Phytocenotic Features of Calluna Vulgaris (L.) Hill. in Ukrainian Polesie

Ecological Questions 31 (2020) 3: 73– 107 http://dx.doi.org/10.12775/EQ.2020.024

Phytocenotic features of Calluna vulgaris (L.) Hill. in Ukrainian Polesie

Oleksandr Lukash1*, Viktor Melnyk2, Hanna Danko1, Oleksandr Rak2, Yurii Karpenko1, Petro Buzunko1

1Department of Ecology and Nature Conservation, T. H. Shevchenko National University “Chernihiv Collehium”, Hetman Polubotko Str., 53, 14013, Chernihiv, Ukraine 2Department of Natural flora, M.M. Gryshko National Botanical Garden of the National Academy of Sciences of Ukraine, Timiriazievska Str., 1, 01014, Kyiv, Ukraine *corresponding author’s e-mail: [email protected]

Received: 21 March 2020 / Accepted: 8 May 2020

The article is dedicated to the memory of the tireless researcher of the Ukrainian Polesie flora and vegetation Professor Tetiana Andrienko (1938-2016)

Abstract. Middle-aged pine and birch-pine forests of the Dicrano-Pinenion suballiance are the most common biotopes of Calluna vulgaris (L.) Hill. in Ukrainian Polesie. This species occurs mostly as solitary plants in forest phytocenoses; rarely its projective coverage can reach 40-45%. C. vulgaris has the highest constancy in phytocenoses of the Peucedano-Pinetum association in Right- bank Polesie. The forests of the Piceo-Vaccinienion uliginosi Seibert in Oberd. (ed.) 1992 suballiance and the Piceion excelsae Pawłowski et al. 1928 alliance are rare habitats of C. vulgaris; they occur only in Right-bank Polesie of Ukraine. Heathlands are rare ecosystems for Ukrainian Polesie, their vegetation is represented by plant communities of the Calluno-Ulicetea Br.-Bl. et Tx. ex Klika et Hadač 1944 and the Nardetea strictae Rivas Goday et Borja Carbonell in Rivas Goday et Mayor Lopez 1966 classes. They are forming in the significantly anthropogenically disrupted territories (e.g. areas after wildfires) or in the areas with intensive human activities, firstly in the military training areas. The phytocenoses of the Calluno-Genistetum pilosae R. Tx. 1937 association occur only in Right-bank Polesie and are the largest heathlands of Ukrainian Polesie in general. For example, in the Rivne region, their area is up to 20 km2. The area of heathlands, belonging to the Calluno-Nardetum strictae Hrync. 1959 association, does not exceed several hectares. The phytocenoses of the Scabioso canescentis-Genistetum Balcerk. et Brzeg 1993 association are represented with small plots in landscapes of pine forests and less frequently in mixed forests on upland terraces of Left-bank Polesie rivers. The characteristic feature of Calluna vulgaris spreading in Right-bank Polesie of Ukraine is its large participation (sometimes with the projective cover up to 80%) and high constancy (92,7%) in phytocenoses of oligotrophic bogs of the Ledo-Sphagnetum magellanici (Malc. 1929) Kästner et Flössner 1933 association. The populations of Calluna vulgaris, explored in the forest phytocenoses of the Cladonio-Pinetum Juraszek 1927 association on the territory of Ukrainian Polesie, are incomplete. Their age spectrum maximum falls on generative individuals. The “Right-bank” population of Calluna vulgaris are characterized by more active proliferation of plants and spreading on new territories. Aging processes dominate among the “left-bank” populations. Generally, environmental conditions of Ukrainian Right-bank Polesie are more favorable to Calluna vulgaris, than conditions of Left-bank Polesie.

Keywords: Calluna vulgaris (L.) Hill., Polesie, plant communities, syntaxonomy, population. 1. Introduction Calluna vulgaris (L.) Hill., European Boreo-temperate element of flora, has a large geographic 74 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko distribution and is dominating in many heath ecosystems. Such a type of vegetation prevails in many heathlands of northern and western Europe. In Ukrainian Polesie Calluna vulgaris is at the 1. Introduction scapes. Poor sandy soils and marshiness favored the con- southern border of its distribution. Thatservation is whatof natural causes vegetation, the affectwhich occupies on the herespecies about distribution in the 2/3 of the area (Andrienko et al., 2006). During the maxi- Calluna vulgarisregion. (L.) Hill., The European purpose Boreo-temperate of our study el- wasmum to glaciations investigate most of the Ukrainian cenotic Polesie features was covered of the by Calluna vulgaris ement of flora, has a large geographic distribution and is glacier. In modern relief glacial forms are most noticeable dominating in habitatsmany heath and ecosystems. its population Such a type structure of veg- in in the Ukrainian west of Polesie. Polesie. etation prevails in many heathlands of northern and west- The Dnieper is the main waterway of Ukrainian Pole- ern Europe. In UkrainianHeathlands Polesie Calluna are vulgaris highly is at the dynamic sie. The habitats,Dnieper divides with Ukrainian strong Polesiebiotic into and Right- abiotic interactions southern border of its distribution. That is what causes the bank (Fig.1: 1a) and Left-bank Polesie (Fig.1: 1b). There affect on the speciesaffected distribution by external in the region.drivers, The despite pur- are their no significant appearance differences as a betweenhomogeneous, Right-bank stablePolesie ecosystem. All of pose of our studythe was factors to investigate studied the cenoticinfluence features diversity of and Left-Bank patterns, Polesie community in the amount structure of weather and elements. general quality of the the Calluna vulgaris habitats and its population structure However, Left-bank Polesie has more continental climate in Ukrainian Polesie.habitat (Fagúndez, 2013). As experiencein comparison of withEuropean Right-bank (in Polesie. particula Karst processesr, Polish) are scientists shows Heathlands are highly dynamic habitats, with strong observed in some regions of Right-bank Polesie. So-called biotic and abiotic(Adamska interactions et affectedal., 2015 by ),external study driv of- mainloess heath“islands” formations are available and in Left-bank accompanying Polesie, where plant communities is ers, despite their appearance as a homogeneous, stable erosion processes take place (Lukash et al., 2018, 2019). ecosystem. Allvery of the important factors studied for influence management, diversity Soilstreatment of Polesie and are highlyalso patternconservation structured. Riseof ofthe the Calluna vulgaris patterns, community structure and general quality of the ground level by only a few decimeters in Polesie usually habitat (Fagúndez,ecological 2013). As systems experience. of European (in causes major changes in soil-forming processes and veg- particular, Polish) scientists shows (Adamska et al., 2015), etation. Sod-podzolic soils are most common. They oc- study of main heath formations and accompanying plant cupy about 70% of the territory of Ukrainian Polesie. The communities is very important for management, treatment sod-podzolic soils vary greatly in mechanical composition, and also conservation of the Calluna vulgaris ecological humidity2. and Study trophicity. area Sandy-loam and clayish sandy- systems. loam soils dominate by mechanical composition. Both pine Ukrainian Polesie is the southern partand deciduous of the Polessianforests are forming lowland on those within types Ukraineof soils. (Fig. 1). Poorly Gray forest soils are common on loamy loessial sediments, defined2. Study relief, area considerable distributionmainly inof Left-bank water- icePolesie. deposits of sand, and high marshiness are According to various literature sources, forest areas Ukrainian Polesiecharacteristic is the southern partfeatures of the Polessian of Polesie low- occupylandscapes. 26-29% ofPoor the territory sandy of Ukrainiansoils and Polesie marshi (An- ness favored the land within Ukraineconservation (Fig. 1). Poorly of natural defined vegetation,relief, con- drienkowhich et occupiesal., 2006). The here considerable about 2/3 distribution of the of area sand (Andrienko et al., siderable distribution of water-ice deposits of sand, and deposits in Ukrainian Polesie causes large areas of pine high marshiness2006) are characteristic. During featuresthe maximum of Polesie land glaciations- forests. Climatic most ofconditions Ukrainian of Polesie Polesie are favorable was forcovered the by glacier. In modern relief glacial forms are most noticeable in the west of Polesie.

FigureFigure 1. Schematic 1. Schematic map of Polesie. map Mapof Polesie.symbols: 1 Map– Ukrainian symbols: Polesie (1a1 – – Right-bank,Ukrainian 1b– Polesie Left- (1a – Right-bank, 1b– Left- bank),bank) 2 – Lublin, 2 Polesie,– Lublin 3 – Belarusian Polesie, Polesie, 3 – Belarusian 4 – Nerissa and Polesie Desna ,Polesie 4 – Nerissa and Desna Polesie

The Dnieper is the main waterway of Ukrainian Polesie. The Dnieper divides Ukrainian Polesie into Right-bank (Fig.1: 1a) and Left-bank Polesie (Fig.1: 1b). There are no significant 2

Phytocenotic features of Calluna vulgaris (L.) Hill. in Ukrainian Polesie 75

growth of deciduous forests, which are local climax zonal lomeshch (2003). Syntax names are ordered according to communities. However, there are differences in distribu- Mucina et al. (2016). tion of deciduous forests between Right-bank and Left- The field populations studies were carried out accord- bank Polesie. For example, the areal of the Carpinion betuli ing to Rabotnov (1964). The Calluna vulgaris ontogenetic Issler 1931 alliance forests includes all Right-bank Polesie stages and evaluation indexes of the general ontogenetic and only part of Left-bank Polesie (within the boundaries status of populations (Index innovation (Iinn ), Index seni- of the Chernihiv region) (Andrienko et al., 2006). lis (Isen ), Index generative (Igen ), Index aetas (I et)) were A high groundwater level and its slow motion favor determined according to Kovalenko (2015). The evalua- spreading of wetland vegetation here. The climate condi- tion indexes of the general ontogenetic status of the popu- tions of Polesie contributed to the development of mires lations are measured in per cent and defined as ratio of and in combination with mineral deficiency of the sur- individuals quantity in different ontogenetic stages to the face formations, have led to sphagnum mires formation total quantity of plants: seedlings (p), juvenile (j), imma-

(Fijałkowski et al., 2002). However, Left-bank Polesie is ture (im) and virginile (v) individuals – Iinn ; old generative characterized by less quantity of bogs. In addition, Left- individuals (g3 ) and senile plants (s) – Isen ; young genera- bank Polesie, compared to Right-bank Polesie, was more tive (g1) and mature generative individuals (g2) – Igen . In- affected by the drainage reclamation in 1970s-1980s. dex aetas is the ratio of Index senilis to Index innovation:

Obviously, the regularities of Calluna vulgaris distri- Iaet = Isen /Iinn. bution in these two regions of Ukrainian Polesie were affected by some differences in natural conditions (including vegetation cover) of Right-bank and Left-bank Polesie. 4. Results and discussion

In Ukrainian Polesie Calluna vulgaris grows mainly 3. Material and methods in the intrazonal boreal-temperate grasslands and heath plant communities, in swamps and also in pine, mixed The materials for the article were collected during the field and spruce forests. The general scheme of the Ukrainian research of the Polesie vegetation in 1995-2019. The field Polesie plant communities with Calluna vulgaris is the fol- study of the vegetation was carried out by geobotanical lowing (note: * – only in Right-bank Polesie, ** – only methods (Korchahin & Lavrenko, 2012). The vegetation in Left-bank Polesie): descriptions were taken during the optimum of vegetation Class: Calluno-Ulicetea Br.-Bl. et Tx. ex Klika et Hadač period in the areas of 400-625 m2 (in forest phytocenoses) 1944 and 100-125 m2 (in herb and dwarf shrub phytocenoses). Order: Vaccinio myrtilli-Genistetalia pilosae Schubert ex The projective coverage of plant communities’ layers and Passarge 1964 coverage of all the species was noted. Cover abundance Alliance: Calluno-Genistion pilosae P. Duvigneaud 1944 scale is the following: + – up to 1%, 1 – 1 – 5%, 2 – 6 – *Association: Calluno-Genistetum pilosae R.Tx. 1937 15%, 3 – 16 – 25%, 4 – 26 – 50%, 5 > 50%. The tables of Alliance: Calluno-Arctostaphylion uvae-ursi Tx. et Preising the floristic structure of the plant communities for Calluna in Preising 1949 vulgaris include the percentage share. **Association: Scabioso canescentis-Genistetum Balcerk. 366 phytosociological authors relevés were done. 297 et Brzeg 1993 geobotanical relevés from Professor Tatiana Andrienko’s Class: Nardetea strictae Rivas Goday et Borja Carbonell field books 1972-1985, which had been transferred to the in Rivas Goday et Mayor López 1966 first author of this article during her lifetime, were ana- Order: Nardetalia strictae Preising 1950 lyzed. 30 published geobotanical relevés, taken on the ter- Boreo-temperate group of alliances ritory of Desniansko-Starogutskyi National Nature Park, Alliance: Violion caninae Schwickerath 1944 were also used (Panchenko, 2013). In total, 693 relevés Association: Calluno-Nardetum strictae Hrync. 1959 were analyzed. Class: Oxycocco-Sphagnetea Br.-Bl. et Tx. ex Westhoff et The documentation has been compiled separately for al. 1946 Right-bank Polesie (total number of relevés 391 including Order: Sphagnetalia medii Kästner et Flössner 1933 247 relevés with Calluna vulgaris) and Left-bank Polesie Alliance: Sphagnion medii Kästner et Flössner 1933 (total number of relevés 302 including 195 relevés with *Association: Ledo-Sphagnetum magellanici (Malc. 1929) Calluna vulgaris). Phytocenotic tables include the relevés Kästner et Flössner 1933 with the projective coverage of Calluna vulgaris equal to Class:Vaccinio-Piceetea Br.-Bl. in Br.-Bl. et al. 1939 or more than 1%. Order: Pinetalia sylvestris Oberd. 1957 Syntaxa were identified according to W. Matuszkiewicz Alliance: Dicrano-Pinion sylvestris (Libbert 1933) W. Ma- (2019), Chojnacka et al. (2010a, 2010b), Bulokhov & So- tuszkiewicz 1962 76 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko

Suballiance: Dicrano-Pinenion Seibert in Oberd. (ed.) of Volodymyr-Volynskyi (the Volyn region). A shrub layer 1992 is formed with Sarothamnus scoparius (L.) W.D.J. Koch Association: Cladonio-Pinetum Juraszek 1927 (the projective cover of 40-60%). Calluna vulgaris domi- Association: Peucedano-Pinetum W. Matuszkiewicz nates in a herb layer and has the projective cover of 50- (1962) 1973 80%. Sarothamnus scoparius is a diagnostic species of the **Association: Veronico incanae-Pinetum Bulokhov et So- Calluno-Genistetum pilosae association and the Calluno- lomeshch 2003 Genistion pilosae alliance. Moss layer is formed with Association: Molinio-Pinetum W. Matuszkiewicz et J. Ma- Pleurozium schreberi (Brid.) Mitt. Polessian communities tuszkiewicz 1973 with Sarothamnus scoparius differ in composition from the Association: Querco-Piceetum (W. Matuszkiewicz 1952) plant communities, described in Poland near the city of W. Matuszkiewicz et Polak 1955 Toruń. The community dominated by common groom – Suballiance: Piceo-Vaccinienion uliginosi Seibert in Oberd. Sarothamnus scoparius covers the areas adjacent to heath- (ed.) 1992 lands Pohlio-Callunetum Shimwell 1973 em. Brzeg 1981 *Association: Vaccinio uliginosi-Pinetum Kleist 1929 and Arctostaphylo-Callunetum R.Tx. et Prsg 1940, as Order: Piceetalia excelsae Pawłowski et al. 1928 well as with grasslands of the gray clubawn grass Sper- Alliance: Piceion excelsae Pawłowski et al. 1928 gulo vernalis-Corynephoretum (R.Tx. 1928) Libb. 1933 *Association: Sphagno girgensohnii-Piceetum Polak 1962 (Chojnacka et al., 2010a). The intrazonal boreo-temperate heath vegetation is The low-altitude heaths of the Atlantic and subconti- represented by plant communities of the Calluno-Ulice- nental regions of temperate Europe, belonging to the Sca- tea (Table 1) and the Calluno-Nardetum strictae classes bioso canescentis-Genistetum association (Table 1, relevés (Table 2). 4-8), are represented only in Left-bank Polesie. They are Relevés 1-3 (Table 1) belong to the Calluno-Geniste- rare and occupy small areas (up to 200 m2) among pine tum pilosae association (the Calluno-Ulicetea pilosae forests on the upland river terraces. The characteristic (Ge- class). The cenoses of the Calluno-Genistetum pilosae as- nista tinctoria L., Scabiosa ochroleuca L.) and differential sociation, which are present only in Right-bank Polesie, (Polygonatum odoratum (Mill.) Druce, Solidago virgaurea occupy the largest areas (up to 20 km2). Calluno-Geniste- L., Vincetoxicum hirundinaria Medik.) species of the asso- tum pilosae encompasses dry heaths represented, as well ciation are present in all the described phytocenoses. Cal- as in Poland (Adamska et al., 2015), by the anthropogenic luna vulgaris grows in curtains; its projective coverage communities, developing on poor sandy soils, as a degen- does not exceed 10%. The plant communities have a moss eration phase of pine and mixed forests. layer (with the projective cover of 25-90%). Pleurozium The heathlands of the Calluno-Genistetum pilosae as- schreberi dominates in the layer. sociation are common on large territories of the artillery The heathlands of Ukrainian Polesie are represented shooting range in Hoshchansk, Kostopil and Rivne dis- by the plant comunities of the Calluno-Nardetum strictae tricts of the Rivne region and on the northern outskirts association (the Nardetea strictae class). They occupy the

Table 1. Floristic structure of the Ukrainian Polesie heath plant communities of the Calluno-Ulicetea class: the Calluno-Genistetum pilosae (relevés 1-3) and the Scabioso canescentis-Genistetum (relevés 4-8) associations

Region Right-bank Polesie Left-bank Polesie Relevé number 1 2 3 4 5 6 7 8 Shrub layer (b) cover [%] 60 40 5 1 5 <1 0 15 Herb and dwarf shrub layer (c) cover [%] 80 50 90 40 35 45 50 25 Mosses layer (d) cover [%] 10 0 30 25 70 80 90 85 Lichens layer (e) cover [%] 0 0 0 0 10 5 0 5 Ch., D. Ass. Calluno-Genistetum pilosae and All. Calluno-Genistion pilosae Sarothamus scoparius 5 4 + . . . . . Genista germanica + + + . . . . . [77]

Region Right-bank Polesie Left-bank Polesie Relevé number 1 2 3 4 5 6 7 8 Ch., D. Ass. Scabioso canescentis-Genistetum Genista tinctoria . . . + + 1 + + Polygonatum odoratum . . . + 1 + + + Scabiosa ochroleuca . . . + + + + + Solidago virgaurea . . . + + + + + Vincetoxicum hirundinaria . . . + + + + + Сh. All. Calluno-Arctostaphylion Arctostaphylos uva-ursi . . . . 1 1 + + Carex praecox . . . + + + . . Peucedanum oreoselinum . . . + + 1 2 + Scorzonera humilis . . . . . + + + Сh. Cl. Calluno-Ulicetea and O. Vaccinio myrtilli-Genistetalia pilosae Calluna vulgaris 80 50 30 10 5 8 10 5 Carex ericetorum + + + + 1 1 + . Dicranum scoparium . . . . + 2 1 + Hieracium umbellatum + + + + + + + + Vaccinium myrtillus 1 + 1 . . . . . Accompanying species Cl. Vaccinio-Piceetea Ajuga reptans . . + . . . . . Cladina rangiferina . . . . 1 1 . . Cladonia uncialis . . . . 1 + . 1 Dicranum rugosum ...... 4 Luzula pilosa . . 1 . . . . . Melampyrum pratense . . . + . . . . Pinus sylvestris (b) 1 1 1 1 1 + + 3 Pinus sylvestris (с) . . . . + 1 1 1 Pleurozium shreberi 2 . 4 3 5 5 5 4 Polytrichum juniperinum . . . 1 2 2 2 3 Pulsatilla patens ...... + Trientalis europaea . . . . + + + + Vaccinium uliginosum . . . . . + + 2 Vaccinium vitis-idaea . . . . + 1 1 1 Cl. Molinio-Arrhenatheretea Achillea submillefolium + ...... Agrostis stolonifera ...... + [78]

Region Right-bank Polesie Left-bank Polesie Relevé number 1 2 3 4 5 6 7 8 Briza media + 1 ...... Carex hirta . + ...... Festuca rubra . . 1 2 . . . . Holcus lanatus . 1 ...... Ranunculus acris . + ...... Sieglingia decumbens . . 3 . . . . . Cl. Nardetea strictae Genista germanica ...... + Luzula campestris . . . . . + + + Lycopodium clavatum . . + . + + + . Nardus stricta . . + . . . . . Pilosella officinarum + + + + + + . + Potentilla erecta . + . . + . . + Veronica officinalis + . . + + + + . Cl. Koelerio-Corynephoretea canescentis Chamaecytisus ruthenicus . . + . . . . . Festuca ovina . . . 2 3 3 3 . Jasione montana + + ...... Koeleria glauca . . . . 2 3 3 . Potentilla argentea + 1 ...... Rumex acetosella . . . + . . . . Solidago virgaurea . . + . . . . . Cl. Crataego-Prunetea Crataegus monogyna + + ...... Galeopsis bifida . . + . . . . . Prunus spinosa . + ...... Rosa canina + + ...... Cl. Epilobietea angustifolii Calamagrostis epigeios + 1 . + . . . . Rubus idaeus . . . . + . . . Sambucus racemosa . . . . + . . . Cl. Festuco-Brometea Centaurea scabiosa . + ...... Euphorbia cyparisias + 1 ...... Cl. Trifolio-Geranietea sanguinei Anemone sylvestris + ...... Galium verum + + ...... Phytocenotic features of Calluna vulgaris (L.) Hill. in Ukrainian Polesie 79

Region Right-bank Polesie Left-bank Polesie Relevé number 1 2 3 4 5 6 7 8 Cl. Erico-Pinetea Daphne cneorum + ...... Cl. Papaveretea rhoeadis Viola arvensis . + ...... Other species Betula pendula (b) + + . + . . . . Betula pendula (с) . + ...... Calamagrostis arundinacea ...... 2 Cerasus mahaleb + ...... Genista tinctoria . . + . . . . . Holcus mollis + + ...... Hypericum perforatum + + . 2 . . . . Populus tremula (b) . + 1 . . . . . Populus tremula (с) . + + . . . . . Pteridium aquilinum . . . + . . . . Pyrus communis (b) + 1 ...... Quercus robur (b) . . 1 . . . . . Veronica chamaedrys . . + . . . . .

Notes. Dates, localities and authors of relevés: 1 – 12.08.2013, the Rivne shooting range, the Hoshchansk, Kostopil and Rivne districts, Rivne region; V. Melnyk. 2 – 21.07.2014, the shooting range near the northern outskirts of the Volodymyr-Volynskyi city, Volyn region; V. Melnyk. 3 – 21.08.1972, the Dubivka forestry, (sq. 36), Volodymyrets district, Rivne region; T. Andrienko. 4 – 15.06.2003, the outkirts of the Lubenets village, Korop district, Chernihiv region; Yu. Karpenko. 5 – 10.06.2018, the Liubetskyi Masyv tract, Liubech forestry (sq. 20), Ripky district, Chernihiv region; O. Lukash, V. Popruha. 6 – 10.06.2018, the Liubetskyi Masyv tract, Liubech forestry (sq. 25), Ripky district, Chernihiv region; O. Lukash. 7 – 10.06.2018, the Liubetskyi Masyv tract, Liubech forestry (sq. 25), Ripky district, Chernihiv region; O. Lukash. 8 – 05.07.1980, the Chudivka forestry, (sq.30), Ripky district, Chernihiv region; T. Andrienko.

areas of 0.5-2 ha, and are distributed on sandy soils. Some- caninae alliance are represented by Juncus squarrosus L. times they represent a stage of a post-pyrogenic succession. (in Right-bank Polesie) or Pimpinella saxifraga L. (in Left- The relevés in Table 2 give the idea of full composition and bank Polesie). Right-bank heathlands are characterized by structure of the heathlands. Calluna vulgaris is a diagnostic high constancy of Sieglingia decumbens (L.) Bernh, but species of the association. Its projective coverage varies it does not dominate. For comparison, this species dom- from 15% to 90%. Nardus stricta is a characteristic spe- inates in communities of the Calluno-Nardetum strictae cies of the Nardetea strictae class. Its role in phytocenoses association in Poland (W. Matuszkiewicz, 2019), but it is is much smaller if compared with Calluna vulgaris. The absent in the relevés of Left-bank Polesie. The left-bank qualitative and quantitative characteristics of these phyto- heathlands are characterized by high constancy of a num- cenoses are different in Right-bank Polesie and Left-bank ber of characteristic species of the Molinio-Arrhenathere- Polesie. Viola canina L. (the diagnostic species of the Cal- tea Tx. 1937 class and its syntaxons. For example, Gen- luno-Nardetum strictae association) in Right-bank Polesie, tiana pneumonanthe L., a regionally rare species of the unlike in Left-bank Polesie, is not a constant component Chernihiv region, is present in all the relevés. of phytocenoses. The characteristic species of the Violion [80] . . . 2 4 + + + + + + + + + + + 21 90 10 40 . . . . . 0 2 0 2 4 + + + + + + + 20 90 30 . . . . . 0 5 1 0 5 + + + + + + + + 19 95 Left-bank Polesie . . . 1 0 1 3 + + + + + + + + + 18 15 80 60 ...... + + + + + + 17 40 80 40 30 ...... 1 2 1 + + + + 16 70 80 60 <1 association ...... 4 + + + + 15 50 45 30 15 ...... + + + + 14 40 55 50 20 ...... + + + + + + 13 70 40 60 25 ...... + + + + + 12 50 45 35 20 ...... + + + + + + + 11 70 15 10 80 ...... + + + + + + 10 60 40 60 35 Calluno-Nardetum strictae class: the Calluno-Nardetum ...... 9 4 1 + + + 15 80 10 40 ...... 5 8 + + + + + + 75 50 40 Right-bank Polesie ...... 7 1 1 + + 50 70 30 10 ...... 6 2 + + + + + + + 80 70 35 ...... 5 0 + + + + + 60 15 70 ...... 1 4 2 1 . + + 40 30 15 ...... 0 1 3 + + + + + + + 55 15 50 ...... 2 4 1 + + + 30 75 50 15 ...... 1 2 + + + + + 60 70 10 15 Region Relevé number Relevé Nardetea strictae of the Nardetea communities plant Polesie heath of the Ukrainian structure Floristic Shrub layer (b) cover [%] Shrub layer Accompanying species Cl. Molinio-Arrhenatheretea submillefolium Achillea Agrostis stolonifera Agrostis Herb and dwarf shrub layer (c) cover [%] Herb and dwarf shrub layer Mosses layer (d) cover [%] Mosses layer D. Ass D. Calluno-Nardetum strictae Calluna vulgaris Viola canina Viola Ch. All. Violion caninae Hypochaeris uniflora Juncus squarrosus Pimpinella saxifraga Pimpinella Ch. O. Nardetalia strictae Ch. O. Nardetalia Hypericum maculatum Nardus stricta Nardus Potentilla erecta Potentilla Lycopodium clavatum Lycopodium Pilosella officinarum Pilosella Veronica officinalis Veronica Ch.Cl. Nardetea strictae dioica Antennaria Luzula campestris Carex pilulifera Carex Table 2. Table [81] ...... 2 . . + + + + 21 ...... 2 1 + + + + + + + 20 ...... + + + + + + + 19 Left-bank Polesie ...... 1 1 + + + 18 ...... 4 . + 17 ...... 1 + + + 16 ...... 1 + + 15 ...... 3 + 14 ...... + 13 ...... 12 ...... + 11 ...... 1 10 ...... 9 + ...... 8 + Right-bank Polesie ...... 7 4 + + ...... 6 4 + ...... 5 + + ...... 4 2 1 + ...... 3 1 3 3 1 + + + + ...... 2 2 3 + + + + + + + ...... 1 5 + Region Relevé number Relevé Alchemilla micans Alchemilla Lythrum salicaria Lythrum Anthoxanthum odoratum Anthoxanthum Carex echinata Carex Carex leporina Carex Carex panicea Carex Centaurea jacea Centaurea Mentha arvensis Molinia caerulea Molinia Cnidium dubium Phleum pratense Poa nemoralis Prunella vulgaris Prunella Coccyganthe flos-cuculi Coccyganthe Deschampsia caespitosa Festuca rubra Festuca Galeopsis pubescens Ranunculus acris Galium uliginosum Ranunculus repens Gentiana pneumonanthe Juncus effusus Juncus tenuis Leontodon autumnalis Lysimachia vulgaris Lysimachia Luzula pallidula [82] ...... 2 . 21 ...... + 20 ...... + 19 Left-bank Polesie ...... 18 ...... 4 + + + + + 17 ...... + + + 16 ...... 2 4 + 15 ...... 2 1 4 + + + 14 ...... 1 5 5 + + + 13 ...... 2 1 3 4 5 2 + + 12 ...... 3 5 1 5 + 11 ...... 4 1 5 4 + + + 10 ...... 9 1 2 + + + ...... 8 1 4 + + + + + + Right-bank Polesie ...... 7 1 3 2 + + + ...... 6 2 5 + ...... 5 2 5 + + ...... 4 + ...... 3 2 + + ...... 2 + + + ...... 1 4 2 + + + Region Relevé number Relevé Trientalis europaea Trientalis Vaccinium vitis-idaea Vaccinium Sieglingia decumbens Sieglingia Polytrichum juniperinum Polytrichum Rhododendron luteum Rhododendron Vaccinium uliginosum Vaccinium Succisa pratensis Trifolium repens Trifolium Viola uliginosa Viola Cl. Scheuchzerio palustris-Caricetea Cl. Scheuchzerio fuscae canina Agrostis Cl. Vaccinio-Piceetea uva-ursi Arctostaphylos Chamaecytisus ruthenicus Dianthus pseudosquarrosus Dicranum rugosum Dryopteris carthusiana Frangula alnus (b) Luzula pilosa Melampyrum pratense Pinus sylvestris (b) Pinus sylvestris (с) Pleurozium shreberi shreberi Pleurozium Polytrichum commune Polytrichum Polytrichum gracile Polytrichum [83] ...... 2 + 21 ...... 20 ...... + 19 Left-bank Polesie ...... + 18 ...... 4 + + + 17 ...... + 16 ...... + 15 ...... 1 + 14 ...... + 13 ...... 2 1 + 12 ...... 1 11 ...... 10 ...... 9 ...... 8 Right-bank Polesie ...... 7 ...... 6 ...... 5 + ...... 4 1 4 + + ...... 3 + ...... 2 4 + + ...... 1 Region Relevé number Relevé Carex echinata Carex Cl. Oxycocco-Sphagnetea polifolia Andromeda Drosera rotundifolia Drosera Carex nigra Carex Juncus articulatus Lycopodiella inundata Lycopodiella Polytrichum strictum Polytrichum Sphagnum compactum Sphagnum cuspidatum Viola palustris Viola Cytisus zingeri Sphagnum fallax Cl. Koelerio-Corynephoretea canescentis Cl. Koelerio-Corynephoretea Corynephorus canescens Jasione montana Koeleria glauca Koeleria Polytrichum piliferum Polytrichum Cl. Alnetea glutinosae Rumex acetosella Oberna bechen Cl. Epilobietea angustifolii Cl. Epilobietea epigeios Calamagrostis Gnaphalium sylvaticum Rubus idaeus Chamaerion angustifolium [84] ...... 21 ...... 20 ...... + + 19 Left-bank Polesie ...... 18 ...... 2 + 17 ...... 5 + + + 16 ...... 5 + + 15 ...... 4 + 14 ...... 4 13 ...... 12 ...... 11 ...... 2 + 10 ...... 9 3 ...... 8 4 Right-bank Polesie ...... 7 4 + + + ...... 1 6 1 + + ...... 5 + + + + ...... 1 4 + + ...... 3 1 + + ...... 2 + + ...... 1 1 3 + + Region Relevé number Relevé Genista tinctoria Betula pubescens (b) Betula Betula pendula (с) Betula Holcus mollis Lycopus europaeus Lycopus Hypericum perforatum Salix aurita Salix Salix cinerea Salix Cl. Isoëeto-Nanojuncetea Gnaphalium luteoalbum Gypsophila muralis Hypericum humifusum Sagina nodosa Cl. Carpino-Fagetea sylvaticae Hypericum montanum Oxalis acetosella Ranunculus auricomus Cl. Phragmito-Magnocaricetea vesicaria Carex Carex vulpina Carex Thyselium palustre Cl. Festuco-Brometea praecox Carex Other species Other capillaris Agrostis Betula pendula (b) Betula [85] ...... + 21

...... 20 ...... 19 Left-bank Polesie ...... + 18 . . . . . + + 17 . . . . + + + 16 ...... 15 ...... 14 ...... 13 . . . . . 2 + 12 . . . . . 2 + 11 . . . . . 1 + 10 ...... 9 2 . . . . . 8 1 + Right-bank Polesie forestry, Kopyshche Zhytomyr Andrienko. forestry, region; T. forestry, Kopyshche Zhytomyr Andrienko. forestry, region; T...... 7 ...... 6 ...... 5 ...... 4 ...... 3 ...... 2 . . . . 1 1 + Region Relevé number Relevé 18.08.2006, between the peaty meadow and pine forest, to the east of the village, Buda Novhorod-Siverskyi Vorobyivska district, Chernihiv region, N 52°22’43,2’’; E 032°31’51,0’’; O. Lukash. E 032°31’51,0’’; – 27.08.1972, the Kurchytsia forestry (sq. district, 57), Zhytomyr Andrienko. Novohrad-Volynskyi region; T. – 25.07.1981, the Polesie Nature Reserve, the Selezivka forestry (sq. 31), Ovruch district, Zhytomyr Andrienko. region; T. – 23.07.1981, the Polesie Nature Reserve, the Selezivka forestry (sq. 19), Ovruch district, Zhytomyr Andrienko. region; T. – 07.06.1978, the dunes, 6 forestry, the km Vysotsk from village, the Dubrovytsia Verbivka district, Rivne Andrienko. region; T. – 11.06.1982, the post-pyrogenic succesion stage, the Polesie Nature Reserve (sq.41), Zhytomyr Andrienko. region; T. – 15.07.2013, the heathland (2 ha), Mokrets tract near the Uhlova Rudnia village, Ripky district, Chernihiv region; O. Popruha. Lukash, V. – – 27.07.1996, the meadow between the swamp and the pine forest, outskirts of the Smolianka village, Kulykivka district, Chernihiv region; O. Lukash. – 11.08.1985, the floodplain of the Prypiat river, near the Nobel lake, the Zarichansk district, Rivne Andrienko. region; T. – 11.08.1985, the floodplain of the Prypiat river, the Zarichansk district, Rivne Andrienko. region; T. – 03.08.1980, the Polesie Nature Reserve, the floodplain of the Kamenka river, the – 23.06.1981, the Polesie Nature Reserve, the Perha Zhytomyr Andrienko. forestry, region; T. – 14.06.1982, the Polesie Nature Reserve, (sq. 50), the Zhytomyr Andrienko. region; T. – 11.06.1982, the post-pyrogenic succesion stage, the Polesie Nature Reserve, (sq.42), the Zhytomyr Andrienko. region; T. – 03.08.1980, the Polesie Nature Reserve, the floodplain of the Kamenka river, the – 12.08.1980, the wasteland, near village, the Berezne Verbivka district, Rivne Andrienko. region; T. 1 2 3 4 Andrienko T. Rivne region; district, 5 – 13.08.1980, the sands, Berezne 6 7 8 9 Phalacroloma annuum Phalacroloma Populus tremula (b) Populus tremula Populus tremula (с) Populus tremula Pteridium aquilinum Pteridium Quercus robur (b) robur Quercus Sorbus aucuparia (b) Veronica chamaedrys Veronica Notes. Dates, localities and authors of relevés: 10 Andrienko. T. Rivne region; district, Berezne – 25.08.1972, the forest area at Malynsk village, 11 Andrienko. T. Rivne region; 12 – 23.08.1972, the Sarny forestry (sq. 147), district, Andrienko. T. Rivne region; 13 – 23.08.1972, the Sarny forestry (sq. 147), district, 14 15 16 17 18 O. Lukash. region; Chernihiv Borzna district, and Sidorivka villages, the Berestovets 19– 19.07.1995, the meadow between 20 21 86 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko

In Right-bank Polesie Calluna vulgaris is a compo- Reserve (the Zhytomyr region). Calluna vulgaris partici- nent of phytocoenoses of dwarf-shrub, sedge and peat- pation in mire phytocenoses varies depending on a stage moss vegetation of the Holarctic ombrotrophic bogs and of mire development. The highest projective coverage of wet heaths on extremely acidic soils. These phytocoenoses Calluna vulgaris (up to 60-80%) is observed in succession belong to the Oxycocco-Sphagnetea class (Table 3). There stages after wildfires. All the forest oligotrophic bogs with are no such biotopes in Left-bank Polesie, as they were de- Calluna vulgaris (Table 3), described by prof. T. Andrien- stroyed in 1970s-1980s as a result of drainage reclamation. ko, were identified, according to W. Matuszkiewicz (2019), Most of plant communities of the Oxycocco-Sphagnetea as belonging to the Ledo-Sphagnetum magellanici associa- class, represented by the Sphagnetalia medii order and the tion, because Ledum palustre L. (the diagnostic species of Sphagnion medii alliance, can be seen in the Polesie Nature this association) was present in all relevés.

Table 3. Floristic structure of the forest bogs plant communities (Ledo-Sphagnetum magellanici association) with Calluna vulgaris

Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Tree layer (a) cover [%] 0 5 0 0 0 50 0 0 40 40 0 0 70 0 40 0 0 Shrub layer (b) cover [%] 50 50 50 60 30 10 40 40 20 75 70 40 20 40 <1 20 50 Herb and dwarf shrub layer (c) 60 75 80 80 65 60 80 60 60 75 70 70 80 60 65 80 90 cover [%]

Mosses layer (d) cover [%] 70 20 20 40 80 60 95 98 90 80 40 70 40 95 98 30 0 D. Ass. Ledo-Sphagnetum magellanici Ledum palustre 1 + + + + 4 1 + 3 1 + + 4 1 1 1 1 Oxycoccus microcarpus ...... + . + . . . + . . . D. Ass. Gr. of the forest bogs (Ch. Cl. Vaccinio-Piceetea) Pinus sylvestris (a) . 1 . . . 4 . . 4 1 . . 5 . 4 . . Pinus sylvestris (b) 3 2 1 1 3 1 4 4 3 4 5 4 + 4 + + + Pinus sylvestris (с) + ...... + + 1 . . . Pleurozium shreberi + . . . . . + . . . 3 . 4 . . . 1 Trientalis europaea . . . . + . . . + ...... + . Vaccinium myrtillus . + . . . + + ...... + Vaccinium uliginosum + 1 + . . + + . . + + . + . . . 2 Vaccinium vitis-idaea 1 + ...... + . 1 . . + . Ch. All. Sphagnion medii and O. Sphagnetalia medii Andromeda polifolia . . + + 1 + + + 1 + + . + 1 2 + . Eriophorum vaginatum . . . + 4 1 4 4 3 1 1 + 1 4 4 4 + Oxycoccus palustris . . 1 . 1 + 3 + 2 3 . . + 1 1 1 . Sphagnum acutifolium 4 . . . . 3 3 . . . 3 ...... Sphagnum centrale ...... + . 5 + . Sphagnum compactum ...... 3 [87]

Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Sphagnum cuspidatum . . 1 4 2 . . 5 5 . . 4 . . . . . Sphagnum fallax . . . . 4 . 5 ...... Sphagnum fuscum . 3 ...... + . 1 . . Sphagnum magellanicum + + + . 3 . . 3 2 . + . . + . 4 + Sphagnum rubellum ...... 5 . + . 5 1 + . Ch. Cl. Oxycocco-Sphagnetea Aulacomnium palustre . . + . . 3 + + . . . 2 . . . . + Drosera rotundifolia + + + + + . . + + + + . + + + + . Salix aurita . + . + + ...... Accompanying species Cl. Alnetea glutinosae Alnus glutinosa (b) + ...... Betula pubescens (a) . . . . . 2 ...... Betula pubescens (b) . . 4 5 . 2 . . . 2 1 2 . . . 3 4 Betula pubescens (с) ...... + . . . . . Salix cinerea + + ...... Cl. Vaccinio-Piceetea Dicranum rugosum ...... 1 . . . . Melampyrum pratense + ...... Polytrichum commune + . . 1 ...... 4 . . . . . Polytrichum gracile . 1 ...... Cl. Scheuchzerio palustris- Caricetea fuscae Carex lasiocarpa . . . 2 1 . . . . + . . . + . . . Carex nigra + + ...... + . . . . . Rhynchospora alba . + . 1 ...... + . Cl. Molinio-Arrhenatheretea Molinia caerulea 2 2 . . . . + . . + 2 4 3 . . . . Juncus effusus 1 ...... + . . . . . Cl. Nardetea strictae Nardus stricta . + ...... Potentilla erecta . + ...... + . . . . . Cl. Calluno-Ulicetea Arctostaphylos uva-ursi ...... + ...... Calluna vulgaris 40 60 65 60 1 5 2 1 10 50 60 25 1 5 10 30 80 88 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko

Relevé number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Cl. Koelerio-Corynephoretea canescentis Astragalus arenarius ...... + ...... Other species Betula pendula (a) ...... + ...... 3 . . Betula pendula (b) 4 4 . . 5 ...... 3 2 + . . Betula pendula (с) . 2 ...... 2 + + . . Populus tremula (b) . 1 + + ...... + . Populus tremula (с) ...... + . Pteridium aquilinum . + ...... Polytrichum alpestre . . 3 . . 4 + 1 2 ......

Notes. Dates and localities of relevés: 1 – 16.08.1980, the Sarny forestry (sq. 9), Sarny district, Rivne region. 2 – 13.08.1980, the post-pyrogenic succession stage, the Berezne district, Rivne region. 3 – 10.08.1980, the post-pyrogenic succession stage, the Pochaiv Reserve (sq. 76), Berezne district, Rivne region. 4 – 10.08.1980, the post-pyrogenic succession stage, the Pochaiv Reserve (sq. 76), Berezne district, Rivne region. 5 – 12.07.1982, the Plotnytsia tract, Chervona Volia forestry (sq. 7), Novohrad-Volynskyi district, Zhytomyr region. 6 – 10.07.1982, the Chervona Volia forestry (sq. 1), Novohrad-Volynskyi district, Zhytomyr region. 7 – 22.10.1974, the Perebrody forestry (sq. 81), Dubrovytsia district, Rivne region. 8 – 27.07.1976, the Rostan forestry (sq. 21), Shatsk district, Volyn region. 9 – 16.07.1973, the Babii Mokh swamp, Zolote forestry (sq. 39), Dubrovytsia district, Rivne region. 10 – 25.06.1981, the Polesie Nature Reserve, the Klitne tract, Kopyshche forestry (sq. 47), Zhytomyr region. 11 – 25.06.1981, the post-pyrogenic succession stage, the Polesie Nature Reserve, the Klitne tract, Kopyshche forestry (sq. 38), Zhytomyr region. 12 – 25.06.1981, the Polesie Nature Reserve, Kopyshche forestry (sq. 26), Zhytomyr region. 13 – 13.06.1982, the swamp forest, the Polesie Nature Reserve (sq. 51), Zhytomyr region. 14 – 12.06.1982, the oligotrophic swamp, the Polesie Nature Reserve (sq. 40), Zhytomyr region. 15 – 12.06.1982, the oligotrophic swamp, the Polesie Nature Reserve (sq. 41), Zhytomyr region. 16 – 11.06.1982, the post-pyrogenic succession stage, the Polesie Nature Reserve (sq. 41), Zhytomyr region. 17 – 07.06.1978, the Vysotsk forestry (sq. 75), near the Verbivka village, Dubrovytsia district, Rivne region. Author of relevés: T. Andrienko.

The European temperate and subboreal pine forests The maximum projective coverage of Calluna vulgaris on nutrient-poor acidic sandy soils (the Vaccinio-Piceetea in plant communities of the Pinetalia sylvestris order asso- class) are typical and most common habitats of Calluna ciations (the Dicrano-Pinion sylvestris alliance) in Right- vulgaris in Ukrainian Polesie. The species occurs in phyto- bank Polesie respectively is: Querco-Piceetum – 45%, cenoses mainly individually. The maximum projective cov- Cladonio-Pinetum – 40%, Molinio-Pinetum – 35%, Peu- erage of Calluna vulgaris was observed in pine, birch-pine cedano-Pinetum – 30%, Vaccinio uliginosi-Pinetum – 10% and oak-pine forests of 30-40 years of age. Tables 4 and (Table 4, relevés 1-26). Calluna vulgaris coverage of 40% 5 provide information on structure and floristic composi- was observed only in the phytocenoses of the Cladonio-Pi- tion of forest phytocenoses with Calluna vulgaris, which netum association (Table 5, relevés 1-5) in Left-bank Pole- projective coverage is over 1%. As it can be seen, all these sie. The projective coverage of Calluna vulgaris in other plant communities were well diagnosed by characteristic plant communities of pine and mixed forests of Left-bank and differentiating species. The relationships between the Polesie does not exceed 5% (Table 5, relevés 6-23). values of tree layer and projective coverage of Calluna vulgaris were not found. [89] ...... 0 0 4 + . . . 50 50 15 75 29 ...... 0 6 . . 0 . 4 + + 45 40 10 65 28 ...... 0 0 4 + 40 50 20 60 27 ...... 0 0 4 50 60 10 50 26 ...... 0 0 . . . 1 . . . . 85 25 20 60 25 Calluna vulgaris in Right- 5 ...... 0 . . 0 . 5 3 90 40 70 24 ...... 0 0 + 75 80 10 60 23 ...... 0 0 + + 70 50 20 70 22 4 ...... 0 0 4 + + . 30 70 30 40 21 ...... 0 . 5 . 0 . 3 5 + 65 20 70 20 ...... 0 0 . 3 . . 3 . 10 70 30 60 19 ...... 0 0 1 1 + + 70 65 60 18 ...... 3 0 0 1 + + . 90 50 10 70 17 ...... 0 . . 0 . 0 4 + + 40 50 50 16 ...... 0 0 . . 1 . 4 . + + 60 55 60 15 ...... 0 0 2 2 + + 10 95 40 40 14 ...... + 0 + 0 + 50 50 30 60 13 ...... + + 0 0 . + 40 50 10 50 12 ...... + 0 + 0 . . . . + + 30 11 60 25 60 2 ...... + + 0 0 . + 75 25 20 60 10 ...... 1 + 0 0 + 1 9 + 80 40 60 ...... + + 0 + 0 1 8 + + 80 75 50 <1 . . . . 5 . . 0 0 . . + 5 . . 7 . . 65 40 50 ...... 1 0 0 + + 6 + + 55 45 25 70 ...... 5 0 0 + 5 + 80 10 40 <1 ...... 1 0 0 + + 4 + 70 40 15 50 . 4 . . 0 0 + . . + . . 3 . . . + 70 15 25 50 ...... 0 0 + + + 0 2 85 20 50 ...... 2 0 + 0 5 1 40 40 20 Floristic structure of the forest plant communities of the Cladonio-Vaccinietalia (relevés 1-26) and Vaccinio-Piceetalia (relevés 27-29) orders with orders 27-29) (relevés Vaccinio-Piceetalia and 1-26) (relevés Cladonio-Vaccinietalia the of communities plant forest the of structure Floristic bank Polesie Relevé number number Relevé Polygonatum odoratum Polygonatum Peucedanum oreoselinum D. Ass. D. Peucedano-Pinetum majalis Convallaria Koeleria glauca Koeleria Cladonia mitis Euonymus verrucosa Syntaxon D. Ass. D. Cladonio-Pinetum ericetorum Carex Corylus avellana Lichens layer (e) cover [%] Lichens Carpinus betulus (c) Mosses layer (d) cover [%] Mosses layer D. Ass. D. Querco-Pinetum, species of mixed forests Carpinus betulus (b) Mosses and lichens layer (d- Mosses and lichens e) cover [%] Frangula alnus Herb layer (c) cover [%] Herb layer Polytrichum commune Polytrichum Shrub layer (b) cover [%] Shrub layer D. Ass. D. Molinio-Pinetum caerulea Molinia Tree layer (a) cover [%] layer Tree Solidago virgaurea Table 4. Table [90] ...... 1 ...... + 29 ...... + . 28 ...... + + 27 ...... + . + + + + + 26 ...... 5 . . . + . . . . + . + . + 25 ...... 1 . + + + 24 ...... + 2 + + + 1 + 23 ...... + 4 + 3 3 . + + + . + 22 ...... + + 4 . . + + + + 21 ...... + + 20 ...... + ...... + 19 ...... 3 . + + + 18 ...... 4 1 . + 17 ...... + . + + 16 ...... 2 . + ...... 15 ...... + + + 14 ...... 1 2 + 13 ...... 3 + 1 . + . + 12 . . . . . 4 + + . . . . . + . . . . . + 11 ...... 3 . + 10 ...... 3 + . + + 9 ...... + + + 1 8 + . . . . 1 + + ...... 7 ...... 3 . + . 6 ...... 1 + 5 ...... 1 4 4 . . . . + 4 ...... 3 ...... 3 + 2 ...... 1 + + 1 uliginosi- Vaccinio Piceo- Relevé number number Relevé Sorbus aucuparia (b) Rhododendron luteum Rhododendron Melampyrum pratense Juniperus communis (b) Luzula pilosa Leucobryum glaucum Ch. All. Ch. Dicrano-Pinion D. SAll. Dicrano-Pinenion Dicranum rugosum Vaccinium uliginosum Vaccinium (c) robur Quercus D. SAll. uliginosi Vaccinienion aurita Salix (b) robur Quercus Oxycoccus palustris (a) robur Quercus Eriophorum vaginatum Stellaria holostea Stellaria Aulacomnium palustre Mycelis muralis Mycelis D. Ass. Vaccinio uliginosi- Ass. D. Vaccinio Pinetum polifolia Andromeda Melica nutans Melica Ch. Ass. Ch. Pinetum Ledum palustre [91] . . . 1 1 + 3 5 1 + . + + . + . . . + . + 29 ...... 1 1 2 + + 2 5 1 + + + + 28 ...... 1 1 + + 3 4 3 + + + + + + + 27 ...... 4 . . . 4 . + . + 26 ...... 4 3 5 + . . . 25 ...... 5 1 5 . + . . . 24 ...... 5 1 5 + . 23 ...... 4 . . 1 . 5 . + 22 ...... + . . . . . + . . 21 ...... 1 1 . . . 20 ...... 3 5 . 2 + 1 + . 19 ...... 4 4 . . . . + 18 ...... 5 5 . . . . 17 ...... 4 1 . . . . 16 ...... 4 . . . 5 . + 15 ...... 3 3 . 14 ...... 4 1 5 + . + . . . + 13 ...... 3 4 . 12 ...... 3 . 4 + + 11 ...... 4 2 3 5 + + 10 ...... 5 4 + + 9 + ...... 4 + + 8 + ...... 1 . . 1 . 4 + 3 . . . 7 ...... 4 5 + 6 ...... 4 5 ...... 3 1 2 1 4 ...... 1 . . 3 . 4 . . . 3 ...... 4 3 2 ...... 1 1 3 1 + - femina Relevé number number Relevé Ch. All. Piceion abietis All. Piceion Ch. and O. Vaccinio-Piceetalia Athyrium filix Sphagnum girgensohnii Betula pubescens (c) Betula Betula pubescens (b) Betula Trientalis europaea Trientalis Ch., Ass. D. Sphagno girgensohnii-Piceetum pubescens Betula Pleurozium shreberi Pleurozium Cladina rangiferina Ch. Cl. Vaccinio-Piceetea Hylocomium splendens Pinus sylvestris (c) Picea abies (b) Picea Pinus sylvestris (b) Picea abies (a) Picea Pinus sylvestris (a) Orthilia secunda Ch., D. O. Cladonio- Vaccinietalia uva-ursi Arctostaphylos Mnium spinosum Monotropa hipopitys Monotropa Mnium affine Chimaphila umbellata Lycopodium annotinum Lycopodium [92] ...... 4 29 ...... 4 . . + 28 ...... 4 . . + + 27 ...... 1 . + 26 ...... 2 . . + . . 25 ...... 1 24 ...... 5 . + 23 ...... 3 . 3 + . + 22 ...... + 4 . . + + 21 ...... + + + + 20 ...... 1 . 3 + 19 ...... 1 . 4 . . 18 ...... 2 . . + 17 ...... 1 1 . 16 ...... 1 . + . . . + 15 ...... 1 . + + + + 14 ...... 4 2 . 1 . 13 ...... + . 4 + . + . + 12 ...... + 1 . 4 2 + 11 ...... + 10 ...... + + 9 + + ...... 1 4 8 ...... + 1 . . 7 . + ...... + . + . 1 6 + + ...... + . 1 . 5 ...... 2 . 4 + ...... 1 . 1 . 1 . 3 ...... 1 2 + ...... 1 + 2 1 multiflora Relevé number number Relevé Festuca rubra Festuca Cl. Molinio- Arrhenatheretea odoratum Anthoxanthum Thymus serpyllum Rumex acetosella Polytrichum piliferum Polytrichum Lembotropis nigricans Lembotropis Lycopodium clavatum Lycopodium Jasione montana Luzula Festuca ovina Festuca Cl. Nardetea strictae Juncus squarrosus Cytisus zingeri Sieglingia decumbens Sieglingia Corynephorus canescens Poa angustifolia Accompanying species Accompanying Cl. Koelerio- Corynephoretea canescentis Chamaecytisus ruthenicus Plantago lanceolata Plantago Vaccinium vitis-idaea Vaccinium Lysimachia vulgaris Lysimachia Vaccinium myrtillus Vaccinium Gentiana pneumonanthe [93] ...... + 1 + + 29 ...... 1 1 . + . + + 28 ...... 1 1 . . + 1 . . . 27 ...... 1 . . . . 26 ...... + . 10 25 ...... 5 . + . + 24 ...... 1 + . . . . 23 ...... 7 . . . . . 22 ...... + . . . 45 21 ...... + 2 . + + + 20 ...... + 5 . . . + . 19 ...... 15 18 ...... + . . . . . 35 17 ...... 5 . . . 16 ...... + 10 15 ...... + + + 1 4 . + 25 14 ...... + 1 1 . . + . 13 ...... 2 . + + . + 12 ...... + 5 . . . + . + . . 11 ...... 5 + . . 10 ...... + . 9 + . 30 ...... 5 . . 8 ...... + . + . + . 7 . + . 40 ...... + + + 6 + 40 ...... + 5 + 5 + ...... 4 30 ...... 7 + . + . . . 3 . . + ...... 2 10 ...... + 1 40 Relevé number number Relevé Cl. Alnetea glutinosae Alnus glutinosa (a) Maianthemum bifolium Holcus lanatus Cl. Quercetea robori- Cl. Quercetea petraea Genista tinctoria Scorzonera humilis Hieracium umbellatum Sarothamnus scoparius Sarothamnus Cl. Scheuchzerio palustris- Cl. Scheuchzerio fuscae Caricetea canina Agrostis Carex ericetorum Carex Oxalis acetosella Cl. Calluno-Ulicetea Calluna vulgaris Cl. Carpino-Fagetea sylvaticae Dryopteris filix-mas Veronica officinalis Veronica Fragaria vesca Potentilla erecta Potentilla Cl. Epilobietea angustifolii Cl. Epilobietea epigeios Calamagrostis Pilosella officinarum Pilosella Salix cinerea Salix Nardus stricta Nardus Carex cinerea Carex [94] ...... 1 + + . 29 ...... + . + . . 28 ...... 1 . . . + + 27 ...... 2 + . . . . . + + 26 ...... + + + ...... + 25 ...... 1 . . . 24 ...... 2 . + . . . . 23 ...... 22 ...... 4 + + 4 . 21 ...... 4 1 . + 4 . + + 20 ...... + 19 ...... 1 3 ...... 18 ...... 2 . . + . . 17 ...... 4 . . 16 ...... 1 5 . . + . . . 15 ...... 3 3 . . . + . . . 14 ...... 3 + . . . 13 ...... 1 2 . . . . 12 ...... 3 + + . 11 ...... 2 . . . . 10 ...... 1 3 + . + 9 ...... + 8 ...... 1 2 . 1 . . . 7 ...... 3 . . 6 ...... 1 + 5 . + ...... 2 4 . 4 ...... 1 . . . . . 3 . + ...... 2 ...... 1 juncella Relevé number number Relevé Betula pendula (c) Betula Betula pendula (b) Betula Betula pendula (a) Betula Other species Other capillaris Agrostis Cl. Phragmito- Magnocaricetea Carex Cl. Trifolio-Geranietea Cl. Trifolio-Geranietea sanguinei Melampyrum nemorosum Populus tremula (c) Populus tremula Cl. Littorelletea uniflorae Cl. Littorelletea Juncus bulbosus Populus tremula (b) Populus tremula Cl. Juncetea maritimi Juncus gerardii Populus tremula (a) Populus tremula Cl. Artemisietea vulgaris Linaria genistifolia Polytrichum alpestre Polytrichum Viola arvensis Viola Malus sylvestris (b) Cl. Papaveretea rhoeadis Cl. Papaveretea segetalis Vaccaria Hypericum perforatum Carex nigra Carex Brachythecium salebrosum Brachythecium [95] . . . 29 . . . 28 . . + 27 . . . 26 . . . 25 . . . 24 . . . 23 . . . 22 . 3 + 21 . . . 20 . . 1 19 . . . 18 . . . 17 , 6 – Sphagno girgensohnii- uliginosi-Pinetum , 5 – Vaccinio . . . 16 . . . 15 . . . 14 Volyn Andrienko. region; T. Volyn . . 1 13 , 4 – Querco-Piceetum . . . 12 . + + 11 3 . . 10 . . . 9 . . 3 8 , 3 – Molinio-Pinetum 7 . . + . . 6 + . . . 5 . . . 4 3 . . . , 2 – Peucedano-Pinetum . . . 2 . . . 1 . Relevé number number Relevé – 08.06.1981, the Ostrivskiy tract, between the Great and Middle lakes, near the Ostrivsk village, Zarichchia district, Rivne Andrienko. region; T. – 26.08.1972, the Horodnytsk near forestry, village, the Zhytomyr Andrienko. Veresovsk region; T. – 24.06.1981, the Polesie Nature Reserve, Kopyshche (sq. forestry, 39), Zhytomyr Andrienko. region; T. – 19.06.1981, the Polesie Nature Reserve, Kopyshche (sq. forestry, 38), Zhytomyr Andrienko. region; T. – 27.08.1972, the (sq. Mala forestry, district, Tsvilia 91), Zhytomyr Andrienko. Novohrad-Volynskyi region; T. – 19.07.1982, the (sq. forestry, Lystvyn 27), Ovruch district, Zhytomyr Andrienko. region; T. – 23.07.1981, the Polesie Nature Reserve, Seleznivske (sq. forestry, 27), Ovruch district, Zhytomyr Andrienko. region; T. – 27.07.1978, the floodplain of the Pripiat river near the Nobel lake, Zarichchia district, Rivne Andrienko. region; T. – 27.08.1972, the (sq. Mala forestry, district, Tsvilia 91), Zhytomyr Andrienko. Novohrad-Volynskyi region; T. – 14.06.1978, the Horodnytsia Novovolynsk forestry, enterprise, (sq. 36), Ivanychi district, – 14.06.1982, the Polesie Nature Reserve, (sq. 50), Zhytomyr Andrienko. region; T. – 25.07.1981, the Polesie Nature Reserve, Seleznivka (sq. forestry, 30), Ovruch district, Zhytomyr Andrienko. region; T. – 11.06.1982, the Polesie Nature Reserve, (sq. 41), Zhytomyr Andrienko. region; T. – 11.08.1991, the Manevychi (sq. forestry, 14), Melnyk Manevychi region; V. district, Volyn – 08.07.1990, the Holovno (sq. forestry, 43), Melnyk Liuboml region; V. district, Volyn – 17.07.1989, the Klitytsk (sq. forestry, 21), Melnyk Kamin-Kashyrskyi region; V. district, Volyn – 27.07.1981, the Polesie Nature Reserve, Selezivka (sq. forestry, 29), Ovruch district, Zhytomyr Andrienko. region; T. – 25.07.1981, the Polesie Nature Reserve, Selezivka (sq. forestry, 29), Ovruch district, Zhytomyr Andrienko. region; T. – 23.07.1981, the Polesie Nature Reserve, Selezivka (sq. forestry, 26), Ovruch district, Zhytomyr Andrienko. region; T. – 06.08.1980, the Polesie Nature Reserve, Kopyshche (sq. forestry, 6), Zhytomyr Andrienko. region; T. – 06.08.1980, the Polesie Nature Reserve, Kopyshche (sq. forestry, 8), Zhytomyr Andrienko. region; T. – 20.10.2012, the Horka tract, near the Svitiaz village, Shatsk region; district, O. Volyn Lukash. – 10.07.1982, (sq. the forestry, district, 5), Chervona Zhytomyr Andrienko. Volia Novohrad-Volynskyi region; T. Rubus caesius Pyrus communis (b) Pteridium aquilinum Pteridium Notes. Dates, localities and authors of relevés: 1 – 28.07.2015, the Rivne Nature Reserve, Ozersk (sq. forestry, 26), Rivne region; О. Rak Andrienko. T. Rivne region; (sq. 14), Dubrovytsia district, 2 – 22.05.1973, the Syra Pohonia reserve, Bilsk forestry, 3 4 5 6 7 8 9 10 – 30.07.1980, the Polesie Nature Reserve, Kopyshche (sq. forestry, 35), Zhytomyr Andrienko. region; T. Andrienko. T. Rivne region; Ostroh district, – 09.08.1972, the Ostroh forestry, 11 12 Andrienko. T. Rivne region; district, (sq. 27), Berezne 13 – 25.08.1972, the Sosnivka forestry, 14 15 16 17 18 19 20 Andrienko. T. Rivne region; (sq. 75), Sarny district, 21 – 14.08.1980, the Sarny forestry, 22 23 24 25 26 27 28 29 1 – Cladonio-Pinetum Plant communities: Piceetum [96] ...... 1 0 0 4 ...... 23 50 40 60 ...... 0 0 2 . . . + . + 22 40 15 40 5 ...... 0 + 21 45 60 15 <1 ...... 1 0 ...... + 20 70 10 25 20 ...... 5 1 0 1 1 . . + + 19 65 25 65 ...... 1 0 4 2 + + 18 55 10 35 60 ...... 3 0 1 . + . . . + . + 17 40 20 45 90 ...... 2 0 . . + + . . . . + 16 90 15 85 <1 ...... 1 2 0 3 + + 15 70 15 70 80 ...... 1 2 0 1 . . . . + . . 14 40 20 15 50 ...... 4 0 3 1 . + . . 13 60 40 30 65 ...... 1 0 + + + + + 12 90 15 95 . . 1 5 1 0 1 1 + . + . . . + . + + 11 45 35 30 ...... 5 5 1 0 1 1 + + + + 10 45 90 Left-bank Polesie Calluna vulgaris in Left-bank 2 ...... 9 0 . . + + + + . + + + 50 15 20 65 . . . . . 8 1 0 1 . + . . . . + + + 90 15 95 <1 ...... 7 2 0 + + + + + 25 10 60 40 ...... 6 2 0 . + + . + + + 30 10 70 50 ...... 5 1 1 3 . . + . + + 35 80 15 <1 ...... 4 5 1 1 + + + 25 35 <1 ...... 3 1 4 . . . . + . + . + 30 10 35 10 <1 ...... 2 1 0 2 + . + + + + 70 25 85 ...... 1 5 1 1 2 1 + 20 50 65 50 Relevé number Relevé Floristic structure of the forest plant communities of order the with Cladonio-Vaccinietalia Solidago virgaurea Tree layer (a) cover [%] layer Tree D. Ass. D. Molinio-Pinetum caerulea Molinia Shrub layer (b) cover [%] Shrub layer Polytrichum commune Polytrichum Herb layer (c) cover [%] Herb layer Frangula alnus Mosses and lichens layer (d-e) cover [%] Mosses and lichens D. Ass. Veronico incanae-Pinetum Ass. D. Veronico Genista tinctoria Mosses layer (d) cover [%] Mosses layer Syntaxon D. Ass. D. Cladonio-Pinetum ericetorum Carex Cladina rangiferina Cladonia uncialis Cladonia gracilis Helictorichon pubescens Helictorichon Jurinea cyanoides Jurinea cyanoides Koeleria glauca Koeleria D. Ass. D. Peucedano-Pinetum majalis Convallaria Peucedanum oreoselinum Polygonatum odoratum Polygonatum Pulsatilla patens Pulsatilla Table 5. Table [97] ...... 1 1 3 3 1 3 + + + + + + 23 ...... 1 3 1 1 . . . . + + + + + + + + + 22 ...... 4 . + + + + + + + + + 21 ...... 5 1 + + + + + + + 20 ...... 5 . . + + + + + + 19 ...... 1 1 5 . . + . . + . + . + + 18 ...... 1 4 2 . + + + + + 17 ...... 4 . . . + 16 ...... 5 1 1 . . + . . . . . + 15 ...... 4 2 . . . . . + + + + 14 ...... 5 . . + 13 ...... 5 . . + 12 ...... 1 4 4 3 + + + 11 ...... 4 . . . + + + + + + 10 ...... 9 4 . . . . . + . . . + ...... 8 5 . + ...... 7 3 . . + + + + + ...... 6 3 ...... + . + ...... 5 4 ...... 4 3 ...... 3 1 4 ...... 2 4 1 5 1 + + + ...... 1 1 3 . Relevé number Relevé Hieracium umbellatum Ch. All. Ch. Dicrano-Pinion Dicranum rugosum Geranium sanguineum Chimaphila umbellata Koeleria grandis Koeleria Dianthus pseudosquarrosus Rubus saxatilis Monotropa hipopitys Monotropa Veronica incana Veronica Polytrichum juniperinum Polytrichum D. Ass. D. Querco-Pinetum, species of mixed forests brizoides Carex D., Ch. O. Cladonio-Vaccinietalia uva-ursi Arctostaphylos Carex sylvatica Carex Pinus sylvestris Carpinus betulus (b) Corylus avellana Euonymus verrucosa Melica nutans Melica Mycelis muralis Mycelis Stellaria holostea Stellaria (a) robur Quercus (b) robur Quercus (c) robur Quercus [98] ...... 1 . . + + + + 23 ...... 1 1 . . + + + + . + + 22 ...... 3 . . + + . + . + + + 21 ...... 1 3 1 1 . . + + + 20 ...... 2 1 5 . . + + + + + . + + + 19 ...... 3 1 1 1 5 + + . + . + + 18 ...... 4 1 1 5 . . + + . + + + + + + + 17 ...... 1 1 . . + . + + 16 ...... 4 3 1 . + . + . + + 15 ...... 1 4 . . . + + + + + + 14 ...... 1 5 . . . . . + + + . + + 13 ...... 3 1 . . + + . + + 12 ...... 4 2 1 4 + + + + + + 11 ...... 5 . . + + + . + + . + + + + 10 ...... 9 1 5 . . . + . + . + + ...... 8 1 1 5 . + + + ...... 7 1 4 . . + + + + + + + + ...... 6 4 4 . . . + . + . + ...... 5 . . . . + + + . + + + ...... 4 . . + + + + + + ...... 3 1 1 . . . . . + . + . . . + ...... 2 1 2 1 4 + + + ...... 1 . Relevé number Relevé Pinus sylvestris (b) Vaccinium myrtillus Vaccinium Pinus sylvestris (c) Vaccinium vitis-idaea Vaccinium Dicranum polysetum Accompanying species Cl. Molinio-Arrhenatheretea submillefolium Achillea Orthilia secunda Agrostis gigantea Agrostis Ch. Cl. Vaccinio-Piceetea Dicranum scoparium Agrostis stolonifera Agrostis Diphasiastrum complanatum Agrostis vinealis Agrostis Diphasiastrum × zeilleri Anthoxanthum odoratum Anthoxanthum Dryopteris carthusiana Hylocomium splendens Luzula pilosa Lycopodium annotinum Lycopodium Melampyrum pratense Pleurozium shreberi Pleurozium Ptilium crista-castrensis Ptilium Pyrola rotundifolia Pyrola Trientalis europaea Trientalis [99] ...... 1 . + + + + + + + + + + + + + + + 23 ...... 1 1 1 1 . . . + + . + 22 ...... 5 . . . . . + 21 ...... 1 ...... 20 ...... 1 1 . . . . + . + + . . 19 ...... 1 . . . . + . + . + . 18 ...... 1 . . . + . . . + 17 ...... + . . + . . + 16 ...... 1 ...... + 15 ...... + 14 ...... + + . . . 13 ...... 1 ...... 12 ...... 1 . . . + . + . . . 11 ...... 2 ...... + . . + + . 10 ...... 9 1 ...... 8 . . . . . + . . + ...... 7 2 ...... + ...... 6 1 ...... + . . + ...... 5 ...... 4 . . . . + ...... 3 ...... 2 1 ...... + . + . . . + ...... 1 1 . . . + Relevé number Relevé Sieglingia decumbens Sieglingia Carex leporina Carex Viola montana Viola Dactylis glomerata Dactylis Cl. Koelerio-Corynephoretea canescentis Cl. Koelerio-Corynephoretea Chamaecytisus ruthenicus Deschampsia caespitosa Corynephorus canescens Dianthus borbasii Eremogone saxatilis Eremogone Festuca rubra Festuca Festuca ovina Festuca Galium boreale Hypochaeris maculata Galium mollugo Jasione montana Leontodon autumnalis Potentilla argentea Potentilla Leucanthemum vulgare Lysimachia vulgaris Lysimachia Plantago lanceolata Plantago Poa angustifolia Poa pratensis Prunella vulgaris Prunella Pteridium aquilinum Pteridium Ranunculus repens [100] ...... 1 1 1 + + + + + + . + + + 23 ...... 1 . + + + + + + + + + + 22 ...... + + + + 21 ...... 1 1 ...... 20 ...... + . + 19 ...... + . . . + 18 ...... + . 17 ...... + . + . . 16 ...... 15 ...... + 14 ...... 13 ...... + . + 12 ...... + . . . + 11 ...... + . + + . . + 10 ...... 9 ...... 8 ...... 7 1 ...... + . . + + ...... 6 ...... + ...... 5 4 ...... 4 3 ...... 3 ...... 2 . . . . + + . . . + . + ...... 1 1 1 ...... (a) (b) Relevé number Relevé Rumex acetosella Geranium sylvaticum Thymus serpyllum Hylotelephium polonicum Hylotelephium Cl. Nardetea strictae dioica Antennaria Melampyrum nemorosum Hypericum maculatum Steris viscaria Lycopodium clavatum Lycopodium Trifolium alpestre Trifolium Nardus stricta Nardus Cl. Carpino-Fagetea sylvaticae Acer platanoides Pilosella officinarum Pilosella Acer platanoides Pimpinella saxifraga Pimpinella Platanthera chlorantha Platanthera Potentilla erecta Potentilla Veronica officinalis Veronica Viola canina Viola Cl. Trifolio-Geranietea sanguinei Cl. Trifolio-Geranietea ramosum Anthericum Clinopodium vulgare Digitalis grandiflora Digitalis Galium verum [101] ...... 1 1 1 1 + + + + + + + + + + 23 ...... 2 1 1 1 . . . . . + + + 22 ...... 3 1 ...... + 21 ...... 2 . . . . . + 20 ...... 2 2 ...... + + 19 ...... 1 2 . . . . . + . + 18 ...... 1 2 1 . . . . . + . + 17 ...... 3 ...... + + 16 ...... 3 ...... + 15 ...... 5 ...... + 14 ...... 3 ...... 13 ...... 3 1 ...... 12 ...... 3 1 1 . . . . + + 11 ...... 2 1 . . . . + + + + 10 ...... 9 2 ...... + . + ...... 8 5 . . . . . + . . + ...... 7 5 1 ...... + ...... 6 5 4 ...... + ...... 5 . . . . . 40 ...... 4 5 ...... + ...... 3 ...... 10 ...... 2 3 ...... + + ...... 1 1 ...... 40 Relevé number Relevé Carex digitata Carex Cl. Papaveretea rhoeadis Cl. Papaveretea Conyza canadensis Carex pilosa Carex Galeopsis tetrahit Oxalis acetosella Scleranthus annus Poa nemoralis Cl. Quercetea pubescentis Cl. Quercetea Lathyrus niger Viola mirabilis Viola Potentilla alba Potentilla Viola riviniana Viola Pulmonaria angustifolia Cl. Calluno-Ulicetea Calluna vulgaris Cl. Quercetea robori-petraea Cl. Quercetea Carex ericetorum Carex Genista germanica Scorzonera humilis Vincetoxicum hirundinaria Vincetoxicum Cl. Epilobietea angustifolii Cl. Epilobietea epigeios Calamagrostis Fragaria vesca Rubus idaeus Salix caprea Salix Sambucus racemosa [102] ...... 1 4 3 + + + + + + 23 ...... 3 1 . . . . + + + + + 22 ...... + . . + 21 ...... 1 3 . . . . . + 20 ...... 1 . . . + + + + + 19 ...... + + + + + + + + 18 ...... 1 ...... + + + 17 ...... 4 . . . . + + + . + 16 ...... 1 2 . . + . + + + 15 ...... + + + 14 ...... 13 ...... + + 12 ...... 1 . . . + + + + + 11 ...... + + + + 10 ...... 9 3 ...... + + + + ...... 8 . . . . . + + + ...... 7 4 3 5 . . . . + + ...... 6 3 . . + . + + ...... 5 ...... + ...... 4 ...... 3 . . . . . + + + ...... 2 3 ...... + + ...... 1 3 3 . . Relevé number Relevé Carex pallescens Carex Malus sylvestris (a) Maianthemum bifolium Malus sylvestris (b) Rubus nessensis Malus sylvestris (b) Cl. Artemisietea vulgaris Geranium robertianum Populus tremula Moeringia trinerva Cl. Crataego-Prunetea Galeopsis bifida Cl. Festuco-Brometea Euphorbia cyparisias Oxycocco-Sphagnetea Ledum palustre Cl. Scheuchzerio palustris-Caricetea fuscae Cl. Scheuchzerio nigra Carex Other species Other capillaris Agrostis Betula pendula Betula Betula pendula (b) Betula Betula pendula (c) Betula Calamagrostis arundinacea Calamagrostis Campanula rotundifolia Hypericum perforatum [103] ...... 1 1 + 23 ...... + + 22 ...... 21 ...... 1 . . 20 . . . . 2 . . . . 19 Buzunko ...... 3 . . 18 ...... 2 . . Rak, P. 17 . . . . . + . + + 16 . . . 1 1 . . . + Danko, O. 15 . . . . . + . + + 14 Danko . , 5 – Querco-Piceetum ...... 13 ...... 12 . . . . . 2 . . . 11 . . . . . + + + + 10 ...... 9 . . + ...... 8 . . . Veronico incanae-Pinetum , 4 – Veronico Lukash . . . 7 . . + + + + . . 6 . . + . + + + . . . . 5 ...... 4 . . . . , 3 – Molinio-Pinetum . . . . 3 ...... 2 . . . + + . . 1 1 . . . + . + , 2 – Peucedano-Pinetum Relevé number Relevé – 18.05.2005, the Nova Zymnytsia tract, Ripky district, Chernihiv region; O. Lukash. – 15.06.1998, the forest near the Drobtsi village, Korop district, Chernihiv Karpenko region; Yu. – 10.07.2016, the pine-forest terrace of the Desna river, near the Sokolivka village, Kozelets district, Chernihiv region; O. Lukash. – 01.07.2003, the Novi Borovychi between forestry, the Turia and Zahrebelna Sloboda villages, Snovsk district, Chernihiv region; O. Lukash. – 17.08.2013, tract, the Liubech Yasenske Ripky forestry, district, Chernihiv region; O. Popruha, Lukash, О. V. Rak. – 15.08.1996, the Horbachi forestry (sq. 19), near the village, Lykholitky Kozelets district, Chernihiv region; O. Lukash. – 02.05.2005, the Kolochivska Dacha tract, Koriukivka district, Chernihiv region; O. Lukash. – 14.08.1996, the Horbachi forestry (sq. 113), Kozelets district, Chernihiv region; O. – 19.07.2019, the pine-forest terrace of the Desna river, near the Krasne village, Chernihiv district, Chernihiv region; H. – 25.08.2018, the Kupshi tract, between the Nevklia and Bezykiv villages, Horodnia district, Chernihiv region; O. Lukash – 25.04.2004, the Zhuklia forestry (sq. 35), Koriukivka district, Chernihiv region; O. Lukash. – 16.08.2019, Mizhrichynskyi Regional Landscape Park, the shooting range, Kozelets district, Chernihiv region; O. Lukash, H. Populus tremula (b) Populus tremula Populus tremula (c) Populus tremula Pyrus communis (a) Pyrus communis (b) Rubus caesius Salix rosmarinifolia Salix Sorbus aucuparia (a) Sorbus aucuparia (b) Veronica chamaedrys Veronica – 05.07.1980, the Chudivka forestry (sq. 30), Ripky district, Chernihiv region; T. Andrienko. T. region; Chernihiv 2 – 05.07.1980, the Chudivka forestry (sq. 30), Ripky district, 3 4 – 11.08.2019, the pine-forest terrace of the Dnieper river, near the Lisove village, Ripky district, Chernihiv region; H. Danko region; Chernihiv Ripky district, near the Lisove village, of the Dnieper river, terrace the pine-forest 5 – 11.08.2019, region, O. Lukash Chernihiv district, 6 – 13.08.2009, the Orlykivka forestry (sq. 101), Semenivka 7 Sumy region, N 52,33425°; EO 33,7469; S. Panchenko, O. Chornous (Panchenko, 2013). (sq. 45, site 8), Seredyna Buda district, 8 – 31.07.2003, the Starohutskyi forest area 9 10 11 region), O. Lukash. Chernihiv (Horodnia district, village Travneve 12 – 12..07.2002, the Myklashevshchyna tract, 13 region, O. Lukash. Chernihiv district, forestry (sq. 60), Semenivka Radomka 14 – 01.08.2010, the Radomska Dacha tract, O. Lukash. region; Chernihiv Horodnia district, villages, Tupychiv and the Nevklia between forestry, 15 – 30.09.2003, the Nevklia Sumy region, N 52,3317°; EO 33,7622; S. Panchenko (Panchenko, 2013). (sq. 65, site 7), Seredyna Buda district, 16 – 14.07.2002, the Starohutskyi forest area O. Lukash. region; Chernihiv district, Semenivka the Masheve and Ivanivka villages, between Spaske tract, 17 – 16.08.1999, the Masheve forestry, 18 19 Sumy region, N 52,3209°; EO 33,8121, S. Panchenko (Panchenko, 2013). (sq. 97), Seredyna Buda district, the Starohutskyi forest area 20 – 11.07.1997, region; O. Lukash. Chernihiv Horodnia district, 21 – 15.07.2010, the Hnizdyshchanska Dacha tract, 22 23 1 – Cladonio-Pinetum Plant communities: Notes. Dates, localities and authors of relevés: 1 104 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko

The ontogenetic structure of Calluna vulgaris in plant spectrum (without seedlings and senile individuals) and is communities of pine forests of the Cladonio-Pinetum as- right-sided, with the peak on virginal and generative indi- sociation wasThe investigated. ontogenetic Two structuremodel cenopopulations of Calluna of vulgarisviduals (Fig.in plant 2: I). communitiesThe density of the of “left-bank”pine forests popula of- the Calluna vulgaris (equal in area, ecological and phytocenot- tion is 120 individuals per m2. It is also incomplete (with- ic Cladonioconditions) -werePinetum selected: association in Right-bank was Polesie investigated. – in the out Two seedlings, model juvenile cenopopulations and senile individuals) of Calluna and vulgarisright- Rivne(equal Nature in area,Reserve ecological (Table 4, relevé and 1) phytocenotic and in Left-bank conditions) sided with werethe peak selected: on old generative in Right individuals-bank Polesie (Fig. 2: – in Polesie – in Mizhrichynskyi Regional Landscape Park II). The difference in indexes of the ontogenetic status as- (Tablethe Rivne5, relevé Nature 1). The resultsReserve of population (Table 4,studies relevé are 1)sessment and in of Left two -populationsbank Polesie indicates – in the Mizhrichynskyi following: the represented in Table 6 and Figure 2. growth conditions for Calluna vulgaris in the Rivne Na- RegionalTable 6 shows Landscape that the density Park of(Table the “right-bank” 5, relevé pop 1).- Theture resultsReserve ofare population more optimal studies and are arefollowed repre bysented more in ulation is 118 individuals per m2. It has incomplete age active growth of plants and settling on new territories. Table 6 and Figure 2.

Table 6. Calluna vulgaris (L.) Hill. populations in Cladonio-Pinetum Juraszek 1927 communities of Ukrainian Polesie Table 6. Calluna vulgaris (L.) Hill. populations in Cladonio-Pinetum Juraszek 1927 communities of

Ukrainian Polesie 2 Index of ontogenetic status Average number of individuals/m 2 Population localization Average number of individuals/m Indexassessment of ontogenetic status Population localization assessment p j im v g1 g2 g3 s Σ Iinn Isen Igen Iaet p j im v g1 g2 g3 s Σ Iinn Isen Igen Iaet Rivne Nature Reserve 0 3 5 30 27 28 25 0 118 32.20 21.19 67.80 0.65 Rivne(Right-bank Nature Polesie) Reserve 0 3 5 30 27 28 25 0 118 32.20 21.19 67.80 0.65 (RightMizhrichynskyi-bank Polesie) Regional MizhrichynskyiLandscape Park (Left-bank Regional 0 0 0 0 2 2 29 29 25 25 24 24 40 40 0 0 120 12025.83 25.8333.33 33.3340.83 40.831.29 1.29 LandscapePolesie) Park (Left- bank Polesie)

% 35

0 I II j im v g1 g2 g3

FigureFigure 2. Age-related 2. Age -structurerelated of s thetructure Calluna ofvulgaris the populationsCalluna invulgaris Rivne Nature population Reserve (I)s and in MizhrichynskyiRivne Nature Regional Reserve Land- (I)

scape Parkand (II). Mizhrichynskyi Denotations: j – juvenile Regional individuals, imLandscape – immature individuals,Park (II) v –. virginile Denotations: individuals; g1j – young– juvenile generative individuals, g – mature generative individuals, g – old generative individuals individuals,2 im – immature individuals,3 v – virginile individuals; g1 – young generative individuals, g2 – mature generative individuals, g3 – old generative individuals

Table 6 shows that the density of the “right-bank” population is 118 individuals per m2. It has incomplete age spectrum (without seedlings and senile individuals) and is right-sided, with the peak on virginal and generative individuals (Fig. 2: I). The density of the “left-bank”

30 population is 120 individuals per m2. It is also incomplete (without seedlings, juvenile and senile individuals) and right-sided with the peak on old generative individuals (Fig. 2: II). The Phytocenotic features of Calluna vulgaris (L.) Hill. in Ukrainian Polesie 105 difference in indexes of the ontogenetic status assessment of two populations indicates the following: the growth conditions for Calluna vulgaris in the Rivne Nature Reserve are more

The “western” population is younger (Iaet = 0.65) than the According to the results of the research, the map of the optimal“eastern” one,and whereare followed the aging byprocesses more areactive prevailing growth Calluna of plants vulgaris and distribution settling in on Ukrainian new territories Polesie biotopes. The

“western(Iaet = 1.29).” Topopulation compare with, is younger the aging (Iofaet the = Calluna0.65) than was the created “eastern (Fig. 3).” one, where the aging processes vulgaris populations in the forest phytocenoses in the east We compared the frequency of the Calluna vulgaris areof Left-bank prevailing Polesie (Iaet (National = 1.29). Nature To compare Park “Desniansko with, the- сonstancyaging of inthe plant Calluna communities vulgaris of Right-bank populations and Left-in the Starohutskyi”), is more intensive: Index aetas (I ) is 1.72 bank Polesie. The results of the comparison are shown forest phytocenoses in the east of Leftaet -bank Polesie (National Nature Park “Desniansko- (Kovalenko, 2015). in Table 7. Starohutskyi”It should be noted), is thatmore the syntaxonomicalintensive: Index affiliation aetas of (I aet) isUndoubtedly, 1.72 (Kovalenko, Calluna 2015).vulgaris has the maximum the CallunaIt vulgarisshould forest be notedhabitats inthat Right-bank the syntaxonomical and Left- сonstancy affiliation frequency of in thethe phytocenoses Calluna vulgaris of the Calluno- forest bank Polesie is different. In particular, rare, as for Polesie, Ulicetea and the Nardetea strictae classes, where it is habitatscommunities in ofRight the Veronico-bank incanae-Pinetumand Left-bank association Polesie isa characteristicdifferent. orIn differentiating particular, (diagnostic)rare, as forspecies. Polesie, The communities(Table 5, relevés of 17-19) the Veronicooccur only inincanae Left-bank-Pinetum Polesie. associationCalluna vulgaris (Table high 5, constancy relevés is17 observed-19) occur in the only plant in Pine forests of the Vaccinio uliginosi-Pinetum associa- communities of the Ledo-Sphagnetum magellanici asso- Lefttion with-bank Calluna Polesie. vulgaris Pine (Table forests 4, relevés of 23-26)the Vaccinio were ciation. uliginosi The Calluna-Pinetum vulgaris association occurrence within pine Calluna forests described only in Right-bank Polesie. The communities of Right-bank Polesie is slightly higher (than in Left-bank vulgarisof the Vaccinio (Table uliginosi-Pinetum 4, relevés 23 association-26) were are described known Polesie).only in TheRight spruce-bank forests Polesie. of the TheSphagno communities girgensohnii- of theonly inVaccinio National Natureuliginosi Park -“Desniansko-Starogutskyi”Pinetum association arePiceetum known association only are rarein biotopesNational for CallunaNature vulgar Park- in Left-bank Polesie (Panchenko, 2013). Calluna vulgaris is. It should be noted that Calluna vulgaris does not grow “Desnianskoas well as Vaccinium-Starogutskyi” uliginosum L. in (diagnostic Left-bank species Polesie of in(Panchenko other spruce ,forests, 2013) which. Cal haveluna an vulgaris “island” distribution as well as Vacciniumthe Piceo-Vaccinienion uliginosum uliginosi L. (diagnostic suballiance) speciesare absent of thein Polesie.Piceo- VaccinienionRather stable position uliginosi and high suballiance) constancy ofare in all 10 relevés of this association. The European boreo- Calluna vulgaris in different phytocenoses of Ukrainian absentmontane inspruce all forests10 relevés of the Piceion of this excelsae association. class (Table The PolesieEuropean is a consequence boreo-montane of high sprucedrought andforests frost ofresis the- Piceion4, relevés 27-29)excelsae occur class only in(Table Right-bank 4, Polesie.relevé sThe 27 -tance29) ofoccur this species only inin combination Right-bank with plasticityPolesie. of Theits phytocenoses are represented by the Sphagno girgensoh- morphological structure (Grubb, 1992). pnii-Piceetumhytocenoses association. are represented The projective by coveragethe Sphagno of Cal -girgensohnii-Piceetum association. The projective coverageluna vulgaris of does Calluna not exceed vulgaris 1%. does not exceed 1%. According to the results of the research, the map of the Calluna vulgaris distribution in Ukrainian Polesie biotopes was created (Fig. 3).

Figure 3. Distribution of Calluna vulgaris (L.) Hill. in the biotopes of Ukrainian Polesie. Map symbols: 1 – border of Ukrainian Figure Polesie,3. Distribution 2 – temperate of and Calluna subboreal vulgarispine forests (L.)on nutrient-poor Hill. in theacidic biotopes sandy soils of(the Ukrainian Vaccinio-Piceetea Polesie. class), Map3 – boreo-temperatesymbols: heathlands1 – border of the of Calluno-Ulicetea Ukrainian Polesie,class, 4 – boreo-temperate2 – temperate heathlands and subboreal of the Nardetea pine strictae forests class, on 5 – dwarf-shrub,nutrient-poor sedge andacidic peat-moss sandy ombrotrophic soils (the raised Vaccinio bogs on extremely-Piceetea acidic soilsclass), (the Oxycocco-Sphagnetea3 – boreo-temperate class) heathlands of the Calluno-Uliceteaclass, 4 – boreo-temperate heathlands of the 31

106 Oleksandr Lukash, Viktor Melnyk, Hanna Danko, Oleksandr Rak, Yurii Karpenko, Petro Buzunko

Table 7. Сonstancy of Calluna vulgaris (L.) Hill. in plant communities of Ukrainian Polesie

Right-bank Polesie Lef t-bank Polesie Number Number Total Total of relevés of relevés Association number Constancy, number Constancy, with with of relevés % of relevés % Calluna Calluna analyzed analyzed vulgaris vulgaris Calluno-Genistetum pilosae 3 3 100 0 0 - Scabioso canescentis-Genistetum 0 0 - 5 5 100 Calluno-Nardetum strictae 17 17 100 4 4 100 Ledo-Sphagnetum magellanici 41 38 92.7 0 0 - Cladonio-Pinetum 67 43 64.2 63 36 57.1 Peucedano-Pinetum 82 63 76.8 93 68 73.1 Veronico incanae-Pinetum 0 0 - 43 38 65.1 Molinio-Pinetum 57 41 71.9 36 25 69.4 Querco-Piceetum 45 18 40.0 48 19 39.6 Vaccinio uliginosi-Pinetum 36 17 47.2 10 0 0 Sphagno girgensohnii-Piceetum 43 7 16.3 0 0 -

5. Conclusions Veronico incanae-Pinetum association is the habitat of Cal- luna vulgaris only in Left-bank Polessie. Calluna vulgaris The heathlands are rare ecosystems in Ukrainian Polesie. has the lowest constancy (57.1%) in the Cladonio-Pinetum Their vegetation is represented by the plant communities association of Left-bank Polesie and the highest constancy of the Calluno-Ulicetea and the Nardetea strictae classes. (76.8%) in the Peucedano-Pinetum association of Right- They are forming in greatly anthropogenically disturbed bank Polesie. Calluna vulgaris constancy does not exceed areas (e.g., areas after wildfires) or in the areas with in- 40% in the oak-pine forests of the Querco-Piceetum associ- tensive human activities (at shooting ranges first of all). ation. The syntaxonomic difference of the Calluna vulgaris The cenoses of the Calluno-Genistetum pilosae associa- forest habitats in Right-bank Polesie and Left-bank Polesie tion, which are present only in Right-bank Polesie, have is primarily revealed at the Piceo-Vaccinienion uliginosi larger areas (up to 20 km2). Other heather communities suballiance level and the Piceetalia excelsae order. The are smaller in size; they usually occupy the area of sev- spruce forests of the Sphagno girgensohnii-Piceetum as- eral hectares (the Calluno-Nardetum strictae association) sociation are rare biotopes for Calluna vulgaris. or are represented by small patches in landscape of pine The Calluna vulgaris populations within the forest forests (less frequently in mixed forests) on upland ter- plant communities of the Cladonio-Pinetum association are races of rivers (the Scabioso canescentis-Genistetum as- characterized by a similar ontogenetic structure: they are sociation). incomplete. There are no sprouts, vegetative reproduction A significant participation (with the projective cover prevails, and age spectrum maximum falls on generative of up to 80%) and high constancy (92.7%) in oligotrophic individuals. However, the Calluna vulgaris cenotic popula- swamps the phytocenoses of the Ledo-Sphagnetum magel- tion in Right-bank Polesie of Ukraine is characterized by lanici association is a characteristic feature of the Calluna a greater activity of plants growth and spreading on new vulgaris distribution in Right-bank Polesie of Ukraine. territories, while aging prevails in the “left bank” popula- The middle-aged pine and birch-pine forests of the Di- tion. crano-Pinenion suballiance are the most common habitats In general, the environmental conditions of Right-bank of Calluna vulgaris in Ukrainian Polesie (mostly, with its Polesie of Ukraine are more favorable to Calluna vulgaris, projective cover of less than 1%). The phytocenoses of the than the conditions in Left-bank Polesie. Phytocenotic features of Calluna vulgaris (L.) Hill. in Ukrainian Polesie 107

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