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Title: Changes in plant species richness in some riparian plant communities as an result of their colonisation by taxa of Reynoutria (Fallopia)
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).
© Adam Mickiewicz University in PoznaÒ (Poland), Department of Plant Taxonomy. All rights reserved.
BRCwww.brc.amu.edu.pl
Changes in plant species richness in some riparian plantcommunities 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 otherways, in the spreading of species outside the limits of their natural distribution range. The aim of the studies undertaken for thepresent report was to estimate the type and scale of threat posed by alien plant species to native plant diversity. The studieswere carried out using the example of Reynoutria (Fallopia) species (knotweeds), which are considered to be invasive in ourcountry as well as in other regions of the world. This is an attempt to determine the impact of these plants on the compositionand diversity of the natural components of the herb layer in floodplain forest. Investigations were performed on permanentstudy plots localised in patches of floodplain forest which have been preserved in the valleys of the So≥a, Bia≥a and Jasienicarivers. The results of the field studies have confirmed the hypothesis that Reynoutria species exert a negative influence on thenative components of the floodplain forest herb layer. This impact is, however, different with regard to the various life formsand ecological habitat groups of the plants. The increase of surface coverd by knotweeds does not significantly influence thedevelopment of early spring geophytes which are able to complete their full life cycle.
Key words: invasive species, Reynoutria, Fallopia, riverside floodplain forests, ecological impact
1. Introduction
The aim of the investigation was to estimate the typeand scope of threat posed by alien plant species to thenative biological diversity, especially in view of the factthat these problems have hitherto been rarely addressedin Poland. The studies were carried out on the exampleof Reynoutria (Fallopia) species (knotweed) which areconsidered to be invasive plants in Poland (Fojcik &Tokarska-Guzik 2000; Tokarska-Guzik 2002, 2005;Tokarska-Guzik & Dajdok 2004) as well as in otherregions of Europe and the world (e.g. Pyöek & Prach1993; Brock et al. 1995; Sukopp & Starfinger 1995;Bailey et al. 1995; Bailey 1999).
Taxa from the genus Reynoutria are conspicuousrhizomatous perennials of large dimensions, spreadingmainly through vegetative processes, which wereintroduced to Europe from Asia at the beginning of the19th century as ornamental plants with numerous usefulcharacteristics (Bailey & Conolly 2000). From sites of
cultivation, these plants have spread both to ruderalhabitats and to sites of natural character (river valleys,scrub margins and broadleaf forests). In Poland, as in otherparts of Central Europe, two species occur at present:Reynoutria (Fallopia) japonica Houtt. [Polygonumcuspidatum; Japanese Knotweed], Reynoutria (Fallopia)sachalinensis (F. Schmidt) Nakai [Polygonumsachalinense; Giant Knotweed], as well as the hybridbetween these two species described in the 1980s: R.◊bohemica Chrtek & Chrtkov·. Reynoutria japonica isfairly widespread over the whole national territory,especially in its southern part, while R. sachalinensis isscattered across the whole of Poland, locally rarer(Tokarska-Guzik 2001; Zajπc & Zajπc 2001). The exactdistribution of the hybrid remains to be investigated indetail (Fojcik & Tokarska-Guzik 2000).
The present study is an attempt to determine theimpact these plants have on the composition anddiversity of natural components of the floodplain forestherb layer.
Biodiv. Res. Conserv. 1-2: 123-130, 2006
EC
OL
OG
Y
124
2. Material and methods
Field studies were carried out during the growingseasons of 2002-2004 in estuary segments of the So≥aand Bia≥a river valleys as well as in the middle sectionof the Jasienica river valley (Fig. 1).
Investigations were performed on permanent studyplots (20 m x 20 m) localised in patches of floodplainforest. The plots were selected to describe varyingdegree of surface coverage by knotweeds. The totalnumber of plots investigated was 36 and in each of them2 phytosociological relevËs were taken in differentgrowth seasons: one in spring and one in late summer.Additionally, within three study plots, measurementsof longitudinal Reynoutria shoot growth rate wereperformed on 30 shoots selected at random. The studieswere supplemented by a detailed inventory of thevascular flora occurring in the investigated sections ofthe river valleys.
3. Results and discussion
A total number of 451 vascular plant species werefound to occur in the study area: 420 species in thesection of the So≥a river valley, 211 in the Bia≥a rivervalley and 160 in the Jasienica stream valley. Thevascular flora of the river valleys investigated ischaracterised by the highest proportion of hemicrypto-phytes which account for 50% species on the So≥a, 51%on the Jasienica and 54.5% on the Bia≥a, respectively.
Fig. 1. Location of investigation plots
Geophytes and therophytes typically have also arelatively large share in the flora. Their high number isdue to the special characteristics of riverside habitats.A higher proportion of therophytes relative to geophytesin the Bia≥a and Jasienica river valleys is linked to theprogressive habitat transformation. Native speciesdominate in the flora of the river valleys investigated,constituting 85% of the Jasienica, 79% of the So≥a and76% of the Bia≥a, respectively. A characteristic featureof the flora and vegetation is the participation of taxafrom the genus Reynoutria, which in the case of theBia≥a river have invaded the river banks on a massivescale. Ecological habitat groups are the indicator whichvaries 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 asmeadow communities (Molinio-Arrhenatheretea) aredominant. On the Bia≥a and Jasienica, ruderal andtherophytic communities occur more frequently.
The composition and diversity of species in the herblayer of the floodplain forests selected for the studywas dependent on the extent to which the investigatedplot was covered by knotweed shoots. The largest num-ber of vascular plant species was recorded from plotswithout any knotweeds (max. 28). In plots with increasingcoverage 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 thenumber of herb layer species (minimum 3 species) andtheir coverage coefficients was seen in plots with a highproportion of knotweed (71%-100%) (Fig. 2; comparealso Table 1 and 2). The number of species recordedon the study plots was different depending on the sea-son of the vegetative period (in spring, this number waslower than in summer). In early spring, spring geophyteswere recorded on the study plots (e.g. Ficaria verna,Symphytum tuberosum, Anemone nemorosa, Dentariaglandulosa, Primula elatior). Their development is not
Fig. 2. Species richness in plots grouped by % cover of Reynoutriaduring different seasons of growth
Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...
125
Spor
adic
spe
cies
: Q
uerc
o-F
aget
ea ñ
Ath
yriu
m f
ilix-
fem
ina
20r;
Sta
chys
syl
vatic
a 25
(+);
Mol
inio
-Arr
hena
ther
etea
ñ F
ilipe
ndul
a ul
mar
ia 4
r; H
erac
leum
sph
ondy
lium
4r;
San
guis
orba
offi
cina
lis 4
(+);
Tar
axac
um o
ffici
nale
25r
;V
eron
ica
cham
aedr
ys 2
5(+)
; Art
emis
iete
a vu
lgar
is ñ
Cir
sium
vul
gare
4r;
Sol
idag
o ca
nade
nsis
25(
+.2)
; Sol
idag
o gi
gant
ea 1
1(1.
2)
Tab
le 1
. Cha
nges
in ri
paria
n pl
ant c
omm
uniti
es h
erb
laye
r com
posi
tion
and
cove
r as
a re
sult
of c
olon
izat
ion
by ta
xa o
f Re
ynou
tria
ñ s
prin
g as
pect
Biodiv. Res. Conserv. 1-2: 123-130, 2006
126
significantly influenced by the growth of knotweedshoots during this period (Fig. 3). These species haveenough time to go through their full life cycle. In thecase of this group of plants, a decrease in the coveragecoefficient can only be seen in plots with a massiveoccurrence of knotweeds (Table 1). In the summer season,
Fig. 4. Cover of species characteristic of the Salicetea purpurea community relative to cover of Reynoutria
Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...
Fig. 3. The average rate of longitudinal growth of Reynoutria shoots measured on a sample of 30 shoots selected randomly
Aegopodium podagraria and Petasites hybridus.Characteristic and differentiating species for the classSalicetea purpureae had their most abundant occurrencein plots without any knotweed and with small andmedium coverage by knotweed plants (Fig. 4). Lack oflight and saturation of the soil with rhizomes on plots
the rapid growth of shoots and especially the developmentof knotweed leaves leads to a conspicuous reduction inthe vegetation of those species for which the peak ofthe life cycle falls in the summer period. The developmentof knotweed plants begins, depending on seasonalcondition variations, in mid-April with the appearanceof shoot tips which develop from subterraneous rhizomes.In the period between end of April and mid-May, rapid(very sudden) longitudinal growth of the shoots occurs(initially between 6 and 12 centimetres per 4 days;subsequently even as much as 40 cm per 4 days; Fig. 3)
with simultaneous development of leaves. In the laterperiod within the growth season, the rate of longitudinalshoot growth is significantly slower (during the firsthalf of June, the shoots reach their maximal height);lateral shoots develop during this time.
On the study plots, the proportion of the mostfrequently recorded species has been compared withregard to the coefficient of coverage by Reynoutriashoots. The following species were included: Ficariaverna, Symphytum tuberosum, Urtica dioica, Rubuscaesius, Phalaris arundinacea, Calystegia sepium,
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127Biodiv. Res. Conserv. 1-2: 123-130, 2006
from group IV (70-100% of Reynoutria) has effectivelyeliminated most characteristic species for this class. Theonly exceptions were Urtica dioica (a rhizome perennial)and Calystegia sepium (a creeper).
In the study plots without any knotweed or with lowcover by knotweed plants, the presence of the charac-teristic species for most of the ecological habitat groupswas confirmed. In these plots, a high proportion of spe-cies from thicket, scrub and forest facies was found.Their number drops conspicuously with an increase inthe share of knotweed plants (group III plots). A highshare of Reynoutria (70-100%) in group IV plots led toa reduction in most ecological habitat groups.
The gradual elimination of herb layer species ismainly caused by the expansion of knotweeds in thehabitat investigated. Their sudden growth at the verybeginning of the vegetative season with a simultaneousdevelopment of large leaves (their size in R. sachalinensisis 15-43 cm long and 10-27 cm broad; after Fojcik &Tokarska-Guzik 2000) causes strong shading of the soiland elimination of other species. Lack of light availabilitynear the surface of the ground is additionally aggravatedby the characteristics zigzag pattern of the leafarrangement along the knotweed stem. Furthermore,knotweed rhizomes thrive in the soil to an extent whichprevents many other species from taking root, germinatingand going through the entire life cycle.
The similar results were published by Sukopp &Sukopp (1988), Schepker (1998) and Schl¸pmann (2000),stated that one of the most serious problems is thatReynoutria modifies or expels the autochthonous veg-etation by shadowing. Investigations conducted byAlberternst (1998) show that the number of plant spe-cies in vegetation transects with Fallopia were lowerthan in transects without the species.
The biological properties of representatives of genusReynoutria cause these plants to have a definite reductionimpact on the native components of plant communities(Tokarska-Guzik & Dajdok 2004; Tokarska-Guzik2005). The enormous potential of these species forspreading by vegetative means, combined with theirrapid growth and a capacity to adapt to diverse or evenextreme habitat conditions, often invading and holdinglarge areas, have resulted in this species earning thestatus of invasive plant and nuisance íweedí (Tokarska-Guzik 2005).
The characteristics influencing the rate and successof alien plant invasions listed by FaliÒski (2004) includehigh fertility and reproductive potential of eachindividual, persistence of seeds (propagules); seedsequipped with devices which allow rapid long-distancetransport; accelerated growth at the juvenile stage, early
maturation to reproduction; resistance to extremeenvironmental conditions; dioecy, polygamy, self-pollination; ability to compete with other species (dueto height, production of allelopathic substances etc.);phenotypical variability, ability to form mutants,polyploids, hybrids with related species. Taxa from thegenus Reynoutria have many of the above-mentionedfeatures. Above all, they belong to the group of plantswith the most efficient mode of vegetative reproductionby rhizome segmentation (Alberternst et al. 1995; Child1999; Fojcik & Tokarska-Guzik 2000). Potentially,a rhizome fragment which is only 1 cm in length and0.7 g in weight may generate a new plant (Brock et al.1995). Any transformations of the environment causedby natural factors (e.g. floods) as well as in particularby anthropogenic factors (earthworks, river regulation)favour the spreading of rhizomes (Pyöek & Prach 1993).Regulation of the Bia≥a river and Jasienica stream aswell as earthworks conducted in the So≥a river valleyhave contributed to a major extent to the colonisationof large areas within the study zone by Reynoutria spe-cies. The growth rate of knotweed plants is especiallyhigh at the beginning of the vegetative season: 43.1 mm/day (Child 1999); 80 mm/day (Seiger 1997). Furthermorethe soil which is thickly occupied by knotweed shoots(down to 7 m in depth) does not allow the germinationof many species. According to the classification of lifestrategies (Grime 1979) knotweeds represent a C-typestrategy, which decisively increases their invasive po-tential (Pyöek & Prach 1993). The following speciesare able to compete successfully with knotweeds evenif the plants are present at high coverage coefficient:Aegopodium podagraria, Urtica dioica (rhizomeperennials with similar mode of growth), balsams:Impatiens parviflora and I. glandulifera (alien annualswith an R-type life strategy and shallow root systems)as well as Calystegia sepium (a creeper).
4. Conclusions
The results of field studies have confirmed thehypothesis that representatives of genus Reynoutriaexert a reducing influence on the native components offloodplain forest herb layer. This impact is, however,different with regard to various life forms and ecologicalhabitat groups of plants. Increase of surface coverageby knotweeds does not significantly influence thedevelopment of early spring geophytes which are ableto go through their entire life cycle. In the case of thisgroup of plants, a decrease in coverage coefficient canonly be seen in plots with massive occurrence of knot-weeds.
128 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...
Tab
le 2
. Cha
nges
in r
ipar
ian
plan
t com
mun
ities
her
b la
yer
com
posi
tion
and
cove
r as
a r
esul
t of
colo
niza
tion
by ta
xa o
f R
eyno
utri
a ñ
late
sum
mer
asp
ect
Num
ber
of th
e re
levè
Day
Mon
thY
ear
Loc
ality
and
num
ber
of p
lot
Cov
er o
f tr
ee la
yer
a1 in
%C
over
of
tree
laye
r a2
in %
Cov
er o
f sh
rub
laye
r b
in %
Cov
er o
f he
rb la
yer
c1 in
%C
over
of
herb
laye
r c2
in %
Cov
er o
f m
oss
laye
r d
in %
Num
ber
of s
peci
es in
the
rele
vèC
over
of
Rey
nout
ria
in %
Rey
nout
ria
japo
nica
Rey
nout
ria
xboh
emic
aR
eyno
utri
a sa
chal
inen
sis
Salic
etea
pur
pure
aeSa
lix f
ragi
lisSa
lix f
ragi
lisSa
lix f
ragi
lisSa
lix a
lba
Salix
alb
aSa
lix a
lba
Salix
alb
aP
opul
us n
igra
Pop
ulus
alb
aSa
lix p
urpu
rea
Salix
vim
inal
isR
ubus
cae
sius
Cal
yste
gia
sepi
umP
hrag
mite
tea
Pha
lari
s ar
undi
nace
aP
hrag
mite
s au
stra
lisC
arex
gra
cilis
Aln
o-U
lmio
nA
lnus
glu
tinos
aA
lnus
glu
tinos
aA
lnus
glu
tinos
aA
lnus
inc
ana
Pad
us a
vium
Urt
ica
dioi
caP
etas
ites
hybr
idus
12
34
56
78
910
1112
1314
1516
1718
1920
2122
2324
2526
2728
2930
3132
11
118
88
1212
1211
1111
118
1212
1211
118
1717
2020
2017
1717
1717
2020
20
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
0909
03
0303
0303
0404
0403
0303
0303
0404
0403
0303
0303
0404
0403
0303
0303
0404
04
5S6S
9S10
S12
S1J
2J3J
3S4S
7S8S
9S4J
5J6J
1S2S
13S
3B7B
7J8J
9J1B
2B4B
5B6B
10J
11J
12J
7050
7060
5060
7050
1060
1060
7070
4060
7050
2010
-50
3030
30-
5-
--
5-
-
--
--
--
-30
-40
5-
--
--
-20
30-
--
-10
55
--
--
-
55
205
55
205
5030
1030
1010
5030
305
2010
-5
2010
105
5
55
100
9080
9090
9080
8090
7090
3060
6090
7080
9010
010
010
090
100
100
100
100
100
100
100
100
100
100
--
--
--
--
8080
9080
9090
8080
9010
090
3040
100
9030
1010
--
--
--
-
10-
-20
--
--
--
--
--
--
--
--
--
--
--
--
--
-
2015
1519
2221
1820
1518
2018
1617
2018
2216
1314
2018
1714
98
115
37
116
Rey
nout
ria
abse
nt -
Gro
up I
up to
40%
- G
roup
II
41 to
70%
- G
roup
III
71 to
100
% -
Gro
up I
V
..
..
..
..
3.5
3.4
3.5
2.2
2.3
..
.3.
54.
55.
55.
55.
5.
..
5.5
5.5
5.5
5.5
5.5
..
.II
Ic 1
..
..
..
..
..
..
..
..
..
..
..
..
2.4
.2.
32.
32.
42.
32.
3.
Ic 1
..
..
..
..
..
..
.2.
33.
53.
4.
..
..
4.5
4.5
4.5
..
..
.5.
55.
55.
5II
a 14.
42.
24.
43.
33.
32.
23.
32.
2.
3.3
.3.
33.
32.
2.
2.2
3.3
3.3
.+
.2.
2.
+2.
2.
..
..
..
IVa 2
..
..
..
..
3.3
.3.
3.
..
2.2
..
.1.
12.
2.
.1.
11.
1.
..
..
..
.b
.+
1.2
+2.
2.
1.2
+3.
32.
2.
1.1
..
2.2
1.1
1.1
1.1
..
.1.
1.
..
..
..
..
.a 1
..
2.2
2.2
1.1
2.2
2.2
1.1
..
.1.
12.
22.
2+
+2.
22.
2.
..
1.1
..
..
..
..
..
III
a 2.
..
..
..
..
.3.
31.
1.
..
..
.1.
1.
..
..
..
..
..
..
b.
..
.+
..
+.
..
.+
+.
.1.
1.
..
..
1.1
..
.+
..
.+
.c
..
.+
..
..
..
..
..
..
..
..
..
..
..
..
..
..
a 1+
2.2
..
2.2
..
.2.
21.
1+
2.2
..
..
2.2
.2.
2.
..
..
..
..
..
..
IIa 1
+2.
2.
.2.
2.
..
..
..
..
..
..
..
..
..
..
..
..
..
Ib
..
..
+.
..
2.2
.1.
12.
2.
..
..
..
..
..
..
..
..
..
.I
b.
..
..
..
..
.2.
2.
..
..
..
..
..
..
..
..
..
..
I+
.2.
32.
23.
32.
21.
23.
31.
2r
+2.
22.
32.
31.
2r
2.2
1.2
.r
.1.
2.
rr
..
..
..
.IV
+.2
1.2
1.2
r1.
2r
+1.
2+
.2+
.2+
.21.
21.
21.
2+
.2+
.2r
1.2
+.2
+.
1.2
+.2
+.
.+
..
++
.II
I
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
..
..
..
..
..
+.
..
..
..
..
..
..
..
..
..
..
I.
..
..
..
..
..
..
..
..
+.
..
..
..
..
..
..
.I
a 1.
..
..
..
..
..
..
..
..
..
..
..
..
.+
..
..
.II
a 2.
..
.+
++
1.1
..
..
1.1
.r
..
..
..
..
..
..
..
.+
.b
..
..
..
..
..
..
1.1
..
..
..
..
..
..
..
..
..
.a 2
..
..
..
..
..
..
2.2
..
..
..
..
..
..
..
..
..
.I
b+
..
..
r.
.2.
2.
.1.
1.
.2.
2.
2.2
.1.
2.
..
1.2
..
..
..
..
.II
3.3
2.2
3.3
2.2
1.2
2.2
3.3
1.2
3.2
3.3
1.2
3.3
2.2
2.2
3.2
3.3
2.2
3.3
2.2
+.2
+.2
3.3
2.2
+.2
1.2
++
.2r
.r
+.2
rV
2.3
3.3
1.2
+2.
3+
1.2
2.3
2.2
2.2
.2.
22.
22.
22.
22.
22.
33.
31.
2r
.1.
21.
2r
..
..
..
..
IV
Constancy
129
Spor
adic
spe
cies
: Que
rco-
Fage
tea
ñ Eq
uise
tum
syl
vatic
um 3
r; T
ilia
plat
yphy
llos
a 1 20
(+);
Mol
inio
-Arr
hena
ther
etea
ñ E
quis
etum
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Biodiv. Res. Conserv. 1-2: 123-130, 2006
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130 Barbara Tokarska-Guzik et al. Changes in plant species richness in some riparian plant communities ...
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