Eur. J. Entomol. 105: 673–680, 2008 http://www.eje.cz/scripts/viewabstract.php?abstract=1384 ISSN 1210-5759 (print), 1802-8829 (online) Orthopteran communities in the conifer-broadleaved woodland zone of the Russian Far East THOMAS FARTMANN, MARTIN BEHRENS and HOLGER LORITZ* University of Münster, Institute of Landscape Ecology, Department of Community Ecology, Robert-Koch-Str. 26, D-48149 Münster, Germany; e-mail: [email protected] Key words. Orthoptera, cricket, grasshopper, community ecology, disturbance, grassland, woodland zone, Lazovsky Reserve, Russian Far East, habitat heterogeneity, habitat specifity, Palaearctic Abstract. We investigate orthopteran communities in the natural landscape of the Russian Far East and compare the habitat require- ments of the species with those of the same or closely related species found in the largely agricultural landscape of central Europe. The study area is the 1,200 km2 Lazovsky State Nature Reserve (Primorsky region, southern Russian Far East) 200 km east of Vladi- vostok in the southern spurs of the Sikhote-Alin Mountains (134°E/43°N). The abundance of Orthoptera was recorded in August and September 2001 based on the number present in 20 randomly placed 1 m² quadrates per site. For each plot (i) the number of species of Orthoptera, (ii) absolute species abundance and (iii) fifteen environmental parameters characterising habitat structure and micro- climate were recorded. Canonical correspondence analysis (CCA) was used first to determine whether the Orthoptera occur in ecol- ogically coherent groups, and second, to assess their association with habitat characteristics. In addition, the number of species and individuals in natural and semi-natural habitats were compared using a t test. A total of 899 individuals of 31 different species were captured, with numbers ranging between 2 and 13 species per plot. Species diversity was higher in semi-natural habitats than natural habitats. There was a similar but non-significant pattern in species density. Ordination analysis indicated four orthopteran communi- ties, which were clearly separable along a moisture and vegetation density gradient. The natural sites in the woodland area of the Lazovsky Zapovednik are characterized by species-poor and low-density orthopteran assemblages compared to the semi-natural sites. But, the natural sites have a higher diversity of habitat specialists. Our findings corroborate the hypothesis that intermediate habitat disturbance levels support particularly species-rich animal communities at high densities. Under such regimes, orthopterans presumably mostly profit from the high diversity in plant species, which generates great structural and microclimatic heterogeneity. INTRODUCTION America, where different aspects of rangeland grass- While natural forests in Europe were to a great extent hopper communities have been studied in detail (e.g. transformed by man into agricultural land and settlement, Kemp et al., 1990; Kemp, 1992a, b; Fielding & Brusven, huge areas of the East Palaearctic are still forested (New- 1993a, b, 1995; Joern, 2004, 2005). Most community ell, 2004; Yan & Shugart, 2005) and thus are important studies in the Palaearctic are for central Europe and dry reference areas for the study of temperate woodland land- and semi-dry grassland habitats (e.g. Fartmann, 1997; scapes. The Far East is one of the three biodiversity hot- Behrens & Fartmann, 2004). Information on the Asian spots in Russia (Venevsky & Venevskaia, 2005) and a part of the Palaearctic is restricted to biogeographic data centre of diversity and endemism of Orthoptera in Eura- (Stebaev et al., 1989; Sergeev, 1998) and detailed studies sia (Sergeev, 1998). of orthopteran assemblages are lacking. Since woodlands Their taxonomy and distribution are well studied, and are usually not considered to be an orthopteran habitat the ease with which they can be sampled and their func- (Theuerkauf & Rouys, 2006) and old forests are rare in tional importance make Orthoptera suitable subjects for central Europe, little is known about habitat selection and ecological and biogeographical studies (Sergeev, 1997; community structure of Orthoptera in natural woodland Lockwood & Sergeev, 2000). Habitat selection in areas in the Palaearctic. Orthoptera is based on a complex combination of dif- We therefore investigated orthopteran communities in ferent and often interrelated environmental factors. Of the natural landscape of the Russian Far East and com- these parameters, the microclimate at oviposition sites, pared the results with observations from the human and which is often affected by vegetation structure, plays a agriculturally dominated landscape of central Europe, crucial role (Uvarov, 1977; Willott & Hassall, 1998). because many taxa occur throughout the Palaearctic (Ser- Sergeev (1997) stressed the suitability of orthopteran geev, 1992, 1997). Hence, orthopteran assemblages in the communities for ecological and biogeographical investi- Lazovsky State Nature Reserve (Primorsky region, Rus- gations. In recent decades many such studies have been sian Far East) were studied to (i) determine their species done in the northern hemisphere. Especially in North composition and abundance in different natural and semi- natural habitats, (ii) analyse orthopteran habitat require- * Present address: Helmholtz-Centre for Environmental Research Leipzig-Halle UFZ, Department of Community Ecology, Theodor-Lieser-Str. 4, D-06120 Halle, Germany. 673 At the coast winters are warmer (average January temperature: –11°C) and summers cooler (average August temperature: 17°C) (Semenchenko, 2003). Sampling of Orthoptera Sampling was carried out on 18 plots representing all the typical orthopteran habitats of the Lazovsky Zapovednik, except floodplains, which were studied by Specht (2004). Nine natural (coastal dunes, semi-dry coastal grasslands, swamps) and nine semi-natural habitats (fallows dominated by Artemisia spp. and meadows) were investigated. The area of the plots was > 2,000–10,000 m2 with a homogenous vegetation structure at every site. Orthopteran densities were recorded in box quadrats (Gardiner et al., 2005) of a total area of 20 m2. From 29/08–15/09/2001 one sample was taken on each plot: The mobile 1 × 1 m (1 m2) and 80 cm high quadrat was randomly placed at twenty different points. Sampling was done in sun- shine at temperatures > 20°C, between 10:00 a.m. and 5:00 p.m. Except for the small Nemobiinae species, which live hidden under stones or in litter on the ground, sampling provided reli- able quantitative data. Most of the specimens were determined in the field and then released. Individuals that could not be iden- tified in the field (Tetrix spp., some Chorthippus spp.) and voucher specimens of each species were collected and identified later. For determination the keys of Bey-Bienko & Mishchenko (1951a, b) and Storozhenko (1986) were used. Nomenclature is based on Storozhenko (1986) and for species that also occur in Europe on Heller et al. (1998). Fig. 1. Study area and location of Lazovsky Zapovednik. Habitat structure ments in relation to habitat structure and microclimate For each plot we measured/estimated fifteen environmental and (iii) compare orthopteran habitat preferences there parameters: inclination, exposure, heights of one (minimum) up to three different vegetation layers (e.g. turf – tall grass – Artem- and in Europe where the same or closely related species isia) and % cover of the following habitat components: total occur. vegetation, field layers, Cyperaceae, Poaceae, herbs, mosses, lit- MATERIAL AND METHODS ter, bare soil, stones and hollows (in swamps). Data analysis Study area Canonical correspondence analysis (CCA) (using CANOCO The study area is located in the Primorsky region (southern 4.51; ter Braak & Šmilauer, 2002), a direct gradient ordination Russian Far East, Fig. 1). The landscape is formed by the technique, was used to determine the organization of orthop- Sikhote-Alin Mountains, stretching from the southwest to the teran species into distinct communities and the relations northeast, parallel to the coastline (average altitude about between habitat structure and species composition (Fielding & 1,000 m a. s. l.). Woodland covers 80% of the Primorsky region Brusven, 1993b, 1995; Palmer, 1993; Szövényi, 2002; Torrusio – taiga in the north, conifer-broadleaved woodlands in the south et al., 2002). Environmental and species data were log arithmi- (Newell, 2004). cally transformed [y’ = ln (y + 1)] to obtain approximately The 1,200 km2 sized Lazovsky State Nature Reserve (i.e. normal distributions and homogenous variances. Species of Lazovsky “Zapovednik”, the official Russian category for this Orthoptera that occurred only on one plot and/or of which protected area) is situated about 200 km east of Vladivostok in < 10 specimens in total were found were not included in the data the southern spurs of the Sikhote-Alin Mountains (134°E/43°N). set (Table 1). Inclination and exposure were not used as vari- It mainly consists of woodlands dominated by Mongolian oak ables in the CCA because only two plots were slightly inclined (Quercus mongolica) with an admixture of Korean pine (Pinus (< 5°); cover (%) of bare soil and stones made up one variable; koraiensis) and various other tree species. The species richness out of the three field layer heights the maximum vegetation of the Zapovednik is impressive (1212 species of vascular height was used in CCA. The statistical validity of the ordina- plants, 57 mammals