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SILVA FENNICASilva Fennica vol. 50 no. 2 article id 1462

Category: research article

www.silvafennica.fiISSN-L 0037-5330 | ISSN 2242-4075 (Online)

The Finnish Society of Forest Science

Natural Resources Institute Finland

Pekka Punttila 1, Olli Autio 2, Janne S. Kotiaho 3, D. Johan Kotze 4, Olli J. Loukola 5,

Norbertas Noreika 4,6, Anna Vuori 3 and Kari Vepsäläinen 6

The effects of drainage and restoration of pine mires

on habitat structure, vegetation and ants

Punttila P., Autio O., Kotiaho J.S., Kotze D.J., Loukola O.J., Noreika N., Vuori A., Vep-säläinen K. (2016). The effects of drainage and restoration of pine mires on habitat structure, vegetation and ants. Silva Fennica vol. 50 no. 2 article id 1462. 31 p.

Highlightsi e aina e s if e is ic c si i n an an asse la es wa s f es c ni ies.aisin e wa e a le level i c fillin an e innin f ees affec e i e c niies si ivel al ea ea s af e e s a f es a i n.

The extent of tree cover, the coverage of Sphagnum sses an e wa e a le level we emajor determinants of ant assemblage structure.

Abstracta i a l ss an e a a i n a e e ain ea s i ive si w l wi e. e a le nea l

80% of peatlands in southern Finland have been drained. There is thus a need to safeguard the e ainin is ine i es an es e e a e nes. n s la a iv al le in an ec s s e san li e an e s ne lan s ecies s a e ec s s e c n i i ns f e i a. e effec s fi e es a i n an s se en ve e a i n s ccessi n n an s weve a e l n e s .e inven ie ee s an s ve e a i n wa e a le level an an s wi i fall a s in nine

mires in southern Finland to explore differences in habitats, vegetation and ant assemblages a n is ine aine ea s a an ecen l es e ea s a ine i es. ee ec e a es in e wa e a le level i c fillin an ec ns c in s a se ee s an s

c in s will ec ve i e ve e a i n an an s. e f n e ic a le es nses in a i as c e is ic c si i n an an asse la e s c e aina e an es a i n.

weve f i e s ecialis an s e es l s we e va ia le an l n e e ni in is nee ec nfi e s ccess f es a i n since ese s cial insec s es a lis e ennial c l nies wi

l n c l n c cles. e c ncl e a es in e wa e a le level an ee s an s c e seeec ve e c a ac e is ic ve e a i n an an asse la es in e s e . is ec ve was

li el en ance eca se aine i es s ill a ac el an ca el an c nnec e nesswas s ill eas na le f i e anis s ec l ni e e es e i es ei e f l cal ef iaf la i ns f nea i es.

Keywords ic i i ive si a e i c in ec l ical es a i n ici ae ine san fens ansf in an ansf e aine i es wa e a le levelAddresses 1 innis nvi n en ns i e . . elsin i inlan 2 Centre for Economic Development, Transport and the Environment in South Ostrobothnia, P.O. Box 252,

aasa inlan 3 nive si f v s l e a en f i l nvi n en alciences . . v s l inlan 4 nive si f elsin i e a en fnvi n en al ciences . . nive si f elsin i inlan 5 nive si fl e a en f i l . . l inlan 6 nive si f elsin i

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

e a en f i sciences . . nive si f elsin i inlanE-mail e a. n ila a is .fiReceived 7 September 2015 Revised 8 December 2015 Accepted 16 December 2015Available at http://dx.doi.org/10.14214/sf.1462

1 Introduction

The Global Peatland Database of the International Mire Conservation Group (IMCG) has estimated that peatlands represent about 3% of the globe’s total land mass, and that at least 80% of peatlands a e l ca e in a eas wi n e n e e a e c l cli a es in an e l . c fe i inal ea lan a ea as al ea een l s in an e l . n inlan ea lan s

c ve f e lan ass w ic is e la es s a e f ea lan s l all vinen e al. .At present, of the 8.8 million ha of peatland about 4.1 million ha remain undrained and 4.6 million ha are drained (Peltola 2014).

s f e innis is ine i es a e si a e in a lan w e eas in s e n inlan nea lf ea lan s ave een aine vinen e al. . aina e f f es an a ic l e

an ea a ves in a e e ain ea s e i e i a an a i a es in inlan aa inen eal. assi e al. in n a a i nal a e f i es an ea lan s .win aina e e si e an c nnec ivi f i es ave as icall ec ease in s e n in

lan w ic as i ai e e ve en an c l ni a i n f i e i a a n a i a s vinene al. . e n e f e lis e s ecies livin i a il an sec n a il in innis i es is

an es ec ivel c isin . f all e lis e s ecies assi e al. . eca sela i ns es n sl wl inc ease a i a l ss a la e s a ial scales ans i an

i e s ecies a e li el e e iencin an e inc i n e il an e al. ans i an vas ainen. e e a n e f i e ass cia e insec s a e a i a s ecialis s i e an an san s ne e inc i n nn . n anal sis f c an es f e lis ca e ies i.e.

e ee f ea f s ecies in inlan e ween assi e al. an assi e al.s we a i e ass cia e s ecies ave ec e e ea ene nl s ecies

less ea ene assi e al. . ls la i ns f an i e i an e s ecies aveecline w ic in ica es an inc easin n e f ea ene i e s ecies vinen e al. .

lea l e e is a nee safe a e e ainin is ine si es an es e e a eec s s e s f ean i es an as a ee f e al f es in f e a e ecs s e s see e ic i i ive si a e ean issi n .is a e weve see s n ealis ic eav es a i n eas es s e c le e ac ss la e

a eas an in a s i e w ile c ensa in f n in e a a i n elsew e e ia e al.see als ia an ilanen .es a i n is a cess w e e e f nc i ns s c es cesses an i ic c ni

ies f a i a s an ec s s e s a e e ne wa s ei is ine s a e cie f c l icales a i n n e na i nal cience lic in in an e lne in a esi e s a e. ea lan es a i n ai s f e ec ve f ec s s e f nc i ns

e n na all f nc i nin an self s s ainin ca n acc la in an n ien e ainin i eec s s e s l vainen e al. asan e e al. a ala e al. . an alsincl e e ec ve f ec l ical e ies e an i an ali f in win anwin wa e s an na all c a in wa e a le levels i.e. na al l e ec vef ea f in ve e a i n e. . Sphagnum sses an e ec ve f s c al c a ac e is ics

(e.g. species composition) and processes (e.g. succession) of the mire biota (Aapala et al. 2008). an i es even wi in e c en c nse va i n a ea ne w f inlan ave een aine f

f es i e es a lis en f c nse va i n a eas. n a a f ese

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

ea lan s ave al ea een es e an a a s ill nee e es e safe a eec l ical val e f e ec e a eas i il e al. .

e effec s f ea lan es a i n n e i a nee f e inves i a i n es eciall na i a s ecialis s in an e l . n s vi e val a le inf a i n w en eval a inlan ana e en ac i ns an assessin l n e ec s s e c an es eca se e a e sensi ivec an es in a i a ali an la a iv al le in an ec s s e s n e sen an a e

n e w an is e a an e al. . e li e an e s ne lan s eciess a e ec s s e s f e anis s. e ac ivi ies f an s an ei nes an ail c ns c i nsa affec e i a c nsi e a l ll le an ils n n ila an il el inen

Finér et al. 2013, and references therein).i es c ise e i a sec n a a i a f a leas a i f e na ive an s ecies

f inlan n ila e al. . n eal a eas a few s ies ave s wn a aina e f i esaffec s i e an s ecies ne a ivel e s llin w e s l inen e al.

n ila an il el inen n wle e n i e an c ni ies an ei asse lcesses a e sca ce l all ve e s l inen e al. l ss llis n e al.

elli an llis n a a elis an a n a c f e al. an e s eal. an e al. . e effec s f i e es a i n an s se en ve e a i n s ccessi nn an s an e insec fa na a e l n e s ai e al. van inen e al.a s e al. l e al. ei a e al. .

i a ai is c a ac e i e iffe ences in i e a i a s an asse la es an e ccence f in ivi al an s ecies a n is ine aine an ecen l es e i es. e f c se nee es i ns in e s e a a e e s essen ial iffe ences in ve e a i n e weenis ine an aine i es an w es a i n affec ese es e an asse la e s ce iffe e ween is ine an aine i es an w es es a i n affec an asse la es

w a e e an s ecies is i e a n is ine aine an es e i es an w a a ee s i an i e c a ac e is ics affec in e an s e s se ves as a aseline

f f e ni in f e s ie i es w en l n e e es a i n s ccess is eval a e .ene all we es if es in e wa e a le level i c fillin an ec ns c in na

all s a se an l w ine s an s eav innin an a ial clea c in will es e a i a sall w ec l ni a i n an ec ve f e ve e a i n an an s f is ine i e a i a s.

2 Materials and methods

2.1 Study design

e s alli s a s il life inlan ana es e innis c nse va i n a ea ne w an ai ses e i e ec s s e s a ave een aine f f es i e es a lis en f c nse vai n a eas. es a i n s ccess is als ni e a ala e al. . e ili e a sa lin lne w es a lis e ni es a i n s ccess f aine ine i es wi e al f e ninthe drained mires to their natural state.

e nine s i es a le a e l ca e in w e i ns in inlan e n a elia anen al inlan al n e e e ween e s e n an i le eal f es ve e a i n nes

i e al. s l in e eccen ic ne Sphagnum fuscum aise s an a lie anneva an l nneva in a le in e s e n aa a i e ne i vi . ei es a e ainl ic li is als wi e an in an s e i e a s a ees ic. ll i es el n a l e a a ne w f na e ec i n a eas a sf e i es a een aine in s s i e es a lis en f na e c nse va i n

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a eas si al e al. . es a i n f a s f all e s ie i es s a e infillin e i c es an a ves in a va in i n f ees in . in a ves ini e an l w we e e ve all ee s s an va in a n s f l in esi esanc es an ee s we e lef e in .

eac i e i e s a f e es a i n cess in ee se a a e s a easwe e selec e e esen iffe en ea en s is ine i e aine i e an es ei e a le . n ne f e i es ein s weve e a ea esi na e e ain aine

was als es e an is i e s vi e ne is ine an w es e a eas n ainea ea a le . eac ea en e i e w ansec s n ave a e a a we ees a lis e wi ee sa lin l ca i ns e ansec . is es l e in si sa lin l ca i ns eea en e i e nine i es ee ea en s w ansec s ee sa lin l ca i ns

sa lin l ca i ns . e ea en s e i e we e n ave a e a a .

Table 1. i e es f e is ine aine an es e ea en s i e s a f es a i n in in e nines i es in en al inlan an e n a elia an ei a i a e c ina es. n e f sa lin l cai ns e esen in e iven i e e al sa lin l ca i ns .

Region/mire Treatment i e es1 Coordinates

Central Finlandie anneva Pristine i

DrainedRestored muIR (3), muRaR(3)

l nneva Pristine aDrained aRestored e a

e n l anv i Pristine IR (1), SR (1), TR (4)DrainedRestored

e n l anv i Pristine LkR (6)Drained aRestored a

e n a eliaRistisuo Pristine LkR (1), RaR (5)

Drained aRestored muIR (4), muPsR (1), muRaR (1)

i as Pristine LkR (1), RaR (5)DrainedRestored s

Rapalahdensuo Pristine LkR (4), RaR (2)Drained aRestored

Tiaissuo Pristine LkR (3), RaR (3)DrainedRestored muIR (4), muRaR (2)

ein s Pristine aes e a muIR (3), muLkR (2), muRaR (1)es e LkR (1), muIR (4), muLkR (1)

1 i e e a evia i ns a e acc in la e al. an n lis ansla i ns a e acc in a ni e al.wa f s ine s w se e s fens w se e ine fens ansf in wa f s ine s

e ansf in i e ll w ine s ansf in in ea e ine i es ansf in w se eine fens s ansf in Carex globularis ine i es a ansf in Sphagnum fuscum s ansf

ing Eriophorum vaginatum ine s i la ine fens a Sphagnum fuscum s all se e ine fensansf e aine i es Eriophorum vaginatum ine s e ic ea f es s.

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2.2 Vegetation, drainage and ant data collection

eac sa lin l ca i n e i e si e e was ec e i es a i n eas es ina le an seve al se s f a a we e c llec e ee s an c a ac e is ics we e ec e f

a circular 100 m2 ee sa lin l in e i le f w ic ee sa lin an ic si e ea a we e ec e f a 2 sapling square. In each corner of the 25 m2 sapling square, (3) e c ve s f vasc la lan ss an lic en s ecies an li e an s face wa e we e es i a e

wi in 2 ve e a i n s a es f s a es e sa lin s a e . e c ve a es we e es i a en a scale . . . a a se s

a ve we e c llec e i es a i n in an a ain in w en we sa lee an s. e e we ainl se e a a c a ac e i e sa lin l ca i ns an vi e

envi n en al va ia les f e an anal ses. i i nall in e wa e a le level wasni e sin las ic es i es len c ia e e as n wa e wells.e es we e se in w si e c ne s f eac 2 sa lin s a e. a e a le e i.e.

is ance f e wa e a le f e i e s face was eas e wi an acc ac f c a wwee in e vals si i es f ea l ne i s in .

ee ci c la ee sa lin l s a i s . 2 we e evenl lace al n eacansec a i a el . a a . eac ee sa lin l ee ei c s ecies

an ei s e n e s we e ec e . e ees we e classifie in si e classes acc in eidiameter at breast height (DBH, d1.3 m classes an c an ei classes .an . ea s an in ees sna s an fallen ea ees l s we e ec e in a si ila waas the living trees (at 1.3 m DBH from the ground for snags and 1.3 m from the butt end for logs).

ee sa lin an s ei c s ecies an ei n e s we e ec e fthe sapling square (25 m2 wi in e ee sa lin l . i in e sa lin s a e e e cen a ec ve s f i e s face a es i.e. ic si e es c lawn an a we eestimated to the nearest 10 percent.

ase n a i i n wle e f e ene al i ance f iffe en envi n en al c a ac eis ics n e cc ence f an s we selec e a se f en ial e lana va ia les in e anal sesn an s. a ia les selec e ela e e e ee f s a in f can la e s f iffe en ei anve e a i n e e cc ence f en ial nes in si es an e cc ence f en ial eand aphid colonies. Also, the most important predictor variables indicating successful restoration we e incl e . e va ia les se incl e ea en a ee level fac is ine ainerestored), and several variables from (2) the 100 m2 ee sa lin l s n e f l w ees. all ees an ea ees e 2 sapling squares (number of tree saplings

an i ns f e ic si e es c lawn an a e 2 vegetation squares e cen a e c ve s f s face wa e an li e e le c ve f Sphagnum spp., the pooled cover f e sses e le c ve f e s se es an asses i.e. n n w ann al lan s ele c ve f l w c wa f s s an e le c ve f all c wa f s s

an s s f e ails see el w ave a e c ve f e f 2 ve e a i n s a es was sean wa e a le e ave a e f e w wells e sa lin s a e .

e e a i n a a we e se c a ac e i e e ic a i a es f e sa lin l ca i nss c a e w l e ec e ain ec l ical a ien s a n e sa lin l ca i ns. e calculated the combined cover of (1) Sphagnum spp. mosses, (2) other mosses, (3) herbs, sedges and asses sens la e al. a n vi e s a e in ea l s in an la e a n

l w wa f s s s ecies icall c in ei Andromeda polifolia L., Calluna vulgaris (L.) Hull, Empetrum nigrum L., Vaccinium myrtillus L., V. vitis-idaea L.) excluding the recumbent V. oxycoccos L. and V. microcarpum c . e . c al . an all wa f s s(Chamaedaphne calyculata (L.) Moench, Ledum palustre L., V. uliginosum . le wi s s(Betula nana L., Salix myrsinifolia alis . a vi e s a e e win seas n.

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n s we e sa le sin i fall a s. ne a a l las ic a wi inne ia e e fan a e f was se in a e esen a ive l ca i n wi in eac 2 ee sa lin

l w ic iel e a s e ea en e i e i.e. a s in al. a in was c n in sf si wee s wi ca c es c llec e an a s ese eve sec n wee e ween a an l

. e ve e s face ensi n an ese ve e a s ca cl f a a l s li n wi a few s f e e en was a e e a s. n s we e i en ifie wi e e s fllin w eife an ec ws i e al. . n e asis f lis e

a a e s e s l inen e al. n ila an il el inen an wn fiele e ience all e f e en l c llec e an s ecies we e an e acc in ei affini f i ea i a s es nse aina e an affini f ine ea f es s n a scale f s n es i eaffini s n es ine f es affini . i i nall w en anal sin e effec s f aina ean es a i n n e n e f i e an s ecies we incl e nl se s ecies assesse in ela es e lis w f inlan e e e f en e a assi e al. livei a il in i es Formica exsecta lan e F. picea lan e F. uralensis s

an Myrmica scabrinodis lan e in a a.

2.3 Statistical analyses

i s we se l li eli a i es s an s al allis n n a a e ic e ian es s e l eva ia i n in e ee s an va ia les an e va ia les c a ac e i in i e a i a s an in ancc ence a n e ea en s. c n l f false isc ve a e in l i le es in e i inal val es we e a s e wi e e in en a ini an e ieli sin e f nc i n.a s e in e ac a e s a s in ve si n . . e ea . i i nall we anal se an cc ence a n iffe en i e es wi s al allis n n a a e ice ian es s inc ease li i e n wle e f a i a ass cia i ns f i e an s. e e cl e

all sa lin l ca i ns n es e i es an all i e es wi less an f sa lin l ca i nsw ic es l e in sa lin l ca i ns in ese anal ses.

ec n we a lie n n e ic l i i ensi nal scalin c a ac e i e iffe encesin ee s an s is ic c si i n an an asse la e c si i n a n e ea en s is ineaine an es e . e ee ina i ns we e e f e sin e ve an c ni

ec l ac a e ve si n . sanen e al. in e ea . e se e ve anf nc i n e a wi n ce w i ensi nal ina i ns wi seve aln e an s a s fin s a le s l i ns. e ina i n i ensi ns a es we e scalealf c an e ni s. e fi e a se f envi n en al va ia les see a ve in ina i ns sin

e ve an f nc i n envfi an es e ei fi wi e a i n es s.e f e ee s an c a ac e is ics e we issi ila i eas e was se

e a a we e e n e f sa lin an ee in ivi als e s ecies e n e f sa lin antree species, the number of trees in three diameter and three height classes, the number of snags in w ia e e classes c c an e n e f l s in ee ia e e classes inthe 100 m2 ee sa lin l s an 2 sa lin s a es. e e cl e va ia les wi less anpositive values out of 162 data entries.

e f e is ic a a e a is issi ila i eas e s a e ansf a i n an isc nsin le s an a i a i n we e se .

e f e an a a e a issi ila i eas e wi an esence a sencea a in e i fall a s was se eca se c ns ecific in ivi als f s cial insec s ca e in ai fall a a e n s a is icall in e en en ni s an n i ec l e ec asse la e s c eel ne e s l inen e al. elli e al. i ins an in en . nl

w e an a a we e incl e in e an e l iva ia e anal ses eca se nl w e sindicate established colonies.

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i we se ene ali e linea i e els l e f nc i n in e l e acage, Bates et al. 2013) in R (R Core Team 2013) to evaluate the effects of treatment and a set of envi n en al va ia les n e cc ence f an s ecies. n esence a sence a a we eele f ll win a in ial e is i i n. i es nes e wi in s e i ns we e incl e as aan fac . ine s ecies cc in in f e a s we e anal se in ivi all .

anal ses we e s ec el selec i n ea en Sphagnum cover, hummock c ve an n e f all ees n win ei a i i e ec e i ance f ein ivi al s ecies anal se . e f ll win e ic va ia les we e weve s ec elselec i n n e f ea ees l s an sna s le li e c ve c ve s f l w 20 cm) and all c wa f s s an e le c ve f e s se es an asses. ese va ia leswe e e ve ne a a i e if ei val es e cee e . an if val es see n aand Anderson 2002) decreased after excluding the particular variable. All predictor variables e ce ea en we e s an a i e e ean an ni va iance c iel e a ee c a a le. i es f w ic n in ivi als f a a ic la s ecies we e ca we e n

incl e in e anal ses since e a e ns i a le f a s ecies f eas ns a a en c nsi e e in is s .

3 Results

3.1 Mire-site types

Prior to starting restoration in 2003, 21.6% of the 162 sample locations represented transformed aine i es w e e e i en ifica i n f e i inal i e e was i ssi le win s ccessi nal

changes. For the 34.0% and 43.8% of the sample locations established in pristine or transforming i e es es ec ivel a le ine i es an ine s we e ina in . f all

l ca i ns an ine fens an ic ine fens a e . . e s c n is ine es we eSphagnum fuscum s a . f all sa le l ca i ns a le an l w se e ine fens

. an e s c n ansf in es we e ansf in wa f s ine s. an ansf in Sphagnum fuscum bogs (muRaR, 10.5%).

3.2 The effects of drainage and restoration on tree stand and vegetation

aina e le a f f l inc ease in e n e f ee s e s ela ive is ine i es ane inc ease in i c e ce e inance f ines an le i e ine i c s an s in eaine i es a le . ee w an s win s c inc ease as e ec e es eciall

e ea e a n ance f la e ees 1.3 7 cm or h 3 m) in drained than in pristine mires. es a i n a ves in s ccessf ll c nve e s an s c e in e s f s e n e ee

si e is i i n an ee s ecies c si i n cl se is ine c n i i ns a le . i ila le e we e e i c sa lin s in e aine an is ine i es w e eas es e i es a

in e e ia e n e s a le . e a n f ea ees was ene all l w an a e si ilaa n e ea en s e a n f s all si e l s was i e in aine an es ethan pristine mires (Table 2).

e ic si e e is i i n f aine an is ine i es iffe e c nsi e a l f nean e e s face f aine i es was al s c le el a le c ve e cw e eas e an alf f e s face f is ine i es was c ve e lawn an a .e ic si e e is i i n in es e i es ese le a f is ine i es e s a e f

c was la e an in is ine i es .

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Table 2. Mean (x ee s an an sa lin c a ac e is ics ic si e e c ve a e i e s face c ve a e anwa e a le levels f e sa lin l ca i ns in is ine aine an es e i es. iffe ences in va ia les a n eea en s we e es e wi s al allis an s es f ea en s n s a in e sa e le e iffe e

si nifican l acc in a s e i i es wi c i ical . . e a s e e i inal val es a . in e a lec n l false isc ve a e in l i le es in sin e e in en a ini an e ieli . ei 1.3 ia e e a eas ei . f e i e s face level . f e en f l s an n e

of sampling locations. Data on living and dead trees are from the 100 m2 ee sa lin l s a a n ee sa lin san ic si e es a e f e 2 sa lin s a es a a n i e s face c ve a e a e f e 2 vegetation s a es an a a n wa e a le e a e f e n wa e wells.

a ia le Treatment Test statistics

is ine aine es ex SE x SE x SE H p adj.

ivin ees .Total number of stems 5.8 1.0 a 23.1 2.5 b . 1.2 a 60.47 <0.0001

. f ines 5.8 1.0 a . 1.3 b 8.0 1.2 a 42.03 <0.0001

. f i c es 0.1 0.0 a 7.4 2.3 b 1.2 0.4 a . <0.0001

. f s e s 1.3 < 7 cm 4.7 0.8 a 13.8 2.3 b 7.1 1.1 a 17.65 0.0007

. f s e s 1.3 c 1.0 0.3 a 8.2 0.8 b 1.8 0.3 a 67.34 <0.0001

. f s e s 1.3 c 0.0 0.0 a 1.1 0.2 b 0.3 0.1 a 32.44 <0.0001

. f s e s . 4.1 0.7 3.5 0.8 3.5 0.8 2.05 1.0000

. f s e s 1.7 0.3 a 12.5 2.0 b . 0.7 c 40.24 <0.0001

. f s e s 0.0 0.0 a 7.0 1.2 b 0.8 0.4 a . <0.0001e f s ecies . 0.1 a 1.8 0.1 b 1.0 0.1 a 53.10 <0.0001

Dead trees:Total number of snags 0.7 0.2 1.1 0.2 . 0.3 . 1.0000

. f sna s 1.3 < 7 cm 0.6 0.1 . 0.2 0.8 0.3 . 1.0000

. f sna s 1.3 c 0.1 0.1 0.2 0.1 0.1 0.0 1.16 1.0000Total number of logs 0.0 0.0 a 0.4 0.2 a 0.6 0.2 a 10.24 0.0354

. f l s 1.3 < 7 cm 0.0 0.0 a 0.4 0.2 a 0.6 0.2 a 11.15 0.0235

. f l s 1.3 c 0.0 0.0 0.1 0.0 0.1 0.0 1.57 1.0000

. f l s 1.3 c 0.0 0.0 0.0 0.0 0.0 0.0 1.70 1.0000

ee sa lin s cTotal number of saplings . 0.3 2.6 0.6 1.7 0.3 1.02 1.0000

. f ines . 0.3 a 1.4 0.4 b 1.2 0.2 ab 8.60 0.0736

. f i c es 0.0 0.0 a . 0.4 b 0.3 0.1 ab 17.14 0.0014e f s ecies 0.7 0.1 . 0.1 0.8 0.1 1.31 1.0000

ic si e es Hummock 46.1 3.6 a . 2.0 b . 4.1 c 71.47 <0.0001awn 37.8 4.0 a 1.1 0.7 b 24.8 3.8 c 58.68 <0.0001

Flark 15.5 3.6 a 0.0 0.0 b 6.3 1.7 a 27.05 <0.0001

i e s face c ve a e Water 0.0 0.0 a 0.0 0.0 a 2.4 1.0 a 14.20 0.0052 Litter 2.4 1.2 a 12.1 3.1 ab 15.5 2.4 b 27.30 <0.0001 Sphagnum spp. . 1.7 a 30.7 3.8 b 46.0 . b . <0.0001 Other mosses 3.1 0.6 a 38.4 . b 22.7 2.8 c 64.53 <0.0001 Herbs, sedges and grasses 14.0 1.1 a 8.2 1.0 b . 2.2 a 22.51 <0.0001

w wa f s s . 1.2 ab 16.6 2.1 a 8.6 1.3 b . 0.0655all wa f s s 3.7 0.6 a 6.2 0.8 ab . 1.0 b . 0.0007

a e a le e c el we i e s face

15.1 1.3 a 38.0 1.5 b 16.0 1.3 a 74.34 <0.0001

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

i ila l e s face c ve f aine i es was a a icall iffe en f is ine i ese ss la e was ina e Sphagnum sses a le in is ine i es in ainei es wi a i e c ve f e sses . iffe ences in e e la e we e cleae le c ve f e s se es an asses was l we an e c ve f all wa f s s i e

in drained than pristine mires (Table 2). Pristine mires had higher cover of Sphagnum mosses but l we c ve f e sses li e an all wa f s s an es e i es a le .

e wa e a le level was l we in e aine a a e f c a le an in epristine (15 cm) and restored (16 cm) mires.

ee s an an sa lin c a ac e is ics f e aine i es iffe e clea l f se fe is ine an es e i es in e ina i n ea en 2 0.205, p < 0.001): samlin l ca i ns f e aine i es we e e i in e ina i n an is la e la e sca ew e eas l ca i ns f e is ine an es e i es we e e lef wi s all sca e i . .aine i es we e c a ac e i e a i c i e an an a n ance f la e ees is ine

an es e i es ines an s alle ees i . . e sca e f e es e sa lin l cai ns weve was la e an a f e is ine nes i . . ive f e a i i selec ean fi e envi n en al va ia les s we si nifican c ela i ns < . wi e

ina i n s ace e n e f all ees wa e a le e an e c ve f c ic si ee a i e val es in e aine i es an elsew e e i . . e val es f le c ve

of Sphagnum sses an se f e s se es an asses we e i e in e is ine an es emires than in the drained ones (Fig. 1 A).

e is ic c si i n f e aine i es iffe e clea l f e is ine an es ei es in e ina i n ea en 2 0.352, p < 0.001, Fig. 2 A). Sampling locations of the aine i es we e s l e i in e ina i n w e eas se f e is ine i es we es l e lef an se f e es e i es s l in e ween i . . a lin l ca i nse i e aine i es an an f e es e nes a i e c ve an cc ence a e f

forest species such as Vaccinium myrtillus, V. vitis-idaea, Pleurozium schreberi (Willd. ex Brid.) Mitt. and Dicranum polysetum w. e an n. w e eas s ecies c a ac e i in is ine i es s c asCarex limosa L., C. pauciflora Lightf., Drosera rotundifolia L., Sphagnum balticum ss w . . . ensenS. fallax . lin . . lin . S. fuscum c i . . lin . S. papillosum Lindb., S. rubellum Wilson and V. microcarpumwe e e lef in ina i n i . en i availa le a http://dx.doi.org/10.14214/sf.1462). Restoration seemed to have increased the covers and occurrence rates of mire species such as Sphagnum russowii Warnst., S. fallax and Polytrichum commune e w. in es e as seen ei l ca i ns in e alf f e ina i n i . en i .

n e f s ecies l ca e in e alf f e ina i n ave enefi e f ecendisturbances during restoration, e.g. Carex globularis L., Polytrichum commune, Sphagnum squar-rosum Crome and Straminergon stramineum ic s. e i . e en s i . en i .

i i nall e c ve f es eciall Eriophorum vaginatum L. seemed to have increased conside a l in es e i es ela ive aine an is ine i es al e e we e n eadifferences in its occurrence rate among the treatments (Appendix 1).

e sa e five a i i selec e envi n en al va ia les as in e ee s an s wesi nifican c ela i ns < . e e an c ela e si ila l wi e ve e a i n ina i n s aceas a ve als five a i i nal va ia les c ela e si nifican l wi e ina i n i .l w wa f s s an li e en e inc ease in c ve wa s e aine i es e ssesthan Sphagnum s . en e inc ease in c ve wa s e aine an es e i es an ec ve f e ic si e e a inc ease wa s e is ine i es an a f lawn wa s

is ine an es e i es. ene all e ina i n eveale a a ien f e we anfe ile c n i i ns c a ac e i e a i c ve f lawn an a ic si es ie an ec n i i ns wi a i c ve f c ic si es i . .

10

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Fig

. 1.

inain

ls

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11

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Fig

. 2.

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12

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Tab

le 3

. M

ean

(xan

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ccence

aes

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13

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

3.3 Ant occurrence among mire-site types

e an a a c ise s ecies w e s f s ecies eens f s ecies anw ales f ne s ecies en i availa le a http://dx.doi.org/10.14214/sf.1462). The most a n an s ecies in e w e a a we e Lasius platythorax eife Formica uralensis, Myrmica ruginodis lan e M. scabrinodis and F. sanguinea a eille an e sf e en l ca s ecies we e M. ruginodis (66% of the 162 sampling locations), M. scabrinodis (65%), L. platythorax Camponotus herculeanus (Linnaeus, 1758) (33%), and Leptothorax acervorum a ici s .

e s ecies an e a i i ave e s n es ine f es affini ies C. herculeanus and M. ruginodis we e e f e en in ansf in an ansf e an in is ine i e es w ea a a le . ecies an e ave e s n es i e affini ies s we e va ia le a e n F.

picea cc e al s e cl sivel in is ine i e es w e eas F. uralensis and M. scabrinodis we e f n f e en l in is ine an ansf in an ansf e i e es a le .e es f e s ecies see e cc e evenl a n e i e es a le . eens f

the forest species M. ruginodis we e e f e en in e ansf in an ansf e an ine is ine i e es w e eas eens f M. scabrinodis s we e si e a e n a le .e n e f all an s ecies i n iffe a n i e es w e eas e n e f i e an

s ecies was i e in is ine i e es an in ansf in an ansf e es an see eec ease wi inc easin win s c a le .

Table 4. Mean (x an e an al f cc ence a es f an w e s an eens an e ean n e ± SE, standard error given in the column fr) of mire ant species and all ant species in pristine, drained and restored mires. iffe ences in e cc ence a e f f an s ecies a n e ea en s we e es e wi l li eli a i es 2, f an iffe ences in e n e f s ecies we e es e wi s al allis an s es f ea en sn s a in e sa e le e iffe e si nifican l f eac e acc in a s e i i es wi c i ical . . ea s e e i inal val es a . in e a le c n l false isc ve a e in l i le es in sin e ein en a ini an e ieli . n e f sa lin l ca i ns al .

Species Treatment Test statistics

is ine aine es ex fr x fr x fr G2 p adj.

n w e s Formica picea 0.44 24 0.02 1 0.02 1 48.63 < 0.0001 Formica uralensis 0.07 4 . 0.08 5 3.76 0.6328 Myrmica scabrinodis 0.74 40 0.52 25 0.67 40 . . Lasius platythorax 0.35 0.50 24 0.60 36 7.13 0.1431 Formica sanguinea 0.22 12 0.13 6 0.23 14 2.44 1.0000 Leptothorax acervorum 0.30 16 0.15 7 0.20 12 3.52 0.6513 Myrmica rubra . 5 0.13 6 0.10 6 0.30 1.0000 Myrmica ruginodis 0.37 20 . 44 0.72 43 37.34 < 0.0001 Camponotus herculeanus 0.06 3 0.67 32 0.30 18 47.23 < 0.0001Ant queens: Myrmica scabrinodis 0.41 22 0.10 5 0.18 11 14.25 0.0061 Myrmica ruginodis 0.04 2 0.42 20 0.28 17 . < 0.0001 Camponotus herculeanus 0.11 6 0.02 1 0.20 12 . 0.0450Mean number of species: H p adj. Mire ants 1.3 0.1 a 0.8 0.1 b 0.8 0.1 b . < 0.0001 All ants 2.7 0.2 a 3.6 0.2 b 3.1 0.1 ab 10.23 .

14

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

3.4 The effects of drainage and restoration on ants

e ean n e f i e an s ecies was i es in is ine i es e n e f all ans ecies was i es in aine i es a le . e i e s ecies Formica picea occurred almost e cl sivel in e is ine i es an e eens f an e i e s ecies Myrmica scabrinodis, we e s f e en in is ine i es is a e n was n se ve f i s w e s a le .e w e s f e f es s ecies Camponotus herculeanus and M. ruginodis an eens f e

la e we e s f e en in aine i es an c n als in es e i es. cc ence a eea e in es e i es nl f C. herculeanus queens (Table 4). The occurrence rates of the rest f e s ecies i n iffe s a is icall si nifican l a n e ea en s a le .

f e nine s ecies anal se in ivi all wi ee we e an e a i i avee s n es i e affini ies an F. picea, F. uralensis and M. scabrinodis w we e an e

a i i ave e s n es ine f es affini ies an M. ruginodis and C. herculeanus), and e e ainin f s ecies e e ene alis s in ei a i a affini ies an L. platythorax,

F. sanguinea and rank 3: L. acervorum, species that occur both in mires and forests but disappear wi ee can cl s e in f es s ccessi n an an M. rubra innae s w icis n wn le a e ee can s a in . el w we f c s n e ve all en s f cc encef s ecies al n e i e f es c n in a e an s a is icall si nifican iffe ences in e

occurrence rate of individual species.

Fig. 3. a is ical es nses f in ivi al an s ecies i e ea en i e s ecialis s ecies ene alis s ecies f es s ecies . s a is icall si nifican . es nses see a le .

15

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

i e s ecialis s ecies s we va ia le es nses ea en i . a le . For-mica picea es n e s a is icall si nifican l an ne a ivel es e si es an e en fF. uralensis s es s ne a ive ass cia i n wi es e i es. n e es in l M. scabrinodis and F. uralensis en e ass cia e wi aine si es. e fai l ene alis s ecies L. platythorax es n e s a is icall si nifican l an si ivel es e si es an e en f F. sanguinea s es s a i ass cia es wi es e si es i . a le . Myrmica rubra behaved like the f es ass cia e s ecies w e eas L. acervorum responded more as a mire specialist (Fig. 3 B).

s e ec e e w f es ass cia e s ecies C. herculeanus and M. ruginodis, occurred s a is icall si nifican l e f en in aine si es less in es e si es an leas f e en l inpristine sites (Fig. 3 C).

Fig. 4. a is ical es nses el c efficien s see a le f in ivi al an s ecies envi n en al va ia lesSphagnum ss c ve n e f all ees c c ve . s a is icall si nifican .

es nses. ecies we e lis e a i i ase n e e ini n an e li e a e e s e s l inen e al.n ila an il el inen f e s i e ass cia e i e s ecialis s ecies a e f eac l

s ecies wi e s n es f es affini ies f es s ecies a e f eac l ene alis s ecies a e l ca ein e ween.

16

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Tab

le 5

. enealise

linea

ie

eles

lsfan

iesecialis

senealissan

fes

secies.

elesiaes

san

ae

san

vales

fe

seaine

ine

els

are

give

n. T

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17

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

n s es n e s a is icall si nifican l a n e f va ia les es e al e s ecies see e iffe in sensi ivi a le . nl e i e s ecialis M. scabrinodis responded si nifican l an si ivel Sphagnum moss cover, although there is a suggestive trend of mire s ecialis s ecies ein si ivel ass cia e wi Sphagnum cover and forest species the opposi e wa i . . e n e f all ees see e affec e a i f s ecies ase ec e a i i e a ili f cc ence f i e s ecialis s ecies en e ec ease ana f f es s ecies inc ease wi an inc ease in e n e f all ees i . . w s ecieses n e s a is icall si nifican l e n e f all ees as e ic e e i e s ecialis

M. scabrinodis ne a ivel an e f es s ecies C. herculeanus si ivel . ne f e s ecieses n e s a is icall si nifican l c c ve al e i e s ecialis F. picea and

the generalist L. platythorax a ea e es n si ivel an e f es s ecies M. ruginodis ne a ivel c c ve i . a le . i i nall a few e va ia les e aine ine final els f nl w s ecies e es nses we e s a is icall si nifican C. herculeanus

s we a ne a ive es nse ea ees an M. ruginodis es n e si ivel l w an allwa f s s an ann al lan s a le .

e an asse la e s c e f e aine i es iffe e f e is ine i es in eina i n ea en 2 0.221, p < . e sca e f e sa lin l ca i ns was

la e in eac f e ee ea en s an e l ca i ns f e es e i es ve la e wi seof both pristine and drained ones (Fig. 5 A). There seems to be a naturalness gradient from the sa lin l ca i ns f is ine i es e i wa s se f aine i es e lef wie l ca i ns f es e i es in e ween i . . even f e fi e envi n en al va i

a les c ela e si nifican l < . wi e ina i n s ace e c ve f Sphagnum sses an a f e ic si e e a inc ease wa s e is ine l ca i ns w e eas e

es f e s a is icall si nifican va ia les we e ela e aina e an c ela e a i a elin e si e i ec i n wa s e aine an es e i es i . . n asse la es iffe e f eac e s c a e e f es e i es see e e l ca e e lef an si ec ela i n wi all ees an e e en i es wi a n an l we ees e e f

the ordination.iffe ences in an asse la es a n e n n is ine i es we e e esse s c a e

s a e le an f es s eciesC. herculeanus and M. ruginodis cc e in e e eavil f es esa lin l ca i ns a e lef f e ina i n w e eas s ecies f en can f es san i es s c as L. platythorax, Leptothorax acervorum, Formica sanguinea and F. exsectac a ac e i e e less ensel f es e sa lin l ca i ns e i i . . en is inei es we e c a ac e i e e cc ence f e i e s ecies F. picea and M. scabrinodis, at the

bottom right of the ordination (Fig. 5 B). Another mire species, F. uralensis weve a enefie f aina e a leas e a il an is l ca e cl se e f es s ecies C. herculeanus

and M. ruginodis in ordination (Fig. 5 B). Indeed, half of the occurrences of F. uralensis we efrom drained mires (Table 4).

4 Discussion

n ene al es l s s we e ic a le es nses in a i a s c e incl in ve e ai n an an asse la es i e aina e an es a i n. is c ncl si n is si ila e es l sin a i e es a i n s ealin wi a n e f s li a inve e a e s ei a e al.

. a en e e es l s a e c nsis en wi e iel f ea s esis al e eal. w ic s es s a if e il e a i a s c e e a an fa na will e ne e i e es a i n es l e in a i ec ve f i e ve e a i n an an asse la es see als

18

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

Fig

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

ei a e al. . weve f i e s ecialis an s ecies e es l s we e va ia le an l n ee ni in is nee e eval a e e s ccess f es a i n. nsi e a le i e la ela ives li a insec s wi s e ene a i n i es is e ec e in e es nse f an s w ic f

l n live e ennial c l nies wi l n c l n c cles.

4.1 Effects of drainage and restoration on mire-habitat structure

es a i n eas es e fillin f i c es eav innin an a ial clea c in f ee s an swe e s ccessf l in a e wa e a le level an s an s c e f es e i es a c ese f is ine i es al eavie c in s f la e ees w l ave een e i e

acc e e wi is ine i es in e s f ee can s a in . e a n f ea w wasene all l w in all i es an e a n f s all l s was nl sli l i e in aine anes e i es an in is ine nes in ica in a ee ali e c e i i n a n es a e in aine i es an a c a se l in esi es felle ees we e n lef e in inrestored mires in large quantities.

e s we a ee s an s c e was e va ia le in e aine an in e is inees e a eas i . w ic f ll we a l f aina e an f es ana e en

ac ivi ies. n e l n n f es ana e en en s eni e s an s c e e. . ec easin va ia i n in s e si e is i i n w e eas f es s an s in is ine i es e i i l i als c e e a e al. .

e is ine ic si e e is i i n was e es a lis e a few ea s af e es a i n ei e s face was al s c le el c ve e e c ic si e e in aine i es

was e ce w i s in es e i es w e eas i was in is ine i es. l a i as c e f e es e i es a clea l c an e wa s is ine c n i i ns in a few ea s af erestoration, the cover of Sphagnum sses was s ill l we si ila l as ea s af e es a i nin aa ale e al. an e c ve f e. . all wa f s s was s ill i e in es e anis ine i es see i . in aine e al. a .

Drainage seemed to have led to similar successional changes of mire vegetation as in earlie s ies inc ease cc ence an c ve f f es s ecies an e isa ea ance eclinef i e s ecies aine e al. a a iainen e al. aa ale e al. aa ale

. i ila l a a i c an e wa s is ine is ic c si i n in es e i es as eense ve in ea lie s ies. e s en f e es nse e en s n e seve i f e a a i nf e aine i es an n i e fe ili . a i es nses ave een es ic e less eavile a e si es wi in es e i es w e eas n nl in c an es ave een e e feavil e a e si es lainen e al. a iainen e al. aa ale e al.aa ale . n a a e a i ec ve a ave een en ance a lac f is e sala ie s as e es e i es we e a s f la e i es wi va in i ns f n i c ea eas. ese li el vi e ef ia f i e s ecies in cl se i i f e aine an es ei es an s e als se ve as en ial a eas f ec l ni a i n f es e i es.

i ila l ea lie s ies s e s ecies e. . Eriophorum vaginatum es n e s n linc easin in c ve lainen e al. a iainen e al. aa ale e al. . ne e an s e s ecies f we ll ws e. . Carex limosa and Scheuchzeria palustris L.,

see e e issin f es e i es in e s e si ila l as ea s af e es a i n inaa ale e al. an in an e s ese s ecies see e n ave c l ni e es ei es even af e alf a cen es i e e e is ence f nea la i ns e al. . aine

et al. (2011) monitored the effects of restoration on vegetation of oligotrophic pine fens in a more n e n a ea in ain eas e n inlan f w ea s. e s we a e ve e a i n f ainei es a n c an e c f is ine c n i i ns in ea s f aina e nl e ical

20

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

ll w s ecies Sphagnum majus ss w . . . ensen S. balticum and Scheuchzeria palustris we e issin . aine e al. weve als f n a e e we e nl in iffe ences ine is ic c si i n ef e vs. af e es a i n in e w ea ni in e i e nl

s a is icall si nifican es a i n ela e c an e was a e c ve f se es inc ease in es eand decreased in pristine fens.

4.2 Ant occurrence in mire-site types

n wle e n e an fa na f i e es is li i e in nen e s . e s wee i ance f ee c ve n e cc ence f an s ecies a n i e es e cc encea es f i e s ecies en e ec ease wi inc easin win s c w e eas f es s eciess we e si e a e n a le .

anal sis f e a a f e s eveale a clea a ien in e c sition of ant assemblages from open fens and rich fens through pine mires and pine bogs to spruce i es n ila e al. . l ss f n a si ila enness a ien wi in an li

ic in e v is ic ssia. e an asse la es f e en cen al a s f ewe e c a ac e i e e en e ic i e s ecies F. picea, F. uralensis and F. forsslundi an e an s e f es s ecies L. platythorax, M. rubra, M. scabrinodis and L.

acervorum w e eas ine f es s f e i e e es we e cc ie f es s ecies C. hercule-anus, L. platythorax, M. rubra and F. polyctena s e . is enness a ien ese les

ina i n es l s i . w ic s we a asse la es a e en en f ea ien we e c a ac e i e e i e s ecies F. picea, M. scabrinodis and F. uralensis. On the e an e asse la es in s a sel f es e a eas wi l w ees we e is in is e s eciesa c l ni e ecen l is e na all en f es a eas e. . L. platythorax, L. acervorum, F.

sanguinea, F. exsecta . inall e e eavil f es e a eas wi all ees we e c a ac e i ef es s ecies a a e ca a le f e sis in e w le c cle f f es s ccessi n i.e.

C. herculeanus and M. ruginodis ie el e al. .

4.3 Drainage and ants

e s ccessi n f ee s an s n aine i es wi inc ease win s c f la e ees aninc ease eci s i e f i c wi ine es l in s a e c n i i ns. is c an e eci a ec l nies f i e an s ecies e i in s n e s e w ic evails in is ine c n i i ns f inei es w e eas s a e le an f es s ecies s c as Camponotus herculeanus and Myrmica rugi-

nodis enefi e si ila l as in ea lie s s ies an s e an asse la es f aine i esiffe e clea l f se f is ine i es e s llin w e s l inene al. n ila an il el inen . e i e s ecialis Formica picea in particular suffered from drainage and this is also expected in the long term for F. uralensis w ic see s enefif e ea l s a es f aina e an M. scabrinodis e s e s l inen e al. .

e s f n a F. picea inhabited the moistest parts of pine mires but F. uralensis the ies si es. n ila an il el inen f n a e cc ence a es f e ee nil in s ecies es ic e i es in ei a a a e i es in n aine is ine i es l we inansf in i es an l wes in ansf e nes F. uralensis), or the species are missing from

transformed mires (F. fennica Seifert, 2000 and F. forsslundi w e eas e s c n s eciesf f es s n ine al s ils e w an F. aquilonia a w s we e si e a e n.i ila es l s ave een aine in s e n we en w e e e sl w in f a eas

has decimated the prevalence of the mire species F. uralensis, F. forsslundi and F. picea in nlne eca e llin w .

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

n a a an s ecies f ie en a i a s e. . en ea l s ccessi nal f es s af e isances an ene alis an s ecies enefi e f e ea l ases f i e aina e. s f

ese s ecies isa ea e wi ee can cl s e als in f es s ccessi n n ine al s ilsn ila e al. in e ea l s a es f aina e e see e lace i e s ecies.

instance, M. ruginodis replaced M. scabrinodis e s l inen e al. al M. scabrinodis a ea e e e esilien an e i e s ecies see als llin w an es eciallLasius platythorax llin w l ss ssi l als Formica fusca Linnaeus, 1758 and F. lemani n i see e e lace F. picea, and in the later successional stages, F. aquilonia e lace e i ial n il in i e s ecies e s l inen e al. n ila anil el inen . en eeless i es s a sel f es e aine en i es n necessa

il vi e an a i a f w an s ecies f e Formica rufa group (F. aquilonia, F. lugubris Zetterstedt, 1838, F. rufa Linnaeus, 1758, F. polyctena and F. pratensis e i s eca sea eal a i c l nies vi in n is en f w an s a e sca ce an e f a in ac ivif w an s a cease f l n e i s f i e in i a i e e e a es in en i esan finall w an s cann fin s i a le el w n ve win e in si es ee en eca sef e i wa e a le level e s l inen e al. . n w e an aine i es e si a i nis iffe en w ic ena les c l ni a i n f w an s w en i e aina e cee s e i ialw an s ecies f ine al s ils als s a ina e ansf e i es n ila an il el inen

. n a a e w an s F. lugubris and F. pratensiswe e issin f is ine i esese s ecies cc e in seven an w sa lin l ca i ns in aine an in ee

an ne l ca i ns in es e i es es ec ivel en i .

4.4 Restoration and ants

e a e n awa e f an e s n e effec s f i e es a i n n an s wi e es l sf e esen s an e es l s f ea lie s ies c nce nin aine i es e e wiinf a i n a e a ec l c e i ive ca aci an is e sal f iffe en an s ecies wecan line s e ene al inci les e a en in acc n w en lannin i e es a i n.

e ewe in f i es an e val s n innin f e win ee s c a e ec nen s in s ccessf l es a i n f i e an s. i wa e a le level even s e ve wine in f c e i s f e e i ial Formica ants of heath forests in mire habitats and thus leaves e e ain f ee f ical i e an asse la es evel wi ei wn c e i sof other territorial Formica ants, such as F. uralensis, F. exsecta and F. forsslundi (see Punttila an il el inen .

n e i an iss e in i e es a i n f an s is e ea en f l in was e i.e.ee s e s s anc es an c s s af e l in in na all eeless s a sel f es ei es w e e ese a e lef e in a ves e n ila e al. .

es i e e fac a e a n f ea ees in s was ene all l w in all ea en se a n f s all l s was s ew a i e in aine an es e i es an in is ine nes

thus providing more nesting sites for forest ants there. Most of the boreal forest ant species can nes in even e i e ea w f es a lis in new c l nies an f an s ecies in ana i a es a la e s a e f an nes s a e l ca e in ea w e. . in nen an en inne

anc an s a ale n ila an aila a c e al. e ss n eal. . s leavin s s felle ees l in esi es in es e i es a en ancec l ni a i n f n n esi e f es s ecies in e es e i e a i a s. ea w vi eseleva e ic a i a s a a ffe e nl s n e se an wa a ve n nes in si esf f es an s. c c n i i ns a e f n in a i a s w e e ense s a in e la e ve e a i neven s s n e s e f e n . n f es e a i a s is a a en s l af e e c ea

22

Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

i n f ee can enin s win ws fi e c in s is a als a e lace af eis ances e. . i c in es a i n in i es.

l e a i l ec ve in ss la e f e es e i es a ve w l inesi es an c s s wi in a few ea s f ll win es a i n e neve eless vi e l iannes in s s a es f a la e va ie f f es an ene alis an s ecies. ese newc e s a la eeven sl w wn e c l n es a lis en f i e s ecialis an s ecies e a i n ess en e is e sin i e an eens a e in c l n f n in an nes si e c e i i nan e e i n f nes in si es s c i i effec s a e e e lifie e. . in e c l ni a i n

an s f s all lan lif islan s f e al ic ea see e s l inen an isa s i . esees f nes in s s a es a e issin al s en i el f is ine en i es an s ei

c ea i n s l e ini i e in e es a i n cess. n s ecies f en s ccessi nal f es sa als e ca a le f e lacin e few e ainin wea ene i e an la i ns f es e

mires. These forest species are also able to inhabit logging residues in the middle of larger restored i es as e is e sin fe ales f s e f ese s ecies a e a e s n ie s e s l inen anisa s i ec ws i e al. . e c l ni a i n f an a essive an e i ial For-

mica s ecies is e en en n e cc ence f s ecies in w se nes s e s a new c l niese a a asi is sswal e ils n llin w eife

ec ws i e al. . ese s ecies can f en ili e i e s ecialis s eciese ainin in e a ea an ene alis s ecies f ine al s ils w ic ave c l ni e e ainei e. s ei c l ni a i n a e en ance in a eas w e e i e an f es s ecies cce e llin w eife . n e i n f l in esi e a ves in is

i le e ees an le e ie s an in w ic e ces e a n f en ial ea w nes insi es f f es an s ecies s ea in e inc ease f l s n e n ve a l n e i e s an.

In restoration planning, the surrounding terrestrial environment should also be taken into acc n in i i ssi le c l ni a i n f f es an s in es e i es. es a i n eff s cane i e ea s w en e e a e n la e e ene a i n c in s n sa lin s an s in cl se

i i e es e a eas. ins ance in s e n we en e e a ica i n f Formica picea f sl wl in s was accele a e c l ni a i n ess e an s se en c e i ivee cl si n Lasius platythorax w ic i i a e f e s n in f es innin c in se f e a few ea s ea lie . n e c a ea e f es was c ve e l in esi esfallen ees an s n e se c s s. ea l all e c s s we e cc ie is inL. platythorax c l nies an ese an s we e als seen in la e an i ies in ss c s in ei es llin w .

n wle e n e is e sal ca aci an c e i ive a ili f i e an s is scansee llin w e s l inen e al. . s i is iffic l e ic e ec l ni a i nf es e i es i e an s. a en l is e sal a n iffe en a i a a c es win efe ales is a e li i e a leas in an l n s an s ecies an la i ns n s eal. e ene all e is e sal ca aci see s va wi el f ve s ndispersers (e.g. L. niger, C. herculeanus see in l nes see e s l inen an isa s i

. e e is evi ence a e is e sal f e i e an Formica picea is wea n nl e weeniffe en i es als wi in i es a elis an a n ees e al. . n a ais s ecies see e e al s e cl sivel c nfine is ine i es. e ec l ni a i n f F.

picea in es e i es a l e en s eavil n e cl seness f eal la i ns. isa e c i ical f es a i n eca se s ccessf l c l ni a i n f e ewe e i es F. picea is

a e e isi e f e s se en c l ni a i n f e e e i e s ecies F. forsslundi and F. uralensis w ic es a lis ei new c l nies e a a asi is in e nes s f F. picea.

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4.5 Habitat disturbance and ant colonization

a i a is ance f en ens new c l ni a i n ssi ili ies f eal an s in f es e a eas.is ances c ea e fav a le cli a ic c n i i ns f s ecies a e i e wa an well li

c n i i ns a leas in e c l ni a i n s a e. a e is ances in a e f es s als enc e i i n f ee e ain f en wi in e e i ial w an c l nies w e eas esec l nies see le a e s alle is ances n ila e al. n ila e al. n ila

n ila e al. va i an a a ainen il el inen e al. n ilaan il el inen va i e al. . n a a 60% of Camponotus herculeanus queens we e f n in e sa les f es e i es. es e i es weve s l e infe i a i af is w in a i in s ecies eca se s f e ees ave een e ve f e e ane c s s a e a l a e ic l ie n e e win i e sses. c c l ni ai n a e s a weve e e laine ssi le a ac iveness f ecen l is e ena eas f ll win es a i n eas es. l in Camponotus eens i ien a e wa s lie ec in f is e a i a a c es si ila l as an e an s ecies iann in ian e al. ils n an n wle sc in el . n f es e

a eas ecen l is e a eas ev i f e i ial w an s Formica rufa group) are presuma l i al a i a s f c l n f n in an i een n e s ave in ee een se ve inf es clea c s f ll win c in an esc i e nin n ila e al. n ila an aila

. ls Lasius platythorax a l enefi e f ecen is ances in es e i es ass es e a a.

is e f es e eal a eas a e ic l c l ni e an an s ecies e i in enc n i i ns in a leas e c l n f n in s a e an s e a i a ec es c w e s c ae nl i ns e ainin f c l n f n in a e nes a asi is an f eens in es a

lis e c l nies c l n in n ila e al. . c c w e c n i i ns weve seen e eac e f en in en is ine i es al ese a e l n live a i a s. wne e ience is a en is ine i es a e f en s a sel in a i e an c l nies. e eas nw a a en l s i a le nes in si es f en e ain nin a i e is n n wn es a l anla e an f ee e ain can e c ea e e e e even s s c as l n e in a wnse an c l nies seve e win e s wi in sn w c ve l we in e ve win e in s vival f

c l nies wil fi es in e ce i nall ea s. e c nness f s c even s i l i a ele e ine e nes ensi f e i e. f is we e e case l n an s se en inlea in l ines c l nies w l inc ease e esilience f i e an c l nies in e l n nas e is f e i a i n f an e ensive nes ne w s l e s alle an a f a sin lenest (Mabelis and Chardon 2005).

On the other hand, ant assemblages in drained mires in our data had become more c w e an in is ine i es as e se la i ns f f es an i e s ecies.Thus, in both pristine and drained mires, nest budding (leading to multinest colonies) and nes a asi is a e s li el as i an es f nes f n in as in s ccessi nal ealf es s. n a a e i f e enc f Myrmica eens M. scabrinodis in pristine and M. ruginodis in aine i es in ica e e cc ence f l n s c l nies w e e eeens a e vin ansfe e a n e nes s f l ines c l nies n ila e al.e e al. . a a we e nf na el ins fficien f a in an se va i ns nnes a asi es eca se sa lin win w was si e e is e sal e i f s s eciesw ic is in e la e seas n.

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4.6 Conclusions

ene all we es e w e e es in si le c cial a i a c a ac e is ics e wa e a lelevel i c fillin an e na all s a se an l w ine s an innin an clea c inw l lea ec l ni a i n an ec ve f e c a ac e is ic ve e a i n an an asse la esf is ine i e a i a s. e e we cann e eval a e l n e es a i n s ccesss e es l s a e c nsis en wi e ic i ns f e iel f ea s esis al e eal. w ic es n necessa il a l e eavil e a e i e ec s s e s. eec ve f s i es was fas eca se e aine i es s ill a an ac el e sface la e f i e s il an ca el c e f ea an i e anis s c l ec l ni e ees e i es ei e f l cal ef ia f la i ns f nea i es see e isc ssi nin lainen e al. asan e e al. . e c nnec e ness f i e a i a s weveas een seve el e ce es eciall in s e n inlan c nse en l ec l ni a i n asec e nli el f an i e s ecialis s ecies w ic ave ec e l call ea ene assie al. . c e an in s ecies in eal a eas a incl e e w e less s cial a asi eMyrmica karavajevi n l i an e e a a asi e M. vandeli n i .Both species require dense colonies of their host ant Myrmica scabrinodis f w ic es eciallwi in e eal ainlan a eas f inlan e s i an a i a is i es e s l inen eal. n ila e al. . e a a i n f i e a i a s an ei f a en a i n a lea

e isa ea ance f ese i l s eciali e a e s cial a asi es an e i e s ecialiss ecies f la e a eas. n inlan e ea is s ac e in e s w e e al s fi es ave een aine an e is ances e ween es e i es an is ine i es sinen ial s ce la i ns a e inc easin wi ec easin c nnec e ness w ic wea ens e

ec l ni a i n s ec s f s ecialis s ecies nce e ave isa ea e e s l inen e al.vinen e al. n ila e al. .

Acknowledgements

e a e a ef l ai a is as aa ale ei anen s v inen eis anen ni en inen ssi ivinen ee in ala aa i i il an nneli i i

f e s alli s a s il life inlan f e a al e i a e e vices ivi alinenof the Centre for Economic Development, Transport and the Environment of central Finland, and

e nen f e nive si f ens f a ici a in in e lannin f e s an fani in an ac icali ies s a in f s a ea selec i n. e an iisa a a i

a inen a i i n an a an iina i a f a ici a i n in e fiel w an i n nenf i en ifica i n f e an sa les an a ici a i n in e ea l s a es f a a cessin anli e a e sea c es. e als an ais a ala as a i is e a nen a ani allana s a i ee ia e inen an a iainen f e innis nvi n en ns i e fin is ensa le scien ific a vice an ac ical el in e c se f s . a a c llec inan an i en ifica i n we e f n e e s alli s a s il life inlan an e s iei es el n w a e ec s a ici a e e s alli s a elian i es an vi in

f es s ea ls in e c ain f e is an ec i n f val a le i ic we lan sin en al inlan .

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Silva Fennica vol. 50 no. 2 article id 1462 · Punttila et al. · The effects of drainage and restoration of pine…

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Supplementary filesAppendix 1 and 2 are available at http://dx.doi.org/10.14214/sf.1462.

Silva Fennica vol 50 no. 2 article id 1462, available at http://dx.doi.org/10.14214/sf.1462 Appendix 1 Mean percentage cover (x̄) and frequency (fr) of vascular plant and moss species in the sampling locations of pristine, drained and restored mires. Abbr. = abbreviations of the names used in the NMDS ordination plots. N = number of sampling locations (total N = 162). Differences in the covers among treatments were tested with Kruskal-Wallis rank sum test (H, df = 2; treatments not sharing the same letter differed significantly according to a posteriori test with critical α = 0.05; test statistics are given for species occurring in > 9% of the sampling locations). We adjusted the original p-values (p adj. in the Appendix) to control false discovery rate in multiple testing using the method in Benjamini and Yekutieli (2001). Species Abbr. Treatment Total Test statistics

Pristine (N = 54)

Drained (N = 48)

Restored (N = 60)

x̄

fr

x̄

fr

x̄

fr

x̄ fr

H p adj.

Andromeda polifolia L. Andpol 1.5 a 54

0.8 b 36

0.9 b 54

1.1 144

26.22 < 0.0001 Aulacomnium palustre (Hedw.) Schwägr. Aulpal 0.3 a 14

1.7 b 30

1.2 b 32

1.1 76

19.19 0.0009

Betula nana L. Betnan 1.3

42

1.8

33

2.5

51

1.9 126

4.51 0.5101 Betula pubescens Ehrh. Betpub 0.0

0

0.0

4

0.1

7

0.0 11

Calamagrostis arundinacea (L.) Roth Calaru 0.0

0

0.0

0

0.0

1

0.0 1 Calluna vulgaris (L.) Hull Calvul 4.4 a 26

2.7 a 13

2.2 a 17

3.1 56

8.12 0.0922

Carex brunnescens (Pers.) Poir. Carbru 0.0

0

0.0

1

0.0

0

0.0 1 Carex canescens L. Carcan 0.0

0

0.0

0

0.2

2

0.1 2

Carex chordorrhiza L. f. Carcho 0.0

1

0.0

0

0.0

0

0.0 1 Carex echinata Murray Carech 0.0

0

0.0

0

0.0

1

0.0 1

Carex globularis L. Carglo 0.0 a 1

0.7 a 12

0.4 a 10

0.4 23

11.62 0.0219 Carex lasiocarpa Ehrh. Carlas 0.1

6

0.0

3

0.0

1

0.1 10

Carex limosa L. Carlim 0.4 a 18

0.0 b 0

0.0 b 0

0.1 18

40.07 < 0.0001 Carex paupercula Michx. Carmag 0.0

0

0.0

0

0.0

2

0.0 2

Carex pauciflora Lightf. Carpau 0.2 a 36

0.0 b 3

0.0 b 11

0.1 50

57.46 < 0.0001 Carex rostrata Stokes Carros 0.1 a 17

0.0 b 0

0.0 b 1

0.0 18

34.19 < 0.0001

Chamaedaphne calyculata (L.) Moench Chacal 0.7

24

0.6

19

0.7

29

0.7 72

0.48 1.0000 Cladopodiella fluitans (Nees) H.Buch Claflu 0.1

3

0.0

0

0.0

0

0.0 3

Dactylorhiza maculata (L.) Soó Dacmac 0.0

1

0.0

0

0.0

0

0.0 1 Dicranum undulatum Schrad. ex Brid. Dicber 0.0 a 4

0.0 a 1

0.1 a 13

0.1 18

11.55 0.0219

Dicranella cerviculata (Hedw.) Schimp. Diccer 0.0

0

0.0

0

0.0

1

0.0 1

Dicranum fuscescens Sm. Dicfus 0.0

0

0.0

2

0.0

2

0.0 4 Dicranum majus Sm. Dicmaj 0.0

1

0.0

8

0.0

2

0.0 11

Dicranum polysetum Sw. ex anon. Dicpol 0.0 a 2

0.8 b 26

0.1 a 17

0.3 45

35.92 < 0.0001 Dicranum scoparium Hedw. Dicsco 0.0 a 0

0.3 a 10

0.0 a 5

0.1 15

13.37 0.0098

Drosera longifolia L. Drolon 0.0

7

0.0

0

0.0

0

0.0 7 Drosera rotundifolia L. Drorot 0.1 a 44

0.0 b 4

0.0 b 9

0.0 57

77.99 < 0.0001

Dryopteris carthusiana (Vill.) H. P. Fuchs Drycar 0.0

0

0.1

3

0.0

0

0.0 3 Empetrum nigrum L. Empnig 3.6

52

4.7

42

4.1

50

4.1 144

0.92 1.0000

Epilobium angustifolium L. Epiang 0.0

0

0.0

0

0.0

1

0.0 1 Equisetum palustre L. Equpal 0.0

0

0.0

0

0.0

1

0.0 1

Equisetum sylvaticum L. Equsyl 0.0

1

0.0

0

0.0

0

0.0 1 Eriophorum vaginatum L. Erivag 8.7 a 52

3.3 b 40

16.6 a 56

10.1 148

39.86 < 0.0001

Festuca pratensis Huds. Fespra 0.0

0

0.0

1

0.0

0

0.0 1 Hieracium Hieracium 0.0

0

0.0

1

0.0

0

0.0 1

Hylocomium splendens (Hedw.) Schimp. Hylspl 0.0

0

0.2

6

0.3

1

0.2 7 Juncus filiformis L. Junfil 0.0

1

0.0

0

0.0

0

0.0 1

Ledum palustre L. Ledpal 0.1 a 15

0.5 ab 14

1.3 b 28

0.7 57

9.12 0.0647 Lepidozia reptans (L.) Dumort. Leprep 0.0

0

0.0

0

0.0

1

0.0 1

Lycopodium annotinum L. Lycann 0.0

0

0.0

1

0.0

0

0.0 1 Melampyrum pratense L. Melpra 0.0

2

0.0

3

0.0

4

0.0 9

Melampyrum sylvaticum L. Melsyl 0.0

1

0.0

3

0.0

1

0.0 5 Menyanthes trifoliata L. Mentri 0.1

6

0.0

0

0.0

1

0.0 7

Molinia caerulea (L.) Moench Molcae 0.0

2

0.0

1

1.6

5

0.6 8 Mylia anomala (Hook.) Gray Mylano 0.3 a 24

0.1 a 11

0.3 a 15

0.2 50

8.13 0.0922

Orthilia secunda (L.) House Ortsec 0.0

0

0.0

1

0.0

0

0.0 1 Picea abies (L.) H. Karst. Picabi 0.0

0

0.0

0

0.1

3

0.0 3

Pinus sylvestris L. Pinsyl 0.2

26

0.0

13

0.1

23

0.1 62

4.36 0.5333 Plagiothecium Plagiot 0.0

0

0.0

4

0.0

1

0.0 5

Pleurozium schreberi (Willd. ex Brid.) Mitt. Plesch 1.3 a 20

31.7 b 43

14.4 c 47

15.2 110

61.24 < 0.0001

Pohlia nutans (Hedw.) Lindb. Pohnut 0.0 a 0

0.0 a 6

0.0 a 9

0.0 15

8.49 0.0846 Polytrichum commune Hedw. Polcom 0.0 a 0

0.6 a 9

1.6 a 12

0.8 21

11.96 0.0194

Polytrichastrum longisetum (Sw. ex Brid.) Pollon 0.0

0

0.0

1

0.0

1

0.0 2

G.L.Sm. Polytrichum strictum Menzies ex Brid. Polstr 1.1 a 29

2.9 ab 31

4.6 b 47

2.9 107

11.36 0.0230

Ptilidium ciliare (L.) Hampe Pticil 0.0

1

0.0

0

0.0

0

0.0 1 Rhynchospora alba (L.) Vahl Rhyalb 0.2

4

0.0

0

0.0

0

0.1 4

Rubus chamaemorus L. Rubcha 2.5 ab 42

4.0 a 39

2.1 b 38

2.8 119

7.26 0.1334 Salix myrsinifolia Salisb. Salmyr 0.0

0

0.0

2

0.0

0

0.0 2

Scheuchzeria palustris L. Schpal 0.0

4

0.0

0

0.0

0

0.0 4 Sorbus aucuparia L. Sorauc 0.0

0

0.0

1

0.0

0

0.0 1

Sphagnum angustifolium (C.E.O.Jensen ex Russow) C.E.O.Jensen Sphang 21.7

41

18.7

42

18.7

52

19.7 135

0.31 1.0000

Sphagnum balticum (Russow) C.E.O.Jensen Sphbal 8.7 a 24

0.0 b 0

1.1 ab 15

3.3 39

29.93 < 0.0001

Sphagnum capillifolium (Ehrh.) Hedw. Sphcap 0.0

0

0.1

1

0.0

1

0.0 2 Sphagnum cuspidatum Ehrh. ex Hoffm. Sphcus 0.0

1

0.0

0

0.3

3

0.1 4

Sphagnum fallax (H.Klinggr.) H.Klinggr. Sphfal 12.9 a 20

0.0 b 0

4.6 a 21

6.0 41

22.80 < 0.0001 Sphagnum fuscum (Schimp.) H.Klinggr. Sphfus 30.3 a 46

7.7 b 20

11.4 b 39

16.6 105

33.11 < 0.0001

Sphagnum girgensohnii Russow Sphgir 0.0

0

0.1

1

0.0

1

0.0 2 Sphagnum magellanicum Brid. Sphmag 4.3 a 44

2.9 ab 30

2.1 b 42

3.1 116

7.45 0.1254

Sphagnum majus (Russow) C.E.O.Jensen Sphmaj 2.9

9

0.0

0

0.2

2

1.0 11 Sphagnum papillosum Lindb. Sphpap 7.5 a 20

0.0 b 0

1.1 b 4

2.9 24

32.39 < 0.0001

Sphagnum pulchrum (Lindb. ex Braithw.) Warnst. Sphpul 0.2

2

0.0

0

0.0

0

0.1 2

Sphagnum riparium Ångstr. Sphrip 0.0

0

0.0

0

0.0

1

0.0 1 Sphagnum rubellum Wilson Sphrub 0.6 a 17

0.0 b 1

0.0 ab 4

0.2 22

23.24 < 0.0001

Sphagnum russowii Warnst. Sphrus 0.7 ab 21

1.2 b 12

6.4 a 29

3.0 62

8.44 0.0846 Sphagnum squarrosum Crome Sphsqu 0.0

0

0.0

0

0.0

3

0.0 3

Sphagnum tenellum (Brid.) Pers. ex Brid. Sphten 0.1

9

0.0

0

0.0

1

0.0 10 Straminergon stramineum (Dicks. ex

Brid.) Hedenäs Strastr 0.0

0

0.0

0

0.0

5

0.0 5 Trichophorum cespitosum (L.) Hartm. Trices 1.6 a 13

0.0 a 2

0.0 a 0

0.6 15

22.04 < 0.0001

Trientalis europaea L. Trieur 0.0

0

0.0

1

0.0

4

0.0 5 Vaccinium microcarpum (Turcz. ex Rupr.)

Schmalh. Vacmic 0.1 a 46

0.1 b 23

0.0 b 37

0.1 106

24.02 < 0.0001 Vaccinium myrtillus L. Vacmyr 0.0 a 3

2.7 b 19

0.6 ab 10

1.0 32

20.41 < 0.0001

Vaccinium oxycoccos L. Vacoxy 0.2

51

0.3

37

0.2

53

0.2 141

0.67 1.0000

Vaccinium uliginosum L. Vaculi 1.6 a 37

3.2 ab 35

5.0 b 48

3.3 120

9.26 0.0635 Vaccinium vitis-idaea L. Vacvit 0.1 a 4

5.7 b 26

0.8 b 21

2.0 51

30.41 < 0.0001

Silva Fennica vol 50 no. 2 article id 1462, available at http://dx.doi.org/10.14214/sf.1462

Appendix 2 Total abundance (ab) and occurrence rate (fr) of ant species collected in the pristine, drained and restored mires. N = number of sampling locations (total N = 162). Species Treatment Total

Pristine (N = 54)

Drained (N = 48)

Restored (N = 60)

ab fr ab fr ab fr ab fr Ant workers: Camponotus herculeanus (Linnaeus, 1758) 6 3 125 32 42 18 173 53 Formica exsecta Nylander, 1846 187 3 3 1 306 1 496 5 Formica fennica Seifert, 2000 0 0 0 0 25 1 25 1 Formica fusca Linnaeus, 1758 1 1 3 3 12 5 16 9 Formica lemani Bondroit, 1917 1 1 4 3 0 0 5 4 Formica lugubris Zetterstedt, 1838 0 0 127 6 3 3 130 9 Formica picea Nylander, 1846 200 24 1 1 2 1 203 26 Formica pratensis Retzius, 1783 0 0 7 2 67 1 74 3 Formica sanguinea Latreille, 1798 795 12 74 6 341 14 1210 32 Formica uralensis Ruzsky, 1895 600 4 1500 9 374 5 2474 18 Harpagoxenus sublaevis (Nylander, 1849) 0 0 1 1 0 0 1 1 Lasius platythorax Seifert, 1991 645 19 772 24 1301 36 2718 79 Leptothorax acervorum (Fabricius, 1793) 28 16 15 7 36 12 79 35 Myrmica lobicornis Nylander, 1846 0 0 6 4 3 1 9 5 Myrmica rubra (Linnaeus, 1758) 10 5 69 6 18 6 97 17 Myrmica ruginodis Nylander, 1846 107 20 953 44 548 43 1608 107 Myrmica scabrinodis Nylander, 1846 723 40 195 25 343 40 1261 105 Total abundance of workers 3303 3855 3421 10579 Total number of species, workers 12 16 15 17 Ant males (M) and queens (Q) Camponotus herculeanus M 2 2 0 0 0 0 2 2 Camponotus herculeanus Q 10 6 1 1 19 12 30 19 Formica aquilonia Yarrow, 1955 Q 1 1 3 2 1 1 5 4 Formica lugubris Q 1 1 4 4 1 1 6 6 Formica pratensis Q 2 2 2 2 1 1 5 5 Formica rufa Linnaeus, 1758 Q 0 0 1 1 0 0 1 1 Formica uralensis Q 4 4 0 0 1 1 5 5 Lasius platythorax Q 0 0 1 1 0 0 1 1 Leptothorax acervorum Q 1 1 0 0 1 1 2 2 Myrmica lobicornis Q 0 0 1 1 1 1 2 2 Myrmica rubra Q 0 0 3 3 0 0 3 3 Myrmica ruginodis Q 2 2 26 20 23 17 51 39 Myrmica scabrinodis Q 35 22 7 5 15 11 57 38 Myrmica sulcinodis Nylander, 1846 Q 0 0 0 0 1 1 1 1 Total abundance of queens 56 49 64 169 Total number of species, queens 8 10 10 13 Total abundance of males 2 0 0 2 Total number of species, males 1 0 0 1 Total abundance of all ant castes 3361 3904 3485 10750 Total number of species, all castes 15 18 17 20