Last Remnants of Riparian Wooded Meadows along the Middle Drava River (Slovenia): Species...

Last Remnants of Riparian Wooded Meadowsalong the Middle Drava River (Slovenia): SpeciesComposition Is a Response to LightConditions and Management

Sonja Škornik & Nina Šajna & Branko Kramberger &

Simona Kaligarič & Mitja Kaligarič

# Institute of Botany, Academy of Sciences of the Czech Republic 2008

Abstract In the present study, we investigated the species richness and speciescomposition in relation to light and management regime in remnants of woodedmeadows within the riparian forests along the middle Drava River in Slovenia. 41plots of still managed and, at different time periods (<5 yrs, 5–15 yrs, >15 yrs),abandoned riparian wooded meadows (RWM) were sampled. In addition tovegetation relevés, light intensity (PAR) was also measured in plots. Within thestill-managed RWM, two floristically distinct types were recognized usingTWINSPAN analysis: meadow-like and forest-like. Light intensity differedsignificantly between types. The CCA of active RWM showed a significant relationbetween species composition and light conditions. The number of species per relevéon active RWM was negatively correlated with light intensity – in contrast to NorthEuropean wooded grasslands. This could be explained by the influence of species-rich riparian hornbeam forests that contribute many understorey species, in contrastto naturally mesotrophic meadows. CCA of both active and abandoned RWMdemonstrated that light was a good predictor of RWM species composition and thatabandonment caused profound changes in floristic composition. The speciesturnover during succession was more pronounced in less shaded meadow-like

Folia Geobot (2008) 43:431–445DOI 10.1007/s12224-008-9024-7

Plant nomenclature For taxa Martinčič et al. (2007) and for syntaxa Mucina et al. (1993)

S. Škornik :N. Šajna :M. Kaligarič (*)Biology Department, Faculty for Natural Sciences and Mathematics, University of Maribor,Koroška 160, SI 2000 Maribor, Sloveniae-mail: [email protected]

B. KrambergerFaculty of Agriculture, University of Maribor, Vrbanska 30, SI 2000 Maribor, Slovenia

S. KaligaričInstitute of the Republic of Slovenia for Nature Conservation, Maribor regional unit,Pobreska cesta 20a, SI 2000 Maribor, Slovenia

RWM where more light-requiring (grassland) species occurred. Species richness wasthe highest in active forest-like RWM. There were no significant differences inspecies richness between active meadow-like RWM and groups of abandonedRWM. The remains of riparian forests stretching along the flood plains of Centraland Eastern Europe are considered one of the most natural ecosystems in theprevailing agricultural landscape. But naturalness could be in many cases only theconsequence of abandonment of ancient land-use practices, like grazing of livestock,cutting between the trees, litter collecting, etc. Examination of the middle Drava Riverin Slovenia suggests the need to recognize the remains of ancient cultural landscape.

Keywords Abandonment . Agricultural management . Multivariate analysis .

Photosynthetically Active Radiation (PAR) . Species richness

Introduction

Wooded meadows throughout Europe represent a type of semi-natural grasslandcharacterized by scattered groups of trees and shrubs (Ellenberg 1996; Hansson andFogelfors 2000). They could be defined also from another point of view – they aresparse natural wooded stands with a regularly mowed herb layer (Kukk and Kull1997). The last definition fits better to the wooded meadows in this study, whichwould be riparian woodland with regular mowing and strewing.

In Germany wooded meadows were called “Laubwiesen” (meadows withdeciduous trees). The distinction between grazed (“Wytweiden”) and mowedwooded meadows (“Studmatten”) was drawn in Switzerland (Ellenberg 1996). Itwas reported that such wooded meadows exist also within riparian forests (Ellenberg1996). These “straw meadows” (mainly for strewing purpose) mainly occupy verywet sites, which are avoided by stock and do not need to be fenced off.

Wooded meadows were once abundant, particularly in Scandinavia (Sweden,Finland, Norway, Danmark) and in the Baltic (Kukk and Kull 1997; Hansson andFogelfors 2000). Less typical wooded meadows could be found also in themountains of Central Europe (Kukk and Kull 1997), e.g., in the Carpathian region(Klimeš 1995, 1997, 1999).

Since the end of the 18th century the abundance of wooded meadows hasdeclined rapidly, mostly owing to agricultural intensification and abandonment(Huhta 1996; Hansson and Fogelfors 2000). Today, only remnants of these meadowsare left, representing a high conservation priority, the result of which should becontrolled management and restoration of wooded meadows (Bakker 1989;Kotiluoto 1998; Paal 1998; Pärtel et al. 1998; Einarsson and Milberg 1999; Klimeš1999; Mitlacher et al. 2002).

A basic ecological study of wooded meadows, focusing especially on lightconditions and species composition was performed in Sweden (Sjörs 1954). Later itwas clearly confirmed that species richness increased with light availability in thewooded meadows of Southern Sweden (Einarsson and Milberg 1999). Large patcheslacking tree cover would be needed to favor light-requiring species, many of whichare rare in certain regions, because most of them tolerate only a small amount ofshade (Einarsson and Milberg 1999). Considering that semi-natural grasslands are

432 S. Škornik et al.

among the most species-rich habitats (e.g., Kull and Zobel 1991; Hansson andFogelfors 2000), it is clear that light-requiring grassland species contribute to thehigh diversity of wooded meadows. What does it mean in numbers? For 328 speciesin Estonian flora, wooded meadows are mentioned as their habitat (Kukk and Kull1997). In wooded meadows the number of species per square meter can exceed 50.The maximum was recorded from Estonia – 68 species as an average on ten 1×1 m2

plots (Kukk and Kull 1997).After abandonment, these meadows became rapidly overgrown with deciduous

shrub and tree species forming a closed canopy that resulted in a strong, rapiddecrease in species number in the herb layer (Mitlacher et al. 2002). 87% of thespecies vanished in the Baltic island of Öland – the mean number of speciesdecreased from 52 species per plot to 18 species (Mitlacher et al. 2002).

Slovenia is the country with highest proportion of Natura 2000 areas (33% of thetotal surface), but wooded meadows do not conform to any of the priority habitats. Noconservation values have officially been attributed to particular landscape relicts sofar, which is a failure that definitely points to the final extinction of the remnants of theriparian wooded meadows (RWM) that we estimate will appear in the next 20 years.

In the present study, we investigated the species richness and species compositionin relation to light and management regime in remnants of wooded meadows withinthe riparian forests along the middle Drava River in Slovenia. Because management –mowing once a year – is nowadays very rare, abandoned RWM were also included inthe study.

Our hypotheses are i) that different types of active RWM exist as a consequenceof different light conditions, ii) that species richness of RWM is positively correlatedwith light availability and iii) in abandoned RWM the number of herb speciesdecreases over time since the point of abandonment.

The present study aims to classify floristically still active RWM and to determinehow light availability and time since abandonment (<5yrs, 5–15 yrs, >15 yrs) affectthe species richness and composition in the herb layer or RWM.

Material and Methods

Study Area

The riparian wooded meadows (RWM) that we studied are located in the lowlandriparian forest along the Drava River in its middle stream between Maribor and theCroatian border near Ormož (NE Slovenia): a belt about 46 km long and no morethan 2 km wide between 46°22′ N, 46°33′ N and 15°43′ E, 16°04′ E, at ca. 200 m a.s.l. (Fig. 1). According to Köppen’s classification (Köppen 1923), the study area’sclimate is of the climatic type Cfb (moderately warm, rainy climate without a dryperiod). The long-term mean of minimum temperatures in the coldest month(January) is -1.5°C and of maximum temperatures in the warmest month (July) is19.6°C. The average annual precipitation is between 900 and 1,000 mm. Theprecipitation is spread rather evenly throughout the year, with a maximum insummer (July and August) and a minimum in winter (January, February) (Žiberna2000). The geological substrate is made up of gravel, sand, clay and loam deposited

Species composition of wooded meadows in relation to light and management 433

in the Pannonian Sea in the Tertiary and river detritus in the Quaternary. On alluvialsubstrates, alluvial and swampy gleyic (eugleyic) soils were developed (Žiberna 2000).

The dominant species in the tree layer are hornbeam (Carpinus betulus),pedunculate oak (Quercus robur), common ash (Fraxinus excelsior), elm (Ulmuslaevis), common alder (Alnus glutinosa), white poplar (Populus alba), black poplar(Populus nigra) and black locust (Robinia pseudacacia), which are characteristictree species of riverine forest communities in the Drava River basin in Slovenia.

Grasslands in the local region belong mainly to mesotrophic meadows from theArrhenatherion alliance (Kramberger and Klemenčič 2003). Grasslands with aninventory of the species-rich Bromion erecti alliance are rare and appear in smallfragments in the nearby hilly region of Haloze (Škornik and Kaligarič 2002).

Interviews with local farmers established that 30 years ago approx. 80% ofwooded meadows in the area were still traditionally managed. Traditionalmanagement means that in spring (March) leaves were raked and litter was usedfor cattle. From the early spring (February–March) until the end of April, cattlegrazed the meadows and then grazing was halted to enable the growing of herbs forhay. Hay-making was done once or twice per year – early in July and late inSeptember. Every year clearings of dry and fallen trees and shrubs were carried outin late autumn/winter. Reforestation with young trees was carried out as necessary tomaintain the traditional tree density.

In August 2005 the total sampling area was scanned in detail using fieldobservation with aerial photos and oral information. All polygons of still-managedRWM and also polygons in different stages of abandoned RWM were recorded. We

Fig. 1 Inset map of Slovenia and neighbouring regions. The more detailed map shows the position ofstudied riparian wooded meadows (RWM) along the Drava River in Slovenia


estimated that at least 80% of the polygons of existent active RWM in the study areawere registered. In our research we examined 18 polygons of active woodedmeadows and 23 polygons of RWM abandoned at different time periods. Thepolygons were adjacent to each other and had similar habitat features (topographyand soil). Therefore, we consider the polygons to present chronosequence in whichwe can study the effects of abandonment on vegetation of RWM. The mean area ofthe polygons was ca. 1,500 m2. On each of the 41 polygons 1 plot of 5×5 m (in thecore area of the polygon, where vegetation is the most homogenous) wereestablished. In sampling procedure of the plot selection, we avoided the 1.5-m-wide outer belt of the polygons. These 41 plots were divided into four groups accordingto the successional stage: i) active (traditionally managed) RWM (18 plots); ii) <5 yrsabandoned RWM (8 plots); iii) 5–15 yrs abandoned RWM (8 plots); iv) >15 yrsabandoned RWM (7 plots). The data for time since abandonment were obtained fromthe land owners.

Vegetation Survey

The species composition of the RWM was recorded by identifying all vascular plantspecies in the plots. Their abundance within plots was recorded on a six-point scale(+ to 5), according to the Braun-Blanquet method (Braun-Blanquet 1964; Westhoffand van der Maarel 1973; Dierschke 1994). Species richness was noted as thenumber of species recorded in each plot. The rough grouping to species occurring ingrassland communities, woodland and shrubland communities and disturbed habitatswere summarized from Ellenberg et al. (1992). 22 species were not connectedexclusively to one of the above-mentioned groups; we classify them as species withindifferent habitat preference. The investigation included the herb-layer and wascarried out from June to September 2006.

Light Recordings

The light situation for each plot was calculated as the percentage of daylightPhotosynthetically Active Radiation (PAR, 400–700 nm) with a PAR quantumsensor (SKP 215, Skye Instruments Ltd. Llandrindod Wells, Wales U.K.). Fivemeasurements were taken at five randomly selected points in each plot beforemowing (in active RWM). The sensor was positioned 30 to 150 cm above the herb-layer canopy, regardless of the canopy height. A measurement was simultaneouslymade in a fully open area. We did light measurements on three occasions during oneday at the peak of the growth season (July 2006): at noon, 2 h before noon and 2 hafter noon. To get representative light recordings, we made them only in cloudyweather, i.e., when thick clouds hid the sun. This reduced the impact of sunlightflecks and eliminated the sharp gradients between shaded and unshaded areas withina plot (Einarsson and Milberg 1999). The light situation (L) in each plot wasdescribed using the following equation: L ¼ 100� L‐2þL0þL2ð Þ=3ð Þ, where L-2 isthe mean value of five recordings 2 h before noon divided by the specific recordingin a fully open area, L0 is the mean value of five recordings at noon divided by thespecific recording in a fully open area, etc. (Einarsson and Milberg 1999).


Statistical Analyses

To classify remnants of still active RWM according to their species composition, thespecies data set was composed of the 18 sample plots (relevés) of mowed RWM. Braun-Blanquet cover-abundance data for the species were converted into a 2 to 9 scale (vander Maarel 1979). This matrix was then subjected to divisive clustering – Two WayIndicator Species Analysis (TWINSPAN, Hill 1979), using WinTWINS version 2.3(Hill and Šmilauer 2005). Additionally, we applied two ordination methods –Detrended Correspondence Analysis (DCA) and Canonical Correspondence Analysis(CCA) (Hill and Gauch 1980; ter Braak 1986; Jongman et al. 1995). DCAwas used toestimate the heterogeneity in the species data. Gradient length for the first DCA axis was3.157, indicating that the unimodal ordinationmethods were suitable for the analysis. Torelate species composition to measured habitat variable (light), CCAwas used.

The second CCA analysis was conducted with all 41 relevés of RWM, to relatevariation in the species data set to both environmental variables – light and timesince abandonment. The last one represented a categorical variable and was re-codedinto four dummy variables - one for each possible level of the factor: i) active; ii) <5 yrsabandoned; iii) 5–15 yrs abandoned and iv) >15 yrs abandoned RWM.

To test whether environmental variables were significantly related to speciescomposition, we used the Monte Carlo permutation test (499 permutations). Effectof rare species was reduced by downweighting. The ordination methods (DCA,CCA) and visualization of their results were carried out using the Canoco andCanoDraw programs (ter Braak and Šmilauer 2002).

The relationship between species richness and light intensity for mowed andabandoned plots were determined using linear (Pearson's) correlation analysis inSPSS 15.0 (SPSS Inc. 2006). To test differences in number of species and lightconditions among four groups of plots (mowed and abandoned in different timeperiods), we used a one-way analysis of variance (ANOVA) (SPSS Inc. 2006), sincethe data were sufficiently normally distributed (Shapiro-Wilks and Lilliefors tests).Differences between treatments were assessed using the Tukey Honest SignificantDifference (HSD) test for unequal sample sizes (Spjotvoll and Stoline 1973).

Results

The total number of vascular plant species recorded in the herb layer of 41 plots(relevés) of studied RWM was 129, with 109 in the still-managed (mean=33±7 s.d.per plot, n=18) and 107 (mean=24±6 s.d. per plot, n=23) in the abandoned RWM.There are 87 common species, 22 species exclusive to the mowed RWM and 20exclusive to the abandoned RWM. All species are listed in Appendix, featured byspecies typical of grassland and deciduous woodland/shrubland (31.0% and 38.7%,respectively), followed by species of disturbed habitats (13.2%) and species withindifferent habitat preference (17.1%).

The TWINSPAN classification split the 18 relevés of still-managed RWM at thefirst cut level into two groups (clusters G1 and G2), which we defined as two mainvegetation types of active RWM. In comparison to group G1, relevés from group G2had a larger number and frequency of woodland and scrubland species (Appendix).


The most abundant are Brachypodium sylvaticum, Galanthus nivalis, Pulmonariaofficinalis, Galeobdolon montanum, Stachys sylvatica, Anemone ranunculoides,Asarum europaeum, Viola riviniana and Cerastium sylvaticum. Most of those specieswere also present in group G2, but with lower frequency. Many grassland species,however, that are frequent in G2 were rare or absent in G1, such as Arrhenatherumelatior, Ranunculus acris, Achillea millefolium agg., Trifolium pratense, Briza media,Trifolium repens, Centaurea jacea and Plantago media.

We found significant differences in light intensity between groups G1 and G2(P<0.001), with higher values in cluster G2 (Table 1). In contrary to the hypothesis,light intensity in mowed RWM was negatively correlated with the number of speciesper plot (R=-0.49, n=18, P<0.05).

The CCA with relevés of mowed RMW and light as the only environmentalvariable (Fig. 2) showed that the species composition of still-managed RWM wassignificantly related to light intensity (eigenvalue=0.33, P=0.002). The specieswith the lowest scores are those characteristically found in shaded plots, such asEuphorbia epithymoides, Galeobdolon montanum, Geum urbanum, Cerastiumsylvaticum, Stachys sylvatica and Paris quadrifolia. Carex hirta, Centaurea jacea,Briza media, Plantago media, Linum catharticum and Lotus corniculatus were someof the species receiving the highest scores in the first CCA, i.e., they are associatedwith plots with higher light intensity.

In the CCA of all 41 vegetation relevés made in active and abandoned RWM,relevés scores of axis 1 (eigenvalue=0.31) were positively correlated with light (R=0.67, P=0.002) and active RWM (R=0.60, P=0.001) (Fig. 3). There were nocorrelations between light and time since abandonment. The ordination biplotshowed species related to plots with higher light intensity abundant particularly instill-active (mowed) RWM (Fig. 3) on the right side. These are mainly species ofgrassland communities such as Leucanthemum ircutianum, Centaurea jacea,Trifolium pratense, Plantago media, Arrhenatherum elatior, Plantago media,Ranunculus acris and Achillea millefolium. On the left side are species that arerelated to plots with low light availability. Species, such as Geum urbanum, Parisquadrifolia, Stachys sylvatica, Galeobdolon montanum and Alliaria petiolata,common for more shaded plots of both active and abandoned RWM are at the

Table 1 Number of relevés, total and mean number of species and mean light intensity on 25 m2 plots infive groups of relevés in the riparian wooded meadows (RWM) along the Drava River in Slovenia

Active Abandoned

G1 G2 <5 yr 5–15 yr >15 yr

Number of relevés 7 11 8 8 7Total number of species 77 89 83 82 70Number of species per relevé:mean±s.d.

38a±7 29ab±4 27ab±6 22b±6 22b±3

Light (PAR) (%): mean±s.d. 14.4a±12.4 51.5b±17.5 23.8ab±17.7 32.8ab±31.4 23ab±24.2

G1 – Twinspan cluster No. 1; G2 – Twinspan cluster No. 2; s.d. – standard deviation. Means bearing thesame superscript letter are not significantly different (determined by the Tukey HSD at P<0.05).


bottom left. At the top left side were species, such as Prunus padus juv., Carex alba,Rubus fruticosus agg. and Corydalis cava, frequent in shaded and more than 5 yrsabandoned RWM. We found two neophyte species Solidago gigantea and Rudbeckialaciniata, which had the highest cover-abundance values on sunlit plots in earlysuccession stages. The number of species per plot for abandoned RWM showed nocorrelation with the light intensity (R=-0.05, n=23, P=0.79).

Discussion

A comparison of vegetation from the herb-layer on still-managed RWM revealedthat there exist two main types of wooded meadows in the area under study and thatlight availability is an important factor in explaining differences in speciescomposition; one type is less and one type more shaded. It is clear that traditionalmaintenance of riparian space developed two types of RWM, one closer to the forestand the other to grassland. This was confirmed with floristic analysis, where specieswith grassland and woodland habitat preferences were distinguished. The type,

Fig. 2 CCA ordination diagram of relevés (n=18, 109 species) of still-managed riparian woodedmeadows (RWM) along the Drava River in Slovenia with light as environmental variable. Eigenvalues:0.33 (axis 1), 0.30 (axis 2); 27.7% of variance in species is explained by both axes. Monte Carlo testsignificant at P<0.005. Shown species have the highest weight. Relevés were divided into two groups (G1and G2) with TWINSPAN analysis: ▪ – relevés from G1; □ – relevés from G2. Abbreviations of speciesare explained in Appendix


closer to the meadows seems to be the very archaical type, a kind of predecessor ofriparian meadows. The more shaded, RWM type closer to the forest is probably theremnant of medieval multifunctional woodland land-use.

Both the less shaded and more shaded RWM types are characterized by relativelyhigh species richness, typical of wooded meadows as described by other authors(e.g., Kull and Zobel 1991; Klimeš 1995, 1997; Klimeš et al. 2001; Mitlacher et al.2002). However, it is interesting to note that species richness for mowed RWMdecreased with increasing light availability, in contrast to what was found for

Fig. 3 CCA ordination diagram of all relevés (n=41, 129 species) and environmental variables (light andtime since abandonment) of active and abandoned riparian wooded meadows (RWM) along the DravaRiver in Slovenia. Number of species (No_Spec) per plot was added as supplementary variable (dashedline) without any effect on the analysis. Eigenvalues: 0.31 (axis 1), 0.15 (axis 2). Shown species have thehighest weight. Abbreviations: No_Spec – number of species per plot; Active – still-managed (mowed)RWM; Aband_1: <5 yrs abandoned RWM; Aband_2: 5–15 yrs abandoned RWM; Aband_3: >15 yrsabandoned RWM. Abbreviations of species are explained in Appendix


Scandinavian and Baltic wooded grasslands (Zobel et al. 1996; Einarsson andMilberg 1999; Mitlacher et al. 2002). Three facts can explain the high speciesrichness of the more shaded RWM type: i) the nutrient-rich riparian space is not afavorable habitat (with exceptions, of course) for oligotrophic, basiphilous andxerophilous species. Therefore, the species-rich contingents of, e.g., semi-drycalcareous grasslands (Bromion erecti) or thermophilous “Saum” vegetation(Geranion sanguinei) are present with only a few species. The species of the FalseOatgrass meadows (Arrenatherion) prevail, but the list of those species is far fromcomplete; ii) the species richness in northern (Scandinavian and Baltic) forests ismuch lower than in Central European hornbeam riparian forests. In the middle DravaRiver those forests are also enriched with a contingent of Illyrian species. Thefloristic mixture and favorable eutrophic conditions result in a species-richunderstorey; iii) the intermediate light level creates a habitat (or mosaics of micro-habitats) that enables both light-requiring and shade-tolerant species to occur(Einarsson and Milberg 1999). Going towards lighter (like in our meadow-like RWM)and darker environments might be associated with decreasing species richness.

The CCA analyses with all relevés also demonstrated that light availability was agood predictor of RWM species composition. Because more plots of meadow-likethan forest-like type of active RWM were sampled the symbol for active RWM liescloser to light-requiring species (grasslands species). Abandonment of mowing isassociated with profound changes in floristic composition. As succession occurs,light-requiring species are prohibited due to the reduction in the light available atground level, caused by shading effect of taller and broadleaved plants. Consequentlymore shade-tolerant species appear. Because many of those species are already presentin the type of active RWM close to forests, the species turnover during succession ismore pronounced in less shaded meadow-like RWM. However, species richnesssignificantly decreased in relation to abandonment only in comparison with the morespecies-rich forest-like type of active RWM. The colonization processes in abandonedwooded meadows appeared to be so slow that they were not recorded in our experiment.

Fresh, nutrient-rich riparian soils together with semi-shaded conditions afterabandonment seem to be ideal for the colonization of two invasive neophyte speciesSolidago gigantea and Rudbeckia laciniata. Their expansion after abandonment isvery rapid. In our study, the clear positive correlation between cover-abundancevalues of S. gigantea and light intensity (R=0.61, P=0.0017, data not shown)suggests that abandoned RWM are very susceptible to colonization until there isenough light. With the establishment of a shrub layer, their cover-abundance declines.

The remnants of riparian forests, stretching along the flood plains of Central andEastern Europe, are considered one of the most natural ecosystems in the prevailingagricultural landscapes. But naturalness of productive flooded forests is often onlythe consequence of abandonment of ancient land-use practices, like livestockgrazing, cutting between the trees, and litter collecting. The wilderness of high-productive riparian forests in Central Europe in many cases reflects the exaggeratedpoint of view of foresters’ interpretations (Accetto 1990; Mlinšek 1996). The woodyspecies and physiognomy are completely natural, but historical management (selectedtree cutting, pasturage, mowing) suggests the need to recognize the remains of a closeto natural ancient cultural landscape.


Conclusions

The results presented here indicate that the species composition of the studiedriparian wooded meadows depends strongly on light availability. There exist twodifferent vegetation types of RWM: the less shaded meadow-like and the moreshaded forest-like type. The latter shows higher species richness as a consequence i)of the intermediate light level that enables both, light-requiring and shade-tolerantspecies to occur and ii) of the species-rich riparian meadows, which contribute agreat number of understorey species. Long-term mowing abandonment causes theloss of many light-requiring grassland species in RWM. Although the riparianhornbeam forests are also very species rich, widespread abandonment produces areduction in floristic and habitat richness at the landscape scale on account of thedifferent species composition in mowed and abandoned RWM.

Appendix

A list of 129 vascular plant species recorded in the herb layer of the 41 relevés ofriparian wooded meadows (RWM) along the Drava River in Slovenia

Active Abandoned

G1 G2 <5 yr 5–15 yr >15 yr

Grassland communitiesValeriana officinalis Val off 43 . . . .Serratula tinctoria Ser tin 14 . . 13 .Thalictrum flavum Tha fla 29 . 13 13 .Arabidopsis thaliana Ara tha 71 9 25 63 43Angelica sylvestris Ang syl 57 9 13 13 29Centaura carniolica Cen car 71 36 50 38 14Ajuga reptans Aju rep 86 73 63 25 43Galium mollugo Gal mol 71 82 75 63 43Cirsium oleraceum Cir ole 57 27 38 25 14Colchicum autumnale Col aut 57 55 88 38 57Heracleum sphondylium Her sph 57 18 38 25 29Brachypodium pinnatum Bra pin 43 91 63 63 71Pimpinella major Pim maj 14 73 25 50 29Anthriscus sylvestris Ant syl 29 18 . . .Filipendula ulmaria Fil ulm 29 9 50 38 14Vicia cracca Vic cra 29 27 25 13 29Orchis militaris Orc mil 29 45 25 . 25Plantago lanceolata Pla lan 29 55 13 13 .Crepis biennis Cre bie 14 9 13 . .Euphorbia cyparissias Eup cyp 14 27 25 13 14Prunella vulgaris Pru vul 14 45 25 . .Ranunculus acris Ran acr 14 91 50 . .Trifolium pratense Tri pra 14 64 . . .Arrhenatherum elatior Arr ela . 100 13 . .Achillea millefolium Ach mil . 73 38 . .Briza media Bri med . 55 . . .Trifolium repens Tri rep . 55 . . .Centaurea jacea Cen jac . 45 . . .


Active Abandoned

G1 G2 <5 yr 5–15 yr >15 yr

Plantago media Pla med . 45 13 . .Bromopsis erecta Bro ere . 36 . . .Lathyrus pratensis Lat pra . 36 . . .Pimpinella saxifraga Pim sax . 27 . . .Trifolium montanum Tri mon . 27 . . .Leucanthemum ircutianum Leu irc . 18 . . .Linum catharcticum Lin cat . 18 . . .Lotus corniculatus Lot cor . 18 . 14Salvia pratensis Sal pra . 18 . . .Agrimonia eupatoria Agr eup . 9 13 . .Knautia arvensis Kna arv . 9 . . .Sanguisorba officinalis San off . 9 . . .Woodland and shrubland communitiesHumulus lupulus Hum lup 14 . . 13 29Euphorbia epithymoides Eup epi 29 . . . .Polygonatum latifolium Pol lat 43 . 13 . .Festuca gigantea Fes gig 43 . . 13 14Geum urbanum Geu urb 43 . . 25 29Paris quadrifolia Par qua 43 . 13 13 29Cerastium sylvaticum Cer syl 71 . 13 13 .Galeobdolon montanum Gal mon 71 . 63 13 .Stachys sylvatica Sta syl 71 . 13 25 29Anemone ranunculoides Ane ran 71 18 13 13 .Asarum europaeum Asa eur 71 18 75 75 71Viola riviniana Vio riv 71 9 . 13 .Brachypodium sylvaticum Bra syl 100 18 75 63 100Galanthus nivalis Gal niv 100 18 13 38 14Pulmonaria officinalis Pul off 100 18 38 63 57Salvia glutinosa Sal glu 86 45 75 75 71Knautia drymea Kna dry 71 55 50 25 29Clematis vitalba Cle vit 71 45 13 63 43Cruciata glabra Cru gla 57 18 75 25 29Dryopteris filix-mas Dry fil 43 9 13 13 .Lunaria rediviva Lun red 43 36 . 25 .Symphytum tuberosum Sym tub 43 27 25 13 29Gagea lutea Gag lut 29 27 13 . 29Fragaria vesca Fra ves 14 9 38 13 14Anemone trifolia Ane tri 14 9 . 13 .Cyclamen purpurascens Cyc pur 14 18 13 13 .Listera ovata Lis ova 14 9 13 . .Primula vulgaris Pri vul 29 82 63 50 14Leontodon incanus Leo inc . 100 13 . .Viola hirta Vio hir . 64 25 13 43Geranium phaeum Ger pha . 36 . .Clinopodium vulgare Cli vul . 18 13 . 14Peucedanum oreoselinum Peu ore . 9 13 . .Carex alba Car alb . 9 13 25 14Corydalis cava Cor cav . 9 13 25 43Rubus fruticosus Rub fru . 9 25 38 43Vinca minor Vin min . 9 . 13 14Euonymus europaea juv. Euo eur . . 38 75 71Ligustrum vulgare juv. Lig vul . . 25 25 43Crataegus monogyna juv. Cra mon . . . 50 29Ulmus laevis juv. Ulm lae . . 13 13 29


Active Abandoned

G1 G2 <5 yr 5–15 yr >15 yr

Fraxinus excelsior juv. Fra exc . . . 13 43Prunus padus juv. Pru pad . . . 13 29Quercus robur juv. Que rob . . . 25 14Acer campestre juv. Ace cam . . . 13 14Rhamnus catharticus juv. Rha cat . . 13 . 14Frangula alnus juv. Fra aln . . . . 29Populus alba juv. Pop alb . . 25 . .Lythrum salicaria Lyt sal . . . 13 .Cornus sanguinea juv. Cor san . . 25 13 29Disturbed habitatsImpatiens glandulifera Imp gla 14 . 25 . 14Eupatorium cannabinum Eup can 29 . 13 13 .Stelaria montana Ste mon 100 . 50 50 14Erigeron anuus Eri anu 71 9 75 25 14Cardamine impatiens Car imp 71 . 25 13 .Alliaria petiolata All pet 86 18 . 25 43Solidago gigantea Sol gig 100 45 75 100 86Aegopodium podagraria Aeg pod 86 100 88 50 100Cruciata laevipes Cru lae 43 18 13 13 14Rudbeckia laciniata Rud lac 57 18 63 63 14Oxalis corniculata Oxa cor 43 9 . . .Carex hirta Car hir 29 18 . 13 .Glechoma hederacea Gle hed 23 73 38 13 14Cardamine hirsuta Car hir . 18 . . .Lamium maculatum Lam mac . 9 . 13 .Pastinaca sativa Pas sat . 9 . . .Urtica dioica Urt dio . . 38 50 .Indifferent habitat preferenceOxalis acetosella Oxa ace 14 . 13 . 14Lysimachia vulgaris Lys vul 43 . 25 13 100Melica nutans Mel nut 71 . 75 38 57Rubus caesius Rub cae 86 . 25 38 43Dactylis glomerata Dac glom 86 82 50 13 .Anemone nemorosa Ane nem 71 64 38 13 29Taraxacum officinale Tar off 71 27 13 . .Ficaria verna Fic ver 43 18 13 . 14Galeopsis tetrahit Gal tet 43 9 25 13 43Hedera helix Hed hel 43 27 . 25 57Lysimachia nummularia Lys num 43 27 . 13 14Symphytum officinale Sym off 43 9 . . 14Vicia sepium Vic sep 14 9 13 . .Cardaminopsis arenosa Car are . 27 . . 29Ranunculus repens Ran rep . 36 . . .Solanum dulcamara Sol dul . 18 . 13 .Sambucus nigra juv. Sam nig . . 13 . 14Robinia pseudacacia juv. Rob pse . . 13 25 29Deschampsia cespitosa Des cae . . 38 13 .Calamagrosits arundinacea Cal aru . . . 25 .Campanula persicifolia Cam per . . 25 . .Dactlylorhiza maculata Dac mac . . 13 13 .

G1 – Twinspan cluster Nr. 1; G2 – Twinspan cluster Nr. 2. Species are grouped according to habitatpreferences by Ellenberg et al. (1992).Species values are percentage frequencies.


Acknowledgements The research for this paper was funded by the program group “Biodiversity” (P1–0078) and the research project “Biodiversity gradient in relation to biogeographic, altitudinal andecological factors” (J1–6577) both headed by B. Kryštufek and granted fby the Slovenian ResearchAgency. We are grateful to our ex-students Helena Rižnar and Melita Caf, who performed somemeasurements and collaborated in the field work. We wish to express our gratitude to ornithologist BorutŠtumberger, an expert on the Drava region, who introduced us to the area of RWM and encouraged us todo further research. We also thank the three anonymous referees for constructive comments on themanuscript.

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Received: 25 September 2007 /Revised: 28 May 2008 /Accepted: 9 June 2008 /Published online: 2 December 2008


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