1,80 0km in latitude an d 3,000km in lo ngitude), beta dive rsity—
which wascharacterized by high turnover—wasnot structured by
drivers associated withspace, the biogeographyofF. sylvatica and
habitat amount. Moreover, increasing nestedness and decreasing
TABLE 1 Z‐valuesandexplaineddevianceofgeneralizedlinearmodels(quasipoissonfamily)withthenumberofspeciesofallspeciesor
Predictor set Predictor All species Fungi specialists Consumer Predator Parasitoids
Space Latitude −1.36 −1. 98 ** −1 .70 ** −0.08 0.93
Longitude −1. 77** −2 .1 2* −1. 8 2** −1 . 0 8 −0.34
Climate Temperature(PC1) −1 .7 2** −1. 90** −1. 92 ** −0.48 −1. 0 0
Temperature(PC2) −0.82 −0.79 0.43 −1.24 0.31
Precipitation(PC1) −1.41 −1. 3 0 −1. 5 7 −0.68 −0.01
Precipitation(PC2) −0.31 0 .13 −0.45 0.74 −1. 49
Habitatamount Forestcover 1. 26 0.94 1.45 0.13 0.13
Samplesize 1.75** 2.20*1.55 2.20*0.17
Explaineddeviance 0.20 0.26 0.22 0.16 0.06
Note. aSignificancelevels:*p<0.05,**p < 0.1
FIGURE 5 Relationshipbetween(a)thenumberofarthropodspeciesperfruitbodysampleandsamplesize,thatisthetotalweightof
(a) (b)
FRIESS E t al.
alpha diversity towards theeast follow not the continental gradi
Post‐glacial dispersal lags have been identified as one of the
driving mechanisms causing patterns of alpha and beta diversity
Svenning, Fløjgaard,&Baselga,2011;Svenning,Normand,&Skov,
guildsinF. fomentarius fruitbodies.Onlypredatoryspeciesshowed
an increased rate in turnover with increasing latitudinal distance:
the rateof change in species composition was highest atlow lati‐
tudes (Supportinginformation Appendix S3,TableS3.6).There are
several potentialexplanations as to why post‐glacialrecolonization
ofthemainhost treespecies appears to be ofminor relevance for
communitiesofarthropods occurring inF. fomentariusfruitbodies.
For instance, species associated with fungal fruitbodies ingeneral
experiments showedadispersal ability of Neomida haemorrhoidalis
and Bolitophagus reticulatus(bothColeoptera;bodylength:6–8mm
and 6 – 7.5mm, respectively; Wagner & Gosik, 2016) of>30km
and>100km, respectively (Jonsson, 2003). Additionally, there is
evidence thatthegenetic distance offungivores doesnotincrease
with geographic distance, indicating the absence of dispersal lim
itation ( Kobayashi & S ota, 2016). Another poss ible explanatio n is
thatalthough European beechis the mainhost of F. fomentarius in
IfF. fomentarius recolonizedEuropewiththelattertreespecies,its
arthropods may have had more time for recolonization and thus
etal.,2016),recolonizationpathwaysmay becomplexandnotwell
TABLE 2 CoefficientsofthreeI‐splines(i.e.,1,2and3)fromtheGDMofoverallbetadiversity,turnoverandnestednessofallarthropod
species.Significant(p < 0.05)ormarginallysignificant(p<0.1)P‐valuesfortheI‐splinesofthepredictorvariablesafter999permutations
Response matrix Predictor set Predictor
Sum of
coefficients P1 2 3
Overallbeta Space Latitude 0.155 0.007 0.003 0.165 0 .11
Longitude 00.067 00.067 0.42
Climate Temperature(PC1) 0 0 0 0 0.99
Temperature(PC2) 0.017 0 0 0.017 0.68
Precipitation(PC1) 0 0 0 0 0.99
Precipitation(PC2) 0.061 0 0 0.061 0.35
Habitatamount Forestcover 00.016 00.016 0.73
Samplesize 0.12 0 0 0.1 2 0. 24
Turnover Space Latitude 0.116 00.054 0.170 0 .24
Longitude 0 0 0.082 0.082 0.46
Climate Temperature(PC1) 0 0 0 0 0.99
Temperature(PC2) 0.004 00.030 0.034 0.65
Precipitation(PC1) 0 0 0 0 0.99
Precipitation(PC2) 0 0 0 0 0.99
Habitatamount Forestcover 00.018 0.001 0.019 0.73
Samplesize 0.121 0 0 0.121 0.25
Nestedness Space Latitude 0.001 0 0 0.001 0.70
Longitude 0.132 0 0 0.132 0.07
Climate Temperature(PC1) 0.013 0 0 0.013 0. 51
Temperature(PC2) 0 0 0 0 0.98
Precipitation(PC1) 0.010 0 0 0.010 0.57
Precipitation(PC2) 0.018 0 0 0.018 0.47
Habitatamount Forestcover 0.05 0 0 0.05 0.27
Samplesize 0 0 0 0 0.97
   FRIESS Et al.
forest s have become rar e or extinct i n western Europ e (Eckelt et
increase of fungicolous arthropod alpha diversity with increasing
longitu de, but in fac t we observed a we ak decrease . Additionall y,
wefoundamarginallysignificant increaseincompositionaldissim
ilarity due to nes tedness with incre asing longitudinal distance of
the overal l arthropo d community. Howeve r,the r ate of change in
explanatory power was low and nestedness did not account for
more than4%of compositionaldissimilarity(Table 2). Forbeetles,
we found an in creased rate in tu rnover and balance d changes of
informationAppendixS4, TableS4.4). In parallel to thegradient of
(Supporting information Appendix S3, TableS3.3). Bothdecreasing
aswellasincreasedbeetleturnoveratlowlongitudesare inconsis‐
tent with t he expected ef fect of histor ic anthropogen ic pressure,
wehavetopointoutthatwewerenotabletocollectF. fomentarius
offruitbodiesofF. fomentarius.Moreover,manyofoursites,alsoin
western Europe,werelocatedinunmanagedforests(Figure1)and
althoug h forest manage ment had no eff ect on overall co mmunity
Environme ntal filtering by climatic dr ivers is often an imp ort‐
ant mecha nism struc turing commun ities (Cadot te & Tucker, 2017;
(Bässleretal.,2010;Mülleretal.,2012;Seiboldetal., 2016).Being
poikilothermic,arthropodsgenerally benefit fromhighertempera
tures (Sc howalter, 2006). However, we foun d a marginally sig nifi‐
explana tion is that frui tbodies are dr ier and thus les s suitable for
thatclimate is ofminorimportance for arthropodsassociatedwith
F. fomentarius despiteconsiderablevariabilityin climaticconditions
ers of biodiversit y (Fahrig, 2013; MacAr thur & Wilson, 1967). In
Europe, hu man activiti es over millennia have re duced the fores ts
and features of old‐growth stands (overmature and dead trees),
2015).Forestcover is only a coarse proxy for theamountofhab‐
itat available tospecies associated with dead wood or fruitbodies
ofF. fomentarius,astheamountof theiractualhabitat—dead wood
orfruitbodiesofF. fomentarius,respectively—canvaryconsiderably
within beech forests depending, for example, on current forest
Müller, 2014). This was als o reflected by t he time need ed to find
tenfruitbodiesofF. fomentarius inthepresentstudy,whichranged
fromminutestodays.Nevertheless,wefound thenumber of con‐
beetle s to increase with f orest cover (70 0m radius aroun d sites).
offruitbody availability on fungicolousbeetlediversity atregional
foundthenumberofarthropodspeciestoincrease withincreasing
notreflecttheabundanceofF. fomentarius atthesites,basedonour
fruitbodiesofF. fomentarius.
videmoredifferenthabitatniches, forexample through different
stages of d ecomposition wi thin and among fr uitbodies (Dajoz et
al., 1966) similarl y as shown for coar se woody debri s (Seibold et
al.,2016).Concerningcommunitycomposition, only the totalbeta
diversity and turnovercomponent of predatory arthropods were
affected by sample size.However, abundance‐baseddissimilarity
in community composition ofb eetleswas affec ted by longitude
and sample size. Here, dissimilarity due to abundance gradients
(analogous to nestedness) increased with sample size. Overall,
this indic ates that loc al habitat amo unt is an impor tant driver of
for fungicolous beetle communities,an impor tant driver of beta
populationsofF. fomentarius arecomprisedoftwosympatriccryptic
species;this hasbeen confirmedbyPristas, Gaperova,Gaper,and
A, has b een suggested to b e prevalent on Europ ean beech whil e
the other,termed genotype B, isadditionally found on other host
species (Judova et al.,2012).Our geneticanalysisofF. fomentarius
supports this, as all but 5 of 36 of our samples—all sampledfrom
renceofgenotype BonEuropeanbeechinthePyrenees,southern
Italy, Belgium andDenmark is a noteworthyresult(Supportingin‐
ongenetic differencesfruitless.Furtherstudiesareneeded to test
the hypothesis that F. fomentarius of genotype B hos ts arthro pod
Inour analyses, we incorporated variables whichare knownto
bestrongdriversoflarge‐scale differencesin communitycomposi‐
FRIESS E t al.
2017).Fur thermore,weaccountedfordifferencesinhabitatspe cial
ization andtrophic level,forestmanagementintensityand biogeo‐
proportions ofvariationin alpha diversity most of the variation in
thecommunity compositionofarthropods occurringinfruitbodies
of F. fomentarius remained unexplained. Although explaining the
full variation in community composition was beyond the scope of
this study, these results appear surprising. We suggest three di‐
rections for future studies. First, future studies investigating the
community composition ofar thropods occurringin fruitbodies of
bracket fungishouldfocus on factors driving communitycomposi‐
tion at local scales.This mayincludethe amount of fruitbodies at
rescueeffects givensufficientpatchconnectivity(Gonzalez,2005;
could inves tigate the ef fects of mic roclimate as me diated by can‐
opyopennessand forestsuccessional stage, whichwereshown to
generate largedifferencesincommunitycomposition in saproxylic
organisms (Hilmers etal., 2018;Seibold et al., 2016).Second, fur‐
ofF. fomentariusonotherhost treespecies, suchas Betula spp. or
Populus spp.,andinvestigatepotentialalternativepost‐glacialrecol
onizationroutes.Third,to betterunderstand scale‐dependency of
F. fomentariusincluding NorthAmericaandEast Asia.Forinstance,
theTenebrionidaeBolitophagus reticulatus isaubiquitousspeciesin
F. fomentarius fromEurope to Korea(Jung,Kim,&Kim, 2007), but
iscompletelyreplacedby itsrelative Bolitotherus cornutus inNorth
America(Matthewman &Pielou,1971),indicating thatthere might
OurresultsshowedthatfruitbodiesofasinglefungusF. fomen
tarius pro vide habi t attoahi ghn u mberofar t h ropo d s,th e rebycon
Conside ring the respon sibility of Europ ean countries t o protect
tion of bra cket fungi as F. fomentarius an inte grated goal of for
tweensites suggestthataprioritizationofcertainregionswithin
nitiesinF. fomentarius. Instead,werecommendthatconservation
should rangefromtheprotectionofforestswhere F. fomentarius
ishighlyabundantand inhabitedbyEurope‐widerarearthropod
reintroductionof thespecies to regions(e.g., in westernEurope)
methodssee Abrego etal., 2016).The example of the region of
Flanders,Belgium,showsthat F. fomentarius isabletorecolonize
ifbee chd eadwoo dan dha bita ttr eesare ret aine d(Vande ke r khove
totrack F. fomentarius populationsrecolonizingsuitablehabitats
due to their high dispe rsal ability ( Vandekerk hove et al., 2011;
Zytynska et al.,2018). In addition topositiveeffects onspecies
associated with its fruitbodies, promoting F. fomentarius will po
tentially help to restore fundamental ecosys tem processes and
natural forest dynamicsinbeechforests as itisthe primaryde
composerofbeech woodand an importantagent of treesenes
cence and de ath. Species a ssociated with b roadleaf dead wo od
andsunnycondi tio nsi nfores t smaya lso benefitfro mga pscreated
when beech trees are killed byF. fomentarius. As F. fomentarius
ecosys tem processes, it c an be considered a key stone modifie r
Wearegratefultoallthosewhohelpe dinthef iel da ndinthelabo
rator y to conduct the s tudy, Svitlana Los for p roviding sampl es
Morinière for laboratory support in DNA barcoding. We thank
Emily Kilham for linguistic revision of the manuscript. Nicolas
Friessreceivedascholarship fromthe RudolfandHeleneGlaser
Foundation organized in the “Stifter verband für die deutsche
Wissenschaf t.”M.MikošandM.Svobodaweresupportedbythe
Czech Universityof LifeSciences, Prague (CIGANo. 20184304)
andbythe institutional project MSMT CZ.02.1.01/0.0/0.0/16_0
Nicolas Friess‐0003‐0517‐3798
Jörg C. Müller‐0002‐14091586
Simon Thorn‐0002‐3062‐3060
Sebastian Seibold‐0002‐7968‐4489
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Nicolas Friessresearchfocusesonlocaltolargescalepat
Jörg C. Müller's re search focuses on forest biodiversity, from
ecological mechanisms toconservation strategies intemperate
Sebastian Seibold's research focuses on the conservation of
biodiversity in forest ecosystems, particularly associated with
dead wood , and the impor tance of biodi versity for eco system
Additional supporting information may be found online in the
How to cite this article:FriessN,MüllerJC,AramendiP,etal.
beechforests.Divers Distrib. 2019;25:783–796. ht tps://d o i .
org /10.1111/ddi.12882
... [37,38]), but usually within one taxonomic group or trophic guild at a time. The studies that have considered all arthropods are either exploratory, i.e. not inferring community patterns based on host traits [17,26,36], or limited to one or two fungal hosts [25,35,39]. However, to get a solid understanding of the effects of host traits in the fungi-arthropod system, we need to assess arthropod communities across different hosts. ...
... The most common families we identified consist mostly (Bolitophilidae, Mycetophilidae and Oppidae) or partly (Chironomidae and Cecidomyiidae) of fungivorous species [27,79,80]. However, fungivores may not necessarily be dominant, as our annotations are largely unresolved at the species level and predators could make up to a third of the faunal composition in fruit bodies [25,39]. At the order level, true flies and oribatid mites were dominant, which we expected based on earlier rearingbased studies on living fruit bodies of wood-decay fungi [17,26,27,34]. ...
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Biological communities within living organisms are structured by their host's traits. How host traits affect biodiversity and community composition is poorly explored for some associations, such as arthropods within fungal fruit bodies. Using DNA metabarcoding, we characterized the arthropod communities in living fruit bodies of 11 wood-decay fungi from boreal forests and investigated how they were affected by different fungal traits. Arthropod diversity was higher in fruit bodies with a larger surface area-to-volume ratio, suggesting that colonization is crucial to maintain arthropod populations. Diversity was not higher in long-lived fruit bodies, most likely because these fungi invest in physical or chemical defences against arthropods. Arthropod community composition was structured by all measured host traits, namely fruit body size, thickness, surface area, morphology and toughness. Notably, we identified a community gradient where soft and short-lived fruit bodies harboured more true flies, while tougher and long-lived fruit bodies had more oribatid mites and beetles, which might reflect different development times of the arthropods. Ultimately, close to 75% of the arthropods were specific to one or two fungal hosts. Besides revealing surprisingly diverse and host-specific arthropod communities within fungal fruit bodies, our study provided insight into how host traits structure communities.
... So far, there have been rather few studies on the relationships between a whole set of TreMs and the abundance and diversity of forest-dwelling species. For example, birds' and bats' species richness was positively related to TreM diversity , while invertebrates showed positive associations with specific TreMs, like fruiting bodies of fungi or cavities (Friess et al., 2019;. Therefore, a better understanding of the links between a comprehensive set of TreMs and forest dwelling taxa may help to identify valuable habitat trees and provide a foundation for monitoring of some aspects of forest biodiversity. ...
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Forests are undergoing through a slow but steady process of forest area change, which happen to be different across countries. Predictions of future scenarios highlight the role of agriculture in shaping land-use change in Europe, with possible local benefits for forests. Thus, the role of management in shaping the forests of the future will increase according to the World’s demand for timber, clean water, biodiversity, clean air and other ecosystem services. Research is testing and developing new management strategies to halt completely biodiversity loss. The threats faced by biodiversity in managed forests constitute the challenges of future forest ecological research. The response of birds to environmental changes is a consequence of the close-knit species-habitat relationships they evolved and are adapted to. Adaptations to the forest environment are structural, behavioural and physiological. The alteration of the forest structures and their spatial allocation across landscapes creates threats to bird populations that need to be taken into account in conservation-oriented forest management. The present thesis aims to assess the numerical response of the bird assemblage to forest structures and its relevance for management. The general aim is focused on close-to-nature forest management and on the retention of structural elements as a conservation tool. The numerical response of the bird assemblage is investigated in terms of species diversity and abundance. The general aim is achieved by focusing on the following research questions: i. Which forest and landscape characteristics best explain the abundance and diversity of forest birds? ii. Which management practices can be developed for the conservation of birds in managed forests? This thesis is carried out within the framework of the ConFoBi Research Training Group (Conservation of Forest Biodiversity in multiple-use landscape of Central Europe). ConFoBi assesses whether and how structural retention measures contribute to the conservation of forest biodiversity in multiple-use landscapes of the temperate zone. The research programme of the present thesis has been implemented in the southern Black Forest, Germany, as a model system for temperate forests. The Black Forest is a forest-dominated low mountain range within a multiple-use landscape typical of central Europe, where 135 quadratic 1-ha-plots where selected within the forested areas. Plots were selected along two major environmental gradients: forest vertical complexity, represented by the number of standing dead trees found in each plot; landscape composition, represented by the amount of forest cover within 25 km2 surrounding each plot. All plots were inventoried by measuring the diameter of every tree (above 7 cm diameter at breast height) and the amounts of laying and standing dead wood. Because habitat trees are an important structural element in retention approaches, for each study plot the 15 largest trees were mapped and their existing microhabitats (such as hollows, cracks, and cavities) quantified. Landscape patterns were described using a range of metrics, including amount of forest cover and landscape metrics. Bird data were collected by employing point counts performed at the centre of each plots and repeated two or three times per sampling season (2017-2018). Abundance and diversity of birds were analysed using hierarchical N-mixture models and generalized linear (mixed) models, respectively. The results highlight that current retention practices are well below the retention levels needed by the bird assemblage to have abundance and diversity similar to those of unharvested forests, assessing the levels around 40-60% of retained trees. Retention practices can, however, be improved by retaining trees of higher ecological importance. Trees with a diameter 20 cm larger than the surrounding trees, with cavities, rot holes or concavities can potentially fulfill the habitat requirments of a large array of species, including woodpeckers and the related cavity-users community. The presence of those trees across the landscape can ensure the continuity of resources, while favouring more natural tree species composition will benefit bird species diversity. The response of the entire bird assemblage to the configuration of forest patches indicates that in landscapes with less aggregated forest patches, the abundance of the bird assemblage can potentially be lower than in landscapes with closer forest patches. I conclude that the establishment and assemble of the bird community is a hierarchical spatial process. Small environmental elements determine the occurrence and observed abundance of populations. The location of such elements determines which species and how many individuals of each species are found across the landscape. Over the entire landscape, the amount of available habitat and its configuration determines how many species can be found at a given site. Therefore, I suggest a hierarchical set of actions should be considered, each benefiting birds at different levels of biological organization and spatial scale. Actions to preserve forest area and avoid fast land-use changes should have higher priority. The provisioning of large trees and deadwood would then ensure that the forests retain those structures and functions that support the establishment of bird populations in optimal habitats.
... For example, the presence of perennial fungal fruiting bodies on European beech (Fagus sylvatica) very likely refers to the tinder fungus (Fomes fomentarius). The fruiting bodies of the tinder fungus was found to serve as habitat for up to 600 different arthropod species (Friess et al., 2019). Whereas the presence of perennial fungal fruiting bodies on silver fir (Abies alba) may refer to Phellinus hartigii, which presumably interacts with different arthropods compared to the tinder fungus (cf. ...
Habitat trees – one of the key elements of integrative retention approaches in European forests – are increasingly studied regarding their benefits to forest biodiversity. In this regard, treerelated microhabitats (TreMs) and deadwood serve as indicators of forest biodiversity. Specific recommendations of amounts of deadwood to be retained in managed forest stands to support related taxa are available, but not for TreMs. Retention of habitat trees aims to increase availability of TreMs in managed forests. To be able to refine selection criteria for habitat trees, I focused in this thesis on factors affecting TreMs on living trees. Therefore, I calculated species specific diameter thresholds bases on TreM occurrences, I investigated the effect of tree maintenance on TreMs in urban areas and I evaluated the short-term value of the retention of habitat tree groups. Chapter 1. The aim of this chapter was to explain TreM occurrence from a qualitative perspective by considering their diversity. Tree diameter at breast height (dbh), tree species, and canopy class were useful predictors of TreM diversity. TreM diversity on broad-leaved trees was on average higher than in conifers of the same diameter. In contrast to TreM abundance their diversity saturated towards higher dbh levels. Those TreM saturation levels were used to derive diameter thresholds. Habitat trees support not only more, but also more diverse, microhabitats in comparison to crop trees. Chapter 2. In this chapter I further developed the findings of chapter one and focused on the calculation of species specific diameter thresholds to precise recommendations for selecting habitat trees. Based on the relation between TreMs and tree diameter as well as TreMs and species I derived diameter thresholds for 18 European tree species (13 broad-leaved, 5 coniferous). Those thresholds refer to statistically disproportionate high levels of TreM richness or abundance. Complementing other aspects that need to be considered during habitat tree selection processes in managed forest stands, I recommend to select habitat trees with or close to a dbh of 70 cm for broadleaves and 86 cm for conifers. The differences of dbh thresholds between broadleaves and conifers as well as between species indicate species specific TreM dynamic. Chapter 3. In the third chapter, I investigated the TreM availability on urban trees along a maintenance gradient in Montréal, Canada. Intensive tree maintenance in urban trees led to levels of certain microhabitats such as cavities and injuries that were comparable to natural, unmanaged forests. Light maintenance of urban trees encouraged more crown deadwood than typical and intensive maintenance levels. My results underline the importance of conserving and maintaining large living trees, especially in urban areas to provide tree microhabitats. These results also demonstrate the important role of intensive tree maintenance in stimulating tree microhabitat development in urban areas. Chapter 4. Here, I addressed the effectiveness of habitat tree group (HTG) retention in forests of Baden-Württemberg, Germany, 10 years after the introduction of the approach. Large living trees (LLTs), standing deadwood and TreMs were significantly more abundant in HTGs than in reference plots. When retaining 5 % of a forest stand area as HTG, old-growth attributes increased significantly at the stand scale: amount of LLTs doubled and its volume almost tripled, and standing deadwood increased on average by 25 %. However, quantities of both attributes remain below recommended minimum thresholds. Retaining 5 % of stand area in HTG had a significantly positive effect on woodpecker cavities, rot holes and exposed heartwood, whereas 15 to 25 % area in HTGs would be required to increase stand level abundance of concavities, exposed sapwood or crown deadwood significantly. Retention of HTGs enriched managed, multiple-use forests with old growth structural attributes. Yet, the selection of HTGs could be made more efficient by focusing on forest patches with high tree volume or low tree density and by further considering snags, tree species mixture and LLTs as well as less vital trees. Overall the findings of this thesis suggest, that common tree attributes (species, diameter, vitality, canopy class, life status) can be used to predict the occurrence of TreMs. Species specific diameter thresholds can help to identify trees with higher levels of TreM richness. The specification of tree attributes in regard to TreMs allow to optimize habitat tree selection procedures. In addition or absence of trees rich in TreMs, tree maintenance could increase structural diversity. Intensively maintained trees providing comparable amounts of TreMs to trees from long-term unmanaged forests emphasize the relevance of artificially induced structures. To consider a holistic view on trees as biodiversity relevant feature throughout the landscape, I propose to expand further TreM research to urban and rural trees. Regardless how TreMs evolved and in which landscape they occur, the relation between TreMs and related taxa needs to be specified to strengthen the biodiversity indicating function of TreMs. Results from current TreM inventories, that followed standardized procedures, allow a approximation of TreM occurrence on a landscape level by linking species and diameter information to observed TreM abundances. To better understand development and persistence of TreMs repeated inventories were needed. Finally, my results allow to refine selection criteria of habitat trees based on the presence of TreMs and the consideration of species specific diameter thresholds. Furthermore, in absence of TreMs or when minimum diameters were not reached, I propose the possibility of artificial TreM creation to be useful for structural enrichment.
... Wood-decaying fungi generally require high volumes of necromass, sometimes exceeding 300 m 3 /ha [23]. Many saproxylic beetles are linked to wood-decaying fungi [7,17,61,62]. The solar influence is essential for the abundance and diversity of saproxylic beetles [28,63,64], and at the same time, sun exposure can compensate for the amount of deadwood [23]. ...
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Due to traditional forest management, the primary goal of which is the production of raw wood material, commercial forest stands are characterized by low biodiversity. At the same time, commercial forests make up the majority of forests in the Central European region, which means a significant impact on the biodiversity of the entire large region. Saproxylic species of organisms are a frequently used criterion of biodiversity in forests. Based upon the analysis of 155 scientific works, this paper defines the fundamental attributes of the active management supporting biodiversity as well as the preservation of the production function. Using these attributes, a model management proposal was created for three tree species, which takes into account the results of research carried out in the territory of the University Forest Enterprise of the Czech University of Life Sciences Prague, since 2019. The optimum constant volume of deadwood in commercial stands was set at 40–60 m3/ha, 20% of which should be standing deadwood. The time framework is scheduled for an average rotation period of the model tree species, while the location of deadwood and frequency of enrichment must comply with the rate of decomposition, the requirement for the bulkiest dimensions of deadwood possible, and the planned time of tending and regeneration operations in accordance with the models used in the Czech Republic. The goal of active management is to maintain the continuity of suitable habitats for sensitive and endangered species. The estimates of the value of retained wood for decomposition can be as high as 45–70 EUR/ha/year for spruce and beech, and about 30 EUR /ha/year for oak.
... For many studies, the area was one part of a larger study (e.g. Bouget Friess et al. 2019;Courbaud et al. 2017;Müller et al. 2015;Larrieu and Cabanettes 2012). However, this forest was sometimes specifically studied for impact of harvesting on soil chemistry (Larrieu et al. 2006), on hoverfly communities , on both deadwood and tree-related microhabitats amount (Larrieu et al. 2012), or on beetle communities (Brin et al. 2010). ...
... Positive correlations between TreMs and several taxa such as bats, birds and to a lesser extent (saproxylic) insects have been shown in earlier studies Basile et al. 2020a), and TreMs are considered valuable biodiversity indicators (Gao et al. 2015). Specific correlations between taxa and TreMs have, for instance, been reported for rare aquatic organisms in water-filled holes in trees (Gossner et al. 2016), arthropod species inhabiting conks of tree-decaying fungi (Friess et al. 2019), as well as cavity nesting birds in tree hollows (Puverel et al. 2019). An overview of associations between TreMs and taxa has been provided by Larrieu et al. (2018). ...
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Habitat trees, which provide roosting, foraging and nesting for multiple taxa, are retained in managed forests to support biodiversity conservation. To what extent their spatial distribution influences provisioning of habitats has rarely been addressed. In this study, we investigated whether abundance and richness of tree-related microhabitats (TreMs) differ between habitat trees in clumped and dispersed distributions and whether the abundance of fifteen groups of TreMs is related to tree distribution patterns. To identify habitat trees, we quantified TreMs in temperate mountain forests of Germany. We determined clumping (the Clark-Evans index), size of the convex hull, diameter at breast height, as well as altitude, slope and aspect of sites for their possible influence on TreMs. We additionally determined the difference in TreM abundance and richness among four options of selecting five habitat trees per ha from 15 candidates: (a) the most clumped trees, (b) five randomly selected and dispersed trees, (c) the single tree with highest abundance or richness of TreMs and its four closest neighbors and (d) a "reference selection" of five trees with known highest abundance or richness of TreMs irrespective of their distribution. The degree of clumping and the size of the convex hull influenced neither the abundance nor richness of TreMs. The reference selection, option (d), contained more than twice the number of TreMs compared to the most clumped, (a), or random distributions, (b), of five habitat trees, while option (c) assumed an intermediate position. If the goal of habitat tree retention is to maximize stand-level abundance and richness of TreMs, then it is clearly more important to select habitat trees irrespective of their spatial pattern.
Interactions among fungi and insects involve hundreds of thousands of species. While insect communities on plants have formed some of the classic model systems in ecology, fungus‐based communities and the forces structuring them remain poorly studied by comparison. We characterize the arthropod communities associated with fruiting bodies of eight mycorrhizal basidiomycete fungus species from three different orders along a 1200‐km latitudinal gradient in northern Europe. We hypothesized that—matching the pattern seen for most insect taxa on plants—we would observe a general decrease of fungal‐associated species with latitude. Against this backdrop, we expected local communities to be structured by host identity and phylogeny, with more closely related fungal species sharing more similar communities of associated organisms. As a more unique dimension added by the ephemeral nature of fungal fruiting bodies, we expected further imprints generated by successional change, with younger fruiting bodies harboring communities different from older ones. Using DNA metabarcoding to identify arthropod communities from fungal fruiting bodies, we find that latitude leaves a clear imprint on fungus‐associated arthropod community composition, with host phylogeny and decay stage of fruiting bodies leaving lesser but still‐detectable effects. The main latitudinal imprint is on a high arthropod species turnover, with no detectable pattern in overall species richness. Overall, these findings paint a new picture of the drivers of fungus‐associated arthropod communities, suggesting that latitude will not affect how many arthropod species inhabits a fruiting body, but rather what species occur in it and at what relative abundances (as measured by sequence read counts). These patterns upset simplistic predictions regarding latitudinal gradients in species richness and in the strength of biotic interactions.
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Owing to climate change, natural forest disturbances and consecutive salvage logging are drastically increasing worldwide, consequently increasing the importance of understanding how these disturbances would affect biodiversity conservation and provision of ecosystem services. In chapter II, I used long-term water monitoring data and mid-term data on α-diversity of twelve species groups to quantify the effects of natural disturbances (windthrow and bark beetle) and salvage logging on concentrations of nitrate and dissolved organic carbon (DOC) in streamwater and α-diversity. I found that natural disturbances led to a temporal increase of nitrate concentrations in streamwater, but these concentrations remained within the health limits recommended by the World Health Organization for drinking water. Salvage logging did not exert any additional impact on nitrate and DOC concentrations, and hence did not affect streamwater quality. Thus, neither natural forest disturbances in watersheds nor associated salvage logging have a harmful effect on the quality of the streamwater used for drinking water. Natural disturbances increased the α-diversity in eight out of twelve species groups. Salvage logging additionally increased the α-diversity of five species groups related to open habitats, but decreased the biodiversity of three deadwood-dependent species groups. In chapter III, I investigated whether salvage logging following natural disturbances (wildfire and windthrow) altered the natural successional trajectories of bird communities. I compiled data on breeding bird assemblages from nine study areas in North America, Europe and Asia, over a period of 17 years and tested whether bird community dissimilarities changed over time for taxonomic, functional and phylogenetic diversity when rare, common and dominant species were weighted differently. I found that salvage logging led to significantly larger dissimilarities than expected by chance and that these dissimilarities persisted over time for rare, common and dominant species, evolutionary lineages, and for rare functional groups. Dissimilarities were highest for rare, followed by common and dominant species. In chapter IV, I investigated how β-diversity of 13 taxonomic groups would differ in intact, undisturbed forests, disturbed, unlogged forests and salvage-logged forests 11 years after a windthrow and salvage logging. The study suggests that both windthrow and salvage logging drive changes in between-treatment β-diversity, whereas windthrow alone seems to drive changes in within-treatment β-diversity. Over a decade after the windthrow at the studied site, the effect of subsequent salvage logging on within-treatment β-diversity was no longer detectable but the effect on between-treatment β-diversity persisted, with more prominent changes in saproxylic groups and rare species than in non-saproxylic groups or common and dominant species. Based on these results, I suggest that salvage logging needs to be carefully weighed against its long-lasting impact on communities of rare species. Also, setting aside patches of naturally disturbed areas is a valuable management alternative as these patches would enable post-disturbance succession of bird communities in unmanaged patches and would promote the conservation of deadwood-dependent species, without posing health risks to drinking water sources.
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The geographical distributions of species associated with European temperate broadleaf forests have been significantly influenced by glacial–interglacial cycles. During glacial periods, these species persisted in Mediterranean and extra-Mediterranean refugia and later, during interglacial periods, expanded northwards. The widespread saproxylic beetle Bolitophagus reticulatus depends closely on European temperate broadleaf forests. It usually develops in the tinder fungus Fomes fomentarius, a major decomposer of broadleaf-wood. We sampled B. reticulatus in sporocarps from European beech (Fagus sylvatica) and Oriental beech (Fagus orientalis) across Europe and the Caucasus region. We analysed mitochondrial gene sequences (cox1, cox2, cob) and 17 microsatellites to reconstruct the geographical distribution of glacial refugia and postglacial recolonization pathways. We found only marginal genetic differentiation of B. reticulatus, except for a significant split between populations of the Caucasus region and Europe. This indicates the existence of past refugia south of the Great Caucasus, and a contact zone with European populations in the Crimean region. Further potential refugia might have been located at the foothills of the Pyrenees and in the Balkan region. Our genetic data suggest a phalanx-wise recolonization of Europe, a reflection of the high mobility of B. reticulatus.
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C’est article constitue un retour d’expérience de gestion à la fois intégrative et adaptative, menée depuis 2005 sur les 17 000 ha d’une forêt de l’Etat de Bavière. L’entreprise forestière d’Etat Ebrach gère environ 17 000 ha de une forêt domaniale dominée par le Hêtre, dans la région du Steigerwald, en Bavière.. Elle a adopté une gestion à double approche, à la fois intégrative et adaptative. Tout en générant environ un million d’euros de bénéfice annuel, ce type desa gestion a pour objectif d’optimiser un large panel de services écosystémiques et d’accroître significativement la capacité d’accueil pour la biodiversité forestière, en particulier les espèces saproxyliques. Après une présentation générale de la forêt et de son histoire, les concepts qui régissent sa gestion sont présentés. Une analyse économique détaillée permet également de quantifier le coût de ce type de gestion. Nous espérons que cet article inspirera les forestiers français soucieux d’améliorer la durabilité de leur gestion.
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1.The successional dynamics of forests – from canopy openings to regeneration, maturation and decay – influence the amount and heterogeneity of resources available for forest‐dwelling organisms. Conservation has largely focused only on selected stages of forest succession (e.g. late‐seral stages). However, to develop comprehensive conservation strategies and to understand the impact of forest management on biodiversity, a quantitative understanding of how different trophic groups vary over the course of succession is needed. 2.We classified mixed mountain forests in Central Europe into nine successional stages using airborne LiDAR. We analysed α‐ and β‐diversity of six trophic groups encompassing approximately 3,000 species from three kingdoms. We quantified the effect of successional stage on the number of species with and without controlling for species abundances and tested whether the data fit the more‐individuals hypothesis or the habitat heterogeneity hypothesis. Furthermore, we analysed the similarity of assemblages along successional development. 3.The abundance of producers, first‐order consumers and saprotrophic species showed a U‐shaped response to forest succession. The number of species of producer and consumer groups generally followed this U‐shaped pattern. In contrast to our expectation, the number of saprotrophic species did not change along succession. When we controlled for the effect of abundance, the number of producer and saproxylic beetle species increased linearly with forest succession, whereas the U‐shaped response of the number of consumer species persisted. The analysis of assemblages indicated a large contribution of succession‐mediated β‐diversity to regional γ‐diversity. 4. Synthesis and applications. Depending on the species group, our data supported both the more‐individuals hypothesis and the habitat heterogeneity hypothesis. Our results highlight the strong influence of forest succession on biodiversity and underline the importance of controlling for successional dynamics when assessing biodiversity change in response to external drivers such as climate change. The successional stages with highest diversity (early and late successional stages) are currently strongly underrepresented in the forests of Central Europe. We thus recommend that conservation strategies aim at a more balanced representation of all successional stages. This article is protected by copyright. All rights reserved.
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The vegan package provides tools for descriptive community ecology. It has most basic functions of diversity analysis, community ordination and dissimilarity analysis. Most of its multivariate tools can be used for other data types as well. The functions in the vegan package contain tools for diversity analysis, ordination methods and tools for the analysis of dissimilarities. Together with the labdsv package, the vegan package provides most standard tools of descriptive community analysis. Package ade4 provides an alternative comprehensive package, and several other packages complement vegan and provide tools for deeper analysis in specific fields. Package provides a Graphical User Interface (GUI) for a large subset of vegan functionality. The vegan package is developed at GitHub ( GitHub provides up-to-date information and forums for bug reports. Most important changes in vegan documents can be read with news(package="vegan") and vignettes can be browsed with browseVignettes("vegan"). The vignettes include a vegan FAQ, discussion on design decisions, short introduction to ordination and discussion on diversity methods. A tutorial of the package at provides a more thorough introduction to the package. To see the preferable citation of the package, type citation("vegan").
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A central hypothesis of ecology states that regional diversity influences local diversity through species-pool effects. Species pools are supposedly shaped by large-scale factors and then filtered into ecological communities, but understanding these processes requires the analysis of large datasets across several regions. Here, we use a framework of community assembly at a continental scale to test the relative influence of historical and environmental drivers, in combination with regional or local species pools, on community species richness and community completeness. Using 42,173 vegetation plots sampled across European beech forests, we found that large-scale factors largely accounted for species pool sizes. At the regional scale, main predictors reflected historical contingencies related to post-glacial dispersal routes, whereas at the local scale, the influence of environmental filters was predominant. Proximity to Quaternary refugia and high precipitation were the main factors supporting community species richness, especially among beech forest specialist plants. Models for community completeness indicate the influence of large-scale factors, further suggesting community saturation as a result of dispersal limitation or biotic interactions. Our results empirically demonstrate how historical factors complement environmental gradients to provide a better understanding of biodiversity patterns across multiple regions.
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Identification of forest stands with priority for the conservation of biodiversity is of particular importance in landscapes with a long cultural and agricultural history, such as Central Europe. A group of species with a high indicator value for the naturalness of forest ecosystems are saproxylic insects. Some of these species, especially within the order Coleoptera, have been described as primeval forests relicts. Here, we compiled a list of 168 “primeval forest relict species” of saproxylic beetles based on expert knowledge. These species can serve as focal and umbrella species for forest conservation in Central Europe. They were selected because of their dependence on the continuous presence of primeval forest habitat features, such as over-mature trees, high amounts of dead wood, and dead wood diversity, as well as their absence in managed Central European forests. These primeval forest relict species showed a moderately strong clumping pattern within the phylogeny of beetles, as indicated by phylogenetic signal testing using the D-statistic. When we controlled for phylogenetic relatedness, an ordinal linear model revealed that large body size and preference for dead wood and trees of large diameter are the main characteristics of these species. This list of species can be used to identify forest stands of conservation value throughout Central Europe, to prioritize conservation and to raise public awareness for conservation issues related to primeval forests.
Fulltext Share Link (until December 15, 2018): *** Abstract: Forests are under pressure from accelerating global change. To cope with the multiple challenges related to global change but also to further improve forest management we need a better understanding of (1) the linkages between drivers of ecosystem change and the state and management of forest ecosystems as well as their capacity to adapt to ongoing global environmental changes, and (2) the interrelationships within and between the components of forest ecosystems. To address the resulting challenges for the state of forest ecosystems in Central Europe, we suggest 45 questions for future ecological research. We define forest ecology as studies on the abiotic and biotic components of forest ecosystems and their interactions on varying spatial and temporal scales. Our questions cover five thematic fields and correspond to the criteria selected for describing the state of Europe’s forests by policy makers, i.e. biogeochemical cycling, mortality and disturbances, productivity, biodiversity and biotic interactions, and regulation and protection. We conclude that an improved mechanistic understanding of forest ecosystems is essential for the further development of ecosystem-oriented multifunctional forest management in the face of accelerating global change.
Habitat restoration aims to improve local habitat conditions for threatened species. While such restorations are widespread, rigorous evaluations of their success are rare. This is especially true of those considering species dynamics. Increasingly, deadwood is a target for forest restoration as many species directly and indirectly depend on this resource. In a broadleaf forest in southern Germany, we explored the effect of landscape-wide deadwood restoration on the population genetic structure of the specialist fungus-dwelling saproxylic beetle Bolitophagus reticulatus. Before 2003, the northern area of this forest was intensively logged for more than half a century, while the southern part was less intensively managed. This drove populations of the host fungus Fomes fomentarius, and consequently the beetle, to local extinction in the northern part. Only after the first decade of restoration were both the fungus and its beetles present across all areas of the forest. Using 17 newly developed microsatellite loci, we show that these beetles exhibit population genetic structuring, mainly influenced by the north-south divide. However, the low degree of isolation-by-distance, and the low relatedness of beetles collected from the same trees or fungus, shows that strong dispersal ability is facilitating the recolonisation of these forests on the scale of tens of kilometers. In another 10 years, it is likely that the population will show even less genetic structuring. Synthesis and applications. Through the recolonisation of the fungus Fomes fomentarius and the fungus-dwelling beetle Bolitophagus reticulatus after deadwood restoration, we demonstrate that, while there are many discussions on the optimal spatial distribution of deadwood, just the presence of deadwood can be sufficient to enable recolonisations of specific species. As long as some relict populations of these species are embedded in a once intensively managed forest, increased deadwood (amount and diversity) anywhere will benefit recolonisation of the habitat. However, increasing deadwood diversity should also be encouraged to benefit even more species.
Isolation of habitats in space and time affects species globally and in a multitude of ecosystems. It is however often difficult to assess the level of isolation from the point of view of the focal species. Indicator species are often used to assess 'conservation value' of habitats. One such approach involves the use of wood-decaying fungal species as indicators of near-natural forests, continuity in old-forest characteristics over time, and/or presence of red-listed species, but not of spatial connectivity. The indicator species were commonly assigned based on expert opinions, but few scientific evaluations have been performed of what these species actually indicate. Building on previous classifications of wood-decaying fungal indicator species on Norway spruce, we hypothesized that indicator species would differ from non-indicator species in how they respond to local temporal connectivity (forest age, the intensity of historical selective logging and the presence of well-decomposed large logs), and to local and landscape-scale spatial connectivity (local forest area, local amount of deadwood and the connectivity to old forest in the surrounding landscape). Based on fungal occurrence data from a fixed number of spruce logs at 28 sites distributed across the Scandinavian Peninsula, we explored the spatiotemporal scales at which the local communities were affected by connectivity. Indicator species showed the strongest response to connectivity of old forest (≥80 years) within 100 km, while non-indicator species depended on connectivity of younger forest (≥40 years) at a smaller spatial scale (≤25 km). Indicator species increased and non-indicator species decreased in total abundance with the increasing age of the local forest stand. Landscape-scale old-forest connectivity was beneficial for indicator species in all sites except those with relatively low amount of deadwood, while non-indicator species showed the opposite pattern. We identify a threshold of around 29 m 3 ha −1 in the amount of spruce logs where indicator species become abundant enough to influence non-indicator species through competitive interactions. There was a pronounced uniformity within each species group in the connectivity responses. We conclude that the studied indicator species indicate high forest age, high amount of resources and, given that the resources are plentiful, also high old-forest connectivity, but they do not indicate a long history without any logging operations or local deadwood continuity. The studied non-indicator species did not indicate any of the studied spatiotemporal connectivity variables. Indicator species are usually red-listed and may continue to decline in the future without habitat restoration efforts.
Limited knowledge of dispersal for most organisms hampers effective connectivity conservation in fragmented landscapes. In forest ecosystems, deadwood-dependent organisms (i.e., saproxylics) are negatively affected by forest management and degradation globally. We reviewed empirically established dispersal ecology of saproxylic insects and fungi. We focused on direct studies (e.g., mark-recapture, radiotelemetry), field experiments, and population genetic analyses. We found 2 somewhat opposite results. Based on direct methods and experiments, dispersal is limited to within a few kilometers, whereas genetic studies showed little genetic structure over tens of kilometers, which indicates long-distance dispersal. The extent of direct dispersal studies and field experiments was small and thus these studies could not have detected long-distance dispersal. Particularly for fungi, more studies at management-relevant scales (1-10 km) are needed. Genetic researchers used outdated markers, investigated few loci, and faced the inherent difficulties of inferring dispersal from genetic population structure. Although there were systematic and species-specific differences in dispersal ability (fungi are better dispersers than insects), it seems that for both groups colonization and establishment, not dispersal per se, are limiting their occurrence at management-relevant scales. Because most studies were on forest landscapes in Europe, particularly the boreal region, more data are needed from nonforested landscapes in which fragmentation effects are likely to be more pronounced. Given the potential for long-distance dispersal and the logical necessity of habitat area being a more fundamental landscape attribute than the spatial arrangement of habitat patches (i.e., connectivity sensu strict), retaining high-quality deadwood habitat is more important for saproxylic insects and fungi than explicit connectivity conservation in many cases.
Tree related Microhabitats (hereafter TreMs) have been widely recognized as important substrates and structures for biodiversity in both commercial and protected forests and are receiving increasing attention in management , conservation and research. How to record TreMs in forest inventories is a question of recent interest since TreMs represent potential indirect indicators for the specialized species that use them as substrates or habitat at least for a part of their life-cycle. However, there is a wide range of differing interpretations as to what exactly constitutes a TreM and what specific features should be surveyed in the field. In an attempt to harmonize future TreM inventories, we propose a definition and a typology of TreM types borne by living and dead standing trees in temperate and Mediterranean forests in Europe. Our aim is to provide users with definitions which make unequivocal TreM determination possible. Our typology is structured around seven basic forms according to morphological characteristics and biodiversity relevance: i) cavities lato sensu, ii) tree injuries and exposed wood, iii) crown deadwood, iv) excrescences, v) fruiting bodies of saproxylic fungi and fungi-like organisms, vi) epiphytic and epixylic structures, and vii) exudates. The typology is then further detailed into 15 groups and 47 types with a hierarchical structure allowing the typology to be used for different purposes. The typology, along with guidelines for standardized recording we propose, is an unprecedented reference tool to make data on TreMs comparable across different regions, forest types and tree species, and should greatly improve the reliability of TreM monitoring. It provides the basis for compiling these data and may help to improve the reliability of reporting and evaluation of the conservation value of forests. Finally, our work emphasizes the need for further research on TreMs to better understand their dynamics and their link with biodiversity in order to more fully integrate TreM monitoring into forest management.