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Barbara Tokarska-Guzik

The Establishment and Spread

of Alien Plant Species (Kenophytes) in the Flora of Poland

W ydawnictwo U niw ersytetu Śląskiego • Katowice 2005

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The Establishment and Spread of Alien Plant Species (Kenophytes)

in the Flora of Poland

To my husband

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PRACE NAUKOWE

UNIWERSYTETU ŚLĄSKIEGO W KATOWICACH

NR 2372

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Barbara Tokarska-Guzik

The Establishment and Spread of Alien Plant Species (Kenophytes)

in the Flora of Poland

W ydawnictwo Uniwersytetu Śląskiego Katowice 2005

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Editor o f the Series: Biologia

Paweł Migula

Reviewers

Bogdan Jackowiak Adam Zając

Cover design: M a re k F ra n c ik Executive Editor: W io le tta T o m a la -K a n ia Technical Editor: B a rb a ra A re n h ó v e l Proof-reader: G ra ż y n a W o jd ała

Copyright © 2005 by W ydawnictwo Uniw ersytetu Śląskiego Ali rights reserved

ISSN 0208-6336 ISBN 83-226-1485-3

Published by

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24,5 + insert. P u blishing sheets: 31,5. P assed to the Printing H ouse in Septem ber 2005. Signed fo r print- ing and p rin tin g fm ish e d in D e ce m b e r 2005.

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Contents

A c k n o w le d g e m e n ts... 9

PART ONE T h e o r e tic a l o v e r v i e w ... 11

1. Introduction. The subject, objectives and the scope of this study: The role o f kenophytes in the flora as representations of the anthropogenic alteration of v e g e t a t i o n ... 11

2. Review of studies on selected aspects of synanthropisation o f the vegetation c o v e r ... 14

2.1. The history of studies on alien plant species in Poland viewed against the situation in Europę as a w hole... 14

2.2. Synanthropisation: the essence o f the process and the role of kenophytes in the changes occurring in the natural environm ent on E a r t h ... 19

PART TWO T e rm in o lo g y a n d m e t h o d o l o g y ... 23

3. Phytogeographical terminology and the classification of synanthropic plants used in Poland . . . 23

4. Materials and m e t h 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 u s e d ... ... . 25

4.3. List of kenophytes and the scope of the inform ation collected in order to characterise them . 26 4.4. Cartogrammes and their a n a l y s i s ... 27

4.5. Use, interpretation and synthesis of d a ta ... 27

PART THREE A n a ly s is a n d s y n th e s is o f d a t a ... 29

5. Geographical and ecological characteristics of the flora of kenophytes in P o l a 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 i g i n ... 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 r o p e r t i e s ... 35

5.1.7. Freąuency and status in the f l o r a ... 36

5.2. Kenophytes in historical accounts of flo ra s ... 40

5.2.1. “O ld” 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 p erio d s... 46

6. Current types of distribution of kenophytes in P o lan d... 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

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6.2. Kenophytes with scattered stations over the whole territory of Poland, with concentrations of morę

frequent stations in some r e g i o 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 (contem porarily) reaching their limit of distribution in P o l a 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 l l e y s ... 64

6.5. Kenophytes associated with urban areas and railway ro u te s ... 66

7. The history of the spread of selected kenophyte species in the territory o f P o la n d ... 67

7.1. The history of the spontaneous spread of cultivated woody plants as the result of “dom esticating” s p e c i e s ... 67

Acer negundo L ... 67

Padus serotina (Ehrh.) B orkh... 68

Ailanthus altissima (M ili.) S w in g le... 71

Clematis vitalba L ... 71

7.2. The history of the spread of useful herbaceous plant species: how m edicinal and decorative plants have established themseWes in the f l o r a ... 74

7.2.1. Exam ples of species o f European o r i g i n ... 74

Cymbalańa muralis P. Gaertn., B. Mey. & Scherb... 74

Digitalis purpurea L ... 74

Echinops sphaerocephalus L ... 78

7.2.2. Е х а т р іе of species of Asian o r i g i n ... 78

Elsholtzia ciliata (Thunb.) H yl... 78

Impatiens glandulifera R o y l e ... 79

Impatiens parviflora D C ... 81

Reynoutria japonica H outt... 84

7.2.3. Examples of species of American o r i g i n ... 86

Echinocystis lobata (E M ichx.) Torr. & A. G r a y ... 86

Rudbeckia laciniata L ... 88

Mimulus guttatus D C ... 88

7.3. The spread o f accidentally introduced plants: how an ephem erophyte turns into a kenophyte. 91 7.3.1. Plants introduced accidentally from various regions of E u r o p ę ... 91

Anthoxanthum aristatum B oiss... 91

Artemisia austriaca Jacq... 91

Bunias orientalis L ... 94

Eragrostis minor H o s t ... 96

Rumex confertus W illd... 96

Salsola kali L. subsp. ruthenica (Iljin) S o ó ... 99

7.3.2. Plants brought accidentally from A s i a ...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 M ichx... 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 determ ine the recent shape of the flora of P o la n d ? ...109

9. Historical aspects of the development of the kenophyte flora of P o l a n d ... 117

9.1. General r e m a r k s ...117

9.2. The effect of historical and economic developm ents on the enrichm ent o f Polish flora by new- c o m e r s ...117

9.3. Cities as “footholds” for further expansion by fresh n e w c o m e r s ... 118

9.4. Historical gardens, botanic gardens, cloister and convent gardens as places of “dom esticating” exotic species prior to their spontaneous e s t a b l i s h m e n t ... 124

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9.5. Immigration periods (peak inflows of k e n o p h y te s )...126

9.6. Migration r o u t e s ...127

9.6.1. Rivers as migration corridors aiding the spread o f k eno ph ytes... 128

9.6.2. The role of humans in the migrations of k e n o p h y te s ... ... ... 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 la n ts ...141

12.1. More remarks on te r m i n o l o g y ... 141

12.2. Consequences of invasions by alien species, legał regulations and m ethods o f com bating the t h r e a t ...144

12.3. Invasive kenophytes in P o l a 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 o f plants o f 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 l u s i o n s ... 155

14. Invasions of alien plant species at the dawn of the 21st century: perspectives for further studies . 157 R e f e r e n c e s ... ...159

S tre s z c z e n ie ...189

Z u s a m m e n fa s s u n g ... 190

A p p e n d ic e s ... 191

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Acknowledgements

This thesis has been written thanks to the help o f many people. I wish to express my warmest gratitude to my Teachers for awakening my in- terest in the problem s o f synanthropisation of plant cover: Professor Dr. hab. K rzysztof Ros- tański, Professor Dr. hab. Adam Zając and Pro­

fessor Dr. hab. Janusz Bogdan Faliński, who sadly passed away recently. I have always expe- rienced great kindness and eagerness during long and insightful scientific discussions on the part o f my Teachers.

I am extraordinarily glad to be able to express my indebtedness to all my Co-w orkers at the Department o f Plant Systematics, as well as to Colleagues from the Departments o f Geobotany and Naturę Protection, as well as Biophysics and Celi Biology, at the University o f Silesia for their help during my investigations and during the preparation o f this thesis for publication.

I wish to thank all Polish Botanists whose data, submitted to the ATPOL database, have made it possible for me to prepare this thesis. I thank those Colleagues who sent their data collected during field investigations directly to me: Dr. Zygmunt Dajdok and Dr. Zygmunt Kącki (University of Wrocław), Dr. hab. Marek Kucharczyk and Dr.

Rafał Krawczyk (University o f Lublin), Dr. Dan Wołkowycki (University o f Białystok), Dr. Marze­

na Środa (University o f Warmia and Mazury) and Dr. Zofia Sotek (University o f Szczecin).

I would like to give my thanks to Professor Dr.

hab. Alicja Zemanek for her critical interpreta- tion o f the 17thcentury work o f Sirenius.

I want to thank Mr. Józef Gajda from the In- stitute o f Informatics o f the Jagiellonian Univer- sity for making his proprietary Computer program available to me and for extensive help in the preparation o f the cartographic section o f my thesis.

The group o f Colleagues to whom I owe a word o f gratitude for steadfast collaboration during lo- calisation o f sites includes Dr. Barbara Fojcik, Dr.

Gabriela Woźniak, as well as MSc. Jadwiga Du- czek (an economist by education) and her daugh- ter Martyna.

I would like to give my special thanks to Professor łan C. Trueman and Dr. Eleanor Cohn (University o f Wolverhampton, UK) who have always shown personal interest in my scientific life and have encouraged me in the field o f my research. Espe- cially Professor łan C. Trueman who has sup- ported me through interesting discussions and helpful comments.

I am grateful to MSc. Maryla Palowska for her extensive and extremely time-consuming technical help in preparation o f the graphical part o f the present work. I wish to thank my Colleagues Dr. Teresa Nowak and MSc. Anna Gawron for in­

sightful reading o f the manuscript and kind en- couragement.

I wish to express my sincere gratitude to Par- ticipants o f discussions conducted during my pre- sentations o f successive stages o f studies on the naturalisation and spreading o f kenophytes in Poland during geobotanical seminars in B iało­

wieża and Katowice.

I give my warm feelings o f gratitude to my Husband Zenek in whom I have always found support, to my daughter Zuzanna and son Kuba, as well as to my Parents, for their patience and constant presence at each moment o f preparation o f this thesis.

In the years 1997-2000, the investigations were conducted as an integral part o f research grant no. 6P04G05312 “The occurrence, history and m odern distribution o f kenophytes (neo- phytes) in Poland” funded by the State Commit- tee for Scientific Research.

2 The Establishm ent. 9

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PART ONE

Theoretical overview

“The life success o f each plant species hinges on not only the capability to settle passively in the places o f 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

The subject, objectives and the scope o f this study:

The role o f kenophytes in the flora as representations of the anthropogenic alteration o f vegetation

The subject o f this book falls within the theme o f the synanthropisation' o f vegetation cover.

Connected representations o f this directional pro- cess occurring on Earth under the impact o f var- ious forms o f human activities, are the processes o f the extinction o f some species and the expan- sion o f others, which have both accelerated in recent centuries and w hich are contributing to changes o f the biological diversity o f entire re- gions, countries or continents. A synthesis o f the role o f humans in the historie changes in land- scape and vegetation cover was presented by

Ko r n a ś (1977a) in a multi-authored book entitled Szata roślinna Polski [The vegetation o f Poland]

and in other detailed papers ( Ko r n a ś 1982, 1983, 1990, 1996). Dynamie change in floras, its scalę and rate - issues which have started to focus the interest o f scientists and conservationists - be- came the main motive for undertaking this study.

Nevertheless, the interest taken by scientists in territorially-expanding plant species o f foreign origin has its roots in ancient times and was con- ceived on the basis of an ever-increasing knowledge

1 Synanthropisation - is the process o f change in plant cover (also in the fauna and the abiotic elem ents o f the en- vironm ent) brought about by hum an im pact (for detailed definition sec Chapter 2.2).

about useful plants, particułarly those that are edible or poisonous, as well as on natural curiosity and a determination to leam about new, exotic species.

Practical considerations were also important and the ambition which drives explorers, both past and contemporary, to search for new plants in newly discovered remote parts o f the world. As early as in the ancient times, the body o f knowledge accu- mulated by naturalists and philosophers such as Theophrastus, Dioscorides and Pliny was impres- sive in terms o f volume and provided a source for copies, adaptations and reprints for the “herbalists”

o f the Middle Ages and Renaissance periods.

The studies o f the floras which accompany people inereased greatly from the beginning o f the 19th century. The oldest works devoted to plants o f foreign origin, however, date back to the 17th century. At this early date, an Italian botanist, Prosper Al p in u s published a work entitled De plantis exoticis (1627), where he gave descriptions o f plants found in Europę but originating from America. Other proofs o f naturalists’ interest in such plants can also be found in the old herbals2.

The phenom enon o f invasion by alien new- comers in their new homelands was also noted by

Darwin(1859) in his work On the origin o f species3, as well as in the diaries o f his joum eys, and in

2 For example, the Wrocław Herbarium collection (WRSL) has one o f the oldest herbaria in Europę which was assem- bled by an Italian Sivius Boccon, dated 1674 ( Ro s t a ń s k j K.

1963). This includes a typical specimen o f Solidago canaden- sis L. - a recent kenophyte (neophyte), distributed through- out Europę, and originating from North America.

3 Among other examples, Darwin described the invasion o f Cardo de Castilla (Spanish Cardoon) Cynara cardunculus L.

brought to Buenos Aires in 1749, and w hich had taken over Argentina, Chile and Uruguay in eight decades ( Cr o s b y 1999).

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letters and research reports. More inform ation te s tif y in g to th e p e r c e p tio n o f th e p h e n o m e n o n a n d th e c o n s e q u e n c e s th e r e o f , c a n b e f o u n d in n u m e ro u s n o te s a n d C o m m u n ic a tio n s p u b lis h e d in p o p u la r s c ie n tif ic jo u r n a l s 4.

The plants of foreign origin appearing in the floras o f many regions o f the world were called

“the vagrants o f our floras” ( Crosby 1999 after Hooker 1864) or “new acąuisitions” ( Kam ieński

1884а & b; Paczoski 1896), although they were sometimes called “newcomers and w aifs” or

“wandering plants” ( Trzebiński 1930; Szu l c z e w-

ski 1931) or - in some special cases - “invaders”

( Elton 1958).

Answers were sought to a number o f essential questions. From where did the alien species arrive in the local flora? Is it an escape from any culti- vation or has it been brought in accidentally?

Which place does it occupy in the new homeland and what consequences result from its arrival?

Nevertheless, the greatest attention was attracted by the spectacular manner of the arrival o f those alien plant species that colonised new territories rapidly and in great num bers5. M any o f those im m igrants soon became burdensom e acquisi- tions in the local flora, sometimes even earning common names reflecting the violent manner o f their invasion. One such е х а т р іе is Elodea canadensis (Canadian W aterweed) originating from North America, which conquered European inland waters in a “blitz” in the second half o f the 19'h century and the beginning o f the 20lh cen- tury, and which was called “the green ghost”

( Fa l iń sk i 2004 after Lons 1910). An Asian

species Im patiens parviflora (Smali Balsam), which dispersed over central Europę as a fugitive from botanic gardens having first established itself in ruderal communities and then succeeded in entering the forests, has been given a nickname

4 E.g. a colum n in Przyroda i P rzem yśl [N aturę and Industry], a w eekly devoted to advancem ent o f the natural sciences and their applications in industry, o f 1872 pub­

lished a note on the appearance o f new plants after the Franco-Prussian War. This note was prepared on the basis o f a study by de Vibraye (1870-1871) presented before the French Academy, in w hich the author describes the emer- gence o f 157 new exotic plants in central France. He at- tributed their presence in a new territory to an accidental introduction o f seeds from A lgeria by the French cavalry.

The author assessed this process as a perm anent change in the flora because “these plants not only w ithstood one o f the most severe w inters but flourished abundantly in the areas once ąuite devoid o f plants. Thus we can be quite sure that it is not a tem porary phenom enon but that essen- tially some o f the regions in France had their plant wild- life augm ented by new flora” .

5 W hile describing the spreading o f Spanish Cardoon, Charles Darwin stated: “I doubt w hether there was any such case in history o f native flora being invaded by an alien species on such a great scalę” ( Cr o s b y 1999).

o f the “pushy M ongoł” ( Fa l iń sk i 2004 after Naumann 1913). Similar associations had been provoked by the invasion o f European plants in other continents. The native Americans o f New England and Virginia called P lantago m ajor (Greater Plantain) “Englishm an’s footprint”, be­

cause in the 17thcentury they believed that this plant grew only “where the aliens set their feet and where it had not been know n before their arrival in this country” ( Crosby 1999).

The migrations o f species occurring as the re­

sult o f human activity which often assumed the characteristics o f massive invasions (“ecological explosions”), and which led eventually to changes in vegetation, fauna and to econom ic dam age, constituted the topie o f a book entitled The Eco- logy o f Invasions by Animals and Plants (1958), by Elton, a British ecologist, whose research in this field is considered to be classic. The date o f the publication o f the book can be regarded as the birth o f ecology of invasion as a new scientific discipline.

Crosby (1999), describing the successful colon- isation of the Globe by Europeans, even presented a hypothesis that the success o f European impe- rialism has an underlying biological and ecolo­

gical background (“ecological imperialism”). The same author, giving examples o f spectacular inva- sions o f the vast spaces o f A ustralia or both Americas, makes ironie comments: “A rapid inva- sion o f species o f European ‘w eeds’ disturbed American naturalists, even though most o f these botanists themselves hailed from the same region as the plants concemed”. Despite the great distance between these continents and Europę, the climate is similar in many regions, providing magnificent conditions for development o f the European col- onists, incłuding plants, animals and people.

The actual scalę o f the exchange o f species o f synanthropic plants between regions o f the world is considerable. The proportion o f alien species naturalised (i.e. permanently established) in some local floras ranges from 20% to even as high as 50%. Particularly dynamie is the exchange b e­

tween Eurasia and North America (JAg er 1988;

Sukopp 1995; Korn a ś 1996; Ja c k o w ia k 1999).

The invasions by plants, anim als or fungi are one o f the most pressing issues o f naturę con­

sidered on a global scalę. Some authors even deem it to be the single most im portant problem in protecting the biodiversity in the 2 Г ' century ( Carlton & Geller 1993; Vito u se k et al. 1996, 1997; Mooney & Hobbs 2000). The International Convention on B iological D iversity contains a special provision calling upon country-signato- ries to fight alien invasive species w hich could be o f danger to native habitats, com m unities or species. These circum stances have contributed

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to an evident increase in the interest in these issues among the theoreticians and practitioners o f naturę conservation.

Studies devoted to the spread o f alien plant species are becom ing alm ost as fundamental a part of the protection o f biological diversity as the compilation of “Red Lists” and “Red Data Books” of rare and endangered species ( Me d w e c- k a- Ko r n a ś & Pię k o ś- Mir k o w a 1997). The lists o f alien species are compiled along with the lists of invasive species which have entered natural and semi-natural communities, or - as weeds - the segetal communities. However, one will not be able to prevent their spread without knowledge o f their biology and habitat reąuirements as well as their geographical ranges o f distribution. For е х ат р іе many plant species took less than two centuries to invade and occupy the whole national territory of Poland ( Za ją c A. & Za ją c M . 2001).

The initiatives taken up in many countries, as well as those undertaken on a European and/or global scalę (e.g. the international programmes Ecology o f Biological Invasion, Global Strategy fo r Invasive Species, Global Invasive Species Programme), have made studies o f alien species, and in particular their extending distribution ranges, their ecology, and the effects exerted - an urgent and indispensable task.

B o th in P o la n d a n d o th e r E u ro p e a n c o u n trie s , stu d ie s o n th e m ig ra tio n s a n d d is trib u tio n o f alien p la n t s p e c ie s h a v e a c e rta in tr a d itio n 6, b e g in n in g fro m stu d ie s d e v o te d to p a rtic u la r sp e c ie s, su ch as, fo r е х а т р і е , Bidens frondosa ( Trzcińska 1961;

LhotskA 1966, 1968), Impatiens glandulifera

(Zając E.U . & Zając A. 1973; Bee r u n g & Per-

rins 1993; Gudżinskas & Sinkevićene 1995; Py-

śek & Prach 1995; Dajdok et al. 2003; Drescher

& Prots 20 03), Iva xanthifolia ( Guzik & Sudnik- Wójcik o w sk a 1989; Gudżin sk as 1991; Jehlik

1998) a n d Reynoutria (Fallopia) japonica (e .g . C o n o l l y 1977; A l b e r t e r n s t et al. 1995; S e ig e r

1997; B a ile y & C o n o l l y 20 0 0 ; C h ild & W a d e 2000; T o k a r s k a - G u z ik in press) o r r e g io n s (e .g . H o l z f u s s 1937; G u t t e 1971; L o h m e y e r & Su-

k o p p 1992; B r a n d e s & S a n d e r 1995; N a t a l i &

J e a n m o n o d 1996; A d a m o w sk i et al. 2 0 0 2 ), u p to s tu d ie s c o v e rin g w h o le c o u n tr ie s (e .g . C le m e n t

& F o s t e r 1994; J e h lik 1998; Z a ją c A. & Z a ją c M.

2 0 0 1 ; E s s l & R a b i t s c h 2 0 0 2 ; P r e s t o n et al.

20 0 2 ; P y ś e k et al. 2002).

The historical impact exerted by humanity on the vegetation cover, and the flora in particular, is best illustrated by examining two phenomena:

the interactions betw een the two groups o f

6 O utside Europę one can find num erous publications devoted to this issue (e.g. com pare literature cited by

Br u n d u et al. 2001 and Ch i l d et al. 2003).

species: native7 and alien, and the com parison between the group o f the oldest com panions o f humans (archaeophytes, so called oldcom ers) and the new er alien types (kenophytes = neo- phytes, so called newcomers).

The issue o f the origin and development o f the distribution ranges o f the oldest group o f alien species occurring in Poland (i.e. archaeophytes) has been taken up by Za ją c in a basie monograph (1979) and in detailed reports ( Za ją c 1983, 1987a

& b, 1988), while the same considerations for more recent newcomers (i.e. kenophytes) should begin to be addressed by the detailed maps included in Distribution Atlas o f Yascular Plants in Poland

( To k a r sk a- Guzik200la & b; Zają c A. & ZającM.

2001), as well as by the present monograph.

The main idea behind the present m ono­

graph is to investigate changes in synanthropic flora of Poland and to provide a synthesis of the know ledge accum ulated to date on the developm ent o f the kenophyte flora o f Po­

land. It is also an attem pt to reconstruct the historie changes in the ranges o f distribution of kenophytes in the territory of Poland. Ad- ditionally, those regions o f Poland w hich are particularly endangered by the excessive inva- sion o f alien species are indicated, and the “in- vasive species”8 are specifically identified in the first ever com prehensive list o f invasive keno­

phytes com piled for Poland.

These aims have been achieved by the follow- ing objectives:

• verifying and updating lists o f kenophytes com­

piled for Poland and presenting an original, comprehensive catalogue o f this group o f spe­

cies, with an associated database o f biological, ecological, geographical and historical attributes and information;

• establishing the first floristic data (first records) for particular species o f Polish kenophytes;

• studying the historie distribution o f kenophytes and attem pting to reconstruct the history o f kenophyte floras on the basis o f distribution maps, applying whenever possible a cartographic interpretation;

• attempting to reconstruct periods o f immigration and spread o f kenophytes (construed as cumu- lations or “migration waves”) showing also how they depended on historie and geographical conditions;

7 Sim ilarly im portant issue is the problem o f “apophy- tisation” o f native species, and its follow ing conseąuences in “ in v asib ility ” o f this group o f species beyoned their natural rangę (cf. C hapter 11 and 12).

8 Invasive species - species o f foreign origin, established in a p rim arily foreign area, producing fertile offspring, often in extrao rd in arily large num bers, dispersing over great distances from parental plants ( Ri c h a r d s o n et al.

2000); for term inology, see also Chapter 3 and 12.

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• identifying and describing the different pattems o f distribution o f kenophytes in Poland;

• reconstructing the history o f the introduction, establishment and spread o f selected species;

• discussing dynamie trends in kenophyte distri- bution, examining routes and pathways o f in- vasion and the factors supporting the conąuest o f various types o f habitats, and identifying areas most vulnerable to invasion (with prac- tical implications for naturę conservation).

2. Review of studies on selected aspects o f synanthropisation o f the vegetation cover

2.1. The history o f studies on alien plant species in Poland viewed against the situation in Europę as a whole

A short review o f the history, research trends and main methods used to study alien plant spe­

cies in Poland was the subject o f one o f the pre- vious paper (Tokarska-Guzik 200la). The present chapter is a further attempt to provide a synthesis o f different aspects o f studies on alien plant spe­

cies in Poland shown on wider perspective.

The discovery o f America by Christopher Co­

lumbus in 1492 boosted the perpetual interest in new, and partially know n plant species. Exotic płants were brought to the collections o f the bo- tanical gardens that were emerging at that time.

As the result botanical gardens were quite often the very spots from where alien species started their spread into new territories, beyond their natural ranges o f distribution. At the same time, together with the introduction o f new plant spe­

cies to garden collections, documentation such as publications and herbaria started to emerge.

One o f the earliest herbal studies devoted, inter 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

tury, saw the publication of a work by Hieronim Spiczyński (1542) under the title O ziołach tutecz- nych у zamorskich у o mocy ich [On herbs native and coming from overseas and their effects].

Information on alien species which the contem- porary botanical science characterised as more recent newcomers, or kenophytes, was included in works by Sir en iu s9 ( Syreń ski 1613) and Kluk

9 For е х а т р іе , S irenius m entioned A co ru s calam us (Sw eet-flag) using old Polish nam e: “ calam us” .

(1786, 1787, 1788). The latter author described several hundred “native wild plants and alien plants which could be o f use in our country” (“rośliny krajowe dzikie oraz i cudzoziemskie, któreby w kraju pożyteczne być mogły”). Most o f the spe­

cies mentioned by Kluk were cultivated at that time (e.g. Aesculus hippocastanum, Artemisia dracuncu- lus, Bryonia alba, Clematis vitalba, Helianthus tuberosus, Hyssopus officinalis, Juglans regia, Robinia pseudoacacia, Rubus odoratus, Sedum album, Sinapis alba) and are now considered to be naturalised in the flora o f our country. For certain species, some details o f their status outside the cultivated state are also included along with a description o f the type o f habitats entered by these species10.

T h e s tu d ie s o f s p e c ie s o f f o r e ig n o r ig in w e re firs t in c lu d e d in a b r o a d ly d e f in e d d is c ip lin e o f s tu d ie s in p l a n t g e o g r a p h y . Prior to D a r w i n ’s studies, i.e. ro u g h ly till th e m id 19th cen tu ry , m o s t o f th e r e s e a r c h a c tiv itie s c o n c e n tr a te d a r o u n d th e c o lle c tio n o f fa c ts p e r ta in in g to th e o c c u r r e n c e o f s p e c ie s a n d th e d if f e r e n tia tio n b e tw e e n th e v e - g e ta tio n la n d s c a p e s o f th e w o r ld ( Kornaś & Med-

w ecka-Kornaś 2002). In th e 18th c e n tu ry , u n d e r th e in flu e n c e o f w o rk c o m p le te d b y C a rl L in n a e u s , th e first flo ris tic a c c o u n ts a p p e a r e d in E u r o p ę , to b e c o n tin u e d in th e c e n tu r ie s th a t f o llo w e d (e.g . Willdenow 1787; Ficinus 1821; Re ic h en b a c h

1842; Peck 1865; Nymann 1878-1882; Schulze

1881; Schm alhausen 1886). T h e d e s c r ip tio n s o f fo re ig n n e w c o m e r s in th e s e flo ra s w e r e a ls o c o u - p le d w ith in itia l a tte m p ts to m a k e in v e n to rie s o f p la n t s p e c ie s o c c u r rin g in E u r o p e a n to w n s , s o m e o f th e m m a d e a s e a rly a s th e b e g in n in g o f th e 1 7 th c e n tu ry (cf. Jackow iak 1990, 1993, 1998a; Sud-

n ik-Wójcikow ska 1987a, 1998a a n d r e f e r e n c e s in th o s e p a p e rs ). T h e c h e c k lis ts o f u r b a n flo ra s a re o f p a r tic u la r im p o r ta n c e in s tu d ie s o f s p e c ie s o f fo re ig n o rig in b e c a u s e to w n s a re u s u a lly th e p la c e s w h e r e th e s e f o r e ig n n e w c o m e r s a p p e a r f o r th e firs t tim e . T h e o ld e s t s tu d ie s o f th is ty p e in P o la n d in c lu d e w o r k s p e r ta in in g to th e W a r s a w re g io n , p u b lis h e d b y B e m h a rd i in 1652 a n d E m d tl in 1730 (Su d n ik-Wójcik ow ska 1987a)".

10 e.g. A corus calam us — regarded by both authors as occurring near w ater; A m brosia a rtem isiifo lia - species described by K luk as o ccu rrin g on sandy sites; D atura stram onium - as early as in the tim es o f these authors, this plant com m only occurred in a w ild state, near fences, on yards and courts; M alva m oschata - found in scrub;

M ercurialis annua - in orchards and grassy sites; Portu- laca oleracea - a plant cultivated in gardens, capable o f spreading on its own throughout garden sites.

11 System atic studies o f urban floras started in Poland as early as at the end o f the 1901 century. T hese types o f studies becam e very com m on in the 1970s. A review o f the studies on the floras o f Central E uropean tow ns and the synthesis o f the m ain findings are presented by Ja c k o w ia k

(1998a) and Su d n i k- Wó j c i k o w s k a (1998a).

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T h e sto rm y h is to ry o f P o la n d , a c o u n try w h ic h p ra c tic a lly w e n t o u t o f e x is te n c e b e tw e e n th e day o f th e a b d ic a tio n o f K in g S ta n is la u s A u g u s tu s (25 N o v e m b e r 1795) till th e d a y it r e g a in e d its in d e p e n d e n c e (11 N o v e m b e r 1918), b e in g n o th in g m o re b u t a n a m e (Davies2001), d id n o t fa v o u r any s y s te m a tic c o lle c tio n o f flo ris tic d a ta . W ith in th a t p e rio d , th e re w e re o n ly flo ris tic s tu d ie s d e v o te d to lo c a l flo ra s (Mattuschka 1776, 1777, 1779;

Krocker 1787, 1790, 1814, 1823; Besser 1809;

Gunther et al. 1824; Adam ski 1828; Schneider

1837; Wim mer 1841; Grabow ski 1843; Ritschl

1850 a n d o th e rs ). T h e s e w e re m o s tly w o rk s b y G e rm a n n a tu r a lis ts a n d p e r ta in e d to th e a re a s w h ic h b e c a m e in c lu d e d in th e a d m in is tra tiv e b o r- d e rs o f P o la n d a fte r W o rld W a r II.

The oldest systematic study o f the flora o f Po­

land is the work by Jakub Waga, who was one of the outstanding Polish botanists o f the first half of the 19th century (RostańskiK. 2 00la). This work, published in 1847, includes “botanical descriptions o f plants, both wild and cultivated in open areas, within the Kingdom o f Poland” (“botaniczne opisy tak dziko jako i hodowanych pod otwartem nie­

bem jawnokwiatowych Królestwa Polskiego roś­

lin”). According to Rostański K. (200la), this first comprehensive study o f the flora o f what was then called the “Congress” Kingdom o f Poland was of the same level and form as other floras of vascu- lar plants from the neighbouring areas o f Prussia, Silesia (Polish), Galicia and Lithuania. Waga

(1847) listed a total o f more than a thousand species o f flow ering plants including several dozens of those currently classified as kenophytes - at that time these were either already established or merely present in cultivation (cf. Chapter 5.2).

The development o f naturalists' studies under- taken in Poland in the second half o f the 19th cen­

tury was associated with the short-lived activities o f Szkoła Główna (a higher education establish­

ment) opened in 1862, from which some botanists graduated: Karo (1867 - Flora o f the Warszawa region, 1881 —Flora o f the Częstochowa region) and Rostafiński, the author o f a 1872 treatise Flo- rae Polonicae Prodromus. At the same time, there were floras of the Pomeranian regions and towns (e.g. Klinggraeff 1848, 1854, 1866), Silesia (Uechtritz 1865; Fiek 1881) and Polish Galicia

(Berdau 1859; Knapp 1872). These publications

are a valuable source for the first record data for many species of kenophytes (cf. Appendix A &

B ). The first half of the 20th century, up until the outbreak o f World War II, saw further regional Floras published where authors, apart from de- scribing native species, also included species o f foreign origin. Particularly noteworthy were the works by German botanists providing information on flora composition and localities for many plant

species from Silesia (Sch u be 1901a, b-1930;

Schalow 1931-1936) and Pomerania (e.g. Abro-

meitet al. 1898-1940). Rich materiał regarding the Silesian flora was summarised in a work by Schube (1903b, 1904), and the Silesian flora was reputed to be one o f the best known floras in Europę of that tim e12 (Sendek 1981). In Galicia (south-eastem Poland), an important work - but unfortunately unfinished - in the field o f floristic research was Conspectus Florae Galiciae Criticus

by Zapalow icz (1906, 1908, 1911).

Another important source o f information was the naturalists’ joumals, which began to be published as early as in the second half o f the 19th century, such as: Wszechświat, Pam iętnik fizjograficzny, Kosmos, Sprawozdania Kom isji Fizjograficznej PAU, Dohrniana, Jahres-Bericht der Schlesischen G esellschaft fu r vaterlandische Cultur. These journals published floristic notes and accounts of bo­

tanical trips across various regions o f contemporary Poland, and also included - apart from the records of native species - new localities for many new alien species, coupled with their probable routes into new territories (Unverricht 1847; Rehm an 1868;

Krupa 1877; Kamieński 1879,1884a & b; Uechtritz

1879, 1880; Łapczyński 1882, 1887, 1888, 1889, 1890; Raciborski 1884, 1885; Błoński 1892;

Cybulski 1894, 1895; Schube 1901-1930; Meyer

1931, 1932; Schalow 1931-1936 and others).

Further systematic floristic inventories were com- pleted in many regions o f Poland in the 1960s and 70s. This period yielded many records and check- lists contributing to local and regional floras.

An outline o f the history o f floristic studies as well as the main currents o f research, taking into account or sometimes devoted exclusively to plants o f foreign origin, are presented in Table 1.

Particularly significant contributions were made by those studies which concentrated on recording the appearance of new species in local floras and gath- ering data on their stations. Articles published in a series Studies o f distribution ranges o f synan- thropic plants by Trzciń sk a (1961), Św iebo d a

(1963); Trzciń sk a-Tacik (1963); Zają c E.U. &

ZającA. (1973) and Guzik & Su d nik- Wó jcik o w-

ska(1989) are pioneering works on the reconstruc- tion of the history o f spread by the synanthropic newcomers. Much attention was also given to the classification o f plants accompanying humans and to compiling checklists o f species o f foreign ori­

gin occurring in Poland (Table 1; Fig. 1).

12 Silesia had its w ildlife particularly well researched even earlier, because the first study o f this area was pub­

lished in the 17,h century by C aspar Sc h w e n c k f e l d (1600).

More Silesian floras were published by Ma t t u s c h k a (1776, 1777, 1779), Kr o c k e r (1787, 1790, 1814, 1823), Wi m m e r

& Gr a b o w s k i (1 8 2 7 -1 8 2 9 ), Fie k (1881), Sc h u b e (1904) and Pa x (1915), after Mu l a r c z y k (2000).

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Table 1. S e l e c t c d p a p e r s c o n c e r n i n g d i f f e r e n t a s p e c t s o f s y n a n t h r o p i s a t i o n a n d s t u d i e s f o c u s s i n g s p c c i f i c a l l y o n a l i e n p l a n t s o c c u r r i n g i n P o l a n d ( i n c h r o n o l o g i c a l o r d e r )

Т у р е o f s t u d y A u t h o r / y e a r

Historical floras (regions, cities & towns)

C o n g r e s s K i n g d o m o f P o l a n d ( K r ó l e s t w o P o l s k i e ) Wa g a 1 8 4 7 ; Ro s t a f iń s k i 1 8 7 2

P o m e r a n i a ( P o m o r z e ) Kl in g g r a e f f 1 8 4 8 , 1 8 5 4 , 1 8 6 6 ; Ab r o m e i tet al. 1 8 9 8 - 1 9 4 0 ; De c k e r 1 9 1 1 ; Mu l l e r 1 9 1 1 ; Ho l z f u s s 1 9 3 7 ; St e f f e n 1 9 4 0 S i l e s i a ( Ś l ą s k ) Wi m m e r 1 8 4 1 ; Gr a b o w s k i 1 8 4 3 ; Fie k 1 8 8 1 ; Sc h u b e 1 9 0 3 b G a l i c i a ( G a l i c j a ) Be s s e r 1 8 0 9 ; Kn a p p 1 8 7 2 ; Za p a ł o w ic z 1 9 0 6 , 1 9 0 8 , 1 9 1 1 B o l e s ł a w i e c t o w n a n d v i c i n i t y ( B y t o m O d r z a ń s k i , J e d l i n a Z d r ó j ,

O ł a w a & W o ł ó w )

Sc h n e i d e r 1 8 3 7

P o z n a ń Ri t s c h l 1 8 5 0

K r a k ó w ( C r a c o w ) a n d s u r r o u n d i n g a r e a Be r d a u 1 8 5 9 ; Ra c ib o r s k i 1 8 8 4 ; Kr u p a 1 8 7 7 , 1 8 7 8 ; Żm u d a 1 9 2 0 W a r s z a w a ( W a r s a w ) a n d s u r r o u n d i n g a r e a Er n d t e l 1 7 3 0 ; Ka r o 1 8 6 7 ; Ła p c z y ń s k i 1 8 8 2 ; Cy b u l s k i 1 8 9 4 , 1 8 9 5 C z ę s t o c h o w a t o w n a n d s u r r o u n d i n g a r e a Ka r o 1 8 8 1

P r z e m y ś l t o w n a n d s u r r o u n d i n g a r e a Ko t u l a 1 8 8 1

B a b i a G ó r a M t . Za p a ł o w ic z 1 8 8 0

T a t r y , P i e n i n y & W e s t e r n B e s k i d y M t s . Be r d a u 1 8 9 0

New alien plant species

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

Acorus calamus, A m aranthus retroflexus, Chamomilla suaveolens, Conyza canadensis, Elodea canadensis, Galinsoga parvi/lora, Impatiens parviflora, Lycium barbarum,

Xanthium spinosum

Ka m ie ń s k i 1 8 8 4 a & b

N e w s p e c i e s r e c o r d e r e d i n t h e W a r s z a w a p r o v i n c e Cy b u l s k i 1 8 9 5

R a r e a n d c a s u a l p l a n t s Tr z e b iń s k i 1 9 3 0

N e w c o m e r s a n d w a n d e r i n g p l a n t s Sz u l c z e w s k i 1 9 3 1

Уегопіса filiform is Ko r n a ś & K u c 1 9 5 3 N e w c o m e r s i n t h e f l o r a o f B i a ł o w i e ż a F o r e s t So k o ł o w s k i 1 9 6 7 , 1 9 7 0

Corydalis lutea Be r n d t 1 9 5 8

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

Bromus carinatus Mi r e k 1 9 8 2 ( 1 9 8 4 )

Уегопіса peregrina Za j ą c M . & Za j ą c A . 1 9 9 0

Eragrostis multicaulis Gu z ik & Su d n i k- Wó j c i k o w s k a 1 9 9 4

Chaerophyllum aureum Ok l e j e w ic z 1 9 9 9

Reynoułria x bohemica Fo j c i k & To k a r s k a- Gu z ik 2 0 0 0 F o r m o r e s e e a l s o a p p e n d i x A a n d В

First localities

L i s t s o f p l a n t s p e c i e s t o g e t h e r w i t h t h e i r l o c a l i t i e s Ko r n a ś 1 9 5 0 , 1 9 5 4 ; Ur b a ń s k i 1 9 5 8 ; Żu k o w s k i 1 9 5 9 ; 1 9 6 0 a & b ; Ta c i k 1 9 6 0 ; Fa b i s z e w s k i & Fa l i ń s k i 1 9 6 3 ; So w a & Wó j c i k- - Ch r o b o k 1 9 6 9 ; Ro s t a ń s k i K . 1 9 6 0 , 1 9 6 1 ; Sc h w a r z 1 9 6 1 ; So w a 1 9 6 2 ; Ha n t z 1 9 6 7 , 1 9 7 2 ; Mi c h a l a k 1 9 6 8 , 1 9 7 1 ; Ko r n ia k 1 9 6 8 ; Tr z c i ń s k a- Ta c i k 1 9 7 1 a ; Mi c h a l a k & Se n d e k 1 9 7 4 - 1 9 7 5 ; Gł o w a c k i 1 9 7 5 ; Wik a 1 9 7 5 ; Ol e s iń s k i & Ko r n i a k 1 9 8 0 K e n o p h y t e s i n t h e f l o r a o f L u b l i n p r o v i n c e Fi j a ł k o w s k i 1 9 7 3

S y n a n t h r o p i c g r a s s e s Ko r n i a k 2 0 0 2

Synanthropic floras

F l o r a s o f c i t i e s : P o z n a ń G d a ń s k S z c z e c i n

Z i e l o n a G ó r a , K o s z a l i n Ł ó d ź

K r a k ó w

Kr a w ie c o w a 1 9 5 1 Sc h w a r z 1 9 6 7 Ćw i k l i ń s k i 1 9 7 0 Ćw i k l i ń s k i 1 9 7 1 So w a 1 9 7 4

Tr z c i ń s k a- Ta c i k 1 9 7 9

C o m p a r i s o n o f t h e u r b a n f l o r a s o n t h e e x a m p l e o f s o m e c i t i e s Kr a w ie c o w a & Ro s t a ń s k i 1 9 7 6 , 1 9 8 1

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Туре o f study Author/ year

F l o r a s o f t o w n s & s e t t l e m e n t s M ow szowicz 1 9 6 0 ; Sk o w r o ń s k a 1 9 6 5 ; Mi c h a l a k 1 9 7 0 ; Sc h w a r z 1 9 7 1 ; Se n d e k 1 9 7 1 ; So w a 1 9 7 1 ; An i o l- Kw ia t k o w s k a 1 9 7 4 ; Ha n t z 1 9 7 4 ; Sz m a j d a 1 9 7 4 ; Cz a p l e w s k a 1 9 7 5 ; Mis ie w ic z 1 9 7 8 ; Se n d e k & Wi k a 1 9 7 9 ; So w a & Na s i ł o w s k i 1 9 7 8 ; We r e t e l n i k 1 9 7 9 ; So w a & Wa r c h o l i ń s k a 1 9 8 0 ; Mi s i e w ic z 1 9 8 1 ; So w a & Wa r c h o l iń s k a 1 9 8 l a , b & c , 1 9 8 4 a & b , 1 9 8 7 ; Ma c i e j c z a k 1 9 8 8 ; Ćw i k l i ń s k i & Ba r t n i k 1 9 9 0 ; To k a r s k a- - Gu z ik & Ro s t a ń s k i 1 9 9 7 , 1 9 9 8

F u l i c a r t o g r a p h i c d e s c r i p t i o n o f t h e u r b a n f l o r a : W a r s z a w a

P o z n a ń J a w o r z n o

Su d n i k- Wó j c i k o w s k a 1 9 8 7 a Ja c k o w ia k 1 9 9 0 , 1 9 9 3 To k a r s k a- Gu z ik 1 9 9 9

Ruderal floras in the rural landscape o f the North Podlasie Lowlands

Wo l k o w y c k i 1 9 9 7

Regions - examples:

W ielkopolska province Gorce Mts.

Tatry Mts.

Wielkopolski National Park Karkonoski National Park

L u b l i n p r o v i n c e

Upper Silesia Industrial Region

eastern part o f the Gniezno Lake District Zaodrze (to the West o f Szczecin)

S ł o w i ń s k i N a t i o n a l P a r k

Sz u l c z e w s k i 1 9 5 1 Ko r n a ś 1 9 5 7 , 1 9 6 6

Ra d w a ń s k a- Pa r y s k a 1 9 6 3 ; Pi ę k o ś- Mir k o w a & Mir e k 1 9 7 8 S z u l c z e w s k i 1 9 6 3 ; Ż u k o w s k i et al. 1 9 9 5

Ro s t a ń s k i K . 1 9 7 7 , 1 9 7 8 Fija ł k o w s k i 1 9 7 8 Se n d e k 1 9 8 1 , 1 9 8 4 Ch m ie l 1 9 9 3 Z a j ą c A . et al. 1 9 9 3 P i o t r o w s k a et al. 1 9 9 7

S e g e t a l f l o r a W n u k 1 9 7 6 ; S o w a & W a r c h o l i ń s k a 1 9 7 9 ; W a r c h o l i ń s k a

1 9 8 1 , 1 9 9 6 ; W n u k et al. 1 9 8 9 ; S i c i ń s k i 1 9 9 7 , 2 0 0 0 ; L a t o w s k i 1 9 9 8 , 1 9 9 9 ; T r z c i ń s k a - T a c i k 1 9 9 6 ; W a r c h o l i ń s k a & S i c i ń s k i 1 9 9 6 ; W a r c h o l i ń s k a & T y s z k o w s k a 2 0 0 0 a l s o J a c k o w i a k &

L a t o w s k i 1 9 9 6 , 2 0 0 1 ; M i s i e w i c z & P i o t r o w s k i ( e d s . ) 1 9 9 6 ; R o l a 1 9 9 6 a n d l i t e r a t u r e c i t e d t h e r e i n

Ruderal plant communities

R e g i o n s Ko r n a ś 1 9 5 2 ; So w a 1 9 7 1

C i t i e s & t o w n s Fij a ł k o w s k i 1 9 6 3 , 1 9 6 7 ; Ro s t a ń s k i K . & Gu t t e 1 9 7 1 ; An i o ł- - Kw ia t k o w s k a 1 9 7 4 ; Kę p c z y ń s k i & Zie n k ie w ic z 1 9 7 4 ; Za ją cE.U.

1 9 7 4 ; Kę p c z y ń s k i 1 9 7 5 ; Cz a p l e w s k a 1 9 8 0 ; Św ię s & Pl e b a n 1 9 8 1 ; Św ię s 1 9 8 3

S p e c i a l h a b i t a t s Cz a p l e w s k a 1 9 8 1

Alien plants in special habitats

Railways and railway stations M e y e r 1 9 3 1 , 1 9 3 2 ; K o r n a ś et al. 1 9 5 9 ; S o w a 1 9 6 6 ; Ć w i k l i ń s k i 1 9 6 8 , 1 9 7 2 a , 1 9 7 4 ; K r a w i e c o w a 1 9 6 8 a ; S e n d e k 1 9 6 9 , 1 9 7 3 ; Z a j ą c E . U . & Z a j ą c A . 1 9 6 9 ; L a t o w s k i 1 9 7 7 ; Ć w i k l i ń s k i 1 9 8 4 —

1 9 8 5 ; W i k a 1 9 8 4

Storę yards (including ballast plants) H e l m 1 8 8 1 ; H o l z f u s s 1 9 3 6 , 1 9 4 1

Sea & river harbors Ro s t a ń s k iK. & Sz o t k o w s k i 1 9 7 3 ; Mis i e w ic z 1 9 7 6 , 1 9 8 5 , 2 0 0 1

Walls We r e t e l n i k 1 9 7 3 , 1 9 8 2 ; Św ie r k o s z 1 9 9 3 ; Ga l e r a & Su d n i k- - Wó j c i k o w s k a 2 0 0 0 a & b

Lists of alien plant species

Kenophytes K o r n a ś 1 9 6 8 b ; Z a j ą c A . et al. 1 9 9 8

Archaeophytes Za j ą cA . 1 9 7 9 , 1 9 8 3 , 1 9 8 7 a & b , 1 9 8 8

Ephemerophytes Ro s t a ń s k i K . & So w a 1 9 8 6 - 1 9 8 7

American trees and shrubs He r e ź n i a k 1 9 9 2

Kenophytes o f American origin So w a & Wa r c h o l iń s k a 1 9 9 4

Anthropophytes M i r e k et al. 1 9 9 5 , 2 0 0 2

Naturalised alien plants - neophytes (excluding archaeophytes) To k a r s k a- Gu z ik 2 0 0 3 a

Terminology & classification

Classification o f synanthropic plants Ko r n a ś 1 9 6 8 а , 1 9 7 7 a & b ; Kr a w ie c o w a & Ro s t a ń s k i 1 9 7 2 ; Mi r e k 1 9 8 l a

Neophytes & neophytism Fa l iń s k i 1 9 6 8 a & b , 1 9 6 9

3 The Establishm ent. 17

Cytaty

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When analysing the percentages of the representatives of 12 most spe- cies-rich families in the flora of the permanently established plants in Poland, as compared to the flora

Aliena- cja od wytworu pracy oznacza, że szeroko rozumiany twór pracy człowieka (nie tylko produkty materialne, ale także instytucje, produkty działalności naukowej,

alien tree species, growing outside their natural range, can establish a mycorrhizal symbiosis with cosmopolitan fungi, both in the natural range of the alien tree species as well