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DISTRIBUTION AND CHROMOSOME NUMBER OF SONCHUS PALUSTRIS IN SLOVAKIA

M. Dudáš1*, P. Eliáš jun.2, D. R. Letz3, Z. Bártová4 and V. Kolarčik1 1Institute of Biology & Ecology, Faculty of Science, P. J. Šafárik University, Mánesova 23 SK-041 54 Košice, Slovakia; E-mail: [email protected]; [email protected], *corresponding author 2Department of Environment and Biology, Slovak University of Agriculture Tr. A. Hlinku 2, SK-949 76, Nitra, Slovakia; E-mail: [email protected] 3Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences Dúbravská cesta 9, SK-845 23 Bratislava, Slovakia; E-mail: [email protected] 4Institute of Geotechnics, Slovak Academy of Sciences Watsonova 45, SK-040 01, Košice, Slovakia; E-mail: [email protected]

(Received: 6 March 2019; Accepted: 16 July 2019)

The distribution of Sonchus palustris in Slovakia was studied using herbarium specimens and literary sources. The herbarium studies, supplemented with targeted field search in the years 2015–2018, revealed 61 new localities and confirmed many other older locations. The species has been recorded in 19 phytogeographical districts and sub-districts of Slova- kia. Most of the records are concentrated in the Podunajská nížina lowland in SW Slovakia and in the Ipeľsko-rimavská brázda region in southern Slovakia. Our results showed that the species is relatively common in different types of wetlands and its re-evaluation in the recent version of the Slovak red list is not needed. Chromosome number data for two new populations in eastern Slovakia (both 2n = 18) were counted. The distribution map is given.

Key words: central Europe, marsh sow-thistle, marsh vegetation, occurrence, phytogeog- raphy, railway embankment

INTRODUCTION

According to Boulos (1973, 1974a, b) there are more than 50 species within the genus Sonchus L. (Asteraceae, Cichorieae, Sonchinae) arranged into three subgenera with centre of distribution in Europe, Northern America and Mid- dle Asia (Meusel and Jäger 1992). Sonchus palustris L. (syn. Sonchus paluster L.) naturally occurs in temper- ate, non-Mediterranean Europe from northwestern England to the Caucasus and Uzbekistan (Lansdown 2014, Meusel and Jäger 1992). It has also been reported from Pakistan (Qureshi et al. 2002). The species is considered as ex- tinct in Switzerland (Bornard et al. 2016). Secondary occurrence was found in North America (Brunton and Crompton 1993). The research based on ITS sequences supports the inclusion of S. palustris to highly polyphyletic subgenus Sonchus, section Maritimi and series Palustres (Kim et al. 2007).

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The oldest literature record of S. palustris was published by Genersich (1798) from northeastern Slovakia from damp places and fens in the Spiš (= Zips) region. 30 years later, Endlicher (1830) published finding in the surroundings of Bratislava (southwestern Slovakia). Hazslinszky (1847) states more precisely the locality in the Spiš region near the village of Lipovce (= Lipovec, Lipócz). Other authors published few data on the species’ occurrence in southwestern (Knapp 1863, Résely 1867, Reuss 1853, Schiller 1864), northeastern (Hazslinsz- ky 1853, Kárpáti 1939), western (Kárpáti 1939, Krzisch 1857, Mikeš 1938) and southeastern (Hazslinszky 1866, Hulják 1942) parts of Slovakia. Neilreich (1866, 1870) summarised the information available till that time on the distribution of S. palustris for the whole territory of historical Austro-Hungary (incl. Slovakia). From the end of World War I to 1970, the distribution of S. palustris in south- ern Slovakia was described by Domin and Podpěra (1928), Hendrych (1963, 1968) and Holub and Moravec (1965) and marginally mentioned in the works of Šomšák (1963), Zahradníková-Rošetzká (1965) and Berta (1970). In the last decade, the research of aquatic and marsh vegetation has be- come a point of interest in most parts of Slovakia (e. g. Kochjarová et al. 2012, Bubíková et al. 2016, and many others). Despite the fact that many regional works deal with aquatic vegetation, the information on the occurrence of Son- chus palustris in natural as well as anthropogenic water habitats are very rare (Balázs 1996, Hrivnák 1999). From the end of 18th till to first quarter of 20th century the data about the distribution of S. palustris were published more or less randomly. Between the years 1960–2005, the published distribution data (and collected herbarium specimens) are more frequent. Exploratory field excursions in southern part of central Slovakia in the basins of Rimava and Slaná Rivers (Kliment et al. 2000, Kochjarová 2010) brought new populations findings. The distribution continues here along the Ipeľ (= Ipoly) and Slaná (= Sajó) rivers to northern and central Hungary, where the species is commonly occurring (cf. Bánkuti and Vojtkó 1995, Bartha et al. 2015, and our observations). In western Slovakia, the occurrence in the Záhorská nížina lowland is tied up to distribution in southern Moravia and eastern Lower Austria. In Moravia, only three areas in the present time around the towns of Hodonín, Mikulov and Tišnov are known (Křísa 2004). Jongepier and Jongepierová (2006) and Perný (2007) mentioned S. palustris from the Biele Karpaty Mts as rare in waterlogged places along streams, mainly in southwestern part of the mountains. However, the detailed extension of the species has not been processed so far in Slovakia. In the latest version of the Slovak red list of ferns and flowering plants of Slovakia (Eliáš et al. 2015) the species is evaluated by the category NT (near threatened).

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This paper provides a detailed distribution of Sonchus palustris in Slova- kia, including historical, as well as recently confirmed locations and reassess- ing its conservation status based on field observation. We analysed chromo- some number in material from two localities in Slovakia.

MATERIALS AND METHODS Collecting of distribution data

The study and field research was carried out during the years 2015–2018. The data concerning the distribution of the species were found only in 14 pub- lic herbaria: BP, BRA, BRNU, HLO, KO, LTM, OL, OLM, NI, SAV, SLO, TM, TTM and W. Herbarium acronyms follow Thiers (2019) and Vozárová and Sutorý (2001) for small local collections. The species’ records were collected also from published and unpublished studies and botanical databases (the file database stored in the Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Bratislava, Slovakia; distribution data from habitats and species mapping of Slovakia (Šefferová Stanová et al. 2015; http://www.biomonitor- ing.sk/); the central database of phytosociological relevés (http://ibot.sav.sk/ cdf/)). The distribution map was designed in the program ArcGIS, version 9.2. The mapping grid follows the grid template described by Niklfeld (1971). A list of revised specimens and other records were compiled according to the di- rectives of the work Flóra Slovenska VI/1 (cf. Goliašová and Michalková 2012).

Karyology

For the karyological studies, the seeds of S. palustris were collected at two localities: (1) Šarišská vrchovina hill area, Hendrichovce (col. M. Dudáš in 2016, voucher specimen deposited in KO: 34307) and (2) Spišské kotliny Ba- sins, Domaňovce (col. V. Kolarčik in 2017). The root-tip meristems of germi- nating seeds were used for chromosome counting. Root tips were pre-treated in 0.002 M aqueous solution of 8-hydroxyquinoline (16 hrs at 4 °C), fixed in the mixture of 96% ethanol and 98% acetic acid (3:1 for 1 hr), then washed with distilled water, macerated in 1 M HCl in water bath (60 °C, 5 min) and washed again. The slides were made using the cellophane square method (Murín 1960). The slides were stained by a 7% solution of Giemsa Stain, Modified Solution, Fluka Analytical, in Sörensen phosphate buffer, dried and observed in a drop of immersion oil. For the observation of chromosomes microscope Leica DM 2500 equipped with camera HDCE-X5 and software ScopeImage 9.0 was used.

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RESULTS Distribution and ecology

Altogether, we have seen over 90 herbarium specimens and collected at least 30 published and unpublished records not supported by herbarium specimens. Our field research revealed 61 new localities and confirmed at least twelve historical locations. The detailed locations of S. palustris in Slova- kia are reported in the Appendix 1. Sonchus palustris has been found in Slovakia altogether in 19 of total of 31 phytogeographical districts and subdistricts. Seven of them are included in the area of the Pannonian flora province (Pannonicum) and twelve in the area of the Carpathian flora province (Carpaticum).

Karyology

Seedlings from the localities of Hendrichovce and Domaňovce (eastern Slovakia) showed 2n = 2x = 18 as somatic numbers, being one of the pairs sat- ellite in both cases.

DISCUSSION Distribution

As the distribution map (Fig. 1) has shown, most of the species’ locali- ties are situated in the southern part of the country (the Podunajská nížina lowland, the Ipeľsko-rimavská brázda Region, southern parts of the Sloven- ský kras Karst, the Košická kotlina Basin and the Východoslovenská nížina lowland) where the species grows in marshes predominantly along railway embankments, streams, fishponds, canals, mining pit-ponds and alluvial meadows. From lowlands and basins of the Pannonicum province in the southern Slovakia the species has penetrated into the neighbouring Carpathian prov- ince, where it is very rare, but locally it may form abundant populations. In the Carpathian province, S. palustris prefers larger warm valleys and basins at lower altitudes. From these sides it always spreads in ditches and wet places along railway embankments and roads. Isolated occurrence in northeastern Slovakia is represented by small local enclaves. One of them, northern of the village of Domaňovce (Spišské kotliny basins) reaches the altitudinal maxi- mum in 439 m a. s. l. in Slovakia. Historical altitudinal maximum, known at the village of Lipovce at the altitude of ca 530 m (Hendrych 1963) has not been confirmed at the present time by us.

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Fig. 1. Distribution map of Sonchus palustris in Slovakia

Fig. 2. Mitotic chromosomes of Sonchus palustris from the locality of Domaňovce, Slovakia, 2n = 2x = 18. Satellite chromosomes are marked with arrows

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Despite a lack of previous data in other areas, during our field research in southern Slovakia almost continual distribution along the railway embank- ments in line Rožňava – Číž – Jesenské – (in length of ca 50–60 km) was found. The plants have always grown here over 3 m high and wind gen- erated by passes of trains significantly contribute to anemochory spreading of the achenes. The railway embankments are associated with many marshes and wet places and this fact supports the spread of the species to further lo- calities along the railways. We noted the species rarely in eastern Lower Austria, mainly along canals, while the most localities were found around the lake Neusiedler See in large areas of Phragmitetum communities in the season of 2018. In the Podunajs­ká nížina lowland, the occurrence is fragmented in small islands, but intensive field study in the future surely brings new population findings. Evident decrease in number of localities was found in the Východoslo­ venská nížina lowland. There were published more than 10 localities of S. palust­ris in the past (Berta 1970, Bogoly 1992, Šomšák 1963). These sites have probably disappeared due to the drainage and riverbed modification of the Latorica River (Šimková et al. 2014). However, we confirmed only two of them (around the town of Kráľovský Chlmec located in oxbow of the Latorica Riv- er) and found only one new (on waterlogging alluvial meadows of the Onda- va River, on an area approximately 1 km2 with thousands of individuals). In Ukraine, the species was recently noted by Woch and Trzcińska-Tacik (2015) in western part of the country located on northern side of the Carpathians Mts, while on the southern side along the Latorica River, there is not known any information about a recent occurrence (V. Sabadosh 2016, pers. comm.), and Chopik and Fedoronchuk (2015) ascribed it as very rare. Isolated occurrence of the species in northeastern Slovakia is represented by three localities. The occurrence in the first locality at the village of Lipovce (Branisko Mts) was given already in the first half of 19th century (Hazslinszky 1847) and it was last time confirmed by Hendrych (1963) around the Salvátor spring. It has not been recently confirmed by us, despite the fact that large area of Phragmitetum community has been growing here until now. In the years 1958–1959 V. Pospíšil collected other Sonchus species here (the speci- mens deposited at BRNM), but his collection does not contain any specimen of S. palustris. The second population was recorded near Domaňovce in an alluvium of the Lodina stream (see also Karyology chapter). The third and most isolated occurrence is represented by a small population south of the town of Bardejov at the Kobyly village. Hazslinszky (1853) published the occurrence “in valleys of the rivers of Poprad and Hornád” (without exact locality). Although some suitable large areas with common reed (Phragmites australis) are present along the Hornád and Svinka Rivers, the occurrence of S. palustris in alluvium of the Poprad River has surely little probability (cf.

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Chrtek and Skočdopolová 1986, Mártonfi 1992). Small number of populations in northeastern part of Slovakia follows rare occurrence in southeastern Po- land, where the most of the species distribution is concentrated in northwest- ern and central part of the country (Zając and Zając 2001).

Karyology

The chromosome number 2n = 18 from two locations (Domaňovce, Hen- drichovce) agree with previously published data from western Slovakia (Má- jovský 1976) and as in material of diverse origin as from Europe and Caucasus (Goldblatt and Johnson 2019) and Northern America (Brunton and Cromp- ton 1993). Therefore it seems, there are no different cytotypes at least within the European distribution range of the species. Karyological analysis of both populations further consistently showed, that one chromosome pair is bear- ing chromosome satellite. Presence of 1–3 satellite chromosomes in karyotype of Sonchus species with 2n = 18 is very common, which may or may not be of taxonomic relevance in particular groups (Mejías and Andrés 2004; Mejías et al. 2012, 2018).

Conservation status

In the last version of the Slovak red list (Eliáš et al. 2015) the species is evalu- ated as near threatened (NT). Considering the common occurrence of this spe- cies in southern Slovakia, conservation status should be reconsidered. On the other hand, although S. palustris is quite common in some regions of Slovakia, its occurrence has declined significantly or the populations have become restricted to small number of individuals in some cases. Probably, the distribution range of S. palustris in Slovakia was underestimated in the past, which led to the over- estimation of the species threat. Methodologically rigorous mapping and demo- graphic analyses for the area required to the accurate evaluation of their conser- vation status (Jensen and Meilby 2012, U.S. Fish and Wildlife Service 2011). Curiously, the species is evaluated as “rare” with category C2b in the Czech Republic (Grulich 2012), while it is protected but not considered to be endangered in Hungary (Király 2007). Finally, S. palustris is ranked in the LC category (least concern) in the global IUCN red list of threatened species. It is widespread with stable populations and does not face any major threats (Lansdown 2014).

*

Acknowledgements – This work was partially supported by grants VEGA 2/0137/17 (Letz), VEGA 1/0047/19 (Eliáš) and internal grant VVGS-PF-2018-751 (Dudáš). We are grateful to

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L. Mártonfiová (Košice, Slovakia) for preparing and imaging the chromosome samples and to V. Sabadosh (Uzhhorod, Ukraine) for his help with literature and distribution data in Ukraine.

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Kochjarová, J., Hrivnák, R. and Oťaheľová, H. (2012): Príspevok k poznaniu výskytu vod- ných a močiarnych rastlinných spoločenstiev na východnom Slovensku. – Bull. Slov. Bot. Spoločn. 34(2): 189–212. (in Slovak with an abstract in English] Krzisch, J. F. (1857): Phanerogame Flora des Oberneutraer Comitates. – Verh. Ver. Naturkde. Presburg 2: 19–108. Křísa, B. (2004): Sonchus L. – mléč. – In: Slavík, B. and Štěpánková, J. (eds): Květena České republiky, Vol. 7. Academia, Praha, pp. 536–540. [in Czech] Lansdown, R. V. (2014): Sonchus palustris. – The IUCN red list of threatened species 2014: e.T19617543A19620971. Website https://doi.org/10.2305/IUCN.UK.2014-1.RLTS. T19617543A19620971.en. [accessed 04 February 2019]. Májovský, J. (ed.) (1976): Index of chromosome numbers of Slovakian flora (Part 5). – Acta Fac. Rer. Natur. Univ. Comen., Bot. 25: 1–18. [in Slovak] Mártonfi, P. (1992): Flóra okresu Stará Ľubovňa. – Univerzita P. J. Šafárika Košice a OÚŽP Stará Ľubovňa. [in Slovak] Mejías, J. A. and Andrés, C. (2004): Karyological studies in Iberian Sonchus (Asteraceae: Lactuceae): S. oleraceus, S. microcephalus and S. asper and a general discussion. – Folia Geobot. 39: 275–291. https://doi.org/10.1007/bf02804782 Mejías, J. A., García del Rey, M. and Silva, J. L. (2012): Variability in prickly sow-thistle (Sonchus asper) from western Mediterranean region. – Bocconea 24: 285–293. Mejías, J. A., Chambouleyron, M., Kim, S. H., Infante, M. D., Kim, S. C. and Léger, J. F. (2018): Phylogenetic and morphological analysis of a new cliff-dwelling species re- veals a remnant ancestral diversity and evolutionary parallelism in Sonchus (Astera­ ceae). – Plant Syst. Evol. 304: 1023–1040. https://doi.org/10.1007/s00606-018-1523-2 Meusel, H. and Jäger, E. J. (1992): Vergleichende Chorologie der zentraleuropäischen Flora. Band. III. – Gustav Fischer Verlag, Jena. Mikeš, J. (1938): Flora des Bezirkes von Bratislava und Malacky. – Vlastived. Sborn. Okr. bratislavského a malackého, Bratislava 3: 29–176. [in Slovak] Murín, A. (1960): Substitution of cellophane for glass covers to facilitate preparation of permanent squashes and smears. – Stain Technol. 35: 351–353. Neilreich, A. (1866): Aufzählung der in Ungarn und Slavonien bisher beobachteten Gefäss­ pflanzen. – W. Braumüller, Wien. Neilreich, A. (1870): Aufzählung der in Ungarn und Slavonien bisher beobachteten Gefäss­ pflanzen. Nachträge und Verbesserungen. – W. Braumüller, Wien. Niklfeld, H. (1971): Bericht über die Kartierung der Flora Mitteleuropas. – Taxon 20: 545– 571. https://doi.org/10.2307/1218258 Perný, M. (2007): Vegetácia Chocholanskej, Melčickej a Kochanovskej doliny v Bielych Kar- patoch. – Bull. Slov. Bot. Spoločn. 29(Suppl. 1): 103–142. [in Slovak with an abstract in English] Qureshi, S. J., Awan, A. G., Khan, M. A. and Bano, S. (2002): Taxonomic study of the ge- nus Sonchus L. from Pakistan. – J. Biol. Sci. 2(5): 309–314. https://doi.org/10.3923/ jbs.2002.309.314 Résely, M. (1867): Zur Flora der Insel Schütt in Ungarn. – Oesterr. Bot. Zeitschr. 17: 52–53. Reuss, G. (1853): Května Slovenska. – F. Lorber, Banská Štiavnica. [in Slovak] Schiller, S. (1864): Correspondenz. Neutra, den 6. November 1864. – Oesterr. Bot. Zeitschr. 14: 386–387. Šefferová Stanová, V., Galvánková, J. and Rizman, I. (eds) (2015): Monitoring rastlín a biotop- ov európskeho významu v Slovenskej republike. Výsledky a hodnotenie za roky 2013–2015. – Štátna ochrana prírody Slovenskej republiky, Banská Bystrica. [in Slovak]

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Šimková, A., Plačková, A. and Molitoris, Ľ. (2014): Všeobecná prírodovedná charakteris- tika okresu Trebišov. – Bull. Slov. Bot. Spoločn. 36(Suppl. 1): 5–25. [in Slovak with an abstract in English] Šomšák, L. (1963): Močiarna vegetácia medzidunových zníženín južnej časti Potiskej nížiny. – Acta Fac. Rer. Natur. Univ. Comen., Bot., Bratislava 8(5–6): 229–302. [in Slovak] Thiers, B. (2019): Index Herbariorum: A global directory of public herbaria and associated staff (continuously updated). – New York Botanical Garden’s Virtual Herbarium, New York. Website http://sweetgum.nybg.org/ih/ [accessed 10 February 2019]. U.S. Fish and Wildlife Service (2011): Astragalus desereticus, Deseret Milk-vetch, 5-year review: summary and evaluation. Utah Field Office. – Ecological Services, West Valley City, Utah 84119, 28 pp. Vozárová, M. and Sutorý, K. (2001): Index herbariorum Reipublicae bohemicae et Reipub- licae slovacae. – Bull. Slov. Bot. Spoločn. Suppl. 7: 1–95. [in Slovak with an introduc- tion in English] Woch, M. W. and Trzcińska-Tacik, H. (2015): High occurrence of rare inland halophytes on post-mining sites in western Ukraine. – Nord. J. Bot. 33: 101–108. https://doi. org/10.1111/njb.00607 Zahradníková-Rošetzká, K. (1965): Geobotanická charakteristika slatinných lúk a pasien- kov (Molinion Koch 1926) na Žitnom ostrove. – Biol. práce SAV, Bratislava 11(5): 5–45. [in Slovak with an abstract in English] Zając, A. and Zając, M. (eds) (2001): Distribution atlas of vascular plants in Poland. – Pra- cownia Chorologii Komputerowej Instytutu Botaniki Uniwersytetu Jagiellońskiego, Kraków, 716 pp. [in Polish and English]

Appendix 1. List of the revised Sonchus palustris localities in Slovakia

For herbarium specimens the collector, year of collection and herbarium acronym are given; herbarium codes follow Thiers (2019). References for published records from sources not listed in the References chapter are given in an abridged form including the page of a particular Sonchus palustris record. For unpublished field records the name(s) of its author(s) and year are given, followed by the notavit (not.) abbreviation. The records are arranged following the phytogeographical division of Slovakia by Futák (1980). In some instances place names are given in the original language in parentheses. Abbreviations: S (south), N (north), E (east), W (west) and its combinations.

Panonnicum

Matricum. 1. Burda Hills: Chľaba, valley of Ipeľ River (Svobodová Bull. Slov. Bot. Spoločn. 11: 22, 1989). 2. Ipeľsko-rimavská brázda Region: Santovka, valley Studená do- lina (Schidlay 1963 SAV). – Santovka, SE, marsh (Svobodová 1970 NI; Svobodová Acta Fytotechn. 23: 12, 1972). – Santovka, Malinovec, Búr stream (David 1984 LTM; Dudáš 2017 KO). – Bajtava, water reservoir Bajtava (Dudáš 2017 not.). – Kamenica nad Hronom, stream Bajtavský potok (Dudáš 2017 KO). – Balog nad Ipľom, alluvium of Galomia stream (Dudáš 2018 not.). – Ružiná, water reservoir Ružiná, NW bank (Belanová 2008 sec. biomonitoring. sk; Letz 2014 SAV). – Lovinobaňa, marsh between the stream Krivánsky potok and Road no. 16 (Dudáš 2016 KO; Letz 2016 SAV). – Veľké Dálovce, marsh Dálovský močiar (Hrivnák 1999: 76). – (Letz 2016 not.). – Čamovce, stream Čamovský potok (Dudáš 2016 KO).

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, S, near stream (Hendrych 1963: 51; Hendrych 1968: 169). – Gemerský Jablonec, marsh (Dudáš 2016 KO). – Dubno, stream (Dudáš 2016 BRNU). – Hajnáčka, Gortva stream (Eliáš jun. 2015 NI; Dudáš 2016 KO). – Ožďany, settlement Ružová osada, alluvium of stream Maštínsky potok, near its mouth into water reservoir Ožďany (Dudáš 2017 KO). – Ožďany, Suchá River (Dudáš 2016 BRNU). – Hodejov, marsh under railway embankment (Dudáš 2017 not.). – Ostrany, alluvium of Blh River near first bridge. – Drienčany, valley of Blh River, wet meadows. – , secondary wetland in village. – Hostišovce, con- fluence of stream Kamenistý potok with unnamed stream inflowing from village, Dolné lúky site (all Kliment et al. 2000: 141). – Rimavská Sobota, Košická cesta Street, Rimava Riv- er, wetland at roadside (Dudáš 2018 not.). – Rimavská Sobota, part Sabová-Rákoš-Vrabčie, near stream Močiar, under Road no. 16 (Dudáš 2016 KO; Letz 2016 SAV). – Hodejov, NE, marsh (Eliáš jun. 2015 NI). – , stream Mačací potok and water reservoir Hostice (Dudáš 2016 KO). – Drňa, stream Drniansky potok (Lepší in Kochjarová 2010: 63). – Drňa, pastures towards Biriň hill (Valachovič 2012 SAV). – Drňa, E, valley Vlčia dolina (Balázs Ochr. Prír 14: 31, 1996). – Chrámec, under Pohanský hrad (Hendrych l. c. 1963, 1968). – Hostice, settlement Kozí výhon [Kecskejáras] (Hendrych l. c. 1963). – settlement Tomášova dolina, near stream (Holub & Moravec 1965: 73). – between village of Drňa and settlement Jalová [Jalová pusta] (Holub & Moravec l. c.). – Drňa (Hendrych l. c. 1963). – Drňa, S (Hen- drych l. c., 1968). – Petrovce, E (Hendrych l. c., 1968). – Petrovce, settlement Voziarsky buk [Szekeresbükk]. – Petrovce, gamekeeper’s cottage Raďaš (Hendrych l. c., 1963). – Petro- vce, stream Mačací potok (Dudáš 2016 KO). – Tornaľa [Šafárikovo], marsh under cemetery (Řehorek Acta Fytotechn. 13: 184, 1966). – Tornaľa, NW, left bank of Slaná River (Trávníček in Kochjarová l. c.). – Teška settlement, Teška stream (Dudáš 2016 KO). – Figa, wet ditch on eastern part of the village (Dudáš 2016 KO). – Figa, marsh in the west part of the village (Dudáš 2016 KO). – Včelince, Kerepec, roadside, in common reed (Dudáš & Bártová 2016 KO). – Rimavská Seč, Blh River, near railway embankment (Dudáš 2016 not.; Dřevojan in Eliáš, Bull. Slov. Bot. Spoločn., 40/2: 182, 2018). – Martinová, water reservoir Martinová (Magic Prehľad odb. výsl. XVII. tábora ochr. prír., 1985, p. 52; Dudáš 2016 KO). – Číž, water reservoir Číž (Rydlo in Kochjarová l. c.; Dudáš & Bártová 2016 KO). – between the villages of Číž and (Svobodová l. c., 1972). – Riečka, N, wetland in field (Dudáš 2018 not.). – Janice, valley under Hill 299 (Hendrych l. c., 1963). – Janice, S, 200 m a. s. l. (Hend- rych l. c., 1968). – Janice, stream Pohanský potok (Neumannová 2002 sec. biomonitoring. sk). – Čikovo settlement, marsh (Holub & Moravec l. c.). – settlement Teplá dolina, near stream (Holub & Moravec l. c.). – Vlkyňa, W, field edge (Boublík et al. in Kochjarová l. c.). – Vlkyňa, stream Vlkyňský potok (Lippl in Kochjarová l. c.). – Vlkyňa, Mozik valley under Hill 195 (Hendrych l. c., 1963: 47). – Gemerská Hôrka, train station (Eliáš jun. & Sádovský in Kochjarová l. c.). – Gemerská Hôrka, near railway on southern part of the village (Rydlo in Kochjarová l. c.). – Gemerská Hôrka, along railway embankment from station towards Slaná River (Dudáš & Bártová 2016 KO). – Starňa, valley of Lapša stream, spring Remetský prameň (Sitášová in Kochjarová l. c.). – Gemer, wetland near railway (Dudáš & Bártová 2016 KO). – Gemerská Panica, train station (Rydlo in Kochjarová l. c.). – between the vil- lages of Čoltovo and Gemerská Panica, marshes between road and railway (Dudáš & Bár- tová 2016 KO). – Starňa, valley of Činča stream (Košťál & Kochjarová in Kochjarová l. c.). – Starňa, forest Panský les (Sitášová in Kochjarová l. c.). – Lenka, stream Hubovský potok (Dudáš & Bártová 2016 KO). – , pond Hubovský rybník (Košťál et al. in Kochjarová l. c.; Dudáš & Bártová 2016 KO). – Neporadza, in the village (Mereďa jun. & Májeková in Kochjarová l. c.). 3. Slovenský kras Karst: Pašková, site Za záhradami (Virók in Virók et al., Flóra Gem.-turn. krasu 2016: 729). – Plešivec, site Breziny (Virók in Virók l. c.; Dudáš 2017

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Unauthenticated | Downloaded 06/10/20 05:49 AM UTC DISTRIBUTION AND CHROMOSOME NUMBER OF SONCHUS PALUSTRIS 19 not.). – Vidová, Slaná River, ditch along Road no. E571 (Dudáš & Bártová 2016 KO; Virók in Virók l. c.). – Slavec, S, marsh (Svobodová l. c. 1989). – Plešivec, crossroad in the village, wet ditch (Dudáš & Bártová 2016 KO; Dudáš in Eliáš jun. Bull. Slov. Bot. Spoločn. 38/2: 295). – Brzotín, Čremošná River [Berzéte, Csermosnya] (Hulják 1941 BP; Hulják 1942: 251). – Brzotín, 1 km SW from train station (= ponds Brzotínske rybníky) (Trávníček 1996 OL; Dudáš 2016 KO). – Brzotín, SE, wetland near railway embankment (Trávníček Bull Slov. Bot. Spoločn. 18: 74, 1996). – Jovice, roadside (Dudáš 2016 BRNU). – Silica, site Čobáňova bučina (Barina in Virók et al. l. c.). – Silica, site Silické jazero (Virók in Virók et al. l. c.). – Jablonov nad Turňou, bus stop near industrial park, wet ditch (Dudáš 2016 not.). Eupannonicum. 4. Záhorská nížina lowland: Suchohrad, Morava, river km 38 (Vorlíčková 1994 sec. biomonitoring.sk). – Jakubov, ponds Jakubovské rybníky (Koso- rínová 1998 sec. biomonitoring.sk). – Plavecký Štvrtok, wet ditch on field edge (Futák 1960 SAV; Futák in Hendrych l. c., 1963). – Holíč, woody fens at the town (Krzisch 1857: 71). – Kopčany, gravel-pit Boričky I. (Dudáš 2017 KO). – Kopčany, bank of gravel pit (Očka 2010 TM). – Gbely [Egbell], Adamov, at Adamov forest (Scheffer 1924 SLO). – Gbely [Eg- bell] (Scheffer in Kárpáti, Borbásia 1/3–7: 64, 1939). – Bilíkove Humence E, Rudava River (Malovcová 1999 sec. biomonitoring.sk). 6. Podunajská nížina lowland: Bratislava (Szép 1907 BRA). – Bratislava, island Pečenský ostrov, marsh meadows (Endlicher 1830: 297; Mikeš 1938: 139). – Svätý Jur, Šúr [Schur] (Endlicher l. c.; Ptačovský 1922 SAV; Holuby s. dato SLO; Holuby Věda Přír. 7: 114, 1926). – Hrnčiarovce, water reservoir (Thomka 1973 TTM). – Veľký Grob, pond Rašelinisko Veľký Grob (Dudáš 2017 KO). – Pusté Uľany, dam (Zlatošová 1994 TTM; Šípošová & Komárová 1996 SAV). – Jurová [Dercsika]. – Gabčíkovo [Böös] (both Resely 1867: 53). – Dunajská Streda (Schiller 1864: 386). – Podolie, stream Obu- chovský potok (Válková 2011 sec. biomonitoring.sk). – Trebatice, Holeška stream (Vavro 1978 HLO). – Jalšové [Jelšové], near Váh River (Vavro 1986 HLO). – Trakovice, Horný Dudváh River (Malovcová 2017 HLO; Malovcová in Eliáš Bull. Slov. Bot. Spoločn. 40/1: 93, 2018). – Tešedíkovo, pond Bystré jazierko (Szabóová 2002 sec biomonitoring.sk). – Dolný Bar, dead oxbow, bog (Zahradníková-Rošetzká 1965: 14). – Kľúčovec, oxbow Kľúčovské rameno (Szabóová 2010 sec. biomonitoring.sk). – Merašice, stream Merašický potok (Kol- lár et al. in Eliáš, Bull. Slov. Bot. Spoločn. 31/2: 119, 2009). – Trnovec nad Váhom, Trnovec stream. – Vlčany, lake Bábske jazero. – Vlčany, dead oxbow Vlčianske mŕtve rameno. – Čalovec, canal Ektovský kanál (all Szabóová 2002 sec. biomonitoring.sk ). – Klokočina, stream in west part of village (Kvapilík 1930 OLM). – Cabaj-Čápor, water reservoir Čápor (Svobodová & Řehořek 1993 NI). – Vrbová nad Váhom, lake Lohótske jazero. – Martovce, locus Gémeš, Tátoš stream (both Szabóová 2002 sec. biomonitoring.sk ). – Martovce, Stará Žitava River (Magic 2003 sec. biomonitoring.sk ). – between the villages of Lándor and Martovce, site Gamota (Zlinská & Kubalová, Acta Envir. Univ. Comen., Bratislava 11: 89, 2001). – Komárno (Reuss 1853: 265). – Svinná, outfield under road, opposite to collective farm (Dudáš 2017 not.). – Dubnička, meadow near Dubnička stream (Bagin 2015 http:// www.fotonet.sk/, ID 124335; Dudáš 2017 KO). – Brezolupy, confluence of streams Hydina and Miezgovský potok (Dudáš 2017 not.). – Nitra, fens at the town. – Golianovo [Lapásg- yarmat] (both Knapp 1863: 156). – Brezolupy, Jerichov, alluvium of Hydina stream (Dudáš 2017 not.). – Partizánske, Veľké Bielice, thermal spring (Štěpánková & Štech in Ambros, Ro- salia, 1996: 129; Eliáš jun. 2002 NI). – Partizánske, Malé Bielice, site Bahná (Řehořek in Am- bros l. c.) = SKÚEV Bielické bahná (Dudáš 2017 KO). – Nevidzany (Svobodová 1993 NI). – Nevidzany, alluvium of stream Podegerský potok over station (David 1986 LTM). – Nev- idzany, pond (David & Vozárová, Biológia 45/5: 445, 1990). – Beša, marsh near train station (Valenta 2000 BRA; Eliáš jun. 2018 NI). – Beša, wetland and wet meadow (Smetanová 2000

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LTM). – Vojnice, marsh ca 2 km towards Mužla (Svobodová l. c., 1989). – Búč, SW part, towards Starý vrch hill (Mereďa jun. 2012 SAV; Eliáš jun. 2017 NI). – Búč, pond Búčsky ryb- ník (Mereďa jun. 2012 SAV). – Bíňa, Kuralanka stream (Svobodová l. c., 1989). – Gbelce, site Parížske močiare (Magic Ochr. Prír 16: 73, 1998). 7. Košická kotlina Basin: Hrhov, bus stop (Dudáš 2017 KO). – Hrhov, ponds Hrhovské rybníky (Sitášová 2007 http://www.fotonet. sk/, ID 8860; Dudáš 2016 KO; Virók in Virók et al. l. c.). – Turňa nad Bodvou, pond Turni- ansky rybník (Dudáš & Bártová 2016 KO; Dudáš in Eliáš, Bull. Slov. Bot. Spoločn. 38/2: 295; Dudáš & Kolarčik 2018 not.). – Turňa nad Bodvou, site Pri dvorníckej ceste (Dudáš 2016 not.; Virók in Virók et al. l. c., p. 730). – Cestice, part Dobogov, Penzión Dubky, alluvium of unnamed stream (Dudáš 2017 not.). – Nižný Lánec, mouth of stream Kanský potok into water reservoir Lánec (Dudáš 2016 not.). – ponds Perínske rybníky, canal between ponds (Dudáš 2016 not.). – Košice, Myslava (Mikoláš 1989 W). – Šebastovce, wet ditch under road (Dudáš 2016 not.). – Haniska, part Grajciar, canal Grajciarsky kanál, wetland in field (Dudáš 2017 not.). – Čaňa, lakes Čanianske jazerá (Dudáš 2017 not.). – Veľká Ida, canal Veľkoidiansky náhon (Dudáš 2017 KO). – Seňa, water reservoir Seňa (Dudáš 2017 KO). – Seňa, alluvium of stream Belžianský potok (Dudáš 2018 not.). – Chym, wet ditch, roadside (Dudáš 2016 not.). 8. Východoslovenská nížina lowland: Zemplínske Hradište, Trnáv- ka River, marsh under dam (Dudáš 2016 KO). – Hraň, between Trnávka River and canal Neľovský kanál, wet meadows (Dudáš 2016 KO, BRNU; Dudáš in Eliáš jun. Bull. Slov. Bot. Spoločn. 38/2: 295). – Ižkovce, forest Spodný les (Berta 1970: 222, tab. 49; Dudáš did not confirm in 2016 and 2018). – dead oxbow Tice, in village near road [Bogoly, (Msc.) depon in ŠOP SR, Správa CHKO Latorica, Trebišov, 1994]. – Boľ, dead oxbow (Dudáš 2016 KO). – Solnička, Tice (Šomšák 1963: 283-285, Tab. 11). – Kráľovský Chlmec, near road in direction to Veľké Kapušany, dead oxbow Tice (Šomšák l. c.). – Plešany, site Veľké jazero, central part (Šomšák l. c., Tab. 10). – Boľ, bog [s. coll. 1984 SAV; Bogoly l. c., 1994; Bogoly (Msc.) de- pon. in Ľudová akadémia „László Mécsa“, Kráľovský Chlmec, 1996]. – Kráľovský Chlmec, dead oxbow Madaraška (Bogoly l. c., 1994; Šimková 2014 photodocumented; Dudáš 2016, 2018 not.). – Strážne, farm Ružový dvor, terrain depression between sand dunes (Šomšák l. c., Tab. 7). – Leles (Hendrych l. c., 1963: 47).

Carpaticum occidentale

Praecarpaticum. 9. Biele Karpaty Mts. (southern part): Melčice-Lieskové, valley Hrad- nianska dolina [Melčická dolina, Kamenná dolina] (Perný 2007: 173). 10. Malé Karpaty Mts.: Dobrá Voda, spring Výtok [Vítek] (Feráková 1972 SLO; Feráková in Májovský et al. 1976: 14, 2n = 18; Vavro 1983 HLO; Dudáš did not confirm in 2017). 12. Tribeč Mts.: Kolačno, valley of stream (Hadinec in Ambros l. c.; Dudáš did not confirm in 2017). 13. Strážovské and Súľovské vrchy Mts.: Uhrovské Podhradie, site Zrubisko (Futák 1960 ined). 14b. Vtáčnik Mts.: Čereňany, part Fančová, wetland (Lustyk in Ambros l. c.) = Čereňany, part Fančová, stream Čereniansky potok (Dudáš 2017 KO). 14c. Kremnické vrchy Mts.: Zvolen, Západ-Tepličky, north of Strážska cesta Street, stream Kováčsky potok (Letz 2016 SAV). – between the villages of Badín and Vlkanová, marsh under R1 highway (Letz 2016 SAV; Dudáš 2017, 2018 not.). 15. Slovenské rudohorie Mts.: Mýtna, stream Dobročský potok (Dudáš 2017 not.). – Sirk, part Červeňany, Východný Turiec River (Fráková 2017 not.). – Striežovce, valley of stream Striežovský potok. – Hrušovo, wet meadows (both Kliment et al. l. c.). – Ratková, Rybník, along Drienok stream (Č. Deyl 1978 OLM). – Rybník, valley of Drienok stream under canyon (Kliment et al. l. c.). – Nandraž, temporary fen over the vil-

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Unauthenticated | Downloaded 06/10/20 05:49 AM UTC DISTRIBUTION AND CHROMOSOME NUMBER OF SONCHUS PALUSTRIS 21 lage (Hajdúk 1969 BRA). – Kameňany, in the village (Lepš in Kochjarová l. c.). – Rožňava, train station Rožňava-mesto, between railway and roundabout crossroad (Dudáš & Bár- tová 2016 not.). – Rožňava, Slaná River, site Slanské lúky, floodplain forest (Dudáš 2017 KO). – Rožňava, stream Hasácky potok (Dudáš 2016 KO). 18. Stredné Pohornádie Valley: Lipovce [Lipovec, Lipócz] (Hazslinszky 1847: 223; Hendrych l. c., 1963). 19. Slanské vrchy Mts.: Nižná Hutka, marsh under Hill 273 (Dudáš 2017 KO; Dudáš in Dudáš et al., Thaiszia – J. Bot. 28/2: 152, 2018). Intracarpaticum. 26b. Spišské kotliny Basins: Domaňovce, confluence of Lodina stream with unnamed left-side tributary (Kolarčik 2016 photodocumented; Dudáš 2018 not.; 2n = 18). Beschidicum orientale. 30a. Šarišská vrchovina hill area: Hendrichovce, wet ditch, roadside and nearest ruderal site (Dudáš 2016 KO; Kolarčik 2018 not.; 2n = 18). – Prešov [Eperjes] (Hazslinszky in Kárpáti l. c.; Hazslinszky s. d. BP, BRA). 30c. Nízke Beskydy hill area: Kobyly, alluvium of Brôdok stream (Dudáš 2016 KO, 2018 not.).

General data (not mapped): Spiš region [= Zips], damp places and fens (Genersich 1798: 57). – valleys of the rivers of Poprad and Hornád (Hazslinszky 1853: 8). – south- ern part of the Zemplín [Zemplén] Comitat (= historical region), depressions (Hazslinszky 1866: 125). – Žitný ostrov [Csallóköz], Dunajská Streda [Szerdahely] (Resely 1867: 53).

Doubtful data (not mapped): Trnava, wet fields and field ditches (Frantová 1947: 172, 207). – Javorníky Mts., Dubie, dam of pond, gravelsands [Falťan, (Msc.) depon. in KFGG, PriF UK Bratislava. 74 pp.]. – Vyšný Žipov, dead oxbow of Topľa River (Terek et al., Zborn. odb. Výsl. XI. Vsl. Tábora ochr. prír., 1988, p. 18; Dudáš did not confirm in 2016) = PP (nature reserve area) Žipovské mŕtve rameno (Hudec & Stano, Nat. Carp. XXXVII: 124, 1996).

Acta Bot. Hung. 62, 2020

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