Biologia 66/6: 1127—1133, 2011 Section Zoology DOI: 10.2478/s11756-011-0115-1

Response of assemblages to management of montane grasslands in the Western Carpathians

Vladimíra Fabriciusová, Peter Kaňuch &AntonKrištín*

Institute of Forest Ecology, Slovak Academy of Sciences, Ľ. Štúra 2,SK-96053 Zvolen, Slovakia; e-mail: [email protected]

Abstract: Montane grassy habitats in the Western Carpathians are relatively well preserved, maintain high species richness and are often important in accordance to the nature conservation policy in Europe. However, knowledge about the impact of farming on the habitat quality there is rather poor. The influence of various management types (permanent sheep pen, irregular grazing, mowing) on Orthoptera diversity and species determining assemblages of these habitats were analysed on 72 plots in Poľana Mts Biosphere Reserve. Altogether, 36 Orthoptera species (15 , 21 ) were found, whereas the highest number of species was found on plots with irregular grazing (28 species), followed by plots with mown grass (17) and permanent sheep pens (14). All four measures of the assemblages’ diversity confirmed significant differences. Using Discriminant Function Analysis, correct classification rate of Orthoptera assemblages was unambiguous according to the type of management. Each form of the management harboured several characteristic species. Thus implications regarding the biodiversity conservation and grassland management were given. Key words: bush-crickets; ; pasture; ecology; nature conservation

Introduction or the influence of plant succession in meadows without any management (Marini et al. 2009). Higher species Various systems used to manage grasslands have a diversity was found in grazed compared to mowed significant influence on plant and communi- meadows in alpine habitats (Wettstein & Schmid 1999; ties, their species richness and abundance (Kampmann Kampmann et al. 2008). In contrast to that, a more in- 2008; Marini et al. 2008). To assess the biodiversity of tense grazing may even have negative influence on some such habitats, several taxonomical groups of inverte- (Kruess & Tscharntke 2002a). The diversity of brates were analysed already, e.g., butterflies, bees, spi- Orthoptera species decreased with more intense man- ders (Balmer & Erhardt 2000; Kruess & Tscharntke agement in semi-natural grasslands and, on the other 2002b; Knop et al. 2006), bugs (Di Giulio et al. 2001) hand, also with the absence of any management at all and orthopterans. Species of Orthoptera are charac- (woody encroachment; Marini et al. 2009). It is appar- teristic inhabitants of grassland habitats in temperate ent that even though some kind of habitat management zones. They are suitable indicators of habitat quality is essential in the case of European grasslands, it can and changes induced by management, as herbal and generally be said, that a decreasing intensity of farm- grassy plants are used for their roosting and shelter- ing activities increases the diversity of this group ing (often prey of birds and reptiles), foraging (primary (Marini et al. 2008). However, the response to habitat and secondary consumers) and reproduction (courtship management is often different in two main ecological and egg laying) (e.g., Báldi & Kisbenedek 1997; Ingrisch and taxonomical Orthoptera units. Bush-crickets (En- &K¨ohler 1998; Kampmann et al. 2008). Besides com- sifera) prefer more cluttered and matured vegetation, mon and widespread species, many other species exhibit while grasshoppers (Caelifera) usually do not (Kruess specific requirements on moisture (Gavlas et al. 2007), & Tscharntke 2002a; Marini et al. 2009). temperature or sun radiation (Kampmann et al. 2008), Montane grassy habitats in the Western Carpathi- they also show a sensitivity to pesticides (Branson et ans are relatively well preserved and maintain a high al. 2006), succession of vegetation (Marini et al. 2009) species richness of Orthoptera assemblages (Kočárek et and habitat management (Marini et al. 2008). al. 2005; Krištín & Hrúz 2005). However, knowledge Several Orthoptera assemblages were analysed re- about the influence of farming on the habitat quality garding the effect of grazing and mowing intensity (e.g., there is rather poor. These habitats are often impor- Wettstein & Schmid 1999; Kruess & Tscharntke 2002a; tant in accordance to the nature conservation policy Gavlas 2003; Batáry et. al 2007; Kampmann et al. 2008) in Europe (i.e., Habitats Directive), whereas their exis-

* Corresponding author

c 2011 Institute of Zoology, Slovak Academy of Sciences 1128 V. Fabriciusová et al.

Fig. 1. Location of the study sites in Poľana Biosphere Reserve, Central Slovakia (white circle – permanent sheep pen; black square – irregular grazing; grey diamond – mowing). tence depends on suitable management (Gavlas 2003). area is well known due to its high diversity of plant and An exemplary area for this phenomenon is the Poľana animal communities (Janišová et al. 2004). Similarly, rich Biosphere Reserve (Central Slovakia), where numer- assemblages of Orthoptera were recorded there (56 species; ous Orthoptera species inhabit grasslands with various Krištín & Hrúz 2005). Most of the valuable montane grass- levels of human influence (Krištín & Hrúz 2005). On lands are managed by irregular grazing or mowing. How- ever, sheep pens are permanently placed on several sites as the other hand, species-specific requirements on habi- well; which represents one specific type of intense manage- tat quality allow us to predict the composition of as- ment with heavy disturbance of soil and plants in that area. semblages which can assist in finding the proper type of management in natural areas (Konvička et al. 2005). Types of habitat management Based on recent studies conducted in less diverse i) Permanent sheep pen. The vegetation on plots of a size and more human-influenced areas of Western Europe, of 0.5–1.0 ha is heavily grazed and trampled by the flock of we may predict that orthopterans occurring on pre- sheep (200–250 individuals) which rests over night or during served natural grasslands in Central Europe will re- the day in movable sheep pens (Fig. 2). This kind of manage- spond more sensitively, also to less intense manage- ment is usually applied from May until September, the pens ment, as the species’ carrying capacity is significantly are mobile and trans-located once in several days; accord- higher there (cf. Wettstein & Schmid 1999; Kruess & ing to the current weather conditions. Our sampled plots Tscharntke 2002a; Kampmann et al. 2008; Marini et al. where on sites with pens located one or two years before the sampling. The vegetation is characterized by predomi- 2008, 2009). Therefore the aim of our study was first, nantly nitrophilous or ruderal species (e.g., Capsella bursa- to analyse the influence of various management types pastoris, Cirsium arvense, Veronica chamaedrys)andsome on diversity of Orthoptera assemblages and second, to bulky grasses, which may occur due to an increased nutrient identify characteristic species which determine assem- level (e.g., Alopecurus pratensis, Poa trivialis). Overall, the blages of managed montane grasslands in the Western plant species-richness is much reduced in those areas. Carpathians. ii) Irregular grazing. Grasslands grazed irregularly by flocks of sheep (∼250 ind.) or by herds of cattle (∼150 ind.) from May till August, however, only 1–3 days per year in to- Material and methods tal. Due to the low intensity of grazing, an encroachment of shrubs and immature trees is evident (Juniperus commu- Study area nis, Picea abies, Crataegus sp., Rosa canina, Rubus idaeus, The study area was placed within the Poľana Mts caldera Salix triandra, Sambucus racemosa). Short grasses domi- in the Western Carpathians (464–1458 m a.s.l., 48◦34–44 nate in understory (e.g., Agrostis capillaris, Festuca rupi- N, 19◦18–38 E; Fig. 1). This Biosphere Reserve is situated cola, Nardus stricta), with a variety of herbs being present in continental climate with an annual temperature of 2.5– (e.g., Hypericum maculatum, Lotus corniculatus, Thymus 7.0 ◦C and an average precipitation of 650–1300 mm. The pulegioides). Orthoptera and management of montane grasslands 1129

Fig. 2. An exemplary study plot with permanent sheep pen occurrence one year previous to the sampling (Photo by V. Hrúz).

Table 1. Characteristics of the study sites with different manage- ment types.

Management Pen Gra Mow

Sites/plots(n) 2/18 3/27 3/27 Altitude (m a.s.l.) 650–780 670–920 580–940 Aspect SE–S S SE–SW Slope (◦) 15–35 20–30 10–20 Vegetation height (cm) <25 45–80 40–50

Explanation: Pen – permanent sheep pen; Gra – irregular graz- ing; Mow – mowing. Vegetation height shows range of maximal vegetation height on all sites. iii) Mowing. Meadows which are mown once a year using a motor rotary drum mower (ZTR 165, and AKPIL) in July Fig. 3. Design of the sampling scheme. Abundance of Orthoptera or August. The grass is cut at ca 5 cm of height and the species was recorded by sweeping the vegetation on the sampled hay is later removed after drying at the site. The habitat is plots (nine per site). dominated by tall or short grasses (Arrhenatherum elatius, Festuca rubra, Agrostis capillaris, Nardus stricta, Trisetum vegetation, were selected in the study area. However, other flavescens). In contrast to the previous form of management, habitat characteristics which may influence the local micro- the habitat is less vertically differentiated, but many flowers climate sustained similar (Table 1). The size of a site ranged are present (e.g., Campanula patula, Cruciata glabra, Hyper- from 0.6 to 1.8 ha. The distance between these sites was at icum perforatum, Leontodon hispidus, Potentilla erecta). least 1 km as the crow flies. A minimal stand of forest (∼0.3 km) isolated the study sites from other grassy habitats. The Data collection largest distance between study sites spanned seven km and Orthoptera species were sampled by sweeping the grass us- the total area covered comprised ca 17.5 km2. Altogether, ing entomological hand-nets (diameter 40 cm) on selected nine plots were sampled per site (72 plots in total; Table 1) transects (Báldi & Kisbenedek 1997). During the season, in a standardized sampling scheme (Fig. 3) and used for the the abundance of species is changing in accordance to their following analyses. Species were determined directly in the phenology and different development cycle. To get informa- field or under a binocular enhancer in the lab, according tion about the carrying capacity of habitats for the individ- to Kočárek et al. (2005), except for Isophya species, which ual species, we controlled the species abundance regularly: were determined according to Heller et al. (2004). once per month during May – September 2008 (2–4 hours per visit). However, only the records of maximum numbers Data analysis of adults from each species were taken into account of this To measure the diversity of Orthoptera assemblages at the period. Eight grasslands (hereafter sites) representing dif- sampled plots, four characteristics were analysed: 1) num- ferent types of management, hence with different quality of ber of species, 2) number of individuals, 3) ratio of Ensifera 1130 V. Fabriciusová et al.

Fig. 4. Diversity measurements of Orthoptera. A – number of species; B – number of individuals; C – ratio of Ensifera vs. Caelifera species; D – Margalef’s richness index) in the sampled plots (size 60 m2) with three different types of management (Pen – permanent sheep pen; Gra – irregular grazing; Mow – mowing). Presented are medians (point), 25–75% quartiles (box) and non-outlier range (whiskers); *** P < 0.001; ** P < 0.01; * P < 0.05. to Caelifera species and 4) Margalef’s richness index (this each). The highest number of species was found on plots index measures species-richness that is adjusted for sample with irregular grazing (28 species), followed by plots size). This data does not follow a normal distribution; hence with mown grass (17) and permanent sheep pens (14). differences in diversity measurements among three types of All four measures of the assemblages’ diversity con- management were tested using a non-parametric Kruskal- firmed significant differences among grazed grasslands Wallis ANOVA. Pairwise testing between the single cate- and mown meadows or plots where a permanent sheep gories was based on Z-statistics. pen occurred in recent past, respectively. Similarly, the To learn how the relative abundance of species dis- number of individuals was higher in plots with irregular criminates among the types of management, in other words, grazing of livestock and more Ensifera species occurred how species respond to habitat management, we performed there. Moreover, using the Margalef’s richness index, a Discriminant Function Analysis (DFA). Although DFA a significant difference was also found between mown is relatively robust to deviations from normality, regarding this analysis, a positive skewed data of the species’ abun- meadows and sheep pens (Fig. 4). The high environ- dance was transformed by a natural logarithm. The species mental value of studied grasslands was acknowledged which rarely occurred at one of the sites only were excluded by the occurrence of several endangered or protected beforehand. This procedure aimed at reducing possible sta- species, i.e., frivaldskyi (Herman, 1871), tistical bias. Visualization of our results was produced in fusca (Pallas, 1773), stridulus (L., a scatterplot for the discriminant functions of a Canonical 1758) (irregularly grazed habitats) and Pseudopodisma Analysis. All statistical computations were performed using nagyi Galvagni et Fontana, 1996 (mown meadows). the software Statistica 7 (StatSoft Inc. 2004). Discrimination of assemblages Results The overall correct classification rate of Orthoptera as- semblages (characterised by the species abundance), Diversity measures which was significantly different from random (Wilks’ λ Altogether, 36 Orthoptera species (15 Ensifera, 21 = 0.008, F50,90 = 17.693, P < 0.0001), was unambigu- Caelifera) were recorded at 72 sampled plots (60 m2 ous (100%) according to the type of management at the Orthoptera and management of montane grasslands 1131

Table 2. Relative importance of each species in the Discriminant Function Analysis (the lower the value of Wilks’ λ,themoreimportant is the species) and standardized coefficients for canonical roots.

Species Wilks’ λ F2,45 P Root 1 Root 2

Psophus stridulus (L., 1758) 0.974 0.606 0.550 0.263 0.104 Phaneroptera falcata (Poda, 1761) 0.670 11.063 0.000 1.010 –0.067 Pholidoptera aptera (F., 1793) 0.543 18.917 0.000 1.180 –0.610 Arcyptera fusca (Pallas, 1773) 0.674 10.875 0.000 0.976 0.181 parallelus (Zetterstedt, 1821) 0.617 13.963 0.000 0.022 –1.064 brachyptera (Ocskay, 1826) 0.803 5.532 0.007 0.334 –0.568 Tetrix bipunctata (L., 1758) 0.625 13.473 0.000 –0.776 0.576 Chorthippus albomarginatus (Degeer, 1773) 0.747 7.630 0.001 –0.294 0.996 Chorthippus dorsatus (Zetterstedt, 1821) 0.878 3.141 0.053 0.039 –0.491 Gryllus campestris L., 1758 0.677 10.724 0.000 0.875 –0.417 Stenobothrus lineatus (Panzer, 1796) 0.817 5.033 0.011 –0.214 0.537 brachyptera (L., 1758) 0.860 3.672 0.033 0.459 –0.180 Decticus verrucivorus (L., 1758) 0.849 3.987 0.025 0.155 0.598 Omocestus viridulus (L., 1758) 0.903 2.417 0.101 0.109 0.531 Omocestus rufipes (Zetterstedt, 1821) 0.878 3.132 0.053 0.487 0.191 Stenobothrus stigmaticus (Rambur, 1838) 0.959 0.957 0.392 –0.308 –0.028 Isophya kraussii Brunner von Wattenwyl, 1878 0.822 4.864 0.012 0.412 0.443 Gomphocerippus rufus (L., 1758) 0.904 2.399 0.102 0.480 0.051 Pholidoptera fallax (Fischer, 1853) 0.910 2.238 0.118 –0.201 0.439 Chorthippus apricarius (L., 1758) 0.897 2.593 0.086 0.132 0.424 Pholidoptera griseoaptera (Degeer, 1773) 0.940 1.435 0.249 0.327 0.172 dispar (Germar, 1834) 0.882 3.025 0.059 0.452 –0.292 Metrioptera bicolor (Philippi, 1830) 0.920 1.953 0.154 0.346 –0.079 Omocestus haemorrhoidalis (Charpentier, 1825) 0.929 1.729 0.189 0.350 –0.297 Tetrix subulata (L., 1758) 0.941 1.423 0.252 –0.381 –0.00h4

Eigenvalues 21.418 4.231 Cumulative proportion 0.835 1.000

sampled plots. The first discriminant function explained 83.5% of the data variation (Table 2), whereas both of the revealed discriminant functions were statistically significant (P < 0.0001). Each form of the management harboured several characteristic species. Plots man- aged by irregular grazing were weighted most heavily by the Pholidoptera aptera, Phaneroptera falcata, Ar- cyptera fusca, Gryllus campestris, Omocestus rufipes, Metrioptera brachyptera and (Ta- ble 2) at the first discriminant function (root 1) towards positive values (Fig. 5). Assemblages of mown mead- ows were positively marked, mostly by Chorthippus al- bomarginatus, Decticus verrucivorus, Tetrix bipunctata and Stenobothrus lineatus at the second function (root 2). On the other hand, mostly species as Chorthippus parallelus, and Chorthippus dor- satus had a negative weight in this function. These species occurred predominantly on plots managed with pens (Fig. 5). Commonly or non-specifically occurring species, in this instance Leptophyes albovittata (Kollar, 1833), Meconema thalassinum (De Geer, 1773), Metrioptera Fig. 5. Discrimination among three different types of manage- ment (Pen, permanent sheep pen; Gra, irregular grazing; Mow, roeselii Hagenbach, 1822, Tettigonia cantans (F¨ussly, mowing) in the canonical analysis scatterplot. Points represent 1775), (L., 1758), Chorthippus sampled plots, species entering the model are displayed in Ta- brunneus (Thunberg, 1815) and Chorthippus mollis ble 2. (Charpentier, 1825), did not enter the model (Wilks’ λ = 0.962–0.998). Species already excluded beforehand the computation of the DFA due to their very rare Discussion occurrence (only one site) were Isophya camptoxypha Fieber, 1853, P. frivaldskyi, P. nagyi and Chorthippus Habitat management and species richness montanus (Charpentier, 1825). We found in natural montane grasslands, that habi- 1132 V. Fabriciusová et al. tat management had a significantly different influence fertilised mown meadows in the Alps (Marini et al. on species diversity according to its kind and intensity. 2008). Despite the differences caused by management, assem- blages of Orthoptera were remarkably rich and diverse Bioindicators and nature conservation comparing to other European circumstances (cf. Kruess Although the structure of an assemblage differs region- & Tscharntke 2002a, b; K¨ohler & Renker 2004; Marini ally due to natural environmental variables (Batáry et et al. 2008). A high ratio of Ensifera vs. Caelifera indi- al. 2007; Kampmann et al. 2008), we found several cates a very low level of human exploitation (Ingrisch characteristic species indicating different types of the &K¨ohler 1998; K¨ohler et al. 2004). Generally, irregu- management within a relatively small area (Table 2). lar grazing had a positive effect on many insect species Species preferring overgrowth and secondary succession in human-influenced habitats (e.g., Gavlas 1999, 2003; of vegetation (P. aptera, P. falcata), species sensitive to Wettstein & Schmid 1999; Krues & Tscharntke 2002b; over-fertilization (G. campestris, A. fusca, P. stridulus), Kampmann et al. 2008; Marini et al. 2008). However, an or species preferring higher moisture (M. brachyptera, increasing intensity of grazing tends to decrease species C. dispar) typically dominated in grasslands with man- diversity (Krues & Tscharntke 2002a). Interestingly, agement of the lowest intensity (Marini et al. 2008, comparing to other less intense types of management 2009; Gavlas et al. 2007). Widespread and common in our study area (meadows mowed once a year), a species of Caelifera (C. parallelus, E. brachyptera, C. positive effect of irregular grazing was observed as well dorsatus) which determined mown meadows or plots (Fig. 4). It is evident that the activity of large mam- with sheep pens in the Poľana Mts are characteristic mal herbivores induces positive disturbances of forests, for other fertilized meadows, which are also used for while patches of new initial open habitats (i.e. various commercial production (Detzel 1998; Marini et al. 2008, grasslands) increase the biodiversity of the temperate 2009). Such dominance of every-topic species in human- zone. However, natural performance of large herbivores influenced farmland was found in other insects as well in habitat management lasted in the late Pleistocene (Di Guilio et al. 2001; Kruess & Tscharntke 2002b). (most of them were extinct). Nowadays some human in- A high intensity of farming resulted in a strong put is highly needed and recently observed high species decrease of the European biodiversity during the last richness of irregularly grazed grasslands is just an imi- decades (Hoste-Danylow et al. 2010). To stop such a tation of the prehistoric situation (Persson et al. 2000; trend, the European Community recently implemented Vera 2000; Konvička et al. 2004, 2005). The result of agro-environmental programmes. The aim is to support such management is vertically and horizontally differ- less intense types of grassland management (Knop et entiated: a mosaic of low and high grasses, herbs, shrubs al. 2006). Timing and frequency of mowing are crucial and trees, as well as spots with bare ground (Gibson for meadow dwelling species (Marini et al. 2008). We 1988; Gavlas 1999; Marini et al. 2010). Irregular graz- found a significant decrease in the abundance (–43%) ing produces various ecological niches at one site, which and the species number (–21%) after mowing the grass is also preferred by various species for feeding, roosting in July and August. To eliminate this negative influ- and reproduction (e.g., Balmer & Erhardt 2000; Kruess ence, mowing the grass in patchy mosaic or at the end & Tscharntke 2002a; Kampmann et al. 2008). of the summer season is suggested (Wettstein & Schmid Extensive grass-mowing and permanent grazing 1999; Gavlas 2003; Hoste-Danylow et al. 2010). While during the whole season are definitely less suitable lacking any kind of management, some mowing in a forms of management (Wettstein & Schmid 1999; longer period of time (3–5 years) is essential to pre- Kampmann et al. 2008). Both forms generate signifi- vent gradual woody encroachment (Marini et al. 2009). cantly decreased roosting possibilities, which increase Though irregular grazing is now believed as the best the predation risk, while motor mowing also threatens grassland management regarding the biodiversity, it is with the direct killing of individuals (Humbert et al. difficult to maintain in montane areas of Slovakia due 2009, 2010). Moreover, insects or their eggs are car- to low economical profits. Hence, an appropriate appli- ried off with the mowed grass and the then bared soil cation of agro-environmental policy (Marini et al. 2008, is overheated and dried out (Belovsky & Slade 1993; 2009) may contribute to the conservation of the unique Konvička et al. 2005). Our discoveries agree with the montane biodiversity in the Western Carpathians. situation in regularly mown alpine meadows where only several species dominated (e.g., C. para- llelus), while bush-crickets had very low individual den- sities (Marini et al. 2008). According to our current Acknowledgements knowledge, the influence of permanent sheep pens on the local diversity of insects has not been tested yet. We thank D. Galvánek for his valuable help with the selec- tion of the study sites and providing plant species data. We We found that the composition of assemblages was also thank Filip, Marek and Adrián Fabricius, as well as V. very poor with dominancy of common Caelifera; even Hrúz for helping in the field and accompaniment. B. Kiehl 1–2 years after departure of livestock from the site is acknowledged for linguistic corrections and valuable com- (Fig. 4). Concentrated livestock causes over-fertilisation ments to the first version of the manuscript. This study was and growth of nitrophilous plants. However, rather supported by a grant of the Slovak Grant Agency (VEGA similar Orthoptera assemblages were also recorded on 2/0157/11). Orthoptera and management of montane grasslands 1133

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