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Abstract: 7KHUHVXOWVRIDQLQYHVWLJDWLRQRIWKHHUJDVLRSK\WHV VSHFLHVGHOLEHUDWHO\LQWURGXFHGWRDUHJLRQIRUFXOWLYDWLRQ RIWKH 8NUDLQH VSHFLHV DUHSUHVHQWHG7KHV\VWHPDWLFOLIHIRUPVDQGmorphological types of plants with respect to soil moisture spectras, origin and degree of naturalisation of species of thɿs JURXSDUHDQDO\]HG$DQQRWDWHGOLVWRIHUJDVLRSK\WHVLVSURYLGHG
Key words: alien species, ergasiophytes, invasive plants, Ukraine
1. Introduction increases and many of the species are characterized by DKLJKGHJUHHRIQDWXUDOL]DWLRQ7KHUHIRUHWKHQHFHVVLW\ %\WKHHQGRIWKHWKFHQWXU\LQYDVLRQVRIDOLHQ RIVSHFLDOFRPSOH[VWXG\RIHUJDVLRSK\WHVDULVHV organisms, including plants, were widely realized as The main aim of the study was to clarify of modern RQHRIWKHPDMRUJOREDOWKUHDWVWRELRGLYHUVLW\ 3\ãHN VSHFLHVFRPSRVLWLRQRIWKH8NUDLQLDQÀRUDHUJDVLR et al %DOGDFFKLQR 3L]]XWR 0RRQH\ SK\WHVDQGWKHLUVRPHFKDUDFWHULVWLFV &OHODQG 5HLFKDUG :KLWH .RZDULN 7KHREMHFWLYHVRIWKHVWXG\LQFOXGHG LQYHQWRU\RI Protopopova et al. &KRUQHVN\ 5DQGDOO ergasiophytes species, and preparation of the annotated 'DYLV 6D[ *DLQHVHWF OLVW JHQHUDOL]DWLRQRIWKHLUV\VWHPDWLFELRORJLFDODQG (UJDVLRSK\WHV VSHFLHVGHOLEHUDWHO\LQWURGXFHGWR ecological characteristics, origin, as well as determina DUHJLRQIRUFXOWLYDWLRQ SOD\DVLJQL¿FDQWUROHV\QDQ WLRQRIWKHGHJUHHRIQDWXUDOL]DWLRQLQWKHUHJLRQ WKURSLVDWLRQWKHSURFHVVE\ZKLFKWKHQDWLYHÀRUDRID UHJLRQEHFRPHVUHSODFHGE\DOLHQVSHFLHV7KHVHVSHFLHV 2. Material and methods generally are delivered into a new region from places where they wereFXOWLYDWHG7KHUHIRUHWKH\DUHSDUWLDOO\ The present research focuses on the ergasiophytes DGDSWHGWRFRQGLWLRQRIDQWKURSRJHQLFKDELWDWV6R RIWKH8NUDLQLDQDOLHQIUDFWLRQÀRUD7KHLQYHVWLJDWLRQV these species can adapt more easy and quickly to new are based on original data obtained by routine surveys FRQGLWLRQV The detection of the new species of alien LQWKHDQDO\VLVRIGDWDLQWKHOLWHUDWXUHDQG invasive plants before they become invasive, with the H[DPLQDWLRQVRIWKHFROOHFWLRQVRIWKH+HUEDULDRI0 aim of preventing their uncontrolled spread is one of *.KRORGQ\,QVWLWXWHRI%RWDQ\1DWLRQDO$FDGHP\RI necessary conditions of the Convention on Biological 6FLHQFHVRI8NUDLQH .: 'RQHWVN%RWDQLFDO*DUGHQ Diversity , the Global strategy on invasive alien 1$6RI8NUDLQH '1= 8]KJRURG1DWLRQDO8QLYHUVLW\ species the European strategy on invasive alien 88 <)HG\NRY\FK1DWLRQDO8QLYHUVLW\ &+(5 , species *HQRYHVL 6KLQH DQGRWKHULQWHUQDWLRQDO )UDQNR/YLY1DWLRQDO8QLYHUVLW\ /: 76KHYFKHQNR SURJUDPPHVDQGGRFXPHQWV,WLVWKHUHVSRQVLELOLW\RI .\LY1DWLRQDO8QLYHUVLW\ .:8 ,,0HFKQLNRY HYHU\FRXQWU\ZKLFKVLJQHGWKHPLQFOXGLQJ8NUDLQH 2GHVVD1DWLRQDO8QLYHUVLW\ 06'8 91.DUD]LQ ,QFRPSLOLQJWKHOLVWRIWKH8NUDLQLDQDOLHQIUDFWLRQ .KDUNLY1DWLRQDO8QLYHUVLW\ &:8 9,9HUQDGVN\L ÀRUDKDVGUDZQDWWHQWLRQWRWKHLQFUHDVLQJUROHRIVSH 7DYULDQ1DWLRQDO8QLYHUVLW\ 6,0) &ULPHDQ%UDQFK cies which have been intentional introduction into the RI$JURWHFKQRORJ\,QVWLWXWHRI1DWLRQDO8QLYHUVLW\RI FXOWXUHDQGODWHUHVFDSHG7KHQXPEHURIHUJDVLRSK\WHV %LRORJLFDO5HVRXUFHVDQG1DWXUDO8VHV &6$8 1LNLWD &+252/2*<
©$GDP0LFNLHZLF]8QLYHUVLW\LQ3R]QDĔ 3RODQG 'HSDUWPHQWRI3ODQW7D[RQRP\$OOULJKWVUHVHUYHG 9LUD93URWRSRSRYD 0\URVODY96KHYHUD (UJDVLRSK\WHVRIWKH8NUDLQLDQÀRUD
%RWDQLFDO*DUGHQ1DWLRQDO6FLHQFH&HQWUH8NUDLQLDQ vascular plants from 2 genera and 92 families VHH $JULFXOWXUDO$FDGHP\RI6FLHQFHV <$/7 $SSHQGL[ RURIWKHDOLHQIUDFWLRQÀRUDDQG &RPSDUDWLYHPRUSKRORJLFDODQGJHRJUDSKLFDOPH RIWRWDOQXPEHURIWKH8NUDLQLDQÀRUD Tendency WKRGVZHUHXVHGLQWKHSUHVHQWLQYHVWLJDWLRQV7D[RQRPLF of increasing number of the Ukrainian alien fraction VWUXFWXUH DFFRUGLQJWR7ROPDFKHY OLIHIRUPV ÀRUD EODFNOLQH DQGHUJDVLRSK\WHVJURXSVSHFLHV JUD\ DFFRUGLQJWR6HUHEU\DNRY DQGPRUSKRORJLFDO OLQH GXULQJ\HDUVLVSUHVHQWHGRQ)LJ W\SHVRISODQWVZLWKUHVSHFWWRVRLOPRLVWXUH DFFRUGLQJ 7KHWD[RQRPLFVSHFWUXPRIOHDGLQJSODFHVRIHUJDVLR WR'LGXNK RIWKHHUJDVLRSK\WHVZHUHDQDO\VHG phytes of the Ukrainie includes 12 families: Asteraceae Characteristics of area species prepared according to VS 5RVDFHDH )DEDFHDH %UDVVLFDFHDH Flora of European part of USSR and Flora of Eastern DQG3RDFHDH 4HDFK $SLDFHDH /DPLDFHDH (XURSH )HGRURY 7]YHOHY 6RODQDFHDH &DU\RSK\OODFHDH 3RO\JRQDFHDH 7KHFODVVL¿FDWLRQRIDOLHQSODQWVSURSRVHG &KHQRSRGLDFHDHDQG0DOYDFHDH HDFK 7KHVH E\.RUQDĞDQGPRGL¿HGE\3URWRSRSRYD DQG OHDGLQJIDPLOLHVFRPSULVHVSHFLHVRURIWKH Moysiyenko ZDVXVHGHJµDJULRHSRHFRSK\WH¶ WRWDO7KUHHRIOHDGLQJIDPLOLHVFRPSULVHVSHFLHVRU – species which fully naturalized in anthropogenic, 2WKHUIDPLOLHVFRQWDLQIURPHLJKWWRRQHVSHFLHV VHPLQDWXUDODQGQDWXUDOSODQWFRPPXQLWLHV,QYDVLYH LQFOXGLQJIDPLOLHVZKLFKDUHSUHVHQWHGRQO\RQHRU and transformer groups determined according Richard WZRVSHFLHV7KHWD[RQRPLFVSHFWUXPRIWKHOHDGLQJ son et al The species names are given according families of ergasiophytes differs from the general WR0RV\DNLQ )HGRURQFKXN VSHFWUDRIWKH8NUDLQLDQDOLHQIUDFWLRQÀRUDDVGRHVWKH UHODWLYHSRVLWLRQRIWKHIDPLOLHV H[FOXGLQJ$VWHUDFHDH 3. Results DQGWKHSUHVHQFHRIQHZRQHV 5RVDFHDH6RODQDFHDH 3RO\JRQDFHDH 6WXGLHVRIDOLHQSODQWVRYHU\HDUSHULRGWHVWL¿HG The taxonomic spectrum of the leading genera of that the process of synanthropisationRIWKHÀRUDVRI VWXGLHGJURXSLVQRWFOHDUO\H[SUHVVHG7KHODUJHVWQXP natural zones )RUHVW)RUHVW6WHSSHDQG6WHSSHDQG EHURIVSHFLHV DUHIURPWKHJHQXVHelianthus/7KH their divisions DQGLQWKH8NUDLQLDQÀRUDLQJHQHUDO genera Opuntia0LOODQGSymphyotrichum1HHVLQFOXGH KDVEHHQFRQVWDQWO\SURJUHVVLQJ:HFDQREVHUYHFOHDU six species each; Acer LAllium LAmaranthus /and tendencies for the number of alien species to increase, Brassica L±¿YH species each; Datura LJuglans L the growth of both µQDWXUDOLVHG¶RUµVWDEOH¶ DJULRSK\WHV Lonicera LPopulus LPrunus LPyrethrum =LQQ, DJULRHSRHFRSK\WHVHSRHFRSK\WHVFRORQRSK\WHV DQG Rosa LSpiraea LTrifolium LDQGVitis L±four µXQVWDEOH¶ ephemerophytes, ergasiophygophytes species each genera include three species each FRPSRQHQWVRIWKHDOLHQIUDFWLRQRIWKHÀRUDJURZLQJ genera – two species each, and 214 – only one species rates of immigration and dispersal, an increasing num each ber and scope of invasions of certain invasive species ,QWKHVSHFWUDRIOLIHIRUPVRIVSHFLHVRIHUJDVLR 3URWRSRSRYDet al. SK\WHVRI8NUDLQH )LJ DVLQWKHDOLHQIUDFWLRQÀRUD The ergasiophyte group of the Ukraine, according DQQXDOVSUHYDLO VSRU EXWWKHLUSDUWLFLSDWLRQ WRRXUGDWDLVUHSUHVHQWHGE\DWOHDVWVSHFLHVRI SHUFHQWDJHLVORZHU7KH second position is occupied by
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herbaceous perennials or $SHFXODULW\RI SRVLWLRQ±PHVR[HURSK\WHV RU DQG[HURSK\WHV the spectra is the high participation of the shrubs and RU 7KHRWKHUJURXSVK\GURSK\WHVK\JURThe other groups, hydrophytes, hygro WUHHVDPRQJWKHPWKHUHDUHVKUXEVWUHHVVKUXE SK\WHVK\JURPHVRSK\WHVDQGK\GURPHVRSK\WHVDUH or tree, 2 sHPLVKUXEV LQtotal VSHFLHVRU unnumerous and presented from six to one species 2WKHUJURXSVGRQRWLQFOXGHQXPHURXVVSHFLHVDQGDUH The species of eUJDVLRSK\WHVRIWKH8NUDLQLDQÀRUD UHSUHVHQWHGE\IURPWKUHHWRRQHVSHFLHV HYDU\LQRULJLQ:LWKUHJDUGWRWKHLUSULPDU\DUHDRI Among the ergasiophyte species of the Ukrainian RULJLQ )HGRURY 7]YHOHY ÀRUDZLWKUHVSHFWWRVRLOPRLVWXUH )LJ [HURPH HUJDVLRSK\WHVDUHGLYLGHGLQWRDQXPEHURIW\SHV VRSK\WHVSUHYDLO VSHFLHVRU 7KH second The basis for this group is presented by origin from position occupied mesophytes or WKHQH[W 1RUWK$PHULFD VSHFLHV WKH0HGLWHUUDQHDQ
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(DVW$VLDDQG$VLD HDFK 7KHQH[WODUJHJURXSVDUH /DQGAilanthus altissima IURP2VQRY\DQVN\LPRGHUQ 6RXWK$PHULFDQ 6RXWK(XURSHDQ &DXFDVLF .UDVQRNXWVN\L3DUNQHDU.KDUNLY Robinia pseudo- :HVW0HGLWHUUDQHDQ DQG:HVW(XURSHDQ acacia/ IURP2GHVDDQG.KDUNLY%RWDQLFDO*DUGHQV LQRULJLQ7KHJURXSRIK\EULGRULJLQLVUHSUHVHQWHGE\ Abutilon theophrasti0HGLN IURP/XEQ\%RWDQLFDO VSHFLHV2WKHUJURXSVGRQRWLQFOXGHQXPHURXVVSH *DUGHQ3ROWDYD5HJ HWF,QWKHth century an essen cies and are represented by from nine to one species, tial role in the cultivation of new plants for agricultural and LQFOXGLQJW\SHVRIDUHDDQGVXEGLYLVLRQZKLFKFRQVLVW technical purposes was taken up by Experimental Stations RQHVSHFLHVRQO\ .KDUNLY3ROWDYD/XEQ\0DVOLYNDHWF ,QUHFHQW\HDUV With respect to the time of immigration, the species the uncontrolled import of wide range of ornamental and RIHUJDVLRSK\WHVRI8NUDLQLDQÀRUDDUHGLYLGHGLQWR YHJHWDEOHSODQWVVHHGKDVEHHQREVHUYHG DUFKDHRSK\WHV VSHFLHV DQGNHQRSK\WHV 6RPH We have attempted to reconstruct the main stages of species have a different status in different natural zones QDWXUDOL]DWLRQRIVRPHHUJDVLRSK\WHV 7DEOH RI8NUDLQHHJWKHVSHFLHVAcer tataricum/ZKLFKLV )RUWKHPRVWSDUWRIWKLVJURXSGDWDDERXWWKHLU¿UVW QDWLYHLQ)RUHVW6WHSSH]RQHVRU Rubus idaeus/QDWLYH FXOWLYDWLRQDUHDEVHQW7KHOLWHUDWXUH 3DOLPSVHVWRY LQ)RUHVW]RQHVDUHNHQRSK\WHVLQ&ULPHD 3URWRSRSRYD 9\QRJUDGRYDet al. 0D\RURYet With regard to their degree of naturalization, the al. HWF +HUEDULDPDWKHULDOVDQGRULJLQDOGDWDFRQ VSHFLHVRIHUJDVLRSK\WHVRIWKH8NUDLQLDQÀRUDPD\EH cerning the naturalization of transformer species shows GLYLGHGLQWRWKHJURXSVHJDJULRSK\WHV DJULR WKDWWKHSHULRGIURPWKH¿UVWQRWHDVHVFDSHGSODQWVWRWKHLU HSRHFRSK\WHV DJULRFRORQRSK\WHV HSRHFR H[SDQVLRQLVIURPRYHU\HDUVWRRUPRUHHJImpa- SK\WHV FRORQRSK\WHV HSKHPHURSK\WHV DQG WLHQVSDUYLÀRUD'&ZDVNQRZQLQFXOWLYDWLRQLQ8NUDLQH HUJDVLRSK\JRSK\WHV 7KHGHJUHHRIQDWXUDOL]DWLRQ IURPLQ/YLYJDUGHQVDQGSDUNVDQGIURP ±LQ of numerous species differs in the various botanical and 'XEO\DQ\SDUN /YLY5HJLRQ $ERXW\HDUVODWHU JHRJUDSKLFDOUHJLRQVRIWKH8NUDLQHHJAntirrhinum WKH¿UVWHVFDSHGSODQWVRIWKLVVSHFLHVZHUHQRWHGLQWKH majus/LVDQHUJDVLRSK\JRSK\WHLQPDMRURIUHJLRQV /YLYDUHDDQGLQWKHɋDUSDWKLDQV7KHDFWLYHGLVWULEXWLRQ RI8NUDLQHDQGDQDJULRSK\WHLQ&ULPHD RIWKLVVSHFLHVZDVEHJLQQLQJ\HDUVDJRDQGFRQWLQXHV The extent of distribution of species of ergasiophytes DWSUHVHQW,Q(XURSHDVDZKROHWKLVSHULRGKDVODVWHG YDULHV7KHGLVWULEXWLRQRIDODUJHSDUWRIWKHHSKHP \HDUVORQJHU1RZLQ3ROLVV\D WKH)RUHVW]RQH WKHVSHFLHV erophytes and ergasiophygophytes has a pattern of a has been noted in plant communities of the Querco-Fagetea single or a few localities; some of them however were Br.%O.HW9OLHJ Robinietea -XUNRH[+DGDþHW6R cultivated over a long period of time and are very com IURQ DQGGalio-Urticetea 3DVV HP.RSHFN\ mon and sporadically distributed throughout the terri FODVVHV /XNDVK DQGLQ)RUHVW6WHSSH]RQHVRI WRU\RIWKH8NUDLQH7KHPDMRULW\RIHSRHFRSK\WHVDQG the Alnetea glutinosae Br.%O. HW57[ FODVVDOVR colonophytes are distributed in more than three botanical .DQLY6WDWH5HVHUYH DQGLQ%XNRY\QD&LV&DUSDWKLDQ DQGJHRJUDSKLFDOUHJLRQVRIWKH8NUDLQH$JULRSK\WHV in Epilobietea angustifolii 7[HW3UVJH[YRQ5RFKRZ are a component of plant communities in one or two 1951, Artemisietea vulgaris /RKP3UVJHW57[LQ UHJLRQV$WHQGHQF\WRDFWLYHVSUHDGLVPRVWHYLGHQWIRU 57[ Polygono-Poetea annuae 5LYDV0DUW DJULRHSRHFRSK\WHVDQGHSRHFRSK\WHV7KHGLVWULEXWLRQ and Galio-Urticetea, and in Querco-Fagetea, of agriophytes is limited by the presence of correspond Robinietea FODVVHV 3URWRSRSRYDet al $VLPLODU LQJSODQWFRPPXQLWLHVVXVFHSWLEOHWRLQYDVLRQ7KXV situation has been observed for Impatiens glandulifera IURPWKHVSHFLHVRIHUJDVLRSK\WHVRU ZHUH 5R\OH3UHVHQWO\WKLVVSHFLHVLQ3ROLVV\DLVDFRPSRQHQWRI FRQVLGHUHGWREHFRPSOHWHO\QDWXUDOL]HG$PRQJWKHPZH plant communities of the Phragmito-Magnocaricetea KDYHLGHQWL¿HGVSHFLHVRUZKLFKPD\EHGH¿QHG .OLNDLQ.OLNDHW1RYDN Molinio-Arrhenatheretea DVLQYDVLYHLQFOXGLQJVSHFLHVRUWUDQVIRUPHUV 57[ Alnetea glutinosae, Bidentea tripartiti 5 The cultivation of plants in Ukraine has an ancient his 7[/RKPHW3UVJ FODVVHV,QEchinocystis lobata WRU\7KHSULPDU\FHQWUHVRIFXOWLYDWLRQZHUHWKHPRQDVWHULHV 0LFK[ 7RUU $*UD\WKHSHULRGRIDGDSWDWLRQZDVDERXW which cultivated mainly medicinal plants, later apothecary \HDUVLQ8NUDLQHDQG±LQRWKHUSDUWVRI(XURSH gardens and private parks where numerous ornamental The period from the beginning of cultivation to an expansion SODQWVZHUHSODQWHG2QHRIWKH¿UVWERWDQLFDOJDUGHQVZHUH in Heracleum sosnowskyi 0DQGHQZDVDERXW\HDUVLQ FUHDWHGLQ/XEQ\ .KDUNLY .UHPHQHWV Solidago canadensis/DQGSenecio cineraria '&±DERXW 1LNLWD 2GHVDDQG1L]K\Q 6RPHDOLHQVSHFLHV 3URWRSRSRYDet al. 3URWRSRSRYD 6KHYHUD which were cultivated there now are widely distributed in DE 9LQRJUDGRYD et al. 7KHVHH[DPSOHV 8NUDLQHHJɋɭFODɫKDHQDɯɚQWKLLIROLD 1XWW )UHVHQ show the different rates at which some species adapt to new IURP.LHY8QLYHUVLW\%RWDQLFDO*DUGHQ Ailanthus FRQGLWLRQV altissima 0LOO 6ZLQJOHDQGBupleurum fruticosum/ 1RZHUJDVLRSK\WHVDUHUHJLVWHUHGLQ8NUDLQHDVFRP IURP1LNLWD%RWDQLFDO*DUGHQ&ULPHD Acer negundo SRQHQWVRIDOPRVWDOOW\SHVRIVHPLQDWXUDODQGQDWXUDO %LRGLY5HV&RQVHUY
Table 1. The main stages of naturalization of ergasiophytes in Ukraine
Species First date of First date of cultivation in Ukraine 'DWHDQGUHJLRQRI The start of active cultivation ¿UVWUHJLVWUDWLRQDV distribution and region in Europe, escaped plants country
Acer negundo (QJODQG 2VQRY\DQVN\L modern VV" The second half of thF .UDVQRNXWVN\L dendrological park )RUHVW6WHSSH]RQH .KDUNLY5HJ ±1LNLWD Bot *DUG Crimea ± Kremenets Bot *DUG Ternopil Reg ± Trykratskyi park of Skarzhynskyi modern 2GHVD Reg ±VelykoAnadolskyi forestry modern 'RQHWVN Reg Ailanthus " 2VQRY\DQVN\L ±1LNLWD Crimea The end of th c, Crimea altissima Bot*DUG ±Simferopol, Sebastopol, Theodosia Ambrosia 1914, YLOO.XGDVKLYND modern 1925, Kiev V V6WHSSH]RQH artemisiifolia *HUPDQ\ 'QLSURSHWURYVN Reg Amorpha (QJODQG V or middle of 19th c2GHVD Probably the V0LGGOH'QLSUR fruticosa parks, and Trykratsky park of middle of th c" Region Skarzhynsky 2GHVD5HJ Bupleurum - 1LNLWD Bot*DUG Alupka 1 6RXWK:HVWHUQ fruticosum &ULPHD Crimea Echinocystis The second Probably ¿UVWKDOIRIth c vill VZHVWHUQUHJLRQVRI lobata half of 19th 'LGLYWVL, Ukraine F " Transcarpathia Elaeagnus 19s, 2GHVD gardens 1925, villForos V, steppe regions of angustifolia Romanie and Sudak Ukraine &ULPHD Heracleum (QJODQG 2VPRORGDIRUHVWU\ PRGHUQ vill V7UDQVFDUSDWKLD mantegazzianum ,YDQR)UDQNLYVN5HJ 2VPRORGD Heracleum 5XVVLD V V, western and northern years of th c, sosnowskyi forest regions of Ukraine Transcarpathia, Polissya )RUHVW]RQH Impatiens Lviv parks Lviv The middle of thF SDUYLÀRUD Switherland )RUHVW6WHSSH]RQHDQG 3ROLVV\D )RUHVW]RQH Impatiens (QJODQG 19s, Transcarpathia and some YLOO2VL\ 19 s, Transcarpathia glandulifera western regions of Ukraine DQG+DQNRY\WV\D Transcarpathia; YLOO Mykhaylivka, Khmelnytskyi 5HJ Helianthus " " " 19 s, Transcarpathia tuberosus Reynoutria (QJODQG " 1929, Rachiv, 19 s, Transcarpathia japonica Transcarpathia Rhamnus 1LNLWD Bot*DUG Probably 19s, Crimea alaternus 6RXWKEDQN Rudbeckia " the middle " and the end of vicinity of The end of the th c laciniata 19th c±the beginning of th c YLOO6HUHGQ\H, Bukovyna, Transcarpathia Transcarpathia, Bukovyna Transcarpathia, YLOO -RUGDQHVKWL Bykovyna Senecio cineraria " First half of 19th cprobably in 195V&ULPHD 6RXWK 1LNLWD%RW*DUG EDQN Solidago (QJODQG The beginning of 19th F VUight bank of canadensis )RUHVW6WHSSH]RQH 9LUD93URWRSRSRYD 0\URVODY96KHYHUD (UJDVLRSK\WHVRIWKH8NUDLQLDQÀRUD
SODQWFRPPXQLWLHVDQGHFRV\VWHPV IRUHVWVWHSSH of the Steppe zone ecotopes is observed, the changing DTXDWLFYHJHWDWLRQHWF HJLQIRUHVWVZHUHJLVWHUHG RIWKHLUZDWHUDQGVDOWUHJLPHVLVQRWHG 9\VKQHYVN\ invasions of Acer negundo, ,PSDWLHQVSDUYLÀRUD; along ,WKDVFDXVHGVXSSUHVVLRQRIVHYHUDOVSHFLHVRI river shores, Heracleum mantegazzianum, Impatiens ORFDOÀRUDDQGIDFLOLWDWHGQDWXUDOL]DWLRQRIVRPHHUJDVLR glandulifera, Echinocystis lobata, Reynoutria japonica, SK\WHVDGDSWHGWRWKHFRQGLWLRQVZKLFKKDYHDULVHQ)RU and several species of Helianthus/IRUPODUJHRIWHQ example, Elaeagnus angustifolia actively consumes PRQRGRPLQDQWSRSXODWLRQV7KHGLVSHUVDORILQYDVLYH water resources and overcompetes many local species; species represents a serious danger to native rare spe LWLVDVDOWWROHUDQWVSHFLHV,WIRUPVVWDQGVWKDWFKDQJH FLHVDQGQDWXUDOSURWHFWHGDUHDVHJWKHPRVWGDQJHURXV the light mode of the herbaceous vegetation layer in DOLHQSODQWLQWKH'DQXEH%LRVSKHUH5HVHUYHLQVDQG\ Steppe zone communities, thus affecting the species habitats – Elaeagnus angustifolium, in Kaniv State FRPSRVLWLRQDQGWKHVWUXFWXUHRISODQWFRPPXQLWLHV 5HVHUYHLQÀRRGSODLQIRUHVW±Acer negundo/DQG This species forms the association Elaeagnetum angus- Amorpha fruticosa/ 3URWRSRSRYDet alD tifoliae &KLQNLQDDQGRFFXSLHVVSHFL¿FHFRQLFKHVLQ RWKHUFRPPXQLWLHVHJHippophoaeta rhamnoidis, thus 4. Discussion SURPRWLQJGHJUDGDWLRQRIDERULJLQDOFRPPXQLWLHV7KH species E. angustifolia shows high invasive ability to The spectrum of ergasiophytes according to origin penetrate many different types of KDELWDWV HFRWRSHV DQG of species are differentiated by large variety, however SODQWFRPPXQLWLHVZKLFKRFFXUVLQGLIIHUHQWÀRULVWLF the success of species naturalization persists in those complexes QDWXUDO SVDPPRSK\WRQSUDWRSK\WRQ groups that in the general range of the Ukrainian alien halophyton, steppophyton, drymophyton, petrophyton, IUDFWLRQÀRUD7KHJUHDWHVWVXFFHVVKDVDVSHFLHVRIWKH litoralophyton, and anthropogenic: aggeratophyton, 1RUWK$PHULFDQDQG(DVW$VLDQRULJLQ$VLQWKHJHQHUDO DQGUXQNDWLRGU\PRSK\WRQ The species occurs mostly VSHFWUXPRIWKH8NUDLQLDQDOLHQIUDFWLRQÀRUDDPRQJ RQVDQG\DQGULYHUVLGHVHPLQDWXUDOHFRWRSHVZLWK ergasiophytes the species of Mediterranean origin are unstable and VSDUVHSODQWFRYHU7KHPRVWGLYHUVLW\RI dominated, however most of them have a low degree plant communities with participation of E. angustifolia RIQDWXUDOL]DWLRQ)HDWXUHRIWKHOLIHIRUPVSHFWUXP is REVHUYHGLQVDQG\ DVVRFLDWLRQV DQGULYHUVLGH is higher percentage of perennials, shrubs and trees DVVRFLDWLRQV ELRWRSHVZLWKVSDUVHYHJHWDWLRQ7KHPDLQ among the ergasiophytes; and in the morphological limiting factors for the species distribution in the studied types of plants with respect to soil moisture spectrum region are excessively humid ecotopes 3URWRSRSRYDet WKHPHVRSK\WHVJURXS LQ DQDJJUHJDWH LVGRPLQDWHG al. compared with the general spectrum of the Ukrainian DOLHQIUDFWLRQÀRUD 5. Conclusion *URZLQJRIWKHVSHFLHVLQFXOWXUHDWWKH¿UVWVWDJHV under the conditions of new region facilitate the natu The above data shows that, the ergasiophyte group, UDOL]DWLRQRISODQWV,QWKHOLWHUDWXUH 9\QRJUDGRYDet al. UHSUHVHQWHGE\VSHFLHVLQ8NUDLQHSOD\VDVLJQL WKHRSLQLRQZDVH[SUHVVHGWKDWWKHVHHGDQGSODQW ficant role in the enrichment of spontaneous flora ing material are often imported not from primary habi 3URWRSRSRYD 6KHYHUDDE The results of WDWVDQGRIWKHSODFHVZKHUHWKHVHVSHFLHVDUHFXOWLYDWHG our studies show that numerous species of this group And that means they are already partly preadapted to the reached a high degree of naturalization and are widely FXOWXUHFRQGLWLRQV1RZDGD\VWKLVLVDOVRHYLGHQFHGE\ GLVWULEXWHG UHGXFLQJWKHWLPHRIQDWXUDOL]DWLRQRIFHUWDLQVSHFLHV ,QFUHDVLQJWKHQXPEHUDQGGLYHUVLW\RIDQWKURSRJHQLF Acknowledgement. :HZRXOGOLNHWRWKDQN0V1DWDOLD and disturbed natural ecotopes that contribute to the 90RUJDUW 0*.KRORGQ\,QVWLWXWHRI%RWDQ\1$6RI emergence of new niches facilitates the introduction 8NUDLQH IRUWHFKQLFDODVVLVWDQFH7KHDXWKRUVDUHJUDWHIXOWR of alien species into synanthropic and disturbed natu an anonymous reviewer for his detailed review of manuscript, UDOSODQWFRPPXQLWLHV)RUH[DPSOHLQ8NUDLQHGXH YDOXDEOHFRPPHQWVDQGVXJJHVWLRQV to changes of climatic conditions the mesophytisation %LRGLY5HV&RQVHUY
References
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KOWARIK & $,%8'=+$.99 .25=+$1.97KH 867$5),1*(5 HGV %LRORJLVFKH,QYDVLRQHQ+HUDXV WUDQVIRUPHUVSHFLHVLQWKHÀRUDRIWKH Bukovyna IRUGHUXQJ]XP+DQGHOQ1HRELRWD &LV&DUSDWKLDQDUHD8NU%RW-RXUQ /8.$6+ 29)ORUDRIYDVFXODUSODQWVRI(DVWHUQ 3527232329$996+(9(5$09 0(/1<.53 3ROLVV\D KLVWRU\RILQYHVWLJDWLRQFRQVSHFWSS The history of introduction and present distribution of 3K\WRVRFLRFHQWHU3UHVV.LHY Elaeagnus angustifolia/LQWKH%ODFN6HD5HJLRQRI 0$<2529 6 5 %2&+.,19'1$6<029,&+<8$ 8NUDLQH&KRUQRPRUVNL%RW-RXUQ 6&+(5%$.29$9$OLHQÀRUDRI0RVFRZDQG 3527232329$996+(9(5$09 026<$.,16/ 0RVFRZ5HJLRQSS7RYDULVFKHVWYRQDXFKQ\NK 'HOLEHUDWHDQGXQLQWHQWLRQDOLQWURGXFWLRQRILQYDVLYH L]GDQL\.0.3UHVV0RVFRZ ZHHGV DFDVHVWXG\RIWKHDOLHQÀRUDRI8NUDLQH MOYSIYENKO ȱȱ 8UEDQÀ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ppendix 1. $QQRWDWLRQOLVWRIHUJDVLRSK\WHVRIWKH8NUDLQLDQÀRUD 7KHOLVWLQFOXGLQJWKHQH[WSRVLWLRQV QDPHRIVSHFLHVIDPLO\OLIHIRUPK\GURPRUSKHFKURQRORJLFDOHOHPHQWV GDWDRI¿UVW SXEOLFDWLRQRU¿[DWLRQ RULJLQGHJUHHRIQDWXUDOL]DWLRQZD\RIGLVWULEXWLRQ LQYDVLYHRUWUDQVIRUPHU UHJLRQ Abutilon theophrasti Medik0DOYDFHDH$QQ;HURPHVR.HQ ($V(SRHF(UJ Acer negundo L$FHUDFHDH7UHHMeso.HQ1 $PAgreSRHFEUJNVHQo; invasive; transformer forest and foreststeppe regions of Ukraine Acer platanoides L$FHUDFHDH7UHHMeso, .HQ Crimea natural species in Ukraine, EurCRO(UJ Acer pseudoplatanus L$FHUDFHDH7UHHMeso.HQ Crimea natural species in Ukraine, EurCauc Tzvelev CRO(UJ Acer saccharinum L$FHUDFHDH7UHHMeso.HQ1 $P(UJDVSKLJ(UJ Acer tataricum L$FHUDFHDH7UHH;HURPHVR, .HQ Crimea natural species in Ukraine, EurCRO(UJ Acorus calamus LAraceae3HUHQQ+\JUR$UFKAs$JU(UJ invasive Aesculus hippocastanum L+LSSRFDVWDQDFHDH, Trees, Meso, .HQ0HG(UJDVSKLJ(UJ Agrostemma githago L&DU\RSK\OODFHDH$QQXeromeso$UFKAnthrop(UJDVSKLJKsenoeUJ Ailanthus altissima Mill Swingle, Simaroubaceae, Shrub, Xeromeso.HQ($V$JUHSRHFErg invasive; transformer Crimea Albizia julibrissin 'XUD]]Mimosaceae, Tree, Xeromeso.HQ Crimea S ETranscaucCRO(UJ Alcea rosea L0DOYDFHDH3HUHQQMesoxero.HQ0HG(UJDVSKLJ(UJ Allium cepa L$OOLDFHDH%LHQQMeso.HQCAs Mayorov et al. or0HG Pyãek et al. (UJDVSKLJ(UJ Allium ¿VWXORVXP L$OOLDFHDH3HUHQQMeso.HQ($V(UJDVSKLJ(UJ Allium porrum L$OOLDFHDHAnnor bLHQQMeso.HQEMed(UJDVSKLJ(UJ Allium sativum L$OOLDFHDH%LHQQMeso.HQMidAs(UJDVSKLJ(UJ Allium victorialis L$OOLDFHDH3HUHQQ;HURPHVR.HQ Crimea natural species in Ukraine Carpathian subAlpCRO(UJ Althaea RI¿FLQDOLV L0DOYDFHDH3HUHQQ+\JURmeso$UFK,UDQTur$JUHSRHF(UJ Amaranthus caudatus L$PDUDQWKDFHDH$QQXeromeso.HQSAm tropic (UJDVSKLJ(UJ Amaranthus caudatus Lsspsaueri -HKOLN$PDUDQWKDFHDH$QQXeromeso.HQSAm(UJDVSKLJ(UJ Amaranthus cruentus L$PDUDQWKDFHDH$QQXeromeso.HQTropAmSAm tropic (SRHF(UJ Amaranthus hypochondriacus L$PDUDQWKDFHDH$QQXeromeso.HQ1$P(UJDVSKLJ(UJ Amaranthus spinosus L$PDUDQWKDFHDH$QQXeromeso.HQSAm(SKHP(UJ Amberboa moschata L '&$VWHUDFHDH$QQXero.HQWAs(UJDVSKLJ(UJ Ambrosia artemisiifolia L$VWHUDFHDH$QQXeromeso.HQ 1$P$JUHSRHFEUJNVHQR invasive; transformer forest steppe and steppe regions of 8NUDLQH Ambrosia WUL¿GD L$VWHUDFHDH$QQXeromeso, .HQ1$P(SKHP(UJ Amelanchier canadensis L Medik5RVDFHDH6KUXEor tree, Meso, .HQ1$P(SKHP(UJ Amelanchier ovalis Medik5RVDFHDH6KUXEMeso.HQ0HG(UJDVSKLJ(UJ Amelanchier spicata Lam KKoch, Rosaceae, Shrub, Meso.HQ1$P$JU(UJ invasive Ammi majus L$SLDFHDHBiennMesoxero.HQ0HG(SKHP(UJ Amorpha fruticosa L)DEDFHDH6KUXEor tree, Meso.HQ1$P$JUHSRHF(UJ invasive; transformer central regions of Ukraine Anacyclus clavatus 'HVI Pers$VWHUDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Anacyclus RI¿FLQDUXP +D\QH$VWHUDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Anemone fasciculata L5DQXQFXODFHDH3HUHQQMeso.HQ Crimea – CaucCROErg Anethum graveolens L$SLDFHDH$QQMeso, .HQMed,UDQTur(UJDVSKLJ(UJ Anredera cordifolia Ten Steenis, Basellaceae, Liane, Meso.HQCand SAm(UJDVSKLJErg Anthericum liliago LAsphodelaceae3HUHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Antirrhinum majus L6FURSKXODULDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ Ukraine $JU Crimea (UJ Apium graveolens L$SLDFHDHAnnor bLHQQMeso.HQWEur Protopopova or unknown Pyãek et al. (UJDVSKLJ(UJ Apocynum cannabinum LApocynaceae3HUHQQMeso.HQ 1$P(UJDVSKLJ(UJ Aquilegia vulgaris L5DQXQFXODFHDH3HUHQQ;HURPHVR.HQ WEur$JU(UJ Armeniaca vulgaris Lam5RVDFHDH7UHHMesoxero, .HQEand CAs(SKHP(UJ Armoracia rusticana Lam *DHUWQMey Scherb%UDVVLFDFHDHH3HUHQQ;HURPHVR.HQ,UDQTurCRO(UJ Aronia melanocarpa Michx Elliot, Rosaceae, Shrub, Meso.HQ1$P(UJDVSKLJ(UJ Aronia ×mitschurinii AKSkvortsov & Brassica napus L%UDVVLFDFHDHAnnor bLHQQXeromeso.HQSEur(UJDVSKLJ(UJ Brassica nigra L W'-Koch%UDVVLFDFHDH$QQXeromeso.HQ0HG(SRHF(UJ Brassica oleracea L%UDVVLFDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Brassica rapa L%UDVVLFDFHDH$QQXeromeso.HQWAs(UJDVSKLJ(UJ Broussonetia papyrifera L Vent, Moraceae, Tree or shrub, Xeromeso, .HQ($V(UJDVSKLJ(UJ Brunnera macrophylla Adams , M-RKQVW%RUDJLQDFHDH3HUHQQ;HURPHVR.HQEur0HG Pyãek et al. (UJDVSKLJ(UJ Buddleja davidii Franch, Buddlejaceae, Shrub, Meso.HQ Crimea AsɋRO(UJ Bupleurum fruticosum L$SLDFHDH6KUXEMesoxero.HQ0HG$JU(UJinvasive; transformer Crimea Calendula RI¿FLQDOLV L$VWHUDFHDH$QQXeromeso, .HQ0HG(UJDVSKLJ(UJ Callistephus chinensis L 1HHV, Asteraceae, Annor bLHQQMeso.HQ($V(UJDVSKLJ(UJ Camelina sativa L Crantz sspsativa%UDVVLFDFHDH$QQXeromeso$UFKunknown Pyãek et al. (SRHF(UJ Campanula medium L, Campanulaceae%LHQQMeso.HQSEur(UJDVSKLJ(UJ Campsis JUDQGLÀRUD Thunb KSchumann, Bignoniaceae, Liana, Meso, .HQ($V(UJDVSKLJErg Campsis radicans L SeemBignoniaceae, Liana, Meso.HQ1$P(UJDVSKLJ(UJ Cannabis sativa LCannabaceae$QQXeromeso.HQ($V(SRHF(UJ Caragana arborescens Lam)DEDFHDH7UHHor shrub, Meso.HQSib(UJDVSKLJErg Carthamus tinctorius L$VWHUDFHDH$QQMesoxero, .HQSAs Mayorov et al. (UJDVSKLJ(UJ Castanea sativa Mill, Fagaceae, Tree, Meso.HQCaucAsia Minor0HG(UJDVSKLJ(UJ Catalpa bignonioides Walt, Bignoniaceae, Tree, Meso, .HQ1$P(UJDVSKLJ(UJ Cedrus atlantica Endl Carrière, Pinaceae, Tree, ;HURPHVR, .HQ Crimea – 1WAfr(UJDVSKLJ(UJ Cedrus deodara 5R[E *'RQ3LQDFHDH7UHHXeromeso.HQ Crimea CAsɋRO(UJ Celtis australis L, Celtidaceae, Tree, Xeromeso.HQ Crimea 0HG(UJDVSKLJ(UJ Celtis caucasica Willd, Celtidaceae, Tree, Xeromeso.HQ Crimea Cauc(UJDVSKLJ(UJ Cenchrus longispinus +DFN Fernald3RDFHDH$QQMesoxero.HQ 1$P$JUHSRHFEUJNVHQR; invasive Centranthus ruber L '&, Valerianaceae3HUHQQ;HURPHVR.HQ Crimea 0HG$JU(UJ Cerastium tomentosum L&DU\RSK\OODFHDH3HUHQQMesoxero.HQ Crimea MedCRO(UJ Cerasus besseyi Bailey Sok5RVDFHDH6KUXEMeso.HQ1$P(UJDVSKLJ(UJ Cerasus tomentosa Thunb Wall5RVDFHDH7UHHor shrub, Meso.HQ($V(UJDVSKLJ(UJ Cerasus vulgaris Mill5RVDFHDH7UHHor shrub, Meso.HQEMedor Balc(UJDVSKLJ(UJ Ceratochloa carinata +RRN Arn Tutin3RDFHDH%LHQQXeromeso.HQ1$P(SKHP(UJ Ceratostigma plumbaginoides Bunge, Plumbaginaceae3HUHQQXeromeso.HQAsCRO(UJ Cercis siliquastrum L&DHVDOSLQLDFHDH6KUXEXeromeso.HQ 0HG(UJDVSKLJ(UJ Chaenomeles japonica Thunb Lindl5RVDFHDH6KUXEMeso.HQ($V(UJDVSKLJ(UJ Chamaemelum nobile L All$VWHUDFHDH3HUHQQMeso.HQ0HG(UJDVSKLJ(UJ Cheiranthus cheiri L%UDVVLFDFHDH3HUHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Chenopodium capitatum L Ambrosi&KHQRSRGLDFHDH$QQXeromeso.HQWEur(UJDVSKLJ(UJ Chenopodium foliosum Asch&KHQRSRGLDFHDH$QQXeromeso.HQEurMedAs(SRHF(UJ Chenopodium schraderianum Schult&KHQRSRGLDFHDH$QQMeso.HQAfr tropic and subtropic (UJDVSKLJ(UJ Chrysanthemum carinatum Schousb$VWHUDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Chrysanthemum coronarium L$VWHUDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Cicer arietinum L)DEDFHDH$QQ.HQMeso0HG(SKHP(UJ Citrullus lanatus 7KXQE 0DWVXP 1DNDL&XFXUELWDFHDH$QQXeromeso, .HQ$IU(UJDVSKLJ(UJ Cleome spinosa -DFT&OHRPDFHDH$QQ;HURPHVR.HQ 6$P(UJDVSKLJ(UJ Clematis ÀDPPXOD L5DQXQFXODFHDH/LDQD;HURPHVR.HQ0HGCRO(UJ Clematis vitalba L5DQXQFXODFHDH/LDQD;HURPHVR.HQSor CEur(UJDVSKLJ(UJ Cnicus benedictus L$VWHUDFHDH$QQ0HVR[HUR.HQSEur(UJDVSKLJ(UJ Colutea arborescens L)DEDFHDHShrub, Meso[HUR.HQ Crimea natural species in Ukraine, EurMed Pyãek et al. CRO(UJ Colutea orientalis Mill)DEDFHDH7UHHXero, .HQ Crimea unknown,$JU(UJ Commelina communis L,ɋommelinaceae$QQMeso, .HQ S($V(SKHPEUJNVHQR Coreopsis JUDQGLÀRUD +RJJ ex Sweet, Asteraceae, Annor bienn;HURPHVR.HQ1$P(UJDVSKLJ(UJ Coreopsis tinctoria 1XWW$VWHUDFHDHAnnor bienn;HURPHVR.HQ1$P(UJDVSKLJ(UJ Coriandrum sativum L$SLDFHDH$QQ;HURPHVR.HQ0HG(UJDVSKLJ(UJ Cosmos bipinnatus Cav$VWHUDFHDH$QQ;HURPHVR.HQ1$PCRO(UJ Cotinus coggygria Scop, Anacardiaceae, Tree or shrub, Xero, .HQ steppe regions of Ukraine natural species in Crimea0HG (UJDVSKLJ(UJ Crocus ÀDYXV Weston,ULGDFHDH3HUHQQ;HURPHVR.HQunknown Pyãek et al. Erglip(UJ Cucumis sativus L&XFXUELWDFHDH$QQMeso, .HQS($V(UJDVSKLJ(UJ Cucurbita maxima 'XFK&XFXUELWDFHDH$QQ Meso, .HQSAm(UJDVSKLJ(UJ Cucurbita pepo L&XFXUELWDFHDH$QQMeso, .HQ1$P(UJDVSKLJ(UJ Cupressus sempervirens L, Cupressaceae, Tree, Mesoxero.HQ Crimea – 0HGCRO(UJ Cyclamen hederifolium AitPrimulaceae3HUHQQMeso.HQ0HGCRO(UJ Cyclamen vernum Sweet, Primulaceae3HUHQQMeso.HQ0HG(UJDVSKLJ(UJ Cydonia oblonga Mill5RVDFHDH6KUXEXeromeso.HQ Crimea WAsCRO(UJ Cymbalaria muralis *DHUWQMey Scherb6FURSKXODULDFHDH%LHQQMesoxero.HQ0HGCRO(UJ Daphne laureola LThymelaeaceae, Shrub, Meso.HQ0HG(UJDVSKLJ(UJ Datura innoxia Mill6RODQDFHDH$QQXeromeso.HQ1$P(SKHP(UJ Datura meteloides '&6RODQDFHDH$QQXeromeso.HQ1$P(UJDVSKLJ(UJ %LRGLY5HV&RQVHUY 41 Datura tatula L6RODQDFHDH$QQXeromeso.HQ1$P(UJDVSKLJ(UJ Dianthus barbatus L&DU\RSK\OODFHDH3HUHQQMeso.HQMiddle Eur(UJDVSKLJ(UJɊɨɝɨɜɢɱ Digitalis lanata Ehrh6FURSKXODULDFHDH3HUHQQMeso, .HQ SEurBalcan, CRO(UJ Digitalis purpurea L6FURSKXODULDFHDH$QQ.HQMeso, Eur0HG(UJDVSKLJ(UJ Diospyros lotus/,(EHQDFHDH7UHH;HURPHVR.HQ Crimea ($VɋRO(UJ Dipsacus sativus L +RQFN'LSVDFDFHDH%LHQQ;HURPHVR.HQunknown Pyãek et al. (UJDVSKLJ(UJ Disphania schraderiana Schult Mosyakin & Clemants&KHQRSRGLDFHDH$QQ;HURPHVR.HQAfr tropic and subtropic Ergasphig(UJ Dracocephalum moldavica L/DPLDFHDH$QQ;HURPHVR.HQ($V(UJDVSKLJ(UJ Duchesnea indica Andr Focke5RVDFHDH3HUHQQ;HURPHVR.HQS($V(UJDVSKLJ(UJ Echinacea purpurea L Moench$VWHUDFHDH3HUHQQ;HURPHVR.HQ1$P(UJDVSKLJ(UJ Echinochloa esculenta ABr +Scholz3RDFHDH$QQ;HURPHVR.HQ($V(UJDVSKLJ(UJ Echinocystis lobata Michx Torr *UD\&XFXUELWDFHDH$QQ0HVR.HQ 1$P$JUHSRHFErg invasive; transformer Transcarpathia Elaeagnus angustifolia L(ODHDJQDFHDH7UHHor shrub, Xeromeso, .HQ0HG$JU(UJ invasive; transformer Black Sea Region Elaeagnus commutata Bernhex Rydb(ODHDJQDFHDH7UHHor shrub, Meso.HQ1AmCRO(UJ Elodea canadensis Michx+\GURFKDULWDFHDHPerenn+ydro, .HQ 1$P$JU(UJinvasive Elodea densa Planch Caspari+\GURFKDULWDFHDHPerenn+\GUR.HQ SAm(SKHP(UJ Elodea nuttallii 3ODQFKRQ +6W-RK Caspari+\GURFKDULWDFHDHPerenn+\GUR.HQ 1$P(SKHP(UJ Eruca vesicaria L Cav%UDVVLFDFHDH$QQ;HURPHVR.HQEMed(SKHPErg Eschscholzia californica Cham3DSDYHUDFHDH$QQMesoxero.HQ1$P(UJDVSKLJ(UJ Eudianthe coeli-rosa L Rchb&DU\RSK\OODFHDHAnnXeromezo.HQ &ULPHD , :0HG(SKHP, .VHQR(UJ Euonymus japonica Thunb&HODVWUDFHDH6KUXE0HVR.HQ($V(UJDVSKLJ(UJ Euonymus sacrosancta Koidz&HODVWUDFHDH6KUXEMeso.HQ($V(UJDVSKLJ(UJ Euphorbia lathyris L(XSKRUELDFHDH$QQMesoxero.HQ0HG(UJDVSKLJ(UJ Euphorbia marginata Pursh(XSKRUELDFHDH$QQMesoxero.HQ1$P(UJDVSKLJ(UJ Fagopyrum esculentum Moench3RO\JRQDFHDH$QQMeso.HQCAs(UJDVSKLJ(UJ Fallopia baldschuanica Regel +ROXE, Polygonaceae, Liana, Xeromeso.HQMiddle As(UJDVSKLJ(UJ Ficus carica L0RUDFHDH7UHHor shrub, Xero.HQAs0HG(SRHF(UJ Flueggea suffruticosa Pall Baillon, Euphorbiaceae, Shrub, Mesoxero.HQ ($VCRO(UJ Foeniculum vulgare Mill$SLDFHDH%LHQQXeromeso.HQ0HGCRO(UJ Fragaria ×ananassa 'XFKHVQH 'XFKHVQH5RVDFHDH3HUHQQMeso.HQhybrid origin(UJDVSKLJ(UJ Fraxinus ornus L2OHDFHDH7UHHMesoxero.HQ0HG$JU(UJ invasive Fraxinus pennsylvanica Marsh2OHDFHDH2OHDFHDH7UHHXeromeso.HQ1$PɋRO(UJ Fritillaria ophioglossifolia Freun Sint/LOLDFHDHPerennMeso, .HQ Crimea CaucCROErg Gaillardia pulchella Foug$VWHUDFHDHAnnor bLHQQMesoxero.HQAmCROErg Gleditsia triacanthos L&DHVDOSLQLDFHDH7UHHXeromeso.HQ1Am(UJDVSKLJ(UJ Glycine max L Merr)DEDFHDH$QQ.HQXeromeso($V(UJDVSKLJ(UJ Grossularia uva-crispa L Millsspreclinata L 'RVWDO*URVVXODULDFHDH6KUXEMeso.HQEur(UJDVSKLJ(UJ Gypsophila acutifolia Fischex Spreng&DU\RSK\OODFHDH3HUHQQXeromeso.HQCauc(UJDVSKLJ(UJ Gypsophila elegans MBieb&DU\RSK\OODFHDH$QQXeromeso.HQCaucAsia Minor or Eur0HG Pyãek et al. (SKHP(UJ Gypsophila paulii Klok&DU\RSK\OODFHDH3HUHQQXeromeso.HQ SEUkraine, 2VWDSNR, Boyko, Mosyakin SCis Black Sea, (UJDVSKLJ(UJ Halimodendron halodendron Pall Voss, Fabaceae, Shrub, Meso, .HQunknown(SKHP(UJ Helianthus annuus Lvarmacrocarpa Lucznik$VWHUDFHDH$QQXeromeso.HQ1$P(SKHP(UJ Helianthus decapetalus L$VWHUDFHDH3HUHQQMeso.HQ1$P(SRHF(UJ Helianthus ×ODHWLÀRUXV Pers$VWHUDFHDH3HUHQQXeromeso.HQ1$P$JUHSRHF(UJ invasive Helianthus rigidus Cass 'HVI$VWHUDFHDH3HUHQQ Xeromeso, .HQ1$P(UJDVSKLJ(UJ Helianthus strumosus L$VWHUDFHDH3HUHQQMeso, .HQ1$PɋRO(UJ Helianthus subcanescens A*UD\ E EWats$VWHUDFHDH3HUHQQMeso.HQ1$P$JUHSRHF(UJ invasive Helianthus tuberosus L$VWHUDFHDH3HUHQQMeso.HQ1$P$JUHSRHF(UJ invasive; transformer Transcarpathia Helichrysum italicum Roth *'RQ ¿O$VWHUDFHDHSshrub, Xero.HQ Crimea – 0HGCRO(UJ Heliopsis scabra 'XQ$VWHUDFHDH3HUHQQXeromeso, .HQ1$PCRO(UJ Helleborus dumetorum Waldst Kit5DQXQFXODFHDH3HUHQQMeso.HQSEur(UJDVSKLJ(UJ Helleborus niger L5DQXQFXODFHDH3HUHQQMeso.HQSEur(UJDVSKLJ(UJ1RWHThe species included in to Red 'DWD Book of Ukraine, according to SMosyakin and MFedoronchuk probably cultivated and HVFDSHG Helleborus viridis L5DQXQFXODFHDH3HUHQQMeso.HQSEur(UJDVSKLJ(UJ Hemerocallis fulva L L, +HPHURFDOOLGDFHDH3HUHQQXeromeso, .HQ($V(UJDVSKLJ(UJ Heracleum mantegazzianum Sommier & Levier$SLDFHDH3HUHQQ+\JUR.HQ Cauc$JUHSRHFErg invasive; transformer Carpathian Heracleum sosnowskyi 0DQGHQ Apiaceae3HUHQQ+\JUR.HQCauc$JUHSRHFErginvasive; transformer Carpathian, Polissya [Forest zone of Ukraine@ Hesperis matronalis L%UDVVLFDFHDHAnnor bLHQQ;HURPHVR.HQ left forest and steppe regions of Ukraine Sub0HG ,O\LQVND et al. or Eurand0HG Pyãek et al. (UJDVSKLJ(UJ Hesperis pycnotricha Borbas & 'HJHQ%UDVVLFDFHDH%LHQQMeso.HQCaucAsmin(UJDVSKLJ(UJ Hesperis sibirica L%UDVVLFDFHDH%LHQQMeso.HQAs(UJDVSKLJ(UJ Hordeum distichon L3RDFHDH$QQXeromeso.HQEMed(UJDVSKLJ(UJ 42 9LUD93URWRSRSRYD 0\URVODY96KHYHUD (UJDVLRSK\WHVRIWKH8NUDLQLDQÀRUD Hordeum jubatum L3RDFHDH$QQMesoxero.HQ1$P(SKHP(UJ Hordeum vulgare L3RDFHDH$QQXeromeso, .HQAs(UJDVSKLJ(UJ Humulus japonicus Siebold & =XFF, Cannabaceae, Liana, Meso, .HQ EAs(UJDVSKLJ(UJ Hypericum calycinum L+\SHULFDFHDH, Shrub, Xeromeso.HQE0HG(UJDVSKLJ(UJ Hyssopus RI¿FLQDOLV L/DPLDFHDH3HUHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Iberis amara L%UDVVLFDFHDH$QQXeromeso.HQ0HG Protopopova (UJDVSKLJ(UJ Iberis pinnata L%UDVVLFDFHDH$QQXero, .HQW0HG(UJDVSKLJ(UJ Iberis umbellata L%UDVVLFDFHDHAnnor bLHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Impatiens balfourii +RRNfBalsaminaceae$QQMeso.HQ W+\PDO(SKHP(UJ Impatiens glandulifera Royle, Balsaminaceae$QQ+\GURmeso.HQ SEAs$JUHSRHF(UJ invasive; transformer Transcarpathia Ipomoea purpurea L Roth, Convolvulaceae$QQXeromeso.HQTropAm(UJDVSKLJ(UJ Iris ×germanica L,ULGDFHDH3HUHQQMeso.HQ Crimea hybrid or EurAs Pyãek et al. CRO(UJ Iris musulmanica Fomin,ULGDFHDH3HUHQQMeso, .HQCaucor Eur Bagrikova (UJDVSKLJ(UJ Iris pallida Lam,ULGDFHDH3HUHQQMeso, .HQ0HG(UJDVSKLJ(UJ Iva xanthiifolia 1XWW$VWHUDFHDH3HUHQQXeromeso.HQ 1$P(SRHF(UJ invasive Jasminum QXGLÀRUXP Lindl2OHDFHDH6KUXEXeromeso.HQ($V(UJDVSKLJ(UJ Jasminum RI¿FLQDOH L2OHDFHDH6KUXEMeso.HQEMed(UJDVSKLJ(UJ Juglans cinerea L-XJODQGDFHDH, Tree, Meso.HQ1$P(UJDVSKLJ(UJ Juglans mandshurica Maxim-XJODQGDFHDH7UHHMeso, .HQAs(UJDVSKLJ(UJ Juglans nigra L-XJODQGDFHDH7UHHMeso.HQ1 $P(UJDVSKLJ(UJ Juglans regia/-XJODQGDFHDH7UHH0HVR.HQ%DOFDQ&As(UJDVSKLJ(UJ Kali collina Pall Akhani & E+Roalson&KHQRSRGLDFHDH$QQXero.HQTur(UJDVSKLJ(UJ Kochia scoparia L Schrad&KHQRSRGLDFHDH$QQMesoxero.HQ,UDQTur(UJDVSKLJ(UJ Koelreuteria paniculata Laxm, Sapindaceae, Tree or shrub, Meso, .HQ Crimea ($VCRO(UJ Laburnum anagyroides Medik)DEDFHDH7UHHs or shrub, Xeromeso.HQSEur(UJDVSKLJ(UJ Lactuca sativa L$VWHUDFHDH%LHQQMeso.HQunknown Pyãek et al. (UJDVSKLJ(UJ Lagurus ovatus L3RDFHDH$QQXero, .HQ Crimea 0HG Pyãek et al. (SKHPKseno(UJ Lallemantia iberica ɆBieb Fisch C AMey/DPLDFHDH$QQMesoxero.HQWAs(UJDVSKLJ(UJ Lamprocapnos spectabilis L Fukuhara Fumariaceae 3HUHQQMeso.HQ($V Mayorov et al. (UJDVSKLJ(UJ Lathyrus odoratus L)DEDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Lathyrus sativus L)DEDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Laurocerasus RI¿FLQDOLV MRoem5RVDFHDH7UHHor shrub, Xeromeso.HQMed,UDQTur$JU(UJ Laurus nobilis L, Lauraceae, Tree or shrub, Xeromeso.HQ0HG(UJDVSKLJ(UJ Lavandula angustifolia Mill/DPLDFHDH3HUHQQXeromeso.HQ0HG Pyãek et al. (UJDVSKLJ(UJ Lavatera trimestris L0DOYDFHDH$QQXeromeso.HQ0HG(UJDVSKLJ(UJ Lens culinaris Medik)DEDFHDH$QQMeso.HQunknown Pyãek et al. 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(UJDVSKLJ(UJ Prunus cerasifera Ehrh5RVDFHDH7UHHor shrub, Xeromeso.HQ0HG(UJDVSKLJ(UJ Prunus divaricata Ledeb5RVDFHDH7UHHor shrub, Xeromeso, .HQAs(UJDVSKLJ(UJ Prunus domestica L5RVDFHDH7UHHor shrub, Xeromeso.HQhybrid Tzvelev (UJDVSKLJ(UJ Prunus insititia L5RVDFHDH6KUXEor tree, Xeromeso.HQAs(UJDVSKLJ(UJ Ptelea trifoliata/, Rutaceae, Tree, Meso.HQ1$P(UJDVSKLJ(UJ Pueraria lobata :LOOG 2KZL)DEDFHDH6KUXEMeso.HQ Crimea S($VCROErg Pyracantha rogersiana A B-DFNV Bean5RVDFHDH6KUXE Meso, .HQ Crimea AsCRO(UJ Pyrethrum cinerariifolium Trev$VWHUDFHDH3HUHQQXeromeso.HQBalcan(UJDVSKLJ(UJ Pyrethrum macrophyllum Waldst Kit Willd$VWHUDFHDH3HUHQQMeso.HQE0HG(UJDVSKLJ(UJ Pyrethrum partheniifolium Willd$VWHUDFHDH3HUHQQMesoxero.HQMed,UDQTur(UJDVSKLJ(UJ Pyrethrum parthenium L Smith$VWHUDFHDH3HUHQQMesoxero.HQWEur(UJDVSKLJ(UJ Pyrus communis L5RVDFHDH7UHHMeso, .HQunknown Pyãek et al. or As Bagrikova $JU(UJKseno Quercus ilex L, Fagaceae, Tree, Mesoxero.HQ Crimea 0HG(UJDVSKLJ(UJ Quercus palustris Moench, Fagaceae, Tree, Meso.HQ1$P$JU(UJ Quercus rubra L)DJDFHDH7UHHMeso.HQ1$P$JU(UJ invasive Raphanus sativus L%UDVVLFDFHDHAnnor bLHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Reynoutria ×bohemica Chrtek & Chrtkova3RO\JRQDFHDH3HUHQQMeso.HQhybrid origin(SRHF(UJ invasive Reynoutria japonica +RXWW3RO\JRQDFHDH3HUHQQMeso, .HQ EAsAgrepoec(UJ invasive; transformer Transcarpathia Reynoutria sachalinensis FrSchmidt 1DNDL3RO\JRQDFHDH3HUHQQMeso.HQ EAsCRO(UJ Rhamnus alaternus L, Rhamnaceae, Shrub, Xeromeso.HQ0HG$JU(UJ invasive; transformer Crimea Rhus typhina L, Anacardiaceae, Tree, Xero.HQ1$P(UJDVSKLJ(UJ Ribes aureum Pursh*URVVXODULDFHDH6KUXEMeso.HQ1$P(UJDVSKLJ(UJ Ribes rubrum L*URVVXODULDFHDH6KUXEMeso.HQWEur(UJDVSKLJ(UJ Ribes spicatum ERobson*URVVXODULDFHDH6KUXEMeso.HQ Crimea QDWXUDOVSHFLHVLQ8NUDLQH, EurAs Pyãek et al. or Eur Bagrikova CRO(UJ Ricinus communis L(XSKRUELDFHDH$QQMeso.HQAfr(UJDVSKLJ(UJ Robinia hispida L)DEDFHDH6KUXEXeromeso.HQ1$PCRO(UJ Robinia pseudoacacia L)DEDFHDH7UHHXeromeso.HQ1$P(SRHF(UJ invasive Rosa alba L5RVDFHDH6KUXEXeromeso.HQhybrid origin(UJDVSKLJ(UJ Rosa centifolia L5RVDFHDH6KUXEXeromeso.HQhybrid origin(UJDVSKLJ(UJ Rosa PXOWLÀRUD Thunb5RVDFHDH6KUXEXeromeso.HQ($V(UJDVSKLJ(UJ Rosa rugosa Thunb5RVDFHDH6KUXEXeromeso, .HQ($VCRO(UJ Rosmarinum RI¿FLQDOLV L/DPLDFHDHSshrub, Mesoxero.HQ0HG(UJDVSKLJ(UJ Rubia tinctorum LRubiaceae3HUHQQMesoxero.HQMed,UDQTurCRO(UJ Rubus idaeus L5RVDFHDH6KUXEMeso.HQ Crimea natural species in Ukraine, Eur$JU(UJ Rubus laciniatus Willd5RVDFHDH6KUXEMeso, .HQ unknown Pyãek et al CRO(UJ Rubus macrophyllus Weihe & 1HHV, Rosaceae, Shrub, Meso.HQ SEof Ukraine, 2VWDSNR, Boyko, Mosyakin CEur Krassovskaya ColKseno(UJ Rudbeckia bicolor 1XWW$VWHUDFHDH3HUHQQMeso.HQ1$P(UJDVSKLJ(UJ Rudbeckia hirta L$VWHUDFHDHAnnor bLHQQXeromeso.HQ1$PCRO(UJ Rudbeckia laciniata L$VWHUDFHDH%LHQQ+\JURmeso.HQ1$P$JUHSRHF(UJ invasive; transformer Transcarpathia, Bukobinian Cis Carpathian Rumex patientia L3RO\JRQDFHDH3HUHQQXeromeso, .HQ0HG(UJDVSKLJ(UJ Sagittaria platyphylla Engelm -*6PLWK$OLVPDWDFHDHPerenn0HVRK\JUR.HQ 1$P(UJDVSKLJ(UJ Salix babylonica L6DOLFDFHDH7UHHMeso.HQCEAsEphem(UJ Salix fragilis L6DOLFDFHDH7UHHVMeso$UFKAsmin$JU(UJ invasive; transformer central regions of Ukraine Salvia RI¿FLQDOLV L/DPLDFHDH3HUHQQXeromeso.HQ0HG(UJDVSKLJ(UJ Salvia splendens Ker *DZO/DPLDFHDH3HUHQQXero.HQSAm(SKHP(UJ Sambucus racemosa L, Sambucaceae, Shrub or tree, Meso, KenWEur(UJDVSKLJ(UJ Santolina chamaecyparissus L$VWHUDFHDH6KUXEMesoxero.HQ0HG(UJDVSKLJ(UJ Santolina virens Mill$VWHUDFHDH6KUXEMesoxero.HQSEur(UJDVSKLJ(UJ Saponaria RI¿FLQDOLV L&DU\RSK\OODFHDH3HUHQQXeromeso.HQ0HG$JUHSRHF(UJ Satureja hortensis L/DPLDFHDH$QQXeromeso.HQ0HG Protopopova or CaucAsMin Mayorov et al. 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Ergasphig(UJ Vitis vulpina L9LWDFHDH/LDQDXeromeso, .HQ1AmCRO(UJ Zea mays L3RDFHDH$QQXeromeso.HQCSAmErgasphig(UJ Zelkova carpinifolia Pall KKoch, Ulmaceae, Tree, Meso.HQ Crimea CaucCRO(UJ Zinnia elegans -DFT$VWHUDFHDH$QQMeso.HQ1$PErgasphig(UJ Zizania aquatica L3RDFHDH$QQ+\GUR.HQ1$PCRO(UJ Zizania latifolia *ULVHE Stapf3RDFHDH3HUHQQ+\GUR.HQ($V$JU(UJ Explanations: life forms, Ann±annual, Bien±biennial, Annbienn±annual or biennial, Perenn±perennial; hydromorphe, +\GUR – hydrophytes, +\JUR – hygrophytes, +\JURmeso – hygromesophytes, Meso – mesophytes, Mesohygro – mesohygrophyte, Mesoxero – mesoxerophytes, Xero – xerophytes, Xeromeso – xeromesophytes; chronological elements, Arch±archaeophytes, Ken±kenophytes; degree of naturalization, Agr±agriophytes, Agr epoec±agrioepoecophytes, Col±colonophytes, Ephem±ephemerophytes, Epoec±epoecophytes, Ergasphig±ergasiophygophytes; origin, Afr± African, Am±American$V±Asia, As0LQ±$VLD0LQRU$V0LQ0LGGOH$V±$VLD0LQRU0LGGOH$VLD%DOF±%DOFDQ%DON&$V±%DOFDQ&HQWUDO $VLDQ&$P±&HQWUDO$PHULFDQ&$V±&HQWUDO$VLDQ&(XU±&HQWUDO(XURSHDQ&DXF±&DXFDVLF&DXF$V0LQ±&DXFDVLF$VLD0LQRU&DXF $V 0LQ0HG± &DXFDVLF$VLD0LQRU±0HGLWHUUDQHXP&DXF(0HG± &DXFDVLF(DVWHUQ0HGLWHUUDQHXP&DXF(XU± &DXFDVLF(XURSHDQ($V± Eastern Asia, (XU±(XURSHDQ(XU$V±(XURSHDQ$VLDQ(XU&DXF±(XURSHDQ&DXFDVLF(0HG±(DVWHUQ0HGLWHUUDQHXP(0HG$V0LQ±(DVWHUQ 0HGLWHUUDQHXP$VLD0LQRU)DU(±Far East, ,UDQ7XU±,UDQ7XUDQLDQ0HG±0HGLWHUUDQHXP0HG$IU$V±0HGLWHUUDQHXP$IULFDQ$VLDQ0HG$V ±0HGLWHUUDQHXP$VLDQ0HG,UDQ7XU±0HGLWHUUDQHXP,UDQ7XUDQLDQ0HG0LGGOH$V±0HGLWHUUDQHXXP0LGGOH$VLDQ0LGGOH$V±0LGGOH$VLD 0LGGOH$V±0LGGOH$VLDQ0LGGOH(XU±0LGGOH(XURSHDQ0LGGOH(XU%DON±0LGGOH(XURSHDQ%DONDQ1$IU±1RUWKHUQ$IULFDQ1:$IU±1RUWKHQ Western African, 1$P±1RUWK$PHULFDQ13DQQ–1RUWKHUQ3DQQRQLDQ6&LV%ODFN6HD±6RXWK&LV%ODFN6HD6$P±6RXWK$PHULFDQ6$V±6RXWK $VLD6($V±6RXWK(DVWHUQ$VLD6(7UDQVFDXF±6RXWK(DVWHUQ7UDVQFDXFDVLFDQ6(XU±6RXWK(XURSHDQ6LE±6LEHULDQVXE$OS±VXEDOSLF 7URS$P±7URSLFDO$PHULFDQ7XU±7XUDQLDQ:$V±:HVW$VLD:$V0LQ±:HVWHUQ$VLD0LQRU:&(XU±:HVW&HQWUDO(XURXSHDQ:0HG± :HVW0HGLWHUUDQHXP:(XU±:HVW(XURSHDQ; way of distribution,(UJ±HUJDVLRSK\WHV(UJNVHQR±HUJDVLRNVHQRSK\WHV