The Establishment and Spread of Alien Plant Species (Kenophytes) in the Flora of Poland

Barbara Tokarska-Guzik

Wydawnictwo Uniwersytetu Śląskiego • Katowice 2005

The Establishment and Spread of Alien Plant Species (Kenophytes) in the Flora of Poland

To my husband PRACE NAUKOWE

UNIWERSYTETU ŚLĄSKIEGO W KATOWICACH NR 2372 Barbara Tokarska-Guzik

The Establishment and Spread of Alien Plant Species (Kenophytes) in the Flora of Poland

Wydawnictwo Uniwersytetu Śląskiego Katowice 2005 Editor of the Series: Biologia Paweł Migula

Reviewers Bogdan Jackowiak Adam Zając

Cover design: Marek Francik Published by Executive Editor: Wioletta Tomala-Kania Wydawnictwo Uniwersytetu Śląskiego Technical Editor: Barbara Arenhóvel ul. Bankowa 12B, 40-007 Katowice www.wydawnictwo.us.edu.pl Proof-reader: Grażyna Wojdała e-mail: [email protected] Copyright © 2005 by First impression. Edition: 200 + 50. Printed sheets: 24,5 + insert. Publishing sheets: 31,5. Passed to the Wydawnictwo Uniwersytetu Śląskiego Printing House in September 2005. Signed for print- Ali rights reserved ing and printing fmished in December 2005. C ena 57 zł ISSN 0208-6336 Czerny Marian. Firma Prywatna „GREG” ISBN 83-226-1485-3 Zakład Poligraficzny ul. Wrocławska 10, 44-110 Gliwice Contents

A cknow ledgem ents...... 9

PART ONE T h eo retica l o v e r v i e w ...... 11 1. Introduction. The subject, objectives and the scope of this study: The role of kenophytes in the flora as representations of the anthropogenic alteration of v e g e ta tio n ...... 11 2. Review of studies on selected aspects of synanthropisation of the vegetation cover...... 14 2.1. The history of studies on alien plant species in Poland viewed against the situation in Europę as a whole...... 14 2.2. Synanthropisation: the essence of the process and the role of kenophytes in the changes occurring in the natural environment on E a r t h ...... 19

PART TWO T erm inology a n d m e th o d o lo g y ...... 23 3. Phytogeographical terminology and the classification of synanthropic plants used in Poland . . . 23 4. Materials and m e th o d s ...... 24 4.1. Selection of species and their s t a t u s ...... 24 4.2. Sources and characteristics of the floristic data used...... 25 4.3. List of kenophytes and the scope of the information collected in order to characterise them . 26 4.4. Cartogrammes and their a n a ly s is ...... 27 4.5. Use, interpretation and synthesis of d ata...... 27

PART THREE A nalysis an d sy n th esis o f d a t a ...... 29 5. Geographical and ecological characteristics of the flora of kenophytes in P o la n d ...... 29 5.1. Proportion of kenophytes in the recent f l o r a ...... 29 5.1.1. General r e m a r k s ...... 29 5.1.2. O r ig in ...... 30 5.1.3. Timing and method of a r r i v a l ...... 31 5.1.4. Systematic g r o u p s ...... 31 5.1.5. Groups of life f o r m s ...... 34 5.1.6. Biological p ro p e rtie s ...... 35 5.1.7. Freąuency and status in the f l o r a ...... 36 5.2. Kenophytes in historical accounts of floras...... 40 5.2.1. “Old” f l o r a s ...... 40 5.2.2. The “oldest” arrivals among the kenophytes and the fairly recent o n e s ...... 46 5.2.3. The most frequent kenophytes in the floras of subsequent historical periods...... 46 6. Current types of distribution of kenophytes in Poland...... 53 6.1. Kenophytes with stations scattered throughout Poland except for in certain regions .... 54 6.1.1. Sisymbrium altissimum g r o u p ...... 55 6.1.2. Diplotaxis tenuifolia g r o u p ...... 56 6.2. Kenophytes with scattered stations over the whole territory of Poland, with concentrations of morę frequent stations in some r e g io n s ...... 56 6.2.1. Bunias orientalis g r o u p ...... 57 6.2.2. Geranium pyrenaicum g r o u p ...... 58 6.2.3. Echinocystis lobata g r o u p ...... 59 6.3. Kenophytes (contemporarily) reaching their limit of distribution in P o la n d ...... 60 6.3.1. Western l i m i t ...... 60 6.3.2. Eastern l i m i t ...... 61 6.3.3. Northern l i m i t ...... 62 6.4. Kenophytes associated with river v a lle y s ...... 64 6.5. Kenophytes associated with urban areas and railway ro u tes...... 66 7. The history of the spread of selected kenophyte species in the territory of Poland ...... 67 7.1. The history of the spontaneous spread of cultivated woody plants as the result of “domesticating” s p e c i e s ...... 67 Acer negundo L...... 67 Padus serotina (Ehrh.) Borkh...... 68 Ailanthus altissima (Mili.) Sw ingle...... 71 Clematis vitalba L...... 71 7.2. The history of the spread of useful herbaceous plant species: how medicinal and decorative plants have established themseWes in the f l o r a ...... 74 7.2.1. Examples of species of European o r ig in ...... 74 Cymbalańa muralis P. Gaertn., B. Mey. & Scherb...... 74 Digitalis purpurea L...... 74 Echinops sphaerocephalus L...... 78 7.2.2. Е хатріе of species of Asian o r i g i n ...... 78 Elsholtzia ciliata (Thunb.) Hyl...... 78 Impatiens glandulifera R o y l e ...... 79 Impatiens parviflora DC...... 81 Reynoutria japonica Houtt...... 84 7.2.3. Examples of species of American o r ig in ...... 86 Echinocystis lobata (E Michx.) Torr. & A. G r a y ...... 86 Rudbeckia laciniata L...... 88 Mimulus guttatus DC...... 88 7.3. The spread of accidentally introduced plants: how an ephemerophyte turns into a kenophyte. 91 7.3.1. Plants introduced accidentally from various regions of E u r o p ę ...... 91 Anthoxanthum aristatum Boiss...... 91 Artemisia austriaca Jacq...... 91 Bunias orientalis L...... 94 Eragrostis minor H o s t ...... 96 Rumex confertus Willd...... 96 Salsola kali L. subsp. ruthenica (Iljin) S o ó ...... 99 7.3.2. Plants brought accidentally from A s ia ...... 101 Sisymbrium altissimum L ...... 101 Veronica persica Poir...... 101 7.3.3. Plants brought accidentally from A m e r i c a ...... 104 Bidens frondosa L...... 104 Chamomilla suaveolens (Pursh) Rydb...... 104 Elodea canadensis Michx...... 107

PART FOUR D i s c u s s i o n ...... 109 8. The proportion and role of alien species in the flora: do kenophytes determine the recent shape of the flora of P o lan d ?...... 109 9. Historical aspects of the development of the kenophyte flora of P o la n d ...... 117 9.1. General r e m a r k s ...... 117 9.2. The effect of historical and economic developments on the enrichment of Polish flora by new- c o m e r s ...... 117 9.3. Cities as “footholds” for further expansion by fresh new com ers ...... 118 9.4. Historical gardens, botanic gardens, cloister and convent gardens as places of “domesticating” exotic species prior to their spontaneous establishm ent...... 124 9.5. Immigration periods (peak inflows of kenophytes)...... 126 9.6. Migration r o u t e s ...... 127 9.6.1. Rivers as migration corridors aiding the spread of kenophytes...... 128 9.6.2. The role of humans in the migrations of kenophytes...... 129 10. Recent distribution ranges of kenophytes and principles affecting the distribution pattern . . . 130 11. Dynamie tendencies in the process of kenophyte expansion in P o la n d ...... 133 12. Plant invasions: the substance of the phenomenon and kenophytes as invasive p lan ts...... 141 12.1. More remarks on te rm in o lo g y ...... 141 12.2. Consequences of invasions by alien species, legał regulations and methods of combating the th r e a t...... 144 12.3. Invasive kenophytes in P o la n d ...... 146 12.4. Threatened regions and h a b i t a t s ...... 150 12.5. Forecasting invasions: potentially invasive s p e c i e s ...... 152 12.6. Finał r e m a r k s ...... 153

PART FIVE Summary, conclusions and the perspectives for studies of plants of ałien origin in Poland against the trends prevailing in Europę and the w o r l d ...... 155 13. Summary and c o n c lu s io n s ...... 155 14. Invasions of alien plant species at the dawn of the 21st century: perspectives for further studies . 157

R e fe r e n c e s ...... 159

S treszczen ie...... 189

Z usam m enfassung...... 190

A p p en d ices...... 191

Acknowledgements

This thesis has been written thanks to the help The group of Colleagues to whom I owe a word of many people. I wish to express my warmest of gratitude for steadfast collaboration during lo- gratitude to my Teachers for awakening my in- calisation of sites includes Dr. Barbara Fojcik, Dr. terest in the problems of synanthropisation of Gabriela Woźniak, as well as MSc. Jadwiga Du- plant cover: Professor Dr. hab. Krzysztof Ros- czek (an economist by education) and her daugh- tański, Professor Dr. hab. Adam Zając and Pro ter Martyna. fessor Dr. hab. Janusz Bogdan Faliński, who I would like to give my special thanks to Professor sadly passed away recently. I have always expe- łan C. Trueman and Dr. Eleanor Cohn (University rienced great kindness and eagerness during long of Wolverhampton, UK) who have always shown and insightful scientific discussions on the part personal interest in my scientific life and have of my Teachers. encouraged me in the field of my research. Espe- I am extraordinarily glad to be able to express cially Professor łan C. Trueman who has sup- my indebtedness to all my Co-workers at the ported me through interesting discussions and Department of Plant Systematics, as well as to helpful comments. Colleagues from the Departments of Geobotany I am grateful to MSc. Maryla Palowska for her and Naturę Protection, as well as Biophysics and extensive and extremely time-consuming technical Celi Biology, at the University of Silesia for their help in preparation of the graphical part of the help during my investigations and during the present work. I wish to thank my Colleagues preparation of this thesis for publication. Dr. Teresa Nowak and MSc. Anna Gawron for in I wish to thank all Polish Botanists whose data, sightful reading of the manuscript and kind en- submitted to the ATPOL database, have made it couragement. possible for me to prepare this thesis. I thank those I wish to express my sincere gratitude to Par- Colleagues who sent their data collected during ticipants of discussions conducted during my pre- field investigations directly to me: Dr. Zygmunt sentations of successive stages of studies on the Dajdok and Dr. Zygmunt Kącki (University of naturalisation and spreading of kenophytes in Wrocław), Dr. hab. Marek Kucharczyk and Dr. Poland during geobotanical seminars in Biało Rafał Krawczyk (University of Lublin), Dr. Dan wieża and Katowice. Wołkowycki (University of Białystok), Dr. Marze I give my warm feelings of gratitude to my na Środa (University of Warmia and Mazury) and Husband Zenek in whom I have always found Dr. Zofia Sotek (University of Szczecin). support, to my daughter Zuzanna and son Kuba, I would like to give my thanks to Professor Dr. as well as to my Parents, for their patience and hab. Alicja Zemanek for her critical interpreta- constant presence at each moment of preparation tion of the 17thcentury work of Sirenius. of this thesis. I want to thank Mr. Józef Gajda from the In- In the years 1997-2000, the investigations stitute of Informatics of the Jagiellonian Univer- were conducted as an integral part of research sity for making his proprietary Computer program grant no. 6P04G05312 “The occurrence, history available to me and for extensive help in the and modern distribution of kenophytes (neo- preparation of the cartographic section of my phytes) in Poland” funded by the State Commit- thesis. tee for Scientific Research.

2 The Establishment. 9

PART ONE

Theoretical overview

“The life success of each plant species hinges on not only the capability to settle passively in the places of its earlier occurrence, but also on the ability to actively conquer such places”. Pa c z o s k i 1933: Podstawowe zagadnienia geografii roślin [The fundamental issues of plant geography]

1. Introduction about useful plants, particułarly those that are edible The subject, objectives or poisonous, as well as on natural curiosity and and the scope of this study: a determination to leam about new, exotic species. The role of kenophytes in the flora Practical considerations were also important and the as representations of the anthropogenic ambition which drives explorers, both past and contemporary, to search for new plants in newly alteration of vegetation discovered remote parts of the world. As early as in the ancient times, the body of knowledge accu- The subject of this book falls within the theme mulated by naturalists and philosophers such as of the synanthropisation' of vegetation cover. Theophrastus, Dioscorides and Pliny was impres- Connected representations of this directional pro- sive in terms of volume and provided a source for cess occurring on Earth under the impact of var- copies, adaptations and reprints for the “herbalists” ious forms of human activities, are the processes of the Middle Ages and Renaissance periods. of the extinction of some species and the expan- The studies of the floras which accompany sion of others, which have both accelerated in people inereased greatly from the beginning of the recent centuries and which are contributing to 19th century. The oldest works devoted to plants changes of the biological diversity of entire re- of foreign origin, however, date back to the 17th gions, countries or continents. A synthesis of the century. At this early date, an Italian botanist, role of humans in the historie changes in land- Prosper A lpin u s published a work entitled De scape and vegetation cover was presented by plantis exoticis (1627), where he gave descriptions K ornaś (1977a) in a multi-authored book entitled of plants found in Europę but originating from Szata roślinna Polski [The vegetation of Poland] America. Other proofs of naturalists’ interest in and in other detailed papers (K ornaś 1982, 1983, such plants can also be found in the old herbals2. 1990, 1996). Dynamie change in floras, its scalę The phenomenon of invasion by alien new- and rate - issues which have started to focus the comers in their new homelands was also noted by interest of scientists and conservationists - be- D arwin (1859) in his work On the origin o f species3, came the main motive for undertaking this study. as well as in the diaries of his joumeys, and in Nevertheless, the interest taken by scientists in 2 For example, the Wrocław Herbarium collection (WRSL) territorially-expanding plant species of foreign has one of the oldest herbaria in Europę which was assem- origin has its roots in ancient times and was con- bled by an Italian Sivius Boccon, dated 1674 (R o sta ń sk j K. ceived on the basis of an ever-increasing knowledge 1963). This includes a typical specimen of Solidago canadensis L. - a recent kenophyte (neophyte), distributed throughout Europę, and originating from North America. 1 Synanthropisation - is the process of change in plant 3 Among other examples, Darwin described the invasion cover (also in the fauna and the abiotic elements of the en- of Cardo de Castilla (Spanish Cardoon) Cynara cardunculus L. vironment) brought about by human impact (for detailed brought to Buenos Aires in 1749, and which had taken over definition sec Chapter 2.2). Argentina, Chile and Uruguay in eight decades (C ro sb y 1999). letters and research reports. More information of the “pushy Mongoł” (F aliński 2004 after testifying to the perception of the phenomenon Naumann 1913). Similar associations had been and the consequences thereof, can be found in provoked by the invasion of European plants in numerous notes and Communications published in other continents. The native Americans of New popular scientific journals4. England and Virginia called Plantago major The plants of foreign origin appearing in the (Greater Plantain) “Englishman’s footprint”, be floras of many regions of the world were called cause in the 17thcentury they believed that this “the vagrants of our floras” (C rosby 1999 after plant grew only “where the aliens set their feet Hooker 1864) or “new acąuisitions” (K amieński and where it had not been known before their 1884а & b; P aczoski 1896), although they were arrival in this country” (C rosby 1999). sometimes called “newcomers and waifs” or The migrations of species occurring as the re “wandering plants” (T rzebiński 1930; Szulczew - sult of human activity which often assumed the ski 1931) or - in some special cases - “invaders” characteristics of massive invasions (“ecological (E lton 1958). explosions”), and which led eventually to changes Answers were sought to a number of essential in vegetation, fauna and to economic damage, questions. From where did the alien species arrive constituted the topie of a book entitled The Eco- in the local flora? Is it an escape from any culti- logy of Invasions by Animals and Plants (1958), vation or has it been brought in accidentally? by E lton, a British ecologist, whose research in Which place does it occupy in the new homeland this field is considered to be classic. The date of and what consequences result from its arrival? the publication of the book can be regarded as the Nevertheless, the greatest attention was attracted birth of ecology of invasion as a new scientific by the spectacular manner of the arrival of those discipline. alien plant species that colonised new territories C rosby (1999), describing the successful colon- rapidly and in great numbers5. Many of those isation of the Globe by Europeans, even presented immigrants soon became burdensome acquisi- a hypothesis that the success of European impe- tions in the local flora, sometimes even earning rialism has an underlying biological and ecolo common names reflecting the violent manner of gical background (“ecological imperialism”). The their invasion. One such ехатріе is Elodea same author, giving examples of spectacular inva- canadensis (Canadian Waterweed) originating sions of the vast spaces of Australia or both from North America, which conquered European Americas, makes ironie comments: “A rapid inva- inland waters in a “blitz” in the second half of sion of species of European ‘weeds’ disturbed the 19'h century and the beginning of the 20lh cen- American naturalists, even though most of these tury, and which was called “the green ghost” botanists themselves hailed from the same region (F aliński 2004 after Lons 1910). An Asian as the plants concemed”. Despite the great distance species Impatiens parviflora (Smali Balsam), between these continents and Europę, the climate which dispersed over central Europę as a fugitive is similar in many regions, providing magnificent from botanic gardens having first established conditions for development of the European col- itself in ruderal communities and then succeeded onists, incłuding plants, animals and people. in entering the forests, has been given a nickname The actual scalę of the exchange of species of synanthropic plants between regions of the world 4 E.g. a column in Przyroda i Przemyśl [Naturę and is considerable. The proportion of alien species Industry], a weekly devoted to advancement of the natural naturalised (i.e. permanently established) in some sciences and their applications in industry, of 1872 pub local floras ranges from 20% to even as high as lished a note on the appearance of new plants after the 50%. Particularly dynamie is the exchange be Franco-Prussian War. This note was prepared on the basis tween Eurasia and North America (JAger 1988; of a study by de Vibraye (1870-1871) presented before the French Academy, in which the author describes the emer- S ukopp 1995; K ornaś 1996; J ackow iak 1999). gence of 157 new exotic plants in central France. He at- The invasions by plants, animals or fungi are tributed their presence in a new territory to an accidental one of the most pressing issues of naturę con introduction of seeds from Algeria by the French cavalry. sidered on a global scalę. Some authors even The author assessed this process as a permanent change in deem it to be the single most important problem the flora because “these plants not only withstood one of in protecting the biodiversity in the 2Г' century the most severe winters but flourished abundantly in the areas once ąuite devoid of plants. Thus we can be quite (C arlton & G eller 1993; V itousek et al. 1996, sure that it is not a temporary phenomenon but that essen- 1997; M ooney & H obbs 2000). The International tially some of the regions in France had their plant wild- Convention on Biological Diversity contains life augmented by new flora”. a special provision calling upon country-signato- 5 While describing the spreading of Spanish Cardoon, ries to fight alien invasive species which could Charles Darwin stated: “I doubt whether there was any such case in history of native flora being invaded by an alien be of danger to native habitats, communities or species on such a great scalę” (C r o s b y 1999). species. These circumstances have contributed to an evident increase in the interest in these species: native7 and alien, and the comparison issues among the theoreticians and practitioners between the group of the oldest companions of of naturę conservation. humans (archaeophytes, so called oldcomers) Studies devoted to the spread of alien plant and the newer alien types (kenophytes = neo- species are becoming almost as fundamental phytes, so called newcomers). a part of the protection of biological diversity as The issue of the origin and development of the the compilation of “Red Lists” and “Red Data distribution ranges of the oldest group of alien Books” of rare and endangered species (M edw ec- species occurring in Poland (i.e. archaeophytes) has ka-K ornaś & P ięk o ś-M irkow a 1997). The lists been taken up by Z ając in a basie monograph of alien species are compiled along with the lists (1979) and in detailed reports (Z ając 1983, 1987a of invasive species which have entered natural & b, 1988), while the same considerations for more and semi-natural communities, or - as weeds - recent newcomers (i.e. kenophytes) should begin to the segetal communities. However, one will not be addressed by the detailed maps included in be able to prevent their spread without knowledge Distribution Atlas of Yascular Plants in Poland of their biology and habitat reąuirements as well (T okarska-G uzik 200la & b; Z ając A. & Z ając M. as their geographical ranges of distribution. For 2001), as well as by the present monograph. ехатріе many plant species took less than two The main idea behind the present mono centuries to invade and occupy the whole national graph is to investigate changes in synanthropic flora of Poland and to provide a synthesis of territory of Poland (Z ając A. & Z a ją c M . 2001). The initiatives taken up in many countries, as the knowledge accumulated to date on the well as those undertaken on a European and/or development of the kenophyte flora of Po global scalę (e.g. the international programmes land. It is also an attempt to reconstruct the Ecology of Biological Invasion, Global Strategy historie changes in the ranges of distribution for Invasive Species, Global Invasive Species of kenophytes in the territory of Poland. Ad- ditionally, those regions of Poland which are Programme), have made studies of alien species, particularly endangered by the excessive inva- and in particular their extending distribution sion of alien species are indicated, and the “in- ranges, their ecology, and the effects exerted - an vasive species”8 are specifically identified in the urgent and indispensable task. first ever comprehensive list of invasive keno Both in Poland and other European countries, phytes compiled for Poland. studies on the migrations and distribution of alien These aims have been achieved by the follow- plant species have a certain tradition6, beginning ing objectives: from studies devoted to particular species, such as, • verifying and updating lists of kenophytes com for ехатріе, Bidens frondosa (T rzcińska 1961; piled for Poland and presenting an original, L hotskA 1966, 1968), Impatiens glandulifera comprehensive catalogue of this group of spe (Z ając E.U. & Z ając A. 1973; B eerung & P er- cies, with an associated database of biological, rins 1993; G udżinskas & S inkevićene 1995; P y- ecological, geographical and historical attributes śek & P rach 1995; D ajdok et al. 2003; D rescher and information; & P rots 2003), Iva xanthifolia (G uzik & S udnik- • establishing the first floristic data (first records) W ójcikowska 1989; G udżinskas 1991; J ehlik for particular species of Polish kenophytes; 1998) and Reynoutria (Fallopia) japonica (e.g. • studying the historie distribution of kenophytes C o n o lly 1977; Alberternst et al. 1995; S eig er and attempting to reconstruct the history of 1997; Bailey & Conolly 2000; Child & Wade kenophyte floras on the basis of distribution 2000; Tokarska-Guzik in press) or regions (e.g. maps, applying whenever possible a cartographic H o lz fu s s 1937; G u tte 1971; Lohmeyer & Su- interpretation; ko p p 1992; Brandes & Sander 1995; N a ta li & • attempting to reconstruct periods of immigration Jeanm onod 1996; A dam ow ski et al. 2002), up to and spread of kenophytes (construed as cumu- studies covering whole countries (e.g. Clement lations or “migration waves”) showing also how & F o s te r 1994; Jeh lik 1998; Z a ją c A. & Z a ją c M. they depended on historie and geographical 2001; Essl & Rabitsch 2002; P r e s to n et al. conditions; 2002; P y śek et al. 2002). The historical impact exerted by humanity on 7 Similarly important issue is the problem of “apophy- the vegetation cover, and the flora in particular, tisation” of native species, and its following conseąuences is best illustrated by examining two phenomena: in “invasibility” of this group of species beyoned their the interactions between the two groups of natural rangę (cf. Chapter 11 and 12). 8 Invasive species - species of foreign origin, established in a primarily foreign area, producing fertile offspring, 6 Outside Europę one can find numerous publications often in extraordinarily large numbers, dispersing over devoted to this issue (e.g. compare literature cited by great distances from parental plants (R ic h a r d s o n et al. B r u n d u et al. 2001 and C h il d et al. 2003). 2000); for terminology, see also Chapter 3 and 12. • identifying and describing the different pattems (1786, 1787, 1788). The latter author described of distribution of kenophytes in Poland; several hundred “native wild plants and alien plants • reconstructing the history of the introduction, which could be of use in our country” (“rośliny establishment and spread of selected species; krajowe dzikie oraz i cudzoziemskie, któreby • discussing dynamie trends in kenophyte distri- w kraju pożyteczne być mogły”). Most of the spe bution, examining routes and pathways of in- cies mentioned by Kluk were cultivated at that time vasion and the factors supporting the conąuest (e.g. Aesculus hippocastanum, Artemisia dracuncu- of various types of habitats, and identifying lus, Bryonia alba, Clematis vitalba, Helianthus areas most vulnerable to invasion (with prac- tuberosus, Hyssopus officinalis, Juglans regia, tical implications for naturę conservation). Robinia pseudoacacia, Rubus odoratus, Sedum album, Sinapis alba) and are now considered to be naturalised in the flora of our country. For certain species, some details of their status outside the 2. Review of studies on selected cultivated state are also included along with a aspects of synanthropisation description of the type of habitats entered by these of the vegetation cover species10. The studies of species of foreign origin were first included in a broadly defined discipline of 2.1. The history of studies on alien studies in plant geography. Prior to Darwin’s plant species in Poland viewed studies, i.e. roughly till the mid 19th century, most of against the situation in Europę the research activities concentrated around the as a whole collection of facts pertaining to the occurrence of species and the differentiation between the vegetation landscapes of the world (K ornaś & M ed- A short review of the history, research trends wecka-K ornaś 2002). In th e 18th century, under and main methods used to study alien plant spe the influence of work completed by Carl Linnaeus, cies in Poland was the subject of one of the pre- the first floristic accounts appeared in Europę, to vious paper (T okarska-G uzik 200la). The present be continued in the centuries that followed (e.g. chapter is a further attempt to provide a synthesis W illdenow 1787; F icinus 1821; R eichenbach of different aspects of studies on alien plant spe 1842; P eck 1865; N ymann 1878-1882; S chulze cies in Poland shown on wider perspective. 1881; Schmalhausen 1886). The descriptions of The discovery of America by Christopher Co foreign newcomers in these floras were also cou- lumbus in 1492 boosted the perpetual interest in pled with initial attempts to make inventories of new, and partially known plant species. Exotic plant species occurring in European towns, some płants were brought to the collections of the bo- of them made as early as the beginning of the 17th tanical gardens that were emerging at that time. century (cf. J ackowiak 1990, 1993, 1998a; S ud- As the result botanical gardens were quite often nik-W ójcikowska 1987a, 1998a and references in the very spots from where alien species started those papers). The checklists of urban floras are their spread into new territories, beyond their of particular importance in studies of species of natural ranges of distribution. At the same time, foreign origin because towns are usually the places together with the introduction of new plant spe where these foreign newcomers appear for the cies to garden collections, documentation such as first time. The oldest studies of this type in Poland publications and herbaria started to emerge. include works pertaining to the Warsaw region, published by Bemhardi in 1652 and Emdtl in 1730 One of the earliest herbal studies devoted, inter (S udnik-W ójcikowska 1987a)". alia, to these alien newcomers was the 15th century work by Jan Stańko, a canon priest in Wrocław and Kraków, entitled Antibolomenum. The next cen 10 e.g. Acorus calamus — regarded by both authors as occurring near water; Ambrosia artemisiifolia - species tury, saw the publication of a work by Hieronim described by Kluk as occurring on sandy sites; Datura Spiczyński (1542) under the title O ziołach tutecz- stramonium - as early as in the times of these authors, this nych у zamorskich у o mocy ich [On herbs native plant commonly occurred in a wild state, near fences, on and coming from overseas and their effects]. yards and courts; Malva moschata - found in scrub; Information on alien species which the contem- Mercurialis annua - in orchards and grassy sites; Portu- laca oleracea - a plant cultivated in gardens, capable of porary botanical science characterised as more spreading on its own throughout garden sites. recent newcomers, or kenophytes, was included in 11 Systematic studies of urban floras started in Poland works by S irenius9 (S yreński 1613) and K luk as early as at the end of the 1901 century. These types of studies became very common in the 1970s. A review of the studies on the floras of Central European towns and the 9 For ехатріе, Sirenius mentioned Acorus calamus synthesis of the main findings are presented by J a c k o w ia k (Sweet-flag) using old Polish name: “calamus”. (1998a) and S u d n ik -W ó jc ik o w s k a (1998a). The stormy history of Poland, a country which species from Silesia (S chube 1901a, b-1930; practically went out of existence between the day Schalow 1931-1936) and Pomerania (e.g. A bro- of the abdication of King Stanislaus Augustus meit et al. 1898-1940). Rich materiał regarding the (25 N o v em b er 1795) till the day it regained its Silesian flora was summarised in a work by independence (11 N o v em b er 1918), being nothing S chube (1903b, 1904), and the Silesian flora was more but a name (D avies 2001), did not favour any reputed to be one of the best known floras in systematic collection of floristic data. Within that Europę of that time12 (S endek 1981). In Galicia period, there were only floristic studies devoted to (south-eastem Poland), an important work - but local floras (M attuschka 1776, 1777, 1779; unfortunately unfinished - in the field of floristic K rocker 1787, 1790, 1814, 1823; B esser 1809; research was Conspectus Florae Galiciae Criticus G unther et al. 1824; A damski 1828; S chneider by Z apalowicz (1906, 1908, 1911). 1837; W immer 1841; G rabowski 1843; R itschl Another important source of information was the 1850 and others). These were mostly works by naturalists’ joumals, which began to be published German naturalists and pertained to the areas as early as in the second half of the 19th century, which became included in the administrative bor- such as: Wszechświat, Pamiętnik fizjograficzny, ders of Poland after World War II. Kosmos, Sprawozdania Komisji Fizjograficznej The oldest systematic study of the flora of Po PAU, Dohrniana, Jahres-Bericht der Schlesischen land is the work by Jakub Waga, who was one of Gesellschaft fur vaterlandische Cultur. These the outstanding Polish botanists of the first half of journals published floristic notes and accounts of bo the 19th century (R ostański K. 200la). This work, tanical trips across various regions of contemporary published in 1847, includes “botanical descriptions Poland, and also included - apart from the records of plants, both wild and cultivated in open areas, of native species - new localities for many new within the Kingdom of Poland” (“botaniczne opisy alien species, coupled with their probable routes into tak dziko jako i hodowanych pod otwartem nie new territories (U nverricht 1847; R ehman 1868; bem jawnokwiatowych Królestwa Polskiego roś K rupa 1877; K amieński 1879,1884a & b; U echtritz 1879, 1880; Ł apczyński 1882, 1887, 1888, 1889, lin”). According to R ostański K. (200la), this first comprehensive study of the flora of what was then 1890; R aciborski 1884, 1885; B łoński 1892; C ybulski 1894, 1895; S chube 1901-1930; M eyer called the “Congress” Kingdom of Poland was of 1931, 1932; S chalow 1931-1936 and others). the same level and form as other floras of vascu- Further systematic floristic inventories were com- lar plants from the neighbouring areas of Prussia, pleted in many regions of Poland in the 1960s and Silesia (Polish), Galicia and Lithuania. W aga 70s. This period yielded many records and check- (1847) listed a total of more than a thousand lists contributing to local and regional floras. species of flowering plants including several An outline of the history of floristic studies as dozens of those currently classified as kenophytes well as the main currents of research, taking into - at that time these were either already established account or sometimes devoted exclusively to or merely present in cultivation (cf. Chapter 5.2). plants of foreign origin, are presented in Table 1. The development of naturalists' studies under- Particularly significant contributions were made by taken in Poland in the second half of the 19th cen those studies which concentrated on recording the tury was associated with the short-lived activities appearance of new species in local floras and gath- of Szkoła Główna (a higher education establish ering data on their stations. Articles published in ment) opened in 1862, from which some botanists a series Studies of distribution ranges of synan- graduated: K aro (1867 - Flora o f the Warszawa thropic plants by T rzcińska (1961), Ś w ieboda region, 1881 —Flora of the Częstochowa region) (1963); T rzcińska-T acik (1963); Z ając E.U. & and R ostafiński, the author of a 1872 treatise Flo- Z ając A. (1973) and G uzik & S udnik-W ójcikow- rae Polonicae Prodromus. At the same time, there ska (1989) are pioneering works on the reconstruc- were floras of the Pomeranian regions and towns tion of the history of spread by the synanthropic (e.g. K linggraeff 1848, 1854, 1866), Silesia newcomers. Much attention was also given to the (U echtritz 1865; F iek 1881) and Polish Galicia classification of plants accompanying humans and (B erdau 1859; K napp 1872). These publications to compiling checklists of species of foreign ori are a valuable source for the first record data for gin occurring in Poland (Table 1; Fig. 1). many species of kenophytes (cf. Appendix A & B). The first half of the 20th century, up until the outbreak of World War II, saw further regional 12 Silesia had its wildlife particularly well researched Floras published where authors, apart from de- even earlier, because the first study of this area was pub scribing native species, also included species of lished in the 17,h century by Caspar S chwenckfeld (1600). foreign origin. Particularly noteworthy were the More Silesian floras were published by M a t t u s c h k a (1776, 1777, 1779), K r o c k e r (1787, 1790, 1814, 1823), W im m e r works by German botanists providing information & G r a b o w s k i (1827-1829), F ie k (1881), S c h u b e (1904) on flora composition and localities for many plant and P a x (1915), after M u l a r c z y k (2000). Table 1. Selectcd papers concerning different aspects of synanthropisation and studies focussing spccifically on alien plants occurring in Poland (in chronological order)

Т у р е o f s tu d y Author/ year Historical floras (regions, cities & towns)

Congress Kingdom of Poland (Królestwo Polskie) W a g a 1 8 4 7 ; R o s t a f iń s k i 1 8 7 2

Pomerania (Pomorze) K l in g g r a e f f 1848, 1854, 1866; A b r o m e it et al. 1 8 9 8 - 1 9 4 0 ; D e c k e r 1 9 1 1 ; M u l l e r 1 9 1 1 ; H o l z f u s s 1 9 3 7 ; S t e f f e n 1 9 4 0

Silesia (Śląsk) W im m e r 1 8 4 1 ; G r a b o w s k i 1 8 4 3 ; F ie k 1 8 8 1 ; S c h u b e 1 9 0 3 b

Galicia (Galicja) B e s s e r 1 8 0 9 ; K n a p p 1 8 7 2 ; Z a p a ł o w ic z 1906, 1908, 1911

Bolesławiec town and vicinity (Bytom Odrzański, Jedlina Zdrój, S c h n e id e r 1 8 3 7 Oława & Wołów)

P o z n a ń R it s c h l 1 8 5 0

Kraków (Cracow) and surrounding area B e r d a u 1 8 5 9 ; R a c ib o r sk i 1 8 8 4 ; K r u pa 1877,1878; Ż m u d a 1 9 2 0

W arszawa (W arsaw) and surrounding area E rn d tel 1 7 3 0 ; K a r o 1 8 6 7 ; Ł a pc zy ń sk i 1 8 8 2 ; C y bu lsk i 1 8 9 4 ,1 8 9 5

Częstochowa town and surrounding area K a r o 1881

Przemyśl town and surrounding area K o t u l a 1881

Babia Góra Mt. Z a p a ł o w ic z 1 8 8 0

Tatry, Pieniny & W estern Beskidy Mts. B e r d a u 1 8 9 0 New alien plant species

Elodea canadensis K a m ie ń s k i 1 8 7 9

Acorus calamus, Amaranthus retroflexus, Chamomilla K a m ie ń s k i 1 8 8 4 a & b suaveolens, Conyza canadensis, Elodea canadensis, Galinsoga parvi/lora, Impatiens parviflora, Lycium barbarum, Xanthium spinosum

New species recordered in the W arszawa province C y b u l s k i 1 8 9 5

Rare and casual plants T r z e b iń s k i 1 9 3 0

Newcomers and wandering plants S z u l c z e w s k i 1931

Уегопіса filiformis K o r n a ś & Kuc 1953

Newcomers in the flora of Białowieża Forest S o k o ł o w s k i 1967, 1970

Corydalis lutea B e r n d t 1 9 5 8

Achillea crithmifolia D ą b r o w s k a 1 9 7 2

Bromus carinatus M ir e k 1982 (1984)

Уегопіса peregrina Z a ją c M . & Z a ją c A . 1 9 9 0

Eragrostis multicaulis G u z ik & S u d n ik -W ó jc ik o w s k a 1 9 9 4

Chaerophyllum aureum O k l e je w ic z 1 9 9 9

Reynoułria x bohemica F o jc ik & T o k a r s k a -G u z ik 2 0 0 0 For more see also appendix A a n d В First localities

Lists of plant species together with their localities K o r n a ś 1950, 1954; U rb a ń sk i 1 9 5 8 ; Ż u k o w sk i 1959; 1960a& b; T a c ik 1 9 6 0 ; F a b is z e w s k i & F a l iń s k i 1 9 6 3 ; S o w a & W ó jc ik - -C h r o b o k 1 9 6 9 ; R o s ta ń sk i K. 1960,1961; S c h w a r z 1 9 6 1 ; S ow a 1 9 6 2 ; H a n t z 1967,1972; M ic h a l a k 1968, 1971;K o r n ia k 1 9 6 8 ; T r z c iń s k a -T a c ik 1 9 7 1 a ; M ic h a l a k & S e n d e k 1974-1975;

G ł o w a c k i 1 9 7 5 ; W ik a 1 9 7 5 ; O l e s iń sk i & K o r n ia k 1 9 8 0

Kenophytes in the flora of Lublin province F ija ł k o w s k i 1 9 7 3

Synanthropic grasses K o r n ia k 2 0 0 2 Synanthropic floras

Floras o f cities: P o z n a ń K r a w ie c o w a 1951 G d a ń s k S c h w a r z 1 9 6 7 S z c z e c in Ć w ik l iń s k i 1 9 7 0 Zielona Góra, Koszalin Ć w ik l iń s k i 1971 Ł ó d ź S o w a 1 9 7 4 K r a k ó w T r z c iń s k a -T a c ik 1 9 7 9

Comparison of the urban floras on the example of some cities K r a w ie c o w a & R o s t a ń s k i 1976, 1981 Туре of study Author/ year

Floras of towns & settlements Mowszowicz 1 9 6 0 ; S k o w r o ń s k a 1 9 6 5 ; M ic h a l a k 1 9 7 0 ; S c h w a r z 1 9 7 1 ; S e n d e k 1 9 7 1 ; S o w a 1 9 7 1 ; A n io l -K w ia tk o w sk a 1 9 7 4 ; H a n t z 1 9 7 4 ; S z m a jd a 1 9 7 4 ; C z a pl e w sk a 1 9 7 5 ; M isiew ic z 1 9 7 8 ; S e n d e k & W ik a 1 9 7 9 ; S o w a & N a s ił o w s k i 1 9 7 8 ; W e r e t e l n ik 1 9 7 9 ; S o w a & W archolińska 1 9 8 0 ; M is ie w ic z 1 9 8 1 ; S ow a & W archolińska 198la, b & c, 1984a & b, 1987; M a c ie jc z a k 1 9 8 8 ; Ć w ik l iń s k i & B a r t n ik 1 9 9 0 ; T o k a r s k a - -G u z ik & R o s t a ń s k i 1997, 1998

Fuli cartographic description of the urban flora: W a r s z a w a S u d n ik -W ó jc ik o w s k a 1 9 8 7 a P o z n a ń J a c k o w ia k 1990, 1993 J a w o r z n o T o k a r s k a -G u z ik 1 9 9 9

Ruderal floras in the rural landscape of the North Podlasie W o l k o w y c k i 1 9 9 7 Lowlands Regions - examples: Wielkopolska province S z u l c z e w s k i 1951 Gorce Mts. K o r n a ś 1957, 1966 Tatry Mts. R a d w a ń sk a -P a r y sk a 1 9 6 3 ; P ię k o ś -M ir k o w a & M ir e k 1 9 7 8 Wielkopolski National Park Szulczewski 1963; Żukowski et al. 1 9 9 5 Karkonoski National Park R o st a ń sk i K. 1977, 1978 Lublin province F ija ł k o w s k i 1 9 7 8 Upper Silesia Industrial Region S e n d e k 1981, 1984 eastern part of the Gniezno Lake District C h m ie l 1 9 9 3 Zaodrze (to the West of Szczecin) Z a j ą c A . et al. 1 9 9 3 Słowiński National Park Piotrowska et al. 1 9 9 7

Segetal flora W nuk 1976; Sowa & W archolińska 1979; W archolińska 1981, 1996; W nuk et al. 1989; Siciński 1997, 2000; Latowski 1998, 1999; Trzcińska-Tacik 1996; W archolińska & Siciński 1996; W archolińska & Tyszkowska 2000 also Jackowiak & Latowski 1996, 2001; Misiewicz & Piotrowski (eds.) 1996; R ola 1996 and literature cited therein Ruderal plant communities

R e g io n s K o r n a ś 1 9 5 2 ; S ow a 1971

Cities & towns F ija ł k o w s k i 1963, 1967; R o s t a ń s k i K . & G u t t e 1 9 7 1 ; A n io ł - -K w ia tk o w sk a 1 9 7 4 ; K ępc zy ń sk i & Z ie n k ie w ic z 1 9 7 4 ; Z a ją c E.U. 1 9 7 4 ; K ę p c z y ń s k i 1 9 7 5 ; C z a p l e w s k a 1 9 8 0 ; Ś w ięs & P l eb a n 1 9 8 1 ; Ś w ięs 1 9 8 3

Special habitats C z a p l e w s k a 198 1 Alien plants in special habitats

Railways and railway stations M eyer 1931, 1932; K ornaś et al. 1959; Sowa 1966; Ćwikliński 1968, 1972a, 1974; Krawiecowa 1968a; Sendek 1969, 1973; Zając E.U. & Zając A. 1969; Latowski 1977; Ćwikliński 1984— 1985; W ika 1984

Storę yards (including ballast plants) Helm 1881; H olzfuss 1936, 1941

Sea & river harbors R o st a ń sk i K. & S z o t k o w s k i 1 9 7 3 ; M is ie w ic z 1976,1985,2001

Walls W e r e t e l n ik 1973, 1982; Ś w ie r k o s z 1 9 9 3 ; G a l e r a & S u d n ik - -W ó jc ik o w s k a 2 0 0 0 a & b Lists of alien plant species

Kenophytes K ornaś 1968b; Zając A. et al. 1 9 9 8

Archaeophytes Z a j ą c A. 1979, 1983, 1987a& b, 1988

Ephemerophytes R o s t a ń s k i K . & S o w a 1 9 8 6 - 1 9 8 7

American trees and shrubs H e r e ź n ia k 1 9 9 2

Kenophytes of American origin S ow a & W archolińska 1 9 9 4

Anthropophytes M i r e k et al. 1995, 2002

Naturalised alien plants - neophytes (excluding archaeophytes) T o k a r s k a -G u z ik 2 0 0 3 a Terminology & classification

Classification of synanthropic plants K o r n a ś 1968а, 1977a & b; K r a w ie c o w a & R o s t a ń s k i 1 9 7 2 ; M ir e k 1 9 8 l a

Neophytes & neophytism F a l iń s k i 1968a & b, 1969

3 The Establishment. 17 Туре of study Author/ year

Dictionary of synanthropisation of plant cover S u d n ik -W ó jc ik o w s k a & K o ź n ie w s k a 1 9 8 8 Origin, history of expansion & the distribution of alien plants

Archaeophytes Z a ją c A . 1 9 7 9

Bidens melanocarpus (= B. frondosa) T r z c iń s k a 196 1

Elsholtzia ciliata (= E. patrini) S w ie b o d a 1 9 6 2

Rumex confertus T r z c iń s k a -T a c ik 1 9 6 3

Artemisia Ż u k o w s k i & P ia s z c z y k 1 9 7 1

Salsola B a r a d z ie j 1 9 7 2

Trifolium patens L o s t e r 1 9 7 2

Mimulus P ięk o ś 1 9 7 2

Corydalis lutea, Cymbalaria muralis, Impatiens glandulifera Z a ją c E .U . & Z a ją c A . 1 9 7 3

Amaranthus F re y 1 9 7 4

Oxalis H a n t z 1 9 7 9

Iva xanthiifolia G u z ik & S u d n ik -W ó jc ik o w s k a 1 9 8 9

Eragrostis pilosa S u d n ik -W ó jc ik o w s k a & G u z ik 1 9 9 6

Oenothera R osta ń sk i K . & T o k a r s k a -G uz ik 1 9 9 8 and literature cited therein

Beckmannia eruciformis F r ey & P a s z k o 2 0 0 0

Veronica peregrina Z a ją c M. & Z a ją c A. 1 9 9 0 ; G u z ik & P a u l 2 0 0 0

Alien grass species in the Silesian Upland T o k a r s k a -G u z ik & N o w a k 2 0 0 1 Threats to protected naturę by alien plant species

Anthropogenic plant communities in Białowieża Forest F a l iń s k i 1 9 6 6 a

Alien plant species in natural com m unities K o r n a ś & M e d w e c k a -K o r n a ś 1 9 6 8

Contribution of alien plant species in the flora of Opawskie K r a w ie c o w a 1 9 6 8 b M o u n ta i n s

Anthropogenic changes in plant cover of Ojców Landscape Park M ic h a l ik 1972, 1974

The naturę of the Pieniny M ts. in face of the com ing changes Z a r z y c k i 1 9 8 2

W eed species from Śnieżnik M assif, the Bialskie and the Złote B rej 2 0 0 1 M ts . General and theoretical aspects

F a l iń s k i 1966b, 1968a, 1969, 1971, 1972, 1998a & b, 2000a; K o r n a ś 1971, 1977b, 1981, 1982, 1983, 1990, 1996; O l a c z e k 1972, 1974, 1982; K r a w ie c o w a & R o s t a ń s k i 1 9 7 6 ; S o w a & O l a c z e k 1 9 7 8 ; T r o ja n 1 9 8 2 ; J a c k o w ia k 1991, 1998a & b, 1999, 2000, 2003; S u d n ik -W ó jc ik o w s k a 1991, 1992, 1998a & b, 2000; S u d n ik -W ó jc ik o w s k a & M o r a c z e w s k i 1 9 9 8 ; T o k a r s k a - G u z ik 2 0 0 l a ; K o r n a ś & M e d w e c k a -K o r n a ś 2 0 0 2 Distribution atlases

Distribution Atlas of Vascular Plants in Cracow Province Z a ją c M. & Z a ją c A. (eds.) 1998

Geobotanical Atlas of the Bug River Valley F a l iń s k i et al. 2 0 0 0

Atlas of distribution of vascular plants in Poland Z a ją c A . & Z a ją c M. (eds.) 2001

Atlas of alien woody species of the Białowieża Primaeval Forest A d a m o w s k i et al. 2 0 0 2

The same broadening of the scope of studies of alien species occurred on a mass scalę and pertaining to alien plants was developed by endangered native vegetation cover (Australia, botanists in other parts of Europę and the New Zealand, South Africa, the western coast wider world. The topics and scope of more than of the United States, Hawaii). The number of 900 papers indexed in Ecological Abstracts studies on these topics is still increasing (up to (1974-1993) were analysed by P yśek (1995). He some 100 publications each year). found that after a period of collecting floristic In recent decades, as a part of Poland’s com- records about the occurrence of species of mitment to international programmes, a certain foreign origin, there was later an evident shift number of studies undertaken in this country of emphasis to the issues of their biology and have concentrated on biological diversity. ecology, as well as a drive to more generał Nevertheless, the topics which are particularly (theoretical) papers. current, as well as those pertaining to the Intensive studies were carried out, particularly recognition of the threats to native biodiversity in those parts of the world where the appearance from invasive or genetically modified plant biology taxonomy and ecology

management

alien plants theoretical N and methodological v studies y lists of invasive p lants

distribution

Fig. 1. Scope of studies pertaining to alien plants developed by botanists in Poland The graph shows the situation referring to the period 1950-2000. Contribution of particular topics and scope of studies in the total number of papers analysed (n = 1074) is indicated in dark grey (significant), light grey (intermediate) and white (Iow or none) species, still represent only a smali proportion Also praiseworthy are studies of special topics of the overall number of studies pursued undertaken by Polish botanists, which have already (W iśniewski 2 0 0 3 ). claimed their place in the overall achievements of biogeographical sciences, such as: - monograph devoted to phytogeographical problems of vegetation in the Gorce Mts. 2.2. Synanthropisation: the essence (K ornaś 1955); of the process and the role - model monograph pertained to anthropogenic of kenophytes in the changes transformations of vegetation in the Białowieża occurring in the natural Primaeval Forest (F aliński 1966a); environment on Earth - pioneering attempts at comparative analysis of floras of towns and settlements (F aliński 1971; K rawiecowa & R ostański 1976); Studies of alien plant species fali into the cur- - methodological studies concerning the spatial rent generał field of research concerning anthro- structure of the flora of major cities (J acko pogenic changes in vegetation. As early as in the wiak 1998a& b; S udnik-W ójcikowska 1998a), 1960s, the discussion was initiated within regular symposia, devoted to various aspects of the including especially the model solution proposed by J ackowiak (1998a & b), who defines synanthropisation13 of vegetation cover (F aliń- the city as a centre of crystallisation in a flo- ski et al. 1998; T okarska-G uzik 200la) (Table 2). ristic-ecological space. Also works by S udnik- W ójcikowska (1998a & b, 2000), confirming 13 A definition of the term “synanthropisation” was pro- the indicative role of the flora with respect to posed by F a l iń sk i (1966b, 1972): “Synanthropisation of vegetation is a part of directional changes occurring on Earth the thermal conditions in an urban area; under the impact of human activities, manifesting themselves - comparisons of the differences among rural flo as replacing specific i.e. endemic components, with non- ras in special areas treated as “environmental specific i.e. cosmopolitan, replacing native i.e. autochtonie islands” such as Mediaeval strongholds (C el- components with newcomers i.e. allochtonic elements, re ka 1999), settlements in agricultural-forest placing stenotopic components by eurytopic ones. In effect, it means replacing primary systems, conditioned by the joint landscapes (W ołkow ycki 2000), and aban- effect of endogenic and exogenic factors, with secondary doned industrial sites and areas (R ostański A. systems conditioned mainly by exogenic factors”. 1998a & b; W oźniak 1998; C ohn et al. 2001); Table 2. Polish symposia on the synanthropisation of plant which do not use the term “synanthropisation”, cover deal essentially with the issues covered by the Conference title Place Year scope of the term: decreasing the diversity of Synanthropisation of plant cover Kraków 1965 naturę and invasion by alien species. These issues Neophytism and apophytism of plant are currently included in the study of the overall cover in Poland Nowogród 1968 changes occurring on Earth and are coupled with Synanthropic flora and vegetation of calls to protect biological diversity (K ornaś & towns connected with their natural M edw ecka-K ornaś 2002). The interest in bio conditions, history and function Wrocław 1970 logical invasions has been boosted recently because Theoretical and methodical basis of the of the threat to native vegetation, but also because studies upon the synanthropisation of of the inereasing likelihood of transgenic organ- the plant cover Białowieża 1971 isms penetrating natural communities in the wake Synanthropisation of plant cover in of developments in genetic engineering (D aehler national parks and naturę reserves Białowieża 1971 & C arin o 2000; Z arzy ck i 2000a; Cronk & Fuller Phytocoenosis degeneration under the 2001). influence of natural and anthropogenic factors Łowicz 1974 Invasion by plants of foreign origin is considered, along with the fragmentation and degradation of Decline and extinctions of the native plant species in Poland Kraków 1976 natural communities, to be one of the leading General problems of synanthropisation Białowieża 1980 threats to global-scale biodiversity (Abbott 1992; Kolar & Lodge 2001). Significant expenditure, American plant species established in Poland Łódź 1992 borne in attempts to control invasive species and results of their invasions, prompted the Scientific Mechanisms of anthropogenic changes of the plant cover Poznań 1999 Committee on Problems of Environment (SCOPE) to initiate a special research programme, called Phytogeographical problems of synanthropic plants Kraków 2000 Ecology of Biological Invasion (1982), which was Invasive species in the flora and fauna then continued under the framework of the Glo of Poland against the background of bal Strategy of Invasive Species project (initiated the conservation of biological diversity Kraków 2001 in 1995). The effect of the SCOPE 37 programme includes a series of book publications covering the S o u rc e s : F a l iń s k i, A d a m o w s k i & J a c k o w ia k (eds.) 1998; Ł a w r y n o w ic z & W archolińska (eds.) 1992; J a c k o w ia k & results of studies devoted to these issues (Groves Ż u k o w s k i (eds.) 2000; Z a ją c A ., Z a ją c M. & Z e m a n e k (eds.) & Burdon 1986; MacDonald et al. 1986; Mooney 2003. & Drakę 1986; Drakę et al. 1989; Di Castri et al. 1990; Mooney & Hobbs 2000). These books - theoretical concepts (models) to interpret the provide enormous lists of references from all over phenomena of ecological and geographical the world. The inereased interest in the issue of bio expansion (Jackowiak 1999; F aliński 2000a, logical invasion has brought about the develop- 2004) (cf. also Table 1). ment of other biological research programmes as Since the end of the 20lh century, the phenom- well as the emergence of specialised intemational enon of synanthropisation, associated with human organisations and research groups. These include population growth, advances of technology, de- GISP - Global Invasive Species Programme and velopment of agriculture, industry and urban ISSG - Invasive Species Specialists Group (opera- centres, has been attracting ever-increasing in- ting under the aegis of the Intemational Congress terest (K ornaś & M edw ecka-K ornaś 2002). It of Naturę Conservation of IUCN), which published has resulted in a number of detailed studies and a list of the most “dangerous” invasive species and reviews, pertaining to many aspects of this issue, a guide to “management” of invasive species important in various regions of the world14. Many (M irek & W ołoszyn 2001). Monographic works authors highlight the fact that numerous papers focusing on various features of biological invasions were published and the specialist journal Biolo 14 The increase in the interest in issues of synanthropi gical Invasions was launched. A number of national sation was particularly great in Western and Central Eu and intemational conferences and seminars have ropę, as a result of the remarkable devastation of vegeta- been held, such as the Slovak conference Invazie tion in these parts of the continent. The transformation of vegetation resulting from human activities was taken up as a invazne organizmy (Eliaś 1997), a conference topie of many scientific conferences and constituted the devoted to Alien Organisms in Germany (D o y le subject of numerous monographs, reviews and theoretical 1999) or the conference organised in 2001 in works (e.g. Thellung 1918-1919; Probst 1 9 4 9 ; S u k o p p Kraków, on the Invasive species in the flora of 1 9 6 2 ; S u k o p p & T r a u t m a n 1 9 7 6 ; K o r n a ś 1 9 8 2 ; O l a c z e k Poland in the context of the protection of biolo 1 9 8 2 ; H o l z n e r et al. 1 9 8 3 ; W it t ig et al. 1 9 8 5 ; J a g e r 1 9 8 8 ; by the Natural Conservation K o w a r ik 1988, 1990; L o h m e y e r & S u k o p p 1 9 9 2 ; P y Sek gical diversity, 1 9 9 3 ; J e h l ik 1 9 9 8 ; F a l iń s k i et al. 1 9 9 8 and others). Committee of the Polish Academy of Sciences. Biologists and ecologists from Germany, at developed into a separate channel of research, a meeting in Berlin in April 1999, founded a also using data from other disciplines of natural research consortium on biological invasions. This sciences (R ejm Anek 1996; D aehler 2001). In group co-ordinates responses to the ever increasing recent years, studies of alien species have dealt problems eaused by the invasion of non-native with the various threats posed to natural vegeta- plants, animals, fungi and micro-organisms. These tion by invasion by alien species (numerous basie “new species” (Neobiota) can threaten the bio- studies devoted to the taxonomy, biology and diversity within existing native species, alter the ecology of alien species as well as to the mech- structure and function of ecosystems and can even- anisms of invasion) and with the methods and tually cause severe economic and human health techniąues to control the spread of invasive spe problems. The Neobiota group initiates and organ- cies (cartographic studies of distribution ranges, izes conferences (e.g. 3rd International Confer- monitoring, “management” and other methods to ence on Biological Invasion Neobiota - From control these species). From among the volumi- Ecology to Control, Bern 2004) and related pub- nous list of papers, the most illuminating are those that attempt to show model descriptions of the lications (K owarik & S tarfinger 2002; S eitz & phenomenon of invasion (S ukopp & S ukopp 1993, K owarik 2003; K uhn & K lotz 2004). An international event of major importance in 1994; F aliński 1998a & c; Jackowiak 1999; L ons- the field is a conference held regularly under the dale 1999), papers devoted to forecasts of invasions (K olar & L odge 2001; P yśek 2001) as well title of Ecology and Management of Alien Plant as those dealing with evolutionary processes re- Invasions, devoted to broadly defined issues of sulting from invasion by alien species (D en Nuss biological invasions (W aal de et al. 1994; Py- et al. 1999; E llstrand & S chierenbeck 2000; śek et al. 1995; B rock et al. 1997; S tarfinger et Z ając A. & Z ając M. 2000; A llendorf et al. 2001). al. 1998; B rundu et al. 2001; C hild et al. 2003). An important contribution to our knowledge of The International Union for the Conservation of invasion has also been made by lists of alien spe Naturę IUCN in February 2001 published Guide- cies and by synthetic studies pertaining to parti- lines for the Prevention o f Biodiversity Loss Caused cular regions, which commonly also provide rich by Alien Invasive Species, focusing the interest of collections of sources and references (e.g. C le- researchers on the development of studies to enable ment & F oster 1994; G udżinskas 1997a, b, с & d, the slowing down or containment of the adverse 1998a, b & с, 1999a & b, 2000a & b; P reston et effects of the invasion by alien species. In the same al. 2002; P yśek et al. 2002; K uhn & K lotz 2003; year under the auspices of Global Invasive Species B otond & B otta-D ukAt 2004). Another easily Programme Global Strategy on Invasive Alien Spe accessible and fast source of information is pro- cies was developed (M cN eely et al. 2001). vided by many websites and home pages present- The role of science is critical in providing the ing both scientific papers and applied research information needed to develop a coordinated Eu- studies, often with maps of growing secondary dis ropean policy (G enovesi 2004). In order to tribution areas and photographs familiarizing respond to these needs a European Strategy on readers with “the perpetrator” and the scalę of the Invasive Alien Species (G enovesi & S hine 2004) phenomena caused by it. has been approved by the Bern convention and However, in spite of a growing body of infor supported by the European Council of Ministers mation accumulated in the last half-century on the (G enovesi 2004). spreading of alien plant species in various cor- As a result of the developments summarised ners of the Earth, many ąuestions have remained above, the issue of invasion by alien species has unanswered.

PART TWO

Terminology and methodology

3. Phytogeographical terminology new terms (T okarska-G uzik 2 0 0 la; P yśek et al. and the classification of synanthropic 2 004). The classifications of synanthropic floras plants used in Poland by various authors differ above all in the criteria adopted as well as in the scope and interpretation of the terms used16. One of the essential aspects of studies devoted In discussions by phytogeographers who study to species of foreign origin is the problem of their the topie of invasiveness, the terminological ques- status within a given flora. tions are regularly addressed, not only for purely The first attempts to provide a typology of semantic reasons, but also for practical purposes species of foreign origin date back to the 19th in order to make a comparative approach possible cen tu ry (C andolle D e 1855; A scherson 1883). (P yśek 1995; Rjchardson et al. 2000; T okarska- The concept of the classification of floras and its -G uzik 2 0 0 la; C hmura & S ierka 2 0 0 4 ; P yśek terminology as adopted in Central Europę was et al. 2004). elaborated by T hellung (1918-1919). This author A comparison of the classification of synan discussed and defined tenns such as “native”, thropic species accepted in Polish literature with “introduced” and “alien” in French, German and those in English-language publications allows the E n g lish (S ukopp 1998). The classification of group of species included in the present study to synanthropic flora proposed by Thellung was be correctly placed within the different systems applied in Poland by many authors15, and modified currently applied (Appendix D). by K ornaś (1968a, 1981), adopting the following The authors of one of the recent publications basie criteria: origin, time of arrival and the aiming at introducing a certain order to the degree to which a particular species is established “invasive” terminology (in particularly devoted to (Fig. 2). According to K ornaś (1981), “plants of invasive plant species) suggested yet another, foreign origin (alien plants) are those species simplified classification in which the status of originating from areas other than that in which a species is determined on the basis of major they are found, which have appeared in new barriers it has to overcome in the process of habitats owing to intentional or unintentional settling in a new territory (Rjchardson et al. 2000; introduction as a result of human activity”. P yśek et al. 2 0 0 4 )17 (cf. also Chapter 12). The In the Polish scientific literature in this field, the proposed classification considers practical implica- first attempt to gather and organize the existing tions connected with the spread of non-native (alien) terms and classifications of synanthropic plants is species beyond their natural ranges, their naturali- the work entitled Słownik z zakresu synantropiza sation in new homelands and the effects on cji szaty roślinnej [Dictionary of tenns used in the naturę and human economic activities. The au field of the synanthropisation of vegetation cover] thors of the above-mentioned papers do not (S udnik-W ójcikowska & K oźniewska 1988). In many current English-language publications, criticism has been directed towards East European 16 Terminologies and definitions in this field of research authors, particularly for introducing a multitude of were compared in a large body of literature by P y ś e k (1995) and in Polish literature by S u d n ik -W ó j c ik o w s k a & K o ź n ie w s k a (1988). 15 Thellung’s classification was first used in Poland by 17 Besides the classification and terminology associated K r a w ie c o w a ( 1 9 5 1 ) in her pioneering work on synanthropic with definitions the authors give also the synonyms for par flora of Poznań. ticular terms. Criteria for Group of species classification

Apophytes Anthropophytes native species alien plant species Origin occurring in man-made habitats Diaphytes naturalisation Metaphytes permanently not permanently established status established /settled Ephemerophytes casual alien plants Ergasiophygophytes kept in cultivation and occasionally escaping

Archaeophytes Kenophytes time of arrival introduced before 1500 introduced after 1500

Epecophytes established in man-made habitats type of plant community Hemiagriophytes invaded penetrating into semi-natural habitats Holoagriophytes penetrating into natural habitats

Fig. 2. Position of kenophytes in the geographical-historical classification of the synanthropic flora ( K o r n a ś 1968a, 1981 after T h e l l u n g 1918/1919; Trzcińska-Tacik 1979) consider the criterion of time (time of immigra- epecophytes), and sometimes penetrating into tion) which although artificial still allows one to semi-natural communities (—> hemiagriophytes) differentiate between processes in the floras or natural communities (—» holoagriophytes) (S u d - which in the Middle Ages went differently com- nik-W ójciko wska & K oźniew ska 1988) (cf. Fig. 2; pared with outcomes in modern times. to compare terminology see also Appendix D).

The species included in this study were selected from two sources: 4. Materials and methods - Kenophytes in the flora of Poland: list, status and origin (Z ając A. et al. 1998); - Flowering Plants and Pteridophytes of Poland, 4.1. Selection of species18 and their a checklist (M irek et al. 2002). status The list elaborated on the basis of these two references required changes and supplements, because the original lists of anthropophytes and Adhering to Thellung’s classification of synan kenophytes were somewhat outdated. For the thropic plants as modified by K o r n a ś (1968a), purpose of the present study, it was therefore this monograph pertains to kenophytes, i.e. spe necessary to create an original and up-to-date cies alien to the natural flora of a given region catalogue of kenophytes occurring in Poland. This (in this case, of Poland), which arrived after the was developed on the basis of regional studies and year 1500 and are now permanently established personal research data (Appendices A and B). (—> metaphytes): in anthropogenic habitats (—» The status of each alien species occurring in Poland has been critically assessed against the 18 The taxa covered in this study include units of var- available historical floras and modern studies ious rank: species, subspecies and hybrid forms (cf. Ap- devoted to the issue of synanthropisation. The pendices A and B). In the text, the whole group of taxa under study are termed “species”, when referred to collec- analysis has also utilised the publications by the tively. following authors: K o r n a ś (1968a & b, 1981), M irek (1981а), R ostański & S owa (1986-1987), included in the geographical, historical and eco- Z ając A. (1979) and Z ając A. et al. (1998). logical analyses of the Polish kenophyte flora In some cases, however, the practical applica- (Chapter 5). This Appendix includes a further tion of the criteria adopted by these authors poses 51 species of which are likely kenophytes but considerable difficulties. This pertains both to the whose status is still under discussion. These species which, according to the present level of species are mostly plants cultivated (planted) in knowledge about their origin, cannot be validly a certain way (mostly tree species), which classified as either native or alien to the flora of manifest a tendency to become “wild” and are Poland19, and to newcomers towards which certain considered established in some regions of Po doubts still exist as to the timing of their arrival land, but are still of uncertain status. Whenever and the degree of their establishment. this group is considered in the analyses, an A species is included in the presented list when appropriate note is made. certain premises have been met: - the species is alien throughout the whole of Poland (if it has even one station in Poland which is deemed to be natural, the species is 4.2. Sources and characteristics not considered as alien); of the floristic data used - hybrids produced by “crossing” a native species with a species of alien origin are treated as alien In this monograph both the author’s own taxa and henceforth included in the list20. records and those obtained by other researchers The list presented does not include the follow- have been used, classified into three groups: ing species of the genus Oenothera: Oe. ammo- - unpublished, phila Focke, Oe. biennis L. s.str. and Oe. rtibri- - published, caulis Kleb., following the opinion of R ostański K. - herbarium records. (1998, 2003), that these have been known in The most significant and voluminous informa Poland (and in Europę) for a long time and they tion on stations has been provided in unpublished have not yet been found in North America. materials sent to the database of the Distribution The taxa which had been previously classified Atlas of Vascular Plants in Poland - ATPOL as kenophytes but have had their status changed (Z ając A. & Z ając M . 2001), by botanists from in the most recent studies were also taken off the all over Poland, and the records collected by the list. These are: Malva alcea L., a species which, according to the newest research should be represent author during floristic studies. In this monograph, the materiał collected by the author garded as an archaeophyte (C elka 1998) and Ver- bascum chaixii Vill. subsp. orientale Hayek consists of floristic records (a total of 4 594 which had previously been included in the list of records), gathered in the course of field studies over a period of more than 10 years, and particu- kenophytes (Z ając A. et al. 1998), but has more recently been classified with the species which larly within the period 1996-2003. The herbarium materials collected during these are not yet established (M irek et al. 2002). The complete list of the species studied is pro- studies have been deposited in the Herbarium of vided in alphabetic order in the concluding part the Department of Plant Systematics of the Sile- of this monograph. The list of kenophytes has sian University (KTU). been divided into two major groups: The records from published sources have been - Appendix A - kenophytes about which the obtained from nearly 1 000 floristic and phytoso- most exhaustive information has been gathered, ciological publications from the last 200 years. including the data on their distribution. This Historical accounts were particularly important Appendix includes 174 species; for the task of reconstructing changes in the dis - Appendix В - kenophytes for which sufficient tribution of individual kenophytes. Available information on distribution has not yet been works by Polish botanists, also by botanists from gathered (75 species); these species have been neighbouring countries undertaking research during the 19th and the beginning of the 20th 19 In the newest edition of the critical checklist of vas- century within the lands of contemporary Poland, cular plants of Poland: Flowering Plants and Pteridophytes were used for this purpose. of Poland - a checklist (M ir e k et al. 2 0 0 2 ) , such species Herbarium collections - both Polish and in have been separately treated as taxon of uncertain status neighbouring countries (Herbaria in Berlin, Prague in the Polish flora, likely to be anthropophytes. and Vienna) - provided some 6% of records on 20 The same treatment has been applied to locally emerg- the occurrence of kenophytes in Poland. Also, in ing new taxa of the genus Oenothera, which are hybrids between species which originated from North America and this case those data which help locate or verify the species which R o s t a ń s k i K. (1998, 2003) regards as the earliest records of particular species were native. particularly important.

4 The Establishment. 25 The ATPOL database has over 66 000 records Pteridophytes o f Poland, a checklist (M irek et al. pertaining to kenophytes. A single record contains 2002), including also the most freąuently used information on one species or a group of species. synonyms. The names of the relevant families are The predominating majority of records comes from provided for all species. unpublished sources (ca. 69%). Published data 2. Biology and ecology constitute ca. 25% of the overall number of records The life form of each species was determined reflecting the enormous volume of modem record- on the basis of the R a u n k ia er system (1905). Out ing compared with herbarium specimens. of more than a dozen, only the basie forms: pha- In the case to analyse the oldest printed sources, nerophytes, chamaephytes, hemieryptophytes, primarily published in Latin, and of older German geophytes, hydrophytes and therophytes were se- (often printed in Gothic type), and Russian sources, lected for further analyses. suitable reference dictionaries (e.g. R ospond 1951) The remaining data on the biology of a species, were consulted in order to translate the geographic such as manner of reproduction, pollination of names and descriptions of sites, and also quite often flowers, dispersal of diaspores and life strategies it was necessary to locate old maps. were compiled from available sources (e.g. T utin The majority of records collected in the data et al. 1964-1986; G r im e 1977, 1979; F r a n k & base are from the last century (over 90%), whilst K lotz 1990) and the author’s own observations. the data collected in the 19th century constitutes some 5%, while bnly 0.2% of records date back 3. Origin, history of expansion and current status to the 18th century. The information about the homelands of indi- In evaluating the ąuality of data available at the vidual species and the time of their introduction start of this research project it should be noted that into Europę, either accidental or for cultivation, only 10-15% of the territory of Poland had then was taken from the literature (the list of refer- been studied more thoroughly (i.e. with a somewhat ences used is provided in the notes explaining the greater number of records per cartogramme unit) abbreviations and symbols used in Appendices A than the remnant part of the country. Thus, it was and B). For each species, the information about necessary to supplement the data, particularly by its first record in Europę was collected (for species examining the oldest records which were then used of European origin this is the first record outside to reconstruct the histories of the spread of various its natural distribution rangę). The information species. Another pivotal element which had to be on the first record in Poland is more detailed, decided was the evaluation of the status of a given indicating the location of the first station and the species at a particular station (i.e. planted or spon- source of the data. taneously) which permits the reconstruction of the For 174 species of kenophytes for which suf- stages of its establishment in the flora of Poland. ficiently comprehensive data have been collected, It must nevertheless be emphasised, that the the numbers of stations are given separately for records of the last 100 years were used as the consecutive periods of time (from 1700 to 1850, primary basis for the interpretation, as being the then 1851-1900, 1901-1950, and 1951-2003), most reliable and comprehensive; it is also con- also the total number of ATPOL sąuares where the sidered that they also allow for a proper assess- species has ever been recorded (Appendix A). ment of the dynamie tendencies in the flora of The dynamie tendencies of species were as- kenophytes occurring in Poland. sessed using the criteria suggested by Z a r zy c k i et al. (2002), but in addition related to the author’s own data on the number of stations analysed in the consecutive 50-year periods and the 4.3. List of kenophytes and the scope number of ATPOL sąuares where the species has of the information collected in been recorded. order to characterise them Based on the number of ATPOL sąuares, it was possible to establish the categories of freąuency in relation to the overall number of sąuares for The alphabetic list of species with their biolo- Poland (n = 3646), i.e. categories 1 to 6 repre- gical, geographical and historical characteristics sent species recorded in the following numbers was compiled in an Excel table and attached to of sąuares the main text as Appendices A and B. The array 1. 0.02 - 1.0% of sąuares gives the following elements of information for 2. 1.1 - 10% of sąuares each species in the order listed below: 3. 10.1 - 20% of sąuares 1. Taxonomy and nomenclature 4. 20.1 - 40% of sąuares The names of species and taxa of hybrid ori- 5. 40.1 - 60% of sąuares gin are adopted from Flowering Plants and 6. 60.1 - 100% of sąuares The number of registered stations were then graph (T o k a rsk a - gu zik 200 lb) and 18 maps in used to set up a detailed scalę of freąuencies co-operation with other authors (C ia ciu ra et al. (adapted to that applied by Z arzycki et al. 2002), 200lb; C za r n a et al. 2001; R o sta ń sk i K. & namely: T o k a r sk a -G u zik 2001). The maps prepared for 1-14 stations -rare the remaining species also include inputs from the 15 - 50 stations -occasional author of this monograph. 51 - 500 stations - occasional, locally frequent The set of distribution maps of Polish keno 501 - 6000 stations -freąuent, locally abundant phytes has been supplemented by five more ori > 6000 stations - abundant (common). ginal maps, prepared for the following species: The current status of the species was deter- Ailanthus altissima (Mili.) Swingle (Chapter 7, mined by listing the habitats which the species Fig. 39), Asclepias syriaca L., Medicago x varia colonises in the area of Poland. Martyn, Sicyos angulata L. and Sisyrinchium ber- The characteristics of the species was also sup- mudiana L. em. Farw. (Appendix C). plemented by information on its invasiveness in In the case of species which were brought to other regions of the world (based on literature data: Poland intentionally as useful plants, some of F ernald 1950; H olm et al. 1979; P errins et al. which are still under cultivation, there have been 1993; S h ev era 1997; C elesti G rapow & B łasi some difficulties in developing maps. Because 1998; J e h l Ik 1998; K o w a rik 1999; L a n d o ld t of the method of collecting information for the 2000; C elesti G ra po w et al. 2001; C ro n k & ATPOL database (the status of a given species at F u ller 2001; F edo rov 2001; U h er ć ik o v A 2001; particular stations was not recorded), and the P yśek et al. 2002); also cited are the most signi- variable descriptions of data in published records ficant published sources containing distribution or herbarium data (in many cases the authors of maps. records did not provide this information), it was impossible to differentiate between symbols on the cartogrammes or to select stations where the given species had appeared spontaneously. It 4.4. Cartogrammes and their analysis should thus be kept in mind that for some species under cultivation, the relevant cartogramme can The cartogrammes were prepared in accordance include both stations at which the species was de- with the reąuirements of the Distribution Atlas liberately introduced and those where the species o f Yascular Plants in Poland - ATPOL, adopting entered unaided. For the same reason (without a 10 x 10 km square as the basie unit. The verifying the data in the field studies), some tree territory of Poland is thus placed in 3 646 such species were excluded from the cartographic part cartogramme fields (taking into account also parts of the study, e.g. Aesculus hippocastanum and of units along the national borders) (Z a ją c A. Quercus rubra, of which it is known that they 1978a & b; Z ając A. & Z ając M. 2001). spread spontaneously but also have a number of A detailed list of stations for individual keno- stations resulting from planned introduction (they phytes is included in the database of software have been included in Appendix B). dealing with kenophytes (ATPOL-KENO), which Two other groups of species were also excluded is an integral part of the ATPOL database. On the from the cartographic part. These are critical basis of the collected data and using an original species which will reąuire separate taxonomic software package called The Regional Atlas of studies, and the species for which the distribution Plants [Regionalny Atlas Roślin] - RAR21, de- data are incomplete and must be verified. veloped by Józef Gajda of the Institute of Informa tion Technology of Jagiellonian University, dis tribution maps were prepared for 174 species of kenophytes occurring in Poland. Most of the 4.5. Use, interpretation and synthesis maps have been published in the Distribution of data Atlas of Yascular Plants in Poland (Z ając A. & Z ając M. 2001), including 59 maps prepared as The analysis of the kenophyte flora was com- original maps by the author of the present mono- pleted for 300 species out of which two groups were separated: a group of 249 species firmly 21 RAR is a software package that operates all functions established in Poland (Appendices A and B), and of a database containing floristic data from a selected 51 species which can currently be deemed to be region of Poland (or from the whole of Poland). The soft established locally (these are mostly cultivated ware enables the administrator to add, delete, or modify records in the database and present them either as maps on plants and those sometime growing in the “wild”; a VDU or as printouts. the species concerned were marked with “?” preceding the species name; cf. Appendix B). In The collected cartographic data were used to every case, the number of species which were draw up an analysis of the contemporary distri- involved in the analysis is provided in the caption bution of kenophytes in Poland and of the typo- supporting the figurę or table. When the graphs logy of their ranges (Chapter 6). The maps have and diagrams were drawn, the principle of “double” been obtained by superimposing individual dis- (or “multiple”) counting was adopted, if two (or tribution maps on one another. On the basis of more) categories are given for the same species. distribution maps for 174 kenophytes the species This principle covers both the origin (e.g. a species have been grouped according to the type of dis whose primary disjunctive rangę includes North tribution in Poland. Comprehensive maps illus- America and Asia has been included in both trating the distribution of groups of species categories of origin), manner of introduction (Chapters 6, 9 and 10) were drawn using the (intentionally introduced and also accidentally options of RAR software (cf. Chapter 4.4). In brought in), and the manner of reproduction. each basie cartogramme unit, an average number The data on the overall number of species in of species from distinguished group occurring the Polish flora were taken from the newest there was calculated. The density of species in a edition of the critical checklist of vascular plants cartogramme field is represented by the size of (M irek et al. 2002), taking into account only the symbol used. Diameter of each circle reflects the spontaneous flora and the cultivated species now number of species in a given cartogramme unit. growing in the “wild” (several hundred species The smallest point corresponds to 1 species in a which are only known as cultivated plants were square (e.g. Figures 25-36 were obtained when thus omitted). Other sources used in the study are the second root of the diameter is taken as the indicated in captions to the relevant tables and measure of the number of species). graphs. For the selected group of 25 species differing The similarity between floras were determined with respect to origin, biology and the represented through cluster analysis conducted by the Ward type of spread within Poland, the histories of their method of minimum variance (M arek 1988), using expansion within Poland were reconstructed and the Statistica 5.0 software package. The results presented in cartogrammes drawn for the consec- obtained are presented in the graphic form of utive time periods (Chapter 7): a dendrogram. 1. prior to 1850 To increase the readability of graphs, some of 2. between 1851 and 1900 them are presented in the form of a logarithmic 3. between 1901 and 1950 function. 4. between 1951 and 2003 PART THREE

Analysis and synthesis of data

5. Geographical and ecological Table 3. Composition of the vascular flora of Poland characteristics of the flora Number of species compiled of kenophytes in Poland Group of species from M i r e k et al. 2 0 0 2 and present author’s sources Native species 2 5 3 7 1 5.1. Proportion of kenophytes Alien species 1 0 1 7 in the recent flora Diaphytes 511 Established aliens 4 6 0 archaeophytes 1 6 0 5.1.1. General remarks kenophytes 300 Species of uncertain status 4 6 Total 3 5 5 4

In the flora of Poland which now includes 3 554 1 According to M ir e k et al. 2 0 0 0 and own sources. Among all taxa, 1 017 species of alien origin have been noted taxa extinct species and probably extinct species are included. to date, amounting to 29% of its composition (Ta Several hundred omamental and useful plants (trees, shrubs and perennials) freąuently cultivated in Poland and listing in ble 3; Fig. 3). Among the alien species, the follow- the critical checklist of vascular plants of Poland are excluded ing categories are distinguished: archaeophytes, or here.

diaphytes 50.2% (14.4%)

native species ą 71.4%

kenophytes species of uncertain 29.5% (8.4%) status 4.5% (1.3%)

archaeophytes “ 15.7% (4.5%) n = 3554 n = 1017 Fig. 3. Participation of alien species in the flora of Poland and composition of Polish alien flora older newcomers (they constitute ca. 16% of all lished (29.5% of alien species and 8.4% ofthe entire alien species and 4.5% of the entire flora) and more flora respectively) and diaphytes, i.e. species not recent newcomers (79.7%), further divided into yet established (50.2% of alien species and 14.4% kenophytes - plants which are permanently estab- of the entire flora) (Fig. 3). The subject of this monograph is a group of 300 kenophytes including species came from different regions of Europę, 9 taxa of subspecies rank, 2 varieties and 25 of including those from Southern Europę (Mediterra- hybrid origin (Appendices A and B). nean or Sub-Mediterranean), from the south-eastem part of Europę, as well as species whose natural distribution ranges are limited to central regions of Europę, particularly the Alps (Fig. 4 & 5). An iden- 5.1.2. Origin tifiable group among the more recent newcomers is one of those North American species which ori The kenophytes occurring in Poland originate ginate from areas with climatic conditions evidently from five continents (Fig. 4). The majority of close to conditions prevailing in Europę (Fig. 4).

Fig. 5. Direction of origin of European elements in the kenophyte flora of Poland Kenophytes coming from western and south- westem Asia also have a relatively large share of the present flora of Poland. In the group of 300 kenophytes covered in the The kenophytes occurring in Poland are from 61 present study, 25 taxa (8%) are species of hybrid families (out of a total number of 188 families in origin. These are the hybrids which emerged the native flora), and from 169 genera. The ma- spontaneously or which owed their existence to jority of the taxa comprise a smali number of humans (i.e. cultivated species now growing in species, namely: 110 genera with a single species, the wild). Particularly worthy of attention are 10 33 with two, 13 with three; these groups combined taxa of hybrid origin within the genus Oenothera, constitute 92% of the genera described. At the whose representatives sometimes form “swarms other extreme, the genera with the greatest num ber of species are: Oenothera - 22, Rosa - 1 1 , of hybrids” (R ostański & S zo tk o w sk i 1973). Populus and Rubus - 6 species each, Amaranthus, Aster and Chenopodium - 5 species each, and Atriplex, Brassica, Bromus, Geranium, Mentha 5.1.3. Timing and method of arrival and Yeronica - 4 species each. It has been found that in the flora of Poland, the most species-rich families are the same fam The kenophytes reached Poland in different ilies which show high proportions of keno historie periods, beginning from the end of the phytes, namely: Asteraceae - 46 species, Ro- 15th century; the older arrivals in this group have saceae - 37, Onagraceae - 23, Brassicaceae - reached an advanced “age” of 400 years: albeit 19, Fabaceae and Poaceae - 14 each (Table 4; their number is few. Most of the kenophytes sup- Fig. 7 & 8). The most species-rich family - plemented the flora of Poland in the 19th century Asteraceae - includes, apart from kenophytes, (Fig. 6), either introduced intentionally or brought eąually numerous archaeophytes and ephemero- in accidentally. phytes (Fig. 8 & 9). The families of Fabaceae,

A n = 300 ajlOOn 2 90- O” 80 S 70- E 60- ^ __

40- 30- 20- 10-

o \ ------T------T------,------T------T-----L ------r------, 1501-1600 1601-1700 1701-1800 1801-1850 1851-1900 1901-1950 1951-2000 Historical period of first record

B n = 174

Historical period of first record in the wild Fig. 6. Recording history of kenophytes in Poland: A - data of first record from cultivation or from the wild have been taken into account, B - exclusively data of first record from the wild have been taken into account Table 4. Families which are richest in genera and species in Lamiaceae, Onagraceae, Polygonaceae, Ama- the kenophyte flora ranthaceae and Poaceae, Brassicaceae, Chenopo- Number Number diaceae and Solanaceae, must evidently be Family [%] [%] of genus of species deemed “synanthropic” with high percentages of Asteraceae 27 16.0 46 5.3 species of alien origin and high proportions of Rosaceae 15 8 .9 3 7 2 .3 kenophytes among them (Fig. 8 & 9). On the Brassicaceae 11 6 .5 19 6 .3 other hand, native species prevail in such fam

Fabaceae 11 6 .5 14 4 .7 ilies as Rosaceae and Cyperaceae and account for all species representing the family of Orchid- Poaceae 9 5 .3 14 4 .7 aceae (53 species). Furthermore, all the species Chenopodiaceae 5 3 .0 13 4 .3 belonging to family Amaranthaceae in Poland Scrophulariaceae 5 3 .0 9 3 .0 are of alien origin (Fig. 8). Lamiaceae 5 3 .0 8 2 .7

Onagraceae 2 1.2 2 3 7 .7

Salicaceae 2 1.2 9 3 .0

A n = 249 B n = 300 % % 20- 20 •

15 15 ■

10 10 •

n n n n n H n n n n n I Hnn rinnin n

CL CO o (0 C O w Cd ? » u- g. 3 (o a < n 2 o - c — 1 u m c a . o c a =3 m a ) o CL m lu m m Oo co ° Fig. 7. The families most frequently represented in the kenophyte flora: A - spectrum for permanently established species, B - spectrum including locally established species (Appendix A & B). Families shown in black differ between figures A & B

700-i

600-

500-

400-

300

200

100-1

JLA Ł j x D. _Ql H 0) 'o. U) o 03 D 0 0. 'w o < 03 c o CL Ic (/> LL 0Ć. CL 03 U) o 03 c 03 3 _] CL c >» L_ C/3 >N < o _c o co D o c Q_ o o 03 c o co 03 0 ok. Q_ E n o 03 < Cć. o C0 o Fig. 8. Number of native and alien species in the 18 most species-rich families in the flora of Poland

Number of species for the Polish flora according to M ir e k et al. 2002 □ Ephemerophytes (casual aliens) n Archaeophytes □ Kenophytes ■ species of uncertain status Q) 8 0 07 8 i 0) $ .07 # © # / 2002 and present author’s sources

8 Qr ir e k M © 8 rS CQ ł / 1 co Number ofspecies for the Polish alien flora according to 80 ■ 70 ■ 10 60 50 ■ 30 40 20 o E

The Establishment.. u» u> Therophytes are either the dominating or co-dominating life form among the anthropophytes. They constitute nearly 70% of all archaeophytes occur- Among the kenophytes studied, hemicrypto- ring in Poland, 60% of ephemerophytes, and more phytes and therophytes predominate (Fig. 10). The than 25% of kenophytes, while in the native flora relatively high proportion of woody plants among they account for some 8%. Hemicryptophytes the kenophytes results from the inclusion within which predominate in the native flora (over 60%) this group of the species listed in Appendix B (rel- also occur in a high proportion among anthropo atively often eultivated plants, returning to the phytes and constitute ca. 30% of kenophytes and “wild” or locally established). Interesting conclu- ephemerophytes, and more than 20% of archaeo sions can be drawn from the analysis of the spec phytes. To summarize, the similarities in the pat- trum of life forms, considered in groups based on tems of frequencies of various life forms among different historical and geographical aspects, all groups of anthropophytes should be emphasised viewed with respect to the whole flora of Poland. and the difference in this respect from the native flora (Fig. 10) should be marked.

1700-1

E 3 therophytes hemicryptophytes Z 1600- □ hydrophytes 0 geophytes -O mm E ■ ■ chamaephytes ■i 1500- = fanerophytes

300 J =

200 -

100 - =

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ARCHAEOPHYTESKENOPHYTESEPHEMEROPHYTES

roX E

c 20 E O 40 -

6 0 -

100 Ward’s method

120 " Fig. 10. A comparison of the participation of life-forms in the native and alien flora of Poland (A) and the similarity between floras (determined through cluster analysis conducted by the Ward method of minimum variance) (B) (which produce suckers; their breaking branches are also capable of taking root). The vegetative manner of reproduction is of essential importance The kenophytes occurring in Poland are mostly pollinated by insects, wind or are self-pollinating n = 300 plants (Fig. 11). The proportions of apomictic plants are fairly Iow. Among kenophytes occurring in Poland, the majority of species reproduce by generative processes, and some species also use various kinds of vegetative reproduction (Fig. 12). Seven spe cies of kenophytes solely use the latter method: the aąuatic plant Elodea canadensis and a rush- community species Acorus calamus (fragments of plants are carried by water or birds), and poplars22 GA/ V V/G V/G? n = 300 Reproduction type Fig. 12. Number of species with different reproduction types in the Polish kenophyte flora: G - generative: V - vegetative

n = 300 3 5 1

30-

20-

15-

10 -

Pollination mode n Fig. 11. Number of species with different pollination modes o o o O O O o o JZ JZ sz sz -C sz -C sz in the Polish kenophyte flora: o o o o o o o o o o o o o o a - apogamic, i - insects, s - self-pollination, w - wind o 2 ■D N 2 ■4—* CL 0 *D c CD 03 o E -Q 03 0 sz Cl) 0 c c 03 22 Poplars are dioecious trees with flowers of either sex. E co Although the seeds are viable only for a short time, they can Dispersal mode germinate in an equally short time. The małe clones are planted much more oftcn than the female clones, bccause the Fig. 13. Participation of species with different seed dispersal latter produce enormous quantities of seeds with cottony tufts. modes in the Polish kenophyte Hora also in the expansion of the perennial plants of n = 174 the genus Reynoutria (knotweed) and the species 70 O)(/) Asclepias syriaca and Artemisia austriaca. ‘o

latter reflecting the role of animals and humans CD -Q in their migrations (Fig. 13). | 50 Among the kenophytes, the species in which great competitive potential (type С life strategy) predominates have adapted to the circumstances 40 - where the impact of stress is Iow, and the competitiveness is limited by disturbances (type C-R strat egy) and mobile pioneer species (type R) (Fig. 14). 30 -

O O o o o O CD CM I I 2 7 CNI T“ o d o o CM CD % of squares

174 В 70 n= Q)C/3 'o0 CL 100% stress-tolerance 60 - Fig. 14. Percentage of species showing particular life Ф .o strategies (G r im e 1979) in the Polish kenophyte flora E (/i= 180): -? 50 С - the competitive strategy, R - the ruderal strategy, S - the stress- tolerant strategy 40 -

5.1.7. Freąuency and status 30 in the flora

20 - Freąuency analysis for the occurrence of keno phytes was undertaken for a group of 174 spe cies for which representative data was obtained 10 - for the whole of Poland (cf. Chapter 4). The allocation of species to freąuency classes was based on both the number of cartogramme с С ^ с 03 ш sąuares where they appear and the number of оз 03 ГО 03 Ф с CD ~o о тз о 3 о с о с о сг ел recorded stations. ZJ — 13 — о —г JD —' 4= S 03 С 03 03 о In the first case, the species recorded in less than 0 с о D о 10% of the total number of ATPOL sąuares are the сг (/) О 03 о most numerous, whereas the species recorded in о more than 60% to 100% of sąuares (i.e. on a large о Frequency scalę, or the whole of Poland) are least numerous (Fig. 15 A). Fig. 15. Freąuency distribution of kenophytes: A - in relation to the number of ATPOL sąuares, В - in relation to the However, freąuency analysis based on the num number of stations. Scalę of freąuencies: 1-14 stations - rare, 15-50 ber of stations points to a significant proportion of stations - occasional, 51-500 stations - occasional, locally freąuent, 501-6000 stations - frequent, locally abundant, > 6000 stations - freąuently occurring (or even locally common) abundant (common) kenophytes, whereas the groups of scattered or rare numbers of population (Table 5). It was found kenophytes are smaller (Fig. 15B). that among the most common species are those The scales adopted permit the determination which were accidentally transported from both of the list of the abundant kenophytes (com- the Americas and Asia and which established mon), both with respect to the type of distribu- themselves in anthropogenic habitats. They are: tion in Poland, as well as, indirectly, to the Chamomilla suaveolens and Conyza canadensis,

Table 5. List of the abundant and most frequent kenophytes in Poland according to the number of 100 km sąuare records and number of localities

No of Way of No of localities 1 Species II Origin introduc- Habitats squares [%] up to year tion 2000

1 Chamomilla suaveolens 2 965 81.3 13 125 1 Am N & Asia E Ul H 2 Conyza canadensis 2 929 80.3 11 600 2 AmN Ul H 3 Galinsoga parviflora 2 726 74.8 10 932 3 Am S & C UI/L H 4 Amaranthus retroflexus 2 379 65.2 7651 6 Am N & Am C I/UI H 5 Yeronica persica 2 204 60.4 7 887 5 Asia SW Ul H 6 Oxalis fontana 2 141 58.7 8 806 4 Am N, Asia E ? Ul H 7 Galinsoga ciliata 2021 55.4 6 777 8 Am C, Am S ? UI/I H 8 Acorus calamus 1 999 54.8 4319 13 Asia C & S I/UI NS 9 Robinia pseudoacacia 1 957 53.7 7 067 7 AmN 1 NSH 10 Senecio vernalis 1 948 53.4 3 932 14 Eur SE & Asia W Ul H 11 Elodea canadensis 1 847 50.7 3 681 15 Am N UI/I NSH 12 Medicago sativa 1 743 47.8 5412 10 Asia SW I SH 13 Impatiens parviflora 1 681 46.1 6 730 9 Asia C & E I NSH 14 Solidago gigantea 1 668 45.7 5 348 11 A m N INSH 15 Juncus tenuis 1 440 39.5 5 332 12 Am N Ul SH 16 Lupinus polyphyllus 1 387 38.0 2 674 22 Am N I NSH 17 Acer negundo 1 379 37.8 3 523 17 A m N I NSH 18 Solidago canadensis 1 254 34.4 3 434 18 Am N I NSH 19 Lycium barbarum 1 224 33.7 2 634 23 Asia E Eur SE I NSH Eur S & W, Afr N 20 Lolium multiflorum 1 174 32.2 2 792 21 ISH & Asia SW 21 Reynoutria japonica 1 158* 31.8 3 004 20 Asia E 1 NSH 22 Erigeron annuus 1 133 31.1 3 557 16 A m N I SH 23 Padus serotina 1 134 31.1 2 564 24 Am N & Am S INS 24 Bidens frondosa 1 068 29.3 3 142 19 AmN UI/I NSH 25 Datura slramonium 1 044 28.6 1 881 29 Am N, Asia? I H 26 Diplotaxis muralis 991 27.2 2 049 27 Eur S & W [Afr.] Ul H 27 Sisymbrium loeselii 976 26.8 2 326 25 Eur SE & Asia C Ul H 28 Rudbeckia ładniała 903 24.8 2251 26 AmN I NSH 29 Sisymbrium altissimum 812 22.3 1 770 30 Eur SE & Asia C Ul H 30 Elsholtzia ciliata 814 22.3 1 352 37 Asia E IH 31 Helianthus tuberosus 778 21.3 1 416 34 AmN I NSH 32 Tanacetum parthenium 734 20.1 1 179 43 Eur SE & Asia SW I H 33 Bryonia alba 728 20.0 1 328 38 Eur E & Asia W I NSH 34 Lepidium densiflorum 724 20.0 1 259 48 AmN Ul H 35 Sinapis alba 716 19.6 1 416 35 Eur S I H 36 Хап tli ium strumarium 712 19.5 1 105 47 Eur/Am N ? Ul H 37 Echinocystis lobata 708 19.4 2 047 28 A m N I NSH 38 Rosa rugosa 701* 19.2 1 299 40 Asia E 1 NSH 39 Impatiens glandulifera 675 18.5 1 574 32 Asia C I NSH 40 Rumex confertus 673 18.5 1 731 31 Eur SE & Asia W Ul SH

Total number of sąuares in Poland = 3646 I 40 most frequent kenophytes according to the number of recorded squares; II - position of kenophytes according to the number of recorded localities: red shading - 10 most frequent species: position 1-10; dark yellow shading - following 10 species: position 11-20; light yellow shading - following 10 species: position 21-30 Abbreviations: I - intentionally, Ul - unintentionally, H human-made habitats (anthropogenic), S - seminatural habitats, N - natural habitats, * indicates that number of squares recorded need to be verified. А В С______О ______Е F __ G А В С D E F G

Chamomilla suaveolens (P u r s h ) r y d b . Conyza canadesis (L.) c r o n q u is t

А В С D E F Q А В С D E F G

А В С D E F G А В С D E F G

Senecio vernalis w a l d s t . & K it . Sisymbrium loeselii L. Fig. 16. Examples of kenophytes showing different degrees of )undance in the Polish flora: abundant and frequent species (after Z a ją c A. & Z a ją c M. 2001, supplemented)

recorded in over 80% of cartogramme sąuares, (although on a local scalę the last species may and Galinsoga parviflora, Amaranthus retrofle- be included within the category of “freąuent”). xus and Veronica persica, recorded in 60-80% Analysing the types of habitats colonised by of ATPOL squares. Among Polish kenophytes all the kenophytes included in the study, it these are also the species for which the highest should be noted that almost half of them limit numbers of stations have been recorded to date. their occurrence to anthropogenic habitats (Fig. The most freąuently occurring species, which 18). Most often, these are species that were are even common in many areas include, among accidentally introduced. The species capable of others: Robinia pseudoacacia, Senecio vernalis, concurrent colonisation of natural and semi- Solidago gigantea or Sisymbrium loeselii (Fig. natural habitats are relatively freąuent and within 16). Sparsely distributed but locally freąuent this group the species intentionally introduced species include for ехатріе: Ambrosia artemisii- by humans predominate (Fig. 19). Least nume- fo lia , Centaurea diffusa, Diplotaxis tenuifolia rous are the species which established them- and Trifolium patens, whereas examples of selves in natural and semi-natural communities, sparsely distributed to rarc spccics might in by-passing the stage of colonising anthropogenic clude: Corydalis lnica, Oxalis dilleni, Miinulus habitats (c.g. Genistella sagittalis, Impaticns moschatus and Impaticns capcnsis (Fig. 17) capcnsis, Lemna turionifera, Miinulus guttatus). nirn-n-r N,111!!! ...... " j. ' i / 1'! : ;; j i — i i r ? v 1 " ~T / N i i - ; ' — V, C ( _r ~ V. Y Y ^ \ ' ~ \

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Diplotaxis tenuifolia (L.) d c . Trifolium patens s c h r e b .

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Corydalis lutea (L.) d c . Impatiens capensis m e e r b .

Fig. 17. Examples of kenophytes showing diffcrent degrees of abundance in the Polish flora: occasional and rare species (Diplotaxis tenuifolia, Trifolium patens and Corydalis lutea after T o k a r s k a -G u z ik 2001b; Impatiens capensis after Z a ją c A. & Z a ją c M. 2001) When the types of colonised habitats are con- serotina and Quercus rubra - to forests. On the sidered together with the manner in which they other hand, the “success” of accidentally intro were introduced into Poland, one may draw the duced species in anthropogenic habitats can be conclusion that the species introduced intention- explained by some earlier adaptations (i.e. in ally by humans show a tendency to colonise their respective homelands) to łive in trans- natural and semi-natural habitats, whereas those formed habitats: namely as a result of their brought in unintentionally occupy anthropogenic apophytic potential23 (cf. Chapter 12). habitats before any other. It seems that an ex- płanation should be sought in the capacity of a species to adapt to the conditions faced in the new homeland. The species brought in on pur- 23 Apophytism - the capacity of a species to migrate pose by humans, remaining under cultivation for from its proper natural habitats to synanthropic communities developing in anthropogenic habitats. S t a r f in g e r a long time, had the opportunity to develop (1998) along with other authors is of the opinion, that the ecotypes adapted to specific environmental apophytism of a species within the limits of its natural conditions, and some of them were introduced rangę may be regarded as an indicator for its later success directly into the “target” habitats, e.g. Padus as an invasive species. n = 300 5.2. Kenophytes in historical accounts w 160 і of floras OCD (D (/)O. 5.2.1. “Old” floras

The available factual data in the form of histo rical floras from the 18lh, 19th and 20th centuries together with preserved herbarium specimens allow only fragmentary reconstruction of the development of the flora with respect to the more recent newcomers into the present territory of Poland. One of the oldest sources is the work by S yreń - ski ( S ir en iu s) (1613), who listed 16 species which are now regarded as relatively recent newcomers, as established in Poland: Acorus calamus (Sir. Vol. I/Chapter 3), Chenopodium botrys (Sir. Vol. III/Chapter 51), Clematis vitalba (Sir. Vol. 1/ Chapter 95 (2), Datura stramonium (Sir. Vol. V/ Chapter 85), Echinops sphaerocephalus (Sir. Vol. III/Chapter 10), Hesperis matronalis (Sir. Vol. III/ Chapter 65), Hyssopus officinalis (Sir. Vol. III/ Chapter 23), Inula helenium (Sir. Vol. I/Chapter 16), Lonicera caprifolium (Sir. Vol. II/Chapter 94), Lycopersicon esculentum (Sir. Vol. V/Chapter 95), Marrubium vulgare (Sir. Vol. III/Chapter 25), Physalis alkekengi (Sir. Vol. III/Chapter 51), Portulaca oleracea (Sir. Vol. IWChapter 88), Tana- cetum parthenium (Sir. Vol. III/Chapter 98),Xan- thium strumarium (Sir. Vol. II/Chapter 78) and Ambrosia artemisiifolia (Sir. Vol. III/Chapter 50). Type of habitat The species mentioned above do not exhaust the Fig. 18. Habitat preferences of kenophytes occurring in list of alien plants included in the Sirenius work, Poland: but the remaining ones still require further studies II - human-made (anthropogenic), S - seminatural, N - natural in the fields of nomenclature and history. % n = 300 Because of an almost complete absence of flo- 100 i ------ristic data from the 17th century, and very scarce 90 ■ data from the 18th century, the reconstruction of historie floras of kenophytes is feasible but mostly only for the last 200-250 years. The authors of old floras, studying different areas now falling within the borders of Poland, have listed a total number of 138 species out of the group of 300 species that have recently es tablished themselves (Table 6; Fig. 20). The flora published in the second half of the 18th century by K luk (1786-1787-1788) included more than thirty kenophyte species. However, most of the species referred to in this work were

------^ ------1------^ ------1------either plants cultivated as medicinal plants, pro- Human-made Natural/ Both types viding industrial raw materials, cultivated for Seminatural food or fodder, or as decorative plants24. Only ■Туре of habitats

24 The Dictionary by K l u k (1786-1787-1788) included □ Unintentional ■ Intentional both native species: “proper native plants” [“proper native Fig. 19. Structurc of the Polish kenophytc flora with respect plants are only those which grow in any corner of our to type of habitats and the presumed type of country, in the wild, unattended by humans”] [“rośliny introduction into the country właściwe kraiowe”] [“rośliny właściwe kraiowe są tylko te, Table 6. Participation of kenophytes in the Floras of Poland and associated areas in different historical periods o

ON T 1 9 5 3

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yreń sk i u 1 8 9 8 - 1 9 4 0 S B A K RosTAFiŃSKi 1872 S Grabowski 1843 W W K łZ co Acorus calamus O ••• • •• •• • • Datura stramonium O •• •• •• • •• • • Portulaca oleracea OO* O O# 09 • • □ •• •• s u b s p . oleracea Inula helenium O • • 09 • •• •• • Marrubium vulgare o om • 9 ••• •• • • Mercurialis annua •• • ••• •• Reseda luteola o* •O • •• • •• Ambrosia artemisiifolia o • • • Malva moschata • • • • • Picris echioides • • •• Amaranthus retroflexus • • •• ••• •• Convza canadensis •• •• •• • •• Senecio vernalis • • •• ••• •• Oxalis stricta • • ••• •• •• Veronica persica • • •• • • Xanthium strumarium o •• •• •• • •• Eragrostis minor • •• •• Cardaria draba • • • □ • • •• Geranium pyrenaicum • • ••• •• • Chenopodium botrys o •• ••• • •• Geranium divaricatum • • • □ •• Sinapis alba o O# 09 • • ••• Bryonia alba o o • •• ••• 09 Onobrychis viciifoiia o» om • •• 09 Medicago sativa • 9 09 •O# 09 • 09 Atriplex hortensis o o * O 09 09 Aster salignus • 9 9 Helleborus viridis • 9 • 9 09 Salix acutifolia • 9 o 09 Syringa vulgaris o o o 09 09 Sedum album o 9 • □ 9 9 9 Lycium barbarum 09 09 o 09 09 Lolium multiflorum 9 09 • 9 09 Rudbeckia laciniata 09 09 • 9 9 09 Xanthium spinosum 9 9 • 9 9 • 9 9 Cymbalaria muralis 9 9 • 9 9 Clematis vitalba o o 9 09 □ 9 9 Lathyrus nissolia 9 9 9 Ulex europaeus 9 9 • 9 O

<------które w któreykolwiek stronie kraju rosną same przez się dziko, bez ludzkiego pielęgnowania”], and “foreign plants, not known earlier” [“rośliny cudzoziemskie nieznaiome”]. The author gave the following description of this group: “Plants of three kinds of those earlier unknown to us will be described here, either those which could be kept in our country as useful plants, or plants whose parts could be used for meals, or as paints, medicines etc., or finally those which display extraordinarily curious aspects”. [“Troiakie rośliny nieznaiome nam znajdą się tu opisane: albo takie, któreby pożyteczne w Kraiu utrzymywane być mogły: albo takie, których iakie części do stołu, lekarstw, farb, etc. zażywamy: albo nakoniec takie, które nadpospolitą osobliwość w sobie mają”.].

6 The Establishment. 41

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Galinsoga parviflora m • • • • • • Impatiens parviflora • • • • • Sisymbrium loeselii • • • • • Solidago canadensis 0 9 • • • • Xanthium albinum • • • • • Bunias orientalis • • □ • • • Echinops sphaerocephalus O • • • • • Erigeron annuus • • • • • Salsola kali s u b s p . ruthenica • • • Helianthus tuberosus O O om om om O» O# Hesperis matronalis s u b s p . matronalis O O • o m • • O# Tanacetum parthenium O O O# m m • • O* Brassica nigra O • • • o* Elsholtzia ciliata • om □ 09 • Digitalis purpurea O o# m o# m o# Diplotaxis tenuifolia • m • m • Calendula arvensis O • • • Dianthus barbatus O o* om □ • om om Hyssopus officinalis O O om □ m om Artemisia austriaca m □ m Asclepias syriaca • m Lysimachia punctata o» m • m om Petrorhagia saxifraga • m m Sicyos angulata o« m om om om Chamomilla suaveolens m m m Elodea canadensis m • m m Juncus tenuis m • m m Solidago gigantea m • m m Diplotaxis muralis m m • m m Aster novi-belgii m m m Medicago x varia m m • m Anthemis ruthenica m m m Atriplex tatarica m □ m m Brassica rapa s u b s p . rapa O m • Geranium bohemicum m m • m Geranium sibiricum m Mentha rotundifolia m om Mimulus guttatus m m m Ornithogalum boucheanum m m om Oxalis corniculata m • m m Polycneum heuffelii m m Rosa glauca m • m m Rosa pimpinellifolia m m m Sedum spurium m • m om Silene conica m • m m o ro ON

7 1 9 5 3

1 8 7 2 1 8 4 3

et et al. 0 Species O n

1 6 1 3 eta/. 1 8 6 8

1 8 9 0 U

1 8 7 2

1 8 4 7 CQ

1 7 8 6 - 1 7 8 8 D immer

aga 1 bromeit luk r a b o w sk i napp erdau o sta fiń sk i zafer yreń sk i O 1 8 9 8 - 1 9 4 0 B S R A S G W Fiek 1881 K K W t/5

Artemisia dracunculus OOO 09 • Bryonia dioica O •• O • O# Amaranthus lividus • • Sisymbrium altissimum ••• • Vicia pannonica • • • Lupinus polyphyllus ••O# Aster novae-angliae •• Aster tradescantii •• Bromus japonicus • • Bromus sąuarossus □ ••• Erucastrum gallicum ••• Silene dichotoma ••• Lonicera caprifolium O OO 09 O# 09 Rubus odoratus 09 o# Vitis vinifera O O 09 o# O Cerasus mahaleb 09 o* O Lonicera tatarica 09 o# 09 Myrrhis odorata OO □ • 9 Galinsoga ciliata • 9 Reynoutria japonica • 09 Anthoxanthum aristatum • 9 Bidens Jrondosa • 9 Lepidium densiflorum • 9 Vicia grandiflora • 9 Amaranthus albus • 9 Artemisia annua • 09 Atriplex oblongifolia • 9 Bidens connata • 9 Centaurea diffusa • 9 Erechtites hieracifolia • 9 Euphorbia humifusa • 9 Kochia scoparia □ • 9 Lepidium virginicum • 9 Physalis alkekengi O □ • 9 Potentilla intermedia • 9 Reynoutria sachalinensis • 09 Sisyrinchium bermudiana • 9 Lycopersicon esculentum OO O o® O Acer negundo o* O Padus serotina O 09 Impatiens glandulifera • Epilobium ciliatum 9 Erigeron ramosus 9 Rumex confertus 9 Alnus rugosa 09 Amelanchier spicata 09 Amorpha fruticosa 09

1 9 5 3

1 8 7 2 1 8 4 3

r- et al. Species "śT 1 6 1 3 00 et et al. 1 8 6 8 1 8 9 0 1 9 0 1 - 1 9 3 0

< 1 8 7 2 1 7 8 6 - 1 7 8 8 o 1881 < immer luk ra b o w sk i napp bromeit o sta fiń sk i erdau y r eń sk i iek chube zafer 1 8 9 8 - 1 9 4 0 S W K G R F K B A S £ S

Corydalis lutea • Hordeum jubatum • Mimulus moschatus • Rosa rugosa • o« Rudbeckia hirta O# 09 Sorbaria sorbifolia 09 09 Aster lanceolatus 9 Beckmannia eruciformis 9 Cuscuta campestris 9 Cuscuta trifolii 9 Erysimum marschallianum 9 Erysimum wahlenbergii 9 Genistella sagittalis 9 Linaria repens 9 Liman austriacum 9 Polycneum majus 9 Rumex longifolius 9 Solidago graminifolia 9 Trifolium patens 9 Vicia dasycarpa 9 Rumex patientia O 09 Amaranthus chlorostachys 09 Cerasus vulgaris 09 Elaeagnus angustifolia 09 Malus domestica 09 Mentha x niliaca 09 Mentha spicata 09 Pyrus communis 09 Prunus domestica 09 Ribes rubrum 09 Robinia pseudoacacia O OOOO O Aesculus hippocastanum O O o O O O Quercus rubra O Juglans regia O O Ailanthus altissima OO Linum perenne □ Oenothera glazioviana □ Parthenocissus inserta O Pinus banksiana o Pinus ni gra o Pinus strobus o Rhus typhina o Rubus laciniatus O Scutellaria altissima □ Symphoricarpos albus o

O - cultivated plant; O # cultivated and escaping from cuItivation / becoming wild; • occurs exclusively in the wild; □ - recorded in the flora of the region but outside the contemporary border of Poland; without shading - species rare or oeeasional at the present time in Poland; yellow shading - species occasional to locally frequent at the present time; red shading - abundant species at the present time. seven species: Acorus calamus, Ambrosia artemisiifolia, Datura stramonium, Inula helenium, Malva moschata, Mercurialis annua and Picris echioides were described by Kluk as those which occurred in the wild and were already established, while three more species: Marrubium vulgare, Portulaca oleracea and Reseda luteola - were described as plants which during his time were often either cultivated or returning to a wild state from cultivation. The number of kenophyte species reported by particular authors, taking into account their increase in successive periods, depended first on the degree of knowledge about flora at that time as well as on the size of the described territory (Fig. 20 & 21). The species which are listed by all or by the majority of the historie authors ąuoted, are those oldest arrivals, which are now common throughout Poland (e.g. Acorus cala mus, Datura stramonium), as well as the plant species which have been cultivated and have then often gone into a wild state (e.g. Hysopus offi- Source of data cinalis and Marrubium vulgare). Fig. 20. Number of kenophytes recorded in historical floras of Poland and associated areas

(1795-1807) (1874-1918) (1939-1945)

1815-1874

1921-1939 from 1945

Fig. 21. Poland’s changing territory (after D avis 2001): A - Republic of Poland - Lithuania (990 000 km2), В - partition o f Poland, С - Duchy o f Warsaw (154 000 km2). D - Congress Kingdom of Poland (127 000 km2), E - Second Republic of Poland (389 720 km2), F - Republic of Poland (312 685 km2) The Flora of the Congress Kingdom of Poland with respect to Diplotaxis tenuifolia sporadically by W aga (1847) reported 10 species of kenophytes listed in the contemporary sources (cf. Appendix A). (including 4 under cultivation), while The Flora It is a plant long used in Europę as a vegetable (and published by R ostafiński (1872) listed as many as still cultivated today), and perhaps at the time it was 55 such species, and the Floras for Silesia and being referred to as a cultivated plant. Pomerania where records were systematically The subseąuent centuries are characterised by a collected over long periods included 62 (A bromeit further inerease in proportion of new arrivals in the et al. 1898-1940), 71 (F iek 1881) and 119 species flora of Poland. The first half of the 19th century was (S chube 1901 a, b—1930) (T able 6; Fig. 20). T he evidently marked by intensified inbound migration guide The Plants o f Poland [Rośliny polskie] pub of alien species, although the highest “migration lished after World War II (S zafer et al. 1953), w ith waves” were in the second half. Throughout the supplements covering those species of alien ori- periods referred too, there is a remarkably high pro gin which the authors regarded as established and portion of species of European origin (chiefly from expanding their ranges in Poland, listed 141 sp e the southem, south-eastem and south-westem part of cies of kenophytes, plus 17 more species as cul- the continent) among the migrants. From the 16th cen tivated plants which are now deemed to be estab tury up to the first half of the 19th century, there was lished (locally at least). an evident predominance of species “flowing into” Poland from various regions of Europę and Asia. The first and second part of the 19lh century showed a marked inerease in the proportion of species origi- 5.2.2. The “oldest” arrivals among nating from both Americas (but particularly North the kenophytes and the fairly America) (Fig. 22; see p. 51). More recently, the recent ones proportion of taxa of hybrid origin, whose emergence has been assisted by humans either directly or indirectly, is on the inerease in the flora of Poland. The compilation of the available historie data In the analysis of life forms of kenophytes, made provides the source for a partial reconstruction of for subseąuent historie periods, one should focus on the historical floras of kenophytes, beginning from the second parts of the 19lh and 20th centuries, when the 17lh century (Table 7). Undoubtedly, such kenophytes displayed the fuli spectrum of life forms species as Acorus calamus, Datura stramonium, (Fig. 23A & B; see p. 52). In the second part of Echinops sphaerocephalus, Marrubium vulgare, the 19th century, therophytes predominated, as they Sisymbrium loeselii and Tanacetum parthenium were mostly brought in accidentally with dynami- were present in the 17lh century flora of Poland. cally developing transportation systems, while the Most of the species listed had been brought into second part of the 20lh century (and particularly its Poland as useful plants (medicinal, food or fodder, last two decades) was the time when many new decorative, honey-yielding or even poisonous)25, species were introduced into cultivation. This perhaps much earlier than indicated by the first phenomenon reflected a growing human interest in records. At the same time, the following species were new species of woody plants and perennials. recorded in Poland: Ambrosia artemisiifolia, Artemi- sia dracunculus, Clematis vitalba, Chenopodium botrys, Hesperis matronalis, Hyssopus officinalis, Mercurialis annua and Portulaca oleracea, again 5.2.3. The most freąuent kenophytes recorded primarily as cultivated plants or those re- in the floras of subseąuent tuming to the wild state. Certain doubts can be raised historical periods

25 e.g. Acorus calamus - a medicinal plant [“the candied root fortifies the stornach against infections and ‘bad’ air” In the descriptions of the Polish flora up until - K l u k 1786] [“korzeń smażony w cukrze na wzmocnie the year 1850, the stations of 50 species of keno nie żołądka, przeciwko zarażeniu i szkodliwemu powie phytes had been recorded, although for most of trzu” - K l u k 1786], Marrubium vulgare - a medicinal plant the species these were the first stations (for 20 also used as a spice, Datura stramonium - a poisonous species) or species whose number of stations did plant [“careless ingestion causes loss of mcmory, mental confusion, indifference of senses, madness, [...] and not exceed 5 (another 20 species). Only 12 spe a complcte loss of the ability to perform in marital affairs” cies had been recorded at between 5-11 stations - K l u k 1786] [“nieostrożne zażycie przynosi utratę pa prior to 1850 (Fig. 24; see p. 52). mięci, pomieszanie rozumu, nieczułość zmysłów, szaleń In the next half-century, data on 3684 stations stwo [...] zupełną utratą sposobności do sprawy małżeńskiej” for 121 species were recorded. In the first half of - K l u k 1786], Echinops sphaerocephalus - a melliferous the 20th century, more records were collected: 147 plant sown by bee-keepers, Tanacetum parthenium - a decorative plant. species at 9378 stations and 174 species at 196 910 Table 7. Records of expansion of kenophytes in the historical floras of Poland and associated areas

First Intro- record Intro- First duction duction record 1/2 2/2 1/2 2/2 1/2 2/2 Species for XVI XVII to Central to for XVIII XVIII XIX XIX XX XX Europę Europę Poland Poland 1613* ??? Chenopodium botrys Ar 1829 1594 XVII 1859 ?? X Hyssopus officinalis 1819 1613* XVI 1613* ? ? X Artemisia dracunculus Ar XVI? 1850 1613* ?? • Ambrosia artemisiifolia 1863 1873 1767 XVIII ?? Mercurialis annua Ar XVI? 1825 • 1613* ?? Portulaca oleracea Ar 1837 •

Clematis vitalba 1663 1883 1613* 1847 ? ? X

XVI XVII ?? Hesperis matronalis 1817 1613* 1837 X 1613* ? Ш ■ ■ ■ Inula helenium 1819 1837 ? ...... 1613* ? ... ■■■■■■ Marrubium vulgare Ar XVI? 1643 • ...... 1597 1652 ? Diplotaxis tenuifolia Ar 1836 1613* 1613* ? Echinops sphaerocephalus 1809 XVI? 1652 • ----- 1561 1613* ? Tanacetum parthenium Ar XVI? 1824 ? 1613* ? •••••••••• ■ ■ ■ ■ Xanthium strumarium Ar 1837 ? = = = = = Datura stramonium 1613* ? ■ ■ ■ ■ 1584 XVI? 1652 • Sisymbrium loeselii 1654 XVI? 1654 ? • 1613* ? ...... Acorus calamus 1557 1577 XVI 1652 • ...... Conyza canadensis 1730 1646 1825 ? • XVIII Picris echioides 1836 1836 • Reseda luteola XVIII Ar 1825 Malva moschata XVIII Ar 1885 Geranium sibiricum 1840 1840 ••

Linaria repens 1825 1825 • ...... Euphorbia humifusa 1813 1846

Helleborus viridis XVIII 1819 1868

Rubus armeniacus XIX? 1843 ••

Beckmannia eruciformis 1837 1837

Geranium divaricatum Ar 1840 Bryonia dioica 1820 Ar 1847 •• Myrrhis odorata XVI 1809 1837

Potentilla intermedia 1652 ? 1652 ? ? ■ ■ ■ ■ 1841 1841 Atriplex tatarica 1820 ■ ■■■■■ Ar 1847

Xanthium spinosum 1681 1849

Digitalis purpurea 1790 1809 ■■■■■■■■ 1862 Cymbalaria muralis 1640 A r? 1837 ...... ■ ■ ■ ■ ■■■■■■■■

Mimulus guttatus 1824 1824 ■ ■ ■ ■ ■■■■■■■■ 1 ...... Amaranthus lividus Ar 1826 ? ... First Intro- Intro- First duction record Species for duction record XVI XVII 1/2 2/2 1/2 2/2 1/2 2/2 to Central to for XVIII XVIII XIX XIX XX XX Europę Europę Poland Poland

Eragrostis minor 1819 Ar 1838 Geranium pyrenaicum 1762 1837

Medicago x varia XIX 1837

Onobrychis viciifolia XVI 1837 1837

Sinapis alba XVII 1824

Bryonia alba Ar XVII 1824

1 7 3 0 ? Helianthus tuberosus 1627 1872 1829 ? Elsholtzia ciliata 1847 1847 1652? Cardaria draba 1675 1837 Ar Sisymbrium altissimum 1780 1843

Rudbeckia laciniata 1615 1787 1787

Erigeron annuus 1700 1830

Lolium multiflorum 1837 1837

Senecio vernalis 1726 1824 XVI Medicago sativa 1819 1832 1801 Amaranthus retroflexus 1783 1814 Yeronica persica 1809 1862

Oxalis fontana 1658 1809

Galinsoga parviflora 1798 1807

Rubus odoratus 1635 1880 1806 1877

Asclepias syriaca XVIII 1855 1872

Sedum album XVII 1868

Lysimachia punctata 1819 1870

Oxalis corniculata 1576 1863

Sicyos angulata 1868 1868

Silene conica 1879 1879

Sedum spurium 1879 1880

Artemisia annua 1871 1881 1871 1881

Anthemis ruthenica 1869 1869

Silene dichotoma 1841 1877

Xanthium albinum 1822 1853

Bunias orientalis 1856 1858 ?? XVIII Diplotaxis muralis 1842 1851 • ••••••••• - Ar Lycium barbarum 1769 1839 1847 1862 Elodea canadensis 1836 1867

Solidago canadensis 1648 1872

Juncus tenuis 1795 1862 First Intro- record Intro- First duction duction 1/2 2/2 1/2 2/2 1/2 2/2 Species for record XVIXVII to Central to for XVIII XVIII XIX XIX XX XX Europę Europę Poland Poland

Solidago gigantea 1758 1853

Impatiens parviflora 1837 1850 XVIII 1836? Robinia pseudoacacia 1601 1824 1806 1868 Chamomilla suaveolens 1850 1862

Echinops exallatus 1897 1897

Rubus allegheniensis 1890 1899 ca.212 Corydalis lutea XVIII 1884 1884 Mimulus moschatus 1868 1879 Oenolhera glazioviana 1864 1866 1879 1682 Oenolhera parvif!ora 1768 1914 1938 Oxalis dillenii 1865 1865

Rubus laciniatus 1770 1885 1859

Yeronica peregrina 1760 1854

Ornithogalum boucheanum ca. XVI 1880

Sisymbrium wolgense 1880 1896

Vicia pannonica 1884 Ar 1884 Petrorhagia saxifraga 1859

Solidago graminifolia XIX 1888

Atriplex oblongifolia 2/2 XIX 1882 ••••••••••

Bidens connata ca. 1874 1865 1895 Centaurea diffusa 1876 1878

Lepidium \irginicum 1697 1860

Artemisia austriaca 1871 1871

Physalis alkekengi 1866 Ar 1613* 1866 Amaranthus chlorostachys 1872 1872

XVIII / Erigeron ramosus 1888 XIX XVIII Kochia scoparia 1811 1872 1805- Anthoxanthum aristatum 1866 1813 Oenolhera depressa 1835 1894

Parlhenocissus inserla 1629 1884 1806 1884

Rumex confertus 1873 1873

Chenopodium strictum XIX 1939 1891 Impatiens glandulifera 1839 1855 1890

Lepidium densiflorum 1883 1888

Vicia dasycarpa 1898

Bidens frondosa 1736 1869

Lupinus polyphyllus 1877 1877

7 The Establishment. 49 First Intro- Intro- First duction record Species for duction record XVI XVII 1/2 2/2 1/2 2/2 1/2 2/2 to Central to for XVIII XVIII XIX XIX XX XX Europę Europę Poland Poland

Acer negundo 1688 1699 1808 1899

Padus serotina 1880 ? 1623 1825 1813 1900 Reynoutria japonica 1823- 1829 1886 1882 Galinsoga ciliata 1853 1876

Lathyrus nissolia 1903 1903

Erechtites hieracifolia 1700 1902

Amaranthus albus 1723 1907

Reynoutria sachalinensis before 1864 1869 1903 Vicia grandiflora 1877 1907

Oxybaphus nyctagineus 1843 1911

Ailanthus altissima 1751 1874 1818 1931

Ambrosia psilostachya 1901 1901

Barbarea intermedia 1908

Sisyrinchium bermudiana 1845 ? 1928 1863 Genistella sagittalis 1928 1929

Melilotus wolgica 1937 1937

Thladiantha dubia 1917 1917

Yeronica filiformis 1780 1838 1936

Oenothera subterminalis 1856 1938

Trifolium patens 1933

Amaranthus blitoides 1893 1911

Eragrostis pilosa XIX 1939 1934 Iva xanthiifolia 1842 1928

Bromus carinatus 1912 1912

Epilobium ciliatum 1891 1917

Rosa rugosa 1841 1950 1913 ?

Echinocystis lobata 1904 1937

Oenothera suaveolens 1805 1961

Oenothera issleri 1949 1958

Oenothera jueterbogensis 1962 1973

Oenothera pseudochicaginensis 1959 1959

Oenothera punctulata 1969 1973

1809? Chaerophyllum aureum 1994

Helianthus decapetalus 1910 1956

Helianthus laetiflorus 1959 1969

Lemna turionifera 1983 1994

Oenothera fallax 1917 1958 First Intro- Intro- First duction record Species for duction record XVI XVII 1/2 2/2 1/2 2/2 1/2 2/2 to Central to for XVIII XVIII XIX XIX XX XX Europę Europę Poland Poland Oenothera royfraseri 1963 1963

Oenothera oakesiana 1614 1962 1962

Heracleum manlegazzianum 1862 1973

Heracleum sosnovskyi 2/2 XX 1980

Oenothera canovirens 1907 1958

Oenothera pycnocarpa 1958 1963

Oenothera victorini 1961 1961

Oenothera wienii 1937 1937

Oenothera paradoxa 1967 1974 ■ ■ ■ ■

Oenothera hoelscheri 1942 1942

In this table only the information from the earliest record is given. Nrs of localities Nrs of localities Nrs of localities Symbol for the species Symbol for the species Symbol for the species Further information is given in Appendices A and B. at the particular time at the particular time at the particular time

Abbreviations used in the table: 1601-2000 Ar - species classified as an archaeophyte in some part of Europę * - occurrence o f species in Poland recorded by SYREŃSKJ (1613) without specification of species status (i.e. whether in cultivation or in wild) X - in cultivation 321-400 4001-6000 ? - probably occurred in the wild, but may have only been cultivated 1 - “old” cultivated plant, probably only in cultivation at that time 11-40 2 - possible doubtful determination of the species at that time 41-80

81-120

121-160

□ Europę □ Asia 0 America N ■ America С & S ■ Africa ■ Anthropog. □ not defined

1501-1700 1701-1800 1801—1850 1851-1900 1901-1950 1951-2000 Historical sequence Fig. 22. Participation of kenophytes of different geographical origin becoming established in Poland in the historical seąuence 1501-ХХ century stations, respectively. The combined number of list of the most frequent kenophytes (this species stations recorded for 174 species of kenophytes prefers certain types of habitats, such as rubble now exceeds 210 000 (Fig. 24; Appendix A). heaps and railway tracks). At the beginning of the The composition of the kenophyte flora ex- 20th century the highest number of stations was pressed in the number of recorded localities has recorded for Amaranthus retroflexus, which is also also changed over the periods studied (Table 8). a species recorded among the most freąuent keno The species most often recorded in the mid-19lh phytes of the last 200 years. Conyza canadensis century was Senecio vernalis, but the rate of its and Chamomilla suaveolens are two species, expansion was slow, hence it “dropped” down the presently common in Poland, which have been n = 249 сл CD ОФ & 4 0 □ ч0 — Therophyte ф 35 □ Hemicryptophyte .о 1 30 □ Geophyte z E Hydrophyte 25 ■ Chamaephyte 20 ■ Phanerophyte

10 Н

5 0 J 1501-1800 1801-1850 1851-1900 1901-1950 1951-2000 Historical sequence

в /7 = 300 шс/5 45 фо <Ло . 40

0) 35 Ą -О є 13 30

25 -

20 -

15 -

10 -

5 - 0 і . J 1501-1800 1801-1850 1851-1900 1901-1950J 1951-2000 Historical sequence Fig. 23. Participation of kenophytes of different life forms becoming established in Poland in the historical seąuence 1 5 0 1 - 2 0 0 0

n= 174 є 1000000 — number of species - ■ nurnber of localities g> 100000

10000

1000

100

1 1850 1900 1950 2000 Historical periods

Fig. 24. Changes in species number and cumulative number of localities in the historical seauence 1 8 5 0 - 2 0 0 0 Table 8. The most frequent kcnophytes in the floras of the four historical periods: before 1850, 1851-1900, 1901-1950. and after 1951 For more explanations see the text up to 1850 1851-1900 Numbers of Numbers of Species Species localities localities Senecio vernalis 11 A maranthus retroflexus 169 Amaranthus lividus 9 Marrubium rulgare 147 Acorus calamus 8 Elodea canadensis 140 Amaranthus retroflexus 8 Onobrychis \iciifolia 140 Biyonia alba 8 Galinsoga panńflora 135 Conyza canadensis 8 Xanthium strumarium 130 Oxalis fontana 8 Datura stramonium 128 Marrubium vulgare 7 Senecio vernalis 119 Datura stramonium 6 Bryonia alba 115 Digitalis purpurea 5 Oxalis fontana 111 Medicago sativa 5 Conyza canadensis 106 Reseda luteola 5 Acorus calamus 88

1901-1950 1951 2000 Numbers of Numbers of Species Species localities localities Onobrychis viciifolia 323 Chamomilla suaveolens 13 125 Amaranthus retroflexus 291 Conyza canadensis 11 600 Silene dichotoma 289 Galinsoga parviflora 10 932 Marrubium vulgare 255 Oxalis fontana 8 806 Chamomilla suaveolens 254 Yeronica persica 7 887 Galinsoga parviJlora 253 Amaranthus retroflexus 7 651 Elodea canadensis 226 Robinia pseudoacacia 7 067 Xanthium strumarium 225 Galinsoga ciliata 6111 Geranium pyrenaicum 220 Impatiens parviflora 6 730 Senecio vernalis 219 Medicago satira 5412 Juncus tenuis 206 Solidago gigantea 5 348 Datura stramonium 205 Juncus tenuis 5 332 Conyza canadensis 206 Acorus calamus 4319 recorded at a relatively high number of stations Upland (particularly the Upper Silesian Industrial compared with other kenophytes occurring in Poland. Region), and - above all - the large urban centres of Szczecin, Gdańsk, Gorzów Wielkopolski, Toruń, Poznań, Łódź, Warszawa, Wrocław, Opole, Lublin, Gliwice, and Kraków (Fig. 25). 6. Current types of distribution of Many kenophytes (69 species) occurring in Po kenophytes in Poland land have stations distributed throughout the country and thus they do not represent any par- This analysis of the distribution of kenophytes ticular type of rangę. These are common species was based on data collected for 174 species (cf. (e.g. Chamomilla suaveolens, Conyza canaden Chapter 4, Appendix A). sis, Galinsoga panńflora, G. ciliata, Tancicetum Migrations of alien plant species which have parthenium and Yeronica persica) as well as spread across Poland sińce the end of the 15th species occurring sporadically, sometimes those century have covered the whole national territory species which are freąuent locally (e.g. Oxalis of contemporary Poland. The distribution map corniculata, Physalis alkekengi and Sinapis alba) representing the density of these species through- and rare species that to-date have only been found out the country does not reveal any areas “free” at single stations. This group also includes spe of these newer arrivals (newcomers), but does cies whose stations are concentrated in certain re show that there are regions where they are con- gions, being reported less often in other re centrated: the Yistula river valley, the Silesian gions. Such a mosaic type of distribution results Fig. 25. Concentration of 174 species of kenophytes in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot (BE49 Wrocław) indicates 126 species per unit. For more explanation see the text principally from local habitat conditions. For exam- The distribution of species such as, for example, Acorus calamus, a species common through- ple, Helianthus tuberosus, Hesperis matronalis, out Poland, is less frequently noted in the regions Hyssopus offwinalis and Marrubium vulgare co- lacking habitats specific to this species (e.g. incides with the areas where they are (or have within the Kraków-Częstochowa Upland which been) often cultivated. is dissected by the Vistula and Oder rivers wa- However, detailed analysis of the distribution tershed, or in the Dynów Foreland which is maps pertaining to individual species has permit- a typical farmland area with a limited area of ted the classification of 105 species of kenophytes riverine or lacustrine bank habitats). into groups representing specific types of distri On the other hand, Amaranthus retroflexus, bution ranges in Poland. a species eąually common in Poland, is rare in elevated mountain locations, in north-eastem Poland and in parts of the Kaszubskie Lake District. The 6.1. Kenophytes with stations scattered main limiting factor for the occurrence of this spe throughout Poland except for cies in the Carpathians and north-eastem Poland in certain regions is climate. The aforementioned areas are also characterised by a Iow level of anthropisation of the environment, and they are largely covered by Two groups of kenophytes are classified here forests, wetlands and bogs. which: Fig. 26. Concentration of 12 species of the Sisymbrium altissimum group in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot indicates 12 species per unit. These species are frequent on the whole territory of the country except the Carpathians

- do not enter the Carpathians - Sisymbrium al Lycium barbarum tissimum group (Fig. 26), Padus serotina - either do not oecur or are rare in the Car Portulaca oleracea pathians and in the north-eastem part of Poland Senecio vernalis - Diplotaxis tenuifolia group (Fig. 27). Sisymbrium altissimum Sisymbrium wolgense Xanthium strumarium 6.1.1. Sisymbrium altissimum group These species are mostly those which have been brought in accidentally from south-eastem Europę and western Asia, but less often from central or This type of distribution is represented in Po eastem Asia, or from both Americas. The feature land by 12 species (Fig. 26): common to all these species is that their occurrence is limited to thermophilous anthropogenic Amaranthus chlorostachys habitats (principally various types of waste lands Amaranthus lividus in urban areas or railway tracks, but also within Anthemis ruthenica fields of root crops). Only two species in this Eragrostis minor group occur outside ruderal and segetal commu- Kochia scoparia nities and also in plant communities of semi- -natural or natural character. These are: Lycium been attributed to accidental or purposeful initial barbarum - a shrub found in thermophilous scrub introductions into built-up areas or on railway and forest edge herb communities (it even forms routes (e.g. Amaranthus blitoides, Ambrosia arte a specific community of Lycietum halimifolii), and misiifolia, Atriplex tatarica, Centaurea diffusa, Padus serotina, most often found in pine forests Diplotaxis tenuifolia and Lepidium virginicum). or mixed forests, oak woods and in forest planta- The species originating from North America (Ama tions with a predomination of pines (actually the ranthus blitoides, Ambrosia artemisiifolia and community into which it was originally introduced Lepidium virginicum) have been accidentally by foresters - cf. Chapter 7). brought into Poland from western or Southern The factors limiting the spread of the species of Europę, where they established themselves ear- this group in the Carpathians are, above all, tem- lier (as indicated by the earliest records). perature26 and overall habitat conditions. Even in In Poland, these three species are associated the cases of species whose oldest stations were principally with urban habitats, railway-related found in the Carpathian Foothills (Amaranthus sites and farmlands. Clematis vitalba, Robinia chlorostachys, Anthemis ruthenica and Kochia pseudoacacia and Vicia grandiflora also colo- scoparia), no further expansion in the Carpathians nise thermophilous grasslands and shrublands. was observed; the expansion has been directed The overall shape of the ranges of these species rather into other upland or lowland parts of Poland. is affected primarily by temperature. The species Sporadic penetration into the Carpathians by some are scattered over the entire national territory, species from this group predominantly follows the except for north-eastern Poland (the Southern part main river valleys (of the Vistula, Dunajec and San of the Old Prussian Upland and the Masurian Lake rivers), even though these mountains are generałly District) and higher sections of the Carpathians. rather accessible (Iow elevations, numerous roads and Apart from larger towns, their densities are also raił routes). The Outer Western Carpathians is the lower in north-westem Poland (the Koszalin Coast region into which at least some of the species con- region and Polanowska Upland) (Fig. 27). The cemed will penetrate in futurę (e.g. Padus serotina), climate of these areas, and particularly of the due to the relatively high density of human popula- Masurian Lake District is cooler, compared with tion and intensity of farming, combined with the other parts of Poland, and the vegetation season proximity of the areas of the Silesian Upland which there is the shortest (200-190 days). are already much disturbed by human activities. This type of distribution rangę also partly reflects the differences between the climatic zones of Europę and their associated landscape and vegetation zones. Furthermore, these areas are the 6.1.2. Diplotaxis tenuifolia group least densely populated parts of Poland and are mostly covered by forests. Amaranthus blitoides Ambrosia artemisiifolia Anthoxanthum aristatum 6.2. Kenophytes with scattered stations Artemisia annua over the whole territory of Atriplex tatarica Poland, with concentrations of Bryonia dioica more freąuent stations in some Centaurea diffusa Clematis vitalba regions Diplotaxis tenuifolia Lepidium virginicum Among the great number of kenophytes distribu- Reseda luteola ted throughout Poland, at least three groups of Robinia pseudoacacia species can be selected which show a markedly Vicia grandiflora higher occurrence in the following regions: - south-west Poland (particularly the Silesia- This group is composed of 13 species, originating Cracow Upland) and south-east Poland (partic from south and south-eastern Europę and from ularly the uplands of Southern Poland: North America, which prefer areas with a relatively Małopolska, Lubelska and Roztocze Uplands) warm climate (Fig. 27). Their spread in Poland has - Bunias orientalis group (Fig. 28); - south-west Poland - Geranium pyrenaicum 26 Average annual temperature in the following vegeta- group (Fig. 29); tion zones according to the altitude above sea levcl fluc- - southem and south-east Poland - Echinocystis tuates from (+8°C) +6°C in the foreland zone to 0°C (-2°C) in the alpine zone. lobata group (Fig. 30). Fig. 27. Concentration of 13 species of the Diplotaxis tenuifolia group in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot indicates 10 species per unit. These species are freąuent on the whole territory of the country except for higher parts of the Carpathians as well as parts of the north-eastem and north-westem Poland

The species classified in these groups are further Reynoutria japonica characterised by concentrations within major Rudbeckia laciniata towns: Gdańsk, Poznań, Łódź, Szczecin, Warsza Sisymbrium loeselii wa, and Wrocław. This group includes 11 species originating from south-eastem Europę and various regions of Asia, as well as from North America. They are mostly 6.2.1. Bunias orientalis group kenophytes which have succeeded in establishing themselves not only in synanthropic communities but also in semi-natural and natural ones. Bunias orientalis Possibly the species of indigenous European Cardaria draba origin expanded in Poland using two routes, Echinops sphaerocephalus gradually expanding their ranges from east to Epilobium ciliatum west, and in addition being accidentally trans- Impatiens parviflora ported by long-distance means of transport, most Juncus tenuis often around the main railway hubs. The recon- Lupinus polyphyllus struction of the stages of the expansion permits Parthenocissus inserta the assumption that the latter of the two methods

8 The Establishment. 57 В В

С С

D D

Е Е

F F

А В С D Е F G Fig. 28. Concentration of 11 species of the Bunias orientalis group in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 x 10 km square). The largest dot indicates 11 species per unit. These species are freąuent on the whole territory of the country, particularly in the Southern Polish Uplands was primarily instrumental, particularly in the presence in a given region of the habitats which initial stages of the dispersion of these species. they prefer (Fig. 28). Apart from various types of ruderal habitats, these kenophytes are found in grasslands, meadows, and pastures. However, the species whose homelands are in 6.2.2. Geranium pyrenaicum group distant continents such as eastem Asia or North America, have mostly been carried intentionally into Europę as cultivated plants (Epilobium ci- Geranium pyrenaicum liatum and Juncus tenuis are the only exceptions). Hercleum mantegazzianum They dispersed, colonising ruderal communities Reynoutria sachalinensis near farmland, and - with the passage of time - Rosa rugosa established themselves in shrublands and various Sedum spurium types of forest communities. Silene dichotoma The regions of concentrations of the Bunias Solidago canadensis orientalis group of species reflect the history of Уісіа dasycarpa their spread in the territory of Poland (there Yicia pannonica being concentrations around the oldest sites reported, and additionally they are related to the Fig. 29. Concentration of 9 species of the Geranium pyrenaicum group in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km sąuare). The largest dot indicates 8 species per unit. These species are frequent in most areas of Poland, particularly in the south-westem part

This group includes 9 species of various origins Medicago sativa and manners of introduction concentrated in Solidago gigantea south-west Poland. The density of their sites Thladiantha dubia probably links with the history of their spread. For most of them this history started with an The origin of the species classified into this accidental or intentional introduction into this group is North America (the only exception is part of Poland. The factors supporting their col- the Asian species, Medicago sativa). The only onisation of new sites were essentially climatic species introduced accidentally is Iva xanthiifo- conditions (long period for growth) (Fig. 29). lia; other species were intentionally introduced by humans as useful plants (mainly as omamental or fodder plants). The common occurrence 6.2.3. Echinocystis lobata group of these species in south-east Poland could perhaps result from the fact that they are fairly often only cultivated in this region. Further Echinocystis lobata spread can be facilitated by habitat conditions: Erigeron annuus the presence of river valleys (particularly in the Erigeron ramosus case of Echinocystis lobata, Erigeron annuus ha xanthiifolia and Solidago gigantea) and the existence of hab- Lolium multiflorum itats preferred by the species of the group А ттттттттт

Fig. 30. Concentration of 8 species of the Echinocystis lobata group in Poland The size of dots shows the number of the species occurring in each cartogramme unit (10 х 10 km sąuare). The largest dot indicates 8 species per unit. These species are freąuent in most areas of Poland, particularly in the south-eastem part

(e.g. ruderal sites, particularly in villages and This group includes 5 species originating from smaller towns) (Fig. 30). south-east Europę and south-west Asia, and one species of North American origin {Lemna turio nifera). Ali these species, except for Heracleum sos 6.3. Kenophytes (contemporarily) nowskyi, have gradually extended their rangę reaching their limit of from east to west (Fig. 31), using various routes distribution in Poland of spread. Artemisia austriaca penetrates mainly along railway routes (cf. also Chapter 7), Elshol tzia ciliata has colonised available ruderal sites 6.3.1. Western limit in built-up areas, where it has also been sown (because of the urban-like transformation of Polish villages, this species has lost its old stations in Artemisia austriaca many localities - cf. also Chapter 7). Rumex con Beckmannia eruciformis fertus has used river valleys (of the Bug and Vis- Elsholtzia ciliata tula rivers) in the initial stages of migration only Heracleum sosnowskyi to continue also along transport routes (cf. also Lemna turionifera Chapter 7). The aforementioned Heracleum sos Rumex confertus nowskyi has been intentionally introduced as Fig. 31. Concentration of 6 species of kenophytes in Poland currently showing a western rangę limit The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km sąuare). The largest dot indicates 5 species per unit. The concentration of these species occurs mainly in the Central and Eastern Polish Lowlands (Southern Podlasie Lowland), Polesye and Western Wolhynia a fodder plant into north-eastem and south-eastem Myrrhis odorata Poland, where it continues to colonise areas near Ornithogalum boucheanum the fields where it was previously cultivated. Petmrhagia saxifraga The areas with an evident concentration of spe Picris echioides cies of this group include the middle and eastern parts Rubus armeniacus of the Polish Lowlands (the South-Podlasie Low Rubus laciniatus land), Polesye and the Wolhynia Upland (Fig. 31). Sedum album Sedum spurium Silene conica Solidago graminifolia 6.3.2. Eastern limit Yicia pannonica

This is one of the larger groups (17 species), Anthoxanthum aristatum showing a common type of rangę in Poland. The Corydalis lutea group covers both those kenophytes which have Digitalis purpurea reached their eastern limit of distribution, and the Cymbalaria muralis species which are still penetrating eastwards and Malva moschata for which Poland is a transit area in their further Mimulus guttatus spread (Fig. 32). Most of species classified into Fig. 32. Concentration of 17 species of kenophytes in Poland currently showing an eastem rangę limit The size of dots shows the number of the species occurring in each cartogramme unit (10 x 10 km sąuare). The largest dot indicates 10 species per unit. The concentration of these species occurs mainly in the Sudety Mts. and their hinterlands this group originate from western, Southern or (cf. Chapter 7), and Sedum spurium, as well as south-east Europę. The kenophytes of Asian or two species of a specific distribution rangę type, North-American origin show a similar type of limited to the Sudety Mts. and Western Pomera rangę of distribution as those of European origin. nia: Mimulus gutattus (cf. Chapter 7) and Myrrhis This implies that they have a similar history of odorata, are particularly noteworthy. establishment and further spread in Europę. They are plants which have been accidentally brought into the western or southem parts of the European continent and have then established there. In most 6.3.3. Northern limit cases their spread in Europę continues from west to east. A large group of species arrived in Poland from Germany and the area which is now Chenopodium botrys the Czech Republic. Erechtites hieracifolia In the “Eastem limit” group of species, those Geranium divaricatum associated with the Sudety Mountains: Cymbalaria Helleborus viridis muralis (cf. Chapter 7), Digitalis purpurea21 Lysimachia punctata Oenothera glazoviana 27 Digitalis purpurea occurs also in western part of the Oenothera subterminalis Carpathians. Trifolium patens Fig. 33. Concentration of 10 species of kenophytes in Poland currently showing a northem rangę limit The size of dots shows the number of the species occurring in each cartogramme unit (10 x 10 km square). The largest dot indicates 5 species per unit. The concentration of these species occurs in the Silesian Uplands, Silesian Lowlands, some regions of Sudety Mts. and some of the Carpathians (Bieszczady Mts.)

Yeronica filiformis The centres of distribution of these species in Yeronica peregrina Poland (as well as outside its borders) are also associated with warmer regions. FIowever, Yero The ten species included in this group have nica filiformis, occurring in Poland in mountains a characteristic type of rangę (Fig. 33). As a rule, and foreland areas, evidently avoids a dry cli- their occurrence is limited to one or several re mate. gions of southem Poland (e.g. Erechtites hiera- Among the kenophytes occurring in Poland it cifolia which shows a concentration of sites in the is difficult to distinguish those which while ex- Silesian Lowland and in the Racibórz Basin; panding from the north or north-east, reach the Trifolium patens, recorded from the Carpathian southem limit of their distribution rangę in Po Foothills and in the adjacent area of the Sando land. This results from the fact that only a few mierz Basin, Yeronica filiformis found in the east- species (cf. Chapter 5) have со т е to Poland from em parts of the Carpathians within the borders of these directions. The routes through which most Poland). These are also the species associated of the North American newcomers arrived in with a specific type of habitat (e.g. Erechtites Poland most often lead through western and hieracifolia is found principally on clearings and southem Europę and not - as one would expect forest edges; Trifolium patens and Yeronica fili - through sea routes from the Baltic Sea. There formis grow principally on moist and moderately is an ехатріе of one species (Beckmannia eru- moist meadows). ciformis) whose proliferation across Poland has probably occurred from the Baltic coast towards Acer negundo the central regions (at least in the early stages of Bidens frondosa the spread). The only species now more abundant Clematis vitalba in the northern part of Poland are Bidens conna- Diplotaxis tenuifolia ta and Oxalis dillenii. Echinocystis lobata Eragrostis albensis Erigeron annuus Oenothera depressa 6.4. Kenophytes associated with river Oenothera x hoelscheri valleys Rumex confertus Salsola kali subsp. ruthenica Solidago canadensis A dozen or so newer arrivals now established Solidago gigantea in Poland manifest an affinity with river valleys. Xanthium albinum The valleys have provided (and still do provide) Xanthium spinosum migration corridors used by alien species in the course of their progress into a new territory. This type of distribution is principally condi- The association with entire river valleys or their tioned by the biological and morphological fea- parts characterises the following kenophytes: tures of the species concemed. In their respective

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Fig. 34. 15 species of kenophytes in Poland currently showing a concentration along the main river valleys (i.e. the riparian corridor plants) The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot indicates 15 species per unit homelands they are also closely associated with In their originał distribution ranges these plants habitats and communities close to rivers (riverine are also associated with river valleys and their woods and shrubs, reed or rush communities, specific habitats along major rivers: sand/mud therophyte communities on sand and gravel alluvia (e.g. Eragrostis albensis) and sand steep alluvial substrates) and they take advantage of banks and scarps (e.g. Oenothera depressa, Sal the pioneering conditions created by rivers sola kali subsp. ruthenica mdXanthium albinum). (alluvia, valley edges, river bank cliffs) (Fig. 34). Their migration and continuing invasion of still further territories in Poland is closely linked with In this group, an additional sub-group can be habitat conditions provided by large rivers. Both the distinguished of 6 species specific to the valleys Vistula and Bug rivers are regarded as still only of two large rivers of Poland: the Vistula and Bug slightly disturbed by humans, and the dynamie and rivers (Fig. 35). These are: diverse natural processes present in this environment support plant migration. Additionally, some Eragrostis albensis anthropogenic factors (river engineering of some Oenothera depressa stages, location of settlements and towns in river Oenothera x hoelscheri valleys, transport routes Crossing rivers, etc.) facil- Rumex confertus itate the migration of plant species both along and Salsola kali subsp. ruthenica across river valleys. These conditions are used by Xanthium albinum alien species which implement the subseąuent

Fig. 35. 6 species of kenophytes in Poland currently showing a concentration specifically along the Vistula and Bug river valleys (i.e. the riparian corridor plants) The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot indicates 6 species per unit

9 The Establishment. 65 phases of their invasion along the river valley (pen- Ambrosia artemisiifolia etrating into a new territory through a corridor Ambrosia psilostachya created by a river). At least some of the species next Atriplex tatarica arrive in adjacent areas, taking over other habitats Amaranthus blitoides (e.g. Rumex confertus and Salsola kali subsp. ru- Cent aurea diffusa thenica - cf. also Chapter 7) or follow the reverse Eragrostis minor course moving from ruderal habitats into riverine Euphorbia humifusa ones (as was probably performed by Oenothera de- Iva xanthiifolia pressa and Oe. x hoelshen*). Linaria repens Melilotus wolgica Oenothera paradoxa 6.5. Kenophytes associated with urban Oxybaphus nyctagineus Parietaria pensyhanica areas and railway routes Potentilla intermedia

Ailanthus altissima This group is represented by 16 species, mostly Amaranthus albus introduced accidentally (less often introduced

Fig. 36. Concentration of 15 species of kenophytes in Poland currently showing an association with urban areas, railways and roads The size of dots shows the number of the species occurring in each cartogramme unit (10 * 10 km square). The largest dot indicates 14 species per unit

28 Oenothera x hoelsheri is a hybrid resulting from hybrid- occurring in Poland both on sandy wastelands and on cliffs isation of Oe. biennis or O. rubricaulis (thus species freąuently along rivers) with North American species Oe. depressa. intentionally) with consignments of cereal grain, 7.1. The history of the spontaneous poultry fodders, soya-beans, oil crops, wool, or spread of cultivated woody plants with garden materials (including those introduced as the result of “domesticating” to botanical gardens) and with ballast. Even species though they originate from various parts of the głobe they share a preference for warm and dry habitats. In their respective homelands they usu- Acer negundo L. [syn.: Negundo aceroides Moench; ally grow in steppes (Atriplex tatarica, Centau- N. fraxinifolia Nutt.; Negundo negundo Karst.] rea diffusa, Melilotus wolgica), prairies and Вох-elder; Ashleaf Mapie deserts (Amaranthus albus, Ambrosia artemisii- Aceraceae folia, Iva xanthiifolia), but also in dry anthropogenic habitats. Biology: woody dioecious plant, anemogamous. In Poland and in the rest of Europę, they occur Winged fruits dispersed by wind, seeds germinate - outside their natural rangę - in the regions with easily. May also spread by suckers. sub-continental climatic features or in habitats Native rangę: North America, where it is one of which meet their ecological requirements, such the most common American mapie species as roadsides, wastelands, and railway embank- (H itchcok et al. 1961). Its rangę extends from the ments. eastem seaboard to the west coast, whilst to the They have found particularly suitable condi- north it reaches Canada and to the south, Gua- tions for development in urban locations and on temala. It has a continuous distribution reaching railway embankments. These sites have provided Califomia to the south-west, Alberta to the north, the stepping stones for their repeated leap- Massachusetts to the north-east, Florida to the frogging into new areas. The distribution of south-west and New Мехісо to the south (L ittle species in this group reflects the location of urban 1971; Scoggan 1978). In its native habitats it centres (particularly large metropolises) and the grows in humid and wet areas along the banks of network of railway routes (Fig. 36; cf. also water bodies, being a dominant component of Chapter 10). humid forests in some areas (M ohlenbrock & V oigt 1959). Secondary rangę: Eurasia reaching as far as western Siberia (Adamowski 1995), w ith the 7. The history of the spread highest concentrations o f stations in Central of selected kenophyte species Europę. Outside P oland it has spread in Saxony, in the territory of Poland Thuringia, in Austria, Czech Republic and Slo- vakia, France, north-eastem Slovenia, in northem and central Italy and in the south-eastem part For the 25 species of kenophytes the probable of the British Isles (Lohmeyer & Sokopp 1992; course of spread within the territory of Poland Bocker & Dirk 1998; B e n k e rt et al. 1998; has been reconstructed and the stages of their ex- Hardtke & Ihl 2000; UherćikovA 2001; Pyśek pansion documented by means of maps. Out of et al. 2002; S ta c e 1997; N ejc 2001; P ig n a tti the group of 174 species for which detailed in- 1982). It is especially abundant along the tribu- formation has been collected to-date (Appendix taries of big rivers (on the Rhine, Dubai, Vistula A), examples of groups of species which have and in southem part of the continent on the Sawa) different biology, origin and manner of introduc- and in cities, e.g. in Warsaw (Sudnik-Wójcikow- tion include: s k a 1987a), Rome (Celesti Grapow 1995), B er - cultivated woody plants with a rangę of lin (Kowarik 1992), Uzhorod (Protopopova & different origins, S h ev era 2002) and Donetsk (Burda 1997). Its - cultivated herbaceous plants with a rangę of current widespread introduced rangę can be at- different origins, tributed to its use on a mass scalę, as a tree grow n - plants accidentally introduced with a rangę of in parks and along boulevards in the 19th and the different origins. first part of the 20th century. In some European The following additional criteria were em- countries it is considered to be an invasive spe ployed: cies (cf. Appendix A). - time of intentional/accidental introduction, History of spread: - types of habitats colonised, Europę: introduced as a decorative plant in 1688 - current status (established or invasive), in the Fulham Garden in England (W ein 1931). - abundance of floristic data. Subsequently it was introduced into the Nether- Introduction and initial phases of colon isation: turn of the 18lh century: flrst plantings in parks and gardens ® first presumably spontaneous localities of occurrence: Wrocław BE49 (BAENITZ, herb. PRC, W, WRSL); Puławy FE03 (BERDAU, herb. LBL) o Kraków DF69 (Boehm 1873) - dubious record: the author did not register the status of the species in this locality; most probably the record refersto a locality fromcultivation naturalisation and spread close to sites of cultivation

Start of invasion phase: transition from ruderal habitats to riverside poplar-willow forest habitats and occupation of "bridgeheads” in river valleys main directions of spread simultaneous further spread near sites of r cultivation and formerly occupied localities as well as migration “out of river valleys" on to ''*■ adjacent anthropogenic habitats

Subsequent phases of invasion: rangę increase and stabilisation by: - migration along rivervalleys - colonisation of further ruderal habitats (fallow land, urban wasteland, railwayterritory) (the map after Z ając A. & Z ając M. (eds.) 2001 - slightly supplemented)

A local example of riparian corridor migration of Acer negundo inthe Bug rivervalley (source: Fauński ełal. 2000)

Fig. 37. Recorded history of the spread of Acer negundo L. in Poland - an example of a species which uses river valleys as spreading corridors lands (1690), Germany and Czechoslovakia Padus serotina (Ehrh.) Borkh. [syn.: Prunus se (1699) (W e in 1931; L o h m e y e r & S u k o p p 1992) rotina Ehrh.] and in Hungary (1872) (B a l o g h 2001). Rum Cherry Poland: probably introduced at the turn of the Rosaceae 18th and 19th centuries (Kornaś 1968b). It is known that the speeies was introduced to Cracow Biology: a tree reaching heights of up to 20 m; in Botanical Garden in 1808 (Hereźniak 1992). Europę usually of shrub-like form. Flowers in race- Subseąuent occurrences of this speeies have been mose inflorescences, pollinated by inseets. Drupe- reported in parks in Krzemieniec (1810) and in type succulent fruits with a fleshy pericarp dispersed Niedźwiedź near Cracow (1813) (Seneta 1994). by fructivorous birds and some mammals. Initially it was planted deliberately, as a fast grow- Native rangę: central and eastem part of North ing tree. The earliest occurrences of this speeies America (Ontario and Quebec and southwards to refer probably to stations where it was first intro Texas and Florida) where it grows in woods and duced, e.g. Sznabel, near 1880 herbarium mate- clearings, floodplains and thickets by roadsides rials, WA - Warsaw gardens (after Sudnik- (C r o n k & F u l l e r 2001) and the northem part of Wójcikowska 1987a). For this reason, an accu- South America (from Мехісо to Guatemala). rate determination of when the speeies first Secondary rangę: central Europę, above all the occurred in the wild is difficult. Undoubtedly, the Netherlands, south-eastem France, Germany, Po stations recorded after World War II are sponta- land and some regions in Austria; reported also neous (Fig. 37). The tree is still grown along in northem Italy, Hungary, Romania, Czech Re- roads and in parks due to its undemanding hab public and England (S t a r f in g e r 1997). itat reąuirements and resi stance to drought and History of spread: frost. Europę: belongs to the earliest tree plants Habitats: willow-poplar carrs, broad-leaved or brought to Europę from North America. 1623 or mixed woods, pine-oak-birch stands and forest 1629 is cited as the oldest date of the introduc- plantations, also anthropogenic habitats: fallow tion, when the tree was grown in the Paris area lands, roadsides, near cottages, rubble heaps, (S t a r f in g e r 1997). Initially grown as a decora- walls, refuse tips, neglected parks and gardens, tive tree in parks, sińce the late 19lh century it has hedges, cemeteries, lawns, urban wastelands, been applied in forestry (such applications as tramway tracks, railway tracks and embankments wood production in poor soils or enriching the and industrial wastelands (spoił heaps and humus layer in forest plantations, especially of sedimentation ponds). coniferous trees). In the first half of the 20th Dynamics: although this speeies has been re century, and in the 1980s, it was planted on a corded in Europę for more than 300 years, it has large scalę in the Netherlands, Germany and in undergone an evident invasion only within the Poland. The first spontaneous stations of this last 100 years, and in Poland only for the last speeies were recorded in a relatively short time 50-60 years. In some regions of Poland from its introduction, after ca. 30 years (K o w a r ik (Wielkopolska) the expansion of this speeies has 1992). Currently, in a number of countries it is been recorded only in the last 30 years considered an invasive speeies entering natural and semi-natural habitats, including protected (Ż u k o w s k i et al. 1995). Currently, it is common in most of the territory of Poland (more than ones (C r o n k & F u l l e r 2001). 3500 stations in 1379 ATPOL sąuares), but rarer Poland: for a long time cultivated in parks and in the north of Poland (Western Pomerania, gardens as a decorative tree, ąuite often planted in Kuiavian region - Kujawy), particularly rare in forest as undergrowth, and subseąuently sowed by birds. In 1813, it was recorded in the collection of the north-east (Warmia and Mazury) and at the garden in Niedźwiedź near Cracow (H e r e ź n ia k higher elevations in the mountains (Tatra Mts., 1992). Although the oldest dates recorded in con- Bieszczady Mts.) (Fig. 37). Reported by K o r temporary Poland only go back to the late 19th n a ś et al. (1996) from the Western Carpathians century (Fig. 38), it may be judged that the spe- as a speeies established in riverine carrs cies started spreading before that period. This although only occurring rarely. assumption is supported by dates referring to east The distribution of the Ashleaf Mapie in Po em Germany, when it was introduced to cultiva- land has a characteristic feature in that it reflects tion in 1796, and the first “wild” station was re the courses of major river valleys (Ż u k o w s k i et al. corded in 1825 (K o w a r ik 1992). In addition, the 1995; F a l iń s k i et al. 2000; Z a ją c A. & Z a ją c M. localisation of subseąuent stations recorded in 2001) (cf. also Chapter 6). It is currently invad- Poland (north-west and south-west of Poland) in ing new sites. an area belonging at that time to Germany, allows Introduction and initial phases of colonisation: tum ofthe 18m century: first plantings in parks and gardens ® first presumably spontaneous locality of occurrence: Bydgoszcz CC26 (Bock 1900)

Naturalisation: naturalisation and spread near sites of cultivation; numerous introductions in cultivated forest plots in many regions of the country have contributed to the naturalisation of this species y - spontaneous spread from sites of cultivation

lnvasion: massive introductions (performed as a part of forest management plans) and simultaneous rapid (for a tree species) unaided spread (the fruits are dispersed by birds), which have jointly led to the occupation of the major part of the country within a period of 50years local rangę limit

(the map after Zając A. & Zając M. (eds.) 2001 - slightly supplemented and modified)

Fig. 38. Recorded history of the spread of Padus serotina (Ehrh.) Borkh. in Poland - an example of a species which owes its naturalisation in the new homeland to man and birds for presumptions on the first stages of the ехрап- Tree-of-heaven, ailanthus, Chinese sumac, stinking sumac sion of this species in Poland and for formulation Simaroubaceae of the hypothesis that the species spread mainly in Poland from west to east, and around sites Biology: tree with polygamous flowers, usually where the species was cultivated and introduced. unisexual, growing rapidly and producing great Habitats: oak-hombeam woods, pine forests and numbers of seeds. Fruits setting as early as be- mixed coniferous forests, pine and oak-pine stands. tween the 10th and 15th year of life. Winged fruits Dynamics: the species has staged a rapid expansion dispersed by wind and water. Capable also of re- in the last half-century, the process being facilitated production by suckers. by foresters who simultaneously introduced it into Native rangę: north-eastem China many forests. Currently, it occurs throughout Poland Secondary rangę: Europę, especially its South except for the Carpathians, rarer also in north-eastern part. Currently, a species is naturalised in the em regions (recorded in 2564 stations in 1134 Mediterranean area, where it spreads from urban- ATPOL sąuares) (Fig. 38; cf. also Appendix A). ised areas along roads, also entering maąuis. In central Europę its spontaneous stations are con- Ailanthus altissima (Mili.) Swingle [syn.: centrated mainly in cities with specific climatic A. glandulosa Desf.] features, for ехатріе in London, Prague, Berlin,

Introduction: tum of the 19th century first plantings in parks and gardens

initial phasesof spread: spontaneous spread from cultivation sites, exclusively near locations where itwas planted

Sites of cultivation of Ailanthus altissima in Poland (source: Pacyniak 1976)

Fig. 39. Recorded history of the spread of Ailanthus altissima (Mili.) Swingle in Poland - an ехатріе of a species making use of urban “heat islands” in its naturalisation process Dresden, Leipzig and other German towns, and on neglected lawns, in hedges, on tramway or in Uzhorod (K u n ic k 1990; S t a c e 1997; H a r d t k e railway tracks and on refuse tips. Outside urban & I h l 2000; P r o t o p o p o v a & S h e v e r a 2002). areas, single stations have been recorded in open Apart from Europę, the synanthropic rangę in- oak-hombeam woods (Ż u k o w s k i et al. 1995) and cludes also Australia, the south-eastem part of beech woods (T o k a r s k a -G u z i k , pers. obser.), North America, and Central and South America where it regenerates both through vegetative and (L a u e n e r 1996). generative processes. Dynamics: In the most recent 20 years the num History of spread: ber of stations in Poland has increased from 6 to Europę: introduced to Europę by Jesuit Pćre 28. The tendency to spread is above all evident d’Incarville in mid-18th century. Introduced into in large towns, and its sustenance or possible in- Great Britain in 1751, by Peter Collinson who re- vasion of new sites will depend principally on cli- ceived the seeds from Pere d’Incarville (Lauener matic factors. The species is not fully frost resis- 1996). At the same time (1760) the tree was also tant and long not yet lignified annual shoots brought into Italy, to the botanical garden in Padua freeze during severe winters. The initial stages of (Pignatti 1982). Subseąuent introductory dates expansion are currently being observed in Poland. are cited by L o h m e y e r & Sukopp (1992): 1780 for Central Europę and Pyśek et al. (2002) who re- Clematis vitalba L. corded 1874 as the flrst date of the occurrence of the species in the area of what is now the T ra v e lle r’s-jo y Czech Republic. In 1902 it was recorded in the Ranunculaceae wild in Germany (Kowarik 1995a). After the Biology: strong climbing plant with shoots up to World War II Ailanthus altissima colonized mins 10 m long; fmits - achenes with flight appara- of bombed towns. For ехатріе, it was recorded tus, which consist of the style elongating after in Berlin (S c h o l z 1957), Stuttgart (Kreh 1955), fertilisation, covered with feathery hairs. and in Poland in the town of Wrocław (Prof. K. Native rangę: central, western and southem Europę Rostański, pers. comm.). (to the north it reaches the Netherlands; in the Due to its adaptive capacity to grow in dry hab- British Isles it is considered a native species in itats, in heavily polluted areas, it was grown along- Wales and southem England (Stace 1997)). Also side other trees in many European and American occurs in northem Africa, Asia Minor and the coast cities. The unpleasant smell of the staminate of the Caspian Sea (Gostyńska-Jakuszewska 1985). flowers growing on separate trees resulted in Secondary rangę: Southern Australia, New a number of trees in Paris and American cities being Zealand, North America; in Europę naturalised in cut down in the early 20* century. Apart from its Ireland, Scotland, Germany, Denmark and in Po natural rangę, once introduced, it rapidly colo- land. Apart from its natural rangę it has the status nises unusable land and all free areas, especially of a widely spread invasive species in many in towns where a hot climate prevails. In many countries, posing a threat to the natural vegeta- cases it becomes a “pest tree” (L a u e n e r 1996). In tion (Cronk & Fuller 2001). Italian towns it is currently one of the most fre- History of spread: quent species of foreign origin (C e l e s t i G r a p o w Europę: used as a decorative climbing plant in & B ł a s i 1998; C e l e s t i G r a p o w et al. 2001). palaces and garden establishments, often spreading L a n d o l d t (2000) describes the rapid expansion of into the wild from these places. For ехатріе, it was this tree in Ziirich, where it was not invasive recorded in Germany as a cultivar in 1663 and 20 before 1980 (in 1988 it was recorded in 29 studied years later (in 1883) in the first “wild” station sąuares, and after 10 years it occurred in as many (K o w a r ik 1995a). After the World War II, the Trav- as 66 sąuares). In warmer Slovakian areas it eller’s-joy occurred in the mins of bombed towns, occurs along the Dubai River, migrating from for ехатріе in Canterbury (UK) (K e n t 1951), in ruderal habitats to forest boundaries (U h e r ć ik o v A western Germany (K r e h 1955), in Gdańsk 2001). Considered as noxious and widely spread (S c h w a r z 1961) and in Wrocław (both towns in “pest” (F e r n a l d 1950; C r o n k & F u l l e r 2001; Poland) (Prof. K. R o s t a ń s k i, pers. comm.). P y ś e k et al. 2002) (cf. Appendix A). Poland: brought into Poland as early as in the Poland: brought to Poland in the early 19th cen 17th century, or even earlier. S ir e n iu s ( S y r e ń s k i tury. Became established in cultivation through- 1613) and subseąuently K l u k (1786) report the out most of Poland, excluding its eastern and Traveller’s-joy as a plant cultivated in Poland (cf. (P a c y n ia k north-eastem part 1976). It has spread Appendix A and Chapter 5.2). Ł a p c z y ń s k i (1889) spontaneously in recent years, in cities where it describes this species as spreading beyond its was previously planted (Fig. 39). managed confines and occurring along the Vis- Habitats: saplings and young specimens usually tula River (Solec, Janowiec, Kazimierz) or “tend- grow under walls in cracks between flagstones, ing to be naturalised”. А В С D 6 Iі О ...... ТТПТТvm г------Г - — т т ------ГГГ— ------~ 7~ГГГГ—г------T T m ггуггтгт "n " 'Т^ТТТТТ Г|

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Subsequent phases of spread: The current distribution reflects the link of this species with rangę increase and stabilisation by colonisation of suitable individual segments of river va lleys (see also Cha pter 6) as well habitats in river valleys and ruderal habitats: aswithwarm semi-natural and anthropogenic habitats: ■f occupation of new localities, especially in the south- the current north-eastern rangę limit of this species is western part of the country delineated by the river valleys of the Vistula and the Bug rivers Fig. 40. Recorded history of the spread of Clematis vitalba L. in Poland — an ехатріе of a species escaping from “romantic” gardens

In subseąuent years this species was recorded tracks and embankments; around garden allot- as growing “wild” near places of cultivation ments and in neglected historical parks. This is across the middle section of the Vistula River and a characteristic species of communities of the Rham- in the western and south-western part of Poland. no-Prunetea class, locally also of the Pruno-Ligus- Habitats: sunny slopes with thermophilous ve- trietum association (Matuszkiewicz 2001). getation, forest edges, principally oak-hombeam, Dynamics: sińce the first records of occurrence, stony sites (Gostyńska-Jakuszewska 1985); also the number of records built up slowly till the on ruderal sites: near cottages, wastelands, railway mid-20th century, whilst a more striking increase in

10 The Establishment. 73 records appeared only after 1950 (Fig. 40; Appen- species is naturalised in many regions where it dix A). Currently, the species occurs freąuently in occupies such habitats as cracks in walls, pavements Western Pomerania and Lower Silesia, reaching and stony and regulated (covered with bricks) river eastwards to Puławy, Kazimierz on the Vistula and banks. Chełm (Fig. 40). Recorded on a single station in Poland: the determination of when the species the Śnieżnik mountain massif (S zeląg 2000). arrived in Poland is difficult (Z a ją c E.U. & Z a ją c A. Recorded to-date from 354 stations in 216 ATPOL 1973). The first citations go back to the first half sąuares. It is currently invading new sites. o f the 19* century, and further more numerous dates go back to the second half of that century (Fig. 41). The first stations for the Ivy-leaved Toadflax were recorded in the Sudety Mts. and in north-westem 7.2. The history of the spread Poland. There is no certainty that the initial dates of useful herbaceous plant refer to plants which had moved into the “wild” and species: how medicinal and naturalised or whether these records refer only to decorative plants have established cultivated plants (Z a ją c E.U. & Z a ją c A. 1973). themselves in the flora Undoubtedly, as stated by the authors referred to above, Cymbalaria muralis spread spontaneously after 1870. A number of authors attribute this pro- 7.2.1. Examples of species of European cess to the plant spreading from sites of cultivation. origin Other authors (Ś w ie r k o s z 1993) State that the expansion of its rangę can be linked to its migration along river valleys. Recent studies conducted in Cymbalaria muralis P. Gaertn., B. Mey. & Lower Silesia support the hypothesis of the anthropogenic origin of the majority of stations of this Scherb. [syn.: C. cymbalaria Wettst.; Linaria cymbalaria (L.) species (Szczęśniak & Świerkosz 2003). M ili.; Antirrhinum cymbalaria L.] Habitats: occurs in secondary habitats, above all Ivy-leaved Toadflax on old walls, less ofiten on rubble, on roadsides and Scrophulariaceae railway tracks and embankments. A species which indicates the Potentillion caulescentis order o f Biology: hemicryptophyte, capable of anchoring crevice-related communities on fairly well-lit lime- on vertical walls, owing to stolons and roots grow- stone substrates (Matuszkiewicz 2001) and dom ing at leaf-bases. After blossom is shed, the pedicel inant in the Cymbalarietum muralis community. elongates and through a negative heliotropism Dynamics: in Poland the species occurs in the mechanism pushes the fruit into the substrate (e.g. Sudety Mts., in Silesia, Pomerania, Mazovia and into a crack in a wali), being an ехатріе of geo- W ielkopolska at 350 stations registered to-date in carpy (B u l iń s k i 2000); the species disperses also 165 ATPOL sąuares (Fig. 41, Appendix A). The through anemo- and anthropochory. number of stations of this species, after an evident Native rangę: southem south-westem and Europę: increase noted in the decades from 1960 to 1980, the Southem Alps, the Dinaric Mts., central and has not maintained this kind of strong tendency. Southern Italy and Sicily (Webb 1972) where it Furthermore, many of the earlier recorded sta grows in rock cracks. It has been also reported in tions have not been confirmed recently, which North Africa and western Asia (W ojewoda 1963). could suggest a gradual retreat of the species. Secondary rangę: central and northem Europę According to S z c z ę ś n ia k and Ś w ie r k o s z (2003), as far as southem Scandinavia; in Eastem Europę this fact should be attributed to the progressively in St. Petersburg; in Ukrainę (F edorov 2001; rarer cultivation of this plant in Poland, as well Protopopova & Shevera 2002). as to the intentional removal of plants from old History of spread: walls during restoration measures. B u l i ń s k i Europę: probably a cułtivar already grown in many (2000) even indicates the necessity for protect- regions of Europę by the early 18* century. Accord- ing its scarce stations in the Pomerania region. ing to Świerkosz (1993, after Segal 1969), this spe cies started migrating from natural habitats in lime- Digitalis purpurea L. stone rock in the Mediterranean basin in ancient Foxglovc times. In central Europę it was recorded in the 17th Scrophulariaceae century, dispersing slowly along the valleys of large rivers, e.g. the Rhine. Lohmeyer & Sukopp (1992) Biology: biennal or annual plant29, characterised by ąuote 1644 as the oldest date for the occurrence of very high production of fine seeds dispersed by wind. the species outside its native rangę in the Nether- lands. An even earlier date, namely 1640, is ąuoted 29 In the original distribution rangę this plant is either by Stace (1997) for the British Isles. Currently, this biennal or perennial (H a n t z 1993; S t a c e 1997). A B c D E F Q i ■

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AB c D EFG Subsequentphases ofspread: The current distribution of this species is concentrated rangę increase and stabilisation mainly in the region of the Sudety Mountains (see also occupation of new localities in the region of their initial Chapter6): concentration a direction of expansion appearing distinctly in the further spread ofthe species east and north-east upthe 1980s Vistula rivervalley . a tendency towards gradual loss of localities (localities not confirmed in current studies in the Lower Silesia by Szczęśniak & Świerkosz 2003) (the map after Z a ją c A. & Z a ją c M. (eds.) 2001 - slightly supplemented and modified) Fig. 41. Recorded history of the spread of Cymbalaria muralis P. Gaertn., B. Mey. & Scherb. in Poland - an example of a species reąuiring specific habitats for naturalisation

Native rangę: south-western, western and central land with isolated stations in the south-western and Europę, including mountainous areas of southem western part of the Scandinavian Peninsula. In cen Europę: Sardinia, Corsica and the Pyrenees. To the tral Europę it is more scarce, occurring primarily north it reaches the British Isles and southem Ice- in Austria, Switzerland and Germany (H antz 1993). Secondary rangę: North America, Southern many and several isolated places in Poland as Australia, New Zealand and North Africa. Ac- areas of the synantrophic occurrence of this cording to M eusel et al. (1978), the so-called species as a kenophyte (neophyte). These au- potential limit of the occurrence of this species thors treat the whole area of Poland to Finland in Europę reaches the eastem limits of the con- in the north and Romania and the former Yugo- tinent. Commonly cultivated and going into the slavia to the south, as potentially an area where wild state in the European part of Russia, on the this species occurs as ephemerophyte. R adwań- south-eastern coast of the Baltic and Crimea ska-P aryska (1950) considered the possibilities (Fedorov 2001). of the artificial origin of the station of D. pur History of spread: purea in the Tatra Mts. and the following hypo- Europę: a plant cultivated for a long time in thesis. She thinks that the distribution of the Europę as a decorative or medicinal plant, includ- Foxglove shown in the map of its rangę suggests ing beyond its natural rangę; entering the wild that the Polish stations could be the eastemmost state in some areas. In the area of former Czecho- outposts of this Atlantic plant. As the climate slovakia it was recorded for the first time in 1790 dried, its rangę might have retreated westwards. (Pyśek et al. 2002). Relict islands remain in Poland in areas with Poland: Cyunel described the distribution of sufficiently high humidity, i.e. above all in this species in Poland as early as in 1965. She montane areas affected by the ocean. presented stations in the mountains, as opposed According to Filinger (1992), the conditions to those in lowlands, as those that most corre- occurring in the Baltic coast region meet these sponded to natural ones. The historie records conditions: he found the presence of the Foxglove from Cracow area provided by Besser (1809), in forests similar to acidophilous beech forest, in Dembosz (1841), B erdau (1859) or Rostafiń the area of the Słowiński National Park. However, ski^ stations (1872) in the Warsaw area were these hypotheses have not been satisfactorily probably ephemeral. The Sudety Mts. and the justified, and it is best understood as an undoubt- Beskid Śląski and Mały Mts. are the primary edly alien species in Poland’s flora dispersed and oldest areas of Poland where the species has from areas of cultivation30. spread (Fig. 42). The plant was already known Habitats: open spruce forests, scrub, felling in these areas in the second half of the 19th sites and windfalls, also grasslands and dwarf century (Hantz 1993). The station on the Klim mountain pine scrub as well as anthropogenic czok mountain, in the springs of the Biała River habitats. In the Sudety Mts. and Beskidy Mts., in the Beskid Śląski is considered the oldest the species forms its own association, Digitali- one in Poland. After that, the species dispersed Epilobietum belonging to the Epilobietea angus- in the Beskid Mały, Tatras, Karkonosze, Orli- tifolii class, for which it is a characteristic spe ckie, Złote and Bystrzyckie Mts. (Radwańska- cies (Wożakowska-Natkaniec 1985; H an tz Paryska 1950; Stecki 1952; P elc 1958; Kucowa 1993). This association occurs in felling sites 1963; C yunel 1965). Radwańska-Paryska left after acidophilous beech forest and fir- (1950) and Cyunel (1965) made attempts to -spruce in the lower montane zone. solve the issues of the origin of the station in Dynamics: in Poland, most of the stations of this the Beskid Mts. It is supposed that seeds of species are concentrated in the western and south- D. purpurea moved to Lower Silesia together western parts of the country (Sudety Mts., Beskid with transported spruce seeds (Cyunel 1965 after Śląski Mts. and Beskid Żywiecki Mts.) (Fig. 42). Bukowiecki 1950). They may also originate In the Karkonosze Mts., the species was recorded from the mountain plant garden located on the as early as the 19th century (Fiek 1881; Schube Klimczok. Remaining stations in Poland, includ- 1903b, 1904), and occurs there at the elevation of ing those from Wielkopolska and the Baltic 850 m a.s.l. (Rostański K. 1977); in the Tatra Mts. coast have their origin in cultivations in gardens - between 1190 and 1240 m a.s.l. (Radwańska- located near houses (Szulczewski 1951; H antz Paryska 1950). Some of the isolated stations in 1987; F ilin ger 1992). It is often cultivated as eastem Poland still retain their ephemeral charac- a decorative and medicinal plant and then ter. The species has to date been recorded on 341 progresses into the wild state, or simply disperses on its own from its previous stations to new areas. 30 This is a difficult plant to assess because it charac- In such areas the decision on the nativeness of teristically appears after disturbance (e.g. felling) inereases the species is difficult (Filinger 1992). M eusel light in woodlands. Once the site goes back to normal, Digitalis purpurea retreats into the seedbank, where it can et al. (1978) consider south-western Sweden, persist for many years without being seen as abundant Denmark, the Netherlands, a part of eastem Ger minutę seeds (G r im e et al. 1988). , , n T i n : rm ■ ■ — ...... TTTTITTTr -iin m i ’ itt m n T iTTHTm , rrn ---!- T) ^ * J s" i w > l v . A X -- -- 7'W - 1. - r s / -A 4 i K \ /

'W Y I? ' ; ______\l f V L \ ^ \ J ) V ? \ i—/ . . 4------) J ____ ... -

i "XX, i : c \ \ __ N V -~ ^ C \ \ J « « l ® ® 1 ------X ____ S' - i ^ ------J \ X \ } \ 1872 \ r ~ (v (...... { ( { ( L,, L \ ( i--. \ k L \ ( - >\ A. X, i \ )\ \ J \ nj ' . : J \ / \ v. ^ X i \ / ^ 'i ^ 1872 > ''-0? 1809? i ; ■< ) X"'' A V , . oa& ) J/ I \ / r~ - - < H- v \ '' m l 59 \ \ - ! V i 18 r' 1 8 5 0 V. ' \ <' 1 9 0 0 ) ) y { ) / r L l l l i A l ł l ...... Yrnn ...... M.A.M

First recorded localities ofoccurrence: Start ofspread: naturalisationandspreadnearsitesofcultivation 0 historical localities from the vicinity of Kraków (Besser 1809) ° localities from the vicinity of Kraków: DF66-69, 78, 79, EF70 (Dembosz1841; Berdau 1859; Rostafiński 1872), Warszawa ED26 (Rostafiński 1872) and Wągrowiec CC50 (Nowicki 1885)thatwere ephemeral in character (escapesfromcultivation; seethetext) ® the oldest localities in the Silesian Beskid the appearance of the species in this regions is liked to its accidental Import with spruce seed or escape from garden cultivation (mountain plant garden atop the mountain of Klimczok)

IM1M m as#ł \ T ^ . > V c » . i • • V \ Ń / • \L • • ■ •> : •• • \ K * - ~ > ( • ~ X ; . • 1 \ • . 1 “S h - i < • • < i ;; •s • • * ¥ L • * * •• • • v a . . •• • / • ) \ = / \ < H T r * , . •r'J / y • hi i \ i Y: l ^ f —N & • t /V <1 cH ( 2000 l z k *r } / L „iirtinn ii 11 i^i ii.±LLLUXU. c Subsequent phases ofspread: The current distribution of the species includes mainly rangę increase and stabilisation the regions ofthe Sudety Mountains, the western Carpathian spread and rangę stabilisation in the Beskidy Mountains and the Baltic Coast, where the appearance of y and Sudety mountain ranges localities of its occurrence may be linked with accidental importation and further spread - with spontaneous rangę further spread of the species east and north-east expansion (see also Chapter6): \ (here the main source of its diaspores are sites ofconcurrentcultivation) areasofconcentrationoflocalities probable direction of arrival of this species probable direction of further spread in Sudety Mts. (the map after Zając A. & Zając M. (eds.) 2001 - slightly supplemented and modified)

Fig. 42. Recorded history of the spread of D igitalis purpurea L. in Poland - an example of a subatlantic species enlarging its rangę of occurrence in an easterly direction stations in 169 ATPOL sąuares (cf. Appendix A). 7.2.2. Ехатріе of species of Asian In many regions of Poland there has been renewed origin interest in this plant as a decorative plant to be used in gardens, which may in the futurę result in an increase in the number of stations. Elsholtzia ciliata (Thunb.) Hyl. [syn.: E. patri- nii (Lcpech.) Garcke; E. cristata W illd.; Sideritis ciliata Thunb.; Mentha partinii Lcpech.] Echinops sphaerocephalus L. Lamiaceae Glandular Globe-thistle Asteraceae Biology: annual plant dispersing through anemo chory, zoochory and anthropochory. Biology: perennial plant dispersing seeds through Native rangę: central part of the former Soviet exozoochory, anemochory and myrmecochory. Union, central and eastem Asia (Grodzińska 1985) Sometimes planted as honey-producing or deco- where this species occurs in the fields, along ri- rative plant. verbanks, along forest roads; also cultivated. Native rangę: south-eastem Europę: especially General distribution has been given by F e d o r o v Pokucie and Podole where it occurs in the Dnie- (2001). ster ravines and its tributaries (Rostański K. Secondary rangę: central Europę (excluding the 1971). British Isles - S ta c e 1997) and North America. Secondary rangę: southem and central Europę, reaching the Caucasus and Siberia (Rostański K. History of spread: 1971). Europę: species once grown, especially by Slavs as a health-giving plant (Krawiecowa 1951), History of spread: progressed to the wild state near sites of cultiva- Europę: plant brought and long cultivated in tion and introduced accidentally into a number of many European regions (sown by beekeepers); European regions, possibly via Poland. Previously going into the wild state near areas where it is recorded in Lithuania by Górski in 1830 (after grown. The level of naturalisation of this species Gudżinskas 1998a). Recorded for the first time in some areas of central Europę led to its con- in Czech Lands in 1853 (Pyśek et al. 2002). sideration as a possibly an indigenous species Poland: first stations evident from the first half (L ohmeyer & Sukopp 1992). of the 19th century (Fig. 44, see p. 80). In 1872 Poland: probably occurred in Poland as early as the plant was found again in the Warsaw area by in 16th century. Referred to for the first time by Rostafiński, and in 1873 it was recorded by Karo S irenius (S yreński 1613) (cf. Chapter 5.2). Be- (,herbarium31: UW and W) in the Łosice area fore the end of the 19th century found in several (eastern Poland). In the foreland of the Car- stations; in the first half of the 20th century the pathians it was collected in 1877 in the Przemyśl number of stations increased to 90. The recon- area (Kotula, herb. KRAM). Up until the end of struction of the pattern of spread of this species the 19th century it was recorded in 65 stations. In allows for its classification as one of the oldest the 1930s it was common in villages, and along of the kenophytes to arrive. It might even be roadsides in the Dynowskie and Przemyskie supposed to have been introduced accidentally Plateau (Batko 1934). In the central part of the or brought into Poland even in earlier centuries Carpathians it was found in the 1950s in the (Fig. 43). Gorce Mts., in Gubałowskie plateau and in the Habitats: slopes, scrub, roadsides, boundary Polica rangę (Guzikowa 1972). In the Pieniny strips, railway embankments, rubble. A charac- Mts. it was found by Zarzycki (1969). In the teristic species of the Onopordenion acanthii sub- Beskid Żywiecki it was less common, up to an alliance and of the Onopordetum acanthii asso- altitude of 500 metres (Białecka 1982). ciation (M atuszkiewicz 2001). Habitats: roadsides, around cottages, ruderał sites. Dynamics: occurring fairly often as early as in Dynamics: scattered across the lowlands, occur the beginning of the 20th century; the number of ring more often in the north-east and east of Po stations have increased significantly in the last land (Fig. 44). An evident increase in the num half-century. Up until the present time it has been ber of stations appeared in the second half of the recorded on 910 stations in all, in 489 ATPOL 20lh century. In subseąuent years the species has sąuares. Currently scattered across the whole of gradually expanded its rangę moving from east Poland, locally rarer, e.g. in mountains (K ornaś to west. Kornaś (1950) reported E. ciliata from et al. 1996; Szeląg 2000) (Fig. 43 and Appen- dix A). Gradually colonises new sites. 31 Acronyms of names of herbaria are introduced in explanations to Appendices A and B. Introduction and first records: Start ofspread: - 17m and 18" centuries: first mentions of the presence of occupation of new localities in south-western Poland and in thisspecies intheterritoryof Poland (Syreński 1613) the upper Vistula valley which were probably located in the vicinity of cultivation sites ® first recorded localities of occurrence: Warszawa ED16 (from S u dn ik-W ó jc ik o w s k a 1987a), Kraków DF69 directionsoffurtherspread (Besser 1809) direction of importation of species from south- eastern Europę

Subsequentspread phases: The current distribution of this species is linked to upland rangę increase areas and to sites of cultivation - a tendency to an increase in the density of localities, mainly (the map after Zając A. & Zając M. (eds.) 2001 - slightly in south-western Poland supplemented) themainmigrationfronts

Fig. 43. Recorded history of the spread of Echinops sphaerocephalus L. - one of the oldest kenophytes in the Polish flora the area around Cracow as a common plant, a total disappearance in some stations, because occurring locally on a massive scalę and expand- of the elimination from the landscape of Polish ing its rangę. Similar characteristics were provided villages and smali towns representing the hab- by G u z ik o w a (1972), i.e. presenting it as a species itats preferred by this species (cf. also Chapter 6). expanding across Poland, with migration from east to west (1352 stations recorded in 814 Impatiens glandulifera Royle [syn.: 7. Roylei ATPOL sąuares). In most recent years there has Walp.] been a tendency for decreasing population num- Indian Balsam bers in the previously known stations and even Balsaminaceae First recorded localities of occurrence: Start ofspread: migration ofthe speciesfromthe easttowardsthe west: ® Warszawa DF16 (from W aga 1847 and Sudnik-Wójci- occupation of new localities, mainly in the north-eastern part kowska 1987a), for these first localities the plant was probably growing after escape from cultivation or ofthe country accidental import with long-range transport ofgoods ^ jg main direction of arrival ofthe species in Poland X migration ofthe species from Ukrainę ------_ contemporary rangę limit direction offurtherexpansion

F G Current distribution ofthe species rangę increase and stabilisation: ■»— a direction of spread appearing distinctly in a tendency to an increase in density of localities in north- the 1980s eastern Poland Currently a tendency to the gradual of localities was furtherspread ofthe species to the west recorded under a whole rangę of species in Poland

Fig. 44. Recorded history of the spread of Elsholtzia ciliata (Thunb.) Hyl. in Poland - an example of a species transiently enlarging its rangę in a westerly direction

B iology: annual plant with high level of seed Native rangę: the Himalayas and eastem India production. Diaspores disperse via two ways: by where it grows in humid riparian forest at an al- autochory32 and by allochory: through wind, titude 1800-3000 m a.s.l. (L h o t sk A & K o peck y animałs and water. 1966).

32 In the case of the Balsams, autochory is implemented following the abrupt release of a tension in the fruit and through the process called ballochory, where a ballistic triggering movements of the pericarp walls (P o d b i e l k o w - mechanism causes throwing out (hurling) of diaspores s k i 1 9 9 5 ) . First recorded localities of occurrence, probably escapes from cultivation: ® the Sudety Mountains: Siodło AD86, Płóczki Dolne AD48, Stępnica AD48 and Płonina BE61 (Schube 1903b)

А В С D E F Q Start ofspread: I Т>ІГП ł 1 1 j Л ж У naturalisation closetocultivation sites: A ь 1 : 1 ] t j ^ L occupation of new localities, especially in the south- \ \ ' western partofthe country И 1 B я \{ f directions ofspread J \ ^ Ш \ _-A | c \ N

) ) г /' \ V г Y г У ' Г J i D \ \ ) ч (V _ (. i X ( E ' . ~ x \ \ Д. )\ ~~І\ ł / ^ \ / I F - 4 _ i 4 - - A : ( у . „ з / ' .. . и ę - 7 ; Г ] к G 1950 ^ t \ - С m А В С D E F Q

Subseq uent phases of spread: rangę increase and stabilisation further naturalisation from cultivation sites and autonomous spread from previously occupied localities

Current wide distribution of the species in the territory of Poland with regionsofclustered occurrences in the Southern partofthe country

Fig. 45. Recorded history of the spread of Impatiens glandulifera Royle in Poland - an ехатріе of an omamental garden plant escaping into ruderal habitats and migrating into riverside habitats

11 The Establishment.. 81 Secondary rangę: central and northem Europę Dynamics: in Poland it is still cultivated and (extending by central Scandinavia). Crossing into the wild state; dispersing spontaneously from newly colonised sites. The number of History of spread: sites began to increase in the 1960s, and a remark- Europę: the history of the expansion of the Indian able growth thereof has appeared sińce the 1970s Balsam in Europę started in the flrst half of the (Fig. 45). At present, it is scattered throughout the 19* century sińce it was cultivated as a decorative national territory (1574 stations in 675 sąuares) and medicinal plant in the gardens of universities, (cf. Appendix A). The regions of its freąuent and convents or monasteries, and later on also in pri- massive occurrence are located in the southem part vate gardens. The first information on the cultiva- of Poland: the Carpathians, Silesian Upland, Kra- tion of this species in Europę dates back to 1839, ków-Częstochowa Upland, the southem part of the from the botanical garden in Kew (Great Britain) Silesian Lowland and Małopolska part of Vistula (LhotskA & Kopecky 1966; Zając E.U. & Zając A. river valley. The species prefers river valleys (par- 1973), and after that it was recorded in Austria: ticularly mountain and foreland rivers), occurring Linz area (1845), Vienna (1871), Innsbruck (1880) often along the upper course of the Vistula and (Drescher & Prots 2003). Odra rivers and their tributaries, e.g. often found Since then it has been grown in the gardens of along the Soła, San, Wisłoka, Skawa and Olza. a number of European countries and it spread to The species still colonises new sites in many re nearby ruderal habitats and next to riparian ones. gions, particularly along rivers. The first spontaneous, wild stations were recorded in England (Middlesex) in 1855 (P errins et al. Impatiens parviflora DC. 1993), in Austria in 1898 (over the Weidling River Smali Balsam near Klostemeuburg) (D rescher & Prots 2003). Balsaminaceae In many countries it is referred to as a serious and widespread weed invading semi-natural or natural Biology: annual plant which produced a high habitats (Cronk & Fuller 2001). In England it was number of seeds. As in the previous species, the given weed status as long ago as 1898 and currently diaspores disperse in two ways: autochorically (as it is considered as the most invasive and common a result of ballochory) and allochorically through species of the genus (Perrins et al. 1993). wind, animals and water. Poland: first stations ofplants which progressed Native rangę: southem Asia, Siberia, Mongolia into the wild State in Poland were recorded by and Turkistan. Schube (1903b) in Lower Silesia in 1890 (Fig. Secondary rangę: central and northem Europę 45). By 1940, spontaneous occurrences of the excluding northem and western Scandinavia. Indian Balsam were recorded in 21 sąuares, mainly in south-westem Poland and an isolated History of spread: Europę: the first map of the synantrophic rangę station in northem Poland, in the Wiślane Marsh- land (Mierzeja Wiślana). After that in the subse- of this species in Europę was developed by Meu- se l et al. (1978), citing the earliest dates for the quent 40 years, this species also expanded in occurrence of this species in Europę: 1834 (Russia), Southern, south-eastem and central Poland as well 1837 (Germany), 1848 (Great Britain). In the as in Pomerania. Few stations had been recorded mid-19th it was already observed in a number of in north-eastem Poland until 1980. localities in western and central Europę. Kamień Habitats: human-made habitats such as: built-up ski (1884b) considered that the species was in- areas, cemeteries, allotments, refuse heaps, urban troduced accidentally by travellers and described wastelands, abandoned fields, and more often in the migration route as follows: “the plant was drainage and roadside ditches. It is also recorded moved to Western Europę by sea, which was a from more natural habitats, namely: scrub, forest considerably longer route than by land, and even edges and most freąuently from riparian habitats today transport by this route is very difficult”. (Tokarska-Guzik 2003a & c). It has been noted Other botanists state that the Smali Balsam is a from the Odra river valley in riverside, and refugee from botanical gardens. periodically in the flooded forests of Alno-Padion, Poland: it was recorded for the first time in but the biggest stands were classified as Impa- Poland in 1850, in the Gdańsk area (M eu sel et tienti-Calystegietum, which prefers semi-shaded al. 1978), whilst subsequent records cite the forest edges, not far from the river (Dajdok et al. Wiślane Marshland (northern Poland) in 1866 1998, 2003). It has been described also from (Klinggraeff 1866). At the same time, the sta poplar-willow carrs Salici-Populetum (Jasnowski tion of the Smali Balsam was noted in the Cra- 1961, Zając E.U. & Zając A. 1973). It forms an cow (Ullepitsch herb. B) and Wrocław areas aggregative community in the association class of (Uechtritz herb. W) (Fig. 46). In the Warsaw Artemisietea vulgaris (Matuszkiewicz 2001). area it was found by Kamieński (1884b) in parks Start ofspread: ® first recorded localities of occurrence: environs of Gdańsk DA80 (MEUSEL et al. 1978), Vistula Żuławy (North Poland) (Klinggraeff 1866) as well as Kraków DF69 (ULLEPITSCH, herb. B) and Wrocław BE49 (UECHTRITZ, herb. W)

А В С D E F G

Subsequent phases ofspread: rangę increase and stabilisation rapid occupation of new localities, especially in south- V western Poland y directionsofspread

А В С D E F Current wide distribution of the species in the territory of Poland with regions of clustered occurrences in the south- western, Southern and south-eastern parts of the country

Fig. 46. Recorded history of the spread of Impatiens parviflora DC. in Poland - a species escaping from botanical gardens, becoming established in ruderal habitats and naturalised in forests as the “obtrusive Mongoł” and gardens. In former floristic papers it was m a n e k & W in n ic k i 1999). It is extending its rangę reported as a species occurring in ruderal places eastward: first recorded in the Ukrainę in 1908 (Klinggraeff 1885; Abromeit et al. 1898), but as (Dr M. S h e v e r a , pers. comm.). In Poland it is an early as in the 1950s it was also found in vari- invasive species (cf. Appendix A). ous types of forests, mainly deciduous ones. Habitats: Forests (oak-hombeam, ash-alder ri- Reynoutńa japonica Houtt. [syn.: Fallopia ja p o parian carrs, willow-poplar carrs, beech forests, nica (Houtt.) Ronse Dccraense; Polygonum cuspidatum Sicbold mixed coniferous forests, oak-pine forests), and & Zucc.; P. zuccarinii Smali; Polygonum sieboldii hort. non DC.; Pleuropterus cuspidatus (Sicb. & Zucc.) Moldenke; P. zuccari anthropogenic habitats: parks, cemeteries, garden nii (Smali) Smali; Tiniaria japonica (Houtt.) Hedberg] allotments, wastelands, cottage yards, refuse tips Japanese Knotweed and railway tracks and embankments. A charac- Polygonaceae teristic species for the Alliańon alliance (Matusz kiewicz 2001); sometimes a separate association Biology: a conspicuous rhizomatous perennial with the predominance of Smali Balsam is dis- plant, dioecious with dioecious or gynomonoe- tinguished as Impatientetum parviflorae. cious flowers, spreading mainly through vegeta- Dynamics: by the end o f the 19th century it had tive processes (Tokarska-Guzik, in press, cf. been reported from 35 localities in 19 ATPOL references therein). sąuares. Massive expansion of I. parviflora started Native rangę: includes Japan, Korea, Taiwan, in 1960s, and up to the present date it has been northem China where it occurs in humid, open recorded in over 6730 localities in 1681 sąuares areas on hills and mountains, on roadsides, and (cf. Appendix A). Zając-Sychowa (1971) de- on the banks of ditches (Tade Zoku 1965; B ailey scribes the species as widespread in the lowlands 1999). In addition, it often occurs in grassy com- and in lower mountain regions (in the Gorce Mts. munities formed by Miscanthus sinensis (Cronk it reaches an elevation of up to 610 m a.s.l., in & F u lle r 2001). It grows on various soils, col- the Sącz region - of up to 480 m a.s.l.), in gar onising even volcanic soils (Ohwi 1965; B a iley dens, near fences, on roadsides, and is also found 1999). upon streams and in humid, shadowy sites. Secondary rangę: extended to Europę, Canada, Currently it is widespread throughout the na- USA, New Zealand and some areas in Australia. tional territory of Poland, although more common Maps developed to date include Europę (Jalas & in the southem part, and rarer in the north-east. Suominen 1988) and some specific European Often found in the Ciężkowice Foothills (West countries: the Czech Republic (Slavik 1986), the ern Carpathians) where it grows in ruderal hab UK (Child & Wade 1999, 2000), Poland (Zając A. itats but also in forests (K ornaś et al. 1996), and & Z ając M. 2001) and the US and Canada in the Beskid Żywiecki Mts. (B iałecka 1982), the (Seiger 1997). Beskid Śląski and Beskid Niski. Accidentally in- It is freąuent in a number of European coun troduced into lower locations in the Tatry Foot tries, more so in the northem and central part of hills, the maximum elevation recorded in the the continent. B eerling et al. (1995) state that its Tatra Mts. is 1150 m a.s.l. (Piękoś-M irkowa & current distribution is determined by climatic fac- M irek 1978). In the Karkonosze National Park tors. The northem boundary demarcates a com- the species was recorded in stations at 950 m bination of factors such as the length of the ve- a.s.l. (Rostański K. 1977). Szeląg (2000) reported getative season and minimum temperatures in win- this species from the Śnieżnik mountain massif ter (B eerling 1993). Water availability in soil and and Bialskie Mts. as often occurring in lower temperature delineate the southem boundary. Al sites, and permanently established in deciduous though in Europę its rangę is contained between forests and scrub. G uzikowa (1972) reconstructed a latitude of 42°N and 63 °N, and its natural rangę the spread of this species in the Pieniny Mts., is between 22°N and 45°N, it is analogous cli referring to the earliest records by Pawłowski matic zone (B eerling et al. 1995). (1925) from Szczawnica and by K ulczyński In some European countries (England, Germa (1928) from the Krościenko locality. Its penetra- ny) it is considered a widespread invasive spe tion into the Pieniny National Park occurred from cies, also entering natural and semi-natural hab the villages, particularly along the tourist trails itats (C ronk & Fuller 2001) (cf. Appendix A). from Szczawnica and Krościenko on Sokolica Mt. In the early 1970s, the species was wide History of spread: spread in the region and not only in ruderal Europę: brought to Europę as a decorative plant, habitats but also in osier beds upon the Dunajec probably by Philippe von Siebold who stayed in and Krośnica rivers and in natural forest habitats Japan from 1823-1829. In 1847, Japanese Knot on the eastem side of the Park. This species does weed won a golden medal award bestowed by not occur in the Bieszczady National Park (Z e- The Society of Agriculture & Horticulture in First recorded localities: ® West Poland: Gniezno CC83 (CYBICHOWSKI herb. POZ), Wrocław BE49 (BAENITZ herb. WU), North Poland: Darzlubie CA48 (G raebner 1894)

A B C D E F O

A B ______C______D______E______F______G Subsequent phases of spread: occupation of new localities, predominantly ___ ' s in the Southern part of the country ' i J 1 f ' directionsoffurther spread * "5 ■ < \ \ J \ i j 7 \ T t L / " " V / — £ ) : \ W i I > '“ a * X \ / , N * \ *. • I V / V h \ \ * J J '1 / D k- > \ > )•* s - i ) \ - r . * • J i ; V__ _ < / , 3 , \ )/ /■ V • : -IV 1 9 5 0 N) ...... I n \r A B C D E F O

The current distribution ofthis species is an effect of fast spread rate, especially in river valleys where it forms compact monospecific phytocoenoses which often occupy extensive areas in the habitats of former willow-poplar forests and thickets. It also occurs commonly in urban areas and railway territory (the map after Zając A. & Zając M f ••• • • • • • • i • •• • - • (eds.) 2001 - slightly supplemented)

ti !!** f r -!*• X ’. .*. v *■. v : ' •Fr

Fig. 47. Recorded history of the spread of Reynoutria japonica Houtt. in Poland - an example of an invasive plant using vegetative reproduction to spread Utrecht, as the most interesting decorative spe- habitats of natural types: on river banks, forest cies of the year (B ailey & C onolly 2000), and edges (particularly of disturbed carrs) and edges as early as in 1848 it was commercially available. of scrub. The re-introduction of R. japonica to a number This species shows wide tolerance towards of European countries was probably launched by types of soil: it has been recorded on soils within the nursery in Leiden (the Netherlands), which the rangę of reaction from pH 3 to pH 8.5 as well offered seedlings for botanical gardens with a as on salinę, polluted or contaminated soils 25% discount. (R ichards et al. 1990). The first receipt of seedlings of Polygonum Dynamics: fairly widespread over the whole na- sieboldii in the Botanical Garden in Kew (En- tional territory, reaches elevations of 750 m a.s.l. gland) from Leiden, was recorded in the cata- in the Karkonosze Mts., in Działy Orawskie - logues on October 9, 1850. Botanical gardens, 535 m a.s.l. and in the Tatra Mts. - 860 m a.s.l. gardeners who sung its praises in professional (Zając A. 1992), or even 1000 m a.s.l. (Piękoś- magazines and private collectors all played an Mirkowa & Mirek 1978). In Poland a total of essential role in the dissemination of this species 3004 stations of this species were identified in (B ailey & C onolly 2000). 1158 ATPOL sąuares33 (cf. Appendix A). The Detailed information on the occurrence of greatest concentrations of these are observed in R. japonica in a wild state in Europę is scarce the southwestem and Southern parts of Poland (B ailey 1999). H egi (1910) published informa (Fig. 47), where apart from anthropogenic hab tion on the application of this plant for stabilis- itats it also enters riparian habitats forming com ing dunes being followed in Helgoland, which pact phytocoenoses. The enormous potential of was published as early as in 1861, and these this species for spreading through vegetative efforts continued. The earliest incidents o f “escapes” means, combined with its rapid growth and a o f R. japonica from cultivation areas were capacity to adapt to diverse or even extreme reported in Germany (the Rhur area) and Great habitat conditions, often invading and holding Britain (Wales) where the plant was introduced large areas, have resulted in this species eaming onto coal and slag heaps (C onolly 1977; B ailey the status of invasive plant and nuisance “weed” 1999). In Germany it was also used by hunters (Tokarska-Guzik in press). It still continues to as camouflage for raised stands (B ailey 1999 colonise new sites, on a massive scalę in many after A lberternst et al. 1995). 1886 is reported regions (cf. also App. A and Chapter 12). as one of the first dates in the UK, when R. japo nica escaped from its areas of cultivation (H ollingsworth & B ailey 2000). Until the late 19thcentury there were only seven stations of 7.2.3. Examples of species of American R. japonica in the British Isles (C onolly 1977). origin Since 1940, the number of the stations has been growing rapidly (C hild & W ade 1999). Echinocystis lobata (F. Michx.) Torr. & A. Poland: the first station for R. japonica in Po- land dates back to the second half of the 19th Gray century. Stations were reported by Cybichowski Wild Cucumber {herb. POZ) in 1882 in Gniezno; by Baenitz in Cucurbitaceae 1893 in Wrocław {herb. WU) and by Graebner (1894) in the same year in Darzlubie (Baltic Biology: annual plant with climbing shoot and Coast) (Fig. 47). Schube (1903b, 1904, 1905, spiny fruits; dispersal involves seeds, fruits and 1908, 1910) reported the location of a dozen of shoots. so stations occurring in Lower and Upper Sile Native rangę: eastem part of North America. sia. These dates may not be complete as R. japo Secondary rangę: Central Europę (absent in nica had undoubtedly more stations, including England) and Asia. western and central Europę, especially in large History of spread: cities, in the first half of the 19th century (Pro- Europę: the plant was brought as a decorative fessor R. Olaczek, pers. comm.). In the 1960s the species at the tum of 19* and beginning of the 20lh number of stations increased to 342 and it con- century. Specimens which had moved into the tinues to grow (Fig. 47). Habitats: within the limits of its secondary dis- 33 The distribution needs certain verification because of tribution rangę it occurs principally in anthropo- probable erroneous records at some stations of the R. x genic habitats, such as roadsides, railway em- bohemica hybrid as R. japonica. Nevertheless, it is defmitely bankments, various urban and industrial waste- the most freąuently recorded species of this genus in the lands, in parks, cemeteries, gardens, but also in Polish flora. Introduction and start of spread: 19’ /20 century - introduction of the species into cultivation ® first recorded localities of occurrence: Gubin AD43 in West Poland (Lademann 1937), Kraków DF69 and a couple of villages in the vicinity of Lublin (localities in FE & FF cartogramme units) >. main direction of arrival ofthis plant from the territories \ of Germany and Ukrainę ^ spontaneous spread near cultivation and naturali- ' sationsites >N riparian corridor migration routes

Subsequentphaseof spread: rapid rangę increase (invasion), especially in the south- eastern part of the country where this plant is also more often cultivated; a distinct link may also be seen between the migration and river valleys

• • •

2000 A local ехатріе of riparian corridor migration of Echinocystis \ lobata in the Bug river valley (source: Fauński et al. 2000)

Fig. 48. Recorded history of the spread of Echinocystis lobata (F. Michx.) Torr. & A. Gray in Poland - an ехатріе of an introduced ornamental plant with invasive properties, or how an introduced plant becomes an invader

“wild” state were recorded for the first time in and several localities in the Lubelskie province. 1904 (M eusel et al. 1992; B alogh 2001); nume- At the same time, especially in the last half- rous stations were found in Austria and Hungary century it was cultivated in many regions, from as early as in the first half of the 20th century whence it spread into the “wild” (for ехатріе in (H eine & Tschopp 1953; Priszter 1958). In the Wrocław it was often recorded on the fences of territories of Poland’s neighbours, the species was garden allotments in the 1960s) (Prof. K. Ros- found to be self-dispersing in 1906 in the former ta ń sk i, pers. comm.). Czechoslovakia (Lohmeyer & Sukopp 1992), and Habitats: willow and willow-poplar carrs on in 1929 it was found in the Ukrainę, where its riverine and lacustrine banks as well as ruderal invasion was observed by Dr M. Shevera (pers. sites: fences, refuse heaps, around cottages, aban- comm.). In Slovakia, on the Dubai, it is considered doned gardens, municipal refuse tips. a potentially invasive species (U herćikovA 2001). Dynamics: the number of its stations began to In Lithuania, it started to spread intensively in the increase only in the second half of the 20th cen 1990s (G udżinskas 1999a). tury, rising from seven sites recorded in the first Poland: it was probably brought into Poland half of the 20th century to 2047 in 708 ATPOL from two directions: from Germany where it has sąuares (cf. Appendix A). Currently, the species been recorded sińce 1922 ( M e u s e l et al. 1992) is widespread in the Southern and south-eastem and from the Ukrainę (Fig. 48). Initially, a dozen parts of Poland, particularly in riparian habitats or so stations were recorded: Kraków-Bronowice (e.g. upon the San, the Yistula rivers, and - increasingly often - along the Oder river; it is A species characteristic for the Rudbeckio-Soli- however rarer in northern parts (Fig. 48) and is daginetum association (Matuszkiewicz 2001). also found in the lower zones of mountains, most Dynamics: as early as at the beginning of the 20th often in the valleys of the Nysa Kłodzka and century, the species was recorded in 78 stations, Biała Lądecka rivers (S zeląg 2000); in the while in the 1950s there were 187 stations. Western Carpathians along the Wisłoka, Ropa and Within recent times, information about as many Biała rivers (K ornaś et al. 1996) (Fig. 48). Still as 2251 stations was noted in 903 ATPOL sąuares colonising new sites. Its spread is of an invasive (cf. Appendix A). It is found throughout the type. territory of Poland, although more rarely in some regions of central and northern Poland. The re- Rudbeckia laciniata L. gions it most freąuently occurs in include the Sudety Mts. and Sudety Foreland, the Wielkopol- Tali Coneflower, Golden Glow ska-Silesian Lowlands, the Silesian-Cracow Asteraceae Upland, the Małopolska Upland, the Carpathian Basins and the Carpathians (Fig. 49). In the Car Biology: conspicuous perennial plant dispersing pathians it reaches the elevations where the major its seeds through anomochory, exochory and settlements are: in Babia Góra Mt. - 750 m a.s.l., myrmecochory. Gorce - 515 m a.s.l., Bieszczady Zachodnie - Native rangę: moderate climatic zone of eastem 720 m a.s.l. (Trzcińska-Tacik 1971 b). and central North America. Secondary rangę: Europę: in the north extending Mimulus guttatus DC. to Sweden, in the south to Corsica and reaching central Russia in eastem Europę. Outside Europę, Monkeyflower the secondary rangę includes also eastem China, Scrophulariaceae New Zealand and Japan (Cronk & Fuller 2001). Biology: a perennial plant expanding generatively History of spread: by minutę seeds dispersed by wind and water and Europę: one of the oldest decorative perennial also by vegetative processes. plants brought into Europę in the early 17,h century Native rangę: western part of North America or even earlier (cf. Appendix A). Its occurrence in from Alaska to northern Мехісо. Paris was recorded in 1615 (Jalas 1993; Francir- Secondary rangę: western and central Europę: kova 2001). Its freąuent cultivation in Europę con- mainly the British Isles, northern France, the tributed to its dispersion. The first stations o f plants Netherlands, Germany, Switzerland, Poland and which “moved into the wild State” were recorded some areas of northern and eastem Europę: Scandi- in 1787 in an area which currently lies within navia, Lithuania and the European part of Russia. Poland (Jalas 1993; FrancirkovA 2001). Currently, it is frequently found in a number of areas in History of spread: Germany, Austria, the Czech Republic and Slova- Europę: in Europę, especially in the western kia. For the last two countries it has been consid- part, a number of species from the M imulus ered an invasive species (cf. Appendix A). genus were grown, including M. guttatus. It is Poland: this species was brought to Poland in the prone to straying into the “wild”; in some areas 18th/19th century (Kornaś 1968b). The first station of Europę it has become naturalised and forms in the Sudety Mts. (1787, cf. Fig. 49) was published a part of natural communities (Piękoś 1972; by Fiek (1881 after Krocker). Subseąuent stations S tace 1997). The first “wild” stations in central recorded in the next half century were also located Europę were recorded in 1824 (Lohmeyer & in this region. In eastem Poland it was recorded by Sukopp 1992), 1847 (B a lo g h et al. 2001) and in Drymmer (1897) in the Lubelskie province and the 1853 (Pyśek et al. 2002). In recent years its first Opoczno, Turek and Sieradz areas. It was Szafer stations have been recorded on the Raba River et al. (1924) who observed the Golden Glow and Dubai in the western part of Hungary moving readily in the “wild” and noted that it could (B a lo g h et al. 2001). be found within scrub on river banks. In subsequent Poland: this species is also grown in some Po- periods this species was recorded more frequently, lish regions (especially in the west) and it strayed especially in south-westem Poland. Trzcińska - from there into the “wild”. The oldest occurrence Tacik (197 lb) characterises this species as common was recorded from the Sudety Mts. (Fig. 50). all over Poland, dispersing without assistance, and This is at the same time the oldest registered date as also present in semi-natural habitats. of the occurrence of this species in Europę Habitats: banks of rivers, streams and ditches; (although it was dispersed in cultivation at that time also enters riparian osier beds and carrs, addition- in other parts of Europę, e.g. in the British Isles). ally also grows in ruderal habitats and in gardens. In the Sudety Mts. it started its occupation of new Introduction and startofspread: Initial phase of spread: ® first records: Świeradów AE67 in the Sudety Mountains ' ' “'sN increase in the density of localities within the occu- (Fiek 1881 afterKrocker); nearLubańAE67 (after Jalas pied territory 1993); subsequent records: Bolesławiec AE28 simultaneous occupation of new localities in other (Schneider 1837) and Bystrzyca BE75 (Fiek 1881 after 4 regions of the country due to popularisation of Krocker) cultivation and the concurrent naturalisation of this ^ spontaneousspread nearcultivationsites plant as well as probable accidental importation of its seeds directionsoffurther spread

Subsequentphasesofspread: The current distribution of this species is linked to the further increase in the density of localities; spread south- historyofitscultivationandescapesfromgardens eastand north map а^ег zając A & Zając M. (eds.) 2001 - slightly supplemented and modified) Fig. 49. Recorded history of the spread of Rudbeckia laciniata L. in Poland - an ехатріе of a popular ornamental plant where naturalisation is due to long and widespread cultivation stations in the second part of the 19th century; at vestigated by P ięk o ś (1972) who recorded the the same period it was recorded in Pomerania and occurrence of this species at 112 stations. Masovia where it was probably introduced acci- Habitats: banks of streams, rivers and lakes, dentally (or, initially on purpose) from Germany as well as along ditches, rare in ruderal habitats. (in neighbouring Lithuania it has been recorded A characteristic species of the association sińce 1931; it is currently spread along the Neris Sparganio-Glycerietum fluitantis (M a t u s z and Niemen Rivers (G u d ż in sk a s 1998a)). The kiew icz 2001). K w iatkow sk i (2003) describes for history of the dispersion of this species was in- the first time for Poland the association Yeronico

12 T he Establishm ent. 89 Start of spread: Initial phaseof spread:

® first record: Kowary BE80 in Sudety Mts (Fiek t n increase in the density of localities within the occu- 1881; ?? herb. WRSL) x v pied territory y" spread in the region of the first record ^ simultaneous occupation of new localities in ' the north of the country ___ w probable direction of arrival of this species in Sudety Mts ------v probable directions of arrival of this species toPoland

A B c D E F G riTTm-n r j r n j r i i Klim s 1 i r y A X? ■ A /Ł _ _'> Sm V* 1 • ••• ^ 7 ~ y ^ • K l : Y \{ B • \ B V. • • • \ - \ / • • c : K J C i c : [ '\ X •• L - \ D / D : : p . f 3 ' ) f E : E :J:/ / V c ( i s • \ j A : \ • / F r . f r / ll F \ a * )/ i - ^ V. \ - 1 G : 2000 G > } y { A B c D EF G Subsequent phasesof spread: The current distribution illustrates the regions of the pre- t n further increase in the density of localities and cre- viously occupied localities ation a rangę encompassing areas in south-western and north-western Poland -»■ directions of further spread

Fig. 50. Recorded history of the spread of Mimulus guttatus DC. in Poland - an example of a species currently having a characteristic rangę type in Poland beccabungae-Mimuletum guttati as a member recorded in 326 stations in 128 ATPOL sąuares of the alliance Sparganio-Glycerion fluitan- (cf. Appendix A). The species is gradually in- tis. Sometimes the species occurs in phyto- creasing the number of its stations, mostly in coenoses of other communities of the classes regions of previous concentrations (Fig. 50). Phragmitetea and Isoeto-Nanojuncetea (K u Rapid expansion of this species has been noted charski 1992). particularly in the Karkonosze Mts. (F abiszew D ynam ics: at present it occurs most often in ski 1985; Fabiszewski & K wiatkowski 2001; Lower Silesia and Pomerania. To date it has been Kwiatkowski 2003). 7.3. The spread of accidentally nessed there a half-century earlier, probably by introduced plants: how the French army stationed after the 1806 Prussian an ephemerophyte turns war in Pomerania and Wielkopolska province. into a kenophyte The reconstruction of the expansion stages in specific periods of the 19th and 20th centuries indicates that the belt of the Central Polish Low 7.3.1. Plants introduced accidentally lands was the main migration route and the spe from various regions of Europę cies migrated to this area from German Łużyce. Habitats: cereal fields, more rare in root crops, stubble fields, sandy areas left out of cultivation, Anthoxanthum aristatum Boiss. [syn.: A. puelii also noted in railway traeks and embankments, as Lecoq & Lamotte] well as on industrial waste heaps. A characteristic species for associations within the alliance Annual Vemal-grass Arnoserido-Scleranthetum (Balcerkiewicz et al. Poaceae 1999). Dynamics: the number of stations has inereased Biology: annual plant, disperses through anemo- markedly in the last 30 years, particularly in the chory, also in agricultural/horticułtural seed mix- central part of Poland. A total of 1031 stations tures (Kuźniewski 1996). have been recorded in 577 ATPOL sąuares to date Native rangę: Western Europę (Atlantic region) (Fig. 51, cf. Appendix A). The area of its ехрап- and northem part of central Europę: to the east sion includes primarily agrocoenoses appearing in it reaches Germany, to the south it reaches South the poorest habitats colonised by Teesdaleo-Arno- ern France via Corsica and Sardinia (M eusel et seridetum minimae (Warcholińska & Siciński al. 1965). 1976). In accordance with the same authors Secondary rangę: central and eastem Europę; (1996), the expansion of this species is facilitated currently a rare ephemerophyte in the British mostly by favourable edaphic and climatic con- Isles, but in the past naturalised in sandy and ditions, as well as methods and pattems of land infertile soils in Surrey and East Suffolk; not seen use ways; they also stated that occurrences of there sińce the 1970s (S tace 1997). A. aristatum have a “destmctive impact on agro History of spread: coenoses as this species eliminates other species”. Europę: it spread out of its original rangę in Napoleonie times (1805-1813). Since then it has Artemisia austriaca Jacq. dispersed in various directions, where it grew Austrian Sagewort exclusively in ruderal and segetal habitats Asteraceae (K uźniewski 1996; K orniak 2002). Poland: introduced in the 19,h century, initially to Biology: a perennial plant which disperses in Po Pomerania and Silesia (Fig. 51). According to land principally through vegetative processes. Warcholińska and Siciński (1976), the species Żukowski and Piaszczyk (1971) suggest that seed had not been reported in central Poland until 1960. development stops short of maturity or that seeds Since 1960 there have been more and more reports maturę only in some years, and that the plant ini of its occurrence in various regions of the country tially colonises sites solely through vegetative (Warcholińska & Siciński 1996). In Warsaw, it processes. had been recorded only once (Zanowa 1964) up Native rangę: eastem and south-eastem Europę until the 1970s, but in the following decade it was (widespread in Podolia, Volhynia and Kiev re found by Sudnik-Wójcikowska (1987a) in several gions), in western and central Asia, Siberia where stations, mainly in cereal fields and non-arable it occurs on steppes, steep slopes and ruderal areas land. In 1975, A. aristatum was recorded in 118 (K ornaś 1968b; Żukowski & Piaszczyk 1971). stations in central Poland. In subsequent years 437 Secondary rangę: central and western Europę. new stations were recorded in the region (W ar History of spread: cholińska & Siciński 1996). Kuźniewski (1996) Europę: it dispersed from the Podolia and Kiev suggests two distinct routes of migration of regions northwards and westwards (Żukowski & Annual Vemal-grass in Poland: a northem route Piaszczyk 1971). In Poland it is gradually reach- from southem areas of the Szczecin province and ing westward to other European countries thereby a southem one - from Lubuskie Lakeland to the extending its rangę. Trzebiński (1930) had already Central and Eastem Polish Lowlands. mentioned that this species was introduced acci Although the first recorded dates of occurrence dentally to Germany and France in a few cases. of Anthoxanthum aristatum go back to the sec- Hardtke & Ihl (2000) present information on a ond half of the 19th century, it was originally wit- single station of this species in Saxony in 1946. First recorded localities of occurrence: early 19m century: probably first undocumented accidental importation in the region of North Poland and Central Poland ® first records: Western Pomorze: Kwidzyń DB52 (Klinggraeff 1866) and Milewo DB60 (A b ro m e it et al. 1898); south-western Poland: environs of Ryczeń BD85 (FIGERT herb. W) and between Rzeszotary and Sześcina BE23 (FIGERT herb. MGS), as well as in Zgorzelec AE35 (H a rd tk e & Ih l2 0 0 0 ) ------■*. probable direction of arrival of this species in Pomerania main direction of arrival ofthis species to Poland

Initial phaseof spread: simultaneous occupation of new localities in the north and in the western part of the country as well as gradual migrationofthe species to the east ^ spread nearformerly occupied localities the main migration front

Subsequent phases of spread: further increase inthe density of localities, mainly in Central Poland, andfurthermigration to the east ***«^ probable direction of further spread

Fig. 51. Recorded history of the spread of Anthoxanthum aristatum Boiss. in Poland - an ехатріе of a species which increases its rangę in an easterly direction Start ofspread: second halfofthe 19'" century: ® first ephemeral accidental introductions near W arszawa (R o stafiń ski 1872); subsequent records: Warszawa ED16, ED26 (Cybulski 1895), environs of Pilawa FD50 (Trzebiński 1930) and Brześć GD14(Pac zo ski1900) ^ directions of arrival of this specles to Poland with railwaytransport

A B C D E F G

A B C D E F G Initial phase ofspread: appearance of new localities in the eastern part of the country as well as migration of the species to the west as a resultof accidental introductions with railwaytransport: subsequent “jumps’’ are linked to the main railway lines (see also the text) direction offurther migration

Subsequent phases ofspread: further increase in the density of localities, mainly in the east ern part of Poland, and further migration to the west Ąmtrn probable direction offurther spread

Fig. 52. Recorded history of the spread of Artemisia austriaca Jacq. in Poland - an example of a species which is enlarging its rangę in a westerly direction, mainly along railway thoroughfares Poland: the oldest reports mention Warsaw (Ro appear in Belgium, Bułgaria, the Czech Republic, stafiński 1872) (Fig. 52). These stations seem to Denmark, Finland, France, Germany, Great Britain, have an casual naturę, sińce Łapczyński (1882) Hungary, the Netherlands, Norway, Poland, did not confirm the occurrence of this species and Romania, Slovakia, Switzerland, Sweden, and the reports it as extinct. It was probably introduced former Yugoslavia. It is also reported as an epe- accidentally after the commissioning of the rail- cophyte (for terminology see Fig. 2 in Chapter way line: the first railway station was launched 3) growing in Estonia (B randes 1992a) and the in Warsaw in 1845; the Petersburski Railway Sta Ukrainę (B urda 1997; Protopopova & Shevera tion was open in 1862, while the Terespolski 2002). Outside Europę, it has been introduced Station followed four years later (Kwiatek & accidentally into North America (Jehlik 1998). Lijewski 1998). Cybulski (1895) reported another History of spread: accidental introduction in Warsaw. Trzebiński Europę: this species was recorded in central (1930) describes subsequent sites: in 1898 from Europę as early as in the beginning of the 19th the Pilawa town; 1910 in Siedlce and in 1922 in century (Hegi 1935-1961; cf. also Appendix A). Puławy. The plant disperses mainly along railway M eusel et al. (1965) published the map of the lines. The “jump” by this species from these sites generał rangę in Europę, giving the earliest dates to Silesia can be also attributed to railway trans within the secondary rangę: western Russia: years port and economic links between what was then 1720, 1790; Southern Scandinavia: 1780; Den Poland and Russia. Kornaś et al. (1959) classify mark: 1790; western Europę: 1814, 1862; En- this species as part of the group of the so-called gland: 1880; and Poland 1888 (date reported for “railway specialist”. Information on the occur Poznań, by Pful after Krawiecowa 1951). Prob rence of this species in railway stations is also ably introduced accidentally by the Russian Army reported by other authors, e.g. Urbański 1958; into France (Paris area, around 1814), Denmark Rostański K. 1960; Nowak 1997. and Germany (Krawiecowa 1951). In Saxony, it Habitats: dry ruderal sites, railway tracks and was recorded in 1867 (Hardtke & Ihl 2000 after embankments, roadsides, grass swards. Wiinsche 1875). In the eastem part of Central Dynamics: the species is gradually extending its Europę its occurrence is concentrated in river rangę towards the west. It survives in many old valleys, e.g. Main, Tauber, Rhine, and Meuse. stations and emerges also in new ones (the num- There, it is one of the species extending its rangę ber of the latter increased particularly in the using roads, rivers and canals channels for mi- 1960s and 1970s). However, the intensity of the gration (Brandes 1991) (cf. Chapter 9.2). expansion is fairly Iow, probably because of In some European countries (e.g. in the Czech features of the biology of its development (Sud- Republic and Slovakia) it qualified as expansive n i k -W ó jc ik o w s k a 1987a). Currently, the overall “ąuarantine weed” spreading on meadows and number of stations exceeds 370 (in 217 ATPOL pastorał land (Jehlik 1998). sąuares) (cf. Appendix A) (Fig. 52). Poland: the first stations registered in Poland go back to the second half of the 19th century from Bunias orientalis L. Pomerania, Lublin Upland, Eastem Carpathians Warty-cabbage and Lower and Upper Silesia. The localities for Brassicaceae the first reported stations suggest two dispersion routes for this species in the eastem part of Po Biology: a perennial plant producing great num- land: it is a species expanding its rangę from east bers of seeds dispersed through anemochory, to west (anthropogenic stations prevailing) and exozoochory (birds, horses), autochory or anthro- via long-range transport, both by land and sea. pochory. Krawiecowa (1951) states that seeds were mostly Native rangę: eastem Europę and western Asia. transferred with fodder and other seed transport. It probably originated from Armenia where it It was also sporadically grown as a fodder plant. grows at an altitude from between 1000-2500 m Perhaps this species had been previously intro a.s.l. up to the sub-alpine vertical zone. From there duced into Poland, a theory which is supported the species dispersed in the European part of the by the existence of relatively numerous stations former Soviet Union, as far as the southem bound- spread all over Poland dated from the second half aries of western Siberia. It grows in forest and of the 19lh century (Fig. 53). The seedlings of this forest-steppe formations, less often in steppes, in species have been tentatively described from the boundaries of fields, in unusable areas and Tuma near Łęczyca and in fossil layers from the ruderal places (Jehlik 1998; Fedorov 2001). early Middle Ages (Sychowa 1985); in addition Secondary rangę: occurs mostly in central and several pieces of information are provided by western European countries. It is known to archebotanical data from Gdańsk. Start ofspread: second half of the 19 century or earlier (suggested by a relatively large number of localities spread out over the whole territory of the country) ® earliest registered record: Gdańsk DA80 (KLINSMANN herb. TRN); subsequent records: Olszanica FG27 (K napp 1869), Lublin FE27 and Chełm GE34 (Rostafiński 1872), Mysłowice DF43 (Uechtritz 1877) as well as localities in Lower Silesia e.g.: Wrocław BE49 and Wielowieś near Wołów BE24 (Uechtritz 1879)

A B O D E F Q

A B C D E F S Initial phase ofspread: appearance of new localities in regions fromwhich the species had been previously recorded as the result of accidental introductions of the species (from various directions, especially from Germany) as well as its migration from the east ^ spread near formerly occupied localities direction of arrival of this species to Poland

Subsequentphases ofspread: further spread from previously occupied localities (in the recent period mainly along railway lines and automobile roadways) as well as gradual rangę increase in the western direction

Fig. 53. Recorded history of the spread of Bunias orientalis L. in Poland - an example of a species using two modes of spread during the increase in its rangę: gradual migration and long-range transport Habitats: roadsides, wastelands, surrounds of tions. Lohmeyer & S ukopp (1992) report this cottages, rubble heaps, railway tracks and em- species as a newcomer (neophyte) spreading bankments, also fields, boundary strips and fal- along the Rhine. low lands; also found in meadows. A species Poland: Its first station in Poland was recorded characteristic regionally for the alliance of Ono- by Grabowski (1843) and Wimmer (1868) in pordion associations (Sychow a 1985). M atusz 1838 and it was then reported by Fiek (1881). In kiewicz (2001) indicates it as a species which the second part of the 19th century this species distinguishes the communities Falcario vulga- was known to grow in 7 stations dispersed in lo- ris-Agropyretum repentis from the class Agro- calities located on the Oder river: in Nowa Wieś pyretea intermedio-repentis. In Central Europę it Wrocławska (W immer 1868; Fiek 1881) and is classified as a species associated with the Arte- Pruszków (F iek 1881); it was also reported in misietea and Molinio-Arrhenatheretea classes of Kraków, on the Vistula river (K napp 1872), in associations (Brandes 1991). Puławy (R ostafiński 1872), Warszawa (C ybul Dynamics: Up until the mid 20th century the ski 1894) and in Bydgoszcz (B ock 1908) (Fig. 54). number of recorded stations gradually increased Rostafiński (1872) still considered it a very rare (cf. Appendix A). An evident increase was noted species. At the beginning of the 20th century, in the last half-century where the species in Szafer (1919) stated that this species is natura- creased from 120 to 1353 stations situated in 567 lised in Silesia. ATPOL sąuares. Currently, it is distributed Habitats: wastelands, roadsides, poorly-managed throughout Poland, including lowland sites in the sites, cracks between flagstones, sport stadiums, Carpathians, e.g. it occurs in massive numbers in railway platforms, storage sites covered with slag, the Zakopane Basin (Piękoś-Mirkowa & Mirek railway tracks. 1978), and fairly often in the Bieszczady Mts. A characteristic species for associations of the (630-740 m a.s.l.), where it has also moved into Eragrostion and Panico-Eragrostietum alliances natural habitats (Zemanek & Winnicki 1999). It (Balcerkiewicz et al. 1999; Matuszkiewicz 2001). is found rarely in the Ciężkowice Foothills (Kor- Dynamics: an evident increase in the number of naś et al. 1996) and Beskid Żywiecki Mts. stations occurred as early as in the first half of where it reaches elevations up to 560 m a.s.l. the 20th century (Fig. 54). In recent times it has (B iałeck a 1982). The species is still expanding been recorded in 1041 stations in 581 ATPOL and is commonly and freąuently found in some sąuares (Tokarska-Guzik 200la; cf. also Appen- regions of Southern and south-eastern Poland dix A). Distributed throughout the national ter- (Fig. 53). ritory, it occurs more freąuently in some regions. It is still colonising new sites, expanding particularly in towns (probably introduced with sand Eragrostis minor Host [syn.: E. poaeoides Р. B.] during pavement renovations). Smali Love-grass Poaceae Rumex confertus Willd.

Biology: annual plant, its smali grain seeds are Russian Dock dispersed by wind and animals. Polygonaceae Native rangę: south-eastern Europę and western Asia. Biology: perennial, dispersing throughout wind Secondary rangę: central and western Europę and animals (by exochory) and by water. (also Great Britain). Native rangę: eastem Europę and central Asia; probably already native in areas along the Dnie- History of spread: ster; it reaches the Tomsk areas in the east (Tacik Europę: it appeared in the central part of 1992). Europę probably in the early 19,h century and at Secondary rangę: Central Europę, towards the that time it also arrived in Poland (K ornaś western part; also recorded in British Isles in 1968b). It was transported with wool, grains, Kent (S tace 1997). In northem-eastern Europę it fodder and hay. To-day, it occurs in a number has spread to Lithuania where it is considered an of regions, but mainly in urbanized and railway invasive species (Gudżinskas 1999b). areas. L a n d o lt (2000) describes the rapid ex- History of spread: pansion of this species in Ziirich, where it was Europę: It was E ich ler and Łapczyński (1892) recorded for the first time in the old part of the who noticed that this species was migrating from city in 1873, but it had not dispersed in a vis- the east to the west and north-west. The history ible way until 1980. By 1989, it had occupied of the expansion in central and western Europę of the 68 sąuares under study, and in the follow- is connected with the history of this species in ing 10 years it occurred at as many as 106 sta Poland (Fig. 55). In addition to the gradual mi- A B C D £ F G First record in a large city: Start of spread: accidental introductions in cities in various regions of ® Wrocław-Gajowice BE49 (W immer 1868; Fiek 1881) the country (see the text); the location of sites of occurrence along main Polish rivers suggests that in the initial phases of spread, water transport may have played an important role ^ spread nearformerly occupied localities

Initial phaseof spread: Subsequentphases of spread: appearance of new localities in regions from which stabilisation and filling in of the rangę: mainly in specific the species was previously recorded as well as new habitats (cracks in pavements and cobblestone-paved town accidental introductions in remote sites (the spatial pattern squares, gravel- or slag-lined sport grounds) in towns and in of spread ofthe species correlates with the location of urban railway areas (paved or slag-lined raił platforms and storage areas and the pattern of communication thoroughfares) sites) ^ directionsof spread

Fig. 54. Recorded history of the spread of Eragrostis minor )st in Poland - an example of a species associated with urban areas within the limits of its secondary rangę gration towards the west, it is also making use of current distribution of this species is the result of long distance transport. the migration via river valleys and railways (among Poland: the first stations of this species were re other things, the first reports on the occurrence were corded on the Bug river in the second part of the from Cracow - K ornaś (1954) and Wrocław - 19'1’ century (Fig. 55). The first stage of intemal R ostański K . (1960), where the stations were found migration in Polish territory was along rivers in railway areas) and macadam roads. Currently, this (T rzciń sk a-T acik 1963 - the author of the first species is penetrating settlements, abandoned fields distribution map of this species in Poland). The and pastorał land (F aliński 2000b).

13 The Establishment. 97 AB c D E F G Start ofspread: second halfofthe 19 century A A 1 1 ® first recorded localities in the Bug river valley: -X Zajęczniki FC98 and Łosice FD18 (KARO, herb :< i -i / : B B KRA) 7 V % direction of arrival of this species to Poland : _ C < \ C w t ) V 183 / / 1873-1 L i D E V \ D \ ( Y u E Z-<- \ E L r' > \ Y i \ S OH \ J F ! 'i-'- s F i n < V / : \ G -W 1900 / G U i u\„,m A B c DEF G

Initial phase ofspread: migration of the species along the Vistula and Bug rivervalleys colonisation of ruderal habitats adjacent to (orbetween)rivervalleys new accidental introductions of the species both in rivervalleys and on railway territory

Subsequent phases ofspread: further migration along river valleys and communication thoroughfares; transfer of the species to new types of habitats: post-agricultural wasteland, meadowsand pasture land Ąmmm gradual rangę increase in a westerly direction

(the map after Z ając A. & Z ając M. (eds.) 2001 - slightly supplemented and modified)

2000 A local example of riparian corridor migration of Rumex confertus in the Bug river valley(source:FAUŃSKiefa/.2000)

Fig. 55. Recorded history of the spread of Rumex confertus Willd. in Poland - an example of a species which has increased its rangę in a westerly direction using river yalleys and later also transport thoroughfares Habitats: semi-natural habitats: meadows, ripar- river basin belonged to a united Poland and ian scrub and ruderal sites: roadsides, railway Lithuania, which were at that time under the tracks and embankments, also in rubble heaps and same political rule (Davies 2001). For some two around cottages. centuries the trade in grain significantly boosted Dynamics: in the last 180 years, this species has the whole economy of the Polish Republic established itself in south-eastern and central (.Rzeczpospolita). Early stations for this species parts of Poland: it is still scattered in the north were located along the Vistula river and its tri- and west (in these regions it might still be an butaries: the Bug and San, which supports the ephemerophyte). In the Carpathians it occurs hypothesis that the first cases of the dispersion frequently in the Beskid Niski Mts. and Western of this species in Poland took place along the Bieszczady Mts., where it reaches elevations of Vistula trade route. The oldest date refers to 690 m a.s.l. (T a c ik 1992; Z e m a n e k & W in n ic k i Gdańsk, a city located strategically at the end 1999). The species is expanding its distribution of the Vistula trade route, which was used for area throughout Poland. In the last half-century, grain exports even in the 17th century. At the the number of recorded stations increased from same time, Gdańsk was connected by a compli- 47 (in 37 ATPOL squares) to 1731 (in 673 cated river network with inland areas. Ali the ATPOL sąuares) (Fig. 55; cf. Appendix A). main tributaries of the Vistula: the Narew, Pili ca, Bug, Wieprz, Wisłoka, Dunajec and San Salsola kali L. subsp. ruthenica (Iljin) Soó were suitable for water transport. Ali the tri butaries had their own ports with storehouses Spiny Saltwort, Prickly Saltwort and shipyards. In the 18th century, the Vistula Chenopodiaceae area was connected with the Warta and Oder via Biology: annual plants dispersing by fine, wind- the Bydgoski Channel (1771), with Prypeć and dispersed seeds. Dniepr by the Królewski Channel (1775-1784), Native rangę: southem part of Russia, Caucasus, and with Szczara and Niemen by the Ogiński Siberia and central Asia. In its native country it Channel (1765-1784) (Davies 2001). These con- grows on sand, steppes and riverine cliffs (Ko- nections made possible the subseąuent stages of m a r o w 1943-1964). the migration of the species which, when the Secondary rangę: central and western Europę. railway developed, started migrating along new routes. The first stations where this species was History of spread: introduced accidentally via the railway transport Europę: According to H eg i (1963-1983) it was were Szczakowa, Lublin and Wrocław (Fig. 56). introduced accidentally to central and western At that time, Szczakowa (the current district of Europę, probably with wool and other raw ma- Jaworzno town) was a railway junction station terials at the beginning of the 19th century. and, at the same time, an Austrian boundary sta H a r d t k e & I h l (2000) found an earlier date of tion serving both directions: to Prussia and the occurrence of this species in Germany, Russia34. In addition, such a pattern of expan- namely 1775. sion is confirmed by early, freąuent reports on Poland: B a r a d z ie j (1972) reports the oldest Po- the occurrence of this species on the Vistula lish stations as existing in the second half of the river, and less frequently on railway areas as 19th century. She was the author of the first map previously noted by Sudnik-Wójcikowska of the distribution of this species in Poland. She (1987a). The earlier accidental introduction can states that the species migrated to Poland by grad- be also contributed by the then trade in salt ually moving further and further westwards. which was developed by the Cistercian monks. B a r a d z ie j (1972) also noticed that the stations of Habitats: inland sands and sand dunes in the Salsola kali subsp. ruthenica clearly occurred interior part of Poland, fields, ruderal sites, road along the valleys of large rivers and on railway sides, wastelands, heaps on industrial properties, lines. and railway tracks and embankments. An analysis of the distribution of the stations A characteristic species of the Salsoletum ru- from the earliest periods of the dispersion of this thenicae association and a distinguishing species species in Poland shows the primary link with for the Corispermo-Brometum association (M a the Vistula river valley (Fig. 56). It can be tuszkiewicz 2 0 0 1 ) . supposed that the Spiny Saltwort was introduced accidentally into the valley even earlier: maybe in the period of trade in grains. The commercial 34 In 1847, the Katowice-Kraków (Cracow) railway line was built across Szczakowa, and in 1848 it was linked with route on the Vistula river was established as Warsaw-Vienna route. In Lublin, the first railway line was early as in the mid-15th century, when the whole opened in 1877 (K w ia t e k & L ij e w s k i 1998). Start ofspread: Initial phase ofspread: first half of the 1T century or earlier (see also Chapter 5.2) migration of the species along the Vistula river ® first recorded localities: Gdańsk DA81 (S chw arz 1967 valley after Oelhaf); Warszawa ED26 (Sudnik-Wójcikowska first localities in which this species appeared 1987a after Erndtel) probably accidentally introduced with railway transport are Wrocław BE49, Szczakowa DF45 and Lublin FE27 probable directions of arrival of this species to \ Poland

А В C D E F G Subsequent phases ofspread: Current distribution of this species reflects the course of further migration along river valleys of the Vistula and its main river valleys as well as communication pathways tributaries simultaneous spread along communication / pathways (especially migration along railway lines)

Fig. 56. Recorded history of the spread of Salsola kali L. subsp. ruthenica (Iljin) Soó in Poland - an ехагпріе of a species which used the Yistula pathway as a “conveyor belt” for further spread

D ynam ics: the species has gradually invaded ment of the Polish flora and its distribution new sites (Fig. 56). In the last 50 years the num- closely reflects the pattem of river valleys (prin- ber has increased from 114 to 901 sites recorded cipally those of the Vistula and its major tribu in 467 ATPOL squares (cf. Appendix A). Cur- taries, and of the Lower Oder river), as well as rently, the species constitutes a permanent ele the outlines of the railway network. 7.3.2. Plants brought accidentally from fied this species as rare, occurring in the north Asia em, central and southem parts of Poland, except for the mountains. Currently, it occurs frequently throughout most of Poland, again except for the Sisymbrium altissimum L. [syn.: S. sinapistrum mountains, and has become common in some Crantz] regions (Fig. 57). In the last half-century the Tali Rocket number of stations recorded increased to 1770 in Brassicaceae 812 ATPOL squares (cf. Appendix A). Gradually invades new sites. Biology: annual plant, perennial in rarer cases, producing many siliąuas. Diaspores (seeds, Veronica per sic a Poir. [syn.: V. Tournefortii Gmel.] fruits or even whole plants - so-called “tum- Common Field-speedwell bleweed” plants) disperse autochorically and hy- Scrophulariaceae drochorically. Native rangę: Asia and south-eastem Europę. Biology: annual plant producing great numbers Secondary rangę: remaining part of Europę and of seed dispersed through wind, water or ants. North America (S y c h o w a 1985). Native rangę: mountains of Asia Minor, north History of spread: em Iran and western part of the Himalayas (M e u Europę: its presence in central and northem Europę s e l et al. 1978). was recorded in the second half of the 18* century Secondary rangę: Central Europę (in the Alps (M eusel et al. 1965), where it was probably intro- up to an altitude of 1600 m a.s.l.), central Asia, duced accidentally with the ballast from ships. North and South America, Southern Australia, Poland: Kornaś (1968b) supposed that the spe- New Zealand and New Guinea. cies arrived in Poland before the end of the 18th History of spread: century. It is confirmed by the oldest dates for Europę: finding this species (Fig. 57). This Iran-Turanian 1885 - this, the oldest date for central species was probably introduced to Gdańsk via Europę reported by M e u s e l et al. (1978), was ballast and grain (Preuss 1928). Its oldest stations recorded in the Karlsruhe botanical garden. A go back to the first half of the 19th century and subsequent date - 1809 - is reported by P y ś e k they were located in the northern part of Poland: et al. (2002). In the British Isles it was recorded in Gdańsk and Toruń. Subseąuent stations dating for the first time in 1825 (S t a c e 1997). On one from the second half of the 19th century were also hand, the species migrated from east to west, and located in the northern part of Poland: in the on the other, it was introduced accidentally into Malbork area (Klinggraeff 1854), in Chełmno various parts of the continent. (Abromeit et. al. 1898 after Wacker 1861), Bra Poland: in the second half of the 19th century it niewo, Bydgoszcz and Kwidzyń (Klinggraeff was known to have numerous stations (Fig. 58), 1866) and in Czarna Grobla near Braniewo which is why it can be supposed that it had been (1868). This species was introduced accidentally introduced accidentally before this time, which is into the interior of Poland by railway transport, also suggested by data from various European to Poznań, Szczakowa (Rehman 1879) and to the regions. The oldest stations are dispersed towards Warsaw area (Sudnik-Wójcikowska 1987a, on the the north and south-east, so allowing the pre- basis of herbarium of Cybulski). sumption that this species was introduced acci Habitats: ruderal weed, found in railroad tracks dentally from various directions: from east and and embankments, industrial sites, wasteland, north, via marinę transport. In the subseąuent rubble, roadsides, and lawns. Regionally reported half-century it dispersed around previously occu- as a characteristic species of the Onopordeta- pied stations, and in following 50 years it occu- lia acanthii order (S y c h o w a 1985). In the clas- pied the remaining parts of Poland. sification published by M atuszkiewicz (2001), it Habitats: cultivated fields, former farmlands, is a characteristic species of the Sisymbrietum garden allotments, ruderal sites (particularly in loeseli association. It also occurs sporadically in moist, shadowed sites). semi-natural communities, e.g. at the edges of A characteristic species for the Polygono-Che- pine-oak stands, and - more often - in xerother- nopodion alliance of associations (M atuszkiewicz mic grasslands. 2001). Associated with soils of high or medium Dynamics: Up until the year 1950 it had 59 level of soil fertility. Often, it appears particularly stations. Their number has begun to grow in the as a weed in cultivated fields in several cultivar 1960s, with a remarkable increase noted in the systems on several types of soils suitable for last 30 years. Sychowa (1985) originally classi- cereals and fodder crops. It occurs, among others, Start ofspread: Initial phase ofspread: first half of the 18m century or earlier (see also Chapter 5.2) ^ spread of the species in the vicinity of sites of initial ® first recorded localities: Gdańsk DA80 (Klinsmann ' accidental introduction (most probably together with 1843) and Toruń DC30 (Klinggraeff 1848) long-range transport of goods) ~n probable directions of arrival of this species to Poland y with marinę transport

A В С D ,.,,Е F G Ш ГИ 'Н"І'"Н"НІ ! І ! І І І І ! І(І І ІИ і II ітіті m г| ікцтгтп 7 A Г ’ґ{ А 4 . j l J 7 Г к , W / І ^ в : ї? \{ в А ' \ л г ~ \

C : : С ( k V /■ V- ...1 S' \ З . л f D ч < D г " ' 'Л І \ ) E j - L) х.; е V > П J \ V. ^ \ / і : / F : r s p V Л: V 7 \ У \ G : 1 9 5 0 V ') ) у ...... 4.М І.іі.аґш 1 1 1 1 1 1 і 1 г Л ш ш т A в с DЕ F а Subsequentphases ofspread: Current distribution of this species shows its association with urban areas and communication pathways. The more ^ further spread from occupied localities common occurrence of the species in the western and —*- consecutive accidental introductions of the species central parts ofthe country can also confirmthe hypothesis with transport of goods (mainly from the area of thatthe mainmigrationfrontwentfromwesttoeast (the map Germany); simultaneous migration of the species after Z a ją c A. & Z a ją c M. (eds.) 2001 - slightly supplemented) from the south-east

Fig. 57. Recorded history of the spread of Sisymbrium altissimum L. in Poland — an illustration of a dominant role of humans in increasing the rangę of a species

in the Yicietum tetraspermae communities, ac- number of stations increased from 84 to over companying winter cereal crops and rape fields 7800, recorded in 2204 ATPOL squares (cf. (W archolińska 1999). Appendix A) (Fig. 58). The species is still ex- D ynam ics: massive expansion of this species panding although the invasion is limited to was recorded in the last half-century when the segetal and ruderal habitats. Start ofspread: second half ofthe 19'” century orearlier(see also Chapter 5.2) ® first recorded localities: Western Pomerania: Chełmno and Świecie CB99 (A b ro m e it etat. 1898 after W a c k e r 1862); Bydgoszcz CC26 (Klinggraeff 1866); Strzelce near Bydgoszcz (K u h lin g 1866); Mazovia-Podlasie Lowland: Warszawa ED16 (Lap- c zyń s k i herb. UW); Polinów near Łosice FD28 (K a ro 1867); probable directions of arrival of this speciesto Poland

A B C D E F G

Initial phase ofspread: ^ spread of the species in the vicinity of sites of initial ' accidental introduction arrival to a new locality most probably together with \ long-range transport of goods

Subsequentphases ofspread: sudden and massive occupation of new localities (mainly in anthropogenic habitats); simultaneous migration of the species fromthesouth-east (the map after Z a ją c A. & Z a ją c M. (eds.) 2001 - slightly supplemented and modified)

Fig. 58. Recorded history of the spread of Yeronica persica Poir. in Poland - an example of invasion by putative repeated accidental importation and simultaneous rangę expansion 7.3.3. Plants brought accidentally from migration of Bidens frondosa along railway tracks. America At that time it was dispersing southwards and eastwards. In Brześć on the Bug, it was discovered in 1955, while Sokołowski (1967) recorded it in the Bidens frondosa L. [syn.: B. melanocarpus Wiegand] Białowieża Forest in 1965. Since 1970, it has been

Beggarticks reported in the Ukrainę (Dr M. S hevera, pers. Asteraceae comm.) where it occurs in urbanized areas (B urda 1997; Protopopova & Shevera 2002). Biology: arrnual plant reproducing generatively. Habitats: banks of inland waters: carrs and ri- Seeds are dispersed in water and by animal and parian alluvia, drying-up margins of lakes and human agents. The warty surface of the fruits ponds, cultivated fields and moist ruderal hab with a crown of awns is covered with downward- itats: roadside ditches, railway tracks and sta pointing hooks facilitating their dispersal. tions, also rubble heaps. It is a component of the Native rangę: North America between the Atlantic therophytic communities of the class Bidentetea and Pacific, from New Foundland and Southern and of forest, Coastal scrub and reed communi Saskatchewan to the north, to Colorado, Califomia ties (Salicion, Phragmition, Glycerio-Sparga- and Мехісо in the south (T r z c iń s k a 1961). nion). A characteristic species of the communi Secondary rangę: Europę and eastem Asia. ties of the alliance Chenopodion fluviatile (M a tuszkiewicz 2 0 0 1 ) . History of spread: Dynamics: to date it has been recorded in 3142 Europę: it appeared in the 18th century in various stations in 1068 ATPOL sąuares (cf. Appendix parts of the continent: France (Botanical Garden A). Distributed throughout Poland, common in in Montpellier) 1762; Italy 1834, 1849, 1861; the Oder and Vistula river valleys as well as along Portugal 1877; and Germany 1894 (Kornaś et al. their tributaries (Fig. 59). In the mountains it 1959; Trzcińska 1961); information on its occur- occurs at lower sites (B iałeck a 1982; K ornaś et rence along the Oder river was reported in 1777 al. 1996; S zelą g 2000), e.g. in the Beskid (K rocker 1790; GruberovA et al. 2001). Loh- Żywiecki Mts. it occurs up to 455 m a.s.l. m e y e r & Sukopp (1992) report the earliest date of (B iałeck a 1982). Still expanding. occurrence of this species in central Europę, i.e. 1736. Currently, it is widespread throughout the Chamontilla suaveolens (Pursh) Rydb. [syn.: continent. UherćikovA (2001) reports it as an in- Matricaria discoidea DC.; M. matricańoides (Less.) Porter] vasive species on the Danube in Slovakia. Poland: probably migrated from Germany to South Pineappleweed ern Poland (it spread in Silesia, where it dispersed Asteraceae along the Oder) and northem Poland (it was known in Pomerania sińce 1897) (Fig. 59). In Poland it was Biology: Annual plant dispersing anemochorically, reported for the first time in Wrocław, by the Oder zoochorically (by endo- and exozoochory) and (Krocker 1790). As late as in 1869 it was found anthropochorically (accidentally introduced via again by Brand, by the Oder, downstream of the transportation over land and water). river at Słubice (Schumacher 1942). Since then the Native rangę: north-west America and eastern occurrence of this species was reported by Graeb- Asia where it grows on the river banks and val- ner (1897), who reported its occurrence in Łęcze leys and on the coast in humid and sandy places near Elbląg and by Ascherson (1898), recording it (S u d n ik - W ó j c ik o w s k a 19 8 7a). by the Vistula river in Ciechocinek. Fiek found this Secondary rangę: Europę. species on the bank of the Oder, near Głogów History of spread: (Schumacher 1942). Beggarticks migrates via two Europę: many authors State that this species routes: along watercourses where it disperses by hy- appeared in the early 19th century. The oldest drochory and epizoochory and as an antropochorous recorded date - 1850 - was published by M eusel plant along railway tracks (Kornaś et al. 1959). The et al. (1992) for Scandinavia. A subseąuent station description of this migration in the earliest part of in the former Czechoslovakia was reported by the invasion was given by Trzcińska (1961), who Pyśek et al. (2002). In 1852, it was found by Braun developed a distribution map on the basis of 101 in the Berlin area: this author considered it a refiigee reported stations. By then it was already a species from the botanical garden (Kamieński 1884a). well established along the Oder river and the Another station was found in Southern Scandina- Vistula basin: downstream in the Toruń and Byd via in the same year. Every few years new stations goszcz areas and upstream in the Kraków area. The of this species are reported in other parts of Eu stations in Upper Silesia and detached from Brześć ropę. The first report on the occurrence of this belong to the group of locatities associated with species in Britain goes back to 1871 (Stace 1997). A B c DE FG Start ofspread: ...... second halfofthe 19"1 century orearlier(see also Chapter5.2) A —'------^ A V p — Hf j ® first recorded localities: Wrocław BE49 (K ro c k e r , ’’ i,‘ i. \ *v / - i' s 1790); Słubice AD02 (Schumacher 1942); Łęcze near - ■4N. t B B Elbląg DA96 (G raebner 1897); Ciechocinek DC51 (Ascherson 1898); Rapocin near Głogów BD82 (Fiek %j i r’ \ r ’ & Schube 1898) ( __ ___A C S <8>1897 : C spread ofthe species nearthe sites ofinitial accidental "w. S-' ' introduction : D ) > J D \ V \1898 i. ( y \ 1777 / f : __ 1 E E '■ \ L k ? ( )\ \ J 7 \ : F : F / "U \ )/ i

/ J

: I G 1900 G \ j r -AS

: : \ xv { A B c D E F G

Initial phase ofspread: -*—-s migration along river valleys, especially along the Odra river first transitions of the species to ruderal habitats \ outside rivervalleys

Subsequentphases ofspread: rapid occupation of new localities: migration along and across river valleys (transition from riverside habitats to ruderal habitats). The current distribution reflects the course of major rivervalleys. (the map after Zająca. & Zając M. (eds.) 2001 - slightly supplemented and modified)

A localexampleofripariancorridor migration of Bidensfrondosa \ in the Bug river valley (source: Fauński etal. 2000)

Fig. 59. Recordcd history of the spread of Bidens frondosa L. in Poland - an example of a species using river valleys in its migrations (migration along and across valleys) However, the oldest studies by G udżinskas (e.g. from Kiev) in 1869; the same author also found ( 1997d) indicate that this species arrived in Eu this species in Białystok and Brześć, and outside ropę earlier, sińce its occurrence was reported by Polish borders in the Mińsk area in Wołyń (where Gilbert in Grodno (Bielarous) in a paper from this species had occurred before the date reported 1782 and in addition it was also mentioned by by G u d ż in sk a s (1997d)). P a c zo sk i (1900), in Jundziłł (1791, 1811) and G órski (1830). a subseąuent publication describes this species as Poland: recorded for the first time in the 19lh a common species in Polesye, in railway facili- century, initially in Lower Silesia (1862), and ties and built-up areas. Due to the fact that the next in Tarnów (1871) and the Kraków area plant was usually found near railway stations the (1878) (Fig. 60). Kamieński (1884a) in his de- author attributed its occurrence to the develop- scription of this species based on the occurrence ment of the railway lines. The invasion by this in Warszawa, stated that “it is likely that this new- species commenced in Poland at the end of the comer to our flora will disperse very ąuickly and 19th and beginning of the 20lh centuries. it soon will grow outside Warszawa”. The spe H abitats: distributed in wastelands, along trans cies dispersed at a fast pace. When Raciborski port routes and in cultivated fields. Associated ( 1885) reported it for the first time (in 1878), this particularly with trampled sites and the initial species was present and abundant at several sta- stages of ruderal communities. tions in Kraków. Paczoski (1895) reports that this D yn am ics: even though it is not one of the plant had been known to the east of Warszawa group of the oldest kenophytes, it has colonised

Start ofspread: secondhalfofthe 19'" century or earlier (see also Chapter5.2); plant introduced accidentally to many regions simultaneously (mostoften in cities); maybe initially planted in bota- nicalgardens ® first recorded localities: Wrocław BE49 (UECHTRITZ herb. WRSL; KNEBEL herb. WU); Tarnów EF67 (Heger1871); Kraków DF69 (Raciborski 1885)

A______В______С______D______E______F______Є

Subsequent phases ofspread: rapid and massive colonisation of anthropogenic habitats in the whole area of the country (the map after Z ając A. & Z ają c M. (eds.) 2001 - slightly supplemented and modified)

Fig. 60. Recorded history of the spread of Chamomilla suaveolens (Pursh) Rydb. in Poland — an ехатріе of a species which became the most common kenophyte in the Polish flora as a result of invasion the entire national territory of Poland in the last History of spread: 150 years (Fig. 60). It is distributed throughout Europę: the naturalisation of Canadian Water lowlands, also in mountain areas as far as the weed in Europę started in the first half of the foothills. The species has also spread rapidly 19th century. The first record in 1836 com es across the Carpathians, being reported from from Ireland (K amieński 1879, 1884a & b; N ow y Sącz (Pawłowski 1919 herb. KRAM ) and D yakowski 1899; S tace 1997). Next, its occur- other localities in the region (P awłowski 1925). rence was recorded in the United Kingdom: in In the Pieniny Mts. from where it was first re 1842 in Scotland and in 1847 in central England. ported by Kulczyński (1928), it occurs freąuently In 1840 it was brought to the Berlin botanical in villages and along roads, in the periphery of garden, where due to its excessive growth some this mountain rangę (G uzikowa 1972). In the was thrown away into the river (K ucharski Tatra Mts. it was found at an elevation of 1500 1992). The peak of its rapid expansion in the m a.s.l. near the Murowaniec mountain shelter central Europę occurred in the period 1859— (Piękoś & M irek 1974), and on the Slovak side 1935 (Lohmeyer & Sukopp 1992). In the British of the Tatra Mts. even as high as at 1770 m a.s.l. Isles, after the mass invasion in the 19th century (R adwańska-P aryska 1963), and later at 1815— and the first half of the 20th century it withdrew, 1830 m a.s.l. near the mountain shelter and ca- superseded by E. nuttallii, also a North A m er ble car station in the surroundings of the Skraj ican species. ne Solisko site (P iękoś-M irkowa & M irek Poland: the first report from Poland (Gdańsk) 1978). In the Karkonosze National Park, the dates from 1867 (A bromeit et al. 1898). The species reached 1540 m a.s.l. occurring along plant was found for the first time in the Vistula River in (D yakowski stations of the road leading to the Śnieżka Mt. (Rostański K. 1876 1899). 30 1977). In the massif of the Śnieżnik Mt. and in the this plant were recorded in then Polish (Congress) Kindom in the period while in Grand Bialskie Mts. it is freąuently found at lower 1878-1897, Duchy of Poznań it was recorded in all counties mountain sites (S zeląg 2000). In the Bieszcza (B łoński 1899). Łapczyński (1882) describes this dy Mts. it is found only occasionally within species and presents a drawing of a specimen ruderal habitats at elevations of 640-750 m a.s.l. collected in Warsaw, and in 1887 reports its (Z emanek & W innicki 1999). In the Mt. Pilsko occurrence in ditches in Sandomierz. A number massif (Beskid Żywiecki Mts.) it was not very of occurrences of this species in many Polish re- freąuent in the early 1980s, being recorded from gions was published in the following years: sites not exceeding 520 m a.s.l. (B iałecka Pomerania and Masuria (A bromeit et al. 1898 1982). In the last two decades, it spread across after Scharlok 1884), in western and Southern the Carpathian Foothills and lower sites in the parts of Poland (U echtritz 1876, 1880; Krupa Carpathians, e.g. in the Ciężkowice Foothills 1882; Raciborski 1884) and in Masuria (R osen- where it occurs freąuently as a component of the bohm 1879; K amieński 1879; Łapczyński 1882) regional flora (K ornaś et al. 1996). (Fig. 61). The species continues to colonise new sites. It Habitats: lakes, old river beds, rivers and stream is one of the most common kenophytes occurring with slow current(s), ponds, clay pits, channels, in Poland, for which data have been obtained drainage ditches. from 13125 stations in 2965 ATPOL sąuares (cf. It occurs in water communities from the class Appendix A and Chapter 5.2) (Fig. 60). Potametea and less freąuently in communities from the alliance Phragmition (K u c h a r s k i 1992 Elodea canadensis Michx. [syn.; Anacharis ca- and literature cited). In eutrophic water this suc- nadensis Planch, E. canadensis Rich.; Helodea\ Philotria ca cessful invader builds its own community Elodee- nadensis Britton.] tum canadensis (M atuszkiewicz 2001). Canadian Waterweed Dynamics: the contemporary distribution map Hydrocharitaceae shows that Canadian Waterweed is abundant on most of the national territory of Poland (except Biology: spreads only through vegetative pro- for mountain regions), although it no longer cesses owing to the fact that only female individ- shows any evident invasion. To date it has been uals were introduced into Europę. The fragments recorded in 3681 stations in 1847 ATPOL sąuares of shoots are transported by water, birds and hu- (cf. Appendix A). mans. Native rangę: North America Secondary rangę: Europę (including northern Scandinavia), New Zealand, Australia (new south Wales, Yictoria), Africa (Fedoroy 2001). Start and initialphaseofspread: mid-19'" centuryorearlier(seealso Chapter5.2); by the tum ofthe 19"1 century, this species already had 140 localities in the waters ofthe Vistula, Odra and Warta rivers as well as in the lakes ofthe Lake Districts ® the first recorded locality: Gdańsk DA80, dates from 1867 (Abromeit et al. 1898). Note that the map describes localities recorded up to 1880.

Subsequent phases of spread: rapid occupation ofnew localities: ^ migrationalong ^ and across river valleys (including by means of ' smaller watercourses, old river beds and water reservoirs)

A B C D E F O The current distribution reflects mainly the presence of water habitats

(the map after Zając A. & Zając M. (eds.) 2001 - slightly supplemented and modified)

| A local example of riparian corridor migration of Bodea \ canadensis in the Bug river valley (source. Fauński etal. 2000) A B C______D______E______F______G Fig. 61. Recorded history of the spread of Elodea canadensis Michx. in Poland - an example of a currently well naturalised species common throughout Poland which owes its ide distribution to water and birds PART FOUR

Discussion

8. The proportion and role of alien cies of alien origin, from 42 species reported for the species in the flora: do kenophytes European part of Turkey to 479 reported for France determine the recent shape of the (W eber 1997a). However, this data so recently flora of Poland? provided for various European countries is now out of date. The comparison made for the Czech flora by Pyśek et al. (2004) indicates that Weber’s fig In the voluminous body of literature devoted ures were an underestimate. For the Polish flora, to alien species of plants, some figures can be W eber (1997a) reported, from analysing Flora found illustrating the scalę of the phenomenon of Europaea, 184 alien species established, including their presence in respective floras. These figures 81 species originating from outside Europę. Weber’s are however usually estimates and hardly com- totals for Poland, when compared with the figures parable because of differences in the terminology obtained in the present monograph, is as for the used, as well as in the intensity and unifor- Czech Republic clearly an underestimate. mity of the research undertaken (P yśek et al. Detailed lists, compiled for Germany by 2002; Tokarska-G uzik 2003a & b; Pyśek et al. Kowarik (1999), show that from among ca. 12 000 2004). For ехатріе, for the United States of species introduced in this area, 417 permanently America, M orse et al. (1995) report 5000 species established species now constitute a part of the of alien origin established within the entire flora overall inventory of the German flora (of 3001 species in all). In Germany, most of the species of of some 17 000 species; Scott (1997) lists more than 1850 alien species for New Zealand, while alien origin colonise disturbed habitats, whereas 228 have been also recorded in natural communi Enomoto (1997) reports 1196 established species in Japan. In the Hawaiian flora, the initial esti- ties and among the latter, 30 species are regarded mate was 900 native plant species and 4000 as a nuisance and reąuiring control measures. The process of exchanging species between introduced species, out of which 870 had estab lished themselves permanently and 91 were ac- various regions of the world has been analysed by JAger (1988). His results point to a particu- corded the status of invasive species (V itousek larly dynamie exchange between Eurasia and et al. 1987). According to the so-called “Tens North America. The estimated figures show an rule” of W illiamson (1993), it is estimated that evidently higher proportion of alien species in the roughly 10% of the total number of alien species North American flora (26%), compared with is capable of permanent establishment in the new Europę (ca. 9%) (Forman 2003)35. homeland, and from among these again only 10% can establish themselves not only in anthropogenic habitats but also enter natural communities. 35 Following other authors F o r m a n (2003) points out that On the basis of Flora Europaea, W eber (1997a) species introduced in mid 19lh century from Europę into America appeared to be more successful at naturalising compiled the estimated data for the continent of than American species introduced to Europę. There are Europę. The list contains 1568 species which have three main hypotheses suggested by author for explaining either expanded beyond their previous ranges (and the phenomenon: 1) European weeds are better competi- these account for 63%) or are species originating tors than their American counterparts; 2) European plant from outside Europę (the remaining 37% which species evolved among greater disturbance than American ones, making it easier for them to establish in a newly Weber classified as “exotic species”). Individual colonised America; 3) the flow of species has been greater countries of Europę differ in the numbers of spe into America than to Europę. Again, it seems elear that the figures showing Generally, the proportions of alien species in the the proportions of alien species in floras will have three countries concemed are similar (Fig. 62). In to be periodically updated. the Czech flora, the combined number of alien The list of alien species compiled by Polish species (1378) is higher compared with Poland researchers for the Polish flora includes more (1017 species), whereas for Germany this figurę than 1000 species (M ir ek et al. 2002; cf. also is lower still (913)36. The proportions of estab Chapter 5.1; Table 3). The list of archaeophytes lished newcomers in the floras of the three coun contains 160 species. The first list of kenophytes tries concemed differ from one another, but these published in the 1960s included 117 species differences stem mainly from discrepancies in the (K o rn a ś 1968b). The list of more recent, but classification applied, as well as from the meth- firmly established arrivals, was checked 30 years odological premises made by the authors. later, and has grown to as many as 251 species The researchers who study the species of alien (Z ając A. et al. 1998). This amounts to ca. 10% origin in various floras report problems connected of the flora of Poland. It can thus be presumed with the appearance of hybrids. These are both that this flora contains a total of over 400 spe- hybrids produced by “Crossing” between two

Poland Germany Czech Republic n = 3554 n = 3656 /7 = 4133

species of species of uncertain status uncertain status 4.5% archaeophytes 4.7% archaeophytes archaeophytes 17.2%

neophytes (= kenophytes) 11.6%

casual aliens 64.7% 50.2% Fig. 62. Comparison of the strueture of the flora of Poland, Germany and Czech Republic (for more explanation see the text) cies of permanently established species (which alien species, often while already in the new represents ca. 15% of the overall flora) (T o k a r - homeland, and the hybrids developing between sk a -G u zik 2003a). alien and native species. The comparison of the floras of Poland, the Among the kenophytes occurring in Poland, only Czech Republic (P y śek et al. 2002) and Germa 25 hybrids have been found (these being mostly ny (K uhn & K lotz 2003) with respect to the pro hybrids produced by Crossing between an alien portions of individual groups of species of alien origin is made difficult because of differences in 36 In their analyses the authors considered only 207 the classification criteria. Moreover, the figures species regarded to be the most freąuent ephemeral new should be also related to geographical location comers into Germany (casual alien plants that are very common in Germany); including the rare casuals would and the natural conditions prevailing in these inerease the number of alien species greatly (K u h n & countries. K l o t z 2003). species and an indigenous one) (cf. Chapter 5.1 and al. 1998). The erosion of the genotype of native Appendices A and B). Nevertheless, both the avail- plants (via backcrossing and introgression) could able guides for the identificatłon of species as well be yet another effect of hybridisation42. as the Flora o f Po land [Flora Polski], provide The most species rich families in the Polish ample information on possible hybridisation be- flora, which also contain species of foreign ori tween alien newcomers and native species37. gin (including many kenophytes), are: Asteraceae The hybridisation of an alien species with (46 species), Rosaceae (37), Onagraceae (23), a native species can result in the production of Brassicaceae (19), Fabaceae (14) and Poaceae a fertile hybrid which is capable of spreading (14)43 (cf. Chapter 5.1.4). Similar proportions faster than its parents38. This happened in the case were reported for the Czech flora by Pyśek et al. of Reynoutria x bohemica39 ( B a il e y et al. 1996; (2002), although in a different order, resulting F o jc ik & T o k a r s k a -G u z ik 2000; M a n d Ak et al. from also adding the most recently arriving ca- 2004), which is found fairly freąuently in some sual (ephemeral) species. regions of Poland. The emergence of species of The comparisons made in this study for var- hybrid origin40 results in various problems of tax- ious systematic groups corroborate with the re- onomic naturę, but above all ecological ones (cf. ports of other authors, who have pointed out that Chapter 12). The invasive characteristics o f Rey the properties enhancing invasiveness are particu- noutria x bohemica present an ехашріе of the so- larly concentrated in some families, namely -called “invasion by hybridisation”41 (W e b e r et Asteraceae, Poaceae, Brassicaceae and Chenopo- diaceae, whereas some other families such as 37 Among others, hybridisation occurs between species Cyperaceae or Orchidaceae lack them completely44 of the genus Amaranthus', Centaurea diffussa crosses with (e.g. RejmAnek et al. 1991; K ornaś 1996; Pyśek C. stoebe, C. rhenana and C. jacea; Digitalis purpurea produces hybrids with D. grandiflora; hybridisation takes et al. 2002). place also between Diplotaxis tenuifolia and D. muralis\ The most species rich family of angiosperms, Echinops exaltatus produces a hybrid with E. sphaeroce- i.e. Asteraceae (ca. 1250-1300 genera with 20000 phalus (= E. x pallenzianus - S t a c e 1997); Epilobium cilia- - 25 000 species) is represented in floras world- tum hybridises with several other species of the genus wide, particularly in regions of moderate or sub- (some of them have been reported as common, for exam- tropical climates (T a k h t a d ż j a n 1987). This fam ple, from the British Isles - S t a c e 1997); Galinsoga ciliata produces a hybrid with G. parvijlora (G. x mixta J. ily provides most of the species of alien origin Mutr.); in Great Britain, hybridisation has been reported within the floras of various regions of the world. between Heracleum mantegazzianum and a native species Poaceae is the second most numerous family H. sphondylium (S t ew a r t & G r a c e 1984); Trifolium pa- among the flowering plants, predominating from tens produces hybrids with T. campestre, and Xanthium albinum with X. strumarium. the ecological perspective and being particularly 38 An ехашріе of a hybrid which, in some parts of important to the human economy. Moreover, this Europę, and in Poland, might be more freąuent compared group represents a major proportion of the flo with its parent species, is provided by a swarm of hybrids ras of most regions worldwide (K ornaś & M ed- described as Medicago x varia, which resulted from the wecka-K ornaś 2002; Forman 2003). In Poland, introgressive Crossing of Medicago sativa with the native grasses constitute 7.3% of the whole flora, and M. falcata. R e b e l e (1988) reported this hybrid from industrial sites in West Berlin with a markedly higher frequen- are one of the three most species rich families, cy (in 39.2% of the areas studied) while the parental spe along with Asteraceae - 12.1% and Rosaceae - cies were found less freąuently (M. sativa in 13.7% and 7.3% (K ornaś & M edwecka-K ornaś 2002; M. falcata in 11.8% of sites). Piękoś-M irkowa & M irek 2002). In all, 298 39 The introduction of taxa of the genus Reynoutria (Fal- species of grasses have been recorded in Poland lopia) into Europę has led to the emergence of a hybrid, Fallopia x conollyana J.P. Bailey, which resulted from the (Frey & Rutkowski 2002), including ca. 155-165 hybridisation of Reynoutria (Fallopia) japonica and Fal lopia baldschuanica (Regel) Hołub. Initially it was found 42 Backcrossing has produced hybrid swarms, e.g. Ca- in the form of seeds, while from Great Britain it has been lystegia sepiurn x C. syhatica, and in the case of Centau reported in the wild State sińce 1986 (B a il e y & C o n o l l y rea jacea x C. nigra, an alteration in the genetic make-up 1984; B a il e y 1988, 2001). of the native species, even though the alien species had died 40 A species of hybrid origin is a much more significant out (C l e m e n t & F o s t e r 1994). phenomenon than a hybrid. The implication is that there 43 These are families which also show the highest num- has been a recovery of fertility and that the new species ber of natural hybrids (according to S t a c e 1975). The will form a self-contained cross-breeding genetic popula- author also lists families such as: Cyperaceae, Salicaceae, tion capable of evolution and adaptation. Scrophulariaceae and others. 41 W e b e r et al. (1998) described a hybrid which resulted 44 The newest findings have modified this opinion, from a cross between an invasive species introduced to indicating for ехатріе that such families as Amarantaceae Califomia from South America, Carpobrotus edulis, with and Cyperaceae are “weedier” than expected (F o r m a n the native C. chiliensis. This hybrid is fully fertile, and the 2003). In Polish native flora Сагех brizoides it certainly authors even foresee a high probability that a genotype appears to be invasive (expanding natives) (S ie r k a & capable of expansion will emerge. C h m u r a 2004). species occurring in the wild or established, plus em and south-eastern parts of Europę (species of almost a hundred of those which are often cul- Mediterranean and Sub-Mediterranean origin) tivated and transitionally move into the wild (cf. Chapter 5.1.2). It seems that this fact could within the lowland regions of Poland (R u t k o w be concerned with the more generał rule once s k i 2 0 0 2 ) . pointed out by Z a ją c A. (1979) with respect to The relationships presented above indicate that archaeophytes, a group again dominated by the grasses of alien origin constitute a considerable species from the same area, namely, that that percentage of the Polish flora, and it also appears predominance is associated with specific waves that the proportion has been increasing over the of human migrations. The “oldest” kenophytes last few decades. Among 251 species of newer (recorded in Poland as early as in 16th century) arrivals recorded in the flora of Poland there are include mostly species originating from these 13 species of grasses (Z a j ą c A. et al. 1998), regions; for some of them even some objections while the first, provisional list of kenophytes as to their non-native status can be raised (cf. (K o r n a ś 1968b) only includes 4 species. The list Chapter 5.2.2, Table 7). of ephemerophytes published in the late 1980s in- Also represented in the flora of Polish keno cluded 662 species in all, with 92 species of phytes are species with natural ranges limited to grasses (R o s t a ń s k i & S o w a 1986-1987). At that the central regions of Europę, particularly species time, this number covered 5 species which are from the Alps (Z a ją c A. et al. 1998). These are now regarded as either fully (Bromus carinatus, mostly species introduced as useful plants or those Eragrostis albensis) or locally established (Bro which were established in Poland as “relics” of mus japonicus, B. sąuarrosus, Hordeum jubatum). certain experiments in pastorał management car- Twenty years ago, the first two were regarded ried out at the tum of the 20th century (M ir e k as being only temporarily and accidentally 1995). brought in; however, they are now extending their The discovery of the Americas and the conse- secondary rangę (T o k a r s k a -G u z ik 2003b). ąuent breaching of the geographical barrier rep In other regions of the world, the participation resented by the Atlantic Ocean, has had an im- of grasses in invasions is also considerable. E n o - portant role in the process of flora exchange. m o t o (1997) lists 29 grass species among 285 During the last five hundred years, the flora of species of alien origin commonly occurring in Poland has been enriched by 112 alien species of Japan. In the flora of Italy (5811 species), 214 American origin, which are currently recognised species are regarded as being alien invasive as naturalised. This group is represented by 36 plants, and this number includes 15 species of families, of which Asteraceae (28 species), Ro- grasses (V ie g i 2001). In the Lithuanian flora, the saceae (13) and Onagraceae (12) are the most grasses contain the third highest number of alien important. The American flora established in species after Asteraceae and Brassicaceae Poland is characterised by the preponderance of (G u d ż in s k a s 1997b). Poaceae (151 species) and long-lived perennial herbs (40), woody plants Asteraceae (142) are represented by much higher (41) and annuals (28). Among 61 species for numbers of introduced species than any other which the first records for Poland are available, families in California (R e j m a n e k et al. 1991). As the majority arrived in Poland in the second half the authors pointed out about Poaceae, “this is of the 19lh and first half of the 20th centuries. Only certainly an extremely successful family in Cali- six species (Chamomilla suaveolens, Conyza fomia”. Among the introduced grass species 95 canadensis, Galinsoga parviflora, Amaranthus are from Eurasia. retroflexus, Oxalis stricta and G. ciliata) are Referring again to the Polish conditions, it is common (occurring in 60-90% of 10 x 10 km noteworthy to stress that most of the species of sąuares; the total number of sąuares for Poland Asteraceae have been introduced as decorative is 3646). Eleven species are abundant (occurring plants, while most species of the family Poaceae in 20-60% of 10 x 10 km sąuares), others are have been introduced accidentally. locally abundant (26 species occurring in 3-20% Because of favourable conditions for the trans of 10 x 10 km sąuares) or rare. The common port of diaspores and the absence of significant American species in Poland grow mainly in dis- barriers preventing expansion, among Polish turbed habitats. Over 50 species belong to kenophytes the groups originating from various a group that is very successful in migrating into parts of Europę predominate (Z a j ą c A. et al. natural and seminatural communities. Robinia 1998). pseudoacacia, Elodea canadensis, Solidago gi- The detailed analysis of species of European gantea, Juncus tenuis, Lupinus polyphyllus, Acer origin broken down by the European region negundo, Solidago canadensis, Padus serotina, where they came from, shows an evident predom- Bidens frondosa, Rudbeckia laciniata, Helianthus inance of species whose homelands are the south- tuberosus, Echinocystis lobata and Quercus rubra are widely distributed species which also city to disperse them over great distances47. For colonise natural habitats. Most of the above- most plants, wind is an essential factor facilitat- mentioned species have been classified as inva- ing long-range transportation of seeds and fruits. sive plants in Poland (cf. Chapter 12.5). It is the same among the kenophytes (cf. Fig. 13 in Chapter 5.1.6). This group has a great vari- In the context of migration, the active move- ability of adaptations to wind dispersal: special ments of plants are of minor importance in their devices which help blow seeds away: wings overall behaviour, even though they have devel- (Acer negundo, Ailanthus altissima), pappus oped a variety of morphological and biological (■Conyza canadensis) etc. Also of significance is adaptations for this purpose (F a l iń s k i 2000b). the weight of seeds and fruits: fine, light seeds Many authors have diseussed those morpho are easily transported over considerable dis logical features of plants which could have pre- tances (Digitalis purpurea, Eragrostis minor). disposed them to become effective colonisers of Another large group among kenophytes includes new habitats, often outside their prime rangę of plants which rely on animals for dispersing distribution (e.g. B a k e r 1974, 1986; N e w s o m e & diaspores, either in the digestive tract (endozoo- N o b l e 1986; K o r n a ś 1990; P y ś e k et al. 1995; chory) or attached to the surface of animals’ S t a r f in g e r 1997; J a c k o w ia k 1999; F a l iń s k i bodies (exozoochory). The former method is 2004)45. As a rule, however, none of the species used by such species as Sisymbrium loeselii, possesses a fuli set of the features which might Padus serotina or Amelanchier spicata. The ex- be considered as enhancing invasiveness. amples of species using the latter method are, The features of a species and its life strategy, for example, species of the genera Bidens and acting in combination with the conditions (most Xanthium (with special protrusions or append- often favourable ones) found in the new environ- ages on the surface of seeds), and Chamomilla ment (disturbed, changed habitats, lack of com- suaveolens (sticky seeds). The plants whose petition from native species, repeated accidental seeds, fruits, leaves or stems can cling on to introduction of diaspores by humans or long-term suitable surfaces, are transported by animals and cultivation ensuring a continuous supply of great humans alike (e.g. Amaranthus albus, Bidens quantities of genetically diverse diaspores, and frondosa, Salsola kali subsp. ruthenica, and sometimes even the introduction of a species into species of the genus Xanthium). Among the a habitat which was a “target habitat”46) enhances zoochores, there is a separate group of myrme- the chances of establishment and further ex- cochores, i.e. the plants whose seeds are trans pansion. The success of the colonisation can also ported and dispersed by ants. Their seeds have be ensured by the plant’s method of reproduction; elaiosomes, which are appendages containing in many cases it will be a vegetative mechanism, sugar, vitamins and other compounds used by and in the case of generative reproduction it will ants as food. Among kenophytes, this manner of perhaps be dioecism, or - as seems to be the case seed dispersion is utilised e.g. by Corydalis - the increasing importance of apomixis and self- lutea, Portulaca oleracea, and species of the fertilisation. genus Euphorbia. Then there are the autochores, Also of importance are, for example, the pro- the self-dispersing plants which have special de- duction of huge ąuantities of seeds and capa- vices to throw their seeds over a certain distance (sometimes even up to 1 m). The species of the genera Impatiens and Oxalis could be given as 45 In the opinions of K o r n a ś (1990) and J a c k o w ia k examples of this group. Another group of kenophytes (1999) the characteristics of plants which proved essential for synanthropie species in the process of their spread are as follows: short life cycle, broad spectrum of tolerance 47 Most abundant alien newcomers produce large ąuan towards living conditions, indifference towards photo- tities of seeds, e.g. a single plant of Conyza canadensis period, lack of special habitat reąuirements during germina- generates ca. 115 000 achenes (K o s t e c k a -M ą d a l s k a 1965), tion, germination spread over the season and prolonged via- a large individual of Bidens frondosa can produce over 500 bility of seeds, rapid growth of seedlings and short juvenile capitulae with over 10 000 seeds (L h o t s k a 1968); this has stage, early reproductive maturity and utilisation of major been corroborated by newer studies which found that it resources for reproduction, self-fertilisation, self-pollina- produces the highest number of capitulae compared with tion or specialised mechanism for cross-pollination, huge the species native to Europę, even as many as 17 700 seeds and uninterrupted (during the growing season) production per plant (G r u b e r o v A et al. 2001). P y ś e k et al. (1995) of seeds, an ability to disperse seeds over great distances, report that a single individual of Heracleum mantegaz- great potential for vegetative reproduction coupled with zianum produces up to 16 000 seeds, while T il e y & P h il ip competitiveness, an ability to develop ecotypes, polyploids (1997) suggest an even higher number of up to 107 000. and hybrids, and, finally, great variability in life strategies. Rudbeckia laciniata produces ca. 1 600 seeds per plant 46 Purposeful introduction of such species as Padus (F r a n c ir k o v A 2001) and Oenothera paradoxa ca. 8 000 serotina, Quercus rubra, Lupinus polyphyllus to forests seeds per plant (T o k a r s k a -G u z ik 1982). Many other exam- provides good examples to illustrate this. ples are given by P odbielkowski (1995). includes anthropochores dispersed either inten- proportion of the species with diaspores cling- tionally or unintentionally by humans. There are ing or sticking with mucilage is relatively high. several special cases of anthropochory, such as Generally, it can be stated that heavy-seed spe speirochory, i.e. dispersal of weeds with the cies (a category which includes barochores and seeding materiał of cultivated plants (among some zoochores) predominate in more maturę other kenophytes, this mechanism is utilised by communities. Their seedlings, provided with Veronica persica), ergasiochory, when diaspores more reserve materiał, stand better chances of are dispersed during tillage (e.g. species of the survival under the enhanced competition prevai- genera Vicia, Oxalis, and Galinsoga) and agesto- ling in dense and multi-storey patches of vegeta- chory, when diaspores are carried by various tion. The group of species with low-motility means of transport. diaspores, “inclined” to stay on the same site, As a rule, most of the kenophytes, however, which include barochores, autochores and the belong to the polychoric group, i.e. the plants heaviest anemochores must “depend on” humans which rely on several different mechanisms to to ensure the transport of diaspores over greater disperse their seeds (cf. Chapter 5.1.6 and Appen- distances (K o r n a ś 1972). dices A and B). The processes of plants’ seed dispersal depend In order to illustrate the degree of flora trans- not only on the morphological and biological fea- formation in a given area, investigators have tures of a given species, but also on the commu- used the proportions of species of alien origin nities where the species finds its niche. In his and their dynamics, presented in their various analysis of dispersal of weeds in cultivated field historical and spatial aspects, e.g. F a l iń s k i 1971; communities in the Gorce Mts., K o r n a ś (1972) S u d n i k -W ó j c ik o w s k a 1987а, 1998a; J a c k o w i a k stated that “(...) they use extensively two utterly 1990, 1998a; C h o j n a c k i & S u d n i k -W ó j c i k o w - different ways to disperse: namely: spontaneous s k a 1994. In connection with progressive synan- seeding brought about by natural factors, and the thropisation, the composition and structure of transport of diaspores by humans (anthropo the flora changes: the number of archaeophytes chory). The ultimate composition of communities remains almost unchanged, but the proportion of results from the combined effect of these two established more recent newcomers increases48 means of dispersal”. (F a l i ń s k i 1971; K o r n a ś 1977a; M i s i e w i c z Apart from confirming the predominant role 1981). of generative reproduction, anemochory and au- Kenophytes, as a group distinguished in the tochory, the attempt of this kind of analysis in geographical/historical classification, are treated the group of kenophytes also indicates the ex- as indicators illustrating the intensity of the pro istence of some specific features revealing them- cess49. The proportion of this group of species in selves in the context of the types of habitats the flora, particularly in urban areas, helps in actually colonised (Fig. 63). In the habitats demarcating the zones of human impact (called occupied by stable communities of a semi-nat- also anthropopressure zones) and has been used ural (meadows and grasslands), or natural (for- by many authors50. ests and scrub) character, the proportion of S u d n i k -W ó j c i k o w s k a (1992) suggested that vegetative methods of reproduction increases. particularly useful in demarcating the zones of This is particularly so in aquatic and riparian human impact in urban areas will be those indi- communities because their specific naturę is conducive to this manner of reproduction. On the other hand, in disturbed habitats or those 48 This correlation is illustrated by the so-called coefficient of flora modernisation: M = epecophytes + agrio- subject to permanent or temporary pressure from phytes/archaeophytes. humans, the proportion of kenophytes following 49 The number of kenophytes in a local flora is also used the vegetative process of reproduction evidently when indices of flora synanthropisation (including so- declines. The great majority of kenophytes called “complex indices of flora synanthropization”) are disseminate their diaspores via anemochory and calculated (J a c k o w ia k 1990; S u d n ik -W ó j c ik o w s k a 1991, autochory, that is by methods characteristic of 1992 and references cited there; U r b is z 1991; S u d n ik - W ó jc ik o w s k a & M o r a c z e w s k i 1993, 1998; M o r a c z e w s k i pioneer communities. However, in more evolved & S u d n ik -W ó j c ik o w s k a 1994). communities which usually have a very complex 50 S u d n ik -W ó j c ik o w s k a (1986, 1998a) in characterising structure, the role of other methods of dissem- the flora of a major town, delineated zones on the basis of ination increases. They will involve endozoo- the percentage proportion of kenophytes in the 1 km2 chory in forest and scrub communities and sąuares of the study area; the spatial diversification of the flora of urban areas has been also illustrated, again in terms epizoochory in meadow and grassland, as well of the proportions of kenophytes, by other authors (e.g. as aquatic communities. Also in other types of T o k a r s k a -G u z ik 2000; K u c h a r c z y k 2003a; M a c ie j c z a k open communities in ruderal habitats, the 2003; W o ł k o w y c k i 2003; Z a j ą c M . & Z a j ą c A. 2003). Type of habitat forests & scrub meadows & grasslands water & riparian railways & roads abandoned fields segetal ruderal

20 40 60 80 E3generative I vegetative I both types

B Type of habitat forests & scrub meadows & grasslands water & riparian railways & roads abandoned fields segetal ruderal

10 20 30 40 50 60 70 80 90 100% Hautochores Hanemochores ■barochores dhydrochores □ epizoochores 0 endozoochores ■ myrmecochores ■ anthropochores

Type of habitat forests & scrub meadows & grasslands water & riparian YZZ/MYl Uli...... lilii! lilii 11 1 1 railways & roads abandoned fields w m w w m segetal ruderal ...... w m iH * im a i m

0 10 20 30 40 50 60 70 80 90 100% E3 seed □ fruit O root ■ rhizome ■ rosette □ stem B plant Fig. 63. Reproduction properties of kenophytes from different habitats: A - manner of reproduction, B - dispersal spectra, C - propagule used ces which describe the proportions of such geo- At the same time (as suggested by the afore- graphical/historical groups as synanthropic new- mentioned author), using these groups of species comers (P4, P5, P9), and permanently established as a tool also allows one to evaluate changes in alien species (PI)51. a flora from a historical perspective (with one

51 P4 = Ep + Ag + Ef + Eg / G x 100% - the percentage the number of euapophytes; Ar - the number of archaeo- of recent anthropophytes in the flora (i.e. within the approach phytes, arrivals established permanently before 15th centu- adopted in this paper, it is a combined proportion of keno ry; Ep - the number of epecophytes, new arrivals established phytes and diaphytes in the flora; in the approach used by permanently in anthropogenic habitats after the 15th century; Anglo-Saxon authors it is the proportion of neophytes) Ag - the number of agriophytes, new arrivals which had P5 = Tn/G x 100% - the percentage of therophytes- established permanently (after the 15th century) in natural and newcomers in the flora semi-natural communities and usually also in anthropo P9 = Ep + Ag/G x 100% - the percentage of kenophytes genic habitats; kenophytes - epecophytes along with agrio in the flora phytes; Ef - the number of ephemerophytes; Eg - the num PI = Ar + Ep + Ag/Ap - the ratio of the total number ber of ergasiophygophytes; Tn - the number of therophyte- of permanently established anthropophytes to the number of newcomers, i.e. therophytes that are also epecophytes, agrio euapophytes where: G - the total number of species; Ap - phytes, ephemerophytes or ergasiophygophytes. reservation: the applied floristic parameters are The author emphasizes that “(...) it is a key only (or may be only) indicators of habitat con- moment for understanding the differences between ditions at a defined point in time). urbanophilous kenophytes and urbanoneutral kenophytes that are widespread in cities”. The issue of the establishment of kenophytes The botanists involved in studies of the changes in various types of habitats is a fairly popular sub- occurring in the structure of urban floras, high- ject of research in Poland (T o k a r s k a -G u z ik light the development of floras specific to such 2003 c and references cited there). The majority urban habitats as old city centres, defence perim- of kenophytes show an ability to adopt to a rel- eter walls, railway stations, tram line tracks, or atively wide rangę of habitats, e.g. Conyza ca- playgrounds, where kenophytes are significantly nadensis and Acer negundo. Only a few species represented (e.g. K unick 1982, 1990; B randes can be named as faithful to a particular type of 1992b, 1995; Świerkosz 1993; K owarik 1995b; habitat: Corydalis lutea and Cymbalaria muralis Jackowiak 1998a & c; Pyśek 1998; S udnik- grow only in crevices in remnants of old walls; W ójcikowska 1998a; G alera & S udnik- Eragrostis minor is recorded on railways, storę W ójcikowska 2000a & b; S hevera 2003; Tokar yards and in the centres of towns between flag- ska-G uzik 2000, 2003c; Z ając M. & Z ając A. stones; Elodea canadensis and Lemna turionifera, 2003). The role of towns and cities in the estab as a hydrophyte, only in water (T o k a r s k a -G u z ik lishment of species of alien origin and in over- 2003c). coming the difficulties of the first stages of ex- Among 300 species of kenophytes which are pansion is still significant (Jackowiak 2003; cf. covered in this monograph, 160 species are as- also Chapter 9.3). The comparison of the share sociated with anthropogenic habitats (so-called of kenophytes in the 19

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Fig. 64. Role of cities and towns in establishing of kenophytes in the flora of Poland (on the basis of the number of the first records)

57 In the 17lb century, the development of Warszawa was 58 At present, Warszawa and Łódź are the largest cities disturbed by Swedish invasions (the “Swedish deluge”) in Poland. Until the 1930s, Wrocław was second only to when the city suffered much damage. Warszawa in the territory within the present Polish borders. species introduction by humans, accidental and The analysis of quantitative changes in the intentional. It was in just these cities where the kenophyte flora within Poland sets out the greatest numbers of “first floristic records” for areas where migration of alien species was kenophytes are located (Fig. 64). Through to the faster and more intensive than elsewhere64 (Fig. beginning of the 19th century, the greatest role 65). These are areas which had been urbanised in establishing newcomers was played by War earlier, thus having denser transport networks szawa and Gdańsk, cities situated on the main connections. commercial route in Poland running along the When considering the role of cities as a “foot- course of the Vistula, but also by some smaller hold” which enables the species, whether acci- cities, such as Wyszogród59 - again situated on dentally brought in or introduced, to expand the Vistula route, and Silesian cities: Wołów60, further into adjacent areas, it is worth noting that Oława61, and Bytom Odrzański62. In the first half the cities had a similar effect on these two groups of the 19th century the role of the city of of plant species. Gdańsk and Kraków stand out Wrocław63 grew as the site for the first records among the four oldest and largest cities in Poland, of kenophytes, as well as the significance of showing opposite tendencies in terms of the other Silesian cities which were then within the means by which plant species were introduced borders of Germany. The political and econo- there (Fig. 66). Gdańsk, being a seaport of par- mic situation at that time evidently favoured the ticular history and tradition, is above all the place settling of new plant species into local floras. of the first records of species brought accidental- The most recent 150-200 years constituted the ly, whereas Kraków, an academic and cultural period of the industrial revolution which oc- centre, perhaps “created” better chances for in curred with variable speed in various cities of tentional introduction. Central Europę. The development of industry A similar mechanism is still operating. In many was coupled throughout by the parallel exten- cases the territory of a city is the destination of sion of transport networks. The inbound migra- the first stage of migration by a foreign newcomer tion of the population into cities accelerated (in Poland, these types of circumstances are ex- causing them to significantly expand their ter- emplified e.g. by Ailanthus altissima - cf. Chap- ritories. As a result, dramatic changes occurred ter 7.1, Parietaria pensylvanica (S a w il s k a & in the habitats and the overall environment M is ie w ic z 1998; G u z ik 2002)). (S u d n i k -W ó j c i k o w s k a 1998a). The cities have performed multiple functions, At that point in time, the Polish lands were being at the same time marketplaces (for farm situated on the main divide of the industrial produce, horses and cattle), industrial centres of map of Europę, separating the highly industri- cloth manufacturing (e.g. Gorzów Wlk., Toruń), alised German lands from backward areas of railway hubs, and sometimes river ports (Byd Russia and Austria-Hungary (D a y ie s 2001). goszcz, Gliwice), thus becoming “open to invasions of alien species, their number unforesee- 59 Wyszogród is one of the oldest defensive strongholds able” (T r e p l 1994). of Mazovia, located on a high bank of the Vistula (its name One should also note the coincidence of dates derives from this situation). The city overlooked the Cros of these first records with the dates of commis- sing of the Vistula and thus developed as a trade centre and sioning new railway connections (cf. Fig. 64 and river port. In the 16th century Wyszogród was the single Appendix A). The railway reached Bole largest centre in Mazovia (Kwiatek & Lijewski 1998). 60 Wołów - a city founded in the 13th century on the site sławiec65 as early as in the first half of the 19lh of a former stronghold; in the 16,h century it became a centre century (in 1845 the city was linked by railway of cloth manufacturing and crafts (K w ia t e k & L ije w s k i 1998). line with Wrocław, then it was extended to 61 Oława - even in the early Mediaeval times, it was a Gubin and Zgorzelec). Wołów is situated on the stronghold and marketplace settlement developing on the trade route from Wrocław to Kraków. In the 16Ih century, Oława had breweries, a paper mili, and the prince’s mint 64 The picture of quantitative changes in the kenophyte (K w ia t e k & L i j e w s k i 1998). flora of various regions of Poland depends much on the 62 Bytom Odrzański - an old stronghold situated at the level of details known about floras of particular regions, Crossing of the Oder river. It recorded a period of dynamie and this in tum is linked with an overall level of economic growth at the tum of the 17th century when the city and cultural development (expressed by the number of developed as a centre of trade and crafts upon the Oder academic centres and various schools where such studies river route ( K w i a t e k & L i j e w s k i 1998). were initiated). 63 Wrocław - at the end of the 18th century the city expe- 65 At the same time, these were old Mediaeval cities or rienced a major boost in its economy: cloth and metal indus- settlements usually located upon rivers or on trade routes tries developed and the Oder inland waterway acąuired major - in the 15th and 16th centuries. Bolesławiec, upon the Bóbr importance as a link with Prussian provinces. In the mid river, was a centre of cloth manufacturing and the single 1900s, the city became connected by railway links with Berlin, largest salt market in Silesia, as well as a market for horses Dresden and Upper Silesia (K w ia t e k & L ije w s k i 1998). and cattle (K w ia t e k & L ij e w s k i 1998). A B C D E F G

Fig. 65. Concentration of 174 species of kenophytes and the expansion ofthe railway network in Poland (C z a pl iń s k i & Ł a d o g ó r s k i 1998) in the historical seąuence 1501 - XX century human settlements. The term “plantae urbanae” was first introduced by Schouw in 1823 (S u k o p p 2002), giving Xanthium strumarium61 as one of the examples. Later, the concept of the formation of local distribution ranges in urban conditions has been further developed e.g. by W it t ig et al. (1985) and J a c k o w ia k (1990, 1998a, b & c). One of the specific features of cities is the “urban 20 climate”, characterised, among other phenomena, by the presence of the “thermal island”68, the over-dryness and pollution of air (S u k o p p & Gdańsk Kraków Warszawa Wrocław W e r n e r 1983; J a c k o w ia k 1998a, b & c; S u d n i k - □ accidentally ■ intentionally W ó jc ik o w s k a 1998a & b, 2000; S u k o p p & W u r z e l 2000). Fig. 66. Role of the Polish greatest cities in the first stages of spread of kenophytes with respect to the The following species are listed as associat presumed type of introduction into the country ed with Central European cities: Ailanthus altissima, Buddleja davidii, Chenopodium botrys, Wrocław - Zielona Góra - Szczecin railway Diplotaxis muralis, and Eragrostis minor (Table line, and this section opened in 1874, whereas 9). Ailanthus altissima, Diplotaxis muralis, and Bytom Odrzański - on the Głogów - Zielona Eragrostis minor are specifically associated with Góra railway line, opened in 1871. The railway the central parts of cities, and are termed “ther line linking Oława with Wrocław is the oldest mal bioindicators” (S u d n i k -W ó j c ik o w s k a 1998a railway line in Poland, which has been in & b). Despite sharing some floristic character- operation sińce 1842. istics, the proportions of individual species of Although the impact of humans within the kenophytes in European cities are differentiated limits of the European cities has a history span- by the climatic conditions resulting from the ning over many centuries, geographical location geographical position of a given city (Table 9). remains the factor of prime importance in terms This specificity has already been noted by of affecting flora composition. The flora of K u n i c k (1982) when he listed alien species urban areas has derived primarily from the typical of cities situated in Eastern Europę, e.g. native species from the nearest environs of the Amaranthus albus, Atriplex tatarica and Iva city. The proportion of species of alien origin, xanthiifolia. On the other hand, in cities situated including the more recent newcomers, which in the western part of the continent, the following usually prevail over older immigrants, fluctuates species are recorded more often: Ailanthus from ca. 12-25% in the cities located in the altissima69 (in Poland, this particular species is south of the continent, to 40-50% in Central at the initial stages of dispersion - cf. Chapter 7), European cities (F a l i ń s k i 1971; P y ś e k 1989; Buddleja davidii (a shrub, established locally in K o w a r ik 1995b; C e l e s t i G r a p o w & B ł a s i 1998; Poland, susceptible to freezing in winter) and cf. also Chapter 8). Apart from the differences Chenopodium botrys (in Poland it is found associated with variable methodologies and primarily in industrial wastelands, such as spoił different time horizons, the overall differences heaps or settlement ponds, and is much rarer in may stem both from the higher proportion of the actual cities). species arriving from North America (similar climate) and Southern Europę which can utilise “urban heat islands”66 offered by the cities situated more to the north, a much less important factor in the cities of Southern Europę (C e l e s t i 67 “In most cases foreign origin is the cause why these G r a p o w & B ł a s i 1998). plants are located only near cities and villages” (S u k o p p Some species have been virtually regarded as 2002 after Schouw). being specifically associated with cities and other 68 In the Central European cities this phenomenon has been observed for over four decades. Urban ecologists have long indicated the importance of flora as a bioindicator of 66 The built-up areas, because of their specific climate the thermal conditions prevailing in a given city (S u d n i k - and the type of habitats, provide favourable conditions for W ó j c i k o w s k a 2000 and references cited there). the expansion of plants and animals from the warmer cli- 69 Ailanthus altissima is also a permanent and freąuent matic zones. In Berlin, for е х а т р і е , some 6 0 % o f alien component of cities situated in the Southern parts of Eu plant and animal species (archaeophytes and neophytes) ropę, e.g. in Rome (C e l e s t i G r a p o w 1995), or Ljubljana с о т е from warmer regions (S u k o pp 2 0 0 2 ). (T o k a r s k a -G u z i k , pers. observ.). Table 9. Species indicated as associated with Central European cities

Locality and author Species examples from Poland examples from other European towns

Acer negundo W arszaw a G alera & Sudnik-W ójcikowska 2000a, b Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997

Ailanthus altissima Łódź Witoslawski 1993 B erlin K unick 1982; BOcker & K owarik 1982; K owarik W arszaw a Sudnik-Wójcikowska & Guzik 1998 & BOcker 1984 Munster, Essen, Dusseldorf W ittig et al. 1985 Katowice, Kraków, Tokarska-Guzik, pers. observ. W rocław Halle, Leipzig G u tte et al. 1987 Berlin, Karlsruhe, Koln, Stuttgart, Wiirzburg K unick 1990 Z urych L andolt 1991 L eipzig G utte 1992 R om a C elesti G rapow 1995

Amaranthus albus Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997

Amaranthus blitoides Łódź Sowa 1960 Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997 Poznań Jackowiak 1993

Atriplex tatarica Poznań J ackowiak 1993 Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997 Warszawa, Lublin Sudnik-W ójcikowska 1998a Buddleja davidii B erlin Kunick 1982 Munster, Essen, Dusseldorf W ittig et al. 1985 Halle, Leipzig Frank & Klotz 1990 Berlin, Cologne, Dusseldorf, Essen, Freiburg, Kunick 1990 Stuttgart

Chenopodium botrys W rocław R ostański 1960 L eipzig G utte 1971 B erlin Sukopp 1971; K unick 1982 Halle, Leipzig F rank & K lotz 1990 M ariupol B urda 1997

Diplotaxis muralis Poznań J ackowiak 1993 Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997 Łódź W itoslawski 1993 W arszaw a C hojnacki* S udnik-W ójcikowska 1994; S udnik-W ójcikowska 1998a

Eragrostis minor Poznań Jackowiak 1993 L eipzig G utte 1971, 1992 Łódź W itoslawski 1991 B erlin Darius & Drepper 1984 W arszaw a Sudnik-Wójcikowska 1998a; G alera & Sudnik- Braunschweig Brandes 1987 Wójcikowska 2000a, b Munster, Essen, Dusseldorf W ittig et al. 1985 K atow ice Tokarska-Guzik, pers. obser. Halle, Leipzig Frank & Klotz 1990 Zurych L andolt 1991 Donetsk, Lugansk, Slavyansk, Mariupol B urda 1997 V ienna J ackowiak 1998c Iva xanthiifolia Poznań U rbański 1955 W arszaw a S udnik-W ójcikowska 1987b Lublin Święs 1993

Parietaria pensyhanica B ydgoszcz Sawilska & Misiewicz 1998; Sawilska et al. 2003 B erlin S ukopp & Scholz 1964 W arszaw a G uzik 2002 9.4. Historical gardens, botanic supplement to the indigenous plants came in the gardens, cloister and convent form of alien species, which were to augment gardens as places of both the biological and spatial structure of a “domesticating” exotic species garden. prior to their spontaneous In 18lh century England, when regular gardens establishment were radically replaced by landscape-type ones, species of alien origin, called “exotic”, were in troduced on a massive, never before seen, scalę Humań settlements in the latter stages of their (S iew n ia k 1989). The peak of this so-called “ex- development have always been accompanied by otica madness” occurred in the 1840s. It was as- plants closely connected with humans and with sociated with expeditions of discovery by arbo- their ever-improving forms of economic activi- riculturalists, e.g. those of D. Douglas along the ties. These plants include cultivated plants, weeds north-eastern coast of North America, and of and ruderal plants. Humans gradually introduced R. Fortune across East Asia. The ехатріе of En increasingly more plant species into cultivation, gland was followed by massive imports of first those used to obtain food and medicines, “exotics” into the Netherlands and France. After then industrial and decorative plants. a certain time delay, these species reached The second half of the 18th century was thus Central Europę. In Poland, these alien species appeared relatively early in gardens because of characterised by K lu k (1 7 8 6 ): “[...] This time wealthy landowners (W o d z ic k i 1824-1828; is so good in housekeeping and thence it looks S iew n ia k 1989). to naturalising much of its native and alien re- The state of our knowledge about the propor- sources in order to need only a little from tions of alien trees and shrubs in historical parks abroad” [“Wiek teraźniejszy bardzo gospodarny, is still far from exhausted. A majority of recorded szuka iak naywięcey w kraju swoim oswoić, aby and published data pertains to the first instance mniey potrzebować z cudzego”]. This author of introducing species into botanic gardens or thus provided not only the lists of native and the more significant parks (M a jd e c k i 1993). alien species accidentally introduced into Poland Relevant materials, if available, are usually and cultivated during those times (cf. Table 6 in scattered or included in sources reaching back Chapter 5.2), but also the plants cultivated in the merely to the 19th century, and difficult to access neighbouring countries which he deemed wor- (H e r e ź n ia k 1992). Also lacking is sufficient thy of acclimatising in Poland (e.g. Asclepias information on nurseries breeding decorative syriaca). plants and directly involved in spreading trees The list of plants cultivated in various histor and shrubs. M a jd e c k i (1993) presents selected ical periods documents the history of the culti- examples of the most freąuently encountered vars, the changing needs of humans, as well as trees and shrubs of alien origin in Polish historical trends prevailing in the art of gardening. Each parks. The list also includes trees and shrubs garden, being a defined spatial system, is shaped established in Poland only recently (Table 10). in terms of functions under certain environmen- Despite having been cultivated for 100-200 tal conditions, but taking into account also the years, some of the species listed in the table time, needs and place of its establishment is have been deemed to be established only in the additionally affected by prevailing tendencies in most recent decades (and some only locally - cf. arts, opinions, customs and beliefs (M a jd ec k i Appendix B). 1993). The historical gardens have preserved till Among the attractive avenue-forming trees our times ample sources of information about used in the Baroque period were the Horse history, fme arts, the use of plant resources and Chestnut Aesculus hippocastanum, and the still naturę in generał. occasionally-used Locust Robinia pseudoacacia; Trends in the way gardening art has shaped which have been known in Europę sińce the 17th gardens have depended on many historical, cul- century; in gardens of that period shrubs were tural and biological factors, but the selection of also planted, such as lilac and spirea. Other spe plant species has also been a significant factor. cies listed in Table 10 include species now com- The basie species structure (plant cover) of any monly occurring in Poland and still expanding, park is usually provided by indigenous elements, such as Acer negundo, Quercus rubra, Robinia matching the habitat conditions of the original pseudoacacia, Fraxinus pennsyhanica and Pa- area where the park was designed. With time, dus serotina. however, plant materiał resources became richer The successful process of establishing many owing to achievements in plant breeding. The tree species was not solely an outeome of their introduction to historical parks70. For ехатріе, The assortment of decorative plants used in the the Black Poplar called Italica (Populus nigra herbaceous layer of gardening arrangements in- ‘Italica’), was initially a hallmark of the parks es- cluded the Dame’s-violet Hesperis matronalis, tablished in the first half of the 19th century and and flower beds were made of Common Foxglove only later did it go into widespread use as a tree Digitalis purpurea, Sweet William Dianthus bar- planted in cemeteries and also to create avenues. batus, Elecampane Inula helenium, and Garden Another essential role of gardens was that of Łupin Lupinus polyphyllus, while the borders of “domesticating” perennial plants imported in or flower beds used Hyssop Hyssopus officinalis der to add decorative and artistic merit. among other species (M a jd e c k i 1993).

Table 10. Trees and shrubs of alien origin the most freąuently encountered in Polish historical parks, contemporarily naturalised in Poland Described as invasive Тахоп Origin Hab. Status in Poland elsewhere XVII-XVIII Acer negundo Am N NSH invasive [rip. agr. urb.] Eur C [rip. agr. & urb.] Acer saccharinum A m N H casual / locally naturalised Aesculus hippocastanum EurSE SH naturalised Elaeagnus angustifolia Eur S, Asia W & С H casual / locally naturalised Hungary; Am N [rip.] Pinus strobus AmN N locally naturalised Czech Rep. Populus nigra “Italica” Anthropog. H locally naturalised Quercus rubra Am N N naturalised / invasive Czech Rep. Robinia pseudoacacia Am N NSH invasive frip. agr. urb.] some regions of Eur C & S Syringa vulgaris EurSE NSH naturalised / relic Czech Rep. 1/2 XIX Ailanthus altissima Asia E H naturalised /pot. invasive [urb.] Eur С & S [urb. & rip.]; Am N Amorpha fruticosa A m N H casual / locally naturalised some regions of Eur С Fraxinus pennsylvanica AmN SH naturalised /invasive Czech Rep. & Hungary Lonicera tatarica Eur SE & Asia С SH locally naturalised Pseudotsuga menziesii AmN N locally naturalised Ptelea trifoliata AmN H locally naturalised Rhus typhina AmN H locally naturalised Symphoricarpos albus Am N NSHnaturalised Czech Rep. Tsuga canadensis AmN N casual / locally naturalised 2/2 XIX Cotoneaster lucidus Asia С NH locally naturalised Mahonia aąuifolium A m N H locally naturalised Czech Republic, Germany Populus berolinensis Anthropog. H locally naturalised *■ Padus serotina Am N & Am S NS invasive [rip. agr. urb.] Eur С [forests] Pterocarya fraxinifolia Asia SW N casual / locally naturalised Rosa multiflora Asia E NSH locally naturalised Rosa rugosa Asia E NSH naturalised /relic Thuja plicata A m N NH casual / locally naturalised

70 European forestry has played an important role in the Botanic gardens, smali gardens at monaster- introduction of exotic species (S ie w n ia k 1989). This author ies and convents, as well as some home gardens is of the opinion that foresters had the following goals: to also became types of “seedling nurseries” for produce more technically valuable wood, to decrease the plants imported for use in medical treatments or vulnerability of forests to pathogens (lack of natural pests) cooking71 (e.g. Artemisia dracunculus, Atriplex and to widen tolerance to habitat conditions. hortensis, Hyssopus officinalis, Marrubium vul- 71 Artemisia dracunculus was used for salads and as a spice, the leaves of Atriplex hortensis were used as spin gare). Some other species were casually brought ach; Hyssopus officinalis was cultivated as a spice or in with imported seed or seedlings, and passed medicinal plant: strewn in baths and also taken intemally through gardens in the first stages of their es “to strengthen nerves”, Marrubium vulgare was applied in tablishment (e.g. Amaranthus albus, Eragrostis the same capacity. multicaulis, Euphorbia humifusa). There has long sińce been a worldwide inter- Chapter 5.2) in the new rangę colonised by it. Such est in the history of introducing exotic plant a conclusion is legitimate if there is a great num- species into parks and gardens. The effects of ber of records for a particular species within a short their “eseape” and establishment outside garden time of its first record (e.g. Echinops sphaerocepha- and park sites places has been freąuently docu- lus, Lycium halimifolium, Reseda luteola, Hesperis mented (e.g. U dvary & F a c sa r 1997; S u k o pp mationalis or Sisymbrium loeselii)72. 2002 and references cited therein). Archaeolog- The second group consists of species intention- ical and ethnobotanical studies permit the recon- ally introduced into cultivation by humans which struction of the history of growing fruit trees, subseąuently become wild. In such cases, the date vegetables and other plants used by humans from of introduction into cultivation is typically more as long ago as the early Neolithic Period. The precise compared with the date when the plant results of these studies can even indicate the becomes wild, which is usually known only as differences in preferences shown towards culti- a rough estimate. Apart from a list of species, the vated species in urban and rural areas (Z eist et oldest floras usually provided only descriptions al. 1991). of the plants and their uses. Information on the course by which they entered the flora in a given area was most often referred to only in gen erał terms. The first “fuli” records, carrying in 9.5. Immigration periods (peak inflows formation suitable for entering into databases and of kenophytes) drawing maps most often pertain to later periods compared with the mostly unnoticed period of The reconstruction of the immigration periods “going into the wild state”. of individual kenophytes and groups (e.g. species with common origin) calls for specific informa- Difficulties in a thorough and accurate recon tion: when the species was accidentally brought struction of the course of expansion, using in, or introduced into cultivation, and when it was a uniform method for the entire group of keno initially recorded as spreading spontaneously (the phytes, do not alter the fact that Poland’s flora “first” record!). Also essential is information on was enriched by newcomers in a whole series of the rate of spread from the first record until recent historical periods. In this process of influx of times (expressed in the numbers of stations in alien species one can even speak of peaks (“mi- subseąuent periods). gration waves”) the highest of which occurred in Differences in the method of gathering data the second half of the 19th century (cf. Chapter and the changes which occurred to the national 5.2). Changes in both the number and proportions territory of Poland throughout the last four cen- of kenophytes of different origins in the series of turies, allow only estimated reconstructions of the “immigration waves” can be linked to the economic periods of “influx” of newcomers into Poland and situation of Poland in the periods identified. their course of expansion. In contrast to Western Europę, the industrial- In most cases, the first known record of a isation of the eastem part of the continent started kenophyte in Poland does not reflect the actual after a significant delay and progressed not timing of its appearance. Unless there were spec- without perturbations (D avies 2001). In the Po- tacular, sudden and massive occurrences of a lish lands, the first rather less advanced industrial newcomer (as was e.g. the case of Canadian manufactories appeared in the first half of the 18th Waterweed Elodea canadensis), a species was century. During the industrialisation of Poland, usually noted after a certain delay. modern accounts distinguish three stages: the For ехатріе, in the case of the species Acorus first, from the 1740s until 1815 (characterised by calamus, described in the 17th and 18th centuries no major changes in economy but significant (S y reń sk i 1613; K lu k 1787) as a common plant, progress in science and technology and the de- growing in inland water courses, lakes and ponds velopment of trade); the second - from 1815 until throughout Poland, the earliest record identifying the outbreak of World War II (called “the first in a specific place dates back only to 1824 (the only dustrialisation”), and the third - post 1945 (called earlier record, which pertains to an urban site in “the second industrialisation stage”), which con- Warsaw dates back to 1652; after S u d n ik - tinues through to today (D ayies 2001). W ó jc ik o w sk a 1987a; cf. Appendix A). It is thus highly likely that many species were 72 In the case of some of the “oldest” arrivals among the accidentally brought in earlier and that the first kenophytes, some doubts exist as to their true status in Poland. For ехатріе, in K o r n a ś ’ s (1968b) opinion, Lycium available records do not correspond with the initial halimifolium and Reseda luteola should be grouped with phase of the expansion of the new species (cf. the oldest arrivals (archaeophytes). Subsequent peaks in the influx of kenophytes But plants also “attempt” long “leaps” of distances can be coupled with the stages of industrialisa- measured in hundreds or even thousands of tion cited above. The gradual rise in the number kilometres (K o r n a ś & M e d w e c k a -K o r n a ś 2002). of species up to the end of the 18lh century, with The effect of these migrations depends on the an evident predominance of species of European biological properties of the plant and, above all, and Asian origin, can be explained by trade links, on the mode of production of offspring and the as well as by wars fought by Poland at that time. methods of dissemination of diaspores, as well as An evident rise in the number of newcomers con- other natural factors. In many cases, anthropogenic tinued throughout the subseąuent periods of the factors are also of significance. 19th century, in the stage of “the flrst industrial - The enormous potential of plants to migrate isation”, with an added tendency towards an has been demonstrated throughout the course of increased proportion of alien species from the the development of natural vegetation cover, and Americas (particularly North America). The most was particularly evident in the Quatemary, i.e. dynamie period of “the first industrialisation” fell following the end of the last glaciation. In the in the years 1864-1918, when Polish industry Holocene, at least sińce the Neolithic Period, was drawn into a wider European market of human beings have become a prime factor in plant migrations (J a c k o w i a k 1999; Z a j ą c A. goods, labour, and Capital (D avies 2001). The railway network expanded, new industrial regions 1979). The discovery of America by Columbus73, developed and stimulated urban growth in adja- which initiated ever-intensifying contacts be cent areas (cf. Fig. 65). The density of the pro tween the continents, contributed to the com- portion (i.e. the concentrations) of kenophytes in mencement of a “global experiment” in which elements of floras are exchanged between regions the floras of individual regions of Poland over separated by natural geographical barriers (K o r this period reflects the distribution of major cit- n a ś 1990; J a c k o w ia k 1999). ies, industrial centres and the links between theni. As regards the group of newer arrivals, these The evidently higher density of kenophytes in the are migrations in the geographical meaning of the south-western, western and northem parts of Poland word. They lead to a widening of the initial rangę arose not only from a higher degree of industrial (this phenomenon pertains to some European spe isation and urbanisation compared with other re cies - cf. Chapters 6 and 7) or mostly result in gions of Poland, but also from many centuries of the appearance and development of a new, sec- traditional links between these areas and Western ondary rangę (F a l iń s k i 2004). Europę. The rate of industrialisation and urbanisa Plants showing only a limited potential for tion was defmitely higher in those areas than re- active migration, apart from developing some maining German. The industrial revolution reached adaptations facilitating dispersion, utilise the Prussia relatively early. The first iron smelter was natural conditions of the colonised regions or opened in the Ruhr basin in the 1780s, and the first make use of the means of transport provided by in Silesia was commissioned in 1794. In 1847, the other species, i.e. animals and humans (cf. Chap- first railway line went to use in Prussia. The most ter 8). important region of modem Poland was established, developed and brought to “economic maturity upon the initiative of Germany within the economic system of Prussia. Throughout the industrial revo- 9.6.1. Rivers as migration corridors lution its links were not oriented towards Poland but aiding the spread of kenophytes towards other parts of Germany” (D a v ie s 2001). Adopting the expression once used by D a v ie s (2001) who suggested that “it is the geography River beds and valleys are migration routes of Poland which stands guilty of determining her used by plants that are often rather easy to doc- past” we might condition (to a certain extent) the ument (K o r n a ś 1990; S u k o p p & T r e p l 1987). results of the reconstruction of past Polish floras These river-related plant migrations, their essence of kenophytes both on the geography and history and importance in ecological and geographical of the Polish Republic. expansions, based on many examples, were re- viewed in detail by F a l iń s k i (2000b).

73 The latest researeh completed in Europę and Ameri 9.6. Migration routes ca shows that Vikings should be regarded as the precur- sors of sailing across the Atlantic Ocean to the New World. Erie the Red of Iceland, who was banished from the island The overall tendency to spread, common to all and thus went on to Greenland where he established two plants, depends on many factors. Most migrations settlements, is considered to be a pioneer of discoveries in are short-distance and are carried out step by step. America (D ł u g o s z 2001). Among the newer arrivals, many authors present This statement pertains to the Bidens frondosa examples of species which use river valleys dur- and Xanthium albinum along the Elbe river. The ing certain stages of their expansion. For example, distribution of the latter species and the associ- LhotskA & Kopecky (1966) refer to the expansion ation of its migrations with river valleys have of Impatiens glandulifera in the Upper Oder river been also illustrated by G u d ż in s k a s (1997d) in basin as well as in the basins of Vratka and Svi- Lithuania where Xanthum albinum is common tava rivers in the present territory of the Czech Re- along the Neris and Neman rivers and in Poland public. Within the borders of Poland, the highest by D a j d o k & K ą c k i (2003) and K u c h a r c z y k number of stations of this species are concentrated (2003b) along the Lower Odra and in the middle in the upper and middle course of the Oder (from course of the Vistula, respectively (cf. also Chap the border with the Czech Republic down to ter 6). In Poland Rumex confertus, a species as Wrocław) (cf. also Chapter 7), which is perhaps sociated with the valleys of the Vistula and Bug related to soil type (rich brown soils) and the forms rivers, has become a model species in this respect of human impact on the environment in this part (cf. Chapters 6 and 7 and references cited there; of the valley ( D a j d o k et al. 2003). Also D r e s c h e r also K r a s ic k a -K o r c z y ń s k a et al. 2003). & P r o t s (2003) highlighted the connection be- B r a n d e s (1991) regards Bunias orientalis as tween the dispersion of I. glandulifera and migra- a species occurring chiefly along rivers (in tions along river valleys, pointing out that the Western Europę its stations concentrate in river “dispersion and establishment of the species along valleys, e.g. the Main, Tauber, Rhine, Meuse), watercourses will depend strictly on the geomor- although some other corridors are also listed, phology of the river and the bank, the duration of such as roads, canals where it grows on banks flood, the speed of water flow and the type of of fertile soils, principally on limestone sub- sediment materiał”. strate (mostly chalky). In Poland, this species Another species spreading along rivers is the has expanded above all along roads and the role Bidens frondosa, which expands its ranges in of river valleys is only secondary. However, in Europę along the Rhein and Elbe in Germany, Poland, similarly to its expansion in Western by canals in the English midlands (P reston et Europę, the species seems “attached” to calci- al. 2002 after Cadbury 1971), and the Loire in um-rich soils and generally to warmer regions France (K eil 1997 and references cited there). of the country (cf. Chapter 6). In Poland also, the first stages of occupying new Other species often referred to in conjunction sites are specifically associated with river val- with rivers are: the Echinocystis lobata, which leys (cf. also Chapters 6 and 7). Further east, originally occurs in riverine forests of North Bidens frondosa is currently expanding along America but is now going into the wild State in the Neman (Lithuanian: Nemunas) (Gudżinskas many European regions and spreading along ma 1997d). jor or middle-sized rivers (G u d ż i n s k a s 1999a; In characterising the distribution of Eragros- D a j d o k & K ą c k i 2003; cf. also Chapters 6 and tis albensis (earlier described as E. pilosa) in the 7), and Mimulus guttatus, which is associated Vistula river valley, S u d n ik -W ó jc ik o w s k a & G u with the Neris and Neman rivers in Lithuania z ik (1996) emphasise the important role played by (G u d ż in s k a s 1998a), whereas in Poland it spreads the natural conditions still prevailing along this, along brooks in the Sudety Mts. (F a b i s z e w s k i the largest river of Poland74. In the spreading of 1985; F a l iń s k i 2000b; K w ia t k o w s k i 2003), as this species the fine and light-weight seeds are well as along some rivers in the Beskidy Mts. and transported by water, thus floods and surges of in northern Poland (e.g. upon the Łupawa river, waters also play a certain role in the process. pers. observ.). S u k o p p (1998) points to the proportion and already permanent presence of some alien species in the summer therophyte communities developing on the draw-down zone of water courses75. 9.6.2. The role of humans in the migrations of kenophytes 74 “In the middle course of the Vistula, where its flow and course are unconstrained, natural habitats still predominate. The river has a slow current here and the width of “Long-range transport” the valley ranges from 1 to 14 km. The Vistula forms river arms, the current often changing direction among nume- One cannot overestimate the role of humans rous sandy banks and holms within the river channel” in creating opportunities for species to reach (S u d n i k -W ó j c i k o w s k a & G u z i k 1996). new territories. Although an “enormous number 75 “It has become elear that agriophytes play a major role of alien newcomers travelled over Atlantic in the floristic structure of therophyte communities, which without any prior intention or knowledge of develop every summer on initially bare riversides as the M an” (K o r n a ś 1996), an eąually large group water level falls” (S u k o p p 1998). was imported by humans on purpose (cf. Chap- laid on sandy embankments where competition ters 5 and 9.4, App. A and B). Unintentionally from other species is non-existent”. This author aided species have utilised enormously diverse enumerated such species as: Anthemis ruthenica, means of transport: with seeding materiał, Chamomilla suaveolens, Artemisia austriaca. The animal fodders, wool, packing stuffs, wood and, issue was then re-addressed by K o r n a ś et al. above all, with ballast earth, bringing with them (1959), emphasising that “the massive transport an entire “bank” of seeds, spores and all other of goods over long distances provides convenient kinds of plants (K o r n a ś 1996) (cf. also Appen- conditions for the transfers of seeds, fruits and dix A). other diaspores”. Accidental introduction has been also facilitat- This manner of spread is utilised by such plants ed by human migrations during past wars and post- as Acer negundo, Artemisia austriaca, Cardaria war periods and by armies marching through76, draba, Centaurea diffusa, Diplotaxis muralis, Era- while in modern times the process is chiefly fa- grostis minor, Impatiens parviflora, Linaria cilitated through the exchange of goods now ef- repens, Potentilla intermedia, Sisymbrium altissi- fected on a global scalę, as well as by dynami- mum, Solidago canadensis, species of the genus cally growing tourism. Oenothera, and in recent decades also by Rumex Once introduced somewhere, the migration of confertus (cf. Chapter 7) (data obtained from au- plants has tended to follow suitable natural con- thor’s own research and that of other authors, e.g. ditions (such as the aforementioned river valleys) U rba ńsk i 1958; R ostański K . 1960; Ć w ik liń sk i as well as making use of the opportunities pro- 1968, 1972a, 1990; S e n d ek 1971; M ich a l a k & vided by humans (through economic develop- S en d ek 1974-1975; S zm a jd a 1974; L ato w sk i ment77, the cutting down of forests, settlement de- 1977; R o sta ń sk i K . et al. 1989; N ow ak 1997; velopment, trade, the construction of roads and W ą so w icz 2003). railways, sea and river ports etc.). Some species have even been deemed to be types of “railway specialists” (K r a w ieco w a Transport routes 1951; K o r n a ś et al. 1959; S e n d ek 1971; J e h lik The possibility of accidental introduction along 1981). Railway type of habitats provides them railway lines was pointed out early by P a c zo sk i with conditions similar to those they select in (1900)78: “Railways are extremely conducive to the accidental introduction of plants which has to their respective homelands (Z a ją c E.U. & Z a ją c A. 1969)79, and further equivalent habitats can be attributed not only to transport and dissipation of seeds, but is also to the fact that the tracks are be found in heaps, rubble dumps, roadsides. Both railways and roads facilitate penetration by adventitious species into communities of indi- 76 S e m p o ł o w s k i (1880-1881) provided such description genous plants or inerease their vertical ranges of the routes and manners of the accidental introduction of Xanthium spinosum to Central Europę: “[...] it is said that in mountains. the Russian armies marching through Vallachia brought it there; according to eye witnesses many characteristic spiny fruit of this plant could be seen entangled in manes and tails, particularly of Cossack horses”. Further expansion has “Even smali countries typically contain been facilitated chiefly by pigs, sheep and the wool of the many different habitats latter: “Spiny fruit of the cocklebur cling easily to the as well as biogeographically distinct subregions, bristles of pigs, particularly Serbian and Hungarian [pigs], each of which may have a uniąue invasion history which have long and curly bristles and the pigs transfer the and be differently susceptible to invasion”. fruit sometimes to very remote locations. This fact is easily Su k o p p 1998 noted because this weed appears in large numbers along the tracks where pig herds have been driven either to Hun- 10. Recent distribution ranges gary or across it, especially at the sites of longer rests. of kenophytes and principles Since the time when railways were used to transport pigs to northern Germany (e.g. to Hamburg), the cocklebur has affecting the distribution pattern been seen along the relevant railway routes. The species reached Vienna and some regions of Germany with contaminated wool”. Initially it was found around wool ware- When considering plants of alien origin, it is houses and cloth factories. In Bukovina it was noted in as difficult to reconstruct the history of their ex- 1830 and because its discovery coincided with a cholera pansion as it is to analyse their contemporary epidemie there, thus the local peasants gave it the verna- distribution ranges. The pattern of the distribu- cular name of “cholera thistle”. 77 In some periods, the accidental introductions of alien plant species were helped to a large extent by the cloth 79 Habitat conditions prevailing at railways sites are spe- manufacturing and food processing industry (grain eleva- cific. Among the essential ones are the predominance of tors, mills). skeletal structures in the substrate, excessive dryness and 78 T h e l l u n g (1918-1919) noted the same. insolation. tion rangę of synanthropic species is affected by data provided by these authors, among the keno- many factors, both historical (time, manner and phyte species which are often found in the Car routes of introduction), as well as those related pathians, 14 kenophytes are plants associated to the biological properties of a given species with anthropogenic habitats (epecophytes), (e.g. life strategies, means and rate of dispersion whereas 23 species (agriophytes) penetrate into of diaspores) and by the specific conditions natural communities. Among the kenophytes a species encounters in the new homeland. found more rarely there, 27 species are epeco Among the factors operating on a large (glo- phytes, while 33 are agriophytes. bal) scalę, climatic factors are all-important80 The number of kenophytes falls in line with (either as limiting or advantageous factors de- the rise in elevation above sea level. Kenophytes pending on the climatic zone from which a new- tend to concentrate in the lower mountain sites comer originates, i.e. on the conditions in which (up to 500 m a.s.l.) which in the Polish Car it grew in its homeland). Other factors of para- pathians means a zone in transition between the mount importance are potential natural barriers foothills and the lower montane zones. The (oceans, seas, mountain ranges, vast forest com- species associated with some extra-zonal hab plexes, deserts). The specific habitat conditions itats such as Juncus tenuis (paths, roads) and found in the new site (land relief, soil types, Solidago gigantea (river and stream banks) can presence of water courses, land use etc.) are only reach the highest elevations attained by keno of secondary importance. phytes. The upper limit of cultivated fields (ca. In the conditions prevailing in Poland, many 700 m a.s.l.) is reached by kenophytes consti- kenophytes find an evident climatic barrier lim tuting field weeds: Conyza canadensis, Oxalis iting their occurrence in the mountains (particu- fontana, Veronica persica, Galinsoga parviflo- larly noticeable in the Carpathians) and in ra and G. ciliata. north-eastem Poland81, as well as in some locally Similar relationships were found in the Sudety cooler regions (cf. Chapter 7). In the moun Mts., the Śnieżnik massif and Bialskie Mts. tains, particularly at higher elevations, there are (S z e l ą g 2000), where rare and very rare species, no kenophytes of Irano-Turanian or Mediterra- with occurrence limited to the lowest elevations, nean origin (they may occur only locally in predominate among some 60 species of keno larger cities or along railway tracks and embank- phytes recorded. The list of kenophytes which are ments, e.g. Sisymbrum altissimum and S. loe- found freąuently and also recorded in sites with selii, Lycium halimifolium, and Datura stramo- elevations of ca. 700 m a.s.l., and sometimes even niurń). 900 m a.s.l. includes: Chamomilla suaveolens, Of the overall number of 123 species of keno Galinsoga ciliata, G. parviflora, Tanacetum phytes recorded in the Carpathians, 42 species parthenium and Yeronica persica - associated have fairly numerous stations in the region, with anthropogenic habitats, and Impatiens parvi- whereas 81 species have only isolated stations flora, Juncus tenuis and Rudbeckia laciniata - (Z a ją c M. & Z a ją c A. 2001). According to the which are also established in natural and semi- natural habitats. 80 The climate may be considered а сотріех of factors On the other hand, one can identify some spe setting the broad limits for plant distribution, while other cific features of the floras in the mountain regions factors, such as geology, soils and competition, will deter- of Poland resulting from the presence of keno mine the presence or absence of a species in a particular phytes. In the Bieszczady Mts. these will be newer area and on a finer regional or local scalę (W e l k et al. arrivals penetrating natural communities: Bunias 2002). For ехатріе Reynoutria (Fallopia) japonica was found to be controlled by two climatic variables - the orientalis, found on roadsides, alluvia, anthropo length of the growing season, measured in day-degrees, and genic habitats and moving on to natural commu the minimum temperature - while for Impatiens glandu- nities (up to 630-740 m a.s.l.); Juncus tenuis lifera only the length of the growing season was critical spreading along paths, up to 630-900 m a.s.l.), (B e e r l i n g 1993; B e e r l i n g et al. 1995). Rudbeckia laciniata, a plant which occurs, some 81 The kenophytes which have also successfully estab- times in massive numbers, on the edges of river- lished themselves in the ruderal floras of the Northern Podlasie Lowland include the very common species Cha- ine woods and scrub (up to 630-750 m a.s.l.) and momilla suaveolens, Amaranthus retroflexus, Galinsoga Yeronica filiformis, not very common but locally parviflora and the rare Datura stramonium (W o ł k o w y c k i abundant (up to 650-800 m a.s.l.) (Z e m a n e k & 1997). Elsholtzia ciliata, a species absent from central and W in n ic k i 1999). northern Poland and even recently withdrawing from ear- In many regions of the western Beskidy Mts., lier stations (cf. Chapter 7) constitutes, along with some species belonging to other geographical/historical groups, particularly in lower sites, the species Digitalis a specific feature of the ruderal floras in villages and smali purpurea, Heracleum mantegazzianum, Impa towns in that region of Poland. tiens glandulifera and Reynoutria japonica, and in the Sudety Mts.: Mimulus guttatus, Impatiens Although in the case of this large group of glandulifera and Reynoutria japonica (F a b is z e w species no specific pattems of distribution can be s k i & K w ia t k o w s k i 2001; K w ia t k o w s k i 2003) detected, nevertheless for one particular sub- undoubtedly belong to this characteristic group group (of 14 species) some centres of concentra of species. tion of stations can be demonstrated, e.g. within There is a lack of detailed studies devoted to the Silesian Upland (and particularly the Upper the autecology of individual species which could Silesian Industrial Region - GOP83) and within elucidate the naturę of the above phenomenon large cities (cf. Chapter 6). The pattern of dis (Z a ją c M. & Z a ją c A. 2001). One pioneering, tribution for these 14 species which at present and so far the only Polish study on this topie, show certain relationships with urban areas, rail- is a monograph by K o r n a ś (1972) devoted to the way lines and roads coincides mostly with the dissemination of weeds in the Gorce Mts., where “heat islands”, or “zones of influence” of urban the author proved, inter alia, that the proportion centres (J a c k o w ia k 2003 after Różański 1979) of anthropophytes fell and the proportion of (Fig. 67). apophytes inereased in line with the elevation Some species show certain pattems of distri above sea level (cf. also Chapter 8). For several bution associated with local habitat conditions species of kenophytes spreading in the Gorce (e.g. the local distribution of Acorus calamus or Mts. as weeds in cultivated fields, maximum Elodea canadensis is determined by the presence elevations were given: Vicia dasycarpa: 560 m of suitable habitats, thus these species will be a.s.l. (average: 499), Galinsoga eiliata: 655 m found on rarer occasions in those regions where a.s.l. (average: 636), Oxalis stricta: 705 m a.s.l. a hydrographical network is less developed, such as the Kraków-Wieluń Upland). (average: 668), Galinsoga parviflora: 730 m There is also a smali group of species which a.s.l. (average: 654), Yeronica persica: 965 m currently form compact ranges limited to a spe a.s.l. (average: 910). cific region of Poland. The group includes, for Specific ecological conditions (type of sub- example: Erechtites hieracifolia (the majority of strate or soil) may locally limit the rangę of a its localities is concentrated in south-western species despite generał climatic conditions being Poland), Solidago graminifolia (Silesian Low- potentially favourable for its occurrence (W e l k land), Trifolium patens (Carpathian Foothills, et al. 2002). In Poland such a relationship is, for Rzeszów Foreland and Małopolska Upland), and example, manifested by Anthoxanthum aristatum, Yeronica filiformis (south-eastern Poland). The a species whose distribution in Poland is deter- current patterns of their distribution are not mined by soil conditions82 (cf. also Chapter 7). simple reflections of climatic conditions, but In the regions potentially less favourable to also bear the marks of the history of their re- dissemination, there might be suitable conditions spective arrivals into Poland. These species were prevailing locally, e.g. in the form of extra-zonal accidentally brought into a single region (or no habitats in a generally dry climate. These might more than a few regions) of Poland and then be: river valleys, wet or shaded sites, northem spread out gradually (P i e t r a s 1970; L o s t e r slopes and mountain habitats, as well as some 1972; G ó r s k i et al. 2003; T o k a r s k a -G u z ik & ruderal habitats in cities. D a j d o k 2004). The majority of kenophytes occurring in Po Only a smali number of kenophytes can be land, however, do not form characteristic distri considered as has recently been reaching any bution ranges over the entire national territory nor limit of their ranges in Poland (cf. Chapter 6). locally. They are mostly ubiąuitous species whose These species (apart from a few exceptions - cf. history of establishment has nevertheless one Chapter 11) still have not completed their mi- common feature. These species were accidentally grations and one may suppose that the area of brought (often repeatedly) or they were (or are) Poland, generally devoid of major barriers, is cultivated in many places in Poland, and have succeeded in spreading from these places in many directions (e.g. Chamomilla suaveolens, Conyza 83 The Silesian Upland and the Upper Silesian Industrial canadensis, cf. Chapters 6 and 7). Area (abbreviated as GOP) are the two most disturbed regions in Poland (T o k a r s k a -G u z ik & R o s t a ń s k i 2001); in terms of the effects on climate, the GOP should be treated 82 W archolińska & S o ń s k i (1996) suggest that Anthoxan- as a single urban-industrial complex constituting a “ther- thum aristatum finds optimum conditions for development mal island” in the atmosphere (K r u c z a ł a 1972). The re on the sandy “bielitza” soils of the weak rye soil or rye- gion^ population amounts to 2.178.400, its population lupine soil complex; another important factor favouring the density (1.720 residents/km2 is the highest in Poland, where concentration of stations in central Poland is the impact of the average is 124 residents/km2) and in the European the oceanie climate. Union (116 persons/km2). Fig. 67. Concentration of kenophytes associated with urban areas and railway routes with respect to the influence of towns (big dark gray circles) on the thermal conditions in the region on the ехашріе of the relation of artificial heat emission (gray spots) to the solar radiation in Poland (source: J a c k o w ia k 2003, significantly changed) not going to be any great obstacle in their fur- based on presumptions (or even speculations)84. ther expansion. Eąually rare are efforts to forecast the limit of As mentioned earlier, some of the species are the synanthropic rangę based on bioclimatolog- currently expanding their ranges in the same con- ical data. An attempt at such a forecast, based tinent (this pertains to species of European ori- on GIS methodology, was made by W elk et al. gin), while others developing disjunctive, second- (2002)85. ary ranges. The species which have expanded their ranges in Europę in recent centuries include, inter alia, Anthoxanthum aristatum (Fig. 68) (cf. 84 These conclusions take into account the origin of diaspores which might be locally adapted to specific con also Fig. 51 in Chapter 7), Artemisia austriaca ditions. Also considered is the fact that it is sometimes only (Fig. 52 in Chapter 7), Clematis vitalba (Fig. 40 random samples of the genetic diversity of the species in Chapter 7) and Rumex confertus (Fig. 55 in which are brought accidentally from their natural ranges. Chapter 7). Finally, whether a species can change its life strategy in P y śe k (2001) states that forecasts and esti- certain circumstances is also discussed. mates pertaining to the distribution of other 85 In that study the authors presented the analysis of the relationship between the distribution within the natural species have been formulated very recently, thus rangę and the spatial interpolation of the average monthly there has been too little time allowed to pass any temperature and precipitation conducted for Alliaria petio- judgement as to their merits. The conclusions lata (Garlic Mustard), a species native to Europę while found in published studies on this topie are being regarded as invasive in North America. natural rangę according to M eusel e t al. 1965;

expanding rangę:

with high number of localities

with Iow number of localities

Fig. 68. Changes in distribution rangę of Anthoxanthum aristatum Boiss. in Europę

11. Dynamie tendencies in the process 49 species; to 1900 - 117 species; to 1950 - 143 of kenophyte expansion in Poland species and to 2000 - 174 species, respectively). The same tendency was apparent for the total number of recorded localities (in the 50 years to As mentioned earlier, the initial stages of an- 1850 - 151 localities; to 1900 - 3 675; to 1950 thropophyte species migrations are linked to the - 9 273 and to 2000 - 196 441 localities, respec- introduction of agriculture (earlier periods are tively) (cf. Chapter 5.2)86. currently impossible to reconstruct; some hope in The number of localities shows an especially this respect may be linked with the results of ar- rapid growth. Nearly 94% of the total number of cheobotanical studies). Since that period, the localities for the above-mentioned 174 species process has continued with varying intensity up have been recorded during the last half-century. to the present time. This fact must be linked mainly to the inerease in The dramatic velocity of these migrations is often stupefying; it usually takes between several 86 It should be taken into account that the presented dozen years to two hundred years for a newly listings refer to data accumulated in the ATPOL database arrived species to fili totally the potential area of on the basis of available sources. The density of kenophyte its occurrence in its new homeland (K o rnaś occurrence presented on maps for consecutive half-cen- 1996) (see also Chapter 7). tury periods during the last two centuries should be regarded only as an approximation when interpreting in terms of The rate of spread in Poland has been recon- the dynamie tendencies of this group of anthropophytes structed for 174 kenophyte species (see also spreading in Poland. The maps which illustrate the earliest Chapter 4 and Appendix A), for which detailed reconstructed periods do not show the presence of some information about the number of localities of oc species due to the process of data acąuisition and presen- currence (historical as well as current) has been tation peculiar for that period of scientific research (lack gathered. In successive 50-year periods, the num of precise data on locality, see also Chapter 4), whereas the maps from subseąuent periods are also affected by differ- ber of new arrivals recorded for the flora of ences in the degree of thoroughness of research conducted Poland grew steadily (in the 50 years to 1850 - in each region of the country. the intensity of studies of the synanthropic flora tion and seed dispersal. Species which show a better in the post-war period (we can thus refer to it as strategy in this respect with regard to the conditions a partly “spurious” increase in the number of lo- found in the new territory of occurrence are usu- calities). In many cases, however, we have data ally characterised by a faster rate of migration. proving that an actual increase in the invasion rate Among 174 kenophyte species for which dy took place for many species. An example for this namie tendencies have been identified, species type of case is Echinocystis lobata - the history with a relatively high number of localities pre- of its arrival and the consecutive phases of occu- dominate, but their distribution is usually limited pation of new localities by this species have been to a specific part of the country (Fig. 69; cf. recorded rather precisely. The history of spread of also Fig 15A & B in Chapter 5 and Chapter 6). this species in the territory of Poland encompasses They are also at the same time those species the whole of the last century when the acqui- which gradually occupy new sites. The least sition of floristic data had proceeded relatively sys- numerous group is formed by common and tematically (with an interruption during World War broadly distributed species (with a very high II), although the intensity of research was variable number of localities). They also predominantly from one region of the country to another (see also include the species which are still spreading: they Chapter 7; Fig. 48). occupy new sites and at the same time in many The migration rate mainly depended not on the cases they increase the number of individuals in mode of translocation from one site to another the populations at all localities (the so-called in- (the spreading of seeds), but rather on the resis- vasive species - see Chapter 12). tance of the environment to colonisation. An W e b e r (1998) has reconstructed the pattem of important factor was the way in which the im- spread shown by 3 species from the genus Solida- migrant species had been introduced. Those spe go in Europę. These species, originally from North cies which spread in anthropogenic habitats (at America, were introduced into Europę as omamen- least in the initial phases of their migrations) and tal plants and as nectar sources for honey produc- which had been introduced simultaneously into tion: S. canadensis (altissima)87 in ca. 1735 (vicin- multiple regions in Poland were characterised by ity of London) and the remaining two species a high invasion rate. This pattem applies inter probably around 1758 (see Appendix A), respec- alia to the following species: Amaranthus retro- tively. The first wild localities were recorded in the flexus, Chamomilla suaevolens, Conyza canaden mid-19th century. S. canadensis and S. gigantea are sis, Galinsoga parviflora and Veronica persica currently common in many regions of Europę and (see also Chapters 5.2 and 7). are considered to be “aggressive” invaders on A u l d & T i s d e l l (1986) have shown that the abandoned fields and river banks, also in protect- increase of total area occupied by an expanding ed areas (S u k o p p 1966; G u z ik o w a & M a y c o c k species is faster when several smali independent 1986; W e b e r 1998; B a l o g h 2001). The compa- populations take part in the expansion, than when rison of data regarding the number of localities, there is one large spreading population. The time starting with 1850s, shows in the case of all spe and mode of introduction are also of importance. cies a continuous tendency to spread, albeit with Subseąuently, natural and anthropogenic factors a varying rate. Solidago gigantea is characterised decide whether a species will spread ąuickly or by the fastest expansion rate, while the slowest one slowly and what type of rangę it will adopt. was recorded for Solidago graminifolia. As no- This hypothesis is proven in the Polish circum- ticed by W e b e r (1998 after H e n g e v e l d 1989), the stances by the following species (in addition to the spread of these species is not reflected in the oc ones listed earlier): Trifolium patens, Yeronica currence of a conspicuous rangę front, but is ef- filiformis and Mimulus guttatus (see also Chapter fectuated according to the model of hierarchie 7), which after having been introduced into a sin diffusion. The spread by large jumps with subse- gle region subseąuently spread gradually in that ąuent local spread in all directions is defined as specific part of the country (the principal factor the hierarchie diffusion model (H e n g e v e l d 1989) that was decisive for the possibility of efficient nat- and might be the most applicable spread mode for invaders introduced as omamentals (W e b e r 1998). uralisation of a species in a given region of the A similar expansion rate has been shown by country was obviously the climate; see also Chap these species in the area of Poland (G u z ik o w a & ter 10). It took another introduction event or the appearance of the species in another region for the species to be able to spread its rangę further, on 87 The taxonomie status of this species in Europę is conditions that the other region also had favour- unclear; S. altissima and S. canadensis are often not distinguished in the literature. On the basis of morphological able conditions for the naturalisation of this spe characteristics it may be inferred that the species occurring cies (see Mimulus guttatus - Chapter 7, Fig. 50). in Europę is S. altissima. However, due to the mentioned The biological properties of a species are also doubts they are both still treated as a single species (W e highly relevant, especially the modes of reproduc- b e r 1997b & c, 1998). Fig. 69. Dynamie tendencies of 174 species of kenophytes occurring in Poland (according to Z a r z y c k i et al. 2002; for more explanation see also Chapter 4) Freąuency in the wild at the territory of the country in relation to the number of localities: 1 - very Iow number of localities (1-20), 2 - Iow number of localities (up to 100), 3 - high number of localities, but with narrower distribution (in one or two regions of the country), 4 - high number of localities in many regions, 5 - common (abundant) in the whole territory. Dynamie tendency: (-2) - high decrease of(in) number of localities, (-1) - decrease in number of(in) localities or decrease in abundancy over existing localities, (+1) - inerease of(in) number of localities, inerease in abundance over existing localities, (+2) - high inerease of localities (colonizing new localities), (-/+) - disappearing of some localities and appearing of new localities, (?) - undefined dynamie tendencies

M aycock 1986) (Fig. 70 & Fig. 71). One might subsp. ruthenica spread significantly more slowly. accept the prognosis of W eber (1998) which sug- It seems that in the case of these species also, gests that these species will continue to spread by the decisive influence on the rate of spread was inereasing both the number of occupied localities the naturę of the wilful or accidental introduction and the number of individuals at each locality. (at at least several dispersed points), and of sec- Only Solidago graminifolia is characterised by ondary impact were the types of habitats occu a slower rate of expansion. In Poland, this species pied by these species as well as their biological was recorded for the first time in Lower Silesia properties. near Niemodlin in 1888 (see Appendix A). Cur- The influence of elements such as the time of rently, its occurrence is limited to south-westem introduction and the biological characteristics of Poland with individual dispersed localities in the a species on the variable rate of spread may be centre of the country (Fig. 70). Nevertheless, illustrated from the example of two closely related during recent years a significant inerease can be species from the genus Galinsoga which are observed both in the number of sites of occur currently common and freąuent kenophytes in rence and in the size of the populations of this nearly the whole area of Poland (Fig. 73). Ga species in regions linked with its longest-lasting linsoga eiliata occupied new localities at a slower presence (Lower Silesia) where it colonises pace than Galinsoga parviflora, presumably be- mainly wet meadows and disused quarries cause the former species had been introduced (T o k a rsk a -G uzik & D a jd o k 2004). later (see Appendix A) and is able to spread its The comparison of rates of spread for selected seeds for shorter distances due to their higher kenophytes (neophytes according to the cited weight. These conjectures are supported by typ- authors) in the area between the Oder and the ical information found in many local floristic Elbe (H ardtke et al. 1 9 8 1 ) and in the territory studies (dating back even to the 1970s) where of Poland has led to similar results (Fig. 72). their authors characterise G. eiliata as: “a rare A faster rate of spread is characteristic for two weed in the initial phase of spreading” species: Sisymbrium loeselii and Rudbeckia lacinia- (B ła szczy k 1959); “rare, only several specimens ta, while Cardaria draba and Salsola kali found” (M azu r et al. 1978); or “with a distinctly 195 0

2000

Solidago canadensis L. Solidago gigantea a it o n Solidago graminifolia (L.) e l l i o t t

Fig. 70. Recorded history of expansion of three species of Solidago in Poland (after T o k a r s k a -G u z ik 200 lb; Z a j ą c A. & Z a j ą c M. 2001)

® i£ 10000-1 - O -43 - - Solidago canadensis a - 1000 > o — Solidago gigantea

A Solidago graminifolia

Year Fig. 71. Increase in the cumulative number of localities of three species of Solidago in Poland Germany 8 140 ro o 120 **—0 jj 100 E z 80

0 up to 1850 1851-1900 1901-1950 1951-1980

Poland jg 2500-1

1 £0 2000 - i— .a0 1 1500 c 0> i 1000- E O 500-

0 up to 1850 1851-1900 1901-1950 1951-1980 Historical sequence

Fig. 72. Comparison of the rate of spread of four alien plant species in Germany (H a r d t k e et al. 1981) and in Poland

lower number of localities” (e.g. S z m a jd a 1974; differences between these two species noted in M a c ie j c z a k 1988; C h m ie l 1993). Poland and other countries can be connected with On the other hand, in Lithuania, according to climate factors and history of introductions of the G u d ź in s k a s (1997d), Galinsoga ciliata is charac species but also can be caused by erroneous iden- terised by a different rate of spread than the one tification of the two species (particularly at the reconstructed for Poland. There, this species is beginning of their spread in Poland). much more “aggressive” than G. parviflora, Locally, however, species which one would although it again started to spread at a later date. expect to realise a similar spreading strategy due to Also apparently in UK, where G. parviflora was their close relationship88 may often increase their introduced into Kew Gardens in 1796, and was first recorded in the wild in 1860 (and was known 88 Often related species are characterised by similar as “the Kew weed”) and has spread steadily geographical origin (e.g. I. glandulifera and I. balfouń which sińce. G. ciliata was first recorded in the wild in also occurs in some regions of Europę are both originally 1909 and has spread more rapidly and now has from the Himalaya mountains) as well as seed dispersal mechanisms (W a d e 1997 and the literature cited therein), a similar rangę to G. parvi/lora (P r e s t o n et al. but their mode and rate of spread may be different in many 2002). The original introductions seem to have regions of their secondary rangę. The above-mentioned been supplemented with later ones from nurser- I. balfouri, even though it was recorded in 1979 on the ies and in wool waste used as an agricultural Thames in London, has not managed to become naturalised in the British Isles, while it is already a naturalised spe fertiliser (a very important source of adventives cies in riverside and ruderal habitats in other regions of in UK; Prof. I.C. Trueman, pers. comm.). The Europę (e.g. in Croatia, pers. observ.). Galinsoga parviflora c a v . Galinsoga ciliata (R a f .) s .f . b l a k e

Fig. 73. Recorded history of expansion of two species of Galinsoga in Poland (after Z a j ą c A. & Z a j ą c M. 2001, supplemented) secondary ranges at a different rate. Perrins et al. White Horehound Marrubium vulgare, ori- (1993) have estimated the rate of invasion for 3 ginally native to the Mediterranean region, has species from the genus Impatiens: for I. glandulifera been cultivated in Poland probably as early as 38 km/year, for/. parviflora 24 km/year and 13 km/ the 16th century. The first locality was recorded year for I. capensis. As a result, these species which from Gdańsk in 1643, the next one - after nearly occur currently in the British Isles are characterised two centuries - was reported from Wyszogród. by different status and distribution pattems. The 3 It is a medicinal plant with a relatively wide ap- species mentioned also occur in Poland, but in the plication and most probably was more often cul- circumstances of our country their distribution and tivated earlier than can be inferred from the status are divergent from the respective character- number of historical localities of occurrence of istics in Britain (Table 11). this plant. In the late 19th and early 20th century Table 11. Comparison of invasive status of three species of it was probably much more freąuent than it is the genus Impatiens from Great Britain (P e r r in s today (Fig. 74). The history of naturalisation of et al. 1993) and Poland Marrubium vulgare in Poland dates maybe all Freąuency and status the way back to the Middle Ages. The status of Species Great Britain Poland this species is uncertain, some authors consider Impatiens invasive, common invasive in S it to be an older arrival, one of the so-called ar- glandulifera chaeophytes (S udnik-WOjcikowska 1987a; Impatiens invasive, local invasive, common Jackowiak 1992; R utkowski 1998). It occurs panńflora currently over the whole area of the country, Impatiens invasive in S & E not invasive, limited although it is not everywhere freąuent (it has not capensis distribution been recorded from some regions, e.g. in the mountains it is rare and occurs only at lower A very smali group among kenophytes is elevations). formed by species which are decreasing their The habit of occurrence of both species: dis area of occurrence (withdrawing species) or the persed localities, usually near sites of old cul- ones which are not currently spreading (they tivation, as well as the type of occupied hab usually persist on the previously occupied local- itats, such as roadsides, old lawn plots, old walls, ities) in the territory of Poland. This group vicinities of allotment gardens, points to a still includes e.g.: Ambrosia artemisiifolia, Coryda- conspicuous link with human activity. Both spe lis lutea, Cymbalaria muralis, Helleborus viri- cies were probably much more common at the dis, Hyssopus officinalis, Lathyrus nissolia, turn of the 19th century (Fig. 75). These species Marrubium vulgare, Mercurialis annua and do not show a tendency to spread, which is Oenothera cruciata - species which were never probably a result of the fact that they are no freąuent in Poland, even in the periods of the longer commonly cultivated. It may, however, recorded increase in the number of new local- be presumed that the recurring increase of in- ities. The progressive decrease of their occur terest in the cultivation of medicinal plants rence may be explained by the elimination of (herbs), observed currently also in smali gar specific habitats which they were linked with, dens, will assure such species as the horehound as in the case of Corydalis lutea and Cymba and the hyssop a permanent presence in the flora laria muralis (see also Chapter 7), as well as the of Poland. ever-diminishing freąuency of their cultivation The data of other botanists also tend to con- in modern times. firm the decrease in occurrence of the kenophyte Common Hyssop Hyssopus offwinale, originat- species cited here. Similarly, Mercurialis annua ing from Southern and eastem Europę and from has been reported as being widespread from the south-western Asia, was introduced into garden territory of Poland by B esser (1809) and B er- cultivation in Poland in the 17lh century (see dau (1859), but, starting from the end of the 19,h Appendix A and Chapter 5.2) or even earlier in century, it has been found more and more rare- monastery gardens. It is a plant which has been ly (R aciborski 1884; Trzcińska-T acik 197la ). cultivated in many countries for a very long time, Gudźinskas (1999a) reports that this species has it is used in the cosmetic industry, it is also become more and more scarce in the Baltic a melliferous and medicinal plant. It may be pre- countries during the last ten years; it still per- sumed that the localities reported during the ini- sists in Klaipeda. tial stage if naturalisation of this species in Po A tendency to decrease their ranges and even land are sites of its escape into the wild near to withdraw from the localities which were oc localities of its cultivation. Currently, it occurs at cupied years ago is being shown also by species dispersed localities in the whole country. which were at some point in time considered to Introduction as cultivated plant Naturalisation and spread closeto sites of cultivation o dubious records: most probably the records refer to ® oldest recorded localities of occurrence in many the localities from cultivation regions; some of them probably refer to sites of its cultivation

Subsequent phases of spread o records dated back to the previous periods of 1851-1900 and 1901-1950 Fig. 74. Recorded history o f cultivation and spread o f Marrubium vulgare L. in Poland - an example o f a species decreasing its area of occurrence

be frequent, at least locally. These species include towns leads to the dwindling of areas which are e.g. Elsholtzia ciliata listed in many old floras occupied by habitats of the Urtico-Malvetum (e.g. Rostafiński 1872; C ybulski 1894; Czyrsz- association which leads to a decrease in the nicówna 1929; K obendza 1930; G rochowski number of sites of occurrence of this species 1931). This species was still spreading westward which is tightly linked with this association (Frey in the early post-war period (see Chapter 7). 1974) not only in Poland, but also in other coun- During the last 20 years, its occurrence has not tries in Europę. been confirmed at many of its earlier localities There has been rather little study of the vari- or the population size has been found to be very ation in rates of spread of alien plant species limited (own data). This conjecture is also (W illiamson et al. 2003). The authors concluded confirmed by data found in some local floras that the rates of spread of species in the same dating from recent years (e.g. Chmiel 1993). genus are both very similar and very different and A similar situation concerns Amaranthus as- that explanation of variations in rate of spread are cendens. The urbanisation of villages and smali likely to remain case by case. Recently, some o -O 3E 50 - z

. ,n -,P- -,FL, .łłłiiiirLY 1 i 1600 1700 1800 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 Years

L □ Marrubium vulgare Hyssopus officinalis

Fig. 75. Comparison of the ratę of spread of Hyssopus officinalis and Marubium vulgare in Poland

long-term studies have been undertaken which use in applicational publications (legał texts, make it possible to tracę the rate of spread for offieial decrees etc.) (T o k a r s k a -G u z ik 2003a & alien species on a varying time scalę. b). These terms are often applied interchangeably, e.g. by T r o ja n (1975): “Invasion or expansion denotes settlement of individuals in new territory which has hitherto not been occupied by any “The species Homo sapiens itself is without question the super invader of all time” population of this species”.

W agner 1993 The authors of the principal Polish academic handbook of plant geography consider invasion to “Humań activities do not only destroy habitats, but be “a spectacular form of massive expansion of they lead - together with climate change - to the spread of species beyond their natural ranges. Alien a recently arrived alien species which appears sud- species may threaten the indigenous flora, completely denly and so abundantly that it can cause signif- change the character of the place they invade, cause icant ecological disturbances and severe economic diseases and be pest organisms”. losses” (K o r n a ś & M e d w e c k a -K o r n a ś 2002)89. “The problem of biological invasions is growing in According to these authors and to other Polish re- severity as global trade and travel accelerate”. searchers (F a l iń s k i 1998a & c; J a c k o w ia k 1999), “Habitat disturbances and biological invasions create plants which inerease their abundance and area contact zones between eon- and heterospecific of occurrence due to human activity (so-called populations which were isolated by distance and/or by the environment”. hemerophilous species) include both native spe

d e n N uss et al. 1999 cies derived from local natural communities and alien species90. 12. Plant invasions: the substance of the phenomenon and kenophytes 89 Already in the earliest Polish phytogeographical publi as invasive plants cations attention has been directed towards the specific phenomena which accompany plant migrations. P a c z o s k i (1900) has indicated the problem of migration rate and the time reąuired by plants to colonise new sites: “some species spread 12.1. More remarks on terminology with a speed nearly as ąuick as lightning, others obviously advance at a turtle’s pace and reąuire a very long time to occupy a very smali space” (cf. also Chapters 2 and 11). In Polish phytogeographical literature, despite 90 Both these groups are encompassed by the common the defined etymology and meaning of terms “mi- term “synanthropic plants” (K o r n a ś & M e d w e c k a -K o r n a ś gration”, “expansion”, “invasion” and the derived 2002; see also Chapter 3 - Terminology). Also in foreign literature some authors tend to accept a similar inclusive terms “expansive”, “invasive” (species), there is concept, e.g. W a d e (1997) mentions that invasions may still a tendency to a rather free interpretation of concern also native species which can also become weeds, these words, additionally compounded by their e.g. Urtica dioica and Typ ha latifolia. Jackowiak (1999) has suggested more specific order to make a comparative approach possible definitions of biological expansion, designating the (Pyśek 1995; R ichardson et al. 2000; T okarska- spreading of a species into anthropogenous hab- G uzik 2001b, 2003b; C hmura & S ierka 2004; itats within its natural rangę as “ecological ехрап- Pyśek et al. 2004)9'. sion”, while he called the spread of a species In the English language literature most stud- outside its natural geographical rangę “chorolo- ies adopt the biogeographical concept of “inva- gical (or territorial) expansion” (Fig. 76). Plant sion” which assumes that this process is a con-

Natural rangę Secondary rangę Native species Native species Native species Alien species Alien species Alien species in natural in seminatural in synanthropic in synanthropic in seminatural in natural

communities communities communities u. communities communities communities © 'C R k_ u. £■ L_ CD O) © © i_ l_ L- n i— N 1 N 1 (0 ra ro -Q -Q "ra n n N 2 15 N 2 "ro N 2 !E 75 w o U o. u o U) O) O) O) E 1 o E 1 o E 1 O) E 1 o o o o o o o o o o Ш Ш O) E 2 E 2 E 2 o E 2 Ш E 2 Ш 00 E 3 E 3 E 3 E 3 E 3 E 3

__ r ----- _____ m I—-— L—— ----- Spontaneous selection of native species under inereasing Selection of alien plant species in colonisation process into anthropopressure seminatural and natural communities Ecological expansion Chorological (territorial) expansion Expansion lnvasion lnvasion as special (spectacular) form of expansion lnvasion as biogeographical process lnvasion as naturalisation of alien species in natural communities (= neophytism)

F ig . 7 6 . Model of ecological and chorological (territorial) expansion of synanthropic plants (according to J a ckow iak 1999) together with “wide” and “narrow” understanding of invasion process: R, N, E - species with different abilities to cross the ecological and biogeographical barriers

migrations expressed as changes in their geograph sequence of intentional or non-intentional human ical rangę and new biological and ecological activity and as such includes only alien species phenomena caused by them have been illustrated (Pyśek 1995; P yśek et al. 2004 and the literature by F aliński (1968а, 1998а & c, 2000a & b, 2004). cited therein)92. In the Polish literature, this view Invasion (from the Latin word invasio = irrup- is represented by C hmura & S ierka (2004). tion, inroad) can be confronted with encyclo- In some publications, examples of an even paedic definitions: narrower understanding of “invasion” may be - armed intrusion into a foreign territory; attack, found, the term being explained as the process of inroad (Dictionary of the Polish Language, naturalisation of an alien species in natural com- PWN); munities (Fig. 76). In this sense, the definition of - spontaneous change of the rangę of a species invasion would correspond partially to the def linked to the initiation of a migration over inition of neophytism suggested by F aliński a relatively long distance simultaneously by (1998a & c). a significant number of individuals and to the These terminological arguments are a result of occupation by the species of areas not previously different approaches to the problem adopted by inhabited by its representatives (Szweykowska various biological disciplines (R ejmanek 1995; & Szweykowski 1993, Botanical Dictionary). P yśek et al. 2004 after R ejmAnek 1995); there is This understanding of the term “invasion” is akin to the term “ecological explosion”, or “territorial ex- 1 Precisely defined terms provide the basis for com- pansion accompanied by a tremendous increase in parative studies; they are also of a practical importance the number of individuals of a species in the newly (preparation of lists of invasive species; conservation of di- occupied territory”, introduced by E lton (1958) versity, combating the threat). It should be stressed here that who is regarded as a key researcher in this field. the value of any study depends especially on the correct In discussions by phytogeographers who study collection of information on the flora of a given area (taxonomy, locality, habitat) and its correct appraisal (status of the topie of invasions, terminological questions a species in a locality, its origin, time and way of arrival). are regularly addressed, not only for purely se- 92 e.g. C e l e s t i G r a p o w et al. 2001: invasive alien spe mantic reasons, but also for practical purposes in cies and expanding natives (apophytes). also a significant impact of the usually anthropo- changes occurring in time and space. Invasion centric viewpoint of researchers93. according to these authors consists of the follow- Analogously, in many studies the terms “nat ing basie phases: uralised”94 and “invasive” are applied inter- - transport of an organism onto its new locality changeably, whereas they are actually represented of occurrence; by two different phases of one continuous process. - naturalisation and increase of population size Aceording to some authors, the process of invasion at the invaded locality; consists of 3 stages: - regional spread from initial successful popula - introduction, tions. - colonisation, Invasion ecology from the population view- - naturalisation (e.g. C ousens & M ortimer 1995). point provides possibilities to explain invasion According to others (e.g. W eb er 1998), the success and the influence of the invasive species spread of an invasive non-native species in an on the existing components of the ecosystem. area where it has never occurred before involves Some authors distinguish species which be- four steps: come naturalised but pose no practical problems - the arrival of the species and the local intro (K o r n a ś 1990; W a d e 1997). The latter author duction of individuals in a habitat, distinguishes (as was done similarly subseąuently - the formation of a persistent founder popula- by R ic h a r d so n et al. 2000) the following tion by growth and reproduction, categories of plant species: alien - established - - deriving of new populations by transport of invasive (pest). The term “invasive” is used for diaspores to safe sites, an alien whose distribution and/or abundance in - rangę expansion by increases in the number a region is increasing, i.e. can be considered as and size of populations. a successful alien. Each of these steps is tightly linked not only to Following the above-mentioned terminology the character and autecological properties of P yśek et al. (2004) suggest definitions of terms a species, but also depends on various facets of associated with plant invasion and place these in human influence. the context of floras. The hierarchical scheme for A commonly accepted mechanism of such the suggested classification of alien plants con invasions is the so-called “enemy release hypoth- sists of after P y śek et al. (2004): esis” (ERH) (K eane & C raw ley 2002). A plant 1. cultivated plants devoid of the burden of natural enemies in the 2. plants outside cultivation new homeland may spread very ąuickly95. An 2.1. casual (not established/naturalised) important factor favouring invasion is also the 2.2. naturalised competition from native species which may be 2.2.1. non-invasive weakened due to the fact that native species still 2.2.2. invasive96 remain in conflict with their own “enemies” (spe- 2.2.2.1. not harmful cialised monophages which are usually non-ex- 22.2.2. transformers97 istent for the potential invasive species in its new 2.2.2.3. weeds98 homeland). The recent European strategy on invasive alien S hea & C hesso n (2002) use population eco- plants uses the definitions agreed by the Confe- logy theory to explain mechanisms of invasion; rence of the Parties to the Convention on Biolo- specifically, they take advantage of the notion of gical Diversity for the purposes of the CBD ecological niche to introduce the concept of “niche opportunity” which defines conditions 96 Definition: Invasive plants are a subset of naturalised favourable to invasion with regard to resources, plants that produce reproductive offspring, often in very natural enemies, abiotic conditions and interac- large numbers, at considerable distances from the parent tions between the listed factors in relation to plant, and thus have potential to spread over a large area. 97 Definition: Transformers - A subset of invasive plants (not necessarily alien) that change the character, con- 93 Invasion ecology has perhaps suffered more than other dition, form or naturę of ecosystems over substantial area. fields, sińce the notion of “invasion” freąuently evokes (Substantial means relative to the extent of that ecosystem.) anthropocentric concepts (aggression, assault, attack, en- Transformers are essentially equivalent with “edificators” croachment, incursion, infringement, intrusion, onslaught, (i.e. edifice builders), a term used in European, especially raid, etc.) (R ic h a rd so n et al. 2 0 0 0 ). Russian literature. Edificators are defined as “environment 94 R ic h a rd so n et al. 2000 have compiled a review of forming plants” (P yśek et al. 2 0 0 4 ). In Polish literature the dictionaries, encyclopaedias and naturalist articles in order term could be compared with neophytes sensu F a l iń s k i to compare the definitions adopted for the term “naturalised”. (1998a & c) as already mentioned in this chapter. 95 It may be significant for some species; for others 98 Definition: Weeds - plants (not necessarily alien) that disturbances in the environment are the most important grow in sities where they are not wanted and which have catalysts. detectable economic or environmental impact or both. Guiding Principles and understand “invasive alien 12.2. Conseąuences of invasions by species” as an alien species whose introduction alien species, legał regulations and and/or spread threaten biological diversity (G en o - methods of combating the threat vesi & S hine 2 0 0 4 ). For practical purposes (taken into account during the creation of national, regional and local The consequences of the migrations of some lists of invasive species), apart from scientific synanthropic plants have proven to be very se- categories, “extrascientifie” criteria of plant ap- rious indeed, sińce the new arrivals have turned praisal" are often used (species that lead to spe- out to be extremely expansive and now domi- cific economic losses, harmful to human and/or nate over large areas occupied at the expense of animal health, etc.). native species (K o r n a ś 1996). The outcome of these processes may be considered in relation to As observed by E ster (1998), the vocabulary introduced into the ecological nomenclature is the following aspects (T o k a r s k a -G u z ik 2002, often without scientific meaning; it contains 2003b): a high dose of the emotional attitude of the author • Natural towards the phenomenon and the introduction of - impact on the biological diversity of the flora an aspect of evaluation into the common mean and fauna at all levels of organisation101 (e.g. ing of the word100. The author quotes also the B r o c k & F a r k a s 1997; I U C N 2000; M a c k et encyclopaedic definition of the term “invasive” al. 2000; S c h e r e r -L o r e n z e n et al. 2000; C r o n k from Webster’s New Encyclopedic Dictionary of & F u l l e r 2001; M c N e e l y et al. 2001; B a l o g h 1993: 2003; F o r m a n 2003; G e n o v e s i & S h in e 2004); Invasion means: “1. (...) entrance of an army - threat to protected areas (e.g. B a l o g h 1996; into a country for conquest; 2. (...) the entrance A d a m o w s k i & K e c z y ń s k i 1998; A d a m o w s k i et or spread of some usually harmful thing”; it al. 1998); means that the process has to do with aggression - changes in the landscape and land use (e.g. and destruction. D’A n t o n io & V i t o u s e k 1992; D’A n t o n io Invasiveness has been predicted on the basis 2000; H o b b s 2000 and literature cited therein) of the biological properties of a species, its eco • Social logical habitat conditions, its generał distribu- - detriment to public health (allergenic plants, tion and information on whether the species “be- stinging plants etc.) (e.g. C a m m et al. 1976; haves” as an invasive species in any region of W a d e et al. 1997); the globe. - creating difficulties or limitations for leisure; Starfinger (1998) and subsequently also F o r - - lowering aesthetic values m a n (2003) state that a possible indicator of later success as an invasive species may be its apo- • Economic phytism within its natural rangę (which is also - need for preparation of plans to combat the threat (e.g. C h il d et al. 1992, 2001; L u k e n & consistent with the chorological expansion model T h ie r e t 1997; C h il d & W a d e 1999, 2000; of J a c k o w ia k 1999; see Fig. 76) (Table 12). B im o v a et al. 2001; - costs of eradication/prevention (e.g. C h il d et 99 For ехатріе such categories of invasive species are al. 1998; C h il d & W a d e 2000; P im e n t a l 2002 given by CalEPPC (the governmental organisation in Cali- and literature cited therein). fornia responsible for monitoring and controlling invasive species): 1. most invasive wildland plants; 2. wildland pests For Poland, examples of species which pose of lesser invasiveness; 3. red alert plants (species with a threat with regard to the aspects listed have potential to spread explosively); 4. species for which more been given later in the present chapter. information is needed; 5. species being considered but not In comparison to other threats to biological lis te d (C a lifo rn ia Е х о т іс P est P lant C o u n cil 1 9 9 9 ). diversity, in most European countries including 100 To underline the scalę of the phenomenon of invasion many authors use sentences like this: “Arundo donax Poland invasive alien species have been given dramatically alters the ecological/successiona! processes in relatively little attention. The reason for this situ- riparian systems (...)” ( B e l l 1 9 9 7 ); “Lepidium latifolium ation is the fact that few countries in Europę have has rapidly spread (...) is an extremely competitive weed” ( Y o u n g et al. 1 9 9 7 ); “Lepidium latifolium (...) aggressively 101 The most significant biological threats include: the invading wetlands and riparian habitats” (Blank & Young replacement of floristically diversified indigenous commu- 1 9 9 7 ); “ Татагіх ramosissima is aggressive competitor (...) nities by monospecific phytocoenoses formed by popula- growing in monoculture stands (...) destroying wetlands tions of the alien species, the direct threat to the native flora and wildlife habitats” ( D u n c a n 1997). The examples men- and fauna leading to elimination of native species, changes tioned have been derived from a single volume devoted to in the habitat, modifications of geomorphological pro biological invasion ( B r o c k et al. 1 9 9 7 ). cesses, as well as generation of a fire hazard. Table 12. Apophytism of sample species described as invasive outside Poland (Europę)

H e m e S p e c ie s Described as invasive S o u rc e ro b y

Acer pseudoplatanus oemp Australia, New Zealand, Oceanie C ronk & F uller 2001 Islands, America S

Agropyron repens * .m ep A m e ric a N L uken & T hieret 1997 (= Elymus repens)

Alliaria petiolata* orne. A m e ric a N L uken & T h ieret 1997

Ammophila arenaria o m e. Australia, New Zealand, some regions L uken & T h ieret 19 9 7 ; C ronk & F uller 2001 of America N (California)

Anthoxanthum odoratum" .mep Chile, Hawaii C ronk & F u ller 2001

Bromus inermis* .mep America N L uken & T h ieret 1997

Bromus tectorum mep America N, Asia E (Japan), Oceanie M ack 1 9 81; L uken & T h ieret 19 9 7 ; C ronk Islands (Tenerife) & F uller 2001

Calluna vulgaris 0 ... New Zealand C ronk & F uller 2001

Carduus nutans .me. Canada, New Zealand C ro nk. & F uller 2001

Cirsium arvense* .mep America N C a lifo rn ia E x o tic P est P lant C o u n cil. 1999

Crataegus monogyna o m e p Australia, New Zealand, America N C ronk & F u ll er 2 0 0 1 ; C a lifo rn ia E x o tic P est P lant C o u n c il. 1999

Cytisus scoparius .m e. Australia, New Zealand, Africa S, C ronk & F uller 2001 Asia (India)

Dactylis glomerata* .m ep H a w a ii C ronk & F uller 2001

Euphorbia esula .mep America N L uken & T h ie re t 1997

Hieracium pilosella .mep Australia, New Zealand S co tt et al. 1 9 90; C ronk & F u ller 2001

Holcus lanatus* .me. Hawaii, New Zealand, America N C ronk & F u ller 2 0 0 1 ; C a lifo rn ia E x o tic P est P lant C o u n c il . 1999

Hypericum perfoliatum* .m e. A m e ric a N C a lifo rn ia E x o tic P est P lant C o u n c il . 1999

Linaria vulgaris* .mep America N L uken & T h ieret 1997

Lythrum salicaria* ome. America N M ałecki et al. 1993; L u ken & T h ieret 1997; C ronk & F u ller 2001

Myriophyllum spicatum o m .. USA C ronk & F u ller 2001

Ranunculus ficaria o m ep A m e ric a N L u ken & T h ieret 1997

Rhamnus cathartica om.. America N L uken & T h ieret 1997

Salix fragilis o m ep New Zealand C ronk & F u ller 2001

Degree of hemeroby: a - ahemerobic, o - oligohemerobic, m - mezohemerobic, e - euhemerobic, p - polyhemerobic, meta - metahemerobic

Scalę of hemeroby for Poznań city after J a c k o w ia k 1993, 1998a & c

* species listed as apophytes appearing in Poland in ruderal and segetal communities (Z a j ą c M. & Z a j ą c A. 1992). had negative experiences with alien species on for the efficient solution of problems posed by a scalę comparable with Australia or USA. Social these species. There is a lack of dedicated laws consciousness of the problems posed by alien with a complex approach to the problem, encom- species is surprisingly Iow in Europę (S o l a r z passing all habitats (terrestrial ecosystems, fresh 2001). Only in recent years have these problems waters and marinę waters), all organisms (plants, been addressed with regard to the whole continent game animals, fish, microorganisms, GMO) and (research programmes, seminars and scientific all branches of the economy (agriculture, marinę conferences; see Chapter 2), and last year has seen and inland fisheries, game hunting, naturę protec the publication of the European strategy on inva- tion). In some countries, no fuli lists of species sive alien species (G e n o v e s i & S h in e 2004). considered to be alien are available102. Also Legał platforms concerning protection against introduction, control and/or combating already- 102 In recent years, actions have been taken for this introduced alien species have hitherto been pre- purpose, ending with the preparation and publication of pared mainly in those parts of the world where lists of alien species, including invasive species, e.g. E s s l the various threats posed by these species were & R a b i t s c h 2002; B o t o n d & B o t t a -D u k A t 2004; also in most conspicuous, i.e. for example in USA, Poland, a research project entitled Alien invasive species in the flora and fauna of Poland in the context of conser- (T o k a r s k a - Canada, Australia and New Zealand vation of biological diversity is in the completion stage by G u z ik 2002, 2003b). The legislation on invasive group of botanists and zoologists and its results should be alien species in European countries is insufficient the publication of the Invasive species data book. regulations are lacking conceming the population synthetic studies regarding individual regions, control and elimination of alien species which which may form a basis for practical action. threaten biodiversity (S o la rz 2001). Legał reg Therefore, lists of invasive species have been ulations may play a significant role at various compiled for many countries and regions. The stages of the process of invasion for a given tentative list of invasive kenophytes occurring in species by preventing or limiting its introduction Poland has been prepared on the basis of the and later on by controlling its spread. following criteria: • the dynamie tendencies of analysed species in Invasions of alien plants are considered to be seąuential time periods (50 years) (i.e. abun- one of the major threats to biological diversity on dance, dominance and expansion rate, also a global scalę, next to the fragmentation and deg- ability to establish in different types of com radation of natural habitats103. In the International munities) - the objective criterion; Convention on Biological Diversity signed in • effects caused in the natural environment, 1992 in Rio de Janeiro, a special stipulation was economy and public health - the subjective cri included exhorting signatory countries to combat terion. alien invasive species which are a threat to native Analysis was performed on 300 species con habitats, communities or species (Art. 8 pt. h). sidered by the author to be recent synanthropic Methods of combating invasive species employed arrivals naturalised in Poland (or merely keno in many regions of the world (in Europę mainly phytes). Eventually, a finał list of 54 invasive in Great Britain) include the following means: species has been selected (including 4 potentially - mechanical - manuał removal, cutting, mow- invasive and 2 post-invasive species). In this ing, rooting out with the use of various eąuip- group, 14 species are limited in their occurrence ment, buming out, usage of screens; to anthropogenous habitats, while others also - chemical - spraying, use of applicator probes; enter semi-natural and natural habitats (Table 13). - biological - grazing, herbivores, pathogens; The invasive species listed for Poland belong to - mixed. 22 families (including 13 families represented by a The accumulated ехрегіепсе related to the single species), with the most amply represented preparation and validation of individual proce- families being: Asteraceae (17 species), Fabaceae, dures, determining the relative effectiveness of Polygonaceae and Scrophulariaceae (4 species separate methods as well as their costs, has been each), Brassicaceae and Poaceae (3 each) as well presented in numerous publications (e.g. R oom as Balsaminaceae, Cucurbitaceae and Apiaceae (2). 1981; S c o tt et al. 1990; H o ld en et al. 1992; The species belonging to this group represent M ałecki et al. 1993; L u k en & T h ieret 1997 and various life forms, with the same share of peren- the literature cited threin; C h ild et al. 1998; nial and annual species (20 and 17 species respec- C h ild & W a d e , 2000; C ro n k & F u l l e r 2001; tively); trees and geophytes are also represented B im o y A et al. 2001; C hild et al. 2001). by the similar number of species (8 and 9 each); Elodea canadensis is the only hydrophyte. The majority of species reproduce generatively (31), while the remaining ones usually take advan- 12.3. Invasive kenophytes in Poland tage of both manners of reproduction (17) and only very few reproduce only vegetatively (1) or with A significant contribution to our knowledge on a predominance of this manner of reproduction (6). invasions is brought by lists of alien species and The plants listed spread their seeds mainly using wind and animals (with a predominance of exochory) with a significant role played by 103 After the problem of invasions had been noticed, methods and directives conceming the elimination of in- myrmecochory and autochory; they also use vasive plants followed. Already at the end of the 19th cen- water as a mode of dispersal and they often take tury one could leam from the work of S em po łow sk i ( 1 8 8 0 - advantage of human assistance. 1881) on Xanthium spinosum that: “It is recommended to A definite majority of this group are effective think in due time about its eradication, before it makes competitor plants (C type strategy - 23 species); itself excessively at home in our fields. A radical mode of a relatively large group is formed also by species with action is plucking or mowing the plant before it is able to produce seeds. An incentive for the extermination of this a mixed strategy of the CR type (11 species) and weed for every landowner should be provided by the of the CRS type (3 species). The only species from deterrent ехатріе showing the extraordinary spread of among 5 species with an R type strategy which has some weeds and parasites such as e.g. the Spring Ground- had a spectacular success in the course of its inva- sel (Senecio vernalis W. et K.) or the dodders (Cuscuta), sion is Chamomilla suaevolens, while two others: against which in some countries the govemment was forced to start a struggle with police decrees about the obligatory Anthoxanthum aristatum and Eragrostis minor limit eradication of these weeds”. the scope of their invasion to very specific habitats Threat

Numbers of loc. Numbers Species Origin Dyn Habitats Scalę Category Invasive elsewhere up to of sq. 2000 landscape leisure health biological biological diversity protected areas economy public

• • •• Acer negundo L. Am N 3526 1379 4(+2) NSH [riparian; Inter-Regional T Eur C; Lithuania urban; abandoned fields] • Ailanthus altissima (Mili.) Swingle Asia E 31 29 2(+l) H [urban] Local pot. inv. Eur C & S; Am N [= A. glandulosa Desf.]

• Amaranthus retroflexus L. Am N & С 7651 2379 5 (+2) H [fields; National W Eur C urban; wasteland]

•• Ambrosia artemisiifolia L. Am N 101 61 2—3(+/-) H [wasteland] Regional pot. inv some regions of Eur; Am N

• Anthoxanthum aristatum Boiss. Eur S 1031 577 3—4(+2) H [fields] Sub-Regional W [= A. puelii Lecoą & Lamotte] ••• Aster lanceolatus Willd. Am N n.c.d 260* ? SH [riparian] Sub-Regional ? T Czech Rep. & Hungary

Aster novi-belgii L. Am N n.c.d 353* ? SH [riparian] Sub-Regional ? T Czech Rep. ••• •• Aster salignus Willd. Am N n.c.d 139* ? SH [riparian] Sub-Regional ? T Czech Rep. & Hungary • • Bidens frondosa L. Am N 3142 1068 4(+2) NSH [riparian; Sub-Regional T Eur C [= B. melanocarpus Wiegand] wasteland] • Bromus carinatus Hook & Am. AmN 1130 404* 3—4(+2) SH Sub-Regional T/W [urban; maedows] • Bryonia alba L. Eur E & Asia W 1328 728 3 -K + l) NSH [riparian] Sub-Regional pot.inv. Czech Rep. •• Bunias orientalis L. Eur SE & Asia W 1353 567 3—4(+2) SH [road banks; Sub-Regional T Czech Rep. & Slovac Rep. grassland]

• • Cardaria draba (L.) Desv. Eur SE 1048 576 3—4(+2) SH [road banks; Sub-Regional T Czech Rep. [= Lepidium draba L.] & Asia SW grassland]

• Chamomilla suareolens (Pursh) Rydb. Am N & Asia E 13125 2965 5(+2) H [urban; fields] National W Eur C [= Matricaria discoidea DC.]

••• Conyza canadensis (L.) Cronąuist AmN 11601 2929 5 (+2) H[urban; National W Eur C [= Erigeron canadensis L.] grassland; fields]

••• Digitalis purpurea L. Eur W 341 169 3(+ l) NSH [forests] Regional T Czech Rep.

••• Echinocystis lobata (F. Michx.) Am N 2047 708 3—4(+2) NSH [riparian; Sub-Regional T Czech Rep. & Slovac Rep.; Torr. & A. Gray wasteland] Hungary Threat

Numbers of loc. Numbers Species Origin Dyn Habitats Scalę Category Invasive elsewhere up to of sq. 2000 landscape leisure biological diversity economy public health protected protected areas

• • ••• Elodea canadensis Michx. A m N 3681 1847 4(+l) NSH [water] Sub-Regional T Eur C

Elsholtzia ciliata (Thunb.) Hyl. Asia E 1352 814 3—4(+/-) H[urban] Sub-Regional p-inv. [ = E. patrini (Lepech.) Garcke]

• • Epilobium ciliatum Raf. A m N 1224 470 3-4(+D NSH [forests; Sub-Regional T Czech Rep. [= E. adenocaulon Hausskn.] wasteland]

Eragrostis minor Host Eur SE & Asia W 1041 581 3-4(+2) H[urban] Sub-Regional NotH Eur C •• Erechtites hieracifolia (L.) Raf. ex DC. Am N & S 124 73 2-3(+ l) NSH [forests] Regional T Hungary

••• Erigeron annuus (L.) Pers. A m N 3557 1133 4(+2) SH [grassland] Sub-Regional T Hungary •• Fraxinus pennsyhanica Marshall A m N n.c.d 179 3(+2) SH [abandoned Regional T Czech Rep. & Hungary fields]

• Galinsoga ciliata (Raf.) S. F. Blake Am C [m] 6777 2021 4-5(+2) H [fields] National W Eur C [ = G. ąuadriradiata Ruiz & Pav.] • Galinsoga parviflora Cav. Am S & C [m] 10932 2726 5(+2) H [fields] National W Eur C

•• • Helianthus tuberosus L. A m N 1416 778 3—4(+2) NSH [riparian; Sub-Regional T some regions of Eur C wasteland]

• • •• Heracleum mantegazzianum Asia C & E 100 74146* 2-3(+2) NSH [riparian; Regional T Eur W, C & N Sommier & Levier road banks; abandoned fields]

• •• Heracleum sosnovskyi Manden. Asia SW [Cauc.] 96 72146* 2(+2) NSH [riparian; Regional T Hungary, Lithuania road banks; abandoned fields]

• •• Impatiens glandulifera Royle Asia C [Himal.] 1574 675 3^1 (+2) NSH [riparian] Sub-Regional T Eur W & C [= I. roylei Walp.] ••• Impatiens parviflora DC. Asia C & E 6730 1681 4-5(+2) NSH [forests] National T Eur C Eur S (warm regions) • Iva xanthiifolia Nutt. A m N 294 150 з (■+■/-) H [wasteland] Regional pot.inv. •• Juncus tenuis Willd. [= J. macer A. Gray] Am N 5332 1440 4—5(+l) SH [meadows] National T Czech Rep.

•• Lupinus polyphyllus Lindl. A m N 2674 1387 4(+l) NSH Sub-Regional T Czech Rep.; Lithuania [forests; grassland] •• Lycium barbarum L. [= L. halimifolium Mili.] Asia E & Eur SE 2634 1224 4(+l) NSH [serub] Sub-Regional T Czech Rep. •• Mimulus guttatus DC. Am N 326 128 3(+2) NS [riparian] Regional T

• Oxalis fontana Bunge [= O. stricta L.] Am N 8806 2141 5(+l) H [gardens] National W

• ••• Padus serotina (Ehrh.) Borkh. Am N & S 2564 1134 4(+2) NS [forests] Sub-Regional T Eur C [= Prunus serotina Ehrh.]

•• Parthenocissus inserta (A. Kem.) Fritsch A m N 558 332 3(+2) NSH [riparian] Regional T some part of Eur C [= P. vitacea (Knerr) Hitchc.]

•••• Quercus rubra L. Am N n.c.d 554* 3-4(+2) N [forests] Sub-Regional T Czech Rep.

Reynoutria x bohemica Chrtek & Chrtkova Anthropog. n.c.d n.c.d. ?(+2) NSH [riparian; Regional ? T Czech Rep. [= R. japonica Houtt. x R. sachalinensis urban] /Sub-Regional ? (F. Schmidt) Nakai]

Reynoutria japonica (Houtt.) Ronse Decraene Asia E 3004 1158* 4(+2) NSH [riparian; Sub-Regional T Eur W & C; Am N var. japonica [ = Fallopia japonica Houtt.] urban]

Reynoutria sachalinensis (F. Schmidt) Nakai Asia E 474 282* 3(+l) NSH [riparian] Regional T Czech Rep. [= Fallopia sachalinesis (F. Schmidt et Maxim) Ronse Decraene]

• ••• Robinia pseudoacacia L. AmN 7067 1957 4-5 (+2) NSH [grassland; National T some regions of Eur scrub & forests] C, Lithuania

•• • Rudbeckia laciniata L. Am N 2251 903 3—4(+2) NSH [riparian; Sub-Regional T Czech Rep. & Slovak Rep. meadows]

• Rumex confertus Willd. Eur SE & Asia W 1731 673 3— 4(+2) SH [riparian; Sub-Regional T Lithuania railway & road banks]

Sisymbrium loeselii L. Eur SE & Asia С 2326 976 3-4(+ l) H [wasteland] Sub-Regional NotH Czech Rep.

Solidago canadensis L. A m N 3434 1254 4(+2) NSH [riparian; Sub-Regional T some regions of Eur C abandoned fields]

Solidago gigantea Aiton. Am N 5348 1668 4-5(+2) NSH [riparian; National T some regions of Eur C [= S. serotina Aiton] abandoned fields]

•• •• Solidago graminifolia (L.) Elliott Am N 46 27 2(+l) NSH [meadows] Local T •

• Trifolium patens Schreb. Eur S 227 54 2-3 (+1) NH [meadows] Regional T •

• Ve ronić a fili fo r mis Sm. Asia SW [Cauc.] 161 69 2-3(+l) SH [meadows] Regional p-inv. Czech Rep.; USA Veronica persica Poir. Asia SW [Cauc.] 7887 2204 5 (+2) H [fields] National W Czech Rep. •

Vicia grandiflora Scop. Eur S & Asia SW 1540 506 3-4(+2) S A [grassland; Sub-Regional T/W • fields]

Species: Latin name and synonym(s); species are arranged alphabetically; species names nomenclature according to MIREK et al. 2002; Origin: E ur- Europę, Asia; Am N - North America; Am S - South America; С - central, E - east, N - north, S - south, W - west; No of loc: number of localities; No of sq: number of ATPOL squares (total number of squares for Poland: 3646); * - indicates that number of squares recorded need to be verified; n.c.d. - not complete data; Dyn - frequency and dynamie tendencies according to ZARZYCKI et al. 2002: Frequency in the wild at the territory of the country: 1 - very Iow number of localities (1-20); 2 - Iow number of localities (up to 100); 3 - high number of localities, but with narrower distribution (in one or two regions of the country); 4 - high number of localities in many regions; 5 - common (abundant) in the whole territory; Dynamie tendency (in brackets): (-2) - high decrease of(in) number of localities; (-1) - decrease in number of(in) localities or decrease in abundance over existing localities; (+1) - inerease of(in) number of localities, inerease in abundance over existing localities; (+2) - high inerease of localities (colonizing new localities); (-/+) - disappearing of some localities and appearing of new localities; ? - undefined dynamie tendencies; Habitats: type of habitats invaded: N - natural; S - seminatural; H - human-made (anthropogenic); [in brackets] impacted ecotopes; Category: according to the classification by PySek et al. 2004: T - transformer; W - weed; NotH - not harmful. and also specific geographical regions in Poland (cf. to occupy new habitats (found more and more Chapter 8); the remaining one: Elsholtzia ciliata has often in protected areas) and it reąuires monitor the status of post-invasive species. ing, in special cases even qualifying for active Most of current invasive kenophytes have elimination (A d a m o w s k i & K e c z y ń s k i 1998), been introduced to Poland intentionally. They Elodea canadensis is a kenophyte which may also are twiee more numerous (36 species) than the have entered the existing ecosystems permanently, casually introduced species. The largest group but which is, however, past the end of its period is formed by species originating from both of rapid spread. Americas (with a predominance of North Amer Most invasive kenophytes are species which ican species - 29), from Asia (9) as well as from pose a threat on the regional scalę (usually in Southern Europę and western Asia (9), so they one or several regions in the country, e.g. An- are mainly arrivals from geographically remote thoxanthum aristatum (cf. also Chapter 7), Bro- areas. The greater part of the species is consid- mus carinatus, Echinocystis lobata, Fraxinus ered invasive also in other regions of Europę or pennsylvanica, Heracleum mantegazzianum) even on the global scalę. (Table 13). Special attention should be paid to those keno phytes which are characterised by high competitive capabilities and which can penetrate into 12.4. Threatened regions and habitats semi-natural and natural communities (T o k a r - s k a -G u z ik 2003a). Eleven (11) species have been deemed to be Invasive kenophytes widespread in forests, es- invasive on the national scalę (Table 13). They pecially in the Southern part of Poland, include are mostly kenophytes linked to anthropogenous the arboreal species Padus serotina and Quercus habitats included in the category of weeds and rubra as well as the herbaceous species Lupinus thus constituting a threat to agricultural areas104. polyphyllus (the spreading of this species is The species which are most often mentioned helped by a continuous supply of diaspores gen- in this context include Amaranthus retroflexus erated by additional sowing) and Digitalis pur- and the two species of genus Galinsoga which are purea (a species which may be considered inva- considered to be troublesome weeds of root crop sive on a local scalę, especially in some regions fields (W n u k 1996; R o l a & R o l a 2002). This of the Carpathian Mountains). group, but considered on a regional scalę, also Species which pose a threat to meadows and includes Anthoxanthum aristatum (K u ź n ie w s k i other grasslands include Solidago canadensis and 1996; W archolińska & S ic iń s k i 1996). Solidago serotina (these species also massively A similar scalę of distribution in Poland is encroach on riverside habitats and on fallow shown by Elodea canadensis and Impatiens fields), species from the genera Aster and Hera parviflora (the latter of which, however, is still cleum, additionally Rumex confertus, Bunias a rarely encountered species in north-eastern orientalis, Bromus carinatus and locally Vero- Poland). These species may justly be considered nica filiformis (which is probably receding in as “dangerous” from the point of view of threat recent times). to the native flora and vegetation. While Impa “Particular habitats such as watersides are the tiens parviflora05 is a species which is still able most endangered ones and are most easily invaded by alien invasive plants and then play a 104 The aceepted prineiple states that dangerous weeds role as a transmitter into other habitats such as are the ones which reach phytosociological constancy lev- scrub and woodland” (T o k a r s k a -G u z i k 2003c). els of IV or V and a high value of the coverage coefficient, Invasive kenophytes which seize this kind of while the species which reach constancy level IV or V habitats, often on a massive scalę, include the buthave a smaller coverage coefficient may be potentially dangerous to agriculture (W n u k et al. 1989). The author species already mentioned from the genus So does not, however, list any kenophytes among “dangerous” lidago, as well as Reynoutria, Impatiens glan- species - although they show up in the releves, they do not dulifera, Rudbeckia laciniata and Heracleum fulfil the standards. Apophytes and archeophytes predom- mantegazzianum. inate in segetal communities - kenophytes have a smaller share. A separate problem is the proportion of inva- 105 In favourable conditions this species may obtain an invasive success while not being a very good competitor. Its sive species in protected areas and the conseąuences success is due according to T r e p l (1984) to its shallow root brought about by their presence. Analysis of sev- system. Contrary to native herbaceous species, the Smali eral dozen publications related to national parks, Balsam can avoid competition with root systems of trees. naturę reserves and other forms of protection has Thus, the species has discovered an unexplored niche (in the sense of unused resources) and is able to colonise it. shown that in many situations of that type, there are no alien species at all106. They mostly encom- However, in many regions of the country due pass areas which were designated to protect either to an insufflcient level of knowledge about synan- multi-specific deciduous forests with a relatively thropic vegetation (even the total number of alien high degree of naturalness or peat-bogs (e.g. C z a r species in the flora is unknown) it is still not n eck a 1978; Łuczycka-Popiel 1989; B r z e g et al. possible to consider the available data as finał or 1995; Sokołowski 1995b, 1996a & b, 1997a; to reliably estimate the degree of invasion of alien Obidziński et al. 1998). The relatively scarce species (Jutrzenka-Trzebiatowski et al. 2002). oceurrence of invasive species in protected areas A generał rule applies that in areas with a larger created in mountainous areas (especially in some extent and thus with a more complicated mosaic regions of the Carpathians) is due to climatic fac- of habitats, the number of alien species is higher. tors which limit their spread (see Chapters 6 and One of the species most often mentioned is 10). In many cases kenophytes are mentioned Impatiens parviflora (e.g. Ćwikliński 1972b; there as sporadic in occurrence with a Iow abun- M ic h a lik 1972; Krawiecowa 1972; S o k o ło w s k i dance (e.g. Krawiecowa 1972; Celiński & Wika 1997b; Adamowski & Keczyński 1998; P isk o r z 1978). & K lim k o 2001). This plant is starting to appear Detailed studies devoted to anthropogenous in massive amounts in those regions in Poland transformations of the flora and vegetation have where it has hitherto been a rare element of the been carried out in the Ojców National Park flora (cf. Chapters 7 and 8), e.g. in mixed forests (M ich a lik 1972, 1974). They have shown the pres- along the Southern edge of Wigry lake (in the ence of ca. 35 species of alien origin, casually Wigry National Park) (Jutrzenka-Trzebiatowski introduced (e.g. Conyza canadensis, Galinsoga et al. 2002). ciliata, G. parviflora, Impatiens parviflora, Bunias Plants which threaten protected areas also in- orientalis, Geranium pyrenaicum and others) as clude species from the genus Solidago which in- well as intentionally planted by humans (e.g. Quer- filtrate meadows and grasslands (Ćwikliński cus rubra, Helianthus tuberosus, Rudbeckia laci- 1972b; see also Chapter 11). A major threat to niata, some species from the genus Aster). protected areas is the migration of synanthropic Similar results were obtained from studies on species along field tracks (Świerkosz 1995) and synanthropisation in the Pieniny National Park tourist trails. (G u zik ow a 1972). This area, in the 1960s char- In Polish national parks, the mąjority of synan acterised by a lower share of anthropophytes in thropic changes occur currently under the pre- comparison with the adjacent territories of Gorce dominant influence of tourism and to a lesser Mts. and Sądecczyzna (Nowy Sącz province), has extent forest management; grazing, pasture and been “opened” for infiltration by recent arrivals meadow management experiments and activities following the building of new communication have also been of historie importance (P ię k o ś- pathways. Next to common kenophytes linked to Mirkowa & Mirek 1978; Mirek & Piękoś- ruderal habitats (mainly in villages and towns), M irk ow a 1987). In this publication, the authors, which are widespread also in other regions of the who have monitored modiflcations in the naturę country, the author of the study lists also species of the Tatra National Park for many years, ex- which enter deep into natural communities (e.g. press their opinion that the most visible effects Impatiens parviflora into riverside willow com are changes in the horizontal and vertical distri- munities on the Dunajec and Krośnica rivers and bution of species. These migrations are made into forest communities Alnetum incanae, Fage- possible by humans who both transport the dia- tum carpaticum, Phyllitido-Aceretum and Cari- spores and create suitable habitats where these ci-Fagetum, as well as Juncus tenuis which species can spread. Sites of occurrence of synan spreads along paths over wet ground). Later stud thropic species in the Tatra mountains include ies in the same area have confirmed further roadsides, roadside ditches, parking lots and their spreading of the aforementioned species and they surroundings, tracks and tracksides of railways, have also turned attention to the rapid spread of clearings cut under tracks of cable cars and chair- lifts, tourist trails, forest glades and mountain species which have escaped from cultivation, meadows. The highest number of synanthropic including: Heracleum sosnovskii, Helianthus species may be found around mountain shelters, tuberosus, Reynoutria japonica, Solidago ca ski-lift stations, chalets and similar sites. nadensis and S. gigantea (Z a rz y ck i 1982, 2000b). Similar relationships have been found by Ros- ta ń sk i (1977, 1978) in the area of the Karkonosze 106 Authors do not however always give a complete flora, National Park. Fabiszewski (1985) summarised while the published phytosociological releves are usually taken on the most typically formed plots. Occasionally, this the threats to naturę in this Park linked to the in- “ruse” of authors appears to be intentional, with the pos- fluences of industry and mass tourism and iden- sible goal to form an argument during legał procedures. tified synanthropic species including alien species (such as Lysimachia punctata and Mimulus gut- families Amaranthaceae, Cyperaceae, Poaceae tatus), which migrate up to the highest peaks in which have hitherto not been considered highly connection with tourist traffic, urbanisation, invasive (as opposed to large families, such as: wastewater and trash littering. Asteraceae, Rosaceae and Fabaceae). The present Despite having a largely preserved natural study has confirmed these results in part. character, the Białowieża Primaeval Forest is also When forecasting the further influx of poten an area where alien species are recorded. Apart tially invasive species into Poland, it is necessary from the already mentioned Impatiens parviflo- to gather information on the behaviour of each ra, cases of naturalisation of cultivated species, species in other regions of Europę. By way of especially trees, are becoming more and more ехатріе, in the Czech Republic and Slovakia (as freąuent. Łuczaj & Adamowski (1991) list among well as in other warm regions of the continent) the most often recorded species: Acer negundo, Ambrosia artemisiifolia is an invasive species Quercus rubra and Cotoneaster lucidus. The which spreads massively along roads, railways latter species in the opinion of the cited authors and in arabie fields. Also, Senecio inaequidensx07 may in the futurę become a permanent compo- (already recorded on first sites of occurrence on nent of a fringe scrub community from the Pru- railway grounds - Guzik J., Pasierbiński A. & netalia order. The degree of encroachment of Rostański A., pers. comm.) and Dittrichia grave- alien tree species in forest communities in the olens may migrate to Poland from the territory Białowieża Primaeval Forest is still however sig- of Germany, as it has happened many times in the nificantly smaller than in analogous communities past for other plants (Radkowitsch 2003). in the western part of Poland. Cynodon dactylon is one of the most common apophytes in the cities of Southern Europę (Celesti Grapow & Błasi 1998), while it has only an ephemerophyte status in Poland; a simi- 12.5. Forecasting invasions: potentially lar situation concerns Eleusine indica which is invasive species naturalised in the Mediterranean basin (U r b isz & U r b isz 2003). Such species as mentioned above can be considered as potentially able to become A reconstruction of the ways and manners of established or even invasive in the futurę. expansion of a species in the past may help in understanding its invasive success and in fore casting further stages of its migration. Of eąual importance in forecasting invasions is autecolo- 12.6. Finał remarks gical research, especially regarding the life strategy, means of reproduction and dispersal and conditions of seed germination of potentially Although studies of alien species in the flora invasive species (W a d e 1997). as well as about the broadly understood process Moreover, useful data in forecasting invasions of synanthropisation of the plant cover have been include not only ecological factors, such as tem- conducted in Poland for a long time (T o k a r sk a - perature, habitat conditions and disturbances, but -G u zik 200la), there is still insufficient multi-as- also information as to whether the species is pect research on alien invasive species, especially invasive in another part of the globe. in the context of the threat they pose to the in- Possibilities and limitations in forecasting fur digenous naturę. In order to limit the invasions ther exchanges of species between various regions of undesirable alien species it is important to know in detail their ecology and distribution of the world have been analysed by Ja ck o w ia k (C h ild et al. 2001). (1999) (earlier also by J a g e r 1988 and Sukopp 1995; see also Chapter 8) on the ехатріе of plants Taking into account the huge variability in definitions of plant invasion and in the evalua- from the family Asteraceae. The results of this tion of the invasive potential of plant species, analysis lead to the conclusion that the exchange Kowarik & Schepker (1998) point to the need for of the flora has not yet been completed. the preparation of speeific case documentation F o rm a n (2003) has published the so-called detailing the impact of invasions on the local Warning list o f species basing upon the previously plant cover and the possible threat they may mentioned relation between the apophytism cause, as well as estimations of the social, eco- (“weediness”) of a species in its homeland and nomic and ecological effects of the invasions. the probability of its invasion into a newly occupied area (compare also Table 12 in Chapter 107 Starting to appear in the Black Country (Central 12.1). The results of her analyses reveal higher England), probably introduced with ornamental plants than expected potential invasive characteristics in imported from Holland (Prof. I.C. Trueman, pers. comm.). A significant element which may lead to lim- able conditions and ąuickly turn into the most iting the spread of invasive plants may be the dis- common weeds. Some synanthropic plants are semination of relevant information. so common and we have grown so used to them “An intensive development of commerce and that we consider them nearly as a part of the the concurrent development of communication local element” (P a c z o s k i 1900). These words routes exert a powerful influence on the spread have not lost their significance in the age of of various plants, often even from very distant globalisation of commerce, the dynamie deve- places. Of course, not all of these introduced lopment of tourism, and in Poland also the cur- plants remain in the locality to which they were rent intensification of the development of re- brought. Some of them, however, find favour- sidential building activity.

PART FIYE

Summary, conclusions and the perspectives for studies of plants of alien origin in Poland against the trends preyailing in Europę and the world

13. Summary and conclusions ing the formation of their ranges (Chapter 10). Many distribution maps have been augmented (Chapter 7) and five new maps have been devel- The objective of this monograph was to sum- oped (Fig. 39 in Chapter 7, and Appendix C). marise the research carried out on the develop- Another reconstruction effort had the aim at ment of the flora of kenophytes within the ter- finding changes in the ranges of kenophytes, with ritory of Poland and to arrive at a synthesis of the the elucidation of possible migration routes relevant knowledge available to date. (Chapter 9). The dynamie trends among keno The intention of the author was also to describe phytes have also been discussed vis-a-vis the the history and directions of studies concerning factors helping them acąuire various types of the newest synanthropic newcomers established habitats (Chapter 11). From the list of keno in Poland, and to provide references to the most phytes, invasive species have been identified important studies and special topics undertaken (a list of invasive kenophytes for Poland has been by Polish botanists, whose work constitutes proposed), opening wider discussion on the cri- a permanent contribution to the achievements of teria adopted for their selection, and indicating biogeographic sciences (Chapter 2; Table 1). those regions of Poland threatened by invasion The result of this attempt is a new list of this (Chapter 12). group of species, considerably broader than that In opinion of the author, the most important which could be found in earlier works and aug- conclusions of this study area are as follows: mented by the inclusion of the ecological and geographical characteristics of the species (Ap- • In the ever-progressing process of the synan- pendices A and B, and Chapters 5.1 and 8). Re- thropisation of vegetation, viewed in the time searching historical sources (“old” floras, herbar- frame of the last flve centuries, the role of newer ium documentation) has allowed the verification arrivals (kenophytes) has been growing. The or determination of the first floristic records of transformation of the composition of the flora particular species of Polish kenophytes (Apendi- occurs at the level of taxonomic, geographical/ ces A and B; also Chapters 5.2, 7 and 9). An historical, biological and ecological structures attempt was also made to reconstruct the periods (even the genetic structure) and its course is where the influx and spread of kenophytes were realised in time and space. most intense, relating these to historical and The kenophytes occurring in Poland originate geographical factors (Chapters 5.2 and 9). from five continents, with a predominance of For a selected group of 25 species the history species from the various European regions (chiefly of their spread in Poland has been reconstructed from its Southern and south-eastern parts) and in detail (Chapter 7). Detailed data on the distri- from North America. Among them, hemierypto- bution of 174 species of kenophytes has been phytes and therophytes predominate. The species used to represent the typology of their ranges intentionally introduced by humans show a ten- within Poland’s borders (Chapter 6), augmented dency to colonise natural and semi-natural habitats, by a discussion on the principal factors influenc- whereas those species introduced accidentally, colonise the anthropogenic habitats before any distribution ranges. Climatic conditions should be others (only in the subseąuent stages of their regarded as the main factors affecting the pattern expansion, do some of them also colonise natural of the rangę while natural conditions and local and semi-natural habitats). anthropogenic factors are the second and third The kenophytes occurring in Poland are mostly most important factors, respectively. insect- and wind-pollinated plants, reproducing At present, a dozen or so species have their by generative means; some of them also imple- eastern limits of distribution within the Polish ment various methods of vegetative reproduction. lands, while several other species reach their Anemochory and zoochory play predominating western and northem limits there. Species closely roles in the expansion of this group of anthropo- linked to rivers, particularly the groups of keno phytes. Among the kenophytes, species of high phytes characteristic of the major rivers of Poland: competitive potential prevail (those with C-type the Vistula, Odra and Bug (Eragrostis albensis, life strategy), together with those adapted to Oenothera depressa, Oenothera x hoelscheri, circumstances where the effect of stress is Iow Rumex confertus, Salsola kali subsp. ruthenica and competition is limited by disturbances (C-R and Xanthium albinum) and the kenophytes as- type life strategy) and mobile pioneer species sociated with cities and railway links between {R type). them represent a specific and distinctive type of distribution. • The reconstruction of the historie floras of kenophytes has permitted the establishment of • The reconstructed courses of immigration and a list of the “oldest” arrivals among this group expansion of individual species have revealed the of alien species. The Polish flora of the 17th cen- factors affecting the rate of spread and direction tury undoubtedly included such species as: of expansion and, ultimately, the current pattern Acorus calamus, Datura stramonium, Echinops of their distribution rangę in Poland. These fac sphaerocephalus, Marrubium vulgare, Tanacetum tors include the manner and naturę (freąuency) parthenium and others. of their introduction. Those species accidentally introduced on many occasions, or those adopted • The historical and economic conditions in as cultivated plants (maintained as cultivars) in Poland exerted a significant impact on the aug- various regions, have spread faster and their mentation of local flora by newcomers. The ranges are larger. Most of the species concerned, migrations of kenophytes had certain culmina- even those which came primarily from very re- tions of influx (called “migration waves”). The mote geographical regions, spread in Poland as highest culmination occurred in the second half an effect of their previous intentional or casual of the 19th century, coupled with the “first indus- introduction into Western Europę (historically trialisation stage”. The culminations differed earlier industrialised and urbanised). from one another by their respective origins. From the beginning of the 16th century through • The rate of expansion shown by kenophytes in to the first half of the 19th century, species of their new homeland depends on their biological European and Asian origin had predominated. properties, historical circumstances (timing and The last 150 years showed a noticeable predom- manner of introduction), and a set of factors inance of species coming from the Americas. (natural and anthropogenic) collectively referred Finally, in the most recent period, the proportion to as the resistance of the environment. A rapid of species of hybrid origin, which owe their rate of expansion has been characteristic for these appearance to humans, either directly or indirectly, kenophytes repeatedly casually introduced to has inereased in the Polish flora. many regions of Poland, and colonising (at least in the initial phase) anthropogenic habitats. This • Kenophyte migrations have covered the whole statement holds for such species as the Chamo- present territory of Poland, and the maps of their milla suaveolens, Conyza canadensis, Galinsoga occurrence reveal the areas of their high concen- parviflora and G. eiliata. trations, namely the Vistula river valley, the Sile- sian Upland and major urban centres. • Among the major factors facilitating the migra Many kenophytes do not show any definite tions of alien species, the following factors can type of rangę. This reflects the history of their be “guaranteed” by humans: arrival into Poland, as well as their mechanisms - elimination of barriers (development of trans of establishment in the local flora (long-term cul- port over great distances, reduction in the size of tivation or multiple accidental introductions into forested areas and wetlands and their fragmenta- many regions). tion); Some of the species of this group can never- - introducing species into cultivation and main- theless be allocated to certain types of definite taining them in cultivation for a long period of time, which helps them to escape into the wild Also in need of more modern studies is the (cultivation involves significant numbers of dia- biology of individual species and their possible spores of great genetic variability being intro- changes (in the methods of pollination, pollina- duced simultaneously, and this process is repeated tors, disseminating methods of entire communi- many times); ties with participating kenophytes, associations - “creating” (often ąuite unintentionally) com- and relationships with other plant species, para- pletely new habitats in the wild which could be sitic and saprophytic fungi, and microorganisms). surrogate habitats for alien newcomers (stone These studies should also be pursued at the level fences and walls, railway tracks and embank- of the genotype. On the population level, studies ments, cracks in flagstones, or utterly artificial on morphological and genetic differentiation are sites such as heaps, new geomorphołogical forms needed along with studies of the evolutionary or areas completely deprived of vegetation); processes operating in the immigrant populations. - applying alien plants in the arrangement of Finally, there is a need to undertake studies on public green areas, as well as in the reclamation the impact of invasive plants on the functions and of degraded areas (this latter measure is some- structure of ecosystems (also involving long-term times implemented on a vast scalę); studies). To date, such studies have only been - developing railway and road transport (creat rarely undertaken in Poland. ing migration corridors); The issue of separating apophytes from anthro- - permanent or periodical interference in the pophytes (K o r n a ś 1981) is still open and await- habitat conditions and structure of native phyto- ing solution. The elucidation of the origin and coenoses (maintaining and extending large-sized status of individual species can still be helped by disturbed habitats, fallow lands, rivers and involving palaebotanical and archaeobotanical streams canalization); methods. - inappropriate forest management (the direct As stated by Welk et al. (2002) “(...) The re- introduction of alien species into forests). search on well-known, non-indigenous European species in North America, and vice versa, • Synthetical studies concerning the expansion of provides us with opportunities for long-term field species of alien origin will provide a theoretical tests because many of the species have had basis to develop checklists of invasive species enough time to reach even the remote parts of which, in turn, will help in planning practical their potential distribution ranges on their ‘new’ measures (prevention and control). continents. With a review of the results of investigations on a large number of species with different life history strategies, life forms and na- 14. Invasions of alien plant species tive rangę types, our understanding of the differ- at the dawn of the 21st century: ent capacities of climatic rangę models for predicting invasiveness could be improved”. perspectives for further studies The subject matter of public and scientific debates on the possible evaluation of hazards Those species of alien origin, particularly in- involving genetically modified plants and their vasive and potentially invasive species will con- release into the environment should also become tinue to attract the interest of taxonomists, eco- topics of futurę scientific studies. Hazards asso- logists, plant geographers, as well as many con- ciated with the gene flow from GM crops to servationists. wildlife species (crossing with close wild rela- The effective protection of biodiversity calls tions)"0 should also become a topie for more for modern taxonomic studies, particularly of studies; it is necessary to develop proper tools critical taxa. In the Polish flora these include the for the assessment of possible hazards (cf. e.g. genera Aster, Helianthus, and Rubusm and hybrid A b b o t 1992; A r n o ld 1997; P o h l-O r f et al. 1998; forms109 in these and other genera.

110 It is also essential to study and chart the distribution 108 Taxonomic studies on this species have already been of wild relatives and hybrids between cultivated plants and completed (Z i e l i ń s k i 2004) their wild relations. This knowledge would allow the as 109 Hybridisation has long-since been recognised as play- sessment of potential. possibilities of hybridisation between ing an important role in the evolution of plants (S t e b b in s GMP and their wild relations; the phenomenon of introgres 1950). In the recent decades the role of anthropogenic hybrid sion is particularly important in respect to the Brassica- isation has increased. Hybrids produced this way, being more ceae, Solanaceae, Poaceae families and some tree species, invasive, can sąueeze out or replace the parental species or e.g. Populus, Salix and Picea. Also needed is more tax- can produce a genetic mix. At present, such a process is even onomic knowledge about cultivated species, the regions of referred to as the “extinction of species by hybridisation and their cultivation, as well as of the distribution of their wild introgression” or “invasion by hybridisation”. relations and potential hybrids. Rieseberg & Carney 1998; den N ijss et al. 1999; biology (including molecular biology and cyto- A lle n d o r f et al. 2001). genetics). Special attention has already been Despite the multitude of studies undertaken in focused on the genetic aspects and methods of the currently developing branch of ecology called reproduction which support the invasiveness of ecology of invasion, many questions are still plants. awaiting answers. An evident and indispensable An additional activity to be coupled with stud tendency leads to precisely planned studies (in- ies should be data gathering (Global Network on cluding long-term projects), employing the modem Тахопоту - a network and data exchange systems), methods and tools of the various disciplines of the exchange and propagation of information. References

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Streszczenie

Tematyka niniejszej pracy mieści się w problema szczególnych gatunków polskich kenofitów (zał. tyce dotyczącej synantropizacji szaty roślinnej. Jednym A i B; także rozdz. 5.2, 7 i 9). Podjęto również pró z przejawów tego ukierunkowanego procesu przemian bę odtworzenia okresów kulminacji napływu i roz zachodzących w wyniku różnych form działalności przestrzeniania się kenofitów, z ukazaniem zależno człowieka na kuli ziemskiej są procesy wymierania ści od czynników historycznych i geograficznych jednych gatunków i rozprzestrzeniania się innych, na (rozdz. 5.2 i 9). silające się w ostatnich stuleciach i przyczyniające się Dla wyselekcjonowanej grupy 25 gatunków odtwo do zmian różnorodności biologicznej w skali regionów, rzono dzieje ich rozprzestrzeniania się na obszarze krajów i kontynentów. kraju (rozdz. 7). Na podstawie zebranych szczegóło Celem niniejszej monografii było ukazanie badań wych danych o rozmieszczeniu dla 174 gatunków nad kształtowaniem się flor nowszych przybyszów kenofitów przedstawiono typologię ich zasięgów synantropijnych zadomowionych na obszarze Polski w granicach Polski (rozdz. 6), a także zweryfikowa (kenofitów) oraz synteza dotychczasowej wiedzy no hipotezy odnoszące się do głównych czynników w tym zakresie. Moim zamysłem było także ukaza wpływających na ich kształtowanie się (rozdz. 10). nie historii i kierunków badań nad tą grupą roślin ob Mapy rozmieszczenia dla wielu gatunków zostały cego pochodzenia, wraz z przytoczeniem najistotniej uzupełnione (rozdz. 7); opracowano ponadto 5 no szych opracowań i zagadnień specjalnych podejmo wych map (rys. 39 w rozdz. 7 oraz zał. C). wanych przez polskich botaników, które na trwałe Dokonano próby rekonstrukcji historycznych wpisane zostały w dorobek nauk biogeograficznych zmian zasięgów kenofitów wraz ze wskazaniem (rozdz. 2; tab. 1). możliwych dróg ich migracji (rozdz. 9). Omówiono Wynikiem podjętych studiów jest opracowanie no ponadto tendencje dynamiczne kenofitów z uwzględ wego, uzupełnionego w stosunku do literatury, wy nieniem czynników sprzyjających opanowywaniu kazu dla tej grupy gatunków, poszerzonego o ich cha różnych typów siedlisk (rozdz. 11). Z listy kenofitów rakterystykę ekologiczno-geograficzną (załączniki wyłoniono tzw. gatunki inwazyjne (propozycja listy A i B oraz rozdz. 5.1 i 8). Dotarcie do źródeł histo inwazyjnych kenofitów dla kraju), jednocześnie ini rycznych (historyczne/„stare” flory, dokumentacja cjując dyskusję nad przyjętymi kryteriami ich selek zielnikowa) umożliwiło zweryfikowanie lub ustale cji, a także wskazano rejony kraju zagrożone inwa nie pierwszych dat florystycznych (znalezisk) dla po zją (rozdz. 12). Etablierung und Ausbreitung gebietsfremder Pflanzenarten (Kenophyten) der Flora Polens

Zusammenfassung

Das Thema der vorliegenden Arbeit gehort zur floristischen Daten (Fundę) fur einzelne Arten der Problematik der Synanthropisierung der Pflanzen- polnischen Kenophyten (Beilagen A, B; Kpt. 5.2, 7 decke, d.h. des Auftretens von wilden Pflanzenarten u. 9) festzulegen. Sie versuchte auch, die Kulmina- in sekundaren Biotopen, in denen die natiirliche Ur- tionsperioden fur Zustrom und Ausbreitung von flora von dem Menschen zerstort worden ist. Eins von Kenophyten wiederzugeben und dereń Abhangigkeit den Symptomen der gezielten Verwandlungen auf der von historischen und geographischen Faktoren zu Erde, die unter der Wirkung von verschiedenartigen zeigen (Kpt. 5.2 u. 9). Man hat die Geschichte der Formen der menschlichen Tatigkeit eintreten, ist das Ausbreitung auf dem polnischen Gebiet von 25 aus- Aussterben von einigen und das Ausbreiten von gewahlten Pflanzenarten wiedergegeben (Kpt. 7). anderen Pflanzenarten, die in den letzten zehn Jah- Anhand der gesammelten genauen Daten iiber die ren stark zugenommen haben und die zur Verande- Anordnung von 174 Kenophytenarten wurde die rung der biologischen Vielfaltigkeit in den Regionen, Typologie ihrer Reichweiten in Polen (Kpt. 6) dar- Landem und auf den Kontinenten beitragen. gestellt und die Hypothesen iiber die wichtigsten Das Ziel der vorliegenden Monografie war, die Faktoren, die fur ihre Gestaltung verantwortlich sind Entstehung von der Flora der neueren synanthro- erórtert (Kpt. 10). Man hat die Anordnungskarten fur pischen und auf dem polnischen Gebiet heimisch wer- viele Pflanzenarten erganzt (Kpt. 7) und iiber 5 neue denden Ankómmlingen (Kenophyten) zu untersuchen Karten erschafft (Abb. 39, Kpt. 7 u. Beilage C). Man und die bisherigen Kenntnisse im dem Bereich zusam- hat sich die Muhe gemacht, historische Veranderun- menzufassen. Die Verfasserin wollte auch die Ge- gen der Reichweiten von Kenophyten zu rekonstru- schichte und die Richtungen der, tiber die Pflanzen der ieren und auf die móglichen Migrationswege hinzu- fremden Herkunft gefuhrten Forsehungen zeigen und weisen (Kpt. 9). Besprochen wurden auch dynami- die wichtigsten, zu Errungenschaften der biogeographi- sche Tendenzen von Kenophyten unter Beriicksich- schen Wissenschaften eingezahlten Monografien der tigung der Faktoren, die die Besetzung von verschie- polnischen Botaniker vorbringen (Kpt. 2; Tab. 1). denen Biotoptypen begiinstigen (Kpt. 11). Man hat von In Folgę der Forsehungen wurde das neue Ver- der Listę sog. invasive Pflanzenarten ausgewahlt (die zeichnis der Pflanzenarten mit dereń ókologisch- Listę von invasiven Kenophyten fur das Gebiet Po geographischer Charakteristik (Beilagen A, B u. Kpt. lens) und die mit der Invasion bedrohten Gebiete 5. 1 u. 8) erschafft. Da es sich der Verfasserin gelun- genannt. Auf diese Weise wurde zur Diskussion iiber gen hat, zu historischen Quellen (historische/„alte” die Auswahlkriterien von Kenophyten der erste Floren, Herbarien) zu gelangen, konnte sie die ersten AnstoB gegeben. Appendices

Abbreviations and symbols used in Appendix A & В

Species - Latin name and synonym(s); species are arranged Origin - native rangę alphabetically; species names nomenclature according to Eur - Europę M ire k et al. 2 0 0 2 Asia H - hybrid origin Am N - North America LF - life form according to R aunkiaer (1 9 0 5 ) Am S - South America M - megaphanerophyte Afr - Africa N - nanophanerophyte С - central Ch - chamaephyte E - east H - hemicryptophyte N - north G - geophyte S - south Ну - hydrophyte W - west T - therophyte Way of INT - way of introduction to the country; hybrids li - clim ber escaped from cultivation are considered “intentionally” p - parasithe Ul - unintentionally —> vector of accidential introduc R - reproduction tion (in brackets): G - generative G - grain V - vegetative S - with seeds of other plants P - pollination mode BS - bird-seeds w - wind SB - soy beans i - insects BA - ballast s - self-pollination W - wool a - apogamic GA - garden materiał Disp - dispersal mode FD - fodder aut - autochory ane - anemochory (wind) В - botanical (as weed in botanical gardens) bar - barochory AN - animals egz - egzochory (epizoochory) RW - railways end - endozoochory P - potatoes myr - myrmecochory (dispersal by ants) I - intentionally —> planting purpose (in brackets): hyd - hydrochory (water) O - ornamental anthr - anthropochory (dispersal by humans) FO - forestry Prop - propagule A - agriculture (incl. food) se. - seed FD - fodder fr. - fruit M - medicinal st. - stem В - botanical (botanical gardens) ro. - root С - cultivation (e.g. for bees, cosmetic industry, rh. - rhizome lawns, landscaping, reclamation) ros. - rosette I/UI - both ways pl. - whole plant First record for Europę - year (if available) or period of LS - life strategy (G rjme 1979) the first record in the wild; for some taxa (mainly woody С - competition, plants) also year of deliberate introduction [I]; S - stress, Ar ? - in some part of Europę considered as archaeo- R - ruderal, CS, CSR, SR phyte (“oldcomer”); in some cases a few known oldest data were given, Anc - from ancient time in cultivation, (+1) - inerease of(in) number of local 1-50 - source (author & year) of information; listed at the ities, inerease in abundancy over end of the table existing localities F irs t record for Poland - year (if available) or period of (+2) - high inerease of localities (col- the first record in the wild; for some taxa (mainly woody onizing new localities) plants) also year of deliberate introduction [I]; in some (-/+) - disappearing of some localities cases a few known oldest data were given and appearing of new localities The oldest locality & source of data for the first record ? - undefined dynamie tendencies for Poland Hab - tape of habitats invaded Locality: region or(and) town N - natural Source: author & year; for herbarium data abbreviation S - semi-natural for particular herbarium was given (in bold). A cro- H - human-made (anthropogenic) nyms for herbaria after M ir e k et al. 1997. Inv. elsewhere - described as invasive elsewhere; in brack В - Botanisches Museum Berlin-Dahlem; K O R - In- ets type of invaded habitat is given stitute of Dendrology, Polish Academy of Sciences, agr. - agricultural, rip. - riparian, urb. - K órnik; KRA - Institute of Botany, Jagiellonian Uni- urban versity; KTC - Department of Botany, Institute of Bio- logy, J. Kochanowski Pedagogical University, Kielce; Maps - published distribution maps (only most KTU - Department of Plant Systematics, University important ones) and distribution map for of Silesia; L BL - Department of Systematics and Poland (in italics) Phytogeography, Institute of Botany, Maria Curie- ** - indicates that distribution map was com- Skłodowska University, Lublin; M GS - Upper Sile- piled exclusively on the herbarium data sian Museum; POZ - Adam Mickiewicz University Imp stud - important studies for Poland in Poznań; PRC - Herbarium at the University of Prague; SZUB - Department of Botany, Szczecin Uni- Sources of the first data for Europę: 1 -H ereźniak 1992; versity; TRN - Institute of Biology and Environment 2 - Lohmeyer & Sukopp 1992; 3 - Hegi 1908-1931; Protection, N. Copernicus University in Toruń; 4 - Lauener 1996; 5 - Pyśek et al. 2002; 6 - Frey W - Herbaria in Wien; WA - Department of Plant 1974; 7 - JehlIk 1998; 8 - Stace 1997; 9 - Kużniew- Systematics and Geography, Institute of Botany, War- ski 1996; 10 - H ardtke & Ihl 2000; 11 - Reichenbach saw University; W RSL - Museum of Natural History, 1842; 12 - Meusel et al. 1965; 13 - Sudnik- University of Wrocław -Wójcikowska 1987a; 14 - M eusel et al. 1992; 15 - Nrs of loc - number of localities in distinguished pe- Hegi 1935-1961; 16 - Meusel et al. 1978; 17 - riods Żukowski & Piaszczyk 1971; 18 - Ascherson & Gra- N rs o f sq - number of ATPOL squares (total number eabner 1902-1904; 19 - Ascherson & Graeabner of squares for Poland: 3646); 1901-1913; 20 - Ascherson & Graeabner 1913; 21 - * - indicates that number of squares recorded Ascherson & Graeabner 1915; 22 - Ascherson & need to be verified Graeabner 1917; 23 - Ascherson & Graeabner 1938; n.c.d. - not complite data 24 - Zając et al. 1998; 25 - Kowarik 1995a; 26 - Guzik Dyn - frequency and dynamie tendencies accord- & Sudnik-Wójcikowska 1994; 27 - Dyakowski 1899; ing to Z a r z y c k i et al. 2 0 0 2 28 - Perring & W alters 1962; 29 - Kornaś 1968b; Frequency in the wild at the territory of 30 - Rostański K. 1998; 31 - Rostański K. & Serwat the country: k a 1968; 32 - G utte & Rostański 1971; 33 - Rostański 1 - very Iow number of localities (1-20) K. & Kloss 1965; 34 - Hantz 1979; 35 - Starfinger 2 - Iow number of localities (up to 1 9 9 7 ; 36 - Misiewicz et al. 1 9 9 6 ; 37 - Bailey & 100) Conolly 2000; 38 - Hollingsworth & Bailey 2000; 3 - high number of localities, but with narrower distribution (in one or 39 - Zieliński 1991; 40 - Zieliński 2004; 41 - two regions of the country) Franc1rkova 2001; 42 - Kucharski 1992; 43 - B a l o g h 4 - high number of localities in many 2 0 0 1 ; 44 - Drescher & Prots 2003; 45 - Adamowski regions et al. 2 0 0 2 ; 46 - Seneta 1994; 47 - Gutte 1997; 5 - common (abundant) in the whole 48 - Seneta & Dolatowski 1997; 49 - Guzik 2002; territory 50 - Ascherson 1866; 51 - Rostański K ., personal inf. Dynamie tendency (in brackets): (specimen from the Natural History Museum in Budap- (-2) - high decrease of(in) number of est); 52 - Krawiecowa 1951; 53 - Lhotska & Kopecky localities 1966; 54 - P e r r in s et al. 1993; 55 - Rostański K. 1982; (-1) - decrease in number of(in) local 56 - G u d ż in s k a s 1997d; 5 7 - G u d ż in s k a s 1997c; 58 - ities or decrease in abundancy G u d ż in s k a s 1998a; 59 - G u d ż in s k a s 1997b; 60 - W eber

over existing localities 1998; 61 - G u d ż in s k a s 2000a; 6 2 - G u d ż in s k a s 2000b. Nrs Nrs Nrs Nrs First First The oldest locality of of of of Nrs Way record Name of species Family LF R P Disp Prop LS Origin record in Poland loc. loc. loc. loc. of Dyn Hab. Inv. elsewhere Maps of INT for for Europęf & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Acer negundo L. Aceraceae M G/V i w ane se. С Am N I [0] 1688 [I]1 1808 [I] Kraków - botanical garden 0 3 30 3526 1379 4(+2) NSH Eur C; L ittle 1971; hyd ro. 16992 1873 ? [I] (H e r eż n ia k 1992); Kra Lithuania Z a j ą c A. & Z a j ą c M. 2001 see also 1899 ków (B o eh m 1873); Wrocław [rip. agr. & urb.] Chapter 7 (Baenitz herb. PRC, W, WRSL); Puławy (Berdau herb. LBL)

Acorus calamus L. Araceae Ну V i w anthr rh. CS Asia С I[M, B] 1557 [I]3 XVI * generał information (after 8 88 146 4319 1999 5(-/+) NS H ulten 1964; H ulten & hyd & S - U l XVI2 1613* S y r e ń sk i 1613); XVIII - K lu k F ries 1986; M eu sel et al. 157718 1652 (1786); Warszawa (S u d n ik - 1965; 1824 -W ó jcikow ska 1987a); Mazow Z a j ą c A. & Z a j ą c M. 2001 sze Lowland: Wyszogród (Z a l ew sk i 1892 after Gawarecki 1824); Warszawa (S zu bert 1824)

Ailanthus altissima (Mili.) Swingle Simaroubaceae M G/V i ane se. С Asia E I [O , M ] 1751 [I]4 1818 [1] [I] (H ereżniak 1992); Wrocław 0 0 3 31 29 2(+l) H Eur С & S original: [= A. glandulosa Desf.] hyd ro. [China] 1780 [I]2' 25 1931 (M ey er 1931) [urb. & rip.] see Chapter 7, Fig. 39 18745 Am N 190225 see also Chapter 7

Amaranthus albus L. Amaranthaceae T g i w ane se. SR Am N Ul 17236 1907 Lublin Upland: Rejowiec 0 0 60 782 379 3(+/—) Н Eur С Jalas & Suominen 1980; egz [W] [G, B] (T r zeb iń sk i 1930) [agr.] F r e y 1974; T o k a r s k a -G u z ik 2001b (1)

Amaranthus blitoides S. Watson Amaranthaceae T g i w ane se. CR Am N Ul [G] 18932 1911 Wielkopolska Lake District: 0 0 8 283 150 3(+l) H Eur С J ala s & S uo m in en 1 9 8 0 ; egz [W] Krosno Odrzańskie; South [agr.] F r e y 1974; T o k a r s k a -G u z ik W ielkopolska Lowland: Żary 20Ólb (2) (D eck er 1 9 1 1 )

Amaranthus chlorostachys Willd. Amaranthaceae T g i w ane se. CR Am С Ul [W] 1872 1872 Carpathian Foothills: Tarnów 0 2 14 425 260 3(+l) Н Eur С Jalas & S u o m in en 1980; [= A. hybridus L.] egz & S (K n a pp 1872) [agr.] F r e y 1974; T o k a r s k a -G u z ik 2001b (3)

Amaranthus liridus L. Amaranthaceae T g i w ane se. CR Eur S I [A] Ar2 1826 Gdańsk (Klinsmann herb. TRN) 9 77 117 72 8 453 3(-/+) Н Eur С J ala s & S u o m in en 1 9 8 0 ; [= A. ascendens Loisel.] egz & Afr N - U l [agr.] F r e y 1974; T o k a r s k a -G u zik end 200lb (4) myr anthr

Amaranthus retroflexus L. Amaranthaceae T g i w ane se. CR Am N I [B] / 17836 1801 O p o le , Gdańsk 8 169 291 7651 2379 5(+2) Н Eur С M e u s e l et al. 1965; H u l t e n egz [W] Ul [BA] 1814 (T h e l l u n g 1914) [agr.] 1968, 1971; J a l a s & anthr & Am С S u o m in e n 1980; H u l t e n & F r ie s 1986; F r e y 1 9 7 4 ; Z a ją c A. & Z a j ą c M. 2001

Ambrosia artemisiifolia L. Asteraceae T g i w ane se. CR Am N 1 8 6 3 7 1613* * generał information: probable 0 11 25 101 61 2 -3 Н some regions M e u s e l et al. 1 9 9 2 ; Ul egz [E & SE] Germany XVIII ref. this species (after S y re ń s k i (+/-) of Eur C, S & E; T o k a r s k a -G u z ik 2001b (5 ) [G, SB, anthr 186514 1873 1613); XVIII - K l u k (1786); Am N BA] Silesian Lowland: Szcze- panowice (Plotel herb. W RSL)

Ambrosia psilostachya DC. Asteraceae H g /v i w ane se. С Am N Ul [G] 1901 1901 Świnoujście (Ruthe 0 0 9 30 21 2 ( + /- ) Н Am N M eu sel et al. 1 9 9 2 ; [= A. coronopifolia Torr. & A. Gray] egz rh. [SE] 19038 herb. SZUB) Tokarska-Guzik 2001b (6 ) anthr

Anthemis ruthenica M. Bieb. Asteraceae T G i ane se. CR Eur SE Ul [G] 1869 1869 Carpathian Foothills: Krządka 0 29 63 408 269 3(+l) Н M eu sel et al. 1992; egz (Jachno 1869) Z a j ą c A. & Z a j ą c M. 2001 anthr Nrs Nrs Nrs Nrs First First The oldest locality of of of of Nrs Way record Name of species Family LF R P Disp Prop LS Origin record in Poland loc. loc. loc. loc. of Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source o f data up to up to up to up to sq. Poland 185 0 19 0 0 195 0 2000

Anthoxanthum aristatum Boiss. Poaceae T G w ane se. R Eur S Ul [FD] 18 0 5 186 6 West Pomerania: Kwidzyn 0 6 4 2 1031 5 7 7 3 - 4 H M e u s e l et al. 1 9 6 5 ; [= A. puelii Lecoą & Lamotte] egz 1 8 1 3 4 (K lin g g r a eff 1 8 6 6 ) (+2) Tokarska-Guzik 2001b (7 ) anthr see also Chapter 7

Artemisia annua L. Asteraceae T G i s ane se. CR Eur SE I [C] 1871 [I] 1871 [I] Wielkopolska Lake District: 0 11 35 3 3 7 154 3(+/-) H Czech Rep. [agr.], M eu sel et al. 1 9 9 2 ; egz & Asia W - U l 1881 1881 Cerekwica (Ż uk o w sk i Slovac Rep. [rip.], Ż u k o w sk i & P ia s z c z y k 1 9 7 1 ; anthr [G. BS, W] 1 8 9 9 7 & P ia szc zy k 1 9 7 1 ) Hungary T o k a r sk a -G u zik 2 0 0 1 b (8 )

Artemisia austriaca Jacq. Asteraceae Ch WG i s ane ros. CS Eur SE Ul [RW] 1871 1871 W a rsz a w a (R o s t a f iń s k i 1 8 7 2 ) 0 8 33 3 7 4 2 1 7 3(+/-) H M eu sel et al. 1 9 9 2 ; egz se. & Asia W 1 9 4 6 10 Ż u k o w sk i & P ia s z c z y k 1 9 7 1 ; anthr see also T o k a r sk a -G u zik 2 0 0 1 b (9 ) Chapter 7

Artemisia dracunculus L. Asteraceae H G/V i s ane se. C Am N І [А. M] X V I'0 XVI[I] * generał information (after 1 10 28 87 59 2(+/-) H M eu sel et al. 1 9 6 5 ; egz rh. & Asia A r17 161 3 * S y r e ń s k i 1 6 1 3 ); XVIII - K l u k Ż u k o w sk i & P ia sz c z y k 1 9 7 1 ; anthr XVIII ( 1 7 8 6 ) as cultivated plant; T o k a r sk a -G u zik 2 0 0 1 b (10) 18 5 0 Poznań (Schoenke herb. POZ)

Asclepias syriaca L. Asclepiadaceae H V/G i ane rh. C Am N [E] I [0] XVIII10 XVIII Kalisz, Lublin 0 4 7 62 52 2(+ l) H some part original: see App. C anthr se. 1 8 5 5 43 1872 (R osta fiński 1 8 7 2 ) o f Eur С & S 19 0 15

Atriplex oblongifolia Waldst. & Kit. Chenopodiaceae T G s i ane se. CR Eur E, Ul [RW] 2/2 XIX10 1882 Toruń (A b r o m e it et al. 1926) 0 4 18 154 100 3 ( + l ) H Czech Rep. M e u s e l et al. 1 9 6 5 ; J a l a s & [= A. oblongifolium Waldst. & Kit.] hyd Asia W S u o m in e n 1 980; anthr & Afr Tokarska-Guzik 2001b (1 1 )

A triplex tatarica L. Chenopodiaceae T G s i ane se. CR Eur S Ul [RW] 1820" 1847 W a rsz a w a (W a g a 1 8 4 7 ) 1 13 36 294 153 3 ( + i ) H Jalas & Suominen 1980; [= A. tataricum L.] hyd & SE, A r H ulten & Fries 1986; anthr Asia C T o k a r sk a -G u zik 2 0 0 1 b (12)

Barbarea intermedia Boreau Brassicaceae H G i s ane se. CR Eur S Ul 1908 West Pomerania: Połczyn 0 0 1 11 9 1(?) H M ire k 1984. 1997; Z a ją c A. fr. & w (Romer 1908) & Z a ją c M . 2 0 0 1

Beckmannia eruciformis Host. Poaceae HG/V w ane se. Eur E I [FD] 18 3 7 1837 south-western Poland: 2 4 9 64 57 2(+ l) SH H u l t e n 1964; H u l t e n & F r ie s egz rh. & S , - U l Wrocław & Bolesławiec 1986; anthr Asia W (S c h n eid er 1 8 3 7 ) F r e y & P aszk o 2 0 0 0 ; Z a ją c A. & Z a ją c M . 2 0 0 1

Bidens connata H. L. Miihl. Asteraceae T G i s egz se. CR AmN [E] I [B] / 18652 14 c a . 18 7 4 Bydgoszcz 0 8 22 148 114 3(+l) NH M e u s e l et al. 1992; [= B. connatus H. L. Miihl.] hyd Ul 1895 (T r z c iń sk a -T acik 19 7 1 a) Tokarska-Guzik 2001b (13) [W, S]

Bidens frondosa L. Asteraceae T G i s egz se. CR Am N [N] Ul 17362 1777 Wrocław (K r o c k e r 1790); 0 4 60 3142 1068 4(+2) NSH Eur C M e u s e l et al. 1992; W a l t e r & [= B. melanocarpus Wiegand] hyd [W, S ] / see also 1869 Wielkopolska Lake District: [rip.] S t r a k a 1970; I [B] Chapter 7 Słubice (Brand after T r z c iń s k a 1 9 6 1 ; Z a j ą c A. & S c h u m a c h e r 1942) Z a j ą c M . 2 0 0 1

Bromus carinatus Hook & Am. Poaceae T H G w ane se. Am N 1 [C, A] 1912 1911 Wielkopolska Lake District: 0 0 3 1130 404* 3-4 NH Z a ją c A . & Z a jąc M . 2 0 0 1 egz 19345 Torzym (D e c k e r 1911) (+2) anthr

Bryonia alba L. Cucurbitaceae H G/V i end fr. C Eur E I [0, M] Ar5' 10 XVII [I] XVIII - Kluk (1786) - only 8 115 169 1328 728 3-4 NSH Czech Rep. M eusel et al. 1992; li hyd st. & Asia W 1824 as cultivated plant; (+D Z a ją c A. & Z ając M . 2 0 0 1 anthr Mazowsze Lowland: Wyszogród (Zalewski 1892 after Gawarecki 1824)

Bryonia dioica Jacq. Cucurbitaceae H G/V i end fr. C Eur S 1 [B. O] 1820'° 1847 West Pomerania: Pomoc near 1 4 13 116 77 2-3 H M e u s e l et al. 1992; [= B. cretica L. subsp. dioica (Jacq.) li hyd st. & w Ar5 Chojnice (H aub 1847 ATPOL (+/-) T o k a r s k a -G u z ik 2 0 0 1 b (14) Tutin] anthr sources) Bunias orientalis L. Brassicaceae H G/V i s ane se. C Eur SE Ul 172012 1858 Gdańsk (Klinsmann herb. 0 49 120 1353 567 3 - 4 SH Czech Rep. M eu sel et al. 1965; H ulten & egz ro. & Asia W [FD. G] Russia W; TRN) (+2) & Slovac Rep. F ries 1986; aut 181452 T o k a r s k a -G u z ik 2 0 0 lb (1 5 ) anthr 18565' 7 186710 see also Chapter 7 Nrs Nrs Nrs Nrs First First The oldest locality o f o f o f o f Nrs Way record Name of species Family LF R P Disp Prop LS O rigin record in Poland loc. loc. loc. loc. of Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Cardaria draba (L.) Desv. Brassicaceae G H G/V i s aut se. CSR Eur SE U l 1652 ? 13 1837 Sudety Mts.: 2 44 174 1048 576 3 -4 SH Czech Rep. M eu se l et al. 1965; H ulten & [= Lepidium draba L.] ane ro. & A sia [G, FD ]/ 167523 Bolesławiec (S ch n eid er 1837) (+2) F ries 1986; egz SW I [0] 172812' 23 T o k arsk a-G u zik 2 0 0 l b (16) anthr 1829* A r5

Centaurea diffusa Lam. Asteraceae T H G i s ane se. CSR Eur SE Ul 18762 1878 Silesian Upland: Szczakowa 0 3 12 178 89 2-3 SH M e u s e l et al. 1992; egz & A sia [BA, G] (Unverricht herb. KRA) (+1) T o k a r sk a -G u zik 2 0 0 l b (17) anthr SW

Chaerophyllum aureum L. Apiaceae H G/V i egz se. C Eur C U l [RW] 1809 ? Dukla (Besser ?); 0 0 0 12 6 1(+1) NH O k l e je w ic z 1999, 2001 & S 1994 Beskidy Mts.: Szczawne (O k l ejew ic z 1999)

Chamomilla suaveolens (Pursh) Rydb. A steraceae T G i s ane se. R Am N Ul 178256 XVII ? Wrocław (Uechtritz h e rb . 0 72 254 13125 2965 5(+2) Н Eur C H u l t e n 1971; M e u s e l e t al. [= Matricaria discoidea DC.] egz & Asia E 1850'4 1862 WRSL; Knebel h e rb . WU) 1992; end 18515 Z a ją c A . & Z a ją c M . 2 0 0 1 anthr 1852'° see also Chapter 7

Chenopodium aristatum L. Chenopodiaceae T g w ane se. ~ Eur E, U l [S] 1941 Szczecin (T r z c iń sk a -T acik 0 0 1 3 3 1(?) H Hungary G ł a z e k , M ir e k & P o ło ń sk a Asia C 1992) 1 9 8 5 ; Z a j ą c A . & Z a ją c M . & E 2 0 0 1 Chenopodium botrys L. Chenopodiaceae T g w ane se. R Asia C I / U l A r5 1613* * - generał information 4 26 34 70 49 2 (+/-) H Hulten 1971; Jalas & Suomi- hyd 1829 (after S y r e ń sk i 1613); Lublin nen 1980; Hulten & Fries 1986; 1837 Upland: Horodło (W aga 1847) T o k ar sk a-G u zik 2 0 0 1 b (18)

Chenopodium strictum Roth Chenopodiaceae T g w ane se. CR Asia C Ul XIX 24 1891 Toruń (Abromeit e t al. 1926) 0 1 1 896 256 3(+/-) Н Hungary P a śn ik 2 0 0 1 * * [= Ch. album L. subsp. striatum hyd 193910 (Kraśan) Murr]

Chenopodium suecicum M urr Chenopodiaceae T g w ane se. CR Am N, Ul 1827 Mazury Lake District: Dobre 96 2-3 H J al a s & S u o m in e n 1980; hyd Eur N, Miasto (ATPOL sources: Sey- (+ /-) H ulten & F ries 1986; Asia N dler 1827) P a śn ik 2 0 0 1 * *

Clematis \'italba L. Ranunculaceae N li g i ane se. C Eur C I [O] 1663 [I]25 1613* * - generał information (after 1 20 43 354 216 3 (+ l) NH New Zealand M e u s e l e t al. 1965; H e g i 1974; egz [m], 188325 XVIII S y r e ń sk i 1613); XVIII - K luk Jalas & Suominen 1989; anthr A sia W see also 1847 (1786) - as cultivated plant; T o k a r sk a -G u zik 2 0 0 1 b (19) & A fr C hapter 7 Lublin Upland: Kazimierz NW (W ag a 1847)

Conyza canadensis (L.) Cronquist Asteraceae T H G i s ane se. CR Am N Ul [S] 16462'14 1730 Warszawa (S udnik-W ójcikowska 8 108 196 11601 2929 5(+2) Н Eur C H u l t ć n 1971; HultEn & Fries [= Erigeron canadensis L.] egz [N] 1825 1987a after Erndtel 1730); [urb. & agr.] 1986; M e u s e l e t al. 1992; 1837 around Gdańsk (R ey g er 1825); Z a ją c A . & Z a ją c M . 2 0 0 1 south-western Poland: Bytom Odrzański, Oława, Wołów (S ch n eid er 1837)

Corydalis lutea (L.) DC. Fumariaceae H G i myr se. CSR Eur C I [B. 0 ] ca.2/2 1884 Sudety Mts.: Bożejów 0 1 5 29 26 2(—1) H M eu sel 1943; J al a s & S u o m i [= Pseudofumaria lutea (L.) Borkh.] [Alps] X V III10 (S chube 1903b) n en 1991; 1884 T o k arsk a-G u zik 2 0 0 1 b (20) 18865

Crepis aurea (L.) Cass. Asteraceae H G i s ane se. Eur C U l [AN] XIX / West Tatra Mts.: Stoły 0 0 0 1 1 l(?) N Z a ją c A . & Z a ją c M . 2 0 0 1 egz [Alps] XX C learing (M irek 1995) m yr 1995 Cymbalaria muralis P. G aertn., B. Scrophulariaceae Ch HG/V i aut se. CSR Eur S I [O] 16408 1837 Sudety Mts.: Zgorzelec, 3 62 181 350 165 3(-l) Н M e u s e l e t a l. 1 9 7 8 ; Mey. & Scherb. ane fr. - U l A r ? Bolesławiec (S chneider 1837 Z a ją c E . U. & Z a ją c A . 1 9 7 3 ; [= Linaria cymbalaria (L.) Mili. anthr see also Z a ją c A . & Z a ją c M . 2 0 0 1 Chapter 7 Datura stramonium L. Solanaceae T G i s ane se. CR Am N I [C] 15842 1613* * - generał information (after 6 128 205 1881 1044 4 (+ /-) Н some part of Eur H u l t e n 1971; Hultżn & Fries egz fr. [SE] -►Ul 1652 S yreński 1613); Warszawa (S ud C & S 1986; M e u s e l e t al. 1978; anthr A sia ? [W, BS, 1825 n ik -W ójcikow ska 1987a); XVIII Z a ją c A . & Z a ją c M . 2 0 0 1 SB] 1837 - K luk (1786); Oliwa (R eyger 1825); Silesian Lowland: Oława, Wołów & Bytom Odrzański (S chneider 1837) Nrs Nrs Nrs Nrs First First The oldest locality o f of o f o f Nrs Way record Name of species Fam ily LF R P Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europę* & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Digitalis purpurea L. Scrophulariaceae H T G i s ane se. CR Eur W I 17905 1809 ? around Kraków (B esser 1809); 5 24 59 341 169 3(+ l) NSH Czech Rep. M eusel e t a l. 1978; Hulten & anthr ros. [0 , M] see also 1862 Beskidy Mts.: Klimczok Fries 1986; - U l [S] C hapter 7 (KOLBENHEYER 1862) C ynuel 1 965; H antz 1993; Z ając A . & Z a ją c M. 1997, 2001

Diplotaxis muralis (L.) DC. Brassicaceae T H G i s ane se. CSR Eur S U l X V III'5 1851 Poznań (R itschl 1851) 0 31 133 2049 991 4 (+ l) H Hulten & Fries 1986; & W [BA, G] 1827” T o k ar sk a-G u zik 2001b (21) [Afr.] 1842 A r5

Diplotaxis tenuifolia (L.) DC. Brassicaceae Ch H G i s ane se. CR Eur S U l 159712 1652 W arszawa (S u d n ik -W ó jc ik o w - 1 36 70 497 245 3(+ l) SH M eusel e t a l. 1965; Hulten & & W [BA, G] England 1836 sk a 1987a); Gdańsk (K l in s - Fries 1986; [Asia, 17682 m a nn 1836) T o k arsk a-G u zik 2 0 0 1 b (22) Afr.] Eur С A r5

Echinocystis lobata (F. Michx.) C ucurbitaceae T li G i end se. CR Am N I [O] 190414-43 1937 Wielkopolska Lake District: 0 0 7 2047 708 3 -4 NSH Czech Rep. M eusel e t a l. 1992; Torr. & A. Gray fr. [E] see also G ubin (L ademann 1937) (+2) & Slovac Rep. T o k a rsk a-G u zik 2 0 0 1 b (23) p i C hapter 7 [rip.], Hungary

Echinops exa!tatus Schrad. A steraceae H G i s egz sę. Eur E 1 [C, O] 1897 1897 Chrośle near Nowe Miasto Lu 0 1 1 9 9 1(7) SH M eusel e t a l. 1992; [= E. c o m m u ta tu s Jur.] ane & Asia W 199510 bawskie (Karslen h e rb . TRN) Z a ją c A . & Z a ją c M . 2 0 0 1

Echinops sphaerocephalus L. Asteraceae H G i s egz se. С Eur E I [C, 0 ] 1613 XVI * - generał information (after 1 25 99 910 489 3 (+ l) SH Czech Rep. M eusel e t a l. 1992; ane & Asia W 1652 1613* S y r e ń s k i 1613); Warszawa Z a ją c A . & Z a ją c M . 2 0 0 1 myr 1809 1652 (S u d n ik -W ójc ik o w sk a 1987a); anthr A r2 1809 K raków (B esser 1809) see also C hapter 7

Elodea canadensis M ichx. Hydrocharitaceae Н у V aut p i Am N U l [BA] 1836®'27 1867 Gdańsk (Abromeit e t al. 1898) 0 140 226 3681 1847 4(+l) NSH Eur C [water] H ulten 1964; H ulten & Fries hyd / see also 1986; egz I [B, 0 ] Chapter 7 Z ając A. & Z a jąc M. 1992, 2001 anthr

Elsholtziu ciliata (Thunb.) Hyl. Lam iaceae T G i ane sę. R Asia E 1 [M] 183058 1829 ? W arszawa (S u d n ik -W ó jc ik o w - 1 79 147 1352 814 3^1 H S wieboda 1 963; To ka rsk a-G uzik [= E. p a tr in i (Lepech.) Garcke] end - U l 1847 1847 ska 1987a; W aga 1847) (+ /-) 2 0 0 1 b (24) myr 18535 anthr see also Chapter 7

18912. i6 Epilobium ciliatum Raf. O nagraceae H G is ane se. С Am N U l [S] 1917 Białowieża Forest (R ubner 0 0 1 1224 470 3-4 NSH Czech Rep. M eu sel e t al. 1978; H ulten & [= E. adenocaulon H ausskn.] [N] 1917) (+ D F ries 1986; Z a ją c A . & Z a ją c M . 2 0 0 1

Eragrostis albensis H. Scholz Poaceae T G w ane se. unclear Ul 0 0 3 301 50 2-3 NH S u d n ik -W ó j c ik o w s k a & G u z ik egz (+1) 1 9 9 6 ; Z a j ą c A . & Z a j ą c M. hyd 2 0 0 1

Eragrostis minor Host Poaceae T G w ane se. R Eur SE Ul 1819'° 1838 Wrocław Gajowice (W immer 1 13 96 1041 581 3 -4 H Eur C [urb.] T o k a r s k a -G u z ik 2 0 0 1 b (25) egz & Asia W [W, FD, A r 1868; F iek 1881) (+2) BS] see also C hapter 7

Eragrostis multicaulis Steud. Poaceae T G w ane se. Asia E Ul [B] 182426 1879 Wrocław (K nebel 1879) 0 1 1 4 4 1(?) H G u z ik & S u d n ik -W ó j c ik o w s k a egz & SE 1 9 9 4 ; Z a j ą c A . & Z a j ą c M. 2 0 0 1 Erechtites hieracifolia (L.) Asteraceae H G i ane se. Am N Ul [S] 17002 1902 Silesian Lowland: Prószków 0 0 32 124 73 2-3 NSH Hungary Croizat 1952; Meusel e t al. Raf. ex DC. & S [N] (Schube 1902) (+1) 1992; C z a r n a , G ó r s k i & T o k a r s k a - -G u z ik 2 0 0 1 ; G ó r s k i, C z a r n a & T o k a r s k a -G u z ik 2 0 0 3

Erigeron annuus (L.) Pers. Asteraceae H T G i s ane se. С Am N I [0 , B] 17002 14 1830 Silesian Lowland: Nowa Kar 2 65 149 3557 1133 4(+2) SH Hungary M eusel e t al. 1992; egz [N] - U l czma upon Odra river (F iek Z a ją c A. & Z a ją c M. 2001 myr 1881) Nrs Nrs Nrs Nrs First First The oldest locality o f o f o f o f Nrs Way record Name of species Family LF R P Disp Prop LS O rigin record in Poland loc. loc. loc. loc. of Dyn Hab. Inv. elsewhere o f INT for M aps for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Erigeron ramosus (Walters) Britton, Asteraceae H T G i s ane se. Am N I [0 ] X V III / 1888 Silesian Lowland: Opole 0 1 28 849 408 3 ( + l) SH Hungary Z a ją c A.

Erysimum wahlenbergii Brassicaceae H G i s aut se. Eur C Ul 1974 Tatry Mts.: near Murowaniec 0 0 0 1 1 1(?) S Z a ją c A. & Z a ją c M. 2001 (Asch. & Engl.) Borbas ane [Carp.] shelter (P iękoś & M irek 1974)

~~Euphorbia humifusa Willd. Euphorbiaceae T G i aut se. R Asia E Ul [B] 181322 1846 Kraków (R ostański 1992) 1 7 13 18 8* 1(+/-) H M eusel et al. 1978 m yr~~

Galinsoga ciliata (Raf.) S.F. Blake Asteraceae T G i s ane se. CR Am C I [B] / 1853'4 1876 Silesian Lowland: Wrocław 0 7 97 6777 2021 4-5 H Eur C Hulten & Fries 1986; Meusel [= G. ąuadriradiata Ruiz & Pav.] egz [m] U l 18662 (Knebel herb. WU), Głogówek (+2) [agr. & urb.] et al. 1992; myr Am S ? [B, G] (R ichter herb. MGS) Z a ją c A. & Z a ją c M. 2001 anthr

Galinsoga parviflora Cav. A steraceae T G i s ane se. CR Am S I [B] / 17982 1807 Budowo near Słupsk 1 135 253 10932 2726 5(+2) H Eur C Hulten & Fries 1986; Meusel egz & C [m] Ul 1863 (Thellung 1915); Wrocław [arg. & urb.] et al. 1992; myr [B, G] (Uechtritz herb. WRSL) Z a ją c A. & Z a ją c M. 2001 antr

Genistella sagittalis (L.) Gams in Hegi Fabaceae Ch G i aut se. CS Eur W U l [S?] 1928* 1929 Carpathian Foothills: Tryńcza 0 0 1 11 11 H?) NS Hegi 1924; Meusel et al. 1965; [= Genista sagittalis L., & S near Przeworsk (N owiński K ażmierczakowa & TUM1DAJOWICZ Chamaespartium sagittale (L.) 1929) 1981; T o k a r sk a -G u zik 2001b P. E. Gibbs] (26)

~~Geranium divaricatum Ehrh. Geraniaceae T G i s aut se. R Eur S Ul Ar'0 1840 Silesian Lowland: near 2 36 55 71 38 2(+ -) H T o k a r sk a -G u z ik 200Ib (27) egz & Asia W W rocław (W immer 1841)~~

Geranium pyrenaicum Burm. f. Geraniaceae H G i s aut se. CSR Eur S Ul / I 176210'16 1837 Sudety Mts.: Bolesławiec 1 46 220 682 396 3 (+ l) SH Czech Rep. M eusel et al. 1978; Hulten & egz (S chneider 1837) Fries 1986; C iac iu ra et al. 2001a, b

Geranium sibiricum L. Geraniaceae H G i s aut se. C Eur E, Ul / I 1840 1840 Sudety Foothills: near 2 4 7 24 20 2 (+ l) H M eu sel e t a l. 1978; M ir e k egz A sia W Dzierżoniów (F iek 1881) 1981b; Z a ją c A. & Z a ją c M . & E 2001

~~Glyceria striata (Lam.) Hitch. Poaceae H G/V w hyd se. Am N Ul 2/2 XX24 1989 Sieraków (Babczyńska-S endek 0 0 0 3 3 1(+1) s Z a ją c A. & Z a ją c M. 2001 egz rh. 195659 & Sendek 1989)~~

~~Helianthus decapetalus L. Asteraceae G V/G i s ane se. Am S I [O] X X 10 1956 Szczecin (ScHEUERMANN 1956) 0 0 0 19 18 1(+1) H T o k a r sk a -G u zik 200Ib (28) egz rh. 191043 m yr anthr~~

Helianthus laetiflorus Pers. H Asteraceae G V/G i s ane se. C Anthro- I [0 ] X X 10 1969 W rocław (Rostański K. 1969) 0 0 0 26 18 1-2 H T o k a r sk a -G u z ik 2001b (29) [= H. rigidus x tuberosus] egz rh. POg- 195943 (+D myr anthr

Helianthus tuberosus L. Asteraceae G WG i s ane se. C Am N I [O, c, 16272 1730 ? near Warszawa (after Sudnik- 0 7 30 1416 778 3-4 NSH some regions of M eusel et al. 1992; egz rh. FO, M] 1872 W ójcikowska 1987a); XVIII - (+2) Eur C Zając A. & Zając M. 2001 myr K luk (1787) - only as cultiva- anthr ted plant; Wielkopolska Low land: Kłódno (Rostafiński 1872)

Helleborus viridis L. Ranunculaceae H G i m yr se. CS Eur C I [0 ] XVIH'° 1868 south-western Poland: Głub 0 17 30 30 26 2(-2) NH Zając A. <£ Zając M. 2001 & w 18195 czyce, Strzelniki, Nowaki near Nysa (W immer 1868)

Heracleum mantegazzianum Apiaceae H G i s ane se. C Asia C 1 [0] 18625 1973 Baltic Coast: near Gryfino & 0 0 0 100 74 2-3 NSH Eur W, C & N T o k a r s k a -G u z ik 2001b (30)* Som m ier & Levier’ hyd & E Pyrzyce (Ć w i k l i ń s k i 1973) 146* (+2) [rip. & urb.] egz anthr

Heracleum sosnowskyi M anden.1 Apiaceae H G i s ane se. Asia SW I [C] 2/2 XX24 1980 West Pomerania: Strzelce Dolne 0 0 0 96 72 2(+2) NSH Hungary T o k a r s k a -G u z ik 2001b (30)* hyd [Cauc.] near Bydgoszcz (Rutkowski 146* egz unpubl.) anthr Nrs Nrs Nrs Nrs First First The oldest locality of o f o f o f Nrs Way record Name of species Family LF RP Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000 Hesperis matronalis L. Brassicaceae H G i s ane se. CS Eur S I [O] X V I2 XVII [I] * - generał information (after 2 43 69 724 510 3(+l) NSH Hulten & Fries 1986; 1791” 1613* Syreński 1613); XVIII - K luk Z a ją c A . & Z a ją c M . 2 0 0 1 18175 1837 ( 1787) - as cultivated plant; Sudety Mts.: Bolesławiec & K up (Schneider 1837)

Hyssopus officinalis L. Lamiaceae Ch G i ane se. CS Eur S I [M, C] (1594)2 XVII[I] * - generał information (after 0 20 45 69 59 2(-l) SH T o k a r sk a-G uzik 2 0 0 l b (31) end ro. & SE, 18195 1613* Syreński 1613); XVIII - K luk myr A sia SW 18292 1859 ( 1787) - only as cultivated & C plant; Carpathian Foothills: Tyniec (B erdau 1859) lmpaliens capensis M eerb. Balsaminaceae T G i aut se. Am N Ul [BA] 182228 1991 Baltic Coast: Trzebieradz, Trze 0 0 0 3 3 U?) N M eusel e t a l. 1978; H u lten & bież & Police (P awlaczyk & Fries 1986; A damowski 1991) P a w la c zyk & A d am o w sk i 1 9 9 1 ; Z a ją c A . & Z a ją c M . 2 0 0 1

Impatiens glandulifera Royle Balsaminaceae T G i aut se. C A sia C I [0, M] 1839 [I]53'44 1890 Sudety Mts.: Siodło, Płóczki 0 9 38 1574 675 3 -4 NSH Eur W & C Z a ją c E. U. & Z a ją c A . 1 9 7 3 ; [= I. r o y le i Walp.] ane [Himal.] 1845 [I]2' 44 Dolne, Stępnica & Płonina (+2) [rip ] T o k ar sk a-G u zik 2 0 0 1 b (32) end 185554' 44 (Schube 1903b) hyd see also C hapter 7

Impatiens parviJlora DC. Balsaminaceae T G i aut se. SR Asia C I [B] 183416 1850 near Gdańsk (Meusel e t a l. 0 54 136 6730 1681 4 -5 NSH Eur C M eusel e t a l. 1978; Hulten & ane & E - U l Russia 1857 1978); near Kraków (Ullepitsch (+2) [decidoues forests] Fries 1986; end 18372 16 h e r b .B ) Z a ją c A . & Z a ją c M . 2 0 0 1 hyd see also Chapter 7

Inula helenium L. A steraceae H G/V i s ane se. C Eur E, I [0 , M] 18195 XVI ? XVI? Zając A. e t a l. 1998; 2 82 168 416 273 3(+l) NSH M eusel e t a l. 1992; egz rh. A sia W 1613* * - generał information (after Z a ją c A . & Z a ją c M . 2 0 0 1 myr & C XVIII Syreński 1613); XVIII - Kluk 1837 (1787); Sudety Mts.: Jedlina Zdrój & Wołów (Schneider 1837)

Iva xanthiifolia Nutt. Asteraceae T G w ane se. CR Am N Ul 184214 1928 Gdańsk (P reu ss 1928) 0 0 2 294 150 3(+ /-) H Eur S M eusel e t al. 1992; egz [B, G, 18587 (warm regions) G u z ik & S u d n ik -W ó jc ik o w sk a myr SB] Germ any [agr.] 1989; Z a j ą c A. & Z ając M . 20 0 1

Juncus tenuis Willd. Juncaceae HG w egz se. CSR Am N U l 1795* 1862 Sudety Mts.: near Zgorzelec 0 37 206 5332 1440 4-5 SH Czech Rep. Hulten 1958; Meusel e t al. [= J. m acer A. Gray] (Uechtritz herb. W) (+ D 1965; Hult£n & Fries 1986; Z a ją c A . & Z a ją c M . 2 0 0 1

Kochia scoparia (L.) Schrad. Chenopodiaceae T G w ane se. CR Eur E I [0 , C] X V III7 1872 Carpathian Foothills: Sokolniki 0 6 35 422 244 3(+D H Czech Rep. & H ulten 1971; anthr & Asia W - U l [G, 18117 (K napp 1872) Slovac Rep. [agr.] Tokarska-Guzik 2 0 0 1 b (33) W, BA] 18195

Lathyrus nissolia L. Fabaceae T G i aut se. CR Eur S U l 1903 1903 Wrocław and surroundings 0 0 13 18 16 1(-1) NS M eusel e t a l. 1965; & W 192110 (S chube 1903b) Z a ją c A . & Z a ją c M . 2 0 0 1

Lem na turionifera Landolt Lem naceae Hy G/V i hyd pi Am N Ul 19832 1994 east & north-east Poland 0 0 0 21 21 2 (+ l) NS W o l f f & L an d o lt 1 9 9 4 ; Z a ją c egz [AN, (W olff & Landolt 1994) A . & Z a ją c M . 2 0 0 1 BA]

Lepidium densiflorum Schrad. Brassicaceae T G i s ane sę. R Am N Ul 188310 1888 Mazury Lake District: between 0 4 67 1259 724 3-4 H M eu sel e t a l. 1965; H ulten & anthr [BS, W, 19045 Korpele and Sawica near (+D F ries 1986; G] Szczytno (Abromeit e t al. 1898) Tokarska-Guzik 2 0 0 1 b (34)

Lepidium virginicum L. Brassicaceae T G i s ane se. R Am N [E] Ul 169712 1860 Baltic Coast: Międzyzdroje 0 3 20 238 146 3(+ /-) H M eu sel e t a l. 1965; H ulten & anthr [W, BS, (H olzfuss 1937) F r ies 1986; G] Tokarska-Guzik 2 0 0 1 b (35)

Linaria repens (L.) Mili. Scrophulariaceae G G i ane se. CS Eur W Ul / 1 1825 1825 Gdańsk Westerplatte 1 3 7 31 26 2 (+ l) H M eu sel e t a l. 1978; H ulten & [= L . striata Lam. & DC.] (S chwarz 1967 after F ries 1986; K linsmann 1825) Wą so w ic z 2001, 2003 Nrs Nrs Nrs Nrs First First The oldest locality o f of o f o f Nrs Way record Name of species Fam ily LF RP Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Lolium multiflorum Lam. Poaceae H T G w ane se. C Eur S I [FD, C] 1837 1837 Sudety Mts.: Bolesławiec 1 11 37 2792 1174 4 (+ l) SH H u ltśn 1964; H u l tśn & F ries egz & W, 18835 (S chneider 1837) 1986; A fr N Z a j ą c A. & Z a j ą c M. 2001 & Asia SW

Lupinus polyphyllus Lindl. Fabaceae H G i aut se. C Am N 1 1877 1877 Silesian-Cracow Upland: Nie- 0 7 55 2674 1387 4 (+ l) NSH Czech Rep. Z a j ą c A. & Z a ją c M. 2001 [W] [0 , FO, 18955 poraz, Carpathian Foothills: Lithuania C] Lucjanowice (K rupa 1877)

Lycium barbarum L. Solanaceae N G/V i s end se. C Asia E, 1 [0 ] 1769 [I]25 1847 [I] [I] as omamental plant (Waga 0 54 80 2634 1224 4(+l) NSH Czech Rep. M e u se l et a l. 1978; [= L. halimifolium Mili.] rh. Eur SE 18392 25 1862 1847); West Pomerania: Świe- Z a j ą c A. & Z a j ą c M. 2001 18705 cie; Chełmno and sourrandings (herb. TRN Wacker 1862)

~~Lysimachia punctata L. Primulaceae H G/V i s aut se. C Eur SE 1 [0] 18195 1870 Grudziądz (A br o m eit et a l. 0 11 36 61 45 2 (+ l) SH T o k a r s k a -G u z ik 2001b (36) rh. 1926)~~

Malva moschata L. M alvaceae H G i s ane se. C Eur W I [0 ] A r10 XVIII XVIII - Kluk (1787); Mazowsze 0 12 102 286 196 ?(+/-) H M eu sel et a l. 1978; H ulten & egz - U l 1885 - Podlasie Lowlands: Płońsk F ries 1986; (Paczoski 1895) T o k a r s k a -G u z ik 2001b (37)

Marrubium vulgare L. Lam iaceae Ch H G i s egz se. CSR Eur S, I [M ] A r5' 10 XVI XVI Zając A. et a l. 1998; 7 147 255 453 315 3(-l) SH M eusel et a l. 1978; Hultśn & ane Asia SW - U l 1613* * - generał information (after Fries 1986; & A fr N 1643 Syreński 1613); Gdańsk 1643 T o k a r s k a -G u z ik 2001b (38) 1824 (Schwarz 1967 after Oelhaf); XVIII - Kluk (1787); Ma zowsze Lowland: Wyszogród (Zalewski 1892 after Gawa- recki 1824)

Medicago sativa L. s. str. Fabaceae H G i ane se. C A sia SW I [FD] XVI29 XVI ? Westerplatte (S chw a rz 1967 5 23 83 5412 1743 4-5 SH Z a j ą c A. & Z a j ą c M. 2001 [= M. s a t i v a L. subsp. s a tiv a ] egz [Cauc.] 18195 1832 after Klinsmann); south-westem (+D anthr 1837 Poland: Bytom Odrzański, Oława, Kup & Jedlina Zdrój (S c h n eid e r 1837)

~~Medicago varia M artyn H Fabaceae H G i ane se. C Anthro- I [FD] XIX15 1837 Wrocław (S chneider 1837) 1 60 86 1132 409 3-4 SH original: see App. C egz POg- - U l (+1) anthr~~

Melilotus wolgica Poir. in Lam. Fabaceae T H G i aut se. Eur E U l [G] 1937 1937 Szczecin Golęcino (H o lzfu ss 0 0 2 13 10 1(+1) H T o k a r s k a -G u z ik 2001b (39) [= M. v o l g ic u s Poir.] & Asia W 1937)

Mercurialis annua L. Euphorbiaceae T G w i aut se. R Eur SW U l 176716 XVI XVI Zając A. et a l. 1998; 2 44 94 143 87 2-3 H M eusel et a l. 1978; Hulten & myr [B, BA] A r5' 10 1825 XVIII - Kluk (1787); Gdańsk (-1) Fries 1986; (Rostański K. 1992) T o k a r s k a -G u z ik 2001b (40)

Mimulus guttatus DC. Scrophulariaceae H Hy G/V i ane se. CS Am N I [0 ] 18242 1824 Sudety Mts.: Kowary (Fiek 1 54 173 326 128 3(+2) NS M eusel et a l. 1987; Hultśn & hyd rh. [W] 18535 1881; ?? herb. WRSL) Fries 1986; see also P ię k o ś 1972; T o k a r s k a -G u z ik C hapter 7 2001b (41)

Mimulus moschatus Douglas ex Scrophulariaceae H G/V i ane se. CS Am N I [0 ] 18685 1879 Baltic Coast: Oliwa (Lutzów 0 3 10 13 11 1(+1) NS P ię k o ś 1972; T o k a r s k a -G u z ik Lindl. hyd rh. [W] herb. TRN) 2001b (42)

Myrrhis odorata (L.) Scop. Apiaceae H G i s egz se. C Eur C 1 [C, M] XVI [I]10 1837 Sudety Mts.: Bolesławiec, 3 10 68 119 76 2-3 NSH M eusel et a l. 1978; Hultśn & ane [Alps] 18095 Jedlina Zdrój, Kup (Schneider (+1) Fries 1986; 1837) Z a j ą c A. & Z a j ą c M. 2001

Oenothera acerviphila Rostański H O nagraceae H G i s ane se. Anthro- U l 1979 Silesia Upland: Brzezinka near 0 0 0 2 2 1 (?) H R o s t a ń s k i K. 2001b [probabl.= Oe. depressa x ammophila] aut POg- Mysłowice town (Rostański herb. KTU)

Oenothera canovirens E.S. Steele Onagraceae H G i s ane se. CR Am N 190751 1958 Wrocław (Rostański herb. 0 0 0 42 26 2 (+ l) H R o s t a ń s k i K.& T o k a r s k a -G u z ik [= Oe. renneri H. Scholz] aut 19535 KTU) 1998, 2001

Oenothera cruciata Nutt. ex G. Don Onagraceae H G i s ane se. Am N I [B] 1826 [I]30 1905 West Pomerania: Trzcianka 0 0 1 2 2 1(-1) H R o s t a ń s k i K.& T o k a r s k a -G u z ik [= Oe. atrovirens auct. Europ.] aut [E] 190531 (Bothe herb. B) 2001 Nrs Nrs Nrs Nrs First First The oldest locality o f o f o f o f Nrs Way record Name of species Family LFRP Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europę" & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Oenothera depressa Greene O nagraceae H G i s ane se. Am N I [B] X IX 2 1894 Warszawa (Cybulski herb. WA] 0 1 3 643 274 3 (+ l) SH R o s t a ń s k i K.& T o k a r s k a -G u z ik [= Oe. salicifolia Desf. ex G. Don aut - U l 183551 1998, 2001 Oe. hungarica Borbas] 19365

Oenothera fallax Renner em. Ros- O nagraceae H G i s ane se. Anthro- I [B] 191751 1958 Wrocław (Rostański herb. 0 0 0 11 9 1 (+ /-) H R o s t a ń s k i K.& T o k a r s k a -G u z ik tański H aut POg- - U l 1958 KTU) 1998, 2001 [probabl.= Oe. glazioviana x biennis] 196 5 196732

Oenothera flaemingina H udziok H O nagraceae H G i s ane se. Anthro- U l 196863 1994 Silesian Upland: Strzyżowice 0 0 0 31 31 2 (+ l) H R o st a ń sk i K. & Wit o sł a w sk i 2001 [probably a hybrid orginated in Central aut POg- (N ow ak herb. KTU) Germany]

Oenothera glazioviana M icheli in Onagraceae H G i s ane se. CR Am N I [B] X IX 2 1879 Silesian Lowland: Sułków 0 1 6 29 23 2 (+ l) H R o s t a ń s k i K. 2001 b M art. aut - U l 186455 (Sintesis herb. W RSL) [= Oe. erythrosepala Borbas] 186630 18905

Oenothera hoelscheri Renner Onagraceae H G i s ane se. Anthro- U l 1942 1942 Włocławek upon Vistula river 0 0 0 397 171 3 (+ l) SH R o s t a ń s k i K . & T o k a r s k a -G u z ik ex Rostański H aut P°g- 197032 (R enner 1942) 1998, 2001 [probabl.= Oe. biennis (or) 19755 Oe. rubricaulis x depressa]

Oenothera issleri Renner ex Rostań O nagraceae H G i s ane se. Anthro- U l 19495 1958 Wrocław (Rostański herb. 0 0 0 7 5 K + /-) H R o s t a ń s k i K . & T o k a r s k a -G u z ik ski H aut POg- KTU) 1998, 2001 [probabl.=Oe. biennis x oakesiana]

Oenothera jueterbogensis H udziok Onagraceae H G i s ane se. Anthro- U l 19625' 1973 Silesian Upland: Gliwice (Ros 0 0 0 6 4 1(+1) H R o s t a ń s k i K . & T o k a r s k a -G u z ik [probabl.= Oe. biennis x ?] H aut P°g- tański K. & Szotkowski 1973) 1998, 2001

Oenothera oakesiana (A. Gray) Onagraceae H G i s ane se. Am N 1 [B] 1614 [I]30'32 1962 Wrocław (Rostański herb. 0 0 0 36 23 2 (+ l) H R o s t a ń s k i K . & T o k a r s k a -G u z ik J.W. Robbins ex S. Watson aut 19625 KTU); Mazowsze - Podlasie 1998, 2001 [= Oe. syrticola Bartlett] Lowlands: Wygoda near Janów Podlaski (Fijałkowski herb. LBL)

Oenothera paradoxa Hudziok H Onagraceae H G i s ane se. Anthro- Ul 196751 1974 Silesian Upland: Gliwice & 0 0 0 218 64 2-3 H R o s t a ń s k i K . & T o k a r s k a -G u z ik [probabl. = Oe. depressa x subter- aut P°g. Katowice (Celiński et al. 1974); (+1) 1998. 2001 minalis] Katowice & Siemianowice Śląskie (Michalak & Sendek 1974 -1975)

Oenothera parviflora L. Onagraceae H T G i s ane se. CR Am N Ul 1682 [I]55 1938 W ałbrzych (R enner 1938) 0 1 2 27 16 1-2 H M eusel et al. 1978; aut 1768 [I]30 (+1) R o s t a ń s k i K . & T o k a r sk a -G u z ik 1914 5 1998, 2001 Oenothera pseudochicaginensis Onagraceae H G i s ane se. Anthro- Ul 1959 1959 Wrocław (Rostański herb. 0 0 0 7 6 !(+ /-) H R o s t a ń s k i K. & T o k a r s k a -G u z ik R ostański H aut P°g- KTU) 1998. 2001 [probabl. = Oe. subterminalis x biennis]

Oenothera punctulata R ostański Onagraceae H G i s ane se. Anthro- Ul 196910 1973 Silesian Lowland: Nysa, Sile 0 0 0 9 8 1(+/-) H R o s t a ń s k i K. & T o k a r s k a -G u z ik & G utte H aut POg- 19725 sian Upland: Gliwice (R os 1998, 2001 [probabl.= Oe. pycnocarpa x biennis] tański K. & Szotkowski 1973)

Oenothera pycnocarpa Atk & Bartl. O nagraceae T H G i s ane se. Am N U l 195832 1963 Baltic Coast: Glinna near Gryfin 0 0 0 50 35 2 (+ l) H R o s t a ń s k i K. & T o k a r s k a -G u z ik in Bartl. aut 19605 (Rostański K. & Tokarska- 1998, 2001 [= Oe. chicaginensis De Vries ex -G uzik 1998) Renner]

Oenothera royfraseri R.R. Gates O nagraceae H G i s ane se. Am N Ul 1963 1963 Sudety Mts.: Turoszów 0 0 0 22 14 1-2 H R o s t a ń s k i K. & T o k a r s k a -G u z ik [= O e. turoviensis Rostański] aut 196910 (Rostański herb. KTU) (+/-) 1998, 2001

~~Oenothera suaveolens Desf. ex Pers. O nagraceae H G i s ane se. Eur S ? Ul 180551 1961 Wrocław, Brzózka Krośnieńska 0 0 0 6 6 1(?) H R o s t a ń s k i K. 2001b aut (Rostański herb. KTU)~~

Oenothera subterminalis R.R. Gates O nagraceae H G i s ane se. Am N Ul 185633 1938 Silesian Lowland: Nowogród 0 0 1 220 91 2-3 H M eusel et al. 1978; [= Oe. silesiaca Renner] aut Bobrzański (R enner 1938) (+ D R o s t a ń s k i K. & K l o s s 1965; R o s t a ń s k i K. & T o k a r s k a -G u z ik 1998, 2001 Nrs Nrs Nrs Nrs First First The oldest locality o f o f o f o f Nrs Way record Name of.species Family LF R P Disp Prop LS O rigin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere M aps o f INT for for Europę’ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Oenothera victorini R.R. Gates O nagraceae Н G i s ane se. Am N Ul 1961 1961 Silesian Lowland: Nysa 0 0 0 49 22 2(+ /-) H R o stań sk i K. & T o k arsk a-G u zik & Catches. in R.R. Gates aut 196732 (Rostański K. 1965) 1 9 9 8 , 2 0 0 1 [= Oe. nissensis Rostański] 19735

Oenothera wienii Renner ex Rostański Onagraceae HG i s ane se. Anthro- Ul 1937 1937 Gdańsk-Stogi (R enner 1937) 0 0 0 116 74 2-3 H R o stań sk i K. & T o kar sk a-G uzik [probabl.= Oe. rubricaulis x aut pog. (+ D 1 9 9 8 , 2 0 0 1 d ep ressa] H

Onobrychis yiciifolia Scop. Fabaceae HG i egz se. C Eur S 1 [FD] XVI [I]10 1837 south-western Poland: Bytom 3 140 323 911 452 3 (+ l) NSH Z a ją c A . & Z a ją c M . 2 0 0 1 [= 0. viciaefolia Scop.] & SE 1837 Odrzański, Oława, Wołów 18525 (Schneider 1837)

Ornithogalum boucheanum Asch. Liliaceae G G i ane se. CSR Eur SE I [0 ] ca . X V I[I]'° 1880 Silesian Lowland: Głogówek 0 3 24 36 29 2 ( - l) SH To k a r sk a -G u zik 2 0 0 1 b (43) m yr ro. (R ichter h e rb . MGS)

Oxalis corniculata L. Oxalidaceae T H G/V i s aut se. R Eur S Ul 157634 1863 Sudety Mts.: Zgorzelec 0 25 42 128 84 2-3 H H ungary Hulten 1971; Meusel e t a l. anthr rh. A sia SW [GA, B] 18525 (H antz 1979) (+1) 1978; Hulten & Fries 1986; H a n t z 1 9 7 9 ; T o k a r sk a -G uzik 2 0 0 1 b (44)

Oxalis dillenii Jacq. Oxalidaceae T H G i s aut se. R A m N Ul [B] 186534 1865 Silesian Lowland: Wrocław 0 1 2 40 31 2 (+ l) H H a n t z 1 9 7 9 ; To k a r sk a -G u zik anthr [E] ( - h e rb . W RSL after Hantz 2 0 0 1 b (45) 1979)

Oxalis fontann Bunge Oxalidaceae G G/V i s aut se. R A m N Ul 165816' 34 1809 Kraków (T rzcińska-T acik 8 111 181 8806 2141 5 (+ l) H M eusel e t a l. 1978; Hulten & [= O . s tr ic ta L.] anthr rh. [E], [P, GA, 182634 1979) Fries 1986; A sia E ? B] 18525 H a n t z 1 9 7 9 ; Z a ją c A . & Z a ją c M . 2 0 0 1

Oxybaphus nyctagineus (M ichx.) Nyctaginaceae G G/V i ane se. Am N Ul / I 18435'7 1911 Wielkopolska Lake District: 0 0 1 6 6 1(?) H Czech Rep. [agr.], C eyn o w a -G ie ł d o n 1 9 8 8 ; Sweet [C] G ubin (D ecker 1911) H ungary T o k a r sk a -G u zik 2 0 0 1 b (46)

Padus serotina (Ehrh.) Borkh. Rosaceae N M G i s end fr. C Am N I 1623 [I]35 1813 [I] Niedźwiedź [I] (H ereźniak 0 1 10 2564 1134 4(+2) NS Eur C [forests] Z a ją c A . & Z a ją c M . 2 0 0 1 [= Prunus serotina Ehrh.] [E] & [0 , FO] 182525' 35 1880 ? 1992); Warszawa (Sudnik-W ój- Am S see also 1900 cikowska 1987a); Bydgoszcz [N] Chapter 7 (B ock 1908)

Parietaria pensyhanica Muhl. U rticaceae TG w ane se. CR Am N Ul 1810 1991 Bydgoszcz (Misiewicz e t a l. 0 0 2 2 2 1(+1) H Z a ją c A . & Z a ją c M . 2 0 0 1 ; ex Willd. [B. GA] 1820 [I]49 1996) G u zik 2 0 0 2 186150 20005

Parthenocissus inserta (A. Kern.) Vitaceae N li G/V i end fr. C Am N I [0] 1629 [I]1 1806 [I] Kraków - botanical garden [I] 0 1 3 558 332 3(+2) NSH some part of Z a ją c A . & Z a jąc M . 2 0 0 1 Fritsch anthr rh. [E] 18842' 25 1884 (H e r eź n ia k 4 9 9 2 ) ; Carpathian Eur C [= P. v ita c e a (Knerr) Hitchc.] st. 19005 Foothills: Tenczyn (R a cibo rsk i 1884)

Petrorhagia saxifraga (L.) Link Caryophyllaceae Ch G i s ane se. CS Eur S U l / I 1859 Kraków (B erdau 1859) 0 5 9 43 36 2 (+ l) N H M e u se l e t a l. 1965; J a l a s & [= Tunica saxifraga (L.) Scop.] egz & SE [0] S u o m in e n 1986; Z a ją c A . & Z a jąc M . 2 0 0 1

Phleum rhaeticum (Humphries) Poaceae H G w ane se. Eur C Ul [AN] XIX / West Tatra Mts.: Stoły 0 0 0 1 1 l(?) N Z a ją c A . & Z a ją c M . 2 0 0 1 Rauschert egz [Alps] XX Clearing (M irek 1995) 1995

Physalis alkekengi L. Solanaceae H G/V i s ane se. C Eur SE I [0 , M] 1866 1613* * - generał information (after 0 4 19 397 286 3 (+ l) NSH M eusel et al. 1978; end fr. & Asia 186710 1866 Syreński 1613); Warszawa T o k a r sk a-G u zik 2 0 0 1 b (47) aut rh. SW A r5 (Karo herb. W U)

Picris echioides L. A steraceae T G i s ane se. CSR Eur S Ul[BA] 1836 XVIII XVIII - Kluk (1787); Gdańsk 0 1 33 60 37 2 (+ /-) NH M eusel e t a l. 1992; [= Helminthia echioides (L.) Gaertn.; egz & A fr N 18615 1836 (Schwarz 1967 after Klinsmann T o k a r sk a-G u zik 2 0 0 1 b (48) Helminthotheca echioides (L.) m yr 187810 h e rb .) Hołub]

Plantago serpentina Ali. Plantaginaceae H G i s ane se. Eur C U l [AN] XIX / West Tatra Mts.: Stoły 0 0 0 1 1 1(?) N Z a ją c A . & Z a ją c M . 2 0 0 1 egz [Alps] XX Clearing (M irek 1995) 1995 Nrs N rs Nrs Nrs First First The oldest locality o f o f o f o f Nrs Way record Name of species Fam ily LFR P Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000 Port ul аса oleracea L. Portulacaceae T G s aut se. R A sia S I [A] Ar2'5 1613* * - generał information (after 4 36 94 216 147 3 (+ l) H some part of Hulten 1971; Jalas & Suominen myr & A fr N XVIII Syreński 1613); XVIII - K luk Eur S 1980; Hulten & Fries 1986; 1837 ( 1787); south-western Poland: T okarska-G uzik 2 0 0 l b (49) Wrocław & Bolesławiec (Schneider 1837)

Potentilla intermedia L. non W ahlenb Rosaceae H G i ane se. CSR Eur NE Ul 1652 ? 1652 ? Warszawa (after Sudnik-W ójci- 1 17 41 207 102 3 (+ l) H Hulten & Fries 1986; myr & Asia N [G, A] 1841 XVIII / kowska 1987a); XVIII/XIX Kor- Z a ją c A . & Z a ją c M . 2 0 0 1 end 189610 XIX naś 1968b; Gdańsk (S chwarz 19035 1841 1961)

Reseda luteola L. Resedaceae H G i s myr se. CS Eur S, I [C] A r2' 5 XVIII XV III - K luk (1788); Gdańsk 5 62 87 299 182 3 (+ l) H M eusel e t a l. 1965; JAger ane A sia W - U l 1825 (R eyger 1825) 1970; Hulten & Fries 1986; T o k arsk a-G u zik 2 0 0 1 b (50)

Reynoutria japonica (Houtt.) Ronse Polygonaceae G V/G? w ane rh. С A sia E I 1823- 1882 Wielkopolska Lake District: 0 3 63 3004 1158* 4(+2) NSH Eur W & С Jalas & Suominen 1979; C hild Decraene var. japon ica i egz st. [B, 0 , C] 1829[I]37 Gniezno (Cybichowski h e rb . [rip. & urb.]; & W ade 1999, 2000; [= Fallopia japonica Houtt.] s myr (se.) 188638 POZ) A m N Z a ją c A . & Z a jąc M . 2 0 0 1 hyd 18925 anthr see also Chapter 7

Reynoutria sachalinensis (F. Schmidt) Polygonaceae G V/G? w ane rh. С Asia E I before 1903 Sudety Mts.: Szklarska Poręba 0 0 16 474 282* 3 (+ l) NSH Jalas & Suominen 1979; Nakai i egz st. [B, 0 , C] 1864[1]37 (S chube 1903b) T o k arsk a-G u zik 2 0 0 1 b (51) [= Fallopia sachalinensis (F. Schmidt s m yr (se.) 18695' 37 et Maxim) Ronse Decraene] hyd anthr

Robinia pseudoacacia L. Fabaceae M G/V i end se. С Am N [E I 1601 [I]1 XVIII [I] XVIII - Kluk (1788) - only as 1 12 39 7067 1957 4-5 NSH some regions of Z a ją c A . & Z a ją c M . 2 0 0 1 ane ro. [0 , M, C] 182425 1806 [I] cultivated plant; Kraków - (+2) Eur C; anthr 18745 1836 botanical garden [I] (Hereźniak Lithuania 1868 1992); Gdańsk-Stogi (Schwarz 1967 after Klinsmann); Mazow sze - Podlasie Lowlands: Tucho- wicz near Łuków (Łapczyński h erb . LBL)

Rosa rugosa Thunb. Rosaceae N G/V i end se. С Asia E I [C] 1841 [I]25 1913 ? Mazury Lake District: 0 0 8 1299 701* 3 -4 NSH Z a ją c A . & Z a ją c M . 2 0 0 1 s anthr fr. 19505 K rzem ity (F uhrer 1913) (+D a ro. 196025

~~Rubus allegheniensis Porter Rosaceae N G/V i end fr. Am N I [C] 189010 1899 Wrocław Zalesie 0 1 1 9 9 K?) NSH Z ie l iń sk i 2 0 0 1 , 2 0 0 4 [E] (Baenitz h e rb . LE)~~

Rubus armeniacus Focke Rosaceae N G/V i end fr. A sia SW I [C] 186040 1843 ? Skarszyn (source: ATPOL); 1 1 2 68 58 2 (+ l) SH Z ie l iń sk i 1991, 2001 , 2 0 0 4 [Cauc.?] 1902 Szczecin (Holzfuss h e rb . PR)

Rubus canadensis L. Rosaceae N G/V i end fr. Am N [E] I 1727[I]' 1811 [I] Krzemieniec - botanical garden 0 0 0 6 6 1(?)NSH Z ie l iń sk i 2001, 2004 196710 1967 [I] (Hereźniak 1992); Parko- szów (Ciaciura h e rb . SZUB)

Rubus laciniatus Willd. Rosaceae N G/V i end fr. Anthro- I [0] 1770 [I]39 1859 Nysa (source: ATPOL); Wro 0 1 2 16 13 K+l) NSH Z ie l iń sk i 1991, 2001, 2004 POg- 188525 1905 cław (Baenitz h e rb . LE)

Rubus odoratus L. R osaceae N G/V i end fr. Am N I [0 ] 1635 [I]' 1806 [I] Kraków - botanical garden 0 4 8 12 11 1(?) NSH Z ieliń sk i 2001, 2004 [E] 18805 1877 [I] (Hereźniak 1992); Książ 1890'° (W acker h e rb . TRN)

~~Rubus xanthocarpus Bureau & Franch R osaceae H G i end fr. Asia E I [O] 19625 1991 Miedzianka (Bróż h e rb . 0 0 0 1 1 K?) H Z ie l iń sk i 2001, 2004 [China] KOR & KTC)~~

Rudbeckia laciniata L. Asteraceae H G G/V a ane se. С Am N I [0] 1615 [I]41 1787 Sudety Mts.: Świeradów (F iek 3 78 187 2251 903 3-4 NSH Czech Rep. M e u s e l e t al. 1992; i egz ro. [E] 178741 1881 after Krocker); Lubań (+ 2 ) & Slovak Rep. Z a ją c A . & Z a ją c M . 2 0 0 1 s myr see also (Jalas 1993) aut Chapter 7

Rumex confertus W illd. Polygonaceae H G w s ane se. Eur SE Ul 1873 1873 Mazowsze - Podlasie Lowlands: 0 4 47 1731 673 3-4 SH Lithuania Jalas & Suominen 1979; egz & Asia W see also Zajęczniki & Łosice (Karo herb. (+2) Trzc iń sk a-T ac ik 1 9 6 3 ; Z a ją c A. hyd Chapter 7 KRA) & Z a ją c M . 2 0 0 1 Nrs Nrs Nrs N rs First First The oldest locality o f o f o f o f Nrs Way record Name of species Family LF R P Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Salsola kali L. Chenopodiaceae T G w i ane se. SR Eur SE Ul 1730 XVII XVII K ornaś 1968b; Gdańsk 2 26 114 901 467 3-4 H M eusel et al. 1965 s.L; H u lten subsp. ruthenica (Iljin) Soó & Asia C [W, BA] 177510 1643 (Schwarz 1967 after Oelhaf); ( + D & Fries 1986; see also 1730 W arszawa (Sudnik-W ójcikow- B a ra d zie j 1 9 7 2 \ To karska-G u zik C hapter 7 ska 1987a after Erndtel) 2 0 0 l b (52)

Sedum album L. Crassulaceae Ch G/V i s ane se. S Eur S I [0] XVII [I] XVIII - Kluk (1788)- only as 0 5 15 59 47 2 (+ l) SH M eusel et al. 1965; Hulten hyd st. & W , 1868 cultivated plant; Sudety Mts.: & Fries 1986 aut rh. A fr N Jordanów Śląski & Mierczyce m yr & Asia W (W lM M ER 1868)

Sedum spurium M. Bieb. Crassulaceae Ch G/V i s ane se. S A sia SW I [0] 18795 1880 Silesian Upland: Marcinkowice 0 15 62 301 230 3(+D H Z a ją c A . & Z a ją c M . 2 0 0 1 hyd st. [Cauc.] (U echtritz 1880) aut rh. m yr

Senecio vernalis Waldst. & Kit. Asteraceae T H G i s ane se. R Eur SE Ul 172614 1824 W arszawa (R ostafiński 1872) U 119 219 3932 1948 4-5 H Hulten & Fries 1986; Hegi egz & Asia W 183057 (+2) 1987; Meusel et al. 1992; m yr 185114 Z a ją c A . & Z a ją c M . 2 0 0 1

Sicyos angulata L. Cucurbitaceae T G i end fr. Am S I [0 ] / 1868 1868 Carpathian Foothills: Krosno 0 13 18 168 101 2-3 H original: see App. C U l [SB] 18805 (K napp 1868) (+D Silene conica L. Caryophyllaceae T G i s ane se. SR Eur S U l 1879 1879 Wielkopolska Lake District: 0 23 76 199 104 2-3 H M eu sel et al. 1965; Jalas & aut & Asia 189210 Czerwieńsk (U echtritz 1879) (+D Suominen 1986; Hulten & SW Fries 1986; Z a ją c A . & Z a ją c M . 2 0 0 1

Silene dichotoma Ehrh. Caryophyllaceae H G i s ane se. R Eur S U l [BS] 18415 1877 Wrocław (U echtritz 1877) 0 30 289 496 335 3 (+ l) H J alas & Suominen 1986; aut & SE, H ult£n & F ries 1986; A sia SW Z a ją c A . & Z a ją c M . 2 0 0 1

Sinapis alba L. Brassicaceae T G i s ane se. CR Eur S I [C, M, XVII [I] XVIII-K luk (1788)-only as 0 18 55 1416 716 3-4 H Hulten & Fries 1986; FD] 1824 cultivated plant; Mazowsze ( + 0 Z a ją c A . & Z a ją c M . 2 0 0 1 Lowland: Wyszogród (Zalewski 1892)

Sisymbrium altissimum L. Brassicaceae H T G i s aut se. CR Eur SE Ul 1780'2 1843 Gdańsk (K l in s m a n n 1843) 2 24 59 1770 812 3 -4 H M eu sel et al. 1965; Hultźn hyd pl. & Asia C [G, BA] 1815* (+D 1971; Hulten & Fries 1986; see also Z a ją c A . & Z a ją c M . 2 0 0 1 C hapter 7

Sisymbrium loeselii L. Brassicaceae H T G i s aut se. CR Eur SE Ul 165412' 15 1654 Gdańsk (Hegi 1935-1961); War 2 31 87 2326 976 3-4 H Czech Rep. M eusel et al. 1965; Hulten & ane & Asia C [BA, G] 1819 5 1824 ? szawa (Sudnik-Wójcikowska (+D Fries 1986; end 1847 1987a); Gdańsk (Schwarz 1967); Z a ją c A . & Z a ją c M . 2 0 0 1 1856 Warta river embancment near Poznań (Lechmann herb. POZ)

Sisymbrium wolgense M. Bieb. Brassicaceae H G/V i s aut se. C Eur SE Ul [RW] 18807 1896 Warszawa (Cybulski herb. WA) 0 1 2 62 40 2 (+ l) H Czech Rep. Z a ją c A . & Z a jąc M . 2 0 0 1 ex E. Fourn. ane ro. Finland [agr.]

Sisyrinchium bermudianu L. em. Farw. Iridaceae H G/V i ane se. CSR Am N I [0] 18353 1928 Sudety Mts.: Jelenia Góra & 0 0 5 22 17 1-2 S Hultśn 1958; [= S. angustifolium Mili.] ro. [E] 1845 ?8 Jeleniec Mały (Schube 1928) (+D original: see App. C 18635

Soli dago canadensis L. A steraceae G H G/V i s ane se. C Am N I [0] 164814 1872 Lublin Upland: Lublin (R os 0 20 60 3436 1254 4(+2) NSH some regions of M eusel et al. 1992; egz rh. [E] 17362 tafiński 1872); Rząska near Eur C G u ziko w a & M a yc o c k 1 9 8 6 ; m yr 183 85 Kraków (K napp 1872) Z a ją c A . & Z a ją c M . 2 0 0 1

Solidago gigantea Aiton. A steraceae G H G/V i s ane se. c Am N 1 [0 ] 175814 1853 Wrocław (Uechtritz herb. 0 40 150 5350 1668 4-5 NSH some regions of M eusel et al. 1992; [= S. serotina Aiton] egz rh. 183014 WRSL) (+2) Eur C G u ziko w a & M a yc o c k 1 9 8 6 ; m yr 18515 Z a ją c A . & Z a ją c M . 2 0 0 1

Solidago graminifolia (L.) Elliott Asteraceae G H G/V i s ane se. c Am N I [O] 175860 1888 Silesian Lowland: Lipno near 0 2 5 46 27 2(+D NSH T o k a r sk a -G u zik 2 0 0 1 b (53) egz rh. [N] XIX2 Niemodlin (Zeidel? herb. myr WRSL) Nrs Nrs N rs N rs First First The oldest locality o f o f o f o f Nrs Way record Name of species Fam ily LF R P , Disp Prop LS Origin record in Poland loc. loc. loc. loc. o f Dyn Hab. Inv. elsewhere Maps o f INT for for Europe+ & source of data up to up to up to up to sq. Poland 1850 1900 1950 2000

Tanacetum parthenium (L.) Sch. Bip. A steraceae H G i s ane se. CSR Eur SE I [O, M] 15612 XVI ? XVI Zając A. et al. 1998; 1 56 91 1179 734 3 -4 H M eusel et al. 1992; [= Chrysanthemum parthenium (L .) egz & Asia 176914 1613* * - generał information (after (+D T o k a r sk a-G uzik 2 0 0 1 b (54) Bernh.] m yr SW A r5' 10 1824 Syreński 1613); XVIII - K luk 1787; Mazowsze Lowland: W yszogród (Z alewski 1892 after Gawarecki 1824)

~~Thladiantha dubia Bunge Cucurbitaceae H V/G i end se. A sia E I [O] 1917 1917 south-eastern Poland: Turka 0 0 5 69 462(+l) SH Z a ją c A . & Z a jąc M . 2 0 0 1 fr. 1939* (K oporska herb. LBL) st.~~

Trifolium patens Schreb. Fabaceae H G i ane se. Eur S Ul [FD] 1933 Carpathian Mts.: Wróblik Szla 0 0 1 227 54 2-3 N H H endrych 1966; egz checki near Rymanów (P iech (+D L o ste r 1 9 7 2 ; T o k a r sk a -G u zik anthr 1939) 2 0 0 1 b (55) Yeronica filiformis Sm. Scrophulariaceae Ch H V/G i s ane st. CSR Asia SW I [B, 0 ] 1780 [I]7 1936 [I]? Baltic Coast: Sopot 0 0 2 161 69 2-3 SH Czech Rep. M eusel et al. 1978; Hulten & hyd se.? [Cauc.] 18387 (L uttschwager 1936) (+D [maedows] Fries 1986; m yr UK USA P ie t r a s 1 9 7 0 ; Z a ją c A . & Z a ją c aut 19385 M . 2 0 0 1 anthr Veronieci peregrina L. Scrophulariaceae T G i s ane se. R Am N Ul [GA] 17602 1854 Kraków Sikomik (- herb. KRA) 0 2 2 21 16 1-2 NSH Hulten 1971; Meusel et al. hyd 18095 (+1) 1978; Hulten & Fries 1986; m yr Z a ją c M . & Z a ją c A . 1 9 9 0 ; T o k a r sk a-G uzik 2 0 0 1 b (56) leronica persica Poir. Scrophulariaceae T G i s ane se. CR Asia SW Ul [G] 180516 1862 West Pomerania: Chełmno; Swie- 0 33 84 7887 2204 5(+2) H Czech Rep. Hulten 1971; Meusel et al. hyd [Cauc.] 18095 cie town sourrandings (Abro- 1978; Hulten & Fries 1986; m yr see also meit et al. 1898 after Wacker Z a ją c A . & Z a ją c M . 2 0 0 1 anthr C hapter 7 1862)

Vicia dasycarpa Ten. Fabaceae T G i aut se. CR Eur S Ul 1898 Toruń (Abromeit et al. 1898) 0 1 2 1302 384 3-4 H Z a ją c A . & Z a jąc M . 2 0 0 1 anthr (+D Yicia grandiflora Scop. Fabaceae TG i aut se. CR Eur S Ul 18775 1907 Silesian Lowland: Kościeżyce 0 0 18 1540 506 3-4 SH Hanelt & Mettin 1970; anthr & Asia & Czepielowice near Brzeg (+2) T o k arsk a-G u zik 2 0 0 1 b (57) SW (S chube 1907)

Vicia pannonica Crantz Fabaceae T G i aut se. CR Eur SE U l / I 1884 1884 Silesian Lowland: Głuchołazy 0 1 49 91 68 2 (+ l) SH T o k a r sk a -G uzik 2 0 0 I b (58) anthr [Pan.] 1893'° (R ichter herb. MGS) A r5

Kanthium albinum (Widder) H. Scholz Asteraceae T G w s egz fr. CR Am N Ul 182242 1853 N ow a Sól (F iek 1881 after 0 40 83 1119 471 3^1 SH some parts of M eusel et al. 1992; [= X. riparium Itzigs. & Hertsch] hyd [S] Franke) (+1) Eur C & S Z a ją c A . & Z ają c M . 2 0 0 1 Xanthium spinosum L. Asteraceae T G w s egz fr. CR Am S Ul 168114 1849 Wrocław (Uechtritz herb. 1 79 129 294 148 3(+/-) H some parts M eusel et al. 1992; hyd [W, SB] 18725 WRSL) of Eur C & S; T o k a r sk a -G u zik 2 0 0 1 b (59) different parts of the w orld A'anthium strumarium L. A steraceae T G w s egz fr. CR Eur / Ul A r5' 10 1613* * - generał information (after 3 130 225 1105 712 3-4 H some parts M eusel et al. 1992; hyd Am N ? [W, SB] 1837 Syreński 1613); Silesian (+ /-) of Eur C & S; Z a ją c A . & Z a ją c M . 2 0 0 1 Lowland: Oława (S chneider Australia; 1837) India; Africa S the Americas A ppendix B. L ist o f Polish kenophytes together with their ecological-geographical characteristic (excluding history o f distribution) This appendix includes 51 species more often cultivated and considered established in some regions of Poland (the species concerned were marked with ”?” preceding the species name).

First First N rs Way Source of historical Described as invasive Name of species Fam ily LF R P Disp LS O rigin record record o f Hab. Dyn M aps of INT data for Poland elsew here for Europę for Poland sq.

~~? Acer saccharinum L. Aceraceae M G i w ane Am N [E] 1 [O] 1725 [I]1 1807 [I] H ereźniak 1992 n.c.d. H 1(+1) XX~~

~~Achillea crithmifolia Waldst. & Kit. A steraceae H G i ane C/CS Eur SE Ul 1886* 2/2 XX Z ając A. et al. 1998 n.c.d. S 1(?) end ?~~

~~? Aesculus flava Sol. ex Hope Flippocastanaceae M G i bar A m N [E] 1 [O] 1764 [I]1 1813 [I] H ereźniak 1992 n.c.d. N 1(?)~~

Aesculus hippocastanum L. Hippocastanaceae M G i bar c Eur SE [m] I [0, M] 1576 [I]45 XVII Adamowski et al. 2002; 620* SH 3 (+ l) anthr 178725 Z ając A. et al. 1998; XVIII - Kluk (1786) - only as cultivated plant

~~Alchemilla rigida Buser Rosaceae H G a egz CS Eur C U l [AN] XIX M irek et al. 2002 6 s 1(?) [ Alps]~~

Alnus rugosa (Du Roi) Spreng. Betulaceae N G w ane c Am N [E] I [0,C ] 1769[I]1 1817 [I] H ereźniak 1992 n.c.d. N 2(?) 18725 1/2 XX ? Amelanchier spicata G.N. Jones R osaceae N G/V i end c Am N [NE] I [O] 1783 [I]' 1820 [I] Hereźniak 1992; n.c.d. SH 2 (+ l) XIX Z ając A. et al. 1998 ? Amorpha fruticosa L. Fabaceae N G i egz c Am N [E & C] I [0 , C] 1724 [I]1 1807 [I] H ereźniak 1992 n.c.d. H 2 (+ l) some regions of Eur C anthr 190743 19325 ? Anaphalis margaritacea (L.) Benth. A steraceae H G/V i ane Am N 1 [0] 18875 XX Z ając A. et al. 1998 n.c.d. SH 1(+1) [= Gnaphalis margaritacea L.] egz anthr ? Aronia melanocarpa (Michx) Elliot R osaceae N G a end c Am N [NE] I [0 , F] ca. 1824 [I] H ereźniak 1992 n.c.d. NSH 1(?) anthr 1688 [I]1

? Aronia prunifolia (Marshall) Rehder H R osaceae N G a end Am N [NE] I [0 , F] 1800 [I]1 1833 [I] H ereźniak 1992 n.c.d. NSH 2(+l) [= A. arbutifolia (L.) Pers. x anthr A. melanocarpa (Michx.) Elliott] Aster lanceolatus Willd. A steraceae H G/V i s ane c Am N [E] I [0] X IX 2 XIX Z ając A. et al. 1998 260* SH 3(+l) Czech Rep. & Hungary M eusel et al. 1992 egz anthr Aster novae-angliae L. A steraceae H G/V i s ane c Am N [E] I [O] XIX2 XIX/XX Zając A. et al. 1998 155* SH 3(+ l) egz anthr Aster novi-belgii L. Asteraceae H G/V i s ane c Am N [E] I [0] X V III2 XVIII Zając A. et al. 1998 353* SH 3(+ l) Czech Rep. M eusel et al. 1992 egz 18505 anthr Aster salignus W illd. H Asteraceae H G/V i s ane c Am N I [O] 17872 XIX Zając A. et al. 1998 139* SH 3 (+ l) Czech Rep. & Hungary M eusel et al. 1992 egz anthr Aster tradescantii L. Asteraceae H G/V i s ane c Am N [E] I [0 ] 17362 XIX Z ając A. et al. 1998 94* H 2 (+ l) M eusel et al. 1992 egz anthr Atriple.y hortensis L. Chenopodiaceae T G s i ane CR A sia C, 1 [0 ] 18725 X V III [I] [I] - K luk (1786) - only n.c.d. H 2 (+ l) hyd Eur ? XIX as cultivated plant; anthr Z ając A. et al. 1998 Atriple.x prostrata Boucher ex DC. C henopodiaceae T G s i ane Eur E Ul 1/2 XX Z ając A. et al. 1998 n.c.d. H 1(?) subsp. polonica (Zapal.) Uotila

~~Brachyactis ciliata (Ledeb.) Ledeb. Asteraceae T G i s ane Asia Ul 1967 2000 Guzik unpubl. 2 H 1(?) egz~~

Brassica elongata Ehrh. Brassicaceae C T G i s ane Eur E Ul XIX ? XIX ? Zając A. et al. 1998 n.c.d. N 1(?) Z a ją c A. & Z a ją c M. 2001 subsp. integrifolia (Boiss.) Breistr. aut & A sia W 19605 Brassica nigra (L.) W.D.J. Koch Brassicaceae T G i s ane CR Eur SW & W I [A]-UI XVI ?2 XVI Z ając A. et al. 1998 286 SH 3 (+ l) Czech Rep. Meusel et al. 1965; aut A r Hultśn & Fries 1986 anthr First First Nrs Way Source of historical Described as invasive Name of species Family LFRP Disp LS O rigin record record o f Hab. Dyn M aps o f INT data for Poland elsew here for Europę for Poland sq. Brassica rapa L. (L.) W.D.J. Koch subsp. rapa Brassicaceae T G i s ane Anthropog. I [A] Ar ? XVI Z ając A. et al. 1998 n.c.d. H 2 (?) aut anthr

~~Brassica rapa subsp. sylvestris (Lam.) Janch. Brassicaceae T H G i s ane CR Eur S Ul 19645 XIX ? Zając A. et al. 1998 n.c.d. H 2(?) aut & A fr N anthr~~

Bromus japonicus Thunb. ex Murr Poaceae TG w ane R Eur S Ul [BS, W] 1839“ 1850 Poznań (K rawiecowa 54 H 2 (+ l) M eu sel et a l 1965; [= B. patulus Mert. & W.D.J. Koch] egz & A sia W A r5 1951 after Ritschl) Hulten 1964; Hulten & anthr Fries 1986 Bromus pseudothominii P.M. Sm. H Poaceae T G w ane Anthropog. Ul XX Zając A. et al. 1998 n.c.d. H 2(?) [= B. hordaceus L. x B. lepidus Holm b.] egz

Bromus sąuarossus L. Poaceae T G w ane CR Eur S Ul [BS W] 1911 Schube 1911 n.c.d. H 2 (+ l) M eusel et al. 1965 egz & A sia SW anthr ? Buddleja daridii Franchet Buddlejaceae N G i ane C A sia I [0 ] 1890 [I]45 XX Adamowski et al. 2002 n.c.d. H H+l) UK; New Zealand anthr [China] 195225

~~Calendula arrensis L. Asteraceae T G i s ane R Eur S I 19015 XVIII K luk (1786); n.c.d. H 2 (+ /-) H u lten & Fries 1986 egz & A sia SW A r10 Z ając A. et al. 1998~~

Calystegia syhatica (Kit.) Griseb. Convolvulaceae G H G i s ane Eur S Ul XIX / XX Z ając A. et al. 1998 n.c.d. H 2(?) H ulten & F ries 1986 li ? Carya cordiformis (Wangerin) K. Koch Juglandaceae M G w bar Am N [C & E] I [0] 1689 [I]' 1820 [I] H ereźniak 1992; n.c.d. N 1(?) Seneta 1994

? Carya ovala (Mili.) K. Koch. Juglandaceae M G w bar Am N [E & C] I [0] 1629 [I]1 1808 [I] H ereźniak 1992; n.c.d. N 1(?) Seneta 1994 Centaurea micranthos S.G. Gmelin ex Hayek Asteraceae H G i s ane Eur SE & EC Ul 2/2 XX Zając A. et al. 1998 n.c.d. H 1(?) [= C. biebersteinii DC.; egz C. stoebe subsp. micranthos Hayek] Cerasus mahaieb (L.) Mili. Rosaceae N G i end C Eur S I [0 , F] 1785[I]25 XV III [I] ? Z ając A. et al. 1998 n.c.d. N 2 (+ l) [= Prunus mahaieb L.] & A sia C 183925 XIX Cerasus vulgaris Mili. subsp. vulgaris Rosaceae M G/V i end Eur SE I [F] Anc Anc Adamowski et al. 2002; n.c.d. N 1(?) [= Prunus cerasus L.] & A sia SW XIX ? Seneta 1994 Chenopodium schraderanum Schult. Chenopodiaceae T G w ane A fr N [m] I / U l 18645 1964 F ijałkowski 1964; 3 H 1(?) K ulpa 1964

~~? Comptonia peregrina (L.) Coult. Myricariaceae N G/V w ane Am N [E] I [0] 1714 [I]1 1813 [I] H ereźniak 1992; n.c.d. N 1(?) Seneta 1994~~

? Cornus alba L. Comaceae N G i s end C Eur E, I [0] 1773 [I]25 1741 [I] Seneta 1994 n.c.d. NH 2(+ D A sia C & E 185725 XIX ? ? Cotoneaster horizontalis Decne Rosaceae N G/V i end C Asia I [0 ] ca. 1870 [I]46 XX Seneta 1994 n.c.d. H 1(+1) [China] 19622

~~? Cotoneaster lucidus Schlecht Rosaceae N G i end A sia C I [0 ] 1840 [I]46 XX Seneta 1994 n.c.d. NH 2(+2)~~

Crataegus flabellata (Bose ex Spach) K. Koch Rosaceae N G i end Am N [NE] I [0 ] 1830[I]46 1928 [I] Seneta 1994 n.c.d. H 1(?) 19935 2/2 XX Crataegus pedicellata Sarg. Rosaceae N M G i end Am N [NE] I [O] 1683 [I]46 1810 [I] Seneta 1994 n.c.d. N 1(+D [= C. coccinea H ort.] XX ? Crocus vernus (L.) Hill Iridaceae GV/G i ane Eur C [m] I [0] XIX Schube 1903; Mirek n.c.d. SH K?) et al. 2002 Cuscuta campestris Yunck. Cuscutaceae Tp G/V i s ane Am N [W] U l 18832' 5' 1939 Piech 1939 29 H 2(+ /-) some countries in Eur: 18982 Hungary & Russia [meadows]

Cuscuta trifolii Bab. & Gibson Cuscutaceae Tp G/V i s ane Eur S U l 1843 1866 K linggraeff 1866 70 H 2 (+ l) some countries in Eur: 18507 [meadows] Dianthus barbatus L.s.s. Caryophyllaceae Ch G/V i ane Eur C & S [m] I [0] 18745 XVI? Zając A. et al. 1998; 95* H 2 (+ l) anthr XVIII - Kluk (1786) First First N rs Way Source of historical Described as invasive Name of species Family LF R P Disp LS Origin record record o f Hab. Dyn Maps o f INT data for Poland elsew here for Europę for Poland sq.

~~? Elaeagnus angustifolia L. Elaeagnaceae N M G i end С Eur S, I [O] 1736 [I]25 1652 [I] H ereźniak 1992 n.c.d. H 1(?) H ungary; A sia W & С 188325 XIX ? Am N [rip.]~~

~~? Elaeagnus commutata Bernh. Elaeagnaceae N G/V i end Am N [E] I [O] 1813 [I]45 XIX Z ając A. et al. 1998 n.c.d. NH 2 (+ l) [= E. argentea Pursh]~~

~~Erucastrum gallicum (Willd.) O.E. Schulz Brassicaceae T H G i ane CR Eur S & W Ul [BA, G] 18675 1936 A scherson & G raebner 31* H 2 (+ l) H ultćn & F ries 1986 1936~~

~~Erysimum marschallianum Andrz. ex m. Bieb. Brassicaceae H G i s aut Eur SE & Asia Ul 2/2 XIX 1985 Zając A. et al. 1998; 11* SH 1(?) [= E. durum J. Presl & C. Presl] ane Rutkowski unpubl.~~

~~Euphorbia maculata L. Euphorbiaceae T G i aut R Am N Ul / I XIX ? 2/2 XIX Z ając A. et al. 1998 n.c.d. H U?) some part of Eur S myr~~

~~Festuca rupicarpina (Hack.) A. Kern. Poaceae H G w ane Eur С U l [AN, S] 1995 M irek 1995 1 s 1(?) Z a j ą c A . & Z a ją c M. 2001 egz [Alps]~~

~~? Fraxinus angustifolia Vahl subsp. angustifolia Oleaceae M G w ane Eur S, I [0 ] XIX ? 1/2 XIX Zając A. et al. 1998 n.c.d. H 1(?) A fr N, A sia W~~

~~Fraxinus pennsylvanica M arshall Oleaceae M G w ane С Am N [С & E] I [O] 1783 [I]1 1817 [I] Z ając A. et al. 1998 179 SH 3(+2) Czech Rep. & Hungary 1/2 XIX~~

Geranium bohemicum L. Geraniaceae T G i s egz CR Eur С [N] Ul [G ?] 180120 1872 K n a p p 1872 4 H l(+/-> ? Red List (Benkert et al. M eusel et al. 1978; aut 1998) Hulten & Fries 1986 Gypsophila perfoliata L. Caryophyllaceae Ch G i ane CS Eur SE, Ul XX Z ając A. et al. 1998 n.c.d. H K+l) A sia W & С

~~Hordę u iii jubatum L. Poaceae T G w ane SR Am N & Asia E 1 [O, G] 189410 XX Z ając A. et al. 1998 n.c.d. H 2 (+ l) Am N H ulten 1964; H ulten & egz F ries 1986~~

~~? Juglans regia L. Juglandaceae M G w bar Asia SW, С & E I [F, M] Ar2- 25 XVIII K luk (1787) - only as n.c.d. SH K+l) end 196825 XIX cultivated plant~~

~~Lactuca tatarica (L.) C.A. Mey. A steraceae HG/V i s egz CS Eur SE U l [G] 18847 1/2 XX Zając A. et al. 1998 12 SH K+l) Hultżn & Fries 1986; ane & A sia W UK M eusel et al. 1992 myr 19002~~

? Larix kaempferi (Lamb.) Carriere Pinaceae M G w ane A sia W I [FO, 0 ] 1861 [I]45 XIX ? Seneta & D olatowski n.c.d. N K+l) [= L. japonica Carriere; [Japan] 1997 L. leptolepis (Siebold & Zucc.) Endl.] Linum austriacum L. Linaceae H G i s aut Eur W & С Ul 18602 XIX ? M irek et al. 2002 n.c.d. NS K?) M eusel et al. 1978 egz Z a j ą c A . & Z a ją c M. 2001

Linum perenne L. Linaceae H G i s aut CS Eur S & E Ul / I [C] XX Z ając A. et al. 1998 17 SH K+1) H ulten 1971; H ulten & egz F ries 1986 anthr ? Lonicera caprifolium L. C aprifoliaceae N li G i end С Eur SE 1 [0] 18095 1613* * - generał information n.c.d. NSH K?) XVIII (after Syreński 1613); K luk (1787) - as cultivated plant; probably also in the wild

~~Lonicera tatarica L. Caprifoliaceae N G/V i end С Eur SE & Asia С I [O] 1752 [I]45 XVIII ? Adamowski et al. 2002; n.c.d. SH 2 (+ l) 186425 Z ając A. et al. 1998~~

Lycopersicon esculentum Mili. Solanaceae T G i s aut CR Am S I [C] XVIII [I]24 1613* * - generał information n.c.d. H 2 (+ l) [= Solanum lycopersicum L.] hyd 18805 XV III [I] (after Syreński 1613); anthr 2/2 XX XVIII - Kluk (1788) - only as cultivated plant; Z ając A. et al. 1998

~~? Mahonia aąuifolium (Pursh) Nutt. Berberidaceae N G i s end CS Am N [W] 1 [0] 1822 [I]25 1839 [I] H ereźniak 1992 n.c.d. H K+l) Czech Rep., Germany 18602' 25 2/2 XX~~

? Malus domestica Borkh. H Rosaceae M G i end A nthropog. I [F, O] Anc Anc n.c.d. SH 2 (+ l) Ar2' 5 First First Nrs Way Source of historical Described as invasive Name of species Family LFR P Disp LS O rigin record record o f Hab. Dyn Maps o f INT data for Poland elsew here for Europę for Poland sq.

Mentha citrata Ehrh. Lamiaceae HG/V i hyd not definie Ul XX ? M irek et al. 2002 n.c.d. H ? subsp. pubescens (Willd.) Tacik H [= M. spicata x aąuatica L.] Mentha niliaca (Juss.) ex Jacq. H Lamiaceae H G/V i hyd not definie I 19765 XIX ? Z ając A. et al. 1998 n.c.d. H ? Mentha rotundifolia (L.) Huds. Lamiaceae H G/V i hyd Eur S I 18465 XIX ? Zając A. et al. 1998 n.c.d. H ?

~~Mentha spicata L. emend. L. Lamiaceae H G/V i hyd C Anthropog. I [C] 18185 XVIII Z ając A. et al. 1998 n.c.d. SH 9 H ulten & F ries 1986~~

~~Oenothera perangusta R.R. Gates Onagraceae H G i s ane Am N Ul XX ? Mirek et al. 2002 n.c.d. H 1(?) aut~~

~~? Oxycoccus macrocarpos (Aiton) Pursh Ericaceae Ch G/V i ane Am N I [C] XX M irek et al. 2002 n.c.d. SH 1(?) end anthr~~

~~? Picea glauca Voss Pinaceae M G w ane Am N [N & NE] I [0] ca. 1700 [I]1 1808 [I] H ereźniak 1992 n.c.d. N 1(?) [= P. alba (Aiton) Link; P. canadensis (Mili.) Britton] end 19766'~~

~~? Picea sitchensis (Bong.) Carriere Pinaceae M G w ane Am N [W] I [0] 1831 [I]' 1876 [I] H ereźniak 1992 n.c.d. N 1(?) end~~

~~Pinus banksiana Lamb. Pinaceae M G w ane Am N [NE] I [FO] 1735 [I]' 1822 [I] H ereźniak 1992: Sud- n.c.d. N H 2(?) 199061 1927 ? nik-W ójcikowska l987a Lithuania~~

? Pinus nigra J.F. Arnold Pinaceae M G w ane Eur S, Afr NW, I [FO, 0 ] 1759 [I]45 XIX [I] Adamowski et al. 2002 n.c.d. N 1(?) Asia W**

~~Pinus strobus L. Pinaceae M G w ane C Am N [NE] I [FO, O] XVI [I]1 ca. H ereźniak 1992 n.c.d. N K?) Czech Rep. 18005 1798 [I]~~

~~Polycneum heuffelii Lang Chenopodiaceae T G w ane Eur E & SE Ul Ar 1879 U echtritz 1880 4 H K-l) Jalas & Suominen 1980~~

Polycneum majus A. Braun Chenopodiaceae T H G w ane SR Eur S & Asia C Ul Ar5 1953 K ornaś 1954 15 H 1-2 (+/-) Jalas & Suominen 1980 ? Populus berolinensis (K. Koch) Dippel H Salicaceae M V w ane Anthropog. I [O, C] 1870 [I]48 2/2 XIX [I] Seneta & Dolatowski n.c.d. H ? [= P. laurifolia Ledeb. x P. nigra L. ‘Italica’] Berlin 1997; Mirek et al. 2002 Populus canadensis M oench H Salicaceae M V w anthr A nthropog. I [0 , C] 1750 [I]45 XX Adamowski et al. 2002; n.c.d. NH ?(+D [= P. x euroamericana (Dode) Guinier; 195221 M irek et al. 2002 P. deltoides Marshall s. 1. x P. nigra L. s.l.] ? Populus candicans Aiton Salicaceae M V w ane Am N [E] I [0] 1755 [I]1 XIX Hereźniak 1992; n.c.d. SH 9 Z ając A. et al. 1998

? Populus ‘NE 4 2 ’ H Salicaceae M V w ane A nthropog. I [0 ] M irek et al. 2002 n.c.d. H ? ? Populus nigra L. ‘Italica’ Salicaceae M V w ane Anthropog. I [0] XVII [I]45 XIX [I] Mirek et al. 2002 n.c.d. H ? ? Populus trichocarpa Torr. & A. Gray ex Hook. Salicaceae M G w ane A m N [NW] I [O] 1892 [I]' XIX ? Hereźniak 1992; n.c.d. H 9 Z ając A. et al. 1998

Prunus cerasifera Ehrh. Rosaceae N M G i end Eur SE, I [F, 0 ] 159445 XIX Adamowski et al. 2002; n.c.d. SH 9 [= P. divaricata Ledeb.] Asia SW & C Z ając A. et al. 1998 Prunus domestica L. subsp. domestica H Rosaceae N M G/V i end Anthropog. I [A] 1594 [I]25 Anc. Adamowski et al. 2002 n.c.d. SH 9 [probabl. = P. cerasifera Ehrh. x P. spinosa L.] 178725 XVI [I] ? A r5 XVIII ?

? Pseudotsuga menziesii (Mirb.) Franco Pinaceae MG w ane Am N [NW] I [0] 1827 [I]1 1833 [I] H ereźniak 1992; Seneta n.c.d. N 9 [= P. douglasii (Sabinę ex D. Don) Carriere; & D olatowski 1997 P. taxifolia (Poir.) Britton ex Sudw.]

~~? Ptelea trifoliata L. Rutaceae NG i w ane Am N [E] I [0] 1704 [I]12'-45 1806 [I] H ereźniak 1992 n.c.d. H 1(+D 1937~~

? Pterocarya fraxinifolia Spach Juglandaceae M G/V i w ane Asia SW [Cauc.] I [0] 1872 [I]45 XIX [I] Adamowski et al. 2002 n.c.d. N 9 Pyrus communis L. H Rosaceae M G i end A nthropog. I [A, 0 ] 1594 [I]25 Anc. Adamowski et al. 2002 n.c.d. NSH ?(+ l) [= P. pyraster (L.) Burgsd. x P. eleagrifolia Pall. x 1787 ?25 XVI [I] ? P. nivalis Jacq.] A r5 X V III ?

? Quercus cerris L. Fagaceae M G w bar C Eur SE, Asia W I [0] 1796 [I]25 XIX [I] Z ając A. et al. 1998 n.c.d. NS 9 end 195725 XX First First Nrs Way Source of historical Described as invasive Name of species Fam ily LF R P Disp LS Origin record record o f Hab. Dyn Maps o f INT data for Poland elsew here for Europę for Poland sq.

Quercus rubra L. Fagaceae M G w bar С Am N [Е] I [FO, .0] 1691 [I]1 1806 [I] Hereźniak 1992; War 554* N 3-4(+2) Czech Rep. end 188725 1924 ? szawa (Sudnik-Wójci anthr 1937 ? kowska 1987a); Wolny herb. MGS

Reynoutria bohemica Chrtek & Chrtkova H Polygonaceae G V/G? w i s ane A nthropog. 1 [O, C] 19425 1/2 XX ? F ojcik & Tokarska-G u- n.c.d. NSH ?(+2) Czech Rep. [= R. japonica Houtt. var. japonica egz - U l zik 2000 x R. sachalinensis (F. Schm idt) Nakai] myr ? Rhododendron ferrugineum L. Ericaceae N G i ane Eur С [m] 1 [0] XIX XIX Z ając A. et al. 1998 n.c.d. SH ? Rhus typhina L. Anacardiaceae N G/V i ane С Am N [С & E] 1 [0] 1602 [I]' 1806 [I] Hereźniak 1992; Ada- n.c.d. H ? aut 1937 mowski et al. 2002 ? Ribes rubrum L. Grossulaceae N G i end Eur [NW] I [F, 0 ] A nc45 Anc Adamowski et al. 2002 n.c.d. SH ? 18095 XIX? ? Rosa acicularis Lindl. Rosaceae N G i s a end Eur NE, 1 [0] XX Zając A. et al. 1998 n.c.d. H ? Asia N & NE

? Rosa blanda Aiton Rosaceae N G i s a end Am N [E] I [0] 1773 [I]45 2/2 XX Adamowski et al. 2002; n.c.d. H 9 Z ając A. et al. 1998 ? Rosa carolina L. Rosaceae N G/V i s a end Am N [E & S] I [0] 2/2 XX Z ając A. et al. 1998 n.c.d. H ? ? Rosa darurica Pall. Rosaceae N G i s a end Asia E I [0] 1/2 XX Zając A. et al. 1998 n.c.d. H ? Rosa foetida Flerrm. Rosaceae N G i s a end A sia SW I [0 ] 18145 1/2 XX Zając A. et al. 1998 n.c.d. H 7

Rosa glauca Pourr. Rosaceae N G/V i s a end С Eur SW [m] I [0] 181445 1/2 XX Zając A. et al. 1998 n.c.d. SH 9 Z ieliński 1981 [= R. rubrifolia Vill.] 18745 Rosa gorenkensis Besser R osaceae N G i s a end Eur SE, Asia W I [0] 9 2/2 XIX Adamowski et al. 2002; n.c.d. NS 9 [= R. glabrifolia auct. non C.A. Mey.] Z ając A. et al. 1998

Rosa multiflora Thunb. Rosaceae N G i s a end Asia E I [0] b e f o r e 2/2 XX Adamowski et al. 2002; n.c.d. NSH 2 (+ l) 186845 Z ając A. et al. 1998 Rosa spinosissima L. Rosaceae N Ch G/V i s a end С Eur S & SE, 1 [C] XVI45 XIX [I] Adamowski et al. 2002; n.c.d. H 9 M eusel et al. 1965; [= R. pimpinellifolia L.] Asia SW & С 2/2 XX Z ając A. et al. 1998 Hulten & Fries 1986 ? Rosa virginiana Herrm. Rosaceae N G/V i s a end Am N [E & N] I [0] 1/2 XIX Z ając A. et al. 1998 n.c.d. H 9

Rudbeckia hirta L. A steraceae H G i s ane CR Am N I [0] 18602 2/2 XIX M irek et al. 2002 n.c.d. H ! ( + / - ) M eusel et al. 1965 egz myr anthr Rumex longifolius DC. Polygonaceae H G w s ane Eur NE Ul 19615 XIX Zając A. et al. 1998 n.c.d. SH 9 Czech Rep. egz hyd 9 Rumex patientia L. Polygonaceae H G w s ane С Eur & Asia I [C] 18615 XVIII [I] Kluk (1788) - only as n.c.d. H Hegi 1958; J a l a s & Suo- egz XIX cultivated plant; Zając A. M1NEN 1979 hyd et al. 1998

Salix acutifolia Willd. Salicaceae N G/V i ane С Eur E & Asia С I [O, C] XVIII Zając A. et al. 1998 154 NSH 2 (+ l) ? Salix cordata M ichx. Salicaceae N G/V i ane Am N I [FO] 2/2 XX Z ając A. et al. 1998 n.c.d. N 9 ? Salix eriocephala Michx. Salicaceae N G/V i ane Am N I [0] XIX10 2/2 XX Zając A. et al. 1998 n.c.d. H 9 Scutellaria altissima L. Lamiaceae H G i ane С Eur S & SE I [C] 19015 1/2 XX Zając A. et al. 1998 n.c.d. SH 9 egz aut Sorbaria sorbifolia (L.) A. Braun Rosaceae N G/V i aut С Asia N & E I [0] 1750 [I]45 XIX Adamowski et al. 2002; n.c.d. NSH 9 anthr 189062 Z ając A. et al. 1998 190425

~~? Spiraea tomentosa L. Rosaceae N G/V i ane Am N [E] I [0] XIXXIX H ereźniak 1992; Dajdok n.c.d. NSH 9 Germany aut Z., Pender K. & Kącki anthr Z. 2003 unpubl.~~

? Spirea chamaedryfoUa L. em. Jacq. Rosaceae N G/V i ane Eur SE, I [0] 1789 [I]45 XIX Zając A. et al. 1998 n.c.d. NSH 9 [= 5. ulmifoilia Scop.] aut Asia NE & С 182662 anthr 19005 First First Nrs Way Source of historical Described as invasive Name of species Family LF RP Disp LS Origin record record of Hab. Dyn Maps o f INT data for Poland elsew here for Europę for Poland sq.

? Spirea pseudosalicifolia Silverside H Rosaceae N V/G i ane A nthropog. 1 [O] XIX ? Zając A. et al. 1998 n.c.d. H 9 [= S. salicifolia L. x S. douglasii Hook.] aut anthr Caprifoliaceae N Hereźniak 1992; 9 (L.) S.F. Blake G/V i end С Am N [NE] I [0] __ 1824 [I] n.c.d. NSH ? Symplioricarpos albus , Czech Rep.

~~[= S. racemosus M ichx.; S. rivularis Suksd.] anthr 00 00 00 40 -J XVIII ? Z ając A. et al. 1998~~

Syringa vulgaris L. Oleaceae N G/V i aut С Eur SE I [0] 1554 [I]45 XVI [I] ? Adamowski et al. 2002; n.c.d. NSH 3(+D C zech Rep. 1787 ?25 XVIII Z ają c A. et al. 1998; Kluk (1788) - only as cultivated plant;

~~? Tbuja plicatu Donn ex D. Don Cupressaceae M G w ane Am N [W] I [O] 1853 [I]1 1826 [I] H ereźniak 1992 n.c.d. NH 9 end~~

? Tsuga canadensis (L.) Carriere Pinaceae M G w ane Am N [E] I [0 ] 1736 [I]' 1813 [I] H ereźniak 1992 n.c.d. N 9 Typha Іахтапіі Lepech. Typhaceae H G/V w ane Eur & A sia I [0] 199647 XX Mirek et al. 2002 15 NSH K + l) Hungary B a ry ła et al. 2005 Ulex europaeus L. Fabaceae N G i aut С Afr N & Eur SW I [O, FO] 177310 XIX Zając A. et al. 1998 n.c.d. SH ? M eusel et al. 1965; m yr 18805 Hulten & Fries 1986

~~Yeronica gentiaitoides Vahl Scrophulariaceae H G/V i s ane Asia SW I [0] 1968 Mirek et al. 2002; n.c.d. s 9 m yr Oklejewicz 1997~~

? Vitis vinifera L. subsp. vinifera Yitaceae H li G/V i s end Eur Asia I [F, O] Ane Anc K luk (1788) - only as n.c.d. SH 9 [= V. vinifera L. A r5 XX cultivated plant Asclepias syriaca l. Medicago x varia M a r t y n Sicyos angulata L. Sisyrynchium bermudiana L. emend. F a rw .

During the process of gathering data on the distribution Among the above-listed species, Medicago x varia routes (on embankments) has been now recorded in dry first half of the 19,h century as a decorative plant. The of kenophytes in Poland, 5 maps were developed for the occurs relatively often and is a species of hybrid ori- meadóws and grasslands, where (being a clonal plant) first sites where it returned to the “wild” were report- species not included in Distribution Atlas of Vascular Plants gin resulting from the C ro s s in g of an alien species M. it colonises large areas. Sisyrinchium bermudiana, ed in 1835 from north-western Germany (after H egi in Poland ( Z a j ą c A. & Z a j ą c M. 2001). These are: sativa with the native M. falcata. In some regions it found in meadows and ruderal habitats, probably has 1909). In Poland, it was first found as recently as in the Ailanthus altissima (Mili.) Swingle (the map for this might even be found more freąuently than the parental more stations but is difficult to find outside the brief first half of the 20lh century in the Sudety Mts. (S chube species is included in Chapter 7: Fig. 39); species (cf. also Chapter 8). The other three species con- flowering period and thus might be overlooked. This 1928). Occurs also in all countries bordering Poland. Asclepias syriaca L. tinue to show increases in the number of stations oc- species, which originated from the eastem part of North The number of sites with the plants returning to the Medicago x varia M artyn cupied. It is worth noting that pre- Sicyos angulata L. Asclepias syriaca, America, occurs also in the Bermudas and in Ireland “wild State” could increase because the plant is currently Sisyrinchium bermudiana L. em. Farw. viously noted around cultivated areas or along railway (S endek & W ik a 1982). It was brought to Europę in the offered by gardening shops.

A ppendix D. A comparison of the terminologies for the classification of synanthropic plants used in studies on plant invasions in Central Europę, in Poland and that proposed by Richardson et al. 2000

~~Proposed Recommended phytogeographical term Term used D efinition D efinition terminology by Definition in Central European in Polish studies Richardson et al. 2000 studies~~

~~A. Apophytes native species occurring in man-made habitats A. Apophytes native species occurring in man-made habitats~~

B. Anthropophytes species introduced by man B. Anthropophytes alien plant species Alien plants Plant taxa in a given area whose presence there is due to intentional or accidential introduction as a result of human activity.

I. Hemerophytes introduced intentionally I. Diaphytes not permanently established Casual alien plants Alien plants that may flourish and even reproduce occasionally in an area, but which do not form self-replacing populations, and which rely on repeated introductions for their persistence.

II. Xenophytes introduced unintentionally II. Metaphytes permanently established /settled Naturalised plants Alien plants that reproduce consistently and sustain populations over many life cycles without direct intervention by humans (or in spite of human intervention); they often recruit offspring freely, usually close to adult plants, and do not necessarily invade natural, semi-natural or human made ecosystems.

1. A rchaeophytes introduced before 1500 1. Archaeophytes introduced before 1500 Naturalised plants that produce reproductive offspring, often in very large numbers, at considerable distances from parent plants (approximate scales: > 100 m; < 50 years for 2. Neophytes introduced after 1500 2. Kenophytes introduced after 1500 taxa spreading by seeds and other propagules; > 6 m/3 years for taxa spreading by roots, rhizomes, stolons, or creeping stems), and thus have the potential to spread over a. Ephemerophytes temporary occurrence, only in man-made habitats Invasive plants1 a considerable area. (not invasive) b. Epecophytes established in man-made habitats a. Epecophytes established in man-made habitats 1 c. Neoindigenophytes penetrating into natural habitats b. Agriophytes penetrating into natural habitats Transformers A subset of invasive plants which change the character, condition, form or naturę of (= agriophytes) (= Neophytes sensu Faliński) ecosystems over a substantial area relative to the extent of that ecosystem.

S o u r c e : K o r n a ś 1981; P y ś e k 1995; T o k a r s k a -G u z i k 2001a; R i c h a r d s o n et al. 2000 1 - The Authors of the cited definition suggest that invasive should be used with reference to the ‘biogeographic/demographic’ status of a species without any connotation of impact; Richardson et al. 2000 also include in their recommended terminology the well-established term for harmful plants - Weeds - plants (not necessarily alien) that grow in sites where they are not wanted and which usually have detectable economic and environmental effects (see also text in Chapter 12); yellow cells - indicates the main correspondence in alien plant terminology between different classifications; red cells - indicates kenophytes and their equivalent groups in the other terminologies cited.