Spider (Arachnida: Araneae) distribution across the timberline in the Swiss Central Alps (Alp Flix, Grisons) and three morphologically remarkable species

2007·6.9 MB·English

by  Patrick Muff

Arachnol.Mitt.34:16-24 Nürnberg,Dezember2007
Spader (Arachnida:Ararteae) distribution across the timberline in the Swiss
Central Alps (Alp F!ix,Grisons) and three morphologically remarkable species
PatrickMuff,Martin H.Schmidt,Holger Frick,Wolfgang Nentwig &Christian Kropf
Abstract:Wecollected6251 adultepigeicspidersfromthedwarf-shrubheathtosubalpineconiferousforest
on Alp Flix (CH,canton Grisons, 1950 m) between May2005 and May2006 using pitfall traps.Total species
richnessandactivitydensityofall speciesdecreasedfromtheopen landtotheforest,althoughthispattern
varied according tofamily.Thedistribution ofthe 102 speciesfound indicatesthatthe small area around a
singletreeatthetimberline provides habitatsforboth open land andforestspiderspeciesaswell as some
possibletimberlinespecialists.FivespecieswerenewtothecantonGr\sor\s:Centromeritabicolor,Centromeritaconcinna,
Hilairaexcisa,Meionetaalpicaand Tallusiaexperta.Three species showed remarkable morphological characteristics
andwereanalysedinmoredetail.WefoundmalesofPelecopsisradicicolawithoutthecharacteristiclongitudinal
depression on the raised carapace.It is shown thatthe males ofMeionetaalpicahave a considerablyvariable
lamellacharacteristica,which isneverthelessdistinctfromthesisterspeciesMeionetaressli.Becausewefound
intermediate forms ofthe head region described for Metopobactrusprominulusand M.schenkeli, respectively,M.
schenkeliisconsideredasyn.nov.ofM.prominulus.Vr\\sstudyshowsthattheknowndistributionandtaxonomic
statusofvariousspidertaxaintheCentralAlpsarestillincompleteandfurtherworkonarthropodsinremote
areasshould bestronglyencouraged.
Keywords:Diversity,dwarf-shrubheath,forest,habitatboundary,Meionetaalpica,Metopobactrusprominulus,Pelecopsis
radicicola,pitfalltraps
Knowledge about species diversity and activity here.Besidesashortdescriptionofthedistribution
across landscapes is indispensable to understand patternofspecies from the openlandto theforest,
howlandscape structure influences habitatquality. wefocusonthreemorphologicallyremarkablespe-
Sincelandscapes existas mosaicsofnumerous dif- cies, two ofthem causingproblems ofidentificati-
A
ferentpatchtypes,knowledgeabouttheecologyof on since theirfirst description. discussion about
habitatedgesisofparticularimportance (e.g. RlES otherfaunisticallynotable species ofthe studysite
etal.2004). Spidersareabundant,species-richand canbe found in FRICKet al. (2006).
knowntorespondsensitivelytoenvironmentaland
structuralconditions,whichmakesthemsuitablefor MaterialandMethods
studyingorganism—habitatrelationships(e.g.WISE Studysite
1993, FOELIX 1996). However, despite extensive Alp Flix (WGS84: 9°38’E, 46°31’N) is partofthe
work,ourknowledgeaboutthedistributionpattern Swiss Central Alps and belongs to the village of
ofspiders across the Central Alps in general and Surin the canton Grisons, Switzerland.The alp is
Switzerland inparticularis stilllimited (MAURER a southwest-exposed terrace of15 km2 at 1950 m
&Hänggi 1990, Hänggi 1993,Thaler 1995, above sealevel.Itis surroundedby3000 m moun-
HÄNGGI 1999, HÄNGGI 2003). A recent study tainpeaks andavalley. Samplingwasconductedin
m
analysing the influence ofenvironment and space a300 longstretchoftimberlineplusfragmentsof
onthedifferentiationofspidercommunitiesacross the adjoining Norway spruce forest (Vaccinio-Pi-
analpinetimberlineinSwitzerland(MUFFetal.in ceion) tothenorthwestandthedwarf-shrubheath
prep.) provided faunistic data, which are reported (Juniperion nanae) to the southeast. Each ofthe
three parts covered approximately3 ha.The site is
located on a small slope inclined slightly towards
PatMruicskeMuUmFFB,erHno,lDgeepraFrRtImCeKn&tCohfriInsvteiratneKbrRaOtPeFs,,NBaetrunraasltrHaisssteor1y5, the forest and is used for occasional cattle grazing
CH-3005Bern,Switzerland,Email:[email protected], throughout the vegetation period. For a more de-
[email protected],[email protected] taileddescriptionofthe studyareaseeHÄNGGI &c
MartinH.SCHMIDT&WolfgangNENTWIG,ZoologicalInstitute, Müller
(2001).
UniversityofBern,Baltzerstrasse6,CH-3012Bern,Switzerland,
Email:[email protected], wolfgang.nentwig@zos.
unibe.ch
SpiderdistributionintheSwissAlps 17
Studydesignandspidersampling meterof6.9 cmandadepthof7.5 cmfilledwitha
Wedifferentiatedbetweenfivehabitatzoneswhich solutionof4%formaldehydeinwaterplusdetergent
represented the whole gradient ofhabitat structu- (0.05% sodium dodecylsulphate, SDS). Eachtrap
res: the open land (dwarf-shrub heath, O), three wascoveredwithaquadrangulartransparentplastic
microhabitats linked to a single spruce tree at the roof(15x 15 cm) fixedbythreewoodenrods 8 cm
timberlineandtheforest(F).Thethreeareasatthe above ground. Due to the proximity ofcattle and
timberlineweredefinedbytheirlocationrelativeto the toxicity of the trapping liquid we fenced off
the tree as: next to the trunk (TT), at the end of each trap with three plastic poles connected with
branch cover (TB) and in the adjoining open area ribbons. The traps were emptied monthly during
outside ofbranch cover (TO). In each ofthe five the snow-freeperiod(May2005 to October2005)
habitatzonesweplaced15pitfalltraps.InO,Fand andthenleftunderthesnowlayeruntilMay2006,
TO thetrapswererandomlypositioned atleast 15 when theywere emptied a final time.
m
apart from each other. For placing the traps in
TB andTT,the tree nearestto the trap inTOwas Identification
chosen. The mean distances between the traps in Onlyadult spiders were identified to species level,
these three microhabitats were 4.1 m (TO -TB), juvenileswere excluded. Identificationwas mainly
4.5 m (TO-TT) and 1.5 m (TB-TT).Thetraps carriedoutusingNENTWIGetal.(2003),ROBERTS
consisted ofwhite plastic cups with an upper dia- (1985, 1987) andWlEHLE (1956, 1960). For ad-
Tab.1:Numberofspecies(andindividuals)accordingtofamilyand season (A)orhabitatzone(B).Lettersbehindindi-
vidual numbersdenoteseasons(excludingwinter)andhabitatssignificantlydifferentfromeachotheraccordingto
pairwiseKruskal-Wallistests(allp<0.05,correctedformultiplecomparisonsafterHolm).June:27.5.-24.6.2005,July:
25.6.-23.7.2005,August:24.7.-21.8.2005,September:22.8.-18.9.2005,October:19.9.-16.10.2005,winter:17.10.2005-
6.5.2006;fordefinitionofhabitatzonesseetext.
(A) June July August September October winter
Linyphiidae 48 (632d) 26 (179b) 25 (113a) 24 (146ab) 25 (365c) 39 (850)
Lycosidae 7(2407d) 8 (658c) 6 (142b) 6 (45a) 6 (20a) 4(8)
Gnaphosidae 7(154d) 6 (46c) 6 (19b) 2 (25b) 4(8a) 3(3)
Thomisidae 7 (54b) 4 (14a) 4 (5a) 2(4a) 2 (5a) 3(9)
Theridiidae 3 (18b) 2 (3a) 2 (22b) 1 (45b) 1 (17b) 2(12)
Philodromidae 1 (5a) 2 (2a) 1 (6ab) 1 (17b) 1 (8ab) 1(22)
Hahniidae 1 (82b) 1 (3a) 1 (4a) 1 (5a) 1 (7a) 1(33)
others 5 (9a) 2 (2a) 4 (6a) 3 (4a) 2 (2a) 5(6)
Total 79 (3361d) 51 (907c) 49(317ab) 40 (291a) 42 (432b) 58 (943)
(B) O TO TB TT F Total
Linyphiidae 34 (251a) 30 (444bc) 33 (423b) 40 (620c) 29 (547bc) 61 (2285)
Lycosidae 7 (1073c) 7 (1089c) 7(788c) 5 (262b) 6 (68a) 8 (3280)
Gnaphosidae 6 (106c) 7 (73c) 7(47b) 3 (10a) 4 (19a) 8 (255)
Thomisidae 6 (32b) 4(25b) 4(27b) 3 (6a) 1 (la) 7(91)
Theridiidae 1 (la) 3 (12ab) 2 (37b) 2 (42b) 1 (25b) 3(117)
Philodromidae 1 (33b) 2 (22b) 1 (5a) 0a 0a 2(60)
Hahniidae 0a 0a 1 (2a) 1 (33b) 1 (99c) 1 (134)
others 4 (10b) 5 (lib) 2(3a) 3 (3a) 2 (2a) 12 (29)
Total 59 (1506c) 57(1676c) 56 (1332bc) 56(976ab) 44 (761a) 102 (6251)
18 PMuff,M.Schmidt,H.Frick,W.Nentwig&Chr.Kropf
ditional literature used see FRICK et al. (2006). low-voltage SEM Hitachi S-3500N for Pelecopsis
Nomenclature followed PLATNICK (2007). The radicicolaandastandardSEM PhilipsXL30FEG
material is stored in the Natural HistoryMuseum forMeionetaalpicaandM. ressli.
ofBern.
Results andDiscussion
Statisticalanalysis Spiderdistributionacrossthealpinetimberline
Forcomparisonsofactivitydensitiesacrossseasons We recorded 6251 adult individuals belonging to
(excludingwinter) andhabitatzonesweusedpair- 102 species of 14 families. Total species richness
wise Kruskal-Wallis tests (k = 9999 Monte Carlo and activity density ofall species decreased from
permutations), because a Levene's test indicated June towards the winter period, with a moderate
thatvariances ofthe species datawere not homo- increaseinOctober(Tab. 1A).Thisgeneralpattern
geneous.These analyses were conducted with the varied at the family level, though. It applied best
programSPSS 14.0forWindows.Allp-valueswere to Linyphiidae,whereas Lycosidae decreased con-
correctedformultiplecomparisonsafterHolm(LE- stantlybothintermsofspeciesandindividualsfrom
&
GENDRE LEGENDRE 1998). June towinter. Both the numberofall species and
individuals decreased from the openland to forest
Morphologicalanalysis (Tab. IB). Species richness ofsingle families did
Forthe analysis ofsome morphological character- not show any clear pattern across the five habitat
istics we used either an optical stereomicroscope zones,althoughinmostfamiliesitappearedtode-
Leica MZ16 for Metopobactrus prominulus, a crease in more shaded habitats.The accumulation
of individuals in open areas, how-
ever,mustbe qualifiedwhenlooking
at single families. While Lycosidae,
Gnaphosidae,Thomisidae and Phi-
lodromidae had higher densities in
open zones, Linyphiidae, Theridii-
dae and Hahniidae preferred more
shaded habitats. Clearly,in the three
habitat zones at the timberline the
speciesmostlyfollowedthesedistinct
trends in activity density across the
openlandandforestinaverygradual
manner(Tab.2).Itis notablethatall
five habitat zones were dominated
by certain species, but no species
(with N > 15 individuals) was found
exclusivelyin onlyone habitat. Nine
specieswere foundwith atleast50%
ofthe individuals in the open land,
six species in the forest and none,
four and six species, respectively, in
thethreezonesatthetimberline(TO,
TB and TT) (Fig. 1). Fig. 1 illust-
rates these species-specific patterns
O TO TB TT F TO TB TT in activity density. For example, the
lycosidAlopecosapulverulenta clearly
Fig.1:Mean number(+SE)ofindividualsofall speciesandofselected preferredtheopenland,whileA. tae-
speciesacrossthefivehabitatzones(datafrom 15 pitfalltrapspooled niatawasfoundmainlyaroundsingle
foreach habitatoveroneyear).Notethateachofthefivespecies
shown isclearlydominantinoneofthehabitats(i.e.occurrencea50%, treesinthetimberline.Otherspecies
exceptforCaradadusavicula:occurrence=38%). (e.g. Scotinotylus alpigena, Cryphoeca
silvicola preferred shaded habitats
)
SpiderdistributionintheSwissAlps 19
withthis,asmostindividualsof
this species were found there.
However, its status as a tim-
berline specialist must be
questioned, since in this study
italsooccurredinconsiderable
numbers in the open land and
inside the subalpine forest (40
Fig.2:HeadregionofthreeadultmalesofPelecopsisradicicola(L.Koch,1872)found and 33 individuals, respec-
onAlpFlix;frontalview. tively) (Tab. 2).
Our findings indicate that
and were found almost exclusively close to the the alpine timberline with its stand-alone trees
tree trunks in the timberline or inside the forest, provides habitats for both open land and forest
respectively. Caracladusavicula has been discussed spider species as well as some possible timberline
as a habitat-specialist ofthe timberline by FRICK specialists.We therebydemonstrate the particular
et al. (2007). Our results are partly in accordance value ofthe timberline and ofheterogeneous, spa-
Fig.3:TipoflamellacharacteristicaofMeionetaressliWunderlich,1973(a,Gastein,A)andMeioneta
alpica(Tanasevitch,2000)(b-i,AlpFlix,CH);dorsalview,leftmalepalp.
20 P.Muff,M.Schmidt,H.Frick,W.Nentwig&Chr.Kropf
dally limited structures in general in maintaining we show the distinction ofthe two species based
divergent invertebrate communities. For a more on the lamella characteristica (Fig. 3). Despite its
detailed description ofthe habitatzones and a full great variability in M. alpica (b-i), it can be well
analyticaldiscussionregardingtheresultsseeMUFF distinguishedfrom thelamellaofM. ressli(a)with
et al. (in prep.). respecttothesizeandshapeofthetwolobes.How-
ever,sinceM.alpicahasonlyrecentlybeendescribed
Someremarkablespecies and the separation by light microscope is rather
Our study revealed one species new to science difficult, it is possible that it has been confused
(Caracladus sp. A,Tab. 2) which will be described withits sisterspeciesM. ressliorM. rurestris(C.L.
elsewhere. Five species were new to the Canton Koch, 1836) in former studies. In the checklist of
Grisons: Centromeritabicolor, Centromeritaconcinna, theSwissspiders(BLICKetal.2004)Meionetaalpica
Hilairaexcisa, Meionetaalpicaand Tallusiaexperta. is notmentionedforSwitzerland,eventhoughthe
Otherremarkable species includedMeionetaorites holotypewasfoundinthecantonUri,Switzerland
and Panamomopspalmgreni, both endemic to the (Tanasevitch 2000).
Alps.The followingthree species showedremark-
able morphological characteristics and were ana- Metopobactrusprominulus(O.P.-Cambridge,1872)
lysed in more detail. =MetopobactrusschenkeliThaler, 1976
AccordingtoTHALER(1976)Metopobactrusschen-
Pelecopsisradicicola(L. Koch, 1872) keli differs from Metopobactrusprominulus by the
THALER(1978)consideredBrachycentrumdelesserti shape ofthe male head region, which is concave
Schenkel, 1925 a synonym of Pelecopsis radicicola and more elevated in profile in M. schenkeli and
since the characteristics of the first species were flat in M.prominulus. Male palps and females are
within the variation range ofthe latter. The only indistinguishable. Since our material comprises
differencewastheabsenceofthemedianlongitudi- individuals with intermediate characteristics (Fig.
naldepressiononthe elevated male headregionin 4),wearenotabletodistinguishthetwospecies,as
B. delesserti. OurmaterialfromAlp Flixcomprises itwas done in former studies (e.g.THALER 1978,
individuals showingpracticallyidentical male and HANSEN 1995, Frick et al. 2006). We therefore
female genital organs, but a great and gradual considerMetopobactrusschenkeli'WwXcx,1976asyn.
variability regarding size and shape ofthe raised nov.ofMetopobactrusprominulus(O.P.-Cambridge,
male head region (Fig. 2).This clearlyemphasises 1872).
the retention ofonly one species as suggested by
Thaler. Conclusion
Our study demonstrates that the known distribu-
Meionetaalpica(Tanasevitch,2000) tion andtaxonomic status ofvarious spidertaxain
In his initialdescriptionTANA-
SEVITCH(2000:211,subAgyne-
ta)characterisedMeionetaalpica
as being very closely related to
Meioneta ressli Wunderlich,
1973 but “well distinguishable
by the narrowed lamella char-
acteristica andlargerofitupper
lobes, almost equal to lower
one”. However, in practice this
differentiation has caused many
problems. Here,we checked 11
males of M. alpica from Alp
Flix,plus fourmales each ofM.
ressli from Ringkogel (Styria, Fig.4:Head regionofsixmalesofMetopobactrusprominulusfromAlpFlix;lateral
A) and Gastein (Salzburg, A), view.Notethegradualincreaseinconcavityfroma{"M.prominulus")tof("M.
respectively. Usingthis material schenkeli").
Spiderdistributionin theSwissAlps 21
theCentralAlpsarestillincomplete.Ourdatacon- Flix(Grisons):MethodenundResultate.-Jb.Naturf.
tributeone species newto science,five species new Ges. Graubünden 110:5-36
totheGrisonsfaunaandhighlightthreeinteresting HANSEN H. (1995): Über die Arachniden-Fauna von
taxonomicquestions.Thisisremarkablebecausethe urbanenLebensräumeninVenedig-III.Dieepigäi-
canton Grisons and Alp Flix in particular belong schenSpinneneinesStadtparkes(Arachnida:Arane-
itno ttheermbsestofstsupdiideedrsar(eTasHiAnLtEhRe S1w9i9s5s,CHenÄtNrGalGAIlp&sc Legaee).n-dBroell.P.Mu&s.LC.ivL.eSgtoe.nNdatr.eVe(n1e9z9i8a):44N:u7m-e3r6ical
MÜLLER2001,Fricketal.2006). Hence,further ecology. 2nd English Edition, Elsevier,Amsterdam.
workon arthropods in remote areas ofthe Alps is Mau85r3eprp.R.&A.HÄNGGI(1990):Katalogderschwei-
stronglyencouraged.
zerischen Spinnen.-Doc.Faun.Helvet. 12:412pp.
MuffP,C.Kropf,H.Frick,W.Nentwig8cM.H.
Acknowledgements
WekindlythankVictoriaSpinas(Sur,CH)forhercon- SCHMIDT (inprep.): Coexistence ofdivergentcom-
munities at natural boundaries: spider (Arachnida:
tinualenthusiasticsupportontheAlpandtheFoundati-
Araneae) diversityacrossthe alpine timberline.
o(Cnh“uSrc,haCtHz)in,seflorAlofpfeFrliixn”g,fpraeretiaccuclaormlmyoJdüartgiPoanu.lWMüellaerre Nentwig W., A. Hänggi, C. Kropf &T. Blick
mostgratefulto Cris Kuhlemeier,ThereseMandel and (2003): Central European Spiders. An internet
WernerGräber(allBern,CH),whoprovidedtheSEMs identification key. V. 08.12.2003. - Internet: http:
//www.araneae.unibe.ch
andperformedtheiroperation.Forvaluablecommentson
anearlierversionofthismanuscriptwethankTheoBlick PLATNICKN.I.(2007):Theworldspidercatalog,version
(Hummeltal,D),AmbrosHänggi(Basel,CH),Christian 8.0.AmericanMuseum ofNatural History.-Inter-
Komposch (Graz,A) andananonymousreviewer. net: http://research.amnh.org/entomology/spiders/
catalog/index.html
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alpinetimberline.-Ann. Zool.Fennici44: 43-57 THALERK.(1978):ÜberwenigbekannteZwergspinnen
HÄNGGIA.(1993):Nachträgezum„Katalogderschwei- aus den Alpen, V (Arachnida; Aranei, Erigonidae).
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Tab.2:Numberofindividualsofall speciesfoundfromMay2005toMay2006onAlpFlixaccordingtohabitatzone
(males/females)and intotal.Numbersarein boldwhereaspeciesoccurredwithatleast50%oftheindividuals(ifN
£ 15).AsterisksdenotespeciesnewtothecantonGrisons.
Species L o TO TB TT F Total
Amaurobiidae
Coelotesterrestris(Wider,1834) 1/1 2
Araneidae
AraneusdiadematusClerck,1757 0/1 1
Hypsosingaalbovittata(Westring,1851) 1/0 1
Clubionidae
ClubionadiversaO.P.-Cambridge,1862 0/1 1
ClubionareclusaO.P.-Cambridge,1863 0/1 1
Dictynidae
Dictynaarundinacea(Linnaeus,1758) 1/0 1
DictynapusillaThorell,1856 1/0 1
Mastigusaarietina(Thorell,1871) 0/1 0/1 0/1 3
Gnaphosidae
Drassodescupreus(Blackwall,1834) 8/4 5/1 1/0 1/0 20
Drassodespubescens(Thorell,1856) 8/3 6/2 5/0 1/0 2/0 27
Gnaphosaleporina(L.Koch,1866) 18/10 1/2 0/1 32
Haplodrassussignifer(C.L.Koch,1839) 15/11 16/9 9/3 5/3 4/4 79
MicariaaeneaThorell,1871 6/1 9/3 19/2 8/0 48
Micariapulicaria(Sundevall,1831) 0/1 1
Zelotessubterraneus(C.L.Koch,1833) 1/0 1
Zelotestalpinus(L.Koch,1872) 19/3 13/5 5/1 1/0 47
Hahniidae
Cryphoecasilvicola(C.L.Koch,1834) 2/0 29/4 69/30 134
Linyphiidae
Agnyphantesexpunctus(O.P.-Cambridge,1875) 0/6 2/14 5/25 52
Agynetacauta(O.P.-Cambridge,1902) 18/13 52/19 21/3 2/0 128
Agynetaconigera(O.P.-Cambridge,1863) 1/0 1/0 2
Anguliphantesmonticola(Kulczyriski,1881) 6/3 3/0 12
AsthenargusperforatusSchenkel,1929 1/0 1/1 1/1 5
Bolephthyphantesindex(Thorell,1856) 1/1 2/1 1/1 1/1 9
Bolyphantesalticeps(Sundevall,1833) 3/0 11/6 2/8 0/1 4/9 44
Bolyphantesluteolus(Blackwall,1833) 11/14 35/26 4/9 2/8 109
Caracladusavicula(L.Koch,1869) 18/22 4/18 17/58 9/17 17/10 190
A
Caracladussp. 0/1 0/1 6/5 13
Centromeritabicolor(Blackwall,1833)* 3/0 3
Centromeritaconcinna(Thorell,1875)* 1/0 1
Centromerusarcanus(O.P.-Cambridge,1873) 1/0 3/0 2/0 3/1 10
Centromeruspabulator(O.P.-Cambridge,1875) 8/1 87/9 74/7 28/4 72/29 319
Ceratinellabrevis(Wider,1834) 1/1 2/0 4
ErigoneatraBlackwall,1833 1/0 1
SpiderdistributionintheSwissAlps 23
Species O TO TB TT F Total
ErigonedentigeraO.P.-Cambridge,1874 1/0 1
Erigonedentipalpis(Wider,1834) 1/0 1/0 1/0 3
Erigonellasubelevata(L.Koch,1869) 2/1 1/1 0/1 6
EvansiamerensO.P.-Cambridge,1900 0/1 1
Gonatiumrubens(Blackwall,1833) 2/2 7/6 3/1 0/1 22
Hilairaexcisa(O.P.-Cambridge,1871)* 0/1 1
Improphantesnitidus(Thorell,1875) 0/2 4/0 53/14 43/20 136
Macrarguscarpenteri(O.P.-Cambridge) 25/4 5/2 3/0 3/0 42
Mansuphantespseudoarciger(Wunderlich,1985) 1/0 3/1 2/0 7/2 2/0 18
MarolehtineniSaaristo,1971 2/0 1/0 1/0 1/0 5
Meionetaalpica(Tanasevitch,2000)* 10/6 1/1 18
Meionetaorites(Thorell,1875) 5/2 3/2 1/0 13
Meionetarurestris(C.L.Koch,1836) 2/6 0/1 9
Metopobactrusprominulus(O.P.-Cambridge,1872) 2/0 2/1 5/1 2/2 15
&
MicrargusalpinusRelys Weiss,1997 0/1 1/0 0/1 3
Microctenonyxsubitaneus(O.P.-Cambridge,1875) 1/0 1
Microlinyphiapusilla(Sundevall,1830) 1/0 1
Miniciamarginella(Wider,1834) 1/1 6/5 1/1 1/0 16
Minyrioluspusillus(Wider,1834) 1/1 2
MughiphantescornutusSchenkel,1927 1/1 2/0 9/10 23
Mughiphantesmughi(Fickert,1875) 1/0 2/1 0/8 13/13 38
Obscuriphantesobscurus(Blackwall,1841) 0/2 2
PanamomopspalmgreniThaler,1973 1/0 1
Panamomopstauricornis(Simon,1881) 1/1 33/23 9/3 70
Pelecopsiselongata(Wider,1834) 7/11 17/10 2/7 54
Pelecopsisradicicola(L.Koch,1872) 2/6 5/11 20/9 16/9 4/2 84
Pityohyphantesphrygianus(C.L.Koch,1836) 1/0 1
Pocadicnemispumila(Blackwall,1841) 1/0 1
PorrhommacampbelliF.O.P.-Cambridge,1894 0/2 0/1 2/2 7
PorrhommapallidumJackson,1913 3/0 4/4 14/17 42
ScotarguspilosusSimon,1913 2/1 5/3 5/1 17
Scotinotylusalpigena(L.Koch,1869) 7/0 75/16 37/11 146
Scotinotylusclavatus(Schenkel,1927) 7/0 38/11 51/7 114
Stemonyphantesconspersus(L.Koch,1879) 0/1 1/0 5/2 9
Tallusiaexperta(O.P.-Cambridge,1871)* 0/1 1
TapinocybaaffinisLessert,1907 2/1 19/1 75/4 92/18 31/7 249
Tenuiphantescristatus(Menge,1866) 2/0 1/0 3
TenuiphantesmengeiKulczyriski,1887 11/27 25/54 9/17 18/10 5/4 180
Tenuiphantestenebricola(Wider,1834) 9/3 12
Thyreostheniusbiovatus(O.P.-Cambridge,1875) 0/3 3
Thyreostheniusparasiticus(Westring,1851) 0/4 4
Tisovagans(Blackwall,1834) 1/0 1
Walckenaeriaantica(Wider,1834) 2/1 3
Walckenaerialanguida(Simon,1914) 3/0 3
Walckenaeriamonoceros(Wider,1834) 1/0 0/1 2
Liocranidae
Agroecaproximo.(O.P.-Cambridge,1871) 3/1 2/0 0/1 7
| | | | ] [
24 PMuff,M.Schmidt,H.Frick,W.Nentwig&Chr.Kropf
Species O TO TB TT F Total
1 | [ | | [
Lycosidae
Alopecosaaccentuata(Latreille,1817) 13/1 8/1 1/1 1/0 26
Alopecosapulverulenta(Clerck,1757) 325/41 220/17 25/7 0/2 2/0 639
Alopecosataeniata(C.L.Koch,1835) 6/2 190/31 118/11 29/3 30/9 429
&
ArctosarenidescensBuchar Thaler,1995 12/3 28/9 7/1 1/2 1/0 64
Pardosablanda(C.L.Koch,1833) 44/22 39/24 10/9 1/1 150
Pardosaferruginea(L.Koch,1870) 1/0 1
Pardosamixta(Kulczynski,1887) 5/8 4/0 0/1 18
Pardosariparia(C.L.Koch,1833) 430/161 366/152 528/69 184/39 20/4 1953
Philodromidae
PhilodromusvagulusSimon,1875 0/1 1
Thanatusformicinus(Clerck,1757) 31/2 18/3 5/0 59
Salticidae
Evarchaarcuata(Clerck,1757) 0/1 1
Talaveramonticola(Kulczynski,1884) 4/1 2/0 7
Sparassidae
Micrommatavirescens(Clerck,1757) 1/1 1/0 3
Theridiidae
Robertaslividus(Blackwall,1836) 1/0 4/0 3/1 9
Robertustruncorum(L.Koch,1872) 5/4 27/6 30/8 17/8 105
Steatodaphalerata(Panzer, 1801) 1/0 2/0 3
Thomisidae
Ozyptilaatomaria(Panzer,1801) 8/1 4/1 3/0 1/0 18
Xysticusaudax(Schrank,1803) 2/1 1/1 4/0 3/0 12
XysticusbifasciatusC.L.Koch,1837 3/0 3
Xysticuscristatus(Clerck,1757) 2/0 2
XysticusgallicusSimon,1875 6/3 3/0 12
Xysticusluctuosus(Blackwall,1836) 6/0 15/0 19/0 40
XysticusmacedonicusSilhavy,1944 0/1 0/2 1/0 4
allfamilies 1112/394 1236/440 1074/258 718/258 515/394 6251