171
Dynamics of the species diversity and 
composition of the ruderal vegetation of 
Slovak and Czech cities
Abstract
This study reports the results of the evaluation of changes in the species diversity 
and composition of ruderal vegetation of three Slovak and one Czech city over 
the time. The dataset of 1489 relevés from five ruderal syntaxa from the cities 
Bratislava, Malacky, Trnava and Brno was used. Data were from two different time 
periods, the older dataset from the years 1960–1982, the more recent dataset from 
the years 2005–2016. The statistical analysis revealed the decrease of Shannon–
Wiener diversity index of all (native + alien) species in the majority of classes of 
ruderal vegetation of cities. The analysis of the changes in the Pielou’s measure of 
species evenness showed that in some of the classes and cities, the species evenness 
remained unchanged, in the other ones it decreased. The percentage of native spe-
cies in the ruderal vegetation did not change over the time, but the percentage of 
invasive alien species in the majority of syntaxa and cities increased significantly. 
In total, 38 invasive taxa were recorded in the ruderal vegetation of cities. Most 
of them are neophytes, therophytes and belong to the family Asteraceae. The 
majority of invasive taxa were recorded in both time periods. Some of the invasive 
species, e.g. Fallopia japonica and Juncus tenuis were recorded only in the more 
recent time period. Our results contribute to the knowledge about biological inva-
sions in the cities.
Izvleček
V članku objavljamo rezultate analize sprememb vrstne pestrosti in sestave rude-
ralne vegetacije treh mest na Slovaškem in enega na Češkem skozi čas. Uporabili 
smo 1489 vegetacijskih popisov petih ruderalnih sintaksonov iz mest Bratislava, 
Malacky, Trnava in Brno. Podatke smo razdelili na dve časovni obdobji, starejši 
podatkovni niz iz obdobja 1960–1982 in novejši niz iz obdobja 2005–2016. S 
statistično analizo smo pokazali, da se je Shannon–Wienerjev diverzitetni indeks 
vseh vrst (domorodnih in tujerodnih) v večini razredov ruderalne vegetacije v 
mestih zmanjšal. Analiza sprememb mere vrstne poravnanosti (Pielou) je pokazala, 
da je vrstna poravnanost v določenih razredih in mestih ostala nespremenjena, v 
določenih pa se je zmanjšala. Odstotek domorodnih vrst v ruderalni vegetaciji se 
ni spreminjal s časom, značilno pa se je tako v sintaksonih kot v mestih povečal 
odstotek invazivnih tujerodnih vrst. Skupaj smo v ruderalni mestni vegetaciji 
zabeležili 38 invazivnih taksonov. Večinoma so neofiti, terofiti in jih uvrščamo v 
družino Asteraceae. Večino invazivnih taksonov smo zabeležili v obeh časovnih 
obdobjih, nekatere invazivne vrste, kot sta na primer Fallopia japonica in Juncus 
tenuis, pa smo smo opazili le v novejšem obdobju. Rezultati naše raziskave prispe-
vajo k poznavanju bioloških invazij v mestih.
Keywords: alien invasive taxa, 
neophyte, Shannon–Wiener 
diversity index, species evenness, 
synanthropic plant communities.
Ključne besede: tujerodne 
invazivne vrste, neofiti, Shannon–
Wiener diverzitetni indeks, vrstna 
poravnanost, sinantropne rastlinske 
združbe.
Received: 1. 7. 2017
Revision received: 11. 1. 2018
Accepted: 14. 1. 2018
1 Department of Botany, Faculty of Natural Sciences, Comenius University in Bratislava, Révová 39, 811 02 Bratislava, Slovakia. E-mail: alenarendekova@gmail.com
* Corresponding author
Alena Rendeková1,*, Karol Mičieta1, Zuzana Randáková1 & Ján Miškovic1 
DOI: 10.1515/hacq-2017-001417/2 • 2018, 171–188
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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Introduction
Urban areas represent strongly anthropogenically altered 
environments. The unique character of urban areas is 
reflected by the well adapted species and communities 
(Sukopp 2002, Forman 2014). There exist considerable 
habitat heterogeneity, and lot of possibilities for species 
immigration in urban areas (Sukopp & Werner 1983). 
All these features cause the general high plant species di-
versity of cities, but on smaller scales the diversity and 
the number of species can vary considerably (Sukopp 
& Werner 1983, Pyšek 1989, 1993, Kühn et al. 2004, 
Schmidt et al. 2014).
The large part of the species composition of the flora 
and vegetation of cities is formed by alien species (Pyšek 
1998, Chytrý et al. 2005, Vilá et al. 2007, Simonová & 
Lososová 2008, Medvecká et al. 2009, 2014). Therefore, 
it is very useful to focus on the research of the alien spe-
cies in the cities. It is important to differentiate among 
various ruderal habitats, since the urban environment 
creates a wide range of different habitat types and there 
is also a large variability of ruderal plant communities 
that colonize them (Jarolímek et al. 1997) and they differ 
in the proportion of alien species (Simonová & Lososová 
2008, Medvecká et al. 2009). 
It is also important to take into the consideration the 
fact, that not all of the alien species represent the same 
threats. Only a small fraction of introduced species can 
survive in new areas for a long time, start to spread rapid-
ly and become invasive (Mack et al. 2000, Pyšek & Tichý 
2001). Hence, it is important to differentiate which alien 
species are casual, naturalised or invasive (Richardson et 
al. 2000), because only the invasive species present an 
actual problem. 
Many researchers focused their studies on the whole 
group of alien plants in urban areas (e.g. Simonová & 
Lososová 2008, Medvecká et al. 2009, Eliáš jun. 2009, 
2011, Jehlík et al. 2013, Király et al. 2014, Aronson et al. 
2015, Ferus et al. 2015, Zisenis 2015, Žabka et al. 2015), 
or various other habitats (Diekmann et al. 2016), but in-
vasive species in ruderal habitats (Tokaryuk et al. 2012) 
or towns (Bomanowska et al. 2017, Štajerová et al. 2017) 
have been rarely the object of special studies. Our study 
is deeper focused on the invasive group of the alien spe-
cies and their representation in cities. 
In the last decades, habitat alteration and introduc-
tions of alien plants have resulted in changes in the Eu-
ropean urban flora and vegetation and typical of these 
trends are an increasing proportion of neophytes (Pyšek 
& Mandák 1997, Chocholoušková & Pyšek 2003, 
Lososová & Simonová 2008, Gregor et al. 2012) and 
the decreasing proportion of native species accordingly 
(Chocholoušková & Pyšek 2003, Lososová & Simonová 
2008) over the time. Human activities that make ruderal 
habitats of studied cities prone to invasions by neophytes 
(e.g. transport of goods and construction activities) and 
the increasing anthropogenic influence in the studied 
cities during the last decades could be reflected by chang-
es in the diversity and species composition, e.g. by an 
increasing proportion of invasive species in the studied 
cities. Hence, it is crucial to pay attention to dynamics 
of the diversity and species composition over the time.
Local diversity (or biodiversity) can be well-expressed 
by the various indicators, among which the Shannon–
Wiener diversity index [H’] is one of the most commonly 
used metric in ecological studies for measuring species 
diversity (Hill 1973, Kent & Coker 1992, Magurran 
1988, 2004). Diversity is sometimes measured also with 
a species evenness index (Stirling & Wilsey 2001). 
The impact of some invasive species on the Shannon–
Wiener diversity [H'] and evenness [J] in various invad-
ed plant communities was studied e.g. in Czech Republic 
(Hejda & Pyšek 2006, Hejda et al. 2009), changes in 
plant species diversity (measured by Shannon index) of 
aquatic ecosystems in the agricultural landscape in West 
Poland in the last 30 years analysed by Gołdyn (2010). 
The forest structure and woody vegetation diversity 
(measured by Shannon index and evenness index) of ri-
parian communities in response to an urbanization gra-
dient was studied in West Georgia, USA (Burton et al. 
2005), the measurement of the effects of invasive species 
Bothriochloa ischaemum on Shannon–Wiener diversity 
diversity was done in central Texas, USA (Gabbard & 
Fowler 2007), effects of an exotic invasive macrophyte 
Urochloa subquadripara on native plant community spe-
cies richness and Shannon diversity in Neotropical reser-
voirs and lakes were evaluated by Michelan et al. (2010), 
but there is no study dealing with invasive species and 
diversity dynamics of ruderal vegetation from Slovakia.
Understanding of behaviour of invasive species in cit-
ies is of crucial importance, because cities can represent 
the sources from which invasive species spread into na-
tive vegetation (Sukopp & Werner 1983). The knowledge 
about the invasive species dynamics can help to mini-
malize their invasions in the future. Therefore, our study 
is focused on the changes in the diversity and species 
composition, especially changes invasive species propor-
tion in cities.
The aims of our study are: (1) to analyze the changes 
in the diversity of all (native + alien) taxa in the various 
syntaxa of ruderal vegetation of cities over the time; (2) 
to analyze the changes in the evenness of all (native + 
alien) taxa; (3) to analyze the changes in the average ratio 
of native taxa; (4) to analyze the changes in the average 
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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percentage of invasive alien taxa (not of all alien taxa, 
because the study is focused only to the invasive group 
of alien taxa). 
Methods 
Study area and relevé data 
In the study, we used 1489 relevés of the ruderal plant com-
munities of the phytosociological classes Polygono-Poetea 
annuae, Sisymbrietea, Digitario sanguinalis-Eragrostietea 
minoris, Artemisietea vulgaris, and Epilobietea angustifolii 
from the cities Bratislava, Malacky, Trnava and Brno. The 
characteristics of studied cities are shown in the Table 1.
The class Polygono-Poetea annuae is comprised of sub-
cosmopolitan therophyte-rich dwarf-herb ruderal veg-
etation, which grows in trampled habitats, e.g. the edge 
of pavements, playgrounds, or parking lots. The class 
Sisymbrietea includes zoo-anthropogenic and modern 
anthropogenic vegetation of disturbed sites, growing in 
cool- and cold-temperate regions of Eurasia. The class 
Digitario sanguinalis-Eragrostietea minoris is comprised of 
thermophilous anthropogenic vegetation rich in grasses 
and summer-annual C4 species, growing in the southern 
nemoral, mediterranean, steppe and semi-desert parts of 
Europe. The class Artemisietea vulgaris includes peren-
nial subxerophilous ruderal vegetation of the temperate 
and submediterranean zones of Europe. The class Epilo-
bietea angustifolii is comprised of tall-herb semi-natural 
and anthropogenic perennial vegetation, which grow on 
wet ruderal areas with high nitrogen content in the soil, 
disturbed forest edges, nutrient-rich riparian fringes and 
in forest clearings in the temperate and boreal zones of 
Eurasia. Described syntaxonomical scheme we used was 
recently proposed by Mucina et al. (2016).
We used data from two different time periods. The old 
dataset consisted of 954 relevés from the years 1960–
1982 and was compiled from relevés published by Eliáš 
(1977, 1978, 1979), Jarolímek (1983) and relevés from 
the Slovak and Czech National Phytosociological Data-
bases (Chytrý & Rafajová 2003, Hegedüšová 2007, Šibík 
2012) which match study area, the time period and the 
selected syntaxa. We selected relevés by the original clas-
sification of authors.
The more recent dataset consisted of 535 relevés from 
the years 2005–2016, out of which 465 were made by 
authors of the study (Rendeková et al. 2014, Rendeková 
2016a, b, c, plus the unpublished relevés made by 
Rendeková) and 70 relevés which match study area, the 
time period and the selected syntaxa were obtained from 
the Slovak and Czech National Phytosociological Da-
tabases (Chytrý & Rafajová 2003, Hegedüšová 2007, 
Šibík 2012).
The phytosociological research in both periods was 
performed according to the methods of the Zürich-
Montpellier school (Braun-Blanquet 1964). In the old 
time period, the old Braun-Blanquet cover-abundance 
scale was used. In the more recent period, the modified 
Braun-Blanquet cover-abundance scale, extended by 2m, 
2a and 2b values (Barkman et al. 1964) was used. The 
area of the data collection in both periods concerns the 
same districts of the studied cities. 
Data analysis
All relevés were imported into the TURBOVEG data-
base (Hennekens & Schaminée 2001) and edited in the 
JUICE 7.0 programme (Tichý 2002). The relevés made 
by the authors of the study were classified into the ruderal 
syntaxa using hierarchical clustering in the programme 
city Bratislava Malacky Trnava Brno
location southwestern  
Slovakia 
southwestern  
Slovakia 
western  
Slovakia 
south-eastern part of the  
Czech Republic 
area 368 km2 25 km2 71,54 km2 230 km2 
approximate population (2016) 426,000 17,300 65,500 378,000
climate moderate to warm, 
continental 
warm, lightly humid 
with mild winters 
warm humid continental and border-
line oceanic with cold winters 
and hot to warm summers 
number of analysed relevés 695 188 209 397
total number of taxa in whole dataset 672 291 313 403
number of invasive taxa in dataset 26 15 17 26
The table was compiled according to the works of Trnka (1998), Kollár & Kollár (2004), Peel et al. (2007), Hrnčiarová et al. (2009), 
Macejka & Marek (2009), Feráková & Jarolímek (2011), Macejka (2015).
Table 1: Characteristic of the studied cities
Tabela 1: Značilnosti preučevanih mest.
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SYN-TAX 2000 (Podani 2001). Various linkage meth-
ods and distance measures were tried. A beta-flexible 
method (β = −0.25) in combination with Wishart’s 
index, a beta-flexible method (β = −0.25) in combina-
tion with Ružička’s coefficient, and the Group Average 
method in combination with Wishart’s index proved to 
be the most effective parameters and were used in most 
of the hierarchical clustering analyses. The assignment of 
the other relevés to syntaxa followed the original assign-
ment made by authors of relevés (Eliáš 1977, 1978, 1979, 
Jarolímek 1983, Chytrý & Rafajová 2003, Hegedüšová 
2007, Šibík 2012). 
Bryophytes and the taxa of vascular plants determined 
only to the genus level were excluded from the analysis 
and the taxa which occurred in more than one layer were 
merged. As different cover-abundance scales have been 
used in the old and the more recent relevés, to make the 
data comparable, the values 2m, 2a, and 2b in both data-
sets were converted to value 2.
The diversity was expressed by the Shannon–Wiener 
diversity index [H'], because this index is one of the most 
commonly used metric in ecological studies for meas-
uring species diversity (Hill 1973, Kent & Coker 1992, 
Magurran 1988, 2004). We also calculated the species 
evenness, as the diversity can be measured also with the 
evenness index (Stirling & Wilsey 2001). 
The Shannon–Wiener index of diversity [H'] (e.g. 
Magurran 1988) and Pielou’s measure of species even-
ness [J=H’/ln(S)], where H’ is the Shannon diversity in-
dex and S is the number of taxa in every relevé] of all (na-
tive + alien) taxa were calculated for every relevé, using 
the JUICE 7.0 programme. 
Then, the taxa in every relevé were divided into native 
and alien. Alien taxa were divided into casual, natural-
ized, and invasive. This division was made according to 
the ‘List of alien vascular plant species of Slovak Repub-
lic’ (Medvecká et al. 2012) for the relevés from Slovak 
cities, and according to the ‘Catalogue of alien plants of 
the Czech Republic’ (Pyšek et al. 2012) for the relevés 
from Brno. Afterwards, the average percentage number 
of native taxa and of invasive alien taxa in each relevé 
was calculated using the JUICE 7.0 programme. We did 
not calculate average percentual number of all alien taxa, 
because the study is focused only to the invasive group 
of alien taxa.
Afterwards, the changes in the average values the Shan-
non–Wiener index of diversity, Pielou’s measure of spe-
cies evenness, percentual number of native taxa, and the 
percentual number of invasive alien taxa between the old 
and the more recent datasets of each class of each city were 
compared using the Main Effects ANOVA analysis in the 
STATISTICA 7.0 programme (Hill & Lewicki 2007).
As explanatory feature, the changes in total proportion 
of recorded invasive alien taxa in the categories of fami-
lies, Raunkiær’s life forms, origin, and residence time be-
tween the old and the more recent period was calculated 
for all relevés of ruderal vegetation (all phytosociological 
classes together). This calculation was made separately 
for the group of Slovak cities (Bratislava, Malacky, Tra-
nava together) and separately for Brno (Czech city). The 
families, Raunkiær’s life forms, origin, and residence 
time of invasive taxa were determined according to the 
‘List of alien vascular plant species of the Slovak Repub-
lic’ (Medvecká et al. 2012) and ‘Catalogue of alien plants 
of the Czech Republic’ (Pyšek et al. 2012). 
As explanatory variable to show if some ecological 
characteristics of the vegetation types have been changed 
the mean Ellenberg indicator values for light, tempera-
ture, continentality, moisture, soil reaction, and nutrients 
 (Ellenberg et al. 1991) based on species presence, were 
calculated, using the JUICE 7.0 programme and the 
changes in the mean Ellenberg indicator values between 
old and recent data were compared using t-test for in-
dependent samples in the STATISTICA 7.0 programme. 
Nomenclature
The nomenclature of the taxa follows Marhold (1998), the 
nomenclature of the syntaxa follows Mucina et al. (2016).
Results
In the majority of the cities, the class Artemisietea vulgaris 
was the class with the highest average values of Shannon–
Wiener diversity index of all (native + alien) taxa and av-
erage values of the Pielou’s measure of species evenness. 
The classes Polygono-Poetea annuae and Digitario sangui-
nalis-Eragrostietea minoris were the classes with the low-
est average values of diversity index and species evenness 
(Figures 1–4).
The highest average percentual number of native spe-
cies was recorded in the class Polygono-Poetea annuae, 
relatively high also in the classes Artemisietea vulgaris and 
Epilobietea angustifolii. The lowest percentual number of 
native species was recorded in the class Sisymbrietea, and 
relatively low also in the class Digitario sanguinalis-Era-
grostietea minoris (Figures 5–6).
The average values of the percentual number of invasive 
species were very high in the class Digitario sanguinalis-
Eragrostietea minoris, especially in the more recent time 
period (Figure 7). The percentual number of invasive 
taxa in this class in the years 2005–2016 in Trnava was 
as high as 28.6%, in Brno 28.9%. In Bratislava, on the 
other hand, the percentual number of invasive taxa in 
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the class Digitario sanguinalis-Eragrostietea minoris was 
the highest (33.7%) in the old time period. Such high 
values were not recorded in any other analysed class. 
Moreover, in the majority of other cities the values in the 
class Digitario sanguinalis-Eragrostietea minoris were also 
relatively high – approximately 15%. The second most 
invaded class was the class Sisymbrietea, where the aver-
age percentual number of invasive species in the major-
ity of cities was more than 10% and only in one city 
(Malacky) it was less than 10% (Figure 7). The classes 
Artemisietea vulgaris and Epilobietea angustifolii were less 
invaded (Figure 8).
In total, 38 invasive taxa were recorded in the ruderal 
vegetation of the studied Slovak and Czech cities (Ta-
ble 2). The majority of them are considered to be invasive 
in both countries. There were 13 taxa, which are invasive 
only in the Czech Republic and 6 taxa, which are invasive 
only in Slovakia (Table 2). The majority of invasive taxa 
were recorded in both time periods. Some of the inva-
sive species, e.g. Fallopia japonica and Juncus tenuis were 
recorded only in the more recent time period (Table 2). 
The majority of the recorded invasive taxa in all cities in 
both periods were neophytes, therophytes and belonged 
to the family Asteraceae (Figures 9–10). Most of the in-
Figure 1: The comparison of the Shannon–Wiener index of diversity 
[H'] in the old and the more recent relevés of the classes Polygono-Poet-
ea annuae, Sisymbrietea, and Digitario sanguinalis-Eragrostietea minoris 
in various cities
Explanation: 
1 – Bratislava, old time period (years 1960–1982); 
2 – Bratislava, more recent time period (years 2005–2016); 
3 – Malacky, old time period (years 1960–1982); 
4 – Malacky, more recent time period (years 2005–2016);
5 – Trnava, old time period (years 1960–1982);
6 – Trnava, more recent time period (years 2005–2016);
7 – Brno, old time period (years 1960–1982);
8 – Brno, more recent time period (years 2005–2016).
Slika 1: Primerjava Shannon–Wienerjevega diverzitetnega indeksa [H'] 
med starejšimi in novejšimi vegetacijskimi popisi razredov Polygono-Po-
etea annuae, Sisymbrietea, in Digitario sanguinalis-Eragrostietea minoris 
v različnih mestih. 
Legenda: 
1 – Bratislava, starejše obdobje (leta 1960–1982); 
2 – Bratislava, novejše obdobje (leta 2005–2016); 
3 – Malacky, starejše obdobje (leta 1960–1982); 
4 – Malacky, novejše obdobje (leta 2005–2016);
5 – Trnava, starejše obdobje (leta 1960–1982);
6 – Trnava, novejše obdobje (leta 2005–2016);
7 – Brno, starejše obdobje (leta 1960–1982);
8 – Brno, novejše obdobje (leta 2005–2016)
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Occurrence in the city and time period
Taxon Family Life  
form
Origin R. t. BA M T Slovak cities  
in total
Brno (Czech  
city in total)
Ailanthus altissima Sim Ph As neo b b b b r
Amaranthus powellii Ama T CAm SAm arch * * * * r
Amaranthus retroflexus Ama T CAm SAm neo b b b b b
Ambrosia artemisiifolia Ast T NAm neo b r r b  
Apera spica-venti Poa T As E arch b b b b ~
Arrhenatherum elatius Poa He E arch * * * * b
Aster novi-belgii agg. Ast He NAm neo b r b  
Atriplex sagittata Ama T As E arch * * * * b
Atriplex tatarica Ama T Af As E arch b o b b ~
Bassia scoparia subsp. scoparia Ama T As E neo * * * * o
Bidens frondosa Ast T NAm neo b b  
Cardaria draba Bra He Af As E arch b r+ b b ~
Cirsium arvense Ast He As E arch * * * * b
Conium maculatum Api T He Af As E arch * * * * o
Conyza canadensis Ast T NAm neo b b b b b
Digitaria ischaemum Poa T E arch * * * * r
Echinochloa crus-galli Poa T As E arch b b b b b
Echinocystis lobata Cuc T NAm neo r r  
Echinops sphaerocephalus subsp. sphaerocephalus Ast He E neo * * * * o
Eragrostis minor Poa T E arch * * * * b
Fallopia japonica Pog G As neo r r  
Galinsoga parviflora Ast T SAm neo b b b b b
Galinsoga urticifolia Ast T CAm SAm neo b b b
Helianthus tuberosus Ast He NAm neo r b r o
Impatiens glandulifera Bal T As neo b b  
Impatiens parviflora Bal T As neo b b b
Juncus tenuis Jun He NAm neo r r ~
Lycium barbarum Sol Ph As neo b b o
Matricaria discoidea Ast T As NAm neo b b b b ~
Negundo aceroides Sap Ph NAm neo b r r b r
Portulaca oleracea subsp. oleracea Por T As E arch * * * * b
Robinia pseudoacacia Fab Ph NAm neo b o b b  
Rumex patientia Pog He E neo b r b ~
Sisymbrium loeselii Bra T Af As E neo * * * * b
Solidago canadensis Ast He NAm neo r r r b
Solidago gigantea Ast He NAm neo b r r b o
Stellaria pallida Car T Af As E arch * * * * b
Stenactis annua Ast T NAm neo b r r b r
Xanthoxalis dillenii Oxa He T NAm neo * * * * r
Explanation: R. t. – residence time; BA – Bratislava city, M – Malacky city, T – Trnava city; 
Ama – Amaranthaceae, Api – Apiaceae, Ast – Asteraceae, Bal – Balsaminaceae, Bra – Brassicaceae, Car – Caryophyllaceae, 
Cuc – Cucurbitaceae, Fab – Fabaceae, Jun – Juncaceae, Oxa – Oxalidaceae, Poa – Poaceae, Pog – Polygonaceae, Por – Portulacaceae, 
Sap – Sapindaceae, Sim – Simaroubaceae, Sol – Solanaceae; T – therophyte, G – geophyte, He – hemicryptophyte, Ph – phanero-
phyte; Af – Africa, As – Asia, E – Europe, CAm – Central America, NAm – North America, SAm – South America; arch – archaeo-
phyte, neo – neophyte; b – taxon was present in the ruderal vegetation of both time periods, o – taxon was present in the ruderal 
vegetation only in the old time period (in the years 1960–1982), r – taxon was present in the ruderal vegetation only in the more 
recent time period (in the years 2005–2016); * – taxon is not considered to be invasive in Slovakia, ~ – taxon is not considered to be 
invasive in the Czech Republic, + – taxon was recorded in the more recent time period in the ruderal flora in Malacky city but not in 
the ruderal community that could be assigned to any phytosociological class of ruderal vegetation by numerical analysis
Table 2: List of recorded invasive taxa and their occurrence in the time periods.
Tabela 2: Seznam invazivnih taksonov in njihovo pojavljanje v obeh časovnih obdobjih.
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vasive taxa recorded in the ruderal vegetation of Slovak 
cities are native to North America and relatively many 
are native to Asia (Figure 9). The majority of invasive taxa 
recorded in Brno come from Europe and many of them 
from Asia (Figure 10).
The statistically significant results of the analysis of the 
changes in the Shannon–Wiener diversity index of all 
(native + alien) taxa over the time showed the decrease 
of the average values of diversity index in the majority of 
classes in almost all of the cities (Figures 1–2). 
The most significant decrease of Shannon–Wiener di-
versity index over time was recorded in the class Artemisi-
etea vulgaris in Trnava [F(1, 55) = 25.7, p < 0.001; F – F 
ratio; p – p value; numbers in brackets behind the F ratio 
represents the values for degrees of freedom], where it has 
decreased from 2.2 to 1.4 and in Brno [F(1, 86) = 17.5, 
p < 0.001], where it has decreased from 2.3 to 1.6 and in 
the class Epilobietea angustifolii in Brno [F(1, 26) = 18.9, 
p < 0.001], where it has fallen from 2.3 to 1.4 (Figure 2).
The average values of the Shannon–Wiener diversity 
index decreased also in the class Artemisietea vulgaris in 
Bratislava [F(1, 277) = 36.6, p < 0.001] and Malacky 
[F(1, 32) = 14.9, p < 0.001] and in the class Epilobietea 
angustifolii in Bratislava [F(1, 163) = 4.4, p = 0.0380] 
and Malacky [F(1, 22) = 4.9, p = 0.038] (Figure 2). The 
decreasing trend was recorded also in the class Polygono-
Poetea annuae in Trnava [F(1, 22) = 5.1, p = 0.034] and 
Brno [F(1, 64) = 10.2, p = 0.002] and in the class Sisym-
brietea in Trnava [F(1, 94) = 11.6, p < 0.001] and Brno 
[F(1, 189) = 14.8, p < 0.001] (Figure 1). 
The changes of the average values of the Shannon–Wie-
ner diversity index in all of the other classes of the ruderal 
vegetation of the analysed Slovak and Czech cities proved 
to be statistically not significant (p > 0.05). 
The analysis of the changes in the Pielou’s measure of 
evenness of all (native + alien) species in the classes of rud-
eral vegetation over the time showed the that in almost 
a half of the classes and cities, the difference in the average 
values of evenness between the old and the more recent 
time period was not statistically significant (p > 0.05), 
the average values of the evenness remain unchanged 
(Figures 3–4). In Brno, for example, there have been no 
significant difference in any of the classes (p > 0.05). Sta-
tistically significant difference have been obtained only for 
some classes and cities, where the decrease of the aver-
age values of evenness was recorded. The most significant 
decrease in the average values of the Pielou’s measure of 
species evenness was recorded in the class Artemisietea vul-
garis in Malacky [F(1, 32) = 48.9, p < 0.001], where it 
has fallen from 0.8 to 0.55 (Figure 4). The average values 
of the  evenness decreased in the statistically significant 
way in the same class also in Bratislava [F(1, 277) = 67.5, 
Figure 2: The comparison of the Shan-
non–Wiener index of diversity [H’] in 
the old and the more recent relevés of the 
classes Artemisietea vulgaris and Epilobi-
etea angustifolii in various cities. Explana-
tion corresponds to those in Figure 1.
Slika 2: Primerjava Shannon–Wiener-
jevega diverzitetnega indeksa [H’] med 
starejšimi in novejšimi vegetacijskimi 
popisi razredov Artemisietea vulgaris in 
Epilobietea angustifolii v različnih mestih. 
Legenda je enaka kot pri Sliki 1.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
Slovak and Czech cities
178
17/2 • 2018, 171–188
↑
Figure 3: The comparison of the Pielou’s 
measure of evenness [J] in the old and 
the more recent relevés of the classes 
Polygono-Poetea annuae, Sisymbrietea, and 
Digitario sanguinalis-Eragrostietea minoris 
in various cities. Explanation corresponds 
to those in Figure 1.
Slika 3: Primerjava vrstne poravna-
nosti po Pielou [J] med starejšimi in 
novejšimi vegetacijskimi popisi razredov 
Polygono-Poetea annuae, Sisymbrietea in 
Digitario sanguinalis-Eragrostietea minoris 
v različnih mestih. Legenda je enaka kot 
pri Sliki 1.
→
Figure 4: The comparison of the Pielou’s 
measure of evenness [J] in the old and 
the more recent relevés of the classes 
Artemisietea vulgaris and Epilobietea 
angustifolii in various cities. Explanation 
corresponds to those in Figure 1
Slika 4: Primerjava vrstne poravnanosti 
po Pielou [J] med starejšimi in novejšimi 
vegetacijskimi popisi razredov Artemisi-
etea vulgaris in Epilobietea angustifolii v 
različnih mestih. Legenda je enaka kot 
pri Sliki 1.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
Slovak and Czech cities
179
17/2 • 2018, 171–188
↑
Figure 5: The comparison of the average 
percentual number of native taxa in the 
old and the more recent relevés of the 
classes Polygono-Poetea annuae, Sisymbri-
etea, and Digitario sanguinalis-Eragrosti-
etea minoris in various cities. Explanation 
corresponds to those in Figure 1
Slika 5: Primerjava povprečnega deleža 
domorodnih vrst med starejšimi in 
novejšimi vegetacijskimi popisi razredov 
Polygono-Poetea annuae, Sisymbrietea in 
Digitario sanguinalis-Eragrostietea minoris 
v različnih mestih. Legenda je enaka kot 
pri Sliki 1.
←
Figure 6: The comparison of the average 
percentual number of native taxa in the 
old and the more recent relevés of the 
classes Artemisietea vulgaris and Epilobi-
etea angustifolii in various cities. Explana-
tion corresponds to those in Figure 1
Slika 6: Primerjava povprečnega deleža 
domorodnih vrst med starejšimi in 
novejšimi vegetacijskimi popisi razredov 
Artemisietea vulgaris in Epilobietea 
angustifolii v različnih mestih. Legenda je 
enaka kot pri Sliki 1.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
Slovak and Czech cities
180
17/2 • 2018, 171–188
p < 0.001] and Trnava [F(1, 55) = 22.7, p < 0.001] and 
in the class Epilobietea angustifolii in Bratislava [F(1, 163) 
= 7.6, p = 0.006] and Malacky [F(1, 22) = 5.0, p = 0.353] 
(Figure 4). The significant decrease of the evenness was 
recorded also in the class Polygono-Poetea annuae in Trnava 
[F(1, 22) = 9.9, p = 0.005] and in the class Sisymbrietea in 
Malacky [F(1, 64) = 11.2, p = 0.001] and Trnava [F(1, 94) 
= 8.2, p = 0.005] (Figure 3).
The analysis of the changes in the average percen-
tual number of native species in the ruderal vegetation 
over the time showed that in the majority of the class-
es and cities there have been no statistically significant 
change between the old and the more recent time pe-
riod (p > 0.05) (Figures 5–6). In the class Digitario san-
guinalis-Eragrostietea minoris in Trnava, the significant 
decrease of the percentual number of native species was 
recorded [F(1, 11) = 9.4, p = 0.011] (Figure 5). The sig-
nificant increase of average percentual number of native 
species was recorded in the class Sisymbrietea in Bratis-
lava [F(1, 164) = 50.4, p < 0.001] and Malacky [F(1, 64) 
= 5.7, p = 0.020] (Figure 5).
The statistically significant results of the analysis of 
the changes in the average percetual number of invasive 
species in the classes of the ruderal vegetation over the 
time showed the increase of the percentual number of 
invasive species in the majority of the classes in almost 
all of the studied Slovak and Czech cities (Figures 7–8).
The most significant increase of the percentual number 
of invasive taxa was recorded in the class Digitario san-
guinalis-Eragrostietea minoris in Trnava [F(1, 11) = 8.0, 
p = 0.017], where it has risen from 10.3% to 28.6% (Fig-
ure 7). A significant increase was also recorded in the 
same class in Brno [F(1, 22) = 17.4, p < 0.001] and in the 
class Sisymbrietea in Trnava [F(1, 94) = 17.4, p < 0.001] 
(Figure 7). The average percentual number of invasive 
species increased also in the class Artemisietea vulgaris 
in all of the studied cities {Bratislava [F(1, 277) = 34.1, 
p < 0.001], Malacky [F(1, 32) = 9.2, p = 0.0048], Trna-
va [F(1, 55) = 17.1, p < 0.001], Brno [F(1, 86) = 9.7, 
p = 0.0025]}and in the class Epilobietea angustifolii in 
Bratislava [F(1, 163) = 22.7, p < 0.001] (Figure 8).
Figure 7: The comparison of the average percentual number of invasive taxa in the old and the more recent relevés of the classes Polygono-Poetea 
annuae, Sisymbrietea, and Digitario sanguinalis-Eragrostietea minoris in various cities. Explanation corresponds to those in Figure 1
Slika 7: Primerjava povprečnega deleža invazivnih vrst med starejšimi in novejšimi vegetacijskimi popisi razredov Polygono-Poetea annuae, Sisymbri-
etea in Digitario sanguinalis-Eragrostietea minoris v različnih mestih. Legenda je enaka kot pri Sliki 1.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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181
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The statistically significant decrease of the percetual 
number of invasive species was recorded only in the in the 
class Polygono-Poetea annuae in Trnava [F(1, 22) = 4.7, p 
= 0.0413] (Figure 7), in the class Digitario sanguinalis-
Eragrostietea minoris in Bratislava [F(1, 38) = 35.8, p < 
0.001] (Figure 7), and in the class Epilobietea angustifolii 
in Brno [F(1, 26) = 5.8, p = 0.0239] (Figure 8).
The changes of the average percentual number of inva-
sive species in all of the other classes of the ruderal vegeta-
tion of the studied Slovak and Czech cities proved to be 
statistically not significant (p > 0.05). 
The analysis of the changes in mean Ellenberg indicator 
values over the time by the t-test for independent samples 
revealed a statistically significant increase in mean Ellen-
berg indicator values for temperature between the old and 
the more recent period (p < 0.05) (Figure 11b). There 
was a slight increase in mean values for light (Figure 11a) 
and nutrients (Figure 11f ) and a slight decrease of mean 
values for moisture (Figure 11d) and soil reaction (Figure 
11a), but these little changes proved to be statistically not 
significant (p > 0.05).
Discussion
The lowest percentual number of native species was re-
corded in the class Sisymbrietea, and relatively low also 
in the class Digitario sanguinalis-Eragrostietea minoris 
(Figures 5–6). On the other hand, these classes were the 
classes with the highest average values of the percentual 
number of invasive species (Figures 7–8). The highest 
percentual number of invasive species was recorded in 
the class Digitario sanguinalis-Eragrostietea minoris, the 
class Sisymbrietea was the second one most invaded. The 
proportion of invasive species in the other three classes, 
especially in the classes Artemisietea vulgaris and Epilo-
bietea angustifolii was lower (Figures 7–8). On the other 
hand, these classes have the higher proportion of native 
species (Figures 5–6). Probably, the alien invasive species 
in the vegetation of classes Digitario sanguinalis-Eragrosti-
etea minoris and Sisymbrietea replaced native species. The 
interesting fact is that in the class Digitario sanguinalis-
Eragrostietea minoris in Trnava, the significant decrease of 
the percentual number of native species over the time was 
recorded (Figure 5) and at the same time, in this class 
and city, the percentual number of invasive taxa increased 
significantly (Figure 7). The increase of invasive alien spe-
cies can be associated with the retreat of native species. 
The increase of invasive species could also play a role in a 
overall decrease of species diversity, because many of the 
plant communities, which were recorded in the more re-
cent time period were formed monodominantly by inva-
sive species and thus had a very low diversity.
Our results agree with the findings of some other au-
thors (McKinney & Lockwood 1999, Olden et al. 2004), 
which indicate the process of homogenization of the biota 
at larger scale. According to these authors, many species 
are declining and are being replaced by a much smaller 
number of expanding species, mainly aliens, and this 
whole process results in a more uniform biosphere with 
lower diversity. 
The similar trend has been recorded in urban floras and 
vegetation. The decrease in species diversity over the time 
Figure 8: The comparison of the average 
percentual number of invasive taxa in the 
old and the more recent relevés of the 
classes Artemisietea vulgaris and Epilobi-
etea angustifolii in various cities. Explana-
tion corresponds to those in Figure 1
Slika 8: Primerjava povprečnega deleža 
invazivnih vrst med starejšimi in 
novejšimi vegetacijskimi popisi razredov 
Artemisietea vulgaris in Epilobietea 
angustifolii v različnih mestih. Legenda je 
enaka kot pri Sliki 1.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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182
17/2 • 2018, 171–188
was recorded in the ruderal vegetation of some European 
cities (Pyšek et al. 2004). Similarly, the results of the anal-
ysis of dynamics of ruderal species diversity in the Harbin 
city in China showed that the number of ruderal species 
decreased in the past half century (Chen et al. 2014). The 
majority of recorded invasive taxa in Slovakia and Czech 
Republic are neophytes, the minority of them are archae-
ophytes (Figures 9–10). In many European urban habi-
Figure 9: The total proportion of the recorded invasive taxa in the ruderal vegetation (all syntaxa together) of Slovak cities in categories of families 
(a), Raunkiær’s life forms (b), countries of orgin (c), residence time status (d).
Explanation: Ama – Amaranthaceae, Ast – Asteraceae, Bal – Balsaminaceae, Bra – Brassicaceae, Cuc – Cucurbitaceae, Fab – Fabaceae, 
Jun – Juncaceae, Poa – Poaceae, Pog – Polygonaceae, Sap – Sapindaceae, Sim – Simaroubaceae, Sol – Solanaceae; T – therophyte, G – geophyte, 
He – hemicryptophyte, Ph – phanerophyte; Af – Africa, As – Asia, E – Europe, CAm – Central America, NAm – North America, SAm – South 
America; arch – archaeophyte, neo – neophyte.
Slika 9: Delež invazivnih taksonov v ruderalni vegetaciji (vsi sintaksoni skupaj) v slovaških mestih in uvrstitev v družine (a), Raunkiærjeve 
življenske oblike (b), izvorna celina (c), čas vnosa (d).
Legenda: Ama – Amaranthaceae, Ast – Asteraceae, Bal – Balsaminaceae, Bra – Brassicaceae, Cuc – Cucurbitaceae, Fab – Fabaceae, Jun – Juncaceae, 
Poa – Poaceae, Pog – Polygonaceae, Sap – Sapindaceae, Sim – Simaroubaceae, Sol – Solanaceae; T – therophyte, G – geophyte, He – hemicrypto-
phyte, Ph – phanerophyte; Af – Africa, As – Asia, E – Europe, CAm – Central America, NAm – North America, SAm – South America; 
arch – arheofiti, neo – neofiti.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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Figure 10: The total proportion of the recorded invasive taxa in the ruderal vegetation (all syntaxa together) of Brno (Czech city) in categories of 
families (a), Raunkiær’s life forms (b), countries of orgin (c), residence time status (d) 
Explanation:
Ama – Amaranthaceae, Ast – Asteraceae, Api – Apiaceae, Bal – Balsaminaceae, Bra – Brassicaceae, Car – Caryophyllaceae, Oxa – Oxalidaceae,  
Poa – Poaceae, Por – Portulacaceae, Sap – Sapindaceae, Sim – Simaroubaceae, Sol – Solanaceae; T – therophyte, He – hemicryptophyte, 
Ph – phane rophyte; Af – Africa, As – Asia, E – Europe, CAm – Central America, NAm – North America, SAm – South America; arch – archaeo-
phyte, neo – neophyte.
Slika 10: Delež invazivnih taksonov v ruderalni vegetaciji (vsi sintaksoni skupaj) v Brnu (Češka) in uvrstitev v družine (a), Raunkiærjeve življenske 
oblike (b), izvorna celina (c), čas vnosa (d).
Legenda:
Ama – Amaranthaceae, Ast – Asteraceae, Bal – Balsaminaceae, Bra – Brassicaceae, Cuc – Cucurbitaceae, Fab – Fabaceae, Jun – Juncaceae, Poa – 
Poaceae, Pog – Polygonaceae, Sap – Sapindaceae, Sim – Simaroubaceae, Sol – Solanaceae; T – therophyte, G – geophyte, He – hemicryptophyte, 
Ph – phanerophyte; Af – Africa, As – Asia, E – Europe, CAm – Central America, NAm – North America, SAm – South America; arch – arheofiti, 
neo – neofiti.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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Figure 11: Changes in the mean Ellenberg indicator values for light (a), temperature (b), continentality (c), moisture (d), soil reaction (e) and 
nutrients (f ) between the old (years 1960–1982) and the more recent (years 2005–2016) time period.
Slika 11: Sprememba povprečnih Ellenbergovih indikatorskih vrednosti za svetlobo (a), temperaturo (b), kontinentalnost (c), vlažnost (d), reakcijo 
tal (e) in hranila (f ) med starejšim (1960–1982) in novejšim (2005–2016) obdobjem.
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
Dynamics of the species diversity and composition of the ruderal vegetation of 
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tats, the trend of increasing proportion of neophytes over 
the time have been observed (Pyšek & Mandák 1997, 
Chocholoušková & Pyšek 2003, Lososová & Simonová 
2008, Gregor et al. 2012), and the proportion of native 
species decreased accordingly (Chocholoušková & Pyšek 
2003, Lososová & Simonová 2008). Over time, signifi-
cant increase in the proportion of neophytes was recorded 
not only in the urban areas, but also in the hardwood 
floodplain forests in the Pannonian Region (Petrášová et 
al. 2013) and in many other habitats in Slovakia (Med-
vecká et al. 2014). 
The increase of invasive species in the ruderal vegetation 
of the studied Slovak and Czech cities (Figures 7–8) can 
be explained by the increase in mean Ellenberg indicator 
values for temperature and light and decrease of values 
for moisture between the old and the more recent period 
(Figure 11), because some of recorded invasive species are 
native to warmer and drier regions (Figures 9–10). In-
creasing drought in cities could be evocated by pavement 
and road construction, and land drainage for the purpose 
of other construction activities. Moreover, the boom of 
building activity itself could contribute to the increase of 
invasive taxa proportion as some invasive neophyte spe-
cies are encouraged by building activity (Kowarik 1990).
There can be several explanation for the fact that the 
classes Digitario sanguinalis-Eragrostietea minoris and 
Sisymbrietea are rich in the invasive alien species and 
the classes Artemisietea vulgaris and Epilobietea angusti-
folii are less invaded (Figures 7–8). The majority of the 
recorded invasive taxa in all cities in both periods were 
therophytes (Figures 9–10). The classes Digitario sangui-
nalis-Eragrostietea minoris and Sisymbrietea are comprised 
of therophyte-rich ruderal vegetation, while the propor-
tion of therophytes in the classes Artemisietea vulgaris 
and Epilobietea angustifolii is lower (Mucina et al. 2016). 
Therefore, the high proportion of the invasive species in 
the classes Digitario sanguinalis-Eragrostietea minoris and 
Sisymbrietea is not surprising. The thermophilous vegeta-
tion of the class Digitario sanguinalis-Eragrostietea minoris 
rich in grasses and other C4 species can host many inva-
sive species also because of the fact that relatively many 
of recorded invasive species belong to the family Poaceae 
and originate from warmer regions (Figures 9–10) and 
therefore, are adapted to the dry conditions.
There can be also other reasons explaining the recorded 
proportion of the invasive species in various classes. The 
resistance of the community to invasion may correlate 
with the structure of the community and with effect of 
disturbance, which creates empty niches and raises the 
probability of colonising by alien species generally (Hobbs 
& Huenneke 1992, Mack et al. 2000). This tendency can 
be seen also in the studies from Czech Republic (Chytrý 
et al. 2005, 2008, Simonová & Lososová 2008) in the 
case of the proportion of alien taxa. These studies confirm 
that habitats with dense vegetation have low proportion 
of alien taxa, while habitats types with open vegetation, 
where the vegetation is sparse due to disturbances, con-
tain many alien taxa. According to these studies, the dis-
turbance regime is the main difference between the most 
invaded and other habitats. All of the habitats most in-
vaded by alien plants experience strong disturbances. For 
ruderal habitats, the impact of disturbance was found to 
be the most significant determinant of invasion (Chytrý et 
al. 2005, 2008, Simonová & Lososová 2008). The record-
ed communities of the classes Sisymbrietea and Digitario 
sanguinalis-Eragrostietea minoris were sparse and many of 
them were recorded in habitats with high level of distur-
bance, e.g. construction sites. Some of the recorded com-
munities of the class Digitario sanguinalis-Eragrostietea 
minoris belong to the alliance Eragrostio-Polygonion aren-
astri which comprises summer-dry trampled ruderal com-
munities on sandy soils of Western and Central Europe 
(Mucina et al. 2016). Trampling is a strong disturbance 
too (Čarni & Mucina 1998). The recorded communities 
of the classes Artemisietea vulgaris and Epilobietea angus-
tifolii were denser and the majority of them occupied not 
so extremely disturbed ruderal habitats.
There are also other reasons for high proportion of the 
invasive species in the class Digitario sanguinalis-Eragros-
tietea minoris and lower proportion of the invasive species 
in the another classes such as Artemisietea vulgaris and 
Epilobietea angustifolii. Communities with higher bio-
diversity (e.g. the class Artemisietea vulgaris in our case) 
are more stable and therefore more resistant to invasions, 
because a community with high biodiversity is unlikely to 
have any vacant niches that cannot be defended from an 
immigrant (Case 1990, Tilman 1997, Mack et al. 2000). 
The class Digitario sanguinalis-Eragrostietea minoris has 
the highest average percentual number of the invasive 
species (Figures 7–8) probably because it has low diver-
sity and average percentual number of native species (Fig-
ures 1–6) and therefore has vacant niches that cannot be 
defended from aliens. The reason why the class Digitario 
sanguinalis-Eragrostietea minoris has low diversity and 
average percentual number of native species can be that 
the communities of this class grow on trampled habitats, 
where only very few species can survive.
The high proprotion of the invasive species could be 
consequence of the vulnerable plant community with poor 
biodiversity (Case 1990, Tilman 1997). Vice versa, lower 
biodiversity could be already a consequence of high pro-
portion of the invasive taxa (Manchester & Bullock 2000, 
Stohlgren et al. 2011). One of the causes of the biodiversity 
loss is the invasion of alien species (Manchester & Bullock 
Alena Rendeková, Karol Mičieta, Zuzana Randáková & Ján Miškovic 
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2000, Stohlgren et al. 2011). The extinctions, for which in-
vasive taxa are responsible, cause the global diversity to be 
reduced much faster than it is recovered (Russel & Black-
burn 2017). Understanding of behaviour of invasive spe-
cies in cities is very important, because cities represent the 
sources from which invasive species spread into native veg-
etation (Sukopp & Werner 1983). Our study confirmed 
the presumption that various human activities in urban 
environment can be reflected by an increasing proportion 
of alien invasive species over the time. These results can be 
useful and helpful in solving nature protection issues.
Acknowledgements
We thank Aaron Fishbone for English grammar proof-
reading of the manuscript. This research was supported 
by the Grant Agency VEGA (Bratislava), Grant No. 
1/0885/16.
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