Acta Botanica 2-2016 - za web.indd ACTA BOT. CROAT. 75 (2), 2016 217 Acta Bot. Croat. 75 (2), 217–225, 2016 CODEN: ABCRA 25 DOI: 10.1515/botcro-2016-0027 ISSN 0365-0588 eISSN 1847-8476 The effect of agricultural landscape type on fi eld margin fl ora in south eastern Poland Małgorzata Wrzesień1*, Bożena Denisow2 1 Department of Geobotany, Institute of Biology and Biochemistry, Maria Curie-Skłodowska University, 19 Akademicka str., 20-033 Lublin, Poland 2 Department of Botany, Laboratory of Horticultural Plants Biology, University of Life Sciences in Lublin 15 Akademicka str., 20-950 Lublin, Poland Abstract – Plant species diversity is threatened in many agricultural landscapes due to the changes it has to undergo. Although the modifi cation of the agricultural landscape pattern is observed across Europe, both ex- tensive and intensive agricultural landscapes still co-exist in Poland. The objective of the study was to exam- ine the fl ora in fi eld margins in intensively and extensively managed agricultural landscapes, located across three regions in SE Poland. The fl ora was compared with respect to species richness, diversity, and evenness indices. Detrended correspondence analysis was employed to characterise variation in species composition. Agricultural landscape type made a higher contribution than the topography or geology to species richness and composition in fi eld margins. Field margins function as important habitats for general vascular plant spe- cies diversity and are useful for the conservation of rare, threatened, endangered or bee plants. A signifi cant decline in species diversity was observed over a distance of 1000 m from the habitat elements. Plants growing on fi eld margins are mainly perennials; however participation of annuals clearly increases in intensive land- scapes. The participation of wind-dispersed species decreased in an open-spaced intensive landscape. Ani- mal-dispersed plants predominated in an extensive landscape with forest islands. Irrespective of landscape type, native species predominated. However, these habitats create the biota and corridors for alien-invasive species as well. Keywords: extensive landscape, intensive landscape, invasive species, vascular plant biodiversity * Corresponding author, e-mail: mseptember@tlen.pl Introduction Agricultural landscape constitutes ca 47% of the area of the European Union (Eurostat 2015) and ca 60% of land in Poland (Central Statistical Offi ce 2014). Since 1970s, the vegetation of the agricultural landscape has been under in- creased anthropogenic pressure in Europe (Stoate et al., 2009, Andreasen and Andresen 2011). Agricultural land- scape structure i.e. ‘a heterogeneous land area composed of a cluster of interacting ecosystems’ is affected by large-scale changes, i.e. the fragmentation of habitats is rising signifi - cantly and the decline of heterogeneity of semi-natural habi- tats is observed (Baudry et al., 2000, Forman and Baudry 1984, Liira et al. 2008). As a result, a variety of small bio- topes – woodlots, hedgerows, ditches or fi eld boundaries – have largely disappeared from agricultural landscape (Rob- inson and Sutherland 2002, Reif et al. 2008). The occurrence of non-crop habitats within cultivated fi eld systems is particularly important. These structures are a buffer against run-off of chemicals from the fi eld into wa- ter, serve to reduce soil erosion, fl oods and pesticide drift, provide breeding and shelter sites, extend food niches for a variety of animals (Marshall and Moonen 2002, Delattrea et al. 2010), and also provide seed banks of many taxa (Du- elli and Obrist 2003, Dajdok and Wuczyński 2008). On a landscape level, non-crop habitats ensure linkages between habitats, maintain landscape diversity (Vickery et al. 2009), and have positive aesthetic effects (Marshall and Moonen 2002). Among non-crop habitats, fi eld margins are of eco- nomic interest for farmers, because these structures harbour organisms such as pollinators and predators of pests, (Her- zon and O’Hara 2007, Denisow and Wrzesień 2007, 2015a, Wrzesień and Denisow 2007, Morelli 2013). In western European countries, fi eld margins have been reduced drastically (Robinson and Sutherland 2002). By contrast, in Central and Eastern Europe with more exten- sive farming the network of fi eld margins is richer (Reif et al. 2008). Since 1990s, radical economic reforms and changes in agriculture sector have occurred in Poland. Currently, both intensive (market oriented) and extensive (self-suffi - WRZESIEŃ M., DENISOW B. 218 ACTA BOT. CROAT. 75 (2), 2016 cient, family-run) farms can be distinguished. The econom- ic changeovers led to environmental changes that are refl - eced in the modifi cation of the agricultural landscape pattern (Wuczyński et al. 2014). One of the specifi c features of the Polish agricultural landscape structure is the co-existence of both extensive (related to more traditional) and intensive (related to modern) landscapes. The aim of the study was to analyse fi eld margin fl ora on the large-scale landscape level in both intensively and extensively managed agricultural landscapes across three regions of Poland. The more specifi c goal was to investi- gate how habitat elements (forests and meadows) have an impact on the species composition, richness and diversity. We established fi eld margins at increasing distances from habitat elements to measure effects of isolation on species diversity. To make the survey more complex we analysed lifespan, dispersal mechanism, geographical status, and synecological groups of species in fi eld margins. Material and methods Study area The survey was conducted in the Lublin province (SE Poland) with about 68.4% of the area (1657.3 ha) covered by farmlands (Central Statistical Offi ce 2014). The study area included the three regions selected due to variability in agricultural landscape types (Kondracki 2002). In each re- gion both extensively and intensively managed landscapes as well as various habitat elements (grasslands, forests) are present. The regions are similar in climatic conditions, yet slightly differ in topography (Fig.1). The Hrubieszów Basin (HB) (50°48’N, 23°53’E) has an almost fl at to gently undulating topography, with elevations of generally less than 220 m above sea level, average annu- al temperature is 7.3 °C, annual precipitation is 600 mm. The soils are brown and chernozem. The natural vegetation of the area is composed of grasslands (Festuco-Brometea, Molinio-Arrhenatheretea) and mesophilous forests (Tilio- Carpinetum). The agricultural landscape is a mosaic of small-acreage fi elds and modern large-acreage fi elds and mean fi eld acreage is 9.2 ha. The main crops are wheat, sugar beets and legume crops. Giełczew Elevation (GE) (51°5’N, 22°58’E) is a plain with gently undulating topography, often with hills > 300 m above sea level, the average annual temperature is 7.5 °C and the annual precipitation is 630 mm. The main soil types are brown and grey-brown or rendzina. The area has an in- tensive agricultural landscape dominated by ≥ 6 ha fi elds (~83% of region area) and only small areas of grassland (Molinietalia, Arrhenatheretalia) and forests (Tilio -Carpi- netum). The common crops are wheat, potatoes, herbs. Lubartów High Plain (LHP) (51°28’N, 22°38’E) is an undulating area, 170–180 m above sea level, average annu- al temperature is 7.4 °C, annual precipitation is 590 mm. Soils are mainly podzolic composed of sands and clays. Landscape structure is dominated by small-scale farming, mean fi eld acreage is 6.3 ha. Forests (Peucedano-Pinetum and Querco-Pinetum), and grasslands (Molinio-Arrhe na the - retea) occur in the landscape matrix. Rye and other cereals predominate among crops. Data collection The fi eld survey was conducted in 2010 and 2011 from late June to mid August. Field margins were defi ned as ho- mogeneous linear structures with vegetation occurring on the outer border of fi elds. For each landscape type, we ran- domly selected 15 transect plots (300 m long and 1.3 to 2.8 meters width), i.e. in each region 45 transect plots have been investigated. In total 135 transect plots were explored. Transect plots were designed according to Dajdok and Wuczyński (2008). The geographic position of each transect plot was re- corded with a differential GPS. The transect plots were cat- egorized based on the type of agricultural landscape. Ac- cording to the habitat types in the surrounding of fi eld margins in each region three agricultural landscapes were selected (1) intensively managed with absence of habitat el- ements at > 1000 m distance from fi eld margins (I); (2) ex- tensively managed with grasslands in the surrounding of fi eld margins (EG); and (3) extensively managed with for- ests in the surrounding of fi eld margins (EF). The grassland and forest habitats were located at < 1000 m distance from transect plots. Vascular plant species were identifi ed in each transect plot. The abundance of plant species was estimated on the basis of the Braun-Blanquet scale (van der Maarel 1979). The syntaxonomic units were described according to Ma- tuszkiewicz (2001), and the nomenclature of vascular plants was based on Mirek et al. (2002). To make the description of fi eld margin fl ora more com- plex we analyzed how landscape type interacts with species characteristics. We compared the distribution patterns of: lifespan (annuals, biennials, perennials) and dispersal mechanism (animal, wind, auto). In addition, geographical status (natives, archaeophytes, i.e. those alien species that arrived prior to 1500, neophytes, i.e. those alien species that Fig. 1. Map of province in SE Poland, showing the study area; A – the location of regions: 1 – Lubartów High Plain (LHP), 2 – Giełczew Elevation (GE), 3 – Hrubieszów Basin (HB). Type of agriculture landscape: B – intensively managed, C – extensively managed with grasslands, D – extensively managed with forests. FIELD MARGIN FLORA IN SE POLAND ACTA BOT. CROAT. 75 (2), 2016 219 arrived after 1500), synecological groups (grassland spe- cies, forest species, synanthropic species) have been con- sidered. All of these categories are hereafter called ‘traits’ in this paper. The relevant data concerning the species char- acteristics were obtained from the LEDA traitbase (Kleyer et al. 2008) and BIOFLOR database (Klotz et al. 2002). Some species were assigned to more than one trait of a set of multistate categorical traits. Data analyses The vegetation on the transect plots was compared with respect to three types of indices, focusing on (i) species richness S = ni, where ni = species i; (ii) species diversity with the Shannon-Wiener index – H’ = −∑pi log2pi, where pi = frequency of the species i; (iii) species evenness with the Pielou index– J’ = H’/lnS, defi ned as the ratio of the ob- served diversity to the maximum diversity, where: S = the number of species and H max= lnS. J’ is constrained between 0 and 1; the less variation in communities between the spe- cies, the higher J’ is. The MVSP package was used to calcu- late the indices (Kovach 2005). The mean and SD (standard deviation) were computed and the values obtained were compared by the Kruskal-Wallis non-parametric test to re- veal the signifi cance of differences in the above-mentioned indices (Stanisz 2007). To characterize the general pattern of variation in species composition within the entire data set of vegetation we used an indirect ordination method, de- trended correspondence analysis (DCA), from CANOCO ver. 5 (ter Braak and Šmilauer 2012). The strength of the re- lation between species diversity and the distance from habi- tat elements was measured with the Pearson’s correlation coeffi cient (r). The Statistica software package version 10 developed by StatSoft Krakow was used for these analyses. Results A total of 376 vascular plant species, belonging to 36 families was recorded on fi eld margins within the three re- gions and agricultural landscape types (Fig. 2). The most abundant were Asteraceae (63 species – 16.7%), Fabaceae (35 species – 9.3%), Poaceae (31 species – 8.2%), Rosaceae (25 species – 6.6%), Lamiaceae (22 species – 5.8%), Ca ry o- phyllaceae (17 species – 4.5%), accounting for 51.3% of species. The most frequent were 32 species (noted on > 80% of transects), e.g. Dactylis glomerata, Elymus repens, Hyperi- cum perforatum, Knautia arvensis, Veronica chamaedrys, Alopecurus pratensis, Berteroa incana, Euphorbia cyparis- sias, Achillea millefolium. The next most frequent 57 spe- cies were noted in 50–80% transects, 251 species were present in 10–50% transects, and 36 species were recorded with a frequency lower than 10% in agricultural landscape. A few rare weeds – e.g. Anchusa arvensis, Cerinthe minor, Consolida regalis, Fumaria vaillantii, Lathyrus tuberosus, Salvia verticillata, Stachys annua, Herniaria hirsuta, Agrosthemma githago, Neslia paniculata, Euphorbia falca- ta, and Bromus secalinus were found (noted on < 5% of transects), including seven species from the Red List of the vascular plants in Poland (Zarzycki and Szeląg 2006) (Adonis aestivalis, Bromus secalinus, Cerasus fruticosa, Ely mus hispidus, Muscari comosum, Myosurus minimus, Potentilla rupestris). Native species (274 species – 72.83%) predominated on fi eld margins under consideration (Fig. 2). Out of total vas- cular fl ora recorded, 102 species – 27.17% were alien spe- cies. The number of alien species was similar among re- gions (Kruskal-Wallis test: H = 5.83, P = 0.122), but differed by the type of agricultural landscape (Kruskal-Wallis test: H = 15.33, P = 0.0005). The highest number of alien species was recorded within traditional landscape with forests in the surrounding of fi eld margins. Most of the alien species were identifi ed as archaeophytes (68 species –18.05%), only 34 neophytes (species – 9.08%) were noted. Most of the archaeophytes were identifi ed as segetal weeds (e.g. Thlaspi arvense, Consolida regalis, Geranium dissectum, Viola arvensis, Fumaria offi cinalis) and they were almost exclusively found in the peripheral zones of margins (= ad- jacent to fi elds). We recorded 19 invasive plant species among neophytes (Tab. 1). The most frequent were Galinsoga ciliata, Eriger- on annuus, Conyza canadensis, Galinsoga parvifl ora, Se- taria pumila, Solidago gigantea, Echinochoa crus-galli, Ama ranthus retrofl exus. The most abundant neophytes, with > 30% of cover were Solidago gigantea and Amaran- thus retrofl exus. The species composition was similar among the re- gions, out of 80% of fi eld margin fl ora was recorded in all regions, however the species composition differed consid- erably among landscape types (Fig. 3). The number of species in the particular transect plots was variable (mean = 97 ± 29.7 SD; ranging from 36 to 160). Species richness noted in fi eld margins differed among the types of agricultural landscape (Kruskal-Wallis test: H = 9.83, P = 0.032), however was similar across regions (Krus- Fig. 2. The number of native and alien (archaeophyte and neo- phyte) species recorded in various type of agricultural landscape, located in SE Poland (mean from three regions). Vertical bars in- dicate standard deviation (+ SD); the values indicated with differ- ent small letters are signifi cantly different between types of land- scapes according to the Kruskal-Wallis test. WRZESIEŃ M., DENISOW B. 220 ACTA BOT. CROAT. 75 (2), 2016 kal-Wallis test: H = 5.83, P = 0.176). The number of species on fi eld margins located in extensively managed landscapes with forest (EF) was approx. 30% higher than in margins located in extensively managed landscapes with grasslands (EG), and approx. 50% higher than in fi eld margins located in intensive landscapes (I) (Fig. 4). No signifi cant relation was observed between species diversity and the habitat elements (forests, meadows) under a distance of 700 m (Fig. 5). At larger distances, over 1000 m from habitat elements, species diversity decreased sig- nifi cantly with increasing distance from the habitat islands. The ratio of perennials to biennials to annuals was ap- proximately 5:1:2 (averaged 74 ± 7.3 and 34 ± 5.59 and 30 ± 18.15 species in the total fl ora, respectively). The share of perennials, biennials and annuals was related to land- scape type but not the region (Kruskal−Wallis test for land- scape effect: H = 9.32, P = 0.035; for region: H = 1.65, P = 0.43). The highest participation of annuals was recorded in intensively managed landscape (Fig. 6). There was no dif- ference in the share of perennial plant species between ex- tensively managed landscapes (EF vs. EG). The dispersal type of species noted within fi eld margins was related to the types of agricultural landscape (Kruskal- Wallis test: H = 0.32, P = 0.035). The number of wind-dis- persed species was the lowest in the modern landscape. Animal-dispersed plants predominated in the traditional landscape with forest islands (Fig. 6). Taking into consideration the synecological groups, the participation of grassland species (Molinio-Arrhenathere- Tab. 1. List of invasive plant species occurring in fi eld margins in SE Poland. The frequency of invasive species in transects depending on landscape type (I – intensively managed landscape, EG – extensively managed landscape with grasslands, EF – extensively managed land- scape with forests) and the habitats under threat. n – total number of transects, A – habitats created by humans, S – habitats partly trans- formed, N – communities of a natural character, Ar – archaeophyte, Ne – neophyte. Asterisk (*) denotes potentially invasive species. Species n % Type of habitats colonized Geographical- historical groupI EG EF Amaranthus retrofl exus 48 43.75 20 43.75 A Ne Aster x salignus 13 6.25 17.5 5.55 A,S Ne Bunias orientalis 31 12.25 20 37.53 A,S Ne Conyza canadensis 98 56.25 86.66 75 A Ne Echinochloa crus-galli 57 37.55 40 50 A,S Ar Echinocystis lobata 14 6.25 13.3 12.54 A,S,N Ne Erigeron annuus 100 75 53.33 93.8 A,S Ne Galinsoga ciliata 121 81.25 86.66 100 A Ne Galinsoga parvifl ora 97 62.55 53.33 100 A Ne Geranium sibiricum* 6 – – 12.55 A Ne Helianthus tuberosus 6 – 6.67 6.25 A,S,N Ne Heracleum sosnovsky 3 – 5.66 – A,S,N Ne Lupinus polyphyllus 28 6.25 6.67 50 A,S,N Ne Rosa rugosa 17 12.5 13.33 12.5 A,S,N Ne Rumex confertus 14 12.5 13.33 6.25 A,S Ne Setaria pumila 68 75 33.33 43.75 A Ar Setaria viridis 27 16.25 13.33 31.25 A Ar Solidago gigantea 66 31.25 40 75 A,S,N Ne Vicia grandifl ora 11 12.6 6.67 5.8 A,S Ne Fig. 3. Ordination diagram of the detrended correspondence anal- ysis (DCA) based on species matrix comprising the fl ora occur- ring in fi eld margins within three regions in SE Poland. Each point refers to the fi eld margins location within agriculture land- scape. White points correspond to intensively managed landscape (I), grey points – extensively managed landscape with grasslands (EG) and black points – extensively managed landscape with for- ests (EF). Regions: HB – Hrubieszów Basin, GE – Giełczew El- evation, LHP – Lubartów High Plain. Eigenvalues: Axis 1 – 0.102, Axis 2 – 0.066. The diagram explains 18.2% of total variance. FIELD MARGIN FLORA IN SE POLAND ACTA BOT. CROAT. 75 (2), 2016 221 tea, Festuco-Brometea), coniferous and mesophilous broad- leaved forest species (Vaccinio-Piceetea, Querco-Fagetea) and synanthropic communities species (Artemisietea, Stel- larietea mediae) were recorded. Irrespective of the agricul- tural landscape type, grassland species predominated and accounted for 40 – 43%. Discussion The fi eld margins in the agricultural landscape of SE Poland function as important habitats for general vascular plant species diversity, which is typifi ed by our research in which 376 vascular plant species were identifi ed, i.e. ap- proximately 1/3 of the regional fl ora (Fijałkowski 2003). This is consistent with a study conducted in Mediterranean region (Bassa et al. 2011) or Finland (Tarmi et al. 2009) and indicates the essentiality of fi eld margins as hotspots of plant species richness in agricultural landscape, irrespective of geographic regions, climatic types or fl ora history. It is well documented that the species richness in fi eld margins is particularly important for wildlife conservation in a crop- land surrounding. It is crucial for agronomic reasons, e.g. many plants that grow in fi eld margins are hosts for insects and spiders that are benefi cial to agriculture by controlling the number of crop pests, i.e. aphids (Marshall 2004). The diversity of species in fi eld margins refl ected the occurrence of grassland and forest islands in their vicinity. The species diversity declined signifi cantly over the 1000 m distance from habitat elements indicating that mid-fi eld islets are valuable sources of diversity in the landscape. Landscape heterogeneity is one of the landscape factors most adequate to explain plant diversity in non-crop habi- tats of agricultural landscapes (Andreasen and Andresen 2011). In several studies the species diversity declined sig- nifi cantly with increasing distance from the nature reserves; however different distances for such a decline have been reported. For example, Kohler et al. (2008) documented a drastic decline in forb species in fi eld margins in the fi rst 75 m from habitat elements. Marshal and Arnold (1995) dem- onstrated that fi eld margin fl ora is strongly infl uenced by Fig. 4. Species richness, Shannon-Wiener diversity index (H’), and Pielou evenness index (J’) calculated for fl ora in fi eld margins located in intensively and extensively managed agricultural landscapes. Vertical bars indicate standard deviation (+ SD); the values indicated with different small letters are signifi cantly different between type of landscapes according to the Kruskal-Wallis test. Fig. 5. Pearrson’s correlation between the diversity of species within fi eld margins located in extensively managed (A) and intensively managed (B) agricultural landscapes. WRZESIEŃ M., DENISOW B. 222 ACTA BOT. CROAT. 75 (2), 2016 location and documented a variety of species from adjacent woodlands. The plant communities of fi eld margins are de- termined by colonization along these linear structures (Marshall and Moonen 2002). Maintenance of diversity re- quires continuous colonization and our results suggest that beyond 1000 m colonization of species from the habitat is- lands can no longer compensate disappearance. Here, only species well adapted to the intensive management practices in the agricultural landscape are able to survive. The species composition across regions was similar, in- dicating that local topography, geology and environmental conditions had far less signifi cance for fi eld margin fl ora than the type of agricultural landscape. According to Aavik et al. (2008) the effects of agricultural landscape structure on fi eld margin fl ora is particularly important. It is accepted that fi eld margin fl ora refl ects the specifi city of habitat, e.g. the soil eutrophication, the increase in nitrogen and other nutrient levels (Kleijn and Verbeek 2000) or the physical disturbance of the soil environment related to agricultural practices (Bassa et al. 2011). We noted a great number of nitrophilous weeds (Urtica dioica, Amaranthus retrofl exus, Artemisia vulgaris, Cirsium arvense, Glechoma hederacea) and frequent occurrences of disturbance-tolerant generalists were also recorded (e.g. Poa pratensis, Rumex acetosa, Achillea millefolium, Elymus repens, Equisetum arvense, Artemisia vulgaris). Interestingly, regardless of the region and the landscape type we found a relatively high share of grassland special- ists in fi eld margins. Lindborg et al. (2014) reported that high grasslands species richness found in linear structures (fi eld margins, road verges) across agricultural landscapes is partly related to transformation of grasslands to crop- fi elds. Indeed, the process has been continuing since the 1960s in the study area (Fijałkowski 2003). Studies from Fig. 6. Boxplots displaying various traits of fi eld margin fl ora depending on the type of agriculture landscape located in three regions in SE Poland (HB – Hrubieszów Basin, GE – Giełczew Elevation, LHP – Lubartów High Plain). Vertical bars indicate standard deviation (± SD); the values indicated with different small letters are signifi cantly different between type of landscapes according to the Kruskal − Wallis test. FIELD MARGIN FLORA IN SE POLAND ACTA BOT. CROAT. 75 (2), 2016 223 other parts of Europe (Estonia, Switzerland) have also re- vealed the role of fi eld margins as alternative habitats for grassland species (Aavik and Lira 2010). We documented a high participation of perennials in fi eld margins; however their number declined in intensively managed landscape. In Poland the margins are allowed to regenerate naturally, therefore the occurrence of long-lived plants refl ects the intermediate stages of ecological succes- sion. Many data have demonstrated the benefi cial effects of perennials on the diversity of many organisms, i.e. insects (Szymkowiak et al. 2014), including pollinators (Faring et al. 2015), butterfl ies (Delattrea et al. 2010), birds (Vickery et al. 2009), small mammals (Żurawska-Seta and Barczak 2012). The effects are due to repeatable food niches, i.e. vegetative organs feed insects, seeds and fruits are suitable for birds (Vickery et al. 2009), nectar and pollen enhance pollinators (Denisow and Wrzesień 2015b). Among peren- nials we noted, i.e. Ranunculus acris, Hypericum perfora- tum, Berteroa incana, Euphorbia cyparissias, Pastinaca sativa, Potentilla argentea, Geranium pratense, species re- garded as particularly important for pollinators (Denisow 2011). Indirectly, the nectar and pollen producing perenni- als observed near entomophilous crops may have positive effects on their yields, as wild fl ower abundance increases the sizes of wild pollinator populations (Meek et al. 2002, Denisow and Wrzesień 2015a). In some EU countries, fi eld margins are exploited in the agri-environmental programs for sowing fl ower-rich seed mixes to counteract the unprec- edented decline in pollinators (Potts et al. 2010). Therefore, we assume that the occurrence of wild bee-fl ora in the sur- rounding of crops should be regarded as important in con- sideration of these habitats as playing a role in the conser- vation of pollinators. The proportion of annual weeds in fi eld margins corre- lated with the type of agricultural landscape. An analogous result was reported by Petersen et al. (2006), Liira et al., (2008) and Lindborg et al. (2014), who found that more an- nuals are present in fi eld margins located in intensely man- aged modern agricultural landscape than in those that are extensively managed. The relationship may refl ect the dif- ferences in farm management, agricultural operations, or differences in herbicide applications followed by large- scale and small-scale farmers. For example, disturbance of fi eld margins, reported from many European countries is more common in modern, intensive farming (Marshall and Moonen 2002). The habitat perturbance can create back- ground, i.e. gaps for colonization of annuals, the r- strate- gist (sensu GRIME, 1974). These species possess the abili- ty to use resources rapidly for successful establishment in changing environmental conditions. The absence of differences in the participation of annual species between fi eld margins located in extensive land- scape indicates that the number of annual weeds was effec- tively reduced by competition from perennials. The signifi - cance of perennial species for the limitation of annual weeds was highlighted by Aavik (2008). We documented that the type of dispersion was signifi - cantly related to landscape type. In accordance with Lind- borg et al. (2014), we found that the share of animal- and auto-dispersed-species increased signifi cantly in extensive- ly managed landscapes with mid-fi eld vegetation islets. Presumably, directional dispersal by biotic agents (animal- or self-dispersal), which delivers seeds less randomly is more effective to enhance colonization in an extensive landscape with different vegetation patches. However, in contrast to our expectations, we noted the lowest share of wind-dispersed species in an open-spaced intensively man- aged landscape. The phenomenon needs more empirical study to be explained. We recorded 3–4 fold more native than alien species. Predominance of native species in fi eld margins was also recorded in agricultural landscapes in other parts of Poland (Dajdok and Wuczyński 2008). Among aliens, the preva- lence of archaeophytes (i.e., those aliens that arrived prior to 1500), over neophytes (i.e., those aliens that arrived after 1500) has been found in our study. The majority of archaeo- phytes were identifi ed as segetal weeds. According to Daj- dok and Wuczyński (2008), weed archaeophytes are noted most frequently in the peripheral areas of fi eld margins, i.e. in zones that adjoin fi elds, and therefore fi eld margins play a minor role in the re-dispersion of weeds into crops. Notwithstanding their positive impact on general spe- cies richness, fi eld margin habitats also create corridors for migration of alien-invasive species. We observed that some of neophytes formed dense patches. Invasive alien species have a particularly devastating impact on native biota and are responsible for the decline of species richness or even extinctions (Vilà et al. 2010). In the regions studied, the cal- careous species (e.g. Adonis aestivalis, Fumaria vaillantii, Stachys annua, Thlaspi perfoliatum, Valerianella dentata) are considered at high risk from invasive plants (Haliniarz and Kapeluszny 2014). In Poland, neophytes from Asia and North America are particularly disadvantageous for native biodiversity (Tokarska-Guzik et al. 2012). Among them, we noted Bunias orientalis and Solidago gigantea. Due to the attractive fl oral reward (nectar and pollen), these species lure a variety of pollinators (Denisow 2011). Therefore, in addition to negative effects on local plant species biodiver- sity such species may induce the collapse of pollination webs and disrupt pollination services of entomophilous crops. We observed strong competition for Apis mellifera between Bunias orientalis and oilseed rape (Brassica na- pus). We frequently noted Amaranthus retrofl exus, Setaria pumila and Galinsoga parvifl ora. These species are known to invade various habitats (ditch banks, grasslands, wood edges) as well as fi elds, vineyards, pastures, orchards in many parts of the world, not only in Europe (Tokarska- Guzik et al. 2012, Daisie 2015). Among the species the geographical distribution of which has expanded and the number of stations substantially increased (approx. 40% since 1970; Latowski et al. 2010, Wrzesień 2010) we re- corded Vicia grandifl ora and Geranium sibiricum. Our results confi rm the fi ndings that fi eld margins are useful for the conservation of biodiversity in the agricultur- al landscape, as well as for plant species currently consid- ered rare, threatened or endangered. In the 1970s, most of these species were common weeds associated with crops. Radical changes in cropping methods and chemical appli- WRZESIEŃ M., DENISOW B. 224 ACTA BOT. CROAT. 75 (2), 2016 cations are responsible for the disappearance of segetal weed species or even a risk of their extinction (Haliniarz and Kapeluszny 2014, Wuczyński et al. 2014). Therefore disappearance of weed species, mainly archaeophytes, is nowadays a common trend in many regions of Poland (Zając et al. 2009) and in Europe (Pinke et al. 2011), where fi eld margins are also recognized as refugial habitats (Hamre et al. 2010, Fahrig et. al. 2015). The presence of rare or red list species has been suggested as an alternative indicator for the evaluation of diversity in agricultural land- scapes (Weibull and Östman 2003). However, our observa- tions indicate that only few rare, endangered or protected species occurred in fi eld margins and consequently, the idea that rare species might indicate the biodiversity in agroeco- systems seems to be untenable. Acknowledgements This research was supported fi nancially by the Ministry of Science and Higher Education of Poland (project OKB/ DS/2) as a part of the statutory activities of Department of Geobotany, Institute of Biology and Biochemistry, Maria Curie-Skłodowska University and the Department of Bota- ny, University of Life Sciences in Lublin. We are grateful to Anna Wesołowska-Zoń for reading the text and for making linguistic corrections. References Aavik, T., Liira, J., 2010: Quantifying the effect of organic farm- ing, fi eld boundary type and landscape structure on the vege- tation of fi eld boundaries. Agriculture Ecosystems and Envi- ronment 135, 178–186. Aavik, T., Augenstein, I., Bailey, D., Herzog, F., Zobel, M., Liira, J., 2008: What is the role of local landscape structure in the vegetation composition of fi eld boundaries? Applied Vegeta- tion Science 11, 375–386. Andreasen, C., Andresen, L. C., 2011: Managing farmland fl ora to promote biodiversity in Europe. CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Re- sources 6, 1–11. Bassa, M., Boutinb, C., Chamorroa, L., Sans, F. X., 2011: Effects of farming management and landscape heterogeneity on plant species composition of Mediterranean fi eld boundaries. Agri- culture Ecosystems and Environment 141, 455–460. Baudry, J., Bunce, R. G. H., Burel, F., 2000: Hedgerows: an inter- national perspective on their origin, function and manage- ment. Journal of Environmental Management 60, 7–22. Central Statistical Offi ce, 2014: Statistical Yearbook of Agricul- ture. Warsaw. Daisie, 2015: European Invasive Alien Species Gateway. Re- trieved from http://www.europe-aliens.org Dajdok, Z., Wuczyński, A., 2008: Alien plants of fi eld margins and fi elds of southwestern Poland. Biodiversity: Research and Conservation 9–10, 19–33. Delattrea, T., Pichancourt, J. B., Burel, F., Kindlmann, P., 2010: Grassy fi eld margins as potential corridors for butterfl ies in agricultural landscapes. A simulation study. Ecological Mod- elling 221, 370–377. Denisow, B., 2011: Pollen production of selected ruderal plant species in the Lublin area. University of Life Sciences Press. Lublin. Denisow, B., Wrzesień, M., 2007: The anthropogenic refuge areas for bee fl ora in agricultural landscape. Acta Agrobotanica 60, 147–157. Denisow, B., Wrzesień, M., 2015a: The habitat effect on the diver- sity of pollen resources in several Campanula spp. – an impli- cation for pollinator conservation. Journal of Apicultural Re- search 54, 1–9. Denisow, B., Wrzesień, M., 2015b: The importance of fi eld mar- gin location for maintenance of food niche for pollinators. Journal of Apicultural Science 59, 27–37. Duelli, P., Obrist, M. K., 2003: Regional biodiversity in an agri- cultural landscape: the contribution of seminatural habitat is- lands. Basic and Applied Ecology 4, 129–138. Eurostat, 2015: Farm structure statistics. Retrieved from http:// ec.europa.eu/eurostat/statistics-explained/index.php/farm_ structure_statistics Fahrig, L., Girard, J., Duro, D., Pasher, J., Smith, A., Javorek, S., King, D., Lindsay, F. K., Mitchell, S., Tischendorf, L., 2015: Farmlands with smaller crop fi elds have higher within-fi eld biodiversity. Agriculture Ecosystems and Environment 200, 219–234. Fijałkowski, D., 2003: Protection of nature and environment of Lublin Voivodship. Lubelskie Towarzystwo Naukowe, Lublin (in Polish). Forman, R. T. T., Baudry, J., 1984: Hedgerows and hedgerow net- works in landscape ecology. Environmental Management 8, 495–510. Grime. P.J., 1974: Vegetation classifi cation by reference to strate- gies. Nature 250, 26–31. Haliniarz, M., Kapeluszny, J., 2014: Rare species of calcicole fl ora in the segetal communities in the Lublin region. Annales Uni- versitatis Mariae Curie-Skłodowska Lublin – Polonia 69, 11– 23 (in Polish). Hamre, L. N., Halvorsen, R., Edwardsen, A., Rydgren, K., 2010: Plant species richness, composition and habitat specifi city in a Norwegian agricultural landscape. Agriculture Ecosystems and Environment 138, 189–196. Herzon, I., O’Hara, R. B., 2007: Effects of landscape complexity on farmland birds in Baltic States. Agriculture Ecosystems and Environment 108, 297–306. Kleijn, D., Verbeek, M., 2000: Factors affecting the species com- position of arable fi eld boundary vegetation. Journal of Ap- plied Ecology 37, 256–266. Kleyer, M., Bekker, R. M., Knevel, I. C., Bakker, J. P., Thompson, K., Sonnenschein, M., Poschlod, P., Van Groenendael, J. M., Klimeš, L., Klimešová, J., Klotz, S., Rusch, G. M., Hermy, M., Adriaens, D., Boedeltje, G., Bossuyt, B., Dannemann, A., Endels, P., Götzenberger, L., Hodgson, J. G., Jackel, A. K., Kühn, I., Kunzmann, D., Ozinga, W. A., Römermann, C., Sta- dler, M., Schlegelmilch, J., Steendam, H. J., Tackenberg, O., Wilmann, B., Cornelissen, J. H. C., Eriksson, O., Garnier, E., Peco, B., 2008: The LEDA Traitbase: a database of life-histo- ry traits of the Northwest European fl ora. Journal of Ecology 96, 1266–1274. Klotz, S., Kühn, I., Durka, W., (eds.) 2002: BIOLFLOR: Eine Da- tenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland. Schriftenr Vegetationsk. 38, 1–334. Kohler, F., Verhulst, J., van Klink, R., Kleijn, D., 2008: At what spatial scale do high quality habitats enhance the diversity of forbs and pollinators in intensively farmed landscapes? Jour- nal of Applied Ecology 45, 753–762. FIELD MARGIN FLORA IN SE POLAND ACTA BOT. CROAT. 75 (2), 2016 225 Kondracki, J., 2002: Regional geography of Poland. Polish Scien- tifi c Publishers PWN, Warszawa (in Polish). Kovach, W. L., 2005: MVSP – A MultiVariate Statistical Package for Windows, ver. 3.1. Kovach Computing Services, Pentra- eth, Wales, UK. Latowski, K., Chmiel, J., Jackowiak, B., Żukowski, W., 2010: Participation of anthropophytes in the segetal fl ora of Wielko- polska. Fragmenta Agronomica 27(3), 103–111. Liira, J., Schmidt, T., Aavik, T., Arens, P., Augenstein, I., Bailey, D., Billeter, R., Bukáček, R., Burel, F., Blust, G., Cock, R., Dirksen, J., Edwards, P. J., Hamerský, R., Herzog, F., Klotz, S., Kühn, I., Le Coeur, D., Miklová, P., Roubalova, M., Sch- weiger, O., Smulders, M. J. M., Wingerden, W. K. R. E., Bug- ter, R., Zobel, M., 2008: Plant functional group composition and large–scale species richness in European agricultural landscapes. Journal of Vegetation Science 19, 3–14. Lindborg, R., Plue, J., Andersson, K., Cousins S. A. O., 2014: Function of small habitat elements for enhancing plant diver- sity in different agricultural landscapes. Biological Conserva- tion 169, 206–213. Marshall, E. J. P., 2004: Agricultural landscapes: Field margin habitats and their interaction with crop production. Journal of Crop Improvement 12, 365–404. Marshall, E.J.P., Arnold, G.M., 1995: Factors affecting fi eld weed and fi eld margin fl ora on a farm in Essex, UK. Landscape Ur- ban Planning 31, 205–216. Marshall, E. J. P., Moonen, A. C., 2002: Field margins in northern Europe: their functions and interactions with agriculture. Agri- culture Ecosystems and Environment 89, 5–21. Matuszkiewicz, W., 2001: Guide to the plant communities of Po- land. Polish Scientifi c Publishers PWN, Warszawa (in Polish). Meek, B., Loxton, D., Sparks, T., Pywell, R., Pickett, H., Nowa- kowski, M., 2002: The effect of arable fi eld margin composi- tion on invertebrate biodiversity. Biological Conservation 106, 259–271. Mirek, Z., Piękoś–Mirkowa, H., Zając, A., Zając, M., 2002: Flow- ering plants and pteridophytes a checklist. In: Mirek, Z. (ed.), Biodiversity of Poland, (vol.1), W. Szafer Institute of Botany, Polish Academy of Sciences, Cracow. Morelli, F., 2013: Relative importance of marginal vegetation (shrubs, hedgerows, isolated trees) surrogate of HNV farm- land for bird species distribution in Central Italy. Ecological Engineering 57, 261–266. Petersen, S., Axelsen, J.A., Tybirk, K., Aude, E., Vestergaard, P., 2006: Effects of organic farming on fi eld boundary vegetation in Denmark. Agriculture Ecosystems and Environment 113, 302–306. Pinke, G., Kiraly, G., Barina, Z., Mesterhazy, A., Balogh, L., Csiky, J., Schmotzer, A., Molnar A., V., Pal, R., W., 2011: As- sessment of endangered synanthropic plants of Hungary with special attention to arable weeds. Plant Biosystems 145, 426– 435. Potts, S. G., Roberts, S. P. M., Dean, R., Marris, G., Brown, M. A., Jones, R., Neuman, P., Settele, J., 2010: Declines of managed honey bees and beekeepers in Europe. Journal of Apicultural Research 49, 15–22. Reif, J., Vorísek, P., Stasny, K., Bejcek, V., Pert, J., 2008: Agricul- tural intensifi cation and farmland birds: new insights from a central European country. Ibis. 150, 596–605. Robinson, R. A., Sutherland, W. J., 2002: Post-war changes in ar- able farming and biodiversity in Great Britain. Journal of Ap- plied Ecology 39, 157–176. Stanisz, A., 2007: Accessible course in statistics Rusing Statis tica Software on ex ample from medicine. Statsoft Polska, Kra- ków. Stoate, C., Baldi, A., Beja, P., Boatman, N. D., Herzon, I., Van Doorn, A., De Snoo, G. R., Rakosy, L., Ramwell, C., 2009: Ecological impacts of early 21st century agricultural change in Europe – a review. Journal of Environmental Management 91, 22–46. Szymkowiak, J., Skierczynski, M., Kuczyński, L., 2014: Are bun- tings good indicators of agricultural intensity? Agriculture Ecosystems and Environment 188, 192–197. Tarmi, S., Helenius, J., Hyvönen, T., 2009. Importance of edaphic, spatial and manage-ment factors for plant communities of fi eld boundaries. Agriculture, Ecosystems and Environment 131, 201–206. Ter Braak, C. J. F., Šmilauer, P., 2012: Canoco reference manual and user’s guide: software for ordination, version 5.0. Micro- computer Power, Ithaca, USA. Tokarska-Guzik, B., Dajdok, Z., Zając, M., Zając, A., Urbisz, A., Danielewicz, W., Hołdyński, C., 2012: Alien plants species in Poland with emphasis on invasive species. Generalna Dyrek- cja Ochrony Środowiska, Warszawa (in Polish). Weibull, A. C., Östman, Ö., 2003: Species composition in agro- ecosystems: The effect of landscape, habitat, and farm man- agement. Basic and Applied Ecology 4, 349–361. Wrzesień, M., 2010: Anthropophytes related to the habitats of rail- way grounds in central eastern Poland, In: Barančoková, M., Krajčí, J., Kollár, J., Belčáková, I. (eds.), Landscape ecology – methods, applications and interdisciplinary approach. Insti- tute of Landscape Ecology, Slovak Academy of Sciences, Bratislava: 625–634. Wrzesień, M., Denisow, B., 2007: The phytocoenoses of anthro- pogenically transformed areas with a great importance for Apoidea. Acta Agrobotanica 60, 117–126. Wuczyński, A., Dajdok, Z., Wierzcholska, S., Kujawa, K., 2014: Applying red lists to the evaluation of agricultural habitat: regular occurrence of threatened birds, vascular plants, and bryophytes in fi eld margins of Poland. Biodiversity and Con- servation 23, 999–1017. Van Der Maarel, E., 1979: Transformation of cover–abundance values in phytosociology and its effects on community simi- larity. Vegetatio 29, 97–114. Vickery, J. A., Feber, R. E., Fuller, R. A., 2009: Arable fi eld mar- gins managed for biodiversity conservation: a review of food resource provision for farmland birds. Agriculture, Ecosys- tems and Environment 133, 1–13. Vilà, M., Basnou, C., Pyšek, P., Josefsson, M., Genovesi, P., Gollasch, S., Nentwig, W., Olenin, S., Roques, A., Roy, D., Hulme, P., and Daisie Partners, 2010: How well do we under- stand the impacts of alien species on ecosystem services? A Pan–European cross–taxa assessment. Frontiers in Ecology and the Environment 8, 135–144. Zając, M., Zając, A., Tokarska-Guzik, B., 2009: Extinct and en- dangered archaeophytes and the dynamics of their diversity in Poland. Biodiversity: Research and Conservation 13, 17–24. Zarzycki, K., Szeląg, Z., 2006: Red list of the vascular plants in Poland. In: Mirek Z., Zarzycki K., Wojewoda W., Szeląg Z., (eds.) Red list of plants and fungi in Poland. W. Szafer Insti- tute of Botany, Polish Academy of Sciences, Kraków. Żurawska-Seta, E., Barczak, T., 2012: The Infl uence of fi eld mar- gins on the presence and spatial distribution of the European mole Talpa europaea L. within the agricultural landscape of northern Poland. Archives of Biological Sciences 64, 971–980.