Decline of rare and specialist species across multiple taxonomic groups after grassland intensification and abandonment

Andreas Hilpold1,*,§, Julia Seeber1,2,§, Veronika Fontana1, Georg Niedrist1, Alexander Rief2,

Michael Steinwandter1, Erich Tasser1, Ulrike Tappeiner1,2

1 Institute for Alpine Environment, Eurac Research, Drususallee 1, 39100 Bolzano/Bozen, Italy

2 Department of Ecology, University of Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria

§ these authors contributed equally as first authors

* corresponding author: [email protected], Institute for Alpine Environment, Eurac

Research, Drususallee 1, 39100 Bolzano/Bozen, Italy

Abstract

Traditionally managed mountain grasslands are declining as a result of abandonment or intensification of management. Based on a common chronosequence approach we investigated species compositions of 16 taxonomic groups on traditionally managed dry pastures, fertilized and irrigated hay meadows, and abandoned grasslands (larch forests). We included faunal above- and below-ground biodiversity as well as species traits (mainly rarity and habitat specificity) in our analyses. The larch forests showed the highest species number (345 species), with slightly less species in pastures (290 species) and much less in hay meadows (163 species). The proportion of rare species was highest in the pastures and lowest in hay meadows. Similar patterns were found for specialist species, i.e. species with a high habitat specificity. After abandonment, larch forests harbor a higher number of pasture species than hay meadows. These overall trends were mainly supported by spiders and vascular plants. Lichens, bryophytes and carabid showed partly contrasting trends. These findings stress the importance to include a wide range of taxonomic groups in conservation studies. All in all, both abandonment and intensification had similar negative impacts on biodiversity in our study, underlining the high conservation value of Inner-Alpine dry pastures.

Keywords (5-8): biodiversity survey, land-use change, multi-taxon study, trait-based approach, biodiversity conservation

1 Introduction

The agricultural landscapes in Central Europe are currently in a state of change. This is especially pronounced for mountain areas, where frameworks for agriculture are, due to steepness, elevation and remoteness, frequently harsher than in lowlands, often leading to the abandonment of farms (Lasanta et al. 2017). Also regions where agriculture has been maintained

(e.g. easily accessible locations on hillsides) show strong changes in agricultural practice via intensification of grassland management (fertilization and irrigation), specialization on a single land-use form (e.g. livestock farming), or the establishment of permanent crops (orchards and vineyards) (Mottet et al. 2006, Graf et al. 2014, Egarter Vigl et al. 2016). Less favoured grasslands such as steep alpine pastures and meadows are continuously abandoned (Niedrist et al. 2009).

In the continental valleys of the Central Alps pastures on south-exposed slopes are usually covered by typical dry grassland vegetation which resemble that of the driest parts of Eastern

Europe. Most prominent are the dry grasslands in the valleys of Aosta (Italy), Valais (Switzerland) and Vinschgau/Val Venosta (Italy). The European Union gave high priority to their conservation by including them in the Habitat Directive “Sub-Pannonian steppic grasslands” (code 6240*,

Lasen and Wilhalm 2004, European Commission 2013). These types of grassland are especially affected by recent land use changes, since their fodder value is even lower than that of other pastoral habitats (Graf et al. 2014). They are, however, rich in floristic and faunistic elements which are extremely rare in Central and Western Europe (e.g., Canthophorus melanopterus,

Tamanini 1982; Omocestus petraeus and Stenobothrus nigromaculatus, Nadig 1991; Astragalus exscapus, Wilhalm 2018) and are in the focus of several Agri-Environmental Measures of the EU. Numerous studies emphasize the outstanding value of dry grasslands for European biodiversity (e.g. Swaay and Warren (2006) for butterflies, Lacasella et al. (2015) for spiders). Dry calcareous grasslands exhibit among the highest alpha diversity of vascular plants on a small scale worldwide (Wilson et al. 2012). These “hotspots of biodiversity” (Cremene et al. 2005) are under constant threat due to land-use changes and ensuing habitat fragmentation (Zulka et al. 2014,

Braschler and Baur 2016, Lengyel et al. 2016). Virtually all Alpine pastures below the treeline will be substituted by forests on the long term, if the grazing impact were reduced (Ellenberg and

Leuschner 2010), or changed into more productive land use forms, such as hay meadows, vineyards or crop fields (Egarter Vigl et al. 2016). In South Tyrol, the analysis of the land-use change revealed two clear trends (Tasser et al. 2009): In the agriculturally favourable areas, grassland and arable land disappeared to a large extent (-59% of the area in 1860) and their place was taken by permanent crops such as apple orchards, vineyards or settlements. Until the 60’s, the south exposed slopes in the Vinschgau Valley were characterized by a mosaic of dry pastures and open wood grasslands. These land-use forms were later largely abandoned (-74%) and today, forests can be found in large parts of this area.

Biodiversity studies on the impact of intensification and abandonment of low-input grasslands and dry pastures in particular are plentiful available. Both land-use changes may lead to the loss of characteristic species due to habitat simplification or degradation (as reviewed by Stoate et al.

2009). In most cases these studies are, however, related to plant diversity (Hector and Bagchi

2007, Hülber et al. 2017, Halada et al. 2017) or to the diversity of one or few groups, very often birds and butterflies (Lessard-Therrien et al. 2018, Zografou et al. 2017, Vermaat et al. 2017,

Ernst et al. 2017). Yet, the study of multiple taxonomic groups is highly informative (Zulka et al. 2014), since patterns of single groups can be very different, partly due to high β-diversity in dry grasslands (Turtureanu et al. 2014). Furthermore, the utilization of species traits for evaluating community diversity patterns can provide more comprehensive insights than simple species numbers, since changes in environmental conditions due to land-use changes might result in the environmental filtering of species (Simons et al. 2016, Cadotte and Tucker 2017).

In this study, we surveyed the species composition and traits of 16 taxonomic groups to study the effect of land-use changes on species diversity on formerly grazed dry grasslands in the

Central Alps. We sampled a land-use transect from dry pastures to either intensively used grassland, i.e. hay meadows, or larch forests as successional stages after abandonment. Special emphasis was put on the conservation value of the three habitats, since although an impoverishment due to intensification and abandonment has been frequently reported for single organism groups, an overall assessment is still missing. To evaluate the conservation value we analyzed the proportion of rare and specialist species. We aimed to answer the following questions: (1) Does intensification or abandonment cause consistent patterns regarding species richness, specificity and rarity among different taxonomic groups? (2) Which taxonomic groups are especially affected by these land use changes? and (3) Does the share of rare and specialist species decline in the intensified or abandoned areas?

2 Material and Methods

Compliance with ethical standards Eurac research has a general permit to conduct scientific research in the LT(S)ER area

Matsch/Mazia. The study design was carefully planned to ensure that no populations were endangered.

2.1. Study area

The Long-term socio-ecological research (LTSER) site Matsch/Mazia is located in the Central

Eastern Alps, in the northernmost part of Italy (Province of South Tyrol, N 46.6840°, E 10.5860°,

Fig. 1 aerial picture, Appendix A2 climate and site parameters). The climate at the study sites is subcontinental with 525 mm mean annual precipitation. The air temperatures range from a minimum of –15.29° to a maximum of +27.79°C with the coldest month average –2.67°C

(January) and the warmest +15.93°C (July).

We selected three habitat types with three replicates each: intensively used hay meadows, dry pastures, and larch forests. All nine sites were within a distance of 3 km and comparable regarding elevation (1500 m a.s.l.), inclination (5-15°) and aspect (southwest).

The three hay meadows (H1, H2, H3) were already identified as hay meadows in old maps of the 19th century. Nevertheless, hay meadows in the investigated area underwent qualitative changes due to an installation of irrigation systems and an elevated fertilizing activity over the last fifty years (Pecher et al. 2017). Following Tomasi et al. (2016) they are not part of the

Directive habitat 6520. They all belong to the class Molinio-Arrhenatheretea and therein to the alliance of Trisetion flavescentis.

The dry pasture sites (P1, P2, P3) are part of the priority habitat Sub-Pannonian steppic grasslands (code 6240*) (Lasen and Wilhalm 2004) of the EU Habitats Directive. They belong to the alliance Stipo capillatae-Poion xerophilae (Br.-Bl. & Richard 1950) within the order

Festucetalia valesiacae (Br.-Bl. & Tüxen ex Br.-Bl. 1950) (Mucina et al. 1993). All pastures were not subjected to substantial land-use-changes over the last 160 years.

The larch forests have been abandoned from pastures within the last 30 (L2) to 160 years (L1).

All of them are successional stages which will on the long term change into a mixed coniferous forest (Larici-Piceetum), a habitat type still very common in the Alps (Tasser et al. 2007). Larch forests of that type are not in the focus of European conservation policies.

2.2 Sampling design

For better comparability of the data, the study was conducted mostly within one week in June

2016 (June 26–30, at the peak of vegetation, before the first cut of the hay meadows) on the sites as described above. End of June is a time period which allows for collecting and identifying a large number of plant and animal species inhabiting sites at this specific elevation. On each of the nine sampling sites a central pole was established, around which the experts erected their required minimum sampling area (Mueller-Dombois and Ellenberg 1974). On two opposing corners within a 5 m distance from the central pole, two pitfall traps were installed. They consisted of plastic cups with 8.5 cm opening diameter, converging on top, filled with 150 ml of propylene glycol as preservative. Additionally, they were covered with a transparent acrylic glass roof to avoid flooding. Pitfall traps were installed two weeks prior to the investigation to collect epigeic (mainly spiders, beetles, and millipedes), which were pre-sorted and passed on to the experts. Arthropods (besides grasshoppers, butterflies and burnets) and earthworms from all surveys were transferred to 75% ethanol and adult specimens were identified to species level using a dissecting microscope by standard identification keys.

The chronological order of taxa to be surveyed was determined according to the degree of destructiveness of the particular method. The following 16 taxonomic groups were surveyed:

(1) Vascular plants (Tracheophyta) were recorded (cover and abundance) within a minimum area of 25 m² for meadows and pastures and 225 m² for the larch forests following the method of

Braun-Blanquet (1964). The minimum areas were chosen following Dierßen (1990). Specimens of species which were difficult to identify in the field were collected and identified in the laboratory.

For the recording of (2) Bryophytes (Bryophyta) a 49 m² minimum area was surveyed and coverage of every species was noted.

(3) Lichens (Lichenes) in meadows and pastures were recorded from five subplots on the ground

(50 × 40 cm) within a 225 m² area around the central pole ( Scheidegger et al. 2002). In forests, five trees were additionally randomly selected within 500 m² around the central pole, from 4 subplots (10 × 50 cm) on each tree all species were recorded (cf. Nascimbene et al. 2010).

(4) Earthworms (Lumbricidae) and (5) millipedes (Diplopoda) were extracted by heat from three soil samples (20 × 20 cm, 15 cm deep, taken randomly within a 100 m² area around the central pole) using a modified Kempson apparatus (Kempson et al. 1963) with propylene glycol as as collection fluid.

(6) Spiders (Araneae) and harvestmen (Opiliones) were hand collected in the field (1 hour per site) within an area of 100 m² around the central pole to obtain a comprehensive list of species.

Additionally, individuals from pitfall traps and soil samples were added to the survey. Adult specimens were identified to species level, juveniles to the highest taxonomic resolution possible.

(7) Oribatid mites (Oribatida) were extracted by heat in a Macfadyen extractor (Block 1966) from four samples (10 cm in diameter, 8-10 cm deep, taken from a 25 m² area around the central pole, including herb, grass, litter, and topsoil); ethanol (75%) was used as collection fluid.

Within a 100 m² area around the central pole, (8) grasshoppers (Orthoptera) were caught by hand and with a sweep net for 20 minutes. Additionally, vocals were used to identify species. A second survey was conducted in the first half of September of the same year, since in June most species had not been adult.

(9) Bugs (Heteroptera) were collected by hand and with a sweep net (1 hour per site), as well as with a modified leaf vacuum (Echo ES, 255-ES, with a gauze bag fitted into the opening) within a

100 m² area around the central pole to obtain a comprehensive list of species. With the leaf vacuum three samples per site were taken by sucking the vegetation a hundred times each.

Additionally, specimens from pit fall traps were identified.

(10) Ground beetles (Carabidae) were collected from pitfall traps and soil samples as mentioned above.

(11) Rove beetles (Staphylinidae) were collected by hand sampling (netting, sifting of ant nests, sifting of leaf litter and detritus) within a 25 m² area around the central pole and from pitfall traps.

Further families (Buprestidae, Catopidae, Coccinellidae, Curculionidae, Elateridae,

Geotrupidae, , Silphidae) with only very few species obtained from pitfall traps and soil samples were also identified. (12) Ants (Formicidae) were recorded via nest counts from 4 m² plots (meadows and pastures) or 9 m² plots (forests) within 100 m² around the central pole (following the sampling design of

Seifert 2017). Individuals from each nest were stored either in 95% ethanol or as dried preparation after identification.

(13) Butterflies (Papilionoidea) and burnets (Zygaenidae) were caught with a sweep net (45 minutes per site) within 400 m² around the central pole, identified to species level, and released.

(14) Birds (Aves) were recorded using a combination of transect and point-count methods (30 min per site). Bird surveys were conducted early in the morning and in the evening, since these are the times when most species are active. We classified bird species into pasture or woodland species according to their potential breeding habitat (information gathered from Niederfriniger et al. 1996). Due to intensive management no species were assigned to the hay meadows.

(15) Reptiles and (16) Amphibians were findings of collaborators during summer field work and were included in the species list (four species).

2.3 Species traits

For each species we assigned three traits including rarity, habitat specificity, and light demand/preference:

Rarity (three categories: rare, frequent, common): A species was defined as rare if the area of occupancy in the province of South Tyrol (Italy) was small. The area of occupancy was deduced from the number of known sites of present occurrence. Main source was the database of the

Museum for Nature South Tyrol and the internet portal www.florafauna.it (Appendix A3a). Due to the current heterogeneous degree of exploration for the single taxonomic groups, the thresholds between the categories were defined individually, but was set to approximately 15 %.

For taxonomic groups with scarce available data we relied on the judgement of experts (Appendix

A3b).

Habitat specificity (three categories: low, medium, high): A species with a high habitat specificity is restricted to a specific habitat type and is therefore a specialist species. The information was retrieved from taxonomic literature (Appendix A3a).

Light demand/preference (three categories: low, medium, high): Plants with low light demand prefer closed forests, plants with medium light demand are associated with shrubs and semi- open areas, plants with a high light demand can be found in open areas (following the Ellenberg indicator value EIV, Ellenberg et al. 1992). The light preference of the was deduced from their preferred habitat, i.e., animals which are usually found in open areas were given a high light preference, while those associated with either closed forests or with high vegetation in open areas were attributed with intermediate to low light preferences.

2.4 Site traits

Important site traits were measured with the “Ecobot” (Wohlfahrt and Tasser 2014), a mobile device which consists of a four-component net radiometer mounted on a hand-held broomstick, an air temperature/humidity sensor in a ventilated radiation shield, a two-dimensional sonic anemometer, a soil temperature and volumetric water content sensor as well as a pair of down- and upward looking multi-spectral sensors. During the survey week, 10 measurements were taken on each site between 10 a.m. and 3 p.m. As a result, meteorological and hydrological standard parameters (air temperature, radiation, soil moisture, etc.) as well as derived vegetation indices such as PRI (photochemical reflectance index) or NDVI (normalized difference vegetation index) were available for the single sites (Appendix A2).

As further important site traits we took into account the Ellenberg indicator values of all vascular plant species (EIV; Ellenberg et al. 1992). The EIVs used were L (light), H (humidity), T

(temperature) and N (nitrogen). We weighted the EIVs with the abundance values of the species occurring in each site.

Finally, three 30 × 30 cm microplots were established in each of the study sites. Plants from each microplot were sorted by functional groups (legumes, grasses, forbs, dwarf shrubs), and the dry aboveground phytomass of each group in the microplot was determined in the laboratory.

2.5 Data analysis To test for differences between proportions of rare and specialist species between habitats, respectively, we employed Chi² tests at significance level p < 0.05. Differences in community composition were evaluated by detrended correspondence analyses (DCA) using the vegan package in R (version 2.5-2, Oksanen et al. 2017).

We used the trait “high light demand/preference” to evaluate whether the shared species of two habitat types could more likely be assigned to the dry pastures (inhabited by species with a high light demand/preference due to low vegetation height) or the converted habitat (hay meadows or larch forests, both offering shelter from direct sun light to most invertebrate groups because of the presence of a dense herb layer shading the soil surface and/or a tree layer).

We used a fourth-corner analysis to assess the relationship between species presence, species traits (rarity, habitat specificity), and site traits (Brown et al. 2014). This approach allows to study environment-trait associations using species presence/absence matrices, environmental data across sites and trait data across species. The site traits used in the analysis were chosen after careful consideration, in the end these were: (1) the standard deviation of the surface temperature, indicating high or low daily temperature variation, (2) total phytomass as proxy for vegetation height and percentage of cover, (3) Ellenberg indicator value L (light) and (4) Ellenberg indicator value H (humidity).

All analyses were conducted using the open-source statistical programming language R (version

3.4.4, R Core Team 2018) in R Studio (version 1.1.383, RStudio Team 2017).

Due to low species numbers (maximum of 5 species) in 4 out of the 16 taxonomic groups studied

(earthworms, millipedes, reptiles, amphibians), we did not include them in the graphical display of our results. However, they are included in statistical analyses on species composition patterns.

3 Results 3.1 Species richness

For the three habitat types we identified 638 species belonging to 16 taxonomic groups (Fig.

1, Appendix A4). The larch forests were the species-richest habitats with 345 species, followed by the dry pastures with 290 species and the intensively used hay meadows with 163 species.

Most species were either vascular plants (140 species), spiders (130 species), or oribatid mites

(87 species).

Within habitat types, the hay meadows had the highest number of species found in all three replicate sites (50.3%), followed by dry pastures (38.6%), and larch forests (34.5%). The high variability in terms of species composition in the three replicates of larch forests was confirmed by the detrended correspondence analysis (as expressed by the distance between replicate sites,

Appendix A1). A correlation between the pairwise distance between replicate sites and their shared species showed no effect of distance on the variability of species composition within habitat type (R = 0.24).

The highest numbers of rare and specialist species were found on the dry pasture (47 and 58 species, respectively), most of them were spiders and vascular plants (Appendix A4, for a full list of rare and specialist species see data repository). Thus, 16% of the species found on the dry pastures were rare, 20% were species with a high habitat specificity. In contrast, in larch forests

7% of the identified species could be considered as rare and 10% as specialist species; hay meadows harbored 4% rare and 4% specialist species.

Of all species found in the hay meadows, 39 (or 24%) were also recorded from the dry pasture

(Appendix A4), mostly spiders (11 species) and bugs (10 species). One spider was a specialist species. The larch forests shared 58 species with the dry pastures, predominantly vascular plants

(18 species), oribatid mites (14 species), and spiders (13 species). Five of these species were rare, seven were specialist species. Hay meadows and larch forests shared 27 species, again mostly spiders (9 species) and vascular plants (6 species). Only 18 species were found in all three habitat types, these were mostly rove beetles (5 species), vascular plants (4 species), and spiders (4 species). One of these rove beetles was a rare specialist (see data repository).

Of the 39 species shared between dry pastures and hay meadows, 21 (54%) had a high light demand (6 of them are vascular plants) and were therefore specified as probable pasture species.

Likewise, of the 58 species shared between dry pastures and larch forests, 20 (35%) had a high light demand (12 of them were vascular plants). The fourth-corner analysis (Fig. 4) demonstrated a high light demand/preference of rare and specialist species, while exhibiting a negative association with high humidity.

3.2 Species trends

The overall trend reflected by all species regardless of their taxonomic affiliation showed a significant decline of rare and specialist species after both management intensification and abandonment (Fig. 2). Broken down to the single taxonomic groups, this trend was solely depicted by spiders (Fig. 3). Most taxonomic groups provided only partial information on the overall trend: a significant decline of both rare and specialist species after intensification was shown by vascular plants, spiders, grasshoppers, bugs, and rove beetles; a significant decline after abandonment was reflected by spiders, ants, and birds. Butterflies supported the trends only for specialist species, some taxonomic groups even followed contrary trends (bryophytes, lichens, ants) or showed no trends at all (oribatid mites, ground beetles).

5 Discussion

We report on a multi-taxon study exploring differences in species diversity and composition on intensified grassland, grazed dry grassland and abandoned pastures covered by larch forest.

By assigning species traits for rarity and habitat specificity, we were able to evaluate the conservation values of the investigated land-use types. Intensively used hay meadows and successional larch forests harbored less rare and specialist species than dry pastures. Based on this, we argue that the intensification and abandonment of low-input dry pastures lead to a trivialization of flora and fauna, even though the overall species diversity in terms of species numbers may not be affected in the same way. In addition, our multi-taxon study confirmed previously reported reasoning (e.g. Zulka et al. 2014, Turtureanu et al. 2014, Simons et al. 2016) that conservation practices should not be deduced from single-species patterns.

5.1 Alpha diversity

Our findings suggest a strong decline of species richness after management intensification but no decline after abandonment, at least during the early succession stages. Due to their uniform treatment (evened surface bare of stones allows for mechanical handling, regular fertilizing and cuts), intensively used hay meadows are poor in micro-habitats, thus providing favorable habitats only for few plant and animal species (Ellenberg & Leuschner 2010, Marini et al. 2008). No bryophytes and lichens were recorded from these sites, since frequent disturbances and fertilization may hamper the establishment of these taxa (Meinunger 1992, Boch et al. 2016).

Early cuttings also prevent birds from using hay meadows as breeding habitat (Müller et al. 2005,

Peer & Frühauf 2009). One exception to the decrease in species richness were ground beetles: their species diversity was much higher in the intensively used hay meadows than in the other two habitat types. Such a positive correlation between management intensity of montane grassland (mowing and fertilizing) and the presence of ground beetles was already shown for the

Swiss Alps by Grandchamp et al. (2005), who explained this fact with the still low enough management intensity in Alpine regions to not reduce species richness. The otherwise lower species numbers in high-input grasslands compared with low-input dry pastures are in line with Newbold et al. (2015), who showed that intensification of pastures lead to a strong decline of biodiversity.

The habitat with the highest overall number of species were, however, the larch forests. They are the most heterogeneous habitats with the lowest number of within-habitat shared species

(i.e. species that can be found in all three replicate sites), and the ordination analysis showed the highest distance between replicate sites (Appendix A1). Larch forests were, due to diverging points of abandonment, rather heterogeneous with either open, pasture-like areas or forest-like shadow-adapted vegetation (mainly with the tor grass Brachypodium rupestre). The presence of many species of open, light habitats (as expressed by a high light preference) showed that dry pasture species had not yet fully disappeared. They seem to survive for a relatively long time and among them we found rare and specialist species usually associated with dry grasslands, for example the bug species Plinthisus pusillus or the western green lizard Lacerta bilineata.

However, species of open habitats will not find suitable habitats in dense forests and will finally disappear (e.g. Pornaro et al. 2013). On the other hand, we already found many species which depend on trees, e.g. many epiphytic lichen and moss species with species specialized in inhabiting specific parts of the larch trees, as well as several tree-breeding bird species.

All three habitat types accommodated many species that were unique to them, making the three habitats important elements for gamma diversity, i.e. the overall landscape biodiversity

(Gámez-Virués et al. 2015). Additionally, synergistic effects as a consequence of high landscape heterogeneity might exist (Fahrig et al. 2011), since predominantly vertebrate, but also some groups (e.g. butterflies) depend on habitat mosaics to cover their exigencies (Law &

Dickman 1998, Slamova et al. 2013).

5.2 Rarity and habitat specificity: losing the special, maintaining the trivial

Dry pastures had the highest share in rare and specialist species. This was true for almost all taxonomic groups and most pronounced in invertebrate groups such as spiders and grasshoppers. The statistically significant overall decrease in rare and specialist numbers was less evident for abandonment than for management intensification. Losses of rare and specialist species and a thus following biotic homogenization have been reported by Clavel et al. (2011),

Fartmann et al. (2012), Simons et al. (2016) and Noordwijk et al. (2017). Reasons for this are manifold: Land-use intensification may act as an environmental filter, leaving smaller and less specialist species, which are better adapted to frequent disturbances such as mowing (Simons et al. 2016). A further important factor affecting species occurrence is local microclimate. A large share of the organisms of the dry pastures possessed a high habitat specificity, and were limited to habitats with bright and dry conditions (Fig. 4). Such a direct dependence on microclimate conditions has been verified for example for butterflies (Bennett et al. 2014) and spiders (Lyons et al. 2018). Only bryophytes and lichens followed a clear opposite tendency: the proportion of rare and specialized species increases with the emergence of trees, providing additional habitats for these taxa (Boch et al. 2016).

At the landscape level, the loss of valuable habitats such as dry pastures leads to a decrease of rare and specialist species, i.e. to trivialization. Stehlik et al. (2007) reported a decrease of specialist species on a larger scale, i.e. a municipality, and connected this change with deterioration and loss of habitats, partly through direct destruction, partly through modification and intensification of agriculture. Also habitat fragmentation and the resulting loss of connectivity caused by land-use changes may lead to the loss of specialists (Zulka et al. 2014).

5.3 Implications for conservation

We conclude that both, abandonment and intensification, have negative impacts on biodiversity since they lead to a trivialization of flora and fauna. We confirm the high conservation value of dry pasture habitats in the Central Alps and in Central Europe and corroborate the priority in conserving the habitat “subpannonic steppe grasslands” (6240*) of the Habitats

Directive. Due to the recent trends in agricultural management many low-input grassland types, which are not in the focus of direct conservation measures, are prone to disappear in the near future.

In contrast, both intensively used hay meadows and successional larch forests do not justify specific conservation measures due to their rather common species composition. However, caution is advised since hay meadows as well as larch forests may encompass specific habitat types which are given a high conservation value (i.e. extensive low-input hay meadows as

Directive Habitat 6520 and subalpine Larch-Scots pine forests as Directive Habitat 9440) and an accurate addressing of the habitat type is key to optimize conservation efforts.

When planning conservation studies usually the question arises which taxonomic group(s) to survey, in many cases existing taxonomic expertise, easy-to-use methods or even financial restrictions impair the selection. In our study most taxonomic groups reflect only partially the overall trend or even showed an opposite tendency. The global trend was best reflected by spiders, followed by vascular plants, while it was controverted by bryophytes and lichens. Pedley et al. (2013) attributed contrasting responses of specialists from different taxonomic groups to differences in dispersal abilities, thus limiting their ability to find suitable new habitats. The interrelated responses of taxonomic groups to disturbances and countermeasures might also lead to situation-dependent patterns (Simons et al. 2016). We believe that the most suitable taxonomic group to reflect larger trends might vary with the habitats studied, pointing to a habitat bias. Therefore, we recommend the use of more than one taxonomic group, as have others done before, but also to give careful consideration to the selection of these groups.

Finally, our results suggest that the sole analysis of alpha diversity has only limited value since it predominantly reflects the inter- and intrasite heterogeneity of the single subplots. In our case the use of species traits for rarity and habitat specificity has proven useful to confirm the high conservation value of dry pastures as already proposed in the Habitats Directive.

Acknowledgements For their help in organizing field campaigns, we thank Thomas Wilhalm, Michele Torresani,

Alessandro Zandonai, Stefano Della Chiesa, the municipality of Mals/Malles and the village community of Matsch/Mazia. We acknowledge the Museum of Nature South Tyrol in

Bozen/Bolzano for data base consultation. This study was conducted at the LTSER platform

LTER_EU_IT_097 - Val Mazia/Matschertal, member of the national and international long term ecological research networks (LTER-Italy, LTER Europe and ILTER). The presented work was made possible by the funding of the Province of Bozen/Bolzano - South Tyrol for the LTSER platform.

Finally, we thank Rachele Carloni for drawing cliparts of plants and animals.

Figure captures

Figure 1: Aerial picture of the research area. H1-3…intensively used hay meadows, P1-3…dry pastures, L1-3…larch forests.

Figure 2: Percentage of rare (left) and specialist species (middle) and total species number (right) in intensively used hay meadows (H), dry pastures (P) and larch forests (L); letters indicate significant differences between habitats based on Chi²-tests at significance level p < 0.05.

Figure 3: Percentage of rare (left) and specialist species (middle) and total species number (right) for each taxonomic group. H…intensively used hay meadows, P…dry pastures and L…larch forests; letters indicate significant differences between habitats based on Chi²-tests at signicance level p < 0.05.

Figure 4: Results of the fourth-corner analysis evaluating environment-trait responses of species.

Environmental traits were the standard deviation of the surface temperature (surfT_sd), total phytomass (pyhto_tot), Ellenberg light indicator value (L), and Ellenberg humiditiy indicator value

(H). Species traits were rarity and habitat specificity. Red squares indicate positive, blue squares negative, and grey squares no associations.

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Figure 1 Aerial picture of the research area. H1–3…intensively used hay meadows, P1–3…dry pastures,

Figure 1: Percentage of rare (left) and specialist species (middle) and total species number (right) in intensively used hay meadows (H), dry pastures (P) and larch forests (L); letters indicate significant differences between habitats based on Chi²-tests at significance level p < 0.05. b b ab a a a vascular plants

b

a a b bryophytes

b

a a a lichens

b b a a a a spiders and harvestmen

a a a oribatid mites

b

b a

a ab a grasshoppers

Figure 2Percentage of rare (left) and specialist species (middle) and total species number (right) for each taxonomic group. H…intensively used hay meadows, P…dry pastures and L…larch forests; letters indicate significant differences between habitats based on χ2-tests at signicance level p < 0.05 b ab ab b a a bugs a

a a a ground beetles

b

ab b ab

a a rove beetles

a b b

c a a ants

b

a a butterflies

a

a

b b birds

L1–3…larch forests

Figure 3> Results of the fourth-corner analysis evaluating environment-trait responses of species. Environmental traits were the standard deviation of the surface temperature (surfT_sd), total phytomass (pyhto_tot), Ellenberg light indicator value (L), and Ellenberg humiditiy indicator value (H). Species traits were rarity and habitat specificity. Red squares indicate positive, blue squares negative, and grey squares no associations. Appendix:

Appendix A1: Community composition based on detrended correspondence analysis (DCA). Appendix A2: Climate and site parameters. surfT_mean … mean surface temperature [°C], surfT_sd … standard deviation of the surface temperature [°C], TempDiff … temperature difference between surface and air [°C], TempDiff_sd … standard deviation of the temperature difference between surface and air [°C], CN … mean C-to-N ratio of the upper soil (15 cm), pH … mean pH value of the upper soil (15 cm), SOM … mean soil organic matter content of the upper soil (15 cm) [%], veg_height … mean height of vegetation [cm], phyto_tot … total phytomass [g], herbs … herb phytomass [g], legumes … legume phytomass [g], grasses … grass phytomass [g], necromass … biomass of necromass [g], NDVI … normalized difference vegetation index, L … Ellenberg Indicator value (EIV) for light, H … EIV for humidity, T … EIV for temperature, N … EIV for nitrogen, H1-3 … intensively used hay meadows, P1-3 … dry pastures and L1-3 … larch forests

temperature parameters soil parameters phytomass parameters Ellenberg Indicator Values site surfT_mean surfT_sd TempDiff TempDiff_sd CN pH SOM veg_height phyto_tot herbs legumes grasses necromass NDVI L H T N H1 23.68 1.32 4.00 1.35 10.18 5.57 15.58 50.6 559.9 101.4 66.5 314.1 77.9 0.81 3.60 3.00 3.20 3.50 H2 21.82 1.17 0.73 1.15 10.68 5.95 12.45 48.9 653.2 115.9 2.2 377.0 158.2 0.86 3.70 2.80 3.20 3.60 H3 23.94 1.64 3.56 1.62 11.19 5.94 14.21 48.6 515.3 166.7 35.2 209.9 103.4 0.82 3.74 2.83 3.22 3.52 P1 32.93 4.39 13.24 4.38 13.83 5.45 5.7 31.8 268.1 11.4 36.6 74.6 145.6 0.73 4.03 1.69 3.80 2.06 P2 32.09 3.99 13.04 3.97 12.5 5.36 8.86 27.8 318.9 143.0 0.5 96.0 79.4 0.74 4.09 1.77 3.43 2.23 P3 30.31 3.88 9.55 3.83 13.39 5.32 8.24 32.0 456.1 62.4 1.9 349.7 26.0 0.70 4.07 2.15 3.16 2.48 L1 18.09 1.12 0.00 1.09 18.65 4.87 11.01 227.9 451.8 19.1 0.0 125.7 296.6 0.73 3.50 2.57 3.27 2.98 L2 24.21 5.48 3.50 5.17 17.76 4.97 8.9 232.0 292.2 2.7 0.0 131.0 115.2 0.74 3.54 2.35 3.37 2.69 L3 19.55 2.18 -0.79 2.62 17.48 5.12 11.31 274.4 341.1 50.3 0.0 65.9 44.0 0.79 3.16 3.02 3.31 3.42 1 Appendix A3a: Literature and database sources for rarity, specificity, and light demand

Organism Source rarity Source habitat specificity Light demand/ group preference Vascular plants Wilhalm & Hilpold given by E. Tasser, using a Ellenberg et al. (1992) (2006) phytosociological database Bryophytes Database Webpage Swissbryophytes From Urmi (2010), cited in Naturmuseum www.swissbryophytes.ch and Webpage Swissbryophytes using further habitat www.swissbryophytes.ch descriptions Lichens http://dryades.units.i http://dryades.units.it/italic/ http://dryades.units.it/italic t/italic/; pers. comm. / J. Nascimbene Earthworms www.florafauna.it, personal experience of personal experience of personal experience J. Seeber; Christian and Zicsi J. Seeber of J. Seeber (1999) Millipedes Pedroli-Christen Pedroli-Christen (1993), Pedroli-Christen (1993), (1993), personal experience of J. personal experience of J. personal experience Seeber Seeber of J. Seeber Spiders www.florafauna.it pers. comm. A. Rief pers. comm. A. Rief and pers. comm. A. Rief Oribatid mites pers. comm H. Schatz Schatz (1983), Weigmann, Schatz (1983), Weigmann, (2006), pers. comm. H. Schatz (2006), pers. comm. H. Schatz Grasshoppers Hilpold et al. 2017) www.orthoptera.ch personal www.orthoptera.ch experience A. Hilpold personal experience A. Hilpold Bugs www.florafauna.it using habitat descriptions in using habitat descriptions and Tamanini (1982) Wagner (1952), Wagner in and Heiss & Hellrigl (1967), Wagner (1967), Wagner (1952), Wagner, (1996), personal Tamanini (1982) (1966), Wagner (1967), experience of A. and personal experience A. Tamanini (1982) Hilpold Hilpold and personal experience A. Hilpold Beetles Database habitat descriptions from habitat descriptions from Naturmuseum “Die Käfer Mitteleuropas”, “Die Käfer Mitteleuropas”, complete list in complete list in http://www.springer.com/ser http://www.springer.com/s ies/7983 eries/7983 Ants Pers. comm. H. C. Pers. comm. H. C. Wagner Pers. comm. H. C. Wagner Wagner and Seifert and Seifert (2007) and Seifert (2007) (2007) Butterflies www.florafauna.it, Huemer (2004), Paolucci Huemer (2004), Paolucci Huemer (2004) (2013) (2013) Birds database Pers. comm. T. Wilhalm and Deduced from breeding Naturmuseum South Niederfriniger et al. (1996), habitat preferences in Tyrol, Niederfriniger Niederfriniger et al. (1996) et al. (1996) Reptiles and www.florafauna.it, personal experience of the personal experience of the amphibians personal experience authors authors of the authors 2 3 Appendix A3b: Experts and publications

Organism Expert(s) Publication group Vascular plants S. Wallnöfer, V. Fontana, G. Niedrist, A. Rief et al. (2017) Hilpold Bryophytes A. Schäfer-Verwimp, P. Mair, T. Kiebacher, S. Mair et al. (2017) Stix Lichens J. Nascimbene Nascimbene, in prep. Earthworms M. Steinwandter, J. Seeber Steinwandter & Seeber (2017) Millipedes M. Steinwandter, J. Seeber Steinwandter & Seeber (2017) Spiders S. Ballini, A. Rief Rief & Ballini (2017) Oribatid mites H. Schatz Schatz (2017) Grasshoppers P. Kranebitter, A. Hilpold Kranebitter & Hilpold (2017) Bugs T. Frieß, A. Hilpold Frieß & Hilpold (2017) Beetles W. Paill, I. Schatz, M. Steinwandter, A. Zanetti Schatz & Zanetti (2017), Steinwandter & Seeber (2017) Ants H. Wagner, F. Glaser, E. Guariento Wagner & Glaser (2017) Butterflies A. Hilpold, B. Stoinschek, E. Guariento Hilpold & Stoinschek (2017) Birds E. Gasser Rief et al. (2017) Reptiles and I. Plasinger, A. Rottensteiner, S. Barbacetto, A. Plasinger et al. (2017) and this publication amphibians Hilpold 4

5

6

7 Appendix A4: Total number of species, number of rare and specialist species, number of shared

8 species between habitats. H…intensively used hay meadows, P…dry pastures and L…larch forests

total species number rare species specialist species number of shared species taxon H P L total H P L H P L HP PL HL HPL vascular plants 32 70 77 140 0 7 3 1 13 10 7 18 6 4 mosses 0 12 21 30 0 1 0 0 1 6 3 lichens 0 9 28 37 0 0 1 0 0 5 spiders 36 71 64 130 2 20 5 2 15 4 11 13 9 4 oribatid mites 13 29 70 87 1 1 3 0 0 0 1 14 4 3 grasshoppers 7 9 1 16 0 1 0 1 4 0 1 bugs 25 42 5 58 1 5 1 1 13 1 10 2 1 carabids 16 3 5 20 1 0 0 0 0 0 1 3 staphylinids 16 18 38 53 1 3 6 1 4 7 4 5 5 ants 1 7 7 15 1 4 0 0 5 0 butterflies 3 7 2 8 0 0 0 0 1 0 2 1 birds 0 8 13 19 0 3 0 0 2 0 2 9 Total 163 290 345 638 7 47 23 6 58 33 39 58 27 18 10 11 References (Appendix)

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75 76 Table A2: List of shared total, rare, and specialist species. H…hay meadows, P…dry pastuers, L…larch

77 forests.

taxonomical family light demand/ humidity H P L group preference HP total shared species Aculepeira ceropegia Arachnida Araneidae 0.25 0.5 1 2 0 Alopecosa trabalis Arachnida Lycosidae 0.5 0.25 3 3 0 Amara aenea Coleoptera Carabidae 0.25 0.25 1 2 0 Aporrectodea caliginosa Oligochaeta Lumbricidae 0.5 0.5 1 1 0 Araeoncus humilis Arachnida Linyphiidae 0.25 0.5 3 1 0 Araniella cucurbitina Arachnida Araneidae 0.5 0.5 3 3 0 Arenaria serpyllifolia Tracheophyta Caryophyllaceae 0.25 0.25 2 1 0 Cerastium holosteoides Tracheophyta Caryophyllaceae 0.25 0.5 3 1 0 Chlamydatus pullus Heteroptera Miridae 0.25 0.25 1 3 0 Chorthippus parallelus Orthoptera Acrididae 0.25 0.5 3 2 0 Coccinella septempunctata Coleoptera Coccinellidae 0.5 0.5 1 1 0 Coenonympha pamphilus Lepidoptera Nymphalidae 0.25 0.5 2 3 0 Derephysia foliacea Heteroptera Tingidae 0.5 0.5 1 1 0 Diplocephalus alpinus Arachnida Linyphiidae 0.5 0.75 1 1 0 Dolycoris baccarum Heteroptera Pentatomidae 0.5 0.5 1 2 0 Erigone atra Arachnida Linyphiidae 0.25 0.5 3 1 0 Erigone dentipalpis Arachnida Linyphiidae 0.25 0.5 3 1 0 Eurygaster maura Heteroptera Scutelleridae 0.25 0.25 1 1 0 Lolium perenne Tracheophyta Poaceae 0.25 0.5 3 1 0 Lygus gemellatus Heteroptera Miridae 0.25 0.25 2 1 0 Lygus pratensis Heteroptera Miridae 0.5 0.5 2 1 0 Lygus rugulipennis Heteroptera Miridae 0.5 0.5 3 1 0 Mermessus trilobatus Arachnida Linyphiidae 0.5 0.5 2 3 0 Oppiella uliginosa Oribatida Oppiidae 0.75 0.5 1 1 0 Pachygnatha degeeri Arachnida Tetragnathidae 0.25 0.5 3 1 0 Philonthus albipes Coleoptera Staphylinidae 0.25 0.5 1 1 0 Philonthus carbonarius Coleoptera Staphylinidae 0.5 0.5 1 1 0 Philonthus cognatus Coleoptera Staphylinidae 0.5 0.5 1 2 0 Plantago lanceolata Tracheophyta Plantaginaceae 0.25 0.5 2 1 0 Plebejus argus Lepidoptera Lycaenidae 0.25 0.25 1 3 0 Rhyparochromus pini Heteroptera Rhyparochromidae 0.25 0.25 3 1 0 Silene vulgaris Tracheophyta Caryophyllaceae 0.25 0.5 3 1 0 Syromastes rhombeus Heteroptera Coreidae 0.5 0.5 1 1 0 Tachinus rufipes Coleoptera Staphylinidae 0.5 0.5 2 1 0 Tiso vagans Arachnida Linyphiidae 0.5 0.5 3 2 0 Trapezonotus arenarius Heteroptera Rhyparochromidae 0.5 0.5 2 1 0 Trifolium repens Tracheophyta Fabaceae 0.25 0.5 3 1 0 Veronica arvensis Tracheophyta Plantaginaceae 0.5 0.5 3 1 0 Xysticus ninnii Arachnida Thomisidae 0.25 0.25 1 2 0 PL total shared species Adoristes ovatus Oribatida Liacaridae 0.75 0.5 0 1 1 Araneus diadematus Arachnida Araneidae 0.5 0.5 0 3 1 Artemisia absinthium Tracheophyta Asteraceae 0.25 0.25 0 1 1 Avenula praeusta Tracheophyta Poaceae 0.25 0.25 0 1 2 Brachypodium rupestre Tracheophyta Poaceae 0.25 0.5 0 3 3 Bromus erectus Tracheophyta Poaceae 0.25 0.25 0 1 2 Bryum capillare agg. Bryophytes Bryaceae 0.5 0.5 0 1 1 Carabodes labyrinthicus Oribatida Carabodidae 0.5 0.5 0 1 3 Carex supina Tracheophyta Cyperaceae 0.25 0.25 0 1 1 Carlina vulgaris Tracheophyta Asteraceae 0.25 0.25 0 1 1 Cerastium arvense Tracheophyta Caryophyllaceae 0.25 0.25 0 2 2 Ceratozetes gracilis Oribatida Ceratozetidae 0.5 0.5 0 1 1 Diapterobates humeralis Oribatida Humerobatidae 0.75 0.5 0 1 1 Drassodes pubescens Arachnida Gnaphosidae 0.5 0.25 0 2 1 Eniochthonius minutissimus Oribatida Eniochthoniidae 0.5 0.5 0 1 2 Eueremaeus oblongus Oribatida Eremaeidae 0.75 0.25 0 2 1 Eueremaeus valkanovi Oribatida Eremaeidae 0.5 0.25 0 2 1 Euphorbia cyparissias Tracheophyta Euphorbiaceae 0.5 0.25 0 2 1 Festuca rupicola Tracheophyta Poaceae 0.5 0.25 0 2 1 Festuca valesiaca Tracheophyta Poaceae 0.25 0.25 0 3 2 Fringilla coelebs Aves Fringillidae 0.5 0.5 0 1 1 Galium pusillum agg. Tracheophyta Rubiaceae 0.25 0.25 0 3 2 Galium verum Tracheophyta Rubiaceae 0.25 0.5 0 3 1 Haplodrassus signifer Arachnida Gnaphosidae 0.5 0.25 0 2 3 Hemileius initialis Oribatida Scheloribatidae 0.5 0.5 0 1 3 Hypnum cupressiforme Bryophytes Hypnaceae 0.5 0.5 0 1 2 Improphantes nitidus Arachnida Linyphiidae 0.5 0.5 0 2 2 Kunstidamaeus tecticola Oribatida Damaeidae 0.5 0.25 0 1 1 Lacerta bilineata Reptilia Lacertidae 0.5 0.25 0 1 1 Linyphia hortensis Arachnida Linyphiidae 0.75 0.5 0 2 1 Lotus corniculatus Tracheophyta Fabaceae 0.25 0.5 0 3 2 Luzula campestris agg. Tracheophyta Juncaceae 0.25 0.5 0 2 1 Maso sundevalli Arachnida Linyphiidae 0.75 0.5 0 1 1 Minyriolus pusillus Arachnida Linyphiidae 0.75 0.25 0 1 1 Ontholestes haroldi Coleoptera Staphylinidae 0.25 0.5 0 1 1 Oribatula interrupta Oribatida Oribatulidae 0.5 0.5 0 1 1 Oribatula tibialis Oribatida Oribatulidae 0.5 0.5 0 3 3 Oxypoda islandica Coleoptera Staphylinidae 0.75 0.75 0 1 1 Parachipteria punctata Oribatida Achipteriidae 0.5 0.5 0 1 1 Pardosa bifasciata Arachnida Lycosidae 0.25 0.25 0 3 1 Pardosa blanda Arachnida Lycosidae 0.25 0.5 0 3 2 Pergalumna formicaria Oribatida Galumnidae 0.5 0.5 0 1 1 Philodromus aureolus Arachnida Philodromidae 0.5 0.5 0 1 1 Phylloneta sisyphia Arachnida Theridiidae 0.5 0.5 0 3 3 Plinthisus pusillus Heteroptera Lygaeidae 0.25 0.25 0 2 1 Poa angustifolia Tracheophyta Poaceae 0.25 0.25 0 2 2 Quedius nemoralis Coleoptera Staphylinidae 0.5 0.25 0 1 1 Sedum rupestre agg. Tracheophyta Crassulaceae 0.25 0.25 0 1 1 Sepedophilus nigripennis Coleoptera Staphylinidae 0.25 0.25 0 1 1 Sepedophilus testaceus Coleoptera Staphylinidae 0.75 0.5 0 1 1 Stenodema laevigata Heteroptera Miridae 0.5 0.5 0 1 1 Syntrichia ruralis Bryophytes Pottiaceae 0.5 0.25 0 3 2 Theridion mystaceum Arachnida Theridiidae 0.75 0.5 0 1 2 Thymus praecox agg. Tracheophyta Lamiaceae 0.25 0.25 0 3 2 Trichoribates berlesei Oribatida Ceratozetidae 0.5 0.5 0 1 1 Turdus merula Aves Turdidae 0.75 0.5 0 1 1 Veronica chamaedrys Tracheophyta Plantaginaceae 0.5 0.5 0 1 3 Xysticus audax Arachnida Thomisidae 0.5 0.5 0 1 3 ML total shared species Achipteria coleoptrata Oribatida Achipteriidae 0.5 0.5 1 0 2 Amphicyllis globus Coleoptera Leiodidae 0.625 0.5 1 0 1 Arrhenatherum elatius Tracheophyta Poaceae 0.5 0.5 2 0 2 Calathus fuscipes Coleoptera Carabidae 0.25 0.25 1 0 1 Calathus melanocephalus Coleoptera Carabidae 0.25 0.25 1 0 1 Caleremaeus monilipes Oribatida Caleremaeidae 0.75 0.5 1 0 2 Cymbaeremaeus cymba Oribatida Cymbaeremaeidae 0.5 0.5 1 0 2 Dactylis glomerata Tracheophyta Poaceae 0.25 0.5 3 0 3 Dendrobaena octaedra Oligochaeta Lumbricidae 0.5 0.5 2 0 2 Heracleum sphondylium Tracheophyta Apiaceae 0.5 0.5 2 0 1 Micrargus subaequalis Arachnida Linyphiidae 0.25 0.25 1 0 2 Myosotis sylvatica Tracheophyta Boraginaceae 0.5 0.5 1 0 1 Neoribates aurantiacus Oribatida Parakalummidae 0.75 0.75 2 0 3 Nicrophorus vespillo Coleoptera Silphidae 0.625 0.5 2 0 1 Ommatoiulus sabulosus Myriapoda Julidae 0.5 0.25 1 0 2 Pardosa lugubris Arachnida Lycosidae 0.5 0.5 2 0 3 Pisaura mirabilis Arachnida Pisauridae 0.5 0.25 1 0 1 Poa pratensis Tracheophyta Poaceae 0.25 0.5 3 0 2 Pocadicnemis pumila Arachnida Linyphiidae 0.75 0.5 1 0 3 Pterostichus strenuus Coleoptera Carabidae 0.75 0.75 3 0 1 Ranunculus repens Tracheophyta Ranunculaceae 0.5 0.5 2 0 2 Sciodrepoides watsoni Coleoptera Catopidae 0.625 0.5 1 0 1 Tenuiphantes mengei Arachnida Linyphiidae 0.5 0.5 1 0 3 Tenuiphantes tenebricola Arachnida Linyphiidae 0.75 0.5 1 0 1 Tenuiphantes tenuis Arachnida Linyphiidae 0.5 0.5 3 0 1 Xysticus erraticus Arachnida Thomisidae 0.5 0.25 1 0 1 Zora spinimana Arachnida Miturgidae 0.75 0.5 1 0 1 HPL total shared species Achillea millefolium agg. Tracheophyta Asteraceae 0.25 0.25 3 1 2 Agyneta affinis Arachnida Linyphiidae 0.5 0.5 2 1 1 Agyneta rurestris Arachnida Linyphiidae 0.25 0.5 3 2 2 Atheta amplicollis / fungi Coleoptera Staphylinidae 0.75 0.75 3 2 3 Chlamydatus pulicarius Heteroptera Miridae 0.25 0.25 1 3 1 Enoplognatha ovata Arachnida Theridiidae 0.5 0.5 1 2 3 Erebia alberganus Lepidoptera Nymphalidae 0.5 0.5 1 1 1 Eupelops torulosus Oribatida Phenopelopidae 0.75 0.5 2 1 2 Liogluta alpestris Coleoptera Staphylinidae 0.5 0.5 1 2 1 Pardosa palustris Arachnida Lycosidae 0.5 0.5 3 3 1 Philonthus decorus Coleoptera Staphylinidae 0.75 0.75 1 1 3 Ranunculus bulbosus Tracheophyta Ranunculaceae 0.25 0.25 1 2 1 Scheloribates laevigatus Oribatida Scheloribatidae 0.5 0.75 2 3 1 Tachyporus abdominalis Coleoptera Staphylinidae 0.5 0.75 1 1 1 Tachyporus chrysomelinus Coleoptera Staphylinidae 0.5 0.5 3 1 1 Taraxacum officinale agg. Tracheophyta Asteraceae 0.25 0.5 3 2 3 Tectocepheus sarekensis Oribatida Tectocepheidae 0.5 0.5 3 3 1 Trifolium pratense Tracheophyta Fabaceae 0.25 0.5 3 1 3 PL shared rare species Carex supina Tracheophyta Cyperaceae See above See above 0 1 1 Kunstidamaeus tecticola Oribatida Damaeidae See above See above 0 1 1 Oxypoda islandica Coleoptera Staphylinidae See above See above 0 1 1 Plinthisus pusillus Heteroptera Lygaeidae See above See above 0 2 1 Sepedophilus nigripennis Coleoptera Staphylinidae See above See above 0 1 1 HPL shared rare and specialist species Liogluta alpestris Coleoptera Staphylinidae See above See above 1 2 1 HP shared specialists species Xysticus ninnii Arachnida Thomisidae See above See above 1 2 0 PL shared specialists species Artemisia absinthium Tracheophyta Asteraceae See above See above 0 1 1 Avenula praeusta Tracheophyta Poaceae See above See above 0 1 2 Carex supina Tracheophyta Cyperaceae See above See above 0 1 1 Hypnum cupressiforme Bryophytes Hypnaceae See above See above 0 1 2 Improphantes nitidus Arachnida Linyphiidae See above See above 0 2 2 Oxypoda islandica Coleoptera Staphylinidae See above See above 0 1 1 Sepedophilus nigripennis Coleoptera Staphylinidae See above See above 0 1 1 78

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