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 sub- species of Arabidopsis halleri in the area studied based on revised herbarium spec- imens 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 ma- terial. 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 de- tailed 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.