©EntomologicaFennica.29August2017 Springtails (Collembola) in meadows, pastures and road verges in Central Finland AtteKomonen*&SaanaKataja-aho Komonen,A.&Kataja-aho,S.2017:Springtails(Collembola)inmeadows,pas- turesandroadvergesinCentralFinland.—Entomol.Fennica28:157–163. Understandingofspeciesdistribution,abundanceandhabitataffinitiesiscrucial for red-list assessment, conservation and habitat management. In Central Fin- land,westudiedCollembolainthreehabitattypes,namelynon-grazedmeadows, pasturesandroadvergesusingpitfalltraps.Altogether,9,630Collembolaindi- vidualswererecorded.Thesebelongedto12families,34generaand60species. Thenumberofspecimenswasclearlyhigherinmeadowsthaninpasturesorroad verges.Thenumberofspecies,however,washigherinmeadowsandroadverges (40and39species,respectively)thaninpastures(33species).Theoverallspe- ciesnumberiscomparabletootherlarge-scalesamplingschemesinsimilarhabi- tats.Werecordedafewabundantspecies(Spatulosminthurusflaviceps,Smin- thurus viridis and Sminthurus nigromaculatus) that have been previously re- cordedfromverydifferentbiotopes.Inconclusion,biodiversityinventoriesof soilfauna,aswellasotherbiota,shouldalsoincludemarginalhabitats,whichof- tenhostpeculiarcommunities. A. Komonen, University of Jyväskylä, Department of Biological and Environ- mentalScience,P.O.Box35,FI-40014UniversityofJyväskylä,Finland;*Cor- respondingauthor’se-mail:[email protected] S.Kataja-aho,UniversityofJyväskylä,NaturalHistoryMuseum,P.O.Box35, FI-40014UniversityofJyväskylä,Finland;E-mail:[email protected] Received15November2016,accepted22December2016 1.Introduction sufficient and geographically limited (Pomme- resche&Løes2014);forexample,very littleis Understandingofspeciesdistribution,abundance known about soil fauna in traditional rural bio- andhabitataffinitiesindifferentenvironmentsis topes,suchaspasturesandmeadows(Huhtaetal. crucialforred-listassessment,conservationand 2010). habitat management. Unfortunately, such infor- Springtails (Collembola) are an important mationislackingfor many invertebrategroups. component of soil meso- and macrofauna, be- InFinland,forestsoilfaunahasbeenratherwell- cause they influence soil decomposition pro- studied,partlyduetoitsimportanceandsensitiv- cesses (Rusek 1975, Petersen 1994, Hopkin itytosilviculture(Huhtaetal.1967,1986,Siira- 1997). Springtails can be abundant and diverse Pietikäinenetal.2001,Huhta&Räty2005,Kata- even in highly modified and disturbed habitats ja-ahoetal.2016).Atthesametime,theknowl- such as agricultural fields (Lagerlöf & Andrén edgeofsoilfaunainnon-forestedhabitatsisin- 1991, Bitzer et al. 2005, Pommeresche & Løes 158 Komonen&Kataja-aho (cid:127) ENTOMOL.FENNICAVol.28 Table1.Studysitecharacteristicsandtotalnumberofindividualsandspecies.Collembolaweresampledin2014 mainlyfrom26th–30thofMayto18th–22ndofJune(=firstperiod).Ifspecimenswereidentifiedalsofromthesec- ondperiod(18th–22ndofJuneto7th–11thofJuly),thepoolednumbersofindividualsandspeciesforbothperiods aregiveninparentheses.Vegetationheightistheaverageoffivestudyplots. Habitat Area pH Soil Vegetation Indiv. Species Sites (ha) fractions height(cm) Meadows Peurala 0.1 5.16 coarsesilt 42 128(277) 11(17) Kirkonmäki 0.3 4.98 finesand 39 103 10 Syysniemi 0.4 4.62 coarsesilt 29 380 10 Vaaru 0.4 4.33 mediumsilt 33 182 10 Vuorela 0.5 4.60 coarsesilt 38 369 15 Liehu 0.5 4.16 coarsesilt 36 287 15 Riihimäki 0.9 4.61 mediumsilt 33 485 11 Karhukorpi 1.0 4.62 finesand 22 1,256 15 Vaateri 1.1* 4.67 coarsesilt 33 309 12 Hettee 2.0 4.45 finesand 22 256 8 Mäentalo 11.9* 4.93 finesilt 39 101(136) 7(7) Harju 32.5* 4.39 mediumsilt 24 78 8 Pastureswithcattle Pekkanen 1.7 4.27 finesand 30 314 7 Ohramaa 5.2 4.16 coarsesilt 30 213 7 Aatula 6.1 3.73 mediumsilt 15 114 6 Kivijärvi 7.6 4.70 coarsesilt 10 193 9 Pasturewithsheep/cattle Haapalehto 8.6 4.28 coarsesilt 14 75(174) 6(9) Pastureswithsheep Vaateri 1.1* 5.18 coarsesilt 39 352 9 Mäentalo 11.9* 4.65 finesilt 33 137(146) 17(17) Harju 32.5* 4.07 mediumsilt 22 217 7 Pastureswithhorses Koivulahti 1.1 4.24 finesand 15 174 7 Huusko 2.6 3.96 coarsesilt 12 524 13 Nuuttila 3.1 4.70 coarsesilt 20 301 13 Suuruspää 6.0 4.98 finesilt 21 52 7 Roadverges Myllypohja na 5.41 coarsesand 23 39(235) 7(17) Multia na 5.45 coarsesand 8 460 15 Petäjävesi na 5.65 coarsesand 10 45 8 Muurame na 5.99 finesand 35 13 3 Paateri na 5.66 coarsesand 19 5 2 Rotkola na 4.75 coarsesand 20 578 22 Klemettilä na 5.09 coarsesand 31 49 1 Valospohja na 4.63 coarsesand 27 106 11 Tammikoski na 5.14 coarsesand 42 21 2 Jukola na 4.65 coarsesand 11 553 14 Havulankangas na 5.22 coarsesand 21 280(386) 18(20) Norola na 5.43 finesand 23 287 12 *Pooledareaformeadowsandpasturesinagivenfarm. na:Notapplicable. 2014).However,differentvegetationtypes(Huh- canaffectcomposition,structureandsmallscale taetal.2005)andsoildisturbances(Kataja-ahoet spatial heterogeneity of collembolan communi- al.2016),aswellasdifferentlevelsofsoilcom- ties. paction(Larsenetal.2004),pollutionornutrient Manyenvironmentalfactorsarelikelytovary loads(Lagerlöf&Andrén1991,Songetal.2016) among meadows, pastures and road verges. For ENTOMOL.FENNICAVol.28 (cid:127) Collembolaintraditionalbiotopes 159 example, animal tramping affects soil compac- weretarmackedandbelongedtoFinnishroadcat- tion,whereasgrazingandexcretionsaffectvege- egory “Local road” or bigger, which ensured at tationandsoilchemistry(Oldén2016).Roadsalt least3mwideverges.Theselectedroadswereall andtrafficpollutioncaninfluencebiotaonroad constructeddecadesago,therewerenosignsof verges (Viard et al. 2004, Owojori et al. 2009). recentrenovations,andtheywereoutsideurban The response of Collembola on environmental areas.Theselectedvergeswereadjacenttoafo- variation can vary among euedaphic and epe- rest,ortoaruderalareaintwocases.InFinland, daphicspecies;euedaphicspeciesaresubsurface- roadvergesaremowedwithmachinery1–2times active,whereasepedaphicspeciesaresurface-ac- yearly;aftermachinemowingvegetationisgen- tive(Eisenbeis&Wichard1987). erally taller than in hand-mowed pastures. The Traditional rural biotopes have high conser- mostcommonplantspeciesinroadvergeswere vationvalue,withmanyrareandthreatenedplant Achilleamillefolium,Festucarubra,Taraxacum and invertebrate species. Their soil fauna, how- spp., Epilobium angustifolium, Hieracium um- ever, is much more poorly known. We studied bellatum, Hieracium vulgata group, Trifolium surface-active Collembola in non-grazed mead- repens, Anthriscus sylvestris, Betula pubescens ows,pasturesandroadvergesinCentralFinland andCerastiumfontanum. tobetterunderstandbiodiversityinthesevaluable habitats,aswellasgainmoreinformationabout thedistribution,abundanceandhabitataffinityof 2.2.Sampling Collembola. At each pasture and meadow, we established a random50mtransectthatranthroughthecenter 2.Materialsandmethods ofthesite.Inroadverges,transectsranalongthe roadandatleastonemeterfromtheedgeofthe 2.1.Studysites asphalt.Alongtransects,five2×2mplotswere placed at 10 m intervals. At two predetermined Thisstudywasconductedinsouthernandmiddle cornersoftheseplots,weplacedonepitfalltrap, boreal vegetation zone in Central Finland. i.e.therewere10pitfalltrapsineachsite.Allpit- Twelvemeadows,pasturesandroadvergeseach falltrapswerecoveredbyaplywoodrooftoex- wereselectedforthestudy(n =36sites,Table cluderainwater;roofsweresome2cmfromthe tot 1). The list of potential meadows and pastures ground.Thepitfalltrapswerefilledwithsaltwa- wasobtainedfromtheCentreforEconomicDe- terforpreservingthematerialandsoaptoreduce velopment, Transport and the Environment of surfacetension.Thepitfalltrapsweresetup26th– Central Finland (ELY Centre). All the selected 30thofMay2014.Theywereemptiedtwice:18th– sitesweremesicordrymeadows,andhadbeen 22nd of June and 7–11th of July; all traps were managedbygrazing(pastures)orhand-mowing catchingequaltimeperiods. (meadows)foryears. Pastureswereafewhect- Pitfall traps sample surface-active inverte- aresinsizewithvarying,butgenerallylowinten- brates(seeGudleifsson&Bjarnadottir2008fora sity,grazingpressure.Meadowshadbeenhand- studyonCollembola).Duetolimitedresources, mowed once annually, and they had not been notallCollembolawereseparatedfromthepitfall treatedwithanimalmanureorartificialfertilizers. material:fromeachpitfallatleastahundreddif- Themostcommon(occurredinmostsites)plant ferent looking specimens were separated for speciesinmeadowsandpastureswereVeronica identificationandallthesewereidentifiedtothe chamaedrys,Agrostiscapillaris,Alchemillasp., speciesifpossible.Thus,thedatadonotwarrant Festucarubra,Fragariavesca,Hypericumma- statisticalcomparisonsandabundancesshouldbe culatum, Poa pratensis, Ranunculus acris, Ru- onlyusedtodisentangleanecdotalandtruehabi- mexacetosaandTaraxacumspp.;thesewereal- tat affinities. Collembola were identified with, mostequallycommoninbothhabitattypes. and the nomenclature follows, Fjellberg (1998, Roadvergeswereselectedasclosetothepas- 2007).Juvenilecollembolans(n=45)wereonly turesandmeadowsaspossible.Allselectedroads identifiedtogenusorfamily. 160 Komonen&Kataja-aho (cid:127) ENTOMOL.FENNICAVol.28 Table2.NumbersofCollembolaindividualsfrompitfalltrapsfrommeadows,pasturesandroadvergesrecorded duringthefirstperiod.Ifspecimenswereidentifiedfromtwoperiods(forperiods,seeTable1),thefirstnumber indicatesthe1stperiod;poolednumbersofindividualsforbothperiodsaregiveninparentheses. Taxa Meadow Pasture Roadverge Total Arrhopalitidae Arrhopalitesprincipalis 1 2 3 Arrhopalitessp. 1 1 Bourletiellidae Bourletiellahortensis 18 18 Deuterosminthurusbicinctus 1 1 Deuterosminthurussp. 0(3) 0(3) Heterosminthurusclaviger 1 1 2 Dicyrtomidae Dicyrtomafusca 2(14) 0(1) 2(15) Dicyrtominaflavescens 0(3) 0(3) Dicyrtomaflavosignata 3(10) 3 6(13) Dicyrtomaminuta 20(27) 12 5(8) 25(47) Dicyrtomaornata 1 1 2 Ptenothrixatra 25 12(13) 37(38) Indet 2 1 3 Entomobryidae Entomobryacorticalis 5 1 2(4) 8(10) Entomobryamarginata 2 0(4) 2(6) Entomobryamultifasciata 2 1(2) 3(4) Entomobryanicoleti 27 6 33 Entomobryanivalis 1 7 16 24 Entomobryasuperba 2 2 Entomobryasp. 1 46(49) 4(6) 51(56) Heteromurusnitidus 27 27 Lepidocyrtuscurvicollis 0(2) 0(3) 0(5) Lepidocyrtuscyaneus 7 41 48 Lepidocyrtuslignorum 2,378(2,454) 451(468) 959(1,031) 3,788(3,953) Lepidocyrtusviolaceus 21(22) 85 87(90) 193(197) Lepidocyrtussp. 1 4 5 Orchesellabifasciata 4 4 Orchesellacincta 8 8 Orchesellaflavescens 30(31) 30 45(49) 105(110) Orchesellaspectabilis 3 3 Orchesellasp. 9 15(16) 24(25) Indet 3 5 8 Hypogastruridae Ceratophysellasp. 57 57(61) 114(118) Choerutinulainermis 10(11) 11 32 53(54) Schoettellaununguiculata 3 1 20(24) 24(28) Isotomidae Desoriatolya 0(1) 0(11) 0(12) Desoriasp. 6(7) 5(6) 1(17) 12(30) Folsomiaquadrioculata 2 4 6 Isotomaanglicana 32 402(404) 50(71) 484(507) Isotomacaerulea 58 214 218 490 Isotomaviridis 121(135) 335(338) 281(322) 737(795) Isotomasp. 35(36) 21 35(39) 91(96) Isotomurusgraminis 22(32) 225 6 253(263) Isotomurusitalicus 1 1 Isotomuruspalustris 5 5 Isotomurussp. 13 21(22) 1 35(36) Parisotomanotabilis 9 2 11 Indet 107 458 113 678 ENTOMOL.FENNICAVol.28 (cid:127) Collembolaintraditionalbiotopes 161 Table2,continued Taxa Meadow Pasture Roadverge Total Katiannidae Gisinianusflammeolus 3 3 Sminthurinusaureus 30 44 74 Sminthurinusnigergroupsp. 7 1 8 Sminthurinusreticulatus 9 1 10 Sminthurinussignatus 32 1 33 Sminthurinussp. 5 3 14 22 Indet 6 1 1 8 Neanuridae Frieseamirabilis 4 4 Neanuramuscorum 14 3 9 26 Pseudachorutesdubius 1 1 Pseudachorutessp. 2 2 Indet 1 1 Onychiuridae Micraphoruraabsoloni 7 1 8 Protaphoruraarmata 4 4 Protaphorurasp. 1 1 Poduromorpha Indet 1 1 Sminthuridae Allacmafusca 18 5 9(14) 32(37) Capraineamarginata 1 1 Sminthurusnigromaculatus 20 20 Sminthurusviridis 8(11) 7 22(26) 37(44) Sminthurussp. 10 10 Spatulosminthurusflaviceps 32 32 Indet 2 5 6 13 Sminthurididae Sminthuridesschoetti 2 2 Sminthuridessp. 4 4 Sphaeridiapumilis 38 5 46(49) 89(92) Symphypleona Indet 9 14 3 26 Tomoceridae Pogonognathellusflavescens 798(827) 148(200) 213(270) 1,159(1,297) Pogonognathelluslongicornis 11(18) 6 5 22(29) Pogonognathellussp. 3(4) 3(5) 6(9) Tomocerusvulgaris 1 1 Total 3,979(4,118) 2,692(2,774) 2,479(2,738) 9,150(9,630) Soilsamplesweretakenfromeachofthefive eraand60species(Table2);12%(1,164individ- studyplotsandpooledforonecompositesample uals)ofthespecimenscouldnotbeidentifiedto tomeasuresoiltypeandpH.Vegetationcomposi- species.Thenumberofspecimensfromthefirst tionandheight(cm)wasmeasuredfromeachof samplingperiod(n=9,150)washigherinmead- thefivestudyplots16thJune–8thJuly2014(Table ows (n = 3,979) than in pastures (n = 2,692) or 1). roadverges(n=2,479).Alltheresultshereinafter concernthefirstsamplingperiodtoallowcom- 3.Results parisonduetoequalsamplingperiodinallsites. Thenumberofspecieswashigherinmeadows(n Altogether, 9,630 Collembola individuals were =40)androadverges(n=39)thaninpastures(n recorded.Thesebelongedto12families,34gen- =33).Thereweresomedifferencesinsoilquality 162 Komonen&Kataja-aho (cid:127) ENTOMOL.FENNICAVol.28 and vegetation: soil pH ranged from 3.73–5.99 one habitat type over the others, Spatulos- and vegetation height 8–42 cm (Table 1). Most minthurusflavicepsbeingpossiblyanexception (78%)oftheindividualswereEntomobryidaeor (seebelow).Fivetoeightuniquespecieswerere- Isotomidae. The most abundant species were cordedfromeachofthethreehabitattypes, but Lepidocyrtuslignorum(41%ofallindividuals), theseweremainlyrepresentedbyonlyafewindi- Pogonognathellusflavescens(13%)andIsotoma viduals. Although we did not identify all speci- viridis(8%). mens, the similarity of the collembolan fauna amongthethreehabitattypescouldbearealpat- tern.Therationaleisthatthedominantvascular 4.Discussion plants were identical among the three habitat types,andvegetationcompositionandstructure The number of collembolan individuals was areknowntoaffectcollembolandensitiesanddi- highest in meadows, whereas pastures and road versity(Salamonetal.2004,Sabaisetal.2011). vergeshadsimilarlylowernumbers.Thenumber Overall,thespeciesfrompastures,meadowsand of species was higher in meadows and road roadvergeswereratherdifferentfromthosethat verges than in pastures. In all road verges soil aretypically caughtby pitfallsfromforestsand fractionwassand,whereasinpasturesandmead- clearcuts. The latter usually include Entomo- ows soil fractions ranged fromsand to fine silt. bryidae,especiallythelargespecies,suchasPo- Althoughsoiltypeinfluencescollembolandensi- gonognathellusandOrchesella(Kataja-ahoetal. ties(Gudleifsson&Bjarnadottir2008),thesedif- 2016). ferencesareunlikelytoexplainthesmallerspe- WerecordedS.flaviceps(n=32individuals) ciesnumberinpastures,becausethesoiltypewas fromseveralroadverges.Previouslythespecies similar in pastures and meadows. However, the has been found only from salt meadows (Fjell- somewhat smaller species number in pastures berg1998).IfS.flavicepsreallyisaspeciesthat may result from animals trampling on pitfall benefitsfromsalt,thenthespilloverofthede-ic- traps, which was commonly observed. Animals ingsaltonroadvergesmaycreateasuitablehabi- mayalsohavecausedsoilcompaction,whichcan tat for the species in Finland, far away from decreasethedensityofcollembolans(Heisler& coastalareas.Moregenerally,theimportanceof Kaiser1995,Larsenetal.2004). road verges for salt-favouring species deserves Itisknownthatcollembolandensitiesinpas- furtherattention. tures and hayfields are highest in late summer Inaddition,SminthurusviridisandS.nigro- (Gudleifsson&Bjarnadottir2008).Eventhough maculatus were recorded in great numbers, the wedidnotidentifyallthespecimensanddidnot former from all studied biotopes and the latter sampleinlatesummer,theoverallspeciesnum- onlyfromroadverges.Althoughtheoccurrence ber is comparable to other large-scale sampling ofthesespeciesinFinlandwasuncertainaccord- studies in similar habitats with roughly similar ingtoFjellberg(1998),laterbothofthesespecies number of identified individuals, but different havebeendiscoveredfromFinland(P.Vilkamaa, samplingmethodology(cf.Huhtaetal.2010). pers.comm.).Inconclusion,ourstudycorrobo- MostoftherecordedindividualswereEnto- rates the results by Huhta et al. (2010) in that mobryidaeorIsotomidae.Indeed,epedaphicspe- many non-forested habitats host apeculiar Col- cies are typified by Entomobryoidea and Sym- lembolafauna.Systematicstudiesinmarginalor phypleona (Hopkin 1997). The most abundant otherwisepoorly studied biotopes arecrucial to species were Lepidocyrtus lignorum, Pogono- get a thorough understanding of species abun- gnathellusflavescensandIsotomaviridis.These dance,distributionandhabitataffinities. wereabundantin allthethreehabitattypes and havebeenrecordedalsoearlieringreatnumbers Acknowledgements.Wethanktwoanonymousreviewers from grasslands (Gudleifsson & Bjarnadottir fortheirsuggestionstoimprovethemanuscriptandKo- neenSäätiöforfundingtheproject“Perinneympäristöjen 2008).RoadvergeshadmoreSminthuridaespe- janiitäkorvaavienkulttuuriympäristöjenhoitobiologisen cies than meadows and pastures, but in general monimuotoisuuden,sosiaalisenkestävyydenjataloudelli- there was no species that would clearly favour sentehokkuudennäkökulmasta”. ENTOMOL.FENNICAVol.28 (cid:127) Collembolaintraditionalbiotopes 163 References Lagerlöf,J.&Andrén,O.1991:Abundanceandactivityof Collembola,ProturaandDiplura(Insecta,Apterygo- ta)infourcroppingsystems.—Pedobiologia35:337– Eisenbeis,G.&Wichard,W.1987:Atlasonthebiologyof 350. soilarthropods.—Springer,Berlin,Germany.437pp. Larsen,T.,Schjonning,P.&Axelsen,J.2004:Theimpact Fjellberg,A.1998:TheCollembolaofFennoscandiaand of soil compaction on euedaphic Collembola. — Denmark.PartI:Poduromorpha.—FaunaEntomolo- AppliedSoilEcology26:273–281. gicaScandinavica35:1–184.Brill,Leiden. Oldén,A.2016:Plantbiodiversityinborealwood-pastures Fjellberg,A.2007:TheCollembolaofFennoscandiaand –impactsofgrazingandabandonment.—PhDThe- Denmark.PartII:EntomobryomorphaandSymphy- sis,Jyväskylästudiesinbiologicalandenvironmental pleona.—FaunaEntomologicaScandinavica42:1– science318.Jyväskylä.46pp. 264.Brill,Leiden. Gudleifsson, B. E. & Bjarnadottir, B. 2008: Springtail Owojori,O.J.,Reinecke,A.J.,Voua-Otomo,P.&Rei- (Collembola)populationsinhayfieldsandpasturesin necke,S.A.2009:Comparativestudyoftheeffectsof northernIceland.—IcelandicAgriculturalSciences salinityonlife-cycleparametersoffoursoil-dwelling 21:49–59. species(Folsomiacandida,Enchytraeusdoerjesi,Ei- Heisler,C.&Kaiser,E.A.1995:Influenceofagricultural seniafetidaandAporrectodeacaliginosa).—Pedo- traffic and crop management on collembolan and biologia52:351–360. microbialbiomassinarablesoil.—BiologyandFerti- Petersen, H. 1994: Areview of collembolan ecology in lityofSoils19:159–165. ecosystemcontext.—ActaZoologicaFennica195: Hopkin, S. P. 1997:Biology ofthe Springtails(Insecta: 111–118. Collembola).—OxfordUniversityPress,NewYork, Pommeresche,R.&Løes,A.-K.2014:Diversityandden- U.S.330pp. sityofspringtails(Collembola)inagrass-cloverleyin Huhta, V., Karppinen, E., Nurminen, M. & Valpas, A. North-westNorway.—NorwegianJournalofEnto- 1967:Effectsofsilviculturalpracticesuponarthropod, mology61:165–179. annelidandnematodepopulationsofconiferousforest Rusek,J.1975:DiebodenbildendeFunktionvonCollem- soil.—AnnalesZoologiciFennici4:87–143. bolenundAcarina.—Pedobiologia15:299–308. Huhta,V.,Hyvönen,R.,Kaasalainen,P.,Koskenniemi,A., Sabais,A.C.W.,Scheu,S.&Eisenhauer,N.2011:Plant Muona,J.,Mäkelä,I.,Sulander,M.&Vilkamaa,P. species richness drives the density and diversity of 1986:SoilfaunaofFinnishconiferousforests.—Acta Collembolaintemperategrassland.—ActaOecologi- ZoologicaFennica23:345–60. ca37:195–202. Huhta,V.&Räty,M.2005:Soilanimalcommunitiesof Salamon, J.-A., Schaefer, M., Alphei, J., Schmid, B. & plantedbirchstandsinCentralFinland.—SilvaFen- Scheu,S.2004:EffectofplantdiversityonCollembo- nica39:5–19. lainexperimentalgrasslandecosystem.—Oikos106: Huhta,V.,Räty,M.,Ahlroth,P.,Hänninen,S.-M.,Mattila, 51–60. J.,Penttinen,R.&Rintala,T.2005:Soilfaunaofdeci- Siira-Pietikäinen,A.,Pietikäinen,J.,Fritze,H.&Haimi,J. duousforestsascomparedwithspruceforestsincen- 2001:Short-termresponsesofsoildecomposercom- tralFinland.—MemorandaSocietatisproFaunaet munitiestoforestmanagement:clearfellingversusal- FloraFennica81:52–70. ternative forest harvesting methods. — Canadian Huhta,V.,Siira-Pietikäinen,A.,Penttinen,R.&Räty,M. JournalofForestResearch31:88–99. 2010:SoilfaunaofFinland:Acarina,Collembolaand Song,L.,Liu,J.,Yan,X.,Chang,L.&Wu,D.2016:Eue- Enchytraeidae.—MemorandaSocietatisproFaunaet daphicandhemiedaphicCollembolasufferlargerda- FloraFennica86:59–82. mages than epedaphic species to nitrogen input. — Kataja-aho,S.,Hannonen,P.,Liukkonen,T.,Rosten,H., EnvironmentalPollution208:413–5. Koivula,M.J.,Koponen,S.&Haimi,J.2016:The Viard,B.,Pihan,F.,Promeyrat,S.&Pihan,J.-C.2004: arthropodcommunityofborealNorwayspruceforests Integrated assessment of heavy metal (Pb, Zn, Cd) respondsvariablytostumpharvesting.—ForestEco- highwaypollution:bioaccumulationinsoil,Gramina- logyandManagement371:75–83. ceaeandlandsnails.—Chemosphere55:1349–1359.