How do plant communities and flower visitors relate? A case study of semi-natural xerothermic grasslands

Damian Chmura, Paweł Adamski, Zygmunt Denisiuk


The paper examines the relationships between the species composition of flower visitors and plants in the semi-natural xerothermic grasslands in southern and central Poland. Thirty 10 × 10 m permanent plots were laid out in total, mainly in nature reserves. The vegetation units studied were classified according to the Braun-Blanquet system; these were phytocoenoses of the Festuco-Brometea classes Inuletum ensifoliae, Adonido-Brachypodietum pinnati and the transitional plant community. Entomological research was performed using the Pollard method within the same plots. A particular site was visited only once and different sites were studied between April and August 2008. We applied, among others, co-correspondence-analysis Co-CA, detrended correspondence analysis (DCA) and redundancy analysis (RDA) to investigate the co-occurrence patterns of plants and flower visitors and their biotopic requirements. We found that the species composition of flower visitors cannot be predicted by floristic composition when the duration of the study is restricted to one day (but under similar weather conditions); however, there is a positive relationship between the species richness of insects and plants and a positive relationship between the number of plant species and the abundance of flower visitors. The Ellenberg moisture index and the cover of meadow species significantly explained the species composition of insects. The three various vegetation units and five dominant xerothermic species, i.e. Adonis vernalis, Anemone sylvestris, Inula ensifolia, Linum hirsutum and Carlina onopordifolia that were studied across time differed in the species richness of insects. Our results demonstrate that possible patterns in the species composition and the assembly rules of flower visitors are not apparent when the Pollard method is applied. Based on the data obtained using this method, the flower visiting assemblages seem not to be driven by competition and they primarily show a tendency to co-occur which can be an artifact. A plant-focused method that included a rarefaction analysis yielded more insightful results and shed more light on the differences between the dominant plants that shape the physiognomy of plant communities in a possible pollination specialization.


calcareous vegetation; Co-CA; null model; entomofauna; species richness; sampling effort

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Cremene C, Groza G, Rakosy L, Schileyko AA, Baur A, Erhardt A, et al. Alterations of steppe-like grasslands in Eastern Europe: a threat to regional biodiversity hotspots. Conserv Biol. 2005;19(5):1606–1618.

Öckinger E, Eriksson AK, Smith HG. Effects of grassland abandonment, restoration and management on butterflies and vascular plants. Biol Conserv. 2006;133(3):291–300.

Dzwonko Z, Loster S. Changes in species composition of an abandoned limestone grassland in relation to climatic conditions. Pol J Ecol. 2011;59(4):687–698.

Dzwonko Z, Loster S. Changes in plant species composition in abandoned and restored limestone grasslands – the effects of three and shrub cutting. Acta Soc Bot Pol. 2008;77(1):67–75.

Wahlman H, Milberg P. Management of semi-natural grassland vegetation: evaluation of a long-term experiment in southern Sweden. Ann Bot Fenn. 2002;39:159–166.

Bąba W, Kurowska M, Kompała-Bąba A, Wilczek A, Długosz J, Szarejko I. Genetic diversity of populations of Brachypodium pinnatum (L.) P. Beauv.: expansive grass in a fragmented landscape. Pol J Ecol. 2012;60(1):31–40.

Tscharntke T, Klein AM, Kruess A, Steffan-Dewenter I, Thies C. Landscape perspectives on agricultural intensification and biodiversity – ecosystem service management. Ecol Lett. 2005;8(8):857–874.

Kremen C, Williams NM, Aizen MA, Gemmill-Herren B, LeBuhn G, Minckley R, et al. Pollination and other ecosystem services produced by mobile organisms: a conceptual framework for the effects of land-use change. Ecol Lett. 2007;10(4):299–314.

Öckinger E, Smith HG. Semi-natural grasslands as population sources for pollinating insects in agricultural landscapes. J Appl Ecol. 2006;44(1):50–59.

Taki H, Kevan PG, Ascher JS. Landscape effects of forest loss in a pollination system. Landsc Ecol. 2007;22(10):1575–1587.

Lopes LE, Buzato S. Variation in pollinator assemblages in a fragmented landscape and its effects on reproductive stages of a self-incompatible treelet, Psychotria suterella (Rubiaceae). Oecologia. 2007;154(2):305–314.

Thomas JA, Bourn N. D, Clarke RT, Stewart KE, Simcox DJ, Pearman GS, et al. The quality and isolation of habitat patches both determine where butterflies persist in fragmented landscapes. Proc R Soc Lond B Biol Sci. 2001;268(1478):1791–1796.

Johnson SD, Steiner KE. Generalization versus specialization in plant pollination systems. Trends Ecol Evol. 2000;15(4):140–143.

Zych M. On flower visitors and true pollinators: the case of protandrous Heracleum sphondylium L. (Apiaceae). Plant Syst Evol. 2006;263(3–4):159–179.

Watts S, Ovalle DH, Herrera MM, Ollerton J. Pollinator effectiveness of native and non-native flower visitors to an apparently generalist Andean shrub, Duranta mandonii (Verbenaceae). Plant Species Biol. 2012;27(2):147–158.

ter Braak CJF, Schaffers AP. Co-correspondence analysis: a new ordination method to relate two community compositions. Ecology. 2004;85(3):834–846.

Chmura D, Adamski P, Denisiuk Z. The plant-pollinator interactions in xerothermic habitats using phytosociological data. In: Chytrý M, editor. 17th International Workshop European Vegetation Survey. Using phytosociological data to address ecological questions, 1–5 May 2008, Brno, Czech Republik. Brno: Masaryk University; 2008. p. 25.

Denisiuk Z, Chmura D, Adamski P. Flowering and generative reproduction in small, isolated populations of endangered monocarpic perennial Carlina onopordifolia Besser (Asteraceae) in Poland. Pol J Ecol. 2009;57(1):89–97.

Westhoff V, van der Maarel E. The Braun-Blanquet approach. In: Whittaker RH, editor. Classification of plant communities. Hague: Junk; 1978. p. 287–399.

Pollard E. A method for assessing changes in the abundance of butterflies. Biol Conserv. 1977;12(2):115–134.

Pollard E, Yates TJ. Monitoring butterflies for ecology and conservation: the British butterfly monitoring scheme. London: Chapman and Hall; 1993.

Tolman T. Butterflies of Europe. Princeton NJ: Princeton University Press; 2001.

De la Barrera E, Nobel PS. Nectar: properties, floral aspects, and speculations on origin. Trends Plant Sci. 2004;9(2):65–69.

Hill MO. Diversity and evenness: a unifying notation and its consequences. Ecology. 1973;54:427–432.

The R project for statistical computing [Internet]. 2008 [cited 2012 Sep 27]; Available from:

ter Braak CJF, Šmilauer P. CANOCO reference manual and CanoDraw for Windows user’s guide: software for Canonical Community Ordination (version 4.5). New York NY: Microcomputer Power; 2002.

Ellenberg H, Weber HE, Düll R, Wirth V, Werner W, Paulißen D. Zeigerwerte der Pflanzen in Mitteleuropa. Scr Geobot. 1992;18:1–248.

Matuszkiewicz W. Przewodnik do oznaczania zbiorowisk roślinnych Polski. Warsaw: Polish Scientific Publishers PWN; 2001.

Gotelli NJ, Entsminger GL. EcoSim: null models software for ecology. Version 7 [Internet]. 2008 [cited 2012 Mar 31]; Available from:

Stone L, Roberts A. The checkerboard score and species distributions. Oecologia. 1990;85(1):74–79.

Gotelli NJ, McCabe DJ. Species co-occurrence: a meta-analysis of J. M. Diamond’s assembly rules model. Ecology. 2002;83(8):2091–2096.[2091:SCOAMA]2.0.CO;2

van Der Werf W, Woldewahid G, Van Huis A, Butrous M, Sykora K. Plant communities can predict the distribution of solitarious desert locust Schistocerca gregaria. J Appl Ecol. 2005;42(5):989–997.

Agrawal AA, Lau JA, Hambäck PA. Community heterogeneity and the evolution of interactions between plants and insect herbivores. Q Rev Biol. 2006;81(4):349–376.

Siemann E, Tilman D, Haarstad J, Ritchie R. Experimental tests of the dependence of arthropod diversity on plant diversity. Am Nat. 1998;152(5):738–750.

Knops JMH, Tilman D, Haddad NM, Naeem S, Mitchell CE, Haarstad J, et al. Effects of plant species richness on invasion dynamics, disease outbreaks, insect abundances and diversity. Ecol Lett. 1999;2(5):286–293.

Hatfield R, Lebuhn G. Patch and landscape factors shape community assemblage of bumble bees, Bombus spp. (Hymenoptera: Apidae), in montane meadows. Biol Conserv. 2007;139(1–2):150–158.

Albrecht M, Duelli P, MüLler C, Kleijn D, Schmid B. The Swiss agri-environment scheme enhances pollinator diversity and plant reproductive success in nearby intensively managed farmland. J Appl Phycol. 2007;44(4):813–822.

Skórka P, Settele J, Woyciechowski M. Effects of management cessation on grassland butterflies in southern Poland. Agric Ecosyst Env. 2007;121(4):319–324.

Fontaine C, Dajoz I, Meriguet J, Loreau M. Functional diversity of plant–pollinator interaction webs enhances the persistence of plant communities. Plos Biol. 2006;4(1):e1.

Ollerton J, Cranmer L. Latitudinal trends in plant-pollinator interactions: are tropical plants more specialised? Oikos. 2002;98(2):340–350.

Gioria M, Bacaro G, Feehan J. Evaluating and interpreting cross-taxon congruence: potential pitfalls and solutions. Acta Oecol. 2011;37(3):187–194.

Martinko EA, Hagen RH, Griffith JA. Successional change in the insect community of a fragmented landscape. Landsc Ecol. 2006;21(5):711–721.

Lopes LE, Buzato S. Variation in pollinator assemblages in a fragmented landscape and its effects on reproductive stages of a self-incompatible treelet, Psychotria suterella (Rubiaceae). Oecologia. 2007;154(2):305–314.

Schaffers AP, Raemakers IP, Sýkora KV, ter Braak CJF. Arthropod assemblages are best predicted by plant species composition. Ecology. 2008;89(3):782–794.

Hopwood JL. The contribution of roadside grassland restorations to native bee conservation. Biol Conserv. 2008;141(10):2632–2640.

Powell JA, Logan JA. Insect seasonality: circle map analysis of temperature-driven life cycles. Theor Popul Biol. 2005;67(3):161–179.

Yurk BP, Powell JA. Modeling the evolution of insect phenology. Bull Math Biol. 2008;71(4):952–979.

Moroń D, Lenda M, Skórka P, Szentgyörgyi H, Settele J, Woyciechowski M. Wild pollinator communities are negatively affected by invasion of alien goldenrods in grassland landscapes. Biol Conserv. 2009;142(7):1322–1332.

Kwaiser KS, Hendrix SD. Diversity and abundance of bees (Hymenoptera: Apiformes) in native and ruderal grasslands of agriculturally dominated landscapes. Agric Ecosyst Env. 2008;124(3–4):200–204.

Tello JS, Stevens RD, Dick CW. Patterns of species co-occurrence and density compensation: a test for interspecific competition in bat ectoparasite infracommunities. Oikos. 2008;117(5):693–702.