Abstract
The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strážovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16 m2. The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.
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References
Balkovič J, Kollár J, Čemanová G, Šimonovič V (2010) Indicating soil acidity using vegetation relevés in spatially limited areas – case study from the Považský Inovec, Slovakia. Folia Geobot 45:253–277
Başnou C, Pino J, Šmilauer P (2009) Effect of grazing on grasslands in the Western Romanian Carpathians depends on the bedrock type. Preslia 81:91–104
Botta-Dukát Z, Chytrý M, Hájková P, Havlová M (2005) Vegetation of lowland wet meadows along a climatic continentality gradient in Central Europe. Preslia 77:89–111
Casas C, Ninot JM (2003) Correlation between species composition and soil properties in the pastures of Plana de Vic (Catalonia, Spain). Acta Bot Barc 49:291–310
Chýlová T, Münzbergová Z (2008) Past land use co-determines the present distribution of dry grassland plant species. Preslia 80:183–198
Chytrý M, Tichý L, Holt J, Botta-Dukát Z (2002) Determination of diagnostic species with statistical fidelity measures. J Veg Sci 13:79–90
Chytrý M, Tichý L, Roleček J (2003) Local and regional patterns of species richness in Central European vegetation types along the pH/calcium gradient. Folia Geobot 38:429–442
Chytrý M, Danihelka J, Axmanová I, Božková J, Hettenbergerová E, Li CF, Rozbrojová Z, Sekulová L, Tichý L, Vymazalová M, Zelený D (2010) Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH. J Veg Sci 21:1125–1137
Corney PM, Le Duc MG, Smart SM, Kirby KJ, Bunce RGH, Marrs RH (2006) Relationships between the species composition of forest field-layer vegetation and environmental drivers assessed using a national scale survey. J Ecol 94:383–401
Critchley CNR, Chambers BJ, Fowbert JA, Bhogal A, Rose SC, Sanderson RA (2002) Plant species richness, functional type and soil properties of grasslands and allied vegetation in English Environmentally Sensitive Areas. Grass Forage Sci 57:82–92
Critchley CNR, Fowbert JA, Wright B (2007) Dynamics of species-rich upland hay meadows over 15 years and their relation with agricultural management practices. Appl Veg Sci 10:307–314
R Development Core Team (2008) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available at: www.R-project.org
Devictor V, Clavel J, Julliard R, Lavergne S, Mouillot D, Thuiller W, Venail P, Villéger S, Mouquet N (2010) Defining and measuring ecological specialization. J Appl Ecol 47:15–25
Ejrnæs R, Bruun HH (2000) Gradient analysis of dry grassland vegetation in Denmark. J Veg Sci 11:573–584
Ellenberg H, Weber HE, Düll R, Wirth W, Werner W, Paulissen D (1992) Zeigerwerte von Pflanzen in Mitteleuropa. Ed. 2. Scripta Geobot 18:1–258
Ewald J (2003) The calcareous riddle: Why are there so many calciphilous species in the Central European flora? Folia Geobot 38:357–366
Fagan, KC, Pywell RF, Bullock JM, Marrs RH (2008) Do restored calcareous grasslands on former arable fields resemble ancient targets? The effect of time, methods and environment on outcomes. J Appl Ecol 45:1293–1303
Fajmonová E (1995) Xerotermná vegetácia v juhozápadnej časti CHKO Strážovské vrchy (Xerothermic vegetation in the southwestern part of Strážovské vrchy Protected Area). Naturae Tutela 3:213–221
Fridley JD, Vandermast DB, Kuppinger DM, Manthey M, Peet RK (2007) Co-occurrence based assessment of habitat generalists and specialists: a new approach for the measurement of niche width. J Ecol 95:707–722
Grace JB (1999) The factors controlling species density in herbaceous plant communities: an assessment. Perspect Pl Ecol Evol Syst 2:1–28
Grime JP (1965) The ecological significance of lime-chlorosis. An experiment with two species of Lathyrus. New Phytol 64:477–87
Grime JP (1979) Plant strategies and vegetation processes. John Wiley & Sons, Chichester
Hájek M, Hájková P (2004) Environmental determinants of variation in Czech Calthion wet meadows: a synthesis of phytosociological data. Phytocoenologia 34:33–54
Hájek M, Hekera P, Hájková P (2002): Spring fen vegetation and water chemistry in the Western Carpathian flysch zone. Folia Geobot 37:205–224
Hájek M, Tichý L, Schamp BS, Zelený D, Roleček J, Hájková P, Apostolova I, Dítě D (2007) Testing the species pool hypothesis for mire vegetation: exploring the influence of pH specialists and habitat history. Oikos 116:1311–1322
Hájková P, Roleček J, Hájek M, Horsák M, Fajmon K, Polák M, Jamrichová E (2011) Prehistoric origin of the extremely species-rich semi-dry grasslands in the Bílé Karpaty Mts (Czech Republic and Slovakia). Preslia 83:185–204
Havlová M, Chytrý M, Tichý L (2004) Diversity of hay meadows in the Czech Republic: major types and environmental gradients. Phytocoenologia 34:551–567
Hejcman M, Klaudisová M, Schellberg J, Paetzold S (2010) The Rengen Grassland Experiment: effect of soil chemical properties on biomass production, plant species composition and species richness. Folia Geobot 45:125–142
Hettenbergerová E, Hájek M (2011) Is species richness of small spring fens influenced by the spatial mass effect? Commun Ecol 12: 202-209
Illyés E, Chytrý M, Botta-Dukát Z, Jandt U, Škodová I, Janišová M, Willner W, Hájek O (2007) Semi-dry grasslands along a climatic gradient across Central Europe: Vegetation classification with validation. J Veg Sci 18:835–846
Janišová M (2005) Vegetation-environment relationships in dry calcareous grassland. Ekológia 24:25–64
Janišová M, Hájková P, Hegedüšová K, Hrivnák R, Kliment J, Michálková D, Ružičková H, Řezníčková M, Tichý L, Škodová I, Uhliarová E, Ujházy K, Zaliberová M (2007) Travinnobylinná vegetácia Slovenska – elektronický expertný systém na identifikáciu syntaxónov (The expert system for identification of syntaxa of grassland vegetation in Slovakia). Botanický ústav SAV, Bratislava
Janssens F, Peters A, Tallowin JRB, Bakker JP, Bekker RM, Fillat F, Oomes MJM (1998) Relationship between soil chemical factors and grassland diversity. Pl Soil 202:69–78
Jongepierová I (ed) (2008) Louky Bílých Karpat (Grasslands of the White Carpathian Mountains). ZO ČSOP Bílé Karpaty, Veselí nad Moravou
Kalusová V, Le Duc MG, Gilbert JC, Lawson CS, Gowing DJG, Marrs RH (2009) Determining the important environmental variables controlling plant species community composition in mesotrophic grasslands in Great Britain. Appl Veg Sci 12:459–471
Karlík P, Poschlod P (2009) History or abiotic filter: which is more important in determining the species composition of calcareous grasslands? Preslia 81:321–340
Kooijman A, Hedenäs L (2009) Changes in nutrient availability from calcareous to acid wetland habitats with closely related brown moss species: increase instead of decrease in N and P. Pl Soil 324:267–278
Maheľ M (1982) Geologická mapa Strážovských vrchov (Geological map of the Stražovské vrchy Mts). GUDŠ, Bratislava
Marhold K, Hindák F (1998) Zoznam nižších a vyšších rastlín Slovenska (Checklist of non-vascular and vascular plants of Slovakia). Veda, Bratislava
Marini L, Scotton M, Klimek S, Isselstein J, Pecile A (2007) Effects of local factors on plant species richness and composition of Alpine meadows. Agric Ecosyst Environm 119:281–288
Marrs RH (1993) Soil fertility and nature conservation in Europe: theoretical considerations and practical management solutions. Advances Ecol Res 24:241–300
Martin MH (1968) Conditions affecting the distribution of Mercurialis perennis L. in certain Cambridgeshire woodlands. J Ecol 56:777–793
Matuszkiewicz W (2007) Przewodnik do oznaczania zbiorowisk roślinnych Polski (A guide for identifying plant communities of Poland). Ed. 3. Wydawnictvo Naukowe PWN, Warszawa
McCune B, Keon D (2002) Equations for potential annual direct incident radiation and heat load. J Veg Sci 13:603–606
Michalcová D, Gilbert JC, Lawson CS, Gowing DJG, Marrs RH (2011) The combined effect of waterlogging, extractable P and soil pH on α-diversity: a case study on mesotrophic grasslands in the UK. Pl Ecol 212:879–888
Öster M, Cousins S, Eriksson O (2007) Size and heterogeneity rather than landscape context determine plant species richness in semi-natural grasslands. J Veg Sci 18:859-868
Pärtel M (2002) Local plant diversity patterns and evolutionary history at the regional scale. Ecology 8:2361–2366
Piqueray J, Bottin G, Delescaille LM, Bisteau E, Colinet G, Mahy G (2011) Rapid restoration of a species-rich ecosystem assessed from soil and vegetation indicators: The case of calcareous grasslands restored from forest stands. Ecol Indicators 11:724–733
Poschlod P, Baumann A, Karlík P (2009) Origin and development of grasslands in Central Europe. In Veen P, Jefferson R, de Smidt J, van der Straaten J (eds) Grasslands in Europe of high nature value. KNNV Publishing, Zeist, pp 15–26
Reitalu T, Johansson LT, Sykes MT, Hall K, Prentice HC (2010) History matters: village distances, grazing and grassland species diversity. J Appl Ecol 47:1216–1224
Rozbrojová Z, Hájek M (2008) Changes in nutrient limitation of spring fen vegetation along environmental gradients in the West Carpathians. J Veg Sci 19:613–620
Rozbrojová Z, Hájek M, Hájek O (2010) Vegetation diversity of mesic meadow and pastures in the West Carpathians. Preslia 82:307–332
Schaffers AP (2002) Soil, biomass, and management of semi-natural vegetation I. Interrelationships. Pl Ecol 158:229–246
Šeffer J, Lasák R, Galvánek D, Stanová V (2002) Grasslands of Slovakia. Final report on National grassland Inventory 1998–2002. DAPHNE – Institute of Applied Ecology, Bratislava
Spiegelberger T, Deléglise C, De Danieli S, Bernard-Brunet C (2010) Resilience of acid subalpine grassland to short-term liming and fertilisation. Agric Ecosyst Environm 137:158–162
SPSS Inc. (2005) SPSS version 14.0. SPSS Inc., Chicago, IL. Available at: http://www-01.ibm.com/software/analytics/spss/
ter Braak CJF, Šmilauer P (2002) CANOCO reference manual and CanoDraw for Windows user’s guide: software for canonical community ordination. Version 4.5. Microcomputer Power, Ithaca, NY
Tichý L (2002) JUICE, software for vegetation classification. J Veg Sci 13:451–453
Tichý L, Chytrý M (2006) Statistical determination of diagnostic species for site groups of unequal size. J Veg Sci 17:809–818
Tichý L, Chytrý M, Hájek M, Talbot SS, Botta-Dukát Z (2010) OptimClass: Using species-to-cluster fidelity to determine the optimal partition in classification of ecological communities. J Veg Sci 21:287–299
Tüxen R, Ellenberg H (1937) Der systematische und ökologische Gruppenwert. Ein Beitrag zur Begriffsbildung und Methodik der Pflanzensoziologie. Mitt Flor.-Soz Arbeitsgem 3:171–184
Tyler G (2003) Some ecophysiological and historical approaches to species richness and calcicole/calcifuge behaviour – contribution to a debate. Folia Geobot 38:419–428
Tzialla CE, Veresoglou DS, Papakosta D, Mamolos AP (2006) Changes in soil characteristics and plant species composition along a moisture gradient in a Mediterranean pasture. J. Environm Managem 80:90–98
Veen P, Jefferson R, de Smidt J, van der Straaten J (eds) (2009) Grasslands in Europe of high nature value. KNNV Publishing, Zeist
Vermonden K, Leuven RSEW, Van Der Velde G, Hendriks AJ, Van Katwijk MM, Roelofs JGM, Lucassen ECHE, Jensen KS, Pedersen O (2010) Species pool versus site limitations of macrophytes in urban waters. Aquatic Sci 72:379–389
Virtanen R, Oksanen J, Oksanen L, Razzhivin VY (2006) Broad-scale vegetation-environment relationships in Eurasian high-latitude areas. J Veg Sci 17:519–528
Wagner V (2009) Eurosiberian meadows at their southern edge: patterns and phytogeography in the NW Tien Shan. J Veg Sci 20:199–208
Wassen MJ, Venterink HO, Lapshina ED, Tanneberger F (2005) Endangered plants persist under phosphorus limitation. Nature 437:547–550
Westhoff V, van der Maarel E (1978) The Braun-Blanquet approach. In Whittaker RH (ed) Classification of plant communities. W. Junk, The Hague, pp 289–399
Whittaker RH (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecol Monogr 30:279–338
Zbíral J (1995–1996) Analýza půd – Jednotné pracovné postupy (Soil analysis – uniform procedures). Ústřední kontrolní a zkušební ústav zemědělský, Brno
Zelený D (2009) Co-occurrence-based assessment of species habitat specialization is affected by the size of species pool: reply to Fridley et al. (2007). J Ecol 97:10–17
Zohlen A, Tyler G (2004) Soluble inorganic tissue phosphorus and calcicole-calcifuge behaviour of plants. Ann Bot (Oxford) 94:427–432
Acknowledgements
We thank Petr Hekera for assistance with sample processing and technical assistance, David Zelený for calculating specialization, Zbyněk Hradílek, Tomáš Berka and Martin Dančák for their help with critical species identification, Jana Smatanová for providing literature, Jarko Solar for map production and Mrs S. Mather for producing the final diagrams. This research was supported by the doctoral grant project GD526/09/H025 and long-term development project of the Czech Academy of Sciences no. RVO 67985939.
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Cachovanová, L., Hájek, M., Fajmonová, Z. et al. Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape. Folia Geobot 47, 349–371 (2012). https://doi.org/10.1007/s12224-012-9131-3
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DOI: https://doi.org/10.1007/s12224-012-9131-3
Keywords
- Community specialists
- Environmental gradients
- Phytosociology
- Soil P
- Soil pH
- West Carpathians
Plant nomenclature
- Marhold and Hindák (1998)