Shifts in ground-dwelling predator communities in response to changes in management intensity in Alpine meadows




Soil, Arachnida, Coleoptera, Formicidae, species diversity


Land-use changes, especially agricultural intensification has increased in the last decades leading to a decrease in biodiversity. In Europe, grasslands have been influenced by humans for centuries and millennia and management intensity has increased since the 20th century. In this small-case study, we investigate how management intensity affects ground-dwelling predators in montane hay meadows in South Tyrol, an Alpine region in Northern Italy, using the pitfall trap method.
As expected, species composition differed significantly when comparing the predator communities of extensive and intensive meadows, with the former supporting a higher predator species richness, and the latter showing higher proportions of frequent and euryoecious species. Regarding their activity densities and Shannon diversity, we did not find clear differences. Investigating selected ecological species traits, we found differences for moisture requirements and ecological tolerance between the two management types, with xerophilous species being more abundant in the extensive meadows, and stenoecious species more abundant in intensive meadows.
In this study, we found management intensity of montane grasslands to have a limited influence on the biodiversity patterns of ground-dwelling predators. However, individual predator groups showed clear reactions to the intensity of management (i.e., decrease or increase in activity density, species richness and Shannon diversity). We conclude that a intensive management of grasslands in combination with local habitat specifics does not lead to a homogenisation of the predatory arthropod community like it was found in other studies. Our study contributes to a better understanding of scarcely investigated predator communities and their diversity in differently managed Alpine grasslands.


Andrey, A., J. Y. Humbert & R. Arlettaz (2016): Functional response of leaf-and planthoppers to modern fertilisation and irrigation of hay meadows. – Basic and Applied Ecology 17(7): 627–637 [].

Assing, V. & M. Schülke (2012): Die Käfer Mitteleuropas, Bd. 4: Staphylinidae (exklusive Aleocharinae, Pselaphinae und Scydmaeninae). – Spektrum Akademischer Verlag, Heidelberg: 560 pp.

Attwood, S. J, M. Maron, A. P. N. House & C. Zammit (2008): Do arthropod assemblages display globally consistent responses to intensified agricultural land use and management? – Global Ecology and Biogeography 17(5): 585–599 [].

Bates, D., M. Mächler, B. Bolker, & S. Walker (2015): Fitting Linear Mixed-Effects Models Using lme4. Journal of Statistical Software 67(1): 1–48 [].

Bell, James R, C. P. Wheater & W. R. Cullen (2001): The implications of grassland and heathland management for the conservation of spider communities: A review. – Journal of Zoology 255(3): 377–387 [].

Billeter, R., J. Liira, D. Bailey, R. Bugter, P. Arens, I. Augenstein, S. Aviron, J. Baudry, R. Bukacek, F. Burel & et al. (2008): Indicators for biodiversity in agricultural landscapes: A Pan-European Study. – Journal of Applied Ecology 45(1): 141–50 [].

Birkhofer, K., M. M. Gossner, T. Diekötter, C. Drees, O. Ferlian, M. Maraun, S. Scheu, W. W. Weisser, V. Wolters, S. Wurst & et al. (2017): Land-use type and intensity differentially filter traits in above- and below-ground arthropod communities. – Journal of Animal Ecology 86(3): 511–20 [].

Birkhofer, K., T. Diekötter, C. Meub, K. Stötzel & V. Wolters (2015): Optimizing arthropod predator conservation in permanent grasslands by considering diversity components beyond species richness. – Agriculture, Ecosystems & Environment 211: 65–72. [].

Blick, T., O.-D. Finch, K. H. Harms,; J. Kiechle; K.-H. Kielhorn; M. Kreuels, A. Malten, D. Martin, C. Muster, D. Nährig, et al. (2016): Rote Liste und Gesamtartenliste der Spinnen (Arachnida: Araneae) Deutschlands. – In: Gruttke, H.; Balzer, S.; Binot-Hafke, M.; Haupt, H.; Hofbauer, N.; Ludwig, G.; Matzke-Hajek, G. & Ries, M. (eds): Rote Liste gefährdeter Tiere, Pflanzen und Pilze Deutschlands, Band 4: Wirbellose Tiere (Teil 2). – Münster (Landwirtschaftsverlag). – Naturschutz und Biologische Vielfalt 70 (4): 383–510.

Chao A, N. J. Gotelli, T. C. Hsieh, E.L. Sander, K. H., R. K. Colwell & A. M. Ellison (2014) Rarefaction and extrapolation with Hill numbers, a framework for sampling and estimation in species diversity studies – Ecological Monographs 84 (1):45–67 [].

Clarke, K. R., P. J. Somerfield & M. G. Chapman (2006): On resemblance measures for ecological studies, including taxonomic dissimilarities and a zero-adjusted Bray-Curtis coefficient for denuded assemblages. – Journal of Experimental Marine Biology and Ecology 330(1): 55–80 [].

Cole, L. J, M. L. Pollock, D. Robertson, J. P. Holland & D. I. McCracken (2006): Carabid (Coleoptera) Assemblages in the Scottish Uplands: The influence of sheep grazing on ecological structure. – Entomologica Fennica 17(3): 229–240 [].

Colla, F., A. Zanetti, E. Guariento, J. Plunger, J. & J. Seeber (2021): Xantholinus audrasi (Coiffait, 1956) (Coleoptera: Staphylinidae), new record for South Tyrol. – Gredleriana: 165–167. [].

De Jong, Y. M., M. Verbeek, V Michelsen, P. de Place Bjørn, W. Los, F. Steeman, N. Bailly, C. Basire, P. Chylarecki, E. Stloukal & et al. (2014): Fauna Europaea - all European animal species on the web. – Biodiversity Data Journal 2: e4034 [].

Dennis, P, M. R. Young & I. J. Gordon (1998): Distribution and abundance of small insects and arachnids in relation to structural heterogeneity of grazed, indigenous grasslands. – Ecological Entomology 23(3): 253–264 [].

Dennis, P., M. R. Young & C. Bentley (2001): The effects of varied grazing management on epigeal spiders, harvestmen and pseudoscorpions of Nardus Stricta grassland in upland Scotland. – Agriculture, Ecosystems & Environment 86(1): 39–57 [].

Di Giulio, M., P. J. Edwards & E. Meister (2001): Enhancing insect diversity in agricultural grasslands: the roles of management and landscape structure. – Journal of Applied Ecology 38(2): 310–19.

Dittrich, A. D. K. & A. J. Helden (2012): Experimental Sward Islets: The effect of dung and fertilisation on Hemiptera and Araneae. – Insect Conservation and Diversity 5(1): 46–56 [].

Eisenhauer, N., S. F. Bender, I. Calderón-Sanou, F. T. de Vries, J. J. Lembrechts, W. Thuiller, D. H. Wall, R. Zeiss, M. Bahram, R. Beugnon et al. (2022): Frontiers in Soil Ecology—Insights from the World Biodiversity Forum 2022. – Journal of Sustainable Agriculture and Environment: 1–17 [].

FloraFaunaSüdtirol (2022): FloraFaunaSüdtirol – Das Portal zur Verbreitung von Tier- und Pflanzenarten in Südtirol. – Naturmuseum Südtirol, Bozen, Italien [].

Fournier, B., F. Gillet, R. C. le Bayon, E. A. D. Mitchell & M. Moretti (2015): Functional responses of multitaxa communities to disturbance and stress gradients in a restored floodplain. – Journal of Applied Ecology 52(5): 1364–1373 [].

Gámez-Virués, S., D. J, Perović, M. M. Gossner, C. Börschig, N. Blüthgen, H. De Jong, N.K. Simons, A.M. Klein, J. Krauss, G. Maier et al. (2015): Landscape simplification filters species traits and drives biotic homogenization. – Nature Communications 6(1): 1–8 [].

Gapp, J. (1994): Rote Liste gefährdeter Tierarten Südtirols. Abteilung für Landschafts- und Naturschutz Autonome Provinz Bozen/Südtirol, Bozen: 420 pp.

Gilgado, J. D., H. P. Rusterholz, B. Braschler, S. Zimmermann, Y. Chittaro & B. Baur (2022): Six groups of ground-dwelling arthropods show different diversity responses along elevational gradients in the Swiss Alps. – PLOS One 17(7): e0271831 [].

Gobbi, M . & D. Fontaneto (2008): Biodiversity of Ground Beetles (Coleoptera: Carabidae) in different habitats of the Italian Po lowland. – Agriculture, Ecosystems and Environment 127(3–4): 273–276 [].

Gobbi, M., D. Fontaneto, N. Bragalanti, L. Pedrotti & V. Lencioni. (2015): Carabid Beetle (Coleoptera: Carabidae) richness and functional traits in relation to differently managed grasslands in the Alps. – Annales de La Societe Entomologique de France 51(1): 52–59 [].

Gossner, M. M., T. M. Lewinsohn, T. Kahl, F. Grassein, S. Boch, S, D. Prati, K. Birkhofer, S.C. Renner, J. Sikorski, T. Wubet (2016): Land-use intensification causes multitrophic homogenization of grassland communities. – Nature 540(7632): 266–269 [].

Grandchamp, A. C., A. Bergamini, S. Stofer, J. Niemelä, P. Duelli & C. Scheidegger. (2005): The influence of grassland management on ground beetles (Carabidae, Coleoptera) in Swiss montane meadows. – Agriculture, Ecosystems and Environment 110(3–4): 307–317 [].

Guariento, E., F. Colla, M. Steinwandter, J. Plunger, U. Tappeiner & J. Seeber (2020): Management intensification of hay meadows and fruit orchards alters soil macro-invertebrate communities differently. – Agronomy 10(6): 767 [].

Guerra, C. A., A. Heintz-Buschart, J. Sikorski, A. Chatzinotas, N. Guerrero-Ramírez, S. Cesarz, L. Beaumelle, M.C. Rillig, F.T. Fernando, M. Delgado-Baquerizo et al. (2020): Blind spots in global soil biodiversity and ecosystem function research. – Nature Communications 11(1): 1–13 [].

Guerra, C. A., M. Berdugo, D. J. Eldridge, N. Eisenhauer, B. K. Singh, H. Cui, S. Abades, F. D. Alfaro, A. R. Bamigboye & F. Bastida (2022): Global hotspots for soil nature conservation. – Nature 610(7933): 693–698 [].

Haddad, N. M., G. M. Crutsinger, K. Gross, J. Haarstad, J. M. H. Knops & D. Tilman (2009): Plant species loss decreases arthropod diversity and shifts trophic structure. – Ecology Letters 12(10): 1029–1039 [].

Hilpold, A., J. Seeber, V. Fontana, G. Niedrist, A. Rief, M. Steinwandter, E. Tasser & U. Tappeiner. (2018): Decline of rare and specialist species across multiple taxonomic groups after grassland intensification and abandonment. – Biodiversity and Conservation 27(14): 3729–3744 [].

Holzinger W., T. Friess, T., C. Komposch & W. Paill (2012): Tierökologische Bewertung von WF-Rotflächen ein und vier Jahre nach Einstieg in die WF-Maßnahme. – Ländlicher Raum - Online-Fachzeitschrift des Bundesministeriums für Land- und Forstwirtschaft, Umwelt und Wasserwirtschaft 2012 (02): 1–15.

Holland, J. M. (2002): The Agroecology of Carabid Beetles. Intercept Limited, Andover: 356 pp.

Humbert, J. Y., J. M. Dwyer, A. Andrey & R. Arlettaz (2016): Impacts of nitrogen addition on plant biodiversity in mountain grasslands depend on dose, application duration and climate: A systematic review. – Global Change Biology 22(1): 110–120 [].

Hunter, M. D. & P. W. Price (1992): Playing chutes and ladders: heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. – Ecology 73: 724–732 [].

Huston, M. & L. Gilbert (1996): Consumer Diversity and Secondary Production. In: Orians, G. H., R. Dirzo & J. H. Cushman (eds): Biodiversity and Ecosystem Processes in Tropical Forests. – Ecological Studies, vol 122. Springer, Berlin, Heidelberg: 33–47 [].

Kahlen, M. (2018): Die Käfer von Südtirol: Ein Kompendium. Naturmuseum Südtirol, Bozen: 602 pp.

Kleijn, D., F. Kohler, A. Báldi, P. Batáry, E. D. Concepción, Y. Clough, M. Díaz, D. Gabriel, A. Holzschuh, E. Knop & et al. (2009): On the relationship between farmland biodiversity and land-use intensity in Europe. – Proceedings of the Royal Society B: Biological Sciences 276(1658): 903–909 [].

Koch, K. (1989): Die Käfer Mitteleuropas: Ökologie. – Goecke & Evers, Krefeld. Band. 1: 440 pp.

Komposch, C. (2009): Rote Liste der Weberknechte (Opiliones) Österreichs. In: Zulka, K. P. (eds.): Rote Liste gefährdeter Tiere Österreichs. Checklisten, Gefährdungsanalysen, Handlungsbedarf. Teil 3: Flusskrebse, Köcherfliegen, Skorpione, Weberknechte, Zikaden. Grüne Reihe des Lebensministeriums (Gesamtherausgeberin Ruth Wallner) – Wien, Böhlau. Band 14/3: 397–483.

Komposch, C. (2022): Spinnentiere und Insekten – Artendiversität, Lebensräume und Bedeutung. In: Wiegele, E., M. Jungmeier & M. Schneider (eds): Handbuch Naturschutzfachkraft. Praktischer Naturschutz für Baustellen, Betriebsgelände und Infrastrukturen. – Fraunhofer IRB Verlag, Stuttgart: 487–514.

Kormann, U., V. Rösch, P. Batáry, T. Tscharntke, K. M. Orci, F. Samu, & C. Scherber (2015): Local and landscape management drive trait-mediated biodiversity of nine taxa on small grassland fragments. Diversity and Distributions 21: 1204–1217 [].

Kotze, D. J., P. Brandmayr, A. Casale, E. Dauffy-Richard, W. Dekoninck, M. J. Koivula, G. L. Lövei, D. Mossakowski, J. Noordijk, W. Paarmann et al. (2011): Forty years of carabid beetle research in Europe–from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation. – ZooKeys 100: 55–148 [].

LAFIS (2020): Amt für Landwirtschaftliche Informationssysteme. – Retrieved from url:

Larsson, S. G. (1939). Entwicklungstypen und Entwicklungszeiten der dänischen Carabiden, Entomologiske Meddelelser 20: 277–560.

Le Provost, G., I. Badenhausser, Y. le Bagousse-Pinguet, Y. Clough, L. Henckel, C. Violle, V. Bretagnolle, M. Roncoroni, P. Manning & N. Gross (2020): Land-use history impacts functional diversity across multiple trophic groups. – Proceedings of the National Academy of Sciences of the United States of America 117(3): 1573–1579 [].

Lessard‐Therrien, M., J. Y. Humbert & R. Arlettaz (2017): Experiment‐based recommendations for biodiversity‐friendly management of mountain hay meadows. – Applied Vegetation Science 20(3): 352–362 [].

Littlewood, N. A., A. J. A. Stewart & B. A. Woodcock (2012): Science into practice - how can fundamental science contribute to better management of grasslands for invertebrates? – Conservation and Diversity 5: 1–8 [].

Marini, L., P. Fontana, M. Scotton & S. Klimek (2008): Vascular plant and Orthoptera diversity in relation to grassland management and landscape composition in the European Alps. – Journal of Applied Ecology 45(1): 361–370 [].

Martens, J. (1978): Spinnentiere, Arachnida: Weberknechte, Opiliones. In: Senglaub, F., H. J. Hannemann & H. Schumann (eds.): Die Tierwelt Deutschland 64. – VEB Gustav Fischer Verlag, Jena: 464 pp.

Matson, P. A, W. J. Parton, A. G. Power & M. J. Swift (1997): Agricultural intensification and ecosystem properties. – Science 277(5325): 504–509 [].

Mathieu, J., A. C. Antunes, S. Barot, A. E. Bonato Asato, M. L. C. Bartz, G. G. Brown, I. Calderon-Sanou, T. Decaëns, S. J. Fonte, P. Ganault, et al. (2022): sOilFauna - a global synthesis effort on the drivers of soil macrofauna communities and functioning: Workshop Report. – Soil Organisms 94(2): 111–126 [].

Maxwell, S. L., R. Fuller, T. M. Brooks & J. E. Watson (2016): Biodiversity: The ravages of guns, nets, and bulldozers. – Nature 536: 143–145 [].

Meyer, S. T., L. Heuss, H. Feldhaar, W. W. Weisser & M. M. Gossner (2019): Land-use components, abundance of predatory arthropods, and vegetation height affect predation rates in grasslands. – Agriculture, Ecosystems & Environment 270: 84–92. [].

Müller-Motzfeld, G. (2004): Käfer Mitteleuropas, Bd. 2: Adephaga I: Carabidae. – Spektrum Akademischer Verlag, Heidelberg: 521 pp.

Nentwig W., T. Blick, D. Gloor, A. Hänggi & C. Kropf (2022): Spiders of Europe. Version 11.2022. Online at url: [].

Nickel, H. & J. P. Hildebrandt (2003): Auchenorrhyncha communities as indicators of disturbance in grasslands (Insecta, Hemiptera)—a case study from the Elbe flood plains (Northern Germany). – Agriculture, Ecosystems & Environment 98: 183–199 [].

Niedrist, G., E. Tasser, C. Lüth, J. Dalla Via & U. Tappeiner (2009): Plant diversity declines with recent land use changes in European Alps. – Plant Ecology 202 (2): 195–210 [].

Oksanen, J., G. L. Simpson F. G. Blanchet, R. Kindt, P. Legendre, P. R. Minchin, R. B. O‘Hara, P. Solymos, M. H. H. Stevens, E. Szoecs & et. al (2022): Vegan: Community ecology package. R package version 2.6-4 [Retrieved from url:].

Pekár, S, J. O. Wolff, Ľ. Černecká, K. Birkhofer, S. Mammola, E. C. Lowe, C. S. Fukushima, M. E. Herberstein, A. Kucura, B. A. Buzzatto, et al. (2021): The World Spider Trait database: a centralized global open repository for curated data on spider traits. – Database Vol. 2021 (2021): baab064 [].

Pfiffner, L. & H. Luka (2003): Effects of low-input farming systems on carabids and epigeal spiders–a paired farm approach. – Basic and Applied Ecology 4(2): 117–127 [].

Power, E. F., D. L. Kelly & J. C. Stout (2012): Organic farming and landscape structure: effects on insect-pollinated plant diversity in intensively managed grasslands. PLOS ONE 7(5): e38073 [].

Purtauf, T., J. Dauber & V. Wolters (2005): The response of carabids to landscape simplification differs between trophic groups. – Oecologia 142(3): 458–464.

R Core Team (2022): R: A language and environment for statistical computing. – R Foundation for Statistical Computing, Vienna, Austria. url:

Robinson, R. A. & W. J Sutherland (2002): Post-war changes in arable farming and biodiversity in Great Britain. – Journal of Applied Ecology 39(1): 157–176 [].

Ryder, C., J. M. R. Mc Donnell & M. Gormally (2005): Conservation implications of grazing practices on the plant and dipteran communities of a Turlough in Co. Mayo, Ireland. – Biodiversity & Conservation 14(1): 187–204 [].

Samu, F., K. D. Sunderland & C. Szinetar (1999): Scale-dependent dispersal and distribution patterns of spiders in agricultural systems: a review. – Journal of Arachnology 27: 325–332.

Schweiger, O., J. P. Maelfait, W. van Wingerden, F. Hendrickx, R. Billeter, M. Speelmans, I. Augenstein, B. Aukema, S. Aviron, D. Bailey, et al. (2005): Quantifying the impact of environmental factors on arthropod communities in agricultural landscapes across organizational levels and spatial scales. – Journal of Applied Ecology 42(6): 1129–1139 [].

Seeber, J., M. Steinwandter, E. Tasser, E. Guariento, T. Peham, J. Rüdisser, B.C. Schlick-Steiner, F. M. Steiner, U. Tappeiner & E. Meyer (2022): Distribution of soil macrofauna across different habitats in the Eastern European Alps. – Scientific Data 9: 632 [].

Seifert, B. (2018): The Ants of Central and North Europe. – lutra Verlags- und Vertriebsgesellschaft, Tauer: 408 pp.

Simons, N. K., M. M. Gossner, T. M. Lewinsohn, S. Boch, M. Lange, J. Müller, E. Pašalić, S.A. Socher, M. Türke, M. Fischer & et al. (2014): Resource-mediated indirect effects of grassland management on arthropod diversity. – PLOS ONE 9(9): e107033 [].

Simons, N. K., W. W. Weisser & M. M. Gossner (2016): Multi‐taxa approach shows consistent shifts in arthropod functional traits along grassland land‐use intensity gradient. – Ecology 97(3): 754–764.

Sjödin, N. E., J. Bengtsson & B. Ekbom (2008): The influence of grazing intensity and landscape composition on the diversity and abundance of flower-visiting insects. – Journal of Applied Ecology 45(3): 763–772.

Socher, S. A., D. Prati, S. Boch, J. Müller, V. H. Klaus, N. Hölzel & M. Fischer (2012): Direct and productivity‐mediated indirect effects of fertilization, mowing and grazing on grassland species richness. – Journal of Ecology 100(6): 1391–1399 [].

Tälle, M., B. Deák, P. Poschlod, O. Valkó, L. Westerberg & P. Milberg (2016): Grazing vs. mowing: a meta-analysis of biodiversity benefits for grassland management. – Agriculture, Ecosystems & Environment 222(4): 200–212 [].

Tappeiner, U., T. Marsoner & G. Niedrist (2020): Landwirtschaftsreport zur Nachhaltigkeit Südtirol. Eurac Research, Bozen: 141 pp.

Thiele, H. U. (1977). Carabid Beetles in Their Environments. A Study on Habitat Selection by Adaptations in Physiology and Behaviour. Springer-Verlag, Berlin, Heidelberg: 369 pp.

Toft, S. & T. Bilde (2002): Carabid Diets and Food Value. In: Holland, J. M (eds): The Agroecology of Carabid Beetles. Intercept Limited, Andover: 81–110.

Tsiafouli, M. A, E. Thébault, S. P. Sgardelis, P. C. de Ruiter, W. H. van der Putten, K. Birkhofer, L. Hemerik, F. T. de Vries, R. D. Bardgett, M. V. Brady & et al. (2015): Intensive agriculture reduces soil biodiversity across Europe. – Global Change Biology 21(2): 973–985 [].

Van den Berg, L. J. L., P. Vergeer, T. C. G. Rich, S. M. Smart, D. Guest & M. R. Ashmore (2011): Direct and indirect effects of nitrogen deposition on species composition change in calcareous grasslands. – Global Change Biology 17(5): 1871–1883 [].

Verberk, W. C. E. P., C. G. E. van Noordwijk & A. G. Hildrew (2013): Delivering on a promise: integrating species traits to transform descriptive community ecology into a predictive science. – Freshwater Science 32(2): 531–547 [].

Vickery, J., J. R. Tallowin, R. Feber, E. Asteraki, P. W. Atkinson, R. J. Fuller & V. K. Brown (2001): The management of lowland neutral grasslands in Britain: effects of agricultural practices on birds and their food resources. – Journal of Applied Ecology 38(3): 647–664 [].

Wickham, H. (2016): Package ggplot2: elegant graphics for data analysis. – Springer-Verlag, New York: 276 pp.

Wood, S. A., D. S. Karp, F. De Clerck, C. Kremen, S. Naeem & C. A. Palm (2015): Functional traits in agriculture: agrobiodiversity and ecosystem services. – Trends in Ecology and Evolution 30(9): 531–539 [].

Woodcock, B. A, S. G. Potts, T. Tscheulin, E. Pilgrim, A. J. Ramsey, J. Harrison-Cripps, V. K. Brown & J. R. Tallowin (2009): Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins. – Journal of Applied Ecology 46(4): 920–929 [].

World Spider Catalog (2022): World Spider Catalog. Version 23.5. – Natural History Museum Bern. Online at url: [].




How to Cite

Plunger, J., Guariento, E., Steinwandter, M. ., Colla, F., Rief, A. ., & Seeber, J. (2022). Shifts in ground-dwelling predator communities in response to changes in management intensity in Alpine meadows. Soil Organisms, 94(3), 149–161.