Title: Changes in richness in some riparian plant communities as an result of their colonisation by taxa of ()

Author: Barbara Tokarska-Guzik, Katarzyna Bzdęga, Dagmara Knapik, Grzegorz Jenczała

Citation style: Tokarska-Guzik Barbara, Bzdęga Katarzyna, Knapik Dagmara, Jenczała Grzegorz. (2006). Changes in plant species richness in some riparian plant communities as an result of their colonisation by taxa of Reynoutria (Fallopia). "Biodiversity Research and Conservation" (vol. 1/2 (2006), s. 123-130).

Biodiv. Res. Conserv. 1-2: 123-130, 2006 BRC www.brc.amu.edu.pl

Changes in plant species richness in some riparian plant communities as a result of their colonisation by taxa of Reynoutria (Fallopia)

Barbara Tokarska-Guzik1, Katarzyna BzdÍga2, Dagmara Knapik & Grzegorz Jencza≥a

Department of Plant Systematics, Faculty of Biology and Environmental Protection, University of Silesia, JagielloÒska 28, 40-032 Katowice, Poland, e-mail: [email protected], [email protected]

Abstract: This study deals with the current issue of the synanthropisation of plant cover which manifests itself, amongst other ways, in the spreading of species outside the limits of their natural distribution range. The aim of the studies undertaken for the present report was to estimate the type and scale of threat posed by alien plant species to native plant diversity. The studies were carried out using the example of Reynoutria (Fallopia) species (knotweeds), which are considered to be invasive in our country as well as in other regions of the world. This is an attempt to determine the impact of these on the composition and diversity of the natural components of the herb layer in floodplain forest. Investigations were performed on permanent study plots localised in patches of floodplain forest which have been preserved in the valleys of the So≥a, Bia≥a and Jasienica rivers. The results of the field studies have confirmed the hypothesis that Reynoutria species exert a negative influence on the native components of the floodplain forest herb layer. This impact is, however, different with regard to the various life forms and ecological habitat groups of the plants. The increase of surface coverd by knotweeds does not significantly influence the development of early spring geophytes which are able to complete their full life cycle.

Key words: , Reynoutria, Fallopia, riverside floodplain forests, ecological impact

1. Introduction cultivation, these plants have spread both to ruderal habitats and to sites of natural character (river valleys, The aim of the investigation was to estimate the type scrub margins and broadleaf forests). In Poland, as in other and scope of threat posed by alien plant species to the parts of Central Europe, two species occur at present: native biological diversity, especially in view of the fact Reynoutria (Fallopia) japonica Houtt. [ that these problems have hitherto been rarely addressed cuspidatum; Japanese Knotweed], Reynoutria (Fallopia) in Poland. The studies were carried out on the example sachalinensis (F. Schmidt) Nakai [Polygonum of Reynoutria (Fallopia) species (knotweed) which are sachalinense; Giant Knotweed], as well as the hybrid considered to be invasive plants in Poland (Fojcik & between these two species described in the 1980s: R. Tokarska-Guzik 2000; Tokarska-Guzik 2002, 2005; ◊bohemica Chrtek & Chrtkov·. is Tokarska-Guzik & Dajdok 2004) as well as in other fairly widespread over the whole national territory, regions of Europe and the world (e.g. Pyöek & Prach especially in its southern part, while R. sachalinensis is 1993; Brock et al. 1995; Sukopp & Starfinger 1995; scattered across the whole of Poland, locally rarer Bailey et al. 1995; Bailey 1999). (Tokarska-Guzik 2001; Zajπc & Zajπc 2001). The exact Taxa from the genus Reynoutria are conspicuous distribution of the hybrid remains to be investigated in rhizomatous perennials of large dimensions, spreading detail (Fojcik & Tokarska-Guzik 2000). mainly through vegetative processes, which were The present study is an attempt to determine the introduced to Europe from Asia at the beginning of the impact these plants have on the composition and 19th century as ornamental plants with numerous useful diversity of natural components of the floodplain forest characteristics (Bailey & Conolly 2000). From sites of herb layer. ECOLOGY

© Adam Mickiewicz University in PoznaÒ (Poland), Department of Plant . All rights reserved. 124 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...

2. Material and methods Geophytes and therophytes typically have also a relatively large share in the flora. Their high number is Field studies were carried out during the growing due to the special characteristics of riverside habitats. seasons of 2002-2004 in estuary segments of the So≥a A higher proportion of therophytes relative to geophytes and Bia≥a river valleys as well as in the middle section in the Bia≥a and Jasienica river valleys is linked to the of the Jasienica river valley (Fig. 1). progressive habitat transformation. Native species dominate in the flora of the river valleys investigated, constituting 85% of the Jasienica, 79% of the So≥a and 76% of the Bia≥a, respectively. A characteristic feature of the flora and vegetation is the participation of taxa from the genus Reynoutria, which in the case of the Bia≥a river have invaded the river banks on a massive scale. Ecological habitat groups are the indicator which varies most between the river valley segments analysed. On the So≥a, species from fertile deciduous forests (Querco-Fagetea) and scrub communities, riverside for- ests and thickets (Salicetea purpureae) as well as meadow communities (Molinio-Arrhenatheretea) are dominant. On the Bia≥a and Jasienica, ruderal and therophytic communities occur more frequently. The composition and diversity of species in the herb layer of the floodplain forests selected for the study was dependent on the extent to which the investigated plot was covered by knotweed . The largest num- ber of species was recorded from plots without any knotweeds (max. 28). In plots with increasing coverage of the invasive species (up to 40% and 41%- 70%), a gradual reduction of the number of other spe- cies was noted. Sudden and extensive decreases in the Fig. 1. Location of investigation plots number of herb layer species (minimum 3 species) and their coverage coefficients was seen in plots with a high Investigations were performed on permanent study proportion of knotweed (71%-100%) (Fig. 2; compare plots (20 m x 20 m) localised in patches of floodplain also Table 1 and 2). The number of species recorded forest. The plots were selected to describe varying on the study plots was different depending on the sea- degree of surface coverage by knotweeds. The total son of the vegetative period (in spring, this number was number of plots investigated was 36 and in each of them lower than in summer). In early spring, spring geophytes 2 phytosociological relevËs were taken in different were recorded on the study plots (e.g. Ficaria verna, growth seasons: one in spring and one in late summer. Symphytum tuberosum, Anemone nemorosa, Dentaria Additionally, within three study plots, measurements glandulosa, Primula elatior). Their development is not of longitudinal Reynoutria growth rate were performed on 30 shoots selected at random. The studies were supplemented by a detailed inventory of the vascular flora occurring in the investigated sections of the river valleys.

3. Results and discussion

A total number of 451 vascular plant species were found to occur in the study area: 420 species in the section of the So≥a river valley, 211 in the Bia≥a river valley and 160 in the Jasienica stream valley. The vascular flora of the river valleys investigated is characterised by the highest proportion of hemicrypto- phytes which account for 50% species on the So≥a, 51% Fig. 2. Species richness in plots grouped by % cover of Reynoutria on the Jasienica and 54.5% on the Bia≥a, respectively. during different seasons of growth Biodiv. Res. Conserv. 1-2: 123-130, 2006 125 25r; Taraxacum officinale 4(+); ñ spring aspect Sanguisorba officinalis 4r; Reynoutria Heracleum sphondylium 4r; 11(1.2) Filipendula ulmaria ñ Solidago gigantea 25(+.2); Molinio-Arrhenatheretea 25(+); Solidago canadensis 4r; Stachys sylvatica 20r; Cirsium vulgare ñ ñ Changes in riparian plant communities herb layer composition and cover as a result of colonization by taxa of Athyrium filix-femina Table 1. ñ ñ Artemisietea vulgaris 25(+); Querco-Fagetea Veronica chamaedrys Sporadic species: 126 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...

significantly influenced by the growth of knotweed Aegopodium podagraria and Petasites hybridus. shoots during this period (Fig. 3). These species have Characteristic and differentiating species for the class enough time to go through their full life cycle. In the Salicetea purpureae had their most abundant occurrence case of this group of plants, a decrease in the coverage in plots without any knotweed and with small and coefficient can only be seen in plots with a massive medium coverage by knotweed plants (Fig. 4). Lack of occurrence of knotweeds (Table 1). In the summer season, light and saturation of the soil with on plots

Fig. 3. The average rate of longitudinal growth of Reynoutria shoots measured on a sample of 30 shoots selected randomly

the rapid growth of shoots and especially the development with simultaneous development of . In the later of knotweed leaves leads to a conspicuous reduction in period within the growth season, the rate of longitudinal the vegetation of those species for which the peak of shoot growth is significantly slower (during the first the life cycle falls in the summer period. The development half of June, the shoots reach their maximal height); of knotweed plants begins, depending on seasonal lateral shoots develop during this time. condition variations, in mid-April with the appearance On the study plots, the proportion of the most of shoot tips which develop from subterraneous rhizomes. frequently recorded species has been compared with In the period between end of April and mid-May, rapid regard to the coefficient of coverage by Reynoutria (very sudden) longitudinal growth of the shoots occurs shoots. The following species were included: Ficaria (initially between 6 and 12 centimetres per 4 days; verna, Symphytum tuberosum, Urtica dioica, Rubus subsequently even as much as 40 cm per 4 days; Fig. 3) caesius, Phalaris arundinacea, Calystegia sepium,

I – Reynoutria absent III – Reynoutria 41%-70% cover

II – Reynoutria up to 40% cover IV – Reynoutria 71%-100% cover

Fig. 4. Cover of species characteristic of the Salicetea purpurea community relative to cover of Reynoutria Biodiv. Res. Conserv. 1-2: 123-130, 2006 127 from group IV (70-100% of Reynoutria) has effectively maturation to reproduction; resistance to extreme eliminated most characteristic species for this class. The environmental conditions; dioecy, polygamy, self- only exceptions were Urtica dioica (a perennial) pollination; ability to compete with other species (due and Calystegia sepium (a creeper). to height, production of allelopathic substances etc.); In the study plots without any knotweed or with low phenotypical variability, ability to form mutants, cover by knotweed plants, the presence of the charac- polyploids, hybrids with related species. Taxa from the teristic species for most of the ecological habitat groups genus Reynoutria have many of the above-mentioned was confirmed. In these plots, a high proportion of spe- features. Above all, they belong to the group of plants cies from thicket, scrub and forest facies was found. with the most efficient mode of vegetative reproduction Their number drops conspicuously with an increase in by rhizome segmentation (Alberternst et al. 1995; Child the share of knotweed plants (group III plots). A high 1999; Fojcik & Tokarska-Guzik 2000). Potentially, share of Reynoutria (70-100%) in group IV plots led to a rhizome fragment which is only 1 cm in length and a reduction in most ecological habitat groups. 0.7 g in weight may generate a new plant (Brock et al. The gradual elimination of herb layer species is 1995). Any transformations of the environment caused mainly caused by the expansion of knotweeds in the by natural factors (e.g. floods) as well as in particular habitat investigated. Their sudden growth at the very by anthropogenic factors (earthworks, river regulation) beginning of the vegetative season with a simultaneous favour the spreading of rhizomes (Pyöek & Prach 1993). development of large leaves (their size in R. sachalinensis Regulation of the Bia≥a river and Jasienica stream as is 15-43 cm long and 10-27 cm broad; after Fojcik & well as earthworks conducted in the So≥a river valley Tokarska-Guzik 2000) causes strong shading of the soil have contributed to a major extent to the colonisation and elimination of other species. Lack of light availability of large areas within the study zone by Reynoutria spe- near the surface of the ground is additionally aggravated cies. The growth rate of knotweed plants is especially by the characteristics zigzag pattern of the high at the beginning of the vegetative season: 43.1 mm/ arrangement along the knotweed stem. Furthermore, day (Child 1999); 80 mm/day (Seiger 1997). Furthermore knotweed rhizomes thrive in the soil to an extent which the soil which is thickly occupied by knotweed shoots prevents many other species from taking root, germinating (down to 7 m in depth) does not allow the germination and going through the entire life cycle. of many species. According to the classification of life The similar results were published by Sukopp & strategies (Grime 1979) knotweeds represent a C-type Sukopp (1988), Schepker (1998) and Schl¸pmann (2000), strategy, which decisively increases their invasive po- stated that one of the most serious problems is that tential (Pyöek & Prach 1993). The following species Reynoutria modifies or expels the autochthonous veg- are able to compete successfully with knotweeds even etation by shadowing. Investigations conducted by if the plants are present at high coverage coefficient: Alberternst (1998) show that the number of plant spe- Aegopodium podagraria, Urtica dioica (rhizome cies in vegetation transects with Fallopia were lower perennials with similar mode of growth), balsams: than in transects without the species. Impatiens parviflora and I. glandulifera (alien annuals The biological properties of representatives of genus with an R-type life strategy and shallow root systems) Reynoutria cause these plants to have a definite reduction as well as Calystegia sepium (a creeper). impact on the native components of plant communities (Tokarska-Guzik & Dajdok 2004; Tokarska-Guzik 4. Conclusions 2005). The enormous potential of these species for spreading by vegetative means, combined with their The results of field studies have confirmed the rapid growth and a capacity to adapt to diverse or even hypothesis that representatives of genus Reynoutria extreme habitat conditions, often invading and holding exert a reducing influence on the native components of large areas, have resulted in this species earning the floodplain forest herb layer. This impact is, however, status of invasive plant and nuisance íweedí (Tokarska- different with regard to various life forms and ecological Guzik 2005). habitat groups of plants. Increase of surface coverage The characteristics influencing the rate and success by knotweeds does not significantly influence the of alien plant invasions listed by FaliÒski (2004) include development of early spring geophytes which are able high fertility and reproductive potential of each to go through their entire life cycle. In the case of this individual, persistence of seeds (propagules); seeds group of plants, a decrease in coverage coefficient can equipped with devices which allow rapid long-distance only be seen in plots with massive occurrence of knot- transport; accelerated growth at the juvenile stage, early weeds.

128 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ... Constancy ñ late summer aspect Reynoutria absent - Group I up to 40% - Group II 41 to 70% - Group III 71 to 100% - Group IV Reynoutria ...... 3.5 3.4 3.5 2.2 2.3 . . . 3.5 4.5 5.5 5.5 5.5 . . . 5.5 5.5 5.5 5.5 5.5 . . . III .....33...... 1.1...... 3.31.1...... 2.2 . 2.2 . 1.1 . . 2.2 2.2 . . 1.1 ...... 3.3 . . . . 1.1 . 3.3 2.2 . 2.2 . . + . 2.3 . + 3.5 ...... 3.4 I ...... +.....II . 2.2. . 2.2 . 2.2 ...... 2.2...... 1.1 + 1.1 . . . 2.2 + 4.5 . . . + 4.5 4.5 1.1 . . . . 1.1 . . 1.1 ...... 2.4 ...... 2.3 1.1 . 2.3 . . . 2.4 5.5 2.3 5.5 . 2.3 5.5 r . . II . . . I . III ...... + ...... +...... I ...... +...... I ------80 80 90 80 90 90 80 80 90 100 90 30 40 100 90 30 10 10 ------1 2 3 4 5 6 7 8 9 1011121314151617181920212223242526272829303132 22...... I + 2.2+2.2..2.2... . .+ 2.2 . . 2.3 . 2.2 3.3 2.2 . 1.2 3.3 2.2 1.1 1.2 + r 2.2 + . 2.2 2.3 . 2.3 1.2 . r . 2.2 1.2 2.2 . . r 2.2 . . 1.2 . . . r . r ...... IV . II 70 50 70 60 50 60 70 50 10 60 10 60 70 70 40 60 70 50 20 10 - 50 30 30 30 - 5 - - - 5 - 3.3 2.22.3 3.3 3.3 2.2 1.2 1.2 2.2 + 3.3 2.3 1.2 + 3.2 1.2 3.3 2.3 1.2 3.3 2.2 2.2 2.2 2.2 . 3.2 3.3 2.2 2.2 2.2 3.3 2.2 2.2 2.2 +.2 2.2 +.2 2.3 3.3 3.3 2.2 1.2 +.2 r 1.2 + . +.2 1.2 r 1.2 r . r . +.2 . r . V . . . . . IV 4.4 2.2 4.4 3.3 3.3 2.2 3.3 2.2 . 3.3 . 3.3 3.3 2.2 . 2.2 3.3 3.3 . + . 2.2 . + 2.2 ...... IV 3.3 3.3 2.3 3.4 3.3 3.3 1.2 2.2 1.2 2.2 4.4 2.2 3.3 2.2 1.2 1.2 2.2 3.3 + . +.2 1.2 + + r . + . . . r . III +.2 1.2 1.2 r 1.2 r + 1.2 +.2 +.2 +.2 1.2 1.2 1.2 +.2 +.2 r 1.2 +.2 + . 1.2 +.2 + . . + . . + + . III 100 90 80 90 90 90 80 80 90 70 90 30 60 60 90 70 80 90 100 100 100 90 100 100 100 100 100 100 100 100 100 100 11 11 09 --2030----1055----- 09 8 03 09 03 5S 8 09 03 6S 09 8 03 9S 30-405------09 10S ------12 03 ------12S ------5 09 04 1J 12 09 04 12 2J 5 04 3J 09 20 11 09 -10--20--- 03 11 3S 5 09 03 20 11 4S 5 09 03 15 11 7S 09 03 15 8S 5 8 09 03 19 9S 20 12 09 04 22 4J 12 5 09 21 04 5J 12 18 04 09 6J 50 11 20 09 30 03 1S 11 09 03 10 15 2S 09 8 13S 03 30 18 3B 09 03 10 17 20 7B 09 03 10 18 17 7J 09 04 50 16 20 8J 09 04 30 17 20 9J 04 20 20 09 30 1B 18 09 03 17 5 2B 09 03 22 17 4B 20 09 03 16 17 5B 10 09 03 6B 13 17 10J - 09 03 14 17 11J 09 04 12J 20 20 5 09 04 18 20 20 04 17 20 10 14 10 9 5 8 5 11 5 5 3 5 7 11 6

1 1 1 2 1 2 1 1 1 2 2 ...... c...+...... b . +b 1.2 . + 2.2 . . . 1.2 + . 3.3 + 2.2 . . 1.1 . . + ...... 2.2 1.1 . 1.1 ....1.1... b...... 1.1 . .b . . + + . . + 1.1 ...... 1.1 . . . r ...... 2.2 . . . 1.1 . . . 1.1 ...... 2.2 + . . 2.2 . . 1.2 . . + . . . 1.2 ...... II ..+.2211...... I ...... I b....+...2.2.1.12.2...... 2.2..... b...... c c a a a a a a a a a in % Changes in riparian plant communities herb layer composition and cover as a result of colonization by taxa of Reynoutria Table 2. Number of the relevè Day Month Year Locality and number of plot Cover of tree layer a1 in % Cover of tree layer a2 in % Cover of shrub layer b in % Cover of herb layer c1 in % Cover of herb layer c2 in % Cover of moss layer d in % Number of species in the relevè Cover of Reynoutria japonica Reynoutria xbohemica Reynoutria sachalinensis Salicetea purpureae Salix fragilis Salix fragilis Salix fragilis Salix alba Salix alba Salix alba Salix alba Populus nigra Populus alba Salix purpurea Salix viminalis Rubus caesius Calystegia sepium Phragmitetea Phalaris arundinacea Phragmites australis Carex gracilis Alno-Ulmion Alnus glutinosa Alnus glutinosa Alnus glutinosa Alnus incana Padus avium Urtica dioica Petasites hybridus Biodiv. Res. Conserv. 1-2: 123-130, 2006 129 ; Chenopodium Lythrum salicaria 21(+.2) 5(+.2); ; Polygonum persicaria ; Lysimachia vulgaris 16r 4r Daucus carota 4r; Robinia pseudacacia ñ

21r Equisetum palustre ñ Cerastium arvense 21r; 17(+.2); Accompying species Molinio-Arrhenatheretea Sonchus arvensis 20(+); Fallopia dumetorum

1 ; a 21(+.2); 11(+.2) Tilia platyphyllos 3r; Solidago gigantea Lycopersicon esculentum 5r; 21(+.2); .r...... I ...... r...... r....rr...... I ....r+.2...... ++...... I ...... I +.2 .. +.2 ...... I ..1.2...... r.... I . ....r... .+...+...... +.2r... I +.2 ...... rr.. r...... 2.2 I ...... r ....r... 1.2 ...... I .r...... 1.2 .+.2...... r . r ..r..rr...... r.r+rr ...... II ...... r ...... rr..rrr. I ..2.3+.2.+.2+...... r...r ...... r...... I ...... +.r. I . . 1.2 . +...... +.2 ....r.r...... I ...... + . +.2 + . . . r ...... r I r r III ...... r.... r...... I rrrrr..r+2r.r .rr....rIII rr....rr+.2.rr..rr .rrr.rrr ...... I ...... rrr .....r...... I ...... 2.3...... 11.+....I ...... 1.1...+...... I ..+..... r rrr..r..r .r...... r+.2..+.2+.2...... r.II 2.3 3.4 1.2 2.2 +.2 +.2 . +.2 . r 2.3 +.2 + + + + 2.3 . . + + + ...... IV rr....rr ...... r...... I .12...rr.. 12...... I r.r...... 1.2...... 1.21.2...... 1.2 1.2 1.2 + 1.2 + 1.2 1.2 3.3 2.3 r 2.2 2.2 2.2 3.3 2.3 3.3 2.3 +.2 + . 2.3 +.2 + + +.2 1.2 r . r r +.2 V 1.2 . . . . r . . 1.2 1.2 r r . . + 1.2 . . . 1.2 . . . . . + . +.2 +.2 . . . II +.2 . r ...... II r 1.2 r ..1.2...... r.r.r r ...... r...... I .+..r+.. 1.2 .+.2...+..+ +.2+.2...... r 1.2 . +.2 +.2 . . r 1.2 . . . 1.2 ...... III Equisetum sylvaticum Equisetum ñ ...... r...... r...... I ...... +.2...... +.2.I .....r...... r...... I

1 1 ...... c...... r...... 11...... 1.1....2.2...... b...... b .b ...... b++...r.. + I . . + ...... + + . 2.2 . 1.2 . . + 1.2 2.2 . 2.2 2.2 1.1 1.2 . . . . 1.1 1.2 1.2 + . . . . . 1.2 + . + . 1.1 ...... 1.1 . III . . . . . I a a Echinocystis lobata ñ ñ Querco-Fagetea Chenopodium polyspermum 21(+.2); Artemisietea vulgaris Lamium maculatum Humulus lupulus Myosoton aquaticum Circaea lutetiana Matteucia struthiopteris Querco-Fagetea Quercus robur Quercus robur Fraxinus excelsior Sambucus nigra Sambucus nigra Cornus sanguinea Euonymus europaea Aegopodium podagraria Athyrium filix-femina Molinio-Arrhenatheretea Poa trivialis Vicia cracca Symphytum officinale Anthriscus sylvestris Filipendula ulmaria Myosotis palustris Artemisietea vulgaris Impatiens glandulifera Impatiens parviflora Artemisia vulgaris Cirsium arvense Rumex obtustifolius Solidago canadensis Aster novi-belgii Tanacetum vulgare Melandrium album Rudbeckia laciniata Accompying species Galeopsis speciosa Poa annua Polygonum hydropiper Oxalis fontana Capsella bursa-pastoris Conyza canadensis Echinochloa crus-galli Galinsoga ciliata Plantago major Sporadic species: album 11(+); 130 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...

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