Eur J Etomol 105: 673 680, 2008 http://wwwejecz/scripts/viewabstractphp?abstract=1384 ISSN 1210-5759 (prit), 1802-8829 (olie) Orthoptera commuities i the coifer-broadleaved woodlad zoe of the Russia Far East THOMAS FARTMANN, MARTIN BEHRENS ad HOLGER LORITZ* Uiversity of Müster, Istitute of Ladscape Ecology, Departmet of Commuity Ecology, Robert-Koch-Str 26, D-48149 Müster, Germay; e-mail: fartma@ui-muesterde Key words Orthoptera, cricket, grasshopper, commuity ecology, disturbace, grasslad, woodlad zoe, Lazovsky Reserve, Russia Far East, habitat heterogeeity, habitat specifity, Palaearctic Abstract We ivestigate orthoptera commuities i the atural ladscape of the Russia Far East ad compare the habitat requiremets of the species with those of the same or closely related species foud i the largely agricultural ladscape of cetral Europe The study area is the 1,200 km 2 Lazovsky State Nature Reserve (Primorsky regio, souther Russia Far East) 200 km east of Vladivostok i the souther spurs of the Sikhote-Ali Moutais (134 E/43 N) The abudace of Orthoptera was recorded i August ad September 2001 based o the umber preset i 20 radomly placed 1 m² quadrates per site For each plot (i) the umber of species of Orthoptera, (ii) absolute species abudace ad (iii) fiftee evirometal parameters characterisig habitat structure ad microclimate were recorded Caoical correspodece aalysis (CCA) was used first to determie whether the Orthoptera occur i ecologically coheret groups, ad secod, to assess their associatio with habitat characteristics I additio, the umber of species ad idividuals i atural ad semi-atural habitats were compared usig a t test A total of 899 idividuals of 31 differet species were captured, with umbers ragig betwee 2 ad 13 species per plot Species diversity was higher i semi-atural habitats tha atural habitats There was a similar but o-sigificat patter i species desity Ordiatio aalysis idicated four orthoptera commuities, which were clearly separable alog a moisture ad vegetatio desity gradiet The atural sites i the woodlad area of the Lazovsky Zapovedik are characterized by species-poor ad low-desity orthoptera assemblages compared to the semi-atural sites But, the atural sites have a higher diversity of habitat specialists Our fidigs corroborate the hypothesis that itermediate habitat disturbace levels support particularly species-rich aimal commuities at high desities Uder such regimes, orthopteras presumably mostly profit from the high diversity i plat species, which geerates great structural ad microclimatic heterogeeity INTRODUCTION While atural forests i Europe were to a great extet trasformed by ma ito agricultural lad ad settlemet, huge areas of the East Palaearctic are still forested (Newell, 2004; Ya & Shugart, 2005) ad thus are importat referece areas for the study of temperate woodlad ladscapes The Far East is oe of the three biodiversity hotspots i Russia (Veevsky & Veevskaia, 2005) ad a cetre of diversity ad edemism of Orthoptera i Eurasia (Sergeev, 1998) Their taxoomy ad distributio are well studied, ad the ease with which they ca be sampled ad their fuctioal importace make Orthoptera suitable subjects for ecological ad biogeographical studies (Sergeev, 1997; Lockwood & Sergeev, 2000) Habitat selectio i Orthoptera is based o a complex combiatio of differet ad ofte iterrelated evirometal factors Of these parameters, the microclimate at ovipositio sites, which is ofte affected by vegetatio structure, plays a crucial role (Uvarov, 1977; Willott & Hassall, 1998) Sergeev (1997) stressed the suitability of orthoptera commuities for ecological ad biogeographical ivestigatios I recet decades may such studies have bee doe i the orther hemisphere Especially i North America, where differet aspects of ragelad grasshopper commuities have bee studied i detail (eg Kemp et al, 1990; Kemp, 1992a, b; Fieldig & Brusve, 1993a, b, 1995; Joer, 2004, 2005) Most commuity studies i the Palaearctic are for cetral Europe ad dry ad semi-dry grasslad habitats (eg Fartma, 1997; Behres & Fartma, 2004) Iformatio o the Asia part of the Palaearctic is restricted to biogeographic data (Stebaev et al, 1989; Sergeev, 1998) ad detailed studies of orthoptera assemblages are lackig Sice woodlads are usually ot cosidered to be a orthoptera habitat (Theuerkauf & Rouys, 2006) ad old forests are rare i cetral Europe, little is kow about habitat selectio ad commuity structure of Orthoptera i atural woodlad areas i the Palaearctic We therefore ivestigated orthoptera commuities i the atural ladscape of the Russia Far East ad compared the results with observatios from the huma ad agriculturally domiated ladscape of cetral Europe, because may taxa occur throughout the Palaearctic (Sergeev, 1992, 1997) Hece, orthoptera assemblages i the Lazovsky State Nature Reserve (Primorsky regio, Russia Far East) were studied to (i) determie their species compositio ad abudace i differet atural ad semiatural habitats, (ii) aalyse orthoptera habitat require- * Preset address: Helmholtz-Cetre for Evirometal Research Leipzig-Halle UFZ, Departmet of Commuity Ecology, Theodor-Lieser-Str 4, D-06120 Halle, Germay 673
At the coast witers are warmer (average Jauary temperature: 11 C) ad summers cooler (average August temperature: 17 C) (Semecheko, 2003) Fig 1 Study area ad locatio of Lazovsky Zapovedik mets i relatio to habitat structure ad microclimate ad (iii) compare orthoptera habitat prefereces there ad i Europe where the same or closely related species occur MATERIAL AND METHODS Study area The study area is located i the Primorsky regio (souther Russia Far East, Fig 1) The ladscape is formed by the Sikhote-Ali Moutais, stretchig from the southwest to the ortheast, parallel to the coastlie (average altitude about 1,000 m a s l) Woodlad covers 80% of the Primorsky regio taiga i the orth, coifer-broadleaved woodlads i the south (Newell, 2004) The 1,200 km 2 sized Lazovsky State Nature Reserve (ie Lazovsky Zapovedik, the official Russia category for this protected area) is situated about 200 km east of Vladivostok i the souther spurs of the Sikhote-Ali Moutais (134 E/43 N) It maily cosists of woodlads domiated by Mogolia oak (Quercus mogolica) with a admixture of Korea pie (Pius koraiesis) ad various other tree species The species richess of the Zapovedik is impressive (1212 species of vascular plats, 57 mammals ad 318 birds), with may rare ad highly edagered species icludig the Amur tiger (Pathera tigris altaica) (Chochrjakow & Schochri, 2002; Newell, 2004) Ope habitats are very rare, ad iclude atural coastal dues ad swamps, parts of the floodplais, screes ad moutai peaks, athropogeic meadows i woodlad-clearigs ear the three rager camps ad a few set-aside fields at the reserve border A mosoo climate with warm, humid summers ad cold, dry witers is characteristic of the study area The average aual precipitatio is 750 850 mm, decreasig from the coast ilad Due to a greater ifluece of cotietal climate the mea temperature ilad is 20 C i Jauary ad 20 C i July August Samplig of Orthoptera Samplig was carried out o 18 plots represetig all the typical orthoptera habitats of the Lazovsky Zapovedik, except floodplais, which were studied by Specht (2004) Nie atural (coastal dues, semi-dry coastal grasslads, swamps) ad ie semi-atural habitats (fallows domiated by Artemisia spp ad meadows) were ivestigated The area of the plots was > 2,000 10,000 m 2 with a homogeous vegetatio structure at every site Orthoptera desities were recorded i box quadrats (Gardier et al, 2005) of a total area of 20 m 2 From 29/08 15/09/2001 oe sample was take o each plot: The mobile 1 1 m (1 m 2 ) ad 80 cm high quadrat was radomly placed at twety differet poits Samplig was doe i sushie at temperatures > 20 C, betwee 10:00 am ad 5:00 pm Except for the small Nemobiiae species, which live hidde uder stoes or i litter o the groud, samplig provided reliable quatitative data Most of the specimes were determied i the field ad the released Idividuals that could ot be idetified i the field (Tetrix spp, some Chorthippus spp) ad voucher specimes of each species were collected ad idetified later For determiatio the keys of Bey-Bieko & Mishcheko (1951a, b) ad Storozheko (1986) were used Nomeclature is based o Storozheko (1986) ad for species that also occur i Europe o Heller et al (1998) Habitat structure For each plot we measured/estimated fiftee evirometal parameters: icliatio, exposure, heights of oe (miimum) up to three differet vegetatio layers (eg turf tall grass Artemisia) ad % cover of the followig habitat compoets: total vegetatio, field layers, Cyperaceae, Poaceae, herbs, mosses, litter, bare soil, stoes ad hollows (i swamps) Data aalysis Caoical correspodece aalysis (CCA) (usig CANOCO 451; ter Braak & Šmilauer, 2002), a direct gradiet ordiatio techique, was used to determie the orgaizatio of orthoptera species ito distict commuities ad the relatios betwee habitat structure ad species compositio (Fieldig & Brusve, 1993b, 1995; Palmer, 1993; Szövéyi, 2002; Torrusio et al, 2002) Evirometal ad species data were log arithmically trasformed [y = l (y 1)] to obtai approximately ormal distributios ad homogeous variaces Species of Orthoptera that occurred oly o oe plot ad/or of which < 10 specimes i total were foud were ot icluded i the data set (Table 1) Icliatio ad exposure were ot used as variables i the CCA because oly two plots were slightly iclied (< 5 ); cover (%) of bare soil ad stoes made up oe variable; out of the three field layer heights the maximum vegetatio height was used i CCA The statistical validity of the ordiatio was tested usig a Mote Carlo permutatio test (ull model: 9,999 urestricted permutatios) This was carried out for every evirometal variable ad all caoical axes (ie the complete model) Oly sigificat variables were icluded stepwise i the model, ad at each step oly the variable that explaied most of the remaiig error variace (maual-forward selectio of CANOCO) was chose No-sigificat variables were those that explaied little of the additioal variace at the time they could be added to the model They also may itercorrelate with other evirometal variables (Storch et al, 2003); which was examied usig Spearma s rak correlatio aalysis 674
TABLE 1 The 14 most commo orthoptera species o the 18 plots i order of their fidelity Species that occurred oly o oe plot ad/or with < 10 specimes i total are ot icluded (see Data aalysis) Distributio i Europe: species that also occur i Europe are idicated by a Exlusiveess: species that are restricted to atural () ad semi-atural (s) habitat types are idicated Species Phaeroptera falcata (Poda, 1761) Polioemobius taprobaesis (Walker, 1869) Tetrix japoica (Bolivar, I, 1897) Oecathus logicaudus Matsumura, 1904 Ruspolia itidula (Scopoli, 1786) Chorthippus maritimus Mishcheko, 1951 Chorthippus hammarstroemi (Miram, 1908) Chorthippus schmidti (Ikoikov, 1913) Omocestus haemorrhoidalis (Charpetier, 1825) Diaemobius fascipes igrofasciatus (Matsumura, 1904) Mecostethus parapleurus (Hagebach, 1822) Teleogryllus iferalis (Saussure, 1877) Oxya maritima Mishcheko, 1951 Pteroemobius itidus (Bolivar, I, 1901) Abbreviatio Phfal Potap Tejap Oelo Ruit Chmar Chham Chsch Omhae Difas Mepar Teif Oxmar Ptit Fidelity (o of occupied plots) 13 12 9 8 5 4 4 4 4 3 3 3 2 1 Sum of specimes 54 270 23 45 31 12 64 13 23 39 16 13 36 116 Desity (id/10 m²) 150 750 064 125 086 033 178 036 064 108 044 036 100 322 Distributio Europe Exclusiveess s s s s T tests (usig SPSS 115) were used to assess sigificat differeces i orthoptera desity ad species umber for the plots at the atural ad semi-atural sites Prior to the aalyses, variables were tested for ormal distributio usig Kolmogorov- Smirov test RESULTS Species richess ad abudace A total of 31 species (9 Tettigoiidae, 5 Gryllidae, 1 Tetrigidae ad 16 Acrididae) ad a sum of 899 specimes were captured Species umber raged from 2 to 13 per plot Phaeroptera falcata ad Polioemobius taprobaesis were the most widespread species occurrig i 13 (72%) ad 12 (67%) of the plots, respectively (Table 1) The total umber of species was higher at semi-atural (mea values ± SE: 911 ± 075) tha atural sites (511 ± 082) (t test, t = 3582, df = 16, P = 001) Similarly orthoptera desity [idividuals (id)/10 m², excludig Nemobiiae] was higher at semi-atural (mea values ± SE: 1744 ± 421) tha atural sites (889 ± 215) However, the differece was ot sigificat (t test, t = 1812, df = 16, P = 0089) I geeral there was a positive relatioship betwee species richess ad overall orthoptera desity (Y = 0735x 3628, R 2 = 027, P < 005) Of the semi-atural sites, youg fallows (N = 4) had the highest species umbers (975 ± 149) ad desities (2850 ± 552 id/10 m²) I cotrast to the total umber of species, that of abudat species restricted to oe of the habitat types was more or less the same (Table 1) Five species occurred exclusively i atural (swamps ad dues: Chorthippus maritimus, Omocestus haemorrhoidalis, Oxya maritima, Pteroemobius itidus ad Ruspolia itidulus) ad four i semi-atural habitats (meadows ad abadoed fields: Chorthippus hammarstroemi, Diaemobius fascipes igrofasciatus, Mecostethus parapleurus ad Oecathus logicaudus) Orthoptera assemblages Five of the te evirometal variables sigificatly cotributed to the CCA ordiatio model The model explaied 60% of the variace i the umber of species ad 96% of the variace i the species-eviromet relatio (total iertia: 308, sum of all caoical eigevalues: 193; Mote Carlo test: F = 404, P 0001) Each of the four caoical axes was highly correlated with oe particular evirometal variable, facilitatig the ecological iterpretatio of each axis (see Table 2 for details) All o-sigificat variables itercorrelated with other evirometal variables (Table 3) ad explaied < 10% additioal variace at the stage whe they would have bee icluded i the model Based o the first two caoical axes four distict commuites ca be separated (Fig 2a): Cover of hollows strogly correlated with the first caoical axis Note that there is o real evirometal moisture gradiet uderlyig this ordiatio patter, because there are oly two plots (swamps) with hollows Alog the first caoical axis species are divided ito two distict mai groups: Taxa that were recorded oly i swamps (atural habitat) ad those oly i mesic to dry habitats (semi-atural ad atural habitats) The swamps have a species-poor orthoptera commuity However, two species, Oxya maritima ad Pteroemobius itidus, were restricted to the wetlads Based o the arragemet of species alog the secod caoical axis it is possible to further discrimiate betwee of the orthoptera commuities i the remaiig habitats The axis is egatively correlated with herb cover The strogest positive correlatios (i order of arragemet) with herb cover are for Mecostethus parapleurus, Chorthippus hammarstroemi, Oecathus logicaudus ad Diaemobius fasciatus They form a secod 675
TABLE 2 Summary of CCA for 14 orthoptera species (N = 755 specimes) ad five sigificat evirometal variables (F-values of Mote Carlo test, ***P < 0001, *P < 005) Evirometal Axis Eigevalue Species-eviromet correlatios Variace explaied (%): Species data Species-eviromet relatio Liear correlatio with cover (%) of: Hollows Herbs Poaceae Field layers Cyperaceae 1 2 3 4 F 100 038 028 020 100 085 088 076 325 124 518 197 100 021 071 046 004 023 042 000 020 89 64 140 105 002 026 015 071 025 048 083 054 006 042 001769*** 210* 215* 219* 215* commuity foud i herb-rich fallows ad meadows (semi-atural habitat) Clearly separated from all other species ad strogly egatively correlated with herb cover is Chorthippus maritimus O the most extreme due sites (atural habitat) with sparse vegetatio ad short swards Chorthippus maritimus is typical of the third species-poor commuity Sporadically a few other species occur at low desity Also, but to a lower extet Omocestus haemorrhoidalis ad Ruspolia itidula are egatively correlated with herb cover Both species occur sytopically i coastal semi-dry grasslads (atural habitat) whe they form part of aother orthoptera commuity Further isights i the relatios betwee species of Orthoptera ad vegetatio structure are idicated by the third ad fourth axes (Fig 2b) The folllowig species are positively correlated with the cover of field layers ad Poaceae: Ruspolia itidula, Phaeroptera falcata ad Polioemobius taprobaesis R itidula was most abudat i Poaceae-rich ad dese coastal grasslads P falcata ad P taprobaesis were foud i early all habitats, except the coastal dues with extesive areas of bare groud ad swamps Fig 2 Biplots for the axes of caoical correspodece aalysis (CCA): a first vs secod axis; b third vs fourth axis Sigificat evirometal/groud cover variables (arrows) ad positio of orthoptera species (crosses) Abbreviatio of species ames (see Table 1) Two variables (hollows, herbs), which do ot correlate with axis three ad four (see Table 2), i e their vectors were close to zero, are ot show i b A group of four species (Chorthippus hammarstroemi, Teleogryllus iferalis, Diaemobius fascipes ad Chorthippus maritimus), which are further from the origi of TABLE 3 Correlatio matrix of Spearma s rak correlatio aalyses (N = 18; ***P < 0001, **P < 001, *P < 005) Evirometal variables Groud coverage Vegetatio total Field layers Herbs Poaceae Cyperaceae Moss Litter Bare soil ad stoes Hollows Vegetatio height (cm) 031 056* 048* 023 023 041 023 014 014 Hollows 039 011 020 001 039 051* 012 045 Bare soil/ stoes 087*** 071** 027 033 053 023 049* Litter 044 067** 037 037 003 036 Groud cover (%) Moss 018 037 033 007 040 Cyperaceae 039 013 020 027 Poaceae 037 053* 004 Herbs 024 049* Field layers 083*** 676
the axis, is strogly egatively correlated with cover of field layers ad Poaceae C hammarstroemi, D fascipes ad T iferalis reach their highest desities i fallows with bare soil The first two are restricted to these sites DISCUSSION Species richess ad abudace Accordig to Stebaev et al (1989) may of the observed orthoptera species are associated with the souther border of the forest zoe ad limited to the Pacific Ocea districts of the Russia Far East with a mosoo climate Numerous species are cocetrated i the study area, i regios of coifer-broadleaved forest ad forest-steppe i the south of the Russia Far East (Stebaev et al, 1989; Sergeev, 1992) The atural sites i the woodlad area of the Lazovsky Zapovedik are characterized by species-poor ad lowdesity orthoptera assemblages compared to the semiatural sites At the semi-atural sites the highest desities ad species umbers were foud o youg fallows The youg set-aside fields studied were characterized by the highest structural heterogeeity What are the reasos for a high orthoptera diversity at the semi-atural sites? Several studies (Fartma & Mattes, 1997; Kruess & Tschartke, 2002; Gebeyehu & Samways, 2003) idicate that orthoptera species richess is highest at sites subject to itermediate disturbace (eg grazig) ad high structural heterogeeity Joer (2005) foud that grasshopper species richess i grasslad is positively correlated with plat species richess ad heterogeeity of vegetatio structure, ad egatively with vegetatio height ad grass biomass Sites subject to a itermediate level of disturbace (like youg fallows) have more plat species (Grime 1973a, b) ad therefore a greater structural heterogeeity Thus, greater resources are available for the coexistece of more orthoptera species (Deis et al, 1998) I our view ovipositio sites ad food resources are the most importat, especially the presece of bare groud for ovipositio ca be a limitig factor May, or eve the majority of orthoptera species lay their eggs i the groud ad prefer bare or sparsely covered groud for ovipositio (Richards & Waloff, 1954) I this study the umber of orthoptera species exclusive to atural ad semi-atural sites is similar (5 vs 4 species) Icludig the orthoptera data of Specht (2004) for the floodplais of the Lazovsky Zapovedik ad thus all the typical atural orthoptera habitats i the study area, five further species are restricted to atural habitats [Bryodemella tuberculata (Fabricius, 1775), Diaemobius csikii (Bolivar, I, 1901), Eireephilus logipeis (Shiraki, 1910), Oedaleus iferalis Saussure, 1884 ad Tetrix teuicoris (Sahlberg, 1893)] Orthoptera assemblages of the atural sites of the Lazovsky Zapovedik are species-poor, but have a greater diversity of highly specialised species I accordace with the results of Joer (2005) there is a positive relatioship betwee overall Orthoptera desity ad species richess, idicatig that there are at least some parameters that promote both species diversity ad desity High orthoptera desities are ofte the result of a trade-off betwee optimal microclimatic coditios o the oe had ad sufficiet food ad low predatio pressure o the other (Fieldig & Brusve, 1992; Gottschalk, 1996; Fartma & Mattes, 1997; Behres & Fartma, 2004) Of crucial importace for Orthoptera abudace are the microclimatic coditios durig egg ad larval developmet (Igrisch, 1979, 1980) I this period most species beefit from high temperatures (va Wigerde et al, 1991a) I the sparse vegetatio of the coastal dues microclimate may be favourable, but food shortage ad easy access for avia predators seem to result i a low orthoptera desity A icrease i vegetatio desity results i lower temperatures i the egg habitat, mostly located ear the soil surface (va Wigerde et al, 1991a) Although, the dese stads of the old set-aside fields should provide eough food ad eemy-free space, the microclimate seems to be ufavourable for may orthoptera species Because the youg ope Artemisia fallows i the Lazovsky Zapovedik provide the best combiatio of requiremets, the orthoptera abudace there is the highest Orthoptera assemblages Presece ad assemblage of orthoptera species are distictly differet amog habitat types This depeds o orthoptera habitat preferece, which is determied by species adaptatio to habitat structure, microclimate ad disturbace itesity (Joer, 1982; Fieldig & Brusve, 1995; Samways, 1997; Szövéyi, 2002) The distributio patter of eurytopic ad steotopic species differ due to their specific habitat requiremets with the latter species characteristic of differet habitat types i the Lazovsky Zapovedik Where possible, the followig compares the preset results (Fig 2, Table 1) with ecological observatios from Europe The eurytopic ad thermophilous species Phaeroptera falcata ad Polioemobius taprobaesis are ot restricted to oe orthoptera commuity ad occur i both atural ad semi-atural habitats i the study area However, their habitat requiremets differ The terricolous, flightless cricket P tabrobaesis is foud i high desities i short-turf vegetatio It is a thermophilous species distributed throughout the Ido-Malaya regio ad the souther Far East (Schmidt 1999) The phytophilous/arbusticolous ad very mobile (capable of flyig) bush-cricket P falcata occurs o those habitats at low desities It prefers tall herbaceous vegetatio ad has a Traspalaearctic distributio (Detzel, 1998a) The followig groups of species have a higher habitat specifity ad form four distict orthoptera commuities: 1 Commuity of coastal dues with bare groud The geophilous Chorthippus maritimus, a far easter siblig species of C biguttulus (Liaeus, 1758) (Krivolutskaya, 1997), occurs i the study area oly i atural ad dyamic habitats with extesive areas of bare groud or stoes, such as coastal dues, floodplais or screes (ow observatio) I the floodplais of the Lazovsky Zapovedik it typically co-occurs with Bryodemella 677
tuberculata ad Eireephilus logipeis (Specht, 2004) C biguttulus, a Eurosiberia ad commo species i cetral Europe, shows differet habitat requiremets: it is phytophilous ad eurytopic with a preferece for semi-dry grasslad (Fartma, 1997; Behres & Fartma, 2004) 2 Commuity of semi-dry coastal grasslad Ruspolia itidula, a Palaeotropic/Mediterraea species is thermophilous, phytophilous/gramiicolous ad especially the immature stages are hygrophilous (Detzel, 1998b) The restrictio of this bush-cricket to coastal grasslad with a moderate maritime climate i the Lazovsky Zapovedik seems to result from these adaptatios R itidula prefers vertical structures ad i the study area it is most abudat (up to 7 idividuals/10 m 2 ) i ecotoes of dese, tall-grass stads, which possibly offer a suitable microclimate ad structure The steotopic Omocestus haemorrhoidalis, a Traspalaearctic species, has similar vegetatio structure prefereces i the Lazovsky Zapovedik ad i cetral Europe It is geo-/phytophilous, gramiicolous, xero- ad thermophilous Microhabitat prefereces i the Russia Far East correspod to those observed by Fartma (1997) i ortheast Germay: ope, short-turf semi-dry grasslad with structural heterogeeity ad bare soil 3 Commuity of herb-rich meadows ad fallows Mecostethus parapleurus ad Oecathus logicaudus are herbicolous ad thermophilous species of the herbrich fallows ad meadows i the study area M parapleurus is distributed alog the souther boudary of the emoral woodlad zoe from the Atlatic Ocea to Japa; i cetral Europe it is a thermophilous species, which prefers wet meadows (Detzel, 1998c) The habitat requiremets of O logicaudus seem to resemble those of the Mediterraea O pelluces (Scopoli, 1763) i cetral Europe: Accordig to Detzel (1998d) O pelluces is a typical species of tall forb-rich plat commuities Chorthippus hammarstroemi is a easter Palaearctic, meso-xerophilous steppe species (Sergeev, 1997) that is abudat i athropogeic habitats, such as fields (Sergeev, 1998) I the Lazovsky Zapovedik the cricket Diaemobius fascipes igrofasciatus occurs sytopically with C hammarstroemi exclusively i recetly set-aside fields, but reaches its highest desity i ope, dry fallows with much bare soil/stoes Hece, this terricolous cricket is cosidered to be thermo- ad xerophilous O the Kuril Islads it coloises soils heated by volcaic activity, close to solfataras ad fumaroles (Krivolutskaya, 1997) It is widely distributed i the temperate regio of East Asia; o the Japaese Islads the souther rage limit is about 30 N (Masaki, 1996) 4 Commuity of swamps The hygrophilous ad phytophilous/gramiicolous Oxya maritima (Catatopiae) occurs together with the terricolous cricket Pteroemobius itidus i the coastal swamps Masaki & Oyama (1963) describe P itidus as a typical species of paddy fields i North Japa The habitat requiremets seem to resemble those of the siblig species P heydeii (Fischer, 1853) with a Holomediterraea origi (Kiechle, 1998) ad a Mediterraea- -southwest Asia distributio (Baur et al, 2006) P heydeii is characterised as hygrophilous, terricolous ad thermophilous; typical habitats i cetral Europe are south-facig lakeshores, swamps or wet meadows (Kiechle, 1998; Witerholler & Bierwirth, 2003) CONCLUSIONS The majority of Orthoptera i the boreal ad temperate forest zoes of the Palaearctic are associated with ope habitats (Sergeev, 1992, 1997) These grasslad habitats ofte are created ad maitaied by disturbace (Theuerkauf & Rouys, 2006), which is especially importat at the microhabitat scale At this level, disturbace creates the microclimatic coditios ad microhabitats ecessary for the survival of eggs ad larval stages This applies to Orthoptera ad other thermophilous isects (eg Lepidoptera: Fartma, 2004, 2006) As for the whole temperate zoe of the Palaearctic (Birks, 2005; Mitchell, 2005), closed forests are the atural vegetatio of most parts of the Lazovsky Zapovedik I the absece of ma, ope habitats are restricted to sites with extreme evirometal coditios Oly sites that are highly disturbed (dues, floodplais), too dry (dues, screes) or wet (swamps) ad too cold (moutai tops) are free of trees, all the itermediate sites are covered with woodlad Due to the origi of the ladscape oe would expect that the hot spots of orthoptera diversity ad desity will be sites created by atural disturbace But, this is ot the case as semi-atural sites subject to athropogeic disturbace (recetly set-aside fields) have the highest species umbers ad abudace The orthoptera commuities of the atural habitats are ot as species-rich as the athropogeic sites, but there are more steotopic species restricted to those habitats Comparable observatios exist for cetral Europe; but disturbace itesity as well as the spatial ad temporal patter i the cultural Europea ladscape are very differet from a atural ladscape Huma activity i cetral Europe creates either extesive ad cotiuously high disturbace (eg itesive croplad) or approximately stable coditios with low disturbace frequecy (eg high forests with log cuttig itervals), which both lead to habitat loss for isects of ope habitats Natural, dyamic disturbace patters ad low-itesity athropogeic maagemet have positive effects o habitats ad populatios of steotopic isects (Sergeev, 1998; Di Giulio et al, 2001), but today such practices are rare i cetral Europe (Fartma, 2006) Hece, our fidigs i the atural ladscape of the Russia Far East cofirm a strategy of ature coservatio that has recetly attracted iterest i cetral Europe That is, the coservatio ad restoratio of atural ladscape dyamic/disturbace (Fick et al, 1998) ad support of athropo-zoogeic maagemet with low itesity (va Wigerde et al, 1991b; Wallis De Vries & Raemakers, 2001; Theuerkauf & Rouys, 2006) for a lastig coservatio of isect biodiversity 678
ACKNOWLEDGEMENTS We are grateful to N Athes (Tübige, Germay), N Hölzel (Müster, Germay), G Köhler (Jea, Germay), M Kovi ka ( eské Bud jovice, Czech Republic) ad two aoymous referees for very helpful commets o a earlier draft of the text Thaks to I Edich (Müster, Germay) who traslated Russia literature ad L Harris (Müster, Germay) for improvig our Eglish REFERENCES BAUR B, BAUR H, ROESTI C & ROESTI D 2006: Die Heuschrecke der Schweiz Haupt, Ber, Stuttgart, Wie, 351 pp BEHRENS M & FARTMANN T 2004: Die Heuschreckegemeischafte isolierter Schieferkuppe der Medebacher Bucht (Südwestfale/Nordhesse) Tuexeia 24: 303 327 BEY-BIENKO GY & MISHCHENKO LL 1951a: Locusts ad grasshoppers of the USSR ad adjacet coutries Vol I Keys to the Faua of the USSR 38: 1 400 [traslated from Russia by Israel Program for Scietific Traslatios, Jerusalem 1964] BEY-BIENKO GY & MISHCHENKO LL 1951b: Locusts ad grasshoppers of the USSR ad adjacet coutries Vol II Keys to the Faua of the USSR 40: 1 291 [traslated from Russia by Israel Program for Scietific Traslatios, Jerusalem 1964] BIRKS HJB 2005: Mid the gap: how ope were Europea primeval forests? 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