Anthropogenic transformation of ground beetle assemblages (Coleoptera: Carabidae) in Bialowieza Forest, Poland: from primeval forests to managed woodlands of various ages

2006·5.7 MB·

by  Jarosław Skłodowski

©EntomologicaFennica. 11 October2006
Anthropogenic transformation of ground beetle assemblages
(Coleoptera: Carabidae) in Bialowieia Forest, Poland:
from primeval forests to managed woodlands of various ages
JaroslawJ.W. Sklodowski
Sklodowski, J. J. W. 2006: Anthropogenic transformation ofgroundbeetle as-
semblages(Coleoptera: Carabidae)inBialowieZaForest,Poland: fromprimeval
forests to managedwoodlands ofvarious ages. —Entomol. Fennica 17: 296—
314.
The interference ofmanintheprimeval forestofBialowieZa(e.g. clear-cutting
andreforestingfertilehabitatswithpineorspruce)bringsaboutchangesincara-
bidassemblages, including areductionintheproportionofindividuals ofrelict
species,reductioninthemeanindividualbiomass(MIB)andinthesumofposi-
tivecharacteristics(SPC;asummaryindicator).Humanimpactinforestecosys-
temscauseslossesofsomerelictspeciesandagainofubiquitousspecies.Icom—
paredthecarabidfaunasofpristineforeststothoseofvariousstagesofclear-cut
originatedsuccession. Clear-cutting causedadrastic decrease inMIB, areduc-
tionintheproportionofbrachypterousspecies,andanincreaseinboththenum-
berofhygrophilous species andtheirproportionintheassemblage.
J. Sklodowski,DepartmentofForestProtectionandEcology, SGGW— Warsaw
Agricultural University, Nowoursynowska 159, 02—766 Warszawa, Poland; E—
mail: [email protected]
Received: 12December2005, accepted23April2006
1. Introduction huntingbyrulingfamilies.Bythefourteenthcen-
tury,theforestwassetasideasaroyalforest,and
Theincreasingresearchinterestonprimevalnat- bythesixteenthcentury300royalwardenswere
uralforestsisreflectedbothinthenumberofpub- employedthere. Theirmaintaskwastokeepthe
lications(e.g.,Rackham 1980,Fosteretal. 1996, forest free fromunwanted intruding, illegal log-
Assmann 1999, Bobiecetal. 2000,Bircks2005) ging andpatching(Wesolowski2005).
andinthe attempts to identify anddefine forests Economic use ofpart ofthe BialowieZa Pri-
resemblingnatural ones, atleastpartially. These meval Forest began as late as the 1920’s. After
forests are referred to as “ancient” or “recent” WorldWar 11, only about 600 km2 ofthe entire
(Assmann 1999) or “semi-natural woodlands” 1,500 km2 BialowieZa Forest were left within
(Pooleetal.2003).BialowieZaPrimevalForestis Polishterritory; therestnowbelongsto Belarus.
theonlylowlandforestareaintemperateEurope The protected zone in the Polish part ofBialo-
where large tracts ofnatural forests, with their wieZaForestincludes anationalparkandnature
typical communities, still remain (Wesolowski reserves, whiletheremainingforests are stillex-
1995, Bobiec etal. 2000). Perhaps surprisingly, ploited for economic purposes and exemplify a
thekeytothemaintenanceofthenaturalnesswas varietyoftypesandintensitiesofanthropicpres-
ENTOMOL. FENNICAVol. 17 ° Anthropogenictransformation ofcarabidassemblages 297
sure. ByanthropicpressureIunderstandanyhu- forestsrespondsto logginginasimilarmanner.
man activity (irrespective ofits temporal or spa- Carabidassemblagesofmature,primevalfor-
tialscale)thatcauseschangestoanecosystemor estsarecharacterizedbyapredominanceoflarge
itsconstituents. Thesechangesareoftendeleteri- andforest-associatedspecies.Theseassemblages
oustothesystem. Thepresenceofbothnaturally include relict species that may not tolerate log-
regenerated and managed forest stands in this ging (Assmann 1999). In this paper, I study the
areaprovidesanopportunitytotracethetransfor- ecologicaleffectsofloggingoncarabidsbycom-
mationofnaturalwoodlandintomanagedforests paring faunas of primeval forests to those ob-
thataretheprevalenttypeofforestatpresent. tainedfromclear-cutoriginated, regenerating (=
Anthropicpressureintimberforestsmaytake timber, ormanaged) forests. Thekey questionis
onvarious forms, fromconstructionoftranspor- tostudytheeffectofclear-cuttingonpristinefor-
tationroutesandsettlementareastoclear-cutting ests, which is ofconsiderablepracticalvalue for
fortimberproduction, orforcreationofopenar- nature conservation. Moreprecisely, I study the
eas forvarious otherpurposes. Perhaps themost followinghypotheses:
profound impact is that of clear-cutting, after
which local macrofaunal assemblages take sev- (1)Clear-cuttingofanaturalforeststand, evenif
eraldozensofyearstoregenerate(Szujecki 1971, limitedinarea,resultsinchangesinthenum-
Szujecki etal. 1983, Szyszko 1983, Sklodowski ber of species, species composition and the
1995, 1997). As naturally regenerated forest structure ofcarabid assemblages (e.g., a sta-
stands arebecoming managed(=timber) forests tistically significant decrease in the propor-
(Wesolowski2005),thequestionariseswhethera tion of relict and hygrophilous species and
clear-cut originated stand can be restored meanindividualbiomass);
(throughnaturalre-growthor)bysimplyplanting (2)Treere-growthofapreviouslyclear-cutstand
anewgenerationoftreesandkeepingthecutting supportsthe(presumablyslow)restorationof
limited in the area. To study the forest-commu- the forest ecosystem, including carabid as-
nityresilience,Iusedgroundbeetles(Carabidae) semblages. However, regenerated assem-
asbioindicators ofenvironmentalchange. blages maybe structurally differentfromthe
Negative effects of clear-cutting on the assemblages ofprimevalforests.
macrofauna, and carabid beetles in particular, (3)Ialsoexaminewhetherforesttype(orsoilfer-
have been earlier reported by Szyszko (1983), tility) ofprimeval habitats has an impact on
Szujecki etal. (1983), Sklodowski (1995),Atle- the structureandresilienceofcarabidassem-
grim et al. (1997), du Bus de Wamaffe and blages.
Lebrun(2004),Koivula(2001,2002),Niemelaet
al. (1993, 2003) and Pontégnie et al. (2005).
These papers reveal that small- (e.g., an area of 2. Material and methods
1,600m2)andlarge-scaleclearcutting(uptotens
orhundredsofha)resultinanincreaseinspecies 2.1. Studyarea and carabidsampling
diversity accompaniedby an influx orincreased
abundanceofspecies associatedwithopenhabi- Thestudywaspartofalargerresearchprojecton
tats and forest-habitat generalists; however, for- the entomo-fauna of the BialowieZa Primeval
est specialists may drastically decrease or even Forest(Szujeckietal. 2001).Thestudywasdone
disappear.Atlegrimetal. (1997)pointedoutthat inthePolishpartoftheBialowieZaPrimevalFor-
single-tree cutting did not cause such profound est(northernPoland)intwostages.First,in1999,
changesincarabidassemblages.Becausesmaller carabid assemblages were studied in the pro-
speciesoftenprevailinloggedareas,theabsence tectedparts ofthe Primeval Forest (Sklodowski
oflargerspecieslogicallymeansareducedmean 2003, 2005). Intotal, 163 plotswere selectedfor
individualbiomass (MIB) ofthe carabid assem- thatstudy.Atthesecondstage, in2003,90 study
blage(Szyszko 1983).Alltheabove-listedresults plotswere setup in stands subjectedto different
wereobtainedinmanagedforests. However, itis degreesofanthropicpressure.Inbothstages,five
lessknownwhetherthecarabidfaunaofprimeval Barber’s pitfall traps were installed at each plot
298 Skiodowski ' ENTOMOLFENNICAV01. 17
Table 1. Studiedforesthabitatsin BialowieZa Forestandtheirabbreviationsused intext. P=primevalforest; C,
MC, MD, Dand DDrefertoforesthabitattypes(seetext); numbers(5, 15,40, 100)refertostandage(years
sinceclear-cutting). p=planted pineands= plantedsprucemonoculture; C=“Centuryf’forest.
Mesic Mesicmixed Mesicmixed Mesic Damp
coniferous coniferous deciduous deciduous deciduous
Primeval P_C P_MC P_MD P_D P_DD
5-y-old plantation C5 MC5 MD5 — DD5
15-y-old plantation C15 MC15 MD15 — DD15
40-y-old plantation C40 MC4O MD4O — DD4O
40-y-old pine monoculture — — MDp D40p —
100-y-old pine monoculture — — MD100p D100p —
100-y-old spruce monoculture — — MD100s D100s —
Old “Century”stand — — MD_C D_C —
andservicedfourtimesatsix-weekintervals,be- was studiedinanalogousforestbiotopes(C,MC
ginningon1May.Fivetraps(0.51glassjarwitha MD,DandDD).IntheC,MC,MDandDDhabi-
plasticfunnel, 12cmindiameter,containing 100 tats, carabidbeetles were studied in 5-, 15- and
ml 70% ethylene glycol) were arranged 15 m 40-years-old plantations growing in areas that
apartalongeachtransect.Astherewere(163+90 hadbeen clear-cut earlier. Additional samplings
=)253plots,thetotalnumberoftrapswas 1265in weredoneinMD andDhabitatsin40- and 100-
bothparts ofthe study. years-old pine monocultures and 100-years-old
BialowieZaPrimevalForestconsistsofamo- monocultures ofspruce planted in clear-cuts of
saic of fertile habitats that have not been sub- previousmulti-speciesprimevalforest,andin80-
jectedtothedestructionandunificationtypicalof years-old stands growing in areas oflarge-scale
managedforests.Twolevelsofforestsoilhumid- clear-cutting carried out during 1927—1928 by a
itywereselectedtostudy:damp(wet;70—99%of Britishcompany“TheCenturyEuropeanTimber
the soil surface under water during spring) and Corporation”, designated here as “Century”.
mesic (fresh; no inundation at any time of the Each variant ofanthropic pressure in the study
year).Withinthemesichabitat,trapswereplaced habitatsC,MC,MDandDDwasreplicatedthree
in four different forest stand types: coniferous times.
(low fertility habitats with a predominance of
Piceaabies,PinussylvestrisandLarixdecidua),
mixedconiferous, mixeddeciduous, anddecidu- 2.2.Ecological-historicaltypes
ous.Thelattertworepresenthighfertilityhabitats ofcarabidbeetles
with a predominance ofbroad-leaved tree spe-
cies. In the mixed coniferous stands, coniferous Foranalyticalpurposes,thecarabidspecieswere
tree species constituted 51—90% of all trees, dividedintovariousgroups(Table2).The“forest
whereas inthe mixed deciduous stands 51—90% relict” species group comprises species that are
ofthethreeswerebroad-leaved. predominantly found in forests and reproduce
The followingprimeval standtypeswere se- there but are seldom or never observed outside
lected for investigation in the mesic habitat: co- closed-forest stands. Species were assigned to
niferous (C), mixedconiferous (MC), mixedde- this group onthebasis ofearlierobservations in
ciduous (MD) and deciduous (D); in the damp Polish forests and literature (Burakowski et al.
habitat, deciduous stands (DD) were included 1973, 1974, Assman 1999, Turin 2000). A high
(Table 1). Each distinguished primeval forest proportionofrelictspeciesinanassemblagepre-
habitatwasrepresentedbyequalnumberofrepli- sumably indicates the presence of little or no
cates (12plots foreachhabitattype). anthropicpressureinagivenhabitat.
The effect ofanthropic pressure on carabids The “brachypterous species” group used in
ENTOMOL. FENNICAVol. 17 ° Anthropogenictransformation ofcarabidassemblages 299
Table2.ANOVAforcarabidassemblagesinfourforesthabitats.Symbolsused in pair-wiseposthoctests: 1 =C,2= MC, 3=
MD,4= DD,andinstandswithdifferentagesofdominanttrees: 1 =5-,2= 15-, 3=40-years—old, and4=(un-aged)primeval
forest(seeTable 1 forabbreviations).
Variable df MS F p LSDposthoctest
Speciesrichness
Habitat 3 59.347 6.535 0.001 1,2<3, 4(p=0.001)
Age 3 18.407 2.027 0.118
Habitat xAge 9 4.938 0.543 0.837
Relicts
Habitat 3 3146.063 14.687 <0.001 4<1,2<3(p<o.oo1);2,3>1(p=o.030)
Age 3 1835.366 8.568 <0.001 4>1,2, 3(p=0.010top<0.001)
Habitat ><Age 9 803.355 3.710 <0.001
M/B
Habitat 3 119846.800 19.914 <0.001 1< 3 (p <0.001); 4 < 1, 2, 3 (p< 0.001); 3 >2 (p< 0.001)
Age 3 105229.200 17.486 <0.001 4 >1, 2, 3 (p< 0.001)
Habitat xAge 9 24089.300 4.003 0.003
SPC
Habitat 3 15574.560 8.743 < 0.001 4< 1,2, 3(p<0.001)
Age 3 5904.950 3.315 0.024 2 > 3 (p = 0.030); 3 <4 (p = 0.003)
Habitat xAge 9 6792.380 3.813 0.001
Hygrophilous
Habitat 3 8440.183 34.025 <0.001 4>1,2,3(p<0.001)
Age 3 1107.472 4.465 0.006 4 <1 (p =0.010); 4 <2 (p =0.003)
Habitat xAge 9 271.367 1.094 0.379
Brachypterous
Habitat 3 3592.541 16.439 <0.001 4<1,2,3(p<0.001)
Age 3 709.437 3.246 0.027 4 > 3 (p = 0.006)
Habitat ><Age 9 999.940 4.576 <0.001
theanalysiswasdistinguishedonthebasisofob- Holarctic species). The classification was based
servedwing lengthandwingmuscles. Brachyp- on observations by the author and Larsson
terous carabids have a low power ofdispersal, (1939),Burakowskietal. (1973, 1974), Szyszko
which makes it difficult for them to leave the (1983), Szujecki etal. (1983) and Turin (2000).
ploughedclearingsquicklyortore-colonizethem Thereasonfortheinclusionofthesefourgroups
lateron. Hence, ahighproportionofbrachypter- in the analyses is their predominance in carabid
ous species in a community indicates an undis- assemblages in mature forests: an abundance of
turbedhabitat(Riberaetal. 2001). these species indicates the recovery ofthe forest
”Hygrophilous species” were classified on ecosystem (Szyszko 1983, Szujecki etal. 1983,
the basis ofearlier observations and data from Sklodowski 1995).
Burakowskietal. (1973, 1974)andTurin(2000).
Ahighproportionofthesespeciesinahabitatin-
dicates highmoisture levelsbutmayalsobedue 2.3.Data analysis
toconsiderableshadingorthepresenceofcoarse
woodydebris andotherfactors. Carabidindividuals were determinedto species,
The SPC index used here (see section 2.3.) andtheirbodylengthwasmeasuredwithanaccu-
was based on four different species groups: au- racy of0.5 mm. This measurewas subsequently
tumn breeders, large zoophages, forest species converted to biomass according to the formula
and European species (i.e., those with limited proposedby Szujeckietal. (1983). Themeanin-
geographical ranges, comparedto Palaearctic or dividualbiomass(MIB)isthequotientofthesum
300 Skiodowski ' ENTOMOL. FENNICAVol. 17
ofbiomassofallindividualsinanassemblageby (terBraak& Smilauer 1997—1999).
theirabundance(Szyszko 1983, 2002). The sum Thehypothesis ofslowregenerationofcara-
ofprogressive characteristics [SPC; Sklodowski bidfaunainregeneratedforeststandswas tested
(1997)] was also used. This metric is correlated usingtwo-wayANOVA [Statistica; StatSoftInc.
withtheageofthestandinquestion [r=+0.93,p (1997)]. The comparisonwas carried out in two
=0.0001,n=76; Sklodowski(1997)] andiscal- habitattypes:MDandD(bothwith24replicates)
culatedas follows: in40- and 100-years-oldpine and spruce stands
and in 80-years-old regenerated broad-leaved
SPC=74.9+ 102 * LOG(standage) (1) stands. Equality ofvariances was verifiedusing
Levene’s test. Aposthoc comparisonofsignifi-
SPC is thus the sum of proportions of those cantdifferencesinANOVAwasconfirmedusing
groups ofcarabids that are dominant in mature the LSD (least significant difference) test. The
foreststands: autumnbreeders, largezoophages, age variants ofregenerated stands (40-, 80- and
forest species andEuropean species (see section 100-years-old)involvedintheslow-regeneration
2.2). Thetwovaluesarecoefficients: 74.9forthe hypothesis were included in Ward analysis of
modelinterceptand102fortheregressionslope. species similarity and DCA gradient analysis.
Priorto the statisticalanalyses, the datawere Similarly, I studied whether the structure and
tested for normality of distributions using abundance of carabid assemblages depends on
Shapiro-Wilk’s statistic [Statistica; StatSoft,Inc. the fertility of the primeval habitats subject to
(1997)]. Species data were log (n) * 100 trans- anthropicpressure.
formedtoimprovenormalityofthedistributions.
Species richness was standardized to the lowest
number of specimens in the samples using 3. Results
Simberloff’srarefactionformula(Krebs 1999).
Two-way ANOVA [Statistica; StatSoft Inc. The first stage ofthe study (year 1999) yielded
(1997)] was used to verify the impact ofclear- nearly 60,000 carabid individuals representing
cuttingontheabundanceandstructureofcarabid 105 species,withafurtherca. 30,000individuals
communities.Datawerecomparedintwogroups. andadditional25speciescapturedduringthesec-
Thefirstcomparisonwasamongareasrepresent- ondstage(year2003). Thus, thetotalnumberof
ingdifferenthabitattypes(C,MC,MDandDD), specieswas 130, accountingforone-fourthofall
with 12 replicates fortheprimeval forests and9 carabid species ever recordedwithinthebound-
replicates foreachofthe differenttypes ofman- aries ofPolandsinceWorldWarII(Burakowski
aged(regenerating)forests. Thesecondcompari- etal. 1973). 50,235 individualsrepresenting 109
sondistinguisheddifferentstandages:5-, 15-and specieswereanalysed.
40-years-oldstands(eachwith 12replicates)and
“indefinite” (primeval) stands (48 replicates).
Equality of variances was verified using 3.1.Effectofclear-cuttingontheabundance
Levene’s test. Aposthoc comparison ofsignifi- and structureofcarabid assemblages.
cantdifferencesinANOVAwasconfirmedusing
the LSD (least significant difference) post hoc Significantdifferencesbetweencharacteristicsof
test.BothANOVAcomparisonsweredoneusing carabid assemblages from primeval forests vs.
datawith 1999and2003 catchespooledforeach those fromyoung, regenerating (5-, 15- and40-
standi.e. analyseswerebasedon standtotals. years-old)standswerefound(Tables2—3).How-
Species-composition similarities were com- ever,ANOVAdidnotindicatedifferencesinspe-
paredusing Ward cluster analysis based on Eu- ciesrichnessbetweenprimevalandregenerating
clidean distances [Statistica; StatSoft Inc. stands,althoughitshoweddifferencesinpercent-
(1997)]. To study the distinctiveness ofcarabid agesofrelictandhygrophilousspeciesandinthe
assemblages ofthe compared habitat types and valuesoftheSPCandMIBindices(Tables2—3).
standages, DetrendedCorrespondenceAnalysis Percentages of relict carabids varied from
(DCA)wascarriedoutusingCANOCOsoftware 15.8%, 36.7%, 44.0% and53.2% (mean38.0%)

ENTOMOL. FENNICAVol. 17 ° Anthropogenictransformation ofcarabidassemblages 303
Table3.ANOVAforcarabidassemblagesintwofertileforesthabitats.Symbolsused in pair-wiseposthoctests: 1 = MD,2=
MC, and instandswithdifferentagesofdominanttrees: 1 =40-,2=80-,3= 100-years-old,and4= (un-aged)primevalforest
(seeTable 1 forabbreviations).
Variable df MS F p LSDposthoctest
Speciesrichness
Habitat 1 1.176 0.103 0.750
Age 3 31.980 2.798 0.050 1 < 2 (p = 0.007); 3 <2 (p = 0,037)
Habitat xAge 3 21.532 1.884 0.147
Relicts
Habitat 1 9.849 0.094 0.760
Age 3 2352.114 21.546 <0.001 4> 1,2, 3(p<0.001)
Habitat xAge 3 495.497 4.749 0.006
M/B
Habitat 1 10017.000 0.895 0.350
Age 3 118475.200 10.584 <0.001 4>1,2,3(p<0.001)
Habitat ><Age 3 25398.900 2.269 0.094
SPC
Habitat 1 334.098 0.277 0.601
Age 3 6280.280 5.213 0.004 1> 2 (p = 0.008); 1 > 3 (p = 0.015); 1 > 4 (p = 0.003)
Habitat xAge 3 1086.107 0.902 0.448
Hygrophilous
Habitat 1 933.267 1.639 0.207
Age 3 3019.507 5.303 0.003 1>2,4 (p=0.001); 1 >3(p=0.010); 2, 3 >4 (p=0.001)
Habitat xAge 3 948.384 1.666 0.189
Brachypterous
Habitat 1 347.705 0.694 0.409
Age 3 1826.846 3.649 0.020 1> 2 (p < 0.001);1>4 (p = 0.030); 2 < 4 (p = 0.007)
Habitat xAge 3 91.362 0.183 0.908
val stands (51.0% vs. 15.6%; LSD test, p= 9.7) stands exceeded the species richness of
<0.001). Also, inthe assemblages inhabitingthe poorerprimeval C and MC stands (6.0 and 7.4;
100-years-old stands, the proportion of hygro- LSDtest,p<0.001; Table2). Theproportions of
philous species was significantly higher than in relict species in the carabid assemblages ofthe
theassemblagesfromprimevalforests(LSDtest, primeval DD habitats was significantly lower
p<0.001). Theproportionofbrachypterous spe- than in the primeval C, MC and MD habitats
cieswasalsolowerintheprimevalthaninthe40- (15.8% vs. 36.7%, 53.2% and 44.0%, respec-
years-oldstands(63.2%vs.77.8%;LSDtest,p= tively;LSDtest,p<0.0001).Alsotheproportion
0.03). ANOVA for the MIB revealed a signifi- ofrelict species was higher in the primeval as-
cant discrepancy ofthe community indices be- semblages offertile MC and MD habitats com-
tween the primeval habitats (297.8 mg/individ— paredtotheirproportioninpoorerprimevalhabi-
ual) and the 40-, 80- or 100-years-old stands tatC(53.2%and44.0%vs. 36.7%; LSDtest,p=
(227.9 [LSD test;p = 0.027], 169.2 [p <0.001], 0.001 and0.03, respectively).
231.8 [p=0.007], respectively). Theproportionofbrachypterous specieswas
alsolowerintheprimevalstandsDDcomparedto
theirproportionintheprimeval stands ofC, MC
3.3.Effectofsoilfertility
and MD (37.4% vs. 70.6%, 63.3% and 69.3%;
onforestcarabid assemblages.
LSDtest,p<0.0001)
Thenumbersofcarabidspecies(rarefactionstan- ANOVAforMIB (Table2) revealedsignifi-
dardized) in fertileprimevalD andDD (9.2 and cant differences in the community indices be-
304 Skiodowski ' ENTOMOL. FENNICAVol. 17
Table4. Dominantandco-dominantspecies,withdominanceindices>5%, inselected MMCand MMDhabitats.
Forcolumn names,seeTable 1.
Species P_MD MD5 MD40 MD100p MD100s MD_C P_MC MC5 MC40
C. hortensis 46.8 — 28.2 10.0 5.0 — 43.3 7.2
P. oblongopunctatus 18.6 — 17.4 13.8 34.2 18.5 10.2 — 21.8
C. glabratus 11.1 7.9 15.8 — — — 13.3 —
P. niger 10.5 17.7 11.4 46.0 27.5 33.0 6.7 21.1 20.6
C. arcensis 6.5 5.5 9.0 5.4 — — 13.2 — 10.1
E. secalis — 12.1 — — — — — —
P. versicolor — 9.6 — — — — — 11.9
H. quadripunctatus — 5.1 — — — — — —
C. caraboides — — — 5.4 — — — —
C. granulatus — — — — 7.0 — — —
P melanarius — — — — 6.1 23.0 — —
C. nemoralis — — — — — 6.4 — —
C. melanocephalus — — — — — — — 15.5 15.6
H. ruflpes — — — — — — — 7.7
tween fertile primeval MD habitats (with the and 100-years-oldpineandsprucemonocultures
highestMIB 303.1 mg/individual) andotherpri- alsoformedaseparateagglomeration,whichcan
meval habitats: MC (218.3), C (178.0) and DD be attributedbothto the considerable shading of
(86.6)(LSDtest,p<0.001).Ontheotherhand,in the litter andthe reduction ofits composition to
theassemblages inhabiting dampprimevalhabi- onecomponent,viz.pineorspruceneedles. Such
tats ofthe type DD, the SPC index was signifi- litterdecomposes more slowlythanmixedlitter.
cantlylowerthanintheassemblagesfromothers Theresultingthicknesswas about 10 cm,proba-
primeval stands oftypes C, MC andMD (184.3 blysupportinge.g.PterostichusnigerandP. me—
vs. 236.1, 242.0 and 260.0, respectively; LSD lanarius(Table4).
test,p<0.001). Also DCA separated the assemblages from
theprimevalstandsoftwoareasonthediagramat
variouslevelsofmoisture(Fig. 1).Thehorizontal
4. Discussion axiscouldberelatedtoanaltitudegradient.How-
ever, the arrangement of all assemblages sug-
4.1. Carabid assemblagevariation gestedagradientofanthropicpressureincreasing
amongthecomparedforesthabitats upwardalongtheverticalaxis,asthescoresofthe
assemblagesoftheyoungestplantationswerelo-
Cluster analysis indicatedfaunal differences be- catedatthe top. Considering also thenumberof
tweenprimevalandregenerating stands (Fig. 2). carabid species disappearing from the primeval
The carabid samples from the primeval stands standswhileharvesting, andthenumberofalien
weregroupedinto separate agglomerations. One species that colonize new plantations on clear-
included the assemblages inhabiting the fertile cuts, theresultis one ofprofoundchanges inthe
MC, MD andD habitats oftheprimeval stands. ecosystem.
However,theDprimevalhabitatassemblageap-
pear located among the regenerated stands sam-
pled within the “Century” area that otherwise 4.2.Effectofclear-cutting andlitterquality
formed a distinct cluster ofsamples. The other on carabid assemblages
group, consisting ofthe assemblages ofthepoor
C habitat and the marshy DD habitat, shared a Niemelaetal. (1993)distinguishedthreegeneral
branchtogetherwiththeyoung5-to40-years-old carabid responses to logging: (1) an increase in
plantations. The carabid assemblages in the 40- abundance of species characteristic of dry and
ENTOMOL. FENNICAVol. 17 ° Anthropogenictransformation ofcarabidassemblages 305
open conditions; (2) an initial decrease offorest (Kimmins 1997). However, according to Ba-
generalist species thatreestablishwith forestre- guette(1993),P. nigerandP. melanarz'usareas-
generation; and (3) canopy closure and disap- sociated with thick litter covering humic soils.
pearance ofsome forest specialists which seem Similar micro-habitat associations with litter
unable to re-establish populations in cut-over have been observed for C. hortensz's (Turin
stands (butsee Spenceetal. 1996). 2000)
In carabid assemblages of clear-cuts, forest Theaboveexplainswhythedestructionoflit-
specialists are apparentlyreplacedby non-forest ter and mosses during top-soil preparation in
(e.g.,AmaraandHarpalus)species,resultingina clear-cuts has negative effects onthe abundance
localincreaseinspeciesdiversity(Beaudryetal. ofe.g. C.hortensisandC.glabratus,whileitmay
1997).Openhabitatcarabids,forexample,areef- not affect the colonization of clear-cuts by P.
ficientincolonizingclear-cuts:thefirstindividu- niger, as showninarecentcapture-recaptureex-
als may colonize small(<0.5 ha) clear-cutopen- periment (Sklodowski 2002). This difference
ings within months (Koivula & Niemela 2003). probablyaccountsforthedominanceofthelatter
Contrary to these findings, the present study speciesincarabidassemblagesinthe5-years-old
showednochangeinthemeannumberofspecies plantations inBialowieza.
following clear-cutting. However, I found few Inspiteofthelackoflitter,P.nigerwasdomi-
non-forest species [P. versicolor, E. secalz's nant in the carabid assemblages of 5-years-old
(MD), C. melanocepnalus andH. rufipes (MC)] plantations,whereastheeurytopicP.melanarl'us,
in the group ofco-dominants in the carabid as- relatively similar interms ofbody size anddiet,
semblages ofthe 5-years-oldplantations. In the was not numerous there. Perhaps the more flat-
15- and 40-years-old plantations, these species tenedbodyofP.nigerallowsittoeasilypenetrate
had apparently been replacedby the forest spe- micro-crevices in furrows createdbyploughing.
cialistsC.nortensz's,C.glabratusandP.oblongo— The slightly stockier Pterostichus melanarl'us
punctatus(Table4). wasaco-dominantinthecarabidassemblagesof
TheforestgeneralistPterostichusnigerwasa mature stands plantedinthe “Century” area and
dominant species in the 5-years-oldplantations. inthepineandsprucemonocultures.Becausethis
However, it has also been reported as being a specieswasrecordedonlyinverylowabundance
dominantspeciesin40-years-oldandolderman- in the primeval stands, the higher abundance in
aged stands (Szyszko 1983, Sklodowski 1995). regeneratingareasapparentlyservesasanindica-
Its locomotor ability resembles those of the tor ofstand disturbance. However, Poole et al.
flightless C. hortensisandC.glabratusthatwere (2003) foundP. melanarl'us more abundantly in
presentinlownumbersintheregeneratingclear- semi-naturalbut“mature” oakwoodlandthanin
cuts. Studies ofmovement patterns of C. hor— oakplantations.
tensis and C. glabratushave indicatedthatthese
species do not enter open areas (Sklodowski
1999,2002,Assmann&Gunther2000).Further- 4.3.Relictandbrachypterouscarabids,
more, C. hortensz'sisassociatedwithhumic soils andclear-cutting
andC.glabratuswithabundantmosses(Niemela
etal. 1996).Also forestlitterisimportantforca- However, perhaps amore importantindicatorof
rabid beetles: litter thickness and composition destructionofprimevalstandsisthedepletionof
stimulate the occurrence of forest-associated relict species (Assmann 1999). Inthis study, the
carabids (Guillemain et al. 1997, Koivula et al. proportion of the ten relict forest species de-
1999,Koivula2001,Maguraetal. 2001,Koivula creased in BialowieZa Forest from 85% to 25%
&Niemela2003).Baguette(1993)associatedthe alongwith increasing anthropic pressure. Clear-
occurrence of P. oblongopunctatus with the cutting,evenifnotparticularlyintenseintermsof
“moder”typeoflitter,aforestfloortypeinwhich area covered, leads to the disappearance ofspe-
thereisadistinctlitteranddecomposinglayerand cies characteristic of older stands, such as L.
humus layer that gradually changes into, and is piceus, C. intricatusandC.glabratus.Inorderto
partly mixed with, the underlying mineral soil complete their life cycle, these species need a