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.