Научная статья на тему 'THE NON-NATIVE WOODY SPECIES OF THE FLORA OF UKRAINE: INTRODUCTION, NATURALIZATION AND INVASION'

THE NON-NATIVE WOODY SPECIES OF THE FLORA OF UKRAINE: INTRODUCTION, NATURALIZATION AND INVASION Текст научной статьи по специальности «Биологические науки»

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Текст научной работы на тему «THE NON-NATIVE WOODY SPECIES OF THE FLORA OF UKRAINE: INTRODUCTION, NATURALIZATION AND INVASION»

Biosystems

Diversity

ISSN 2519-8513 (Print) ISSN 2520-2529 (Online) Biosyst. Divers., 2019, 27(3), 276-290 doi: 10.15421/011937

The non-native woody species of the flora of Ukraine: Introduction, naturalization and invasion

R. I. Burda, S. N. Koniakin

Institute for Evolutionary Ecology, National Academy of Sciences of Ukraine, Kyiv, Ukraine

Article info

Received 16.07.2019 Received in revised form

14.08.2019 Accepted 15.08.2019

Institute for Evolutionary Ecology, National Academy of Sciences ofUkraine, Lebedeva st., 37, Kyiv, 03143, Ukraine. Tel.: +38-097-245-99-18. E-mail: riburda@ukr.net, ser681@ukr.net

Burda, R I., & Koniakin, S. N. (2019). The non-native woody species of the flora of Ukraine: Introduction, naturalization and invasion. BiosystemsDiversity, 27(3), 276-290. doi:10.15421/011937

The process of invasion, naturalization, dispersion and invasive activity of non-native woody species in 5 regional floras, 5 urban floras and over 30 floras of the protected areas is discussed. It has been established that 182 non-native species out of 95 genera and 45 families are currently at different naturalization stages in the spontaneous flora of Ukraine. In terms of life-forms, they may be divided as follows: trees - 41%, shrubs - 35%, trees/shrubs - 15%, lianas - 9%. Most species spread both via seed dispersal and the vegetative way - 56%, using only seed dispersal - 42%, only the vegetative way - 2%. According to the preliminary rating of species-wise invasive activity, 71 species (40%) have naturalized completely, among them 20 invasive species, including 12 transformer species, 29 potentially invasive species, and 22 naturalized species which demonstrate no invasive tendencies. The initial stages of invasion of 111 species are as follows: introduction - 9%, survival - 36%, adaptation of the reproductive sphere - 28%, establishment, formative of local populations - 27%. In terms of dispersion, non-naturalized species are divided as follows: rare - 45%, local - 16%, sporadic - 6%, unique - 12%, spreading under control beyond collections and expositions in botanical gardens and arboretums - 21%. Twelve transformer species, the greatest threat to local diversity, are Acer negundo, Ailanthus altissima, Amorpha fruticosa, Bupleurum futicosum, Elaeagnus angustifolia, E. rhamnoides, Fraxinus ornus, Parthenocissus quinquefolia, P. vitacea, Robinia pseudoacacia, Rhamnus alaternus, Salix x blanda and 8 invasive proper species: Berberis aquifolia, Colutea orientalis, Daphne laureola, Prunus cerasifera, P. serotina, Quercus ilex, Viburnum tinus, Vitis vinifera. The list of alien species, most widespread in 50 regions of Europe, includes Robinia pseudoacacia (42 regions), Ailanthus altissima (40), Acer negundo (38), Prunus cerasus (34), Quercus rubra (34), Rosa rugosa (34), Prunus domestica (31). The mitigation of the impacts of these species on local biodiversity is possible via the restoration of local native plant communities, land use organization, strict selection of introduced species prior to the introduction, culture of planting management, preventive measures and extending awareness and sharing of information about plant invasions. Therefore, Ukraine's spontaneous flora is notable for the active process of naturalization of non-native woody species with considerable involvement of invasive alien species. This is the first and preliminary evaluation of the invasive activity of woody species in Ukraine's flora. The manifestations of the global tendency of increased involvement and invasive activity of alien woody species in domestic flora have been confirmed. These conclusions are also relevant for elaborating the system of preventive, containing and mitigating measures regarding plant invasions in Ukraine.

Keywords: invasive alien species; tree; shrub; liana; species invasive activity.

Introduction

It is hard to overestimate the role of woody plants in creating plant cover and nutrient cycling, transfer of energy and information in the biosphere. They are dominants of plant community complexes and nuclei of consortia in ecosystems. While improving the environment, humans have long tried to draw woody plants nearer, to use all their diversity in order to satisfy intellectual, aesthetic, recreational and utilitarian needs. Since the end of the 18th century, Ukraine has had the oldest introduction centers -arboretums "Alexandria" and "Sofiivka", and somewhat younger collections of world flora treasures - the Nikitsky Botanical Garden, the arboretum of the biosphere reserve "Askania Nova", academic and university botanical gardens. In addition to their decorative, educational, aesthetic value and favourable impact on the environment, the establishment of the reserves of alien woody species (so called exotic plants) is relevant for greenery building, forestries and protective forestation, etc. However, the accumulation of exotic plants in cultivation is a remote risk of further invasions, naturalization, some of them penetrate to natural plant communities, which causes changes in the composition, structure and functions of the latter and poses a threat to the local gene fund.

Information on the distribution of alien plant species in the world is dramatic. The Global Naturalized Alien Flora (GloNAF) database contains 13,168 species or 3.9% of the extant global flora of vascular

plants. These alien species have secondary regions (van Kleunen et al., 2015; Pysek et al., 2017a, b). At least 75% and 93% of the world's naturalized alien flora is grown in home and botanical gardens, respectively (van Kleunen et al., 2018). The role of evolutionary changes during biological invasions must be understood (Blackburn et al., 2014; Zenni et al., 2017). This is important for mitigating the impacts and threats of penetration of woody plants into the natural flora (Pollegioni et al., 2013; Hirsch et al., 2017; Gaskin et al., 2017).

The article starts the analysis of diversity of the woody species, introduced to Ukraine, in order to find those which have invaded native or native-related plant communities, formed stable self-restoring local populations in them, become naturalized, revealed the ability to invade, formed secondary ranges and their own ecological niches, become invasive; it aims to determine the level of invasive activity of alien woody species and cryptic (hidden) threats of non-native woody species at the initial stages of migration.

Materials and methods

Let us define the main terms used in this article. Non-native (adventive, non-indigenous, exotic) species are the species whose occurrence in a specific territory is not related to the processes of natural florogenesis. These have overcome the geographical barrier due to human activity. Among them, we

distinguish alien species - completely naturalized, which created secondary ranges, and non-naturalized species, which are undergoing the first stages of migration. Spontaneous species are those, the occurrence and distribution of which in a specific territory does not depend on humans.

There is no separate accessible database in Ukraine, related to a relevant group of vascular plants in human life - woody species, intentionally introduced into the culture. In order to form the total list and the volume of cultivated woody species, we have had to review current "Catalogues" of collection funds of the botanical gardens (Kondratyuk, 1988; Kokhno, 1997; Grevtsova, 2000; Solomakha, 2007; Kolisny-chenko et al., 2011) and arboretums (Kosenko, 2000). We also used some articles about live botanical collections and spontaneous dispersion of woody exotic species from collections and expositions within introduction centers. The invasive activity of alien species in Ukraine's spontaneous flora was studied beyond introduction centers in native and approximately native community complexes within protected areas (Ba-ranovski et al., 2016; Lykholat et al., 2017, 2018a, 2018b). Special attention was paid to studying urban floras. We also noted the results of evaluating the invasive activity in regional floras, floras of agrolandsca-pes and floras of the nature reserve fund of different zones (Burda, 2003, 2007, 2018). Only non-native and spontaneously spreading species of Ukraine's flora have been considered. Sometimes native species, specified in the mentioned articles, have not been included in the analyzed lists (Acerplatanoides L., A. tataricum L., Sambucus nigra L. and Staphylea pinnata L.).

Non-native woody species, found in Ukraine's flora, have been characterized by 6 categories, traditional for invasive botany. The degree of their invasive activity has been described in accordance to the categories, implemented by the European Botanical Gardens Consortium (Mayorov et al., 2013). These are three categories of alien species: transformer species - a group of species, changing the conditions and nature of ecosystems in their larger area, they pose a threat to environmental safety of the region; invasive species - naturalized species, capable of having great amounts of progeny, due to which they spread rapidly across large distances from their paternal plants, are capable of invading and persisting in large areas, have invaded local community complexes, they affect local species, their communities and ecosystems; potentially invasive species - also naturalized species, which may become invasive upon future increase in their number, they do not have any visible impacts on local diversity; species, which have become naturalized, but have not demonstrated invasive activity. Additional consideration has been given to non-naturalized species in the process of naturalization, which are sometimes restored via seed dispersal or in a vegetative way near the area of native species, but are not capable of further invasion yet (a small number of diaspores, no functional connections). These plants do not demonstrate expressed tendencies of naturalization, their presence depends on constant stable input of diaspores from cultivated plants, though they do reproduce in some cases.

The species of the abovementioned groups are characterized in accordance to traditional classifications, used by Ukrainian authors in the abovementioned articles, which requires some elaboration (Didukh et al., 2000). The category "degree of naturalization" has 6 descriptors: аgrio-phyte - naturalized species which invaded native and native-related habitats; hemiagriophyte - species naturalized mostly in habitats which are close to native or degraded ones; epoecophyte - species naturalized only in anthropogenically transformed habitats; ergasiophyte - species intentionally introduced by humans which sometimes escape the cultivation limits, but are usually in the immediate proximity to the place of cultivation; colonophyte, non-naturalized species, unstable element of flora, related to cultivation, somewhat adjusted to new conditions of survival, capable of periodic formation of seeds or reproduction in the vegetative way; ephemerophyte - also non-naturalized species, unstable element of flora, which periodically occur in the places of cultivation of introduced woody species or among secondary habitats and vanish with time, as they are accidental migrants. The following species categories are distinguished in terms of occurrence: common species, widely and abundantly spread in the whole territory; local species, known in the whole territory, but abundant only in some locations; sporadic species, which occur frequently, but with low abundance, rare species, which are

present in 3-5 (7) locations, unique species, found in 1-2 locations, and controlled species, spread via seed dispersal or in the vegetative way beyond collections and expositions within the botanical gardens and arboretums. Three categories have been distinguished in terms of life-forms: a tree - a perennial woody plant with one well-formed stem and a crown, formed by side branches; a shrub - a perennial woody plant with multiple stems coming out of one source, the habitus does not have one well-formed main stem, and a liana - a perennial woody plant with a long stem, not capable of maintaining the vertical position without any additional support.

The names of the species are presented according to "The Plant List" database. The authors of all the names of the species, mentioned in the text, are given in Tables 5 and 6. The volumes of families of Mag-noliophyta (Angiosperms) and the sequence of their location in Tables 2, 5 and 6 are in accordance to "Synopsis of Families and Orders of Angiosperms of the Flora of Ukraine" (Mosyakin, 2013).

The non-native woody species of the flora of Ukraine, escaping from cultivation in botanical gardens and arboretums

Over 50 centers ofplant introduction are officially registered in Ukraine (Cherevchenko, 2011), and 10 of them are over 130 years old (Table 1). In addition, there are private commercial centers which import decorative plants, including trees, shrubs, and lianas, for sale. The attempts at finding and summarizing the diversity of live collections of the botanical gardens and arboretums have failed due to the absence of any unified database in free access mode. Our centers of plant introduction are mainly located in the temperate zone. They have been working for many years within creative cooperation of the Council of Botanical Gardens and Arboretums of Ukraine, exchanging their methods and seeds.

Taking into consideration these two circumstances, it becomes clear that the collections of woody plants in the introduction centers are somewhat duplicated. It would be reasonable to consider the richest collections of woody plants, collected in the arboretum "Sofivka", the M M Hryshko National Botanical Garden, the O. V. Fomin Botanical Garden, and the arboretum of the Nititsky Botanical Garden, which is located in conditions, approximated to those of the eastern Mediterranean (Table 1).

However, it is common knowledge that "every garden is famous for its own collections". For instance, the National Arboretum "Sofiiv-ka" is known for its rich collections of the genera Corylus - 28 species, Fagus - 21, Picea - 41, 100 species of lianas and over 320 varieties of roses (Kosenko, 2000). The O. V. Fomin Botanical Garden collected the most abundant generic complexes of Juniperus, Pinus, Cotoneaster, Forsythia, Lonicera, Magnolia, Rhododendron, Spiraea (Solomakha, 2007) and the Nikitsky Botanical Garden owns luxurient collections of genera Cupressus, Berberis, Cotoneaster, Lonicera, Philadelphus, Quer-cus, Spiraea, Thuja and Viburnum (Plugatar et al., 2015). As this statement reflects reality, it is noteworthy that the volume of the collections has neither changed much nor decreased for many years. For instance, the statistics for the arboretum of the Nikitsky Botanical Garden as of 1970 stated the presence of 1,704 species, including 689 trees, 879 shrubs, 136 lianas (Kormilitsin & Golubeva, 1970). In 1971 the collection of the National Botanical Garden contained 2,054 species, including Angiospermae - 1,884 species and Gymnopsermae - 170 species (Rubtsov & Gordienko, 1971). The collections, exceeding 1,000 woody species, have been collected in each of 8 introduction centers. It was recorded that in 1994 there were 1,655 species, 32 varieties, 547 forms of introduced trees, shrubs, and lianas (Kokhno, 1994). It should be also noted that the ratio between trees and shrubs (different life-forms) is not significant. It may be balanced (arboretum "Alexandria", the Botanical Garden of the National University of Dnipro, the Donetsk Botanical Gardens), or the number of species whose life-form is a shrub may exceed the number of species belonging to the tree life-form (arboretum "Sofiivka", the Kryvyi Rih Botanical Garden), or the ratio between the species with these life-forms is reverse (the Botanical Garden of Odesa University). Thus, the volumes of the richest modern collections and their age permit the assumption that these collections may be a source of dispersing non-native woody species into the domestic flora, completely established quite a long time ago.

Table 1

The collection funds of woody plants in the leading Ukrainian centers of plant introduction

Center of plant introduction Year of Source Number of species

foundation total tree* shrub* liana*

The arboretum "Alexandria", NAS of Ukraine 1793 Galkin et al., 2017 1282 538 680 64

The National Arboretum "Sofiivka", NAS of Ukraine 1796 Kosenko, 2000 2400 800 1600

Botanical Garden of V. N. Karazin Kharkiv National University 1804 Alokhin et al., 2019 781 - - -

The Nikitsky Botanical Garden 1812 Plugatar et al., 2015 1690 - - -

The arboretum "Trostyanets", NAS of Ukraine 1834 Medvedyev & Ilyenko, 2018 623 - - -

O. V. Fomin Botanical Garden of Taras Shevchenko National Universi- 1839 Solomakha, 2007 1828 923

ty of Kyiv - -

Botanical Garden of Ivan Franko L'viv National University 1852 Prokopiv, 2004 787 - - -

Botanical Garden of Odesa of I. I. Mechnikov National University 1867 Filatova et al., 2014; Slyusarenko et al., 2017 1025 611 341 73

Botanical Garden of Yuriy Fedkovych Chernivtsi National University 1877 Cherevchenko, 2011 1150 - - -

The arboretum of F. J. E. Falz-Fein "Askania Nova" Nature Biosphere 1887 Rubtsov et al., 2012 1114

Reserve, NAAS of Ukraine

Botanical Garden of National University of Life and Environmental Sciences of Ukraine, Kyiv 1928 Kolisnychenko et al., 2011 604 - - -

Botanical Garden of Oles Honchar Dnipro National University 1933 Kvasha et al., 2010 389 159 147 83

M. M. Hryshko National Botanical Garden, NAS of Ukraine 1935 Kokhno, 1997 1025 - - -

The Donetsk Botanical Garden, NAS of Ukraine 1964 Kondratyuk, 1988 868 457 411 -

The Kryvyi Rih Botanical Garden, NAS of Ukraine 1980 Grevtsova, 2000 661 245 393 23

Note: "-" - marks: data is absent in source.

Only two published "Catalogues" of collection funds present systematized information about spontaneous dispersion ofwoody species from collections and expositions (Kondratyuk, 1988; Solomakha, 2007) (Table 2). Some fragments of the data are stated in special publications. After twenty years of introductory testing, 15 woody species, including 8 non-native cultivated ones, had natural regrowth in the Donetsk Botanical Garden of NAS of Ukraine (Kondratyuk, 1988). After 10 years, 54 species, including 13 aboriginal species, were found to be spontaneously distributing; 48 species had stable natural regrowth (Burda et al., 1998). It was noted

Table 2

The list of non-native species, spontaneously distributing in the centers of plant introduction

that the species of genus Clematis, in particular, C. vitalba, spread far from the place of initial introduction, while remaining in the garden. Later it was additionally reported that 19 non-native species had spontaneously spread within the boundaries of the garden (Eremenko & Ostapko, 2011, 2014). The total number amounted to 67 species. The location of C. vitalba was found 4 km from the garden and was interpreted as "escaping" from its collections (Ostapko et al., 2013). We place this assumption in question as C. vitalba was found on the roadside between the cities of Donetsk and Makiivka, which bordered summer cottages.

Family Taxon* Life-form Spread Center of introduction

Pinophyta

Pinaceae *Larix decidua MU tree seed Donetsk Botanical Garden

Cupressaceae *Thuja occidentalis L. 1ree/shrub seed Botanical Garden of Ivan Franko National University of L'viv

Magnoliophyta

Berberidaceae Berberidaceae Berberidaceae Berberis aquifolium Pursh. *B. thunbergii DC. B. vulgaris L. shrub shrub shrub seed, vegetative way seed, vegetative way seed, vegetative way Donetsk Botanical Garden Arboretum "Alexandria" Donetsk Botanical Garden

Ranunculaceae Ranunculaceae Ranunculaceae "Clematis gouriana Roxb. ex DC. *C. jackmanii T. Moore. *C. ligusticifolia Nutt liana liana liana seed, vegetative way seed, vegetative way seed, vegetative way Donetsk Botanical Garden Donetsk Botanical Garden Donetsk Botanical Garden

Ranunculaceae Ranunculaceae *C. tangutica (Maxim.) Korsh. C. vitalba L. liana liana seed, vegetative way seed, vegetative way Donetsk Botanical Garden Arboretum "Alexandria", Donetsk Botanical Garden

Ranunculaceae *C. viticella L. liana seed, vegetative way Arboretum "Alexandria"

Hamamelidaceae *Parrotia subaequalis (Hung T. Chang) R M. Hao & H. T. Wei tree/shrub seed M. M. Hryshko National Botanical Garden

Grossulariaceae *Ribes americanum Mill. shrub seed Donetsk Botanical Garden

Grossulariaceae R. aureum Pursh shrub seed Donetsk Botanical Garden

Grossulariaceae R europaea (L.) Mill. shrub seed Donetsk Botanical Garden

Grossulariaceae R. rubrum L. shrub seed Donetsk Botanical Garden

Grossulariaceae R. spicatum Robson shrub seed Donetsk Botanical Garden

Vitaceae Vitaceae "Ampelopsis aconitifolia Bunge *A. delavayana var. glabra (Diels & Gilg) C. L. Li liana liana seed, vegetative way seed, vegetative way Donetsk Botanical Garden Donetsk Botanical Garden

Vitaceae Vitaceae Vitaceae *A. bodinieri (H. Lev. & Vaniot) Rehder *A. glandulosa var. brevipedunculata (Maxim.) Momiy Parthenocissus quinquefolia (L.) Planch. liana liana liana seed, vegetative way seed, vegetative way seed, vegetative way Donetsk Botanical Garden Donetsk Botanical Garden Botanical Garden of Oles Honchar National University of Dnipro, Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Vitaceae Vitaceae Vitaceae P. vitacea (Knerr) Hitchc. *Vitis coignetiae Bull. ex Planch V. vinifera L. liana liana liana seed, vegetative way seed, vegetative way seed, vegetative way Donetsk Botanical Garden Donetsk Botanical Garden Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Fabaceae Fabaceae Amorpha fruticosa L. Caragana arborescens Lam. shrub shrub seed, vegetative way seed, vegetative way Arboretum "Alexandria", Botanical Garden of Oles Honchar National University of Dnipro, Donetsk Botanical Garden Arboretum "Alexandria", Donetsk Botanical Garden

Fabaceae Gleditsia triacanthos L. tree seed Botanical Garden of Oles Honchar National University of Dnipro, Donetsk Botanical Garden

Fabaceae Gymnocladus dioica (L.) K. Koch tree seed Arboretum "Alexandria", Donetsk Botanical Garden

Family Taxon* Life-form Spread Center of introduction

Fabaceae Halimodendron halodendron (Pall.) Voss shrub seed Donetsk Botanical Garden Arboretum "Alexandria", Donetsk Botanical Garden,

Fabaceae Robinia pseudoacacia L. tree seed, vegetative way O. V. Fomin Botanical Garden, Botanical Garden of Ivan Franko National University of L'viv

Fabaceae R viscosa Vent. tree seed, vegetative way Arboretum "Alexandria"

Rosaceae Cerasus tomentosa (Thunb.) Wall. ex T. T. Yu & C. L. Li shrub seed Donetsk Botanical Garden

Rosaceae Crataegus submollis Sarg. shrub seed Arboretum "Alexandria"

Rosaceae Crataegus sp. shrub seed Donetsk Botanical Garden

Rosaceae Malus domestica Borkh. tree seed, vegetative way Donetsk Botanical Garden

Rosaceae Mespilus germanica L. shrub seed, vegetative way Donetsk Botanical Garden

Rosaceae Physocarpus opulifolius (L.) Maxim. shrub seed, vegetative way Arboretum "Alexandria"

Rosaceae Prunus armeniaca L. tree seed, vegetative way Donetsk Botanical Garden

Rosaceae P. cerasus L. tree seed, vegetative way Donetsk Botanical Garden

Rosaceae P. cerasifera Ehrh. tree/shrub seed, vegetative way Arboretum "Alexandria"

Rosaceae P. mahaleb L. shrub seed Donetsk Botanical Garden

Rosaceae P. serotina Ehrh. tree/shrub seed, vegetative way Donetsk Botanical Garden

Rosaceae Rosa sp. shrub seed, vegetative way Donetsk Botanical Garden

Rosaceae *Rubus odoratus L. shrub seed, vegetative way Arboretum "Alexandria"

Rosaceae Sorbaria sorbifolia (L.) A. Braun shrub seed, vegetative way Arboretum "Alexandria", Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Rosaceae Spiraea x billardii Hérin shrub seed, vegetative way Donetsk Botanical Garden

Rosaceae S. chamaedryfolia L. shrub seed, vegetative way Arboretum "Alexandria"

Rosaceae S. salicifolia L. shrub seed, vegetative way Arboretum "Alexandria"

Elaeagnaceae Elaeagnus angustifolia L. tree/shrub seed, vegetative way Donetsk Botanical Garden

Elaeagnaceae E. rhamnoides (L.) A. Nelson shrub seed, vegetative way Donetsk Botanical Garden

Elaeagnaceae E. umbellata Thunb. tree seed, vegetative way Arboretum "Alexandria"

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Ulmaceae Celtis australis L. tree seed M. M. Hryshko National Botanical Garden

Ulmaceae C. occidentalis L. tree seed Donetsk Botanical Garden

Ulmaceae Ulmus pumila L. tree seed, vegetative way Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Moraceae Morus alba L. tree seed Donetsk Botanical Garden

Fagaceae Quercus rubra L. tree seed Arboretum "Alexandria", Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Fagaceae *Q. macranthera Fisch. & C.A Mey. ex Hohen. tree seed M. M. Hryshko National Botanical Garden

Juglandaceae Juglans mandshurica Maxim. tree seed Donetsk Botanical Garden

Juglandaceae J. nigra L. tree seed Arboretum "Alexandria"

Juglandaceae J. regia L. tree seed Arboretum "Alexandria", Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Celastraceae Celastrus angulata Maxim. liana seed, vegetative way Donetsk Botanical Garden

Celastraceae C. flagellaris Rupr. liana seed Donetsk Botanical Garden

Celastraceae C. orbiculatus Thunb. liana seed Arboretum "Alexandria", Donetsk Botanical Garden

Salicaceae Populus balsamifera L. tree seed, vegetative way Donetsk Botanical Garden

Salicaceae P. x canadensis Moench tree seed, vegetative way Donetsk Botanical Garden

Salicaceae P. trichocarpa Torr. & A. Gray ex Hook. tree seed, vegetative way Donetsk Botanical Garden

Anacardiaceae Rhus glabra L. tree/shrub seed, vegetative way Botanical Garden of Ivan Franko National University of L'viv

Anacardiaceae *R typhina L. tree vegetative way Arboretum "Alexandria", Donetsk Botanical Garden, Botanical Garden of Ivan Franko National University of L'viv

Anacardiaceae *R. typhina 'Laciniata' tree vegetative way Arboretum "Alexandria"

Anacardiaceae Toxicodendron pubescens Mill. shrub seed, vegetative way Arboretum "Alexandria"

Anacardiaceae T. radicans (L.) O. Kuntze shrub seed, vegetative way Donetsk Botanical Garden Arboretum "Alexandria", Botanical Garden of Oles Hon-

Aceraceae Acer negundo L. tree seed char National University of Dnipro, Donetsk Botanical Garden, O. V. Fomin Botanical Garden, Botanical Garden of Ivan Franko National University of L'viv

Aceraceae *A. platanoides L. f. atropurpurea 'Krimson King A. saccharinum L. tree seed Donetsk Botanical Garden

Aceraceae tree seed Donetsk Botanical Garden

Hippocasta naceae Aesculus hippocastanum L. tree seed Arboretum "Alexandria", Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Simaroubaceae Ailanthus altissima (Mill.) Swingle tree seed, vegetative way Arboretum "Alexandria", Botanical Garden of Ivan Franko National University of L'viv, Donetsk Botanical Garden

Cornaceae Cornus alba L. shrub seed, vegetative way Donetsk Botanical Garden

Hydrangeaceae Philadelphus coronarius L. shrub seed, vegetative way Donetsk Botanical Garden

Solanaceae Lycium barbarum L. shrub seed, vegetative way Arboretum "Alexandria"

Oleaceae Fraxinus ornus L. tree seed Nikitsky Botanical Garden

Oleaceae F. pennsylvanica Marshall tree seed Donetsk Botanical Garden

Oleaceae Syringa vulgaris L. shrub seed, vegetative way Arboretum "Alexandria", Donetsk Botanical Garden, O. V. Fomin Botanical Garden

Bignoniaceae *Campsis grandiflora (Thunb.) K. Schum. liana vegetative way Arboretum "Alexandria"

Bignoniaceae *C. radicans (L.) Seem. liana vegetative way Arboretum "Alexandria"

Caprifoliaceae Lonicera caerulea L. shrub seed Donetsk Botanical Garden

Caprifoliaceae L. caprifolium L. shrub seed Arboretum "Alexandria"

Caprifoliaceae L. tatarica L. shrub seed Arboretum "Alexandria", Donetsk Botanical Garden

Caprifoliaceae Symphoricarpos albus (L.) S.F. Blake shrub seed, vegetative way Donetsk Botanical Garden

Apiaceae Bupleurum fruticosum L. shrub seed, egetative way Nikitsky Botanical Garden

Aracaceae *Aralia elata (Miq.) Seem. shrub seed, vegetative way Botanical Garden of Ivan Franko National University of L'viv

Note: * - the species whose dispersion is known only beyond the framework of collections and expositions in botanical gardens and arboretums.

The inspection of spontaneous spreading of intentionally introduced species from O. V. Fomin Botanical Garden of Taras Shevchenko National University of Kyiv resulted in issuing "The preliminary list of the species of the O. V. Fomin Botanical Garden, spreading beyond their place of cultivation" (Solomakha, 2007). This list contains 245 species, including 6 species - trees (Acer negundo, Aesculus hippocastanum, Juglans regia, Quercus rubra, Robinia pseudoacacia and Ulmus pumila), 2 species - shrubs (Sorbaria sorbifolia and Syringa vulgaris), 2 species -lianas (Parthenocissus quinquefolia and Vitis vinifera).

45 woody species, including 13 aboriginal ones, are spontaneously spreading using seed dispersal and vegetative reproduction within the arboretum "Alexandria". Some non-native species do not reproduce via seed dispersal, but reproduce actively via vegetative reproduction: Camp-sis grcmdifom, C. radicans, Rhus typhina, R typhina 'Laciniata', Rubus odoratus, Syringa vulgaris, Toxicodendronpubescens (Doyko et al., 2014).

Woody plants, capable of reproducing spontaneously via either seed dispersal or vegetative reproduction, were found among intentionally introduced plants in the Botanical Garden of the Ivan Franko L'viv National University. These were four species of trees (Acer negundo, Ailanthus altissima, Robinia pseudoacacia and Thuja occidentalis) and three species of shrubs (Aralia elata, Rhus glabra, R. typhina) (Borsu-kevych & Prokopiv, 2014). Among spontaneous species of the flora at the Botanical Garden of Oles Honchar National University of Dnipro, there are four known woody species (Acer negundo, Amorpha frutico-sa, Gleditsia triacanthos and Parthenocissus quinquefolia). While analyzing the results of long-term observations, the authors noted the tendency towards the increase in their number and degree of naturalization within the garden (Tarasov et al., 1998). The "Catalogue" of plants of the M. M. Hryshko National Botanical Garden (Kokhno, 1997) does not have any data about seed dispersal. Occasionally there were some publications about seed dispersal of woody plants from its collections and expositions. Shinder (2013) considers the botanic-geographical area "Caucasus" to be the source of distribution of Caucasian species into anthropogenically altered locations within the garden. Celtis australis escaped the boundaries of the "Caucasus" area. Some Caucasian species have established stable spontaneous cenopopulations in the created simulated communities of this exposition. For instance, a rare species Quercus macranthera dominates in the growing stock of the planted dry oakery, constantly forming abundant natural self-sown plants. Periodic formation of abundant natural self-seeding of Parrotia subaequalis was recorded (Doroshenko et al., 2013).

The naturalization of plants in the oldest arboretums of the Nikitsky Botanical Garden must have occurred at the beginning of the 20th century. Without highlighting the collections ofthe garden, Stankov (19241925) noted the remarkable presence of the cultural element of the Mediterranean flora in the plant cover of the South Coast of Crimea. He mentioned 55 non-native species, which added unique Mediterranean charm to the landscape. S. S. Stankov observed the naturalization of the following 22 species in the Crimea personally: Ailanthus altissi-ma, Berberis aquifolium, Bupleurum fruticosum, Cercis siliquastrum, Clematis flammula, Cydonia oblonga, Elaeagnus angustifolia, E. rhamnoides, Ficus carica, Fraxinus ornus, Ilex aquifolium, Laburnum anagyroides, Madura pomifera, Morus alba, Olea europaea, Prunus armeniaca, P. cerasifera, P. cerasus, P. domestica, P. domestica subsp. insititia, Rhamnus alaternus and Spartium junceum. This list was later supplemented with Laurus nobilis, Lonicera caprifolium, Viburnum ti-nus, and S. S. Stankov doubted their going out of cultivation (Bagriko-va, 2013). Obviously, we do not have any evidential materials to prove that all of these species have spread within the South Coast of Crimea after escaping the Nikitsky Botanical Garden. On the one hand, this institution had supplied many exotic trees, shrubs, and lianas to the market, being the only center of plant introduction in the Crimea, on the other hand, in different times, even with active dendrological collections of the Garden, amateurs of the garden art brought decorative woody plants to the Crimea, which could have spread beyond the locations of cultivation in different corners of the South Coast of Crimea. There is only one direct statement about two species (Bupleurum fruticosum, Fraxinus ornus), spreading within the nature reserve "Mys Martian" from the adjacent plant communities ofthe Nikitsky Botanical Garden (Bagrikova et al., 2014).

Therefore, 92 non-native woody species out of 49 genera and 28 families are spreading in collection areas and expositions or beyond them, but not escaping the boundaries of introduction centers in 6 abo-vementioned botanical gardens and arboretums. Table 2 presents 6 families, covering 5 or more species: Rosaceae - 17, Vitaceae - 8, Fabaceae - 7, Ranunculaceae - 6, Anacardiaceae - 5 and Grossulaceae - 5. Spontaneous distribution of most species has been noted only in one garden (73), in 5 gardens there is a noted distribution of only Acer ne-gundo, and in 4 - Robinia pseudoacacia. Half of these species have long been naturalized and become common elements of spontaneous flora. However, spontaneous distribution of 23 species is known only in controlled conditions of the botanical gardens and arboretums. For instance, these are Aralia elata, Parrotia subaequalis, Vitis coignetiae, Rhus typhina, R. typhina 'Laciniata', Rubus odoratus. Thus, the assumption, expressed by us regarding the centers of initial introduction of plants in Ukraine as the sources of distributing non-native woody species, has not been confirmed.

The non-native woody species in the urban flora of Ukraine

The degree of invasive activity of alien woody species in the framework of urban flora plays an important role in determining their part in the spontaneous flora of Ukraine. The main centers of naturalization of non-native species are cities with their suburbs, which unite parks, public gardens, green zone plantings, botanical gardens and arboretums. We have analyzed the lists of four recently thoroughly studied urban floras of such cities as Kyiv (Mosyakin & Yavorska, 2002), Kryvyi Rih (Kuchrevskyi & Shol, 2009), Uzhhorod (Protopopova & Shevera, 2002), Kharkiv (Zvyagintseva, 2015), and combined floras of Slo-viansk, Donetsk, Luhansk, and Mariupol, which have formed within industrial Donbas (Burda, 1997). These cities are in different regions of Ukraine and differ in the degree of environmental urbanization. Concluding the consideration of urban floras, we would like to highlight that the lists of alien species coincide in the main part (Table 3). This fact allowed us to combine woody species into a unified list to determine the degree of their naturalization. The authors of urban floras often added aboriginal species to them. They are absent in the combined list: Betula pendula Roth, B. pubescens Ehrh., Lonicera xylosteum L., Rubus idaeus L., Sambucus racemosa L., Sorbus aucuparia L., Tilia platy-phyllos Scop., Viburnum lantana L., etc. Consideration was also given to the list of flora in Chernihiv, where the degree of naturalization was not indicated (Zavyalova, 2010). This urban flora contains about 50 non-native woody species, and the combined list of urban floras of Ukraine was added Phellodendron amurense and Spiraea x vanhouttei therefrom. The total list of alien woody species of 6 urban floras of Ukraine contains 80 species, 1 subspecies, and 1 hybrid (Table 3, 4). The picture of naturalization of non-native species in urban floras is very rich. They differ in terms of structure, degree of naturalization, species composition, and all of these are mutually related. The richest floras are noted for Kyiv, Kharkiv, Chernihiv, and the urban flora of Kryvyi Rih is not far behind them.

The degree of naturalization is also different. The only species, evaluated in all the floras as agriophyte, is Salix x blanda, and the one close to it is Acer negundo. The following species occur in one urban flora only: Ficus carica, Rubus macrophyllus, Toxicodendron radicans, Vitis labrusca. Agriophytes are concentrated in Kharkiv, hemiagriophytes and colonophytes - in Kyiv, ergasiophytes - in Kharkiv and Kryvyi Rih. Thus, urban floras have a high degree of naturalization of non-native woody species.

The course of invasion of non-native woody species in the regional floras and in the floras of protected areas

The regional studies, conducted by the specialists, headed by V. V. Protopopova, defined the following transformer species in the flora of 5 regions: Polissia - Robinia pseudoacacia (Protopopova et al., 2015); Bukovyna Cis-Carpathian region - Acer negundo, Robinia pseudoacacia (Protopopova et al., 2010); Middle Dnipro region - Acer negundo, Amorpha fruticosa, Robinia pseudoacacia (Protopopova et al.,

2014); North Black Sea region - Amorpha fruticosa, Elaeagnus angus-tifolia and Salix x blanda (Protopopova et al., 2009); South Coast of Crimea - Ailanthus altissima, Bupleurum fruticosum, Fraxinus ornus and Rhamnus alaternus (Protopopova et al., 2012). The authors distinguished 39 invasive species, including 13 woody species, in the flora of the Crimea. Robinia pseudoacacia is mentioned in four regions, Acer negundo, Amorpha fruticosa - in three, Elaeagnus angustifolia - in two and Salix x blanda - in only one region. According to our observations, the features of transformer species in the floras of 14 territories, which are subject to special protection in the Forest Steppe, are remarkable for 7 species: Acer negundo, Elaeagnus angustifolia, E. rhamnoides, Par-thenocissus quinquefolia, P. vitacea, Robinia pseudoacacia and Salix x blanda. They occurred in at least 9 out of 14 floras, except for E. rham-noides, noted in 3 floras only (Burda et al., 2015b). The study of alien flora species in 30 territories of the nature reserve fund in different natu-

ral zones revealed the same 7 species of transformers.These are three trees: Acer negundo, Robinia pseudoacacia and Salix x blanda; 2 shrubs: Elaeagnus angustifolia and E. rhamnoides and 2 lianas: Parthenocissus quinquefolia and P. vitacea (Burda et al., 2015a). While determining the "transformer species" status, the main feature is stated as the ability to change the ecosystem completely. Five invasive species proper have been distinguished in the same place: Amorpha fruticosa, Morus alba, Prunus cerasus, P. semtina and P. virginiana. The floras of the nature reserve fund contain 16 woody species, which have naturalized (according to the scheme, accepted in this article, this notion is close to the term, further used by us, - "potentially invasive species"). In addition to the mentioned species, let us mention 31 more species, related to the ones which reproduce spontaneously, sporadically or in a single way in the reserve areas.

Table 3

The degree of naturalization and the occurrence of non-native woody species in some urban floras of Ukraine

Species Kyiv Kryvyi Rih Kharkiv Uzhhorod Donbas Flora Degree of naturalization

Salix x blanda agriophyte agriophyte agriophyte agriophyte agriophyte 5 5 agriophytes

Acer negundo agriophyte hemiagriophyte agriophyte hemiagriophyte agriophyte 5 3 agriophytes: 2 hemiagriophytes

Berberis aquifolia agriophyte ergasiophyte agriophyte ergasiophyte 0 4 2 agriophytes: 2 ergasiophytes

Prunus cerasifera 0 ergasiophyte agriophyte agriophyte 0 3 2 agriophytes: 1 ergasiophyte

Amorpha fruticosa agriophyte hemiagriophyte ergasiophyte ergasiophyte ergasiophyte 5 1 agriophyte: 1 hemiagriophyte: 3 ergasiophytes

Elaeagnus angustifolia 0 hemiagriophyte agriophyte ergasiophyte epoecophyte 4 1 agriophyte: 1 hemiagriophyte: 1 epoecophyte: 1 ergasiophyte

Prunus serotina agriophyte ergasiophyte 0 0 0 2 1 agriophyte: 1 ergasiophyte

Parthenocissus vitacea hemiagriophyte 0 agriophyte 0 0 2 1 agriophyte: 1 hemiagriophyte

Cydonia oblonga 0 0 agriophyte 0 0 1 1 agriophyte

Prunus domestica subsp. insititia 0 0 0 agriophyte 0 1 1 agriophyte

Robinia pseudoacacia hemiagriophyte hemiagriophyte ergasiophyte ergasiophyte ergasiophyte 5 2 hemiagriophytes: 3 ergasiophytes

Lycium barbarum hemiagriophyte hemiagriophyte epoecophyte epoecophyte epoecophyte 5 2 hemiagriophytes: 3 epoecophytes

Ulmus pumila hemiagriophyte hemiagriophyte ergasiophyte 0 0 3 2 hemiagriophytes: 1 ergasiophyte

Caragana arborescens hemiagriophyte ergasiophyte ergasiophyte 0 ergasiophyte 4 1 hemiagriophyte: 3 ergasiophytes

Acer saccharinum hemiagriophyte ergasiophyte ergasiophyte 0 0 3 1 hemiagriophyte: 2 ergasiophytes

Quercus rubra hemiagriophyte ergasiophyte ergasiophyte 0 0 3 1 hemiagriophyte: 2 ergasiophytes

Ptelea trifoliata hemiagriophyte ergasiophyte 0 0 0 2 1 hemiagriophyte: 1 ergasiophyte

Elaeagnus commutata 0 hemiagriophyte 0 0 0 1 1 hemiagriophyte

Morus alba epoecophyte ergasiophyte colonophyte 0 epoecophyte 4 2 epoecophytes: 1 ergasiophyte: 1 colonophyte

Gleditsia triacanthos colonophyte ergasiophyte epoecophyte ergasiophyte 0 4 1 epoecophyte: 2 ergasiophytes: 1 colonophyte

Fraxinus pennsylvanica epoecophyte epoecophyte ergasiophyte 0 0 3 2 epoecophytes: 1 ergasiophyte

Elaeagnus rhamnoides 0 epoecophyte epoecophyte ergasiophyte 0 3 2 epoecophytes: 1 ergasiophyte

Lonicera tatarica colonophyte epoecophyte epoecophyte 0 0 3 2 epoecophytes: 1 colonophyte

Populus deltoides colonophyte epoecophyte epoecophyte 0 0 3 2 epoecophytes: 1 colonophyte

Parthenocissus quinquefolia colonophyte epoecophyte 0 ergasiophyte 0 3 1 epoecophyte: 1 ergasiophyte: 1 colonophyte

Ailanthus altissima epoecophyte colonophyte ergasiophyte 0 0 3 1 epoecophyte: 1 ergasiophyte: 1 colonophyte

Spiraea salicifolia epoecophyte 0 ergasiophyte 0 0 2 1 epoecophyte: 1 ergasiophyte

Symphoricarpos albus colonophyte 0 epoecophyte 0 0 2 1 epoecophyte: 1 colonophyte

Parthenocissus tricuspidata 0 0 epoecophyte 0 0 1 1 epoecophyte

Cornus sericea 0 0 epoecophyte 0 0 1 1 epoecophyte

Juglans regia ephemerophyte ergasiophyte ergasiophyte ergasiophyte 0 4 3 ergasiophytes: 1 ephemerophyte

Prunus armeniaca colonophyte ergasiophyte ergasiophyte 0 ergasiophyte 4 3 ergasiophytes: 1 colonophyte

P. cerasus colonophyte ergasiophyte ergasiophyte 0 ergasiophyte 4 3 ergasiophytes: 1 colonophyte

Syringa vulgaris colonophyte colonophyte ergasiophyte ergasiophyte 0 4 2 ergasiophytes: 2 colonophyte

Vitis vinifera ephemerophyte ergasiophyte ergasiophyte 0 ergasiophyte 4 3 ergasiophytes: 1 ephemerophyte

Aesculus hippocastanum 0 ergasiophyte ergasiophyte ergasiophyte 0 3 3 ergasiophytes

Ribes uva-crispa 0 ergasiophyte ergasiophyte 0 ergasiophyte 3 3 ergasiophytes

Malus domestica colonophyte ergasiophyte ergasiophyte 0 0 3 2 ergasiophytes: 1 colonophyte

Prunus mahaleb colonophyte ergasiophyte 0 0 ergasiophyte 3 2 ergasiophytes: 1 colonophyte

Catalpa bignonioides 0 0 ergasiophyte ergasiophyte 0 2 2 ergasiophytes

Juglans nigra 0 ergasiophyte ergasiophyte 0 0 2 2 ergasiophytes

Populus bolleana 0 ergasiophyte ergasiophyte 0 0 2 2 ergasiophytes

P. nigra var. italica 0 ergasiophyte ergasiophyte 0 0 2 2 ergasiophytes

Aronia melanocarpa colonophyte 0 ergasiophyte 0 0 2 1 ergasiophyte: 1 colonophyte

Malus baccata colonophyte 0 ergasiophyte 0 0 2 1 ergasiophyte: 1 colonophyte

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Celtis occidentalis colonophyte ergasiophyte 0 0 0 2 1 ergasiophyte: 1 colonophyte

Lonicera caprifolium colonophyte 0 0 ergasiophyte 0 2 1 ergasiophyte: 1 colonophyte

Rhus typhina 0 colonophyte ergasiophyte 0 0 2 1 ergasiophyte: 1 colonophyte

Aesculus flava 0 0 ergasiophyte 0 0 1 1 ergasiophyte

Castanea sativa 0 0 0 ergasiophyte 0 1 1 ergasiophyte

Crataegus sanguinea 0 0 ergasiophyte 0 0 1 1 ergasiophyte

Juglans cinerea 0 ergasiophyte 0 0 0 1 1 ergasiophyte

Populus suaveolens 0 0 ergasiophyte 0 0 1 1 ergasiophyte

Persica vulgaris 0 ergasiophyte 0 0 0 1 1 ergasiophyte

Prunus virginiana 0 0 ergasiophyte 0 0 1 1 ergasiophyte

Species_Kyiv_Kryvyi Rih_Kharkiv_Uzhhorod_Donbas Flora_Degree of naturalization

P. domestica 0 ergasiophyte 0

Tilia americana 0 0 ergasiophyte

Colutea arborescens 0 ergasiophyte 0

Cornus alba 0 ergasiophyte 0

Philadelphus coronarius 0 0 ergasiophyte

Salix babylonica 0 0 ergasiophyte

Ribes aureum 0 ergasiophyte 0

R. rubrum 0 0 0

Rosa rugosa colonophyte 0 ergasiophyte

Robinia neomexicana 0 0 ergasiophyte

Robinia viscosa 0 ergasiophyte 0

Juglans mandshurica colonophyte 0 0

Amelanchier ovalis colonophyte 0 0

Berberis thunbergii colonophyte 0 0

B. vulgaris colonophyte 0 0 Populus balsamifera colonophyte 0 0 Quercus palustris colonophyte 0 0 Sorbaria sorbifolia colonophyte 0 0 Spiraea douglasii colonophyte 0 0 Physocarpus opulifolius colonophyte 0 0 Cotoneaster melanocarpus colonophyte 0 0 Clematis jackmanii colonophyte 0 0

C. viticella colonophyte 0 0 Ficus carica ephemerophyte 0 0 Rubus macrophyllus 0 0 0 Toxicodendron radicans ephemerophyte 0 0 Vitis labrusca_ephemerophyte_0_0

Tbtaí 49 44 49

Note: here and in Tables 4 and 7 "0" - the species is absent.

Table 4

The degree of naturalization of woody exotic species in some urban floras of Ukraine

Degree of naturalization Kyiv Kryvyi Rih Kharkiv Uzhhorod Donbas Total*

Agriophyte 5 1 7 3 2 10

Hemiagriophyte 8 7 0 1 0 8

Epoecophyte 4 5 8 1 3 12

Ergasiophyte 0 28 33 13 9 36

Colonophyte 27 3 1 0 0 12

Ephemerophyte 5 0 0 0 1 4

Total 49 44 49 18 15 82

Note: * - this column presents the data about the number of species in specific

categories of the total list.

68 exotic species have spontaneously spiead within the South Coast of Crimea. These aie 27 agiiophytes: Acer negundo, Ailanthus altissima, Berberis aquifolia, Bupleurum fruticosum, Buxus sempervirens, Cercis si-liquastrum, Clematis flammula, Colutea orientalis, Cydonia oblonga, Daphne laureola, Elaeagnus angustifola, Ficus carica, Fraxinus ornus, Laburnum anagyroides, Lonicera etrusca, Lycium barbarum, Malus domestica, Platycladus orientalis, Prunus cerasifera, P. dulcis, Ptelea trifo-liata, Pueraria montana var. lobata, Pyrus communis, Quercus ilex, Rhamnus alatemus, Viburnum tinus and Vitis vinifera, 40 colonophytes: Abies pinsapo, Amorpha fruticosa, Buddleja davidii, Caragana arbores-cens, Castenea sativa, Catalpa bignonioides, Cedrus atlantica, C. deoda-ra, Celtis australis, C. caucasica, Cladrastis kentukea, Colutea arbores-cens, Cotoneaster glaucophyllus, Cupressus sempervirens, Dyospyros lotus, Gleditsia triacanthos, Koelreuteria paniculata, Prunus laurocerasus, Laurus nobilis, Lonicera caprifolium, L. standishii, L. tatarica, Maclura pomífera, Morus alba, M. nigra, M. rubra, Olea europaea, Parthenocis-sus quinquefolia, Prunus armeniaca, P. domestica, P. vulgaris, Pyracan-tha rogersiana, Ribes aureum, R. rubrum, R. spicatum, Robinia pseudoacacia, Sophora japonica, Spartium junceum, Syringa vulgaris and Zel-kova carpinifolia and ephemerophyte Prunus cerasus.

18 exotic woody species penetrated the flora of the Yalta Mountain-Forest Natural Reserve. All of them are agiiophytes - 13 species, and colonophytes - 5 species (Bagrikova & Bondarenko, 2015). A number of alien woody species form local populations of normal type in the native communities of "Mys Martian" nature reserve: Bupleurum fiuticosum, Buxus sempervirens, Fraxinus ornus, Laburnum anagyroides

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 ergasiophyte 1 1 ergasiophyte

0 0 2 1 colonophyte

0 0 1 1 ergasiophyte

0 0 1 1 ergasiophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 colonophyte

0 0 1 1 ephemerophyte

0 ephemerophyte 1 1 ephemerophyte

0 0 1 1 ephemerophyte

0_0_1_1 ephemerophyte

18 15 - -

and Prunus cerasifera (Bagrikova et al., 2014). In addition to the data about naturalization of plants in the botanical gardens, cities and nature reserve fund, we have considered articles about their separate records (Tyshchenko et al., 2013; Burda, 2014). It has also been noted that Acer negundo and Robinia pseudoacacia are viewed as diagnostic species of the synanthropic Robinietea class, stable components of Salicetea pur-pureae class, capable of intruding into the phytocenoces of Querco-Fagetea class causing structural disruption. Elaeagnus angustifolia and E. commutata are transformer species in Steppe and Circum-Pontic regions, conditioning the colonization of river basins, changing the grass cover of salinized coastal depressions (Abduloyeva & Karpenko, 2009). Ailanthus altissima, Celtis occidentals, Juglans cinerea, J. mandshurica, J. regia, Prunus serotina, P. virginiana, Ribes uva-crispa, spread spontaneously from forest culture in the nature reserve territories of the Forest-Steppe, also in the habitats of G type (forests and shrubs), Acer negundo and Amorpha fruticosa are involved in the cenoses of Rham-no-Prunetea class (Pashkevych & Burda, 2017).

The invasion of alien woody species into spontaneous flora in Ukraine

Therefore, 182 non-native woody species (172 species, 1 subspecies, 4 varieties and 5 hybrids) have been distinguished in the spontaneous flora of Ukraine (Tables 5, 6). This list is obviously incomplete. Only the articles in general access have been considered. The archive data on thorough studies of the course of initial introduction are unaccessible. Herbarium labels do not always highlight the origin of herborized samples, collected in the botanical gardens and arboretums. On the one hand, the compilation of a more precise list of woody exotic species is hindered by the absence of a unified inventory of collection funds and database of the course of naturalization of these species, and, on the other hand, the naturalization process is not over. On the contrary, it is getting stronger. Plant invasion is an extremely dynamic process. However, the presented list gives a general idea about the composition and character of non-native woody trees in the domestic flora.

71 alien completely naturalized plants have been differentiated (66 species, 1 subspecies, 1 variety and 3 hybrids). They belong to 47 genera and 28 families of two divisions - Pinophyta and Magnoliophyta (Table 5). This group presents the highest risk for the local diversity of species: 12 of them are transformers, 8 are invasive proper, 29 - potentially invasive, 22 - naturalized species, whose invasive activity has not been manifested. In terms of life-forms, the list of alien woody species contains

30 species - trees, 26 species - shrubs, and 8 species - lianas, 7 species de- velop their life-form as a tree or a shrub, depending on their environment.

Table 5

The invasive activity of alien woody species in spontaneous flora of Ukraine

Family

Taxon

Life-form

Spread

Degree of naturalization*_Occurrence Invasive activity

Pinophyta

Cupressaceae Platycladus orientalis (L.) Franco

tree/shrub

seed

agriophytec

naturalized

Berberidaceae Berberis aquifolia Pursh

Berberidaceae B. vulgaris L.

Ranunculaceae Clematis flammula L.

Buxaceae Buxus sempervirens L.

Grossulariaceae Ribes uva-crispa L.

Vitaceae Parthenocissus quinquefolia (L.) Planch.

Vitaceae P. tricuspidata (Siebold & Zucc.) Planch.

Vitaceae P. vitacea (Knerr) Hitchc.

Vitaceae Vitis vinifera L.

Fabaceae Amorpha fruticosa L.

Fabaceae Caragana arborescens Lam.

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Fabaceae Cercis siliquastrum L.

Fabaceae Colutea orientalis Mill.

Fabaceae Gleditsia triacanthos L.

Fabaceae Laburnum anagyroides Medik.

Fabaceae Pueraria montana var. lobata (Willd.)

Sanjappa & Prade

Fabaceae Robinia pseudoacacia L.

Rosaceae Amelanchier spicata (Lam.) K. Koch

Rosaceae Cydonia oblonda Mill.

Rosaceae Malus domestica Borkh.

Rosaceae Physocarpus opulifolius (L.) Maxim.

Rosaceae Prunus armeniaca L.

Rosaceae P. cerasus L.

Rosaceae P. cerasifera Ehrh.

Rosaceae P. domestica L.

_ P. domestica subsp. insititia (L.) Rosaceae

Bonnier & Layens

Rosaceae P. dulcis (Mill.) D.A. Webb

Rosaceae P. serotina Ehrh.

Rosaceae P. virginiana L.

Rosaceae Pyrus communis L.

Rosaceae Rosa rugosa Thunb.

Rosaceae Sorbaria sorbifolia (L.) A. Braun

Elaeagnaceae Elaeagnus angustifolia L.

Elaeagnaceae E. rhamnoides (L.) A. Nelson

Elaeagnaceae E. commutata Bernh. ex Rydb.

Rhamnaceae Rhamnus alaternus L.

Ulmaceae Ulmus pumila L

Moraceae Ficus carica L.

Moraceae Morus alba L.

Fagaceae Quercus ilex L.

Fagaceae Q. rubra L.

Celastraceae Celastrus scandens L.

Salicaceae Populus balsamifera L.

Salicaceae P. bolleana Lauche

Salicaceae P. x canadensis Moench

Salicaceae P. deltoides Marshall

Salicaceae P. nigra var. italica Münchh

Salicaceae P. suaveolens Fisch.

Salicaceae P. richocarpa Torr. & A. Gray ex Hook.

Salicaceae Salix x babilonica L.

Salicaceae S. x blanda Andersson

Anacardiaceae Toxicodendron radicans (L.) O. Kuntze

Aceraceae Acer negundo L.

Rutaceae Ptelea trifoliata L.

Simaroubaceae Ailanthus altissima (Mill.) Swingle

Tiliaceae Tilia americana L.

Thymelaeaceae Daphne laureola L.

Cornaceae Cornus alba L.

Cornaceae C. sericea L.

Hydrangeaceae Philadelphus coronarius L.

Solanaceae Lycium barbarum L.

Oleaceae Fraxinus americana L.

Oleaceae F. ornus L.

Magnoliophyta

shrub seed, vegetative way

shrub seed, vegetative way

liana seed, vegetative way

shrub seed

shrub seed, vegetative way

liana seed, vegetative way

liana seed, vegetative way

liana seed, vegetative way

liana seed, vegetative way

shrub seed, vegetative way

shrub seed

tree/shrub seed

shrub seed

tree seed

shrub seed

liana seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree/shrub seed, vegetative way

tree seed, vegetative way

shrub seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree/shrub seed, vegetative way

tree seed, vegetative way

tree/shrub seed, vegetative way

tree seed

tree/shrub seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

shrub seed, vegetative way

shrub seed, vegetative way

tree/shrub seed, vegetative way

shrub seed,vegetative way

tree seed, vegetative way

shrub seed, vegetative way

tree seed, vegetative way

tree/shrub seed

tree seed

tree seed

tree seed

liana seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

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tree seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

tree seed, vegetative way

shrub seed, vegetative way

tree seed

shrub seed, vegetative way

tree seed, vegetative way

tree seed

shrub seed, vegetative way

shrub seed, vegetative way

shrub seed, vegetative way

shrub seed, vegetative way

shrub seed, vegetative way

tree seed

tree seed

agriophyte epoecophyte agriophytec agriophytec ergasiophyte colonophytec, epoecophyte epoecophyte agriophyte agriophytec, epoecophyte agriophyte, colonophytec colonophytec, hemiagriophyte agriophytec agriophytec colonophytec, epoecophyte agriophytec

agriophytec

colonophytec, agriophyte agriophyte agriophyte agriophyte epoecophyte colonophytec, epoecophyte ephemerophytec, epoecophyte agriophyte colonophytec, ergasiophyte

agriophyte

agriophytec agriophyte ergasiophyte agriophyte ergasiophyte epoecophyte agriophyte agriophyte hemiagriophyte

agriophytec hemiagriophyte

agriophytec, ephemerophyte colonophytec, epoecophyte agriophytec emiagriophyte epoecophyte ergasiophyte ergasiophyte epoecophyte epoecophyte ergasiophyte ergasiophyte ergasiophyte ergasiophyte agriophyte ergasiophyte agriophyte agriophytec, hemiagriophyte agriophytec, epoecophyte ergasiophyte agriophytec ergasiophyte epoecophyte ergasiophyte agriophytec, epoecophyte epoecophyte _agriophytec_

local invasive

sporadic naturalized

sporadic potentially invasive

local potentially invasive

local potentially invasive

common transformer

rare naturalized

common transformer

common invasive

common ttransformer

common naturalized

rare naturalized

local invasive

sporadic naturalized

local potentially invasive

local potentially invasive

common transformer

sporadic potentially invasive

common potentially invasive

common potentially invasive

common naturalized

common naturalized

common naturalized

common invasive

common naturalized

common potentially invasive

local potentially invasive

common invasive

common potentially invasive

common naturalized

common naturalized

common potentially invasive

common transformer

common transformer

rare potentially invasive

local trnsformer

common potentially invasive

local potentially invasive

unique

common potentially invasive

local invasive

local potentially invasive

rare potentially invasive

rare naturalized

common naturalized

common naturalized

common naturalized

common naturalized

common naturalized

common naturalized

common naturalized

common trnsformer

local potentially invasive

common transformer

common potentially invasive

common trnsformer

common naturalized

local invasive

common potentially invasive

common potentially invasive

common naturalized

common ins potentially invasive

common naturalized

local transformer

Family Taxon Life-form Spread Degree of naturalization* Occurrence Invasive activity

Oleaceae F. pennsylvanica Marshall tree seed epoecophyte common potentially invasive

Oleaceae Syringa vulgaris L. shrub seed, vegetative way colonophytec, ergasiophyte common naturalized

Viburnaceae Viburnum tinus L. shrub seed, vegetative way agriophytec local invasive

Caprifoliaceae Lonicera etrusca Santi shrub seed agriophytec local potentially invasive

Caprifoliaceae L. tatarica L. shrub seed colonophytec, epoecophyte common naturalized

Caprifoliaceae Symphoricarpos albus (L.) S. F. Blake shrub seed, vegetative way epoecophyte common naturalized

Apiaceae Bupleurum fruticosum L. shrub seed, vegetative way agriophytec local transformer

Note: * - here and in Table 6 interlinear "c" marks "Crimea".

The prevailing majority of alien woody species have both seed dispersal and vegetative reproduction (56 species). Vegetative reproduction was not observed in nature for 15 species. In terms of the naturalization degree of alien woody species, 35 species are agriophytes, 5 -hemiagriophytes, 16 - epoecophytes, 18 - ergasiophytes, 15 - colono-phytes and 1 - ephemerophyte.

A number of species in specific regions have acquired different degrees of naturalization. For instance, Ficus carica is an agriophyte in the flora of the South Coast of Crimea, but an ephemerophyte - in the urban flora of Kyiv, Ailanthus altissima and Lycium barbarum are agriophytes in the Crimea, and in the rest of the territory they are epoecophytes. At the same time, Caragana arborescens and Robinia pseudoacacia have not naturalized completely on the peninsula, and have the status of colono-phytes, while on the mainland they are hemiagriophyte and agriophyte, respectively, etc. The following species spread as agriophytes only on the Crimean Peninsula: Bupleurum frvticosum, Buxus sempervirens, Clematis flammula, Cercis siliquastrum, Colutea orientalis, Daphne laureola, Laburnum anagyroides, Lonicera etrusca, Platycladus orientalis, Prunus dulcis, Pueraria montana var. lobata, Quercus ilex andRhamnus alaternus, etc.

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Table 6

Non-native naturalizing woody species of spontaneous flora of Ukraine

The naturalization non-native woody species in plant communities in Ukraine

There are 111 non-native woody species whose naturalization has not completed yet (107 species, 2 varieties, and 2 hybrids), out of 65 genera and 35 families, 2 divisions - Pinophyta and Magnoliophyta (Table 6). In terms of life-forms, these are trees (45 species), shrubs (38 species), lianas (19 species), 9 have the life-form ofboth a tree and a shrub, they have seed dispersal (61), vegetative way of reproduction (4), or spread via both ways (46). The degree of occurrence of non-native species is reflected by the following spectrum: 7 - sporadic, 17 - local, 51 - occurring in 3-5 (7) localities, 13 - unique and 23 species which have dispersed by seed dispersal or vegetative way, having escaped the collections and expositions only in botanical gardens and arboretums. The species of this group are mainly ephemerophytes by the degree of naturalization. However, some of them are already acquiring the status of colonophytes: Allbizia julibrissin, Sophora japonica, Spartium junce-um, while Ficus carica and Juglans regia on the South Coast of Crimea are even agriophytes (Bagrikova, 2013).

Family Taxon Life-form Spread Degree of naturalization Occurrence Stage of naturalization

Pinophyta

Pinaceae Abies pinsapo Boiss. tree seed colonophytec rare reproduction

Pinaceae Cedrus atlantica (Endl.) Manetti ex Carrière tree seed colonophytec rare reproduction

Pinaceae C. deodara (Roxb. ex D. Don) G. Don tree seed colonophytec rare reproduction

Pinaceae Larix decidua Mill. tree seed ephemerophyte controlled reproduction

Cupressaceae Cupressus sempervirens L. tree seed colonophytec rare reproduction

Cupressaceae Thuja occidentalis L. tree/shrub seed ephemerophyte controlled reproduction

Magnoliophyta

Aristolochiaceae

Lauraceae

Berberidaceae

Magnoliaceae

Ranunculaceae

Ranunculaceae

Ranunculaceae

Ranunculaceae

Ranunculaceae

Ranunculaceae

Platanaceae

Hamamelidaceae

Grossulariaceae Grossulariaceae Grossulariaceae Grossulariaceae Grossulariaceae Vitaceae

Vitaceae

Vitaceae

Vitaceae

Vitaceae Vitaceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae

Aristolochia macrophylla Lam. Laurus nobilis L. Berberis thunbergasiophyteii DC. Liriodendron thunbergii DC. Clematis gouriana Roxb. ex DC. C. jackmanii T. Moore. C. ligusticifolia Nutt C. tangutica (Maxim.) Korsh. C. vitalba L. C. viticella L.

Platanus acerifolia (Aiton) Willd.

Parrotia subaequalis (Hung T. Chang)

R.M. Hao & H.T. Wei

Ribes americanum Mill.

R. aureum Pursh

R. europaea (L.) Mill.

R. rubrum L.

R. spicatum Robson

Ampelopsis aconitifolia Bunge

A. delavayana var. glabra (Diels & Gilg)

C.L. Li

A. bodinieri (H. Lev. & Vaniot) Rehder A. glandulosa var. brevipedunculata (Maxim.) Momiy

Vitis coignetiae Bull. ex Planch

V. labrusca L.

Allbizia julibrissin Durazz

Cladrastis kentukea (Dum. Cours.) Rudd

Colutea arborescens L.

Gymnocladus dioica (L.) K. Koch

Halimodendron halodendron (Pall.) Voss

liana

tree/shrub shrub tree liana liana liana liana liana liana

tree

tree/shrub

shrub shrub shrub shrub shrub liana

liana

liana

liana

liana liana tree shrub shrub tree shrub

seed seed

seed, vegetative way seed, vegetative way seed, vegetative way seed, vegetative way seed, vegetative way seed, vegetative way seed, vegetative way seed, vegetative way

seed

seed

seed seed seed seed seed

seed, vegetative way

seed, vegetative way

seed, vegetative way

seed, vegetative way

seed, vegetative way seed, vegetative way seed, vegetative way seed seed seed seed

ephemerophyte unique records

colonophytec rare reproduction

ephemerophyte controlled survival

ephemerophyte unique records

ephemerophyte controlled establishing

ephemerophyte controlled establishing

ephemerophyte controlled establishing

ephemerophyte controlled establishing

colonophyte rare establishing

ephemerophyte controlled establishing colonophytec

unique records

ephemerophyte

ephemerophyte controlled reproduction

ephemerophyte colonophytec ephemerophyte colonophytec colonophytec ephemerophyte

ephemerophyte

ephemerophyte

ephemerophyte

ephemerophyte

colonophyte ephemerophytec colonophytec colonophytec ephemerophyte colonophyte

controlled rare rare rare rare controlled

controlled

controlled

controlled

controlled rare rare

sporadic

rare rare

local

reproduction reproduction reproduction reproduction reproduction survival

survival

survival

survival

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survival reproduction

survival establishing survival survival establishing

Family

Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Rosaceae Rosaceae

Rosaceae

Rosaceae

Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae

Rosaceae

Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Ulmaceae Ulmaceae Ulmaceae Ulmaceae Moraceae Moraceae Moraceae Fagaceae

Fagaceae

Fagaceae

Juglandaceae

Juglandaceae

Juglandaceae

Juglandaceae

Juglandaceae

Juglandaceae

Celastraceae

Celastraceae

Celastraceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Anacardiaceae

Aceraceae

Aceraceae

Hippocastanaceae

Hippocastanaceae

Sapindaceae

Rutaceae

Tiliaceae

Cistaceae

Hydrangeaceae

Hydrangeaceae

Ebenaceae

Eucommiaceae

Oleaceae

Scrophulariaceae

Paulowniaceae

Bignoniaceae

Bignoniaceae

Taxon

Robinia hispida L. R. neomexicana A. Gray R. viscosa Vent. Sophorajaponica L. Spartium junceum L. Amelanchier ovalis Medik. Aronia melanocarpa (Michx.) Elliott Cerasus tomentosa (Thunb.) Wall. ex T.T. Yu & C.L. Li Chaenomeles japonica (Thunb.) Lindl. ex Spach

Cotoneaster glaucophyllus Franch. C. lucidus Schlecht. C. melanocarpus Fisch. ex A. Blytt Crataegus coccinea L. C. sanguineus Pall. C. submollis Sarg. Malus baccata (L.) Borkh. Mespilus germanica L. Persica vulgaris Mill. Prunus laurocerasus L. P. mahaleb L. P. padus L. P. vulgaris Schur

Pyracantha rogersiana (A. B. Jacks.) Coltm.-Rog.

Rubus macrophyllus Weihe & Nees

R. odoratus L.

Spiraea x billardii Hérin

S. chamaedryfolia L.

S. douglasii Hook.

S. salicifolia L.

S. x vanhouttei (Briot) Zabel

Celtis australis L.

C. caucasica Willd.

C. occidentalis L.

Zelkova carpinifolia (Pall.) K. Koch Maclura pomifera (Raf.) C. K. Schneid. Morus nigra L. M. rubra L. Castanea sativa Mill.

Quercus macranthera Fisch. & C. A. Mey. ex Hohen.

Q. palustris Moench Juglans ailanthifolia Carrière J. cinerea L. J. mandshurica Maxim. J. nigra L. J. regia L.

J. subcordiformis Dode Celastrus angulata Maxim. C. flagellaris Rupr. C. orbiculatus Thunb. Rhus glabra L. R. typhina L. R. typhina L. 'Laciniata' Toxicodendron pubescens Mill. Acer platanoides L. f. atropurpurea Krimson King' A. saccharinum L. Aesculus flava Sol. A. hippocastanum L. Koelreuteria paniculata Laxm. Phellodendron amurense Rupr. Tilia begoniifolia Steven Poncirus trifoliata (L.) Raf. Deutzia scabra Thunb. Philadelphus microphyllus A. Gray Diospyros lotus L. Eucommia ulmoides Oliver. Olea europaea L. Buddleja davidi Franch. Paulownia tomentosa (Thunb.) Steud.

Catalpa bignonioides Walter

Campsis grandiflora (Thunb.) K. Schum.

Life-form Spread Degree of naturalization Occurrence Stage of naturalization

tree seed, vegetative way colonophyte local establishing

tree seed, vegetative way colonophyte local establishing

tree seed, vegetative way colonophyte local establishing

tree seed colonophytec unique survival

shrub seed, vegetative way colonophytec sporadic establishing

tree/shrub seed ephemerophyte local establishing

tree seed ephemerophyte local establishing

shrub seed ephemerophyte rare reproduction

shrub seed, vegetative way ephemerophyte rare reproduction

shrub seed colonophytec rare reproduction

shrub seed ephemerophyte local reproduction

shrub seed ephemerophyte local reproduction

shrub seed ephemerophyte rare reproduction

shrub seed ephemerophyte rare reproduction

shrub seed ephemerophyte rare reproduction

tree seed ephemerophyte rare reproduction

shrub seed, vegetative way ephemerophyte rare reproduction

tree seed ephemerophyte rare reproduction

shrub seed, vegetative way colonophytec local establishing

shrub seed ephemerophyte rare reproduction

tree/shrub seed ephemerophyte rare reproduction

tree/shrub seed, vegetative way colonophytec sporadic establishing

shrub seed, vegetative way colonophytec rare reproduction

shrub seed, vegetative way ephemerophyte unique records

shrub seed, vegetative way ephemerophyte controlled survival

shrub seed, vegetative way ephemerophyte rare survival

shrub seed,vegetative way ephemerophyte rare survival

shrub seed, vegetative way ephemerophyte rare survival

shrub seed, vegetative way colonophyte rare survival

shrub seed, vegetative way ephemerophyte rare survival

tree seed colonophytec rare survival

tree/shrub seed colonophytec rare survival

tree seed ephemerophyte rare survival

tree seed colonophytec rare survival

tree seed, vegetative way colonophytec rare survival

tree seed colonophytec sporadic establishing

tree seed colonophytec sporadic establishing

tree seed colonophytec unique establishing

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tree seed ephemerophyte controlled establishing

tree seed ephemerophyte rare reproduction

tree seed ergasiophyte local establishing

tree seed ergasiophyte local establishing

tree seed ergasiophyte local establishing

tree seed ergasiophyte local establishing

tree seed ergasiophyte sporadic establishing

tree seed ergasiophyte local establishing

liana seed, vegetative way colonophyte local survival

liana seed ephemerophyte local survival

liana seed ephemerophyte local survival

tree/shrub seed, vegetative way colonophyte rare survival

tree vegetative way colonophyte controlled survival

tree vegetative way colonophyte controlled survival

shrub seed, vegetative way ephemerophyte rare survival

tree seed ephemerophyte controlled survival

tree seed colonophyte sporadic survival

tree seed ephemerophyte unique records

tree seed ergasiophyte rare esablishing

tree seed colonophytec rare survival

tree seed ephemerophyte unique records

tree seed, vegetative way ephemerophyte rare survival

tree/shrub seed ephemerophyte rare reproduction

shrub seed, vegetative way ephemerophyte rare survival

shrub seed, vegetative way ephemerophyte rare reproduction

tree seed, vegetative way colonophytec rare survival

tree seed, vegetative way ephemerophyte unique records

tree seed, vegetative way colonophytec rare establishing

shrub seed, vegetative way colonophytec unique records

tree seed ephemerophyte unique records

tree seed, vegetative way colonophytec ephemerophyte unique survival

liana vegetative way colonophyte controlled establishing

Family Taxon Life-form Spread Degree of naturalization Occurrence Stage of naturalization

Bignoniaceae C. radicans (L.) Seem. liana vegetative way colonophy1e controlled establishing

Aquifoliaceae Ilex aquifolium L. 1ree seed, vegetative way ephemerophyte unique records

Caprifoliaceae Lonicera caerulea L. shrub seed ephemerophyte rare survival

Caprifoliaceae L. caprifolium L. shrub seed colonophytec rare survival

Caprifoliaceae L. standishii Jacques shrub seed colonophytec rare survival

Caprifoliaceae Weigela florida (Bunge) A DC. shrub seed, vegetative way ephemerophyte rare survival

Aracaceae Aralia elata (Miq.) Seem. shrub seed, vegetative way ephemerophyte controlled survival

A general overview of the course of invasion of woody species into the spontaneous flora of Ukraine

According to the results of our determination and analysis of the group of non-native woody species, this component of the spontaneous flora of Ukraine has a rather diverse taxonomy: 182 species, belonging to 95 genera and 45 families. These include 71 alien species, which have completely naturalized. Let us compare: the global database of invasive trees and shrubs contains 751 alien species (434 trees and 317 shrubs); the flora of Europe has 134 of them (Rejmánek & Richardson, 2013). It is evident that it has no significant relevance for the course of invasion whether the life-form is a tree or a shrub. The involvement of trees in the total list slightly exceeds the involvement of shrubs, and as for alien species - they are even (Table 7). The most intense distribution has been noted mainly for the species remarkable both for seed dispersal and vegetative reproduction, though most non-naturalized species have only seed dispersal. Mostly common and rare species are noted in the total list in terms of occurrence and distribution, which is explained by high involvement of these very species among alien and non-naturalized ones.

The course of invasion of non-native woody species, which are at the initial stages of migration - introduction, acclimatization, survival, adaptation of reproductive sphere and establishment (Blackburn et al., 2011) - and have not naturalized yet, takes place after overcoming the geographical barrier as a result of human activity. Continuing their introduction, they cross the barriers of controlled cultivation, a barrier to survival and adaptation of reproductive sphere, and start the formation and establishment of populations. According to our observations, only 30 of them have established local populations, have cryptic (hidden) invasion risk, and the rest of the non-naturalized species do not manifest any invasive activity. First of all, noteworthy are vegetatively mobile shrubs of Rhus typhina, R typhina 'Laciniata', Rubus odoratus and lianas Ampelopsis aconitifolia, A. delavayana var. glabra, A. bodinieri, A. glandulosa var. brevipedunculata, Campsis grandiflora, C. radicans, Clematis gouriana, C. jackmanii, C. ligustiafolia, C. tangutica, C. viti-cella and Vitis coignetiae. The global pattern of plant invasions demonstrates that under favourable conditions the species of these life-forms invade a territory fast and retain there hold on it. The studies on the naturalization in forests and parks of Kyiv demonstrated that 6 species of the genus Juglans (J. ailanthifolia, J. cinerea, J. mandshurica, J. nigra, J. regia, J. subcordiformis) (Burda & Koniakin, 2018) and Aesculus hippocastanum establish self-reproductive local populations. As for further successful or failing seed dispersal of trees (Acer platinoides f. atropurpurea Krimson King', Celtis australis, Larix decidua, Quercus macranthera), it is too early to forecast anything. The mentioned ornamental form of Acer platanoides is present in some parks of Kyiv, but there were no reports about self-dispersal. Non-naturalized and not high trees or shrubs (Aralia elata, Parrotia subaequalis, Thuja occidentalis) are still undergoing the processes of acclimatization and adaptation of reproductive sphere. Such shrubs as Berberis thunberii and Ribes ame-ricanum do not have abundant seed dispersal. Thus, the group of non-native woody species combines the species with hidden or non-expressed invasive activity. At first sight, they are completely undynamic, almost invisible in flora, plant communities, ecosystems and landscape. However, at the impact of some factors, for instance, factors of time, favourable conditions of reproduction, sharp change in environment, etc., their activity may be revealed. Most species, which have not naturalized, have not acquired sufficient vegetative development for at least some invasive activity, so they remain passive in terms of this feature for some time.

The picture of invasive activity of alien species is quite different. The species, which have formed secondary ranges within Ukraine, are known as invasively active species, at least within Europe and Northern 2A8s6ia. Transformer species in the spontaneous flora of Ukraine are revealed as the most wide spread in almost 40 regions of Europe: Robinia pseudoacacia (42 regions), Ailanthus altissima (40), Acer negundo (38) (Lambdon et al., 2008). Some of our non-native plant species are among the more than "100 worst" alien species in Europe (Nentwig et al., 2018). There are Pueraria lobata var. montana (rank 10 and total impact sum 29), Robinia pseudoacacia (13 and 28), Prunus serotina (59 and 17), Elaeagnus angustifolia (67 and 16), Rosa rugosa (76 and 13) and Buddleja davidii (80 and 11 respectively).

Table 7

The diversity of non-native woody species in spontaneous flora of Ukraine

Feature _Number of species_

total alien on-naturalized

Taxonomic diversity

of species 182 71 111

of genera 95 47 65

of families 45 28 35

Life-form

tree 75 30 45

shrub 64 26 38

tree or shrub 16 7 9

liana 27 8 19

Spreading

seed dispersal 77 15 62

vegetative way 4 0 4

both way 101 56 45

Occurrence

common species 44 44 0

local species 34 17 17

sporadic species 11 4 7

rare species 57 6 51

unique species 13 0 13

controlled species 23 0 23

*Degree ofnaturalization

agriophyte 35 35 0

hemiagriophyte 5 5 0

epoecophyte 16 16 0

ergasiophyte 23 15 8

colonophyte 42 0 42

ephemerophyte 61 0 61

*Invasive activity

transformer species 12 12 0

invasive species proper 8 8 0

potentially invasive species 29 29 0

naturalized species with no manifested 22 22 0

invasive activity

* Non-naturalized species, such stages of naturalization continue

introduction 10 0 10

survival 40 0 40

adaptation of reproduction sphere 31 0 31

establishment 30 0 30

Note: * - the highest level has been accepted for species, which is quite different by the degree features at the local level.

The expansion of Amorpha fruticosa is observed in neighbouring countries. In the flood plain of Kuban near the town of Temriuk A. fruti-cosa formed compact clumps, pushing out native plant communities of hygro- and hydrophytes. There were cases when the traditional shrub life-form ofA. fruticosa was changed to the tree life-form. The height of trees was 6 m with the trunk diameter of 15 cm (Shvydkaya & Kudino-va, 2013). About two dozen alien woody species of the spontaneous 2019, 27(3)

flora of Ukraine have spread in 25 and more regions as the most common alien species of Europe: Prunus cerasus (34), Quercus rubra (34), Rosa rugosa (34), Prunus domestica (31), P. cerasifera (30), Aesculus hippocastanum (30), Pyrus communis (30), Syringa vulgaris (30), Malus domestica (29), Berberis aquifolia (28), Parthenocissus quinquefolia (27), Symphoricarpos albus (27), Juglans regia (26), Populus x canadensis (26), Vitis vinifera (26) (Lambdon et al., 2008).

Thus, coming back to the issue on the source of diaspores in the course of woody plant invasions in Ukraine, we would like to highlight a probable role of interstate migrations of invasive woody species. It is also possible that some migrations go both ways. At present, Ukraine's botanical gardens and arboretums have abundant and rather old collections. Some introduction centers own over 2,000 species of trees and shrubs. During their history (from 1793 till 2019), these institutions have passed on many woody exotic species for creation of forest cultures, use by communal services, to private amateurs, etc. In our country, woody species, used in gardening by the botanical gardens and arboretums after initial introductory testing, are planted to create current greenery of the cities along with the local species. They ensure comfortable life of local residents, promote optimization of microclimate in agriophyteoecosystems, protect railways and highways from unfavourable weather effects, resist water and wind erosion of soils, etc. In this situation, reports about immediate facts of woody species escaping the botanical gardens and arboretums which have introduced them, and establishing spontaneous plantings, are almost absent. We have managed to find only one abovementioned case of the "escape" of Clematis vitalba from the Donetsk Botanical Garden of NAS of Ukraine, on the mainland. This species is usually estimated by specialists as an unstable element of flora (colonophyte), remarkable for potential invasive activity, conditioned by the ability of both fast vegetative reproduction, and easy dispersal of seeds in the wind. As for the Crimean Peninsula, there is a registered fact of native penetration of Bupleurum fruticosum and Fraxinus ornus into the nature reserve "Mys Martian" from the adjacent territory of the Nikitsky Botanical Garden, where both species were naturalized long time ago (Bagrikova et al., 2014). However, the question about the nature of Fraxinus ornus in the Crimea is still under discussion. P. S. Pallas considered F. ornus on the South Coast of Crimea, in particular, in the plant communities on Mys Martian, to be an aboriginal East Mediterranean species, even 18 years prior to the establishment of the Nikitsky Botanical Garden. This thought coincides quite well with the opinions of some modern botanists. However, taking into consideration successful seed dispersal of F. ornus in modern plant community complexes, it seems reasonable to accept it as a transformer species on the South Coast of Crimea (Kish et al., 2009; Protopopova et al., 2012). The consequences of invasions of woody trees from the introduction centers seem to be negligible on the background of the events in domestic spontaneous flora. This conclusion is in agreement with the status of naturalization for woody plants from collections and expositions of the N. V. Tsytsyn Main Botanical Garden, RAS. Its specialists have not observed any fact of an introduced woody species escaping the garden in 70 years of introduction tests (Yatsenko & Vino-gradova, 2018). They have described the following course of migration: 2,400 species have been involved in the introduction experiment; 1,317 species (55%) have overcome the ecological barrier and adjusted to new natural and climatic conditions; 66 species have overcome the reproductive barrier (5%); 12 species have actually "left the arboretum" and invaded natural forest cenoses within the territory of the Garden (8%). It should be noted that among 66 species, which have spread beyond the limits of cultivation, 10 species are aboriginal in the flora of Ukraine, and the rest are mostly mentioned in Tables 5 and 6.

Thus, the botanical gardens and arboretums of Ukraine as centers of plant introduction are neither direct sources of naturalization nor sources of spontaneous distribution of non-native woody species. The assumption about these institutions being the centers of initial introduction is surely reasonable.

In addition, the fact of seed dispersal, made by this or that species within the collection or under conditions, approximated to natural ones, is not a guarantee of its invasion in the nearest future. The way in which a plant migrates within a new territory is rather complicated, there may

be "boom" and "bust" situations, or other occasions (Blackburn et al., 2011). There have been many situations when the invasion of some species started with seed dispersal and ended with it. For instance, Par-rotia subaequalis was introduced into the arboretum of the M. M. Hryshko National Botanical Garden in 1950, the plants first blossomed in 1975 and the mature seeds had 98% germination. Abundant seed dispersal was observed the following year, however, the seedlings were soon eliminated. In the following years the plants of P. subaequalis blossomed, had seeds, but there was no seed dispersal (Doroshenko et al., 2013). In this respect, noteworthy are observations of the spreading of woody trees in the arboretum of the Donetsk Botanical Garden (Eremenko & Ostapko, 2011). Three tendencies of distribution were noted under the controlled conditions of the arboretum during 10 years of observations. As expected, the species with the positive tendency include most invasive species: Acer negundo, Ailanthus altissima, Par-thenocissus quinquefolia, potentially invasive: Morus alba, Ribes uva-crispa, and naturalized Gleditsia triacanthos, Berberis vulgaris and Juglans regia. The latter species sometimes establishes separate local self-reproducing populations in Donetsk on the background of abundant cultivation on private land. Decreasing dispersion was demonstrated by Berberis aquifolia, while Caragana arborescens, Ptelea trifoliata, Robinia pseudoacacia had a stable tempo of spreading.

In 55 years of observations, 227 alien woody species out of 73 genera and 31 families with seed dispersal were revealed in green plantings of Saint Petersburg. Among these, only about 10 species have become invasive and even threats to aboriginal plant communities (Firsov & Byalt, 2015). For instance, these are species: Cornus saricea, Aronia mitschurinii A. Scvorts. et Maitull., Acer negundo, Amelanchier spicata.

262 species of vascular plants, which escaped the collections and expositions, were determined in the Tsytsyn Main Botanical Garden, RAS and the Botanical Garden of Lomonosov of the Moscow State University (Mayorov et al., 2013). Aralia elata and Symphoricarpus albus, capable of independent distribution in botanical gardens of Ukraine, are among them. Contrary to the abovementioned, in the arboretum of the Tsytsyn Main Botanical Garden, natural forest communities of the Garden were invaded from the outside greenery of the city by Acer negundo, Cotoneaster lucidus, Malus domestica (Yatsenko & Vinogra-dova, 2018).

The phenomenon of naturalization of non-native species near collections and expositions of the botanical gardens and arboretums is of global character. Let us supplement the description of the experience of botanical gardens of Ukraine, Moscow and Saint Petersburg with the facts of Caragana arborescens, going out of cultivation within the Yakutsk Botanical Garden of the Institute of Biological Problems of Cryo-lithozone, the Siberian Division, RAS (on dry coastal shores of a lake, sometimes), which belongs to hemiagriophytes in Ukraine, and Sorba-ria sorbifolia (in a birch forest, near collection plots), which is an epo-ecophyte in our flora (Nikolaeva & Danilova, 2019).

Thus, important sources of naturalization of non-native woody species are green areas of cities, forest cultures, protective plantings of different intended purposes, private households, etc. The management of woody species invasion, the pragmatic unified framework for biological invasions (Blackburn et al., 2011), should be based on the global strategy on invasive alien species. The course of invasion of non-native woody species in the domestic flora dictates the following scheme of actions: preventing invasions, eradication - at the stages of survival and adaptation of the reproductive sphere, containment - after the species has overcome the barrier of reproduction and dispersal, and mitigation -on the final stages of invasion.

Conclusion

This is the first and preliminary determination and evaluation of non-native woody species in Ukraine's spontaneous flora. The data, presented in the article, demonstrate the active course of naturalization of non-native woody species and invasion of alien species in the spontaneous flora. The tendencies towards global increase of involvement and invasive activity of alien woody species in the spontaneous domestic flora have been confirmed.

The analysis of the occurrence, naturalization, distribution and the estimation of invasive activity of non-native species of trees, shrubs, and lianas in floras of 5 regions, 5 urban floras, over 30 floras of protected areas demonstrated as follows: there are 182 species from 95 genera and 45 families (75 trees, 16 trees or shrubs, 64 shrubs, and 27 lianas) are at different stages of naturalization in the spontaneous flora.

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71 species have completely naturalized, 20 of which are invasive and pose the highest threat to local diversity (12 transformer species and 8 invasive proper ones). The rest of the naturalized species are potentially invasive (29 species) or do not demonstrate invasive activity (22 species).

111 species have overcome the geographical barrier due to human activity. These are undergoing the initial stages of invasion: survival, acclimatization, adaptation of reproductive sphere, establishment and formation of local populations.

In Ukraine, there are about 50 botanical gardens and arboretums -centers of initial introduction of plants, 2 of them have been working since the end of the 18th century, and 8 - since the 19th century. The collections of 1 arboretum exceed 2,000, 2 botanical gardens have about 2,000 species, and the number of woody species in 4 botanical gardens and 1 arboretum exceeds 1,000 species. There is information about controlled spreading of 23 woody species beyond collections and expositions of the introduction centers. There was only one described case of spontaneous escape of 2 invasive woody species beyond the introduction center (Bupleurum fruticosum and Fraxinus ornus) from the Nikitsky Botanical Garden to the adjacent nature reserve "Mys Martian".

The mitigation of the effects of invasive alien woody species on local biodiversity requires restoration of local natural plant community complexes, organization of land use and culture of taking care of woody plantings. Our conclusions are relevant for employees of introduction centers, nature protection bodies, communal services, and state quarantine while elaborating the system of preventive, radical (eradication, limitation of assortment), containment and mitigation of impacts of plant invasions.

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