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Reciprocal contamination by invasive plants:


Analysis of trade exchange between Slovakia
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Article in Biologia July 2015


DOI: 10.1515/biolog-2015-0102

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Biologia 70/7: 893904, 2015
Section Botany
DOI: 10.1515/biolog-2015-0102

Reciprocal contamination by invasive plants:


analysis of trade exchange between Slovakia and Romania

Peter Ferus1*, Culita Srbu2, Pavol Eli jr.3, Jana Konpkov1, uba uriov3,
Costel Samuil2 & Adrian Oprea4
1
Mlyany Arboretum SAS, Institute of Forest Ecology SAS, Vieska nad itavou 178, SK-95152 Slepany, Slovakia; e-mail:
peter.ferus@savba.sk
2
Department of Plant Science, University of Agricultural Sciences and Veterinary Medicine, Mihail Sadoveanu Alley 3,
700490 Iasi, Romania
3
Department of Botany, Slovak Agricultural University, Tr. A. Hlinku 2, SK-94976 Nitra, Slovakia
4
Botanical Garden, University Alexandru Ioan Cuza, Dumbrava Rosie st. 7-9, 700 487 Iasi, Romania

y
Abstract: In this work, potential contamination by invasive plant propagules as a result of trade exchange between Slovakia

p
and Romania, was assessed. National lists, describing biology and ecology of 30 worst invasive plant taxa, were formulated,
and trading in period 2006-2010 between countries analysed. Using norms for commodity impurity level, information on
co
species habitat occupancy and literature data dealing with seed/fruit attachment on roads we calculated then potential
invasive plant propagule export (PE) for each taxon. We found three fold higher total good export from Slovakia than in op-
posite direction, increasing export of commodities potentially containing invasive plant propagules exported from Romania
to Slovakia and rise of road compared to railway transport. PEs for Slovak invasive plant taxa were one-two orders higher
than those for Romanian ones. Potentially most exported taxa for Slovakia were: Amaranthus sp., Ambrosia artemisiifolia,
Galinsoga sp., Kochia scoparia and Sorghum halepense (tens to hundreds tonnes each). And these could mostly be exported
's
from Romania: Amaranthus sp., Ambrosia artemisiifolia, Artemisia annua, Conyza canadensis, Cuscuta campestris, Datura
stramonium, Erigeron annus, Galinsoga sp., Iva xanthiifolia, Kochia scoparia, Lycium barbarum, Sorghum halepense, Veron-
or

ica persica and Xanthium orientale subsp. italicum (units to tens tonnes each). High PE was signicantly associated with
cereals export. Our formula for PE is applicable for any inter- and intra-continental trade exchange.
Key words: invasive plants; Slovakia; Romania; trade; transport; potential introduction.
th

Introduction viding us fresh water, food and bres or recreation (Bin-


Au

imelis et al. 2007; Pejchar & Mooney 2009). Gurevitch


Invasive taxa are dened as taxa spreading in areas & Padilla (2004) discussed alien caused species extinc-
where they are not native. They represent a subset tions and concluded that this problem is system-species
of naturalized taxa producing reproductive ospring, specic.
often in very large numbers, at considerable distances In the past, there were some attempts to calculate
from the parent plants and thus have the potential to how much invasive organisms cost. Pimentel (2009) re-
spread fast over a large area (Rejmnek 2000; Pyek & ported for the USA that total damages caused by them
Richardson 2006). in environment and human health as well as control
In Europe more than 5700 alien plant species expenses are yearly of 120 billion USD. Other work,
was recognized (Lambdon et al. 2008). As indicate focussed on alien plants, referred to 344 million USD
Stohlgren et al. (2011), compared to North America for 10 years lasting chemical ght against 12 species in
with 40.1%, this continent has only 2.1% of alien species the UK (Williamson, 1996). Scalera (2010) presents a
among the 536 most widespread species of its ora. De- 15 years overview on project number and their budget-
spite of this disproportion, they can cause many com- ing by European Union. From 1992 to 2006 EU spent
plications to European man and nature. We can distin- more than 132 million EUR for almost 300 LIFE and
guish between direct eects of invasive plants on human FP projects.
wellbeing, which are of public concern (health prob- Huge direct and indirect negative eects of non-
lems, road system and articial waterways injury, plant native ora spontaneous dispersal on man activities and
control costs), from indirect ones (often not of public interests led to concentration of research on causal re-
concern), through alteration of ecosystem services, pro- lationships explaining plant invasion success and en-

* Corresponding author


c 2015 Institute of Botany, Slovak Academy of Sciences
894 P. Ferus et al.

abling prediction of future events. During the past two E1 Dry grasslands
decades, works reconstructing introduction pathways E1.94 Inland dune pioneer grassland
(Jehlk 1998; Mack & Lonsdale 2001; Kowarik & von der E2 Mesic grasslands
Lippe 2007; Meyerson & Mooney 2007; Hulme 2009), E3 Seasonally wet and wet grasslands
outlining socio-economic (Vil` a & Pujadas 2001; Lin et E4 Alpine and subalpine grasslands
E5 Woodland fringes and clearings and tall forb habitats
al., 2007; Pyek et al. 2010; Essl et al. 2011), ecolog-
E5.1 Anthropogenic tall-forb stands
ical (Davis et al. 2000; Rejmnek 2000; Lockwood et E5.4 Moist or wet tall-herb and fern fringes and meadows
al. 2005) and biological pre-requisites (Kolar & Lodge E5.5 Subalpine moist or wet tall-herb and fern stands
2001; Pyek & Richardson 2007) of successful inva- E5.6 Anthropogenic forb-rich stands
sion, accumulated. Scientic literature on possible con- E6 Inland salt steppes
sequences of climate change for invasive species also F3.1 Temperate thickets and scrub
appears (Thuiller et al. 2007; Hulme 2014). F9.1 Riverine scrub
All the knowledge in this area should lead to cre- G1 Broadleaved deciduous woodland
ation of a sophisticated control system anchored in the G1.C Highly articial broadleaved deciduous forestry
plantations
state legislative, which would minimalize further intro-
G4 Mixed deciduous and coniferous woodland
duction of already invasive taxa in general (Hulme et al.
G5.2 Small broadleaved deciduous anthropogenic wood-
2009). One of the most important moments in this re- lands
spect is integrating the introduction pathways into pol- G5.6 Early-stage natural and semi-natural woodlands and
icy (Hulme et al. 2008). Although there is information regrowth
on principal import routes and invasion success for most H5.6 Trampled areas

y
of invasive plant taxa based on database data (Pyek I Regularly or recently cultivated agricultural, horticul-
et al. 2011) and we can model transfer risk for some tural and domestic habitats

p
invasive organisms (Sellens et al. 2007), a huge gap in I1 Arable land and market gardens
detailed information on rates of individual species in- co I1.5 Bare tilled, fallow or recently abandoned arable land
J Constructed, industrial and other articial habitats
troduction still remains (Hulme 2009).
J4 Transport networks and other constructed hard-
Since the European control system is still under surfaced areas
construction and executive organs do not inspect goods J6.1 Waste resulting from building construction or demo-
coming in to be able to express the total infestation by lition
invasive plant propagules, on example of Slovakia and
's
Romania we tried to dene at least potential transport Statistical data on trade and transport
of invasive plant propagules between them, carried out Raw statistical data on trade exchange (export/import) and
or

by trade exchange. transport means utilized for period 20062010 were provided
by Statistical Oce of the Slovak Republic (personal com-
munication; Benkoviov 20072011) and National Statisti-
Material and methods cal Institute of Romania (Voineagu 20072011). Concretely,
th

Characterization of the worst invasive plant taxa we were interested in total export/import between Slovakia
A list of thirty most dangerous (spreading) invasive plant and Romania, export/import of goods potentially contain-
species (ecological nomenclature according to Richardson et ing invasive plant propagules (categorised by Statistical Of-
Au

al. 2000) in Slovakia and Romania, respectively, was com- ce of the Slovak Republic) 1. cereals; 2. other seeds,
pleted using actual national lists of invasive plants (Med- fruits and industrial as well as medicinal plants + straw
veck et el. 2012; Gojdiov et al. 2002; Eli 2009; Srbu and volume fodder (shortly other seeds); 3. vegetables, ed-
& Oprea 2011), as well as personal evidence of participat- ible plants, roots and tubers (shortly root crops); 4. wood
ing botanists. Taxonomic nomenclature follows the classical and wooden stu (shortly wood); 5. living animals; and 6.
work Flora Europea (Tutin et al. 19641980) and Greuter eece. There was a suggestion that every category fully con-
(2003). In the lists we also indicate important plant char- sists of goods with probability to be infested by seeds/fruits
acteristics, like families they belong to, geographical ori- of other plant species. We omitted category living plants
gins, characters of the plants in relation to human activities because of common soil substrate sterilisation as well as cat-
(hemerophyte/xenophyte, dened by Rikli 1904), life forms, egory fruits, nuts and citrus peels because they are clean
modes of reproduction and seed distribution (Table 1). and packed in paper boxes without straw bedding.
For description of habitats they reside in, EUNIS
nomenclature available on internet (http://eunis.eea.europa. Potential export of invasive plant propagules
eu/habitats.jsp), was applied. Taxa analysed in this work Potential export of invasive plant propagules (PE, in tonnes)
occupy following habitats, as derived from their presence from Slovakia to Romania and vice-versa was calculated for
in distinct plant alliances (Davies et al. 2004), with speci- respective taxa in both national lists using formula compris-
cations for Slovakia (Jehlk 1998) and Romania (Srbu & ing good and transport means contamination:
Oprea 2011), respectively:
C1 Surface standing waters 
x

y

C2 Surface running waters PE = (pn cn Cn ) + (lk tk ak T SWl )


C3 Littoral zone of inland surface waterbodies n=1 k,l=1

C3.1 Species rich helophyte beds


D5 Sedge and reedbeds, normally without free-standing where p normative for maximal foreign matter content
water in a commodity; c correction of the normative in order
D6 Inland saline and brackish marshes and reedbeds to approximate weed seed/fruits portion; C amount of
Reciprocal contamination by invasive plants 895
Table 1. Biological characteristics and residence habitats of thirty most dangerous invasive plant taxa from Slovakia and Romania.
Abbreviations: SK Slovakia, RO Romania; Ace Aceraceae, Ama Amaranthaceae, Api Apiaceae, Ast Asteraceae, Bal
Balsaminaceae, Chen Chenopodiaceae, Cuc Cucurbitaceae, Cus Cuscutaceae, Fab Fabaceae, Hyd Hydrocharitaceae, Jun
Juncaceae, Phyt Phytolaccaceae, Poa Poaceae, Pol Polygonaceae, Sim Simaroubaceae, Scr Scrophulariaceae, Sol Solanaceae,
Vit Vitaceae, Am America, As Asia, Cau- Caucassus, Eur Europe, Him Himalaya, Md Mediterranean, N North, S South,
SW South-West, E East, C Central; h hemerophyte, x xenophyte; Ph- phanerophyte, tree, Ph- h phanerophyte-shrub,
Ph- phanerophyte-liana, G geophyte, H hemicryptophyte, HH hydrophyte, Ht hemitherophyte, T therophyte, sex sexual,
veg vegetative; ane anemochory, ant anthropochory, bar barochory, end endozoochory, epi epizoochory, hyd hydrochory
and myr myrmecochory. For description of EUNIS habitat codes see Material and methods.

Taxon Country Family Origin H/X Life form Reprod. Dispersal EUNIS habitats in Slovakia
mechanisms vs. Romania

Acer negundo L. SK, RO Ace Am N h Ph- sex+veg ane, ant C3, D5, E5.4, E5.6, G1,
G1.C, G4, G5.2, I1.5, J4,
J6.1/E5.1, G1, G1.C, I1.5,
J4, J6.1
Ailanthus altissima SK, RO Sim As E h Ph- sex+veg ane, ant E5.6, G1, I1.5, J4, J6.1/E1,
(Mill.) Swingle E5.1, G1.C, I1.5, J4, J6.1
Amaranthus albus L. SK, RO Ama Am N(C) x T sex ane, end, ant C3, E5.1, E5.6, H5.6, I1,
I1.5, J4/C3, E1, E1.94, E5.1,
I1, I1.5, J4, J6.1
Amaranthus powellii SK, RO Ama Am N x T sex ane, end, ant C3, E5.1, E5.6, I1, I1.5, J4,
S.Watson J6.1/C3, E1, E5.1, H5.6, I1,
I1.5, J4, J6.1

y
Amaranthus retroflexus SK, RO Ama Am N x T sex ane, end, ant C3, E5.1, E5.6, H5.6, I1,
L. I1.5, J4, J6.1/C3, E1, E2,

p
E5.1, E5.4, E6, G1, H5.6, I1,
co I1.5, J4, J6.1
Ambrosia artemisiifolia SK, RO Ast Am N x T sex ane, epio, anto C3, E5.6, H5.6, I1, I1.5,
L. J4, J6.1/C3, D5, D6, E5.1,
H5.6, I1, I1.5, J4, J6.1
Amorpha fruticosa L. RO Fab Am N h h
Ph- sex+veg ane, ant C3, E1, E5.4, E6, G1, G1.C,
G5.2, I1.5, J4, J6.1
's
Artemisia annua L. SK, RO Ast As h T sex ane C3, E5.1, E5.6, H5.6, I1.5,
J4, J6.1/C3, E1, E5.1, F3.1,
G1.C, G5.2, H5.6, I1, I1.5,
J4, J6.1
or

Aster lanceolatus Willd. SK Ast Am N h H sex+veg ane, epi, ant E5.1, E5.4, F9.1
Aster salignus Willd. SK Ast Am N h H sex+veg ane, epi, ant E5.1, E5.4, F9.1
Conyza canadensis (L.) RO Ast Am N x T sex ane, ant C3, E1, E1.94, E2, E5.1,
th

Cronq. E5.4, F3.1, F9.1, G1, G1.C,


G5.2, G5.6, H5.6, I1, I1.5,
J4, J6,1
Cuscuta campestris RO Cus Am N x T sex ant(bar, end, hyd) E1, E1.94, E2, E5.1, I1, I1.5,
Au

Yunck. J4, J6.1


Datura stramonium L. RO Sol Am x T sex ant, ane, hyd C3, E5.1, I1, I1.5, J4, J6.1
Echinocystis lobata (F. SK Cuc Am N h T sex hyd, ant C3, D5, D5.1, E2, E5.4,
Michx.) Torr. et A. Gray F9.1, G1
Elodea canadensis SK Hyd Am N x HH veg(sex) hyd, epi C1, C3, C3.1, D5.1
Michx.
Elodea nuttallii RO Hyd Am N x HH veg(sex) hyd C1, C2
(Planch.) H.St John
Erigeron annuus (L.) SK, RO Ast Am N x Ht(T-H) sex+veg ane(hyd) C3, E2, E5.4, E5.6, G1, I1.5,
Pers. J4, J6.1/C3, D5, D6, E1,
E1.94, E2, E3, E5.1, E5.3,
E5.4, F3.1, F9.1, G1, G1.C,
G5.6, H5.6, I1, I1.5, J4, J6.1
Galinsoga parviflora SK, RO Ast Am S x T sex+(veg) ane, ant, hyd,epi C3, E1.1, I1, I1.5, J4,
Cav. J6.1/C3, E2, E3, E5.1, E5.4,
F9.1, G1, G5.6, H5.6, I1,
I1.5, J4, J6.1
Galinsoga quadriradiata SK, RO Ast Am C x T sex ane, ant, hyd, epi C3, E5.4, E5.6, I1, I1.5, J4,
Ruiz et Pav. J6.1/C3, E2, E5.1, F9.1, I1
Helianthus tuberosus L. SK, RO Ast Am N h G sex+veg hyd, ant C3, D5.1, E5.1, E5.4,
E5.6/E5.1, E5.4, F9.1, G1.C,
G5.2
Heracleum mantegazz- SK Api Cau h H sex anemo, hydro E5.4, E5.6, G1
ianum Sommier et
Levier
896 P. Ferus et al.
Table 1. (continued)

Taxon Country Family Origin H/X Life form Reprod. Dispersal EUNIS habitats in Slovakia
mechanisms vs. Romania

Impatiens glandulifera SK, RO Bal Him h T sex antropo, hydro C3, D5, E2, E5.4, E5.6,
Royle G1/E5.1, E5.4, F9.1, G1
Impatiens parviflora DC. SK Bal As C h T sex ant, myr? C3, E5.4, E5.6, G1
Iva xanthiifolia Nutt. SK, RO Ast Am N x T sex ant, bar, ane C3, E5.1, E5.6, I1.5, J4,
J6.1/C3, E5.4, I1, I1.5, J4,
J6.1
Juncus tenuis Willd. RO Jun Am N x G sex+veg ane, ant, end C3, E2, E3, E4, E5.4, F9.1,
G1, H5.6, I1.5, J4, J6.1
Kochia scoparia (L.) SK, RO Chen As, Eur E h T sex ane, ant(hyd) C3, E6, E6.2, I1, I1.5, J4,
Schrad. J6.1/E5.1, H5.6, I1, I1.5, J4,
J6.1,
Lycium barbarum L. SK, RO Sol As E h h
Ph- sex+veg ant, end E5.1, E5.6, F3.1, I1.5, J4,
J6.1/E1, E5.1, F3.1, G1.C,
G5.2, I1, I1.5, J4, J6.1
Panicum miliaceum L. SK Poa As SW x T sex ant, end, bar I1.5, J4, J6.1
subsp. ruderale (Kitag.)
Tzvelev
Parthenocissus inserta RO Vit Am N h Ph- sex+veg ant, end E5.1, G1, G1.C, G5.2
(A. Kerner) Fritsch

y
Phytolacca americana L. SK Phyt Am N h T sex end, ant C3.5, E5.6, G1, G4, G5.6,
I1.5, J4, J6.1

p
Reynoutria bohemica SK, RO Pol Eur C h G (sex)+veg ant, ane, hyd D5.1, E5.4, E5.6, I1.5, J4,
Chrtek et Chrtkov co J6.1/D5.1, E5.4, E5.6, I1.5,
J4, J6.1
Reynoutria japonica SK Pol As E h H sex+veg ant, ane, hyd D5.1, E5.4, E5.6, I1.5, J4,
Houtt. J6.1
Robinia pseudacacia L. SK Fab Am N h Ph- sex+veg antropo, anemo E5.6, G1, G1.C, G5.2
Rudbeckia laciniata L. SK, RO Ast Am N h H sex ant, hyd(ane) C3, E5.4, F9.1, G1/C3, E2,
E5.1, E5.4, G1
's
Solidago canadensis L. SK, RO Ast Am N h H sex+veg ant, ane D5, E5.4, E5.6, G1, G1.C,
G5.2/E5.1, E5.4, F3.1, G1,
G5.6, I1.5, J4, J6.1
or

Solidago gigantea Aiton SK, RO Ast Am N h H sex+veg ant, ane C3, D5, E5.4, E5.6, G1,
G1.C, G5.2/E5.1, E5.4,
F3.1, G1, G5.6, I1.5, J4,
th

J6.1
Sorghum halepense (L.) SK, RO Poa Md x G sex+veg ant, ane, bar, hyd, end I1, I1.5, J4/E1, E5.1, I1,
Pers. I1.5, J4, J6.1
Veronica persica Poir. RO Scr As SW x T sex ant, myr C3, E1, E1.94, E2, E5.1,
Au

E5.5, F9.1, G1.C, G5.2, I1,


I1.5, J4, J6.1
Xanthium orientale RO Ast Am N x T sex epi, bar C3, D6, E1, E2, E5.1, E6,
L. subsp. italicum F9.1, G1, H5.6, I1, I1.5, J4,
(Moretti) Greuter J6.1
Xanthium spinosum L. RO Ast Am S x T sex epi, bar C3, E1, E2, E5.1, E6, H5.6,
I1, I1.5, J4, J6.1

exported commodity (t); l length of the transport journey (potatoes and sugar beet), where high portion of soil con-
(km); t turning coecient, describing how many times taminated by weed seeds is expected, we analysed works
transport means had to turn to deliver concrete amount of on eld seed bank. Upper 0.15 m thick part of brown soil
a commodity; a propagule attachment coecient (pcs of prole (specic weight of 1.3 t m3 ) in elds of southern
seeds/fruits per km); T SW thousand seed/fruit weight Slovakia was composed from 0.0088% by weed seeds (de-
(g); n commodity; k transport means; l invasive plant rived from Lacko-Bartoov & Krolk 2001). Since there
taxon. is no normative for wood purity, we applied that for wood
Norms for acceptable weed seed contamination of com- chips, and dened the portion of seeds in beech forest (most
modities originating in the eld, grassland or forest are listed common in Slovakia and Romania) soil (0.0023%; derived
in Table 2. Since they mainly dene course impurities level, from Godefroid et al. 2006). Only one work deals with weed
using available literature we tried to ascertain the weed seed seeds/fruits attached on sheep eece, in detail (Olson et al.
content in them. For cereals of 9597% purity we derived 1997). We utilized it for calculation of the infestation level
1.56% of total harvested matter, which can be attributed to (0.0043%). Living animals could also carry invasive plant
weed seeds (Barac et al. 2013). For other seed crops (legumes propagules in their digestive tract. Taking into account aver-
and oil crops) 1.72% was approximated. In case of root crops age cattle (cow and bull) weight of 900 kg, volume of the cat-
Reciprocal contamination by invasive plants 897
Table 2. Normative for maximal impurity (inorganic and organic impurities as well as weed seeds) content and maximal weed seed
content in traded commodities (Sources: www.sutn.sk, Oravec et al. (2012), www.amsem.ro, www.incs.ro), as well as invasive plant
taxa from national lists, usually growing in respective crop group(s) (EUNIS habitat I1; Sources: Lka et al. (1995); Kohaut (2001);
Anghel et al. (1972); Srbu and Oprea (2011)), grasslands (EUNIS habitat E1, E1.94, E2, E3, E4, E6) and forests (EUNIS habitat G1,
G1.C, G4, G5.2, G5.6). * for allimetary usage.

Max. impurity content (%) Max. weed seed content (%) Species growing in
Commodity
SK RO SK RO SK RO

Cereals 35 12 0.10.4* 410 pcs Amaranthus sp., A. all residing in I1 habitats except of
per artemisiifolia, Galin- X. spinosum
5001000 g soga sp., K. scoparia, S.
sample halepense
Legumes 0.53 2 0.1 Amaranthus sp., S. Amaranthus sp., C. canadensis, D.
halepense stramonium, E. annuus, Galinsoga
sp., S. halepense, V. persica, X.
orientale subsp. italicum
Oil crops 18 12 515 pcs Amaranthus sp., A. all residing in I1 habitats except of
per 1000 g artemisiifolia, Galin- A. annua
sample soga sp., K. scoparia, S.
halepense
Potatoes 10 1 0.1 Amaranthus sp., A. all residing in I1 habitats except of
artemisiifolia, Galinsoga A. artemisiifolia, A. annua, I. xan-
sp., S. halepense thiifolia, K. scoparia, L. barbarum,

y
X. spinosum
Sugar beet 1525 3 0.5 Amaranthus sp., A. all residing in I1 habitats except of

p
artemisiifolia, Galinsoga A. albus, A.artemisiifolia, I. xan-
co sp., S. halepense thiifolia, K. scoparia, L. barbarum,
S. halepense, X. spinosum
Fleece 36 E. lobata, E. annuus, G. all taxa except of A. negundo, A.
parviflora, I. glandulifera, artemisiifolia, D. stramonium, E.
K. scoparia nuttallii, H. tuberosus, Impatiens
sp., I. xanthiifolia, K. scoparia, R.
x bohemica, Solidago sp.
's
Wood chips 0.3 A. negundo, A. altissima, all taxa except of A. albus, A.
E. lobata, E. annuus, H. powellii, A. artemisiifolia, C.
mantegazzianum, Impa- campestris, D. stramonium, E. nut-
or

tiens sp., Ph. americana, talii, G. quadriradiata, I. xanthiifo-


R. pseudacacia, R. lacini- lia, K. scoparia, R. x bohemica, S.
ata, Solidago sp. halepense, X. spinosum
th
Au

tle digestive tract about 320 litres, Pleasant and Schlathers propagules per one km. Fluctuations in road occupancy by
(1994) results on 75100 weed seeds (of average TSW 2.10 g) seeds/fruits during year were excluded (Zwaenepoel et al.
per tonne of manure (specic weight 0.75 t m3 ) and fatal (2006) found the highest vehicle mud and seed contami-
eect of ensiling and digestion process on weed seed viability nation in January and late summer). Thousand seed/fruit
(Blackshaw & Rode 1991), only 0.0002% of animal weight weights of species accompanying transport networks (EU-
can be represented by viable weed seed (because of no litera- NIS habitat J4) are listed in Table 3. Amount of propagules
ture data as well as relatively small importance, propagules gained on roads was expressed for each plant taxon as the
on animal hoofs and in straw bedding were omitted). Nor- only one contaminant. We did not take into account po-
matives and coecients were then applied for calculation of tential dierences in the invasive plant list composition of
commodity contamination by propagules of taxa residing in transit country between Slovakia and Romania Hungary.
following habitats: I1, E1, E2, E3, E4, E6, G1, G1.C, G4, Since cargo trains do not move fast (max. 90 km h1 ), rail-
G5.2 and/or G5.6 (each as the only one contaminant). yards are kept clean from taller vegetation, carriage surfaces
Regarding transport, we suppose that goods were ex- often oer low deposit area for seeds/fruits, which can be
changed between capitals (Bratislava and Bucharest). Thus, uplifted from rails or torn down especially from trees by
the shortest road connection between them, leading through air turbulence caused by train and/or wind when light and
Gy or, Budapest, Kecskemt, Szeged, Arad, Drobeta-Turnu dry, and since no research was done on the role of railway
Severin, Craiova, Pitesti, is of 1063 km. Turning coecient freight transport in weed redistribution, we did not take
(t) of was calculated from the total goods amount and ca- it into account. Although studies conducted in all coun-
pacity of a truck (approx. 26 t). Propagule attachment coef- tries downstream Danube revealed presence of both Elodea
cient (a) was derived from a study on diverse vehicle clear- species (Oahelov et al. 2007; Rth et al. 2003; Vukov
ing techniques (Rew 2011). On a US army personnel carrier et al. 2008; Sarbu et al. 2006), there is no evidence on
of 3 axles (7 m in length) were attached 0.4 propagules per the role of ships in their fragmentation and redistribution
km of dry paved roads (in wet conditions vehicles markedly (Nehring 2005; Leuven et al. 2009). Therefore, we omitted
lose attached seeds/fruits with travelled distance (Taylor et this transport means as a source of invasive plant propag-
al., 2012)). Thus, for a truck of 5 axles we count with 0.67 ules.
898 P. Ferus et al.
Table 3. Thousand seed/fruit weight (g) of taxa residing in areas along road communications (EUNIS habitat J4).

Taxon Thousand seed/fruit weight (g) Reference

Acer negundo L. 34.26 Csontos et al. (2007)


Ailanthus altissima (Mill.) Swingle 38.49 Csontos et al. (2007)
Amaranthus albus L. 0.32 Moravcov et al. (2010)
Amaranthus powellii S.Watson 0.53 Moravcov et al. (2010)
Amaranthus retroflexus L. 0.45 Moravcov et al. (2010)
Ambrosia artemisiifolia L. 4.87 Moravcov et al. (2010)
Amorpha fruticosa L. 12.96 our determination
Artemisia annua L. 0.03 http://www.kew.org
Conyza canadensis (L.) Cronq. 0.04 Moravcov et al. (2010)
Cuscuta campestris Yunck. 0.80 Moravcov et al. (2010)
Datura stramonium L. 7.63 Moravcov et al. (2010)
Echinocystis lobata (F. Michx.) Torr. et A. Gray 284 Tor
ok et al. (2013)
Erigeron annuus (L.) Pers. 0.04 Moravcov et al. (2010)
Galinsoga parviflora Cav. 0.19 Moravcov et al. (2010)
Galinsoga quadriradiata Ruiz et Pav. 0.22 Moravcov et al. (2010)
Helianthus tuberosus L. 4.89 Moravcov et al. (2010)
Heracleum mantegazzianum Sommier et Levier 15.18 Moravcov et al. (2010)
Impatiens glandulifera Royle 9.54 Moravcov et al. (2010)
Impatiens parviflora DC. 5.69 Moravcov et al. (2010)
Iva xanthiifolia Nutt. 0.99 Moravcov et al. (2010)
Juncus tenuis Willd. 0.01 Moravcov et al. (2010)

y
Kochia scoparia (L.) Schrad. 1.22 Moravcov et al. (2010)
Lycium barbarum L. 1.29 our determination
Panicum miliaceum L. subsp. ruderale (Kitag.) Tzvelev 3.96 Tor
ok et al. (2013)

p
Parthenocissus inserta (A. Kerner) Fritsch 34.05 Tor
ok et al. (2013)
Phytolacca americana L. co 11.88 Tor
ok et al. (2013)
Reynoutria bohemica Chrtek et Chrtkov 1.57 estimated from Csontos et al. (2007)
Reynoutria japonica Houtt. 1.66 Tor
ok et al. (2013)
Robinia pseudacacia L. 22.60 Tor
ok et al. (2013)
Rudbeckia laciniata L. 2.66 Csontos et al. (2007)
Solidago canadensis L. 0.07 Moravcov et al. (2010)
Solidago gigantea Aiton 0.18 Moravcov et al. (2010)
's
Sorghum halepense (L.) Pers. 5.17 our determination
Veronica persica Poir. 0.53 Moravcov et al. (2010)
Xanthium orientale L. subsp. italicum (Moretti) Greuter 326 our determination
or

Xanthium spinosum L. 116 our determination


th

Results ily, and originating in North-America dominated in it.


As in Slovak list, all of them are neophytes, but num-
National lists of 30 most dangerous invasive plant
Au

ber of xenophytes slightly exceeded number of hemero-


species phytes. Almost the same situation was found when
In the Slovak list of the worst invasive plants (Table 1) analysing life forms, modes of reproduction and types
we found species belonging particularly to Asteraceae of spreading assistance. Regarding the EUNIS habitat
family (12), and originating from North-America. All composition, we found marked dierences from Slovakia
of these species were introduced to Slovakia after New on rst ve places: 1. Anthropogenic tall-forb stands
World discovery (neophytes), from which almost two (E5.1) 25 taxa, 2. Bare tilled, fallow or recently aban-
thirds escaped from the culture (hemerophytes). This doned arable land (I1.5), Transport networks and other
list was dominated by therophytes (almost one half of constructed hard-surfaced areas (J4) and Waste result-
the taxa), species reproducing exclusively sexually or ing from building construction or demolition (J6.1) 24
combining sexual and vegetative reproduction (27), and taxa, 3. Arable land and market gardens (I1) 18 taxa,
most of them spreads using man as a vector. Among 4. Littoral zone of inland surface waterbodies (C3) 17
EUNIS habitats dominated 1. Anthropogenic forb-rich taxa, and 5. Dry grasslands (E1) 14 taxa.
stands 20 taxa (EUNIS habitat code E5.6), 2. Lit-
toral zone of inland surface waterbodies (C3) 19 taxa, Export/Import
3. Bare tilled, fallow or recently abandoned arable land Total Slovak export into Romania as well as Romanian
(I1.5), Transport networks and other constructed hard- export into Slovakia for period 20062010 exhibited rel-
surfaced areas (J4) 18 taxa, 4. Waste resulting from atively large uctuations (Fig. 1A) but in general Slo-
building construction or demolition (J6.1) 16 taxa and vakia exported approximately three fold more goods
5. Moist or wet tall-herb and fern fringes and meadows than Romania (round 600.000 t). In export of com-
(E5.4) 15 cases. modities potentially containing invasive plant propag-
The Romanian list is from two thirds identical with ules from Slovakia into Romania we observed decreasing
the Slovak one. Similarly, species from Asteraceae fam- tendency (Fig. 1B; from ca. 67 to 15 thousand tonnes).
Reciprocal contamination by invasive plants 899

both directions markedly decreased (from 73 to 24%


and from 47 to 30%, respectively). River transport var-
ied in range of per cents and aircraft cargo had only
marginal importance. In case of goods potentially con-
taining invasive plant propagules, the rise of road trans-
port and fall of railway transport were much sharper
(Fig. 3C, 3D). Utilization of river transport was casual.

Potential export of invasive plant propagules


Amounts of invasive plant propagules potentially trans-
ported from Slovakia to Romania in the studied period
uctuated markedly but in general were one-two orders
higher than those transported in the opposite way (Ta-
ble 4). However, in 2010 potential propagule exchange
balanced on approx. 600 t. Second, number of plant
taxa mostly imported to Slovakia was twofold higher
than that imported to Romania (17 : 8 taxa). Following
species could mainly leave Slovakia (tens to hundreds
tonnes each): Amaranthus sp., Ambrosia artemisiifolia,
Galinsoga sp., Kochia scoparia and Sorghum halepense.

y
And these could be mostly exported from Romania:
Amaranthus sp., Ambrosia artemisiifolia, Artemisia an-

p
nua, Conyza canadensis, Cuscuta campestris, Datura
stramonium, Erigeron annus, Galinsoga sp., Iva xan-
co
thiifolia, Kochia scoparia, Lycium barbarum, Sorghum
halepense, Veronica persica and Xanthium orientale
subsp. italicum (units to tens tonnes each). On the
Fig. 1. Development of Slovak-Romanian trade exchange (A) and other hand, potential for transfer of Aster sp., Elodea
export/import of goods potentially contaminated by alien plant canadensis and Helianthus tuberosus by trade from Slo-
's
propagules (B) in the period 20062010. vakia and Elodea nuttallii back was zero.
or

On the other hand, Romanian export grew continuously Discussion


from about 13 to 46 thousand tonnes.
In both countries export of living animals reached Globally, number of invasive plant species is a func-
th

the largest extent in the middle of studied period, tion of international trade (Westphal et al. 2008). Ge-
and Romanian export was only a third of the Slo- ographic and other socio-economic measures were not
vak one (Fig. 2A). Fleece export from Slovakia showed found to be important determinants of a countrys de-
Au

rather decrease (Fig. 2B). Romania exported much gree of biological invasion. Study of Vil`a & Pujadas
lower amounts of this commodity, except of year 2010, (2001) for Mediterranean countries showed that alien
when it jumped to 850 t. Decrease of Slovak root crops plants density can be best explained by human devel-
export stabilized at 500 t (Fig. 2C) but Romanian ex- opment index and imports from socio-economic param-
port showed increasing tendency. Export of other seeds eters, and transport network density with the percent-
from Slovakia into Romania peaked in 2008 (Fig. 2D; age of protected areas from land-use variables. Results
ca. 1100 t) and then fell to the starting value. Larger from modelling look very similar (Pyek et al. 2010).
export of this commodity from Romania was seen in Essl et al. (2011) add that number of established aliens
2010, only. Slovak export of cereals reached maximum is more closely related to socioeconomic activity from
in 2008 (approx. 17000 t) and then fell on 4000 t level the year 1900 than year 2000, although the majority of
(Fig. 2E). Compared to Slovakia, amount of cereals ex- species introductions occurred during the second half
ported from Romania was for most of the 5-years period of the 20th century.
negligible but jumped to 5500 t in 2010. Wood export Taxa included in the Slovak/Romanian national
of Slovakia declined gradually from almost 66000 t to list are already invasive. Moreover, eight of them
10000 t (Fig. 2F). For Romania we found an opposite (Ailanthus altissima, Ambrosia artemisiifolia, Echino-
trend. cystis lobata, Elodea canadensis, Heracleum mantegazz-
ianum, Impatiens glandulifera, Reynoutria japonica
Goods transport and Robinia pseudacacia) belong to the 100 worst Eu-
More and more goods were exported from Slovakia us- ropean alien organisms (DAISIE, http://www.europe-
ing road transport in period 20062010 and the same aliens.org/) and other six (Ailanthus altissima, Am-
situation was observed for Romania (from ca. 20 to 68% brosia artemisiifolia, Impatiens glandulifera, Reynou-
and from 44 to 70%, respectively (Fig. 3A, 3B). On tria japonica, Sorghum halepense and Xanthium spino-
the other hand, use of railways for trade exchange in sum) to the 100 worst world alien organisms
900 P. Ferus et al.

p y
co
's
or

Fig. 2. Slovak export/import of articles probably containing alien plants propagules into/from Romania: Living animals (A), wool (B),
root crops (C), other seeds (D), cereals (E) and wood (F) in the period 2006-2010.
th
Au

Fig. 3. Participation of road, railway and river transport in total Slovak export (A)/import (B) as well as in export (C)/import (D)
of goods potentially contaminated by alien plant propagules from 2006 till 2010.
Table 4. Potential exchange of invasive plant propagules between Slovakia and Romania by trade for period 2006-2010. For calculation procedure see Material and methods.

Potentially exported from Slovakia (t) Potentially exported from Romania (t)
Taxon Country
2006 2007 2008 2009 2010 2006 2007 2008 2009 2010

Acer negundo L. SK, RO 0.1128 0.2099 0.4029 0.2687 0.3435 0.0796 0.1108 0.1033 0.0679 0.1235
Ailanthus altissima (Mill.) Swingle SK, RO 0.1262 0.2357 0.4524 0.3018 0.3858 0.0897 0.1244 0.1160 0.0762 0.1749
Amaranthus albus L. SK, RO 1.2697 261.5027 289.3497 70.2889 67.8751 0.2114 1.2642 0.5285 1.3395 37.1653
Amaranthus powellii S.Watson SK, RO 1.2704 261.5040 289.3522 70.2905 67.8772 0.2118 1.2649 0.5291 1.3399 37.1660
Amaranthus retroflexus L. SK, RO 1.2701 261.5035 289.3513 70.2899 67.8764 0.2125 1.2652 0.5296 1.3408 37.1684
Ambrosia artemisiifolia L. SK, RO 1.2841 261.5303 289.4029 70.3245 67.9207 0.2214 1.2788 0.5420 1.3468 37.1444
Amorpha fruticosa L. RO 0.0311 0.0422 0.0396 0.0263 0.0849
Artemisia annua L. SK, RO 0.0001 0.0002 0.0004 0.0002 0.0003 0.2116 1.2638 0.5284 1.3400 37.1669
Aster lanceolatus Willd. SK 0 0 0 0 0
Aster salignus Willd. SK 0 0 0 0 0
Conyza canadensis (L.) Cronq. RO 0.2116 1.2638 0.5284 1.3400 37.1669
Au
Cuscuta campestris Yunck. RO 0.2125 1.2658 0.5299 1.3405 37.1670
Reciprocal contamination by invasive plants

Datura stramonium L. RO 0.2277 1.2877 0.5504 1.3538 37.1545


Echinocystis lobata (F. Michx.) Torr. et A. Gray SK 0.0044 0.0018 0.0024 0.0008 0.0007
Elodea canadensis Michx. SK 0 0 0 0 0
th
Elodea nuttallii (Planch.) H.St John RO 0 0 0 0 0
Erigeron annuus (L.) Pers. SK, RO 0.0059 0.0047 0.0030 0.0012 0.0016 0.2116 1.2638 0.5284 1.3400 37.1669
Galinsoga parviflora Cav. SK, RO 1.2706 261.5045 289.3484 70.2880 67.8744 0.2120 1.2643 0.5289 1.3403 37.1675
Galinsoga quadriradiata Ruiz et Pav. SK, RO 1.2694 261.5021 289.3486 70.2881 67.8741 0.2106 1.2632 0.5275 1.3389 37.1641
Helianthus tuberosus L. SK, RO 0 0 0 0 0 0.0009 0.0005 0.0008 0.0010 0.0027
or
Heracleum mantegazzianum Sommier et Levier SK 0.0044 0.0018 0.0024 0.0008 0.0007
Impatiens glandulifera Royle SK, RO 0.0044 0.0018 0.0024 0.0008 0.0007 0.0009 0.0005 0.0008 0.0010 0.0027
Impatiens parviflora DC. SK 0.0044 0.0018
's 0.0024 0.0008 0.0007
Iva xanthiifolia Nutt. SK, RO 0.0031 0.0060 0.0116 0.0077 0.0099 0.2125 1.2663 0.5304 1.3392 37.1307
Juncus tenuis Willd. RO 0.0014 0.0009 0.0016 0.0032 0.0401
Kochia scoparia (L.) Schrad. SK, RO 1.2477 261.5033 289.3569 70.2918 67.8763 0.2130 1.2670 0.5311 1.3397 37.1315
Lycium barbarum L. SK, RO 0.0041 0.0078 0.0151 0.0101 0.0129 0.2145 1.2678 0.5321 1.3408 37.1710
Panicum miliaceum L. subsp. ruderale (Kitag.) Tzvelev SK 0.0125 0.0241 0.0463 0.0310 0.0396
co
Parthenocissus inserta (A. Kerner) Fritsch RO 0.0795 0.1101 0.1027 0.0675 0.1593
Phytolacca americana L. SK 0.0420 0.0740 0.1413 0.0937 0.1195
SK, RO 0.0050 0.0095 0.0184 0.0123 0.0157 0.0040 0.0051 0.0047 0.0031 0.0421
Reynoutria bohemica Chrtek et Chrtkov
Reynoutria japonica Houtt. SK 0.0053 0.0101 0.0194
p 0.0130 0.0166
Robinia pseudacacia L. SK 0.0044 0.0018 0.0024 0.0008 0.0007
Rudbeckia laciniata L. SK, RO 0.0044 0.0018 0.0024 0.0008
y 0.0007 0.0013 0.0005 0.0008 0.0010 0.0392
Solidago canadensis L. SK, RO 0.0044 0.0018 0.0024 0.0008 0.0007 0.0011 0.0007 0.0010 0.0011 0.0029
Solidago gigantea Aiton SK, RO 0.0044 0.0018 0.0024 0.0008 0.0007 0.0013 0.0011 0.0013 0.0014 0.0033
Sorghum halepense (L.) Pers. SK, RO 1.2851 261.5322 289.4063 70.3268 67.9237 0.2225 1.2798 0.5430 1.3490 37.1824
Veronica persica Poir. RO 0.2127 1.2654 0.5299 1.3410 37.1687
Xanthium orientale L. subsp. italicum (Moretti) Greuter RO 0.9603 2.3127 1.5043 1.9767 38.3163
Xanthium spinosum L. RO 0.3084 0.3733 0.3473 0.3483 3.1917

Total 10.5196 2092.6786 2316.0460 563.1343 544.0488 4.9894 23.3748 10.7418 24.0450 636.7658
901
902 P. Ferus et al.

(GISD, http://www.issg.org/database). ranthus sp., Ambrosia artemisiifolia, Artemisia annua,


Global trade is dominated by dry cargo (46%), fol- Conyza canadensis, Cuscuta campestris, Erigeron an-
lowed by oil and oil products (31%), coal and min- nuus, Galinsoga sp., Iva xanthifolia, Kochia scoparia,
eral ores (18%) and nally grain (4%) (Hulme 2009). Sorghum halepense and Xanthium spinosum.
We also found dry cargo prevalence in trade exchange Analysis of mud attached on vehicles revealed pe-
between Slovakia and Romania (in 2010 it was 37.7 riodicity in plant seed/fruit load. Zwaenepoel et al.
and 35.1%, respectively) but because of no litera- (2006) refer to January with July and August as the
ture data, analysis of package contamination by inva- richest months on mud and propagules on roads round
sive plant propagules was omitted. Slovak-Romanian Ghent, Belgium. Khan et al. (2013) found the largest
business on commodities with potential directly to number of seeds per vehicle in autumn and the lowest
contain seeds/fruits of invasive plants did in aver- in winter for Queensland, Australia. Most frequently
age 6.6 and 11.8%, respectively, and was based on they adhere to bumpers and do not persist there in wet
wood and cereals exchange. Because of relatively high conditions (Rew 2011; Taylor et al. 2012; Zwaenepoel et
contamination of cereals was allowed, potential ex- al. 2006). However, amounts of seeds/fruits collected by
port/import of invasive plant propagules was strongly trucks on roads are of relatively low importance com-
dependent on the fact whether concrete plant taxon pared to those contaminating some commodities.
occupied I1 habitats and what was the marketed No work considering the role of railyard weed
amount of cereals. Thus, potentially most exported contamination in their long-distance re-distribution is
plant taxa from Slovakia were:Amaranthus sp., Am- available. Implementation of this transport means can
brosia artemisiifolia, Galinsoga sp., Kochia scoparia be seen in risk assessments for invasion of exotic insects

y
and Sorghum halepense; and from Romania: Amaran- (Colunga-Garcia et al. 2009) and diseases associated
thus sp., Ambrosia artemisiifolia, Artemisia annua, with them (Thomas et al. 2014).

p
Conyza canadensis, Cuscuta campestris, Datura stra- Book of Jehlk (2013) represents a large com-
monium, Erigeron annus, Galinsoga sp., Iva xanthi- pendium of exotic ora in European harbours. Al-
co
ifolia, Kochia scoparia, Lycium barbarum, Sorghum though Elodea species are present in many parts of
halepense, Veronica persica and Xanthium orientale Danube river ux, there is no mention in literature on
subsp. italicum. The larger taxon number for Roma- their fragmentation and spread neither by cargo ships
nia can probably be related to dierent socio-economic nor boats. Our analysis of a cargo ship in Bratislava
and natural background. docks brought no light into this problem. However, as
's
In former Czechoslovakia foreign plant species ap- described by Barrat-Segretain et al. (2002), they are
peared especially through so called Pannon (from able to regenerate from low internode segments, as well,
or

Hungary and Romania), Eastern (from Ukraine) and and strong ship generated waves can also provide stem
Elbe routes (Jehlk 1998). Gates for Romanian alien segments (Ali et al. 1999).
ora have ever been Black Sea harbours Sulina and In summary we can conclude that:
th

Constansa (Anastasiu et al. 2009, 2011). As commu- 1. In the period 20062010, total Slovak export to
nicated by the Bratislava cargo harbour representative Romania was approximately three fold higher than in
and Slovak railway info-service, freight amount trans- the opposite direction. Commodities potentially con-
Au

ported from South-East to Slovakia by Danube was in taminated by invasive plant propagules (in average 6.6
2010 only 7.7% of total shipments, and railway import and 11.8% from total export, respectively) showed de-
coming from Ukraine was the largest (56.6%) in this creasing tendency for Slovakia and rising one for Ro-
year (goods crossing the Slovak-Hungarian border rep- mania.
resented only 3.3%). These data are in accordance with 2. Transport of both goods categories was more and
increasing road transport between Slovakia and Roma- more dominated by road compared to railway shipment.
nia as well as worldwide (Hulme 2009). River cargo was found to be of lower importance and
Roads have important ecological eects on vege- applied particularly for freight of not expected direct
tation. Relatively rich roadside plant communities of- infestation by plant seeds/fruits.
ten represent reservoirs of biodiversity in intensively 3. Potentially most exported from Slovakia to
exploited landscape. Regular mowing, higher nitrogen Romania were propagules of: Amaranthus sp., Am-
deposit from exhaust gases, fast water drainage and brosia artemisiifolia, Galinsoga sp., Kochia scoparia
high temperatures as well as salts, herbicides and other and Sorghum halepense tens to hundreds tonnes
chemicals favour resistant aggressive weeds. Woody each.
plants (often foreign taxa) are also planted along roads 4. In opposite direction could mainly be trans-
to reduce erosion, control snow accumulation, support ported seeds/fruits of: Amaranthus sp., Ambrosia arte-
wildlife, reduce headlight glare or enhance aesthetics misiifolia, Artemisia annua, Conyza canadensis, Cus-
(Forman & Alexander 1998). And, long-distance trans- cuta campestris, Datura stramonium, Erigeron annus,
port by vehicles provides further supply of native and Galinsoga sp., Iva xanthiifolia, Kochia scoparia, Ly-
foreign plant propagules (von der Lippe & Kowarik cium barbarum, Sorghum halepense, Veronica persica
2006). and Xanthium orientale subsp. italicum (units to tens
As listed by Jehlk (1998), extreme conditions of tonnes each).
road verges can stand especially species like: Ama- 5. Because of relatively high amount of traded ce-
Reciprocal contamination by invasive plants 903

reals and acceptable impurities content, the highest po- Blackshaw R.E. & Rode L.M. 1991. Eect of ensiling and rumen
tential propagule export in both directions was associ- digestion by cattle on weed seed viability. Weed Sci. 39: 104
108.
ated with cereals. This fact should be taken into ac- Colunga-Garcia M., Haack R.A. & Adelaja A.O. 2009. Freight
count by the control of invasive plant import in the transportation and the potential for invasions of exotic insects
future. in urban and periurban forests of the United States. J. Econ.
6. Our formula for calculation of potential invasive Entom. 102: 237246.
Csontos P., Tams J. & Balogh L. 2007. Thousand-seed weight
plant propagule export can be utilized for any inter- records of species from the ora of Hungary. II. Dicotyledop-
and intra-continental trade exchange. sida. Studia Bot. Hung. 38: 179189.
Davies C.E., Moss D. & OHill M. 2004. EUNIS habitat classica-
tion revised 2004. European Environment Agency, European
Acknowledgements Topic Centre on Nature Protection and Biodiversity.
Davis M.A., Grime P. & Thomson K. 2000. Fluctuating resources
This work was supported by the project of international in plant communities: a general theory of invasibility. J. Ecol.
science-technical cooperation between Slovakia and Ro- 88: 528534.
mania no. SK-RO-0013-10 (in Romania project ANCS- Eli P. 2009. Biotick invzie. VES, SPU Nitra, 192 pp.
Essl F., Dullinger S., Rabitsch W., Hulme P.E., Hubler K., Jarok
UEFISCDI Romania, PN II CAPACITATI). Special thanks
V., Kleinbauer I., Krausmann F., K uhn I., Nentwig W., Vil` a
to Mr. Peter Heidinger, providing us most of the statistical M., Genovesi P., Gherardi F., Desprez-Loustau M.L., Roques
data, presented here, and to Mr. Juraj Bohnsky, speaker A. & Pyek P. 2011. Socioeconomic legacy yields an invasion
of the Slovak Shipping and Ports Co., for very helpful ap- debt. PNAS 108: 203207.
proach and communication of internal data on river freight Forman R.T.T. & Alexander L.E. 1998. Roads and their major
transport, as well as Mr. Marcel Minich from Slovak Rail Co. ecological eects. Annu. Rev. Ecol. Syst. 29: 207231.
info-service for assertive completion of data le on railway Godefroid S., Phartyal S.S. & Koedam N. 2006. Depth distri-

y
cargo transport. We also very appreciate valuable comments bution and composition of seed banks under dierent tree
layers in a managed temperate forest ecosystem. Acta Ecol.
to our work from prof. Phillip E. Hulme from New Zealand
29: 283292.

p
and checking our English by Mrs. Ivana Kocisk. Gojdiov E., Cvachov A. & Karasov E. 2002. Zoznam
nepvodnch, invznych a expanzvnych cievnatch rastln
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