Habitat associations and seasonal activity of carabid beetles (Coleoptera: Carabidae) in Dongling Mountain, North China

2006·0.16 MB·

by  Xiao-Dong Yu

©EntomologicaFennica.16June2006
Habitat associations and seasonal activity of carabid beetles
(Coleoptera: Carabidae) in Dongling Mountain, North China
Xiao-DongYu,Tian-HongLuo&Hong-ZhangZhou*
Yu,X.-D.,Luo,T.-H.&Zhou,H.-Z.2006:Habitatassociationsandseasonalac-
tivityofcarabidbeetles(Coleoptera:Carabidae)inDonglingMountain,North
China.—Entomol.Fennica17:174–183.
Habitatdistributionandseasonaloccurrenceofcarabidbeetlesweredetermined
usingpitfalltrapsin1999and2000inthetemperateforestzoneoftheDongling
Mountain,NorthChina.Eightsitesdifferinginvegetationandmoisturewerese-
lectedsoastorepresentfourhabitattypes.Carabidassemblagesofthesixfor-
estedhabitats(lowland,uplandandcoppice)weremoresimilartoeachotherthan
tothetwoshrubassemblages.Lowlandforesthadthehighestspeciesrichness,
andcoppiceforesthadthehighestdiversity(H’)andequitability(J).Ofthe41
speciescaught,the18mostabundantspeciesweredividedintofourdistribution
types:habitatgeneralists,forestgeneralists,forestspecialists,andshrub(orcop-
pice)specialists.MeancatchesofallbeetlesshowedclearpeaksfromMaytoAu-
gust in nearly all habitats. The catches of the six most abundant species were
moreorlesspositively correlatedduringthetwostudy years,suggestingtheir
similarhabitatpreferences.
X.-D.Yu,T.-H.Luo&H.-Z.Zhou*,InstituteofZoology,ChineseAcademyofSci-
ences, Beijing 100080, P. R. China; *correspondent author’s e-mail: zhouhz
@ioz.ac.cn
X.-D.Yu&T.-H.Luo,GraduateSchool,ChineseAcademyofSciences,Beijing
100039,P.R.China
Received8June2005,accepted22November2005
1.Introduction 1992,Niemelä&Halme1992).Previousstudies
havesuggestedthatthedistributionsofcarabids
Theheterogeneousdistributionofresourcesand are generally structured with respect to micro-
abioticconditionsaffectthespatialandtemporal habitatpatcheswithinforestsandevenwithinthe
occurrence of carabid beetles (Wiens 1976, movementrangeofindividualbeetles(Wallin&
Niemelä & Halme 1992, Niemelä et al. 1992). Ekbom1988,Epstein&Kulman1990,Niemelä
Associatedwithseveralscalesofenvironmental etal.1988,1992,Niemelä&Halme1992,Nie-
heterogeneity, species responses may be deter- melä&Spence1994).
minedbydifferentfactorsatthevariouslevelsof Thereexistmany quantitativestudiesonthe
environmental heterogeneity (Addicott et al. distributionofcarabidbeetlesinvarioushabitats
1987).Carabidbeetlesshowanon-randomspa- inEuropeandNorthAmerica,andassociationsof
tialdistributionbothwithinandbetweenhabitats particular species with broad habitat categories,
(Lindroth1961–1969,Thiele1977,Wiens1989, such as “forests” or “fields”, have been estab-
Epstein & Kulman 1990, Niemelä et al. 1988, lishedgenerallyforcarabidsinthecontextoftax-
ENTOMOL.FENNICAVol.17 (cid:127) HabitatassociationsofcarabidsinChina 175
onomicandfaunisticwork(Lindroth1961–1969, precipitation amounts to 611.9 mm, 78% of the
Thiele 1977, Esau & Peters 1975, Liebherr & annualrainfalloccurringbetweenJuneandAu-
Marhar1979,Epstein&Kulman1990,Niemelä gust(Chen&Huang1997).
&Halme1992,Niemeläetal.1992).However, Eight sites were sampled, each representing
for Chinese carabid species quantitative studies vegetation types commonly found in the area.
ofthewholeassemblagesarescarce,exceptfora FivesiteswerelocatedatBeijingForestryEco-
few studies on the ecology of agricultural envi- system Research Station (BFERS, ca. 114 km
ronments or taxonomy (Yu et al. 1980, Huang west of Beijing), at 800–1,600 m a.s.l. At that
1990,Huetal.1998).Studiesofassemblagesof area, oak forests (Quercus liaotungensis) repre-
forest carabids are particularly rare, although sentthemostextensiveforesttype,anativeup-
someexcellentstudiesonhabitatdistributionand landforestassociation.Insomelowlandandval-
molecular evolution, using particular species or ley sites, walnut forests (Juglans mandshurica)
groups ofcongeners, do exist (Kimet al. 2000, andmixedbroad-leavedforestsrepresentthetyp-
Yuetal.2001,2002,2004,Suetal.2003,2004). icalnativeforests,thelatterbeingmostlydomi-
Theexcessivelanduseduringrecentdecades nated by Acer mono, Betula dahurica, Juglans
hasresultedinadrasticdecreaseinthetotalforest mandshurica, Fraxinus rhynchophylla, Populus
cover and an increasingly skewed size distribu- davidiana, Quercus liaotungensis, Sorbus pau-
tionofforestremnants.Thisfragmentationpro- huashanensis,Tiliamongolica,T.mandshurica,
cessaccountsformuchofthelossofbiodiversity etc. In addition, in some areas where the native
andratesofspeciesextinctionallovertheworld forestshavebeenlogged30–40yearsago,non-
(Didhametal.1996).Thetemperateforestzone, native coniferous plantations of larch (Larix
alargephytogeographicalzoneinChina,hashis- principis-rupprechtii) and pine (Pinus tabulae-
torically been one of the most extensive forests formis)dominate.Atthetimeofthestudy,allof
within China. However, at present that area is thefiveforestpatcheswithintheBFERSareahad
heavilyimpactedbyhumans. aclosedcanopywithtreeheightof8–15m(upto
Comparedwithstudiesdoneatthesamelati- 20 m). The three other sites were located at
tudesworld-wide,NorthChinesestudiesonthe Liyuanling,ca.10kmSEfromBFERS,at800–
assemblages of carabid beetles in the temperate 1,100ma.s.l.Here,muchofthenativevegetation
deciduous broad-leaved forests are rare, and isoakforestthathasbeenclear-cutforfarmland
someofthespeciesmayhavebecomeextinctbe- useatleast50–60yearsago.However,duringthe
foretheriseofconcern.Thepurposeofthepres- 1990s, farming activities at the study area were
entstudywastogainbasicecologicalknowledge aborted and the forest vegetation was restored;
on factors affecting carabid beetle abundances. the former clear-cuts sprouted again and devel-
We examined the spatial and temporal distribu- opedtooakcoppicestandsthatcanberegardedas
tion of carabids in temperate forests of North youngforests(coppiceforests).Theuplandand
China,anddescribedthespeciescompositionof valleyareasattheLiyuanlingarea,ontheother
carabidassemblagesovermultiplehabitattypes. hand, are dominated by vitex shrubs (Vitex
negundo var. heterophylla) and almond shrubs
(Prunusarmenicavar.ansu).Allofthecoppice
2.Materialandmethods andshrubsitestostudyhadaclosedcanopy,with
theaveragetreeheightbeing2–3m(upto5m).
2.1.Studysite Correspondingtovariationinvegetationand
soilmoisture,eightplotswereestablishedinthe
ThisstudywasconductedattheDonglingMoun- core of each of the eight sites representing four
tain(40º00’N,115º26’E),whichisapartofthe habitat types: three upland forests (oak forest,
Taihang Mountain Ranges near Beijing, North larch plantation and pine plantation), two low-
China. Thestudy areaisassociatedwithmoun- land forests (walnut forest and mixed broad-
tain brown soil and temperate monsoon climate leaved forest), one oak coppice forest, and two
with average annual temperature being +4.8º C shrublands (dominated by vitex and almond
(January –10.1º C, July +18.3º C). The annual shrubs).Eachofthestudiedforestsorshrublands
176 Yuetal. (cid:127) ENTOMOL.FENNICAVol.17
Soilmoisture dry dry-medium dry-medium dry-medium dry medium-wet wet wet
ayers. Litter thin(<1cm)thin(ca.1–2cm) medium-thick(ca.5–8cm) medium(ca.3–5cm)medium(ca.3–5cm)medium-thick(ca.5–8cm) thick(ca.8–10cm)thick(ca.8–10cm) a,etc.
ntforeststructurall Herbaceous sparse(ca.10%)sparse-moderate(ca.30%) moderate-dense(ca.50%) sparse-moderate(ca.15%)sparse-moderate(ca.10%)moderate(ca.40%) moderate-dense(ca.60%)dense(ca.80%) mongolicaT.mandshuric,
e a
columnsrefertodiffer Shrub moderatetodense(ca.60%)dense(ca.80%) dense(ca.90%) sparse(ca.20%)sparse(ca.20%)moderate(ca.40%) moderate-dense(ca.60%)moderate(ca.50%) SorbuspauhuashanensisTili,
dLitter dense ngensis,
baceousan Canopy – – – moderate-(ca.60%)dense(ca.70%)dense(ca.70%) moderate(ca.55%)moderate(ca.45%) dianaQ.liaotu,
er avi
H a) sd
Table1.Characteristicsofeachplot(vegetationtype).Canopy,Shrub, Habitat/LocationSlope,elevationHabitatPlot ShrubsVitexVSuplandSE-facingslope,Vitexshrubs(negundoheterophyll15º,1,100mvar.PrunusASlowlandSW-facingslope,Almondshrubs(armeniacaansutoupland5º,1,070mvar.)Coppiceforest(youngforest)QuercusOClowlandNW-facingslope,Oakcoppices(liaotungensistoupland10º,1,050m)UplandforestsQuercusOFuplandSE-facingslope,Oakstand(liaotungensis25º,1,250m)PinusPPuplandSW-facingslope,Pineplantation(tabulaeformis20º,1,190m)LarixLPlowlandtoNW-facingslope,Larchplantation(principis-rupprechtiiupland20º,1,140m)LowlandforestsMFlowlandNE-facingslope,Mixedbroad-leaved2º,1,225mforest*JuglansWFlowlandSW-facingslope,Walnutforest(mandshurica5º,1,160m) AcermonoBetuladahuricaJ.mandshuricaFraxinusrhynchophyllaPopulu*Dominatedby,,,,
ENTOMOL.FENNICAVol.17 (cid:127) HabitatassociationsofcarabidsinChina 177
wasover4ha,andthedistancebetweenadjacent richness(S)wasthenumberofspeciesforsam-
siteswas1–2km.Thecharacteristicsofeachsite pleswithoneormoreindividuals(Pielou1975).
areshowninTable1. After logarithmic transformation, Detrended
CorrespondenceAnalysis(DCA)wasappliedto
comparethecommunitystructureamongdiffer-
2.2.Sampling entsites(Pielou1984)usingthePASTpackage
(Hammer et al. 2001). Associations between
Carabidbeetleswerecapturedusingpitfalltraps. abundant species were analyzed with non-para-
Althoughpitfalltrapsarebiasedtoactiveforms metric Spearman correlation (Sokal & Rohlf
andarethereforenotadirectmeasureofabsolute 1981)usingSPSS(1997).
populationdensity,thismethodisusefulinmoni-
toring and assessing local population changes
(Baars 1979, Spence & Niemelä 1994). Traps 3.Results
wereplasticbeveragecups(400ml,depth9cm,
mouthdiameter7.5cm)filledwithca.100mlof
3.1.Speciesrichnessanddiversity
trappingfluid(vinegar:sugar:alcohol:water–
10ml:5g:5ml:20ml)(Yuetal.2004).Asmall
Forty-onespeciesin14generaofcarabidswere
hole (diameter ca. 0.2 cm) was drilled on each
collected(Table2).Fourspecies,eachaccount-
trapca.2.5cmbelowthemouthtoallowrainwa-
ingfor>10%ofthetotalcatch,wereconsidered
ter to flow out. Because of that, some small
dominant:Carabusmanifestus,Carabuscrasse-
carabids (such as Bembidion and Trechus) may
culptus,PterostichusacutidensandPterostichus
have been lost from the samples. The collected
subovatus. These species altogether accounted
specimens were preserved in 70% alcohol for
for62%ofthetotalcatch.Twootherspecies,viz.
identification.
Pristosia sp.1 and Reflexiphodrus refleximargo,
SamplingwasconductedfromApriltoOcto-
accountedfor8%and7%oftotalspecimens,re-
ber(coveringthegrowingseason)in1999.Pitfall
spectively.Alltheotherspeciescollectedrepre-
trapswereplacedontwotransectsthatwereran-
sented<5%ofthetotalcatch.
domly arranged within each plot; however, the
Inbothyears,thelowlandforesthostedmore
distancebetweenadjacenttransectswasatleast
speciesthandidtheotherhabitattypes,whereas
100 m. Each transect had 55 traps placed 2 m
the shrubland sites had fewer species than the
apart. The traps were serviced for every 3 days
other habitat types. Moreover, diversity was
each month. We continued sampling using the
highest in the coppice forests and lowest in the
samedesignin2000duringMay,JulyandSep-
shrub sites. The catches varied species specifi-
tember.
cally among the studied habitat types and be-
tween the two years. However, in terms of
carabidspeciesabundances,thecatchesforboth
2.3.Dataanalysis
yearsweremoreevenincoppiceforeststhanin
theotherhabitattypes(Table3).
Theactivitydensityofcarabidsindifferentplots
(habitats)wasvisuallyexaminedusingtheaver-
age number (standardized catches to 100 trap-
3.2.Carabidcommunitysimilaritiesamong
days)ofbeetlescapturedperplot*trappingday.
thestudiedhabitats
Species diversity was calculated with Shan-
non-Wienerinformationindex(Pielou1975):
TheDCAindicatedstrongclusteringsofhabitat
(cid:2)s typesoverthetwoyears(Fig.1).Basedontheoc-
H'=– P lnP
i i currenceandabundanceofcarabidassemblages,
i(cid:1)1
twoshrubsiteshadsimilarcarabidassemblages
where P is the proportion of the sample repre-
i withineachyear;ontheotherhand,theothersix
sented by ith species (i = 1–S). Equitability was
(all forested) sites including five mature forests
representedbyJ=H’/H (Pielou1975).Species
max and 1 coppice forest formed another cluster of
178 Yuetal. (cid:127) ENTOMOL.FENNICAVol.17
Table2.Ecologicaldistributionof41speciesofcarabidbeetlescollectedinpitfalltraps,1999–2000.Thespecies
areclassifiedintofourtypes.
Species Shrubs Coppice Upland Lowland Total Per-
forests forests cent
VS AS OC PP OF LP WF MF (%)
Habitatgeneralists
Carabussmaragdinus 22 51 67 10 4 2 12 1 169 4.01
Carabusvladimirskyi 11 41 6 8 7 8 5 1 87 2.06
Pristosiasp.3 – 4 6 – 4 4 21 5 44 1.04
Forestgeneralists
Carabusmanifestus 8 1 152 42 233 355 229 68 1088 25.79
Reflexiphodrusrefleximargo – – 27 44 39 122 31 20 283 6.71
Pterostichusacutidens – 1 75 – 81 147 5 190 499 11.83
Pterostichussubovatus – 1 52 1 106 257 8 50 475 11.26
Pristosiasp.1 1 13 66 4 42 134 13 56 329 7.80
Pterostichusfortipes 5 6 39 9 32 1 33 5 130 3.08
Carabusscultipennis – 4 36 5 29 8 25 – 107 2.54
Carabuscrasseculptus – – 4 76 68 226 105 90 569 13.49
Carabussui – – 9 25 39 7 6 5 91 2.16
Pristosiasp.2 – 1 1 1 11 – 22 26 62 1.47
Forestspecialists
Carabuscanaliculatus – – – – 2 12 1 1 16 0.38
Pterostichusinterruptus – – – – – 18 – 3 21 0.50
Pterostichusadstrictus – – – – 5 – – 75 80 1.90
Shrubandcoppicespecialists
Carabusgranulatus – 7 59 – – 1 7 – 74 1.75
Carabusbrandti 10 3 10 – – – – 1 24 0.57
Otherspecies
Agonumnitidum – – – 2 – 1 – – 3 0.07
Amarasp.1 – 1 – 1 – 1 1 – 4 0.09
Amarasp.2 – 1 – – – – – 2 3 0.07
Asaphidionsp. – – – 1 – – – – 1 0.02
Badisterpictus – – – – – – – 1 1 0.02
Bembidionsp.2 – – – – – 2 – – 2 0.05
Carabushemmeli – – – – – 1 – – 1 0.02
Chlaeniusposticalis – 1 – – – – – – 1 0.02
Curtonotussp.1 – – – – – – – 1 1 0.02
Curtonotussp.2 – 1 – – – – – – 1 0.02
Curtonotussp.3 – – – – – 1 – – 1 0.02
Curtonotussp.4 – 1 – – – – – – 1 0.02
Cymindisdaimio 6 – – – – – – – 6 0.14
Harpalusbungii – 1 – 1 – – – – 2 0.05
Harpaluscoreanus – 2 – – – – – – 2 0.05
Harpaluscrates – 5 – – – – – – 5 0.12
Harpalusquadripunctatus – – 1 – – – 1 4 6 0.14
Harpalustinctulus – – – 2 – – – – 2 0.05
Harpalustschiliensis – 1 – – – – – – 1 0.02
Notiophilussp. – – – 7 5 3 3 – 18 0.43
Pristosiasp.4 – – – – – – – 1 1 0.02
Pterostichuslaticollis – – – – – – – 7 7 0.17
Pterostichusmicrocephalus – – 1 – – – – – 1 0.02
Moisture LOW HIGH
ENTOMOL.FENNICAVol.17 (cid:127) HabitatassociationsofcarabidsinChina 179
Table3.Carabidspeciesrichness,diversityandequitabilityinthestudiedvegetationtypesandduringthetwo
studyyears.
Habitattype Richness(S) Diversity(H') Equitability(J)
1999 2000 1999 2000 1999 2000
Shrubs 14 7 1.90 1.37 0.79 0.74
Coppiceforests 16 14 2.28 2.23 0.82 0.84
Uplandforests 16 13 1.94 1.94 0.71 0.75
Lowlandforests 18 15 1.94 1.96 0.67 0.74
sites, indicating rather similar carabid assem- 3.3.Carabiddistribution
blages.However,theyoftenshowedremarkable
variation in placing over the two-dimensional We regarded a species as being abundant if the
DCAspacewithinthegroupofforestedsitesand catchwas>10individualsinatleastonehabitat
betweenthetwoyears(Fig.1). typeoverthewholetrappingseason(onaverage
Amongthesixmostabundantspecies,C.ma- morethanoneindividualpercollection)andifit
nifestus, C. crasseculptus, P. subovatus and P. comprised>5%ofallthebeetlescaptured(modi-
acutidensshowedsignificantabundancecorrela- fiedfromNiemeläetal.1992).Accordingtothis
tionswitheachother.Morespecifically,P.subo- rule of thumb and based on their occurrence
vatus was positively correlated with C. mani- amongthehabitats,wedividedthe18abundant
festusoverthetwoyearsandandpositivelywith speciesintofourgroups(Table2).
C.crasseculptusin2000.Reflexiphodrusreflexi-
margo was positively correlated with the two
above-listedCarabusspecies,andPristosiasp.1 3.4.Seasonalactivity
was positively correlated with the two Ptero-
stichusspeciesandCarabusmanifestusin2000 The mean catches (n / 100 traps * day) with
(Table4). pooledcarabidsindicatedadistinctiveabundance
peakinAugustinthevitexshrubsites(Fig.2a),
relativelyevenabundancefromMaytoAugustin
Fig.1.DetrendedCorrespondenceAnalysisforthecarabidsamples.Axes1and2respectivelyexplain37.4%
and16.4%ofthetotalvariationin1999and38.1%and8.8%in2000.Shrubs=VS(vitexshrubs)andAS(al-
mondshrubs);Coppice=OC(oakcoppice);Uplandforests=PP(pineplantation),OF(oakforest)andLP(larch
plantation);Lowlandforests=WF(walnutforest)andMF(mixedbroad-leavedforest).
180 Yuetal. (cid:127) ENTOMOL.FENNICAVol.17
Table4.Spearmanrankcorrelationcoefficients(pvalue)fortheabundanceofthesixcarabidspeciesthatmade
up>5%ofthetotalcatchin1999and2000.
Species Year C.crasse- P.subovatus P.acutidens R.reflexi- Pristosiasp.1
culptus margo
C.manifestus 1999 0.731(0.040) 0.711(0.048) 0.611(0.108) 0.611(0.108) 0.214(0.610)
2000 0.868(0.005) 0.898(0.002) 0.659(0.076) 0.802(0.017) 0.756(0.030)
C.crasseculptus 1999 0.461(0.251) 0.657(0.077) 0.639(0.088) –0.024(0.955)
2000 0.800(0.017) 0.630(0.094) 0.861(0.006) 0.691(0.058)
P.subovatus 1999 0.849(0.008) 0.545(0.162) 0.602(0.114)
2000 0.859(0.006) 0.614(0.105) 0.908(0.002)
P.acutidens 1999 0.392(0.337) 0.587(0.126)
2000 0.319(0.441) 0.975(<0.001)
R.refleximargo 1999 0.072(0.866)
2000 0.442(0.273)
thealmondshrubsites(Fig.2b),andaJulypeak 4.Discussion
inalloftheforestsites(Fig.2c–g)exceptinthe
walnutforest,wherethemeancatchesremained 4.1.Speciesabundanceandcomposition
highfromJunetoAugust(Fig.2h).
Atthespecieslevel,Carabusmanifestushad Our results indicate that although carabid spe-
two peaks: in the early season and middle-late cies-level responses to habitat type varied be-
season(Fig.3a),whereasCarabuscrasseculptus, tweenthetwoyears,carabidassemblagesofthe
P. acutidens, P. subovatus and Pristosia sp. 1. five mature forests and the coppice forest were
showed a main peak in the middle season (Fig. moresimilartoeachotherthantothoseinthetwo
3b–e). Finally, Reflexiphodrus refleximargo shrublandsites(Fig.1).Thisresultwasconsistent
showedasimilarabundancepatternasdidC.ma- withthemicroclimateandedaphiccharacteristics
nifestus(Fig.3f). ofeachhabitat.AsindicatedinTable1,thetwo
Fig.2.Seasonalactivity-
densityofcarabidbee-
tlesintheeighthabitat
typesin1999.–a.Vitex
shrubs.–b.Almond
shrubs.–c.Oakcop-
pice.–d.Oakforest.
–e.Pineplantation.
–f.Larchplantation.
–g.Mixedbroad-leaved
forest.–h.Walnutfor-
est.
ENTOMOL.FENNICAVol.17 (cid:127) HabitatassociationsofcarabidsinChina 181
Fig.3.Meancatch
(±S.E.)ofthesixmost
abundantspeciesthatall
comprised>5%oftheto-
talcatchin1999.
shrubland sites were both characterized by dry work (Lindroth 1961–1969, Thiele 1977),
soil,thinleaf-litterlayer,andsparseherbs.Con- carabids were associated with broad habitat
trarytothis,thesixforestedhabitatssharedvery types,suchasforestandhrubhabitats.However,
differentcharacteristics:moistsoil,thickleaflit- most of 18 abundant species were common in
ter,andmoderatetodenseshrublayer.Thus,the severalhabitats,andjustafewwererestrictedto
differences between forests and shrub sites in eitherforestorshrubsites.Onthefinerscale,our
terms of habitat characteristics might determine resultsalsoshowedthatforestsarepatchyenvi-
thedistributionofcarabidsoverthewholestudy ronmentsforcarabids,asthereweresomespecies
area. Our previous research on Carabus assem- restrictedtoonlyoneortwohabitattypesstudied.
blages in thesamestudy areaalso supports this Niemeläetal.(1992)reportedthatcarabidspe-
hypothesis(Yuetal.2002). ciesmayhaveabundancevariationamongdiffer-
Ourresultsalsoindicatedthatthecoppicefor- enthabitattypesandthatmostofthespeciesare
esthadhigherdiversityandequitabilitythanhad habitat specialists in this sense; they also sug-
mature forests and shrub sites, and that the for- gestedthatsoilmoisturecanbeanimportantde-
ested sites with closed canopy supported more terminant of carabid habitat associations. How-
carabidspeciesandindividualsthandidthetwo ever, wefoundarathersimilarhabitatselection
shrubsitesthatwererelativelyopen.Alsothisre- among carabid species, as most of the carabids
sultcanbeexplainedbytheedaphiccharacteris- appearedtobeforestgeneralists.Thisdifference
tics of different habitats. Except for the aspects maybeattributedtodifferencesinedaphicchar-
similar to mature forests, the oak coppice stand acteristicsamonghabitats(Niemeläetal.1992).
hadlowercanopycoverandhigherstructuralhet- Thereareonlyafewdetailedreportsoncarabid
erogeneitythandidthematureforests;thehetero- fauna among different habitats of North China;
geneitymightcontributetothehigherdiversityof therefore,wecouldnotdirectlycomparethede-
carabidassemblagethere.Jenningsetal.(1986) tails of carabid distributions. Thus, further re-
alsosuggestedthatcarabidscanbecommonlyas- searchoncarabidfaunainthisrespectareclearly
sociatedwithclosedthanopenhabitats. needed.
4.2.Carabiddistributionandspatialscale 4.3.Temporaldistribution
Carabids responded to habitat structural differ- In extra-tropical regions, the factors regulating
ences in varying ways depending on the spatial the life cycles of carabid species may include
scale in question. As was expected fromearlier temperatureandphotoperiodicism(Thiele1977,
182 Yuetal. (cid:127) ENTOMOL.FENNICAVol.17
alsothankDr.Ke-PingMa,Xian-MingGao,WeiWang,
Qing-KangLiandMs.Wen-JieYan(InstituteofBotany),
ourcolleaguesinResearchStationofInstituteofZoology,
forfieldandtechnicalassistance,andprovidingmeteoro-
logical data. We also acknowledge Dr. Scott L. Bearer
(CenterforSystemsIntegrationandSustainability,Michi-
gan State University) for linguistic comments on the
manuscript,Dr.JariNiemeläandDr.TeijiSotaforkindly
sendingreprints.Thisstudywassupportedpartlybythe
StateKeyBasicResearchandDevelopmentPlanfromthe
Chinese Ministry of Science and Technology
(G2000046801),theNationalNaturalScienceFoundation
Fig.4.Monthlymeantemperature(ºC)andrelative
of China (30570215), a grant for the senior author (no.
humidity(%)inthestudiedregionin1999.Meteoro- D/3684-1)oftheInternationalFoundationforSciences(I.
logicaldatawerecollectedfromBeijingForestryEco- F.S.,Stockholm,Sweden)andtheNationalScienceFund
systemResearchStation(40º00’N,115º26’E,1,200 for Fostering Talents in Basic Research (NSFC-
ma.s.l.).Columns=temperature,curve=relativehu- J0030092).
midity.
Sota 1985, 1994, Lövei & Sunderand 1996), as References
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