PUBLISHED: DECEMBER 2003

The Nonnative Flora of the Kyiv (Kiev) Urban Area, Ukraine: A Checklist and Brief Analysis

by Sergei L. Mosyakin and Oksana G. Yavorska

M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereshchenkivska St., Kyiv (Kiev) 01601 Ukraine

Published online December 24, 2002

Abstract

In this paper, an annotated checklist of the nonnative flora of the city and suburbs of Kiev (Kyiv in Ukrainian transliteration), Ukraine, is presented in tabular form. For each taxon, the following data are provided: occurrence, generalized distribution in the area, degree of naturalization, time of immigration, mode of immigration, and geographical origin. The total nonnative flora (past and present) of the Kiev Urban Area (KUA) consists of 536 species of vascular plants belonging to 297 genera and 71 families. It is the most diverse nonnative flora of any urban region in Ukraine. The modern nonnative flora includes 356 species of 207 genera and 62 families. The stable component of the flora consists of 198 species of 147 genera and 51 families. Ephemerophytes (175 species, or 49.2%) and epoecophytes (99 species, or 28.2%) clearly prevail among Kiev's modern nonnative flora. The percentage of species introduced before the end of the 19th century, including archaeophytes (19.6%) and kenophytes (31%), equals that of the eukenophytes (49.4%), the species that were introduced during the 20th century. Leading roles are played by species native to Mediterranean (254 species, or 50.3%) and North American (99 species or 16.3%) floristic regions. Interestingly, North American taxa form the largest portion of the group of species that successfully naturalized in the 20th century; they are followed by plants from eastern Asia. The nonnative plants in the modern urban flora of Kiev continue to gain in importance. This is evident from the high numbers of newcomers that arrived during the last 20 years of the 20th century.

Introduction

The Kiev City Agglomeration (Kiev Urban Area, KUA) is situated in the central part of the eastern European plains, at the border of the forest and forest-steppe physiographic and vegetation zones. Ukraine's capital, the city of Kiev (Kyiv in the Ukrainian-based transliteration), is the natural center of this urbanized area. It is surrounded by several satellite towns and smaller settlements, including Irpin, Brovary, Boryspil, Vyshgorod, and Boyarka. Kiev and its satellites are located on both banks of the Dnipro (Dnieper) River. The area of Kiev within its official administrative borders is 824 square kilometers; its population is currently about 2,616,000 people (according to the 2001 census; other estimates that consider recent migration patterns place it at close to 3 million).

Seminatural and man-made habitats are well represented in KUA, and the region's altered or disturbed plant communities are formed mostly by synanthropic plant species. During their long history in central Ukraine, and the Kiev area in particular, humans have greatly promoted the immigration of nonnative plants through migration, war, trade, agriculture, urbanization, and other activities. The oldest fossil human remains in Kiev date to the late Paleolithic era. In Neolithic times, the area was already home to well-developed agriculture, cattle breeding, and various trade crafts.

By the ninth century A.D., Kiev had become an important political and economic center of eastern Europe, with extensive political and trade contacts in the Baltic and Mediterranean regions, in central and western Europe, western and central Asia, and the Caucasus, as well as in other adjacent and distant areas. In the 18th century, Kiev began to develop as an industrial city. Continued development resulted in the formation of a large urbanized area with a dramatically transformed flora and vegetation.

Nonnative plants are an important component of any modern urban flora. The present checklist of Kiev's nonnative flora was compiled using literature data (Bortnyak, 1978a, 1978b; Bortnyak, et al., 1992; Kotov, 1979; Mosyakin, 1990, 1991a, 1991b, 1995, 1996; Protopopova, 1973, 1991; Yavorska & Mosyakin, 2001; and many others), herbarium collections (mainly the collection of the National Herbarium of Ukraine), and data from our recent field studies (in particular, Mosyakin's collections and observations made in 1985–2002 and Yavorska's collections and studies of 1998–2001). However, because of the dynamics of the process of synanthropization of the flora in the Kiev region, the list cannot be regarded as the final one. It only generalizes the most recent stage of nonnative plant study and remains open for further additions and corrections. The number of nonnative species will grow not only as a result of their casual dispersal and immigration but also due to the deliberate introduction into cultivation of new plants, some of which undoubtedly will be able to adapt to, or naturalize in, the man-altered habitats of KUA.

Families, genera within families, and species within genera are arranged in our table alphabetically. The nomenclature mainly follows the checklist of vascular plants of Ukraine (Mosyakin & Fedoronchuk, 1999). The terminology on synanthropic floras and nonnative plants used in the article follows that of European publications. Please note that this terminology is not commonly used in English-speaking countries. However, explanations and discussion of it can be found in many useful publications (Thellung, 1918; Jäger, 1988; di Castri, 1989; Kornas, 1968, 1990; Protopopova, 1973, 1991; and many others). C. Lambelet-Haueter (1990, 1991) presents especially detailed historical overviews. A good North American introduction to traditional European-style studies of nonnative floras can be found in V. Muhlenbach's classic article (1979) on nonnative plants inhabiting the railroads of St. Louis, Missouri. G. Nesom (2000) provides a more recent discussion on categories of alien plants.

The central and eastern European sources on urban floristics are too numerous to be listed here extensively; therefore we cite only selected publications directly related to the nonnative flora of Kiev or to the methods used in this study. Additional references can be found in the cited books and articles.

Categories and Abbreviations

For the checklist of nonnative species, the following categories and abbreviations were used:

Occurrence of species in the modern nonnative flora of KUA (column 1)
C—Common: plants that are widespread and abundant in the territory of the city and/or in adjacent towns and villages.
D—Disappeared: plants that were reported in early surveys but are now considered extirpated (or in some cases eradicated) in the territory.
E—Ephemera-ergasiophytes: plants introduced by man that occasionally escape beyond cultivation but usually do not persist except in the immediate vicinity of their area of cultivation. These plants do not show pronounced trends toward naturalization, and their occurrence depends on a reliable (stable) source of dissemination (e.g., deriving from plants that are extensively and/or permanently cultivated in the area concerned).
L—Local: plants that only occur in some parts of the city but may be locally abundant.
R—Rare: plants that occur in three to five (sometimes up to seven) localities.
S—Sporadic: plants that occur almost everywhere in the city but not in abundance.
U—Unicates: plants collected in one to three localities in KUA during recent decades but not registered during our 1996–2001 floristic survey of the area.

Distribution of species (column 2)
B—Plants found within the town of Boryspil.
Bo—Plants found within the town of Boyarka.
Br—Plants found within the suburb Brovary.
Ir—Plants found within the town of Irpin.
K—Plants found within the city of Kiev only.
KUA—Plants found within the territory of the city of Kiev and its suburbs.

Degree of naturalization (column 3)
Agr—Agriophytes: naturalized in natural and seminatural habitats.
Col—Colonophytes: epoecophytes that occur in the area in one to several stable colonies but which show little or no trend toward further expansion.
Eph—Ephemerophytes: nonnaturalized species, occasional immigrants, or waifs.
Epo—Epoecophytes: naturalized in man-made and disturbed habitats.
Hagr—Hemiagriophytes: naturalized mostly in seminatural or disturbed habitats.

Note: For extinct species, the degree of naturalization is not indicated.

Time of immigration to KUA (column 4)
arch—Archaeophytes: plants that immigrated before the end of the 15th century.
eu-A—Eukenophytes-A: plants that immigrated in the first half of the 20th century.
eu-B—Eukenophytes-B: plants that immigrated after World War II to the end of the 1970s.
eu-C—Eukenophytes-C: plants that immigrated during the last 20 years.
ken—Kenophytes: plants that immigrated between the 16th century and the end of the 19th century.

Mode of immigration (column 5)
The above groups are segregated according to the traditional classification (see Lambelet-Haueter, 1990, 1991; Protopopova, 1991, et al.), modified by N.A. Vyukova (1985). For archaeophytes, the mode of immigration to KUA is not specified due to the lack of scientifically reliable data. However, we specify the immigration status for a few archaeophytes (14 species of cultivated plants and some specialized weeds of crops) in those rare cases when it is reliably known from archaeobotanical and historical sources. Those species whose nonnative status in the territory of research is questionable (especially when it is unclear whether borders of the native range of a species probably cover, or at least closely approach, our territory) are also provisionally listed below, but their names in column 1 are preceded by a question mark (?).

Erg—Ergasiophytes: plants that were intentionally introduced and cultivated by man, and then spread from places of their cultivation.
Xen—Xenophytes: plants introduced unintentionally.
X-Erg—Xeno-ergasiophytes: plants cultivated outside of the studied area but unintentionally introduced to Kiev.

Initial (original, native) ranges of nonnative species (column 6)
Afr—African; Afr-Sas—African–south Asian; anthr—of anthropogenic origin (taxa that emerged and evolved in man-made habitats); As—Asian; AsM—Asia Minor; CAm—Central American; CaAs—central Asian; Cauc—Caucasian; CEu—central European; EAs—eastern Asian; EMed—eastern Mediterranean; hybr—species of hybrid origin; Ir-An—Irano-Anatolian; Ir-Tr—Irano-Turanian; Med—Mediterranean; Med-Cas—Mediterranean–Irano-Turanian (or Mediterranean–central Asian); Med-Ir-An—Mediterranean–Irano-Anatolian; n/a—initial range uncertain (34 species); NAm—North American; NMed—North Mediterranean; Pon—Pontic; S-Eas—southeastern Asian; SAm—South American; SAs—south Asian; SEu—southern European; Sib—Siberian; WEu—western European; WMed—western Mediterranean.

Checklist of Nonnative Plants of the Kiev Urban Area

Families and Species 1 2 3 4 5 6
ACERACEAE            
Acer negundo L. C KUA Agr eu-A Erg NAm
Acer pseudoplatanus L. S KUA Epo eu-B Erg WEu
Acer saccharinum L. S KV Hagr eu-B Erg NAm
AMARANTHACEAE            
Amaranthus albus L. C KUA Epo eu-A Xen NAm
Amaranthus blitoides S. Watson C KUA Epo eu-B Xen NAm
Amaranthus blitum L. (= A. lividus L.) R K Eph ken Erg S-EAs
Amaranthus caudatus L. R KUA Eph ken Erg SAm
Amaranthus cruentus L. (incl. A. paniculatus L.) E KUA Eph eu-A Erg CAm
Amaranthus hybridus L. s. str. R K Eph eu-C Xen CAm
Amaranthus hypochondriacus L. E K Eph eu-C Erg NAm
Amaranthus palmeri S. Watson U K Eph eu-C Xen NAm
Amaranthus powellii S. Watson L KUA Epo eu-C Xen CAm
Amaranthus retroflexus L. C KUA Epo ken Xen NAm
Amaranthus rudis Sauer U K Eph eu-C Xen NAm
Amaranthus spinosus L. U K Eph eu-C Erg SAm
Amaranthus tuberculatus (Moq.) Sauer U K Eph eu-C Xen NAm
ANACARDIACEAE            
Cotinus coggygria Scop. L K Hagr eu-B Erg Med
Toxicodendron radicans (L.) O. Kuntze E K Eph eu-C Erg NAm
APIACEAE            
? Aethusa cynapium L. C KUA Epo arch   WEu
Anethum graveolens L. L KUA Epo ken Erg Med-CAs
Anthriscus cerefolium (L.) Hoffm. R K Eph eu-B Xen Med
Bifora radians M. Bieb. R K Eph eu-C Xen Med
Bupleurum rotundifolium L. R KUA Eph eu-A   Med-Ir-An
Carum carvi L. E KUA Eph ken Erg n/a
Caucalis platycarpos L. D K Eph eu-A Xen Med
? Conium maculatum L. C KUA Epo n/a Xen Med-CAs
Coriandrum sativum L. E KUA Eph ken Erg Med
Foeniculum vulgare Mill. E KUA Eph ken Erg Med
Heracleum mantegazzianum Sommier & Levier L K Col eu-C Erg Cauc
Levisticum officinale Koch E KUA Col ken Erg Med-CAs
Pastinaca sativa L. E KUA Eph ken Erg Med-CAs
Pastinaca umbrosa Steven ex DC. S K Epo ken Xen Med-CAs
Petroselinum crispum (Mill.) A.W. Hill E KUA Eph ken Erg Med
Turgenia latifolia (L.) Hoffm. U K Eph eu-C Xen Med
APOCYNACEAE            
Apocynum cannabinum L. L KUA Col eu-C Erg n/a
Vinca minor L. L KUA Hagr ken Erg n/a
ARACEAE (incl. ACORACEAE)            
Acorus calamus L. S KUA Agr arch   S-EAs
ASCLEPIADACEAE            
Asclepias syriaca L. S K,Ir Epo ken Erg NAm
ASPHODELACEAE            
Anthericum liliago L. E K Eph ken Erg Med
ASTERACEAE            
Achillea micrantha Willd. D K Eph? eu-A Xen Pon
Acroptilon repens (L.) DC. R K Col eu-C Xen Ir-Tr
Ageratum houstonianum Mill. R KUA Eph eu-B Erg CAm
Ambrosia artemisiifolia L. S KUA Epo eu-B Xen NAm
Ambrosia trifida L. U K Eph eu-C Xen NAm
Anthemis arvensis L. S KUA Epo arch   NMed
Anthemis cotula L. S KUA Eph arch   EMed
? Artemisia absinthium L. S KUA Epo arch   Med-CAs
Artemisia annua L. C KUA Hagr eu-A Xen EAs
Artemisia argyi Levéillé & Vaniot R K Col eu-C Xen EAs
Artemisia dracunculus L. R K Eph eu-C Xen Sub
Artemisia glauca Pallas ex Willd. R K Eph eu-C Xen Sub
Artemisia rubripes Nakai D K Eph eu-C Xen EAs
Artemisia selengensis Turcz. ex Besser D KUA ? eu-C Xen CAs
Artemisia sieversiana Willd. L KUA Epo eu-C Xen Med-CAs
Artemisia tournefortiana Reichb. L K Col eu-C Xen Ir-Tr
Artemisia umbrosa (Turcz. ex Besser) Pamp. R K Eph eu-C Xen EAs
Aster lanceolatus Willd. S KUA Eph ken Erg NAm
Aster novae-angliae L. E KUA Eph ken Erg NAm
Aster novi-belgii L. E KUA Eph ken Erg NAm
Aster x salignus Willd. E KUA Eph ken Erg NAm
Bellis perennis L. E K Eph ken Erg n/a
Bidens connata Muehl. ex Willd. L K Hagr eu-C Xen NAm
Bidens frondosa L. S K Agr eu-B Xen NAm
Calendula officinalis L. E KUA Eph ken Erg Med
Carduus acanthoides L. L KUA Epo arch   NMed
Carduus nutans L. L KUA Epo arch   NMed
Centaurea cyanus L. S KUA Epo arch   Med
Centaurea depressa M.Bieb. R K Eph eu-C Xen Ir-Tr
Centaurea diffusa Lam. R K Epo eu-A Xen Med-CAs
Cichorium endivia L. D K ? ken Erg Med-CAs
? Cichorium intybus L. C KUA Hagr arch   Med-CAs
Cirsium ciliatum (Murr.) Moench U K Eph eu-C Xen Kv
Conyza canadensis (L.) Cronq. (= Erigeron canadensis L.) C KUA Hagr ken Xen NAm
Coreopsis grandiflora Hogg ex Sweet E K Eph eu-C Erg NAm
Cosmos bipinnatus Cav. E KUA Eph eu-C Erg CAm
Galinsoga parviflora Cav. C KUA Epo ken Erg SAm
Galinsoga urticifolia (Kunth) Benth. (G. ciliata auct.) R K Eph eu-C Xen SAm
Grindelia squarrosa (Pursh) Dunal L KUA Col eu-B Xen NAm
Helianthus annuus L. R KUA Eph ken Erg NAm
Helianthus decapetalus L. S KUA Col eu-C Erg NAm
Helianthus rigidus (Cass.) Desf. E K Eph eu-C Erg NAm
Helianthus subcanescens (A.Gray) E. Watson S KUA Col eu-C Erg NAm
Helianthus tuberosus L. L KUA Col ken Erg SAm
Helianthus x laetiflorus Pers. S KUA Col eu-C Erg NAm
Inula helenium L. E KUA Eph eu-A Erg n/a
Iva xanthiifolia Nutt. C KUA Epo ken Erg NAm
Lactuca sativa L. E KUA Eph ken Erg n/a
? Lactuca serriola L. S KUA Epo arch   Med-CAs
Lactuca tatarica (L.) C.A.Mey. L KUA Col eu-B Xen Ir-Tr
Lepidotheca suaveolens (Pursh) Nutt. C KUA Epo ken Xen NAm
Matricaria recutita L. C KUA Epo ken Erg WEu
Onopordon acanthium L. C KUA Epo arch   Med-CAs
Phalacroloma annuum (L.) Dumort. C KUA Epo ken Xen NAm
Phalacroloma septentrionale (Fern. et Wieg.) Tzvelev C KUA Epo eu-B Xen NAm
Pyrethrum partheniifolium Willd. E K Eph eu-B Erg Med
Pyrethrum parthenium (L.) Smith E K Col eu-A Erg WEu
Rudbeckia hirta L. (incl. R. bicolor Nutt.) L KUA Eph ken Erg NAm
Rudbeckia laciniata L. R KUA Col ken Erg NAm
Senecio viscosus L. S KUA Epo eu-B Xen CEv
? Senecio vulgaris L. S KUA Epo arch   Med
Silphium perfoliatum L. L KV Col eu-C Erg NAm
Solidago canadensis L. L KUA Hagr ken Erg NAm
Solidago serotinoides A.Löve & D.Löve (S.gigantea auct.) L KUA Col eu-B Erg NAm
Sonchus arvensis L. L KUA Epo arch   Med
Sonchus asper (L.) Hill S KUA Epo arch   Med
Sonchus oleraceus L. S KUA Epo arch   Med
Tagetes erecta L. E KUA Eph ken Erg CAm
Tagetes patula L. E KUA Eph ken Erg CAm
Tripleurospermum inodorum (L.) Sch.Bip. R KUA Epo arch   EMed
Xanthium albinum (Widder) H.Scholz C KUA Agr eu-B Xen WEu
Xanthium ripicola Holub L KUA Epo eu-B Xen CEv
Xanthium spinosum L. U KUA Eph ken Xen SAm
Xanthium strumarium L. s. str. R KUA Epo arch   Ir-Tr
BALSAMINACEAE            
Impatiens glandulifera Royle (I. roylei Walp.) S K Hagr eu-B Erg S-EAs
Impatiens parviflora DC. C KUA Agr ken Erg Med-CAs
BERBERIDACEAE            
Berberis thunbergii DC. E KUA Col eu-C Erg EAs
Berberis vulgaris L. S KUA Col ken Erg n/a
Mahonia aquifolium (Pursh) Nutt. L KV Agr eu-B Erg NAm
BORAGINACEAE            
Anchusa officinalis L. C KUA Epo arch   Med
Argusia sibirica (L.) Dandy R K,Ir Eph eu-A Xen Med-CAs
Borago officinalis L. R K Epo ken X-Erg Med
Buglossoides arvensis (L.) I.M.Johnst. L KUA Epo arch   Med-CAs
Cynoglosum officinale L. L KUA Epo arch   Med
Echium plantagineum L. D KUA ? eu-A Xen Med
Lappula patula (Lehm.) Menyh. R K Epo eu-A Xen Med-CAs
Lappula squarrosa (Retz.) Dumort. L KUA Epo arch   Med-CAs
? Myosotis arvensis (L.) Hill C KUA Epo arch   Med-CAs
Symphytum asperum Lepechin E K Col ken Erg WMed
BRASSICACEAE            
Armoracia rusticana P.Gaertn., B.Mey. & Scherb. L KUA Agr arch X-Erg Ir-Tr
Brassica campestris L. C KUA Epo arch X-Erg Med-CAs
Brassica juncea (L.) Gzern. L KUA Eph eu-B Xen Ir-Tr
Brassica napus L. R KUA Eph ken Erg SEu
Brassica nigra (L.) W.D.J.Koch S KUA Epo eu-A X-Erg Med
Brassica rapa L. R KUA Eph ken Erg Med
Bunias orientalis L. S KUA Epo ken Xen EMed
Camelina alyssum (Mill.) Thell. D KUA ? arch   ant
Camelina microcarpa Andrz. L K Epo arch   Med-CAs
Camelina sativa (L.) Crantz L KUA Eph arch   ant
? Capsella bursa-pastoris (L.) Medik. C KUA Agr arch   n/a
Capsella orientalis Klokov (C. bursa-pastoris) aggr.) U K Eph eu-B Xen Pon
Capsella rubella Reut. (C. bursa-pastoris aggr.) D K ? eu-B Xen Med
Cardaria draba (L.) Desv. C KUA Epo ken Xen Med
Cardaria pubescens (C.A.Mey.) Jarm. R K Eph eu-C Xen n/a
Chorispora tenella (Pall.) DC. L KUA Epo ken Xen Med-CAs
? Descurainia sophia (L.) Webb. ex Prantl C KUA Epo arch   Med-CAs
Diplotaxis muralis (L.) DC. L KUA Eph eu-A Xen SEu
Diplotaxis tenuifolia (L.) DC. C KUA Epo ken Xen Med
Eruca vesicaria (L.) Cav. (= E. sativa Mill.) R KUA Eph arch X-Erg Med
Erucastrum armoracioides (Czern. ex Turcz.) Cruchet D KUA ? eu-A Xen Ir-Tr
Erysimum cheiranthoides L. L KUA Epo arch   Med-CAs
Erysimum repandum L. S KUA Epo arch   Med-CAs
Euclidium syriacum (L.) R.Br. U K Eph eu-C Xen Ir-Tr
Goldbachia laevigata (M.Bieb.) DC. U K Eph eu-C Xen CAs
Hesperis matronalis L. E KUA Eph ken Erg Ir-Tr
Hirschfeldia incana (L.) Lagr.-Foss. L K Col eu-C Xen EMed
Iberis amara L. E KUA Eph ken Erg Med
Iberis umbellata L. E KUA Eph ken Erg Med
Isatis tinctoria L. D K ? ken Xen Med-CAs
Lepidium campestre (L.) R.Br. R K Eph arch   Med
Lepidium densiflorum Schrad. C KUA Epo ken Xen NAm
Lepidium latifolium L. s.l. (incl. L. affine Ledeb.) L KUA Col ken Xen Ir-Tr
Lepidium perfoliatum L. L KUA Epo ken Xen Med-CAs
Lepidium ruderale L. S KUA Epo arch   Med-CAs
Lobularia maritima (L.) Desv. E KUA Eph ken Erg Med
Matthiola bicornis (Sibth. & Sm.) DC. E KUA Eph ken Erg Med
Matthiola incana (L.) R. Br. E KUA Eph ken Erg Med
Matthiola longipetala (Vent.) DC. E KUA Eph ken Erg Med
Myagrum perfoliatum L. R K Eph eu-C Xen Med
Neslia paniculata (L.) Desv. R K Eph arch   ant
? Raphanus candidus Worosch. R KUA Epo eu-B Xen hybr
Raphanus raphanistrum L. S KUA Epo arch   Med
Raphanus sativus L. R KUA Eph ken Erg Med
Rapistrum perenne (L.) All. R KUA Col eu-A Xen Med
Rapistrum rugosum (L.) Bergeret R KUA Eph eu-A Xen Med
Sinapis alba L. S KUA Eph arch X-Erg Med
Sinapis arvensis L. S KUA Eph arch   Med
Sinapis dissecta Lag. R K Eph eu-A Xen Med
Sisymbrium altissimum L. C KUA Epo ken Xen Med-CAs
Sisymbrium loeselii L. C KUA Epo ken Xen Med-CAs
? Sisymbrium officinale (L.) Scop. C KUA Epo arch   Med
Sisymbrium orientale L. R K Eph eu-C Xen Med
Sisymbrium volgense M.Bieb. ex Fourn. S KUA Epo eu-A Xen Pon
Thlaspi arvense L. C KUA Epo arch   Ir-Tr
CAESALPINIACEAE            
Gleditsia triacanthos L. R KUA Col ken Erg NAm
CAMPANULACEAE            
Campanula medium L. E KUA Eph eu-C Erg n/a
CANNABACEAE            
Cannabis sativa L. s.l. (incl. C. ruderalis Janisch.) C KUA Epo arch   Ir-Tr
CAPRIFOLIACEAE            
Lonicera caprifolium L. L K Col eu-A Erg Med
Lonicera tatarica L. L KUA Col eu-A Erg Sub
Symphoricarpos albus (L.) S.F.Blake L KUA Col eu-B Erg NAm
Viburnum lantana L. L K Col eu-A Erg Med
CARYOPHYLLACEAE            
Agrostemma githago L. U K Eph arch   ant
Dianthus barbatus L. E KUA Eph eu-B Erg Med
Dianthus euponticus Zapał E K,Ir Eph eu-B Xen EMed
Gypsophila perfoliata L. L K Eph eu-B Xen Ir-Tr
Lychnis chalcedonica L. S KUA Epo ken Erg As
Oberna cserei (Baumg.) Ikonn. D K ? eu-A Xen Med
Petrorhagia saxifraga (L.) Link E K Eph ken Erg Med
Saponaria officinalis L. S KUA HAgr ken X-Erg Med
Scleranthus annuus L. S KUA Epo arch   WMed
Silene armeria L. R K Eph ?   n/a
Silene dichotoma Ehrh. R K Eph eu-A Xen WEu
Silene gallica L. S KUA Epo eu-B Xen WMed
Silene pendula L. S KUA Epo ken Erg Med
Spergula arvensis L. S KUA Epo ken Xen Med
Spergula maxima Weihe D KUA ? arch   ant
Spergula morisonii Boreau R K,Br Eph eu-B Xen WEu
Vaccaria hispanica (Mill.) Rauschert U KUA Eph arch   S-EAs
CHENOPODIACEAE            
Atriplex hortensis L. L KUA Epo ken Erg Ir-Tr
Atriplex micrantha C.A. Mey.(=A. heterosperma Bunge) R K Eph ken Xen Ir-Tr
Atriplex rosea L. R KUA Eph eu-B Xen Med
Atriplex sagittata Borkh. (= A. nitens Schkuhr) C KUA Epo arch   Ir-Tr
? Atriplex tatarica L. C KUA Hagr arch   Med-CAs
Ceratocarpus arenarius L. O K Eph eu-C Xen Med-CAs
Chenopodium berlandieri Moq. R K Eph eu-C Xen NAm
Chenopodium botrys L. R KUA Eph eu-A X-Erg Med-CAs
Chenopodium chenopodioides (L.) Aellen R K Eph eu-C Xen Med-CAs
Chenopodium ficifolium Smith S KUA Epo eu-A Xen EAs
Chenopodium foliosum Asch. U K Eph eu-C Erg Med
Chenopodium glaucophyllum Aellen U K Eph eu-C Xen NAm
Chenopodium hybridum L. S KUA Epo n/a   Res
Chenopodium missouriense Aellen U K Eph eu-C Xen NAm
Chenopodium murale L. U K Eph arch   Med
Chenopodium opulifolium Schrader ex DC. S KUA Epo arch   Med
? Chenopodium polyspermum L. S KUA Epo arch   n/a
Chenopodium pratericola Rydb. U K Eph eu-C Xen NAm
Chenopodium probstii Aellen R K Eph eu-C Xen n/a
Chenopodium reticulatum Aellen U K Eph eu-C Xen n/a
Chenopodium schraderianum Schultes R K Eph eu-C Xen Afr
Chenopodium striatiforme J. Murr S KUA Epo n/a Xen Med
Chenopodium strictum Roth C KUA Epo n/a Xen NAm
Chenopodium urbicum L. R K Eph n/a Xen Med-CAs
Chenopodium vulvaria L. D K ? ken Xen Med-CAs
Corispermum declinatum Steph. ex Iljin R K Eph eu-C Xen EAs
Corispermum pallasii Steven (C. leptopterum auct.) S K Epo eu-C Xen Sub
Corispermum redowskii Fisch. ex Steven R K,Br Eph eu-B Xen Sub
Kochia scoparia (L.) Schrader s.l. S KUA Epo ken Erg Ir-Tr
Polycnemum arvense L. R K Eph ken Xen Med-CAs
Salsola collina Pallas L K,Ir Epo eu-C Xen Med-CAs
Salsola tragus L. s. str. (= S. ruthenica Iljin) C KUA Epo ken Xen Ir-Tr
Spinacia oleracea L. S KUA Eph ken Erg Med-CAs
COMMELINACEAE            
Commelina communis L. S KUA Epo eu-A Erg S-EAs
Tradescantia virginiana L. E K Eph eu-C Erg NAm
CONVOLVULACEAE            
Calystegia spectabilis (Brummitt) Tzvelev (C. inflata auct.) E K Eph eu-C Erg NAm
Ipomoea hederacea (L.) Jacq. E K Eph eu-C Xen CAm
Ipomoea purpurea (L.) Roth S KUA Eph eu-B Erg SAm
CRASSULACEAE            
Sedum rupestre L. (incl. S. reflexum L.) R K Col eu-B Erg Kv
Sedum spurium M. Bieb. R K Col eu-A Erg Kv
CUCURBITACEAE            
Bryonia alba L. S KUA Epo ken Erg Med-Cas
Citrulus lanatus (Thunb.) Matsum. & Nakai E KUA Eph ken Erg NAm
Cucurbita pepo L. E KUA Eph ken Erg NAm
Echinocystis lobata (Michx.) Torr. & A.Gray S KUA Agr eu-B Erg NAm
Sicyos angulata L. E K Col ken Erg NAm
Thladiantha dubia Bunge E K Col eu-A Erg EAs
CUSCUTACEAE            
Cuscuta campestris Yunck. S KUA Epo eu-B Xen NAm
Cuscuta gronovii Willd. ex Roemer & Schultes R K Eph eu-B Xen NAm
EQUISETACEAE            
Equisetum ramosissimum Desf. R KUA Col eu-B Xen n/a
EUPHORBIACEAE            
Euphorbia dentata Michx. L K Epo eu-C Xen NAm
Euphorbia falcata L. L K Epo arch   Med-CAs
Euphorbia helioscopia L. S KUA Epo arch   Med
Euphorbia marginata Pursh R KUA Eph eu-C Erg NAm
Euphorbia peplus L. S KUA Epo arch   Med
Flueggea suffruticosa (Pallas) Baillon L K Col eu-C Erg EAs
FABACEAE            
Amorpha fruticosa L. L KUA Agr eu-B Erg NAm
Astragalus onobrychis L. R KUA Col ken Erg Med-CAs
Caragana arborescens Lam. L KUA Hagr eu-B Erg Sub
Caragana frutex (L.) K.Koch E KUA Col eu-B Erg Ir-Tr
Lathyrus odoratus L. R KUA Eph ken Erg Med
Lathyrus sativus L. E KUA Eph eu-C Erg Med
Lathyrus tuberosus L. L KUA Hagr ken Erg Med-CAs
Lens culinaris Medik. E KUA Eph ken Erg Med
Lupinus polyphyllus Lindl. L KUA Hagr eu-C Erg NAm
Medicago minima (L.) Bartal. U KUA Eph ken Xen n/a
Medicago sativa L. C KUA Epo ken Erg AsM
Onobrychis viciifolia Scop. R KUA Eph ken Erg SEu
Ornithopus sativus Brot. R KUA Eph eu-C Erg WMed
Phaseolus vulgaris L. E KUA Eph ken Erg SAm
Pisum sativum L. E KUA Eph ken Erg Ir-Tr
Robinia pseudoacacia L. C KUA Hagr eu-A Erg NAm
? Trifolium hybridum L. S KUA Epo eu-A Erg Med
Trifolium incarnatum L. E KUA Eph eu-A Erg EMed
Trifolium resupinatum L. R K Eph eu-C Xen Med-CAs
Trifolium sativum (Schreber) Crome R KUA Eph eu-A Erg WEu
Trigonella caerulea (L.) SÈr. L KUA Eph ken Erg Med
Vicia angustifolia Reichard L KUA Eph ken Xen Med-CAs
Vicia faba L. E KUA Eph ken Erg Ir-Tr
Vicia hirsuta (L.) S.F.Gray L KUA Eph arch   WMed
Vicia sativa L. L KUA Eph ken Erg hybr
Vicia tetrasperma (L.) Schreber S KUA Eph arch   Med
Vicia villosa Roth s.l. (incl. V.sordida Waldst. & Kit.) R KUA HAgr arch   Med
FAGACEAE            
Quercus palustris Moench E K Col eu-C Erg NAm
Quercus rubra L. (= Q. borealis Michx.) S KV Hagr eu-B Erg NAm
FUMARIACEAE            
Fumaria officinalis L. S KUA Epo arch   Med
Fumaria schleicheri Soy.-Willem. L KUA Epo arch   Ir-Tr
Fumaria vaillantii Loisel. L KUA Epo eu-A Xen Med-CAs
GERANIACEAE            
Erodium cicutarium (L.) LíHÈr. C KUA Eph arch   Med-CAs
Geranium dissectum L. L KUA Epo arch   Med
Geranium molle L. D K ? ken Xen Med
Geranium pusillum L. S KUA Epo arch   Ir-An
Geranium pyrenaicum Burm. f. L K Epo ken Xen Med
Geranium sibiricum L. C KUA Epo ken Erg Ir-Tr
HEMEROCALLIDACEAE            
Hemerocallis fulva (L.) L. E KUA Eph eu-B Erg EAs
Hemerocallis lilioasphodelus L. E KUA Eph eu-B Erg EAs
HYDROCHARITACEAE            
Elodea canadensis Michx. C KUA Agr ken Xen NAm
HYDROPHYLLACEAE            
Phacelia tanacetifolia Benth. E K Eph ken Erg NAm
IRIDACEAE            
Iris florentina L. E KUA Eph ken Erg Med
Iris germanica L. E KUA Eph ken Erg WEu
Limoniun meyeri (Boiss.) O.Kuntze R K Col eu-C Xen Med
Sisyrinchium septentrionale Bicknell R K Eph ken Erg NAm
JUGLANDACEAE            
Juglans mandshurica Maxim. L KUA Col eu-C Erg EAs
Juglans regia L. S KUA Eph eu-C Erg EAs
JUNCACEAE            
Juncus tenuis Willd. (= J. macer S.F.Gray) C KUA Hagr ken Xen NAm
LAMIACEAE

           
? Ballota nigra L. s.l. C KUA Hagr arch   Med
Dracocephalum thymiflorumL. D K ? ken Erg EAs
Elsholtzia ciliata(Thunb.) Hyl. S K Epo eu-A Xen EAs
Ga leopsis ladanum L. L KUA Epo arch   NMed
Lamium album L. s.l. L KUA Epo arch   CAs
Lamium amplexicaule L. L KUA Epo arch   Med-CAs
? Lamium purpureum L. C KUA Hagr arch   Med
Marrubium vulgare L. R KUA Eph arch   Med-CAs
Melissa officinalis L. E KUA Eph ken Erg n/a
Mentha arvensis L. E KUA Eph ken Erg n/a
Mentha spicata L. E KUA Epo ken X-Erg Med
Mentha x piperita L. E KUA Eph ken Erg n/a
Nepeta cataria L. L KUA Epo arch   EMed
Salvia reflexa Hornem. R K Eph eu-C Xen NAm
Sideritis montana L. R K Epo ken Xen Med
Stachys annua (L.) L. S KUA Epo arch   WMed
LINACEAE            
Linum ustitatissimum L. E KUA Eph arch Erg As
MALVACEAE            
Abutilon theophrastii Medik. L K Epo ken Erg EAs
Alcea rosea L. E KUA Col ken Erg n/a
Hibiscus syriacus L. E K Eph eu-C Erg n/a
Hibiscus trionum L. D K ? eu-B Xen Med
Malva crispa (L.) L. L KUA Epo ken Erg EAs
Malva excisa Reichenb. L KUA Eph arch X-Erg Pon
Malva mauritiana L. E KUA Eph ken Erg Med
Malva neglecta Wallr. C KUA Epo arch X-Erg Ir-Tr
Malva pusilla Smith. C KUA Epo arch X-Erg Res
Malva sylvestris L. L K Hagr arch X-Erg Med
Sida rhombifolia L. U K Eph eu-C Xen S-EAs
MORACEAE            
Morus alba L. S KUA Epo ken Erg EAs
NYCTAGINACEAE            
Oxybaphus nyctagineus (Michx.) Sweet S KUA Epo ken X-Erg NAm
OLEACEAE            
Fraxinus lanceolata Borkh. R K Col eu-B Erg NAm
Fraxinus pennsylvanica Marshall S K Epo eu-B Erg NAm
Ligustrum vulgare L. E KUA Col eu-C Erg n/a
Syringa vulgaris L. E K Col ken Erg EMed
ONAGRACEAE            
Epilobium ciliatum Raf. s.l. (=E. adenocaulon Hausskn.) C K Agr eu-B Xen NAm
Oenothera biennis L. s.l. L KUA Epo ken Xen NAm
Oenothera laciniata Hill O K Eph eu-C Xen NAm
Oenothera rubricaulis Klebahn C KUA Hagr eu-A Xen WEu
Oenothera oakesiana (A. Gray) S. Watson & Coulter R K Eph ken Xen NAm
Oenothera villosa Thunb. s.l. C K Epo eu-B Erg NAm
OROBANCHACEAE            
Orobanche cernua Loefl. R KUA Eph ken Xen ant
Phelipanche ramosa(L.) Pomel (Orobanche ramosa L.) L K Eph ken Xen Med-CAs
OXALIDACEAE            
Xanthoxalis dillenii (Jacq.) Holub L KUA Epo eu-B Xen NAm
Xanthoxalis stricta (L.) Small C KUA Hagr ken Xen NAm
PAPAVERACEAE            
Glaucium corniculatum (L.) J. Rudolph R B Eph ken Xen Med
Papaver dubium L. S KUA Eph arch   Med-CAs
Papaver ocellatum Woronow U K Eph eu-C Xen Ir-Tr
Papaver rhoeas L. C KUA Epo arch   Med-CAs
Papaver somniferum L. D KUA ? ken ErgMed EMed
POACEAE (= GRAMINEAE)            
Agropyron cristatum (L.) P.Beauv. s. str. R K Eph eu-C Xen Ir-Tr
Agropyron pectinatum (M.Bieb.) P.Beauv. R KUA Eph arch   Ir-Tr
Alopecurus myosuroides Huds. L K Eph eu-C Xen Med-Ir-An
Anisantha sterilis (L.) Nevski (= Bromus sterilis L.) R K Eph eu-B Xen Med-Ir-An
Anisantha tectorum (L.) Nevski (= Bromus tectorum L.) S KUA Epo arch   Med-CAs
Apera spica-venti (L.) P.Beauv. C KUA Agr arch   n/a
Arrhenatherum elatius (L.) J.Presl & C.Presl R KUA Eph ken Erg WEu
Avena fatua L. U KUA Eph arch Xen Ir-Tr
Avena nuda L. D KUA ? ken Erg WEu
Avena sativa L. L KUA Eph arch Erg SEu
Avena strigosa Schreb. D K ? eu-A Xen ant
Beckmannia syzigachne (Steud.) Fern. U K Eph eu-C Xen NAm
Bromus arvensis L. C KUA Epo arch   NMed
Bromus commutatus Schrad. D KUA ? ken Xen WEu
Bromus hordeaceus L. C KUA Hagr arch   NMed
Bromus japonicus Thunb. S KUA Epo ken Xen Med
Bromus secalinus L. D KUA ? arch   ant
Bromus squarrosus L. S KUA Epo ken Xen Med-CAs
Cenchrus longispinus (Hack.) Fern. S K Epo eu-C Xen NAm
Ceratochloa carinata(Hook. & Arn.) Tutin L K Col eu-C X-Erg NAm
Ceratochloa cathartica (M.Vahl.) Herter R K Eph eu-C X-Erg SAm
Cynodon dactylon (L.) Pers. R K Col eu-B Xen Med-CAs
Digitaria ciliaris (Retz.) Koeler U K Eph eu-C Xen Med
Digitaria pectiniformis (Henrard) Tzvelev R K Eph ken Xen Med
Digitaria sanguinalis (L.)Scop. C KUA Agr arch   S-EAs
Echinochloa colona (L.) Link U K Eph eu-C X-Erg SAs
? Echinochloa crusgalli(L.) P.Beauv. s.l. C KUA Hagr arch   S-EAs
Echinochloa esculenta(A.Br.) H.Scholz U K Eph eu-C X-Erg EAs
Echinochloa frumentacea Link U K Eph eu-C Xen S-EAs
Echinochloa microstachya (Wiegand) Rydb. L K Col eu-C Xen NAm
Echinochloa oryzicola (Vasing.) Vasing. U K Eph eu-C Xen EAs
Echinochloa oryzoides (Ard.) Fritsch U K Eph eu-C Xen S-EAs
Echinochloa wiegandii(Fassett) McNeill & Dore U K Eph eu-C Xen NAm
Eleusine indica (L.) Gaertn. U K Eph eu-C Xen Afr-SAs
Elymus sibiricus L. U K Eph eu-C Xen EAs
Elymus trachycaulus (Link) Gould & Schinners R K Eph eu-C X-Erg NAm
Eragrostis cilianensis (All.) Vign. ex Janchen R K Eph eu-C Xen Med
Eragrostis minor Host C K Hagr ken Xen Med-CAs
Eragrostis multiflora Steudel R K Eph eu-C Xen S-EAs
Eragrostis pectinacea (Michx.) Nees S KUA Epo eu-C Xen NAm
Eragrostis pilosa (L.) P.Beauv. C KUA Hagr ken Xen EAs
Eremopyrum orientale (L.) Jaub. & Spach R K Eph eu-C Xen Ir-Tr
Hordeum distichon L. R KUA Eph ken Erg Ir-Tr
Hordeum jubatum L. L K Col eu-C Xen NAm
Hordeum leporinum Link R K Eph eu-A Xen Med
Hordeum murinum L. L K Col eu-A Xen Med-CAs
Hordeum vulgare L. E KUA Eph ken Erg Ir-Tr
Lolium multiflorum Lam. S KUA Eph ken Erg Med-CAs
Lolium persicum Boiss. & Hohen. R K Eph eu-C Xen Ir-Tr
Lolium remotum Schrank D KUA ? arch   ant
Lolium temulentum L. R KUA ? arch   NMed
Panicum capillare L. s.l. L K Eph eu-C Xen NAm
Panicum dichotomiflorum Michx. R K Eph eu-C Xen NAm
Panicum miliaceum L. s.l. E KUA Eph ken Erg EAs
Phalaris canariensis L. E K Eph ken Erg Med
Puccinellia hauptiana V.Krecz. U K Eph eu-C Xen CAs
Puccinellia nuttalliana (Schult.) A.Hitchc. U K Eph eu-C Xen NAm
Puccinellia poecilantha(K.Koch) Grossh. R K Eph eu-C Xen Ir-Tr
Puccinellia tenuissimaLitv. ex V.Krecz. R K Eph eu-C Xen Med-CAs
Sclerochloa dura (L.) P.Beauv. L K Eph arch   Med-CAs
Secale cereale L. E KUA Eph arch Erg AsM
Secale sylvestre Host R K Eph eu-B Xen n/a
Setaria adhaerens (Forssk.) Chiov. R K Eph eu-C Xen Afr-SAs
Setaria faberiF.Herrmann s.l. U K Eph eu-C Xen EAs
? Setaria glauca(L.) P.Beauv. (S. pumila auct.) C KUA Hagr arch   S-EAs
Setaria italica(L.) P.Beauv. S KUA Eph ken Erg EAs
Setaria pycnocoma(Steud.) Henrard ex Nakai L KUA Epo eu-C Xen EAs
Setaria verticilliformis Dumort. R K Eph arch   Med
? Setaria viridis (L.) P.Beauv. S KUA Hagr arch   Med-CAs
Sorghum bicolor (L.) Moench s. str. U K Eph eu-C X-Erg SAs
Sorghum cernuum (Ard.) Host U K Eph eu-C X-Erg SAs
Sorghum halepense (L.) Pers. L K Col eu-C X-Erg Med
Sorghum saccharatum (L.) Moench R K Eph ken X-Erg SAs
Sorghum sudanense (Piper) Stapf E K Eph eu-C Xen Afr
Trisetum sibiricum Rupr. R K Eph n/a   n/a
Triticum aestivum L. R KUA Eph ken Erg Ir-Tr
Triticum durum Desf. R KUA Eph ken Erg Ir-Tr
Zea mays L. E KUA Eph ken Erg CAm
POLEMONIACEAE            
Phlox paniculata L. E KUA Col eu-B Erg NAm
Phlox subulata L. E KUA Col eu-B Erg NAm
POLYGONACEAE            
Fagopyrum esculentum Moench S KUA Eph ken Erg EAs
Fagopyrum tataricum (L.) P. Gaertn. R KUA Eph ken Xen EAs
? Fallopia convolvulus (L.) A. Lˆve. C KUA Epo arch   As
Persicaria orientali (L.) Spach E K Eph eu-C Erg S-EAs
Polygonum ramosissimum Michx. U K Eph eu-C Xen NAm
Reynoutria japonica Houtt. S K Epo eu-C Erg EAs
R. sachalinensis (F.Schmidt ex Maxim.) Nakai R K Epo eu-C Erg EAs
Rumex longifolius DC. R KUA Epo ken X-Erg n/a
Rumex patientia L. R KUA Eph ken Erg Med
Rumex stenophyllus Ledeb. S K Epo eu-B Xen Med-CAs
Rumex triangulivalvis (Danser) Rech.f. L K Col eu-C Xen NAm
PORTULACACEAE            
Portulaca grandiflora Hook. E KUA Eph eu-C Erg hybr
Portulaca oleracea L. S K Epo arch   Med-CAs
PRIMULACEAE            
Anagallis arvensis L. C KUA Epo arch   Med-CAs
RANUNCULACEAE            
Adonis aestivalis L. U K Eph eu-A Xen Med-CAs
Adonis annua L. D K ? ken Xen Med
Adonis flammeus Jacq. D K ? ken Xen Med-CAs
Aquilegia vulgaris L. E KUA Col ken Erg WEu
Clematis jackmannii Moore E K Col eu-C Erg hybr
Clematis viticella L. E K Col eu-C Erg Kv
Consolida ajacis (L.) Schur E KUA Eph ken Erg n/a
Consolida orientalis (J.Gay) Schroedinger U K Eph eu-C Xen Med
Consolida paniculata (Host) Schur U K Eph eu-C Xen Med
Consolida regalisS.F.Gray S KUA Epo arch   Med-CAs
Nigella arvensis L. E KUA Eph eu-B Xen EMed
Nigella damascena L. E KUA Eph ken Erg Med
Nigella sativa L. E K Eph ken Erg Med-CAs
Nigella segetalis M.Bieb. U K Eph eu-C Xen EMed
Ranunculus arvensis L. U K Eph eu-C Xen Med-CAs
RESEDACEAE            
Reseda lutea L. S KUA Epo eu-A Xen Med
Reseda luteola L. D KUA ? arch X-Erg Med
ROSACEAE            
Amelanchier ovalis Medik. E KUA Col eu-B Erg Med
Armeniaca vulgaris Lam. E KUA Col ken Erg EAs
Aronia melanocarpa(Michx.) Elliot s.l. R KUA Col eu-C Erg NAm
Cerasus mahaleb (L.) Mill. E KUA Col eu-B Erg EMed
Cerasus vulgaris Mill. E KUA Col eu-B Erg EMed
Cotoneaster melanocarpus Fisch. ex Blytt R KUA Col eu-B Erg Ir-Tr
Fragaria x ananassa(Duchesne) Duchesne E KUA Col ken Erg hybr
Malus baccata (L.) Borkh. E KUA Col ken Erg Sub
Malus domestica Borkh. E KUA Col ken Erg n/a
Padus serotina (Ehrh.) Ag. (= Prunus serotina Ehrh.) L KUA Agr eu-B Erg NAm
Physocarpus opulifolius (L.) Maxim. L K Col eu-B Erg NAm
Potentilla longifolia Willd. ex Schlecht. R K Col eu-C Xen As
Potentilla paradoxaNutt. ex Torr. & A.Gray S K Epo eu-C Xen As
Potentilla tergemina Soj·k D K ? eu-C Xen EAs
Poterium polygamum Waldst. & Kit. L KUA Epo eu-B Xen Med-CAs
Poterium sanguisorba L. S KUA Epo eu-B Xen Med-CAs
Rosa rugosa Thunb. E KUA Col ken Erg EAs
Sorbaria sorbifolia(L.) A.Braun E KUA Col ken Erg n/a
Spiraea chamaedryfolia L. E K Col ken Erg n/a
Spiraea douglasii Hook. s.l. L K Col eu-C Erg NAm
Spiraea salicifolia L. L KUA Epo eu-C Erg Sub
RUBIACEAE            
? Galium exoletum Klokov U K Eph eu-B Xen WEu
? Galium spurium L. R K Eph eu-A Xen ant
Sherardia arvensis L. R KUA Eph arch   WMed
RUTACEAE            
Ptelea trifoliata L. L K Hagr eu-B Erg NAm
SALICACEAE            
Populus balsamifera L. L K Col eu-C Erg NAm
Populus bolleana Lauche L K Col eu-C Erg Med-CAs
Populus deltoides Marshall L K Col eu-C Erg Med-CAs
Salix fragilis L. C KUA Agr arch   Med-CAs
SCROPHULARIACEAE            
Antirrhinum majus L. E KUA Eph ken Erg Med
Veronica cardiocarpa(Kar. & Kir.) Walp. L K Epo eu-C Xen Ir-Tr
Veronica filiformis Smith L K Eph eu-C Erg Kv
Veronica hederifolia L. S KUA Epo eu-A Xen Med
Veronica persica Poir. L KUA Epo ken Xen Ir-Tr
Veronica polita Fr. S KUA Epo arch   Med-CAs
SIMAROUBACEAE            
Ailanthus altissima (Mill.) Swingle R K Epo eu-C Erg EAs
SOLANACEAE            
Datura stramonium L. L KUA Epo ken Xen As
Hyoscyamus niger L. S KUA Epo ken Xen Ir-An
Lycium barbarum L. S K,Bo Hagr arch Erg EAs
Lycopersicon esculentum Mill. (Solanum lycopersicum L.) E KUA Eph ken Erg SAm
Nicandra physalodes (L.) P.Gaertn. E K Epo ken Erg SAm
Nicotiana alata Link & Otto E KUA Eph ken Erg Cam
Petunia x atkinsianaD. Don. ex Loudon E KUA Eph ken Erg hybr
Physalis alkekengi L. E K Eph eu-C Erg Med
Physalis ixocarpa Brot. ex Hornem. E KUA Eph eu-C Erg CAm
Solanum alatum Moench L KUA Epo ken Xen WEu
Solanum carolinense L. R K Eph eu-C Xen NAm
Solanum schultesii Opiz R K Eph ken Xen Med
Solanum tuberosum L. E KUA Eph ken Erg SAm
THYMELAEACEAE            
Thymelaea passerina(L.) Coss. & Germ. D KUA ? arch   Med-CAs
TYPHACEAE            
Typha laxmannii Lepech. L K Col eu-B Xen Med-CAs
ULMACEAE            
Celtis occidentalis L. E K Col eu-C Erg EAs
Ulmus pumila L. S K Hagr eu-C Erg EAs
URTICACEAE            
Parietaria officinalis L. E K Eph ? Erg Med
Urtica urens L. S KUA Epo arch   Med
VERBENACEAE            
Verbena officinalis L. S KUA Epo arch   Med-Ir-An
VITACEAE            
Parthenocissus inserta (A.Kerner) Fritsch C KUA Hagr eu-B Erg NAm
Parthenocissus quinquefolia (L.) Planch. L K Col eu-B Erg NAm
Vitis labrusca L. L KUA Eph eu-C Erg n/a
Vitis vinifera L. L KUA Eph eu-C Erg n/a
ZYGOPHYLLACEAE            
Tribulus terrestris L. L K Eph eu-A Xen Med

Results and Analysis

According to the data, the total nonnative flora (past and present) of KUA consists of 536 species of vascular plants belonging to 297 genera and 71 families. The flora has the highest degree of diversity of any urban nonnative flora in Ukraine. The modern nonnative flora, confirmed for the area in 1997–2001, consists of 356 species belonging to 207 genera and 62 families. We did not include in this group plants that are now considered extinct in the territory of research (28 species), recent unstable introductions (occasionally introduced during the last 20 years), plants not confirmed with new collections during the 1997–2001 study (48 species), and Ephemero-ergasiophytes (104 species). The stable component of the modern nonnative flora includes only effectively naturalized species (agriophytes, hemiagriophytes, and epoecophytes). This component is represented by 198 species belonging to 147 genera and 51 families.

The observations indicate that 63 species (17.7% of the total number of modern nonnative plants) are widespread over practically the whole territory of KUA, and that they are stable and abundant components of disturbed habitats. The majority of the nonnative species analyzed (108 species, or 30.3%) occur infrequently, 99 species (27.8%) occur locally, 86 species (24.2%) are sporadic, and at least 37 species from these groups display pronounced trends toward further dispersal and invasions within the studied area.

In examining the taxonomic structure of the nonnative flora, we found that, unlike the native regional flora, just a few families contain a considerable portion of the species (Asteraceae, Poaceae, and Brassicaceae comprise 38.8% of species; 6 leading families comprise 54.1%; and 15 leading families, 76.7%). This is typical of other nonnative floras (Protopopova, 1991; Vyukova, 1985; Mosyakin & Yavorska, 2001, etc.). In its taxonomic spectrum, the studied flora proved to be more similar to other nonnative floras of the Palaearctics than to the native flora of the Kiev region, where it is physically located. In contrast to the native flora of KUA, the role of the families Brassicaceae, Chenopodiaceae, and Polygonaceae is somewhat elevated, as is the role of Poaceae, Asteraceae, and Rosaceae. The families Asteraceae, Onagraceae, Rosaceae, and Polygonaceae play leading roles in forming the stable component of the alien flora.

Our analysis of nonnative plants by their degree of naturalization has shown that ephemerophytes (175 species, or 49.2%) and epoecophytes (99 species, or 28.2%) clearly prevail in the structure of the modern nonnative flora of Kiev. Smaller numbers of species are represented by colonophytes (42 species, or 11.7%) and agriophytes (21 species, or 5.7%). The group of typical agriophytes is smallest (19 species, or 5.2%). Thus, ephemerophytes and epoecophytes are over three times more numerous than agriophytes, hemiagriophytes, and colonophytes combined, which again is rather typical of nonnative floras. These data testify to instability in the present nonnative component of KUA's flora as it goes through an intensive period of formation. The nonnative fraction of the Kiev flora is characterized by a much higher level of instability in species composition compared to floras of satellite cities. This is due to increased numbers of recently introduced ephemerophytes and ergasiophytes in KUA. Nevertheless, there is some degree of stabilization in the species composition. The ratio of stable to unstable components in the structure of the modern nonnative flora is 54% to 46%, respectively.

The percentage of species introduced before the end of the 19th century, which includes archaeophytes (19.6%) and kenophytes (31%), is roughly equal to that of eukenophytes (49.4%), species that were introduced during the 20th century. According to the obtained data, the last century saw the number of nonnative species swell at an ever-increasing rate. Though eukenophytes-C immigrated to our territory within the shortest time span (20 years) of the three eukenophyte subgroups, they constitute 55.8% of the total number of eukenophytes (subgroups A and B constitute 17.3% and 26.9%, respectively).

In mode of immigration, xenophytes (47.8%) and ergasiophytes (46.5%) dominate. Xeno-ergasiophytes constitute only 5.7% of the whole nonnative flora. Taking into account the increased role (both in the number of species and their growing participation in plant communities) of ergasiophytes in the structure of our nonnative flora and vegetation, it would be wise to pay more attention to the naturalization ability of newly introduced cultivated plants. There are numerous cases of uncontrollable invasions by some of these plants in our city and adjacent areas.

In the formation of the total nonnative flora of KUA, the leading role is played by species native to the ancient Mediterranean (254 species; or 50.3%) and North American (99 species, or 16.3%) floristic regions. The American group (122 species, or 21.5%) includes taxa native to both North and South America. Much fewer of our nonnative species immigrated from Asia (81 species, or 19%), Europe (22 species, or 4.7%), Caucasus (6 species, 1.3%), and other regions (3.2%). Interestingly, North American taxa form the largest portion of the group of species that successfully naturalized in the 20th century (36% of all eukenophytes). They are followed in that group by eastern Asian plants (26% of all eukenophytes).

Nonnative plants are gaining in importance in the modern urban flora of Kiev. In general, in the second half of the 20th century, the number of new nonnative plants increased tenfold compared with the number of taxa registered from the end of the 19th to the middle of the 20th century. The number of completely naturalized species at least doubled. Finally, since more than half of the century's nonnative species immigrated only within the last two decades, we can expect further growth in the number of new nonnative plants.

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Glossary

Agriophytes: Naturalized in natural and seminatural habitats.

Archaeophytes: Plants that immigrated before the end of the 15th century.

Colonophytes: Epoecophytes that occur in the area in one to several stable colonies but which show little or no trend toward further expansion.

Ephemerophytes: Nonnaturalized, occasional immigrants, or waifs.

Epoecophytes: Naturalized in man-made and disturbed habitats

Ergasiophytes: Plants that were intentionally introduced and cultivated by man, and then spread from places of their cultivation.

Eukenophytes: Plants that immigrated in the 20th century.

Hemiagriophytes: Naturalized mostly in seminatural or disturbed habitats.

Kenophytes: Plants that immigrated between the 16th century and the end of the 19th century.

Palaearctic: Found in the arctic regions of the Old World.

Xenophytes: Plants introduced unintentionally.

Xeno-ergasiophytes: Plants cultivated outside of the studied area but unintentionally introduced.

Synanthropic: Living in close association with humans.