J Insect Conserv (2011) 15:633–642 DOI 10.1007/s10841-010-9362-7 ORIGINAL PAPER Impacts of succession and grass encroachment on heathland Orthoptera Jens Schirmel • Jasmin Mantilla-Contreras • Irmgard Blindow • Thomas Fartmann Received: 21 April 2010 / Accepted: 23 October 2010 / Published online: 10 November 2010 Ó Springer Science+Business Media B.V. 2010 Abstract For the conservation of biodiversity, heathlands density was highest in grey dunes. Threatened and specia- present important ecosystems throughout Europe. The for- lised species were restricted to the young stages grey dunes merly widespread habitats are nowadays restricted to small and dwarf-shrub heath. Hence, in order to maintain a high and isolated remnants. Without land use heathland vegeta- diversity of Orthoptera in heathlands, maintaining different tion undergoes succession and, in addition, the increasing successional stages is of critical importance and this should amount of atmospheric nitrogen deposition has resulted in an be integrated into heathland management practices. encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Keywords Biodiversity Á Calluna heath Á Eutrophication Á Orthoptera in a coastal heathland on the Baltic island of Global change Á Microclimate Á Vegetation structure Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf- Introduction shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong European heathlands are important ecosystems for the differences among the successional stages. Orthoptera spe- conservation of biodiversity (Webb 1998) and thus pro- cies richness was highest in transitional stages. The high tected by the EU Habitats Directive (Ssymank et al. 1998). proportion of grasses offer favourable habitat conditions for Due to intensification of land use and afforestation these graminivorous, chorto- and thamnobiont species. Orthoptera formerly widespread habitats are today restricted to small and isolated remnants (Rose et al. 2000; Webb 1998). As a man-made habitat the presence and quality of heathlands J. Schirmel (&) Á I. Blindow strongly depend on traditional land uses like sheep grazing, Biological Station of Hiddensee, University of Greifswald, sod cutting and burning. Without land use or management, Biologenweg 15, 18565 Isle of Hiddensee, Germany the depletion of nutrients is inhibited and heathland vege- e-mail: [email protected] tation undergoes succession towards shrub or tree domi- I. Blindow nated vegetation (Britton et al. 2001; Gimingham 1972; e-mail: [email protected] Webb 1998). In addition, the increasing amount of atmo- J. Mantilla-Contreras spheric nitrogen deposition during the last decades resulted Department of Landscape Ecology, Institute of Botany in an eutrophication of nutrient-poor heathland ecosystems and Landscape Ecology, University of Greifswald, (Heil and Diemont 1983; Remke et al. 2009a), causing Grimmerstr. 88, 17487 Greifswald, Germany e-mail: [email protected] encroachment of grasses like Deschampsia flexuosa and Molinia caerulea (Britton et al. 2001; Marrs 1993; Roem T. Fartmann et al. 2002). As shown for heathlands and dune ecosystems Department of Community Ecology, Institue of Landscape in the Baltic Sea region, even low nitrogen inputs can led to Ecology, University of Mu¨nster, Robert-Koch-Str. 26, 48149 Mu¨nster, Germany such grass encroachment, especially of Carex arenaria e-mail: [email protected] (Remke 2003; Remke et al. 2009a, b). 123 634 J Insect Conserv (2011) 15:633–642 The responses of open terrestrial ecosystems to succes- how does the Orthoptera composition change? (3) Which sional processes have been studied for several invertebrate are the most important drivers for the shifts in Orthoptera groups indicating both positive (butterflies: Balmer and composition? Erhardt 2000) and negative effects on diversity (Orthop- tera: Ko¨hler and Kopetz 1993; Marini et al. 2009). While most studies concentrate on grassland ecosystems (e.g. Materials and methods Balmer and Erhardt 2000; Baur et al. 2006; Kruess and Tscharntke 2002a, b; Marini et al. 2009;O¨ ckinger et al. Study site 2006; Sko´rka et al. 2007), the knowledge of successional effects on insects in heathlands is rare. Moreover, effects of The study was conducted in a coastal heathland on the eutrophication on arthropods have poorly been analyzed in Baltic island of Hiddensee (54°320N, 13°50E) in north- general (Cuesta et al. 2008; Sjursen et al. 2005) and par- eastern Germany in spring/summer 2008. Mean annual ticularly the impact of grass encroachment in heathland temperature in this region is 7.5°C and mean annual pre- ecosystems is hardly known. cipitation is 547 mm (Reinhard 1962). Orthoptera are highly sensitive to environmental chan- We chose an area of approx. 2.5 km2 in the centre of the ges like grazing, mowing and abandonment (Ba´ldi and island were the five main stages of heathland succession Kisbenedek 1997; Fartmann and Mattes 1997; Samways could be found. The number of replicates per stage corre- 1997). They have an important position in food webs of sponded approximately to their respective area fraction in open ecosystems where they often are both the main the study area. In total, 31 plots were established with a invertebrate consumers (Curry 1994) and food source size of 500 m2 each and a homogeneous vegetation struc- (Samways 1997). ture (for the representing stage type). The following suc- The present study investigates the effects of succession cessional stages were studied: (1) open and sparsely and grass encroachment, as the main recent threats to vegetated grey dunes (GD) dominated by Corynephorus heathland ecosystems, on Orthoptera in a coastal heath at canescens, Carex arenaria and cryptogams (N = 6). (2) the Baltic Sea island of Hiddensee, Germany. The study the main (dominant) vegetation type, dwarf-shrub heath aims to assess changes in Orthoptera species richness and (DH) with dominance of Calluna vulgaris, Empetrum assemblage structure along a successional gradient. In nigrum, Salix repens and Erica tetralix as less frequent addition, we try to determine the impact of the potential species (N = 10). (3) Grassy heath (HG; N = 7), where drivers (microclimate, vegetation, soil humidity) of these heather species are regularly displaced by tall graminoids, changes. Based on our results, we give recommendations especially by C. arenaria. This species, usually a pioneer for the management of heathland ecosystemes. The fol- plant of acidic coastal dunes at the Baltic Sea (Remke et al. lowing questions are addressed: (1) How do the succes- 2009b), can build up very dense and high ([1 m) stands sional stages differ in their abiotic conditions and within heathland vegetation (Fig. 1; Remke 2003; own vegetation composition? (2) Do Orthoptera species rich- observations). In the following, we therefore distinguish ness and density differ along the successional gradient, and between ‘‘tall C. arenaria’’ as the eutrophic form occuring Fig. 1 Typical stands of a dwarf-shrub heath and b heath encroached by tall Carex arenaria within the study area of the coastal heath on the Baltic isle of Hiddensee, Germany 123 J Insect Conserv (2011) 15:633–642 635 in grassy heath (HG) and ‘‘small C. arenaria’’ as the typical of 1 m all optically and acoustically observed Orthoptera pioneer form occuring in grey dunes (GD). (4) Unmanaged were counted (total observed area per plot: 70 m2). Indi- heath encroached by shrubs like Betula pendula, B. pu- viduals which could not be determined by observation were bescens or Prunus serotina (HS; N = 4). (5) Young birch caught (by sweep net or hand) and released after determi- forest (BF), which had developed due to the lack of land nation in the field. The loops had a distance of about 5 m to use during the last 30–40 years (N = 4). each other to minimize double counting. Surveys were conducted at three dates during main activity period of Vegetation adult Orthoptera between the end of July and the end of August under favorable weather conditions (warm, sunny Vegetation was sampled three times during May and and calm days) between 10:00 and 17:00 h (CET ? 1). August 2008 in all plots. In each plot, two sub-plots were Prior to starting transect counts all plots were visited at established with a specific size depending on the vegetation least two times (beginning of July and mid-July) for a type: 25 m2 in grey dunes (GD) and dwarf-shrub heath qualitative survey in order to detect the species spectrum of (DH), 50 m2 in grassy heath (HG) and heath with shrubs each plot. Species were determined according to Bellmann (HS) and 100 m2 in birch forest (BF). In each sub-plot, we (2006), and the scientific nomenclature followed Coray and recorded cover of total vegetation (CTV), trees (CTR), Lehmann (1998). shrubs (CSH), grasses (CGR), tall C. arenaria (CTC), small C. arenaria (CSC), cryptogams (CCR) and bare soil Data analyses (CBS). Species cover was estimated according to the scale of Londo (1976, 1984). Density of field layer was esti- Orthoptera species were classified into the life-forms geo- mated in % for the vegetation heights 0–10, 10–20, 20–30, bionts (Oedipoda caerulescens, Myrmeleotettix maculatus, 30–40, and 40–50 cm (Sundermeier 1998) and summed up Tetrix undulata), geo-chortobionts (Chorthippus biguttu- for further analyses (DFL).