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Abstract

Alien flora of the Czech Republic is presented. In Appendix 1, 1378 alien taxa (33.4% of the total flora) are listed with information on the taxonomic position, origin, invasive status (casual, naturalized, invasive; a new category post-invasive is introduced), time of immigration (archaeophytes vs. neophytes), habitat type invaded (natural, seminatural, human-made), vegetation invaded (expressed as occurence in phytosociological alliances), mode of introduction into the country (accidental, deliberate), and date of the first record. Number of phytogeographical as well as biological and ecological attributes were compiled for each species in the database; its structure is presented in Appendix 2 as a suggestion for similar work elsewhere. Czech alien flora consists of 24.1% of taxa which arrived before 1500 (archaeophytes) and 75.9% neophytes. There are 891 casuals, 397 naturalized and 90 invasive species. Of introduced neophytes, 21.9% became naturalized, and 6.6% invasive. Hybrids contribute with 13.3% to the total number of aliens, and the hybridization is more frequent in archaeophytes (18.7%) than in neophytes (11.7%). If the 184 hybrids are excluded from the total number of aliens, there are 270 archaeophytes and 924 neophytes in the Czech flora, i.e. total of 1195 taxa. Accidental arrivals account for 53.4% of all taxa and deliberate introduction for 46.6%; the ratio is reversed for neophytes considered separately (45.5 vs. 54.5%). Majority of aliens (62.8%) are confined to human-made habitats, 11.0% were recorded exclusively in natural or seminatural habitats, and 26.2% occur in both types of habitat. Archaeophytes and neophytes occur in 66 and 83 alliances, respectively, of the phytosociological system. Flora is further analysed with respect to origin, life histories, life forms and strategies. Only 310 species (22.4% of the total number of all alien taxa) are common or locally abundant; others are rare, based on a single locality or no longer present. The following 19 taxa are reported as new for the Czech alien flora: Agrostis scabra, Alhagi pseudalhagi, Allium atropurpureum, Bromus hordeaceus subsp. pseudothominii, Carduus tenuiflorus, Centaurea xgerstlaueri, Centaurea nigra x phrygia, Cerastium xmaureri, Gilia capitata, Helianthus strumosus, Hieracium pannosum, Hordeum leporinum, Oenothera coronifera, Papaver atlanticum subsp. mesatlanticum, Parietaria pennsylvanica, Polypogon fugax, Rodgersia aesculifolia, Sedum pallidum var. bithynicum, Sedum stoloniferum; these represent results of our own field research as well as of herbaria search, and unpublished data from colleagues. Other 44 taxa are reported as escaping from cultivation for the first time. Twenty two archaeophytes are listed in the Red List of the Czech flora.
Catalogue of alien plants of the Czech Republic (2nd edition):
checklist update, taxonomic diversity and invasion patterns
Nepůvodní flóra České republiky: aktualizace seznamu druhů, taxonomická diverzita a průběh invazí
Petr P y š e k1,2, Jiří D a n i h e lk a1,3, Jiří S á d lo1, Jindřich C h r t ek Jr.1,4,
Milan C h y t r ý3, Vojtěch J a r o š ík2,1, Zdeněk K ap l an1, František K ra h u l ec1,
Lenka M o r a v c o v á1, Jan P er g l1, Kateřina Št a j e r o v á1,2 & Lubomír T i c h ý3
1Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice,
Czech Republic, e-mail: pysek@ibot.cas.cz, chrtek@ibot.cas.cz, kaplan@ibot.cas.cz,
krahulec@ibot.cas.cz, moravcova@ibot.cas.cz, pergl@ibot.cas.cz, stajerova@ibot.cas.cz;
2Department of Ecology, Faculty of Science, Charles University in Prague, Viničná 7, CZ-
128 44 Prague, Czech Republic, e-mail: jarosik@cesnet.cz; 3Department of Botany and
Zoology, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic, e-mail:
danihel@sci.muni.cz, chytry@sci.muni.cz, tichy@sci.muni.cz; 4Department of Botany,
Faculty of Science, Charles University in Prague, Benátská 2, CZ-128 01 Prague, Czech
Republic
Pyšek P., Danihelka J., Sádlo J., Chrtek J. Jr., Chytrý M., Jarošík V., Kaplan Z., Krahulec F.,
Moravcová L., Pergl J., Štajerová K. & Tichý L. (2012): Catalogue of alien plants of the Czech
Republic (2nd edition): checklist update, taxonomic diversity and invasion patterns. – Preslia 84:
155–255.
A complete list of all alien taxa ever recorded in the flora of the Czech Republic is presented as an
update of the original checklist published in 2002. New data accumulated in the last decade are
incorporated and the listing and status of some taxa are reassessed based on improved knowledge.
Alien flora of the Czech Republic consists of 1454 taxa listed with information on their taxonomic
position, life history, geographic origin (or mode of origin, distinguishing anecophyte and hybrid),
invasive status (casual; naturalized but not invasive; invasive), residence time status (archaeophyte
vs neophyte), mode of introduction into the country (accidental, deliberate), and date of the first
record. Additional information on species performance that was not part of the previous catalogue,
i.e. on the width of species’ habitat niches, their dominance in invaded communities, and impact, is
provided. The Czech alien flora consists of 350 (24.1%) archaeophytes and 1104 (75.9%) neo-
phytes. The increase in the total number of taxa compared to the previous catalogue (1378)is due to
addition of 151 taxa and removal of 75 (39 archaeophytes and 36 neophytes),importantpart of the
latter being the reclassification of 41 taxa as native, mostly based on archaeobotanical evidence.The
additions represent taxa newly recorded since 2002 and reported in the national literature; taxa
resulting from investigation of sources omitted while preparing the previous catalogue;
redetermination of previously reported taxa; reassessment of some taxa traditionally considered
native for which the evidence suggests the opposite; and inclusion of intraspecific taxa previously
not recognized in the flora. There are 44 taxa on the list that are reported in the present study for the
first time as aliens introduced to the Czech Republic or escaped from cultivation: Abies concolor,
A. grandis,A. nordmanniana,Avena sterilis subsp. ludoviciana,A. ×vilis,Berberis julianae,
B. thunbergii,Bidens ferulifolius,Buddleja alternifolia,Buglossoides incrassata subsp. splitgerberi,
Buxus sempervirens,Corispermum declinatum,Cotoneaster dielsianus,C. divaricatus,Euphorbia
myrsinites,Gleditsia triacanthos,Helleborus orientalis,Hieracium heldreichii,Koelreuteria pani-
culata,Lonicera periclymenum,Lotus ornithopodioides,Malus baccata,M. pumila,Miscanthus
sacchariflorus,Morus alba,Muscari armeniacum,Paeonia lactiflora,Pennisetum alopecuroides,
Pinguicula crystallina subsp. hirtiflora,P. grandiflora subsp. rosea,Podophyllum hexandrum,
Pyracantha coccinea,Rhodotypos scandens,Rumex patientia × R. tianschanicus ‘Uteuša’, Salix
cordata,Sarracenia purpurea,Sasa palmata ‘Nebulosa’, Scolymus maculatus,Spiraea japonica,
Preslia 84: 155–255, 2012 155
Tagetes tenuifolia,Thuja occidentalis,Trifolium badium,Vaccinium corymbosum and Viburnum
rhytidophyllum. All added and deleted taxa are commented on. Of the total number of taxa, 985 are
classified as casuals, 408 as naturalized but not invasive, and 61 as invasive. The reduction in the
number of invasive taxa compared to the previous catalogue is dueto a more conservative approach
adopted here; only taxa that currently spread are considered invasive. Casual taxa are strongly over-
represented among neophytes compared to archaeophytes (76.7% vs 39.4%), while naturalized but
non-invasive taxa follow the reversed pattern (18.8% vs 57.4). However, these two groups do not
significantly differ in the proportion of invasive taxa. Of introduced neophytes, 250 taxa (22.6%) are
considered vanished, i.e. no longer present in the flora, while 23.3% became naturalized,and 4.5%
invasive. In addition to the traditional classification based on introduction–naturalization–invasion
continuum, taxa were classified into 18 population groups based on their long-term trends in
metapopulation dynamics in the country, current state of their populations, and link to the propagule
pressure from cultivation. Mappingthese population groups onto the unified framework for biologi-
cal invasions introduced by Blackburn et al. in 2011 made it possible to quantify invasion failures,
and boom-and-busts, in the Czech alien flora. Depending on inclusion criteria (whether or not
extinct/vanished taxa and hybrids are considered), alien taxa ever recorded in the Czech Republic
contribute 29.7–33.1% to the total country’s plant diversity; taking into account only naturalized
taxa, a permanent element of the country’s flora, the figure is 14.4–17.5%. Analysis of the dates of
the first record, known for 771 neophytes, indicatesthat alien taxa in the flora have been increasing
at a steady pace without any distinct deceleration trend; by extrapolating this data to all 1104 neo-
phytes recorded it is predicted that the projected number would reach 1264 in 2050. Deliberate
introduction was involved in 747 cases (51.4%), the remaining 48.6% of taxa are assumed to have
arrived by unintentional pathways. Archaeophytes are more abundant in landscapes, occupy on
average a wider range of habitat types than neophytes, but reach a lower cover in plant communities.
The alien flora is further analysed with respect to representation of genera and families, origin and
life history.
K e y w o r d s: abundance, alien flora, checklist, casual, cover in plant communities, Czech Repub-
lic, exotic species, geographic origin, habitat niche, hybridization, impact, introduction–naturaliza-
tion–invasion continuum, invasive plants, life history, naturalized, non-native species, residence
time, taxonomy
Introduction
The last decade was a period of intensive research on biological invasions in Europe (see
Pyšek & Hulme 2011 for review), an important part of which represented the collation of
regional data on alien plant species. With the exception of the UK (Clement & Foster
1994, Ryves et al. 1996, Preston et al. 2002), complete checklists of alien floras for Euro-
pean countries only started to appear at the beginning of the 2000s (Essl & Rabitsch 2002,
Klotz et al. 2002, Reynolds 2002). The first comprehensive checklist of alien plants in the
Czech Republic was published 10 years ago as a part of the Catalogue of alien plants of the
Czech Republic (Pyšek et al. 2002). It provided information on 1378 alien taxa andstimu-
lated development of the associated database CzechFlor, held at the Institute of Botany AS
CR in Průhonice. These data, together with other datasets resulting from recent research,
have been used for a number of analyses of plant invasions in the country that addressed
issues such as species invasiveness (Kubešová et al. 2010, Moravcová et al. 2010), associ-
ations with pollinators (Pyšek et al. 2011a), habitat invasibility (Chytrý et al. 2005, 2008a,
2009b, Sádlo et al. 2007), rates of spread and range filling (Williamson et al. 2005, 2009,
Pyšek et al. 2011c), interaction of traits, propagule pressure and residence time in affect-
ing invasion success (Pyšek et al. 2009b), pathway efficiency (Pyšek et al. 2011b), and risk
assessment (Křivánek & Pyšek 2006, Chytrý et al. 2009b). In addition, data on native
156 Preslia 84: 155–255, 2012
species that are also part of the CzechFlor database provided basis for analyses of the per-
formance of central-European species as aliens in other parts of the world (Pyšek et al.
2009a, Phillips et al. 2010, Stohlgren et al. 2011). Within the DAISIE and ALARM
(Settele et al. 2005) projects, the data from the 2002 catalogue were part of the pan-Euro-
pean dataset that was used to analyse invasion patterns at the continental level, including
cross-taxonomic evaluation of the role of macroeconomic and demographic factors in
determining regional levels of invasion (Pyšek et al. 2010b, Essl et al. 2011), distribution
of alien species in habitats (Pyšek et al. 2010a), assessment of ecological and economic
impacts of alien species in Europe (Winter et al. 2009, Vilàet al. 2010) and risk-assessment
for plants based on habitat mapping (Chytrý et al. 2008b, 2009a, 2012).
These studies clearly indicate the value of complete national or regional checklists for
understanding invasions. This started to be fully recognized in the 2000s and resulted in
a call for pan-European inventory of invasive species within the European framework
programmes; until then there was some information on alien floras available for European
countries (Weber 1997), but the quality of data was highly variable (Pyšek 2003). The
DAISIE project (2004–2008) made it possible to organize and develop this line of research
based on extensive international cooperation in Europe (DAISIE 2009). The project
assembled available data on alien plants for 48 European countries and regions, which
until then were scattered in a variety of published and unpublished accounts and data-
bases. For some countries DAISIE collected the first comprehensive checklists of alien
species based on primary data (Lambdon et al. 2008), and established an online database,
the European Invasive Alien Species Gateway (DAISIE 2008). At the same time it stimu-
lated elaboration of comprehensive alien species checklists in individual countries, a pro-
cess that still continues, and yielded new plant data for e.g. Belgium (Verloove 2006),
Estonia (Ööpik et al. 2008), Italy (Celesti-Grapow et al. 2009), Greece (Arianoutsou et al.
2010), and most recently Slovakia (Medvecká et al. 2012).
The Czech Republic, a central-European country with an area 78,864 km2, 10.3 million
inhabitants, and a human population density of 131 inhabitants per km2, is prone to plant
invasions due to historical and geographical factors: location on the crossroads of the con-
tinent, many natural or human-created migration routes opening possibilities for coloniza-
tion, and long-lasting human influence that further diversified the naturally diverse and
heterogeneous landscape mosaic (see Pyšek et al. 2002 for details). These features,
together with a strong botanical tradition and in-depth knowledge of plant communities
(Chytrý 2007, 2009, 2011) make the country a suitable model for studying regional pat-
terns of plant invasions. In the last decade since the publication of the previous catalogue
a wealth of information on alien species has been accumulated, which created a need for
a revision of the original checklist.
The aim of the present paper is to update and improve the original checklist of alien
plant taxa in the Czech Republic (Pyšek et al. 2002) by incorporating new data accumu-
lated in the last decade, reassessing the status of taxa resulting from improved taxonomic
knowledge, and wherever needed, correcting errors which can hardly be avoided in such
a comprehensive work. We also provide additional information that was not part of the
previous catalogue, including the width of species’ habitat niches, their dominance in
invaded communities and their impacts. Changes from the 2002 version are documented
so that the reasoning behind them can be followed.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 157
Methods
Data sources
The basis for the present checklist was the Catalogue of alien plants of the Czech Republic
published a decade ago (Pyšek et al. 2002). For historical data, the compilation of both the
previous and current checklist relied on an outstanding tradition of the floristic research in
the Czech Republic dating back to the second half of the 18th century (reviewed in detail in
Pyšek et al. 2002). Already in the 19th century, a series of floras and species lists were pub-
lished, covering the present territory of the Czech Republic (see Krahulec 2012 for a review
of the history of botanical research), and recognizing plants by geographic origin; thesepro-
vide valuable information about the occurrence of plants at those times and residence times
of neophytes (Pohl 1809–1814, Presl & Presl 1819, Opiz 1823, 1852, Rohrer & Mayer
1835, Makowsky 1863, Oborny 1886, Formánek 1887–1897). The wealth of information on
alien plants can be found especially in the remarkable works by Čelakovský (1868–1883,
1882–1894), who recognized the alien status and origin of some plants present in the Czech
flora and commented in considerable detail on their distribution. The recognition of alien
plants continued in floras and specialized studies in the 20th century (e.g. Polívka
1900–1904, Laus 1908, Domin 1917, 1918, 1919, Dostál et al. 1948–1950, 1954, 1958,
1989). Since the 1960s, systematic attention started to be paid to plants, including aliens, in
specific human-made habitats (ports, railways, oilseed or wool processing factories, grain
silos, mills, rubbish tips, arable land, etc.) thanks to a specialized research section estab-
lished at the Institute of Botany, Průhonice, in the 1960s. This work yielded several focused
compendia (e.g. Hejný et al. 1973) and provided a basis for systematic recording of alien
plants (e.g. Jehlík 1986, 1998a).
The Flora of the Czech Republic, with eight of nine planned volumes published up to
now (Hejný & Slavík 1988–1992, Slavík 1995, 1997a, 2000, Slavík & Štěpánková 2004,
Štěpánková 2010) and the Key to the flora of the Czech Republic (Kubát et al. 2002), served
as a fundamental information source for this checklist. Other recent sources included
national floristic literature, namely that published in the journals of the Czech Botanical
Society (see References). During the last decade, new records for the flora of the Czech
Republic have been systematically reported in an annually published series, Additamenta ad
floram Reipublicae Bohemicae, which has thus far yielded 10 summarizing accounts
(Hadinec et al. 2002, 2003, 2004, 2005, Hadinec & Lustyk 2006, 2007, 2008, 2009, 2011,
2012). The series, initiated and edited by J. Hadinec, in cooperation with František
Procházka and Pavel Lustyk, proved a valuable source because it not only reports new finds
but also critically re-evaluates status of particular species and provides additional data on
their distribution.
For archaeophytes, a strong tradition of Czech archaeobotanical research provided
a solid basis for evaluation of species origin and immigration status. Main sources include
the works of E. Opravil and V. Čulíková (see References), the results of which are now
available in the Archaeobotanical database of the Czech Republic (CZAD; Archaeologi-
cal Institute AS CR 2011).
Other data sources included unpublished information provided by many colleagues
(see Acknowledgments), herbarium collections to verify some literature reports (namely
PR, PRC, BRNU and PRA; codes follow Thiers 2012) and our own floristic field records
from 2002–2012.
158 Preslia 84: 155–255, 2012
The data presented here and in the previous catalogue (Pyšek et al. 2002) are organized
in the working database CzechFlor held at the Institute of Botany AS CR, Průhonice.
Classification of taxa: invasion status
This work focuses on alien species (synonyms: adventive, exotic, introduced, non-indige-
nous, non-native) in the Czech Republic which we define as species present in the region
because human actions enabled them to overcome fundamental biogeographical barriers
(i.e. human-mediated extra-range dispersal); they occur in the area as a result of inten-
tional or accidental introduction by humans, or of a spontaneous spread from other regions
where they were introduced by humans. Crosses resulting from hybridization with one or
both alien species involved are considered alien (Pyšek et al. 2004a). We define native
species (synonym: indigenous species) as those that have evolved in a given area or that
arrived there by natural means (through range expansion) without any intentional or acci-
dental intervention of humans from an area where they are native (Pyšek et al. 2004a).
We classified species according to the stage they reached along the introduction–natu-
ralization–invasion continuum (INIC) that describes how species proceed in the invasion
process by overcoming geographical, environmental and biotic barriers (Richardson et al.
2000, 2011, Richardson & Pyšek 2006, Blackburn et al. 2011). Based on this concept we
use the following terms to describe the invasion status:
(i) Casual species are those alien species that do not form self-sustaining populations
in the invaded region; they may flourish and reproduce occasionally in an area but their
persistence depends on repeated introductions of propagules.
(ii) Naturalized species (synonym: established species) form self-sustaining popula-
tions for several life cycles without direct intervention by people, or despite human inter-
vention; they often recruit offspring freely, usually close to adult plants and their persist-
ence does not depend on ongoing input of propagules.
(iii) Invasive species are a subset of naturalized species; they form self-replacing popu-
lations over many life cycles, produce reproductive offspring, often in very large numbers
at considerable distances from the parent and/or site of introduction, and have the potential
to spread over long distances. In addition to this definition, we introduce the
metapopulation criterion to separate invasive species from naturalized, to account for the
historical population dynamics of the treated taxa (see the next section).
We included in the list all taxa that were reported to occur at least once in the wild,
while those kept exclusively in cultivation are not considered. For escapees from cultiva-
tion, a plant was included in the list if it reproduced on its own outside the space where it
was sown or planted (Pyšek et al. 2002). In plants reproducing by seed, germination out-
side such space was considered as an escape from cultivation. A plant reproducing
clonally was considered as an escape from cultivation only if it survived winter and per-
sisted in a given site until the following growing season.
Compared to the previous catalogue (Pyšek et al. 2002), we adopted a more conserva-
tive approach; if there were doubts about a species’ origin status and no strong evidence to
consider it alien, it was not included in the list; this conservative approach resulted in
removing some species that were listed in the previous catalogue (see Appendix 1).
The classification of casual vs naturalized status is especially difficult for woody plants
reproducing in the parks or gardens where they are planted; in some cases this happens
Pyšek et al.: Catalogue of alien plants of the Czech Republic 159
over a large area and for decades (e.g. many trees and shrubs in the Průhonice Park near
Prague where there is a long-term systematic recording of regeneration). Here we aimed at
adopting the criterion of reproduction over several generations (Richardson et al. 2000)
which puts the time criterion in a different perspective than that applied for non-woody
taxa. Such taxa are therefore mostly classified as casual. Also, the majority of hybrids are
considered casual, with the exception of stabilized hybrids that include some naturalized
(e.g. Medicago ×varia,Helianthus ×laetiflorus,Mentha ×rotundifolia and Oenothera
spp.) or invasive taxa (e.g. Reynoutria ×bohemica,Populus ×canadensis and Symphyo-
trichum ×versicolor).
Unlike the previous catalogue (see Pyšek et al. 2002 and their Appendix 1), we do not
explicitly label taxa as locally naturalized. In the present paper this can be inferred from
the combination of invasion status and regional abundance category in Appendix 2. In the
same vein, taxa are not labelled as post-invasive since this status is included in the classifi-
cation using the population groups (see below).
Classification of taxa into categories based on long-term population dynamics and histor-
ical link with cultivation: incorporating the unified framework for biological invasions
In addition to traditional classification scheme dividing species into three basic categories
along the INIC (Richardson et al. 2000, Richardson & Pyšek 2006, Pyšek & Richardson
2010) here we attempt for an even finer classification based on the population approach
emphasized by Blackburn et al. (2011). The basis for this classification are the criteria of
reproduction and survival applied against the background of the metapopulation
approach. This makes it possible to separate species that survive in a single or few popula-
tions in a spatially restricted area from those that spread and form metapopulations over
large areas.
Another important point to emphasize is that we refer to the population history viewed
from the current perspective, i.e. the state in which the populations of a given species
exist at present. Therefore, invasions that proved unsuccessful in proceeding along the var-
ious stages of the INIC (see Blackburn et al. 2011 and their Fig. 1) are reflected in the cur-
rent classification, and in changes of invasion status compared to the previous treatment
(Pyšek et al. 2002). From this it follows that some taxa that were previously classified as
naturalized are moved to the casual category (reflecting ‘invasion failure’), and some taxa
previously considered invasive are now classified as naturalized (reflecting ‘boom and
bust phenomenon’; sensu Blackburn et al. 2011). These shifts among the INIC categories
reflect not only changes in species’ behaviour in the past decade but also the more conser-
vative approach adopted for the current classification. Another principle we follow is that
of the highest stage achieved at the population level; individual populations of an alien
species may occur in a region in different stages of the INIC; early in the process, some can
be naturalized while others are still casual (e.g. Essl et al. 2009), whereas later on, some
can be invasive while others not (e.g. Meyerson et al. 2010a, b, Saltonstall et al. 2010).
Therefore, if some of the populations of a species reached the naturalized or invasion
stage, the species is classified as such in Appendix 2.
Therefore, the rationale of classification of alien species into finer groups (termed
‘population groups’) is based on the following criteria (Table 1):
160 Preslia 84: 155–255, 2012
(A) Sustainability of populations of the species in the target region of the Czech
Republic; here we distinguish between (i) species existing as non-self-sustaining popula-
tions or occasionally recorded individuals, corresponding to Blackburn et al.’s (2011) cat-
egories B3+C2, and the casual stage of Richardson et al.’s (2000) framework; the reason
for lumping the categories B3 (defined as individuals transported beyond limits of native
range, and directly released into novel environment) and C2 (individuals surviving in the
wild in location where introduced, reproduction occurring, but population not self-sus-
taining) is that from records in floristic literature it is impossible to infer whether the pres-
ence of the plant is due to a direct introduction of a propagule into the region or a result of
a temporary reproductive event within the region; (ii) species occurring in self-sustaining
populations; these populations can be numerous and widespread but remain isolated
(C3+D1+D2, naturalized species – lumping due to insufficient knowledge about whether
the populations recruit from the original point of introduction and whether those spread far
from it reproduce in new locations); and (iii) species that currently form numerous and
persistent metapopulations widespread over large areas (Blackburn et al.’s 2011 category E).
(B) Historical population dynamics is used to classify species according to the high-
est stage they reached in the invasion process combined with the current state. We distin-
guish whether or not the most successful populations of unsuccessful species have estab-
lished and were surviving in the region before decline to the current levels of occurrence;
successful species are classified based on the tendency for spread, with respect to whether
this trend occurred in the past or is still valid (Table 1). Employing this criterion, i.e. focus
on the current status of species’ populations and processes that resulted in the present
state, is the reason why the correspondence with the categories of Blackburn et al. (2011)
is, however, not automatically translated into those of the introduction–naturaliza-
tion–invasion continuum. This concerns those species classifiedas D1, D2 and considered
invasive in Blackburn et al.’s (2011) scheme (self-sustaining population in the wild, with
individuals surviving, or also reproducing, a significant distance from the original point of
introduction), populations of which no longer exhibit dynamic spread and are currently
stabilized (Groups 7, 9, 11 in Table 1), or even decline in the Czech Republic (Group 6).
We also do not consider as invasive those species that only start to exhibit symptoms of the
beginning spread (Groups 8, 10, 12). Adhering to a conservative approach, these species
are still considered as naturalized. Nevertheless, they merit particular attention in terms of
monitoring as they are likely to become invasive in the near future. Only those species that
are currently spreading are classified as invasive (Groups 14, 16, 18; Table 1).
(C) Link to populations in cultivation. The above criteria are employed against the
background of species’ planting histories in the region. Here we separate species into (i)
those that have never been cultivated (corresponding to contaminant and stowaway path-
ways of introduction according to Hulme et al. 2008; Appendix 2), hence unsupported by
the propagule pressure from planted populations; (ii) those in which the peak of planting
intensity was in the past and at present the planting ceased or is only of marginal impor-
tance; and (iii) those that are still commonly kept in cultivation, be it for horticultural or
agricultural purposes. For the cultivated speciesthis criterion refers to the degree of conti-
nuity of propagule pressure. The time frame over which this criterion applies is the last ca
200 years for which period the information on the frequency of planting can be inferred.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 161
162 Preslia 84: 155–255, 2012
Table 1. – Classification of the alien flora of the Czech Republic into population groups (PG) based on the current population state and their connectivity, trends in their long-term
dynamics, and link to cultivated populations as a source of propagule pressure in the past and present. See text for details. The population groups are referred by numbers pre-
sented in Appendix 2, with the INIC (introduction–naturalization–invasion continuum) status indicated and number of species shown in parentheses. The link to the unified inva-
sion framework (Blackburn et al. 2011) is indicated by their categories that are relevant to the given population state shown in parentheses; note that some of their categories
referring to the invasion stage such as D1, D2, E (Blackburn et al. 2011; their Fig. 1) are classified as naturalized because the focus here is on the present state and approach
adopted is conservative. Taxa in these categories may have reached the invasion stage in the past but their populations are stabilized and no longer spread. Link to standard classi-
fication of the INIC categories (Richardson et al. 2000) is indicated by coloured shading. The scheme also separates groups of taxa introduced by unintentional pathways (con-
taminant, stowaway), marked “none” in the Cultivation column, from those introduced deliberately (release, escape; Hulme et al. 2008, Pyšek et al. 2011b).
Populations Cultivation Introduction & Failure Establishment &
Failure
Establishment & No
trend
Starting spread Ongoing spread
(a) Not self-sustaining
(B3, C2)
(a1) None PG1:
casual (395)
PG2:
casual (45)
(a2) Past PG3:
casual (17)
(a3) Ongoing PG4 & 5:
casual (501 & 28)
(b) Self-sustaining
(C3, D1, D2)
(b1) None PG6:
naturalized (54)
PG7:
naturalized (40)
PG8:
naturalized (43)
(b2) Past PG9:
naturalized (36)
PG10:
naturalized (11)
(b3) Ongoing PG11:
naturalized (65)
PG12:
naturalized (31)
(c) Metapopulations
(E)
(c1) None PG13:
naturalized (100)
PG14:
invasive (28)
(c2) Past PG15:
naturalized (8)
PG16:
Invasive (9)
(c3) Ongoing Group 17:
naturalized (19)
PG18:
invasive (24)
Total taxa 924 116 268 85 61
Residence time status
Based on the residence time, i.e. the time since the arrival of a species to the territory of the
present Czech Republic, we distinguish archaeophytes (taxa introduced before the discov-
ery of America, approx. 1500 A. D.) and neophytes (taxa introduced after that date), fol-
lowing the concept traditionally used in European studies on plant invasions (e.g. Holub &
Jirásek 1967, Pyšek et al. 2002, 2004a). When evaluating residence time status of hybrids,
we followed that of the alien parent; therefore, crosses of archaeophytes with native are
considered archaeophytes, and hybridization with neophytes involved are classified as
neophytes regardless of the status of the second parent.
For neophytes, we determined the year of the first record in the Czech Republic that is
used to infer the minimum residence time, i.e. the time for which the species is known to be
present (Rejmánek 2000, Pyšek & Jarošík 2005, Richardson & Pyšek 2006); this character-
istic is important in evaluation of invasion status since it indicates how much time the species
had to colonize suitable habitats (Williamson et al. 2009, Gassó et al. 2010), go through a lag
phase (Kowarik 1995, Crooks 2005) or build relationship with native biota (Pyšek et al.
2011a). As pointed out above, the reliability of the years of first records crucially depends on
the intensity of floristic research in the past (see Pyšek et al. 2002 for discussion).
Species traits: taxonomic affiliation and life history
Taxonomic affiliation of taxa to families follows the approach of the Angiosperm Phylogeny
Group Classification: APG III (Stevens 2001 onwards, Angiosperm Phylogeny Group
2009), and Smith et al. (2006) for ferns. This classification system incorporates data from
molecular, chemical and morphological phylogenies in an attempt to represent the latest
thinking on angiosperm evolution, and in a few lineages (e.g. Scrophulariales) it differs
markedly from the traditional system.
The following life histories were assigned to the species: annual, biennial, perennial,
semishrub, shrub, tree, fern, aquatic and parasitic (see Appendix 2).
Geographic origin
Taxa were classified according to their geographic origin (native range) at the level of con-
tinents (parts of Europe other than the Czech Republic, Africa, Asia, North America
including Mexico, Central America, South America, and Australia). Unlike the previous
catalogue (Pyšek et al. 2002), we distinguished the Mediterranean region as a separate
region of origin, covering respective parts of southern Europe, northern Africa and west-
ern Asia from Turkey and Israel to Afghanistan. This broad definition of the Mediterra-
nean region corresponds to the Mediterranean, Submediterranean and Oriental Floristic
Subregions according to Meusel et al. (1965). The region delimited in this way is very con-
venient for plant invasion studies as it includes the areas of origin of Neolithic agriculture.
Indications of Europe, Asia and Africa in Appendix 2 refer to their parts other than the
Mediterranean region in this delimitation.
Hybrids and species that originated through recent hybridization are listed as a special
origin category and we employed classification based on how species originated in terms
of their evolutionary history. This approach acknowledges that some did not evolve natu-
rally, but under human influence, do not have a natural home range, and their original hab-
Pyšek et al.: Catalogue of alien plants of the Czech Republic 163
itat is unknown (Kühn & Klotz 2003). Especially for many archaeophytes, native ranges
are not known or are highly uncertain, and some archaeophytes are regarded as alien
throughout their known global range. These taxa, termed anecophytes (homeless plants;
Zohary 1962) could be cultivated plants that escaped to the wild or plants that co-evolved
with human land uses such as agriculture (Kühn & Klotz 2002, 2003, Kühn et al. 2004). In
our treatment, we follow the more conservative approach and label as anecophytes mostly
those species that evolved in cultivation, or species occurring in the wild but with their
region of origin being unknown.
Regional abundance
Type of regional abundance in the landscape was estimated for each taxon using the fol-
lowing scale: single locality, rare, scattered, locally abundant, and common across the
whole Czech Republic. A special category termed ‘vanished’ relates to the taxa for which
no records have been known for a long period, and where it is highly improbable that they
would appear again (Pyšek et al. 2002).
Occurrence in habitats
The previous catalogue provided information on the occurrence of alien species in phyto-
sociological alliances, different types of landscapes and with respect to landuse (Pyšek et
al. 2002). Here we use extensively revised data from the database of species occurrences in
88 major habitat types of the Czech Republic as defined by Sádlo etal. (2007), which cor-
respond to phytosociological alliances or groups of alliances. All four levels of species
affinity to the habitats as defined by Sádlo et al. (2007: 305) are taken into account, i.e.
a species is considered as occurring in a habitat even if the habitat is outside its ecological
optimum, but the species is occasionally found there.
Cover in plant communities
To obtain the data on the cover of alien species in plant communities, we used vegetation
plot observations (phytosociological relevés) stored in the Czech National Phytosocio-
logical Database held at the Department of Botany and Zoology, Masaryk University,
Brno (Chytrý & Rafajová 2003, EU-CZ-001 according to Dengler et al. 2011). At the time
of data extraction (April 2012) the database contained 88,215 relevés from plots smaller
than 1000 m2with an indication of plot size and geographical coordinates. Of these,
41,582 relevés contained at least one alien species. To reduce oversampling of some areas
or some vegetation types, we selected only one relevé from a group or relevés assigned to
the same phytosociological alliance within the same grid cell of 1.25 longitudinal × 0.75
latitudinal minutes, i.e. approximately 1.5 × 1.4 km. This stratified resampling yielded
16,033 relevés containing 437 alien species, which were used to quantify species cover.
Only species occurring in at least 25 relevés were evaluated to avoid inaccuracies resulting
from small sample size. For these species, mean percentage cover across all relevés in
which the species was present was calculated.
Impact
To provide the first insights into the impacts of alien plant species in the Czech Republic,
we used the data gathered by the DAISIE project (DAISIE 2008, 2009) and indicated
164 Preslia 84: 155–255, 2012
those species on our list for which an ecological and/or economic impact is reported in the
literature (Vilàet al. 2010). With a few exceptions indicated in Appendix 2, this classifica-
tion has not been done specifically for the Czech Republic but refers to any region in
Europe, meaning that species labelled as exerting impact may not do so in this country.
Statistical analysis
To test whether there are differences between species numbers according to their invasion
status, life histories, abundances and origins, their counts were analysed by row × column
contingency tables, using generalized linear models with log-link function and Poisson
distribution of errors (e.g. Crawley 1993: 231–237). To ascertain for which species the
counts are lower or higher than would be expected by chance, adjusted standardized resid-
uals of G-tests were compared with critical values of normal distribution (Řehák &
Řeháková 1986). The estimates of yearly accumulations of neophytes, including projected
total numbers in 2050, were assessed from linear regressions of cumulative numbers that
started in the year 1800.
Results and discussion
Diversity of alien flora
The alien flora of the Czech Republic consists of 1454 taxa, made up by 350 archaeo-
phytes (24.1%) and 1104 neophytes (75.9%; Table 2, Appendix 2), which represent addi-
tion to ca 2945 native taxa known from the country (using a preliminary estimate from
Danihelka et al. 2012) and form 33.1% of the total plant diversity ever recorded there.
Although similar figures for individual countries are subject to variation resulting not only
from composition of floras but also from the variable depth of their knowledge, intensity
of research into alien species, or whether apomictic species are included in comparisons
(see Williamson 2002, Pyšek et al. 2002 and discussion therein), the proportion given here
seems to reasonably reflect situation in countries with detailed knowledge of their floras.
Subtracting species that are assumed to be vanished among alien (277 taxa, Appendix 2)
and extinct from native flora (153 taxa in theRed List categories A1 and A2; Danihelka et
al. 2012) yields a figure of 29.7% of aliens contributing to the plant diversity currently
occurring in the Czech Republic.
Table 2. – Numbersof all alien taxa in the Czech Republic, including hybrids, cross-tabulated across invasion sta-
tus and immigration time. Note that invasive taxa are subgroup of naturalized. Overall, the observed counts of
alien taxa (in bold) highly significantly (χ2= 193.56; df = 2; P < 0.0001) differ from counts expected by chance
(values in parentheses). Statistically highly significant deviations of individual counts from counts that can be
expected by chance are expressed by asterisks (*** P < 0.001); numbers in parentheses not followed by any sym-
bol do not differ from randomly expected values.
Naturalized
Casual Naturalized non-invasive Invasive Total
Archaeophytes 138 (235.5)*** 201 (97.6)*** 11 (14.8) 350
Neophytes 847 (748.5)*** 207 (310.3)*** 50 (47.2) 1104
All aliens 985 408 61 1454
Pyšek et al.: Catalogue of alien plants of the Czech Republic 165
If we further exclude 94 hybrids recorded from the total number of alien taxa, and com-
pare this figure with the current native species diversity without 575 hybrids (Danihelka et
al. 2002), the proportion of alien taxa 32.8%. The hybrids between neophytes and native
taxa, and between two neophytes, are more frequent than hybrids involving archaeo-
phytes. Overall, neophytes are involved in 58 hybrid combinations, archaeophytes in 42
and native species in 56 (Table 3).
Finally, considering only permanently present taxa, i.e. 469 naturalized aliens (includ-
ing both non-invasive and invasive) and the native flora without extinct representatives,
yields 14.4% contribution of alien flora to the current plant diversity, or 17.5% if hybrids
are excluded from native flora. This proportion is probably a more realistic measure of the
level of invasion of the country’s species pool than is usually given in overall figures based
on all species ever recorded, including casuals, because it better reflects the threat from
alien species’ impacts and potential for invasion debt to operate (Essl et al. 2011).
Table 3.– Numbers of hybrids in the alien flora classified according to the origin and residence time status of their
parental species. Note that the total number of hybrids across the three groups (n = 94) does not correspond to the
sum of numbers within the groups involved because all combinations are displayed row-wise. Anecophytes are
listed as species of unknown origin, the majority of which originated by hybridization in cultivation. Hybrids of
native species are not relevant (n.r.) for this comparison.
× Archaeophyte × Neophyte × Native Total within group
Archaeophyte 13 6 23 42
Neophyte 6 19 33 58
Native 23 33 n.r. 56
Hybrids total 94
Anecophytes 105
Hybrids and anecophytes total 199
Changes to the 2002 checklist
Compared to the first checklist (Pyšek et al. 2002), 75 taxa were removed (39
archaeophytes and 36 neophytes). The majority of these changes resulted from reclassify-
ing some taxa as native (41 taxa) where evidence for their alien origin was not convincing
enough under the conservative approach adopted in the present paper; they were mostly
archaeophytes but there are also six neophytes with alien status which appeared doubtful
based on recently published evidence: Agropyron pectinatum,Crocus heuffelianus,
Epilobium dodonaei,Senecio rupestris,Teucrium scorodonia and Viola tricolor subsp.
curtisii. For nine taxa previously classified as deliberately introduced casuals, the evi-
dence for escaping from cultivation was ambiguous. Other deletions relate to 10 taxonom-
ically unjustified taxa now omitted from the Czech flora, and 16 cases are doubtful records
previously only reported in the literature that cannot be considered as proven without her-
barium evidence, or taxa that were erroneously determined by the collector. All deleted
species are dealt with in detail in Appendix 1.
In total, 151 taxa not listed in Pyšek et al. (2002) are included, representing additions to
the alien flora of the Czech Republic. This includes taxa newly recorded since 2002 and (i)
reported in the literature (e.g. Convallaria majalis var. transcaucasica,Darmera peltata,
166 Preslia 84: 155–255, 2012
Dittrichia graveolens,Euphorbia agraria,Galium murale,Geranium purpureum,Gratiola
neglecta,Hypericum annulatum,Legousia pentagonia,Pimpinella peregrina and Stachys
setifera), including two volumes of the Flora of the Czech Republic published in this period
(Slavík & Štěpánková 2004, Štěpánková 2010) that report taxa missing from previous cata-
logue (e.g. Cichorium endivia,Egeria densa and Filago pyramidata); (ii) additions resulting
from investigation of sources omitted from the previous catalogue (e.g. Euphrasia
salisburgensis,Herniaria incana,Rumex longifolius subsp. sourekii,Trifolium badium and
Xerochrysum bracteatum), including some herbarium materials (e.g. Centaurea carniolica,
C. transalpina and Corispermum declinatum); (iii) redetermination of previously reported
taxa (e.g. Eriochloa punctata,Gilia achilleifolia,Hieracium sp. ex H. heldreichii agg.,
Rodgersia pinnata and Spiraea hypericifolia subsp. obovata); (iv) reassessment of some
taxa traditionally considered native for which the evidence suggests the opposite (e.g.
Eragrostis pilosa,Lathyrus hirsutus,Lilium bulbiferum,Matricaria chamomilla and Sorbus
austriaca); (v) intraspecific taxa previously not recognized in the flora (e.g. Avena sterilis
subsp. ludoviciana). Accounts on the newly added alien species in the Czech flora are given
in Appendix 1, with respective references.
In total, 44 taxa are reported in the present study for the first time as aliens introduced to
the Czech Republic or escaping from cultivation (Appendix 1): Abies concolor,A. grandis,
A. nordmanniana,Avena sterilis subsp. ludoviciana,A. ×vilis,Berberis julianae,
B. thunbergii,Bidens ferulifolius,Buddleja alternifolia,Buglossoides incrassata subsp.
splitgerberi,Buxus sempervirens,Corispermum declinatum,Cotoneaster dielsianus,
C. divaricatus,Euphorbia myrsinites,Gleditsia triacanthos,Helleborus orientalis,Hiera-
cium heldreichii agg., Koelreuteria paniculata,Lonicera periclymenum,Lotus ornitho-
podioides,Malus baccata,M. pumila,Miscanthus sacchariflorus,Morus alba,Muscari
armeniacum,Paeonia lactiflora,Pennisetum alopecuroides,Pinguicula crystallina subsp.
hirtiflora,P. grandiflora subsp. rosea,Podophyllum hexandrum,Pyracantha coccinea,
Rhodotypos scandens,Rumex patientia × R. tianschanicus ‘Uteuša’, Salix cordata,Sarra-
cenia purpurea,Sasa palmata ‘Nebulosa’, Scolymus maculatus,Spiraea japonica,Tagetes
tenuifolia,Thuja occidentalis,Trifolium badium,Vaccinium corymbosum and Viburnum
rhytidophyllum.
Finally, compared to the previous version of the catalogue (Pyšek et al. 2002), 134
names were changed due to nomenclatural reasons or development in taxonomic opinion;
these changes are summarized in Electronic Appendix 1.
Transitions along the introduction–naturalization–invasion continuum
Among the 1454 taxa, 985 (67.7%) are classified as casual, 408 (28.1%) as naturalized but
non-invasive, and 61 (4.2%) as invasive (Fig. 1, Table 2). Among casual taxa, 86.0% are
neophytes and 14.0% archaeophytes, the corresponding figures being 50.7 and 49.3%,
respectively, for naturalized, and 82.0 and 18.0% for invasive taxa. From this it follows
that casual taxa are strongly over-represented among neophytes, and naturalized among
archaeophytes (Table 2, Fig. 1), a pattern previously illustrated for the Czech flora by
Pyšek et al. (2002) and also valid for neighbouring Slovakia (Medvecká et al. 2012). Inter-
estingly, the observed numbers of neither archaeophytes nor neophytes differ from those
expected by chance, indicating that there is no difference between the two groups in the
proportion of species that reach the invasion stage (Table 2, Fig. 1).
Pyšek et al.: Catalogue of alien plants of the Czech Republic 167
Data on neophytes provide insights into the transition rates along INIC, i.e. how large
a proportion of species reach the subsequent stages of the invasion process (Fig. 2); this
proportion cannot be calculated for archaeophytes because information on casual species
from the initial periods of introduction is missing (Pyšek et al. 2002). Of the total number
of 847 recorded casual neophytes, 250 (29.5%) have not been recorded for a long period of
time and are therefore considered vanished (96 of them were only known from a single
locality), and 597 (70.5%) are currently present as casuals. Of the 1104 neophytes, 257
(23.3%) became naturalized, and 50 (19.5%) of the naturalized are considered invasive
(Fig. 2).
The approach we adopt takes into account invasion failures, represented by dotted
arrows in Fig. 2 that indicate reversed directions in the invasion process. This makes it pos-
sible, by using finer classification based on the assessment of long-term population
dynamics and its comparison with the current stage (Table 1), to map the number of taxa
onto the unified framework of biological invasions (Blackburn et al. 2011). Four types of
unsuccessful invasions can be recognized, depicted in Fig. 3 and based on population
groups described below: (i) casual taxa that failed to establish, never forming self-sustain-
ing populations (PG 1+4+5); (ii) taxa that formed self-sustaining populations in the past
but declined so that this is no longer the case (PG 2+3); (iii) taxa present for a long time
with populations surviving in the landscape; although they are still considered naturalized,
their invasion obviously failed because they are rare and their decline is likely to continue
(PG6); (iv) naturalized species that form stabilized metapopulations in the wild, some of
them reached the invasion stage in the past but their current occurrence indicates that they
declined; therefore they are considered as representatives of the boom and bust phenome-
non (PG 13+15+17; Fig. 3).
168 Preslia 84: 155–255, 2012
39.4
76.7
67.7
57.4
18.8
28.1
3.1 4.5 4.2
0
10
20
30
40
50
60
70
80
90
Archaeophytes Neophytes All aliens
Casual
Naturalized
Invasive
Percentage of taxa
Fig. 1. – Representation of taxa according to invasion status (casual, including vanished taxa; naturalized but non-
invasive; invasive) among archaeophytes, neophytes and all aliens in the flora of the Czech Republic. See Table 2
for the numbers of taxa and statistics.
Overview of population groups
(a) Not self-sustaining populations or individuals
(a1) No link to cultivation
Group 1. Introduction and failure. Unintentionally introduced taxa that were only
recorded as individuals or in small populations, mostly occasionally, and are reported
from a single or few locations; they are classified as casuals and a significant proportion
(186 of 395 in total) are considered vanished, i.e. recorded in the past and not observed for
a long time since the last record. The vast majority of taxa in this group (364) are neo-
phytes, and many occasionally recorded hybrids (75) also fall here. Typical examples
include Alhagi maurorum,Chloris virgata,Cakile maritima,Conyza triloba and Scleroblitum
atriplicinum.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 169
Introduced: 1104
Casual
surviving: 597
Casual
vanished: 250
Naturalized
non-invasive: 207
Naturalized
invasive: 50
Casual: 847 Naturalized: 257
100%
Invasion success
77% 23%
Course of invasion (time)
19%
54%
23%
4%
Fig. 2. – Transition rates in alien flora of the Czech Republic, shown for neophytes,along the introduction–natu-
ralization–invasion continuum (INIC). For each category, the number of taxa is given and the height of the bar
with the associated number indicates the percentage of the total number of 1104 neophytes recorded that reached
that stage. Casuals are divided into those that survive (70.5% ofthe total number of casuals) and that are consid-
ered vanished (29.5%), naturalized into non-invasive (80.5%) and invasive (19.5%). Invasion failures at different
stages of the INIC are represented by dotted arrows and quantified in Fig. 3.
Group 2. Establishment and failure. This group includes almost exclusively archaeo-
phytes (37 of 45 in total) that were surviving in the landscape for centuries or millennia,
formed self-sustained populations in the past, some of them might have been even invasive
at some stage, but now they have declined or are even considered vanished (22 taxa). In the
previous catalogue, they were mostly classified as naturalized, often post-invasive (Pyšek
et al. 2002); the change in classification of these taxa resulted from the focus on the current
state adopted in the present treatment and the fact that they no longer occur in populations
that can be considered self-sustaining. The group includes some red-listed archaeophytes
(e.g. Agrostemma githago,Atriplex rosea,Heliotropium europaeum,Lolium remotum
and Scandix pecten-veneris; Holub & Procházka 2000), but also neophytes (e.g. Cnidium
silaifolium and Xanthium spinosum), and refers to the invasion failure in the sense of
Blackburn et al. (2011).
(a2) Past link to cultivation
Group 3. Establishment and failure. A group of 17 taxa that are either archaeophytes or
neophytes introduced long ago, mostly in the 19th century, weresurviving due to weak but
continued propagule pressure from cultivated populations in the past but never formed
self-sustaining population in the wild. Since the planting has ceased or its intensity
strongly decreased, they are currently declining or have already vanished (13 taxa). Exam-
ples include Camelina sativa,Chenopodium foliosum,Dracocephalum moldavica,Madia
sativa,Pyrus nivalis,Stachys affinis or Trigonella foenum-graecum.
170 Preslia 84: 155–255, 2012
Geography
Captivity or
Cultivation
Survival
Reproduction
Dispersal
Environmental
Stage
Prevention
Eradication
Containment
Management Mitigation
Invasion Failure “Boom and Bust”
E:
62
(PG14+16+18)
61
(PG14+16+18)
4.2%
55
(PG6)
(PG6)
61
(PG2+3)
(PG2+3)
E:
126
(PG13+15+17)
(PG13+15+17)
C2
C1 C3 D1, D2C0
B3
B1, B2
A
Barrier
8.7%
Alien
Alien
Casual
Casual Naturalized/Established
Naturalized/Established
Terminology
Invasive
Invasive
Transport Introduction Establishment Spread
1454
taxa
1454
taxa
85
(PG8+10+12)
5.8%
141
(PG7+9+11)
9.7%
924
(PG1+4+5)
924
(PG1+4+5)
62
54
127
3.7%
4.3%
63.5%
Fig. 3. – Population groups (PG) of alien taxa in the Czech flora (see text for details and Table 1 for overview)
mapped onto the unified framework for biological invasions (Blackburn et al. 2011; the background figure
reprinted with permission from Elsevier Limited). Population groups correspondingto casual , naturalized but
not invasive , and invasive taxa are distinguished by different colours. Number of taxa and percentages of
the total of 1454 are indicated for each stage. Note that the groups do not match precisely the casual–natural-
ized–invasive areas at the top of the scheme due to distinguishing taxa that correspond to invasion boom and bust
(taxa that spread in the past, formed metapopulations but their spread ceased, therefore are at present considered
naturalized rather than invasive; PG13+15+17).
(a3) Ongoing link to cultivation
Group 4 & 5. Introduction and failure. An escape from cultivation analogous to Group 1.
Group 4 includes 501 casual taxa, mostly neophytes (458), that rely on continued input of
propagules from planted populations. Usually they are planted as garden ornamentals and
the link between planted populations and those in the wild is very close. In terms of abun-
dance, these taxa are at best scattered (339 are rare, 109 reported from a single site) and 56
are vanished. Examples include Convolvulus tricolor,Dahlia pinnata,Dasiphora fruticosa
and Ficus carica. Some woody plants that escaped from cultivation have close link with
planted populations, but have not formed (yet) long-sustaining populations due to long
generation time (e.g. Celtis occidentalis,Crataegus persimilis and Paulownia tomentosa)
or limited ability to establish permanently (e.g. Abies grandis and Platanus ×hispanica)
are included in this group. Some taxa previously classified as naturalized by Pyšek et al.
(2002) were reassigned to this group (e.g. Allium tuberosum,Helleborus viridis,
Othocallis siberica,Polygonatum latifolium and Sedum rupestre subsp. erectum), includ-
ing some shrubs surviving in single or a few locations (e.g. Alnus rugosa,Ribes odoratum
and Rubus canadensis).
Group 5 is defined based on the same principles, the difference being current rather mas-
sive propagule pressure from large-scale planting for agricultural or horticultural pur-
poses. It includes 28 taxa, with archaeophytes prevailing (21) but neophytes also repre-
sented, and examples include Allium cepa,Anethum graveolens,Helianthus annuus,
Triticum aestivum or Zea mays. There are 18 anecophytes in this group.
(b) Self-sustaining isolated populations
(b1) No link to cultivation
Group 6. Establishment and failure. This group includes 54 archaeophytes that were
introduced independently of cultivation, survived in the landscape for centuries or even
millennia and although their populations are declining, they still survive in the wild as rare
or scattered. The majority of them occur in warm regions and it is assumed that many of
them were invasive at some stage in their invasion history (classified as naturalized post-
invasive in Pyšek et al. 2002), often as weeds of arable land. Examples include Ajuga
chamaepitys subsp. chamaepitys,Anagallis foemina,Bifora radians and Ranunculus
arvensis. A subset in this group are taxa confined to habitats associated with breeding
domestic animals in villages, e.g. Chenopodium vulvaria,Lepidium coronopus,Marru-
bium peregrinum and Sclerochloa dura.
Group 7. Establishment and no trend. The group consists of 40 taxa, most of them
archaeophytes (21) but also old neophytes are represented (19), most of them introduced
in the 19th century. The taxa from this group occur mostly as scattered or rare but without
a significant trend for decline or spread. Examples include: Brachypodium rupestre,
Genista sagittalis,Crepis capillaris,Geranium molle,Papaver dubium,Pastinaca sativa
subsp. urens and Potentilla intermedia.
Group 8. Starting spread. A group comprising almost exclusively neophytes (40 of 43 in
total), mostly introduced in the 20th century, that have formed self-sustaining populations
and exhibited signs of starting spread in the last decades. The majority of them were classi-
fied as naturalized in the previous catalogue (Pyšek et al. 2002), but there are also 11 taxa
that were in the casual stage at the beginning of the 2000s and their dynamics in the last
decade justifies reassessment, e.g. Abutilon theophrasti and Senecio inaequidens. The
Pyšek et al.: Catalogue of alien plants of the Czech Republic 171
group includes also taxa that formed a small but abundant and persisting population that is
currently prevented from further spread by the barrier of unsuitable habitats (Corisper-
mum pallasii) or those that were introduced fairly recently and had not time yet to fully
manifest their invasion potential (Agrostis scabra,Dittrichia graveolens and Panicum
miliaceum subsp. agricola).
(b2) Past link to cultivation.
Group 9. Establishment and no trend. An escape from cultivation analogous to Group
7. This group includes 36 taxa, mostly neophytes (27), that form stabilized self-sustaining
populations in the wild as a result of past planting, ranging from rare to common in abun-
dance (e.g. Calystegia pulchra,Hesperis matronalis subsp. matronalis,Saxifraga hostii
subsp. hostii and Viola suavis), but also archaeophytes with the same characteristics
(Glycyrrhiza glabra,Lilium bulbiferum and Myrrhis odorata).
Group 10. Starting spread. This group includes 11 taxa, nine of them being naturalized
neophytes that exhibit signs of starting spread and are likely to become invasive in the
future, e.g. Dipsacus strigosus and Duchesnea indica. Compared to previous catalogue
(Pyšek et al. 2002), Azolla filiculoides and Bromus carinatus that were assessed as casual,
appear in this category. The group also includes two archaeophytes, Bryonia dioica and
Galega officinalis.
(b3) Ongoing link to cultivation
Group 11. Establishment and no trend. A group of 65 taxa with early introduced neo-
phytes prevailing (57 taxa, for the majority of them the first record is available from the
19th century), that occur as rare or scattered but have formed self-sustaining populations
with ongoing support of propagule pressure from cultivated populations. Examples
include Alcea rosea,Lychnis coronaria and Matteuccia struthiopteris. Compared to pre-
vious classification (Pyšek et al. 2002), 25 taxa considered as casual then are now consid-
ered to form self-sustaining populations, e.g. Arabis procurrens,Eranthis hyemalis and
Erysimum cheiri. Populations of some taxa are likely to start spread in the future, being
currently still constrained by a short residence time (e.g. Elaeagnus commutata).
Group 12. Starting spread. This group includes 31 taxa, all but one neophytes, that are
still more or less widely planted and exhibit the signs of beginning spread, e.g. Colutea
arborescens,Fallopia aubertii,Hordeum jubatum and Pinus nigra. Based on the marked
dynamics in the last decade, some of them were reclassified from the casual category in
Pyšek et al. (2002) to naturalized, e.g. Buddleja davidii (first reported to escape from culti-
vation in 2000), Aesculus hippocastanum,Symphyotrichium laeve or Sagittaria latifolia.
The group also includes several taxa formerly classified as invasive for which this classifi-
cation is not (yet) justified using the conservative approach adopted here: they are
Amorpha fruticosa,Cytisus scoparius subsp. scoparius,Galeobdolon argentatum,
Mahonia aquifolium,Physocarpus opulifolius,Rhus typhina or Sedum hispanicum.
(c) Invasive metapopulations
(c1) No link to cultivation
Group 13. Establishment and no trend. A group of 100 unintentionally introduced taxa
with occurrence stabilized during centuries or millennia of presence in the target region,
consisting mostly of archaeophytes (87 taxa). The examples include many common weeds
172 Preslia 84: 155–255, 2012
of agricultural land and ruderal taxa such as Anagallis arvensis,Anthemis arvensis,
Chenopodium strictum,Convolvulus arvensis,Euphorbia peplus,Lamium purpureum,
Lapsana communis subsp. communis,Malva neglecta and Thlaspi arvense. Majority of
taxa (68) were assumed to be post-invasive in Pyšek et al. (2002). Sixteen species previ-
ously classified as invasive were reassigned into this naturalized category, e.g. Apera
spica-venti,Atriplex oblongifolia,Bryonia alba,Epilobium adenocaulon,Matricaria
discoidea,Rumex thyrsiflorus,Tripleurospermum inodorum and Veronica persica.
Group 14. Spread. This group includes 28 taxa that became invasive following uninten-
tional introduction. Most of them are neophytes (20), e.g. Amaranthus powelii,Ambrosia
artemisiifolia,Bidens frondosus,Conyza canadensis,Cuscuta campestris,Rumex alpinus,
but invasive archaeophytes are also represented, e.g. Atriplex sagittata,Cirsium arvense,
Echinochloa crus-galli and Portulaca oleracea subsp. oleracea. Apparently, annual
weeds prevail with some exceptions such as Bunias orientalis, whereas both other invasive
groups (16 and 18) consist mainly of robust perennials and woody taxa, the differences
reflecting life histories associated with unintentional vs deliberate pathways of introduc-
tion (Pyšek et al. 2011b).
(c2) Past link to cultivation
Group 15. Establishment and no trend. Group of eight taxa, both archaeophytes (e.g.
Cymbalaria muralis and Spergula arvensis subsp. sativa) and neophytes (e.g. Acorus cal-
amus and Elodea canadensis), with the same features as Group 13 but supported in their
naturalization by past cultivation, and no longer spreading. Elodea canadensis,Mimulus
guttatus,Tanacetum vulgare and Veronica filiformis have been reclassified from invasive
status (Pyšek et al. 2002) to naturalized.
Group 16. Spread. Nine taxa that still spread and the naturalization and invasion of which
has been supported by planting that was most intensive in the past; they are all early intro-
duced neophytes classified as invasive already in the previous catalogue (Pyšek et al.
2002): Ailanthus altissima,Angelica archangelica subsp. archangelica,Echinops sphaero-
cephalus,Heracleum mantegazzianum,Impatiens glandulifera,I. parviflora,Lycium
barbarum and Telekia speciosa. The only exception is Asclepias syriaca, previously clas-
sified as naturalized; this species started to spread in the last decade, especially in southern
Moravia.
(c3) Ongoing link to cultivation
Group 17. Establishment and no trend. A group of 19 taxa, consisting of 12 archaeo-
phytes and 7 neophytes that are still commonly planted at present and form stabilized
metapopulations in the wild. Examples include Armoracia rusticana,Lolium multiflorum,
Prunus cerasus and Trifolium hybridum. Twelve taxa were classified as post-invasive by
Pyšek et al. (2002) and four considered as invasive in this source were reassessed (Digi-
talis purpurea,Melilotus albus,M. officinalis and Viola odorata) and included in this
group of naturalized taxa.
Group 18. Spread. A group of 24 invasive taxa that are currently spreading were sup-
ported by planting throughout their invasion history, including the present time. There are
only two archaeophytes, Arrhenatherum elatius and Prunus cerasifera, while the vast
majority of species in this group are neophytes that started to appear in the wild in the 19th
century. The examples include many major plant invaders in the Czech Republic such as
Pyšek et al.: Catalogue of alien plants of the Czech Republic 173
Acer negundo,Helianthus tuberosus,Lupinus polyphyllus,Pinus strobus,Prunus serotina,
Quercus rubra,Reynoutria ×bohemica,R. japonica var. japonica,Robinia pseudacacia,
Solidago canadensis and S. gigantea. All taxa in this group but Prunus cerasifera were
classified as invasive already in Pyšek et al. (2002). Although taxa confined to eutrophic
ruderal habitats generally prevail in this group, those preferring nutrient-poor soils (such
as Pinus strobus,Prunus serotina, and Quercus rubra) are also present.
Taxonomic composition
Alien taxa in the Czech flora are representatives of 586 genera and 107 families (Appendix
2). The genera richest in taxa (including hybrids and anecophytes) among all aliens are
Amaranthus (24 taxa), Oenothera (23) and Trifolium (19) but there are marked differences
between neophytes and archaeophytes in this respect: Oenothera,Amaranthus,Trifolium,
Rumex,Solanum,Rubus and Centaurea are most represented genera among neophytes,
whereas Vicia,Prunus,Veronica,Atriplex,Bromus,Viola and Chenopodium among
archaeophytes (Table 4).
Overall, neophytes belong to 508 and archaeophytes to 184 genera; exclusively
‘archaeophytic genera’ (with only archaeophytes among their alien taxa) that include at
least three alien representatives are Arctium (7 taxa), Spergula (4), Anthriscus,Marrubium,
Myosotis,Polycnemum,Pyrus,Sonchus and Valerianella (3).
Families most represented in alien flora (Table 5) are Asteraceae (198 taxa; 13.6% of the
alien flora), Poaceae (152; 10.5%) and Brassicaceae (101; 6.3%); apart from minor changes
in the numbers of taxa resulting from the above described additions and deletions, the pat-
tern of richness at the level of most represented families is the same as reported in detail in
Pyšek et al. (2002). Some major changes in the richness of families in the current treatment,
compared to Pyšek et al. (2012; e.g. Amaranthaceae 76 vs 25 taxa, Scrophulariaceae 5vs
39), are attributed to the different classification system used here (Stevens 2001 onwards,
The Angiosperm Phylogeny Group 2009). All but one (Linaceae) of the total number of 107
families included contain at least one neophyte representative, while archaeophytes origi-
nate from only 42 families. The families richest in neophytes are Asteraceae,Poaceae,
Rosaceae,Fabaceae and Brassicaceae (Table 5), which together contain 485 taxa and
account for 43.9% of all neophytes. Asteraceae,Poaceae and Brassicaceae also rank high
among archaeophytes, but there are also other families that are rich in archaeophytes (e.g.
Apiaceae,Caryophyllaceae,Plantaginaceae and Boraginaceae;Table5).
Temporal trends and pathways of introduction
The data on the first record in the studied region, known for 771 neophytes, allow to recon-
struct the increase in the number of taxa introduced into the Czech Republic over the last
three centuries, although it is clear that the reliability of data on residence times decreases
towards the past (Lambdon et al. 2008). The numbers of new taxa recorded in particular
years reflect peaks associated with specific events such as the increased interest in plants
of human-made habitats in the 1970s, linked to the establishment of a working group at the
Institute of Botany (Hejný etal. 1973, Pyšek 2001, Pyšek et al. 2003, 2011b), or the publi-
cation of the first catalogue of Czech alien plants (Pyšek et al. 2002). However, when the
cumulative number of the first species records is plotted against time, the trend suggests
a rather steady increase of four alien arrivals per year since the beginning of the 19th century
174 Preslia 84: 155–255, 2012
without any distinct decelerating trend and a projected total number of 1264 taxa in the
year 2050. Fifty per cent of the present known taxa were recorded up to 1935, 60% up to
1957, 70% up to 1963, 80% up to 1973, and 90% up to 1997 (Fig. 4). This indicates that
the number of alien taxa recorded in the Czech Republic will be increasing at a similar rate
in the near future, corresponding to a trend reported for Europe (Hulme et al. 2009) and
creating an invasion debt (Essl et al. 2011).
As to the pathways of introduction into the country, deliberate introduction was
involved in 747 of the 1454 taxa (51.4%). Most deliberate introductions resulted from
ornamental or horticultural plantings (see Pyšek et al. 2002 for detailed analyses of plant-
ing purposes). The remaining 48.6% of taxa are assumed to have arrived by unintentional
pathways, i.e. mostly as contaminants of commodities or stowaways (Hulme et al. 2008,
Pyšek et al. 2011b). The ratio of deliberate and unintentional introduction is reversed in
archaeophytes and neophytes, with 30.7% of the total number of taxa deliberately intro-
duced among the former and 57.9% among the latter.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 175
Table4. – Genera with the highest diversity of alien taxa in the Czech flora, cross-tabulated according to immigra-
tion time and invasion status. The 23 genera represented by at least 10 alien taxa are shown. Other taxon-rich gen-
era include Avena,Cirsium,Hordeum,Malva,Papaver,Setaria,Silene,Sisymbrium,Symphyotrichum (8 alien
taxa), Brassica,Camelina and Fumaria (7 alien taxa). Hybrids are included. Cas – casual; natur – naturalized
non-invasive; inv – invasive.
Archaeophytes Neophytes Total
Genus cas natur inv cas natur inv archaeophytes neophytes all aliens
Amaranthus 11 1642 2 22 24
Oenothera 16 7 0 23 23
Trifolium 16 3 0 19 19
Chenopodium 15 92 6 11 17
Rumex 11 3 3 0 17 17
Viola 43 82 7 10 17
Bromus 25 81 7 9 16
Solanum 1141 11516
Centaurea 1 1 11 2 2 13 15
Vicia 26 61 8 7 15
Rubus 95 0 14 14
Allium 31 81 4 9 13
Artemisia 12 73 3 10 13
Euphorbia 49 4913
Epilobium 11 1 0 12 12
Geranium 453 4812
Lepidium 462 4812
Veronica 732 7512
Atriplex 331 4 7 4 11
Prunus 251 2 1 8 3 11
Eragrostis 181 1 9 10
Lathyrus 11 62 2 8 10
Sedum 64 0 10 10
Life histories and regions of origin
Among all aliens, 43.3% are annuals, 33.1% perennials, 10.8% biennials, 8.5% shrubs or
semishrubs, and 4.3% trees. Archaeophytes and neophytes demonstrate a highly signifi-
cant difference in the distribution of life histories: the former are more often annuals
(56.4% vs 38.8% among neophytes) or biennials (17.0% vs 8.6%) and less often
perennials (18.2% vs 38.3%) or shrubs and trees (8.5% vs 14.3%; Fig. 5).
The main donors of alien plants to the Czech Republic are the Mediterranean region
(34.6%), other parts of Europe (19.4%), other parts of Asia (13.1%) and North America
(12.6%). The contribution of other regions (Central America, South America, Africa,
Australia) does not exceed 4%. The region of origin could not be assigned for 199 taxa,
a group consisting of 105 anecophytes and 94 taxa of hybrid origin (Fig. 6). The data on
origins confirm the well-known difference between archaeophytes and neophytes in terms
176 Preslia 84: 155–255, 2012
Table 5.– Families with the highest diversity of alien taxa in the Czech flora, cross-tabulated according to immi-
gration time and invasion status. The 29 families represented by at least 10 alien taxa are shown. Hybrids are
included. Cas – casual; natur – naturalized but non-invasive; inv – invasive. The classification of families follows
that of Angiosperm Phylogeny Group: APG III (Stevens 2001 onwards, Angiosperm Phylogeny Group 2009).
Archaeophytes Neophytes Total
Family cas natur inv cas natur inv archaeophytes neophytes all aliens
Asteraceae 18 26 1 114 22 17 45 153 198
Poaceae 14 20 4 99 15 38 114 152
Brassicaceae 10 22 50 17 2 32 69 101
Rosaceae 7 10 1 54 19 1 18 74 92
Fabaceae 5 11 58 15 2 16 75 91
Amaranthaceae 9 11 1 42 8 5 21 55 76
Lamiaceae 12 9 30 9 21 39 60
Apiaceae 14 6 2 18 2 1 22 21 43
Onagraceae 29 8 0 37 37
Solanaceae 3 30 3 1 3 34 37
Caryophyllaceae 6 7 1 14 5 14 19 33
Plantaginaceae 2 12 12 6 14 18 32
Polygonaceae 21 1856 3 29 32
Boraginaceae 47 172 111930
Papaveraceae 3 11 10 2 14 12 26
Ranunculaceae 2 3 15 4 5 19 24
Malvaceae 3 4 13 2 7 15 22
Geraniaceae 5 9 3 5 12 17
Violaceae 43 82 7 10 17
Amaryllidaceae 31 92 4 11 15
Asparagaceae 1122 11415
Euphorbiaceae 5 10 5 10 15
Crassulaceae 95 0 14 14
Cucurbitaceae 32 411 5 6 11
Orobanchaceae 12 71 3 8 11
Saxifragaceae 10 1 0 11 11
Campanulaceae 91 0 10 10
Iridaceae 271 2810
Rubiaceae 13 6 4 6 10
Pyšek et al.: Catalogue of alien plants of the Czech Republic 177
0
200
400
600
800
1000
1200
1650 1700 1750 1800 1850 1900 1950 2000
neophytes
2050
0
10
20
30
40
50
60
70
80
90
100
Cumulative number of taxa
Estimated total number of taxa
Number of taxa
Number of reported in the year
Cumulative number of neophytes
Fig. 4. Temporal trends in the alien flora of the Czech Republic in the last 200 years based on neophytes with
known year of the first report (n = 771). Also shown is extrapolatedtrend for the total number of taxa (n = 1104),
and numbers of taxa reported in particular years (right axis).
16
(19.2)
20
(36.4)
**
77
(140.9)
***
72
(43.6)
***
239
(180.7)
***
55
(55.7)
119
(105.5)
.
466
(408.1)
**
105
(126.4)
*
472
(523.3)
**
0
10
20
30
40
50
60
Annual Biennial Perennial Shrub Tree
Archaeophyte
Neophyte
Percentage of the total number of taxa
Fig. 5. – Representation of life histories among alien taxa in the Czech Republic. Taxa with multiple life histories were
considered in each category so the sum of the numbers of taxa (shown on top of the bars) does not match the total num-
bers of archaeophytes and neophytes. Overall, the observed counts of alien taxa highly significantly (χ2= 94.25; df = 4;
P < 0.0001) differ from counts expected by chance (values in parentheses). Statistically significant deviations of indi-
vidual counts from counts that can be expected by chance are expressed by the number of asterisks (*** P < 0.001; ** P
< 0.01; * P < 0.05) and marginal significance by a dot (. P < 0.1); numbers in parentheses not followed by any symbol
do not differ from randomly expected values. Semishrubs are included within shrubs. Excluded from these statistics
are 4 ferns (all neophytes), 11 aquatic species (all neophytes) and 11 parasitic species (3 archaeophytes, 8 neophytes).
of source regions (e.g. Pyšek et al.2002, 2004b, 2005, Chytrý et al. 2005, 2008a, b): more
than a half (52.7%) of archaeophytes originate from the Mediterranean region (the figure
increases to 64.5% if anecophytes and hybrids are excluded), which is, however, also the
most frequent donor of neophytes (28.7%). The contribution of other parts of Europe and
Asia to the total number of taxa is slightly higher for neophytes than for archaeophytes,
19.9% vs 17.8% and 14.2% vs 10.1%, respectively (Fig. 7).
Since archaeophytes, by definition, have not arrived from overseas, it is plausible to
compare their regions of origins with those of neophytes if Americas and Australia are
excluded. The difference between archaeophytes and neophytes in such a comparison is
still statistically highly significant (χ2= 45.057; df = 3; P < 0.0001). Highly significantly
(P < 0.001) more archaeophytes originated in the Mediterranean region (231 vs 180.5
expected counts), but highly significantly less (P < 0.01) in the other parts of Asia (44 vs
67.9), significantly (P < 0.05) less in the other parts of Europe (78 vs 100.1) and margin-
ally significantly less (P < 0.1) in Africa (5 vs 9.6). Conversely, neophytes originated in the
Mediterranean region were significantly less represented (385 vs 436.5) and those from
the other parts of Asia marginally significantly more represented (190 vs 164.1).
178 Preslia 84: 155–255, 2012
Europe; 345
Mediterranean region;
616
Africa; 34
Asia; 234
North America; 224
Central America; 42
South America; 67
Australia; 19
hybrids; 94
anecophytes; 105
Fig. 6. – Proportional contribution of the world regions to the alien flora of the Czech Republic. Region names are
followed by numbers of taxa native to that region. Note that native distribution regions extend over more than one
area, therefore the sum of taxon numbers exceeds the total of 1454 recorded in the presentstudy. Europe, Asia and
Africa refer to parts of these continents outside the Mediterranean region. Taxa originated through hybridization
and anecophytes are shown separately.
Regional abundance, habitats and cover in plant communities
Archaeophytes are generally more abundant in the field, which reflects that they were pro-
vided with more time in the target region (Pyšek et al. 2002, 2004b, 2011a). Of the total
number of archaeophytes, 22.0% are considered common (highly significantly more than
expected by chance), 2.9% locally abundant and 28.5% scattered (highly significantly more
than expected by chance). This pattern strikingly contrasts with that found for neophytes.
Only 2.9% of neophytes (35 taxa) are classified as common (highly significantly less than
expected by chance) and 3.0% locally abundant, 8.1% scattered (highly significantly less
than expected) while as many as 86.0% occur in low-abundance categories (rare, single
locality or vanished; with the last two categories occurring highly significantly or signifi-
cantly, respectively, more often than expected by chance); the corresponding figure for
archaeophytes being 46.6%, with these categories significantly or highly significantly
underrepresented. Two hundred and twelve neophytes (17.7%) are only known from a sin-
gle locality (compared to only five archaeophyte hybrids;Appendix 2) and 250 (22.6%) are
labelled as vanished (compared to only 27 archaeophytes, i.e. 7.7%) (Fig. 8).
The contrasting patterns in the occurrence of both immigration status groups,
archaeophytes and neophytes, translate into those of the breadth of their habitat niches,
expressed as the number of habitats of the total of 88, occupied by 497 taxa that could be
classified according to their habitat affinities (Sádlo et al. 2007). Archaeophytes occupy
on average more habitats (9.5±9.0, mean±S.D., n = 244) than neophytes (6.4±6.1,
n = 253), and 31.6% of them occur in more than 10 habitats (compared to only 17.8% of
Pyšek et al.: Catalogue of alien plants of the Czech Republic 179
0
10
20
30
40
50
60
Europe
Medit erranean
region
Afri ca
Asia
North America
Central Americ a
South Ameri ca
Austral ia
hybri d
anecophyt es
Percentage of the total number of taxa
Archaeophytes
Neophytes
Fig. 7. – Distribution of archaeophytes and neophytes in the Czech Republic according to their origin. Taxa origi-
nating from multiple regions as designated here are included in each region. See text for the results of statistical
analysis.
neophytes; Fig. 9). Ten archaeophytes and only three neophytes (Conyza canadensis,
Epilobium adenocaulon and Impatiens parviflora) grow in a wide range of habitats
exceeding 30 (see Sádlo et al. 2007: their Table 2). The species with the broadest habitat
niche of all alien taxa in the Czech Republic is an archaeophyte, Arrhenatherum elatius,
occurring in 62 of 88 habitats (see Appendix 1 for comments on its classification).
The covers that alien taxa reach in plant communities in the Czech Republic yield
a completely opposite picture of neophyte vs archaeophyte comparison (Fig. 10). Neo-
phytes are shifted towards high-cover categories, reaching on average 8.5% cover (n = 48),
markedly more than archaeophytes (4.7%, n = 131). The first five taxa with highest aver-
age covers are all neophytes: Acorus calamus 39% (recorded in n = 293 vegetation plots),
Elodea canadensis 35% (n = 412), Helianthus tuberosus 26% (n = 62), Heracleum
mantegazzianum 26% (n = 27) and Reynoutria japonica var. japonica lumped with Rbohe-
mica 26% (n = 51). Other neophytes with a high cover are Impatiens glandulifera (18%, n
= 302), Solidago gigantea (17%, n = 99), Echinocystis lobata (14%, n = 33) and Pinus
nigra (13%, n = 33).
180 Preslia 84: 155–255, 2012
78
(25.6)
***
10
(10.4)
101
(44.8)
***
133
(158.3)
*
5
(49.1)
***
27
(62.7)
***
35
(87.4)
***
36
(35.6)
97
(153.2)
***
566
(540.7)
212
(167.9)
***
250
(214.3)
*
0
5
10
15
20
25
30
35
40
45
50
Common Locally
abundant
Scattered Rare Single Vanished
Archaeophytes
Neophytes
Percentage of the total number of taxa
Fig. 8. – Distribution of alien taxa in the Czech Republic in abundance categories. The sum of the numbers of
taxa, shown on top of the bars, exceeds the total numbers of archaeophytes andneophytes as some taxa occurred
in a single location and disappeared; they are included in both ‘single’ and ‘vanished’ categories. Overall, the
observed counts of alien taxa highly significantly (χ2= 312.392;df = 5; P < 0.0001) differ from counts expected
by chance (values in parentheses). Statistically significant deviations of individualcounts from counts that can be
expected by chance are expressed by the number of asterisks (*** P < 0.001; * P < 0.05); numbers in parentheses
not followed by any symbol do not differ from randomly expected values.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 181
0
50
100
150
200
250
<10 11–20 21–30 31–40 >40
Number of habitats
Archaeophytes
Neophytes
Number of taxa
Fig. 9. – Frequency distribution of the numbers of habitats (n = 88) in which alien taxa are recorded, shown sepa-
rately for archaeophytes (n = 244) and neophytes (n = 253).
0
20
40
60
80
100
<10 10–20 20–30 30–40
Average cover in vegetation plots (%)
Archaeophytes
Neophytes
117
34
12
9
2
32
Percentage of the total number of taxa
Fig. 10. – Frequency distribution of coversof alien taxa in plant communities in the Czech Republic. Only taxa for
which data from at least 25 plots are available were included. Numbers of taxa in each cover class are shown on
top of the bars.
Although this comparison must be taken with caution because the vegetation plots were
sampled in a subjective, preferential way, average plot sizes for individual taxa differ and
there is also great variation in the number of plots from which the data are derived, the dif-
ferences between the two groups of aliens are robust enough to indicate that neophytes are
on average more successful in colonizing plant communities and often forming monodo-
minant stands (see also Chytrý et al. 2008a).
Impact
A thorough assessment of impacts of plant invasions inthe Czech Republic is still missing
which reflects the fact that studies summarizing information on impacts across alien floras
of large regions are still rare despite intensive research in the last few years (Parker 1999,
Gaertner et al. 2009, Pyšek & Richardson 2010, Vilàet al. 2010, 2011, Winter et al. 2009,
Pyšek et al. 2012). Based on data on impacts of alien plants in Europe summarized by the
DAISIE project (DAISIE 2009, www.europe-aliens.org), there are 133 taxa on the list of
Czech alien plants that were documented in the literature to exert ecological impacts
and/or economic impacts in some parts of Europe (Appendix 2), some of them also in the
Czech Republic (Hejda et al. 2009). These data make it possible to highlight taxa that
already impose ecological impacts but also those that can become threat in the future.
The group of taxa with documented ecological impacts covers 33 taxa that are classi-
fied as invasive in the present study, and includes most of the major invaders in the Czech
Republic, some of them threatening seminatural habitats (e.g. Acer negundo,Ailanthus
altissima,Helianthus tuberosus,Heracleum mantegazzianum,Impatiens glandulifera,
Impatiens parviflora,Lupinus polyphyllus,Lycium barbarum,Pinus strobus,Prunus
serotina,Reynoutria japonica var. japonica,R. sachalinensis,R.×bohemica,Robinia
pseudoacacia,Rudbeckia laciniata,Solidago canadensis and S. gigantea) but also nox-
ious weeds of arable land (e.g. Amaranthus retroflexus and Galinsoga parviflora) or spe-
cies affecting human health (Ambrosia artemisiifolia). Besides these taxa, already exert-
ing impacts in the Czech Republic, the 113 taxa with ecological impacts in Europe include
45 that we currently classify as naturalized; some of them belong to population groups that
exhibit symptoms of starting spread and their impact in the near future is likely (e.g.
Abutilon theophrasti,Lepidium virginicum and Senecio inaequidens). Finally, for 35 taxa
that occur as casual in the Czech Republic ecological or economic impact is documented
from elsewhere in Europe; this group includes some noxious invaders (e.g. Elodea
nuttalii,Rosa rugosa and Solidago graminifolia) that should be monitored to enable early
action should their population dynamics change (Appendix 2).
Notes on the classification of taxa
The present update of the 10 years old data yielded a number of changes to the taxa listed,
and their invasion and residence time statuses. These changes are due to several reasons.
First, they reflect the real changes in species’ behaviour and their invasion dynamics over
the last decade. Second, the interest in and knowledge of alien plants has improved consid-
erably as a result of intensive research in biological invasions in the Czech Republic dur-
ing this period. Third, the more conservative approach towards what should be considered
native or alien also brought about changes in the species list, and finally, introducing the
182 Preslia 84: 155–255, 2012
population-based approach to the classification of taxa adopted here (Blackburn et al.
2011) resulted in shifts in invasion status.
The main change in approach relative to the previously used scheme concerns a strict
focus on the current state of a taxon’s populations in a region. This allowed us to take into
account and quantify categories that refer to unsuccessful invasions – the ‘invasion failure’
and ‘boom and bust’ phenomena as defined by Blackburn et al. (2011). This is reflected
namely in classifying taxa that formed self-sustained populations in the past, some
assumed to have been invasive (and labelled post-invasive in Pyšek et al. 2002), as casual,
suggesting the reversed trajectory along the INIC (Fig. 2). Although they would not be
classified as casuals, should the criterion of relying on repeated introduction of
propagules, which is part of the traditionally accepted definition, be strictly followed
(Richardson et al. 2000, 2011), we believe that the criterion of population self-
sustainability is a more important one, reflecting closely the population dynamics in both
directions along the INIC. This approach is further supported by the fact that many of
these taxa are red-listed or missing for a long time, which strongly argues against self-
sustainability of their populations. This group includes also many archaeophytes that have
never been planted indicating that their occasional occurrence is due to long-term survival
in and occasional germination from seed banks.
Consequently, the number of invasive taxa is substantially smaller than in the previous
catalogue (50 neophytes and 11 archaeophytes in the present study compared to 69 neo-
phytes and 21 archaeophytes, respectively, in Pyšek et al. 2002). A decrease this dramatic
is due to the newly adopted conservative approach; unlike in the previous account, the
emphasis here was on ongoing spread as a major criterion. The lower numbers do not
mean that the problems with invasive plants in the Czech Republic are diminishing; rather
the opposite is true as indicated by species that started to spread recently. In conclusion, we
believe that the more rigorous approach to separating invasive species from naturalized
makes the current assessment of species status more comparable with other parts of the
world, especially those that experience serious problems with invasions, and forms
a sounder basis for managing plant invasions at the national scale.
See http://www.preslia.cz for Electronic Appendices 1,2
Acknowledgments
The work was supported by grant no. P504/11/1028 (Czech Science Foundation), long-term research develop-
ment project no. RVO 67985939 (Academy of Sciences of the Czech Republic) and institutional resources of
Ministry of Education, Youth and Sports of the Czech Republic. P.P. acknowledges the support by the Praemium
Academiae award from the Academy of Sciences of the Czech Republic. We thank Vladimír Řehořek and Laura
Meyerson for their comments that greatly improved the manuscript. We thank following colleagues for consulta-
tions and/or providing us with unpublished data, and permission to use them: Jiří Burda, Věra Čulíková, Vít
Grulich, Jiří Hadinec, Rudolf Hlaváček, Jan W. Jongepier, Petr Kočár, Petr Koutecký, Radka Kozáková, Petr
Krása, Martin Křivánek, Karel Kubát, Martin Lepší, Petr Petřík, Petr Pokorný, Uwe Raabe, Olga Rotreklová,
Jaroslav Rydlo, Lukáš Sekerka, Pavel Sekerka, Ota Šída, Milan Štech, Jan Štěpánek, Bohumil Trávníček, Jiří
Uher, Adam Veleba, Václav Větvička and Jiří Zázvorka. Laura Meyerson kindly improved our English. Technical
help from Zuzana Sixtová is acknowledged.
Pyšek et al.: Catalogue of alien plants of the Czech Republic 183
Souhrn
Práce přináší úplný seznam nepůvodních taxonů zaznamenaných na území České republiky; je aktualizací a dopl-
něním předchozího seznamu publikovaného v roce 2002. Zahrnuje nové údaje shromážděné za poslední desetiletí
a přehodnocuje zařazení a status některých druhů, vyplývající z rozvoje taxonomického poznání. Nepůvodní
flóra České republiky zahrnuje 1454 taxonů, které jsou uvedeny v Apendixu 2 s informacemi o taxonomické pří-
slušnosti, životní formě, oblasti původu, invazním statusu (zda jde o druh přechodně zavlečený, naturalizovaný
avšak neinvazní,nebo invazní), charakteru výskytu v krajině, době zavlečení (archeofyt nebo neofyt), způsobu in-
trodukce do země a u neofytů o datu prvního nálezu. Oproti původnímukatalogu je uveden počet typů biotopů, ve
kterých se druh vyskytuje, pokryvnost v rostlinných společenstvech a impakt. Podíl zavlečených druhů v české
flóře je značný: tvoří jej 350 (24,1%) archeofytů a 1104 (75.9%) neofytů. Nárůst počtu taxonů oproti původnímu
katalogu, který uváděl 1378 taxonů,vyplývá z toho, že bylo přidáno 151 taxonů. Celkem 75 (39 archeofytů a 36
neofytů) bylo naproti tomu vypuštěno; značná část tohoto počtu jde na vrub přeřazení 41 taxonů mezi původní
druhy, a to vesměs na základě archeobotanických dokladů. Přírůstky na seznamupředstavují taxony nově objeve-
né a uvedené v botanické literatuře od roku 2002,taxony zařazené na základě excerpce dříve opominutých zdrojů
či revize zdrojů použitých, nebo přehodnocenístatusu některých taxonů tradičně považovaných za původní. V ně-
kterých případech jde o infraspecifické taxony, které nebyly dříve v české flóře rozeznávány. Seznam obsahuje 44
taxonů, které jsou uváděny pro Českou republikupoprvé jako zavlečené, nebo proněž je podán první důkaz o je-
jich zplaňování: Abies concolor,A. grandis,A. nordmanniana,Avena sterilis subsp. ludoviciana,A. ×vilis,Berbe-
ris julianae,B. thunbergii,Bidens ferulifolius,Buddleja alternifolia,Buglossoides incrassata subsp. splitgerberi,
Buxus sempervirens,Corispermum declinatum,Cotoneaster dielsianus,C. divaricatus,Euphorbia myrsinites,
Gleditsia triacanthos,Helleborus orientalis,Hieracium heldreichii,Koelreuteria paniculata,Lonicera pericly-
menum,Lotus ornithopodioides,Malus baccata,M. pumila,Miscanthus sacchariflorus,Morus alba,Muscariar-
meniacum,Paeonia lactiflora,Pennisetum alopecuroides,Pinguicula crystallina subsp. hirtiflora,P. grandiflo-
ra subsp. rosea,Podophyllum hexandrum,Pyracantha coccinea,Rhodotypos scandens,Rumex patientia ×R. ti-
anschanicus ‘Uteuša’, Salix cordata,Sarracenia purpurea,Sasa palmata ‘Nebulosa’, Scolymus maculatus,Spi-
raea japonica,Tagetes tenuifolia,Thuja occidentalis,Vaccinium corymbosum aViburnum rhytidophyllum.Ko-
mentáře ke všem přidaným nebo vypuštěným taxonům jsou uvedeny v Appendixu 1. Z celkového počtu 1454 ta-
xonů je jich 985 klasifikováno jako přechodně zavlečené, 408 jako naturalizované a 61 jako invazní. Úbytek in-
vazních taxonů oproti původnímu katalogu je důsledkem konzervativnějšího přístupu: za invazní jsou považová-
ny pouze ty taxony, které se v současnosti šíří. Mezi neofyty převládají přechodně zavlečené taxony (76,7 % ze
všech neofytů, ale jen 39,4 % archeofytů), mezi archeofyty naturalizované (57,4 % versus 18,8 % neofytů).Po-
kud jde o podíl invazních druhů,není mezi oběma skupinami statisticky průkazný rozdíl. Z celkového počtu 1104
neofytů jich 250 vymizelo (byly pozorovány pouze jednou nebo několikrát a z lokalit vymizely nebo nebyly za-
znamenány po dlouhou dobu); 23,3 % jich zdomácnělo a 4,5 % se stalo invazními. Vedle tradiční klasifikace po-
stavení druhu v invazním procesu byly taxony klasifikovány do 18 populačních skupin, definovaných na základě
dlouhodobých trendů v metapopulační dynamice, současného stavu populace na území ČR a přísunu diaspor
z kultury. Tato podrobná klasifikace umožnila kvantifikovat, v jaké fázi invazního procesu dochází ke „ztrátám“
a jak jsou tyto ztráty velké. Podle toho, zda zahrneme dosrovnání zavlečené a původníflóry specifické kategorie
taxonů (vymizelé a vyhynulé, křížence), tvoří nepůvodní taxony 29,7–33,1 % z celkové flóry. Pokud vyčíslíme
podíl pouze pro zdomácnělé, tedy trvale přítomné složky zavlečené flóry, dospějeme k 14,4–17,5 %. Analýza
roků prvního nálezu, který je k dispozici pro 771 neofytů, ukazuje, že nepůvodní druhy přibývají v květeně České
republiky stálým tempem; extrapolujeme-li tato data na všechy neofyty, lze předpovědět, že do roku 2050 by je-
jich počet měl vzrůst na 1264. Přes polovinu taxonů (747, tj. 51,4 %) bylo zavlečeno alespoň zčásti prostřednic-
tvím kultury, zbývajících 48,6 % neúmyslně. Archeofyty jsou obecně v krajině hojnější a obsazují širší spektrum
stanovišť než neofyty; ty naopak dosahují v průměru většípokryvnosti v invadovanýchspolečenstvech. Práce dále
analyzuje složení nepůvodní flóry z hlediska příslušnosti k rodům a čeledím, oblasti původu a životní formy.
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Revision received 8 June 2012
Accepted 11 June 2012
Pyšek et al.: Catalogue of alien plants of the Czech Republic 201
Appendix 1. – Comments on taxa that represent changes against the previous Catalogue of alien plants of the
Czech Republic (Pyšek et al. 2002).
Changes of names, difficult cases and corrections of earlier misidentifications
Compared to the previous version of the catalogue (Pyšek et al. 2002), 124 names were changed due to nomencla-
tural reasons or development in taxonomic opinion (Electronic Appendix 1). Additional seven taxa are listed
under a different name due to the reidentification; their names refer to the same taxa which were erroneously
determined in 2002 or their taxonomic classification has changed. These taxa are commented below and represent
additions to the alien flora of the country.
Azolla filiculoides was listed as A. caroliniana in Pyšek et al. (2002), based on treatment in the Flora of the
Czech Republic (Křísa in Hejný & Slavík 1988). The taxonomy of the New World Azolla has been controversial
for a long time. The number of distinguished species varied and differentcharacters were used for their identifica-
tion. However, Evrard & Van Hove (2004) in their recent thorough investigation based on morphological, molec-
ular and physiological data concluded that only two species can be distinguished taxonomicallyin America. They
revealed that the type specimen of A. caroliniana belongs to the species described earlier as A. filiculoides, and
the fern usually identified as A. caroliniana by many authors should be correctly named A. cristata. Although
both species were recorded as introduced in Europe, only A. filiculoides is widespread, whereas A. cristata was
apparently documented only from the Netherlands. Plants recently collected in the Czech Republic are identical
with A. filiculoides (coll. and det. Z. Kaplan, PRA, rev. C. Van Hove). The other species, A. cristata (A. caroliniana
auct.), has apparently never occurred in the country as introduced or escaped.
Corispermum pallasii was listed in Pyšek et al. (2002) as C. leptopterum. However, recent taxonomic studies
revealed that the European plants are conspecific with the Siberian ones, described much earlier as C. pallasii
(Mosyakin 2003). Vymyslický & Grulich (2004), reporting on their find of Corispermum from Ivančice, distr.
Brno, suggested that southern Moravian plants correspond to C. canescens, which is native to Hungary. However,
based on a careful re-examination of specimens from BRNU and PR (J. Danihelka), we believe that all
Corispermum specimens so far collected in the Czech Republic, with the only exception of C. declinatum (see
below), most likely belong to C.pallasii. The earliest documented record of this species is from 1933 (ex herb.
F. Hrobař, PR). At present, C. pallasii occurs in two populations consisting of thousands of plants in sand pits near
Bzenec, southern Moravia, from where it is spread with traded sand to other places.
Eriochloa punctata. Three Eriochloa species were reported in the literature from the Czech Republic:
E. ramosa from a wool-processing factory Mosilana in Brno (Dvořák & Kühn 1966, Grüll 1979) and E. punctata
from railway station in Brno (Grüll 1979); these two species are given in the Flora of Dostál (1989), who in addi-
tion lists E. procera, all as casual wool aliens introduced to Brno. Actually, the names E. procera and E. ramosa
refer to the same taxon, with the former accepted as its correct name (Zuloaga & Morrone 2003, Shouliang &
Phillips 2006). The plant reported as E. ramosa by Dvořák & Kühn (1966), collected by F. Kühn in 1960, was
deposited in BRNU in 1972 under the name E. punctata; obviously, J. Dvořák re-determined the plant before
depositing it in the herbarium. Comparison of the specimen collected by F. Kühn in 1960 and another specimen
collected by F. Grüll in 1965 (reported by Grüll 1979)has shown that both of them very likely represent the same
species, most probably E. punctata, as already suggested by J. Dvořák (rev. J. Danihelka). Consequently, the spe-
cies listed as E. procera in Pyšek et al. (2002) is in fact E. punctata, the same as found by Grüll (1979).
Gilia achilleifolia. Another species of the genus, G. multicaulis, is listed in Pyšek et al. (2002), based on
a note in the Flora of the Czech Republic (Křísa in Slavík 2000) that it is planted and rarely escapes from cultiva-
tion, without further details. In 2005, two flowering plants of G. achilleifolia were reported growing in the
Stárkovský les forest near Lanžhot, southern Moravia, on a forest clearing along a road, together with Legousia
pentagonia. They were probably introduced to the site with forestry v