Distribution, Chromosome Numbers and Nomenclature Conspect of Arabidopsis Halleri (Brassicaceae) in the Carpathians

Biologia, Bratislava, 61/1: 41—50, 2006 Section Botany DOI: 10.2478/s11756-006-0007-y

Distribution, chromosome numbers and nomenclature conspect of Arabidopsis halleri (Brassicaceae) in the Carpathians

Martin Kolník1,3 & Karol Marhold1,2

1Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14,SK–84523, Bratislava, Slovakia; tel.: +421-2-5942 6128,fax:+421-2-5477 1948, e-mail: [email protected], [email protected] 2Department of Botany, Charles University, Benátská 2,CZ–12801, Praha 2, Czech Republic 3Institute of Biodiversity Conservation and Biosafety, Faculty of Agrobiology and Food Resources, Agricultural university, tr. A. Hlinku 2,SK–949 76 Nitra, Slovak Republic

Abstract: Arabidopsis halleri represents an important model species for the study of phytoremediation. In the Carpathians it is represented by three subspecies: A. halleri subsp. halleri, A. halleri subsp. tatrica and A. halleri subsp. dacica.Allthree subspecies are diploid with chromosome number 2n = 16. They differ mainly in indument of flower parts, colour of petals and in the position of the longest leaf on stem. A. halleri subsp. halleri occurs in the Eastern and Southern Carpathians and in the northern and eastern part of the Western Carpathians, subsp. tatrica is endemic of the Western Carpathians and subsp. dacica occurs in the Eastern and Southern Carpathians most probably extending in its distribution further to the south to the Balkan mountains. Full synonymy of these three taxa and also a fourth European subspecies, A. halleri subsp. ovirensis is presented including the information on type specimens. Lectotypes are selected for several names. List of the examined herbarium specimens is given as well. Key words: Arabidopsis, chromosome numbers, Cruciferae, distribution, taxonomy, typification

Introduction Arabidopsis halleri is reported from mountains of Central Europe extending to the SW Alps, N part of the Recent molecular studies showed that the traditional Balkan Peninsula and eastern Asia. Three subspecies taxonomical circumscription of genera Arabidopsis and are usually recognised within Arabidopsis halleri: A. Arabis is to a large extent artificial. Both genera were halleri subsp. halleri usually reported from Austria, the revealed to be polyphyletic, consisting of several in- Czech Republic, Germany, Switzerland, N and Central dependent lineages (Koch et al., 1999, 2003). Al- Italy, Poland, Slovenia, Croatia, Romania and Ukraine; Shehbaz et al. (1999) listed 59 binomials, previously A. halleri subsp. ovirensis from Austria, Italy, Slove- assigned to Arabidopsis, which are currently classified nia, Ukraine, Serbia, Albania and Romania; A. halleri in 14 different genera. On the other hand, taxa for- subsp. gemmifera from Russian Far East, NE China, merly treated in the genus Cardaminopsis (including Korea, Japan and Taiwan (Jones & Akeroyd, 1993; Arabidopsis halleri) are now to be included in the Jalas & Suominen, 1994; Al-Shehbaz & O’Kane, genus Arabidopsis, as well as several taxa, which were 2002). Populations from the Slovak Republic and S originally described within Arabis. At present, follow- Poland, especially those from the Tatry Mts., have of- ing the treatment by O’Kane & Al-Shehbaz (1997), ten been reported as intermediate between subsp. hal- the genus Arabidopsis consists of nine species, some of leri and subsp. ovirensis (e.g. by Jones & Akeroyd, which are further divided into subspecies. All Arabidop- 1993; Jalas & Suominen, 1994). They were described sis species in the circumscription of these authors are as a separate variety (Arabis halleri var. tatrica)by indigenous to Europe, with the ranges of two species Pawlowski (1930) and subsequently transferred to the extending into northern and eastern Asia and North subspecies rank as Cardaminopsis halleri subsp. tatrica America (Al-Shehbaz & O’Kane, 2002). In our cur- by Měsíček (2002). Although not so widely used as rent project we concentrate on the taxonomic revision closely related Arabidopsis thaliana, A. halleri is an im- of this genus in the Carpathian mountain range com- portant model species for the study of phytoremedia- paring also the Carpathian populations with those from tion. It is tolerant to high soil concentrations of zinc, the Eastern Alps. Previous studies on this genus (e.g. cadmium and lead and hyperaccumulates zinc and cad- Koch et al., 1999) concentrated mostly on phylogenetic mium (Bert et al., 2000, 2002, 2003; Van Rossum aspects, but thorough study combining molecular, mor- et al., 2004 and references therein). Numerous studies phometric and karyological data is still missing. have been devoted to this species, but only seldom at-

c 2006 Institute of Botany, Slovak Academy of Sciences 42 M. Kolník &K.Marhold

Fig. 1. Map of distribution of three subspecies of Arabidopsis halleri in the area studied based on revised herbarium specimens and own collections. + A. halleri subsp. halleri,#A. halleri subsp. tatrica, * A. halleri subsp. dacica. tention has been paid to its infraspecific variation and fixed in freshly prepared mixture of 96% ethanol and 99% to the correct infraspecific identity of the material stud- acetic acid (1:1) for 1 h at room temperature. Root tips ◦ ied (e.g. Bert et al., 2000; Fabiszewski, 1986). There- were stored at 75% ethanol at 4 C. Before squashing, the fore, results of the study by Kolník & Marhold (2005 root tips were rinsed in water for 10 min, then hydrolysed and in prep.) as well as the results presented here are in a mixture of concentrated hydrochloric acid and ethanol (1:1) for 2.5 min and rinsed in water for 10 min. Squashes important to clarify the taxonomic complexity of A. were made in a drop of 45% acetic acid under a celophane halleri. square (MURÍN, 1960; MARHOLD et al., 2002) and stained Based on the detailed morphometric, molecular in 10% solution of Giemsa stock dye in Sörensen phosphate and karyological evaluation of the Carpathians popu- buffer for 1.5 h. Details on localities of the analysed plants lations of A. halleri, using comparative material from are included in Table 1. the Eastern Alps and Japan, we confirmed the existence Morphological descriptions. Morphological descriptions of four subspecies within this species in Europe and one presented here are based on the results of detailed morpho- additional subspecies in E Asia (Kolník & Marhold, metric studies of populations of A. halleri collected through- 2005 and in prep.). Here we present nomenclature syn- out the Carpathian mountain range (KOLNÍK &MARHOLD, thesis of the Carpathian taxa, A. halleri subsp. halleri, 2005 and in prep.). Values in morphological descriptions rep- resent 5% and 95% percentiles, with 1% and 99% percentiles A. halleri subsp. tatrica,andA. halleri subsp. dacica, in brackets. Most important differentiation characters of the with the morphological descriptions and detailed distri- Carpathian subspecies of A. halleri are presented in Table 2. bution and karyological data. Nomenclature and kary- Distribution data. To assess the geographic distribution ological data on A. halleri subsp. ovirensis from the of the studied taxa we have studied herbarium specimens Eastern Alps are provided as well. from the following herbaria (herbarium acronyms follow HOLMGREN et al., 1990): BP, BRA, BUC, BUCA, CL, Material and methods KO,KRA,KRAM,LI,LW,LWS,LWKS,PR,PRC,PRA, SAV, SLO, W, WU, ZV. The list of localities is arranged Nomenclature. For each taxon nomenclatural (≡)and following the phytogeographical division of the Slovak Re- taxonomic (=) synonyms are presented, together with public (FUTÁK, 1966), Ukrainian Carpathians (ČOPYK et misidentifications and invalid names (–) reported mainly al., 1977), Polish Carpathians (KONDRACKI, 1981) and ad- from the area studied. For each name, beneath the heading ministrative districts of Romania. “Ind. loc.”, information from the protologue is cited con- cerning the distribution and localities of the original material. Data from the labels of the type specimens are pre- Taxonomic consp ect of Arabidopsis halleri in sented beneath the “Type” headings. Nomenclatural notes Central Europe. are included in some cases. Chromosome numbers. The diploid status of A. halleri Arabidopsis halleri (L.) O’Kane & Al-Shehbaz, was verified throughout the Carpathians. As is typical in Novon 7: 325, 1997. Brassicaceae, the small size of chromosomes prevents detailed karyotype studies. For chromosome number determi- ≡ nation we used seeds taken from herbarium specimens of Arabis halleri L., Sp. Pl. ed. 2: 929, 1763. plants collected in the field. Seedlings were taken in the mid- Ind. loc.: “Habitat in Harcynia ad Clausthal, locis dle of the day and after pretreatment in saturated solution humectis.” of 8-hydroxyquinoline for 3–5 h at 4 ◦C, the root tips were Lectotype (Al-Shehbaz in Cafferty & Jarvis Distribution, chromosome numbers and nomenclature conspect of Arabidopsis halleri 43

Table 1. Localities of populations of four subspecies of Arabidopsis halleri, from which chromosome numbers were analysed.

Taxon Locality Locality 2n/number code of analysed plants

A. halleri H-UC Ukraine, Ľvivska oblasť, Kozevo, beside a road, 48◦57 N, 023◦20 E, 18.5.2002, coll. KOLNÍK, 16/2 subsp. KAGALO,SYČAK halleri H-UI Ukraine, Ivano-Frankivska oblasť, Myslivka, beside a road, 48◦46 N, 023◦40 E, 23.5.2002, 16/2 coll. KOLNÍK,KAGALO,SYČAK H-UE Ukraine, Ľvivska oblasť, Staryj Sambir, along a road, 49◦20 N, 023◦01 E, 21.5.2002, coll. 16/2 KOLNÍK,KAGALO,SYČAK H-UJ Ukraine, Ivano-Frankivska oblasť, Vyškiv, meadow in village, 48◦43 N, 023◦11 E, 23.5.2002, 16/3 coll. KOLNÍK,KAGALO,SYČAK H-RC Romania, Bistrit¸a, Mt. Bˆarg˘au,s.Tihut¸a, 47◦14.995 N, 025◦01.418 E, alt. 1155 m, 22.5.2003 16/1 H-RD Romania, Bucovina, Mt. Bistrit¸iei, 47◦27.611 N, 025◦21.318 E, alt. 1046 m, 22.5.2003 cca 16 H-AUH Austria, Lower Austria, Zellenbach W of Guttenstein, along a stream, 47◦52.400 N, 16/1 015◦50.448 E, alt. 521 m, 29.4.2004, coll. KOLNÍK,MATSCHINGER H-AUK Austria, Lower Austria, Hohenberg castle near Mariazel, 47◦54.240 N, 015◦37.440 E, alt. cca 16 584 m, 29.4.2004, coll. KOLNÍK,MATSCHINGER H-KO Slovakia, Bukovské vrchy Mts., Kolbasov, on meadows, 49◦00.395 N, 022◦22.999 E, alt. 307 16/2 m, 1.6.2004 H-ITA Italy, prov. Trentino, Val di Breguzzo, 46◦02.142 N, 10◦38.183 E, alt. 1176 m, 8.6.2003, coll. 16/4 MARHOLD,LIHOVÁ,SLOVÁK

A. halleri O-AUO Austria, Carinthia, Ebriach, Mt. Hoch Obir, 46◦30.090 N, 014◦30.707 E, alt. 1557 m, 16/5 subsp. 25.8.2004, coll. KOLNÍK,MATSCHINGER ovirensis

A. halleri D-RUME Romania, F˘ag˘ara¸sMts.,Bˆalea Lac glacial lake, N slopes of Mt. Negoiu, 45◦35.918 N, 16/2 subsp. 024◦37.305 E, alt. 2095 m, 10.7.2003 dacica D-RUMF Romania, F˘ag˘ara¸s Mts., Saua Caprei glacial lake, 45◦36.118 N, 024◦37.687 E, alt. 2274 m, 16/1 10.7.2003

A. halleri T-BTK Slovakia, Belianske Tatry Mts., Biele pleso-Kopské sedlo, 49◦13.466 N, 020◦13.522 E, alt. 16/2 subsp. 1646 m, 21.6.2002 tatrica T-JDK Slovakia, Nízke Tatry Mts., Jánska dolina valley, along a path direct Mt. Krakova Hoľa, 16/2 49◦01 N, 019◦39 E, alt. 1050-1200 m, 19.6.2002 T-TZO Slovakia, Západné Tatry Mts., Mt. Osobitá, Teplý žľab, 49◦15.549 N, 019◦42.455 E, alt. 16/2 1204 m, 27.6.2002 T-MODH Slovakia, Belianske Tatry Mts., Monkova dolina valley, 49◦15.024N, 020◦13.670E, alt. 1200 16/3 m, 2.7.2003 T-KL Slovakia, Slovenský Raj Mts., Kopanecké lúky, 48◦53.435 N, 020◦16.747 E, alt. 1016 m, 16/7 19.6.2003 T-BKR Slovakia, Turzovská vrchovina Mts., Biely Kríž, 49◦29.516 N, 018◦32.516E, alt. 857 m, 16/3 7.7.2004 T-PLDH Poland, Tatry Mts., Mt. Kopa Kondraczka, 49◦14.221 N, 019◦55.879 E, alt. 1955 m, 2.8.2003, 16/8 coll. KOLNÍK,WALUSIAK,ONETTE

2002:531): “Sisymbrium [palustre album] foliis ≡ Arabis stolonifera (Scop.) Hornem., Hort. Bot. imis Barbareae reliquis”inHaller, Opusc. Bot.: Hafn. 2: 618, 1815. [“t. 1, f. 1”], 1749. = Arabis tenella Host, Fl. Austriac. 2: 273, 1831. Epitype (Al-Shehbaz in Cafferty & Jarvis Ind. loc.: “Habitat in Styria superiore in humidis 2002:531): Herb. Linn. No. 842.11 (LINN). alpium judenburgensium.” ≡ Cardamine halleri (L.) Prantl, Exkurs.Fl.Bay- Lectotype (designated here): “Arabis tenella” ern, ed. 2: 229–230, 1884. (s. a., Host,W–herb.Host). ≡ Cardaminopsis halleri (L.) Hayek, Fl. Steiermark 1: = Arabis halleri var. trachytica Fritsch, Verh. Zool.- 479–480, 1908–1911. Bot. Ges. Wien 44: 311, 1895. = Cardamine stolonifera Scop., Fl. Carniol., ed. 2: 229, Ind. loc.: “Bosnia orientalis. In pratis montis 1772. Kvarac prope Srebrenica, solo trachytico et cal- Ind. loc.: “ Habitat in Districtu Vipaccensi.” careo (?), 400–500 m s. m., Julio mense (Wettstein). Lectotype (designated here): Scop.,Fl.Car- Styria inferior. In silvis prope Cilli, solo cal- niol. ed. 2, tab. 39, ﬁg. 818, “Cardamine stolonife- careo(?), ca. 600 m s. m., 3.VIII. ﬂor. (Rechinger, ra”, 1772. R.).” 44 M. Kolník &K.Marhold

Type not known. spread on the stem, petiolate, blades of mid- = Arabis halleri subsp. transsilvanica Klášt., Preslia dle and lower leaves ovate to oblong or lance- 8: 29, 1929 non Schur 1866, nom. illeg. olate, coarsely toothed or rarely lobed, gradu- Ind. loc.: “Montes Svidovec: In Alneto viridis in ally reduced in size upwards the stem. Leaves Polonina Sterešora sub monte Bližnica, 1540 m s. with simple, forked and 3-forked trichomes, grad- m. sparse (H. m. 1595); in pratis ad cacumen mon- ually reduced in density upwards the stem, rarely tis Bližnica, 1820 m s. m. sparse (H. m. 1601, 1604); glabrous. The second lowermost stem leaf (0.7–)1– in saxosis ad cacumen montis Dragobrat, 1750 m 3.7(–5.5) × (0.3–) 0.4–1.6(–2.1) cm, the upper- s. m. sparse (H. m. 1998, 2005). – Montes Černá most one (0.4–)0.5–2.5(–3) × (0.1–)0.1–0.5(–0.7) Hora: in fastigio montis Pozyževská, 1650 m s. m. cm. Distance from the stem base to the longest (H. m. 2082) in valle alpino inter montes Gutin stem leaf (0.4–)0.7–11(–15.8) cm. Basal leaves ro- et Tomnatek, in pratis ad ripam rivuli 1800 m s. sulate, pinnatifid to lyrate-pinnatifid, maximum m. (H. m. 2100); in arenosis ad ripam rivi Breben- length of basal rosette leaves (1–)1.5–5.5(–7.5) jeskul sub monte Tomnatek, 1000 m s. m. sparse cm, length of their blades (0.5–)1.5–5.5(–7.5) cm, (H. m. 2113).” number of teeth on their blades (1–)2–6(–8). Lectotype (designated here): Svidovec, trav- Flowers (6–)8–33(–37), petals white, lilac, rarely natý svah od vrcholu Bližnice do kotle, 1800 m purplish, (4.1–)4.5–7.5(–8.1) × (1.9–)2.2–3.8(–4.1) (5.7.1926, Klášterský 1601,PR). mm. Sepals green to greenish purple, usually = Arabis halleri var. valida Zapal., Rozpr. Wydz. with few simple and forked trichomes, some- Mat.-Przyr. Akad. Umiejetn., DzialB,NaukiBiol. times densely hairy with small 3-forked trichomes, 52, Ser. B.: 46, 1912. rarely glabrous, wider sepals (1.5–)1.9–3.2(–3.5) × Ind. loc.: “Rospuntowa kolo Chrzanowa (Kru- (0.9–)1.1–1.9(–2.1) mm, narrower sepals (1.5–)1.7– pa).” 2.8(–2.9) × (0.6–)0.7–1.4(–1.5) mm. Siliquae lin- Holotype: Rospuntowa, Chrzanow, (24.7.1876, ear, replum constricted between seeds, length of Krupa 27.590, KRAM 114327). siliquae (1.2–)1.4–2.6(–3) cm, length of pedun- = Arabis halleri var. prelukiensis Zapal., Rozpr. cle (0.5–)0.6–1.2(–1.3) cm. Flowering and fruiting Wydz. Mat.-Przyr. Akad. Umiejetn., DzialB, May-August. Nauki Biol. 52, Ser. B.: 45, 1912. Chromosome number (see Table 1). 2n = Ind. loc.: “Mi¸edzi obu Czeremoszami blizko ich 2x =16. Zród˙ lowisk: Preluki na skalach wapiennych kolo Ecology. Mountain slopes, grasslands, meadows, 1550 m (Woloszczak).” forest edges, forests, mostly on shaded places. It is Lectotype (designated here): Preluk (1886, able to persist and compete in closed grass commu- Woloszczak 70.276, KRAM 114348). nities. It is indifferent to substrate, and occurs from = Arabis halleri β. pilifera Beck, Fl. Nieder-Osterreich¨ acid stones and bogs to limestone and dolomite. II/1: 458, 1892. It is found also on metal-contaminated soils. In Ind. loc.: “in Wiesen, an Waldrändern, etwas the Carpathians it is found from the colline to feuchten steinigen Stellen, vornehmlich auf Schie- alpine belt, in the altitudes from 200 to 1900 m ferder Voralpen, vom Wechsel und Gloggnitz, ent- a.s.l. lang der Steierischen Grenze zerstreut bis Göstling.” Distribution. In Belgium, France, Germany, Swi- Type not known. tzerland, Austria, Italy, Czech Republic, Poland, = Arabis halleri var. sudetica Dostál,Čas.Nár.Mus., Slovakia, Slovenia, Croatia, Bosnia and Herzegov- Odd. Přír. 60: 61, 1936. ina, Serbia, Ukraine, Romania, Moldova. In the Ind. loc.: “Montes Krkonoše.” Carpathian area it is common in the Eastern and Type not known. Southern Carpathians, in the Western Carpathi- – Arabis halleri subsp. eu-halleri Hayek, Repert. Spec. ans it occurs only in their northern and eastern Nov. Regni Veg. Bein. 33/1: 401, 1927, nom. inval. part. In the Polish Carpathians it occurs in the (Art. 24.3). Beskid S˛adecki Mts., Beskid Niski Mts., and in the – Arabis halleri subsp. eu-halleri András., Borbásia 2: Bieszczady Zachodnie Mts. In Slovakia it is found 21, 1940, nom. inval. (Art. 24.3). in the Slanské vrchy Mts., Nízke Beskydy Mts., Vihorlat Mts., Bukovské vrchy Mts. In Ukraine Arabidopsis halleri subsp. halleri it is documented from all phytogeographic districts of the Carpathians except from districts Description. Perennial herb, stoloniferous, (7–) of Vulkanični Karpaty and Zakarpats’ke pered- 10–53(–70) cm tall, with few branches in the up- hir”ja. From Romania we have seen specimens per part of stem. Stem not pruinous, densely from the districts of Maramure¸s, Suceava, Hune- hairy with simple, forked and 3-forked trichomes, doara, Bihor and Hargita, it can be, however, ex- rarely glabrous. Length of leafy part of stem (4–) pected also in some other districts of this coun- 6–41(–53) cm. Stem leaves (3–)5–20(–24), equally try. Distribution, chromosome numbers and nomenclature conspect of Arabidopsis halleri 45

Arabidopsis halleri subsp. tatrica (PAWL. ) Kol- hairy than mountain and alpine plants. The sec- ník, comb. nov. ond lowermost stem leaf (0.6–)1.1–4.2(–4.7) × (0.3–)0.4–1.5(–1.9) cm, the uppermost one (0.5–) ≡ Arabis halleri var. tatrica Pawl. Acta Soc. Bot. Polo- 0.7–3.1(–4.2) × (0.1–)0.2–0.8(–1) cm. Distance niae 7: 129–130, 1930–1931. from the stem base to the longest stem leaf (1.6–) Ind. loc.: “... w postaci zupelnie typowej na calej 2.6–17(–27) cm. Basal leaves rosulate, with weakly przestrzeni od Krzesanicy po Kop¸e Kondracka w developed lateral lobes to pinnatifid, maximum wysoko´sci od 1950 po 2128 m.” length of basal rosette leaves (1.1–)1.5–5.7(–6.5) Lectotype (designated here): Tatry Zach., cm, length of their blades (0.5–)0.6–3.0(–4.5) cm, Krzesanica 2128 v murawach (26. 7. 1928, Paw- number of teeth on their blades (0–)1–5(–7). Flow- lowski, KRAM 320705). ers (8–)11–35(–38), petals white, rarely lilac, (3.8–) = Arabis halleri var. carstica Dostál, Čas. Nár. Mus., 4.4–7.1(–7.5) × (1.8–)2.0–3.4(–3.6) mm. Sepals Odd. Přír. 60: 61, 1936. green to greenish purple, glabrous, rarely with few Ind. loc.: “ad declivia aprica graminosa in regione simple trichomes, small 3-forked trichomes never Slovenský Kras dicta et in convallibus rivi Hornád present, wider sepals (1.6–)1.9–3.2(–3.5) × (0.9–) inter oppida Obyšovce et Košice.” 1.1–2.0(–2.1) mm, narrower sepals (1.5–)1.7–2.9 Lectotype (designated here): Regio Slovenský (–3.3) × (0.7–)0.8–1.4(–1.6) mm. Siliquae linear, Kras: in nemore xerophytico supra opp. Turna, alt. replum constricted between seeds, length of sili- ca. 450 m s. m., solo calcar. (16. 5. 1933, Dostál, quae (0.8–)1.1–2.5(–3) cm, length of peduncle PR). (0.4–)0.5–0.9(–1.1) cm. Flowering and fruiting = Arabis halleri var. devestita Zapal., Rozpr. Wydz. May-August. Mat.-Przyr. Akad. Umiejetn., DzialB,NaukiBiol. Chromosome number (see Table 1). 2n = 52, Ser. B.: 45, 1912. 2x =16. Ind. loc.: “Na zachód od Babiej góry... wTat- Ecology. Mountain slopes, grasslands, meadows, rach... w krainie kosodrzewu Czarnej Hory.” forest edges, forests, mostly on shaded places. It is Lectotype (designated here): Gluchaczki na able to persist and compete in closed grass com- zachód od Babiej góry, (28.4.1906, Zapalowicz munities. It is indifferent to substrate, from acid 221.39, KRAM 114338). stones and bogs to limestone and dolomite. It is Nomenclature note: in the original circumscription found also on metal-contaminated soils. In the it included both A. halleri subsp. halleri and subsp. Carpathians it is found from colline to alpine belt, tatrica, while by the present typification it becomes in the altitudes from 400–2000 m a.s.l. synonym of subsp. tatrica only. Distribution. Endemic of the Western Carpathi- – Cardaminopsis halleri subsp. ovirensis auct. non. ans. Distribution center of A. halleri subsp. tatrica (Wulfen) Thell. 1919: Dostál et Červenka is located in Slovakia, where it occurs through- 1991. out the Western Carpathian mountains (it is very – Cardaminopsis halleri subsp. tatrica (Pawl.) Dos- common in the Slovenské rudohorie Mts., Murán- tál, Folia Mus. Rerum Nat. Bohemiae Occid., ska planina Mts., Slovenský raj Mts., Nízke Tatry Bot., 21: 6, 1984, nom. inval. (Art. 33.3, 33.6). Mts., Západné Tatry Mts., Vysoké Tatry Mts., – Cardaminopsis halleri subsp. tatrica Pawl. ex Dos- Belianske Tatry Mts., and occurs seldom e.g. in tál, Seznam Cév. Rostlin Kv. Československé, the Poľana Mts., Štiavnické vrchy Mts., Chočské 92, 1982; Dostál, Nová Květena ČSSR 1: 354, vrchy Mts., Pieniny Mts., Západné Beskydy Mts., 1989; Dostál et Červenka Veľký Kľúč Urč. Vyšš. Slovenský Kras Karst, and in the Spišské vrchy Rastl. 1: 335, 1991, nom. inval. (Art. 33.3). Mts.), it was found also in the eastern part of – Cardaminopsis halleri subsp. slovaca Měsíček in the Czech Republic (Moravsko-Slezské Beskydy Dostál, Nová Květena ČSSR 1: 354, 1989, nom. Mts.), in the southern part of Poland (the Beskid prov., pro syn. Zywiecki˙ Mts., Pieniny Mts., Kotlina Orawsko- Description. Perennial herb, stoloniferous, (9–) Nowotarska Basin, and in the Tatry Mts), and 12–52(–60) cm tall, usually not branched. Stem northern part of Hungary (Ágtelek). not pruinous, usually with simple trichomes in leafy part of stem and glabrous in inflorescence. Arabidopsis halleri subsp. dacica (Heuff.) Kol- Length of leafy part of stem (6–)7–38(–50) cm. ník, comb. nov. Stem leaves (3–)4–17(–23), equally spread on the stem, blades of middle and lower leaves ≡ Arabis ovirensis var. dacica Heuff. Verh. Zool.-Bot. ovate to oblong or lanceolate, rarely toothed, Ges. Wien 8: 52, 1858. longest leaf usually in the middle part of stem. Ind. loc.: [In Banatu Temesiensii]. Stem leaves usually only with few simple tri- Nomenclature note: No original material relevant chomes, the uppermost one usually glabrous. to this name was found in the herbarium BP, where Plants from populations in lower altitude are more specimens from the Heuffel’s herbarium are de- 46 M. Kolník &K.Marhold

posited (Somlyai, pers. comm.). Lectotype (designated here): In Alpibus Ovi- – Arabis ovirensis var. dacica Heuff.exGriseb.et rensibus supra Ebriach... (s. a., Wulfen W). Schenk in Arch. Naturgesch. 18: 307, 1852 (pro ≡ Cardaminopsis halleri subsp. ovirensis (Wulfen) syn.). Thell.inHegi Ill. Fl. Mitt.-Eur., 4/1: 424, 1919. – Arabis halleri subsp. ovirensis auct. non (Wulfen) ≡ Cardaminopsis ovirensis (Wulfen) Jáv. Magyar Klášt.: Klášt. Preslia 8: 30, 1929. Fl.: 436, 1924. Description. Perennial herb, stoloniferous, with ≡ Arabis halleri subsp. ovirensis (Wulfen) Klášt. plenty repent stolons, (4–)5–23(–26) cm tall, usu- Preslia 8: 30, 1929. ally not branched. Length of leafy part of stem – Cardaminopsis ovirensis (Wulfen) O. Schwarz (1.8–)2.8–14(–16.5) cm, stem not pruinous, usu- Mitt. Thüring. Bot. Ges. 1: 101, 1949 (isonym). ally with simple trichomes in leafy part of stem. The circumscription of this taxon and its delimi- Stem leaves (3–)4–10(–12), usually with few sim- tation in respect of the populations of A. halleri ple and forked trichomes, equally spread on the subsp. halleri in the Alps is not fully clarified and stem, longest leaf usually in the lower third to will require detailed morphometric study of the the middle part of stem, blades of lower stem Eastern Alpine populations. Material included in leaves orbicular to ovate entire or crenate. The sec- our study was insufficient to make final conclu- ond lowermost stem leaf (0.6–)0.7–2.5(–3) × (0.2–) sions. Nevertheless, our studies showed that ma- 0.3–0.9(–1.2) cm, the uppermost one (0.4–)0.4–1.9 terial from the type locality of A. halleri subsp. (–2.2) × (0.1–)0.2–0.4(–0.5) cm. Distance from the ovirensis in Austria is well different from the pop- base to the longest stem leaf (0.5–)0.9–6.5(–8.5) ulations occurring in the Eastern and Southern cm. Basal leaves rosulate, entire or with weakly de- Carpathians. veloped lateral lobes, their maximum length (0.7–) 0.9–3.5(–4) cm, length of their blades (0.3–)0.4– 1.7(–2) cm, number of teeth on their blades 0– Acknowledgements 4(–5). Flowers (7–)7–21(–27), petals lilac to pur- We would like to thank the curators of the herbaria BP, × ple, rarely white, (4.9–)5.3–7.6(–8.1) (2.5–)2.7– BRA, BUC, BUCA, CL, KO, KRA, KRAM, LW, LWS, 4.5(–4.7) mm. Sepals green to greenish purple, LWKS, PR, PRC, PRA, SAV, SLO, W, WU, ZV for allowing glabrous, rarely with few simple trichomes, small us to study herbarium specimens, L. SOMLYAI for informa- 3-forked trichomes always absent, wider sepals tion on herbarium material deposited in BP, J. KUČERA,A. (1.9–)2.0–3.2(–3.4) × (1.2–)1.3–2.5(–2.7) mm, nar- KAGALO,N.SYČAK,E.WALUSIAK,M.ONETTE,C.BIT¸A rower sepals (1.7–)1.8–2.9(–3.0) × (0.7–)0.8–1.4 and M. MATSCHINGER for their assistance in the field, and (–1.6) mm. Siliquae linear, replum constricted be- J. LIHOVÁ for useful comments. The grant agency VEGA, tween seeds. Flowering and fruiting May-August. Bratislava, Slovakia (grant. No. 3043) and the German Aca- n demic Exchange Service DAAD (project 323-PPP to K. M.), Chromosome number (see Table 1). 2 = are acknowledged for financial support. 2x =16. Ecology. Alpine slopes, grassy screes, alpine grasslands, stony soils, substrate on metamorphic and References acid bedrock never on alkaline limestone and AL-SHEHBAZ, I.A., O’KANE,S.L.&PRICE, R.A. 1999. Generic dolomite. Rare in montane belt, its distribution placement of species excluded from Arabidopsis (Brassi- center is from high montane to alpine belt, in the caceae). Novon 9: 296–307. Carpathians it is found from 1000 to 2400 m a.s.l. AL-SHEHBAZ,I.A.&O’KANE, S.L. 2002. Taxonomy and phy- Distribution. Eastern and Southern Carpathi- logeny of Arabidopsis (Brassicaceae), pp. 1–22. In: The Arabidopsis ans and the Balkan Peninsula. In the Carpathi- Book, American Society of Plant Biologists, http://dx.doi.org/10.1199/tab.0001. ans it occurs in high mountains in Romania, in BERT,V.,MACNAIR, M.R., DE LAGUERIE,P.,SAUMITOU- the Vl˘adeasa Mts., Rodna Mts., F˘ag˘ara¸sMts., LAPRADE,P.&PETIT D. 2000. Zinc tolerance and accu- T¸ arcului Mts., Retezat Mts., Bucegi Mts., and in mulation in metallicolous and nonmetallicolous populations Arabidopsis halleri 146: the Siriului Mts. The distribution of this taxon, of (Brassicaceae). New Phytol. 225– 233. based on the specimens cited here is apparently BERT,V.,BONNIN,I.,SAUMITOU-LAPRADE,P.,DE LAGUERIE, incomplete and should be subject of the further P. & PETIT, D. 2002. Do Arabidopsis halleri from nonmetalli- study. colous populations accumulate zinc and cadmium more effec- tively than those from metallicolous populations? New Phy- 155: Arabidopsis halleri ovirensis Wulfen) tol. 47–57. subsp. ( BERT,V.,MEERTS,P.,SAUMITOU-LAPRADE,P.,SALIS,P., O’Kane & Al-Shehbaz Novon 7: 325, 1997. GRUBER,W.&VERBRUGGEN, N. 2003. Genetic basis of Cd tolerance and hyperaccumulation in Arabidopsis halleri. ≡ Arabis ovirensis Wulfen in Jacq. Collectanea 1: Plant Soil 249: 9–18. CAFFERTY,S.&JARVIS, C., eds. 2002. Typification of Linnean 196, 1786. 51: ... plant names in Brassicaceae (Cruciferae). Taxon 529–537. Ind. loc.: “ cum Ovirenses supra ebriacum in ČOPYK, V.I., KOTOV,M.I.&PROTOPOPOVA, V.V. 1977. Viz- Valle Junonia percurrerem alpes”. načnik roslyn Ukrajinskich Karpat. Naukova dumka, Kyjiv. Distribution, chromosome numbers and nomenclature conspect of Arabidopsis halleri 47

FABISZEWSKI, J. 1986. Heavy metal tolerance of Cardaminopsis data. In: ANONYMOUS (ed.), XVII International botanical halleri (L.) HAYEK populations in the Polish Tatra Mts. Acta congress, Vienna, Austria, Europe, Abstracts, p. 399. Soc. Bot. Poloniae 55: 431–428. KONDRACKI, J., 1981. Geografia fizyczna Polski. Ed. 4. Państwo- FUTÁK, J. 1966. Fytogeografické členenie Slovenska, pp. 535–538. we wydawnictwo naukowe, Warszava. In: FUTÁK, J. (ed.), Flóra Slovenska 1, Veda, Bratislava. MARHOLD,K.,LIHOVÁ,J.,PERNÝ,M.,GRUPE,R.&NEUFFER, HOLMGREN, P.K., HOLMGREN,N.H.&BARNETT, L.C. 1990. B. 2002. Natural hybridization in Cardamine (Brassicaceae) Index Herbariorum. Part I: The Herbaria of the World. Ed. in the Pyrenees: evidence from morphological and molecular 8. Regnum Veg. 120: 1–693. data. Bot. J. Linn. Soc. 139: 275–294. JALAS,J.&SUOMINEN, J. 1994. Atlas florae europaeae 10 MĚSÍČEK, J. 2002. 21. Cardaminopsis (C.A. MEY)HAYEK., pp. (Sisymbrium to Aubrieta). The Committee for Mapping the 338–415. In: GOLIAŠOVÁ,K.&ŠÍPOŠOVÁ, H., (eds), Flóra Flora of Europe and Societas Biologica Fennica Vanamo, Slovenska V/4, Veda, Bratislava. Helsinky. MURÍN, A. 1960. Substitution of celophane for glass covers to JONES, B.M.G. & AKEROYD, J.R. 1993. 41. Cardaminopsis L., faciliate preparation of permanent squashes and smears. Stain pp. 351–352. In: TUTIN, T.G., BURGES, N.A., CHATER, A.O., Technol. 35: 351–353. EDMONDSON, J.R., HEYWOOD, V.H., MOORE, D.M., VALEN- O’KANE,S.L.&AL-SHEHBAZ, I. A. 1997. A synopsis of Ara- TINE D.H., WALTERS,S.M.&WEBB, D.A. (eds), Flora eu- bidopsis (Brassicaceae). Novon 7: 323–327. ropaea 1, Psilotaceae to Platanaceae, Ed. 2., Cambridge Uni- PAWLOWSKI , B. 1930. Zapiski florystyczne z Tatr III. Acta Soc versity Press, Cambridge. Bot. Poloniae 7: 127–137. KOCH,M.,AL-SHEHBAZ,I.A.&MUMMENHOFF, K. 2003. Molec- VAN ROSSUM, F.R., BONNIN,I.,FÉNART,S.,PAUWELS,M., ular systematics, evolution and population biology in the PETIT,D.&SAUMITOU-LAPRADE, P. 2004. Spatial genetic mustard family (Brassicaceae). Ann. Missouri Bot. Gard. 90: structure within a metallicolous population of Arabidopsis 151–171. halleri, a clonal, self-incompatible and heavy-metal-tolerant KOCH,M.,BISHOP,J.&MITCHELL-OLDS, T. 1999. Molecular species. Molec. Ecol. 13: 2959–2967. systematics and evolution of Arabidopsis and Arabis.Plant Biol. 1: 529–537. Received Sept. 25, 2005 KOLNÍK,M.&MARHOLD, K. 2005. Infraspecific differentiation Accepted Nov. 10, 2005 of Arabidopsis halleri based on AFLP and morphological

Appendix Ukraine. 1. Prykarpattja. Kaližskij rajon: Zawadka (s. a., REHMAN, LWS 41820, LW 041620). 2. Schidni Herbarium specimens studied Beskidy j Nyz‘ki Polonyny Mts. Perečinskij rajon:

Lanušove, beach wood (20.5.1991, KAGALO,SYČAK,LWKS

ÖÓÔ×× Ð Ð Ö Ð Ð Ö subsp. 03827); polonina Runa, on its top (Klášterský, PR 24527). Poland. 513. Zewn˛etrzne Karpaty Zachodnie Mts. Velikobereznianskij rajon: Mt. Stinská, 1000 m (8.1994, 513. 54. Beskid S˛adecki Mts., Dzwonkówka (2.6.1965, TASINKEVIČ, LWS N16960); Rosypanec, 1000 m (19.7.1995, Pelc, KRA 246418). 513.71. Beskid Niski Mts.,pasmo TASINKEVIČ, LWS 41880); Velké Berezné (19.5.1929, MA- Bukowicy, turning to Kamienia (21.5.1961, JASIEWICZ, ZEPA, PR 24528). Voloveckij rajon: Volovec, Mt. Pikuj (s. GRODZINSKA´ , KRAM 060695); Janów (14.5.1950, DO- a., SLOBODICA, LWKS 11260, 16791); Bereg, Kiszbiszta, BRZANSKA´ , KRAM 115460); Szczawnica, Nowy S˛acz, 1000 Mt. Pikuj (20.6.1909, THAISZ, PR; 21.7.1910, MARGIT- m (1897, WOLOSZCAK , KRAM 114815, 114839). 522. TAI, PR); Mt. Velký vrch (12.8.1919, DOMIN,PRC); Beskidy Wschodnie Mts. 522.12. Bieszczady Za- Mt. Vysoky Kamen, beach wood (6.8.1992, KAGALO, chodnie Mts., Jureczkowa, in forest (13.5.1971, PIEKO˛ ´S, SYČAK, LWKS 04630); Mt. Stih (20.6.1949, s. coll., LWS TUMIDAJOWICZ, KRAM 206551); Mt. Szeroki Wierch, 41847, 41846); Volovec (26.4.1948, MALINSKY, LWS 41844; 970 m (21.5.1986, PAWLUS,JASIEWICZ, KRAM 443238, 22.6.1949, MELINČUK, LWS 41849); Boržavska polonina 443239); Liszna, 625 m (22.5.1986, PAWLUS,JASIEWICZ, Mts., Mt. Plai, 1334 m (12.5.1947, JUSKIN, LW 041633, KRAM 443250); Mt. Polonina Weltlińska (5.1973, Pelc, 041634); Sinevirske ozero lake (17.8.1947, JUSKIN,LW KRA 246413); Mt. Polonina Caryńska, 1110 m (20.8.1956, 041630); Beskid stancia (31.5.1947, JUSKIN, LW 041631). JASIEWICZ, KRA 004746; 10.7.1982, KUC, KRAM 000924); Svaľavskij rajon: Svaľava, beach wood (3.5.1949, s. coll., Wolosate, 740 m (20.5.1960, JASIEWICZ, KRA 004745). LWS 41848); beside Bystrá stream (4.5.1947, JUSKIN,LW Slovakia. 19. Slanské vrchy Mts. Körök hegy, Slanec 041632); Polonina Boržava Mts., Mt. Velký vrch, 1500 (5.1938, KLÁŠTERSKÝ et DEYL,PR).20. Vihorlat Mts. m (28.7.1937, KLÁŠTERSKÝ, PR 24512). 3. Gorgany Bystrý, Mt. Pliška (28.7.1937, JIRÁSEK, PRC); Stakčín, Mts. Mižhirskij rajon: Koločava, beach wood (7.6.1994 Kováčikovo, 280 m (29.6.1957, SOJÁK, PR 561323–561326). KAGALO, LWKS 05731); Sinevir, Mt. Polonina Krásna, 30c. Nízke Beskydy Mts. Humenné, Olšinkov, 650 m 1500 m (7.1932, LÁSKA, PR); Koločava, Mt. Negrovec, (14.7.1974, CHRTEK,KŘÍSA,PRC).31. Bukovské vrchy 1600 m (30.6.1932, LÁSKA, PR), 1700 m (13.8.1936, Mts. Beside a road between the villages of Zboj and Uličské DEYL, PR), 1650 m (13.8.1936, DEYL,LÁSKA,PR), Krivé, 321 m (16.5.2004, KOLNÍK,SAV);Ruské,Mt.Kuz- 1400 m (28.6.1932, LÁSKA, PR). Tiačivskij rajon: Mt. mov Bezkyd, 1050 m (17.7.1973, MÁJOVSKÝ, SLO); Runina, Sverdel (8.5.1954, VAJNAH, LWS 41829); Mt. Polonina Mt. Riaba skala, 1193 m (18.6.1969, s. coll., BRA; 4.1957, Krasna (22.5.1955, VAJNAH, LWS 41834); Velka Uhoľka, MÁJOVSKÝ, SLO); Runina, saddle below Mt. Ďurkovec, 1000 m (8.8.1976, TASENKIEVIČ, LWS N16597), 750 m 1000 m (1.8.1957, SOJÁK, PR 651310); Ulič, pasture NE of (12.9.1971, SMOŇKO, LWS N16596), 800 m (22.8.1980, the village, 440 m (5.5.1970, GALLO, BRA); Nová Sedlica, SMOŇKO, LWS N16961). Rožniatinsikij rajon: Gorgany Mt. Príkry, 800–850 m (17.8.1957, HEJNÝ,PR);Mt.Czolo, Mts., Doza, 480 m (19.5.1938, ÚRODON´,KRAM);Ne- 1000 m (3.8.1957, SOJÁK, PR 561308, 561309); Mt. Stinská prova, 1430 m (11.8.1934, ÚRODON´, KRA 004745); Pere- (18.8.1988, MARHOLD,SAV). hinska (22.5.1935, FOTINUK, LWS 41819). Nadvirnianskij 48 M. Kolník &K.Marhold rajon: Ivanofrankivsk (22.5.1986, KAGALO, LWKS 06308). Zachodnie 514.12. Pieniny Mts., Hombark, 800 m Bohorodčanskij rajon: Maňava (18.6.1965, BERKO,LWS (1.6.1961, GRODZINSKA´ , KRAM 221374); Pieniński Pas 41810 – 41812). 4. Svydovec Mts. Rachivskij rajon: Skalkowy klippen belt (5.5.1968, GRODZINSKA´ ,KRAM Rachiv, allong the river Tisa, 440 m (29.4.1937, PUL- 221373). 514.11. Kotlina Orawsko-Nowotarska Basin. CHART, PRC); Svydovec, Mt. Bliznica, 1800 m (5.7.1926, Nowy Targ, Lasek, valley of stream Syraczki, 620 m KLÁŠTERSKÝ, PR), in glacial valley (13.7.1932, MAR- (6.5.1966, GRUZIKOWA, KRAM 069099); Podwilk, 710 m TINEC, PR 24503, 24504), on top, 1860 m (5.7.1926, (19.6.1962, GRUZIKOWA, KRAM 069075, 069076, 069098). KLÁŠTERSKÝ, PR, PRC), 1200 m (31.5.1948, MALINSKY, Tatry Mts., Tatry (s. a., leg. Rehman, LWS 41852–41854); LWS 41845; 15.6.1954, MALINSKY, LWS 41806; 5.6.1983, 514.52. Tatry Zachodnie Mts., Przel¸ecz Liliowe saddle, MALINSKY, LWS 41696), 1500 m (13.7.1932, ŽOFÁK, 1950 m (16.6.1929, PAWLOWSKI , KRAM 320709); Dolina PRC); Mt. Todjaska, 1700–1750 m (19.7.1999, MRÁZ,JU- Chocholowska valley, Polana Huciska glade, 1020–1050 RKOVIČOVÁ); Mt. Polonina Sterešora, 1520 m (14.7.1926, m (5.5.1948, PAWLOWSKI , KRAM 320694, 036354; KRA KLÁŠTERSKÝ, PR); Mt. Dragobrat, rock on top, 1750 m 012863); MalaL¸ aka glade, 1170 m (15.5.1946, PAWLOWSKI , (29.6.1929, KLÁŠTERSKÝ, PR; 14.7.1972, LAZEVNA,LWS KRAM 320695); Polana Bialy Potok glade, 940 m (5.5.1948, 41677, 41803); Rachov, Mt. Dumen, 1100 m (14.5.1937, PAWLOWSKI , KRAM 320691); Mt. Krzesanica, 2128 m PULCHART, PR); Mt. Polonina Šumnevska (7.5.1954, MA- (16.7.1928, PAWLOWSKI , KRAM 320705), 2128 m (1.7.1932, LINSKY, LWS 41695). 5. Čornohora Mts. Rachivskij ra- PAWLOWSKI , KRAM 320706); Mt. Kopa Kondracka, 1950 jon: Čorna hora Mts. (6.1954, ROZENKO, LW 041593), m (25.7.1928, PAWLOWSKI , KRAM 320708); Hala Olczyska 1000 m (15.6.1959, ŽDAROVA, LW 041594); Mt. Hov- glade, 1100 m (3.5.1948, PAWLOWSKI , KRAM 320693, erla, Prut (20.7.1995, KAGALO,SYČAK, LWKS 07333), 036898), (KRA 012864); Hala Kondratowa glade (27.7.1955, Mt. Šešul (16.5.1957, VAJNAH, LWS 41830); Mt. Stoly TACIK, KRAM 389334). 514. 53. Tatry Wschodnie (13.6.1955, VAJNAH LWS 41827, 41828); on sand banks Mts., under glacial lake Morskie Oko, 1400 m (22.8.1926, of stream Brebenjeskul, 1000 m (6.7.1929, KLÁŠTER- PAWLOWSKI , KRAM 320711); Zakopane, on meadow, SKÝ, PR, PRC); Mt. Tomnatek, 1900 m (7.1938, DEYL, 860 m (28.6.1968, PIE¸KO´S, KRAM 89249), (10.5.1911, PR); Mt. Požiževska, 1650 m (3.7.1929, KLÁŠTERSKÝ, WISTLEINOWA,STECKI, LW 041628, 041622); Ku´znice PR; 11.6.1928, TULIGLOVIČ, LWS 41826, 41838, 41842; (29.5.1955, URBANSKA´ , KRA 012874, 012875; s. a., leg. 13.6.1962, ŠEVČUK, LWS 41697; 22.6.1960, KOLIŠČUK, Miczyński, KRAM 005500), Koziniec (29.5.1921, s. coll., LWS 41686; 28.6.1978, TASINKEVIČ, LWS N6693), 1830 m KRAM 005499); Mt. Ciemniak (9.7.1961, s. coll., KRA (3.7.1978, TASINKEVIČ, LWS N6694; 8.7.1921, TULIGLOVIČ, 72949); Dolina Spadowiec valley, 975 m (1.6.1961, RA- LWS 41807); Cibulnik (21.6.1973, LAZEVNA, LWS 41813– JCHEL, KRA 112565). 41817); Kvasi, Mt. Menčil, 1300 m (3.6.1956, s. coll., LWS Slovakia. 3. Slovenský kras Karst. Turňa (1.6.1941, 41685; 11.6.1955, ŠIMOVA, LW 046426, 046427; 1.7.1985, ZSÁK, BRA; 13.7.1941, BRECHER, BRA; 22.6.1941, ZSÁK, VOLHIN,KARDAŠ,HINDA, LW 041597); Kvasi, Mt. Šešul BRA); 450 m (16.5.1933, DOSTÁL, PR 24511, PRC); Turni- (1.7.1985, VOLHIN, Kardaš, Hinda, LW 041596; 23.6.1984, anska plošina plateau (17.5.1958, KLÁŠTERSKÝ,MĚSÍČEK, ZAGUĽSKI, LW 041598). Nadvirnianskij rajon: Vorochta ŽERTOVÁ, PR 584514); Zádiel (25.5.1967, MIKOLÁŠ,KO (4.6.1986, DANILIK, LW 041589); Mt. Turkul (28.7.1909, 12261); Zádielska dolina valley (23.5.1958, SOJÁK,PR s. coll., LWS 41808); Ľuča saddle, beach wood (5.1933, 288661; 28.5.1964, VOJTUŇ, KO 2192); Zádiel, Mt. Havranie PRIHRODSKI, LWS 41805). 6. Čyvčyno-Grynjavs‘ki skaly, 800 m (14.5.1923, DOSTÁL, PRC); Lúka nad Borkou hory Mts. Vrchovinski rajon: Mt. Čivčin (13.6.1958, VA- (25.5.1967, VOJTUŇ, KO 2447); Pramenište (14.9.1970, VO- JNAH, LWS 41831); vicinity of chalet Diana (1.6.1930, JTUŇ, KO 1259). 14d. Poľana Mts. Mt. Poľana, Sihla, LÁSKA,PRC).7. Marmaros‘ki Aľpy Mts. Rachivskij ra- meadows, 800 m (15.6.1981, ZAHRADNÍKOVÁ,SAV),on jon: Barlebaška, Mt. Pop Ivan, 1600 m (7.1938, DEYL PR), meadows, 700 m (13.5.1981, JASIČOVÁ, SAV), on meadow 1560–2026 m (7.1928, DEYL,PR);Marmaroš,Trebušany, near the village, 700 m (13.5.1981, JASIČOVÁ et ZAHRAD- Gutin Mts., saddle between Mt. Pop Ivan and Mt. Pietros NÍKOVÁ, SAV); Mt. Predná Poľana, 900 m (20.5.1985, (8.1933, DEYL, PR 24526). ZAHRADNÍKOVÁ,SAV).14e. Štiavnické vrchy Mts. Romania. Maramure¸s. Baia Mare (19.5.1951, 26.9.1954, Hronská Breznica (5.1920, TRAPL,PRC).15. Sloven- s. coll., BUC 344195–344207, 272990, 272991). Suceava. ské Rudohorie Mts. Čuntava-Masná, on pasture, 1100 paraul Boulu stream, 560 m (30.4.1986, LUCESCU,BUC m (27.5.1959, MANICA, ZV 7703–7705); Hnilec, Babiná, 273492); Bukovina, Dadula, valley of river Kârlibaba 700 m (9.6.1977, HAJDÚK, BRA); Slovinky W, Čierny Bo- (6.1911, J.B.B., KRAM 483479, LWS 41839, 41840, LW cian chalet, beside stream (30.5.1986, MÁJOVSKÝ,SLO); 041619); Obcina Mestec˘ani¸s Mts., Moldova Sulita, Lucina- Mníšany NE, Mt. Skalka, 730 m (10.5.1971, MÁJOVSKÝ, G¯aina, 1170 m (18.6.1997, NEGREAN, BUC 153968). Hune- SLO); Zdychava NE, Zdychavské lesy, 700 m (8.5.1971, doara. Valea Boholt, 10.7.1961 (s. coll., BUC 263829– MÁJOVSKÝ, SLO); Vlachovo, Mt. Zadný Ďuránov (3.1991, 263831); Petro¸sani, Mt. Piule, 200 m (25.7.1956, CSÜRÖS, HAJDÚK, BRA); Hnilecké hory Mts., Galmus, Mt. Slovinská CL). Bihor. Vadu Crisului, 270 m (11.5.1962, NEACS¸U, skala, 900 m (18.5.1958, MARSA, BRA), 1000 m, (14.5.1961, BUC 260901); Cri¸sana, Stâna de Vale, 1100 m (1.5.1963, MARSA, BRA); Revúca, Mt. Kohút (24.7.1898, s. coll., PR BORZA, BUC 101565); Szatmár, Nagybánya, 220–230 m 24518). 16. Muránska planina Mts. Tisovec, (30.5.1927, (18.7.1913, JÁVORKA,BRA).Hargita. Hargita Mts., Mt. VRANÝ, PRC); Veľká lúka meadow, 900 m (26.6.1948, Hargita, 1700 m (11.6.1983, GROZA, LI 151392). HENDRYCH, PR), 960 m (s. a., Dvořák, BRA); Zámok

huntig chalet, 850–880 m (15.5.1949, HENDRYCH,PR);

ÖÓÔ×× Ð Ð Ö ØØÖ OJÁK subsp. Mt. Veľká Stožka, 1100–1250 m (21.5.1958, S ,PR Poland. 513. Zewn˛etrzne Karpaty Zachodnie 513.51. 230798), (8.7.1970, VOJTUŇ, KO 1260); Stožka – Hrdzavá Beskid Zywiecki.,˙ Mt. Babia Góra, Zawoja (1.6.1970, valley, (17.6.1970, FUTÁK,ZAHRADNÍKOVÁ, SAV); Studňa, OCHYRA, KRAM 227398). 514. Centralne Karpaty 1100–1250 m (21.5.1958, SOJÁK, PR 230806; 11.6.1970, Distribution, chromosome numbers and nomenclature conspect of Arabidopsis halleri 49

FUTÁK,ZAHRADNÍKOVÁ, SAV), 1200 m (10.7.2001, VALA- chalet, 1680 m (4.8.1930, HROBAŘ, PRC); Králička, 1650– CHOVIČ, SAV); Martinová dolina (14.5.1970, FUTÁK,SAV); 1700 m (3.7.1931, SILLINGER, PRC); valley of Bystrá river Muráň castle ruin, 800 m (18.7.1978, VAŠÁK,BRA);Mt. (5.1931, TRAPL, PRC); Liptovská Porúbka, railway em- Muráň, 1400 m (6.1928, s. coll., SLO); Muránska Huta, bankment, 660 m (9.5.1973, HORNÍČKOVÁ,BRA);Čierny spruce forest W of the village, 800 m (30.6.1961, SCHID- Váh (2.5.1963, GRODKOVSZKY, BRA; 27.5.1965, ČER- LAY, SAV; 24.5.1958, SOJÁK, PR 288662). 17. Slovenský VENKA, SLO), 800 m (13.5.1973, VARTÍKOVÁ,SLO),Nižný Raj Mts. Glac, 900 m (13.6.1930, SUZA,SOCIL,PR),900 Chlmenec, 750 m (12.5.1973, VARTÍKOVÁ, SLO); valley of m (13.6.1930, SUZA,SOCIL, SLO), 900 m (13.6.1936, SUZA river Čierny Váh, Záturňa, 1100 m (5.6.1974, VARTÍKOVÁ, et SOCH, ZV 4897; 14.7.1944, NÁBĚLEK, SAV; 13.7.1944, SLO); valley of river Čierny Váh, Svarín, 700 m (19.5.1972, FUTÁK, SLO); Kláštorisko (14.7.1944, FUTÁK,SLO);val- VARTÍKOVÁ, SLO); Mt. Muránik not far of the village ley of Biela voda stream (17.4.1944, FUTÁK,SLO);Vernár Čierny Váh, 950 m (9.7.1974, VARTÍKOVÁ, SLO), 800 m (24.5.1970, VOJTUŇ, KO 1262, 1263); Malý Sokol gorge (6.6.1974, VARTÍKOVÁ, SLO); Mt. Kolibisko, not far of the (14.8.1958, SOJÁK, PR); Stratená, meadow W of the village, village Čierny Váh, 1000 m (5.6.1974, VARTÍKOVÁ,SLO); 800 m (18.5.1963, FUTÁK,SAV);betweenMt.Malčekov Čierny Váh, path from the Torysa stream valley to Mt. vrch and Mt. Havrania skala, 1120 m (14.8.1958, SO- Holica, 1400 m (27.7.1972, VARTÍKOVÁ,SLO);Malužiná, JÁK, PR); Píla, 800 m (27.7.1947, KLÁŠTERSKÝ,PR); Mt. Skribňovo, 1100 m (5.6.1974, VARTÍKOVÁ,SLO);Šver- Dedinky, meadows beside a railway (23.5.1973, BERNÁ- movo, wet meadows, 700 m (13.6.1969, DVOŘÁK,BRA); TOVÁ, BRA); meadows uppon the Hnilec river, 950 m Švermovo [Telgárt], valley of river Hron, NE (5.7.1984, VOJ- (1.6.1978, DVOŘÁK, BRA); Mt. Gačovská skala, 1100 m TUŇ, KO); Telgárt, Mt. Skalka (6.1932, HAJNÝ,PRC).23a. (6.7.1947, KLÁŠTERSKÝ, PR); Vernár, meadows on the hill Západné Tatry Mts. Zuberec, Mt. Osobitá (30.6.1949, Javorina, 1000 m (30. 5.1966, MICHALKO, SAV); Novoveská FUTÁK, SLO), S slope, 1500–1600 m (7.8.1959, SOJÁK, Huta, conifer wood beside the Dubnica stream, 600 m PR), along a stream, 900 m (4.7.1929, DOSTÁL,PRC), (11.5.1958, s. coll., BRA). 18. Stredné Pohornádie Mts. Teplý žľab, 1300 m (15.7.1965, FUTÁK, SAV), 400 m Malá Lodina, Brezovice, Mt. Čierna hora (7.7.1937, MAR- (6.1967, DOSTÁL, PR 361175); Zuberec, Mt. Slatín, 1975 GITTAI, PRC); Veľká Lodina (1.5.1998, MIKOLÁŠ,KO m (9.8.1959, SOJÁK, PR 565394); Oravice, beside hunting 27554); Košice, Bankov (1.6.1971, VOJTUŇ, KO 1261); Ja- chalet (25.5.1972, SCHWARZOVÁ, SLO); Juráňova dolina hodná, 600 m (14.5.1931, s. coll., PRC). 21b. Malá Fa- valley, 890 m (14.7.1965, FUTÁK, SAV); Zverovka, mead- tra Mts. Mt. Chleb (5.6.1968, FUTÁK,ZAHRADNÍKOVÁ, ows (14.6.1973, BERNÁTOVÁ, BRA); Roháčska dolina val- BRA). 21c. Veľká Fatra Mts. Gaderská dolina valley, ley, 1100 m (8.1926, DOSTÁL, PRC); Látaná dolina val- 800 m (1978, HAJDÚK, BRA); Blatnica, Blatnická dolina ley, 1100 m (8.7.1930, DOSTÁL, PRC); Mt. Babky, 1200 valley (28.6.1978, HAJDÚK,BRA).21d. Chočské vrchy m (31.7.1919, DOSTÁL, PRC); Huty, Mt. Biela skala, 1400 Mts. On conﬂuence of rivers Orava and Váh, 410 m m (31.5.1952, DOSTÁL, PRC); Mt. Sivý vrch, 1200 m (11.6.2003, Dítě, herb. Dítě); Mt. Prosečné uppon Svorad (8.1926, DOSTÁL, PRC), (2.6.1979, HUBOVÁ,ZAHRAD- plateau (15.6.2002, Dítě, herb. Dítě); Oppid. Witow, na NÍKOVÁ, SAV); Jalovecká dolina valley, beside a stream, hutách (8.5.1894, Truchlý, BRA). 22. Nízke Tatry Mts. 912 m (26.6.1998, VALACHOVIČ,BRA).23b. Vysoké Vyšná Boca (22.5.1985, ZAHRADNÍKOVÁ,SAV),slopes Tatry Mts., (1892, Trusch, LW 041580; s. a., Heindrich, above the wilage (22.5.1985, ZAHRADNÍKOVÁ,SAV);Nižná LW 041615, 041616; 5.1922, GRESCHIK,SLO);Tomanova Boca (23.4.1964, VOJTUŇ, KO 2448–2450); Tále, meadow dolina valley, 1500 m (14.7.1951, ŠOREK, PR 264967); beside the hotel Partizán (15.5.1985, DEÁKOVÁ, BBZ); Čer- Mt. Stôly (24.7.1984, MUCINA, SAV); Podspády, along tovica, W slope above a saddle, 1400 m (5.6.1953, BENČAŤ, Javorinka stream (23.7.1925, KLÁŠTERSKÝ,PR);Štrb- MAGIC,MANICA, ZV 19037), 1230 m (27.5.1964, MANICA, ské Pleso, 1400 m (11.6.1976, CHRTEK,Deylová,PR); ZV 1378), saddle, 1240 m (30.4.1961, MANICA, ZV 4893); Bielovodská dolina valley (10.7.1925, KLÁŠTERSKÝ,PR Kráľova hoľa, in direction towards Liptovská Teplička vil- 24514); below glacial lake České pleso (8.7.1925, KLÁŠTER- lage, 1210 m (7.6.1988, MARHOLD, SAV); Veľký Brunov, SKÝ, PRC); Podtisovky (20.6.1925, KRAJINA,PRC);Ze- hunting chalet, 1050 m (5.6.1974, VARTÍKOVÁ,SLO); lené pleso glacial lake, 1577 m (11.7.1925, KLÁŠTER- Mt. Červená skala (21.5.1985, ZAHRADNÍKOVÁ,SAV);Po- SKÝ, PR); above the Zelené pleso glacial lake, 2150 m horelá, on border of a road (21.5.1985, ZAHRADNÍKOVÁ, (15.7.1925, KRAJINA, PRC); dolina Bielej vody Kežmarskej SAV); Machnatá (8.7.1970, FUTÁK,ZAHRADNÍKOVÁ,SAV); valley, 1200 m (3.6.1969, CHRTEK,KŘÍSA,PRC).23c. Studňa (8.7.1970, FUTÁK,ZAHRADNÍKOVÁ,SAV);Va- Belianske Tatry Mts. Javorina (5.6.1972, HODOVAL, jskovská dolina valley (19.5.1986, ZAHRADNÍKOVÁ,SAV); BRA); on wood between villages Javorina and Podspády Brezno, Bystrá dolina valley, beside the stream, 600 m (22.6.1925, KRAJINA, PRC); Ždiar, Mt. Havran, 1800– (30.4.1963, GREŠTIAK, ZV 9786); Svidovská dolina val- 2050 m (19.8.1967, VAŠÁK, PR; 20.7.1929, DOMIN,PRC); ley, 900 m (7.1983, HORNÍČKOVÁ, BRA); Vajskovská Mt. Ždiarska Vidla, 1600 m (13.8.1954, KLÁŠTERSKÝ, dolina, beside a hunting chalet (20.5.1986, MARHOLD, PR), N slope (30.7.1998, MIKOLÁŠ, KO 27752; 20.7.1951, ZAHRADNÍKOVÁ, SAV); Mt. Ďumbier (5.8.1919, DOMIN, KNEBLOVÁ, PR 252843, 252844; 18.7.1925, SILLINGER,PR PRC), (14.7.1896, KUPČOK, PR 24517), 2000 m (7.1935, 24515; 18.7.1925, SILLINGER, PRC); between Mt. Široké NÁBĚLEK, SAV, BRA; 13.6.1947, FUTÁK, SLO), Príslop pole and Mt. Ždiarska Vidla, 2000 m (22.6.1969, HUBOVÁ, (2.7.1952, MÁJOVSKÝ,SLO);betweenMt.Chopokand SAV); Tristárska dolina valley, 1550 m (25.7.1928, KRA- Mt. Ďumbier (10.8.1959, KÁŠTERSKÝ,PR);Mt.Krakova JINA,PRC);valleybetweenMt.HavranandMt.Nový hoľa, 1650 m (20.6.1967, JASIČOVÁ, Zahradníková, SAV; (11.9.1919, DOSTÁL, PRC); under the saddle Ždiarske 22.7.1928, CEJP, PRC); Mt. Krúpova hoľa (10.8.1956, sedlo (12.6.1971, HODOVAL, BRA); Ždiar, on meadows SOJÁK, PR 563564), 1927 m (18.8.1958, SOJÁK,PR); (20.5.1969, HAJDÚK, BRA); Mt. Bujačí (7.1937, ŠMARDA, Demänovská dolina valley, Repisko, Mt. Siná, 1500 m PR); Mt. Nový (13.8.1954, KLÁŠTERSKÝ, PR; 12.8.1959, (18.4.1958, KLÁŠTERSKÝ,PR);besideŠtefánikovachata SOJÁK, PR 565054); Mt. Jatky, N slope, 1900 m (22.7.1925, 50 M. Kolník &K.Marhold

KLÁŠTERSKÝ, PR); Mt. Hlúpy vrch, 1900 m (7.1938, DEYL, Bihor. Vl˘adeasa Mts., Cri¸sana, Stâna de Valle valley, PR); Mt. Belianska kopa, 1700 m (8.6.1970, DOSTÁL,PR 1100m (1.5.1936, BORZA, BUC 101565); Cri¸ana, Budureasa, 368651); Mt. Kopa, 1700 m (19.6.1924, KRAJINA,PRC), 1000 m (9.5.1990, GROZA, BUCA 160449-160452); Cluj, 1740 m (25.6.1915, TIMKO,KÜMERLE,PRC);Mt.Komíny, Vl˘adeasa Mts., 1300–1460 m (14.5.1950, POP, BUC 122346, 1620 m (19.7.1939, PAWLOWSKI , KRAM 320696); Červená 120354, KRAM, 962708). Cara¸s. T¸ arcului Mts., Mt. T¸atcu, Hlina (8.1926, KRAJINA, PRC); Dolina Siedmich prameňov supra Groapa Bistrit¸ii, 2000–2200 m (25.7.1936, NYÁRÁDY, valley, Jelenia lúka (15.5.1955, HADAČ, PR); Javorinka, 950 CL); Krassó-Szörény, Mt. Szarkó (26.6.1905, LENGYEL,BP m (23.7.1925, DOMIN,KRAJINA, PRC); Zadné Meďodoly 353443). Hunedoara. Retezat Mts., Mt. Gurazlata, beside valley, beside a path, 1500 m (22.6.1994, UHLÍŘOVÁ, Zenoga lake, ca 2000 m (29.8.1938, KÁRPÁTI,BP);Valea VOZÁROVÁ, BRA); Mt. Skalné Vráta, 1600 m (27.6.1969, Arm˘asarului, 1950 m (8.6.1996, NEGREAN, BUC 153431); HUBOVÁ,SAV).24. Pieniny Mts. (22.7.1926, s. coll., LW Mt. Bucura, above the lake, 2200 m (3.7.1925, NYÁRÁDY, 041606); Červený Kláštor, Mt. Kláštorná hora, SE slopes, CL); Mt. Papu¸sa, 1800–2100 m (29.7.1929, NYÁRÁDY,CL); 450 m (12.9.1988, MANICA, ZV 19291). 26a. Liptovská Custura (9.8.1937, SULMA, KRAM 271250); Custura, 1800 kotlina Basin. Liptovský Hrádok, on alluvium of river m (12.7.1988, GROZA, BUC 163352). Oltenia. Vˆılcea, Belá, 640 m (30.6.1976, HORNÍČKOVÁ, BRA); Pribilina, F˘ag˘ara¸sMts.,Mt.Zânoaga, 1600 m (25.5.1969, POPESCU, Rakytiny, meadow, 750–800 m (20.4.1972, HORNÍČKOVÁ, BUC 126709, PRC, KRAM 095691). Sibiu. F˘ag˘ara¸sMts., BRA); Lány, meadows, 800 m, (5.1983, HORNÍČKOVÁ, Mt. Piatra Rosie, 2400 m (28.7.1959, BORHIDI,BP); BRA); Potôčky, 660 m (1982, HORNÍČKOVÁ,BRA).26b. Bâlea glacial lake (5.1952, ALEXANDRU, BUC 151097, Spišské kotliny Basins. Meadows beside the road to the 151098), 1950–2000 m (22.7.1914, TUZSON, 208370 BP), Lendak village, 760 m (18.5.1963, FUTÁK,SAV).27b. Ja- 2100 m (2520–21.7.1920, ZSÁK, BP 73058); Árpas, 2100 vorníky Mts. Kysucká vrchovina Highland, Biely Kríž – m (24.7.1914, TUZSON, BP); Bulea Tal valley, 2150–2250 Jančíkovci, 780 m (5.6.1999, VALACHOVIČ,SAV).28. Zá- m (14.8.1916, s. coll., W 2512), 1820–1900 m (22.7.1914, padné Beskydy Mts. Oravské Beskydy Mts., Oravská TUZSON, 208367, 28368 BP). Arge¸s. F˘ag˘ara¸s Mts., Saua Polhora, beside the stream Polhoranka, 770 m (20.6.1976, Caprei glacial lake (11.7.1967, s. coll., BUC 108078); “Valea MIGRA, SLO); Vydrovka, 745 m (20.6.1976, MIGRA,SLO); Domni”, ca 1900–2000 m (23.7.1914, TUZSON, 208373 BP), Slaná voda, 780 m (18.5.1976, MIGRA, SLO); beside hunt- ca 2000 m (23.7.1914, TUZSON, 208371 BP). Prahova. ing chalet Roveň, 885 m (8.6.1976, leg. MIGRA, SLO); Bor- Sinaia, Bucegi Mts., Colt¸i (19.6.1929, GRINTESCU,BUC sučia dolina valley, beside hunting chalet Borsučie, 786 m 44697); Valea Chalae¸sti (30.5.1967, BUC 108112; s. a., (14.11.1976, MIGRA, SLO); alluvium of Vonžovec stream, DALECIU, LI); Bucegi Mts. (s. a., Prodan, BUC 16024; beside hunting chalet Vonžovec, 751 m (19.6.1976, MIGRA, 14.6.1890, LEONHARD, PRC; 27.7.1902, DEGEN, 73095 BP); SLO); Liesek, Pohanka (24.4.1894, Truchlý, BRA); Oravské Valea Serbului, Mt. Omul, 2100–2110 m (9.7.1931, DOMIN, Veselé, Mt. Talagova Grapa, 802 m (18.5.1976, MIGRA, KRAJINA,PRC).Buz˘au. Siriului Mts., Cura muntelui SLO); Tvrdošín, zákruta, (13.5.1894, Truchlý, BRA); Si- (1.6.1957, DILMER, LI 149612).

hoť (20.4.1894, Truchlý, BRA). 29. Spišské vrchy Mts.

ÖÓÔ×× Ð Ð Ö ÓÚÖ Ò×× Spišská Magura Mts., Osturňa, on a crest veldt, 1100 m subsp. (25.4.1959, HAJDÚK, BRA); Ružbachy, beside the Lazňanka Austria. Fl. Exs. Austro-Hung., no. 601, loc. classicus stream, 730 m (18.5.1971, GALLO, SAV); Mt. Magurka, 950 (JABORNEGG, PRC, WU); Carinthia, valle Junonia, Mt. m (9.6.1969, CHRTEK,Křísa,PRC);LevočskévrchyMts., Obir, 1400–2150 m (JABORNEGG,BRA);Carinthia,Mt. Mt. Ihla (11.6.1994, TOPERCER,KOCHJAROVÁ, BBZ). Obir (6.1864, JABORNEGG, PRC); Carinthia, Raibl, on 30b. Čergov Mts. Čergovský Minčol, SSE slope, 1050 m mountain meadows, 950 m (6.1885, JABORNEGG,PRC); (23.5.1996, KLIMENT,SAV). Carinthia, Raibl (21.5.1904, BECK,PRC);Carinthia,Mt.

Obir, 2000 m (11.7.1880, PREISSMANN,PRC).

ÖÓÔ×× Ð Ð Ö subsp. Romania. Maramure¸s. Rodna Mts., Cornaia supra izv. Repede, 1800–1900 m (29.7.1948, NYÁRÁDY, BUC 151096).