Hacquetia 15/1 • 2016, 93-100	Dol: 10.1515/hacq-2016-0007
Cyperus glomeratus L. - rediscovered in Slovakia
DE GRUYTER
OPEN
Zuzana Melečkova1, *, Daniel Dite1, Pavol Elias Jr.2 & David Schmidt3
Keywords: Eleocharition ovatae, Danube, Pannonian lowland, temporary wetland habitats, eutrophication.
Ključne besede: Eleocharition ovatae, Donava, Panonska nižina, občasni mokriščni habitati, eutrofikacija.
Abstract
Two new and one historical site of Cyperus glomeratus were discovered in SW Slovakia along the left bank of the river Danube. After 60 years, this is the second record of the species in the country. We provide details about the historical and current distribution of C. glomeratus in Slovakia based on herbarium revision and field survey on the Slovak section of the river Danube. According to the phytosociological data collected from the new locality (Cenkov) and from the confirmed historic locality (Sturovo), the stands with C. glomeratus were identified with less developed, ruderalized form of the association Cyperetum micheliani.
Izvleček
Odkrili smo novo in potrdili že znano rastišče vrste Cyperus glomeratus na levem bregu Donave na jugozahodu Slovaške. Po 60 letih je to drugo nahajališče te vrste v državi. Na osnovi revizije herbarijskega materiala in terenskih pregledov območja ob Donavi na Slovaškem smo prikazali nekdanjo in trenutno razširjenost vrste C. glomeratus na tem območju. Sestoje z vrsto C. glomeratus smo uvrstili v asociacijo Cyperetum micheliani na osnovi fitocenoloških popisov z novega rastišča (Čenkov) in potrjenega znanega nahajališča (Sturovo).
Received: 13. 3. 2015 Revision received: 4. 11. 2015 Accepted: 7. 11. 2015
1	Institute of Botany of the Slovak Academy of Sciences, Dubravska cesta 8, Bratislava, Slovakia. E-mail: daniel.dite@savba.sk, zuzana.meleckova@savba.sk * Corresponding author
2	Department of Botany, Slovak University of Agriculture, Tr. A. Hlinku 2, 949 76 Nitra, Slovakia. E-mail: pavol.elias.jun@gmail.com
3	Department of Botany, University of West Hungary, Bajcsy-Zsilinszky u. 4, 9400 Sopron, Hungary. E-mail: david.schmidt@emk.nyme.hu
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Cyperusglomeratus L. — rediscovered in Slovakia
Introduction
The concept of the genus Cyperus L. is very diverse; therefore the number of species is not fixed. The genus contains many species, in the European flora there are recognized 27 species (DeFilipps 1980). According to Marhold & Hindak (1998) in Slovakia six species are known: C. flavescens, C. fuscus, C. glomeratus, C. longus, C. micheli-anus (syn. Dichostylis micheliana) and C. pannonicus (syn. Acorellus pannonicus).
Cyperus glomeratus L. [syn. C. aureus Georgi, Chlorocy-perus glomeratus (L.) Palla, Pycreus glomeratus (L.) Hayek] is an annual, rarely perennial short-lived flatsedge (gal-ingale) species from the Cyperaceae family. Rhizomes are absent or they are rarely creeping, stems 10-80 cm, mainly solitary or rarely caespitose. Leaves are 2-10 mm wide, shorter than to exceeding the stems. Bracts are 2-6, exceeding the inflorescence. Inflorescence is a compound umbel or a sessile, dense head; rays up to 10 cm. Spikelets 5-12 x 1-1.5 mm, linear, with 8-20 flowers, ascending to patent. Glumes (1.5-)2-2.3 x 0.5 mm, linear or linear-lanceolate, obtuse, pale- or reddish-brown, with greenish keel. Stamens are 3. Nuts are 3/5 as long to as long as the glumes, linear-oblong, dark brown (DeFilipps 1980).
Cyperus glomeratus is thermophilous, pioneer plant of lowland river banks, lakeshores, marshes, rarely wet meadows (Dostal & Cervenka 1992). It grows secondarily on rice fields and ditches. The species prefers raw gravel or sandy soils with muddy surface resulting from periodical inundation. In paleobotanical studies it is an indicator of former warm areas of the interglacial periods in Eurasia as it was found in several macrofossils (Binka & Nitychoruk 2003, Durnikin & Zinovyeva 2014) including sites north of the present northern limit of its distribution range (Lancucka-Srodoniowa 1979, Lesniak 1994).
The distribution range of C. glomeratus covers SE part of central Europe (Hungary, Slovakia), SE Europe (the former Yugoslavia, Romania, Bulgaria, Greece and Turkey), Ukraine, the western part of Russia, Caucasus, NW Iran, Turkmenistan, Uzbekistan, Pakistan, N China and Japan (Schulze-Motel 1980). In Central, SW and W Europe the species is considered as a casual alien (Spain, France - Verloove 2014, the Nederland - Stolwijk 1991, Switzerland, Germany - Schulze-Motel 1980, Austria -Essl & Rabitsch 2002, the Czech Republic - Dostal & Cervenka 1992, Pysek et al. 2012) or invasive (Italy -Celesti-Grapow et al. 2009).
In the Pannonian lowland C. glomeratus is autoch-tonous; it occurs sporadically: in Hungary it is more frequent in the western part of the country along larger rivers like Danube, Raba, Drava and Mura and in the eastern part of the country along the lower sections of
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Tisza and Maros rivers (Kiraly et al. 2009). C. glomeratus in lowland Serbia (Vojvodina) is common along the Danube and Sava rivers. Outside of the Pannonian lowland, Zlatkovic et al. (2005) report it only from a few localities on sandy river banks of Južna Morava and beside the moderate number of floristic data (Jovanovic & Bartula 1997, Petric et al. 2010, Stojkovic 2012) and field observations it is more widespread than it can be concluded from the literature (Peric in litt.). More to the north, like in Slovakia it is rather rare.
In this contribution we present historic and recent occurrence of Cyperus glomeratus in Slovakia. As the species is frequently reported from more locations on the right side of the Danube, e.g. in Györ, Hungary (Polgar 1941, Schmidt & Bauer 2005), we compare the vegetation composition of the stands with this species on each side of Danube within the NW part of the Pannonian lowland.
Material and methods
The field observation was carried out along 82 km section of the left bank of the Danube river (from 1792 to 1710 fluvial kilometers) in Slovakia (Podunajska nižina lowland) in year 2014 and the survey was repeated in 2015. Special attention was given to active and former gravel pits and open gravel beaches with potential habitats of semi-aquatic pioneer vegetation of flat depressions in Kližska Nema, Vel'ké Kosihy, Zlatna na Ostrove, Čenkov, Sturovo and Chl'aba. Phytosociological relevés were sampled according to the Zürich-Montpellier approach using the adapted nine-grade Braun-Blanquet's scale (Barkman et al. 1964) and were stored in the TURBOWIN database (Hennekens & Schaminée 2001). The nomenclature of plant taxa are according to the database of Euro+Med plantbase (2006).
Data regarding the distribution of the species in Slovakia were obtained from herbaria BP, BRA, BRNU, NI, PR, PRC, OLM, SAV, SLO and from the Database of vascular plants deposed in the Institute of Botany, Slovak Academy of Sciences in Bratislava. Herbarium abbreviations are according to Vozarova & Sutory (2001).
Results and discussion
Historical occurrence of Cyperus glomeratus in Slovakia
In Slovakia, the exact distribution of Cyperus glomeratus is not clarified, since most of the data are published in determination keys on the Czechoslovak flora and the source data are not available. The oldest literature mentions that
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Cyperus glomeratus L. — rediscovered in Slovakia
C. glomeratus was found in some years in sandy bottoms of the Danube in the Zitny ostrov area near Bratislava ("Schutt Insel"), but without exact locality (Csader 1856). The second reference gives Pantocsek (1907) from large surroundings of Bratislava city and later Polivka et al. (1928) from the Podunajska nižina lowland („Velky Zitny ostrov"), but exact data are not given in these reports.
Dostal (1958) confirms the species from the same area but also without closer localisation. Doubtful data provides Majovsky (1959) from SE Slovakia. He reports C. glomeratus from the Tice oxbow near Hrušov settlement on the Vychodoslovenska nižina lowland in the Tisa river basin (Database of vascular plants deposed in the Institute of Botany). The locality was documented by two herbarium specimens (Majovsky & Zaborsky 1961 SLO). This data was used in the actualized Czechoslovak determination key (Dostal 1989) where two locations are mentioned: Streda nad Bodrogom, still in the Vychodoslovenska nižina lowland (SE Slovakia) and Sturovo in the Podunajska nižina lowland (SW Slovakia). Herbarium material of C. glomera-tus collected from SE Slovakia (Streda nad Bodrogom and Keresztur settlements) by Majovsky & Zaborsky we revised as Bolboschoenus maritimus agg. and Cyperus michelianus. To the second cited site (Podunajska nižina) Dostal (l.c.) notes that this observation was valid until 1953 originating from Hejny (1960) who observed the species in 1953
on the Danube river bank in Sturovo by the Boži kopec (in different part of the text Hejny stated this location as Obid near Sturovo). The latest determination key (Dostal & Červenka 1992) includes only Podunajska and Vycho-doslovenska nižina lowlands in general, without closer data. In the following years there was not paid attention to the confirmation of these localities. Recent floristic studies from the above mentioned areas (e.g. Jehlik et al. 2005, Martonfi et al. 2014) have not reported the species.
Recently confirmed locations
We found Cyperus glomeratus in the town of Sturovo (locality "Boži kopec"), on the left bank of the Danube in a former gravel depot (Fig. 1). This is a confirmation of the first reliable record in Slovakia found by Hejny in 1953. Artificial shallow depressions approx. 20 m from the shoreline were flooded and at the beginning of September the muddy pools were occupied by pioneer semi-ruderal stands of annual hygrophytes. Cyperus glomeratus was developed sporadically in three shallow ponds each with an average area of 20 m2. The estimated population of the species was up to two hundred fertile individuals. The vegetation can be characterized by the following two relevés:
17°
18°
19°
21°
22°
49°
48°
67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101
Figure 1: Historical and recent occurrence of Cyperus glomeratus L. in Slovakia: O — doubtful location, • — recently confirmed locations. Slika 1: Nekdanja in recentna nahajališča pojavljanja vrste Cyperus glomeratus L. na Slovaškem: O — dvomljiva lokacija, • — nedavno potrjeno nahajališče.
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Relevé 1: Stúrovo, Bozí kopec, left bank of the Danube river, former gravel depot, shallow pool, water depth 10 cm, 16m2, E1: 20% E0: 0; open water: 90%; 112 m a. s. l., 47°46'55.0"; 18°42'20.4", Melecková, Díte & Eliás jun., 16. 9. 2014.
E1: Trifolium repens 2a, Cyperus glomeratus 1, Echi-nochloa crus-galli 1, Juncus articulatus 1, Ambrosia arte-misiifolia +, Conyza canadensis +, Cyperus fuscus +, Cheno-podium ambrosioides +, Persicaria lapathifolia +, Plantago major +, Populus alba juv. +, P. canadensis juv. +, Potentilla supina +, Setaria viridis +, Solidago gigantea +, Tripleuro-spermum inodorum +.
Relevé 2: Stúrovo, Bozí kopec, left bank of the Danube river, former gravel depot, shallow pool, water depth 5 cm, 16m2, E1: 25% E0: 0; open water: 100%; 112 m a. s. l., 47°46'55.0"; 18°42'20.4", Melecková, Díte & Eliás jun., 16. 9. 2014.
E1: Cyperus fuscus 2a, Echinochloa crus-galli 1, Juncus articulatus 1, Veronica anagallis-aquatica 1, Typha angusti-folia 1, Ambrosia artemisiifolia +, Cyperus glomeratus +, Epilobium tetragonum +, Lemna minor +, Persicaria lapathifolia +.
The second, new site to Slovakia was found 12 km west of Stúrovo, between the settlements Muzla and Cenkov in the area of active, extensively used gravel pit on the left bank of the Danube. The population was poor: two robust fertile specimens were observed on raw gravel with thin silt exposed in the last year. The following relevé was recorded:
Relevé 3: Muzla, Cenkov, left bank of the Danube, active gravel pit, shallow dried depression, 16m2, E1: 20%, E0: 10%; 105 m a. s. l., 47°46'16.54"; 18°32'57.10", Melecková & Díte, 16. 10. 2014.
E1: Echinochloa crus-galli 2a, Agrostis stolonifera 1, Plantago major 1, Salix fragilis 1, Sonchus arvensis 1, Cyperus fuscus +, Cyperus glomeratus +, Persicaria lapathifolia +, Poa annua +, Polygonum aviculare +, Populus x canadensis juv. +, Ranunculus sceleratus r.
In presented relevés the stands are two-layered, on the surface are typical dwarf plants like Cyperus fuscus and Trifolium repens, in the upper layer besides Cyperus glomeratus, ruderal hygrophytes prevail (Persicaria lapathifolia and Echinochloa crus-galli). Each recorded species composition is regarded as an early succession stage of desiccating pools after the river retreats.
We identified these relevés as a transition vegetation between the classes Isoëto-Nano-Juncetea Br.-Bl. et Tüxen ex Br.-Bl. et al 1952 and Bidentetea tripartitae Tüxen et al. ex von Rochow 1951 (Sumberová 2011). Regarding
the heterogeneity and the high amount of ruderal ni-trophilous species the vegetation cannot be determined to an exact association, it can be considered as a strongly ruderalized stage of the association Cyperetum miche-liani (Horvatic 1931) within the alliance Eleocharition ovatae Philippi 1968. The vegetation reported by Hejny (1960) on the same locality had similar initial and ruderal character which he considered as a coenosis of Rorippa sylvestris-Rumex crispus.
In 2015, when large parts of river shores were exposed during extreme droughts, we found a new location of Cyperus glomeratus in the vicinity of Sturovo, in Kamenica nad Hronom settlement. It created sporadical micro-populations in the dense stands of young Salix fragilis-shoots in the vegetation of Cyperetum micheliani association. We assume that Cyperus glomeratus was in years of favorable climatic and ecological conditions continuously present on these localities in the last decades as well.
Coenological and ecological remarks to Cyperus glomeratus
C. glomeratus is not a major diagnostic species for particular syntaxonomical unit. It is a component of a large variety of azonal vegetation and it occurs in more associations belonging to even different classes. Yet, the optimum of the species is found in the vegetation of the alliances Na-no-Cyperion Koch ex Libbert 1932, Eleocharition ovatae and Verbenion supinae Slavnic 1951 of the order Nanocy-peretalia Klika 1935, class Isoeto-Nanojuncetea (Brullo & Minissale 1998).
Within this order and class, Csiky & Purger (2008) include stands with C. glomeratus along the Drava river in SW Hungary into the association Polygono-Eleocharite-tum ovatae Eggler 1933; in the Tisza river basin in SE Hungary Bodrogkozy (1982) includes the species in the associations Cypero fusci-Juncetum bufonii Soo et Csuros 1944 and Cyperetum micheliani, each community belongs into the alliance Eleocharition ovatae.
The alliance Eleocharition ovatae is oligotrophic amphibious vegetation in temporary pools on river deposits or fishponds, characterized by therophytes, rarely accompanied by hemicryptophytes and dwarf geophytes which grow on periodically flooded soils (Sumberova 2011). Crucial factors for development are the short vegetation period and the periodically moist habitat. After floods, raw alluvial deposits and organic silty material are left behind. In desiccating habitats due to gradual evaporation large amount of carbonate accumulate (Borhidi et al. 2012). Cyperus glomeratus is well adapted to the fluctuating ecological circumstances. Such habitat conditions were present in our sites, but the eutrophication was high.
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Figure 2: Vegetation of desiccating pools with Cyperus glomeratus L. in Šturovo (a) and Čenkov (b) in SW Slovakia on the left bank of the Danube Slika 2: Vegetacija izsušenih uleknin z vrsto Cyperus glomeratus L. na nahajališčih Šturovo (a) in Čenkov (b) na jugozahodnem Slovaškem na levem bregu Donave.
On the opposite side of the Danube in Hungary, where are analogous ecological circumstances, C. glomeratus is relatively common in the favourable years (Barina & Schmidt 2004, Schmidt 2014). We present the following relevé from Gyôr from a nonstandard area of 2 x 2 m for the purpose of comparison the vegetation with the newly discovered locations in Slovakia.
Relevé 4: Gyôr, between Likocs and Bacsa settlements on the muddy bank of the Mosoni-Duna river, 2 x 2 m, E1: 70%, E0: 0; 47°43'33"; 17°41'08", 111 m a. s. l. ; Schmidt. 9. 9. 2003.
E1: Salix alba juv. 4, Persicaria dubia 3, Cyperus fuscus 2, Chenopodium rubrum 2, Cyperus glomeratus 1, Rorippa palustris 1, Potentilla supina 1, Bidens frondosus 1
Ranunculus sceleratus 1, Atriplexpatula +, Chenopodium glaucum +, Dichostylis michelianus +, Gnaphalium uligi-nosum +, Veronica beccabunga +, Amaranthus blitum +, Urtica dioica +, Digitaria sanguinalis +, Plantago major +, Capsella bursa-pastoris +,
Poa annua +, Rumex obtusifolius +, Barbarea stricta +, Butomus umbellatus +, Amaranthus albus +, Batrachium trichophyllum +.
Vegetation sampled in Hungary is more species-rich. These stands are enriched with several Nanocyperion species such as Cyperus michelianus or Gnaphalium uligino-sum. Species of trampled, wet habitats are also abundant, together with nitrophilous plants of the alliance Biden-tion tripartitae. Other accompanying species of Cyperus glomeratus outside of the relevé area were Limosella aquat-ica, Potamogeton pectinatus and Potentilla supina. The species composition is heterogeneous due to the rough terrain and high amount of organic silt. Like relevés from Slovakia, this relevé can be also regarded as a less typical association of Cyperetum micheliani. High cover of young Salix alba shoots and the higher species richness indicate latter stages of succession. In Slovakia, observed vegetation had more pioneer character, since the total cover of vascular plants did not reach higher than 25%, in two relevés stagnant water was still present.
Nitrophilous and ruderal species are frequent in both sides of the Danube, since stands are developed in anthropogenic, sometimes eutrophic habitats. Extraction of gravel and sand from the river terraces, like in Cenkov is suitable for Cyperus glomeratus, as it needs constant distur-
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bance in order to create pioneer surface without high plant cover. In such sites ruderalisation is high, but according to our observations the species is resistant to this stress.
Succession of these temporal communities is fast, the pioneer stands usually alters to the vegetation of the alliances Salicion triandrae or Phragmition australis (Valachovic et al. 2001). Cyperus glomeratus has short life cycle and it is not adapted to the strong competition of perennials and in these communities is less typical (Borhi-di et al. 2012). However, the understory of willow gallery forests can offer suitable conditions, as it occurs in such places along the lowland section of Drava river (Kevey et al. 2008). In southern parts of Europe, for instance in Italy, it is quite abundant in vegetation with Phragmition australis (Biondi et al. 2009).
Cyperus glomeratus has strong spreading potential if ecological conditions are optimal (humid early-vegetation period followed by dry and warm late-summer). If the circumstances are not suitable, the vegetation does not develop at all within the growing season (Csiky & Purger 2008).
Distribution and threats
Barina and Riezing (in litt.) report Cyperus glomeratus from at least five locations on the right bank of the Danube in Hungary between the towns Komarom and Esztergom, very close to the discovered localities. Despite the relatively frequent occurrence in Hungary along the whole section of the Danube (Barina & Schmidt 2004, Schmidt 2014), the species is far rarer on the opposite side, in Slovakia.
Rivers are important transporters of diasporas in cultivated landscape (Jehlik et al. 2005) where several exotic species can spread, for instance Cyperus strigosus in Bulgaria (Tzonev et al. 2003) or Cyperus eragrostis in Slovenia (Dakskobler & Vres 2009). Except such neophytes, river-banks often provide places for rare native species. To similar habitats as of C. glomeratus are bound other rare species of Cyperaceae, which are reported in Slovakia very rarely (e.g. Dostal 1989, Dostal & Cervenka 1992). Here belong Scirpus radicans (Dite & Elias 2013), Schoenoplec-tus triqueter (Sucha 1992, Dite et al. 2016), Schoenoplectus pungens (Ondrasek 2006) and Carex bohemica (Dite et al. 2015) which recent distribution is restricted to one or two localities. About Schoenoplectus mucronatus and Cyperus longus, another species of periodically flooded habitats, are no recent data at all.
Redirection of river beds and their consecutive erosion and eutrophication and invasion of alien species in riparian habitats reduce the suitable conditions of C. glomeratus. In the IUCN Red List of threatened species it is
classed as least concern (LC) (Kavak 2014). In the actual Red List of the Slovak flora C. glomeratus is included as critically endangered (CR) (Elias jun. et al 2015) like in Switzerland (Welten & Sutter 1982). In the surrounding countries, C. glomeratus is regarded either as common species and it is not included in Red lists (Hungary — Kiraly 2007, Ukraine — Didukh 1999) or casual alien (Austria — Essl & Rabitsch 2002, the Czech republic — Pysek et al. 2012). In Croatia it is classed as vulnerable (VU) (Nikolic & Topic 2005) and in Slovenia as rare (R) (Anonymus 2002). It is protected by law only in Serbia (Anonymus 2010).
Acknowledgements
We thank to Zoltan Barina, Daniel Pifko, Norbert Riezing, Ranko Peric, Filip Verloove and Laura Celesti-Grapow for their comments on the distribution of Cyperus glomeratus in South and Central Europe, to the staff of visited herbaria for their technical help. The study was supported by the grants VEGA No. 2/0003/12 and 1/0083/16.
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