CARYOLOGIA Vol. 64, no. 1: 55-66, 2011

*Corresponding author: e-mail: melahat.ozcan@yahoo.com

Chromosome reports in some Cirsium (Asteraceae, Cardueae) taxa from north-east Anatolia

Ozcan* Melahat, Sema Hayırlıoglu-Ayaz and Huseyın Inceer

Department of Biology, Faculty of Science, Karadeniz Technical University, 61080 Trabzon, Turkey.

Abstract — This paper comprises mitotic chromosome numbers of 37 accessions representing 24 taxa and 1 interspecifi c hybrid of Cirsium from north-east Anatolia. Twelve counts are new to science, and eighteen counts and one new ploidy level are reported for the fi rst time from Turkey. Other counts are the confi rmations of lim-ited or unique previous data. In the present study, the chromosome numbers of the studied taxa are determined as 2n = 34, 36 and 68 respectively. Moreover, B chromosomes are reported in two taxa. The relevance of the results are also discussed in relation to the available data for each of the analysed taxa.

Key words: Asteraceae; Cardueae; chromosomes; Cirsium; hybrid; Turkey.

INTRODUCTION

Taxonomic classifi cation of plant species is initially based on morphological characteristics. Because these features vary, other characters such as chromosome number and chromosome morphology are increasingly used in plant tax-onomy. These data are also important to eluci-date the origin, speciation and phlogenetic re-lationships of plants (STEBBINS 1971; CAI et al. 2004; PAVLOVA and TOSHEVA 2005). Cirsium is a large and widespread genus of complex taxono-my (HARRINGTON 1954; KEIL and TURNER 1993; CRONQUIST 1994) and its relationship with pu-tatively allied genera Carduus L. and Cnicus L. in the tribe Cardueae. (DAVIS and PARRIS 1975; CHARADZE 1998).

The genus Cirsium Miller is represented more than 250 taxa or variable number de-pending on the authors (SMITH 1977; ZOMLEFER 1994). This perennial, biennial or rarely annual, spiny species distributed in Europe, North Afri-ca, Siberia, Central Asia, West & East Africa and

Central America. Most of this species richness is concentrated in the mountains of southern Eu-rope and Caucasia (WERNER 1976; MEUSEL and JÄGER 1992; GARCIA - JACAS et al. 2001).

According to recent studies, this genus is rep-resented 77 taxa at the level of species, subspecies and variety in Turkey (DAVIS and PARRIS 1975; DAVIS et al. 1998; GUNER et al. 2000; DAŞKIN et al. 2006; YILDIZ and DIRMENCI 2008; YILDIZ et al. 2009a,b; 2011). Twenty eight of these taxa are endemic to Turkey resulting in an endemism ra-tio of 36%. Endemic species are most frequently found in the section of Epitrachys DC. (DAVIS and PARRIS 1975).

Cirsium is a typical example of a genus with a high affi nity to form interspecifi c and intersub-specifi c hybrids. Under certain circumstances, the sterility barriers between closely related spe-cies of Cirsium break down, permitting hybridi-zation and backcrossing. Hybrids also exhibits wide interfertility limits which translate into a large number of interspecifi c and intersubspecif-ic hybrids. Several hybrids have been described from Caucasus and adjoining regions of Asia minor (DAVIS and PARRIS 1975; WERNER 1976; CHARADZE 1998; BUREŠ et al. 2004; SEGARRA - MORAGUES et al. 2007).

Different chromosome counts have been re-ported in the genus Cirsium till now. However

OZCAN, HAYIRLIOGLU-AYAZ and INCEER56

the most frequent chromosome number with in the genus Cirsium is diploid 2n = 34 and tetra-ploid 2n = 68. In addition to these numbers, other counts such as 2n = 16, 18, 20, 22, 51, 56, 102 are also known (AISHIMA 1934; ARANO 1957; 1963; FRANKTON and MOORE 1961; 1963; FEDOROV 1969; MOORE and FRANKTON 1962a; b; 1963; 1966; 1967; 1969; 1974; TONIAN 1981a; b; c; 1982; GOLDBLATT 1981; 1984; 1985; 1988).

The taxonomy of Cirsium has received much attention. However, cytological data for the ge-nus are scarce. Taking all papers into account, few cytological data have been noted different countries and only one cytotaxonomic study has been represented from Turkey, so far. The present paper continues our previous cytotaxo-nomic investigations, which involved 12 taxa of Cirsium distributed in Turkey. Twenty four taxa and one hybrid were studied in the course of the present work.

MATERIAL AND METHODS

Material collection - Plants for chromosome counts of the genus Cirsium were collected from 37 localities in the mountains of north-east Ana-tolia, Turkey in 2007-2009. The localities are; Giresun: Yavuzkemal (M. Ozcan 209, 212, 213), Egribel (M. Ozcan 220, 221) and Şebinkarahisar (M. Ozcan 216), Gümüşhane: Köse (M. Ozcan 191, 192, 196, 199, 201), Kelkit (M. Ozcan 203), and Gezge village (M. Ozcan 171), Bayburt (M. Ozcan 191) and Kop Dagı (M. Ozcan 289), Tra-bzon: Akçaabat (M. Ozcan 242), and Çaykara (M. Ozcan 254, 303), Rize: Ispir (M. Ozcan 250, 251, 252), Çamlıhemşin (M. Ozcan 187, 189, 223, 224), Ikizdere (M. Ozcan 268, 276, 277), Artvin: Ardanuç (M. Ozcan 159, 290), Murgul (M. Ozcan 262), Borçka (M. Ozcan 281, 278), Şavşat (M. Ozcan 230, 231, 232), and Yusufeli (M. Ozcan 248). The examined plant materials are grouped into sections and arrangated phy-logenetical order (Table 1). Voucher specimens are deposited in the herbarium at the Karadeniz Technical University, Department of Biology (KTUB).

Chromosome preparations - Mitotic chromo-somes were studied in root meristematic cells. Root tips of 1-2 cm were cut off and pre-treated with saturated solution of 1-Bromonapthalene at 4ºC temperature for 16 h, fi xed in fresh Carnoy’s solution (3:1 ethanol - glacial acetic asid) for 24 h at 4ºC, rinsed in distilled water, hydrolyzed in 5 N HCI for 4-5 min at room temperature (OZ-

CAN et al. 2008) and stained with lacto-propionic orcein for at least 2-3 hours at room tempera-ture. Squashing was done in a drop of acetic acid 45% and slides were permanent using Entellan. The best metaphase plates were photographed using Leica DM 4000 microscope with digital camera attachment Leica DFC 490.

To assess the existence of previously pub-lished chromosome counts in the studied spe-cies, we used the chromosome number databases of plant chromosome numbers cited by OZCAN et al. (2008), as well as the online chromosome number databases, Index to Plant Chromosome Numbers (Missouri Botanical Garden, http://mobot.org/W3T/search/ipcn.html) and Index to Chromosome Numbers in Asteraceae (Info-Lib Index to Chromosome Numbers in Aster-aceae, http://lib.kobe-u.a.c.jp/infolib/meta_pub/G0000003asteraceae_e).

RESULTS AND DISCUSSION

The chromosome numbers of 37 accessions of Cirsium plants were determined in this study. All sections and taxa follow the classifi cations by DAVIS and PARRIS (1975). The analysed spe-cies, their chromosome numbers and accessions numbers are given in Table 1. These results in-clude the fi rst chromosome count for 11 taxa and 1 hybrid (*), new ploidy level in 1 taxa (++), and new chromosome counts for 5 taxa (+) from Turkey. The results obtained for each species in relation to previous studies and the cytological information available for the taxa are discussed.

Sect. Epitrachys DC.C. bulgaricum DC - It is a biennial species

and Euxine element that grows in Pinus forests, open mountain woods, roadsides, and meadow areas with the altitudes of 1100-1600 m.

Our counts as 2n = 34 from two populations are the fi rst in material from Turkey (Table 1, Fig. 1). Two different chromosome counts have been reported in literature till now. Chromo-some number of this species was reported by KUZMANOV et al. (1983) from Bulgaria as 2n = 30, and by FAVARGER (1969) from France as 2n = 34. Our results are in accordance with FAVARGER (1969). The diverging karyological results may be arised from intraspecifi c karyological differ-entiation. Such differentiations are also present in literature (HAYIRLIOGLU-AYAZ and INCEER 2006; OZCAN et al. 2008).

CHROMOSOME REPORTS IN SOME CIRSIUM (ASTERACEAE, CARDUEAE) TAXA FROM NORTH-EAST ANATOLIA 57

Figs. 1-12 — Somatic metaphases. (1) Cirsium bulgaricum 2n = 34. (2) C. sommieri 2n = 34. (3) C. osseticum 2n = 34. (4) C. caucasicum 2n = 34. (5) C. adjaricum 2n = 34. (6) C. cephalotes 2n = 34. (7) C. macrobotrys 2n = 34. (8) C. kos-melii 2n = 34. (9) C. lappaceum subsp. lappaceum 2n = 34. (10) C. lappaceum subsp. lappaceum 2n = 34 + 1B. (11) C. lappaceum subsp. anatolicum 2n = 34. (12) C. rigidum 2n = 34. Arrow indicates B-chromosome. Bars: 10 µm.

OZCAN, HAYIRLIOGLU-AYAZ and INCEER58

C. sommieri Petrak - It is an endemic, peren-nial species and Ir.-Tur element that grows in the forest, rocky slopes, roadsides, cultivated and waste places with the altitudes of 60- 2100 m.

The chromosome number of this species is found as 2n = 34 from three populations (Table 1, Fig. 2). Our results confi rm only one previous count reported by OZCAN et al. (2008).

C. osseticum (Adams) Petrak - This species is a perennial that grows on rocky slopes, in Picea forest and roadsides with the altitudes of 900-1500 m.

This species is a diploid and possesses 2n = 34 chromosomes (Table 1, Fig. 3). Only one pre-vious chromosome report was carried out by TONIAN (1982) from Armenia as 2n = 34 in the species. Our count seems to be the second for this species and corroborates the former report.

C. caucasicum (Adams) Petrak - It is a peren-nial species and Euxine element that grows in roadsides and open banks in Picea forest with the altitudes of 1300-2300 m.

The chromosome number of this species is found as 2n = 34 (Table 1, Fig. 4). According to our result, this is the fi rst cytological report for this species. In our study, in addition to normal cells with 2n = 34, some aneuploid cells with 2n = 36 are also observed. The aneuploids in this species found here represent simply gain of normal chromosomes, as is to be expected of products of the irregular meiosis. Similar situa-tion was reported by OZCAN et al. (2008) in C. munitum (Bieb.) Fischer.

C. adjaricum Somm. & Lev. - A perennial spe-cies and Euxine element that grows in pastures, roadsides and meadow areas with the altitudes of 580-1800 m.

In our study, it is determined somatic chro-mosome number as 2n = 34 (Table 1, Fig. 5). Ac-cording to our knowledge, this is the fi rst report for this species distributed in north-east Anato-lia and Georgia.

C. cephalotes Boiss. - It is a perennial species and Euxine element that grows on slopes, fallow fi elds and roadsides with the altitudes of 1600-2450 m.

This species is a diploid and has 2n = 34 chro-mosomes (Table 1, Fig. 6). To our knowledge, this is the fi rst count for this species distributed in Transcaucasia and Bulgaria.

C. macrobotrys (C. Koch) Boiss. - This species is a biennial that grows in pastures, fallow fi elds, roadsides and waste lands with the altitudes of 1300-2450 m. It has involucres, directly attached to the stem.

Chromosome number of this species is found as 2n = 34 (Table 1, Fig. 7). Previous count in the species was reported by OZCAN et al. (2008) as 2n = 34. This is the second count for this Turkish species. Our result agrees with former one.

C. kosmelii (Adams) Fisch. ex Hohen - This is perennial species that grows on rocky slopes, roadsides and stony pastures with the altitudes of 2100-2300 m.

In the present study, the chromosome number of this species is found as 2n = 34 (Table 1, Fig. 8). This is the fi rst chromosome count for this species, distributed in Transcaucasia.

C. lappaceum (Bieb.) Fischer subsp. lap-paceum - A many stemmed and perennial plant grows in pastures, roadsides and limestone sub-strates with the altitudes of 1500-2000 m. It has whitish corollas and loose rasemose panicles.

It is determined that somatic chromosome number is 2n = 34 (Table 1, Fig. 9 and 10) and also one B chromosome is detected in some cells (55 % of counted cells). Till now, only tetraploid cytotype has been known for this taxon (NOU-ROOZI et al. 2010) from Iran. Our count is the fi rst as diploid level and fi rst B chromosome re-port in this taxon, and also the fi rst B chromo-some report in the genus Cirsium from Turkey.

C. lappaceum subsp. anatolicum Petrak - A many stemmed and perennial plant grows in subalpine pastures, roadsides and marl hills with the altitudes of 1100-2000 m.

Two populations of this taxon were studied in this study and both have the same chromo-some number as 2n = 34 (Table 1, Fig. 11). These are also the fi rst counts for this taxon.

C. rigidum DC - This species is a perennial and Ir.-Tur element that grows on stony pas-tures and eroded slopes with the altitudes of 450-560 m.

The somatic chromosome number of this species is 2n = 34 (Table 1, Fig. 12). According to our knowledge, this is the fi rst report for this species which is distributed in north-east Anato-lia, Azerbaijan and Armenia.

CHROMOSOME REPORTS IN SOME CIRSIUM (ASTERACEAE, CARDUEAE) TAXA FROM NORTH-EAST ANATOLIA 59

Figs. 13-20 — Somatic metaphases. (13) C. aggregatum 2n = 34. (14) C. vulgare 2n = 68. (15) C. echinus 2n = 34. (16) C. hypoleucum 2n = 34. (17) C. pseu-dopersonata subsp. pseudopersonata 2n = 34. (18) C. pseudopersonata subsp. kusnezowianum 2n = 34. (19) C. obvallatum 2n = 34. (20) C. rhizocephalum subsp. rhizocephalum 2n = 34. Bars: 10 µm.

C. aggregatum Ledeb. - It is a perennial species and Euxine element that grows on marshy mead-ows and steep moist hayfi elds with the altitudes of 2000-2150 m. It has sessile involucres at the apex.

This species possesses 2n = 34 chromosomes (Table 1, Fig. 13). This is also the fi rst count for this species, distributed in north-east Anatolia and Georgia.

OZCAN, HAYIRLIOGLU-AYAZ and INCEER60

TABLE 1 — Somatic chromosome numbers (2n), accessions number and previous chromosome counts of Cirsium taxa investigated.

Taxa 2n Accessions no. Previous counts +C. bulgaricum 34 M.Ozcan 209

M.Ozcan 242 Favarger 1969; n = 17 (France)Kuzmanov et al. 1983; 2n = 30 (Bulgaria)

C. sommieri 34 M.Ozcan 191 M.Ozcan 251 M.Ozcan 285

Ozcan et al. 2008; 2n = 34 (Turkey)

+C. osseticum 34 M.Ozcan 224 Tonian 1982; 2n = 34 (Armenia)* C. caucasicum 34, 36 M.Ozcan 159

M.Ozcan 232–

* C. adjaricum 34 M.Ozcan 281 –* C. cephalotes 34 M.Ozcan 289 – C. macrobotrys 34 M.Ozcan 252 Ozcan et al. 2008; 2n = 34 (Turkey)* C. kosmelii 34 M.Ozcan 230 –++C. lappaceum subsp. lappaceum 34+0-1B M.Ozcan 203 Nouroozi et al. 2010; 2n = 68 (Iran)* C. lappaceum subsp. …anatolicum 34 M.Ozcan 196

M.Ozcan 216–

* C. rigidum 34 M.Ozcan 290 –* C. aggregatum 34 M.Ozcan 303 – C. vulgare 68 M.Ozcan 212

M.Ozcan 199 M.Ozcan 268

Löve and Löve 1961; 2n = 34 (Sweden)Tischler 1950; 2n = 56-60, (Germany)Morton 1977, Dempsey et al. 1994; 2n = 68 (British Isles) Kuzmanov et al. 1991; 2n = 34 (Bulgaria)Krasnikov et al. 2003; 2n = 60 (Russia)Ozcan et al. 2008; 2n = 34, 60, 68 (Turkey)Nouroozi et al. 2010; 2n = 68 (Iran)

C. echinus 34 M.Ozcan 220 Tonian 1981c; 2n = 34 (Armenia)Ozcan et al. 2008; 2n = 34 (Turkey)

* C. hypoleucum 34 M.Ozcan 223 – C. pseudopersonata subsp. pseudop-ersonata

34 M.Ozcan 221 M.Ozcan 262

Ozcan et al. 2008; 2n = 34 (Turkey)

* C. pseudopersonata subsp. kusn-ezowianum

34 M.Ozcan 278 –

+C. obvallatum 34 M.Ozcan 189M.Ozcan 231

Poghosian et al. 1971; 2n = 34 (Armenia)Tonian 1981b; 2n = 34 (Armenia)

+C. rhizocephalum subsp. rhizo-cephalum

34 M.Ozcan 201 Podlech and Bader 1974; 2n = 34 (Afghanistan)Tonian 1981b; 2n = 34 (Armenia)

C. rhizocephalum subsp. sinuatum 34 M.Ozcan 254 Ozcan et al. 2008; 2n = 34 (Turkey)* C. pubigerum var. ...glomeratum 68+0-3B M.Ozcan 187 –* C. pubigerum var. caniforme 68 M.Ozcan 248

M.Ozcan 277–

+C. elodes 34 M.Ozcan 171 M.Ozcan 250

Tonian 1981a; 2n = 34 (Armenia)Nouroozi et al. 2010; 2n = 68 (Iran)

C. arvense subsp. vestitum 34 M.Ozcan 213 M.Ozcan 192

Van Loon and Kieft 1980; 2n = 34 (Yugoslavia)Kuzmanov et al. 1981; 2n = 68 (Bulgaria)Ozcan et al. 2008; 2n = 34 (Turkey)

* C. adjaricum x C. trachylepis 34 M.Ozcan 276 –

*: First chromosome count, ++: New ploidy level, +: New record from Turkey

CHROMOSOME REPORTS IN SOME CIRSIUM (ASTERACEAE, CARDUEAE) TAXA FROM NORTH-EAST ANATOLIA 61

C. vulgare (Savi) Ten. - This is a widespread and biennial species that grows in roadsides banks, streamsides, slopes, drains and open pas-tures with the altitudes of 10-1950 m.

It is determined that the somatic chromo-some number is 2n = 68 (Table 1, Fig. 14). Sev-eral chromosome counts are present in literature for these widely distributed common weeds. Chromosome count of this species previously reported by MORTON (1977) and DEMPSEY et al. (1994) as 2n = 68 in British isles, by KUZMANOV et al. (1991) in Bulgarian population and LOVE and LOVE (1961) in Swedish population as 2n = 34, by KRASNIKOV et al. (2003) as 2n = 60 and FEDOROV (1969) as 2n = 56, 60, 68 in Russian populations, by TISCHLER (1950) as 2n = 56-60 in Germanian population, by OZCAN et al. (2008) as 2n = 34, 60, 68 in Turkish populations, and by NOUROOZI et al. (2010) as 2n = 68 in Iranian population. The tetraploid x = 17 based cytotype is found from two populations in this study. This is the second count from Turkey and is in agreement with pre-vious counts.

Sect. CirsiumC. echinus (Bieb.) Hand.-Mazz. - It is a bien-

nial species and Ir.-Tur element that grows on rocky slopes and roadsides with the altitudes of 1200-2100 m.

The chromosome number of this species is found as 2n = 34 (Table 1, Fig. 15). To our knowledge, this is the third count for this spe-cies. The earlier reports from Armenia (TONIAN 1981a) and Turkish populations (OZCAN et al. 2008) and that presented here agree with them.

C. hypoleucum DC. - It is a perennial species and Euxine element that grows on shady banks, among grass and in forests with the altitudes of 900-1200 m.

This species is a diploid and has 2n = 34 chro-mosomes (Table 1, Fig. 16). To our knowledge, this is the fi rst count for this species distributed in Turkey and Georgia.

C. pseudopersonata Boiss & Bal. subsp. pseu-dopersonata - An endemic species and Euxine element that grows in Picea forest, shady banks, streamsides and alpine meadows with the alti-tudes of 1450-2240 m. It has generally densely fl occose-arachnoid hairs in the lower leaf surface.

In the present study, it is determined chro-mosome number as 2n = 34, from two different populations (Table 1, Fig. 17). To our knowl-edge, this is the second count for species, en-

demic in the north-east Anatolia. It agrees with the only previous report (OZCAN et al. 2008).

C. pseudopersonata subsp. kusnezowianum (Somm. & Lev.) Petrak - This is a perennial spe-cies and Euxine element that grows by shady banks, stream sides and alpine meadows with the altitudes of 1400-1600 m. It has short scattered or subglabrous hairs in the lower leaf surface.

It possesses 2n = 34 chromosomes (Table 1, Fig. 18). This is also the fi rst count for this taxon which is distributed in north-east Anatolia and Georgia.

C. obvallatum (Bieb.) Fischer - It is a peren-nial species and Hyrcano-Euxine element that grows on rocky igneous slopes, near streamsides, meadowy place and roadsides with the altitudes of 2000-2400 m.

Two populations of this species were stud-ied in this study and both have the same chro-mosome number as 2n = 34 (Table 1, Fig. 19). POGHOSIAN et al. (1971) and TONIAN (1981b) previously reported the chromosome number of its as 2n = 34. Our fi rst count, based on Turkish plants, is in accordance with these counts.

C. rhizocephalum C.A. Meyer subsp. rhizo-cephalum - This is a perennial, deeply pinnatifi d leafy, stoutly spiny and dwarf taxon that grows on mountain pastures, damp turf and by stream with tha altitudes of 2700-3000 m.

We found the chromosome number of this taxon as 2n = 34 (Table 1, Fig. 20). PODLECH and BADER (1974) who studied plants from Af-ghanistan and TONIAN (1981b) who studied plants from Armenia reported the chromosome number of its as 2n = 34. Our count, from Tur-key, agrees with former reports.

C. rhizocephalum subsp. sinuatum (Boiss.) Davis & Parris - This is a perennial, subentire-simple leafy, soft spiny and dwarf taxon that grows on mountain pastures, damp turf and by stream with the altitudes of 2700-3000 m.

Our studied taxon possesses 2n = 34 chromo-somes like the other subspecies of C. rhizocepha-lum (Table 1, Fig. 21). Our counts corroborate the only former count cited in OZCAN et al. (2008).

C. pubigerum var. glomeratum (Freyn & Sint.) Davis & Parris - It is a perennial and variable taxa with in habit and leaf shape that grows in forests and near streamsides with the altitudes of 2000-2100 m.

OZCAN, HAYIRLIOGLU-AYAZ and INCEER62

The chromosome number of this species is determined as 2n = 68 and also three B chromo-somes are detected in some cells (Table 1, Fig. 22). This is the fi rst chromosome count and fi rst B chromosome report in this taxon.

C. pubigerum var. caniforme Petrak - It is per-ennial and variable taxa with in habit and leaf shape that grows in forests and near streamsides with the altitudes of 600-1400 m.

Chromosome number of this taxon from two different populations is determined as 2n = 68 (Table 1, Fig. 23). To our knowledge, this is the fi rst count for this taxon, distributed in N. Ana-tolia and N. Iraq.

C. elodes Bieb. - This is a perennial species and Ir.-Tur element. It has tall and continu-ously winged stems that grows in water mead-ows, streamsides and spring with the altitudes of 1300-2400 m

Our counts from two populations show that it is diploid species with chromosome number as 2n = 34 (Table 1, Fig. 24). This species was initially described as diploid (2n = 2x = 34) from Armenia (TONIAN 1981a). However, tetraploid cytotype (2n = 4x = 68) was recently detected in Iranian material (NOUROOZI et al. 2010). Our count is the fi rst report from Turkish population and is in accordance with the previous report from Armenia.

Figs. 21-26 — Somatic metaphases. (21) C. rhizocephalum subsp. sinuatum 2n = 34. (22) C. pubigerum var. glomeratum 2n = 68 + 3B. (23) C. pubigerum var. cani-forme 2n = 68. (24) C. elodes 2n = 34. (25) C. arvense subsp. vestitum 2n = 34. (26) C. adjaricum x C. trachylepis 2n = 34. Arrow indicates B-chromosome. Bars: 10 µm.

CHROMOSOME REPORTS IN SOME CIRSIUM (ASTERACEAE, CARDUEAE) TAXA FROM NORTH-EAST ANATOLIA 63

Sect. Cephalonoplos (Necker) DC.C. arvense subsp. vestitum (Wimmer &

Grab.) Petrak - It is a dioecious, perennial and widespread plant that grows on roadsides, river-banks, cultivated land, meadow areas and steppe with the altitudes of 990-1500 m.

The chromosome number of this taxon, counted from two different populations, is found as 2n = 34 (Table 1, Fig. 25). Chromosome count of this taxon was previously reported by VAN LOON and KIEFT (1980) with 2n = 34 and by KUZMANOV et al. (1981) with 2n = 68. These results indicate that this taxon is cytologically heterogeneous, showing populations with dif-ferent ploidy levels. Our counts in two popula-tions agree with previous counts from Yugosla-via (VAN LOON and KIEFT, 1980). According to our previous count (OZCAN et al. 2008) and this present count (totaly four populations), the only diploid cytotype is present in Turkey.

C. adjaricum x C. trachylepis - This is a peren-nial hybrid plant and allied with the two paren-tal species and displays intermediary situations. It grows in open slopes and roadsides with the altitudes of 2000-2100 m.

Chromosome number of this hybrid is deter-mined as 2n = 34 like parental species (Table 1, Fig. 26). This is the fi rst chromosome number record for this interspecifi c hybrid. It was report-ed that hybrids of diploid (2n = 34) species of the genus Cirsium generally have the same chromo-some numbers as the parental species (ROTREK-LOVÁ et al. 2004). The same hybrid chromosome numbers were reported by CZAPIK (1958) in Cir-sium canum x C. oleraceum, by LAVRENKO et al. (1991) in Cirsium heterophyllum x C. palustre, by BUTTLER (1989) in Cirsium oleraceum x C. palus-tre and by ROTREKLOVÁ et al. (2004) in Cirsium oleraceum x C. rivulare as their parental species. This situation was detected BUREŠ et al. (2004) in many samples using fl ow cytometry.

Several basic chromosome numbers such as x = 8, 9 in the genera of Achillea (SHEIDAI et al., 2009) and Artemisia (PELLICER et al. 2007), x = 9 in the genera of Tripleurospermum (INCEER and HAYIRLIOGLU-AYAZ 2010) and Hieracium (COŞKUNCELEBI and HAYIRLIOGLU-AYAZ, 2006), x = 12 in the genus Bidens (GROMBONE-GUARATI-NI et al., 2006), x = 9, 10,11,12,13 in the genus Cousinia (SUSANNNA et al. 2003; GHAFFARI et al. 2006; DJAVADI and ATTAR 2010), and x = 17 in the genus Jurinea (DOĞAN et al. 2009) have been re-ported in the family Asteraceae. In the present study, the mitotic cell of most of Cirsium taxa

examined were determined as 2n = 34 chromo-somes with the basic number of x = 17. In three taxa (C. vulgare, C. pubigerum var. caniforme and C. pubigerum var. glomeratum), however, tetraploidy was also observed. In Turkey, this genus generally presents low level of variation in chromosome number in most of the species like Jurinea (DOĞAN et al. 2009) and Bidens (GROM-BONE-GUARATINI et al. 2006). Diploid chromo-some number, as 2n = 34, is generally reported by several authors in Cirsium species (FRANKTON and MOORE 1961; MOORE and FRANKTON 1963; OWNBEY 1968; OWNBEY et al. 1975). However, a reduction in gametic number from 17 to 9 run into in extreme cases in America has been also reported by several authors (FRANKTON and MOORE 1961; MOORE and FRANKTON 1963; OWNBEY 1968; OWNBEY et al. 1975). It is noted that the reduction, accompanied by transloca-tion between non-homologous chromosomes is an important evolutionary mechanism in plants and has played a signifi cant role in speciation (OWNBEY et al. 1975; DABEYDEEN 1980; OTTO and WHITTON 2000). On the other hand, it is claimed that Cirsium is paraphyletic and paralel mutations Cirsium with Carduus L. (FRANKTON and MOORE 1961) or Notobasis Cass. are coming into being (HÄFFNER and HELLWIG 1999). How-ever, in Turkish Cirsium taxa, the reduced num-bers have not been encountered so far.

According to the present study and our pre-vious chromosome reports (OZCAN et al. 2008), the mitotic cell of most of taxa examined possess 2n = 34 chromosome, whatever their geographi-cal locality and their taxonomic position. More-over, B chromosomes 0-3 were observed in C. lappaceum ssp. lappaceum and C. pubigerum var. glomeratum, in this study. It has been reported by several authors that this accessory chromo-somes (B-chromosomes) like to the A-chro-mosomes. Therefore it is diffi cult to determine correct chromosome number of the species (FRANKTON and MOORE 1963; DABYDEEN, 1980). B-chromosomes have been also reported in some other Cirsium species like C. muticum Michx. (1 B-chromosome) and C. acaule (L.) Scop. (6 B-chromosome) by MOORE and FRANKTON (1962a), and FRANKTON and MOORE (1963). We believe this study will play a positive role to enlighten the taxonomically complex genus. On the other hand, to evaluate ultimately taxonomic position of this genus in Turkey, it is necessary to carry out further chromosomal studies on Cirsium taxa. No detailed karyotype analyses present in literature. Some karyological investigations have

OZCAN, HAYIRLIOGLU-AYAZ and INCEER64

been also continued on Turkish Cirsium taxa by OZCAN (2009). A combination of detailed karyo-type analysis with molecular studies is needed in order to overcome taxonomic problems of Cir-sium plants.

Acknowledgements — The authors would like to thank Mr. Uzeyir OZCAN for his kindly help during the collection of samples and Mr. Aykut SAGLAM for technical help in the prepar-ing of manuscript. This research was supported by the Research Fund of Karadeniz Technical University (Project number: 2006.111.004.4). We should also like to thank anonymous review-ers for valuable helps on improving an earlier draft of the paper.

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Received June 17th 2010; accepted January 22th 2011