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Effects of Habitat and Landscape Features on Grassland Orthoptera on fl Oodplains in the Lower Reaches of the Tisza River Basin

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Effects of Habitat and Landscape Features on Grassland Orthoptera on fl Oodplains in the Lower Reaches of the Tisza River Basin EUROPEAN JOURNAL OF ENTOMOLOGYENTOMOLOGY ISSN (online): 1802-8829 Eur. J. Entomol. 113: 60–69, 2016 http://www.eje.cz doi: 10.14411/eje.2016.007 ORIGINAL ARTICLE Effects of habitat and landscape features on grassland Orthoptera on fl oodplains in the lower reaches of the Tisza River Basin ATTILA TORMA1 and MIKLÓS BOZSÓ 2 1 Department of Ecology, University of Szeged, Közép Fasor 52, Szeged H-6726, Hungary; e-mail: [email protected] 2 Plant Health and Molecular Biology Laboratory, Directorate of Plant Protection, Soil Conservation and Agri-environment, National Food Chain Safety Offi ce, Budaörsi u. 141-145, Budapest H-1118, Hungary; e-mail: [email protected] Key words. Orthoptera, Caelifera, Ensifera, dispersal, diversity, fl ooding, niche breadth, reproductive potential Abstract. The Tisza River Basin is an important area as it is a green corridor in which there are highly endangered habitats and a high level of biodiversity. The patterns in the species richness of invertebrates and the environmental conditions affecting these patterns are poorly studied in the grassy habitats in the lower reaches of the Tisza River Basin. The present study focuses on the effects of fl ooding, habitat and landscape features on the species richness of orthopterans at 24 grassland sites in two different landscapes. The relations between the explanatory variables and the pattern of diversity of orthopterans with different life-history traits were studied, using ordination and Generalized Linear Mixed Models. Although the infl uential factors for the different trait groups differed, we suggest that landscape features are the most important in shaping orthopteran assemblages, whereas habitat characteristics and fl ooding have comparatively little effect. Habitat characteristics affected only the non-xerophilous and Ensifera species and only the species richness of non-xerophilous orthopterans in fl ooded and non-fl ooded sites differed. We emphasize that even in countries where there are still considerable areas of high value natural grasslands, such as Hungary, non-protected meadows, linear grassy habitats (dikes, ditch banks, road verges, etc.) need more attention and should be given higher priority in the conservation of invertebrates. INTRODUCTION the remaining grasslands are surrounded by a highly modi- In their natural state, riverine landscapes are character- fi ed landscape consisting of arable fi elds and forest planta- ized by mosaics of various habitat patches. Due to their tions, often of non-native trees. Despite these changes, the high heterogeneity and connectivity (Naiman et al., 2005), Tisza River Basin (TRB) still includes highly endangered they can support a diverse fl ora and fauna (Gregory et al., habitats, which are important ecological areas and green 1991; Zwick, 1992; Ward et al., 1999). However, many Eu- corridors (L. Gallé et al., 1995; Rádai, 1995) with a high ropean rivers are restricted to narrow riverbeds bordered level of biodiversity (Sommerwerk et al., 2009). The in- by dikes and the majority of fl oodplain habitats have been vertebrate fauna in the TRB is well-documented (see e.g. transformed into agricultural land (Tockner et al., 2009), L. Gallé, 2005, 2008), but there are only a few studies on causing a severe decline in biodiversity (Godreau et al., some taxa, e.g. spiders (Araneae) (R. Gallé et al., 2011) 1999). and true bugs (Heteroptera) (Torma & Császár, 2013), The River Tisza is the largest tributary of the Danube which indicate the environmental conditions that are likely and its catchment includes most of the Carpathian Moun- to infl uence the species richness of grassland arthropods tains covering approximately 157,000 km2 (Sommerwerk in this human-modifi ed riverine landscape. To the best of et al., 2009). The regulation of the Tisza in the 19th century our knowledge, patterns in the diversity of assemblages of caused profound changes; a considerable amount of the Orthoptera in relation to their life-history traits and envi- former fl oodplain has since never been fl ooded. However, ronmental conditions have not been studied in this region. on this non-fl ooded part of the former fl oodplain (so-called Orthopteran assemblages are known to depend on differ- “historical fl oodplain”) there were several high value ent and often interrelated environmental factors. Vegeta- habitats, i.e. pastures, woody pastures and hay-meadows, tion has a great infl uence on many invertebrates, including in this extensively used mosaic landscape until the 1950s orthopterans (Batáry et al., 2007; Poniatowski & Fartmann (Deák, 2007; Sendzimir et al., 2008). During the social- 2008, 2010). Vegetation is related to soil and microclimate ist era, intensifi cation of agriculture resulted in a decrease conditions, which are also important for orthopterans (Wil- in the area of these grasslands (Deák, 2007). Nowadays, lott & Hassall, 1998; Gardiner & Dover, 2008). In ripar- Final formatted article © Institute of Entomology, Biology Centre, Czech Academy of Sciences, České Budějovice. An Open Access article distributed under the Creative Commons (CC-BY) license (http://creativecommons.org/licenses/by/4.0/). 60 Torma & Bozsó, Eur. J. Entomol. 113: 60–69, 2016 doi: 10.14411/eje.2016.007 ian landscapes, fl ooding and land use pressure have a pro- sweep netting was carried out along three, 50 m long, fi xed tran- nounced infl uence in shaping assemblages of Orthoptera sects in 2009. To avoid periods of fl ooding, sweep netting was (Dziock et al., 2011). The majority of species of Ortho- carried out in summer. The fi rst samples (30 May–2 June) were ptera are associated with open grassy habitats; therefore, collected before mowing and the second samples (22–24 July) when the vegetation started to regrow after mowing. For the data for these species the amount of grassland in a landscape is analyses, the sweep netting samples were pooled separately for important (Marini et al., 2008; Badenhausser & Cordeau, transects and periods, resulting in a total of 24 statistical samples 2012). The “habitat amount hypothesis” (Fahrig, 2013) (one sample per site). postulates that patch size and patch isolation effects are Assessments of explanatory variables both due mainly to the sample area effect, thus patch size and isolation can be replaced with a single variable, the Habitat and landscape features were assessed at each site (Ap- pendix 1). Habitat characteristics included features of the vegeta- amount of habitat. tion and soil water content. The vegetation was sampled in three, In order to determine the main factors affecting orthop- 1 × 1 m quadrats along each transect. Mean data for the quadrats terans, we tested the effects of (1) habitat characteristics were used to defi ne variables at the sites sampled. To characterize (soil moisture and vegetation structure and diversity), (2) the structure of vegetation, the average height of the vegetation, landscape features (amount of grassland habitat, landscape the total cover of vegetation at 10 and 40 cm above the ground structure) and (3) fl ooding (fl ooded vs. non-fl ooded sites) and cover of litter were recorded. To characterize the richness of on the species composition and richness of assemblages of vegetation, the total number of species of plants and of only the Orthoptera. The effect of landscape composition on assem- dicotyledonous plants were recorded in the quadrats. Soil sam- blages of Orthoptera is often scale-dependent (e.g. Marini ples were taken from the top 10 centimetres close by the coe- nological quadrats. The percentage of gravimetric water in soil et al., 2009a), thus we also aimed to determine the appro- samples was measured. priate spatial scale for assessing the amount of grassland To assess the amount of habitat we measured the percentage of habitat. As species with different life history traits often the area covered with grassland in a radius of 100, 250, 500 and need different environmental conditions, the effects on 750 m around each site using ArcView 3.11 GIS software. species richness of various life-history traits were tested Life-history traits separately. Dispersal ability, niche breadth and reproductive potential MATERIAL AND METHODS were the traits considered, because they are hypothesized to be key determinants of species persistence (Kotiaho et al., 2005). As Study sites and sampling design a measure of dispersal ability, the mobility index (Reinhardt et Assemblages of Orthoptera were studied at 24 sites in two al., 2005) was used. However, mobility is not a constant trait for different landscapes. The landscapes were located on the same orthopterans; it may differ between and within populations (e.g. side of the river in Csongrád County, Hungary, and were selected Endo, 2006; Poniatowski & Fartmann, 2009). To reduce the effect based on intensity of land use and landscape structure. The hete- of this potential variability, broad mobility classes: sedentary, in- rogeneous landscape (HET) was situated near the town of Szeged termediate disperser and mobile species, were identifi ed. Further, (approx. 46°17´34 ̋ N, E 20°12´45 ̋ E) and it consisted of a mosaic intermediate dispersers were excluded from the analyses, as they of various habitats. The percentage of the area covered by inten- are often the species whose classifi cation is uncertain (cf. Marini sively managed arable fi elds was high (58.3 ± 3.4%, mean ± SE et al., 2009a; 2012). The mean number of ovarioles is a rough within a radius of 500 m around the sites). Small patches of mead- measure of the reproductive potential of females (Reinhardt et ows with trees and abandoned fi elds were embedded in the matrix al., 2005) and is generally coded into three categories: low (< 10), of arable fi elds. Numerous trees and bushes also occurred along intermediate (11–25) and high (> 25) (cf. Dziock et al., 2011). road verges, but continuous forest occurred only near the river. However, this trait is proportional to body size and the phylog- The percentage of the area covered by forest habitats, including eny of the species (Dziock et al., 2011), and Ensifera species are single trees and bushes, was 18.6 ± 5.6%.
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