35 Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia Abstract The aim of the study is to characterise herbaceous vegetation (meadows and ruderal communities) remaining after several decades of protection and compare it to the vegetation of currently managed local sites in the Central Forest Reserve, Tver Oblast, Russia. Cluster analysis of the communities was based on 209 relevés, while their ecological features were analysed using phytoindication assessment. The analyses revealed four types of herbaceous communities: managed mesic meadows, abandoned mesic meadows, tall-herb meadowsweet communities and ruderal tall-herb communities. These four types differ in management, floristic composition and ecological conditions as well as in coenotic and functional group shares (including forbs, graminoids and woody species). The occurrence of these species groups determines the current state of the herbaceous communities. Our study revealed that mesic meadows have retained all the key meadow features for more than 25 years without any management, although their area has shrunk and shares of coenotic and functional groups have changed. The observed herbaceous communities encompass around 40% of the reserve flora including four red list species and 16 alien species. Izvleček V članku smo opisali zeliščno vegetacijo (travišča in ruderalne združbe), ki se je ohranila po nekaj desetletjih zaščite in jo primerjali s sedanjo vegetacijo na gospodarjenih rastiščih v Osrednjem gozdnem rezervatu, Oblast Tver, Rusija. S klastrsko analizo smo klasificirali 209 vegetacijskih popisov, njihove rastiščne značilnosti pa smo analizirali s pomočjo fitoindikacije. Z analizo smo ugotovili 4 tipe zeliščne vegetacije: gojena vlažna travišča, opuščena vlažna travišča, visoka steblikovja brestovolistnega oslada in ruderalna visoka steblikovja. Štiri rastlinske združbe se razlikujejo po načinu gospodarjenja, vrstni sestavi in rastiščnih razmerah kot tudi glede na delež cenotskih in funkcionalnih skupin (delež zelišč, trav in lesnatih vrst). Pojavljanje teh skupin vrst označuje trenutno stanje zeliščnih združb. Z raziskavo smo pokazali, da so vlažna travišča ohranila vse ključne značilnosti travišč tudi po 25. letih brez kakršnega koli gospodarjenja, vendar se je njihova površina zmanjšala, deleži cenotskih in funkcionalnih skupin pa so se spremenili. V obravnavanih združbah smo zabeležili 40% celotne flore rezervata, vključno s štirimi vrstami z rdečega seznama in 16 tujerodnih vrst. Keywords: abandonment, coenotic groups, Indicator Species Analysis (ISA), mainland meadows, species richness. Ključne besede: opuščanje, cenotske skupine, analiza indikatorskih vrst (ISA), celinska travišča, vrstna pestrost. Received: 28. 2. 2017 Revision received: 20. 6. 2017 Accepted: 18. 12. 2017 Co-ordinating Editor: Jacqueline Loos 1 Department of Geobotany, Biological faculty, Lomonosov Moscow State University, Leninskye Gory, 1-12, 119234, Moscow, Russia. E-mail: valentinka_bo@mail.ru, sciapoda@mail.ru * Corresponding author Oxana Cherednichenko1, Valentina Borodulina1,* DOI: 10.1515/hacq-2017-001517/1 • 2018, 35–59 O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 36 17/1 • 2018, 35–59 Introduction Palaearctic grasslands are known for their high biodiver- sity in many taxa (Habel et al. 2013, Dengler et al. 2014, Dengler et al. 2016). In particular, European grasslands are among the most diverse ecosystems in the world (Wil- son et al. 2012, Dengler et al. 2014). Extensively used mainland meadows are considered to be the most species- rich in small spatial scales (Kull & Zobel 1991). Today many of the grassland ecosystems are threatened by the change of land use towards intensification, abandonment and afforestation. Under these changing conditions, the current state of grasslands is still poorly studied. Hence our research is aimed at filling this knowledge gap. Tradi- tionally, the term “grassland (vegetation)” is understood as formation dominated by grasses (or graminoids), usu- ally with a single-layered structure and sometimes (in case of wooded savannas or savannoid vegetation) with an open woody plant cover (modified after Rutherford et al. 2006, reviewed by Mucina et al. 2016). However, in later studies (Dengler et al. 2014) on palaearctic grasslands, steppes are considered as dry grasslands and meadows as mesic grasslands. According to Mucina et al. (2016), meadow (vegetation) is a plant formation dominated by grasses (or other graminoids) and herbs usually found in humid to mesic habitats. Most European meadows are of anthropogenic origin, dependent on regular management such as mowing (Mucina et al. 2016); therefore, they are considered as semi-natural communities. Semi-natural grasslands are an essential part of the cul- tural landscape of Europe and resulted from centuries or millennia of low-intensity land use since the beginning of the Neolithic period (Pott 1995, Poschlod et al. 2009, Ellenberg & Leuschner 2010, Hejcman et al. 2013, re- viewed by Dengler et al. 2014). Such grasslands are mainly used for grazing or hay-making, but might also represent successional stages of abandoned arable fields. Moreover, grasslands provide essential ecosystem services (Bruun & Ejrnæs 1998, Gibson 2009, Zavaleta et al. 2010, reviewed by Moeslund et al. 2013) and are important habitats for many light-demanding and nitrophobic red list plant spe- cies (Meadows of Nechernozemie 1984, Bulokhov 2001, Semenishchenkov 2009, Dengler et al. 2014). The pres- ence of a wide range of redlist species makes extensively managed meadows biodiversity hot-spots. Mainland meadows of the East European forest zone are formed and maintained by human activities. In most studies, anthropogenic impact and soil pH have been shown to be the most important determinants of local vegetation patterns in grasslands (Saunders et al. 1991, Hansson & Fogelfors 2000, Stevens et al. 2004, Maskell et al. 2010, Reitalu et al. 2010, reviewed by Moeslund et al. 2013), although some studies also mention the role of other abiotic factors and topography (Moeslund et al. 2013). The impact of abiotic and biotic factors on species assemblage composition (abundance and species rich- ness) is of major interest in conservation ecology (Habel et al. 2013). Semi-natural grasslands were widespread in Europe at the turn of the XX century. During the last century, due to changes in land use, the area of semi-natural grass- lands decreased across whole Europe (van Dijk 1991, Rūsiņa 2006, Zelnik & Čarni 2013, Dengler et al. 2014). Not only has the area declined dramatically, but their con- dition has also suffered due to factors such as fragmenta- tion, improper management and using synthetic fertilisers and pesticides, increased cutting frequency and lowered cutting height in meadows, higher stocking rates on pas- tures as well as frequent re-sowing with artificial seed mix- tures (van Dijk 1991, Myśliwy & Bosiacka 2009, Dengler et al. 2014). Further sources of threat include eutrophica- tion through airborne nitrogen deposition and, in some cases, biotic invasions (Janišová et al. 2011). Grassland loss through transformation into arable land, quarries or settlements and infrastructure also occurred; however, it is generally of lower importance than the twin threats of intensification and abandonment (Valkó et al. 2012). This trend leads to a strong decrease in biodiversity. Successional changes in grassland communities, with many red-listed species disappearing from them, is still taking place. Large areas of grasslands become overgrown with shrub and woody vegetation, which has negative consequences for conservation and restoration of mead- ow communities (Falińska 1999, Korobeynikova 2002, Helm et al. 2006, Gornov & Evstigneev 2011, Evstigneev 2012, Budzhak et al. 2016). Nowadays species-rich semi- natural grasslands, also termed High Nature Value grass- land (Veen et al. 2009), are under threat and included in the European Red List of Habitats (2016). Today many of the European grassland ecosystems of high conservation value are threatened by the change of the very land use that formerly created and maintained them, i.e. replace- ment of traditional management systems with intensifica- tion, abandonment, afforestation etc. (Valkó et al. 2012, WallisDeVries et al. 2002, Öckinger et al. 2006, Veen et al. 2009, reviewed by Habel et al. 2013). According to van Dijk (1991), important efforts have been made for the protection of threatened habitats and species over the past one hundred years of nature conser- vation. However, grassland and arable land vegetation has been neglected in the development of effective conser- vation actions (van Dijk 1991). Traditional biodiversity conservation started with nature reserves from which all kinds of human impact (including grazing) were excluded. O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 37 17/1 • 2018, 35–59 In the case of European grasslands, strict exclusion of hu- man land use most often leads to a loss of grassland area due to succession into scrubland and forest (Pärtel et al. 2005). They do not fit the usual protection systems be- cause they are maintained by anthropogenic influence. Therefore, meadows in the territories of reserves are also under threat. However, the actual drivers of local grass- land plant diversity patterns are not fully understood, and hence, conserving and managing these semi-natural habi- tats is challenging (Moeslund et al. 2013, Zelnik & Čarni 2013, Habel et al. 2013). The aim of our study carried out in NW Russia is to survey grasslands remaining after several decades of pro- tection regime in the Central Forest Nature Reserve. We addressed the following questions: 1) How strong is the variety of grassland vegetation in the reserve and adjacent territories; 2) How variable are different vegetation types in terms of species richness, species diversity, type of man- agement and site conditions; 3) What are the differences in functional and coenotic structure of the floristic com- position in different community types. Materials and methods Study area The research was carried out in the territory of the Cen- tral Forest State Nature Biosphere Reserve (CFR), Tver Oblast, Russia (Figure 1).The Central Forest Reserve was established in 1931. The core area comprises 21,348 ha, the transition area 46,061 ha (Minyaev & Konechnaya 1976). The reserve is located in the SW part of the Valdai Upland within the main Caspian-Baltic watershed of the Russian plain (Latitude: 56° 26' – 56° 39' N, Longitude: 32° 29' – 33° 01' E). The relief is mostly flat, with only low and generally gentle slopes of riverbanks and streams. The soils are sod-podzolic and gley-podzolic (Minayeva & Shaposhnikov 1999). The climate is humid continental (Minayeva & Sha- poshnikov 1999). The mean annual rainfall for the period 1963–2014 is 760 mm (510 to 1050 mm in different years). The mean January temperature is −8.6 °C (the ab- solute minimum is −39.4° C), and the mean July temper- ature is +16.9 °C (the absolute maximum is +36.5 °C). Long-term climatic data were obtained from the meteo- rological station “Forest reserve” (http://www.clgz.ru/ page.php?al=sci_monitor), which is available as database from the archive of the Central Forest Reserve. The territory of the Central Forest Reserve belongs to the mixed coniferous forest zone. Forests represent the prevailing vegetation type: spruce forests cover 47% of the whole area, mixed secondary forests 40%. Boggy pine forests occupy 9%, oligotrophic and mesotrophic mires 4%. The area of meadows is much smaller and occupies less than 1% of the territory (Kurayeva et al. 1999). According to floristic studies of Konechnaya (2012), 43 red-listed species were noted in the territory of the reserve. These species are listed in the IUCN Red List of Threatened Species of Russia (Red Data Book of the Rus- sian Federation) (three species) and the IUCN Red List of Threatened Species of Tver Oblast (Red Data Book of Tver Oblast) (40 species). Object of research The object of our research is herbaceous vegetation (meadows and ruderal communities) under protection regime in the CFR and under agricultural use in adjacent territories. All biosphere reserves must contain zones free of human interference (core area), buffer zones common- ly used for cooperative activities compatible with sound ecological practices such as educational and research ac- tivities and transition areas where restricted agricultural use is allowed (http://www.unesco.org/). Thus, the grass- lands of the core area have not been in use due to the protection regime since the early 1960s. In the grasslands of the transition area, agricultural use is allowed, but most of them are abandoned nowadays. In addition to the pro- tected herbaceous vegetation, we explored managed grass- lands to identify features associated with management. In the core area of the CFR, herbaceous vegetation occurs only as small patches in the places of former settlements (villages, farmsteads, forest huts). Most of Figure 1: Location of the study area. The red dot indicates the reserve location. The red line is the border of Tver Oblast. Slika 1: Lokacija preučevanega območja. Rdeča točka prikazuje lokacijo rezervata. Rdeča črta je meja Oblasti Tver. O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 38 17/1 • 2018, 35–59 the grasslands were abandoned 30–60 years ago. After that, the vegetation of hay meadows and pastures began to change. Some communities still look like grass-forb meadows. Others have completely changed into shrub or forest vegetation. Large areas are covered by ruderal tall-herb stands with dominance of forbs (Urtica di- oica, Anthriscus sylvestris, Chamaenerion angustifolium etc.). These communities can be found in the places of abandoned houses, backyards and vegetable gardens in former villages. In the transition area and the (one kilo- metre) buffer zone, meadows occupy large areas around villages. Most of them are abandoned, others are used at present. The managed meadows are comparatively small and characterised by mixed extensive usage (mowing and grazing). They were studied in the vicinities of CFR headquarters and around inhabited villages situated in both buffer zone and transition area of the reserve and in adjacent territories. Data collection The data discussed below was collected in 2013 and 2014. The two years were similar in climatic conditions: The annual rainfalls were 715.6 mm and 725.7 mm, re- spectively. The mean January temperatures were −9.8 °C and −10.0 °C, the mean July temperatures +17.0 °C and +18.3 °C (respectively), which is close to the long-term average. Our study was based on 209 relevés of herbaceous vegetation. We made 88 relevés in six sites (Ovsyaniki, Mezha, Krasnoye, Starosel’e, Zapovedniy, Fyodorovskoe) in the South of the reserve in 2013 and 111 relevés in nine sites (Moshary, Kruglaya Luka, Trozhkov Lug, Bol’shoe Makarovo, Shlyuz, Gusevka, Osinovka, Pogorelka, Gor- bunovka) in the North and 10 relevés (Zapovedniy) in the South of the reserve in 2014 (Figure 2). Thus, we in- vestigated most of the meadows in the core area of the CFR. The plot size of each relevé was 100 m2, which is considered appropriate for grassland vegetation (Mirkin & Naumova 2012, Baranova et al. 2016, Belonovskaya et al. 2016, Budzhak et al. 2016). Our relevés repre- sent almost all herbaceous communities in the reserve according to physiognomy and land use type. Samples were placed using the method of preferential selection of sites considered typical and homogenous (Gauch 1982). The relevés were compiled in homogenous appearing contours of vegetation along the visible gradients of the relief, as a rule from the edge to the centre of the meadow to cover the entire diversity of plant communities of each site. The distance between the plots depended on the size of the homogeneous plant community contour: the smaller the contours, the smaller the distance between Figure 2: Location of the explored sites in the territory of the reserve (no frame – explored in 2013; purple frame – explored in 2014). Slika 2: Lokacija preučevanih rastišč na območju rezervata (brez okvirja – preučevano leta 2013; vijoličast okvir – preučevano leta 2014). O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 39 17/1 • 2018, 35–59 the plots. Nevertheless, the minimum distance between neighbouring plots was 10 m, as was the minimum dis- tance to the nearest forest edge. If there were single trees in the meadow, the plots were placed to exclude the in- fluence of tree roots. Shading during the day took place on abandoned sites in some cases due to the small area of these sites. To identify specific features of herbaceous vegetation under protection regime, we took relevés of abandoned communities in the core area as well as managed mead- ows in buffer zone, transition area and adjacent territo- ries. Within the sample plots we collected the following data: (1) species composition of vascular plants; (2) spe- cies abundance (according to the scale of Braun-Blanquet (1964)); (3) total live vegetation cover (as area percentage); (4) height of the herbage layer, (5) presence and character of disturbance, (6) management pattern, (7) geodata (us- ing a Garmin GPS), (8) micro- and mesorelief features. Species names are given after Cherepanov (1995). Especially, we put stress on the presence of two particu- lar groups of species as related to the type of community and the management: red list species and alien (including invasive) ones. The list of alien and invasive species fol- lows Vinogradova et al. (2011). The list of red list spe- cies follows the Red Data book of the Russian Federation (plants and fungi) (2008) and the Red Data Book of Tver Oblast (2002). Data analysis Classification of vegetation was carried out by clus- ter analysis (flexible beta method using PC-ORD v.6 software, β = −0.25). To check the significance of the differentiated clusters, we then proceeded with a Multi-Response Permutation Procedure based on Eu- clidian (Pythagorean) dissimilarity distance measure (Peck 2010, McCune & Mefford 2011). Diagnostic spe- cies of the vegetation units (clusters) were identified us- ing Indicator Species Analysis (phi coefficient (Tichý & Chytrý 2006)) with randomisation test realised in the PC-ORD v.6 software (Peck 2010, McCune & Mefford 2011). Species with phi values above 0.25 were consid- ered to be diagnostic and those with values above 0.5 as highly diagnostic (Chytrý 2007). The site conditions were characterised by calculating cover-weighted averages of Landolt’s and Ramensky’s eco- logical indicator values (Ramensky et al. 1956, Landolt et al. 2010) for each plot using EcoScale v.5 software (Grokh- lina & Khanina 2015). We choose four Landolt indicators for the analysis (F – soil moisture, N – soil nitrogen pool, L – light, R – soil reaction) and one Ramensky indicator (PD1 – grazing intensity for the forest zone). The functional composition of community types was studied by calculating shares of forbs and graminoids in the total species number for each relevé. The coenot- ic composition was analysed according to Ulanova & Zhmylev (2014). Coenotic groups are species assemblages confined to main vegetation types of the region (Yurtzev & Kamelin 1991). We used the system of coenotic groups proposed by Ulanova and Zhmylev for the Moscow re- gion. They reveal coenotic affiliation of 1209 species of vascular plants. As a result, 13 coenotic groups could be distinguished: forest species – grow only or predomi- nantly in forest communities (including floodplain wil- lows); meadow species – only or predominantly in mead- ow communities (including steppe meadows and large glades); mire species – only or predominantly in mires of different types; aquatic species – only or predominantly in water and riparian communities (including temporary ponds, streams and springs); weed species – only or pre- dominantly in synanthropic communities; forest-mead- ow – in forests and meadows; forest-mire – in forests and mires; meadow-aquatic – grow in meadows and riparian water communities; meadow-mire – in meadows and mires; aquatic-mire – in riparian water communities and mires; weed-forest – in synanthropic and forest communi- ties; weed-meadow – in synanthropic and meadow com- munities; weed-mire species – in synanthropic and mire communities. The distribution of species recorded in our study among these groups can be found in Appendix 1. In this analysis, only the native fraction of the flora (257 species) was included; the alien species became the object of another analysis. In total, the flora of the inves- tigated meadows included species from 12 out of 13 coe- notic groups. There were no species of the aquatic species group. In addition, 12 species (four of which belonged to the genus Pilosella and two to the genus Dactylorhiza) were absent in the species list of Ulanova & Zhmylev (2014). For the analysis we have chosen the coenotic groups that are represented in all types of communities and have more than 25 species. Thus, we used the four coenotic groups of forest, meadow, forest-meadow and weed-meadow species. We did not include weed species in the analysis because they were poorly represented in two types of communities. Diversity was assessed using Simpson diversity index (1-D) (Magurran 2004), which was calculated in PC- ORD v.6. Species richness was estimated as the total number of vascular plant species per plot (100 m²). We used Kruskal-Wallis tests with post-hoc multiple compar- isons of mean ranks in Statistica v.10 to compare numeri- cal parameters of clusters revealed: ecological indicator values, species richness, Simpson diversity index, herbage height, shares of coenotic and functional species groups. O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 40 17/1 • 2018, 35–59 Results Diversity of herbaceous communities Cluster analysis revealed four herbaceous community types, which differed by physiognomic, ecological and topological criteria (Table 1). The significance of species composition differences between the groups was found by Multi-Response Permutation procedure: A = 0.14, p < 0.0001 (A = chance-corrected within-group agree- ment). Appendix 2 shows diagnostic species for each type of the herbaceous communities according to the results of the indicator species analysis. Cluster 1 – managed mesic meadows (MMM) (Fig- ure 3a). All MMMs are located in the buffer zone and in the transition area (Fyodorovskoe site), where they are situated around villages and in adjacent territories (Moshary site). This type comprises shortgrass commu- nities with dominance of grasses and meadow forbs on flat places and upper parts of gentle slopes. Diagnostic species were: Potentilla anserina, Leontodon autumnalis, Taraxacum officinale, Plantago major, Cynosurus cristatus etc. (Appendix 2). We found a total of 177 species of vascular plants (Table 1) in the type. These meadows are characterised by mixed usage with moderate grazing and irregular mowing. Some sites are currently overgrazed. Cluster 2 – abandoned mesic meadows (AMM) are mostly located in the core area and in the buffer zone and confined to habitats similar to the previous (Figure 3b). In the past these communities were mown, but this us- age was ceased more than 25 years ago (in some cases 50–60 years ago). However, until now these communi- ties still appear as typical meadows with dominance of forbs and grasses. Diagnostic species were: Viola canina, Trollius europaeus, Acetosa pratensis, Hieracium umbel- latum, Potentilla erecta etc. (Appendix 2). In AMM we found 182 species of vascular plants (Table 1). Cluster 3 – meadowsweet tall-herb communities (MTC) (dominated by Filipendula ulmaria) form tiny patches lo- cated in the core area and in the buffer zone (Figure 3c). These communities developed on wet and mesic soils in small relief depressions and along temporary streams, probably after abandonment of hayfields. Diagnostic species were: Filipendula ulmaria, Viola palustris, Galium palustre, Crepis paludosa, Cirsium palustre etc. (Appen- dix 2). In MTC we found 145 species of vascular plants (Table 1). Cluster 4 – ruderal tall-herb communities (RTC) are located in the core and in the buffer zone of the reserve and occupy habitats heavily disturbed in the past, such as places of abandoned houses and kitchen gardens. They are currently not in use and totally covered with tall herbs, with ruderal and nitrophilous species like Anthriscus sylvestris, Dactylis glomerata, Cirsium arvense, Urtica dioica, Chamaenerion angustifolium as the main dominants (Figure 3d). We consider the above species as diagnostic (Appendix 2). In most cases RTCs are associ- ated with natural disturbance caused by wild boars root- ing for plant rhizomes. In RTC communities we found 156 species of vascular plants (Table 1). Table 1: Recognised herbaceous community types. Tabela 1: Obravnavane zeliščne združbe. Community type Number of relevès in cluster Total number of species Herbage height, mean ± s.e., cm Presence and character of disturbance Current state Management pattern Managed mesic meadows 51 177 58±4 Low-disturbed Managed Mowing, grazing Abandoned mesic meadows 79 182 75±2 - Abandoned Former hayfields and pastures Meadowsweet tall-herb communities 24 145 100±8 - Abandoned Former hayfields and pastures Ruderal tall-herb communities 55 156 140±4 Soil disturbance: rooting by wild boars Abandoned Former kitchen gardens O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 41 17/1 • 2018, 35–59 There are some differences in herbage height between MMM, MTC and RTC. In the first case, herbage is sig- nificantly shorter than in the last two cases. The RTC are significantly different from all other types of communi- ties, and the herbage is the tallest here. Abandoned me- sic meadows are intermediate between MMM and MTC and not significantly different from these types.Our phy- toindication approach revealed ecological features of the community types studied (Table 2). MTCs are character- ised by the highest values of soil moisture according to the Landolt scale (from wet to very wet).Values for the other community types are significantly lower (p <0.01) and not very different from each other; they all correspond to moderately wet soils (Landolt et al. 2010). Comparatively high values on the soil nitrogen pool scale are character- istic of the RTCs, which can be considered as moderately rich. The values for the other types range from poor to moderately rich soils, and their mean ranks do not differ significantly between each type, but they do differ sig- nificantly from the RTCs (p <0.01).The highest values of the light scale are characteristic of the MMMs. Indeed, managed meadows usually occupy large open spaces. All three types of abandoned communities are characterised by lower light availability because they usually have small areas located in glades etc. In addition, the tall herbage of MTCs and RTCs itself produces shade, which increases the proportion of shade-tolerant species in its near-sur- face layer. Nevertheless, all medians are between 3 and 4 (semi-shaded and open sunny places), so there are only slight differences, although they are statistically signifi- cant (p <0.01). Also, differences in the soil reaction scale were revealed. The lowest value on the soil reaction scale was observed in the MMMs (slightly acid) and the high- est in the RTCs (neutral). The AMMs and MTCs occupy an intermediate position: The former showed no differ- ences from MMMs, and the latter none from RTCs. In general, all values range from slightly acid to neutral soils (pH 4.5–7.5) according to the Landolt scale. Figure 3: Photos of community types: a) Managed mesic meadow mowed with a small tractor; b) Abandoned mesic meadow: a young spruce tree (Picea abies) on herbaceous background; c) Meadowsweet tall-herb community; d) Ruderal tall-herb community. Photos: O.V. Cherednichenko. Slika 3: Fotografije rastlinskih združb: a) gojena vlažna travišča, košena z majhnim traktorjem; b) opuščena vlažna travišča: mlade smreke (Picea abies) v ozadju; c) visoko steblikovje z brestovolistnim osladom; d) ruderalno visoko steblikovje. Fotografije: O. V. Cherednichenko. a b dc O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 42 17/1 • 2018, 35–59 The comparison of the types according to the grazing intensity scale (Ramensky et al. 1956) also revealed signif- icant differences (p<0.01, Table 2). For AMMs moderate influence of grazing was typical. For the other commu- nity types, the scale indicated a very weak effect or lack of grazing influence. Species richness and species diversity of herbaceous communities The flora of herbaceous communities investigated in the Central Forest Reserve consists of 273 species of vas- cular plants (from 163 genera and 50 families), that is 46.1% of its whole flora, which consists of 592 species (Konechnaya 2012). Moreover, the flora of the explored herbaceous communities accounts for 17.3% of the Tver Oblast checklist (1579 species) (Notov 2005). The following graph shows no significant differences in species richness (Figure 4) between MMM and AMM (medians are 47 and 48, respectively). The abandoned mesic meadows have the highest values of this parameter, which exceed by far those of MTC and RTC (medians are 39 and 27, respectively). The latter type is significant- ly poorer in species due to the dominance of tall herbs. The maximum number of species (65) per 100 m2 was observed in MMM. The minimum number of species (9) per 100 m2 was noted in RTC. The analysis of species di- versity demonstrates that MMM and AMM have higher diversity than MTC and RTC. At the same time there is no significant difference in species diversity between MMM and AMM. Ratio between graminoids and forbs The share of graminoid species (with respect to the to- tal number of species per relevé) is significantly higher in MMM than in other types. The smallest fraction of graminoid species is found in RTC. Abandoned mesic meadows and meadowsweet communities are not signifi- cantly different by this parameter (Figure 5a). The share of forbs in MMM is significantly lower than in all tall- herb types (MTC and RTC), but not significantly differ- ent from the fraction of forbs in AMM (Figure 5b). Ratio of coenotic groups For the analysis we selected some coenotic groups well represented in number of species in our dataset, i.e. for- est, meadow, forest-meadow and ruderal-meadow species group. As shown in Figure 6, the composition of coenotic groups differs significantly between all community types. Though the largest part of the species totals is always Table 2: Ecological indicator values for community types revealed. Landolt scale: F – soil moisture, N – soil nitrogen pool, L – light, R – soil reaction; Ramensky scale: PD1 – grazing intensity. Superscripts refer to significant differences between groups (p < 0.01, Kruskal-Wallis test). Tabela 2: Ekološke indikatorske vrednosti za posamezne združbe. Landoltove vrednosti: F – vlažnost, N – dušik v tleh, L – svet- loba, R – reakcija tal; Vrednosti po Ramenskem: PD1 – intenzivnost paše. Nadpisane vrednosti prikazujejo statistično značilne razlike med skupinami (p < 0.01, Kruskal-Wallis test). Community type F N L R PD1 MMM 2.99 A 3.17 A 3.60 A 2.86 A 3.81 A AMM 3.11 A 3.08 B 3.43 A 2.88 AB 3.32 B MTC 3.62 B 3.19 C 3.27 A 2.97 BC 3.27 C RTC 3.12 A 3.63 C 3.24 B 2.96 C 3.02 BC Figure 4: Species richness in four community types: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Capitals refer to significant differences between groups (p<0.05, Kruskal-Wallis test). Slika 4: Vrstna pestrost v štirih združbah: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Črke kažejo na statistično značilne razlike med skupinami (p<0.05, Kruskal-Wallis test). O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 43 17/1 • 2018, 35–59 Figure 6: Coenotic groups in four community types: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Capitals refer to significant differences between groups (p<0.05, Kruskal-Wallis test). Slika 6: Cenotske skupine v štirih združbah: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Črke kažejo na statistično značilne razlike med skupi- nami (p<0.05, Kruskal-Wallis test). Figure 5: Graminoid (a) and forb (b) diversity in four community types: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Capitals refer to significant differences between groups (p<0.05, Kruskal-Wallis test). Slika 5: Pestrost trav (a) in zelišč (b) v štirih združbah: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Črke kažejo na statistično značilne razlike med skupinami (p<0.05, Kruskal-Wallis test). ba ba made up of meadow species (median from 30 to 45%), there are significant differences in the share of the group between different community types. Thus we can divide the studied types into two groups by the ratio of meadow species in the herbage: In MMM and AMM, the partici- pation of meadow species is remarkably higher than in RTC and MTC (Figure 6a). The share of forest species ranges from 2% to 11%, but it is significantly lower in the MMM versus the three other types, which are all abandoned and appear similar by this parameter (Figure 6b). Forest-meadow species (Figure 6c) are best represented in AMM (median 24%), obviously due to the spreading of these species in the absence of agricultural use. Also, a high share of forest-meadow spe- O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 44 17/1 • 2018, 35–59 cies is characteristic of RTC (but its share is significantly lower than in AMM). The lowest share of these species occurs in MMM (median 16%). The largest participa- tion of weed-meadow species is found in MMM (median 14%) (Figure 6d). This group is comparatively well repre- sented in RTC, too (median 7%), its share being signifi- cantly lower than in MMM, but significantly higher than in AMM and MTC (medians 5% and 4%, respectively). Here, the high share of weed-meadow species in managed meadows and ruderal communities suggests a stronger disturbance intensity than in the two other groups. Thus, the spectra of the coenotic groups indicate both the in- tensity of disturbance and the successional status of the herbaceous communities investigated. Encroachment of trees A total of 29 woody species were observed in the four community types. The AMM are the richest in woody species (18 species, 72% frequency). MMM and RTC are similar on the number of woody species and their occurrence: 10 species, 25% frequency in MMM and 10 species, 33% frequency in RTC. MTC occupies an inter- mediate position by these parameters between the previ- ous two groups and AMM (13 species, 58% frequency). Thus, the frequency of woody species is rather low in dis- turbed habitats (MMM and RTC). The most frequently occurring woody species are not the same in different habitats. Salix caprea and Salix myrsinifolia have the highest frequency (10% each) in MMM; all the other species account for only 2–5%. The AMM are of particular interest for the highest frequency of Malus domestica (32%). Two other typical trees here are Betula pendula (25%) and Picea abies (20%). In MTC the common meadow willow Salix myrsinifolia (33%) has the highest frequency, along with some moisture- demanding species: Alnus incana, Betula pubescens, Salix aurita (21% each). In RTC Malus domestica is also com- paratively frequent (15%), while the occurrence of other species does not exceed 5%. Alien and invasive species Sixteen alien species occurred in the community types, including invasive plants (according to the list of Vino- gradova et al. (2011)) and cultivated species (Table 3). The number of alien species was highest in AMM, where we found ten species (six invasive and four cultivated). In MMM only seven species (five invasive and two cultivat- ed) were recorded, but here their frequency is the highest among all studied community types. The alien species number in RTC was rather low (three invasive and one cultivated species), while in MTC no alien species were observed at all (Table 3). Red list species In the herbaceous community types in the Central For- est Reserve, we found only four red list species recorded by Konechnaya (2012). Of those, Dactylorhiza baltica (D. longifolia) is listed in the Red Data Book of the Rus- sian Federation. The others (Salix phylicifolia, Coeloglos- sum viride, Gymnadenia conopsea) are in the Red Data Book of Tver Oblast (2002). All red list species were observed only in abandoned communities. In AMM we found three red list orchids (Gymnadenia conopsea, Coe- loglossum viride, Dactylorhiza baltica), while one species (Salix phylicifolia) is associated with MTC. dc Figure 6c and 6d O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 45 17/1 • 2018, 35–59 Discussion Our study revealed the coenotic, floristic and functional group diversity of herbaceous vegetation in the Central Forest Reserve to be connected with ecological condi- tions, productivity and type of management. Until now, the diversity of the reserve’s meadows had not been stud- ied. Our results supplement data on species diversity of mainland meadows in the forest zone. Species richness on studied mesic meadows is higher than on mainland meadows in other regions of the forest zone of northern and central Russia (Martynenko 1989, Rabotnov 1984, Marakulina & Degteva 2008). The results we have obtained indicate a high level of di- versity in the studied herbaceous communities and their importance for biodiversity conservation in the CFR because the meadow flora makes up more than 40% of the reserve’s whole flora, and the species richness in the studied meadows is comparatively high. Overgrowing of meadows under protection regime is also observed in some other reserves of the forest zone of Russia, for exam- ple in the Nizhnesvirsky Nature Reserve (Tikhodeeva et al. 2016) and in the Polistovsky Nature Reserve (Chered- nichenko 2014). Thus, meadows in nature reserves of the forest zone require a special protection regime, and an investigation of its features is needed to develop adequate conservation methods. Nevertheless, our study confirms that mesic meadows are quite stable systems, which retain their characteris- tics for a long time. In our case, more than 25 years af- ter abandonment, these communities still remain to be meadows. According to Tikhodeeva et al. (2016), woody plants appear and take root on meadows when there is no haymaking for 11–20 years or more; after that, meadows can become overgrown in different ways. If meadows are overgrown by single trees, they retain their structure and species composition for a long time (Tikhodeeva et al. 2016). However, after abandonment the functional com- position of meadows changes in course of time. Factors determining species richness are complicated and often act in opposite directions. An overview by Dengler et al. (2014) discusses the impact of the main leading factors: the availability of mineral nutrients and other soil conditions, topography, climate, intensity of disturbance etc. In spite of the supporting effect of in- creasing nutrient availability on biomass production and a positive productivity–diversity relationship, nega- tive effects of excess nutrients (especially phosphorus, nitrogen and potassium) on species diversity have been reported for different types of grassland vegetation (Jans- Table 3: Frequency (%) of alien species in the four community types. Invasive species are given after Vinogradova et al. (2011). Community types: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Tabela 3: Frekvenca (%) tujerodnih vrst v štirih združbah. Invazivne vrste so podane v skladu z Vinogradova et al. (2011). Združba: 1 – MMM, 2 – AMM, 3 – MTC, 4 – RTC. Alien species State Community type 1 2 3 4 Conyza canadensis Invasive - 1 - - Epilobium adenocaulon Invasive 14 1 - 2 Festuca arundinacea Invasive - 3 - 2 Juncus tenuis Invasive 35 4 - - Lepidotheca suaveolens Invasive 2 - - - Lolium perenne Invasive 12 - - - Malus domestica Invasive 2 32 - 15 Petasites hybridus Invasive - - - 2 Populus balsamifera Invasive - 1 - - Cornus alba Cultivated 2 - - - Dianthus barbatus Cultivated - 1 - - Lychnis chalcedonica Cultivated - 1 - - Populus longifolia Cultivated - 1 - - Rosa rugosa Cultivated - 1 - - Symphyotrichum sp. Cultivated - - - 1 Triticum aestivum Cultivated 4 - - - Number of species 7 10 0 4 O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 46 17/1 • 2018, 35–59 sens et al. 1998, Austrheim 2002, Hejcman et al. 2010, Merunková & Chytrý 2012 reviewed by Dengler et al. 2014). Our results also prove that ruderal nitrogen-rich communities are poorer in species than mesic meadows. Apart from site conditions, land use type and inten- sity can influence species diversity and floristic composi- tion (Wellstein et al. 2007, Dengler et al. 2014). In their investigation of mesic meadows in Germany, Wellstein et al. (2007) have shown that site conditions may have greater influence on species composition than manage- ment type. They explained this by the low-intensity us- age of these meadows. In our case, too, species richness and diversity and participation of graminoids and forbs in AMM were more similar to MMM than to other abandoned types (moister MTC and RTC on nitrogen- richer soils). However, there remain some differences in species composition between MMM and AMM, re- flected in coenotic group spectra and in grazing intensity according to Ramensky’s indicator values. Some features that managed and abandoned mesic meadows have in common can be explained by the fact that the present-day diversity patterns are more af- fected by former management systems than by present ones (Helm et al. 2006, Reitalu et al. 2014 reviewed by Dengler et al. 2014). We can assume that our aban- doned meadows bear some traces of the past manage- ment. The former hayfields and pastures also retain some traits of managed meadows, e.g. a significant proportion of graminoids and typical meadow forb species. Rud- eral communities are associated with high levels of dis- turbance in the past and differ greatly from the “real” meadows (grasslands) of the present. Their prevailing disturbance caused by wild boars in essence mimics the specific features of their former use – digging, natural “plowing” and “weeding”. The meadows in and around the Central Forest Re- serve are generally under haphazard management. They are mostly used as irregular hayfields and pastures for moderate grazing. However, we have seen a few inten- sively used sites. In our analysis we did not divide the meadows by the usage intensity, but considered all man- agement types as one group – one reason why we could not find any strong connection between management patterns and either species diversity or richness of mead- ows. Species numbers and Simpson index of MMM do not significantly differ from those of AMM, though some studies have shown that the highest species diver- sity is observed at an average level of grazing – something between heavy grazing and abandonment (Dengler et al. 2014). However, the coenotic group composition of the four community types showed significant differences. With a similar share of meadow species in MMM and AMM, there is a greater share of weed-meadow species in MMM, while forest and forest-meadow groups were better represented in AMM. An increase in the share and diversity of forb species after cessation of agricultural usage has been shown for different types of grasslands in the broad-leaved forest and the forest-steppe zone (Evstigneev 2012, Ronkin & Savchenko 2016). We observed the same trend in case of mesic mainland meadows of the northern forest zone. In abandoned mesic meadows, the share of forb species is significantly greater than in the managed ones, with a higher share of graminoids. As mentioned above, tree encroachment poses a seri- ous problem in abandoned grasslands. The appearance of woody species (trees and shrubs) favours the further in- vasion of forest herbs. This process can occur both in dry and wet grasslands and tends to lower biodiversity levels (Helm et al. 2006, Vassilev et al. 2011, Evstigneev 2012, Valkó et al. 2012, Ronkin & Savchenko 2016). The aban- doned meadows in the Central Forest Reserve are no exception. We find the highest woody-species diversity (and frequency) in all abandoned types of communities (mesic and moist with meadowsweet) that have experi- enced the lowest disturbance level. In disturbed commu- nities, encroachment of trees is not going on so fast. Also, abandoned mesic meadows are of particular interest for the highest frequency of Malus domestica (32%), which may be caused by the activities of brown bears that feed on apples in abandoned villages and spread the seeds (Zhiryakov 1980, Ogurtsov 2012, 2016). The presence of trees and shrubs in managed meadows is caused by their low-intensity usage in some sites. It is possible to put the differences in coenotic group composition in connection with some changes in habitats caused by tree encroachment. Our dataset demonstrates an increased role of forest and forest-meadow species in all abandoned (and currently undisturbed) meadows in the study area. Remarkably, the total number of species in abandoned and managed mesic meadows is almost identical (177 and 182 species, respectively), the species richness per 100 m2 does not differ significantly, and the proportion of meadow species is equally high. This sug- gests that after abandonment the vascular plant diversity remains at the same level due to a kind of compensation: a significant increase of forest and forest-meadow spe- cies on the one hand and a significant decrease of weed- meadow species on the other hand, while the number of meadow species remains at the same level. We could have expected that as the number of tree species has in- creased, the proportion of meadow species must have declined, and the overall species diversity has decreased in the course of a new forest community formation. In O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 47 17/1 • 2018, 35–59 fact, after 25 (in some places even more) years of lack of any management the abandoned meadows still remain to be grasslands and retain a significant proportion of meadow species. Chytrý et al. (2005) showed that grasslands generally appeared to be quite resistant against vascular plant inva- sions, with average cover values of neophytes for differ- ent grassland types being only 0.1–1.7%. In the studied meadows of the Central Forest Reserve, there is a pal- pable fraction of alien species (6% of the total number of species), although their average coverage has nowhere exceeded 1% so far. Over the past 40 years, a number of alien species (Fe- stuca arundinacea, Epilobium adenocaulon, Malus dome- stica) have naturalised in local plant communities and also increased their presence in the reserve itself. Some species remained in the places of decorative plantings and did not spread farther. But their artificial popula- tions are quite stable and have endured in surrounding grasslands for a long time (more than 40 years), so the threat from their spreading and invasion still remains. One cultivated species, Cornus alba, spreads from hedge- rows, another, Triticum aestivum, was found in a dis- turbed MMM near an agricultural field. Some species (Juncus tenuis, Lepidoteca suaveolens) are mainly associ- ated with disturbances. Three species (Epilobium ad- enocaulon, Juncus tenuis and Lepidoteca suaveolens) have been present in the reserve over decades and already been mentioned by Minyaev & Konechnaya in 1976. Epilo- bium adenocaulon and Juncus tenuis used to be rare in the past (Minyaev & Konechnaya 1976), but are nowa- days widespread in semi-natural and ruderal communi- ties (Konechnaya 2012). Lepidoteca suaveolens has long since been widespread in disturbed habitats (Minyaev & Konechnaya 1976) and remains so these days (Kone- chnaya 2012). The frequency of L. suaveolens in our rel- evès is low since it prefers heavily disturbed habitats, not meadows. An invasive transformer species, Festuca arun- dinacea, was once cultivated in grass mixtures. Now it occurs in abandoned meadows, but its frequency in the studied communities is extremely low (1%), in contrast to a tendency to spread massively over most of the ter- ritory of Tver Oblast (Vinogradova et al. 2011). Thus, regular monitoring is needed as one of the activities to control alien species in the reserve. Grasslands are habitats of various red list and valuable species (Bruun & Ejrnæs 1998, Gibson 2009, Zavaleta et al. 2010, reviewed by Moeslund et al. 2013). Disap- pearance of red list species may, as some experts predict (Evstigneev 2012), follow meadow abandonment. The same author believes that red list species persist most suc- cessfully in hand-mown herbaceous communities. This might explain the absence of red list species in our RTC because these communities probably emerge in the sites of destroyed houses and abandoned vegetable gardens in former villages. Most likely, many red list species are un- able to survive in modern tractor-mown meadows with quite intensive usage (Evstigneev 2012). Conclusions We have identified four types of herbaceous communi- ties in the reserve. They are all different in management, productivity (herbage height), floristic composition and ecological conditions. Generally, herbaceous vegetation in the Central Forest Reserve features high ecological, flo- ristic and phytocoenotic diversity. Encompassing around 40% of the reserve flora, the meadows also include four red list species, for which grasslands make the most suit- able habitats. Mesic meadows are relatively stable ecosystems. For more than 25 years without any management, they have retained all the key meadow features, although their area has shrunk and the spread of alien species increased. In the long run, however, meadows under protection regime require special maintenance activities because they can not last on their own, and eventually, the loss of the veg- etation will cause a significant loss of biodiversity of the entire protected area. Monitoring of the populations of red list and invasive species is highly required, as well as regular mowing, preferably by hand. Acknowledgments The authors are grateful to the administration and staff of the Central Forest State Nature Biosphere Reserve, and also Veronika Gorik and Maria Nosova for their help in field work. We thank two anonymous reviewers of the manuscript as well as Orsolya Valkó and Jacqueline Loos for their helpful comments and suggestions for improve- ments. Special thanks to Aiko Huckauf for linguistic ed- iting. 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Bulletin of Moscow Society of Naturalists 85 (2): 20–29. [in Russian]. O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 51 17/1 • 2018, 35–59 Appendix 1: Frequency (%) of species in the four community types and coenotic affiliation. Abbreviations: MMM – managed mesic meadows, AMM – abandoned mesic meadows, MTC – meadowsweet tall-herb communities, RTC –ruderal tall-herb com- munities.* – alien species, R – red list species. Dodatek 1: Frekvenca (%) vrst v štirih združbah in njihova cenotska pripadnost. Okrajšave: MMM – gojena vlažna travišča, AMM – opuščena vlažna travišča, MTC – visoka steblikovja z brestovolistnim osladom, RTC – ruderalna visoka steblikovja.* – tujerodne vrste, R –vrste z rdečega seznama. Species Coenotic group MMM AMM MTC RTC Acetosa pratensis meadow 47 94 83 13 Acetosella vulgaris weed 24 - - - Achillea millefolium meadow 96 94 33 42 Aconitum septentrionale forest - 1 - - Aegopodium podagraria forest 2 9 13 22 Agrostis gigantea meadow 10 3 - 2 Agrostis stolonifera meadow-mire 8 - - - Agrostis tenuis forest-meadow 98 97 29 53 Ajuga reptans forest-meadow 4 18 4 - Alchemilla vulgaris forest-meadow 90 91 50 51 Alnus incana forest-mire 6 15 21 - Alopecurus geniculatus meadow-mire 8 - - - Alopecurus pratensis meadow - 6 - 4 Angelica sylvestris forest-meadow 14 82 79 56 Anthemis tinctoria weed-meadows 1 - - - Anthoxanthum odoratum meadow 92 89 17 - Anthriscus sylvestris forest-meadow 27 41 33 89 Arctium tomentosum weed 2 - - 7 Artemisia vulgaris weed-meadows 8 1 - 18 Asarum europaeum forest - - 4 - Athyrium filix-femina forest 2 1 4 4 Barbarea vulgaris weed 24 - - - Betula pendula forest 2 25 4 4 Betula pubescens forest-mire - 10 21 - Bistorta major meadow 10 37 50 11 Botrychium lunaria forest-meadow 2 - - - Briza media meadow 45 75 17 5 Bromopsis inermis weed-meadows - - - 5 Calamagrostis canescens mire - 11 42 - Calamagrostis epigeios weed-meadows 6 20 54 18 Calamagrostis phragmitoides indefinite - - 4 2 Caltha palustris mire 2 - 17 - Campanula glomerata meadow 16 8 - - Campanula latifolia forest - - - 7 Campanula patula meadow 55 77 29 24 Campanula rapunculoides weed-meadows - 3 - 7 Capsella bursa-pastoris weed 4 - - 2 Carduus crispus weed 6 - - 7 Carex acuta aquatic-mire 2 1 4 2 Carex appropinquata mire - - 4 - Carex brunnescens forest-mire - - 4 - Carex cespitosa mire - - - 4 Carex contigua meadow 63 19 - 4 O. 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Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 52 17/1 • 2018, 35–59 Species Coenotic group MMM AMM MTC RTC Carex echinata mire - - 8 - Carex flava meadow-mire 18 4 21 - Carex hirta weed-meadow 27 11 4 5 Carex juncella indefinite - 6 8 - Carex leporina forest-meadow 90 35 42 4 Carex nigra meadow-mire 37 28 50 - Carex pallescens forest-meadow 67 89 58 7 Carex panicea meadow-mire 10 3 13 - Carex vesicaria aquatic-mire 4 3 25 - Carum carvi weed-meadow 59 8 - - Centaurea jacea meadow 80 47 17 7 Centaurea phrygia meadow 37 97 83 60 Cerastium holosteoides meadow 92 37 4 - Chaerophyllum aromaticum forest 10 15 - 31 Chamaenerion angustifolium weed-forest - 15 21 58 Chenopodium album weed - - - 2 Chrysaspis spadicea meadow - 3 - - Chrysosplenium alternifolium forest - 1 8 5 Cirsium arvense weed 18 6 4 42 Cirsium heterophyllum forest-meadow - 27 54 11 Cirsium oleraceum forest-mire 2 6 33 13 Cirsium palustre forest-mire 6 14 67 7 Cirsium vulgare weed 18 - - - Coccyganthe flos-cuculi forest-meadow 20 51 67 5 Coeloglossum viride forest-meadow - - 4 - Comarum palustre mire 2 - 8 - Convolvulus arvensis weed 2 - 4 - Conyza canadensis* weed - 1 - - Cornus alba* indefinite 2 - - - Corylus avellana forest - - - 2 Crepis paludosa forest 2 29 83 13 Cuscuta europaea forest-meadow - - - 4 Cynosurus cristatus meadow 65 9 - - Dactylis glomerata meadow 45 72 38 98 Dactylorhiza balticaR indefinite - - 4 - Dactylorhiza fuchsii indefinite 2 3 8 2 Dactylorhiza maculata forest-mire 8 10 17 - Deschampsia cespitosa forest-meadow 59 90 58 35 Dianthus barbatus* indefinite - 1 - - Dianthus deltoides meadow 2 - - - Dryopteris carthusiana forest - - 4 2 Dryopteris expansa forest - 1 - - Dryopteris filix-mas forest - - - 2 Echium vulgare weed-meadow 1 - - - Elymus caninus forest - - - 2 Elytrigia repens weed-meadow 47 24 8 71 Epilobium adenocaulon* weed 16 1 - 2 Epilobium hirsutum weed-mire 2 - 4 - Epilobium palustre mire 2 1 21 - Equisetum arvense weed-meadow 51 52 54 29 O. 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Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 53 17/1 • 2018, 35–59 Species Coenotic group MMM AMM MTC RTC Equisetum fluviatile aquatic-mire - - 8 - Equisetum palustre forest-meadow 10 - 29 4 Equisetum pratense forest-meadow - 1 - 11 Equisetum sylvaticum forest 6 18 42 16 Eriophorum polystachyon mire 2 - - - Erysimum cheiranthoides weed - - - 2 Euphorbia virgata weed-meadow - - - 2 Euphrasia brevipila meadow 24 5 - - Fallopia convolvulus weed - - - 2 Festuca arundinacea* weed - 3 - 2 Festuca pratensis meadow 100 90 50 51 Festuca rubra meadow 86 66 21 11 Filipendula ulmaria meadow-mire 20 48 96 45 Fragaria vesca forest-meadow - 4 - 4 Frangula alnus forest - 18 - 4 Galeopsis bifida weed 24 20 8 51 Galeopsis speciosa weed 39 28 25 69 Galium mollugo forest-meadow 27 25 8 24 Galium palustre meadow-mire - - 46 - Galium uliginosum mire 47 84 75 25 Geranium palustre forest-mire 22 68 75 76 Geum rivale meadow-mire 41 46 92 42 Geum urbanum weed 2 - - - Glechoma hederacea forest-meadow - 1 8 4 Gnaphalium uliginosum weed 2 - - - Gymnadenia conopseaR meadow - 9 4 - Heracleum sibiricum weed-meadow 2 15 - 25 Hieracium umbellatum forest-meadow 12 44 4 - Hierochloe odorata meadow 2 5 21 2 Hylotelephium triphyllum meadow - 1 - - Hypericum maculatum forest 65 95 71 58 Juncus articulatus weed-mire 16 1 - - Juncus bufonius weed-mire 2 - - - Juncus compressus weed-meadow - 1 - - Juncus conglomeratus meadow-mire 4 6 4 - Juncus effusus meadow-mire 35 24 46 - Juncus filiformis meadow-mire 10 27 46 - Juncus tenuis* weed 35 4 - - Juniperus communis forest - 1 - - Knautia arvensis meadow 29 77 21 38 Lamium maculatum forest - - 4 5 Lathyrus pratensis meadow 67 54 79 51 Lathyrus sylvestris forest - 1 - 5 Leontodon autumnalis meadow 76 1 - - Leontodon hispidus meadow 24 44 - - Lepidotheca suaveolens* weed 2 - - - Leucanthemum vulgare meadow 80 75 4 22 Linaria vulgaris weed-meadow - - - 2 Linum catharticum  indefinite 2 - - - Listera ovata forest 2 13 4 - O. 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Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 54 17/1 • 2018, 35–59 Species Coenotic group MMM AMM MTC RTC Lolium perenne* weed 12 - - - Lonicera xylosteum forest - - - 1 Luzula multiflora meadow 51 65 29 - Luzula pallidula forest-meadow 2 10 - - Luzula pilosa forest - - - 2 Lychnis chalcedonica* indefinite - 1 - - Lysimachia vulgaris forest-mire 22 15 42 18 Maianthemum bifolium forest - 1 - 2 Malus domestica* indefinite 2 32 - 15 Matteuccia struthiopteris forest - - - 2 Melampyrum nemorosum forest 14 89 54 71 Melandrium album weed-meadow 2 - - - Melandrium dioicum forest - 4 4 16 Mentha arvensis weed-meadow 27 8 17 2 Mercurialis perennis forest - - 4 - Milium effusum forest - - - 2 Myosotis arvensis weed 12 - - 4 Myosotis palustris forest-mire 24 43 25 - Nardus stricta meadow 2 6 8 - Oberna behen weed-meadow 2 - - - Odontites vulgaris weed-meadow - 4 - 4 Omalotheca sylvatica forest 6 11 - - Ophioglossum vulgatum meadow - 3 25 4 Padus avium forest - 1 - 2 Persicaria amphibia meadow-aquatic 2 - - - Persicaria minus weed-meadow 2 - - - Petasites hybridus* weed - - - 2 Phalaroides arundinacea meadow-mire - 3 8 4 Phleum pratense meadow 98 91 50 49 Picea abies forest - 20 13 5 Pilosella caespitosa indefinite 4 28 - 2 Pilosella flagellaris indefinite 4 - - - Pilosella officinarum forest-meadow - 11 - - Pilosella onegensis indefinite 12 18 13 2 Pilosella scandinavica indefinite 2 - - - Pimpinella saxifraga meadow 41 34 - 9 Pinus sylvestris forest - 1 - - Plantago lanceolata meadow 73 49 4 4 Plantago major weed 71 1 - 2 Plantago media meadow 18 1 - 2 Platanthera bifolia forest 18 57 29 5 Poa angustifolia meadow 76 84 21 35 Poa annua weed 12 - - - Poa compressa weed 2 - - - Poa nemoralis forest - 1 - - Poa palustris meadow-mire - 8 4 7 Poa pratensis meadow 47 8 4 4 Poa trivialis meadow 33 27 46 18 Polemonium caeruleum forest-meadow - 29 54 60 Polygala vulgaris meadow 6 24 - - O. 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Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 55 17/1 • 2018, 35–59 Species Coenotic group MMM AMM MTC RTC Populus balsamifera* indefinite - 1 - - Populus longifolia* indefinite - 1 - - Populus tremula forest 6 5 8 - Potentilla anserina weed-meadow 75 - - - Potentilla erecta forest-meadow 53 95 75 18 Potentilla goldbachii indefinite - 1 - 4 Potentilla intermedia weed 10 - - - Prunella vulgaris weed 88 47 17 2 Ptarmica vulgaris meadow 6 - - - Ranunculus acris meadow 100 85 75 20 Ranunculus auricomus meadow 20 42 88 9 Ranunculus cassubicus forest - - 17 2 Ranunculus flammula meadow 10 - - - Ranunculus repens aquatic-mire 88 20 25 7 Raphanus raphanistrum weed 8 - - - Rhinanthus minor meadow 25 13 - 2 Rhinanthus serotinus weed-meadow 4 - - - Ribes nigrum forest - - - 1 Rorippa palustris weed-mire 6 - - - Rosa majalis forest - 1 - 2 Rosa rugosa* indefinite - 1 - - Rubus idaeus weed-forest - - 4 13 Rumex confertus weed-meadow 2 1 - - Rumex crispus weed-meadow 31 4 - 2 Rumex longifolius meadow 2 1 8 2 Rumex obtusifolius weed-forest 2 4 4 9 Sagina procumbens weed 14 - - - Salix aurita mire 2 18 21 - Salix caprea forest 10 9 4 - Salix cinerea mire 6 1 4 - Salix myrsinifolia meadow-mire 10 16 33 2 Salix pentandra mire 4 - 4 - Salix phylicifoliaR indefinite - - 4 - Salix rosmarinifolia meadow-mire - - 4 - Salix starkeana meadow-mire - 8 4 2 Scirpus sylvaticus aquatic-mire 6 5 33 9 Scrophularia nodosa forest - - 8 9 Scutellaria galericulata forest-mire - - 25 - Selinum carvifolia forest - 3 - - Solanum dulcamara forest-mire - - 13 - Solidago virgaurea forest-meadow 2 18 - 2 Sorbus aucuparia forest - 1 - - Stachys palustris weed-mire 4 5 - 4 Stachys sylvatica forest 2 1 - 2 Stellaria graminea meadow 92 95 54 60 Stellaria holostea forest - 11 21 5 Stellaria nemorum forest - 3 - 7 Succisa pratensis forest-meadow 33 73 46 2 Symphyotrichum sp.* indefinite - - - 1 Taraxacum officinale weed-meadow 80 6 - 4 O. 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Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 56 17/1 • 2018, 35–59 Species Coenotic group MMM AMM MTC RTC Thalictrum aquilegifolium forest - 8 29 27 Thalictrum flavum meadow-mire - - 4 2 Thalictrum lucidum forest-meadow 2 9 - 9 Thlaspi arvense weed - - - 2 Tilia cordata forest - 1 - 2 Trientalis europaea forest - - 4 - Trifolium hybridum weed-meadow 55 38 - 2 Trifolium medium forest-meadow 57 82 21 78 Trifolium pratense weed-meadow 82 25 - 2 Trifolium repens weed-meadow 82 24 - 7 Tripleurospermum inodorum weed 18 - - - Triticum aestivum* indefinite 4 - - - Trollius europaeus meadow 12 81 67 29 Urtica dioica weed-forest 8 1 17 67 Valeriana officinalis meadow 6 11 54 9 Veronica chamaedrys forest-meadow 82 99 83 75 Veronica longifolia meadow - 8 13 11 Veronica officinalis forest - 9 4 - Veronica scutellata mire 2 - 4 - Veronica serpyllifolia weed-meadow 20 - - - Viburnum opulus forest - 1 - - Vicia cracca meadow 94 86 63 80 Vicia sepium meadow 29 34 46 47 Vicia villosa weed 2 - - - Viola arvensis weed 4 - - - Viola canina forest-meadow 6 42 4 9 Viola epipsila forest-mire - 1 4 - Viola nemoralis indefinite - 15 4 - Viola palustris forest-mire - 15 67 2 Viola tricolor weed - - - 9 Total number of species 177 182 145 156 O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 57 17/1 • 2018, 35–59 Appendix 2: Diagnostic species of different community types according to phi coefficient. Only species with phi >0.25 are shown; phi coefficient values for the highly diagnostic species are highlighted in bold. Monte Carlo test of significance of the observed maximum indicator value for each species, with 999 randomisations, including p-values. Dodatek 2: Diagnostične vrste različnih združb glede na fi koeficient. Prikazane so samo vrste s koeficientom fi >0.25; fi koeficien- ti za pomembne diagnostične vrste so prikazani krepko. Monte Carlo test statistične značilnosti maksimalne indikatorske vrednosti za posamezno vrsto z 999 slučajenjem vključno z p-vrednostmi. Herbaceous community type Phi Mean Standard deviation p-value Managed mesic meadows (MMM) Potentilla anserina 0.829 0.089 0.047 0.0002 Leontodon autumnalis 0.827 0.09 0.046 0.0002 Taraxacum officinale 0.777 0.089 0.046 0.0002 Plantago major 0.772 0.089 0.046 0.0002 Cynosurus cristatus 0.659 0.089 0.047 0.0002 Cerastium holosteoides 0.655 0.088 0.045 0.0002 Ranunculus repens 0.654 0.087 0.045 0.0002 Trifolium pratense 0.649 0.088 0.045 0.0002 Trifolium repens 0.632 0.087 0.046 0.0002 Carum carvi 0.626 0.089 0.046 0.0002 Carex leporina 0.57 0.088 0.045 0.0002 Prunella vulgaris 0.541 0.088 0.045 0.0002 Carex contigua 0.529 0.09 0.047 0.0002 Poa pratensis 0.488 0.09 0.047 0.0002 Juncus tenuis 0.477 0.089 0.046 0.0002 Centaurea jacea 0.456 0.088 0.045 0.0002 Barbarea vulgaris 0.433 0.09 0.049 0.0002 Acetosella vulgaris 0.433 0.091 0.049 0.0002 Rumex crispus 0.421 0.09 0.047 0.0002 Plantago lanceolata 0.41 0.088 0.046 0.0002 Festuca rubra 0.402 0.088 0.044 0.0002 Veronica serpyllifolia 0.393 0.091 0.048 0.0002 Ranunculus acris 0.373 0.086 0.044 0.0002 Cirsium vulgare 0.372 0.093 0.049 0.0002 Tripleurospermum inodorum 0.372 0.092 0.048 0.0002 Euphrasia brevipila 0.339 0.09 0.047 0.0004 Trifolium hybridum 0.339 0.088 0.045 0.0004 Sagina procumbens 0.326 0.091 0.051 0.0004 Juncus articulatus 0.318 0.091 0.047 0.0016 Plantago media 0.315 0.091 0.048 0.0004 Festuca pratensis 0.309 0.086 0.043 0.0002 Leucanthemum vulgare 0.303 0.087 0.044 0.0006 Poa annua 0.302 0.09 0.052 0.0022 Lolium perenne 0.302 0.089 0.052 0.0028 Epilobium adenocaulon 0.29 0.092 0.046 0.0022 Phleum pratense 0.287 0.086 0.043 0.0002 Ranunculus flammula 0.275 0.092 0.049 0.001 Potentilla intermedia 0.275 0.092 0.048 0.002 Mentha arvensis 0.272 0.091 0.048 0.0036 Abandoned mesic meadows (AMM) Viola canina 0.519 0.089 0.046 0.0002 Trollius europaeus 0.469 0.087 0.045 0.0002 O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 58 17/1 • 2018, 35–59 Herbaceous community type Phi Mean Standard deviation p-value Acetosa pratensis 0.464 0.087 0.045 0.0002 Hieracium umbellatum 0.459 0.088 0.045 0.0002 Potentilla erecta 0.45 0.087 0.044 0.0002 Briza media 0.446 0.088 0.046 0.0002 Succisa pratensis 0.445 0.089 0.046 0.0002 Knautia arvensis 0.415 0.087 0.044 0.0002 Platanthera bifolia 0.407 0.089 0.047 0.0002 Anthoxanthum odoratum 0.404 0.088 0.045 0.0002 Carex pallescens 0.397 0.088 0.045 0.0002 Centaurea phrygia 0.39 0.086 0.044 0.0002 Melampyrum nemorosum 0.387 0.087 0.044 0.0002 Pilosella sp. 0.387 0.09 0.047 0.0002 Leontodon hispidus 0.383 0.089 0.046 0.0002 Galium uliginosum 0.362 0.087 0.044 0.0002 Deschampsia cespitosa 0.356 0.088 0.043 0.0004 Angelica sylvestris 0.345 0.086 0.044 0.0002 Frangula alnus 0.345 0.091 0.048 0.0004 Polygala vulgaris 0.345 0.09 0.047 0.0004 Malus domestica 0.336 0.089 0.046 0.0004 Betula pendula 0.335 0.089 0.047 0.0006 Campanula patula 0.328 0.088 0.045 0.0004 Agrostis tenuis 0.313 0.086 0.043 0.0002 Hypericum maculatum 0.31 0.087 0.043 0.0002 Alchemilla vulgaris 0.308 0.088 0.044 0.0006 Solidago virgaurea 0.299 0.09 0.047 0.0008 Myosotis palustris 0.297 0.09 0.046 0.0012 Poa angustifolia 0.282 0.087 0.044 0.0002 Ajuga reptans 0.271 0.091 0.047 0.0016 Meadowsweet tall-herb communities (MTC) Viola palustris 0.624 0.091 0.048 0.0002 Galium palustre 0.623 0.09 0.047 0.0002 Crepis paludosa 0.611 0.089 0.046 0.0002 Cirsium palustre 0.578 0.088 0.047 0.0002 Ranunculus auricomus 0.541 0.088 0.045 0.0002 Filipendula ulmaria 0.489 0.087 0.045 0.0002 Valeriana officinalis 0.483 0.088 0.046 0.0002 Calamagrostis canescens 0.458 0.09 0.047 0.0002 Scutellaria galericulata 0.447 0.09 0.052 0.0002 Geum rivale 0.422 0.087 0.045 0.0002 Cirsium heterophyllum 0.398 0.089 0.046 0.0002 Calamagrostis epigeios 0.383 0.088 0.046 0.0004 Ophioglossum vulgatum 0.367 0.092 0.047 0.0006 Carex vesicaria 0.367 0.092 0.048 0.0008 Equisetum palustre 0.364 0.091 0.048 0.0004 Epilobium palustre 0.36 0.091 0.05 0.0004 Scirpus sylvaticus 0.343 0.089 0.048 0.0008 Coccyganthe flos-cuculi 0.339 0.088 0.045 0.0006 Caltha palustris 0.334 0.092 0.047 0.0004 Ranunculus cassubicus 0.334 0.093 0.049 0.001 Cirsium oleraceum 0.332 0.089 0.047 0.0004 O. Cherednichenko & V. Borodulina Biodiversity of herbaceous vegetation in abandoned and managed sites under protection regime: a case study in the Central Forest Reserve, NW Russia 59 17/1 • 2018, 35–59 Herbaceous community type Phi Mean Standard deviation p-value Juncus filiformis 0.332 0.089 0.046 0.001 Solanum dulcamara 0.311 0.098 0.047 0.0012 Equisetum sylvaticum 0.294 0.09 0.048 0.0042 Hierochloe odorata 0.287 0.09 0.047 0.003 Bistorta major 0.272 0.089 0.046 0.0044 Carex nigra 0.265 0.089 0.046 0.0054 Betula pubescens 0.257 0.09 0.048 0.0094 Juncus effusus 0.254 0.088 0.046 0.007 Ruderal tall-herb communities (RTC) Urtica dioica 0.645 0.09 0.047 0.0002 Chamaenerion angustifolium 0.481 0.089 0.047 0.0002 Anthriscus sylvestris 0.457 0.088 0.044 0.0002 Cirsium arvense 0.38 0.089 0.047 0.0002 Elytrigia repens 0.373 0.088 0.045 0.0004 Dactylis glomerata 0.37 0.087 0.044 0.0002 Galeopsis bifida 0.325 0.09 0.046 0.0002 Galeopsis speciosa 0.316 0.087 0.045 0.0004 Polemonium caeruleum 0.315 0.088 0.045 0.0006 Rubus idaeus 0.285 0.092 0.048 0.0016 Viola tricolor 0.269 0.092 0.048 0.0026 Artemisia vulgaris 0.264 0.09 0.047 0.0034