Habitat affinities of 228 boreal Finnish spiders: a literature review

2004·0.27 MB·

by  Katja Matveinen-Huju

©EntomologicaFennica.17September2004
Habitat affinities of 228 boreal Finnish spiders:
a literature review
KatjaMatveinen-Huju
Matveinen-Huju,K.2004:Habitataffinitiesof228borealFinnishspiders:aliter-
aturereview.—Entomol.Fennica15:149–192.
Informationaboutthehabitataffinitiesofspidersisnecessaryinecologicalstud-
ies.Ifocusedontwofactorsinhabitatsofspiders:lightintensityandmoisture.
First,thehabitatswereclassifiedaccordingtothesefactors.Canopycoverand
treespeciescompositionwereusedtoindicatetheamountofsunlightreaching
theground.Habitatmoisturewasdeterminedindirectlyfromthevegetation.Sec-
ond,thespiderspecieswereclassifiedaccordingtowhatkindofahabitatthey
werefoundintheliterature.ThispaperupdatesandcontinuesHuhta’s(1971)
classification, including 228 boreal spider species, based on Finnish literature
fromborealregion.
K. Matveinen-Huju, Department of Biological and Environmental Sciences,
P.O.Box65,FI-00014UniversityofHelsinki,Finland;E-mail:katja.matveinen
@helsinki.fi
Received8April2003,accepted21January2004
1.Introduction beapplicabletoFinland(Huhta1971,Lehtinenet
al.1979).Theupdatedclassificationnowcovers
In ecological studies, it is useful to know the 228 of the ca. 620 species found in Finland
autecology of the species studied (Shrader- (Marusik&Koponen2000).Thespecieschosen
Frechette&McCoy1993).Thisinformationcan for this literature review are such that can be
beusedasabasisforhypothesistesting.Classifi- caught with pitfall traps. The species list is not
cationofthespeciesaccordingtotheirhabitataf- comprehensive, but contains species that I have
finities is common for arthropods (e.g. Huhta collected in my ecological studies (K. Matvei-
1971, Niemelä & Halme 1992, Koivula 2001). nen-Huju,unpublisheddata).
However, often classifications are quite rough The nomenclature follows Platnick (2003).
(forestspecies,openhabitatspecies,andgeneral- Other names used in the publications reviewed
ists), and e.g. moisture is usually not taken into are given in parentheses after each species; au-
account.Huhta(1971)publishedahabitat-affin- thorsfortheseothernamesaregivenonlywhen
ity classificationfor198spiders.Here,Iupdate theydifferfromtheauthorsofthenamesusedin
and continue this classification, emphasizing Platnick(2003).
Finnishliteraturewheneveritisavailableforthe
focalspecies.Speciesmayoccurindifferenthab-
itatsindifferentpartsoftheirrangebecausesuit- 2.Habitatclassification
able microclimatic conditions may vary among
types of habitat (e.g. Huhta 1971), and conse- Theoccurrenceofmostspidersisstrictlylimited
quentlytheCentralEuropeanliteraturemaynot byphysicalconditions,suchastemperature,hu-
150 Matveinen-Huju (cid:127) ENTOMOL.FENNICAVol.15
midity,wind,andlightintensity,andbybiologi- C=Shadyhabitats(spruceforests,spruce
calfactors,suchasvegetationtype,foodsupply, mires)
competitionandpredation(Foelix1982).Icon-
centratedonlightintensityandmoisture,because Moisture
Iassumedthesefactorstobeamongthemostim- 1= Dryhabitats(e.g.clear-cuts,drymeadows,
portantabioticfactorscharacterizingthehabitats forestswithagroundlayeroflichens)
ofspiders. 2= Medium-moisthabitats(e.g.meadowsof
mediumhumidity,forestswithaground
layerofforestmosses,likeHylocomium
2.1.Classificationofhabitats splendens,andgroves)
3= Moisthabitats(e.g.bogs,mires,moist
Light intensity varies with the amount of radia- meadows,forestswithagroundlayerof
tionreachingthegroundlayer.Theamountofra- Sphagnummosses)
diationreachingthegrounddependsontheden-
sity of canopy cover, and tree species composi-
tion (Geiger 1965). In Norway spruce (Picea 2.2.Definitionofaspecies’habitataffinity
abies) stands, 4%–40% of the radiation reaches
the ground, whereas in birch (Betula spp.) or Aspecies was classified according to its occur-
Scotspine(Pinussylvestris)standstheamountis rence in different habitats, using above levels
higher: for birch 20%–30% and for pine 22%– fromAtoCand1to3.Thespeciesindifferentto-
40%.Fordeciduoustrees,theamountofradiation wardslightintensitywerenotedasABC,andthe
reachingthegroundvarieswiththeseason:inthe speciesindifferenttowardsmoistureas123.Note
summer deciduous forests may be as dark as thatHuhta(1971)usedDinsteadofABCand4
spruceforests,butwhentherearenoleavesabout insteadof123.
50%oftheradiationreachestheground(Geiger I concentrated on published studies done
1965). Here, I considered deciduous forests as withinthehemiborealandborealforestzonesin
light forests, because the abundance/activity Finland,citedinthetextforeachspecies.Asmy
peaksofmanyspeciesareinspringandautumn focuswasonborealzone,Ididnotconsiderstud-
(Huhta 1971, Itämies & Ruotsalainen 1984, ies done in the northern parts of Finland. One
Niemeläetal.1994). should thus note that a given species might be
It is more difficult to find accurate informa- foundindifferenthabitatsinLapland,compared
tionaboutthemoisturelevelsneartheground.I to the boreal regions of Finland. For example,
usedaruleofthumbbasedonvegetation:forests MarosublestusismoreeurytopicinLaplandthan
with lichens at the bottom layer are dry, those intheborealregion(Saaristo1971).Ialsoused
with forest mosses, such as Hylocomium onefieldguidefromCentralEurope(Heimer&
splendens, are medium-moist, and those with Nentwig1991),becauseitcontainedmostofthe
Sphagnummossesaremoist.Iconsideredclear- reviewedspecies,andaselectionofpublications
cutsasdry,andgrovesasmedium-moisthabitat, fromSweden (Holm1947, Holm& Kronestedt
even though there might be considerable varia- 1970,Kronestedt1990).However,fortheactual
tionwithinthem. classificationsIignoredthesepublicationsifthey
Thehabitatclassesareasfollows.Theclasses werenotconsistentwiththeFinnishstudies.
usedforlightintensityaremostlythesameasin Animalsmoveandcanbeoccasionallyfound
Huhta(1971),butformoisturetheclassbound- inhabitatswheretheydonotliveorbreed.There-
ariesprobablydiffer. fore,itwasdifficulttoknowwhetherthefindings
were from the preferred habitats or represented
Lightintensity occasional ones. Whenever possible (Palmgren
A= Openhabitats(e.g.clear-cuts,meadows, 1964,1977a,Huhta1965,1971),Icalculatedthe
fields,openbogs) percentagesofindividualsfoundineachhabitat
B= Semi-openhabitats(e.g.lightforests,for- type,andthefocalspecieswasconsideredtohave
estedges,pinebogs,bushes) anaffinityforacertainhabitatclassifmorethan
ENTOMOL.FENNICAVol.15 (cid:127) Habitataffinitiesofborealspiders 151
75%oftheindividualswerecaughtthere.More biotopes, suchasshoresandbogs(e.g. Myrica-
weightwas putin thosestudies wherethefocal Molinia bogs) (Palmgren 1972, 1974b). It has
species was caught abundantly. The literature alsobeencaughtinlightforests(e.g.rockypine
usedinthisreviewfor228spiderspeciesispre- forests),roadsidesorclearedforests,andLedum
sented in section 3, and the habitat-affinity bogs(Palmgren1974b,1977a).
classes, based on this literature, are listed in
Appendix1. Araneussturmi(Hahn,1831)(Ateasturmi)
Araneus sturmi lives in trees and bushes (espe-
cially pines) (Palmgren 1964, 1972, 1974b,
3.Descriptionsofthehabitataffinities 1977a,Lehtinenetal.1979,Heimer&Nentwig
of228spiderspecies 1991).Ithasmorerarelybeencapturednearfor-
est ground (e.g. Calluna and Cladonia in light
Fromnowon,ifIreportaspeciesbeingcaught pine forests, Vaccinium myrtillus, VT, MT)
’near the ground’, it means in the ground-layer (Palmgren 1972, 1974b, Biström & Väisänen
and/orintheloweststrataofthevegetation.With 1988), peatlands (e.g. open bogs, Ledum bogs)
’dryish’Imeandryhabitatsandthedriesthabitat (Palmgren 1972, 1977a), meadows (Palmgren
typesinthemedium-moistclass.Theforesttypes 1972, 1977a), and roadsides or cleared forests
andtheirabbreviationsfollowCajander(1949). (Palmgren1977a).
Cercidiaprominens(Westring,1851)
3.1.Araneidae Cercidia prominenslivesin open or semi-open,
dryishhabitatsneartheground(Palmgren1972,
AraneusdiadematusClerck,1757 1974b). It has been found in forests (e.g. rocky
Araneusdiadematuslivesintreesandbushes(es- pineforests,VT,mixedforests;alsoMT,spruce
peciallyinconifers)(Lehtinen&Kleemola1962, mires)(Palmgren1964,1972,1977a,Biström&
Palmgren1972,1974b,1977a,Koponen1996).It Väisänen1988,Heimer&Nentwig1991),bogs
has more rarely been caught near the ground in (Ledum bogs, Myrica-Molinia bogs; with the
forests (e.g. rocky pine forests, VT, MT, hazel higher parts generally dry) (Palmgren 1972,
groves)(Huhta1971,Palmgren1972,1977a,Bi- 1977a), meadows (e.g. dry meadows, near
ström & Väisänen 1988), meadows (Palmgren ditches) (Palmgren 1972, 1977a, Heimer &
1977a), Ledum bogs (Palmgren 1977a), road- Nentwig 1991), small islands (Palmgren 1972),
sidesorclearedforests(Palmgren1977a),andin and roadsides or cleared forests (Palmgren
walls of wooden houses or in firewood piles 1977a).
(Palmgren 1977a). According to Heimer and
Nentwig(1991),thespecieslivesinforestedges Cyclosaconica(Pallas,1772)
andgardens. Cyclosaconicalivesmainlyintrees(Heimer&
Nentwig 1991), predominantly in spruces and
Araneusnordmanni(Thorell,1870) veryrarelyinpinesandjunipers(e.g.MT,OMT,
Araneusnordmannihasbeencapturedmostlyin spruce mires) (Palmgren 1964, 1972, 1974b,
spruce forests and on spruce trees (Palmgren 1977a).Ithasalsobeenencounteredinthelower
1974b, 1977a, Biström & Väisänen 1988, Väi- strataofforests(e.g.moss,Vacciniummyrtillus,
sänen&Biström1990);accordingtoHeimerand Calluna) (Palmgren 1964, 1972, 1974b, 1977a,
Nentwig(1991),itlivesintreecrowns.However, Huhta 1971, Pajunen et al. 1995). Afew speci-
Lehtinenetal.(1979)includedA.nordmanniin menshavebeenfoundinLedumbogs,too(Palm-
mirefauna. gren1977a).
AraneusquadratusClerck,1757 Gibbaranea omoeda (Thorell, 1870) (Araneus
Araneusquadratuslivesinmeadows,especially omoedus)
moistones(Palmgren1972,1974b,1977a,Hei- Gibbaranea omoeda lives predominantly in
mer & Nentwig 1991), and in other moist open spruces(e.g.MT,sprucemires)(Palmgren1972,
152 Matveinen-Huju (cid:127) ENTOMOL.FENNICAVol.15
1974b, 1977a, Lehtinen etal. 1979). Ithasalso ows(e.g.nearditches,shoremeadows;onlyone
beencaughtinotherconiferoustrees(Palmgren individual in dry ones) (Palmgren 1964, 1972,
1977a, Heimer & Nentwig 1991), near the 1977a). However, according to Heimer and
ground in forests (e.g. rocky pine forests, VT, Nentwig(1991),C.lutescensliveswithinshore-
MT, OMT, mixed forests) (Huhta 1971, Palm- linevegetationandpeatlands.
gren1977a,Palmgren&Biström1979,Biström
&Väisänen1988,Pajunenetal.1995).Oneindi- ClubionaneglectaO.P-Cambridge,1862
vidual has been reported even in a Ledum bog Clubiona neglecta has been caught mostly in
(Palmgren1977a). meadows, mainly dry ones (Palmgren 1972,
1977a), and meadow-like semi-open vegetation
Hypsosingasanguinea(C.L.Koch,1844)(Singa onsand(Palmgren1972).Ithasalsobeenfound
sanguinea) in edges of brooks in spruce or mixed forests
Hypsosingasanguineahasbeencaughtinforests; (Palmgren1977a),andindeciduousbushesand
mainly light ones (e.g. CT, rocky pine forests, herbs(Palmgren1943).AccordingtoHeimerand
mixed forests; but also MT, junipers) (Huhta Nentwig(1991),itlivesinmoisthabitatsintrees
1965, 1971, Palmgren 1972, 1974b, 1977a, Bi- andbushes,andonshoresunderstones.
ström&Väisänen1988,Pajunenetal.1995).It
hasalsobeencaughtinpeatlands(Ledumbogs, ClubionasubsultansThorell,1875
openbogs),andmeadows(Palmgren1977a).Ac- Clubiona subsultans has been caught mostly in
cording to Heimer and Nentwig (1991), H. forests,insomestudiesespecially inconiferous
sanguinealivesinwarm,sunnyhabitats. trees,andamongthemespeciallyinspruces(e.g.
CT,coastalPineta,VT,MT,OMT,mixedforests,
hazel groves, peaty forests) (Palmgren 1943,
3.2.Clubionidae 1972,1977a,Huhta1965,1971,Lehtinenetal.
1979,Palmgren&Biström1979,Biström&Väi-
ClubionacaerulescensL.Koch,1867(Clubiona sänen1998,Heimer&Nentwig1991,Pajunenet
coerulescens) al.1995).Ithasmorerarelybeencaughtinbogs
Clubionacaerulescenswascaughtinforests(e.g. (Ledum bogs, Myrica-Molinia bogs) (Palmgren
mixedforests,MTwithnotveryclosedgrowth, 1972),andinmeadows(Palmgren1977a).
deciduoustreesandbushes;alsospruceforests)
(Palmgren 1943, 1964, 1977a, Heimer & Nent- ClubionasubtilisL.Koch,1867
wig1991),andLedumbogs(Palmgren1977a). Clubionasubtilislivesinmoisthabitats(Heimer
&Nentwig1991):bogs(mainlyMyrica-Molinia
ClubionakulczynskiiLessert,1905 bogs, also open ones, shore bogs) (Palmgren
Clubiona kulczynskii has been caught in forests 1943, 1972), and moist meadows (Palmgren
(e.g. spruce or spruce-dominated forests, mixed 1972).Ithasalsorarelybeencaughtindrymead-
forests)(Palmgren1964,Koponen1995,Pajunen ows(Palmgren1972,1977a).
et al. 1995). Singletons have been found near a
spring(Palmgren1943)andinamoistmeadow ClubionatrivialisC.L.Koch,1843
(Palmgren 1972). According to Heimer and Clubiona trivialis is most commonly associated
Nentwig(1991),however,C.kulczynskiiisaspe- with moderate tree cover and dryish habitats. It
ciesofraisedbogs. has been found in light forests (e.g. coastal
Pineta,CT,VT,mixedforests,coniferoustrees)
ClubionalutescensWestring,1851 (Lehtinen&Kleemola1962,Huhta1965,1971,
Clubiona lutescens has mainly been caught in Palmgren1972,1977a)andLedumbogs(lessin
light forests (e.g. deciduous forests, mixed for- Myrica-Molinia bogs) (Palmgren 1972). It has
ests,birch-mixedmires,forestedges;morerarely more occasionally been caught in meadows, at
shady forests) (Palmgren 1943, 1964, 1972, least dry ones (Palmgren 1972, 1977a), and in
1977a, Lehtinen & Kleemola 1962, Lehtinen et skerries(Palmgren1972).
al.1979).Ithasalsobeencaughtinmoistmead-
ENTOMOL.FENNICAVol.15 (cid:127) Habitataffinitiesofborealspiders 153
3.3.Corinnidae Mastigusa arietina (Thorell, 1871) (Tuberta
arietina,Tetrilusarietinus)
Phrurolithusfestivus(C.L.Koch,1835) Mastigusaarietinahasmostlybeenfoundinant
Phrurolithusfestivushasbeencaughtinlightfor- nests (Formica rufa coll., Lasius fuliginosus) at
ests(e.g.hazelgroves,mixedforests,CT,VT,a least in dark spruce forests (Palmgren 1972,
forest fringe at a shore, overhanging grass in 1977b,Heimer&Nentwig1991).Accordingto
scarps; more rarely MT, junipers) (Palmgren Heimer and Nentwig (1991), it lives also under
1972,1977a,Biström&Väisänen1988,Heimer bark. Huhta (1971) caught one individual in a
&Nentwig1991,Pajunenetal.1995),peatlands burnt-overarea.
(e.g.openbogs,Myrica-Moliniabogs,pinebogs)
(Palmgren1972,Koponenetal.2001),meadows 3.5.Gnaphosidae
(mostlydryones)(Palmgren1972,1977a),lawns
andheaths(Heimer &Nentwig1991, Koponen Drassodespubescens(Thorell,1856)
2000), skerries (Palmgren 1972), shores (Lehti- Drassodespubescenshasbeenfoundinforests,
nen&Kleemola1962,Heimer&Nentwig1991, mainlydryandlightones(e.g.rockypineforests,
Koponen 2000), dry open habitats (Lehtinen et CT,VT,mixedforests;morerarelysprucemires)
al. 1979), under Cladonia on rocks (Palmgren (Palmgren 1964, 1972, 1977a, Huhta 1971,
1943),andclear-cuts(Huhta1971). Heimer & Nentwig 1991, Pajunen et al. 1995),
butalsoindrytomedium-moistmeadows(Huhta
1971, Palmgren 1972, 1977a), grasslands and
3.4.Dictynidae heaths(Heimer&Nentwig1991),clear-cutsand
burnt-overs (Huhta 1971), shores (Lehtinen &
Dictynaarundinacea(Linnaeus,1758) Kleemola1962),andpeatlands(e.g.openbogs,
Dictyna arundinacea is mostly associated with Calluna bogs, Myrica-Molinia bogs, Ledum
sites of moderate tree cover, but with varying bogs,Alnusswamps,Phragmitesfens,apeathar-
moisture.Ithasbeencaughtinlightpineforests, vesting area) (Koponen 1968, 1979, 2002a,
mostlydryones(e.g.rockypineforests,VT,for- 2002b,Palmgren1972,1977a,Heimer&Nent-
estedges)(Palmgren1964,1972,1977a,1977b, wig1991,Koponenetal.2001).
Biström & Väisänen 1988, Heimer & Nentwig
1991), peatlands (e.g. open bogs, Myrica- Drassyllus praeficus (L. Koch, 1866) (Zelotes
Molinia bogs, Ledum bogs) (Palmgren 1972, praeficus)
1977a), and meadows (Palmgren 1964, 1977a, According to Heimer and Nentwig (1991),
Heimer&Nentwig1991).Itlivesmainlynearthe Drassylluspraeficuslivesindryandlightsites,
groundand–morerarely–intrees,e.g.junipers, e.g.rockyhabitatsandlawns.MostoftheFinnish
pinesandspruces(Palmgren1964,1972,1977a). findingshavebeeninshoresandheaths(Kopo-
nen2000).Otherfindingsincludefen-likeAlne-
DictynapusillaThorell,1856 tum (Palmgren 1972), a Myrica-Sesleria bog
Dictynapusillahasmostfrequentlybeencaught (Palmgren 1972), a spruce-pine forest of VT
in coniferous trees (junipers, pines, spruces) (Palmgren 1972), grass on rocks (Palmgren
(Lehtinen & Kleemola 1962, Palmgren 1972, 1972),averydrysublapidicol(Palmgren1964),
1977a,1977b).Ithasalsooccasionallybeencap- anddryopenhabitats(Lehtinenetal.1979).
turedinpineandspruceforestsneartheground
(Huhta1971,Palmgren1972,1977a,1977b),in Drassyllus pusillus (C. L. Koch, 1833) (Zelotes
Ledumbogs(Palmgren1977a),andinmeadows pusillus)
(Palmgren 1977a). According to Heimer and According to Heimer and Nentwig (1991),
Nentwig(1991),thespecieslivesinherbs,bushes Drassylluspusilluslivese.g.inforests,meadows,
andtreesinsunnyhabitats. heaths,andsand.Ithasbeencaughtinpeatlands
[e.g. open bogs; in Lithuania also pine bogs; a
xerophilous mire species in Koponen (1968)]
(Koponen1968,Koponenetal.2001,Koponen
154 Matveinen-Huju (cid:127) ENTOMOL.FENNICAVol.15
2002b), dry heaths (Koponen 2000), clear-cuts tinen and Kleemola (1962) caught one female
andburnt-overs(Huhta1971),forests(Pajunenet amonglitter.
al. 1995), and under Cladonia on rocks (Palm-
gren1943). Haplodrassusmoderatus(Kulczyn’ski,1897)
Haplodrassus moderatus has been caught in
Gnaphosabicolor(Hahn,1833) more or less open peatlands (e.g. open bogs,
Gnaphosabicolorhasbeencaughtmostlyinlight Myrica-Molinia bogs, fens; in Heimer &
forests(CT,rockyPineta,VT,mixedforests,old Nentwig (1991) also peaty forests) (Palmgren
forests; more rarely shady forests) (Palmgren 1972,Koponen1978,2002a,2002b,Lehtinenet
1943, 1964, 1972, 1977a, Huhta 1965, 1971, al. 1979, Heimer &Nentwig 1991, Koponen et
Heimer&Nentwig1991,Koponen1995,Paju- al. 2001), drift-sand shores and shore meadows
nenetal.1995),butithasalsobeenabundantly (Palmgren 1943), and other medium-moist to
foundinclear-cutsandburnt-overs(Huhta1965, moistmeadows(onlyrarelydryones)(Palmgren
1971). G. bicolor has also been caught in grass 1972, 1977a, Heimer & Nentwig 1991), fields
ribbons on small islands (Palmgren 1972), (Palmgren1943),andunderloosebarkoffence
Myrica-Molinia bogs (Palmgren 1972), mead- stakes(Palmgren1943).
ows(Palmgren1964,Huhta1971),verydrysub-
lapidicol(Palmgren1964),anddryopenhabitats Haplodrassussignifer(C.L.Koch,1839)
(Lehtinen et al. 1979). Other habitat types in Haplodrassussigniferhasmostlybeencaughtin
Heimer and Nentwig (1991) include rocky clear-cuts and burnt-overs (Huhta 1971), but it
steppes. has also been found in light forests (e.g. hazel
groves,mixedforests,coastalPineta,VT,rocky
GnaphosamicropsHolm,1939 pine forests; rarely shady ones) (Huhta 1971,
Gnaphosa microps has been caught mostly in Palmgren1972,1977a,Lehtinenetal.1979,Bi-
peatlands, such as open bogs (Lehtinen et al. ström & Väisänen 1988, Heimer & Nentwig
1979, Heimer &Nentwig 1991, Koponen etal. 1991,Koponen&Niemelä1994,Koponen1995,
2001,Koponen2002a,2002b),butoneindivid- Pajunenetal. 1995), peatlands[e.g. openbogs,
ual has also been found in a clear-cut (Huhta Myrica-Molinia bogs, pine bogs; a xerophilous
1965).AccordingtoPalmgren(1943),G.microps mire species according to Koponen (1968)]
is a rare northern species. Other habitats in (Palmgren1972,1977a,Koponenetal.2001,Ko-
Heimer and Nentwig (1991) include light ponen2002a,2002b),meadows(Palmgren1943,
heather-birchforestsandmeadows. 1972,Huhta1971),lawnsandheaths(Heimer&
Nentwig1991),dryopensublapidicol(Palmgren
Gnaphosamontana(L.Koch,1866) 1977a),Pleuroziumonskerries(Palmgren1972),
The few findings of Gnaphosa montana have androcks(Palmgren1943).
been from forests (e.g. under the bark of trees,
VT)(Palmgren1943,1972,Huhta1971,Heimer Haplodrassussilvestris(Blackwall,1833)
&Nentwig1991,Pajunenetal.1995),clear-cuts Most of the few findings of Haplodrassus
and burnt-overs (Huhta 1971), very dry sub- silvestrishavebeeninforests(aforestfringeata
lapidicol(Palmgren1964),theouterwallsandin- shore, birch forests or birch-mixed stands of
sideahouse(Palmgren1977a),anddryopenhab- OMT) (Palmgren 1977a, Heimer & Nentwig
itats(Lehtinenetal.1979). 1991). Two individuals have been reported in
meadows(Huhta1971,Palmgren1977a).Other
Haplodrassuscognatus(Westring,1861) habitattypesinHeimerandNentwig(1991)in-
ThefewfindingsofHaplodrassuscognatushave cludedrylawnsandbogs.
mostlybeenfromtrees(underbark,instems;e.g.
oaks)(Palmgren1943,1972,1977a,Lehtinen& Haplodrassussoerenseni(Strand,1900)
Kleemola 1962, Koponen 1996). According to Haplodrassus soerenseni seems to be a forest
Heimer and Nentwig (1991), however, the spe- species (e.g. CT, rocky pine forests, VT, mixed
ciesliveswithinleaflitterinforests;indeed,Leh- forests,MT)(Huhta1965,1971,Palmgren1972,
ENTOMOL.FENNICAVol.15 (cid:127) Habitataffinitiesofborealspiders 155
Palmgren&Biström1979,Biström&Väisänen Zeloteslatreillei(Simon,1878)
1988,Koponen1995,Heimer&Nentwig1991, Zeloteslatreilleihasbeencapturedinlightforests
Pajunen et al. 1995). It has also been caught in (e.g.oldpineforest,VT,underloosebark)(Palm-
peatlands(e.g.openbogs,pinebogs)(Heimer& gren1943,1972,1977a,Huhta1971,Heimer&
Nentwig1991, Koponenetal. 2001), meadows Nentwig 1991, Koponen 1995, Pajunen et al.
(Huhta 1971, Palmgren 1977a), and clear-cuts 1995),understonesandlichens(Palmgren1943,
andburnt-overs(Huhta1971). Lehtinen&Kleemola1962),peatlands[e.g.pine
bogs, open bogs; a xerophilous mire species in
MicariaaeneaThorell,1871 Koponen (1968)] (Koponen 1968, Heimer &
Micariaaeneahasbeenfoundinforests(e.g.VT, Nentwig 1991, Koponen et al. 2001, Koponen
overhanging grass in rock scarps) (Huhta1971, 2002b), meadows (Huhta 1971), dry open
Palmgren1977a,Pajunenetal.1995),clear-cuts sublapidicol(Palmgren1964,1977a),andclear-
andburnt-overs(Huhta1971),meadows(Huhta cutsandburnt-overs(Huhta1971).
1971),andshores(Palmgren1943).
Zelotespetrensis(C.L.Koch,1839)
Micariapulicaria(Sundevall,1831) Zelotespetrensishasbeenfoundinlightforests
In most studies Micaria pulicaria has mainly (e.g. a polluted dry pine forest with almost no
been caught in open habitats: clear-cuts and vegetation) (Palmgren 1943, 1977a, Heimer &
burnt-overs(Huhta1971),drytomoistmeadows Nentwig1991,Koponen&Niemelä1994,1995),
(Palmgren 1943, 1972, 1977a, Huhta 1971), on rocks in shores (Palmgren 1943), dry mead-
shores (Palmgren 1943, 1964, 1972), treeless ows (Palmgren 1977a), heaths and lawns
skerries(Palmgren1972),openpeatlands(Myri- (Heimer & Nentwig 1991), clear-cuts (Huhta
ca-Molinia bogs, open bogs) (Palmgren 1972, 1971), and dry open habitats (Lehtinen et al.
1977a), and roads (Palmgren 1943). However, 1979).
Palmgren(1977a) caughtitinlightforests(e.g.
rockypineforests,aforestfringeatashore,over- Zelotessubterraneus(C.L.Koch,1833)
hangingmossinscarps)asabundantlyasinopen Zelotessubterraneuswasfoundinshores(Palm-
habitats,andPajunenetal.(1995)caughtafew gren 1943, 1972, Lehtinen & Kleemola 1962,
individuals in forests. Heimer and Nentwig Koponen 2000), open rocks (Palmgren 1972),
(1991)statethatM.pulicarialivesinwell-lithab- verydrysublapidicol(Palmgren1964),dryopen
itats. habitats(Lehtinen etal. 1979), andforests(e.g.
rockyforests,coastalPineta,VT,mixedforests,
MicariasilesiacaL.Koch,1875 overhanginggrassinscarps,stemsofpines,afor-
Micaria silesiaca has been caught in dry open estfringeatashore;alsoMT)(Palmgren1943,
habitats (e.g. caves, among loose rocks) (Lehti- 1972,1977a,Palmgren&Biström1979,Heimer
nen et al. 1979), dry heaths (Koponen 2000), &Nentwig1991,Koponen1995).Otherhabitats
burnt-overs(Huhta1971),andforests(Pajunenet mentioned in Heimer and Nentwig (1991) are
al.1995). heaths,lawnsandbogs.
Zelotes clivicola (L. Koch, 1870) (Zelotes
clivicolus) 3.6.Hahniidae
Zelotesclivicolahasbeencaughtinforests(e.g.
CT, VT) (Huhta 1965, 1971, Palmgren 1972, Cryphoecasilvicola(C.L.Koch,1834)
1977a, Heimer & Nentwig 1991, Koponen & Cryphoecasilvicolahascommonlybeenfoundin
Niemelä 1994, Pajunen et al. 1995), peatlands shady forests,bothintreesandneartheground
(e.g. Myrica-Molinia bogs) (Palmgren 1972, (MT, OMT, less often spruce mires) (Palmgren
Heimer&Nentwig1991),rockyhabitats(Palm- 1964, 1972, 1977a, 1977b, Huhta 1965, 1971,
gren1964),meadows(Huhta1971),andheaths Palmgren&Biström1979,Biström&Väisänen
(Heimer&Nentwig1991),butmostabundantly 1988,Väisänen&Biström1990,Koponen1995,
inclear-cutsandburnt-overs(Huhta1971). 1999,Pajunenetal.1995).Ithasmorerarelyoc-
156 Matveinen-Huju (cid:127) ENTOMOL.FENNICAVol.15
curredinlightforests(VT,coastalPineta,mixed 3.7.Linyphiidae
forests,hazelgroves)(Huhta1965,1971,Palm-
gren 1972, 1977b, Biström & Väisänen 1988, Abacoproecessaltuum(L.Koch,1872)
Koponen 1995, Pajunen et al. 1995), peatlands Abacoproecessaltuumhasmostlybeenfoundin
(Koponen 1978), and clear-cuts (Huhta 1965, light forests (e.g. a forest fringe at a shore, VT,
1971).HeimerandNentwig(1991)didnotsepa- coastalPineta,mixedforests;rarelyinthedriest
ratethetypesofconiferousforests. foresttypes)(Huhta1965,1971,Palmgren1972,
1977a, Koponen 1995, Pajunen et al. 1995). It
Hahnianava(Blackwall,1841) hasmorerarelybeencaughtinshadyforests(e.g.
Hahnianavahasbeencaughtinlightforests(e.g. MT) (Palmgren 1977a, Palmgren & Biström
CT, spruce-pine forests, coastal Pineta, rocky 1979), near meadow ditches (Palmgern 1977a),
pineforests)(Palmgren1964,1972,1977b,Paju- androcks(Lehtinen&Kleemola1962,Palmgren
nenetal.1995),butalsoinmeadows(mostrarely 1972, 1976). A. saltuum has an affinity for airy
moistones)(Palgmren1972,1977b),islandsand groundstructure(Palmgren1976,1977a),andit
skerries (e.g. in grass ribbons, Pleurozium) has been caught within mosses and grasses
(Palmgren 1972), and very dry sublapidicol (Palmgren1972,1976).
(Palmgren1964).
Agnyphantesexpunctus(O.P-Cambridge,1875)
HahniaononidumSimon,1875(Hahniamengei (Lepthyphantesexpunctus)
Chyzer&Kulczyn’ski,1897) Agnyphantes expunctus has mostly been caught
Hahnia ononidum seems to be a forest species in trees, at least spruce(Palmgren 1964, 1977a,
(mainly not very dry ones, e.g. MT, VT, mixed Heimer&Nentwig1991).Ithasalsobeencaught
spruce-pineforest;butalsorockypineforest,CT) inthelowerstrataofforests(e.g.oldsprucefor-
(Huhta 1965, 1971, Palmgren 1972, 1977b, ests, spruce-deciduous mixed forest, forest
Palmgren&Biström1979,Biström&Väisänen edges) (Palmgren 1964, Koponen 1995). A. ex-
1988, Heimer & Nentwig 1991, Pajunen et al. punctusisanorthernspecies(Palmgren1977a).
1995), although Huhta (1971) caught it quite
abundantlyalsoinclear-cuts. Agynetacauta(O.P-Cambridge,1902)
Agynetacautahasbeencaughtinforests(e.g.CT,
HahniapusillaC.L.Koch,1841 VT, MT, OMT, mixed forests, hazel groves)
MoistureseemstobemoreimportantforHahnia (Huhta1965,1971,Palmgren1972,1975,1977a,
pusillathanthelightintensity(Palmgren1972). Heimer&Nentwig1991,Koponen1995,Paju-
AccordingtoPalmgren(1977b),thespeciescan nenetal.1995),andpeatlands(e.g.sprucemires,
befoundindrierhabitatsmainlyinspringandau- Ledum bogs, Myrica-Molinia bogs, open bogs)
tumn (Palmgren 1977b). H. pusilla has been (Palmgren 1964, 1972, 1975, 1977a, Koponen
caughtindifferenttypesofpeatlands(e.g.open 1978, 2002a, 2002b, Heimer & Nentwig 1991,
bogs,Myrica-Moliniabogs,Ledumbogs,spruce Koponenetal.2001).Ithasalsobeencaughtin
mires)(Palmgren1964,1972,1977a,1977b,Ko- meadows (Palmgren 1964, Huhta 1965, 1971),
ponen1978,Lehtinenetal.1979),butithasalso sandyshores(Palmgren1964),andclear-cutsand
beencapturedinforests(e.g.hazelgroves,MT, burnt-overs(Huhta1965,1971).
VT,drypineforests,coastalPineta,oaks)(Palm-
gren1964,1972,1977a,1977b,Huhta1965,Bi- Agynetaconigera(O.P-Cambridge,1863)
ström & Väisänen 1988, Väisänen & Biström Agynetaconigerahasbeencharacterisedasbeing
1990,Pajunenetal.1995,Koponen1996).Afew aforestspecies,beingcaughtmostlyinconifer-
findingshavebeenfromclear-cuts(Huhta1971), ous forests (e.g. CT, VT, coastal Pineta, MT,
medium-moist to moist meadows (Palmgren OMT,oldforests,hazelgroves)(Palmgren1964,
1972), and very dry sublapidicol (Palmgren 1972,1977a,Huhta1965,1971,Palmgren&Bi-
1964). ström1979,Heimer&Nentwig1991,Koponen
1995, 1999, Pajunen et al.1995). Some of the
findings have been from peatlands (e.g. pine
ENTOMOL.FENNICAVol.15 (cid:127) Habitataffinitiesofborealspiders 157
bogs, open bogs, Myrica-Molinia bogs, spruce (Palmgren 1977a), and peatlands (e.g. Myrica-
mires) (Palmgren 1972, Koponen et al. 2001), Molinia bogs) (Palmgren 1972, Koponen 1978,
meadows and shore meadows (Huhta 1971, Heimer &Nentwig1991). However, ithasalso
Palmgren1972,1977a),edgesofquagmiresand been caught in spruce forests of OMT (Huhta
brooks (Palmgren 1977a), clear-cuts (Huhta 1971),andinmixedforests(Palmgren1964).
1971),androckyshelves(Palmgren1972).Itcan
be caught in moss (Heimer & Nentwig 1991), Anguliphantes angulipalpis (Westring, 1851)
“förna”,orgrass(Palmgren1975),butitmayalso (Lepthyphantesangulipalpis)
ascendhigher(Palmgren1972,1975,1977a). Anguliphantes angulipalpis has mainly been
caughtinforests(e.g.fen-likeandshoreAlneta,
Agynetaolivacea(Emerton,1882) deciduousforests,mixedforests,aforestfringeat
ThebiologyofAgynetaolivaceaisinsufficiently ashore,coastalPineta,VT,drypineforests,MT,
known (Heimer & Nentwig 1991). It has been spruce mires, junipers) (Palmgren 1964, 1972,
foundinforests(e.g.anolddensespruceforest,a 1975,1977a,Huhta1971,Lehtinenetal.1979,
mixedforestofbirch,pineandspruce)(Koponen Palmgren & Biström 1979). Some individuals
1995,Pajunenetal.1995). have also been captured in clear-cuts (Huhta
1971), edges of fields (Palmgren 1972, 1975),
AgynetaramosaJackson,1912 grassribbonsinsmallislands(Palmgren1972),
Agynetaramosahasmainlybeencaughtindiffer- andmeadows(Palmgren1977a).A.angulipalpis
enttypesofforest(especiallymossinconiferous liveswithinlitter(Heimer&Nentwig1991).
forests,e.g.VT,coastalPineta,MT,OMT,spruce
mires,mixedforests,hazelgroves)(Huhta1971, Asthenarguspaganus(Simon,1884)
Palmgren 1972, 1977a, Lehtinen et al. 1979, Asthenarguspaganushasmainlybeencaughtin
Palmgren&Biström1979,Koponen1995,1999, forests (especially spruce forests, mixed forests
Pajunenetal.1995).Someindividualshavebeen with spruce, or spruce mires) (Palmgren 1964,
caught in peatlands (e.g. open bogs) (Palmgren 1976,1977a,Biström&Väisänen1988,Heimer
1972,1977a,Koponen1978,Heimer&Nentwig &Nentwig1991,Koponen1995,1999,Pajunen
1991), and clear-cuts (Huhta 1965, 1971). An- et al. 1995). Lehtinen et al. (1979) caught A.
otherhabitattypeinHeimerandNentwig(1991) paganusinabog,butstatedthatitinhabitsmires
wastrees. onlyinperipherallocations.Itisaneasternspe-
cies(Palmgren1977a).
Agynetasubtilis(O.P-Cambridge,1863)
Agynetasubtilishasmainlybeenfoundindiffer- Bathyphantesgracilis(Blackwall,1841)
enttypesofforest(e.g.CT,VT,MT,OMT,spruce Bathyphantes gracilis has been caught in moist
mires,oldforests;alsodeciduousforests)(Palm- habitats:meadows(morerarelydrytomedium-
gren1964,1972,1975,1977a,Huhta1965,1971, moistones)(Huhta1971,Palmgren1972,1977a,
Palmgren & Biström 1979, Heimer & Nentwig Heimer&Nentwig1991),andpeatlands(Sphag-
1991,Koponen1995,1999,Pajunenetal.1995). num in small depressions in semi-open forest,
Someindividualshavealsobeencaughtinclear- spruce mires, open bogs, Myrica-Molinia bogs,
cuts(Huhta1965,1971),Ledumbogs,openbogs, fen-like Alneta) (Koponen 1968, 1978, 2002a,
and moist meadows (Palmgren 1972, 1975, 2002b, Palmgren, 1972, 1977a, Lehtinen et al.
1977a,Heimer&Nentwig1991). 1979, Koponen et al. 2001). It has also been
caught in smaller numbers in clear-cuts (Huhta
Allomengeascopigera(Grube,1859) 1971),andforests(Coryleta,MT,mixedforests)
Allomengeascopigerahasmostlybeenfoundin (Palmgren 1972, 1977a, Heimer & Nentwig
moistsites:shoreandfen-likeAlneta(Palmgren 1991).
1972,1975,Lehtinenetal.1979),shores(Palm-
gren 1972, 1975), moist meadows (Palmgren Bathyphantesnigrinus(Westring,1851)
1972,1975,1977a),nearditchesinforests(Palm- The most typical habitat of Bathyphantes
gren 1977a), edges of quagmires or brooks nigrinus varies among different studies: hazel
158 Matveinen-Huju (cid:127) ENTOMOL.FENNICAVol.15
groves(Palmgren1972,1975),bogswithshrubs quite abundantly in meadows (Palmgren 1964,
(Myrica, Ledum, small birches etc.) and some 1972, 1977a, Huhta 1971, Heimer & Nentwig
cover ofscattered trees (Palmgren 1972, 1975), 1991). It has also been captured in peatlands
moist meadows (Palmgren 1977a), and spruce (Myrica-Molinia bogs, Ledum bogs) (Palmgren
mires (Palmgren 1977a, Heimer & Nentwig 1972,1977a)andclear-cuts(Huhta1971).
1991).B.nigrinuswasalsocaughtindrytome-
dium-moist meadows (Palmgren 1964, 1972, Bolyphantesluteolus(Blackwall,1833)
1977a), other types of forests (e.g. MT, mixed Bolyphantes luteolus has been caught in light
forests, forest edges) (Palmgren 1964, 1972, pinestands(e.g.coastalPineta)(Palmgren1972,
1977a), and roadsides or cleared forests (Palm- 1975), Elymus arenarius on sandy beaches
gren1977a). (Palmgren 1972, 1975), small islands and
skerries (Palmgren 1972, 1975), dry meadows
Bathyphantesparvulus(Westring,1851) (Palmgren 1972, 1975), and bogs (e.g. open
The habitat distribution of Bathyphantes par- bogs)(Koponen1978,Koponen2002b).
vulus resembles that of B. gracilis (Palmgren
1972, Heimer & Nentwig 1991): moist to me- Centromerusarcanus(O.P-Cambridge,1873)
dium-moist meadows (more rarely dry ones) Centromerusarcanushasbeencaughtinforests
(Palmgren1964,1972,1975,1977a,Huhta1971, (e.g. CT, coastal Pineta, VT, MT, OMT, mixed
Lehtinenetal.1979,Heimer&Nentwig1991), forests, spruce mires, hazel groves) (Palmgren
and peatlands (e.g. open bogs, Myrica-Molinia 1964, 1972, 1975, 1977a, Huhta 1965, 1971,
bogs, Alnus swamps, Phragmites fens, Calluna Palmgren&Biström1979,Biström&Väisänen
bogs,apeatharvestingarea,Sphagnuminbirch- 1988, Väisänen & Biström 1990, Heimer &
mixed forest and depressions) (Palmgren 1972, Nentwig1991,Koponen1995,1999,Pajunenet
1977a, Koponen 1979, 2002b, Lehtinen et al. al.1995),butthecatchesinclear-cutsandburnt-
1979).Ithasalsobeencaughtinclear-cuts(Huhta overshavealsobeenreportedhigh(Huhta1971),
1971), treeless skerries and forests (e.g. pine- and it has also been regular in peatlands (e.g.
birchforests)(Palmgren1972,1977a,Pajunenet Alnusswamps,Phragmitesfens,Callunabogs,a
al. 1995). Other habitat types mentioned in peatharvestingarea,sprucemires,Ledumbogs,
HeimerandNentwig(1991)wereforests. Myrica-Molinia bogs, open bogs) (Palmgren
1964,1972,1975,1977a,Koponen1978,1979,
Bolephthyphantes index (Thorell, 1856) (Boly- 2002a,2002b,Lehtinenetal.1979,Koponenet
phantesindex) al. 2001). Few findings exist in dry to moist
Bolephthyphantesindexhasbeenfoundinconif- meadows,sandyshores,treelessskerries,andco-
erousforests(e.g.Sphagnuminsmalldepressions niferous trees (Palmgren 1964, 1972, 1975,
insemi-openforest,rockypineforests,VT,MT; 1977a).
also junipers, spruces) (Palmgren 1964, 1977a,
Huhta1971,Biström&Väisänen1988,Pajunen Centromerussylvaticus(Blackwall,1841)
et al. 1995), and peatlands (Ledum bogs, open Centromerussylvaticushasbeenfoundinforests
bogs, Betula nana) (Palmgren 1972, 1975, (e.g.CT,coastalPineta,VT,mixedforests,hazel
1977a).However,accordingtoHeimerandNent- groves; rarely spruce forests of MT, OMT)
wig (1991), B. index lives e.g. in tree roots and (Palmgren1964,1972,1975,1977a,Huhta1971,
cleftsofrocks. Heimer & Nentwig 1991, Pajunen et al. 1995),
dry to moist meadows (Huhta 1971, Palmgren
Bolyphantesalticeps(Sundevall,1833) 1972, 1975, 1977a), clear-cuts and burnt-overs
Bolyphantes alticeps has mainly been caught in (Huhta 1971), and peatlands (e.g. open bogs,
forests (e.g. CT, coastal Pineta, mixed forests, Myrica-Molinia bogs, Ledum bogs) (Palmgren
VT,forestedges,spruceforests,sprucemires,ha- 1972,1977a).
zel groves) (Lehtinen & Kleemola 1962, Palm-
gren1964,1972,1975,1977a,Huhta1965,1971, Ceratinellabrevis(Wider,1834)
Heimer & Nentwig 1991), but sometimes also Ceratinellabrevisisrathereurytopic,althoughit