UDK 630* 188 (001)
Izvorni znanstveni članci – Original scientific papers
Šumarski list, 3–4 (2015): 155–169
nEW InvASIvE FOREST COMMUnITIES In THE
RIPARIAn FRAGILE HABITATS – THE CASE
STUdY FROM RAMSAR SITE CARSKA BARA
(vOJvOdInA, SERBIA)
NOVE INVAZIVNE ŠUMSKE ZAJEDNICE POPLAVNIH OSJEtLJIVIH
StANIŠtA – StUDIJA SLUČAJA IZ RAMSARSKOG PODRUČJA
CARSKA BARA (VOJVODINA, SRBIJA)
Vera BAtANJSKI1, Eva KABAŠ2, Nevena KUZMANOVIĆ2, Snežana VUKOJIČIĆ2, Dmitar LAKUŠIĆ2,
Slobodan JOVANOVIĆ2
Summary
The phytosociological investigation of habitats with highly invasive tree species Acer negundo L. and Fraxinus pennsylvanica Marshall was performed in Ramsar site Carska bara (Vojvodina, Serbia). A total of 107 species were noticed within 32 relevés. Recorded relevés are georeferenced and analysed in detail. The results of the relevant numerical analyses suggest the existence of two floristically and coenologically well defined groups of stands defined
as the forest communities: Rubo caesii–Aceretum negundi ass. nova and Carici otrubae–Fraxinetum pennsylvanicae
ass. nova. The increasing dispersal rate of the invasive trees is detected as a problem amongst many fragile wet
habitats across Serbia and SE Europe, alerting their urgent and effective control.
KEY WORDS: invasive trees, Acer negundo, Fraxinus pennsylvanica, forest communities, wetland.
InTROdUCTIOn
UVOD
It is known that invasive alien species are the second risk
factor of biodiversity endangering, right after habitat destruction (Brennan and Withgott, 2011). According to the
European strategy on invasive alien species, they are one of
the biggest challenges in the preservation of biodiversity in
Europe (Genovesi and Shine, 2003). Negative consequences
of the presence and spreading of the invasive species were
analyzed by numerous authors (Elton, 1958; Drake et al.,
1989; Di Castri et al., 1990; Williamson, 1996; Starfinger et
al., 1998; Tilman, 1999; Parker et al., 1999; Hejda et al., 2009;
Pyšek and Richardson, 2010). Threats to the autochthonous
biodiversity and degradation of the natural habitats become
the most prominent when the invasive species become naturalized and form stable communities. Field studies and
experiments confirmed that the presence of the alien species can be a threat to the native species, primarily due to
the negative effects of the competition (Vilà and Weiner,
MSc Vera Batanjski: University of Belgrade, Faculty of Biology, Department of Plant Ecology and Geography; Institute of Criminological and Sociological Research,
Gračanička 18, 11000 Belgrade, Serbia, Corresponding author: vera.batanjski@gmail.com
2
MSc Eva Kabaš, PhD Nevena Kuzmanović, PhD Snežana Vukojičić, PhD Dmitar Lakušić, PhD Slobodan Jovanović: Institute of Botany and Botanical Garden
"Jevremovac", Faculty of Biology, University of Belgrade, Takovska 43, 11000 Belgrade, Serbia, ekabas@bio.bg.ac.rs, nkuzmanovic@bio.bg.ac.rs, sneza@bio.bg.ac.rs,
dlakusic@bio.bg.ac.rs, sjov@bio.bg.ac.rs
1
156
Šumarski list, 3–4, CXXXIX (2015), 155–169
2004). This can lead to the homogenization of ecosystems
and decrease in autochthonous species diversity (D’Antonio
and Vitousek, 1992). In the final outcome, this can results
in "novel ecosystems" (Hobbs et al., 2006; Hobbs et al.,
2009). A large number of invasive plants are the trees
(Binggeli, 1996).
by using the green ash. Nowadays, this species is widely
present, and concentrated in lowland river valleys and
marshy or saline areas in Hungary, some of which are close,
or bordering Serbia (Tiborcz et al., 2011). F. pennsylvanica
inhabit the same wet places, such as our study area in the
northern Serbia.
Twenty three alien tree species have been identified in forest
ecosystems in Serbia (Medarević et al., 2008), 17 of which are
invasive (Grbić et al., 2007). Due to their aggressive expansion abilities, some of them represent a serious threat to the
natural regeneration and survival of the native trees. This fact
especially refers to Robinia pseudoacacia L., Acer negundo L.,
Ailanthus altissima (Miller) Swingle and shrub Amorpha fruticosa L. However, more precise data on the distribution of
the invasive woody species in Serbia still do not exist, particularly the ones related to protected areas. Except for the
few recent literature sources on the topic of alien or invasive
plants or communities in Serbia (Vrbničanin et al., 2004; Jarić
et al. 2011; Lazarević et al., 2012; Anačkov et al., 2013), the
data related to the protected Ramsar areas are still lacking
(Panjković and Stojšić, 2001; Čavlović et al., 2011).
According to Török et al. (2003) Hungary represents the
gateway for invasions into the rest of the Central and Eastern Europe, and one of the corridors for invasions are precisely the wet habitats (gallery forests, disturbed bogs and
marshes). Since our study area is a floodplain, suitable for
A. negundo and F. pennsylvanica, we consider that analyzed
invasive tree species could spread from Hungary to the
northern Serbia across these corridors.
In the light of the fragility and vulnerability, as well as the
rarity of the wetlands (Smart, 1997), alarmingly large presence of highly invasive species was observed, some of which
are highly naturalized, such as the North American trees:
Acer negundo L. and Fraxinus pennsylvanica Marshall. The
species Acer negundo was introduced to Europe in 17th century (Mędzycki, 2011), and Fraxinus pennsylvanica was introduced in 18th century (Csiszár and Bartha, 2008). Since
then, they have spread across the Europe, especially in the
riparian habitats (DAISIE, 2013).
Since the literature sources for our teritorry provided no
data on the exact time two analyzed invasive species have
arrived to Serbia we looked up for these information in
neighboring Hungary, which covers the biggest part of the
Pannonian plain, where our study area belongs.
After its introduction, A. negundo became quite a popular
garden tree and in the second half of 19th century it was
widely recommended for planting, as a wind-break and
shelter-belt tree across the Western, Central and Eastern
parts of Europe (Mędzycki, 2011). In Hungary, it has been
known since the second half of the 19th century. It was
widely planted on flood areas of the Great Plains from
where it escaped and established on riparian forests and
black locust plantations, mainly along the rivers (lower
Danube valley), marshy areas near Lake Balaton, and also
on dry sandy soils of the Pannonian Plain (Udvardy, 2008).
A. negundo inhabit wet places, such as our study area in the
northern Serbia.
F. pennsylvanica was known since the very beginning of the
20th century in Hungary, when there were attempts of converting willow-poplar gallery forests to hardwood stands
Despite all the negative impacts the invasive plant species
can exibit, paradoxically, the level of the exploration of invasive plant communities, especially those dominated by
woody species, is small. However, except for the ass. Sambuco nigrae–Aceretum negundo Exner 2004, described in
the potential broad-leaved woodland areas of Austria
(Exner and Willner, 2004), there are no other literature data
on the communities built up by the two observed invasive
woody species. Invasive species Acer negundo and Fraxinus
pennsylvanica have established stable communities in the
Special Nature Reserve Carska bara (Vojvodina, Serbia),
which are analyzed and described in this paper.
MATERIALS And METHOdS
MAtERIJALI I MEtODE
Study area – Područje istraživanja
The investigated area Carska bara is located on the alluvial
plain between the rivers Tisa and Begej, in the central Banat (Vojvodina, Serbia), southwest of Zrenjanin city. It belongs to the UTM Grid zone 34T, UTM square 10x10 km
DR50 and DR51 (Figure 1), presenting one of the preserved
floodplains in this part of Serbia. According to the pedological map of Vojvodina (Naugebauer et al., 1971) and ArcGIS 10.2 Software, the alluvial saline soil is mostly present
within the study area. The area belongs to the temperate
climate zone with emphasized continental characteristics
(Stevanović and Stevanović, 1995; Kovačev, 2010). Habitats
are presented as the mosaic of the wetlands, ponds, (salt)
marshes, wet meadows, steppes and forests, intersected by
rivers, canals and dikes. Over 230 bird species (including
all European heron species and cormorants), 17 of which
are internationally important were recorded here (Puzović
et al., 2009). Total of 277 taxa of the higher plants (on species and subspecies level) are noticed in the study area, two
of which are the Pannonian endemics – Aster tripolium L.
subsp. pannonicus (Jacq.) Soó and Cirsium brachycephalum
Juratzka. Also some relict species of marshy flora and of
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 157
age of 112.07 m2. All data were georeferenced using a GPS
device eTrex Vista C (Garmin). The plant material was deposited in the Herbarium of the University of Belgrade –
BEOU (Theirs, 2014).
data analysis – Analiza podataka
Figure 1. Map of Ass. Rubo caesii–Aceretum negundi ass. nova (white
relevés' points) and Ass. Carici otrubae–Fraxinetum pennsylvanicae
ass. nova (black relevés' points) in the study area Carska bara (Vojvodina, Serbia).
Slika 1. Karta Ass. Rubo caesii–Aceretum negundi ass. nova (bijelo
označena polja) i Ass. Carici otrubae–Fraxinetum pennsylvanicae ass. nova
(crno označena polja) u području istraživanja Carska bara (Vojvodina, Srbija).
xerothermic steppe flora are present within (Perić, 2010).
Regarding the phytogeographical affiliation, the study area
belongs to the Pannonian province of the Pannonian-Vlach
subregion and the Pontic-Southsiberian floristic-vegetation
region (Stevanović et al., 1999). The studied area is located
in the wider area of the potential steppe and forest-steppe
vegetation (Jovanović et al., 1986). The diversity of the habitats enables high species diversity, hence the variety of rare,
endangered or vulnerable animals and plants could be
found here. Because of these natural characteristics, Special
Nature Reserve Carska bara is recognized as the area of the
international importance, so it is included in the list of Ramsar sites, Important bird area (IBA), Important plant area
(IPA), Emerald and ASCI area (Hlavati – Širka et al., 2013),
which are of the particular importance for the conservation
of nature.
vegetation sampling – Uzorkovanje vegetacije
The phytosociological investigation of the selected riparian
sites with the high presence of the invasive trees in the area
of Carska bara (Vojvodina, Serbia) was conducted in the
period 2011 – 2013. Thirty two relevés were made according to Braun – Blanquet (1964) methodology. The size of
sampling plots was adjusted to the minimum areal size as
proposed by Mueller – Dombois & Ellenberg (1974). It varied from 25 to 200 m2 for wild growing forest sites, in aver-
After the transformation of Braun – Blanquet combined
alpha-numeric scale into a completely numerical scale as
proposed by Van der Maarel (1979), the classification of
phytocoenological relevés was done. In addition to our own
32 relevés, four relevés of association Sambuco nigrae–Aceretum negundo (from different localities of Austria: Karlhof, Pamhagen, Teichhof, District Neusiedl am See (Exner
and Willner, 2004) were also included in the analysis and
used for comparison. Relevés were classified using Jaccard
(1928) similarity and group average clustering, for further
coenological characterization and differentiation. All the
analyses were done in PcOrd 6.0 software (Mccune and
Mefford, 2011). The names of the newly described associations are harmonized with The International Code of Phytosociological Nomenclature (Weber et al., 2000), while the
nomenclature and EUNIS codification of primary vegetation of riparian habitats is compliant with Davies et al.
(2004).
In this paper we used the concept of dominant and diagnostic species proposed by Chytrý et al. (2002), Chytrý and
Tichý (2003) and Tichý and Chytrý (2006). Fidelity was
calculated for two target groups, distinguished in the cluster analysis i.e. those in which two mentioned woody invasive species were highly present. Using the statistical measures of fidelity, we quantified concentrations of species
occurrences in groups of classified sites in order to determine diagnostic species (Chytrý et al., 2002). In order to
determine dominant species, the coverage index (Ic) was
calculated according to Lausi et al. (1982).
Species with Phi-coefficient values higher than 0.40 have
been considered diagnostic. Species with cover ≥ 25% in a
minimum 5% of the relevés for any association have been
accepted as dominant.
Nomenclature of plant taxa, with a few exceptions, follows
the Flora Europaea Database (Tutin et al., 2001). All taxa
with authors’ names quoted in the paper are given in the
tables (Table 1 and Table 2).
2
2
2
3
1
4
2
1
2
1
4
1
3
2
2
1
4
+
r
1
1
2
1
+
1
+
2
+
2
r
r
1
1
r
1
1
r
2
r
1
+
+
+
+
+
r
1
2
+
2
1
1
2
+
r
2
3
1
2
1
2
1
+
2
1
2
1
1
r
+
1
r
3
1
4
2
r
+
+
+
+
1
r
4
3
2
+
+
1
+
2
+
2
2
r
r
3
4
1
1
2
2
1
+
2
+
+
r
+
+
+
3
1
1
r
+
1
1
r
1
1
+
1
1
2
+
1
3
+
2
r
r
r
r
r
3
1
2
2
2
+
+
1
2
r
r
2
1
4
1
1
r
r
1
2
4
+
r
r
+
1
3
r
r
r
+
2
1
r
r
1
r
2
1
2
r
2
r
r
r
1
Cover
index
according to
Lausi
et al.
(1982)
(Ic)
Fidelity index (Ф)
2
2
Frequency (%)
DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR50 DR50 DR51 DR51
75 75 75 75 75 74 74 74 74 74 74 75 75 75 75 78 75 73 75 76
SE SE
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
3
10
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
90 70 80 95 80 60 40 30 60 30 40 60 80 95 80 100 100 70 60 70
10
/
80 40 80 50 40 30 60 30 30 20 30
/
80 70 50 70
/
60
10 70 20 10 20 60 40 30
5
30 40 10 50 20 50 30 90 40
/
/
90 70 50 95 20 40 40 30 10 20 40 60 80 95 40 80 70 40 70
5
25 30 100 25 100 100 50 50 100 50 25 50 50 25 100 100 200 100 100 200
5
6
8
7
9
13 15 16 17 18 14 19 20 21 30 11 22 23 10 12
1
2
3
4
5
6*
7
8
9
10 11 12 13 14 15 16 17 18 19 20
100
80
66,67
37,22
0,40
0,26
55
22.22
0.44
40
18.33
0.48
60
55
55
50
50
35
35
35
30
30
25
25
20
20
20
20
20
20
20
15
17.78
15.56
11.67
20.00
10.00
16.67
8.33
6.11
10.00
7.78
7.22
3.33
11.67
8.89
7.78
7.22
6.67
4.44
2.22
5.00
0.10
-0.08
0.24
-0.06
0.32
-0.40
0.26
0.18
-0.08
0.13
0.36
0.07
0.02
-0.06
0.02
-0.19
0.10
-0.19
0.32
0.04
Šumarski list, 3–4, CXXXIX (2015), 155–169
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
Phytogeogra- General cover (%)
Life
phical
forms
tree layer cover (%)
elements
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m 2)
Relevé number from cluster
Relevé No.1
Dominant taxa
P
ADV
Acer negundo L.
NP
EAz
Rubus caesius L.
Diagnostic taxa
H
KOSM
Dactylis glomerata L.
Fraxinus angustifolia Vahl subsp. oxycarpa (Bieb.
P
EAz
ex Willd.) Franco & Rocha Afonso
Other taxa
G
SE
Iris pseudacorus L.
H
SE
Symphytum officinale L.
H
EAz
Poa trivialis L.
HOL
Urtica dioica L.
H
t
EAz
Bromus sterilis L.
P
ADV
Fraxinus pennsylvanica Marshall
H
EAz
Cirsium arvense (L.) Scop.
H
EAz
Arrhenatherum elatius (L.) Beauv. ex J. & C. Presl
G
MED-SUBMED Aristolochia clematitis L.
P
ADV
Prunus cerasifera Ehrh.
H
EAz
Arctium lappa L.
H
HOL
Festuca rubra L.
G
KOSM
Elymus repens (L.) Gould
G
KOSM
Phragmites australis (Cav.) trin. ex Steudel
P
ADV
Gleditsia triacanthos L.
SH
HOL
Humulus lupulus L.
HydG
EAz
Carex riparia Curtis
t
KOSM
Galium aparine L.
t
HOL
Lapsana communis L.
H
SE
Carex hirta L.
158
table 1. Diagnostic table of new association Rubo caesii–Aceretum negundi ass. nova from SNR Carska bara in Serbia. (*holotypus)
tablica 1. Dijagnostička tablica nove asocijacije Rubo caesii–Aceretum negundi ass. nova iz SRP Carska bara u Srbiji. (*holotip)
Frequency (%)
Cover
index
according to
Lausi
et al.
(1982)
(Ic)
Fidelity index (Ф)
DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR50 DR50 DR51 DR51
75 75 75 75 75 74 74 74 74 74 74 75 75 75 75 78 75 73 75 76
SE SE
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
3
10
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
90 70 80 95 80 60 40 30 60 30 40 60 80 95 80 100 100 70 60 70
10
/
80 40 80 50 40 30 60 30 30 20 30
/
80 70 50 70
/
60
10 70 20 10 20 60 40 30
5
30 40 10 50 20 50 30 90 40
/
/
90 70 50 95 20 40 40 30 10 20 40 60 80 95 40 80 70 40 70
5
25 30 100 25 100 100 50 50 100 50 25 50 50 25 100 100 200 100 100 200
5
6
8
7
9
13 15 16 17 18 14 19 20 21 30 11 22 23 10 12
1
2
3
4
5
6*
7
8
9
10 11 12 13 14 15 16 17 18 19 20
1
+
1
1
1
+
1
1
r
1
+
r
r
1
+
1
r
r
3
2
r
3
1
1
1
1
2
r
1
1
1
1
+
1
+
1
+
+
+
+
r
1
+
r
r
r
3
2
2
2
2
2
2
1
1
1
15
15
15
15
15
15
10
10
10
10
10
10
10
10
10
10
10
10
10
10
5
5
5
5
5
5
5
5
5
5
4.44
4.44
3.89
3.33
3.33
2.78
6.67
4.44
3.33
3.33
3.33
3.33
3.33
2.78
2.78
2.22
2.22
2.22
1.67
1.11
3.89
2.78
2.78
2.78
2.78
2.78
2.78
1.67
1.67
1.67
0.27
0.04
0.27
0.14
-0.43
0.14
-0.04
0.07
0.22
0.22
-0.04
-0.13
-0.13
0.07
-0.13
0.22
0.22
0.07
0.22
0.07
0.15
-0.46
-0.22
0.15
0.15
0.15
-0.03
-0.22
-0.22
-0.13
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 159
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
Phytogeogra- General cover (%)
Life
phical
forms
tree layer cover (%)
elements
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m 2)
Relevé number from cluster
Relevé No.1
H
SE
Scutellaria columnae All.
H
EAz
Glechoma hederacea L.
H
EAz
Picris hieracioides L.
H
EAz
Polygonum mite Schrank
S
EAz
Solanum dulcamara L.
P
EAz
Quercus robur L.
G
EAz
Phalaris arundinacea L.
H
HOL
Carex pseudocyperus L.
HydG
ADV
Acorus calamus L.
P
ADV
Morus nigra L.
H
EAz
Calamagrostis epigejos (L.) Roth
H
KOSM
Calystegia sepium (L.) R. Br.
t/H
ADV
Erigeron annuus (L.) Pers.
H
KOSM
Carduus acanthoides L.
H/t
EAz
Lactuca serriola L.
H
EAz
Carex cespitosa L.
t
KOSM
Solanum nigrum L.
t
KOSM
Torilis arvensis (Hudson) Link
t
KOSM
Lolium temulentum L.
H
EAz
Alopecurus pratensis L.
St
ADV
Echinocystis lobata (Michx) torrey & A. Gray
H/HydG
EAz
Lycopus europaeus L.
NP
SE
Crataegus monogyna Jacq.
H
MED-SUBMED Carex divulsa Stokes
H
EAz
Dipsacus laciniatus L.
G
MED-SUBMED Glycyrrhiza glabra L.
H
EAz
Festuca rupicola Heuffel
HydG
KOSM
Typha latifolia L.
t
ADV
Bidens frondosa L.
P
EAz
Salix alba L.
Frequency (%)
Cover
index
according to
Lausi
et al.
(1982)
(Ic)
Fidelity index (Ф)
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
5
1.67
1.67
1.67
1.67
1.67
1.67
1.11
1.11
1.11
1.11
1.11
1.11
1.11
1.11
1.11
1.11
0.56
0.56
0.56
0.56
0.56
0.56
0.56
0.56
0.56
0.56
-0.03
-0.03
0.15
0.15
0.15
0.15
-0.52
-0.13
-0.22
-0.13
-0.03
-0.03
-0.03
0.15
0.15
-0.03
-0.46
-0.13
-0.03
0.15
0.15
0.15
0.15
0.15
-0.03
-0.03
1 Relevés 1 – 20: Serbia, Carska bara: 45° 16' 887" N – 20° 25' 152" E, 45° 16' 863" N – 20° 25' 066" E, 45° 16' 953" N – 20° 25' 318" E, 45° 16' 863" N – 20° 25' 066" E, 45° 16' 852" N – 20° 24' 824" E, 45° 16' 818" N – 20° 25' 310"
E, 45° 16' 754" N – 20° 25' 308" E, 45° 16' 672" N – 20° 25' 289" E, 45° 16' 553" N – 20° 25' 241" E, 45° 16' 517" N – 20° 25' 230" E, 45° 16' 800" N – 20° 25' 338" E, 45° 16' 140" N – 20° 25' 120" E, 45° 16' 104" N – 20° 25' 119" E,
45° 15' 695" N – 20° 24' 693" E, 45° 15' 670" N – 20° 24' 668" E, 45° 16' 912" N – 20° 26' 025" E, 45° 14' 529" N – 20° 23' 069" E, 45° 14' 255" N – 20° 22' 819" E, 45° 15' 963" N – 20° 25' 016" E, 45° 16' 029" N – 20° 25' 102" E.
Šumarski list, 3–4, CXXXIX (2015), 155–169
DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR51 DR50 DR50 DR51 DR51
75 75 75 75 75 74 74 74 74 74 74 75 75 75 75 78 75 73 75 76
SE SE
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
/
3
10
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
90 70 80 95 80 60 40 30 60 30 40 60 80 95 80 100 100 70 60 70
10
/
80 40 80 50 40 30 60 30 30 20 30
/
80 70 50 70
/
60
10 70 20 10 20 60 40 30
5
30 40 10 50 20 50 30 90 40
/
/
90 70 50 95 20 40 40 30 10 20 40 60 80 95 40 80 70 40 70
5
25 30 100 25 100 100 50 50 100 50 25 50 50 25 100 100 200 100 100 200
5
6
8
7
9
13 15 16 17 18 14 19 20 21 30 11 22 23 10 12
1
2
3
4
5
6*
7
8
9
10 11 12 13 14 15 16 17 18 19 20
1
1
1
1
1
1
+
+
+
+
+
+
+
+
+
+
r
r
r
r
r
r
r
r
r
r
160
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
Phytogeogra- General cover (%)
Life
phical
forms
tree layer cover (%)
elements
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m 2)
Relevé number from cluster
Relevé No.1
H
SE
Galium mollugo L.
P
EAz
Prunus avium L.
G/HydG
KOSM
Polygonum amphibium L.
P
ADV
Malus domestica Borkh.
P
EAz
Salix fragilis L.
H
HOL
Pastinaca sativa L.
H
EAz
Carex otrubae Podp.
H
EAz
Cucubalus baccifer L.
H
EAz
Ranunculus repens L.
H
KOSM
Cirsium vulgare (Savi) ten.
H/t
KOSM
Daucus carota L.
t
ADV
Conyza canadensis (L.) Cronq.
St/t
KOSM
Fallopia convolvulus L.
H
EAz
Thalictrum flavum L.
t/H
ADV
Lactuca sativa L.
H
EAz
Vicia cracca L.
HydG
HOL
Glyceria maxima (Hartman) Holmberg
H
EAz
Festuca pratensis Hudson
H
KOSM
Agrostis stolonifera L.
H
EAz
Lychnis flos-cuculi L.
t
EAz
Lamium purpureum L.
H
MED-SUBMED Prunella laciniata (L.) L.
H
SE
Aremonia agrimonoides (L.) DC.
H
KOSM
Plantago major L.
H
KOSM
Rumex crispus L.
P
ADV
Morus alba L.
DR51
78
/
0
90
80
60
2
200
26
2
DR51
78
/
0
100
70
50
90
200
29
3
DR51
76
/
0
90
60
70
10
100
27
4
DR51
75
/
0
80
/
80
5
100
31
5
DR51
78
/
0
100
90
50
10
200
36
6
DR51
78
/
0
100
70
40
100
200
25
7
DR51
78
/
0
100
60
20
100
200
28
8*
DR51
78
/
0
90
60
80
5
200
35
9
DR51
76
SE
5
100
10
100
100
50
32
10
DR51
76
SE
5
100
/
100
100
50
33
11
4
3
4
3
3
4
3
3
3
4
4
+
73
27.27
0.73
+
+
73
26.26
0.19
Cover
index
according to
Lausi et
al.
(1982)
(Ic)
Fidelity index (Ф)
Phytogeographical elements
ADV
DR51
76
/
0
90
90
10
5
20
24
1
Frequency (%)
Life forms
P
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
General cover (%)
tree layer cover (%)
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m2)
Relevé number from cluster
Relevé No. 2
100
82.83
0.63
Dominant taxa (! also diagnostic)
Fraxinus pennsylvanica Marshall!
H
EAz
Carex otrubae Podp.!
2
H
SE
Symphytum officinale L.
+
S
EAz
Solanum dulcamara L.
H/HydG
EAz
Lycopus europaeus L.
HydG
HOL
Glyceria maxima (Hartman) Holmberg
t
EAz
Polygonum hydropiper L.
1
+
3
+
r
1
4
1
3
+
+
1
+
r
Diagnostic taxa
r
r
+
1
r
r
r
r
+
r
r
r
1
+
r
+
73
12.12
0.55
r
+
64
12.12
0.66
r
64
10.10
0.66
27
4.04
0.45
+
r
r
r
+
Other taxa
SH
HOL
Humulus lupulus L.
t
KOSM
Galium aparine L.
H
KOSM
Lythrum salicaria L.
H
EAZ
H
H
H
KOSM
Poa trivialis L.
r
2
1
1
Sonchus arvensis L.
3
r
45
17.17
0.26
+
r
36
6.06
0.13
r
r
27
5.05
0.26
1
r
27
7.07
-0.23
2
MED-SUBMED Althaea officinalis L.
KOSM
1
r
18
6.06
0.37
r
18
4.04
0.37
r
r
18
2.02
0.37
1
Cichorium intybus L.
H
EAz
Lysimachia vulgaris L.
r
18
2.02
0.24
H
EAz
Cirsium arvense (L.) Scop.
r
2
r
18
6.06
-0.10
Lactuca serriola L.
+
r
18
3.03
0.08
Typha latifolia L.
+
r
18
3.03
0.15
H/t
EAz
HydG
KOSM
H
EAz
Epilobium hirsutum L.
r
9
1.01
0.25
H
EAz
Epilobium parviflorum Schreber
r
9
1.01
0.25
t
HOL
Senecio congestus (R Br) DC.
r
9
1.01
0.25
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 161
Table 2. Diagnostic table of new association Carici otrubae–Fraxinetum pennsylvanicae ass. nova from SNR Carska bara in Serbia. (*holotypus)
tablica 2. Dijagnostička tablica nove asocijacije Carici otrubae–Fraxinetum pennsylvanicae ass. nova iz SRP Carska bara u Srbiji. (*holotip)
H
HOL
Urtica dioica L.
1
P
ADV
Prunus cerasifera Ehrh.
+
H
EAz
Cucubalus baccifer L.
r
H
KOSM
Calystegia sepium (L.) R. Br.
r
H
EAz
Eupatorium cannabinum L.
H
EAz
Glechoma hederacea L.
P
ADV
Acer negundo L.
EAz
Rubus caesius L.
G
G
t
MED-SUBMED Aristolochia clematitis L.
Phragmites australis (Cav.) trin. ex
KOSM
Steudel
ADV
Bidens frondosa L.
H
HOL
Galium palustre L.
t/H
ADV
Erigeron annuus (L.) Pers.
ADV
Gleditsia triacanthos L.
ADV
Amorpha fruticosa L.
H
ADV
Aster lanceolatus Willd.
P
ADV
Celtis occidentalis L.
NP
HOL
H
SE
H
SE
Rubus idaeus L.
Epilobium lanceolatum Sebastiani &
Mauri
Ballota nigra L.
H
EAz
Leonurus marrubiastrum L.
H
EAz
Verbascum lychnitis L.
G
KOSM
Cynodon dactylon (L.) Pers.
DR51
78
/
0
100
70
50
90
200
29
3
+
2
DR51
76
/
0
90
60
70
10
100
27
4
DR51
75
/
0
80
/
80
5
100
31
5
DR51
78
/
0
100
90
50
10
200
36
6
r
1
2
DR51
78
/
0
100
70
40
100
200
25
7
DR51
78
/
0
100
60
20
100
200
28
8*
1
1
DR51
78
/
0
90
60
80
5
200
35
9
+
DR51
76
SE
5
100
10
100
100
50
32
10
DR51
76
SE
5
100
/
100
100
50
33
11
r
r
1
+
+
3
2
2
+
r
r
1
1
+
+
1
+
2
r
0.02
1
55
18.18
27
6.06
0.03
+
18
3.03
0.24
+
18
3.03
0.08
+
9
2.02
0.25
9
2.02
-0.09
30.30
-0.34
1
1
55
15.15
-0.21
1
45
10.10
0.17
36
11.11
0.23
27
6.06
0.34
18
3.03
0.37
18
4.04
0.08
18
4.04
-0.02
9
7.07
0.25
9
7.07
0.25
9
3.03
0.25
9
2.02
0.25
9
1.01
0.25
r
+
+
+
3
3
1
+
r
r
r
r
r
0.11
73
1
+
0.25
r
2
+
1.01
18.18
+
+
+
9
64
+
+
+
Cover
index
according to
Lausi et
al.
(1982)
(Ic)
9
1.01
0.25
9
1.01
0.25
9
1.01
0.25
9
1.01
0.25
Šumarski list, 3–4, CXXXIX (2015), 155–169
P
NP
DR51
78
/
0
90
80
60
2
200
26
2
Frequency (%)
Iris pseudacorus L.
Phytogeographical elements
ADV
Life forms
EAz
t/H
NP
DR51
76
/
0
90
90
10
5
20
24
1
Fidelity index (Ф)
162
H
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
General cover (%)
tree layer cover (%)
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m2)
Relevé number from cluster
Relevé No. 2
Valeriana officinalis L.
DR51
78
/
0
100
70
50
90
200
29
3
DR51
76
/
0
90
60
70
10
100
27
4
r
DR51
75
/
0
80
/
80
5
100
31
5
DR51
78
/
0
100
90
50
10
200
36
6
DR51
78
/
0
100
70
40
100
200
25
7
DR51
78
/
0
100
60
20
100
200
28
8*
DR51
78
/
0
90
60
80
5
200
35
9
DR51
76
SE
5
100
10
100
100
50
32
10
r
DR51
76
SE
5
100
/
100
100
50
33
11
Cover
index
according to
Lausi et
al.
(1982)
(Ic)
Fidelity index (Ф)
DR51
78
/
0
90
80
60
2
200
26
2
Frequency (%)
Papaver rhoeas L.
Phytogeographical elements
EAz
Life forms
SE
t
DR51
76
/
0
90
90
10
5
20
24
1
9
1.01
0.25
9
1.01
0.25
0.10
P
EAz
Salix cinerea L.
1
9
3.03
H
SE
Pimpinella major (L.) Hudson
+
9
2.02
0.10
t
EAz
Chenopodium polyspermum L.
9
1.01
0.02
G
KOSM
Elymus repens (L.) Gould
+
9
2.02
-0.17
H
HOL
Festuca rubra L.
+
9
2.02
-0.21
H
KOSM
Rumex crispus L.
H
EAz
Bromus inermis Leysser
H
HOL
Carex pseudocyperus L.
H
SE
Carex hirta L.
r
r
r
1
2
9
1.01
0.10
9
1.01
0.10
9
3.03
0.02
9
5.05
-0.09
2Relevés 1 – 11: Serbia, Carska bara: 45° 16' 007" N – 20° 25' 041" E, 45° 15' 489" N – 20° 24' 435" E, 45° 15' 535" N – 20° 24' 472" E, 45° 16' 109" N – 20° 24' 111" E, 45° 15' 596" N – 20° 24' 592" E, 45° 15' 617" N – 20°
24' 588" E, 45° 15' 570" N – 20° 24' 516" E, 45° 15' 770" N – 20° 24' 732" E, 45° 15' 721" N – 20° 24' 729" E, 45° 16' 213" N – 20° 24' 500" E, 45° 16' 186" N – 20° 24' 438" E.
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 163
t
UtM Grid zone 34t
Altitude (m)
Exposition
Slope (°)
General cover (%)
tree layer cover (%)
Shrub layer cover (%)
Herb layer cover (%)
Relevé area (m2)
Relevé number from cluster
Relevé No. 2
Bromus commutatus Schrader
164
Šumarski list, 3–4, CXXXIX (2015), 155–169
RESULTS
REZULtAtI
Syntaxonomical treatment – Sintaksonomska
interpretacija
Classification – Klasifikacija
Ass. Rubo caesii–Aceretum negundi ass. nova hoc loco
(Holotypus Table 1, reléve 6, Figure 3)
Cluster analysis clearly differentiated three groups of
relevés (Figure 2). Cluster A consists of 20 relevés dominated by Acer negundo, while cluster B includes 11 relevés
dominated by Fraxinus pennsylvanica. The third cluster
(C) consists of four relevés and represents the stands of
the ass. Sambuco nigrae–Aceretum negundo from Austria.
Although Acer negundo is dominant species in this association, its relevés are separated and allocated from those
of cluster A and B. In the relevé 34, Prunus spinosa L. is a
dominant species, hence this relevé is singled out and has
no relevance in characterization of described invasive forest communities.
From a total of 107 taxa that are present in clusters A and
B, 50 appears only in cluster A, and 27 only in cluster B,
while 30 species are common for both groups. The most
common and abundant species in both analyzed groups
are: Acer negundo, Rubus caesius L., Fraxinus pennsylvanica, Urtica dioica L., Symphytum officinale L., Aristolochia clematitis L. and Iris pseudacorus L.
Based on the results of the cluster analysis and also the
analysis of the dominant and diagnostic species, we consider the first two groups (clusters A and B) specific and
different enough, to be defined as associations.
Dominant species: Acer negundo L.(1 – 4) (Ic = 66.67) and
Rubus caesius L.(+ – 3) (Ic = 37.22)
Diagnostic species: Fraxinus angustifolia Vahl. subsp. oxycarpa (Bieb ex Willd) Franco & Rocha Afonso (Φ = 0.48),
Dactylis glomerata L. (Φ = 0.44).
Diagnosis: Ass. Rubo caesii–Aceretum negundi is developed
on marshy, gley (eugley), hypogley, saline soils, on altitudes
between 73 – 78 m on a flat surface, rarely on small slopes
(up to 10°). Individuals of the dominant species Acer negundo are predominantly present in the canopy layer, up to
15 m high, and also in the shrub and herb layers. Total cover
in different stands of this association is 30 – 100 % (average
69.5 %). Total of 80 taxa are present in all 20 relevés, average number of species per relevé is 13. Since the dominant
and nominal species Acer negundo, is invasive, the association has an invasive character itself. The fact that individuals of this species are present in shrub and herb layers, indicate that this community is in progradation, and the final
stage will, most probably, result in the development of the
monodominant forest. The other invasive species also
present in this association are: Bidens frondosa L., Echinocystis lobata (Michx) Torrey & A. Gray, Fraxinus pennsylvanica and Gleditsia triacanthos L.
Ass. Carici otrubae–Fraxinetum pennsylvanicae ass. nova
hoc loco (Holotypus Table 2, reléve 8, Figure 4)
Dominant species: Fraxinus pennsylvanica Marshall(3 - 4) (Ic
= 82.83) and Carex otrubae Podp.(r - 2) (Ic = 27.27)
Diagnostic species: Carex otrubae (Φ = 0.73), Lycopus europaeus (Φ = 0.66), Glyceria maxima (Hartman) Holmberg
Figure 2. Results of cluster analysis (cluster A – relevés with the dominant species Acer negundo L.; cluster B – relevés with the dominant
species Fraxinus pennsylvanica Marshall; cluster C – relevés of the ass.
Sambuco nigrae–Aceretum negundo Exner 2004)
Slika 2. Rezultati klaster analize (klaster A – snimci s dominantnom vrstom
Acer negundo L.; klaster B – snimci s dominantnom vrstom Fraxinus pennsylvanica Marshall; klaster C – snimci asocijacije Sambuco nigrae–Aceretum negundo Exner 2004)
Figure 3. the Ass. Rubo caesii–Aceretum negundi ass. nova
Slika 3. Nova asocijacija Rubo caesii–Aceretum negundi
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 165
Figure 4. the Ass. Carici otrubae–Fraxinetum pennsylvanicae ass. nova
Slika 4. Nova asocijacija Carici otrubae–Fraxinetum pennsylvanicae
(Φ = 0.66), Fraxinus pennsylvanica Marshall (Φ = 0.63),
Solanum dulcamara (Φ = 0.55), Polygonum hydropiper L.
(Φ = 0.45).
Diagnosis: Ass. Carici otrubae–Fraxinetum pennsylvanicae
is developed on marshy, gley (eugley), hypogley, saline soils,
on altitudes between 73 - 78 m on a flat surface, up to 5° inclination. Dominant species Fraxinus pennsylvanica can
reach height up to 10 m in the canopy layer, while younger
individuals are present also in the lower layers. Total cover
varies from 80 to 100 %, averaging about 95 %. Total of 57
taxa are present in all 11 relevés. The average number of species per stand is 14. Since Fraxinus pennsylvanica, the dominant and nominal species is invasive, the association has an
invasive character itself. The fact that individuals of the dominant invasive species are present in shrub and herb layers,
indicate that this community is in progradation, and the final
stage will result in the development of monodominant forest, as well as the first association. The other invasive species
present in this association are the following: Acer negundo,
Amorpha fruticosa, Aster lanceolatus Willd., Bidens frondosa,
Celtis occidentalis L. and Gleditsia triacanthos.
Ecology and synchorology of associations – Ekologija
i sinhorologija asocijacija
Stands of new associations Rubo caesii–Aceretum negundi
and Carici otrubae–Fraxinetum pennsylvanicae are so far
known only from the investigated area, on the former floodplain of Carska bara in northeastern Serbia. There are typical alluvial landforms, both natural and anthropogenically
conditioned, due to frequent man interventions. The geological structure is dominated by sediments of Neogene age
and pedological structure by alluvial saline soil and gleys,
marshy black soil and smonitsa and solonchak's solonetz.
Mean annual temperature is 11.6 °C. The coldest month is
January with the mean monthly temperature of 0.1 °C,
while August is the hottest month with the mean temperature of 22 °C. The mean temperature of winter months is
above zero, which is regularly the case in recent years. The
average of annual rainfall is 609.8 mm. In the vegetation
period (April – October), mean annual amount of precipitation is 68.0 mm in the monthly average. The potential
steppe and forest-steppe vegetation (alliances Festucion
rupicolae Soó 1940, and Aceri tatarici–Quercion Zólyomi et
Jakucs 1957) are determined by these climatic conditions
and geographic location of the wider surrounding area. The
potential vegetation in the riparian zone of this area is presented by the azonal vegetation of hygrophile flooded forests of the alliances Salicion albae Soó 1940 (EUNIS G1.1)
and Fraxinion angustifoliae Pedrotti 1970 (EUNIS G1.2)
(Davies et al., 2004). However, the natural vegetation has
been greatly disturbed, providing the space for the establishment and spread of invasive plant communities.
Syntaxonomical scheme – Sintaksonomska shema
Class Robinietea Jurko 1963 ex Hadač et Sofron 1980
Order Chelidonio–Robinietalia Jurko ex Hadač et Sofron
1980
Alliance non defined
Ass. Rubo caesii–Aceretum negundi Batanjski et S.
Jovanović ass. nova hoc loco
Ass. Carici otrubae–Fraxinetum pennsylvanicae Batanjski et S. Jovanović ass. nova hoc loco
dISCUSSIOn
RASPRAVA
According to Stojšić (2010), the most common forest community in the riparian habitats of the investigated area is
Salicetum albae pannonicum Parabućski (1965) 1972. There
are also Populetum nigro–albae Slavnić 1952 and the fragmented communities Fraxino–Quercetum roboris Rudski
(1940) 1949. The native riparian forest vegetation was under strong anthropogenic pressure (before it was protected),
which resulted both in occurrence and the spreading of the
invasive species, especially woody ones. Negative direction
of secondary succession of native forest vegetation, and the
substitution of the primary wet meadow habitats in riparian zone continues today. Accordingly, the natural forest
communities Populetum nigro–albae and Fraxino–Quercetum roboris are extensively replaced by new community of
invasive species Fraxinus pennsylvanica. On the other hand,
Acer negundo has expanded even more and increasingly occupies the habitats of the native community Salicetum albae
pannonicum (next to local waters), and some habitats of ass.
Populetum nigro–albae, forming the stable stands of new
invasive community. Similar situation was observed in the
middle course of the Vistula river in Poland (Künstler,
1999). However, the substitution of communities is not so
166
strictly divided by habitats, and observed invasive woody
species frequently occure together in the analyzed area of
Carska bara.
In the light of the anthropogenic interventions and frequent
inundation of these habitats, the competitive characteristics
of the analyzed invasive species are highlighted, which explain the establishment of their communities. This led to
the formation of "novel ecosystems", which are, in this study,
formed due to the occurrence of invasive woody plants, that
prevent the growth and regeneration of pre-existing plants
and thus reduce the potential for system redevelopment
(Hobbs et al., 2006; Hobbs et al., 2009). Acer negundo occupies the flood plain forests of the willow and poplar in
disturbed conditions (due to vegetative growth), which prevents their natural regeneration (Rosario, 1988; Künstler,
1999). Possible limiting factor is the frequency and duration of flooding (Mędrzycki, 2011). In contrast, Fraxinus
pennsylvanica has tolerance to flooding and is characterized
by extremely rapid growth (Gucker, 2005; Kremer et al.,
2006), due to which it can successfully suppress the native
species from competition, establishing thus the stable community. Acer negundo and Fraxinus pennsylvanica often
cenotically associate with different kinds of willow and poplar in their homeland (Csiszár and Bartha, 2008; Udvardy,
2008). Such situation was observed and analyzed in this reserve. As these communities are similar in composition and
structure to those in their native conditions, it should not
be surprising that analyzed invasive woody species are not
only naturalized, but cenotically stabilized in the study area.
Description and classification of the communities of some
woody alien species is done by several authors (Jurko, 1963;
Hadač and Sofron, 1980; Zerbe, 2003; Exner and Willner,
2004; Sîrbu and Oprea, 2011), argumenting why non-native
species can build communities which can be very stable and
rich in characteristic species as native communities. Since
many ruderal or segetal communities dominated by nonnative plants are identified in the literature as valid, there is
no reason why non-indigenous communities of woody
plants should not be (Hadač and Sofron, 1980), like in the
case of two new communities described in this paper.
The question of the syntaxonomical affiliation of two newly
described associations is challenging because these communities have not been described previously. As it was already mentioned, there is only one described association in
Europe so far, in which species Acer negundo is dominant.
It is Sambuco nigrae-Aceretum negundo described by Exner
and Willner (2004), within potential broad-leaved woodland areas of Austria. It was classified within the class Robinietea. Accordingly, we have also classified our two newly
described communities in the class Robinietea. This class
includes anthropogenic tree communities, colonizing disturbed habitats such as deforested lands, selvedges, agricultural and industrial fallows etc. Only two alliances were de-
Šumarski list, 3–4, CXXXIX (2015), 155–169
scribed within this class. Alliance Chelidonio–Robinion
Hadač & Sofron 1980 occurs at loamy, mesic, eutrophic
soils, or on solidified screes, usually on Southern, Eastern
or Western slopes, and the other one Balloto nigrae–Robinion Hadač & Sofron 1980, develops at relatively poor sandy,
dry soils, mainly in lowlands. Since the newly described invasive woody associations were found in anthropogenically
disturbed habitats in the plains, it is reasonable to classify
them within the class Robinietea. Diagnostic and dominant
species of Robinietea class are: Robinia pseudoacacia, Chelidonium majus L., Sambucus nigra L., Poa nemoralis L. and
Impatiens parviflora DC. (Hadač and Sofron, 1980; Chytrý
and Tichý, 2003). The fact that invasive species is the dominant one (Robinia pseudoacacia), suggests that the class
Robinietea includes communities whose occurrence and establishment are conditioned by anthropogenic activities. In
compliance with this standpoint and currently available
data on higher syntaxonomic categories, we have classified
both new invasive communities within the mentioned class.
We agree with Chytrý and Tichý (2003) that Robinietea class
is insufficiently described in relation to diagnostic species
throughout its geographic range. However, the diagnostic
species and the habitat types of two newly described communities in this work, suggest that they cannot be classified
to neither of the two currently available alliances of Robinietea class. For this purpose, the authors consider that an
introduction of new syntaxa at the level of alliance or order,
including human-caused azonal communities, could be the
possible solution.
Another possible solution is further research, which would
determine whether some of the communities currently belonging to Robinietea class, could be assigned to other
higher syntaxa in accordance with native distribution of the
dominant species, as considered by Exner and Willner
(2004).
Considering the fact that described invasive forest communities represent a major threat to indigenous biodiversity
an immediate action plan is necessary for minimizing its
negative effects on the autochthonous flora and fragile
riparian habitats of Ramsar site and Special Nature Reserve
Carska bara. These results are extremely alarming, especially within the internationally important and protected
area.
ACKnOWLEdGMEnT
ZAHVALA
This study was supported by the Serbian Ministry of Education, Science and Technological development (Grant No.
173030). The authors wish to thank the management team
of SNR Carska bara for technical help during field investigation.
BATAnJSKI, v. et al.: NEW INVASIVE FORESt COMMUNItIES IN tHE RIPARIAN FRAGILE HABItAtS – tHE CASE StUDY FROM RAMSAR SItE CARSKA BARA... 167
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SAžETAK
Invazivne vrste, kao drugi faktor rizika ugrožavanja nativne biološke raznolikosti, jedan su od najvećih izazova
u očuvanju biološke raznolikosti u Europi prema Europskoj strategiji o invazivnim vrstama. Prijetnje autohtonoj biološkoj raznolikosti i degradacije prirodnih staništa postaju najistaknutije kada invazivne vrste postanu naturalizirane i formiraju stabilne zajednice. To može dovesti do homogenizacije ekosustava i smanjenja
autohtone raznolikosti vrsta, a konačni ishod može rezultirati u nastanku novih ekosustava. Veliki broj invazivnih biljaka su drveće. Dvadeset tri strane vrste drveća su identificirane u šumskim ekosustavima Srbije,
od kojih je 17 invazivno. Unatoč svim poznatim negativnim utjecajima koje invazivne biljne vrste mogu imati,
paradoksalno je što je razina istraživanja invazivnih biljnih zajednica, posebice gdje dominiraju drvenaste
vrste, mala. Alarmantno velika prisutnost vrlo invazivnih sjevernoameričkh drvenastih vrsta Acer negundo i
Fraxinus pennsylvanica zabilježena je u zaštićenom močvarnom području Carska Bara. One su uspostavile
stabilne šumske zajednice koje su analizirane i opisane u ovome radu. U znanstvenoj literaturi nedostaju podaci o stabiliziranim zajednicama izgrađenim od dve promatrane drvenaste invazivne vrste.
Istraživano područje Carska Bara nalazi se na aluvijalnoj ravnici između rijeka Tise i Begeja, u središnjem Banatu (Vojvodina, Srbija) (Slika 1). Zbog svojih prirodnih svojstava, uživa status specijalnog rezervata prirode,
ramsarskog područja, važnog područja za ptice (IBA), važnog područja za biljke (IPA), kao i Emerald i ASCI
područja.
Fitocenološka istraga odabranih priobalnih šumskih staništa provedena je u razdoblju od 2011. do 2013. godine. Trideset i dva snimka sakupljena su prema Braun-Blanquet metodologiji. Svi podaci su georeferencirani.
Osim vlastitih, 4 fitocenološka snimka slične asocijacije iz Austrije uključena su u analizu i korištena za usporedbu. Snimci su klasificirani i grupirani u klaster, za daljnju cenoekološku karakterizaciju i diferencijaciju.
Sve analize su rađene u PcOrd 6,0 softveru. Korišten je koncept dominantnih i dijagnostičkih vrsta.
Klaster analizom jasno se izdvajaju tri skupine snimaka (Slika 2). Klaster A se sastoji od 20 snimaka u kojima
dominira Acer negundo, a klaster B uključuje 11 snimaka gdje dominira Fraxinus pennsylvanica. Treći klaster
(C) od 4 snimka predstvalja posebnu skupinu sastojina ranije opisane zajednice Sambuco nigrae–Aceretum
negundo, s područja Austrije. Na temelju rezultata svih analiza, smatramo da su prve dvije skupine (klastera
A i B) specifične i dovoljno različite, te da se mogu definirati kao nove asocijacije.
Ove nove asocijacije Rubo caesii–Aceretum negundi (Tablica 1) (Slika 3) i Carici otrubae–Fraxinetum pennsylvanicae (Tablica 2) (Slika 4) do sada su poznate samo iz istraživanog područja. Tu je prirodna vegetacija
uveliko poremećena i sekundarna sukcesija vegetacije primarnih vlažnih staništa nastavlja se i danas. Prirodne
šumske zajednice Populetum nigro–albae i Fraxino–Quercetum roboris opsežno su zamijenjene novom zajednicom invazivne vrste Fraxinus pennsylvanica. S druge strane, Acer negundo se proširio i sve više zauzima
staništa autohtone zajednice Salicetum albae, a negdje i staništa ass. Populetum nigro–albae, formirajući stabilne sastojine nove invazivne zajednice. Međutim, zamjena zajednica nije tako strogo podijeljena po tipovima
staništima, te se promatrane invazivne vrste često nalaze zajedno.
Definiranje potpune sintaksonomske pripadnosti dvaju novih opisanih zajednica je u ovom trenutku teško,
jer one nisu nigdje prethodno opisane. Njihove dijagnostičke vrste i tipovi staništa ukazuju na to da se one ne
mogu svrstati niti u jednu od dve trenutno opisane sveze Robinietea klase. Kako ove opisane invazivne šumske
zajednice predstavljaju veliku opasnost za autohtonu biološku raznolikost, situacija je izuzetno alarmantna s
obzirom na činjenicu da je istraživano područje međunarodno važno i zaštićeno.
KLJUČNE RIJEČI: invazivne drvenaste vrste, Acer negundo, Fraxinus pennsylvanica, šumske zajednice, poplavne
zone