Table Of ContentAMETHODTODEVELOPAN 'INDICATOR VALUE' SYSTEM FOR SPIDERS
USINGCANONICAL CORRESPONDENCE ANALYSIS (CCA)
RALPHPLATEN
Platen, R. 1993 11 1i: A method to develop an 'indicator value' system for spiders using
Canonical Correspondence Analysis (CCA) Memoirs ofthe Queensland Museum 33(2):
621-627 Brisbane, ISSN0079-8835.
MultivariateCanonicalCorrespondenceAnalysis(CCA)isusedtodenvc'indicatorvalues'
forspiders,similartothoseusedforplants.Thedatasetconsistsofactivityabundancevalues
ofspiders, sampledbypitfall trappinginvarioushabitattypes(mires,woods,drygrassland)
indie Berlinarea. Lightexposure, soil moistureandtemperature werealso measuredat the
sites. Speciesscores are plottedas a function ofenvironmental factors withinanordination
diagram. Themethodusedtodetermine the indicatorvalue fromthis ordinationdiagram is
presented- The system ofindicatorvalues isregardedasasuitable method toevaluate sites
andareaseasily.Advantagesand limitationsarediscussed.
Mil Hilfc der multivariaten statisiischen Methodc Kanonische Korrespondcnz Analyse
(CCA) wcrden Zcigcrwerte fur Webspinnen, ahnlich denen fur Pflanzen ermiitell. Die
Entw'icklungdiesesZeigcrwertsysternsunddessenAnvvenduiigwirdirnPrinzipt»cschrieben.
Doi verwendete Daicnsatz besteht aus Aktivitatsabundanzwcrtcn von Spinncn, die mil
Bodcnfallen in unterschiedlichen Biotoptypen (Mooren. Waldern und Trockenrasen) im
QcblelvonBerlingelangenwurden.DieandenStandoncngcmcsscncnabioiischcnFaklorcn
Licht, Tcmperatur und Bodenfeuchte werden mil in die CCA einbezogen. An Hand von
,«iclcn wird der Weg erlaulert, Zeigerwene aus Ordinationsdiagrammen iu ermhieln.
Mit Hille ciniger Arten werden Anwendungsbereich und Beschrankungcn des Zeiger-
wcrtsyslems aufgezeigt und diskutiert. Das Zeigerwertsystem wird als eine brauchbare
Methodc betraehtet, urn Standorte und Untersuchungsgebiete relativ leicht mit Hilfe der
Spinncnzubewcrtcn. [jAraneae, indicatorvalue, multivariateanalysis.
RalphPlaten,InstitutfiirBodenzoologieundOkologie,FreieUniversitatBerlin, Tietzenweg
85-87, W-lOQOBerlin-45, Germany; 12January 1993
Ecosystems change under anthropogenic in- been developed, e.g. lor soil organisms by
flucnecsfasterthan theirstructuresandfunctions Wodarz et a!. (1992), for epigaeic predatory
..in be analysed. It is therefore difficult to make arthropods (spiders and ground beetles) by
well-founded comments about their ability to Haenggi (1987) and Platen (1989. 1992), for
withstandexternalpressure,oraboutpossibilities spiders by Martin (1991) and for ground beetles
for theirprotection orrenaturalisation. Thecom- by Mossakowski and Paje (1985). Some ofthese
plex ecological questions this deficit poses will evaluation systems describe the ecological be
require field work involving as many environ- haviourofthe species in the field very preo
mental factors and groups of organisms as pos- (Martin, 1991), or allow a differentiated evalua-
sible. tionofsitseorareas ofstudv (Mossakowski and
Paje, 1985; Haenggi, l987;*P!aten, 1989). Some
GOALS evaluation systems, however, have the disad-
vantage that lengthy calculations are necessary
A firststep is the description oftheecological foTsynopticevaluationfordifferentsites or a
behaviourofspecies inthefield. A furtherstepis (Wodarz et al.. 199*; Haenggi, 1987). In other
to derive evaluations for the sues, biotopes or cases parameters are used in the caleiil.itu.os
areas of study from the ecological behaviour of whicharc noistabictoriimeand/orlocality. such
the species. For example when establishing as a Iow locaI abundance of a species, or the
whetheranareashouldbeprotected, torpurposes numbers of individuals of a species caught in a
w1.h1 epnlansntiundgyianngdtbhieotcohpaen-gmeasnaagtetmheentsiatsesweulnldears yt.coanrs(yMsotsesmaskmoewnstkiioanneddPcaajne,al1s9o85o)n.lyThbee eapvpallau:a.-!
anthropogenic influence, an efficient evaluation ,0Cal1Y whcre* as a result °fintensivefieldwork,
sysiem wVich beyond that is easy to handle will theecological behaviourofspecie* atoogahiotie
l.'c necessary. gradients isknown.
A numberofevaluation systems have recently A much simplermethod would be theapplica-
1 1
622 MEMOIRS OFTHEQUEENSLANDMUSEUM
liOfl of an indicatorvalue system similar to thai RESULTS
forplantsofEllenbergetal. 991).Itwouldthen
no longer be necessary to redetermine and re- OrdinationDiagrams
evaluatetheecologicalbehaviourofaspecies for The CCA results with abiotic factors light ex-
eachlocalinvestigation, sincethiswouldalready
be contained in the key values Nor would the posure and temperature, ns wrll n£ Soil water
evaluation involvecomplicated calculations. content, are shown graphically (Figs 1C.C2)A. The
My aim has been to develop just audi an in- ahonrdiztohnetvaelrtaixciaslctoortrheesspoencdosndtoCtCheAfiarxsits.TheaIxiIsI
dicatorvalue system forspiders species of spider are represented by an *X\
togcthcrwith an abbreviation ofthe name as Far
MATERIAL AND METHODS as possible. Using CANOPLOT itwas alsopos-
sible In determine the coordinates and the name
ofa species which could not be presented unam-
Data biguously in the diagram.
The data consisted ofthe activity abundances Initially the axes of the site factors light ex-
of spider species. These were determined using posure and soil water content are extended
ground traps in the Berlin area for open and beyond the origin (Fig. I). The factor along the
woodedsites in moors, in various types offorest t'heenvairrroonwm.enTthaeloarxiegsi'nimncarrekassetshienmtheeadnirveacltuieonfoofr
andforheathlandandsemi-dryanddrymeadows.
The investigation period was a full yearin each the entire data set Species whose position lies
case. Activity abundance is defined by between thearrowheadofan environmentaI aX1
and the origin have a larger weighted mean.
Hrweehpiyrcdehseehmnaatsnedbne(bey1n95ttr3hee)sapdsaisatsmheeedtneaurbmoobrfedretrholefinipeintd(fiawvlhildiutcarhlaspi.)s Wthheeproesitthieonoroifgitnheisspbeectiewse,enitsthweeiagrhrtoewdhemaedananidv
smallerthan the overall mean. Forinterpretation
withinacertainperiodoftime.Parallel tothetrap
catches the following abiotic factors were also a perpendicular is projected for each species in
turn onto the environmental axis according to
measured:
their sequence (cf. Jongman ei at, 1987). X\s-
The soil water content (measured as the per- ticus nitwit and Thanatm arenarius occupy the
centage by volume of water in the upper soil brightest sites, and Pardasa agrestis and
dlaeysecrr)i,betdhebyligFhntenedxpo(1s9u6r1e),usainndgtthheeemffeetchtiovde BSpaercyipehsyamtaexptrrareemnesseotfhtehweaarxmeessrtepsirteessen[tRgl,imiItsi
temperature after Pallmannn et td (1940). The ofthearea foratwo-factorsystem,and thusform
sites aredescribed in detail in Platen * 19S9». the start and end points of the indicator value
scale. The distance between these points is
measured and divided into five equal parts. The
General
species are then noted for each class with the
The activity abundance of the spider species appropriateindicatorvalue.
and the measurements ofthe abiotic factors are The determination ofindicatorvaluesforthree
analysed using Canonical Correspondence
factors requiresat leasttwo ordination diagrams.
pAnraolgyrsaims C(CACNAO;CJoOngVmerasnieotnat3...1019(SB7r)a,auks,in1g98t8h.e Initiallyanindicatorvalueisassignedforallthree
individual factors, then for all combinations of
1991). The results ofthis analysisCaArNc dOiDspRlaAyeWd twofactors.Theresultalwaysremainedthe same.
as ordination diagrams using For the representation Of all three factors in an
(Smilauer, 990
1 ordination diagram the class of a species cmi.
Before running CCA the spider data had been however, change, in some cases considerably,
maskedaccordingtodominanceinaformalway: since the relative spatial distances ofthe species
species whichdid nothave anactivityabundance in Ihrce-factor constellations cannot be repre-
ofatleast ]% a:asitewereremovedfromthedata sented in a two-dimensional coordinate system
set. This meant that of the original 281 S| wiihoui distortion. For the determination ol in-
species only 111 remained for the further dicator values from the data of individual en-
anaFluyrstihs.ermore a transformation of the WW data avniroopntmienmtuaml.variables the solution (Figs 1 . 2 ) is
was carried out. Instead ofthe abundance values Moisture is strongly negative correlated with
theirsquare roots wereused. the 1st CCA-Axis (Inter-setcorrelation: -0.919)
INDICATOR VALUESFROMCANONICALCORRESPONDENCE ANALYSIS 623
Temperature
1st
CCA-Axis
V
FIG. 1. CCA ordination diagram wilh 1 1 1 spider species represented by an and environmental variables
moistureandtemperaturerepresented byarrows. The partofthearrowstoderive indicatorvaluesaredevided
inlo fiveparts("M1-M5 andTI-T5 respectively). Forfurtherexplanation sec text.
(Fig. I) which means that the horizontal species cannot claim to be comprehensive or generally
distribution isbestexplainedby lightand less by valid. Some examples will show the similarities
temperature (Inter-set correlation with 1st CCA- and differences between the indicatorvalues and
Axis: 0.26, with 2nd CCA-Axis: 0.538). Hence, other methods of determining ecological be-
the vertical species distribution is bestexplained haviour.
along the 2nd CCA-Axis. The distribution of Xysticus ninnii is centered
The Inter-set correlation between light and 1st exclusively on dry meadows. Fi, L5» and T4
CCA-Axis is0.955 which meansthatthedata set reflect this ecological behaviourwell.
again is best explained by this abiotic factor. Diplocephaluspermixius: Occurring mostly in
Temperature is strongly correlated with 2nd wei alder forest-habitats characterised by high
CCA-Axis (Inter-set correlation with 1st CCA- soil water content, low light exposure and low
Axis: 0.0367, with 2nd CCA-Axis; 0.6177). temperatures. This is expressed with adequate
precision by the indicator values F5, LI and T2.
Indicator Values(Table I) Diplocephaluspicinus: F2, LI and Tl charac-
The last two columns contain details of the terise its habitat preferences, namely shadowy
ecological type and habitat type in which the sites with low pH in dry mixed forests.
species predominantly occurs in the Berlin area Pardosa agrestis: In this case the indicator
(after Platen et fli, 1991). The data are intended value does not reflect the ecological behaviour,
only to demonstrate the principleofthis method. as a result ofthe inadequacy ofthe data set. The
Inviewofthelimiteddata setthe indicatorvalues species occurs mostly on arable farmland and
624 MEMOIRSOFTHEQUEENSLAND MUSEUM
2nd
CCA-Axis
Temperature
Xy« -.inn
Light
IstCCA-
Axis
V
FIG. 2.CCAordinationdiagramwith 1J1 spiderspeciesrepresented byan andenvironmentalvariableslight
and temperature representedby arrows. The partofthe arrows to derive indicatorvalues arc divided into Five
parts (L1-L5 andTUTS respectively). Forfurtherexplanation seetext.
otheropen, ratherdry habitat types (Platenetal., evaluation systems (Mossakowski and Paje,
1991).WhileT5describes thebehaviourin terms 1985;Haenggi, 1987; Platen, 1989).
of temperature, the indicated soil water content Some reservations concerning the applications
(F5) is too high and the light exposure too low ofindicatorvaluesarenecessary. Asalreadyem-
(LI). This is because in the data set it only oc- phasised by Ellenberg et al (1991), indicator
curredononesite(aformermoorwhichwas still values describe the ecological behaviour of
wet in comparison to the sites on mineral soils), species in a multiple system ofbiotic and abiotic
so that as an outsider it had an extreme position factors,fromwhichthosechosen in thispaperare
in the ordination diagram. The data set did not regarded as thekeyabioticfactorsforthespiders.
include arable farmland sites. Theydo not, however, describe their physiologi-
cal optima.
Indicator values for animals underlie more
Comments
restrictions as those for plants because animals
This method is a relatively easy one to deter- are mobile and often change their habitat for
mine indicator values for spiders and other soil overwintering(Schaefer, 1976). Therefore in the
arthropods (cf. Platen, 1992). strict sense they are valid only for Adults which
Indicator values have the advantage that with does not change theirhabitat within theirperiod
only a few values the ecological behaviour of a of maturity. As juvenile animals were not in-
species can be characterised. Mean indicator cluded in the data set this holds true for this
valuescanbecalculated,andintheabsenceofthe investigation.
measurementsorvegetationsurveystheyallowa Indicator values should never be used in the
rough quantitative assessment ofenvironmental same way as measurements. They are ordinal,
variables for a investigation site, habitat type or and are not suited foruse in statistical (including
an area of study. The use of indicator values multivariate) methods requiring higher scale
obviatestheneedforcomplicatedcalculationsfor data.
the evaluation of sites, as required by some The indicator values determined above arc al-
INE>ICATOR VALUESFROMCANONICALCORRESPONDENCEANALYSIS 625
ways valid only for thedata sel used. Since they program forcanonical community ordination b)
were limited to only a few lypes ofbiotope the [partial |dcucndcdj [canonical correspondence
|
resultsabovecanonlyberegardedasbeingafirst analysis, pnncip.il components analysis and
approlxyimdaetpieonnd.enTtheonretsuhletstyopfetahnedannaulmysbiesraoref rreepdourntdLanWcyA aSnKa-f0y2s.lgGr(oVcerpsiLoannd2b.o1)u1w.wtTesckhunnidcsa,l
Wageningcn
habitats types and of the frequency with which 1990, Update notes: CANOCO Version 3.10.
various species occur there. The combination nf Manuscript. Agricultural Mathematics Croup,
a wet. light site (F5, L5) is not represented by an Wageningcn.
indicatorvalue,although it is relevantfora num- hLLhNBERG.H-.WEBER. RE.. DULL,R.,WIRTH,
ber of species (Drepanotylus uncatus* AnttsUa V., WERNER. W. & PALESSEN, D.
clegans) (Table 1). However,sincealmost2/3 of Zcigerweru wan PI'lan/cn inMiUcleutopa.Scrii*a
the 30 sites investigated were we*, and most GeoboUmcitXVIFl: 1-248.
speciesoccurred with almostthe samefrequency prifnd,D.T.C 1961, Asimple method oi measuri
inwethabitats,thesespeciesgroupedelo.se Uithe integrated light values in the held. Ecology 12
origin Species which occur frequently, but only 577-580.
at one or two dry sites with very high Hgbl HANGGI, A. !987. Die Spinnenfauna d*r
posure are far ftoin ihc origin, so thai there is a FcuclllgehietC lies Grolkn Mooscs, Kt. Bern. 2
higher differentiation ofthe axis over the bright Die Bcurteilung des Nalurschut/.wcrtes natur
nahcr Standoric anhand der Spinnenl'ann.v Mii-
range.
Agenerally validindi'catQi value s> stem would tBceirlounNg.cFn d4e4:r N15a7u-i1r8f5o.rsehcnden Gcscllschaft in
need toanalyseall known spider specks ofGer- HEYDEMANN. B. 1953. Agrarokologischc
many or Central Europe for all existing types of Frobleroatik, dnrgetan an Untcrsuchungcn tlbcr
biotope{abioticfactorcombinations)inone data die Tierweh der Bodenoberfluche der Kultnr-
set.fromwhichtheindicatorvaluescouldthe felder. (Thesis Kiel). 4^ Pp
derived. The scale could then be expanded, or JONGMAN, R.H.G., BRAAK, C.J.F TER & TONCr
otherenvironmental factors, such as the biotope EREN.O.RRfcds,), 1987 'DaMmilysiMncom-
structure,couldbeincluded.Aproblemwouldbe munity and landscape ecology.' (Pudoc Books:
the large numberofmeasurements required Wageningcn).
A furtherproblemisthattheCCAonlydepicts MARTIN, D. 1991. Zur Autokologie der Spinn. r
speciescorrectlyintheordinationdigram jj (atAaru3scshtnaitdtau:ngAruanndeaPera)f1crCcbnazrvaekrthearlitsetnikepdiegraHiasbcihtcr
have an unimodal response curve alonga factor
gradient (Jongman et ai,t 1987), By plotting the MOSSSApiKnOneWnSarKtIen,.ADr.ac&hnPoAloJgEi.aFM.it1t9e8i5l.unEgienn 1B;e5w-e2i6-.
frequenciesof species at all habitat types sorted tungsverfahren von Raumeinhctlcn an Hand der
accordingtothe lc.els ofa factor, ii is possible10 Carabidenbestande.Verhandluneen,Gesellschaft
determine bi-modul, multi-modal or continuous Okologic in Bremen 1983 13: 747-750.
responses. The coordinates of all (he habitats PALLMANN, H., EICHENBERGER, E. & HASEL-
where theSpeciesoccurswithhighfrequencycan ER. A. 1940. EincMetnode derTempcraturmev-
be entered in the ordination diagram, making it sung bei okologischen oder bodenkundlichcn
possible to recognise a corresponding range of Untersuchungen. Berichte. Schweizerischc
occurrence on the environmental axis. For this BotanischcGcscllschaft50: 337-362.
species, asis thecase withsome plantspecies, an PLATEN, R. 1989. Struktur der Spinnen- und
indifferent response to this factorfor the species LaufkaTcrfannn (Araeh.; Araneida, Col..
or to give a range of the indicator value (cf. Carabidae) amhropoden bccmtlutitcr Moorstan-
Ellcnbergefa/.. 199])may be possible. dorle in Berlin (West); laxonomischc, raumtichc
und zcillicbc Aspckte. (Thesis. Techmsciie
ACKNOWLEDGEMENTS Univcrsitat Berlin)470pp.
PLATEN, R MOR1TZ, M. & BROEN, B.v 1991
lf
I.isle der Webspinnen- und WcberkncchiarU-n
toFtohraindkeaFsl,ocrriiatnicBiesmmmaenrdldeiisnc-uLswsixo,nN1uwroeumlbderligk,e (RAaruamchc*; Arounneidd.i,ihOipcilioAnuisdaw)crdtesunBgerlifnieirr
Prof. Lasco Mucins, Vienna, and Prof. Gcrd Ktturech«lzzwecke(RcrteLi8ie) Pp 243-275 Eh
Weigrnann, Berlin A.Auliagen,R.Platoand H.Sukopp(eds). 'Rote
Listen ||er gefahrdeten Pfian/en und Tic
BcHin\(LiindsLlutisentvv.; Umwcltforsch. S6).
1TFRATURECTTED
I PLATEN, R. !992. Die Lutwieklung eines Zeigw-
wcrtsystcms fur Lnufkiifer (Col.; Carabidae) ml
BRAAK, C.J.F. TGR 1988. CANOCO -a FORTRAN Hilfccmer"CanonicalCorrespondence:Analysts"
626 MEMOIRS OFTHEQUEENSLANDMUSEUM
(CCA). Verhandlungen der Gesellschaft fiir SMILAUER, P. 1990. CANODRAW. A companion
Okologie21: 321-326. program to CANOCO for publication-quality
SCHAEFER, M. 1976. Experimentelle Untersuchun- graphicaloutput. (Ithaca).
genzumJahreszyklusundzurUberwinterungvon WODAR2,E., AESCHT,E. &FOISSNER, W. 1992.
Spinnen (Araneida). Zoologische Jahrbiicher, Der Bodenbiologische Index (BI) - ein quantita-
Systematik(Okologie),GeographicundBiologie tives MaB fiirdie Bqdenqualitat. Verhandlungen
103: 127-289. derGesellschaftfiirOkologie21: (inpress).
TABLE 1. Indicatorvalues forsoil watercontent(F), light(L), temperature (T) of 111 common spiderspecies
ofvarying types of habitat. ET=Ecological type: h = hygrophilic, (h) = weakly hygrophilic, x=xerophilic,
(x)=weakly xerophilic, eu= euryoecious open-space dwellers, w=forest type, (w) = also in open spaces,
hw=sparse forest species, (h)w=inhabits mesophilic deciduous forests, (x)w=inhabits forest on acid soil,
h(w)=dependingontypeofpreferredhabitat: inhabitsunwoodedwethabitatsorsparseforest. -=nopreferred
habitats. Family (C): Ag, Agelenidae; Dy, Dysderidae;Gn,Gnaphosidae: Ha,Hahniidae;Li,Linyphiidae; Lc,
Liocranidae; Ly, Lycosidae; Ph, Philodromidae; Pi, Pisauridae; Sa, Salticidae; Te, Tetragnathidae; Tr,
Theridiidae;Tm,Thomisidae;Zo,Zoridae.
SPECIES F L T ET \c SPECIES F L T ET C
| | | |
WetForests W.cucullata(C.L.Koch) 2 2 (x)w Li
PachygnathalistenSundevatl 5 3 hw Te W.dysderoides(Wider) 4 3 (x)w Li
Bathyphantesapproximates(O.P.C.) 4 3 h(w) Li IV.monoceros(Wider) 2 2 2 (x)w Li
B.nigrinus(Westring) 5 3 hw Li Euryopisflavomaculata(C.L.Koch) 4 4 (x)(w) Tr
Diplocephaluspermixtus(O.P.C.) 5 2 h(w) Li TrochosaterricolaThorell 3 2 2 (x)(w) Ly
Diptostylaconcoior(Wider) 4 .'; (h)(w) Li Xerolycosanemoraiis(Westring) 2 2 (x)(w) Ly
Gonatiumrubellum(Blackwall) 3 2 hw Li Cicurinacicur(Fabricius) 4 3 (x)(w) Ag
Porrhommapygmaeum(Blackwall) 4 3 h(w) Li Agroecabrunnea(Blackwall) 4 3 (w) Lc
Walckenaeriaatrotibialis(O.P.C.) 4 2 (w) Li Haplodrassussoerenseni(Strand) 1 1 (x)w Gn
Piratahygrophilus(Thorell) 5 3 h(w) Ly Zelotessubterraneus(C.L.Koch) 3 3 (x)(w) Gn
Deciduousforests Ozyptilapraticola(C.L.Koch) 2 2 (x-)w Tm
Centromerussylvaticus(Blackwall) 4 2 (h)w Li Watersidesites
Ceratinellabrevis(Wider) 3 2 (h)w Li Gnathonariumdentation(Wider* 15 j i 14 |h |Li
Diplocephaluslatifrons(O.P.C.) 3 2 (h)w Li Moors
Gongylidiumrufipes(Sundevall) 3 2 (h)(w) Li Agynetacauta(O.P.C.) 5 3 h(w) Li
Lepthyphantespallidus(O.P.C.) 2 2 (h)(w) Li DiplocephalusdentatusTullgren 5 3 h(w) Li
L.tenebricola(Wider) 4 2 (h)w Li Drepanotylusuncatus(O.P.C.) 5 3 h Li
/...zimmermanniBertkau 2 1 (h)w Li Erigonellaignobilis(O.P.C.) 5 3 h Li
Micronetaviaria(Blackwall) 3 2 (h)w Li LepthyphantesmengeiKulczynski 5 3 h(w) Li
Nerieneclathrata(Sundevall) 4 3 (h)w Li Lophommapunctatum(Blackwall) 5 3 h Li
Pardosalugubris(Walckenaer) 4 3 (h)w Ly Notioscopussarcinatus(O.P.C.) 5 3 h Li
Drymixedforests Oedothoraxgibbosus(Blackwall) 5 4 h Li
Harpactearubicunda(C.L.Koch) 1 1 (X)W Dy Silometopuselegans(O.P.C.) 5 5 h Li
Abacoproecessaftuum(L.Koch) 2 1 (x)w Li Tallusiaexperta(O.P.C.) _> 4 (h) Li
Centromeritaconcinna(Thorell) 2 2 (x)(w) Li WalckenaeriaalticepsBlackwall 5 4 h(w) Li
Centromeruspabuiator(O.P.C.) 3 2 (x)(w) Li W.kochi(O.PC) 5 4 h Li
Diplocephaluspicinus(Blackwall) 2 1 (x)w Li W.nudipalpis(Westring) 5 4 h Li
Gonatiumrubens(Blackwall) 2 1 (x)w Li W.vigilax(Blackwall) 5 3 h Li
L(eWpetshtyrpihnagntesangulipalpis 2 1 (x)w Li Arctosaleopardus(Sundevall) 5 3 h Ly
Hygrolycosarubrofasciata(Ohlert) 5 3 h Ly
LepthyphantesJlavipes(Blackwall) 2 1 (x)w Li Pardosapullata(Clcrck) 5 3 h Ly
Macrargusrtifus(Wider) 4 2 (x)w Li !' latitans(Blackwall) 5 3 h Ly
Minyrioluspusillus(Wider) 2 1 (x)w Li P.piraticus(Clerck) 5 3 h Ly
PanamomopsmengeiSimon 1 1 (x)w Li P.piscatorius(Clerck) 5 3 h Ly
Tapinocybainsecta(L.Koch) 2 2 (x)w Li
P.tenuitarsisSimon 5 4 h Ly
WalckenaeriaacuminataBlackwall 4 3 (x)w Li
Trochosaspinipalpis(F.O.P.C) 5 3 h(w) L>
INDICATORVALUES FROMCANONICALCORRESPONDENCEANALYSIS 627
TABLE 1. continued
SPECIES F L T ET C SPECIES F L T ET C
Dolomedesfimbriatus(Clerck) 5 I 3 h Pi Tegenariaagrestis(Walckenaer) 4 1 4 (x) Ag
Antisteaelegans(Blackwall) 5 1 4 h Ha Heathland
Reeds Triccalutetiana(Simon) 1 3 2 (x) Ly
Baryphymapratense(Blackwall) |5 |l J5 [h JLi Zeloteslatreillei(Simon) 2 1 1 (x) Gn
WetMeadows Drygrassland
Allomengeascopigera(Grube) 3 1 2 h Li PachygnathadegeeriSundevall 2 2 3 eu Te
A. vidua(L.Koch) 4 1 2 h Li Meionetabeata(O.P.C.) 1 4 3 X Li
Ceratinellabrevipes(Westring) 4 ! 3 h Li Trichopternacito(O.P.C.) 1 5 3 X Li
Erigonellahiemalis(Blackwall) 4 1 3 (h)(w) Li Troxochrusscabriculus(Westring) 2 1 1 X Li
Oedotboraxfuscus(Blackwall) 4 1 2 eu Li Typhochrestusdigitatus(O.P.C.) 1 5 3 X Li
0.retusus(Westring) 5 1 4 eu Li Steatodaphalerata(Panzer) 1 5 3 X Tr
Pelecopsismengei(Simon? 3 1 1 h Li Alopecosacuneata(Clerck) 2 3 3 X Ly
Pardosapalustris(Linn6) 1 4 3 eu Ly A.pulverulenta(Clerck) 4 1 3 eu Ly
P.prativaga(L.Koch) 4 1 3 eu Ly Xerolycosaminiata(C.L.Koch) 1 4 4 X Ly
Cough-grasssites Hahnianava(Blackwail) 1 4 3 X Ha
Centromeritabicolor(Blackwall) 2 |3 |3 |(x)(w) Li Agroecaproximo(O.P.C.) 4 1 3 (x) Lc
Ruderalsites Zeloteselectus(C.L.Koch) 1 5 3 X Gn
Baihyphantesparvulus(Westring) 5 I 3 eu Li Zlongipes(L.Koch) 2 4 5 X Gn
Pocadicnemispumila(Blackwall) 4 1 3 eu Li ThanatusarenariusL.Koch 1 5 4 X Ph
Stemonyphanteslineatus(Linn6) 4 2 3 (x) Li XysticuskochiThorell 1 4 3 X Tm
Trochosaruricola(DcGcer) 3 2 3 eu Ly X.ninniiThorell 1 5 4 X Tm
Arablefields Aelurillusv-insignitus(Clerck) 1 5 4 X Sa
Bathypbantesgracilis(Blackwall) 4 1 3 eu Li Phlegrafasciata(Hahn) 1 3 2 X Sa
Erigoneatra(Blackwall) 3 2 3 eu Li Noobvioushabitatpreferences
E.dentipalpis(Wider) 4 2 3 eu Li Cnepbalocotesobscurus(Blackwall) 3 1 3 eu Li
Pardosaagrestis(Westring) 5 1 5 (x) Ly Zoraspinimana(Sundevall) 4 1 3 eu Zo
