Bull.Br.arachnol.Soc.(2001)12(1),1–9 1 Spider (Arachnida: Araneae) communities of (Framenauetal.,1996a).Themobilebroodcareofwolf riparian gravel banks in the northern parts of the spidersappearstobefavouredinanenvironmentwhich European Alps is characterised by frequent inundation (Framenau, 1995; Churchill, 1998). Randolf Manderbach Today man-made weirs, dams, and embankments control most floodplains in Central Europe (Baxter, DepartmentofAnimalEcology, FacultyofBiology, 1977; Petts et al., 1989). These constructions have Philipps-UniversityMarburg, changedthenaturalriversystemdynamicsandaffect,in 35032Marburg,Germany particular, the occurrence of vegetation-sparse or -free gravel banks, which are now mainly restricted to the and upper catchment of large alpine rivers. Only remnant gravel banks can be found in the lower floodplains Volker W. Framenau (Fabrice & Michel, 1992; Plachter, 1993). The number DepartmentofZoology, and size of gravel banks in small streams seems to be UniversityofMelbourne, similarly reduced (Hering et al., 1997). Parkville3052,Victoria,Australia Whileanumberofstudieshaveinvestigatedthespider communities of floodplains, they have not included Summary riparian gravel banks (e.g. Uetz, 1976; Moring & Acomprehensiveinvestigationofthegravelbankspider Stewart, 1994; Bell et al., 1999). Only one study com- faunaof10riversandstreamsinthenorthernpartsofthe prehensively characterises a whole floodplain spider European Alps provided an account of 45 species in 13 community (Steinberger, 1996). This study provides a families. The most common species were Pardosa wagleri, detailed account of the habitat requirements of most ErigoneatraandOedothoraxagrestis.Incombinationwith six previously published studies on spiders of flood plain spiderspeciesencounteredattheLechRiverinAustria, habitats, a set of 60 species, that are ‘‘common or domi- but was limited to one river. Most studies that nant’’onripariangravelbanks,wasidentifiedandgrouped have specifically examined gravel banks, have mainly in four categories: riparian, open land, forest, and euryo- been limited to single rivers (e.g. Plachter, 1986; ecious. Pardosa wagleri, Pardosa saturatior, Pirata knorri, Dröschmeister, 1994; Hering, 1995; Smit, 1997; Heidt andOedothoraxagrestiswereclassifiedasstenoeciousripar- ian species. Altitude, floodplain width and level of human et al., 1998; Steinberger, 1998; Zulka et al., 1998). Only impacthadnoinfluenceonthenumberofspeciesrecorded two autecological studies have dealt with spider species ineachriversection.Thestenoeciousriparianspeciesdonot thatarecharacteristicofripariangravelbanks(Albert& appeartobesuitableindicatorsforanaturalriversystem, Albert, 1976; Framenau et al., 1996a,b). sincethereisnoevidencethattheirnumbersareaffectedby Our study provides a comprehensive account of the the artificial reduction in number and size of floodplain gravelbanks. spider fauna of Central European gravel bank flood plains. Samples taken at ten different rivers have been incorporated with previously published studies to deter- Introduction mineasetofthemost‘‘commonordominant’’riparian InCentralEurope,alpineriversarecharacterisedbya spider species. These are classified according to habitat highly variable discharge rate, with peak floods occur- requirements. Collections from river sections with and ring mainly in spring. The sediment load in the upper withouthumanimpactaimedtodemonstratetheeffects catchment of these rivers consists mainly of large rocks of hydrological engineering on the composition of and gravel. Constrictions in the river floodplain cause the spider communities, and the suitability of riparian thisgraveltoaccumulate,therebyforminglargebraided spiders as indicators of a natural river system is gravel beds. During a peak flood, this gravel is moved discussed. and vegetation destroyed (e.g. Aulitzky, 1980; Karl & Mangelsdorf, 1975; Mangelsdorf & Scheuermann, Methods 1980). Floods therefore pose a threat to the survival of many typical organisms that inhabit this environment. The ecological characterisation of the riparian gravel However, these organisms are also dependent on floods bank spider fauna is based on a survey of 10 alpine tomaintainspecifichabitatqualities,suchasasubstrate rivers and streams and a review of six major previous made up of loose sand and gravel, and no or only a studies including one conducted outside the alpine sparse vegetation cover. region. A specifically adapted arthropod community inhabits Fourteen floodplain sections belonging to 10 rivers this highly dynamic environment. Among the most and streams in three main river systems in the northern dominant members of the riparian gravel bank fauna parts of the European Alps were investigated during are spiders, and in particular wolf spiders (Lycosidae) three surveys in May/June, July/August and October (Manderbach & Reich, 1995; Smit et al., 1997; 1995(Fig.1,Table1).Thesesectionsrepresentedawide Steinberger, 1996, 1998; Manderbach, 1998). They are varietyfromlargeriverstosmallstreamswhichdiffered thought to have evolved a number of characteristics not only in the amount and size of gravel bars but also in response to this dynamic river environment, such inthelevelofhumanimpactondischargeandsediment as the use of polarised light for orientation (Papi, load. The sample programme included the largest 1959; Papi & Tongiorgi, 1963) and seasonal movements naturalbraidedriversectioninthenorthernpartsofthe 2 Spidersofripariangravelbanks Species which were found in more than 10% of all riversectionsorwhichrepresentedmorethan5%ofthe total catch in one section were called ‘‘common or dominant’’species.Onlythesespecieswereconsideredin describingthetypicalgravelbankspidercommunitiesof alpine rivers. These species were grouped in one of four categories: riparian, open land, forest and euryoecious. The categories ‘‘riparian’’ and ‘‘open land’’ comprise three sub-categories each. The categorisation followed standard publications on the ecology of Central European spider species (Maurer & Hänggi, 1990; Heimer & Nentwig, 1991; Hänggi et al., 1995). Results A total of 1,698 spiders were collected in 14 river sections, including 474 (28%) adult spiders belonging to 45 species in 13 families (Table 2). Overall, they com- prised 15% of all arthropods encountered (Fig. 2). Five families(Amaurobiidae, Anyphaenidae, Philodromidae, SalticidaeandTetragnathidae)wererepresentedonlyby juvenilespiders.Mostspecies(22)belongedtothefamily Linyphiidae. More than half of all species (26; 58%) were found in only one river section. The wolf spider Pardosawagleriwasthemostcommonspecies,foundin 11 of 14 sections. Erigone atra and Oedothorax agrestis Fig. 1: Sampling sites. Hornbach (1), Streimbach (2), Lech near were represented in 9 river sections. The spider com- Forchach(3),Stillach(4),Ostrach(5),IsarnearSchröfeln(6), munities were dominated by only a few species. Five Halblech (7), Iller near Rubi (8), Isar near Vorderriss (9), species (Erigone atra, E. dentipalpis, Oedothorax Schwarzenbach(10),IllernearSonthofen(11),Arzbach(12), agrestis, Pardosa wagleri and Pirata knorri) represented IsarnearArzbach(13),LechnearLangweid(14). more than 75% of the total catch of all adult spiders. We found no evidence that altitude (ANOVA, European Alps (Lech River near Forchach: Fig. 1[3]), F =0.93, p>0.5), floodplain width (ANOVA, 11,2 large rivers considerably affected by weirs and embank- F =0.5, p>0.5), or the level of human impact 3,10 ments (e.g. Lech River near Langweid [14]) and small (ANOVA, F =0.898, p>0.1) had an influence on the 2,11 streams in a more or less natural condition (e.g. number of species found in any of the river sections. Schwarzenbach [10]). Each river was assigned to one The largest number of adult spiders per sample was of three categories depending on the level of human foundattheSchwarzenbach,thelowestattheIllernear impact: natural condition, minor and major human Sonthofen (Table 3). Floodplain width and level of impact (Table 1). humanimpactshowednoinfluenceonthetotalnumbers Thecompositionofthespidercommunitieswasinves- of adult spiders encountered, nor on the numbers of tigated by standardised time samples (Herold, 1928, any of the typical gravel bank species P. wagleri, P. 1929; Andersen, 1969; Plachter, 1986). On vegetation- saturatior, P. knorri and O. agrestis. Altitude appeared freegravelbanksinastripabout0.5–1mwidealongthe to affect only the numbers of adult P. saturatior water edge, the fine substrate was combed with fingers (ANOVA, F =4.904, p<0.05), a species that is 1,12 and larger stones were turned over. All arthropods, restricted to higher elevations. except Apterygota, were caught: smaller ones with an Incombinationwiththepreviouslypublishedstudies, aspirator, larger ones with forceps. The specimens were 60 ‘‘common or dominant’’ species were identified and transferred into 70% ethyl alcohol. Each floodplain wereconsideredinthecharacterisationoftheEuropean section was sampled for a total of 120min per survey. spider communities of riparian gravel banks (Table 4). Thissamplingtimewasdividedinto6periodsof20min onconsecutivegravelbanks.Thelengthsofthesampled Discussion areas varied between 5 and 25m depending on the substrate and the number of animals encountered Spider communities of riparian gravel banks (details in Manderbach, 1998). Sixpreviousstudiesonfloodplainspidercommunities Riparian species in which data were obtained by ‘‘hand-collecting’’ (time Thespidercommunityontheshorelineofriversdoes or area limited) were included in the analysis (Table 4). not appear to be very specific. Only 10 (17%) of the Within these studies, data not obtained on vegetation- ‘‘common or dominant’’ species have their main distri- free gravel banks or by a different method (e.g. pitfall bution on the banks of rivers and streams and seem to trapping) were not taken into account in our study. be almost exclusively ‘‘riparian’’. However, seven of R.Manderbach&V.W.Framenau 3 these ten species can occasionally be found in habitats that share similar qualities with riparian gravel banks, such as rocky outcrops and gravel pits. These may not be located in river floodplains and therefore lack the proximity of running water. Therefore, the riparian species can be divided into three groups depending on thedegreeoftheiraffinitytorunningwater:stenoecious riparianspecies(withastrongaffinitytorunningwater), species of gravel bars (with limited affinity to running water) and species of lake shores. Only Pardosa saturatior, Pardosa wagleri, Pirata knorri and Oedothorax agrestis can be classified as stenoecious riparian species. They seem to be restricted to the shores of running waters, where they can reach very high densities. Pardosa wagleri is the commonest speciesonripariangravelbanksandwasfoundineleven ofthe14investigatedriversections.Aboveanaltitudeof approximately 900m it is accompanied by its slightly larger sibling species Pardosa saturatior (Barthel & Fig.2: Compositionoftheriparianarthropodcommunitiesingravel Helversen, 1990). Both species coexist on gravel banks bedfloodplainsoftheEuropeanAlps(n=11,645). between 980m and 1000m. Oedothorax agrestis has not only been recorded on gravel banks. Single indi- viduals have been found in other floodplain habitats, species of this group prefer rocky areas without veg- e.g.alderwoodland(Nentwig,1983)andonmuddysoil etation cover. These habitat qualities are maintained by (Palmgren, 1976). Steinberger (1996) found this species the dynamic water regime in gravel bed floodplains but mainly near the edge of a river, but a few individuals can also be found in other circumstances. Janetschekia were also collected in muddy areas at some distance monodon has been collected in gravel habitats near the fromthewater.However,itsaffinitytowaterappearsto edgeofglaciers(Thaler,1969).Clubionasimilisisgener- be strong. ally known from ‘‘moist areas with low vegetation’’ Five species (Janetschekia monodon, Clubiona similis, (Heimer & Nentwig, 1991). Steinberger (1996) encoun- Arctosacinerea,CaviphantessaxetorumandDiplocentria teredC.similisonxerothermicgravelbankswithsparse mediocris) can regularly be found on riparian gravel vegetation. Caviphantes saxetorum seems to migrate banksbuttheydisplayonlyalimitedaffinitytorunning seasonally between areas with large stones near the water.Theyhavealsorepeatedlybeenrecordedinareas water in September and dry areas covered by small without adjacent streams or rivers. It appears that the stones in June (Cooke & Merrett, 1967). However, this No.in River Altitude Description Floodplain Human Fig.1 (location) (m)a.s.l. width impact 1 Hornbach 1000 Largebraidedsectionabovegorge(dam) >100m 1 2 Streimbach 980 Mainlyonlyonechannel;highwatervelocity 10–50m 0 3 Lech 910 LargestbraidedriversectioninnorthernEuropeanAlps;largegravelbanks;large >100m 0(e) (nearForchach) areaswithsparsevegetationbetweengravelbanksandforest 4 Stillach 900 Largebraidedsectionabovegorge >100m 0(e) 5 Ostrach 860 Singlestreambed;highinclination 10–50m 0 6 Isar 810 Largebraidedfloodplainsection;largegravelbanks;vegetationcoverincreasing 10–50m 1(w) (nearSchröfeln) overrecentyears;belowKrünWeir 7 Halblech 770 Undividedstream;smallgravelbanks;coarsesubstrateandsludge;algalgrowth 10–50m 2(ee) 8 Iller 770 LargestgravelbanksalongIllerRiver;bedrockoneastbank;adjacentforest 51–100m 2(e) (nearRubi) 9 Isar 770 Largest,mostnaturalbraidedfloodplainsectioninGermanAlps;largegravel >100m 0(w) (nearVorderriss) banks;belowKrünWeir 10 Schwarzenbach 760 SmalltributarytoIsarRiverbelowSylvensteinReservoir;smallgravelbanks; <10m 0 completelyshadedbyriparianforest 11 Iller 720 Small,isolatedgravelbanks;coarsesubstrateandsludge,relictsofriparianforest 10–50m 2(ee) (nearSonthofen) 12 Arzbach 700 SmalltributarytoIsarRiver;smallgravelbanks;partlyshadedbyriparianforest <10m 0 13 Isar 660 Large,isolatedgravelbanks;coarsesubstratewithsludge;relictsofriparian 10–50m 2(ee,r) (nearArzbach) forest;algalgrowth;belowSylvensteinReservoir(regulationofdischarge) 14 Lech 440 OldriverbedofLechRiver;riverbedonlypartlyfilledwithslowrunningwater; 51–100m 2(ee) (nearLangweid) largegravelbanks;coarsesubstrateandsludge;algalgrowth;mainwaterflow redirectedabovethissection Table 1: Characterisation of sampled river sections. Human impact: 0=natural condition, 1=minor human impact, 2=major human impact, e=embankment(ee=onbothsides),w=weirupstream,r=reservoirupstream. 4 Spidersofripariangravelbanks Arzbach Halblech Hornbach IllernearRubi IllernearSonthofen IsarnearArzbach IsarnearSchröfeln IsarnearVorderriss LechnearForchach LechnearLangweid Ostrach Schwarzenbach Stillach Streimbach Nesticidae Nesticuscellulanus(Clerck,1757) — — — — — — — — — — — 1— — Theridiidae Robertusneglectus(O.P.-Cambr.,1871) — — — — — — — — — — — — — 1 TheridionvariansHahn,1833 — — — — — 1— — — — — — — — Linyphiidae Araeoncushumilis(Blackwall,1841) — — — — — — — — — 1— — 1— Bathyphantesgracilis(Blackwall,1841) — 1 — — — — — — — 1— — — — Bathyphantesnigrinus(Westring,1851) — — — 1— — — — — — — — — — Caviphantessaxetorum(Hull,1916) — 1 — — — — 1— — — — — — — Centromerusincilium(L.Koch,1881) — — — — — — — 1— — — — — — Centromerussylvaticus(Blackwall,1841) — — — — — 1— — — — — — — — Ceratinopsisromana(O.P.-Cambr.,1872) — — — — — — — 1— — — — — — Diplocephaluscristatus(Blackwall,1833) — 1 2 1 —1 — — — — — — — — Diplocephaluslatifons(O.P.-Cambr.,1863) — — — 2— — — — — — — — — — ErigoneatraBlackwall,1833 — 2 4 2 1 2 —2 1 7 1 — — — Erigonedentipalpis(Wider,1834) — — 3 2 —2 1 —2 3 — — — — Gnathonariumdentatum(Wider,1834) — — — — — — — — — — — 1— — Janetschekiamonodon(O.P.-Cambr.,1872) — — 1— — — 1— 4— 1 1 2 1 Lepthyphantespulcher(Kulczyn´ski,1881) — — — — — — — — — — — — — 1 Meionetarurestris(C.L.Koch,1836) — — — — — — — 1— 1— — — — Oedothoraxagrestis(Blackwall,1853) 14 37 26 4 — 3 2 — — —12 24 18 — Oedothoraxapicatus(Blackwall,1850) — — — — — — — — — 1— — — — Oedothoraxretusus(Westring,1851) — 3 — 1 — — 1 2 — — 1— 1 1 Porrhommaconvexum(Westring,1851) — — 2 1 — — — — — — — 2— — Prinerigonevagans(Audouin,1826) — — — — — — — — — 1— — — — Tisovagans(Blackwall,1834) — — — — — — — — — — — — — 1 Araneidae AraneusdiadematusClerck,1757 — — — — — — — 1— — — — — — Araniellaopisthographa(Kulczyn´ski,1905) — — — 1— — — — — — — — — — Lycosidae Arctosacinerea(Fabricius,1777) — — — — — — 4 3 1 — — — — — Arctosamaculata(Hahn,1822) — — — — — — — — — — — 2— — Pardosaamentata(Clerck,1757) — 1 — — — — — — — — — — 5— Pardosariparia(C.L.Koch,1833) — — — — — — — 1— — — — — — PardosasaturatiorSimon,1937 — — 2— — — — — — — — — 5 3 Pardosawagleri(Hahn,1822) 27 — 1 4 5 1 2 16 11 — 36 8 — 2 Pirataknorri(Scopoli,1763) 2 4 — 1 — — 2— — — 36 2 4 2 Trochosaruricola(DeGeer,1778) — — — — — — 1— — — — — — — Agelenidae Histoponatorpida(C.L.Koch,1834) — — — — — — — 1— — — — — — Cybaeidae Cybaeustetricus(C.L.Koch,1839) — — 1— — — — — — — — 2— — Hahniidae Cryphoecasilvicola(C.LKoch,1834) — — 1— — — — — — — — — — — Amaurobiidae Coelotesterrestris(Wider,1834) — — — 1— — — — — — — — — — Liocranidae Agraecinastriata(Kulczyn´ski,1882) 1 — — — — 1— — — — 1 1 —1 Clubionidae ClubionaneglectaO.P.-Cambr.,1862 — — — — — — — 1— — — — — — ClubionasimilisL.Koch,1867 — — 2— — 1— — 1— — 3— — ClubionatrivialisC.L.Koch,1843 — — — — — — 1— — — — — — — Gnaphosidae Zeloteslatreillei(Simon,1878) — — — — — — — — — — — — — 1 Zelotessubterraneus(C.L.Koch,1833) — — — — — — — — — — — 1— — Thomisidae Diaeadorsata(Fabricius,1777) — — — — — — — 1— — — — — — Table2: Numbersofadultspiderscollectedonthe14investigatedriversectionsinthenorthernpartsoftheEuropeanAlps.Nomenclaturefollows Platnick(1998). R.Manderbach&V.W.Framenau 5 speciescanalsobefoundindry,urbanhabitats(Maurer (cf. Ru˚zˇicˇka, 1978). A further six species (10%) do not & Hänggi, 1990). Arctosa cinerea has been recorded prefer any specific substratum, moisture or vegetation from disused open brown coal mines (Hussein, 1998). cover but can be found in all types of open areas Other non-riparian records for this species include (indifferent open land species). gravel pits and sandy and pebble beaches of the North and Baltic Seas (Framenau, 1995). Forest-dwelling species The wolf spider Pirata piraticus is predominantly found in river and stream sections with reduced water Eight species (13%) occur mainly in forests and simi- velocity. Its typical habitats are the shores of lakes with larly shaded habitats. Forest-dwelling species occur amoreorlessdensecoverofvegetation(Renner,1986). regularly on banks of small streams where the adjacent woodland reaches the banks and facilitates the invasion of these spiders. Species of open areas Thirty-three (55%) of the species listed as ‘‘common Euryoecious species or dominant’’ on river shores are generally known from openareaswithalowcoverofherbaceousvegetation.In Six species (10%) do not show any particular habitat the floodplains of larger rivers, these habitats are fre- preference and can therefore be characterised as gener- quently found on the lower terraces which are only alists. Erigone atra and E. dentipalpis are among these occasionally submerged (e.g. Moor, 1958; Müller & species and occasionally attain high numbers on river Bürger, 1990). Here, the soil moisture can vary consid- banks. erably within a short distance depending on the water carrying capacity and the drainage quality of the soil. Thesuitabilityofriparianspidersforindicatingchangesin The ‘‘common or dominant’’ open land species were floodplain dynamics categorised by their preference for different degrees of humidity: xerophilous, mesophilic (hydrophilous) and In the European Alps, gravel bank spider communi- indifferent species. tiesappeartobedominatedbyafewriparianspecialists. Six species (10%) prefer open, very dry habitats Except for Pirata piraticus, all the species that we (xerophilousspecies).Theseareasarefoundparticularly classified as stenoecious riparian are restricted to in the lower floodplains of gravel bed rivers and are vegetation-free gravel banks. The reduction in number often characterised by gravelly and stony soil surfaces and size of gravel banks due to human interference with rather sparse vegetation cover. The category of might therefore be expected to result in a decline in mesophilic (hydrophilous) species comprises the largest the number of riparian species and their densities. numberofspecies(21;35%).Speciesinthisgroupprefer Thus, riparian spiders have the potential to be used as intermediate to moist areas, which are typically covered indicators by comparing their diversity and density in bydensegrass(PoaceaeandCyperaceae).Spidersofthe disturbed and undisturbed river systems. genus Oedothorax are particularly common, with Oedo- Arthropods, and in particular spiders, have com- thoraxretususbeingdominantonnumerousriverbanks. monly been suggested as ecological indicators (Clausen, Plachter(1986)recordedOedothoraxfuscusinunusually 1986; Helsdingen, 1991; Kremen et al., 1993; Platen, largenumbers;amisidentificationofthefemalesappears 1993; Malt, 1995; Neet, 1996; Churchill, 1998). The to be likely, since the sibling species O. agrestis is development of indicator systems requires the clear commoner throughout all the other considered studies definitionof(a)whatenvironmentalimpactorchangeis River Oedothorax Pardosa Pirata Pardosa All agrestis wagleri knorri saturatior species Hornbach 1.44&4.31 0.06&0.24 0.11&0.32 2.5&4.72 Streimbach 0.11&0.47 0.11&0.32 0.17&0.38 0.78&1 Lech(nearForchach) 0.61&0.98 1.11&1.18 Stillach 1&2.64 0.22&0.55 0.28&0.67 2&4.33 Ostrach 0.67&1.41 2&3.27 0.17&0.51 3.06&4.53 Isar(nearSchröfeln) 0.11&0.32 0.11&0.32 0.11&0.47 0.89&1.41 Halblech 2.06&5.41 0.22&0.43 2.78&5.94 Iller(nearRubi) 0.22&0.55 0.22&0.43 0.06&0.24 1.17&1.34 Isar(nearVorderriss) 0.89&2.11 1.72&3.27 Schwarzenbach 1.33&2 0.44&1.04 3.44&6.48 6&7.91 Iller(nearSonthofen) 0&0 0.28&0.96 0.33&0.97 Arzbach 0.78&1.67 1.5&3.67 0.11&0.32 2.44&4.99 Isar(nearArzbach) 0.25&0.62 0.08&0.29 1.08&2.28 Lech(nearLangweid) 2.5&1.05 Table3: Mean(&s.e.)numberofadultspiders(stenoeciousriparianspeciesandallencounteredspecies) per20minsample(n=18samplesperriversection,exceptIsarnearArzbach,n=12,andLech nearLangweid,n=6).Numbersinboldtyperepresentthethreehighestvaluesforeachspecies. 6 Spidersofripariangravelbanks High-montane Montane Altitude(m)a.s.l. 1000 980 930 910 900 980 970 820–1010 890 860 860 850 840 820 820 810 780 770 770 770 770 760 Riversection(Reference) Hornbach(a) LechVorderhornb./Häselg.(b) LechForchach—Stanzach(b) LechnearForchach(a) Stillach(a) Streimbach(a) Elmau(c) Linder(c) UpperHalblech(d) LechHöfen—Weissenbach(b) Ostrach(a) Neidernach1(c) LechPflach—Lechaschau(b) LechMusau—Pinswang(b) Neidernach2(c) IsarnearSchröfeln(a) IsarnearVorderrissII(e) IsarnearVorderrissI(a) IsarnearVorderrissIII(c) IllernearRubi(a) Halblech(a) Schwarzenbach(a) Riparianspecies PardosasaturatiorSimon,1937 4 8 — —14 21 10 8 — — — — — — — — — — — — — — (a) Pardosawagleri(Hahn,1822) 2 1 5 55 — 14 — — 23 6 65 8 1 6 6 13 28 52 19 19 — 7 Oedothoraxagrestis(Blackwall,1853) 58 1 — —50 — 5 16 12 1 22 8 7 + — 13 — — 4 19 74 22 Pirataknorri(Scopoli,1763) — 16 1 — 11 14 57 68 22 3 5 56 4 + 18 13 16 — 48 5 8 57 Janetschekiamonodon(O.P.-Cambr.,1872) 2 1 5 20 6 7 — — 3 3 2 —1 6 — 6 — — — — — 1 ClubionasimilisL.Koch,1867 4 3 3 5 — — — — 3 4 — — 2 9 — — — — — — — 3 (b) Arctosacinerea(Fabricius,1777) — — + 5 — — — — — 1—— — — — 25 11 10 11 — — — Caviphantessaxetorum(Hull,1916) — — 2 — — — — — 1 6 — — 1 2 — 6 — — — — 2 — Diplocentriamediocris(Simon,1884) — — 5 — — — — — — 1— — — 1— — — — — — — — (c) Piratapiraticus(Clerck,1757) — — — — — — — — — — — — 1 + — — — — — — — — Speciesofopenareas PardosatorrentumSimon,1876 — 2 11 — — — — — 3 4 — — + 7 — — — — — — — — Phrurolithusfestivus(C.L.Koch,1835) — + — — — — — — + 1 — — — — — — — — — — — — xerophilous Drassylluspumilus(C.L.Koch,1839) — + 1 — — — — — 1 — — — — 1— — — — — — — — HeliophanuspatagiatusThorell,1875 — — + — — — — — 3 — — — — 5— — — — — — — — PhrurolithusminimusC.L.Koch,1839 — — — — — — — — — — — — — — — — — — — — — — Myrmarachneformicaria(DeGeer,1778) — — — — — — — — — — — — — — — — — — — — — — Oedothoraxretusus(Westring,1851) — 24 15 — 3 7 — — 3 26 2 —3 21 0 6 6 16 6 7 5 6 — Oedothoraxfuscus(Blackwall,1834) — — — — — — — — — — — — — — 6—15 — — — — — Pardosaamentata(Clerck,1757) — 1 1 — 14 — — — 3 2 — — 10 2 — — — — — — 2 — Agraecinastriata(Kulczyn´ski,1882) — 1 — — — 7— — 1 1 2 — + 1 — — — — — — — 1 Oedothoraxapicatus(Blackwall,1850) — — — — — — — — — — — — — — — — 3— — — — — Porrhommaconvexum(Westring,1851) 4 1 + — — — — — 3 — — — 1 1 — — — — — 5— 2 Bathyphantesnigrinus(Westring,1851) — — — — — — — — + — — — — 0— — — — — 5— — Bathyphantesgracilis(Blackwall,1841) — — — — — — — — — — — — 1 1 — — — — — — 2 — Trochosaruricola(DeGeer,1778) — — + — — — — — — — — — — 1— 6— — 4— — — mesophilic Collinsiadistincta(Simon,1884) — — — — — — — — — — — — 1 + — — 1— — — — — (hydrophilous) PachygnathaclerckiSundevall,1823 — — — — — — — — — — — — 1 — — — — — — — — — TrochosaterricolaThorell,1856 — — — — — — — — 1 — — 8 0 — — — — — — — — — Oedothoraxgibbosus(Blackwall,1841) — — — — — — — — — — — — 2 + — — — — — — — — Piratahygrophilus(Thorell,1872) — — — — — — — — + — — — — — — — — — — — — — PiratalatitansBlackwall,1833 — — — — — — — — — — — — + — — — — — — — — — Antisteaelegans(Blackwall,1841) — 1 — — — — — — + — — — — — — — — — — — — — Porrhommapygmaeum(Blackwall,1834) — 1 — — — — — — — — — — — — — — — — — — — — Pelecopsismengei(Simon,1884) — — — — — — — — — 1— — 5 1 — — — — — — — — Tisovagans(Blackwall,1834) — — + — — 7— — — — — — — — — — — — — — — — Hypommabituberculatum(Wider,1834) — — — — — — — — — — — — 2 — — — — — — — — — Prinerigonevagans(Audouin,1826) — — — — — — — — — — — — — — — — — — — — — — Diplocephaluscristatus(Blackwall,1833) 4 8 1 — — — 5— 1 1 — — 4 3 — — — — — 5 2 — Meionetarurestris(C.L.Koch,1836) — — 0 — — — — 2 2 1 — — — 2 6 —1 3 4 — — — indifferent Araeoncushumilis(Blackwall,1841) — + 1 — 3 — — — + 1 — — — — — — — — — — — — Micariapulicaria(Sundevall,1832) — — — — — — — — — — — — — — — — — — — — — — TheridionvariansHahn,1833 — + — — — — — — — — — — — — — — — — — — — — Zeloteslatreillei(Simon,1878) — — — — — 7— — — — — — — — — — — — — — — — Euryoeciousspecies Erigoneatra(Blackwall,1841) 9 3 15 5 — —10 — 1 5 2 4 3 5 35 — — 6—10 4 — Erigonedentipalpis(Wider,1834) 7 3 9 10 — — — — 1 3 — 4 4 9 12 6 — — 4 10— — Zelotessubterraneus(C.L.Koch,1833) — + + — — — — 2 1 — — — — + — — — — — — — 1 Araniellacucurbitina(Clerck,1757) — + 1 — — — — — + — — — + + — — — — — — — — Pardosalugubris(Walckenaer,1802) — 2 + — — — — — 1 — — — — — — — — — — — — — ClubionatrivialisC.L.Koch,1843 — — — — — — — — — 1—— — — — 6 — — — — — — Forest-dwellingspecies Diplocephaluslatifrons(O.P.-Cambr.,1863) — + + — — — — — — 1— — — — — — — — — 10 — — Centromerussylvaticus(Blackwall,1841) — — — — — — — — — — — — — — — — — — — — — — Lepthyphantestenebricola(Wider,1834) — — — — — — — — — — — — — + — — — — — — — — Coelotesterrestris(Wider,1834) — — — — — — — — — — — — — — — — — — — 5— — Robertusneglectus(O.P.-Cambr.,1871) — — + — — 7— — — 1— — — — — — — — — — — — Diaeadorsata(Fabricius,1777) — + — — — — — — — —— 8 — — — — — 3— — — — Diplostylaconcolor(Wider,1834) — — — — — — — — 0 — — — — — — — — — — — — — PhilodromuscollinusC.L.Koch,1835 — — + — — — — — — — — — — — 6— — — — — — — ? Leptorhoptrumrobustum(Westring,1851) — — — — — — — — — — — — + — — — — — — — — — Lessertinellakulczynskii(Lessert,1909) — 1 2 — — — — — — 8— — + 7 — — — — — — — — Lepthyphantespulcher(Kulczyn´ski,1881) — — — — — 7—— — — — — — — 6 — — — — — — — %otherspecies: 4 20 21 0 0 0 14 4 14 19 0 4 16 22 0 0 8 19 0 5 0 6 totalnumberofindividuals: 45 208 242 20 36 14 21 50 503 144 55 25 338 315 17 16 74 31 27 21 50 108 Table4: Dominance(%)ofthe‘‘commonordominant’’spiderspeciesofripariangravelbanksinCentralEurope.NomenclaturefollowsPlatnick (1998).‘‘+’’=0–0.5.Riparianspecies:a=specieswithstrongaffinitytorunningwater,b=specieswithlimitedaffinitytorunningwater, c=speciesoflakeshores.References:(a)thisstudy,(b)Steinberger(1996),(c)Hering(1995),(d)Dröschmeister(1994),(e)Plachter(1986), (f)Smit(1997),(g)Heidtetal.(1998). R.Manderbach&V.W.Framenau 7 Montane Submontane Colline 720 700 660 650 588 540 290–464 430 410 257–520 320–485 300–455 212–440 440 475 518 367/380 359 320 440 200 160 238–255 253–300 172–295 245 (e) IllernearSonthofen(a) Arzbach(a) IsarnearArzbach(a) IsarsouthBadTölz(e) IsdarsouthPuppling(e) IsarnorthGrünwalderBrücke LinspherBach(f) IsarsouthMarzling(e) IsarnearVolkmannsdorf(e) Orke(f) Perf(f) ElbrighäuserBach(f) Treisbach(f) LechnearLangweidII(e) Isarnordwestl.Ismaning(e) IsarMünchen(e) IsarnearNiederaichbach(e) IsarnearMamming(e) DonaunearPondorf(e) LechnearLangweidI(a) Ain(g) Tagliamento(g) Lahn(f) Eder(f) Allna(f) Rhône(g) Pardosasaturatior — — — — — — — — — — — — — — — — — — — — — — — — — — Pardosawagleri 83 61 8 43 30 — — — — — — — — — — — — — — — 9 26 —— — — Oedothoraxagrestis — 32 23 — — — 69 — — 63 13 79 80 2 — — — — — — 14 — 23 1 79 — Pirataknorri — 5 — 14 13 7 14 2 8 15 25 5 5 28 — — — — — — — — 9 18 — — Janestschekiamonodon — — — — — — — — — — — — — — — — — — — — — 16 — — — — Clubionasimilis — — 8— — — — — — — — — — — — — — — — — 6 3— — — — Arctosacinerea — — — 4 5—— — — — — — — — — — — — — — 1 3— — — — Caviphantessaxetorum — — — — — — — — — — — — — — — — — — — — — — — — — — Diplocentriamediocris — — — — — — — — — — — — — — — — — — — — — 2 —— — — Piratapiraticus — — — — — — — — — — — — — — — — — — — — 1— 1 1— — Pardosatorrentum — — — — — — — — — — — — — — — — — — — — — — — — — — Phrurolithusfestivus — — — — — — — — — — — — — — — — 2 2 — — 1— — — — 7 Drassylluspumilus — — — — — — — — — — — — — — — — — — — — 1 2— — — — Heliophanuspatagiatus — — — — — — — — — — — — — — — — — — — — 1 2— — — — Phrurolithusminimus — — — — — — — — — — — — — — — — — — — — 15 — — — — — Myrmarachneformicaria — — — — — — — — — — — — — — — — — 11 — — — — — — — — Oedothoraxretusus — — — 41 8 61 5 33 6 6 — 3 — 1 — 40 9 — 43 — 2 3 22 49 2 — Oedothoraxfuscus — — — 82 1 17 + 37 44 + — + + 27 47 12 35 17 — — 1— 1 1— — Pardosaamentata — — — 4— — 2 — — 1 8 + —2 2 5 5— — 5— 1— 11 2 — — Agraecinastriata — 2 8 6 —— — — — — — — — — — — 6— — — 6 7— — — 7 Oedothoraxapicatus — — — — — — + 1 — + 6 — —1 — — —15 — 7 12 — 74 3 7 Porrhommaconvexum — — — — — — + — — — — + — — — — — — — — — — — — — 7 Bathyphantesnigrinus — — — — — 1 — —22 + — + — 8 — — 3—19 — — — — — — — Bathyphantesgracilis — — — — — — 1— — — 2 + 2 — — — — — — 7 — — 1— 1 — Trochosaruricola — — — — — — — — — + — — — — — — — 7— — + 2 1 — 1 — Collinsiadistincta — — — — — — — — — — — — — — 28 5 — — 5— 1— 4— — — Pachygnathaclercki — — — — — — + 2 — + 2 — — — — — — — 5— — — 1 3— — Trochosaterricola — — — — — — + — — — 2— — — — — — — — — + — 1— — — Oedothoraxgibbosus — — — — — — 1— — — 2 + — — — — — — — — — — — — 1 — Piratahygrophilus — — — — — — + — — + 2 2 — — — — — — — — — — — — 2 — Piratalatitans — — — — — — — — — + — — — — — — 2— — — — 18 1 — — — Antisteaelegans — — — — — — + — — — — — + — — — — — — — 1— — — — — Porrhommapygmaeum — — — — — — + — — + — + — — — — — — — — + — — — — — Pelecopsismengei — — — — — — — — — — — — — — — — — — — — — — — — — Tisovagans — — — — — — — — — — — + — — — — — — — — — — — — — — Hypommabituberculatum — — — — — — — — — — — — — — — — — — 19 — — — — — — — Prinerigonevagans — — — — — — — — — — — — — — — — — — — 7 6 — — — — — Diplocephaluscristatus — — 8 8 5 8—5 1 3 + 2 + — 9—10 18 4 — — 6— — — — 7 Meionetarurestris — — — — — — + — — + — — — — — — — 4— 7 2 —2 2— — Araeoncushumilis — — — — — — — — — — — — + — — — — — — 7 1 3 1 —1 7 Micariapulicaria — — — — — — — — — — — — — — — — 8— — — — — 1— — — Theridionvarians — — 8— — — — — — + — — — — — — — — — — — — — — — — Zeloteslatreillei — — — — — — — — — — — — — — — — — — — — — — — — — — Erigoneatra 17 — 15 1 5 2 1 9 4 2 21 1 1 6 — 10 11 20 5 47 2 — 8 14 9 13 Erigonedentipalpis — —15 — 2 3 1 2 1 2 4 0 1 5 — 19 — 15 — 20 5 3 3 2 2 33 Zelotessubterraneus — — — — — — — — — — — — — — — — — — — — — — 1— — — Araniellacucurbitina — — — — — — — — — — — — — — — — — — — — — — — — — — Pardosalugubris — — — — — — — 2— — — 0— — — — — — — — — — — — — — Clubionatrivialis — — — — — — — — — — — — — — — — — — — — — — — — — — Diplocephaluslatifrons — — — — — — — — — + 2 0 4 — — — — — — — — — — — — — Centromerussylvaticus — — 8— — — + — — + 4 1 +1 — — — — — — — — — — — — Lepthyphantestenebricola — — — — — — + — — + — + + — — — 2— — — — — — — — — Coelotesterrestris — — — — — 1 + — — — — 1 1 — — — — — — — — — — — — — Robertusneglectus — — — — — — — — — — — — — — — — — — — — — — — — — — Diaeadorsata — — — — — — — — — — — — — — — — — — — — — — — — — — Diplostylaconcolor — — — — — — — — — — — — — — — — — — — — — — — — — 7 Philodromuscollinus — — — — — — — — — — — — — — — — — — — — — — — — — — Leptorhoptrumrobustum — — — — — — + — — 2 2 1 + — — — — — — — — — — — — — Lessertinellakulczynskii — — — — — — — — — — — — — — — — — — — — — — — — — — Lepthyphantespulcher — — — — — — — — — — — — — — — — — — — — — — — — — — 0 0 0 7 0 0 7 6 2 8 2 8 6 8 0 0 6 4 0 0 7 10 22 2 0 6 44 13 72 56 103 976 86 85 614 48 1142 342 96 36 42 66 46 21 15 362 61 176 97 126 15 Table4:Continued. 8 Spidersofripariangravelbanks to be indicated, (b) what effect this change has on the ANDERSEN, J. 1969: Habitat choice and life history of Bembidiini indicator organism (e.g. morphological change, reduc- (Col., Carabidae) on river banks in central and northern Norway.Norskent.Tidskr.17:440–453. tion in numbers), and (c) how this change can be AULITZKY,H.1980:Preliminarytwo-foldclassificationoftorrents. accurately measured (New, 1995). The environmental Interpraevent4:285–309. impact of hydrological engineering can be easily estab- BARTHEL,J.&HELVERSEN,O.von1990:Pardosawagleri(Hahn lished by comparing historic data of discharge patterns 1822) and Pardosa saturatior Simon 1937, a pair of sibling and river morphology with current information. How- species (Araneae, Lycosidae). Bull. Soc. europ. Arachnol. vol. ever, the consequence of these changes on floodplain horssérie1:17–23. BAXTER,R.M.1977:Environmentaleffectsofdamsandimpound- spider populations appears to be rather unclear and a ments.A.Rev.Ecol.Syst.8:255–283. methodology to determine differences is difficult to BELL,D.,PETTS,G.E.&SADLER,J.P.1999:Thedistributionof implement. spidersinthewoodedriparianzoneofthreeriversinwestern The restriction of Pardosa saturatior to rivers and Europe.Regul.Rivers:Res.&Mgmt15:141–158. streams above 800m makes it exempt from human CHURCHILL, T. 1998: Spiders as ecological indicators in the interference. Pardosa wagleri is predominantly found Australian tropics. Family distribution patterns along rainfall and grazing gradients. In P. Selden (ed.), Proceedings of the aboveanaltitudeof500m,alsosuggestingageographi- 17th European Colloquium of Arachnology, Edinburgh 1997: cally restricted distribution. Its presence below 200m at 325–330.BritishArachnologicalSociety,BurnhamBeeches. the Ain and Tagliamento Rivers (Heidt et al., 1998) CLAUSEN,I.H.S.1986:Theuseofspiders(Araneae)asecological suggeststhathumanimpactmayhaveamoreimportant indicators.Bull.Br.arachnol.Soc.7:83–86. influence on its distribution than ecological preferences, COOKE, J. A. L. & MERRETT, P. 1967: The rediscovery of Lessertiella saxetorum in Britain (Araneae: Linyphiidae). since the impact of dams, weirs and embankments is J.Zool.,Lond.151:323–328. more prominent in the lower catchment. However, DRÖSCHMEISTER, R. 1994: Die Spinnenfauna der Kies- und P. wagleri does not seem to be affected by changes in Schotterbänke des nordalpinen Wildbaches Halblech the floodplain dynamics. It is abundant in floodplains (LandkreisOstallgäu).Ber.naturw.Ver.Schwaben98:61–70. with both minor (Lech near Forchach [910m], Isar FABRICE, M. & MICHEL, D. 1992: Die letzten naturnahen near Vorderriss [770m]) and extensive human impacts Alpenflüsse. Kl. Schr. Intern. Alpenschutzkonv. CIPRA 11: 1–69. (Iller near Sonthofen [720m], Isar south of Bad FRAMENAU,V.1995:PopulationsökologieundAusbreitungsdynamik Tölz [650m]).Therefore, its use as an indicator species von Arctosa cinerea (Araneae, Lycosidae) in einer alpinen appears to be limited. Oedothorax agrestis and Pirata Wildflusslandschaft.M.Sc.thesis,Philipps-UniversityMarburg. knorri are common not only on gravel banks in FRAMENAU, V., DIETERICH, M., REICH, M. & PLACHTER, the European Alps but also by small streams in low H. 1996a: Life cycle, habitat selection and home ranges of Arctosa cinerea (Fabricius, 1777) (Araneae: Lycosidae) in a mountainous areasbothwithandwithouthumaninter- braided section of the Upper Isar (Germany, Bavaria). Revue ference (Smit, 1997). There was no significant corre- suisseZool.vol.horssérie1:223–234. lation between the intensity of human impact and FRAMENAU, V., REICH, M. & PLACHTER, H. 1996b: Zum abundance of these two species in our study. Wanderverhalten und zur Nahrungsökologie von Arctosa It is difficult to establish a standardised sampling cinerea(Fabricius,1777)(Araneae:Lycosidae)ineineralpinen Wildflusslandschaft.Verh.Ges.Ökol.26:369–376. methodtoquantifypotentialdifferencesinspiderdiver- HÄNGGI, A., STÖCKLI, E. & NENTWIG, W. 1995: Habitats of sity and abundance between rivers with and without centralEuropeanspiders.Miscneafaun.helv.4:1–459. hydrological constructions. Pitfall traps near the river HEIDT, E., FRAMENAU, V., HERING, D. & MANDERBACH, bankareverylikelytobeflooded,andhandsamplesare R. 1998: Die Spinnen- und Laufkäferfauna auf ufernahen subjecttovariationdependingontheindividualinvesti- SchotterbänkenvonRhône,Ain(Frankreich)undTagliamento gator(Plachter,1986;Dröschmeister,1995).Differences (Italien)(Arachnida:Araneae;Coleoptera:Carabidae).Ent.Z., Frankf.a.M.108:142–153. in the abundance of gravel bank spiders might also be HEIMER, S. & NENTWIG, W. 1991: Spinnen Mitteleuropas: Ein a consequence of differences in habitat structure, e.g. Bestimmungsbuch.Berlin,PaulParey. size composition of gravel. Our knowledge of habitat HELSDINGEN, P. J. van 1991: Can Macrothele calpeiana preferences for any of the riparian species is still (Walckenaer) (Araneae, Hexathelidae) be used as a bio- inadequatetointerpretanyvariationinspiderdensities. indicator?Bull.Soc.neuchâtel.Sci.nat.116:253–258. 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