291
Abstract
Cyperus esculentus (yellow nutsedge) is an alien species that started to spread in 
arable fields in Slovenia and has already become a noxious weed. In this study we 
present the spatio-temporal pattern of this invasion in Slovenia. Species was first 
recorded in 1980 and currently two hotspots are evident (Ljubljana Basin and 
Posočje). Habitat preferences of Cyperus esculentus and the floristic compostion 
of invaded plant communities were studied. These communites were compared 
to similar communities in Slovenia and to so far described vegetation types with 
dominating Cyperus esculentus in Europe. Based on these analyses we described 
new weed association, namely Digitario sanguinalis-Cyperetum esculenti. 
Izvleček
Cyperus esculentus (užitna ostrica) je tuejrodna vrsta, ki se v Sloveniji širi 
na obdelanih tleh in je že postala škodljiv plevel. V raziskavi predstavljamo 
prostorsko-časovni vzorec razširjanja v Sloveniji. Vrsto so prvič opazili leta 1980 
in trenutno sta očitni dve vroči točki pojavljana (Ljubljanska kotlina in Posočje). 
Preučili smo rastiščne značilnosti in vrstno sestavo rastlinskih združb, v katerih 
se pojavlja vrsta Cyperus esculentus. Te združbe smo primerjali s podobnimi v 
Sloveniji in z dosedaj opisanimi vegetacijskimi tipi, v katerih prevladuje Cyperus 
esculentus v Evropi. Na osnovi teh analiz smo opisali novo plevelno asociacijo 
Digitario sanguinalis-Cyperetum esculenti.
Key words: alien plant, agriculture, 
plant communities, yellow nutsedge, 
vegetation.
Ključne besede: tujerodna rastlina, 
kmetijstvo, rastlinske združbe, 
užitna ostrica, vegetacija.
Received: 7. 4. 2020
Revision received: 3. 1. 2021
Accepted: 29. 1. 2021
1 ZRC SAZU, Institute of Biology, Novi trg 2, 1000 Ljubljana, Slovenia. E-mail: branko.vres@zrc-sazu.si, urban.silc@zrc-sazu.si
*
 
Corresponding author
Branko Vreš1

 & Urban Šilc1
,
*

DOI: 10.2478/hacq-2021-0012 20/2 • 2021, 291–302
Invasion, distribution and habitat 
affiliation of Cyperus esculentus,  
a new weed in Slovenia

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Branko Vreš & Urban Šilc
Invasion, distribution and habitat affiliation of Cyperus esculentus, a new weed 
in Slovenia
Introduction
Many alien plants are expanding their areal and are in-
vading arable land (Weber & Gut 2005). Alien plants are 
particularly successful and several have already become 
invasive and noxious weeds with significant ecological 
impact and economic damage (Pimentel et al. 2001).
For understanding the invasion pattern of an invasive 
alien plants it is important to know the spatio-temporal 
history of their invasion and which habitats are predomi-
nately colonized. Successful invasive alien plants can be 
part of many plant communities and can even change 
their habitat affiliation over time (Essl et al. 2009). Knowl-
edge of these invasion processes is particularly important 
for preventing current spread of invasive alien plants and 
to develop strategies to predict and stop future invasions. 
Recently early warning systems were established to detect 
the occurrence of potential invasive alien species (Sloveni-
an forestry institute 2016–2020) and the reconstruction 
of the historical spread of alien species can be a valuable 
assistance to such efforts (Essl et al. 2009). 
Cyperus esculentus or yellow nutsedge, a species from 
the Cyperaceae family, is an invasive alien plant in most 
parts of Europe. It is a perennial weed that reproduces 
by seeds and underground tubers. Its origin is uncertain, 
but it originates most likely from the Mediterranean and 
Southwest Asia. Today, it is widely distributed in western 
and southern Europe as well as in parts of Central Europe 
(Follak et al. 2016). Cyperus esculentus has C4 photosyn-
thetic pathway (Li et al. 1999) and this among other traits 
(high tuber production, allelopathy, vegetative reproduc-
tion) contributes to its competitiveness (Brown 1999, 
Follak et al. 2016).
Cyperus esculentus can cause up to 80% loss of yield 
in field crops or vegetables (Lešnik & Vajs 2017). It was 
listed as the 16
th
 worst weed in the world (Holm et al. 
1991). The management of Cyperus esculentus is very dif-
ficult and usually integrated methods (the combination 
of cultural, mechanical and chemical control options) 
are recommended (Bohren & Wirth 2013, Follak et al. 
2016). Trials of herbicide efficacy show that the herbi-
cides available on the Slovenian market cannot guaran-
tee completely reliable chemical control on arable land 
(Lešnik & Vajs 2017). Yellow nutsedge is also common 
on arable land near water bodies and safeguard zones 
where only a narrow selection of herbicides is available 
(Lešnik & Vajs 2017). 
Plant communities dominated by Cyperus esculentus 
were first described from riverbanks and gravel bars from 
several parts of Europe (Wisskirchen 1995, Felzines & 
Loiseau 2005, Lastrucci et al. 2012, Pellizzari 2020). Al-
tough Cyperus esculentus is known as a noxious weed on 
arable land for a long time, the vegetation with yellow 
nutsedge of invaded crop field has been rarely sampled so 
far (Fragner 2010) and as far as we know, it was never ana-
lysed on a large scale. Novel ecosystems have new com-
binations of species and arise through human action or 
environmental change (deliberate or inadvertent) (Hobbs 
et al. 2006). Arable vegetation could hardly qualify as 
novel ecosystem as it is intensively managed for centuries, 
but in recent decades we can observe drastic changes of 
weed vegetation species composition and abundance of 
neophytes due to changed agricultural practices (Richner 
et al. 2015). 
The aims of our study were: 1) to present the current 
distribution of Cyperus esculentus in Slovenia, and 2) to 
analyse the habitats and plant communities it invaded in 
Slovenia and as well as in Europe.
Methods
For the analysis of the current distribution of Cyperus es-
culentus in Slovenia we used data of flora mapping (ac-
cording to Ehrendorfer & Hamann 1965) that is stored 
in the Flovegsi database (Seliškar et al. 2003). Moreover, 
we collected all available distribution data from various 
sources (published and grey literature, unpublished data, 
internet). Specimens collected in the field are deposited 
in the herbarium of the Institute of Biology ZRC SAZU 
(LJS).
The vegetation in one hotspot of spread in Slovenia 
(Ljubljana basin) was sampled according to the Braun-
Blanquet method (Braun-Blanquet 1964) between 2015 
and 2020. Furthermore we collected all available relevés 
with dominant species Cyperus esculentus from Europe 
(Table 2) and stored them in the Turboveg database 
(Hennekens & Schaminée 2001).
We transformed the combined cover-abundance values 
into percentage and then log transformed it. For numeri-
cal comparisons we performed Non-metric multidimen-
sional scaling (NMDS) ordinations. Statistical analyses 
were made using Juice (Tichý 2002) and R software (R 
Development Core Team 2012). Diagnostic species were 
determined by comparison of all relevés and phi fidelity 
index (Chytrý et al. 2002).
The nomenclature of species is according to Martinčič 
et al. (2007) except for species Amaranthus emarginatus 
Salzm. ex Uline & Bray and Panicum laevifolium Hack., 
and syntaxa according to Šilc & Čarni (2012) and Mucina 
et al. (2016). In the classification of species into phytoso-
ciological groups (groups of diagnostic species) we mainly 
refer to the EuroVegChecklist (Mucina et al. 2016).

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in Slovenia
Figure 1: Distribution of Cyperus esculentus in Slovenia.
Slika 1: Razširjenost vrste Cyperus esculentus v Sloveniji.
Figure 2: Cumulative number of occurrences and occupied quadrants 
of Cyperus esculentus since its introduction in Slovenia. 
Slika 2: Kumulativno število pojavljanj in kvadrantov z vrsto Cyperus 
esculentus od prve zabeležbe v Sloveniji.
Results
Distribution and spatio-temporal spread pattern in 
Slovenia 
We gathered 99 records of Cyperus esculentus in Slovenia 
from 1980 to 2020 and this comprise 21 quadrants of 
the floristic mapping grid (Figure 1). Rapid increase in 
number of records and quadrants is evident after 2006 
(Figure 2).
Currently there are two major »hotspots« of invasion: 
in the Posočje region and in central Slovenia, however 
there are also some individual locations scattered across 
eastern Slovenia (Figure 1). 
Out of 99 records, 79 (80%) had habitat informa-
tion. Most invaded habitats in Slovenia were arable fields 
(66%), followed by riverine vegetation (7%), while Cype-
rus esculentus is rare in ruderal sites (4%) and grasslands 
(3%). Among arable fields maize was most frequently in-
vaded, but yellow nutsedge occurred also among potato, 
pumpkin, and cereals.
Phytosociologial affiliation and habitat preferences 
In the Ljubljana Basin, that is one of the two hotspots 
of its spread in Slovenia vegetation on 24 mostly arable 
fields was sampled. The average number of species per 
plot was 14 (ranging from 6 to 26 species). Cyperus es-
culentus dominated vegetation is found mainly on maize 
fields that are the most common fields in Ljubljansko 
Barje. Stands were dominated by Cyperus esculentus and 
characteristic plant species of three classes of anthropo-
genic vegetation—i.e., Digitario sanguinalis-Eragrostietea 
minoris, Papaveretea rhoeadis and Sisymbrietea, prevail as 
accompanying species (Table 1). Their presence indicates 
weed character of this vegetation type.
Figure 3: NMDS spiderplot of relevés of the three syntaxa from 
Slovenia. Numbers indicate centroids of particular vegetation cluster: 
1 – Digitario sanguinalis-Cyperetum esculenti, 2 – Bidentetea, 3 – weed 
vegetation (only Atriplici-Chenopodietalia sensu Mucina (1993)). 
Slika 3: NMDS graf popisov treh sintaksonov iz Slovenije. Številke 
predstavljajo centroide posameznih vegetacijskih klastrov: 1– Digitario 
sanguinalis-Cyperetum esculenti, 2 – Bidentetea, 3 – weed vegetation 
(samo Atriplici-Chenopodietalia v skladu z Mucina (1993)). 
The comparison of the relevés from Slovenia, origi-
nally classified into the Bidentetea (seasonally flooded 
nutrient-rich alluvia, banks and heavily nutrient loaded 
anthropogenic habitats) and Atriplici-Chenopodietalia 
(arable crops, gardens and waste places), and the relevés 

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in Slovenia
of plant communities dominated by Cyperus esculentus 
shows higher similarity with weed vegetation of cultivat-
ed land, although position in the graph is intermediate 
(Figure 3). 
tietea and Papaveretea classes are present, but in stands 
from wet, natural habitats species of classes Bidentetea and 
Phragmitetea are more abundant.
Plant community on arable fields is negatively charac-
terised by absence of species characteristic for Bidentetea 
class (e.g. Bidens frondosa, B. cernua, Xantium italicum). 
Weed stands are differentiated by typical weeds Veronica 
persica, Setaria pumila and Digitaria sanguinalis (T able 2). 
Based on the floristic composition and particular habi-
tat differences, we are of the opinion that Cyperus escu-
lentus dominated plant community presents new plant 
association within Digitario-Eragrostietea class.
Syntaxonomical scheme of Cyperus dominated 
communities:
Bidentetea Tx. et al. ex von Rochow 1951
Bidentetalia Br.-Bl. et Tx. ex Klika et Hadač 1944
Chenopodion rubri (Tx. In Poli et J. Tx. 1960) 
Hilbig et Jäger 1972
Cyperetum esculenti Wisskirchen 1995
Characteristic species: Cyperus esculentus
Differential species: Xanthium italicum, 
Amaranthus tuberculatus, Portulaca oleracea, 
Polygonum lapathifolium, Eragrostis pectinacea, 
Cyperus glomeratus, Bidens frondosus
Digitario sanguinalis-Eragrostietea minoris Mucina, 
Lososová et Šilc 2018
Eragrostietalia J. Tx. ex Poli 1966
Eragrostion Tx. in Oberd. 1954
Digitario sanguinalis-Cyperetum esculenti ass. 
nova hoc loco
Holotypus: relevé 7 in Table 1. 
Characteristic species: Cyperus esculentus
Differential species: Equisetum palustre, 
Digitaria sanguinalis, Setaria pumila, Veronica 
persica, Galinsoga ciliata
Discussion
Cyperus esculentus was first reported for Slovenia already in 
the 1980s by Gabrijel Seljak on maize fields near Kobarid 
(Dakskobler & Čušin 2002, Čušin 2006) and the first 
published locality was nearby in 1993 (Jogan & Podob-
nik 1995). The time of the first introduction in Slovenia 
is comparable to other western and southern countries 
(e.g. France, Italy) and Follak et al. (2016) erroneously 
indicated year 1999 as first occurrence in Slovenia due 
to reports published in local journals. In Europe, a rapid 
spread of species was observed since 2005 (Follak et al. 
1 Digitario-Eragrostietea 
a. Digitario-Cyperetum esculenti, Slovenia (this paper)
b. weed vegetation, Austria (Fragner 2010)
2 Isoëto-Nanojuncetea
c. Cyperus eculentus-(Isoëto-Nanojuncetea), Poland  
(Dajdok et al. 2007)
3 Bidentetea
d. Cyperetum esculenti, Italy (Lastrucci et al. 2012)
e. Cyperetum esculenti, Italy (Otolini 2013)
f. Cyperus eculentus community, France (Felzines &  
Loiseau 2005)
g. Cyperetum esculenti, France (Wisskirchen 1995)
h. Cyperetum esculenti, France (Cornier 2002)
i. Cyperetum esculenti, France (SMEAG 2011)
j. Cyperetum esculenti, Italy (Pellizarri 2020)
Figure 4: NMDS spiderplot of relevés with dominant Cyperus 
esculentus from Europe. Classification of the syntaxa is according to 
original authors.
Slika 4: NMDS graf popisov s prevladujočo vrsto Cyperus esculentus iz 
Evrope. Uvrščenost sintaksonov se ujema z izvorno uvrstitvijo avtorjev.
We also compared Cyperus esculentus dominated plant 
communities from other parts of Europe (Table 2). 
NMDS analysis shows great similarity between stands 
(Figure 4). Stands with Cyperus esculentus from (1) arable 
land (a-b) and (2 and 3) on wet riverbanks and gravel 
(c, e-f) are floristically very similar. In both vegetation 
types plant species characteristic for Digitario-Eragros-

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in Slovenia
2015), while in Slovenia this was delayed for a few years 
(Figure 2). The reason for time lag can also be attributed 
to differences in sampling efforts. In Slovenia, records 
were from the Ljubljana basin, while the north-eastern 
part is less invaded. In addition, we can expect spread 
to NE Slovenia from the neighbouring Styria (Austria), 
which is one of the invasion hotspots in Central Europe 
(Follak et al. 2015).
The status of naturalisation of the species in Slovenia 
changed rapidly from casual to invasive species (sensu 
Richardson et al. 2000). In 2001, Cyperus esculentus was 
not mentioned even as rare based on a surey of arable 
fields (Lešnik 2001) and it was also (erroneously) listed 
on the Red List as a vulnerable species (Anonymous 
2002). Few years later Lešnik (2009) mentioned it as 
present in smaller extent on arable land. In 2017, yel-
low nutsedge was already recognised as a species with the 
highest impact of noxious effects in agricultural produc-
tion systems (Lešnik & Vajs 2017). The species is now 
listed on the black action list of Slovenia (Grudnik et al. 
2015). In Europe, it is listed on the EPPO List of invasive 
alien plants (EPPO 2002), and on national black lists 
in other European countries (e.g. in Switzerland, INFO-
FLORA 2020).
Alien species initially first colonize disturbed habitats 
with low competition from other plant species, such as 
ruderal sites or riparian habitats (Jogan & Vreš 1998, 
Šilc 2002, Dakskobler & Vreš 2009, Dakskobler et al. 
2019). Later they can invade also arable fields. Such pat-
tern was observed in Slovenia with Ambrosia artemisiifo-
lia. First, it was found in ruderal sites and along roads 
(Šilc 2002), later it occupied arable land where it became 
an economically important weed (Lešnik & Vajs 2017). 
In Slovenia, Cyperus esculentus invasion began with the 
colonization of arable land (maize fields), where the spe-
cies is still most abundant. Other habitats, such as ruder-
al and riverine vegetation, are currently less invaded. In 
other parts of Europe, the invasion process was similar, 
pre-1990 records were mostly from arable land, and also 
recent records were from arable land (Follak et al. 2015). 
Other habitats, like riparian sites were by the opinion of 
Follak et al. (2015) colonized later from cultivated land. 
However, first reports of plant communities were from 
stands along water bodies (Wisskirchen 1995).
The pathways of introduction of Cyperus esculentus in 
Europe are very diverse and related to human activities. 
Soils transport, ploughing, and handling of crop waste are 
probably the main drivers for the species spread within 
and between fields (Follak et al. 2016). Lešnik (2009) 
pointed out that used growth substrate from nurseries 
that is used as compost, and cultivation in small gardens 
are one of the possible vectors of introduction and spread. 
As Cyperus esculentus is found mainly on maize fields in 
Central Slovenia, a possible reason could also be also un-
certified maize seeds (Dancza et al. 2004), but this was 
not confirmed (Follak et al. 2016).
Plant communities with Cyperus esculentus in Europe 
were so far studied sporadically, and contrarily to prevail-
ing floristic records in arable fields, they were sampled 
mostly along river banks and gravel bars. Riparian habi-
tats were colonized by Cyperus esculentus mainly along 
the rivers Rhine and Elbe rivers (Wisskirchen 1995), 
Tiber (Lastrucci et al. 2012) and Po (Pelizzarri 2020). 
Stands were classified as Cyperetum esculenti (class Bi-
dentetea). Relevés of weed vegetation with Cyperus escu-
lentus are rare, only from Austria (Styria) (Fragner 2010) 
and Slovenia (this paper). In both cases habitats are very 
similar: disturbed, often wet, therefore both vegetation 
types are floristically very similar with many common 
weed pioneer species (e.g. Echinochloa crus-galli, Setaria 
pumila, Polygonum lapathifolium) and therefore are tran-
sitional between Bidentetea and Digitario-Eragrostietea 
classes. Floristic differences between Bidentetea and Che-
nopodietalia albi are very small, as many weed species 
have their natural origin on wet, periodically flooded 
river banks (Mucina 1993). Many relevés from Fragner 
(2010) with lower abundance of Cyperus esculentus are 
also transitional to other thermophilous weed commu-
nites like Echinochloo-Setarietum.
Character species of Cyperetum eculenti is almost with 
all authors only Cyperus esculentus (Wisskirchen 1995, 
Pelizzarri 2020). Wisskirchen (1995) mentions also Po-
lygonum lapathifolium and Echinochloa crus-galli with 
higher constancy. Xanthium italicum also characterises 
riparian stands. On the other hand, arable stands of Digi-
tario sanguinalis-Cyperetum esculenti also dominated by 
Cyperus esculentus as character species but are character-
ized also by typical weed species while pioneer river bank 
species are missing.
Rapid spread of Cyperus esculentus is expected also in 
the future in many European countries, particularly with 
global warming (Simpson 2011). Based on the Slovenian 
experience with Cyperus esculentus spread and taking into 
consideration the short time from establishment of the 
species to becoming noxious weed, we can also expect 
other alien Cyperaceae species to invade and colonize 
Central Europe (Lešnik 2009). Beside Cyperus esculentus 
also C. rotundus L., C. iria L. and C. congestus Vahl. can 
potentially spread (Lešnik & Vajs 2017), while in Slo-
venia C. eragrostis is already established in ruderal sites 
(Dakskobler & Vreš 2009).

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Invasion, distribution and habitat affiliation of Cyperus esculentus, a new weed 
in Slovenia
Acknowledgements
Tatjana Čelik, Igor Dakskobler, Filip Küzmič and Fed-
erica Paradiso helped us in the field work. Igor Dakskob-
ler and Brane Anderle are co-authors of the distribution 
map. This work was financed by the Slovenian Research 
Agency (research core funding No. P1-0236).
Branko Vreš  https://orcid.org/0000-0002-7504-0416
Urban Šilc  https://orcid.org/0000-0002-3052-699X
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(Mucina et al. 2016) Layer 1 2 3 4 5 6 7* 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 fr %
Altitude 290 290 290 291 290 289 289 289 289 290 290 290 305 350 316 307 306 288 288 288 288 288 288 288
Relevé area (m
2
) 9 20 3 32 22 35 25 25 9 50 20 50 25 100 25 25 25 25 100 400 10 25 25 25
Class Cover herb layer (%) 80 80 90 90 90 90 90 80 80 50 90 70 80 70 80 65 65 80 80 90 80 70 80 100
CHE Cyperus esculentus C 3.2 4.5 3.4 5.5 3.4 5.5 4.5 5.5 3.4 3.4 5.5 4.4 4,5 4.5 1.1 2.1 4.4 2.1 5.5 4.5 3.4 2.1 2.4 1.1 24 100
Digitario sanguinalis-Eragrostietea minoris
DIG Digitaria sanguinalis C 3.1 + + 1.1 + 2.4 1.1 1.1 1.1 1.1 1.1 + + 4.4 16 67
DIG, BID Echinochloa crus-galli C 2.2 + 2.2 2.2 + + 1,1 + 2.1 3.1 2.4 1.1 + 1.1 14 58
DIG Panicum capillare (incl. P . barbipulvinatum) C 2.1 1.1 2.2 + 2.1 1.1 + 3.4 + 1.1 5.5 10 42
DIG Setaria pumila C + + 1.1 + + + 1.1 + 1.1 9 38
DIG Amaranthus lividus C + + + 3 13
DIG, POL Polygonum aviculare C + + + 3 13
DIG Portulaca oleracea C + 1 4
DIG Panicum miliaceum subsp. agricola C + 1 4
Papaveretea rhoeadis
PAR Veronica persica C + 1.1 1.1 1.1 + + + + 2.1 3.4 + 11 46
PAR Galinsoga ciliata C 2.1 + + 1.1 1.1 2.1 1.1 2.4 8 33
PAR, BID Chenopodium polyspermum C + + 1.1 2,1 + + + + 8 33
PAR, SIS Cirsium arvense C + + + + 1.1 + 6 25
PAR Polygonum lapathifolium subsp. pallidum C + + + + 2.1 2.4 6 25
PAR, SIS Stellaria media C + + + 3 13
PAR Capsella bursa-pastoris C + + + 3 13
PAR Brassica napus C 1.1 1.1 + 3 13
PAR, SIS Sonchus asper C + + 2 8
PAR Galinsoga parviflora C + + 2 8
PAR Lamium purpureum C + + 2 8
PAR, SIS Convolvulus arvensis C + + 2 8
PAR, SIS Matricaria perforata C + + 2 8
PAR Apera spica-venti C + 1 4
PAR Sinapis alba C + 1 4
PAR Solanum nigrum C + 1 4
Sisymbrietea
SIS Chenopodium album C + 1.1 + + 1.1 + + + + + 10 42
SIS Amaranthus retroflexus C 2.1 1.1 + + 4 17
SIS Atriplex patula C + + 2 8
SIS Conyza canadensis C 1.1 1 4
SIS Geranium pusillum C + 1 4
Bidentetea
BID Polygonum lapathifolium subsp. lapathifolium C 2.4 2.1 1.1 1.1 + + 1.1 + + + 10 42
BID, PAR Polygonum persicaria C 1.1 + + + + 5 21
BID Polygonum mite C + 1.1 + 1.1 4 17
BID Bidens tripartita C + + + 3 13
BID Rorippa sylvestris C + + + 3 13
BID Bidens frondosa C 1.1 2.1 + 3 13
BID Rorippa palustris C + 1 4
BID Ranunculus sardous C + 1 4
Epilobietea angustifolii
EPI, PHR Calystegia sepium C + + + 1.1 + + 6 25
EPI Myosoton aquaticum C + + + 3 13
EPI Urtica dioica C + + 2 8
EPI Solidago gigantea C + 1 4
EPI Verbascum nigrum C 1.1 1 4
EPI Echinocystis lobata C + 1 4
Artemisietea vulgaris
ART, EPI Erigeron annuus subsp. annuus C + + + + 4 17
ART, GER Silene latifolia subsp. alba C + + + + 4 17
ART Elytrigia repens C + + 2 8
ART; MOL Rumex obtusifolius C + + 2 8
ART, SIS Equisetum arvense C + 1 4
ART Medicago lupulina C + 1 4
ART Medicago sativa C + 1 4
ART, MOL Plantago lanceolata C + 1 4
Molinio-Arrhenatheretea
MOL Equisetum palustre C 1.1 1.1 2.1 1.1 2.2 2.1 1.1 1.1 1.1 1.1 1.1 1.1 1.1 2.1 2.1 + + 17 71
MOL, ISO Plantago intermedia C + + + + 2,1 + 2.1 + 2.2 + + 11 46
MOL Carex hirta C 2.2 1.1 + + + + + 7 29
MOL Lolium perenne C + + 2.4 3 13
MOL Trifolium repens C + + + 1.1 1.1 + 1.2 7 29
MOL Agrostis stolonifera C 1.2 1 4
MOL Dactylis glomerata s.str. C + 3.4 2 8
MOL Festuca pratensis C + 1.1 2 8
MOL Galium mollugo C + + + 3 13
MOL Hypericum tetrapterum C + 1 4
MOL Leucanthemum ircutianum C + 1 4
MOL Lotus corniculatus C + + 2 8
MOL Poa pratensis C + 1 4
MOL Poa trivialis C + + + + + 5 21
Table 1: Phytosociological table of Digitario sanguinalis-Cyperetum esculenti.
Tabela 1: Fitocenološka tabela asociacije Digitario sanguinalis-Cyperetum esculenti. 

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(Mucina et al. 2016) Layer 1 2 3 4 5 6 7* 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 fr %
Altitude 290 290 290 291 290 289 289 289 289 290 290 290 305 350 316 307 306 288 288 288 288 288 288 288
Relevé area (m
2
) 9 20 3 32 22 35 25 25 9 50 20 50 25 100 25 25 25 25 100 400 10 25 25 25
Class Cover herb layer (%) 80 80 90 90 90 90 90 80 80 50 90 70 80 70 80 65 65 80 80 90 80 70 80 100
CHE Cyperus esculentus C 3.2 4.5 3.4 5.5 3.4 5.5 4.5 5.5 3.4 3.4 5.5 4.4 4,5 4.5 1.1 2.1 4.4 2.1 5.5 4.5 3.4 2.1 2.4 1.1 24 100
Digitario sanguinalis-Eragrostietea minoris
DIG Digitaria sanguinalis C 3.1 + + 1.1 + 2.4 1.1 1.1 1.1 1.1 1.1 + + 4.4 16 67
DIG, BID Echinochloa crus-galli C 2.2 + 2.2 2.2 + + 1,1 + 2.1 3.1 2.4 1.1 + 1.1 14 58
DIG Panicum capillare (incl. P . barbipulvinatum) C 2.1 1.1 2.2 + 2.1 1.1 + 3.4 + 1.1 5.5 10 42
DIG Setaria pumila C + + 1.1 + + + 1.1 + 1.1 9 38
DIG Amaranthus lividus C + + + 3 13
DIG, POL Polygonum aviculare C + + + 3 13
DIG Portulaca oleracea C + 1 4
DIG Panicum miliaceum subsp. agricola C + 1 4
Papaveretea rhoeadis
PAR Veronica persica C + 1.1 1.1 1.1 + + + + 2.1 3.4 + 11 46
PAR Galinsoga ciliata C 2.1 + + 1.1 1.1 2.1 1.1 2.4 8 33
PAR, BID Chenopodium polyspermum C + + 1.1 2,1 + + + + 8 33
PAR, SIS Cirsium arvense C + + + + 1.1 + 6 25
PAR Polygonum lapathifolium subsp. pallidum C + + + + 2.1 2.4 6 25
PAR, SIS Stellaria media C + + + 3 13
PAR Capsella bursa-pastoris C + + + 3 13
PAR Brassica napus C 1.1 1.1 + 3 13
PAR, SIS Sonchus asper C + + 2 8
PAR Galinsoga parviflora C + + 2 8
PAR Lamium purpureum C + + 2 8
PAR, SIS Convolvulus arvensis C + + 2 8
PAR, SIS Matricaria perforata C + + 2 8
PAR Apera spica-venti C + 1 4
PAR Sinapis alba C + 1 4
PAR Solanum nigrum C + 1 4
Sisymbrietea
SIS Chenopodium album C + 1.1 + + 1.1 + + + + + 10 42
SIS Amaranthus retroflexus C 2.1 1.1 + + 4 17
SIS Atriplex patula C + + 2 8
SIS Conyza canadensis C 1.1 1 4
SIS Geranium pusillum C + 1 4
Bidentetea
BID Polygonum lapathifolium subsp. lapathifolium C 2.4 2.1 1.1 1.1 + + 1.1 + + + 10 42
BID, PAR Polygonum persicaria C 1.1 + + + + 5 21
BID Polygonum mite C + 1.1 + 1.1 4 17
BID Bidens tripartita C + + + 3 13
BID Rorippa sylvestris C + + + 3 13
BID Bidens frondosa C 1.1 2.1 + 3 13
BID Rorippa palustris C + 1 4
BID Ranunculus sardous C + 1 4
Epilobietea angustifolii
EPI, PHR Calystegia sepium C + + + 1.1 + + 6 25
EPI Myosoton aquaticum C + + + 3 13
EPI Urtica dioica C + + 2 8
EPI Solidago gigantea C + 1 4
EPI Verbascum nigrum C 1.1 1 4
EPI Echinocystis lobata C + 1 4
Artemisietea vulgaris
ART, EPI Erigeron annuus subsp. annuus C + + + + 4 17
ART, GER Silene latifolia subsp. alba C + + + + 4 17
ART Elytrigia repens C + + 2 8
ART; MOL Rumex obtusifolius C + + 2 8
ART, SIS Equisetum arvense C + 1 4
ART Medicago lupulina C + 1 4
ART Medicago sativa C + 1 4
ART, MOL Plantago lanceolata C + 1 4
Molinio-Arrhenatheretea
MOL Equisetum palustre C 1.1 1.1 2.1 1.1 2.2 2.1 1.1 1.1 1.1 1.1 1.1 1.1 1.1 2.1 2.1 + + 17 71
MOL, ISO Plantago intermedia C + + + + 2,1 + 2.1 + 2.2 + + 11 46
MOL Carex hirta C 2.2 1.1 + + + + + 7 29
MOL Lolium perenne C + + 2.4 3 13
MOL Trifolium repens C + + + 1.1 1.1 + 1.2 7 29
MOL Agrostis stolonifera C 1.2 1 4
MOL Dactylis glomerata s.str. C + 3.4 2 8
MOL Festuca pratensis C + 1.1 2 8
MOL Galium mollugo C + + + 3 13
MOL Hypericum tetrapterum C + 1 4
MOL Leucanthemum ircutianum C + 1 4
MOL Lotus corniculatus C + + 2 8
MOL Poa pratensis C + 1 4
MOL Poa trivialis C + + + + + 5 21

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(Mucina et al. 2016) Layer 1 2 3 4 5 6 7* 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 fr %
MOL Potentilla reptans C 1.2 + 2 8
MOL Ranunculus flammula C + 1 4
MOL Mentha verticillata C + 1 4
MOL Trifolium pratense C + + 2.4 1.2 4 17
MOL Lolium multiflorum C 2.2 1 4
MOL Lychnis flos-cuculi C + 1 4
MOL Phleum pratense C 1.1 1 4
MOL Ranunculus repens C + + 1.2 3 13
MOL Rumex conglomeratus C + + 2 8
MOL Rumex crispus C + + + 3 13
MOL T araxacum sect. Ruderalia C + + 2 8
MOL Trifolium hybridum C 1.1 1 4
MOL Verbena officinalis C 1.1 1 4
MOL, PHR Lysimachia vulgaris C + 1 4
MOL; PHR Stachys palustris C + 1 4
Other species
PHR Lythrum salicaria C 1.1 + 2.1 + + 5 21
POL Plantago major s. str. C + + 2.1 + 4 17
POP Rubus caesius C + + 2 8
PHR Mentha aquatica C + 1 4
POL Poa annua C + 1 4
Arctium sp. C + 1 4
Geranium sp. C + 1 4
Sonchus sp. C + 1 4
FRA Salix cinerea B + 1 4
ISO Veronica catenata C + 1 4
ISO, SCH Juncus articulatus C + 1 4
Mentha sp. C + 1 4
ISO Cyperus fuscus C + 1 4
Rumex x pratensis C + 1 4
PHR Scutellaria galericulata C 1.1 1 4
POP Scrophularia nodosa C + 1 4

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Appendix to Table 1
Number of relevé; authors; date; locality; habitat descrip-
tion; altitude (m); coordinates (WGS84).
1: 20080709/08; Vreš B. & Čelik T.; 9.7.2008; Sloveni-
ja, Ljubljansko barje, Brest: along dirt road and ditch 
Peščenek ca. 700 m north from village; pumpkin field.; 
290; 45.97924; 14.4888;
2: 20180709/16; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Ig, ca. 1 km NE from village, left bank 
of river Ižica (Iščica); withinin and margin of maize field; 
290; 45.96918; 14.53979;
3: 20150630/1; Šilc U. & Vreš B.; 30.6.2015; Slovenija, 
Ljubljansko barje: Ig, ca. 1 km NE from village in vicinity 
of bridge over river Ižica (Iščica); within and margin of 
maize field; 290; 45.96724; 14.53607;
4: 20180709/11; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Ig, along the road Ig-Škofljica between 
streams Želimeljščica and Dremavščica, ca. 2 km NE 
from settlement; abandoned field, along the ditch; 291; 
45.96508; 14.54596;
5: 20180709/01; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Ig, east along Ižanska road, ca. 1 km 
N from Ig and ca. 700 m east from Iška Loka; along the 
ditch between maize fields; 290; 45.97203; 14.52572;
6: 20180709/02; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Iška Loka, ca. 1.7 km north from vil-
lage, along the road Iška Loka – Ižanka (Črna vas); mar-
gin and in the maize field; 289; 45.98541; 14.51765;
7: 20180709/03; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Iška Loka, ca. 1.2 km north from the 
village, along the road Iška Loka – Ižanka (Črna vas); 
abandoned field; 289; 45.98454; 14.51696;
8: 20180709/04; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Iška Loka, ca. 1 km north from the vil-
lage, along the road Iška Loka – Ižanka (Črna vas); maize 
field; 289; 45.98404; 14.51625;
9: 20180709/05; Šilc U. & Vreš B.; 9.7.2018; Slovenija, 
Ljubljansko barje: Iška Loka, ca. 350 m north from the 
village, east along the road Iška Loka – Črna vas; maize 
field; 289; 45.97763; 14.5172;
10: 20180702/1; Šilc U. & Vreš B.; 2.7.2018; Slovenija, 
Ljubljansko barje, Brest: Nature Reserve Iški morost (Mali 
deli), along the dirt road and channel Peščenek ca. 1.3 
km north from the village; maize field; 290; 45.98476; 
14.48702;
11: 20180702/2; Šilc U. & Vreš B.; 2.7.2018; Sloveni-
ja, Ljubljansko barje, Brest: Gmajne, long dirt road ca. 
850 m north from village; margin of maize field; 290; 
45.98053; 14.48855;
12: 20180702/3; Šilc U. & Vreš B.; 2.7.2018; Slovenija, 
Ljubljansko barje, Brest: Gmajne, along the dirt road ca. 
600 m north from the village; maize field, at the margin 
of the ditch; 290; 45.97865; 14.48922;
13: Šilc U. & Vreš B.; 31.7.2019; Slovenija, Loka pri 
Mengšu, millet field; 305; 46.14940; 14.55247;
14: 30.7.2020; Vreš B.; Slovenija, Gorenjska, Smlednik, 
maize field; 350; 456176, 5113930;
15: 10.9.2020; Vreš B.; Slovenija, Gorenjska, Zgornje 
Pirniče, buckwheat field; 316; 456375, 5110550;
16: 16.9.2020; Vreš B.; Slovenija, Gorenjska, Rača, aban-
doned field; 307; 473910, 5110130;
17: 16.9.2020; Vreš B.; Slovenija, Gorenjska, Rača, aban-
doned field; 306; 473725, 5110145;
18: 8.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Črna 
vas, hemp field; 288; 460440, 5094950;
19: 8.7.2020; Vreš B.; Slovenija, Ljubljansko barje, 
Črna vas, wet depression on a rape field; 288; 461900, 
5095550;
20: 8.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Črna 
vas, abandoned rape field; 288; 461962, 5095440;
21: 6.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Sinja 
Gorica, edge of soya field; 288; 448220, 5092208;
22: 6.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Blat-
na Brezovica, wheat field; 288; 448310, 5091920;
23: 6.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Blat-
na Brezovica, sown grassland; 288; 448308, 5091944;
24: 6.7.2020; Vreš B.; Slovenija, Ljubljansko barje, Blat-
na Brezovica, maize field; 288; 448288, 5091690.

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Table 2: Synoptic table of plant communities dominated by Cyperus esculentus.
Tabela 2: Sinoptična tabela rastlinskih združb z vrsto Cyperus esculentus.
Column number 1 2 3 4 5 6 7 8 9 10
Number of relevés 24 20 2 2 2 1 11 4 2 14
CHE              Cyperus esculentus                                100 100 2 2 2 1 100 100 2 100
DIG Digitario-Eragrostietea
Panicum capillare                                 42 . . . 1 1 100 25 1 .
Digitaria sanguinalis 58 25 . . 1 1 18 75 . 7
Setaria pumila 38 40 . . . 1 9 . 2 .
Portulaca oleracea 4 . . . 2 1 36 . . 86
Chenopodium ambrosioides                          . . . . . 1 36 . . .
Amaranthus emarginatus                            . . . . . . 82 50 . .
Polygonum aviculare                               13 . . . . . . 25 . .
Panicum laevifolium                               . 20 . . . . . . . .
Setaria viridis . . . . 2 . . . . .
Panicum dichotomiflorum                           . . . . 2 . . . . 29
Setaria macrocarpa . 10 . . . . . . . .
Amaranthus lividus                               13 . . . . . . . . .
BID Bidentetea
BID, DIG         Echinochloa crus-galli 58 55 2 1 1 1 100 25 2 43
Polygonum lapathifolium (incl. subsp. lapatifolium) 42 15 . . 1 . 100 50 2 29
Bidens frondosa                              13 . . 2 1 . 91 50 2 14
Rorippa palustris                                 4 . . . 1 . 9 50 2 .
Polygonum persicaria 21 5 . . . . 64 . 1 7
Polygonum hydropiper                             . . 2 . 2 . 18 75 . .
Bidens tripartita                     13 . 1 . . . 9 . . .
Xanthium italicum                                 . . . 2 2 . 91 . . 93
Rorippa sylvestris                                . . . . . . 82 75 . .
Bidens cernuus . . . . . . . 75 1 .
Chenopodium polyspermum                           33 . . . . . 64 . . .
BID, SIS         Xanthium strumarium . . . . . . . 2 .
Pulicaria vulgaris . . . . . . 25 . .
Bidens radiatus                                   . 1 . . . . . . .
Polygonum mite 17 . . . . . 9 . . .
Polygonum dubium                               . . . 1 . . . . .
Chenopodium rubrum . . . . . . 50 . .
Solanum lycopersicum . . . . . 18 . . .
Polygonum lapathifolium subsp. brittingeri . . . . . 9 . . .
Rumex maritimus                                   . 1 . . . . . . .
Corrigiola litoralis . . . . . . 25 . .
Other species
ALN              Lythrum salicaria                                 21 . . 1 . . 27 50 2 .
ALN              Lycopus europaeus                                 . . . 1 . . 9 25 1 7
ART              Equisetum arvense                                 4 . . 1 . . 9 . 1 .
EPI              Calystegia sepium                                 25 10 . . . . 9 . 1 .
ISO              Cyperus fuscus                                    4 . 2 2 . . 9 25 . 21
MOL              Lysimachia vulgaris                               4 . . 1 . . 9 25 1 .
MOL              Agrostis stolonifera 4 . 2 1 . . . 50 . .
SIS              Chenopodium album 42 20 . . 1 . 9 . . .
 1. Digitario-Cyperetum esculenti, Slovenia (this paper)
 2. weed vegetation, Austria (Fragner 2010)
 3. Cyperus eculentus-(Isoëto-Nanojuncetea), Poland  
(Dajdok et al. 2007)
 4. Cyperetum esculenti, Italy (Lastrucci et al. 2012)
 5. Cyperetum esculenti, Italy (Otolini 2013)
 6. Cyperus eculentus community, France (Felzines &  
Loiseau 2005)
 7. Cyperetum esculenti, France (Wisskirchen 1995)
 8. Cyperetum esculenti, France (Cornier 2002)
 9. Cyperetum esculenti, France (SMEAG 2011)
10. Cyperetum esculenti, Italy (Pellizzari 2020)