OHaCqiietia 18/2 . 2019,289-312 DOL 10.2478/hacq-2018-0013 Vsciendo Kenophytes in different forest types of Ukraine Viktor Onyshchenko1 ® Key words: forest, vegetation, alien plants, kenophytes, invasion, habitats. Ključne besede: gozd, vegetacija, tujerodne rastline, kenofiti, invazija, habitati. Received: 13. 4. 2018 Revision received: 4. 12. 2018 Accepted: 5. 1. 2019 Abstract The article aims to 1) describe the distribution of non-native species of vascular plants (only kenophytes, i. e. naturalized species introduced after the 15 th century) in different types of forests and different biogeographical regions of Ukraine; 2) compare proportions of kenophytes in forests of different areas; 3) detect statistically significant changes in the occurrence of kenophytes over the last 80 years. The material consists of 2701 relevés sampled in 1990-2018. They were taken from Ukrainian phytocoenological publications and databases. In Ukraine, as in other European countries, the highest proportion of kenophytes (percentage of species number per relevé) is in floodplain forests (up to 9.1% in willow and poplar forests). The lowest proportion is characteristic for bog woods (0%) and most types of mountain forests. Among biogeographical regions, the highest values were found in the flatland areas of the Steppic region and the forest-steppe part of the Continental region. The most frequent species are Impatiensparviflora (predominantly in broadleaved woods, absent in relevés from the steppic biogeoregion) and Erigeron canadensis (pine woods on sand). Comparison with 1466 older relevés sampled in 1950—1989 allows us to make a conclusion that the proportion of kenophytes increased at least in one habitat type (oak-hornbeam forests). Izvleček V članku smo želeli 1) opisati razširjenost tujerodnih rastlinskih vrst (samo keno-fitov, to je naturaliziranih vrst, ki so se pojavile po 15. stoletju) v različnih gozdnih habitatnih tipih in v različnih biogeografskih regijah v Ukrajini; 2) primerjati deleže kenofitov v gozdovih v različnih območjih; 3) prikazati statistično značilne spremembe v pojavljanju kenofitov v zadnjih 80. letih. Podatke predstavlja 2701 popis, ki so bili narejeni v obdobju med 1990 in 2018. Uporabili smo fitoceno-loško literaturo in podatkovne baze iz Ukrajine. V Ukrajini, kot tudi v ostalih državah v Evropi, je najvišji delež kenofitov (odstotek vrst na popis) v poplavnih gozdovih (do 9,1% v vrbovih in topolovih gozdovih). Najnižji odstotek je značilen za barjanske gozdove (0%) in večino habitatnih tipov gorskih gozdov. Med bioge-ografskimi regijami je najvišji delež v nižinskih predelih stepske regije in gozdne stepe v kontinentalni regiji. Najbolj pogosti vrsti sta Impatiens parviflora (prevladuje v listopadnih gozdovih, odsotna je v popisih iz stepske biogeografske regije) in Erigeron canadensis (v borovih gozdovih na pesku). S primerjavo 1466 starejših popisov, narejenih med leti 1950 in 1989 smo prišli do zaključka, da se je delež kenofitov povečal vsaj v enem habitatnem tipu (hrastovo-gabrovi gozdovi). 1 M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Ukraine. E-mail: v.a.onyshchenko@gmail.com 107 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Introduction Phytosociological relevés are an important source of information about ecological preferences of species, their geographical distribution and dynamics. In recent decades some studies were published that show geographical and ecological patterns of alien species on the basis of phyto-coenological data. The study areas were the Czech Republic (Chytry at al. 2005), Catalonia (Vila et al. 2007), the Basque Country (Campos et al. 2013), Slovakia (Med-vecka et al. 2014) and Slovenia (Kuzmic & Silc 2017). Chytry et al. (2008) performed a comparison of plant invasions for three areas of Europe (the Czech Republic, Catalonia, Great Britain) that represent three biogeo-graphical regions (Continental, Mediterranean, Atlantic). Estimations in the aforementioned articles were made for classes and alliances of vegetation or for EUNIS level 2 habitats (Davies et al 2004, EUNIS habitat ...). According to these studies the lowest proportions of aliens in all regions are in bogs, mires, alpine and subalpine grasslands, heaths on poor soils. The highest proportions are found in synanthropic vegetation, marine coastal habitats, standing inland waters (only kenophytes), sedge and reed beds, riverine shrubs. Wagner et al. (2017) published the assessment of plant invasions in European forests that is based on the analysis of 83,396 relevés from the European Vegetation Archive (Chytry at al. 2016) sampled in 1970-2015. The assessment was made for habitat types of the revised version of EUNIS habitat classification (Schaminée et al. 2014). These habitat types approximately correspond to level 3 or 4 habitats of the official EUNIS habitat classification. All analyses were performed only for kenophytes (naturalized species introduced voluntarily or involuntarily after the 15th century). Archaeophytes (introduced before the end of the 15th century) were excluded from consideration because the status of archaeophytes is poorly known in some countries. The highest proportion of kenophytes was found in riparian woods. The lowest proportions were in mountain forests and mire woodlands. The most frequent kenophytes of European woods turned out to be Impatiensparviflora (21.1% of total frequency of kenophytes), Padus serotina (8.8%), Robinia pseudoacacia (7.5%). The highest number of occurrences of kenophytes belongs to phanerophytes and therophytes (for native species hemicryptophytes and phanerophytes). Species originating in Europe constitute 47% of the total number of kenophytes in European woods, and their total frequency is 22% of total frequency of kenophytes. The distribution of relevés used in the article by Wagner et al. (2017) has a strong geographic bias: Ukraine, Belarus, Moldova, Georgia, Armenia, Azerbaijan and Eu- 290 ropean Russia occupy 49% of the study area but they are represented only by 1% of the total number of relevés. We made the assessment of occurrences of kenophytes in forests of Ukraine using a higher number of relevés. A more narrow geographical scope allows using the results for comparisons between regions of Europe. The analysis for the territory of Ukraine, even without comparison with other countries, may reveal some geographical patterns because Ukraine comprises parts of four biogeo-graphical regions and measures about 1300 km in length. At present there is no information in literature about the level of invasion in forests of the Steppic biogeographical region of Europe. In other bioregions of Europe, estimations were carried out only at higher levels of habitat classifications. Presence of 40 alien invasive species within plant communities of Ukraine at the level of alliances of vegetation without quantitative estimations was described in article by Abduloyeva & Karpenko (2009). The aim of the present study is to describe the frequency of kenophytes in forest types and different biogeographi-cal regions of Ukraine, and to compare the results with published data on the subject from other areas. Materials and methods The study area includes the entire territory of Ukraine. According to the data on the European Environment Agency website (Europe's biodiversity ... 2018) Ukraine comprises parts of four European biogeographical regions: Alpine, Continental, Pannonian and Steppic. Two regions (Continental and Steppic) were subdivided according to the geobotanical subdivision of Ukraine in the National Atlas of Ukraine (2008) (Figure 1). The relevés for the analysis were taken from the database EU-UA-006 "Vegetation Database of Ukraine and Figure 1: Biogeographical regions of Ukraine. Slika 1: Biogeografske regije Ukrajine. O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Adjacent Parts of Russia", unpublished database of relevés from the National Nature Park "Holosiivskyi"(Kyiv city) and numerous Ukrainian publications (Andrienko et al. 2004, Bairak 1996, Budzhak & Onyshchenko 2004, Chorney et al. 2005, Derzhypilsky et al. 2011, Didukh 1996, 2003, Didukh et al. 2003, Fitsailo 2003, Gomlya 2005, Goncharenko 2001, 2003, Klimuk et al. 2006, Konishchuk 2003, Kuzemko 2001, Kuzemko & Chorna 2002, Lukash & Onyshchenko 2006, Melnyk & Korinko 2005, Onyshchenko 1998, 2002, 2007, 2009, 2013a, 2013b, Onyshchenko & Budzhak 2003, Onysh-chenko &t Lubinska 2006, Onyshchenko & Lukash 2004, Onyshchenko & Sidenko 2002, Onyshchenko & Yuglichek 2010, Onyshchenko et al. 2007, 2009, 2015, 2016, Orlov & Yakushenko 2005, Panchenko 2013, Panchenko & Onyshchenko 2003, Ralo & Onyshchen-ko 2008, Shelyag-Sosonko et al. 2003, Shevchyk et al. 1996a, 1996b, 1997, I. Solomakha et al. 2006, 2015, V. Solomakha et al. 2004, 2016, Soroka 2008, Vorobyov et al. 2002, Yakushenko 2003, Yuglichek & Onyshchen-ko 2003). All relevés selected for estimation of modern occurrence of kenophytes were sampled in 1990—2018. Each relevé was assigned to a habitat of the revised version of the EUNIS habitat classification (European Red List of Habitats, Schaminée et al. 2014) (Table 1). Some relevés were excluded from the analysis: 1) relevés representing highly artificial woods (plantations of species out of their natural range or in clearly unnatural conditions and natural woods of non-native species), 2) relevés that have no species data obtained in period May 15 — August 31, 3) relevés that could not be unequivocally assigned to either habitat type. The area of the relevés varies from 100 to 2500 m2, predominantly 400-900 m2. This parameter was not used in relevé filtering. Some analyses showed that the effect of plot size on the proportion of species is negligible (Chytry et al. 2005, Vilà et al. 2007). The final number of relevés was 2701. In order to compare the current frequency of kenophytes with their frequency in previous decades 1466 relevés sampled in 1950-1989 were used (Bukhalo 1962, Grygora et al. 2005, Vorobyov et al. 1997, Vorobyov et al. 2008). Old relevés were not sampled in the same areas as the relevés from 1990-2018. This reduces the possibility of comparison of two sets of data. The names of species follow the database Euro+Med (2006-2018). Residence statuses of species (whether a species is a kenophyte or not) were taken from Ukrainian publications (Protopopova 1991, Mosyakin & Yavorska 2003, Bagrikova 2013, Solomakha et al. 2015). Some species are kenophytes only in a part of their distribution area in Ukraine (Acer pseudoplatanus, Arrhenatherum elatuis, Berberis vulgaris, Corynephorus canescens, Fagus sylvatica, Ligustrum vulgare, Picea abies, Spergula moriso-nii, Vinca minor). They were involved into the analysis Table 1: Codes, names of habitats and number of relevés. Tabela 1: Šifra, ime habitanega tipa in število popisov. Habitat Habitat name Number of relevés Number of code before 1990 relevés since 1990 G1.1 Temperate and boreal softwood riparian woodland - 84 G1.2a Alnus woodland on riparian and upland soils - 93 G1.2b Temperate and boreal hardwood riparian woodland - 95 G1.4 Broadleaved swamp woodland on non-acid peat 215 85 G1.5 Broadleaved bog woodland on acid peat 316 - G1.6a Fagus woodland on non-acid soils - 459 G1.6b Fagus woodland on acid soils - 37 G1.7a Temperate and submediterranean thermophilous deciduous woodland 66 151 G1.8 Acidophilous Quercus woodland - 26 G1.Aa Carpinus and Quercus mesic deciduous woodland 630 911 G1.Ab Ravine woodland - 90 G3.1a Temperate mountain Picea woodland - 121 G3.1b Temperate mountain Abies woodland - 44 G3.2 Temperate subalpine Larix, Pinus cembra and Pinus uncinata woodland - 5 G3.4a Temperate and continental Pinus sylvestris woodland 120 388 G3.4b Temperate and submediterranean montane Pinus sylvestris-Pinus nigra woodland 80 17 G3.A Picea taiga woodland 27 17 G3.Da Pinus mire woodland 44 78 All habitats 1498 2701 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine only in the cases when they met outside their old distribution area. For each of the species its constancy in every habitat type, constancy in total set of relevés, constancy in Ukrainian forests (mean for habitats types with weighting proportionally to habitat area) and proportion of species occurrences in the sum of occurrences of all kenophytes were calculated. Results Selected relevés were referred to 17 habitat types of the modified EUNIS classification (Table 1). Mean proportion of kenophytes for all relevés is 1.13%, mean weighted proportionally to habitat areas is 1.02%. A total of 62 kenophytes are present in the dataset. Their distribution along habitat types is shown in Table 2. The most frequent kenophytes in Ukrainian woods are Impatiens parviflora (20.6% of the sum of kenophytes constancies), Erigeron canadensis (14.8%), Prunus serotina (9.4%), QQuercus rubra (7.9%), Acer negundo (7.7%), Erigeron annuus (3.8%), Robinia pseudoacacia (3.4%). Impatiens parviflora is the most frequent kenophyte in the Continental, the Alpine and the Pannonian biogeographical regions. It is not present in forest relevés from the Steppic biogeographical region. Erigeron annuus is present only in relevés from the Continental and the Alpine biogeo-graphical regions. The largest average proportion of kenophytes in relevés (9.1%) is peculiar to G1.1 "Floodplain Salix- and Populus-dominated woods" (Figure 2). Other habitats with high proportions of kenophytes are G1.2a "Alnus woodland on riparian and upland soils" (2.3%) and G1.2b "Temperate and boreal hardwood riparian woodland" (3.1%). Kenophytes are not present in relevés of coniferous bog woodlands (G3.Da), acidophilous beech forests (G1.6b), mountain pine forests (G3.2, G3.4b). A very low proportion is in G3.1a "Temperate mountain Picea woodland" (0.03%). Other habitats have intermediate values of this Table 2: Constancy of kenophyte species in habitat types of Ukraine (habitats types without kenophytes were excluded). Tabela 2: Stalnost kenofitov v habitatnih tipih v Ukrajini (habitatni tipi brez prisotnih kenofitov niso prikazani). G1.1 G1.2a G1.2b G1.4 G1.6a G1.7a G1.Aa G1.Ab G3.1a G3.1b a .4 3. G Constancy in dataset (%) '% of total kenophytes' occurrences Constancy with weighting of plots proportionally to habitat areas (%) Number of relevés 84 93 95 85 459 151 911 90 121 44 388 Number of kenophytes species Acer negundo 39.3 2.2 16.8 11.8 . 4.0 2.4 . . 1.0 3.41 11.7 2.35 Acer pseudoplatanus* 1.1 3.2 1.2 . 0.4 . 0.33 1.1 0.27 Acer saccharinum . 0.3 0.04 0.1 0.09 Aesculus hippocastanum 1.1 0.2 . 0.11 0.4 0.09 Ambrosia artemisifolia . 2.1 0.30 1.0 0.73 Amelanchier ovalis 3.2 . 0.3 0.15 0.5 0.21 Amelanchier spicata 1.1 6.0 . 1.3 0.56 1.9 0.65 Amorpha fruticosa 35.7 1.2 . . 0.5 1.22 4.2 0.65 Armeniaca vulgaris 1.2 6.5 0.26 0.9 0.27 Arrhenatherum elatius* 2.0 0.11 0.4 0.05 Berberis vulgaris* 2.0 . 1.5 0.33 1.1 0.60 Bidens frondosus 16.7 6.5 1.1 7.1 . 1.00 3.4 0.55 Bryonia alba 1.2 0.04 0.1 0.02 Cannabis sativa 1.1 0.7 0.07 0.3 0.06 Caragana arborescens 1.1 2.1 1.3 0.7 . . . 0.3 0.45 1.5 0.40 Corynephorus canescens* . 1.5 0.22 0.8 0.54 Cuscuta campestris . 0.3 0.04 0.1 0.09 Digitaria aegyptiaca . 0.3 0.04 0.1 0.09 Echinocystis lobata 3.6 4.7 . 0.26 0.9 0.07 Eragrostis minor . 1.3 0.19 0.6 0.45 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine G1.1 G1.2a G1.2b G1.4 G1.6a G1.7a G1.Aa G1.Ab G3.1a G3.1b a .4 3. G Constancy in dataset (%) '% of total kenophytes' occurrences Constancy with weighting of plots proportionally to habitat areas (%) Number of relevés 84 93 95 85 459 151 911 90 121 44 388 Erigeron annuus 7.1 2.1 2.4 0.4 2.6 1.6 2.2 . . 0.8 1.33 4.6 0.96 Erigeron canadensis 13.1 4.3 2.1 5.3 1.2 . . 9.0 2.26 7.7 3.74 Fagus sylvatica* 0.2 0.07 0.3 0.05 Fraxinus americana 1.2 0.04 0.1 0.02 Gleditsia triacanthos 0.7 0.2 0.11 0.4 0.07 Hemerocallis lilioasphodelus 1.2 0.04 0.1 0.02 Hordeum murinum s. l. 1.1 0.04 0.1 0.04 Hyoscyamus niger 1.1 6.0 0.52 1.8 0.25 Impatiens glandulifera 1.1 0.04 0.1 0.04 Impatiens parviflora 3.6 21.5 46.3 15.3 5.9 18.5 8.2 16.7 0.8 4.5 0.8 8.55 29.3 6.11 Iva xanthiifolia 1.1 0.04 0.1 0.04 Ligustrum vulgare* 0.7 0.04 0.1 0.02 Lonicera tatarica 2.6 0.1 0.19 0.6 0.09 Morus alba 13.1 0.7 0.2 0.52 1.8 0.24 Morus nigra 2.4 0.07 0.3 0.03 Oenothera biennis 8.3 . 1.3 0.44 1.5 0.57 Oenothera rubricaulis 0.7 . 0.3 0.07 0.3 0.11 Oxalis dillenii 2.4 0.07 0.3 0.03 Oxalis stricta 0.1 0.04 0.1 0.03 Partenocissus quinquefolia 1.2 0.1 0.07 0.3 0.04 Parthenocissus inserta 2.1 . 0.3 0.11 0.4 0.17 Physocarpus opulifolius . 0.5 0.07 0.3 0.18 Picea abies* 0.7 0.22 0.8 0.15 Prunus cerasifera 2.0 0.11 0.4 0.05 Prunus serotina 1.1 2.1 4.6 0.2 . 5.4 1.22 4.2 2.20 Quercus rubra 2.2 2.1 1.3 2.3 1.2 . . 0.8 1.15 3.9 1.01 Robinia pseudoacacia 1.2 3.2 2.1 0.2 2.0 1.4 1.2 . 2.3 2.1 1.22 4.2 1.39 Saponaria officinalis 2.6 0.15 0.5 0.07 Secale sylvestre 1.2 0.04 0.1 0.02 Senecio viscosus . 0.5 0.07 0.3 0.18 Sorbus intermedia 0.7 0.04 0.1 0.02 Spergula morisonii* . 0.5 0.07 0.3 0.18 Spiraea chamaedryfolia 1.2 . 0.04 0.1 0.00 Swida alba 1.2 1.2 0.2 0.11 0.4 0.04 Symphoricarpos albus . 0.5 0.07 0.3 0.18 Triticum aestivum 1.1 0.04 0.1 0.04 Ulmus pumila 1.2 0.04 0.1 0.02 Veronica hederifolia 2.6 0.3 0.26 0.9 0.15 Vinca minor* 1.1 1.1 0.4 . 0.3 0.26 0.9 0.28 Vitis vinifera 1.2 0.04 0.1 0.02 Xanthium orientale subsp. 1.2 0.04 0.1 0.02 riparium Xanthium rupicola 2.4 0.07 0.3 0.03 Xanthium strumarium 2.4 0.07 0.3 0.03 * — species is native in a part of the Ukrainian territory. 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Figure 2: Mean proportions of kenophytes in relevés of different habitats in Ukraine (with standard deviation) and Europe. Data on Europe are from the article by Wagner et al. (2017). Slika 2: Povprečni deleži kenofitov v popisih v različnih habitatnih tipih v Ukrajini (s standardnim odklonom) in v Evropi. Podatki o pojavljanju v Evropi so iz članka Wagner et al. (2017). Figure 3: Mean total covers of kenophytes in relevés of different habitats in Ukraine (relevés without kenophytes were included in the calculation with cover of kenophytes = 0). Slika 3: Povrečna skupna pokrovnost kenofitov v popisih v različnih habitatnih tipih v Ukrajini (popisi brez kenofitov so bili vključeni v izračun s pokrovnostjo kenofitov=0). index. Proportions of kenophytes in habitats in different areas are presented in Figures 4—6. Major kenophytes of habitat G 1.1 are Acer negundo (constancy 39%), Amorpha fruticosa (36%), Bidens fron-dosus (17%). These species often have high cover (Figure 3). Mean cover of Acer negundo in relevés of G 1.1 with the presence of the species is 8%, Acer negundo — 7%, Bidens frondosus — 3%. These species are frequent also in other types of hygrophilous broadleaved forests (G1.2a, G 1.2b), however the most frequent kenophyte in these habitats is Impatiens parviflora (constancy 22% and 46%, mean cover 20% and 18% respectively). The typical habitats of G1.4 "Broadleaved swamp woodland on non-acid peat" are probably poor in kenophytes. The available relevés represent mainly forests that are transitional to G1.2a. The most frequent kenophyte of mesic eutrophic broadleaved forests (G1.6a, G1.Aa, G1.Ab) is Impatiens parviflora (constancy in these habitats 5%, 8%, 17%, mean cover 0.1%, 7%, 1% respectively). In habitat G1.7a "Temperate and submediterranean thermophilous deciduous woodland", Impatiens parviflora also holds the first position (constancy 19%, mean cover 4%). In addition, this habitat has significant constancies of Acer negundo, Amelannchier spicata, Erigeron canadensis, Prunus serotina. The most frequent kenophytes of habitat G3.4a "Temperate and continental Pinus sylvestris woodland" are Erigeron canadensis (9%) and Prunus serotina (5%, only in relevés from Kyiv). The relevés from mountain regions have the smallest proportion of kenophytes (Table 3). In the flatland part of Ukraine, higher proportions are characteristic of the Steppic biogeographical region and the forest-steppe part of the Continental biogeographical region. Table 3: Mean proportion of kenophytes in plots from different biogeographical regions of Ukraine. Tabela 3: Povprečni delež kenofitov na popisnih ploskvah v različnih biogeografskih regijah Ukrajine. Biogeographical region Mean % of kenophytes in relevé Number of relevés Continental: deciduous forest 0.53 1276 Continental: forest-steppe 2.46 657 Pannonian 1.02 26 Alpine 0.31 530 Steppic: flatland 3.13 180 Steppic: Crimean Mountains 0.00 32 All biogeoregions 1.13 2701 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Figure 4: Proportions of kenophytes in geographical groups of releves (total and in habitats G1.1, G1.2a, G1.2b, G1.4. G1.6a, G1.6b, G1.7a). Slika 4: Delež kenofitov v geografskih skupinah popisov (skupno in v habitatnih tipih G1.1, G1.2a, G1.2b, G1.4. G1.6a, G1.6b, G1.7a). 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Figure 5: Proportions of kenophytes in geographical groups of relevés (habitats G1.8, Gl.Aa, Gl.Ab. G3.1a, G3.1b, G3.2, G3.4a, G3.4b). Slika 5: Delež kenofitov v geografskih skupinah popisov (habitatni tipi G1.8, Gl.Aa, Gl.Ab. G3.1a, G3.1b, G3.2, G3.4a, G3.4b). 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Figure 6: Proportions of kenophytes in geographical groups of relevés (habitats G3A, G3.Da) Slika 6: Delež kenofitov v geografskih skupinah popisov (habitatni tipi G3A, G3.Da) Discussion In general, the distribution of proportions of kenophytes by different forest types of Ukraine is similar to that of the same forest types of entire Europe presented in the article by Wagner et al. (2017). Habitats with the highest number of relevés and largest occupied areas (G1.6a, G1.A1, G3.4a) have close values in Ukraine and Europe. Most kenophyte-rich alluvial, moist and swampy eutrophic forests (G1.1, G1.2a, G1.2b, G1.4) in Ukraine have almost twice as high proportions of kenophytes than forests of the same types from Europe. The main discrepancies are in thermophilous deciduous forests G1.7a, acidophilous beech forests G1.6b and acidophilous oak forests G1.8. A lower frequency of kenophytes in Ukrainian forests of G1.6b can be explained by the fact that this habitat is rare in flatland areas of Ukraine and that all relevés were sampled in the mountains. The proportion of kenophytes in habitat G1.7a from Ukraine is higher than in the same habitat in Europe because in Ukraine this habitat is presented only by relevés from the flatland part of Ukraine with a high proportion of relevés from urbanized areas. The mean value for all relevés is somewhat higher for Ukraine (1.1%) than for Europe (0.9%). One of the reasons for this difference may be that Ukrainian relevés were sampled more recently. Some data from literature show that the proportion of non-native species in forests of southern Europe is lower than in forests of more northern areas of the Atlantic and the Continental biogeographical regions. According to the results of Chytry et al. 2008 the proportion of neophytes in deciduous and mixed deciduous-coniferous forests of Catalonia (the Mediterranean biogeoregion) is much lower (0.2%) than in the same group of forests in the Czech Republic (the Continental biogeoregion, 1.0%) and Great Britain (the Atlantic biogeoregion, 3.1%). For coniferous forests, these values are 0.0%, 0.4% and 24.8% respectively. Low proportions of neophytes were found also in Slovenian forests (0.5% in deciduous and 0.1% in coniferous forests) (Kuzmic & Silc 2017). In Ukraine, the lowest proportion (0.0%) was found in the Crimean Mountains that are the southernmost part of the country. This area formally belongs to the Steppic biogeoregion, but its vegetation has more in common with that of the northern parts of the Mediterranean biogeoregion. According to the floristic data (Bagrikova 2013, Yena 2012), in Crimea, frequent kenophytes of Ukrainian forests are absent or rare, less naturalized and more recently established. The "true" steppic zone (the flatland part of the Steppic biogeoregion), on the other hand, has the highest proportion of kenophytes (3.1%). In the relevés from the Steppic biogeoregion including the Crimean Mountains, the most frequent forest kenophyte of Ukraine and Europe Impatiensparviflora is not present. In order to detect changes in the proportion of keno-phytes over the last few decades, 1498 older relevés sampled in 1950-1989 were analyzed. They correspond to 8 habitat types. For most of them, a comparison of proportions of kenophytes cannot be made for two reasons: 1) absence of relevés of the habitat sampled in one of two time periods (only "new" or only "old" data), 2) "old" and "new" relevés represent different areas. Only habitat G1.Aa "Carpinus and Quercus mesic deciduous woodland" has large number of "old" and "new" relevés from the same area. This area comprises central and western parts of the forest-steppe subregion and adjacent areas of the deciduous forests region of Western Ukraine. Two variants of selection of "new" relevés from this area were used: with and without relevés from Kyiv that have a high proportion of kenophytes. The results of the comparisons 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Table 4: Comparison of kenophyte proportions in "old" and "new" relevés of habitat G1.Aa "Carpinus and Quercus mesic deciduous woodland". Tabela 4: Primerjava deležev kenofitov v "starih" in "novih" popisih habitatnega tipa G1.Aa "mezofilni listopadni gozdovi vrst rodov Carpinus in Quercus ". Number of Mean % of Standard Mann-Whitney relevés kenophytes deviation U test Datasets old new old new old new p-level "Old" and "new" data without relevés from Kyiv 520 429 0.261 0.430 0.879 1.174 0.0163 "Old" and "new" data with relevés from Kyiv 520 481 0.261 0.812 0.879 1.831 0.0000 are in Table 4. The data does not satisfy the assumption of normality so it was tested with nonparametric MannWhitney U test with adjustment for t[[ies. The null hypothesis about equal kenophyte proportions in "old" and "new" relevés was rejected at the 1.63% significance level when "new" data did not include relevés from Kyiv and at the 0.00% level for dataset with relevés from Kyiv. In other words, the increased proportion of kenophytes is statistically reliable in both variants of comparison. In fact, the result is a little less certain because of possible differences in data sampling by different researchers. Viktor Onyshchenko ©, https://orcid.org/0000-0001-9079-7241 References Abduloyeva, O. 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[in Ukrainian]. 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine Appendix All forest habitats Biogeographical region and subregion Habitat type Geographical group of releves Latitude Longitude Radius (km) Year(s) Number of plots Mean % of kenophytes Number of plots in subregion * steppic: Crimean mountains G1.6a Ai-Petri 44,499 34,026 2 2004-2005 9 0,00 32 * steppic: Crimean mountains G1.Aa Anharskyi Pass 44,791 34,341 4 2005 5 0,00 32 * continental: forest-steppe G1.Aa Bernashivka 48,587 27,489 2 2002 1 0,00 657 * continental: deciduous forest G3.4a Bile Ozero 51,506 25,728 4 2004 15 0,00 1276 * alpine * Bukovyna alpine 48,197 25,694 32 1995-2005 86 0,73 530 alpine G1.6a Bukovyna alpine 48,214 25,664 29 1995-2005 59 0,60 530 alpine G1.Aa Bukovyna alpine 48,181 25,734 25 2000-2005 22 0,88 530 alpine G3.1b Bukovyna alpine 48,214 25,664 29 1995-2005 5 1,68 530 * continental: deciduous forest * Bukovyna deciduous 48,325 25,880 10 2003-2005 21 0,52 1276 continental: deciduous forest G1.6a Bukovyna deciduous 48,317 25,797 5 2003-2005 14 0,52 1276 continental: deciduous forest G1.Aa Bukovyna deciduous 48,325 25,880 10 2004-2005 3 0,00 1276 continental: deciduous forest G1.Ab Bukovyna deciduous 48,317 25,797 10 2003-2005 4 0,93 1276 * continental: forest-steppe * Bukovyna forest-steppe 48,046 25,946 28 2000-2005 21 0,25 657 continental: forest-steppe G1.6a Bukovyna forest-steppe 48,024 25,901 25 2000-2005 7 0,75 657 continental: forest-steppe G1.Aa Bukovyna forest-steppe 48,048 25,969 26 2000-2005 11 0,00 657 continental: forest-steppe G3.1b Bukovyna forest-steppe 48,024 25,901 29 1995-2005 3 0,00 530 * continental: deciduous forest * Cheremskyi 51,526 25,539 5 1998-2001 46 0,00 1276 continental: deciduous forest G3.4a Cheremskyi 51,526 25,539 5 1998-2001 29 0,00 1276 continental: deciduous forest G3.Da Cheremskyi 51,526 25,539 5 1998-2001 9 0,00 1276 continental: deciduous forest G3.A Cheremskyi 51,526 25,539 5 1998-2001 8 0,00 1276 * alpine G3.1a Chornohora 48,076 24,646 13 2002 7 0,00 530 * continental: forest-steppe G1.Aa Chornukhynshchyna 50,206 33,018 9 1997 25 0,00 657 * continental: forest-steppe G1.Aa Chornyi Lis 48,774 32,581 6 2001 41 0,00 657 * alpine * Chyvchyny 47,849 24,838 16 2002, 2003 20 0,21 530 alpine G3.1a Chyvchyny 47,849 24,838 16 2002, 2003 18 0,21 530 alpine G1.6a Chyvchyny 47,849 24,838 16 2002, 2003 2 0,21 530 * steppic: flatland G1.1 Danube delta 45,404 29,634 23 2003 7 10,16 180 * alpine G1.A4 Dubyno 48,544 22,803 5 2016 4 0,00 530 * alpine * Gorgany 48,441 24,344 8 2005 53 0,11 530 alpine G1.2a Gorgany 48,441 24,344 8 2005 5 0,00 530 alpine G1.6a Gorgany 48,441 24,344 8 2005 7 0,84 530 alpine G3.1a Gorgany 48,441 24,344 8 2005 27 0,00 530 alpine G3.1b Gorgany 48,441 24,344 8 2005 9 0,00 530 alpine G3.2 Gorgany 48,441 24,344 8 2005 5 0,00 530 * continental: forest-steppe G1.Aa Hadiach 50,333 34,027 22 2011-2016 40 1,03 657 * continental: deciduous forest * Halych 49,140 24,620 25 2010-2011 23 1,89 1276 continental: deciduous forest G1.1 Halych 49,083 24,667 9 2010-2011 3 7,89 1276 continental: deciduous forest G1.2b Halych 49,182 24,597 5 2010-2011 4 0,00 1276 continental: deciduous forest G1.6a Halych 49,076 24,600 9 2010-2011 6 0,56 1276 continental: deciduous forest G1.Aa Halych 49,163 24,620 11 2010-2011 10 1,64 1276 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine * alpine * Hutsulshchyna alpine G1.1 Hutsulshchyna alpine G1.2a Hutsulshchyna alpine G1.6a Hutsulshchyna alpine G1.6b Hutsulshchyna alpine Gl.Aa Hutsulshchyna alpine G3.1a Hutsulshchyna * continental: forest-steppe G1.Aa Ichnia * continental: forest-steppe G1.4 Irdyn * continental: deciduous forest G1.Aa Ivano-Frankivsk * continental: forest-steppe * Kaniv continental: forest-steppe G1.1 Kaniv continental: forest-steppe G1.7a Kaniv continental: forest-steppe G1.Aa Kaniv continental: forest-steppe G3.4a Kaniv * steppic: flatland * Kherson steppic: flatland G1.1 Kherson steppic: flatland G1.2a Kherson steppic: flatland G1.2b Kherson steppic: flatland G1.7a Kherson * continental forest-steppe * Khmelnytskyi continental forest-steppe G1.7a Khmelnytskyi continental forest-steppe G1.Aa Khmelnytskyi * continental forest-steppe * Khorol continental forest-steppe G1.1 Khorol continental forest-steppe G1.2a Khorol continental forest-steppe G1.4 Khorol continental forest-steppe G1.Aa Khorol * continental deciduous forest * Korostyshiv continental deciduous forest G1.7a Korostyshiv continental deciduous forest G1.Aa Korostyshiv continental deciduous forest G1.Ab Korostyshiv * continental forest-steppe G1.7a Kotovsk * continental forest-steppe * Kremenchuk continental forest-steppe G1.1 Kremenchuk continental forest-steppe G1.4 Kremenchuk * continental deciduous forest G1.Aa Kremenets * continental deciduous forest * Kyiv west continental deciduous forest G1.7a Kyiv west continental deciduous forest G1.2a Kyiv west continental deciduous forest G1.2b Kyiv west continental deciduous forest G1.4 Kyiv west continental deciduous forest G1.Aa Kyiv west continental deciduous forest G3.4a Kyiv west * continental forest-steppe * Kyiv south continental forest-steppe G3.4a Kyiv south continental forest-steppe G1.7a Kyiv south continental forest-steppe G1.4 Kyiv south 48,335 24,953 21 2006-2008 73 0,75 530 48,339 25,280 5 2008 3 5,34 530 48,335 24,953 21 2006-2008 11 2,10 530 48,335 24,953 21 2006, 2007 20 0,17 530 48,335 24,953 21 2006, 2007 9 0,00 530 48,335 24,953 21 2006, 2008 18 0,69 530 48,335 24,953 21 2006, 2007 12 0,00 530 50,719 32,317 28 2004 6 0,00 657 49,496 31,819 3 1996 3 0,00 657 48,972 24,683 10 2006-2007 7 0,29 1276 49,775 31,535 14 1990-1995 53 4,91 657 49,719 31,569 2,5 1990-1995 15 16,17 657 49,855 31,549 1,7 1990-1995 9 0,00 657 49,717 31,500 4 1990-1995 26 0,67 657 49,855 31,549 1,7 1990-1995 3 0,00 657 46,483 32,291 54 1999-2010 60 3,13 180 46,482 32,676 25 2000-2010 12 4,29 180 46,484 32,144 42 1999-2010 16 2,49 180 46,484 32,144 42 1999-2010 8 1,39 180 46,484 31,911 21 1998-2009 24 3,56 180 49,418 27,052 12 2004 28 1,23 657 49,325 27,007 0,5 2004 3 1,15 657 49,418 27,052 12 2004 25 1,24 657 50.0580 33.6073 67 2001-2004 60 2,77 657 50.0580 33.6073 67 2001-2004 8 8,21 657 50.0580 33.6073 67 2001-2004 11 5,37 657 50.0580 33.6073 67 2001-2004 8 1,35 657 50.0580 33.6073 67 2003-2004 33 0,93 657 50,270 29,095 25 2001-2004 23 0,48 1276 50,270 29,095 25 2001-2004 10 0,42 1276 50,270 29,095 25 2001-2004 6 0,00 1276 50,270 29,095 25 2004 7 3,65 1276 47,807 29,493 12 2007 15 2,25 657 49,053 33,428 9 2000-2005 24 7,85 657 49,053 33,428 9 2000-2005 17 9,84 657 49,053 33,428 9 2000-2005 7 3,02 657 50,074 25,655 2 1999 7 0,00 1276 50,506 30,302 10 2015-2017 48 4,91 1276 50,506 30,302 10 2015-2018 14 4,42 1276 50,506 30,302 10 2015-2019 9 5,83 1276 50,506 30,302 10 2015-2020 9 5,75 1276 50,506 30,302 10 2015-2021 5 2,53 1276 50,506 30,302 10 2015-2022 3 5,84 1276 50,506 30,302 10 2015-2022 8 6,84 1276 50,302 30,544 23 2003-2017 154 4,35 657 50,291 30,577 9 2003-2017 33 4,71 657 50,333 30,541 8 2008-2014 4 1,99 657 50,333 30,541 8 2003-2015 15 4,42 657 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine continental: forest-steppe Gl.Aa Kyiv south continental: forest-steppe Gl.2a Kyiv south continental: forest-steppe Gl.2b Kyiv south * continental: forest-steppe G3.4a Kyiv plato * steppic: Crimean mountains Gl.6a Laspi * continental: deciduous forest * Lviv continental: deciduous forest G1.6a Lviv continental: deciduous forest G1.Aa Lviv * alpine * Mala Uholka alpine G1.6a Mala Uholka alpine G1.Ab Mala Uholka * continental: deciduous forest * Medobory continental: deciduous forest G1.6a Medobory continental: deciduous forest G1.Aa Medobory continental: deciduous forest G1.Ab Medobory * alpine * Mukachevo alpine alpine G1.6a Mukachevo alpine alpine G1.Aa Mukachevo alpine * pannonian G1.Aa Mukachevo pannonian * continental: forest-steppe * Mykhailivka continental: forest-steppe G1.7a Mykhailivka continental: forest-steppe G3.4a Mykhailivka * alpine * Mykulychyn alpine G1.6a Mykulychyn alpine G3.1a Mykulychyn alpine G3.1b Mykulychyn * continental: deciduous forest G1.Aa Panivetska Dacha * continental: deciduous forest * Perebrody continental: deciduous forest G1.4 Perebrody continental: deciduous forest G3.4a Perebrody continental: deciduous forest G3.Da Perebrody * continental: forest-steppe G1.7a Poltava * continental: deciduous forest G1.Aa Radomska Dacha * continental: forest-steppe * Ros continental: forest-steppe G1.1 Ros continental: forest-steppe G1.4 Ros continental: forest-steppe G3.4a Ros * continental: deciduous forest * Roztochia continental: deciduous forest G1.2a Roztochia continental: deciduous forest G1.4 Roztochia continental: deciduous forest G1.6a Roztochia continental: deciduous forest G1.7a Roztochia continental: deciduous forest G1.8 Roztochia continental: deciduous forest G1.Aa Roztochia continental: deciduous forest G1.Ab Roztochia continental: deciduous forest G3.4a Roztochia * continental: deciduous forest G3.A Rys * steppic: flatland G3.4a Samara 50,333 30,541 8 2003, 2009 53 3,89 657 50,333 30,541 8 2003-2016 7 4,20 657 50,333 30,541 8 2003-2016 42 4,88 657 49,941 30,673 60 1996-1991 8 0,00 657 44,419 33,756 3 2004-2005 5 0,00 32 49,827 24,025 12 1992-1997 17 2,37 1276 49,827 24,025 12 1992-1997 10 3,09 1276 49,827 24,025 12 1992-1997 7 1,34 1276 48,262 23,632 6 2005-2006 34 0,08 530 48,275 23,658 4 2005-2006 31 0,00 530 48,245 23,618 4 2016 3 0,93 530 49,282 26,164 15 1995-1998 80 0,24 1276 49,282 26,164 15 1995-1998 4 0,00 1276 49,282 26,164 15 1995-1998 49 0,36 1276 49,282 26,164 15 1995-1998 27 0,07 1276 48,473 22,740 3 2003-2004 20 0,63 530 48,478 22,746 3 2003-2004 10 0,98 530 48,469 22,740 6 2003-2004 10 0,28 530 48,399 22,670 3 2003-2004 12 0,23 26 49,609 31,509 7 1998-2003 17 0,00 657 49,609 31,509 7 2003 5 0,00 657 49,609 31,509 7 1998-2003 12 0,00 657 48,401 24,594 8 2002 14 0,89 530 48,384 24,574 8 2002-2003 6 2,20 530 48,403 24,570 6 2002 4 0,00 530 48,392 24,613 4 2002 4 1,79 530 48,619 26,594 3 2004 14 0,30 1276 51,689 27,100 12 2004 19 2,16 1276 51,689 27,100 12 2004 3 0,00 1276 51,689 27,100 12 2004 8 5,13 1276 51,689 27,100 12 2004 8 0,00 1276 49,552 34,632 1 1997 2 0,00 657 51,948 32,483 5 2002 28 0,00 1276 49,721 30,403 90 1998-2000 43 3,00 657 49,721 30,403 90 1999-2000 15 6,72 657 49,721 30,403 90 1999-2000 13 1,15 657 49,556 30,885 52 1998-2000 15 0,88 657 50,001 23,668 20 1990-2000 179 0,25 1276 50,001 23,668 20 1996, 2000 13 0,28 1276 50,001 23,668 20 1990-1996 12 0,00 1276 50,001 23,668 20 1990-1999 44 0,38 1276 50,001 23,668 20 1996, 2000 9 0,00 1276 50,001 23,668 20 1990-1998 6 0,00 1276 50,001 23,668 20 1990-2000 40 0,31 1276 50,108 23,701 4 1996, 2004 2 0,00 1276 50,001 23,668 20 1990-1998 53 0,21 1276 51,261 25,384 1,5 1997 6 0,00 1276 48,656 35,329 17 2003 49 5,17 180 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine * continental: forest-steppe * Sestrynivska Dacha continental: forest-steppe Gl.2b Sestrynivska Dacha continental: forest-steppe Gl.Aa Sestrynivska Dacha * continental: deciduous forest * Shepetivka continental: deciduous forest Gl.2a Shepetivka continental: deciduous forest Gl.2b Shepetivka continental: deciduous forest G1.4 Shepetivka continental: deciduous forest Gl.7a Shepetivka continental: deciduous forest G1.8 Shepetivka continental: deciduous forest Gl.Aa Shepetivka continental: deciduous forest G3.4a Shepetivka continental: deciduous forest G3.Da Shepetivka * alpine * Skole alpine Gl.2a Skole alpine Gl.6a Skole alpine G1.6b Skole alpine Gl.Aa Skole alpine Gl.Ab Skole alpine G3.lb Skole * steppic: flatland Gl.Aa Snizhne * continental: deciduous forest G3.4a Somyno * continental: deciduous forest * Stara Huta continental: deciduous forest Gl.l Stara Huta continental: deciduous forest Gl.2a Stara Huta continental: deciduous forest G1.2b Stara Huta continental: deciduous forest G1.4 Stara Huta continental: deciduous forest Gl.7a Stara Huta continental: deciduous forest G1.8 Stara Huta continental: deciduous forest Gl.Aa Stara Huta continental: deciduous forest G3.4a Stara Huta continental: deciduous forest G3.Da Stara Huta * steppic: Crimean mountains Gl.Aa Staryi Krym * continental: forest-steppe * Sumy continental: forest-steppe Gl.2b Sumy continental: forest-steppe G1.4 Sumy continental: forest-steppe Gl.7a Sumy continental: forest-steppe Gl.Aa Sumy continental: forest-steppe G3.4a Sumy * steppic: flatland * Sviati Hory steppic: flatland Gl.7a Sviati Hory steppic: flatland Gl.Aa Sviati Hory steppic: flatland G3.4b Sviati Hory * alpine * Synevyr alpine Gl.2a Synevyr alpine Gl.Ab Synevyr alpine Gl.6a Synevyr alpine G1.6b Synevyr alpine G3.la Synevyr 49,736 28,886 3 2003-2004 6 1,01 637 49,736 28,886 3 2003-2004 S 0,74 637 49,736 28,886 3 2003-2004 1 2,33 637 30,230 26,908 36 1999-2001 39 0,00 1276 30,230 26,908 36 1999-2001 3 0,00 1276 30,230 26,908 36 1998-2001 6 0,00 1276 30,230 26,908 36 1999-2001 3 0,00 1276 30,230 26,908 36 1999-2001 3 0,00 1276 S0,2S0 26,908 36 1999-2001 4 0,00 1276 30,230 26,908 36 1999-2001 1S 0,00 1276 30,230 26,908 36 1997-2001 6 0,00 1276 30,230 26,908 36 1997-2001 19 0,00 1276 49,033 23,3S9 17 1990-2001 83 0,14 330 49,03S 23,3S9 17 1990-2001 8 0,66 330 49,0SS 23,3S9 17 1990-2001 49 0,14 330 49,0SS 23,3S9 17 1990-2001 S 0,00 330 49,0SS 23,3S9 17 1990-2003 9 0,00 330 49,0SS 23,3S9 17 1990-2003 3 0,00 330 49,0SS 23,3S9 17 1990-2001 11 0,00 330 48,03S 38,783 1 2003 11 1,46 180 31,414 26,916 8 2004 13 0,00 1276 S2,277 33,S67 21 2012 176 0,43 1276 S2,333 33,343 1 2012 4 7,03 1276 S2,297 33,331 13 2012 2 0,00 1276 S2,242 33,364 S 2000-2010 16 1,16 1276 S2,292 33,341 12 1998-2010 12 0,37 1276 S2,277 33,S67 21 2001-2012 6 1,87 1276 S2,339 33,763 7 1997-2010 16 0,00 1276 S2,328 33,732 S 1997-1999 30 0,00 1276 S2,276 33,620 18 1997-2012 72 0,19 1276 S2,342 33,698 10 1997-2012 18 0,00 1276 43,021 33,072 4 2007 13 0,00 32 S0,678 34,839 SS 1996-2002 39 0,12 637 S0,678 34,839 3S 1996-2002 S 0,00 637 S0,678 34,839 3S 1996-2002 4 0,00 637 S0,678 34,839 3S 1996-2002 19 0,38 637 S0,678 34,839 3S 1996-2002 20 0,00 637 S0,678 34,839 3S 1996-2002 11 0,00 637 48,9S6 37,644 23 2006 44 0,12 180 48,988 37,493 11 2006 3 0,31 180 48,988 37,493 11 2006 24 0,1S 180 49,029 37,493 7 2001 17 0,00 180 48,S08 23,703 16 1990-2016 96 0,00 330 48,S08 23,70S 16 1990-2016 S 0,00 330 48,S08 23,70S 16 1990-2016 2 0,00 330 48,S08 23,70S 16 1990-2016 22 0,00 330 48,S08 23,70S 16 1990-2016 14 0,00 330 48,S08 23,70S 16 1990-2016 49 0,00 330 290 O Hacquetia 18/2 • 2019, 289-312 Viktor Onyshchenko Kenophytes in different forest types of Ukraine alpine G3.1b Synevyr 48,508 23,705 16 1990-2016 4 0,00 530 * steppic: flatland Gl.7a Tarutyno 46,206 29,146 4 2007 4 5,21 180 * continental: deciduous forest G3.A Tkhoryn 51,415 28,317 15 1999 3 0,00 1276 * continental: deciduous forest * Tsuman 50,926 25,884 9 2002-2003 15 0,00 1276 continental: deciduous forest G1.7a Tsuman 50,875 25,965 0,5 2002-2003 3 0,00 1276 continental: deciduous forest G1.Aa Tsuman 50,926 25,884 8 2002-2003 12 0,00 1276 * continental: forest-steppe G1.Aa Uman 48,719 30,243 5 2004-2005 22 1,81 657 * continental: forest-steppe G1.Aa Vasyshchevo 49,826 36,278 1 2006 8 0,00 657 * continental: deciduous forest * Verkhniobuzki Lisy 49,914 24,941 9 2002-2006 387 0,17 1276 continental: deciduous forest G1.6a Verkhniobuzki Lisy 49,914 24,941 9 2002-2005 143 0,11 1276 continental: deciduous forest G1.Aa Verkhniobuzki Lisy 49,914 24,941 9 2002-2006 223 0,17 1276 continental: deciduous forest G1.Ab Verkhniobuzki Lisy 49,914 24,941 9 2002-2005 21 0,48 1276 * pannonian G1.Ab Vynohradiv-Shaian 48,131 23,227 13 2016 14 1,69 26 * alpine * Vyzhnytsia 48,192 25,208 8 1999-2004 38 0,00 530 alpine G1.2a Vyzhnytsia 48,192 25,208 8 2002-2004 3 0,00 530 alpine G1.6a Vyzhnytsia 48,192 25,208 8 1999-2004 11 0,00 530 alpine G1.6b Vyzhnytsia 48,192 25,208 8 2004 9 0,00 530 alpine G1.Ab Vyzhnytsia 48,192 25,208 8 2002 3 0,00 530 alpine G3.1a Vyzhnytsia 48,192 25,208 8 2002 4 0,00 530 alpine G3.1b Vyzhnytsia 48,192 25,208 8 1999-2004 8 0,00 530 * steppic: flatland G1.7a Zaporizki Balky 47,817 35,201 35 2003-2006 5 1,66 180 * continental: deciduous forest G1.7a Zhukiv Khutir 50,483 30,080 2 1996 13 3,68 1276 * continental: forest-steppe G1.Aa Zhuravlivska Dacha 48,597 28,714 4 2003 21 1,70 657 * continental: deciduous forest * Zhytomyrska 50,840 28,334 81 2000-2003 77 0,06 1276 continental: deciduous forest G3.4a Zhytomyrska 50,840 28,334 81 2000-2003 53 0,09 1276 continental: deciduous forest G3.Da Zhytomyrska 50,840 28,334 81 2000-2002 24 0,00 1276 290 90S BL BL BL BL Title string o o E K O O O O O O O O O O O O Ci ■í ir B Cl CI n o o ci ci ci m m p c¡ Habitat type 4x 4x G\ \D CO \D Lati-tude oj OJ oj OJ OJ OJ oo oo 00 ^ o\ \£> Longitude Radius (km) to to to to to to to to Uï No. of releves •--J ON \£> \£> Mean % of kenophytes d > _ o E2 cr PT sT c S s- 3 a. s s. g 03 3 3 2_ Mean for dataset Mean for geographical groups "B S ° g o '"O _ S c CT02 Br pi I P Mean for European dataset to to \£> \£> NJ ^ NJ - ÏÏ 3 b' 3 p B B ! m S o ^ ^ S 3 S g- n> o o- 2- o « i-h X m ^ ^J ^J ---J 3Ure.D[Q JO SodXl 1S9JOJ lU3J3JJTp UT S9}XtjdoU3]>[ 0^U31|DI|SXUQ JOl^TyY ZIÍ-G2Z '6\0Z • Z/81 Bpanbora JJ q continental: forest-steppe G1.2b Kyiv south 50,333 30,541 8 2003-2016 continental: deciduous forest G1.2b Kyiv west 50,506 30,302 10 2015-2020 continental: forest-steppe G1.2a Sestrynivska Dacha 49,756 28,886 3 2003-2004 continental: deciduous forest G1.2b Shepetivka 50,250 26,908 36 1998-2001 continental: deciduous forest G1.2b Stara Huta 52,242 33,364 5 2000-2010 continental: forest-steppe G1.2b Sumy 50,678 34,859 55 1996-2002 G1.4 continental: forest-steppe G1.4 Irdyn 49,496 31,819 3 1996 continental: forest-steppe G1.4 Khorol 50.0580 33.6073 67 2001-2004 continental: forest-steppe G1.4 Kremenchuk 49,053 33,428 9 2000-2005 continental: deciduous forest G1.4 Kyiv west 50,506 30,302 10 2015-2021 continental: forest-steppe G1.4 Kyiv south 50,333 30,541 8 2003-2015 continental: deciduous forest G1.4 Perebrody 51,689 27,100 12 2004 continental: forest-steppe G1.4 Ros 49,721 30,403 90 1999-2000 continental: deciduous forest G1.4 Roztochia 50,001 23,668 20 1990-1996 continental: deciduous forest G1.4 Shepetivka 50,250 26,908 36 1999-2001 continental: deciduous forest G1.4 Stara Huta 52,292 33,541 12 1998-2010 continental: forest-steppe G1.4 Sumy 50,678 34,859 55 1996-2002 G1.6a steppic: Crimean mountains G1.6a Ai-Petri 44,499 34,026 2 2004-2005 alpine G1.6a Bukovyna alpine 48,214 25,664 29 1995-2005 continental: deciduous forest G1.6a Bukovyna deciduous 48,317 25,797 5 2003-2005 continental: forest-steppe G1.6a Bukovyna forest-steppe 48,024 25,901 25 2000-2005 alpine G1.6a Chyvchyny 47,849 24,838 16 2002, 2003 alpine G1.6a Gorgany 48,441 24,344 8 2005 continental: deciduous forest G1.6a Halych 49,076 24,600 9 2010-2011 alpine G1.6a Hutsulshchyna 48,335 24,953 21 2006, 2007 steppic: Crimean mountains G1.6a Laspi 44,419 33,756 3 2004-2005 continental: deciduous forest G1.6a Lviv 49,827 24,025 12 1992-1997 alpine G1.6a Mala Uholka 48,275 23,658 4 2005-2006 42 4,88 204,96 657 9 5,75 51,75 1276 5 0,74 3,70 657 6 0,00 0,00 1276 16 1,16 18,56 1276 5 0,00 0,00 657 85 300,00 1,46 1,19 0,57 0,9 3 0,00 0,00 657 8 1,35 2,22 657 7 3,02 21,14 657 5 2,53 12,65 1276 15 4,42 66,3 657 3 0,00 0,00 1276 13 1,15 14,95 657 12 0,00 0,00 1276 3 0,00 0,00 1276 12 0,57 6,84 1276 4 0,00 0,00 657 459 650,00 0,34 0,56 0,21 0,3 9 0,00 0,00 32 59 0,60 35,40 530 14 0,52 7,28 1276 7 0,75 5,25 657 2 0,21 0,43 530 7 0,84 5,88 530 6 0,56 3,33 1276 20 0,17 3,44 530 5 0,00 0,00 32 10 3,09 30,90 1276 31 0,00 0,00 530 o ffi p Ci 'S rt» h-" • oo ¡o to o to 00 VD I OJ s sr >2 O 3! < a' B-B r, a o S continental: deciduous lorest G1.6a Medobory 49,282 26,164 15 1995-1998 alpine G1.6a Mukachevo 48,478 22,746 3 2003-2004 alpine G1.6a Mykulychyn 48,384 24,574 8 2002-2003 continental: deciduous lorest G1.6a Roztochia 50,001 23,668 20 1990-1999 alpine G1.6a Skole 49,055 23,359 17 1990-2001 alpine G1.6a Synevyr 48,508 23,705 16 1990-2016 continental: deciduous lorest G1.6a Verkniobuzki Lisy 49,914 24,941 9 2002-2005 alpine G1.6a Vyzhnytsia 48,192 25,208 8 1999-2004 G1.6b alpine G1.6b Vyzhnytsia 48,192 25,208 8 2004 alpine G1.6b Hutsulshchyna 48,335 24,953 21 2006,2007 alpine G1.6b Skole 49,055 23,359 17 1990-2001 alpine G1.6b Synevyr 48,508 23,705 16 1990-2016 G1.7a continental: torest-steppe G1.7a Kaniv 49,855 31,549 1,7 1990-1995 steppic: flatland G1.7a Kherson 46,484 31,911 21 1998-2009 continental: torest-steppe G1.7a Khmelnytskyi 49,325 27,007 0,5 2004 continental: deciduous lorest G1.7a Korostyshiv 50,270 29,095 25 2001-2004 continental: torest-steppe G1.7a Kotovsk 47,807 29,493 12 2007 continental: deciduous torest G1.7a Kyiv west 50,506 30,302 10 2015-2018 continental: torest-steppe G1.7a Kyiv south 50,333 30,541 8 2008-2014 continental: torest-steppe G1.7a Mykhailivka 49,609 31,509 7 2003 continental: torest-steppe G1.7a Poltava 49,552 34,632 1 1997 continental: deciduous torest G1.7a Roztochia 50,001 23,668 20 1996, 2000 continental: deciduous torest G1.7a Shepetivka 50,250 26,908 36 1999-2001 continental: deciduous torest G1.7a Stara Huta 52,277 33,567 21 2001-2012 continental: torest-steppe G1.7a Sumy 50,678 34,859 55 1996-2002 steppic: flatland G1.7a Sviati Hory 48,988 37,493 11 2006 steppic: flatland G1.7a Tarutyno 46,206 29,146 4 2007 continental: deciduous forest G1.7a Tsuman 50,875 25,965 0,5 2002-2003 4 0,00 0,00 1276 10 0,98 9,8 530 6 2,20 13,19 530 44 0,38 16,90 1276 49 0,14 6,81 530 22 0,00 0,00 530 143 0,11 16,30 1276 11 0,00 0,00 530 37 80,00 0,00 0,00 0,00 0,9 9 0,00 0,00 530 9 0,00 0,00 530 5 0,00 0,00 530 14 0,00 0,00 530 151 200,00 1,94 1,51 0,83 0,4 9 0,00 0,00 657 24 3,56 85,44 180 3 1,15 3,45 657 10 0,42 4,20 1276 15 2,25 33,75 657 14 4,42 61,88 1276 4 1,99 7,96 657 5 0,00 0,00 657 2 0,00 0,0 657 9 0,00 0,00 1276 3 0,00 0,00 1276 6 1,87 11,22 1276 19 0,38 7,22 657 3 0,51 1,53 180 4 5,21 20,84 180 3 0,00 0,00 1276 o ffi p Ci 'S rt» h-" • oo ¡o to o to 00 VD I OJ s sr >2 O 3! < a' B-B r, o- 5" a o S steppic: flatland G1.7a Zaporizki Balky 47,817 35,201 35 2003-2006 continental: deciduous torest G1.7a Zhukiv Khutir 50,483 30,080 2 1996 G1.8 continental: deciduous torest G1.8 Stara Huta 52,339 33,763 7 1997-2010 continental: deciduous torest G1.8 Shepetivka 50,250 26,908 36 1999-2001 continental: deciduous torest G1.8 Roztochia 50,001 23,668 20 1990-1998 GLAa steppic: Crimean mountains Gl.Aa Anharskyi Pass 44,791 34,341 4 2005 continental: torest-steppe GLAa Bernashivka 48,587 27,489 2 2001 alpine GLAa Bukovyna alpine 48,181 25,734 22 2000-2005 continental: deciduous torest GLAa Bukovyna deciduous 48,325 25,880 10 2004-2005 continental: torest-steppe GLAa Bukovyna torest-steppe 48,048 25,969 26 2000-2005 continental: torest-steppe GLAa Chornukhynshchyna 50,206 33,018 9 1997 continental: torest-steppe GLAa Chornyi Lis 48,774 32,581 6 2001 continental: torest-steppe GLAa Hadiach 50,333 34,027 22 2011-2016 continental: deciduous torest GLAa Halych 49,163 24,620 11 2010-2011 alpine GLAa Hutsulshchyna 48,335 24,953 21 2006, 2008 continental: torest-steppe GLAa Ichnia 50,719 32,317 28 2004 continental: torest-steppe GLAa Kaniv 49,717 31,500 4 1990-1995 continental: torest-steppe GLAa Khmelnytskyi 49,418 27,052 12 2004 continental: torest-steppe GLAa Khorol 50.0580 33.6073 67 2003-2004 continental: deciduous torest GLAa Korostyshiv 50,270 29,095 25 2001-2004 continental: deciduous torest GLAa Kremenets 50,074 25,655 2 1999 continental: deciduous torest GLAa Kyiv west 50,506 30,302 10 2015-2022 continental: torest-steppe GLAa Kyiv south 50,333 30,541 8 2003, 2009 continental: deciduous torest GLAa Lviv 49,827 24,025 12 1992-1997 continental: deciduous torest GLAa Medobory 49,282 26,164 15 1995-1998 alpine GLAa Mukachevo alpine 48,469 22,740 6 2003-2004 pannonian GLAa Mukachevo pannonian 48,399 22,670 3 2003-2004 continental: torest-steppe GLAa Panivetska Dacha 48,619 26,594 3 2004 5 13 1,66 3,68 26 16 0,00 4 0,00 6 0,00 800,00 0,00 0,00 0,00 1,4 8,30 47,84 0,00 0,00 0,00 180 1276 1276 1276 1276 911 1800,00 0,64 0,71 0,28 0,8 5 0,00 0,00 32 1 0,00 0,00 657 22 0,88 19,36 530 3 0,00 0,00 1276 11 0,00 0 657 25 0,00 0 657 41 0,00 0 657 40 1,03 41,2 657 10 1,64 16,40 1276 18 0,69 12,40 530 6 0,00 0 657 26 0,67 17,42 657 25 1,24 31 657 33 0,93 30,69 657 6 0,00 0,00 1276 7 0,00 0,00 1276 3 5,84 17,52 1276 53 3,89 206,17 657 7 1,34 9,38 1276 49 0,36 17,40 1276 10 0,28 2,8 530 12 0,23 2,76 26 14 0,30 4,2 657 O ffi p Ci •i rt» h-" • 00 ¡o to o to 00 VD I OJ s sr 2 o a < B r, o- 5" a o S continental: deciduous lorest Gl.Aa Radomska Dacha 51,948 32,483 5 2002 continental: deciduous lorest Gl.Aa Roztochia 50,001 23,668 20 1990-2000 continental: deciduous lorest Gl.Aa Ivano-Frankivsk 48,972 24,683 10 2006-2007 continental: torest-steppe Gl.Aa Sestrynivska Dacha 49,756 28,886 3 2003-2004 continental: deciduous lorest Gl.Aa Shepetivka 50,250 26,908 36 1999-2001 alpine Gl.Aa Skole 49,055 23,359 17 1990-2003 steppic: flatland Gl.Aa Snizhne 48,035 38,783 1 2003 continental: deciduous lorest Gl.Aa Stara Huta 52,328 33,732 5 1997-1999 steppic: Crimean mountains Gl.Aa Staryi Krym 45,021 35,072 4 2007 continental: torest-steppe Gl.Aa Sumy 50,678 34,859 55 1996-2002 steppic: flatland Gl.Aa Sviati Hory 48,988 37,493 11 2006 continental: deciduous lorest Gl.Aa Tsuman 50,926 25,884 8 2002-2003 continental: torest-steppe Gl.Aa Uman 48,719 30,243 5 2004-2005 continental: torest-steppe Gl.Aa Vasyshchevo 49,826 36,278 1 2006 continental: deciduous torest Gl.Aa Verkniobuzki Lisy 49,914 24,941 9 2002-2006 continental: torest-steppe Gl.Aa Zhuravlivska Dacha 48,597 28,714 4 2003 Gl.Ab alpine Gl.Ab Dubyno 48,565 22,824 3 2016 continental: deciduous torest Gl.Aa Korostyshiv 50,270 29,095 25 2001-2004 alpine Gl.Ab Mala Uholka 48,245 23,618 4 2016 continental: deciduous torest Gl.Ab Medobory 49,282 26,164 15 1995-1998 continental: deciduous torest Gl.Ab Roztochia 50,108 23,701 4 1996, 2004 alpine Gl.Ab Skole 49,055 23,359 17 1990-2003 alpine Gl.Ab Synevyr 48,508 23,705 16 1990-2016 continental: deciduous torest Gl.Ab Verkniobuzki Lisy 49,914 24,941 9 2002-2005 continental: deciduous torest Gl.Ab Bukovyna deciduous 48,317 25,797 5 2003-2005 pannonian Gl.Ab Vynohradiv-Shaian 48,131 23,227 13 2016 alpine Gl.Ab Vyzhnytsia 48,192 25,208 8 2002 G3.1a alpine G3.1a Chornohora 48,076 24,646 13 2002 28 0,00 0,00 1276 40 0,31 12,40 1276 7 0,29 2,03 1276 1 2,33 2,33 657 15 0,00 0,00 1276 9 0,00 0 530 11 1,46 16,06 180 30 0,00 0,00 1276 13 0,00 0 32 20 0,00 0,00 657 24 0,15 3,6 180 12 0,00 0,00 1276 22 1,81 39,82 657 8 0,33 2,64 657 223 0,17 38,80 1276 21 1,70 35,7 657 90 5,00 0,50 0,41 0,07 0,8 4 0,00 0,00 530 7 0,43 3,01 1276 3 0,93 2,78 ^30 27 0,07 2,00 1276 2 0,00 0,00 1276 3 0,00 0,00 530 2 0,00 0,00 530 21 0,48 9,98 1276 4 0,93 3,72 1276 14 1,69 23,66 26 3 0,00 0,00 530 o ffi o •g O) <-h h-' • P 00 to to C' to 00 VC I LU B 5" ■% S. Jf O Ii 3 a s- ri O ? 500,00 0,03 0,03 0,00 0,1 0,00 0,00 530 alpine alpine alpine alpine alpine alpine * alpine continental: lorest-steppe alpine alpine alpine alpine alpine alpine * continental: deciduous lorest continental: deciduous lorest continental: lorest-steppe continental: deciduous forest continental: lorest-steppe continental: lorest-steppe continental: lorest-steppe continental: deciduous lorest continental: lorest-steppe continental: deciduous forest steppic: flatland continental: deciduous lorest continental: deciduous lorest G3.1a Chyvchyny 47,849 24,838 16 2002, 2003 G3.1a Gorgany 48,441 24,344 8 2005 G3.1a Hutsulshchyna 48,335 24,953 21 2006, 2007 G3.1a Mykulychyn 48,403 24,570 6 2002 G3.1a Synevyr 48,508 23,705 16 1990-2016 G3.1a Vyzhnytsia 48,192 25,208 8 2002 G3.1b G3.1b Bukovyna alpine 48,214 25,664 29 1995-2005 G3.1b Bukovyna lorest-steppe 48,024 25,901 25 2000-2005 G3.1b Gorgany 48,441 24,344 8 2005 G3.1b Mykulychyn 48,392 24,613 6 2002 G3.1b Skole 49,055 23,359 17 1990-2001 G3.1b Synevyr 48,508 23,705 16 1990-2016 G3.1b Vyzhnytsia 48,192 25,208 8 1999-2004 G3.2 G3.2 Gorgany 48,441 24,344 8 2005 G3.4a G3.4a Bile Ozero 51,506 25,728 4 2004 G3.4a Cheremskyi 51,526 25,539 5 1998-2001 G3.4a Kaniv 49,855 31,549 1.7 1990-1995 G3.4a Kyiv west 50,506 30,302 10 2015-2022 G3.4a Kyiv south 50,291 30,577 9 2003-2017 G3.4a Kyiv plato 49,941 30,673 60 1996-1991 G3.4a Mykhailivka 49,609 31,509 7 1998-2003 G3.4a Perebrody 51,689 27,100 12 2004 G3.4a Ros 49.556 30,885 52 1998-2000 G3.4a Roztochia 50,001 23,668 20 1990-1998 G3.4a Samara 48,656 35,329 17 2003 G3.4a Shepetivka 50,250 26,908 36 1997-2001 G3.4a Somyno 51,414 26,916 8 2004 18 0,21 3,85 530 27 0,00 0,00 530 12 0,00 0,00 530 4 0,00 0,00 530 49 0,00 0,00 530 4 0,00 0,00 530 44 130,00 0,35 0,50 0,00 0,1 5 1,68 8,40 530 3 0,00 0,00 657 9 0,00 0,00 530 4 1,79 7,14 530 11 0,00 0,00 530 4 0,00 0,00 530 8 0,00 0,00 530 5 0,18 0,00 0,00 0,00 0 5 0,00 0,00 530 388 2700,00 1,41 1,45 0,05 1,4 15 0,00 0,00 1276 29 0,00 0,00 1276 3 0,00 0,00 657 8 6,84 54,72 1276 33 4,71 155,43 657 8 0,00 0,00 657 12 0,00 0,00 657 8 5,13 41,04 1276 15 0,88 13,20 657 53 0,21 11,13 1276 49 5,17 253,33 180 6 0,00 0,00 1276 13 0,00 0,00 1276 o ffi p Ci 'S rt» h-" • oo ¡o to o to 00 VD I OJ s sr >2 O 3! < a' B-B r, o- 5" a o S continental: deciduous forest continental: forest-steppe continental: deciduous forest * steppic: flatland * continental: deciduous forest continental: deciduous lorest continental: deciduous lorest continental: deciduous lorest continental: deciduous lorest continental: deciduous lorest continental: deciduous lorest continental: deciduous lorest G3.4a Stara Huta 52,276 33,620 18 1997-2012 G3.4a Sumy 50,678 34,859 55 1996-2002 G3.4a Zhytomyrska 50,840 28,334 81 2000-2003 G3.4b G3.4b Sviati Hory 49,029 37,493 7 2001 G3.A G3A Cheremskyi 51,526 25,539 5 1998-2001 G3A Rys 51,261 25,384 1,5 1997 G3A Tkhoryn 51,415 28,317 15 1999 G3.Da G3.Da Cheremskyi 51,526 25,539 5 1998-2001 G3.Da Perebrody 51,689 27,100 12 2004 G3.Da Shepetivka 50,250 26,908 36 1997-2001 G3.Da Stara Huta 52,342 33,698 10 1997-2012 G3.Da Zhytomyrska 50,840 28,334 81 2000-2002 72 0,19 13,90 1276 11 0,00 0,00 657 53 0,09 4,98 1276 17 12,00 0,00 0,00 0,00 0,1 17 0,00 0,00 180 17 0,50 0,00 0,00 0,00 0 8 0,00 0,00 1276 6 0,00 0,00 1276 3 0,00 0,00 1276 78 500,00 0,00 0,00 0,00 0 9 0,00 0,00 1276 8 0,00 0,00 1276 19 0,00 0,00 1276 18 0,00 0,00 1276 24 0,00 0,00 1276 o ffi p Ci 'S rt» h-" • oo ¡o to o to 00 VD I OJ s sr >2 O 3! < a' B-B r, o- 5" a 3