Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 72(1): 45-53, 2019 Firenze University Press www.fupress.com/caryologiaCaryologia International Journal of Cytology, Cytosystematics and Cytogenetics ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.13128/cayologia-250 Citation: İ. Genç, M. Firat (2019) Kar- yological study of the genus Gundelia (Compositae) in Turkey. Caryologia 72(1): 45-53. doi: 10.13128/cayolo- gia-250 Received: 13th june 2018 Accepted: 4th December 2018 Published: 10th May 2019 Copyright: © 2019 İ. Genç, M. Firat. This is an open access, peer-reviewed article published by Firenze University Press (http://www.fupress.com/caryo- logia) and distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All rel- evant data are within the paper and its Supporting Information files. Competing Interests: The Author(s) declare(s) no conflict of interest. Karyological study of the genus Gundelia (Compositae) in Turkey İlker Genç1,*, Mehmet Firat2 1 Department of Pharmaceutical Botany, Faculty of Pharmacy, İstanbul University, İstanbul, Turkey 2 Department of Biology Education, Faculty of Education, Van Yüzüncü Yıl University, Van, Turkey * Corresponding author: gencilker1@gmail.com Abstract. Karyotypes in 12 taxa of Gundelia are compared, based on Feulgen-stained somatic metaphase chromosomes. The karyotypes of G. anatolica, G. asperrima, G. cilicica, G. colemerikensis, G. dersim, G. glabra, G. komagenensis, G. mesopotamica, G. munzuriensis and G. vitekii are described for the first time. Karyological analyses indicate relationships among the species with respect to their asymmetry indices. All Gundelia species studied were diploid with 2n = 2x = 18 chromosomes. All karyotypes are symmetrical, consisting of metacentric and submetacentric chromosomes. The sub- metacentric chromosomes of all the investigated specimens contain a secondary con- striction. Three chromosome types were identified according to the position of the sec- ondary constrictions. The chromosomes ranged in size from 2.00 µm to 7.02 µm. The total haploid chromosome length (THL) varied from 24.97 μm (G. asperrima) to 42.56 μm (G. rosea). To determine the karyological relationships among taxa, PCoA (Princi- pal Coordinate Analysis) with six uncorrelated parameters was performed. Keywords. Chromosome number, karyotype asymmetry, secondary constriction, Cichorieae, Scolyminae. INTRODUCTION Gundelia L. was first recorded in one of the earliest natural history col- lections made in the Near East by the German physician, botanist and trav- eller, Leonhard Rauwolf (1535–1596) at 16. century. However, the plant was first evaluated in Silybum then Eryngium groups. About 125 years later, in the early years of the 18th century, Joseph Pitton de Tournefort saw the plant in the natural habitat. Moreover, he concluded that the plant should be called Gundelia (Hind 2013). Finally, the plant was named Gundelia tourneforti by Linnaeus, in accordance with the binomial nomenclature (Linnaeus 1753). The infrafamilial position of Gundelia has also changed over time (Hind 2013). The last tribal position of genus Gundelia was Cichorieae Lam. & DC., subtribe Scolyminae Less., along with Catananche L., Hymenonema Cass., and Scolymus L. (Kilian et al. 2009). The genus grows in the semi-desert areas, and it is distributed in the Mediterranean to Central/Eastern Asia (Karis et al. 2001). 46 İlker Genç, Mehmet Firat Some different species and varieties have been described over the years, although many authors have treated Gundelia as monospecific and the wide vari- ation in corolla colour was considered unrelated to gross morphology (Komarov 1961; Kupicha 1975; Fein- brun Dothan 1978; Meik le 1985; Rechinger 1989). However, in the last decades, researches on Gundelia increased. Numerous new species have been published as a result of research on live and more abundant mate- rials. The genus is currently represented by 16 species, of which 12 (10 endemic) in Turkey (Vitek et al. 2010, 2014, 2017; Nersesyan 2014; Armağan 2016; Fırat 2016, 2017a, 2017b, 2017c; Vitek and Noroozii 2017; Vitek 2018). These are Gundelia anatolica Fırat, G. asperrima (Trautv.) Fırat, G. cilicica Fırat, G. colemerikensis Fırat, G. dersim Vitek, Yüce & Ergin, G. glabra Mill., G. kom- agenensis Fırat, G. mesopotamica Fırat, G. munzurien- sis Vitek, Yüce & Ergin, G. rosea M.Hossain & Al-Taey (non-endemic), G. tournefortii L. (non-endemic), and G. vitekii Armağan. The chromosome numbers of Cichorieae range between 2n = 14x = 126 chromosomes in Sonchus (Beuzenberg and Hair 1984; Dawson 2000) and 2n = 2x = 6 in some species of Crepis (Ikeda 1988; Gupta and Gill 1989; Dimitrova and Greilhuber 2001). The basic chromosome number in the majority of the subtribes is x = 9, or a descending series starting with x = 9 to x = 3. Subtribe Scolyminae has the basic chromosome num- bers x = 9 and 10 (Kilian et al. 2009). The only chromo- some number Gundelia is 2n =18 (Waisel 1962; Al-Taey and Hossain 1984; Ghaffari and Chariat-Panahi 1985; Nersesyan and Nazarova. 1989; Ghukasyan and Janjug- hazyan 2015). Fig. 1. Synflorescences of the studied taxa. (a) G. anatolica; (b) G. asperrima; (c) G. cilicica; (d) G. colemerikensis; (e) G. dersim; (f ) G. glabra; (g) G. komagenensis; (h) G. mesopotamica; (i) G. munzuriensis; (j) G. rosea; (k) G. tournefortii; (l) G. vitekii. 47Karyological study of the genus Gundelia (Compositae) in Turkey This study aimed to determine the chromosome numbers and karyomorphology of al the 12 Gundelia species occurring in Turkey (Figure 1). MATERIALS AND METHODS Twelve Gundelia species were analysed in this study. A list of examined specimens is provided in Table 1. All endemic taxa were collected from their type localities. Voucher specimens were deposited in the Herbarium of University of Van Yüzüncü Yıl (VANF) and in a private herbarium (Herb. M. Fırat). For karyological observa- tions, four to eight individuals were used for each spe- cies in this study. Mitotic metaphase cells of root tips were obtained from germinated seeds which were col- lected in natural habitats from Turkey. Mitotic chromosomes were prepared from root tips and pre-treated with 0.002 M 8-Hydroxyquinoline at +4 °C for 24 h. Roots were fixed for a minimum of 2 h in absolute ethanol:glacial acetic acid, (3:1,v/v), hydro- lysed at 60 °C in 1 N HCl for 12 min. and stained with the Feulgen method. Finally, root tips were squashed in 1% aceto-orcein. Permanent slides were prepared with entellan mounting medium. Microphotographs of good quality metaphase plates were taken using an Olym- pus BX53 (Tokyo, Japan) microscope equipped with a high-resolution digital camera. Metaphase observations and chromosome measures were made using the image analysis systems KAMERAM (ARGENİT Microsystems, İstanbul, Turkey). The somatic chromosome number and karyotype details were studied in five to eighteen well-spread metaphase plates from different individuals; mean values were used for the analysis. Chromosome pairs were identified and arranged on the basis of their length and any other evident kar- yomorphological features. The nomenclature used for describing karyotype composition followed Levan et al. (1964). To determine the karyological relationships among taxa, we performed a PCoA (Principal Coordi- nate Analysis) with six uncorrelated parameters as sug- gested by Peruzzi and Altinordu (2014). These param- eters are chromosome number (2n), basic chromosome number (x), total haploid length (THL), mean centro- meric asymmetry (MCA), coefficient of variation of chro- mosome length (CVCL), and coefficient of variation of centromeric index (CVCI) (Paszko 2006; Peruzzi et al. 2009; Zuo and Yuan 2011, Peruzzi and Eroğlu 2013). The software Past 3.03 (Hammer et al. 2001, Hammer 2018) was used to perform this analysis. Mitotic metaphase chromosomes are given in Figure 3. Idiograms of these taxa are arranged in order of cen- tromere position and then decreasing the length of hom- ologue chromosome pairs (Figure 4). In this study, chromosome types were determined according to the position of the secondary constrictions of Gundelia chromosomes for the purpose of chromo- some comparison. General description of these chromo- Table 1. Localities and voucher specimens of Gundelia taxa examined in the present study. Taxa Locality Collector (M. Fırat) /voucher G. anatolica Fırat Kırıkkale: Delice province, Tuzkayası region, dry steppe, 700 m, 10 July 2017 33866 G. asperrima (Trautv.) Fırat Erzurum: Palandöken mountain, mountain slope, steppe, 2444 m, 30 July 2017 33845 G. cilicica Fırat Mersin: Erdemli province, Tozlu village, open forrest, 1460 m, 11 July 2017 33868 G. colemerikensis Fırat Hakkâri: Hakkâri Province (Colemerîk) Berçelan plateau, open erode region and steppe, 2284 m, 7 July 2017 33861 G. dersim Vitek, Yüce & Ergin Tunceli (Dersim): Ovacık, c. 11.7 km SW Ovacık, 1.9 km NE Ziyaret (fountains of river Munzur), 1300 m, 28 July 2017 33872 G. glabra Mill. Bayburt: Kop mountain arround, near Kop Village, 1897 m, 19 July 2017 33867 G. komagenensis Fırat Adıyaman: Kahta Province, Nemrut mountain, rocky steppe, 1445 m, 30 July 2017 33783 G. mesopotamica Fırat Mardin: 2-3 km from Mardin to Nusaybin (Nisêbîn), eroded slopes, aride steppe, 807 m, 11 July 2017 33865 G. munzurensis Vitek, Yüce & Ergin Tunceli (Dersim): Ovacık, c. 2 km SW Ovacık, near road Ovacık plain, 1275 m, 28 July 2017 33871 G. rosea M.Hossain & Al-Taey Hakkari: Şemdinli Province, Sad Mountain, Derîyê Kera region, meadows and stony slopes, 1662 m, 7 July 2017 33862 G. tournefortii L. Hatay: Reyhanli district, near Syria (Aleppe) border, 121 m, 9 July 2017 33864 G. vitekii Armağan Tunceli (Dersim): c. 8-9 km N of Tunceli, mountain slope NW of Tüllük Bucaği, rocky steppe, 1760 m, 28 July 2017 33870 48 İlker Genç, Mehmet Firat some types is given below, followed by the karyotype description. Type A: Longest metacentric chromosomes with two constrictions, secondary constrictions in the distal posi- tion of the long arm. Type B: Submetacentric chromosomes with two con- strictions, secondary constrictions nearly in the median position of the long arm. Type C: Submetacentric chromosomes with two constrictions and secondary constrictions located very close to the centromere on the short arm. RESULTS All species showed basic chromosome number x = 9 and diploids with 2n = 18 (Figures 3 and 4). Chro- mosome measurements and karyotype formula of the twelve analysed species are indicated in Table 2. Total haploid length, asymmetry indices, chromosome types and flower number within the partial synflorescences in the middle part are summarised in Table 3. Secondary constrictions were observed at the long or short arms of all submetacentric chromosomes, and in the distal regions of the long arms of some of the longest Table 2. Karyotype formula according to Levan et al. (1964) and measurements of the investigated taxa. (SC: the shortest chromosome length; LC: the longest chromosome length; p: mean long arm length; q: mean short arm length; SD: standard deviation; m: metacentric; sm: submetacentric). SC–LC q (μm) Mean (±SD) p (μm) Mean (±SD) p+q Mean (±SD) Karyotype formula G. anatolica 2.90 – 5.15 1.64(±0.30) 2.19(±0.44) 3.83(±0.65) 16 m + 2 sm G. asperrima 2.00 – 4.01 1.19(±0.23) 1.58(±0.42) 2.77(±0.60) 14 m + 4 sm G. cilicica 2.77 – 5.35 1.67(±0.35) 2.14(±0.46) 3.81(±0.76) 16 m + 2 sm G. colemerikensis 2.40 – 4.90 1.43(±0.33) 1.89(±0.50) 3.32(±0.74) 14 m + 4 sm G. dersim 2.78 – 5.78 1.66(±0.40) 2.18(±0.50) 3.84(±0.83) 16 m + 2 sm G. glabra 2.86 – 5.61 1.70(±0.35) 2.25(±0.56) 3.96(±0.82) 14 m + 4 sm G. komagenensis 3.10 – 6.08 1.79(±0.37) 2.42(±0.61) 4.21(±0.87) 14 m + 4 sm G. mesopotamica 2.73 – 5.27 1.55(±0.36) 2.08(±0.47) 3.63(±0.71) 14 m + 4 sm G. munzurensis 2.28 – 4.43 1.33(±0.26) 1.77(±0.43) 3.10(±0.63) 14 m + 4 sm G. rosea 3.30 – 7.02 2.03(±0.49) 2.70(±0.58) 4.73(±1.01) 16 m + 2 sm G. tournefortii 2.31 – 4.45 1.33(±0.30) 1.68(±0.37) 3.02(±0.61) 16 m + 2 sm G. vitekii 2.49 – 5.04 1.45(±0.31) 1.95(±0.51) 3.40(±0.74) 14 m + 4 sm Table 3. Karyo-morphometric parameters, symmetry indices, Chromosome types and cephaloid flowers number for investigated taxa (THL: total haploid length; MCA: mean centromeric asymmetry; CVCL: coefficient of variation of chromosome length; CVCI: coefficient of variation of centromeric index). THL MCA CVCL CVCI Chromosome Types Cephaloid. Flowers numbers G. anatolica 34.48 14.13 16.87 9.73 _/B/_ 6 G. asperrima 24.97 13.08 21.58 9.83 A/B/C 3(–4) G. cilicica 34.26 12.06 20.01 7.44 _/_/C 6(–7) G. colemerikensis 29.84 13.03 22.42 10.75 _/B/C (3–)5(–6) G. dersim 34.52 13.27 21.57 9.56 A/_/C 6–7 G. glabra 35.60 13.27 20.84 9.94 _/B/C 3–5(–6) G. komagenensis 37.90 13.95 20.56 11.94 _/B/C 3(–4) G. mesopotamica 32.64 14.38 19.70 12.44 A/B/C 6–7 G. munzurensis 27.87 13.27 20.24 10.21 A/B/C 3–5 G. rosea 42.56 14.40 21.42 8.12 A/_/C (6–) 7–8 G. tournefortii 27.16 11.41 20.20 8.83 A/_/C (5–)6(–7) G. vitekii 30.57 13.76 21.64 11.82 A/B/C 3(–5) 49Karyological study of the genus Gundelia (Compositae) in Turkey metacentric chromosomes (Figure 2). Moreover, three chromosome types were determined according to the position of the secondary constrictions (Figure 2). The chromosomes ranged in size from 2.00 µm to 7.02 µm. G. asperrima showed the smallest mean chromosome length (2.77 µm), while G. rosea the biggest (4.73 µm). Similarly, the smallest mean short arm length (q) was observed in G. asperrima (1.19 μm) and the largest mean long arm length (p) was observed in G. rosea (2.70 μm). The idiograms of the analysed species are shown in Figure 3. Fig. 2. Chromosome types according to the position of the second- ary constrictions (indicated by arrows). Fig. 3. Somatic chromosomes (2n = 18) in the studied taxa. (a) G. anatolica; (b) G. asperrima; (c) G. cilicica; (d) G. colemerikensis; (e) G. der- sim; (f ) G. glabra; (g) G. komagenensis; (h) G. mesopotamica; (i) G. munzuriensis; (j) G. rosea; (k) G. tournefortii; (l) G. vitekii. Scale bars 3 μm. 50 İlker Genç, Mehmet Firat Fig. 4. Haploid idiograms in the studied taxa. (a) G. anatolica; (b) G. asperrima; (c) G. cilicica; (d) G. colemerikensis; (e) G. dersim; (f ) G. glabra; (g) G. komagenensis; (h) G. mesopotamica; (i) G. munzuriensis; (j) G. rosea; (k) G. tournefortii; (l) G. vitekii. 51Karyological study of the genus Gundelia (Compositae) in Turkey The chromosomes with secondary constriction ranged from 2 to 6 in number. The total haploid chro- mosome length (THL) varied from 24.97 μm (G. asper- rima) to 42.56 μm (G. rosea). Karyotypes of the analysed species exhibit a pre- dominance of metacentric chromosomes. However, one or two submetacentric chromosomes were detected in each taxon. Due to the prevalence of metacentric pairs and to the absence of strong differences between small- er and larger chromosomes, asymmetry indices were in general low. However, some species show a tendency to have karyotypes distinct on asymmetry grounds: Gundelia rosea, with relatively high intrachromo- somal (MCA) and G. tournefortii with low intrachromo- somal asymmetry, also, G. vitekii with high interchro- mosomal (CVCL) and G. anatolica low interchromosom- al asymmetry. DISCUSSION Karyotype data for all taxa are reported for the first time in the present study with the exception G. tournefortii and G. rosea. The present investigation on Gundelia supports earlier data about 2n = 2x = 18 (Waisel 1962; Al-Taey and Hossain 1984; Ghaffari and Chariat-Panahi 1985; Nersesyan and Nazarova 1989; Ghukasyan and Janjughazyan 2015). The article by Ners- esyan and Nazarova (1989) is the most detailed work published on the karyology of Gundelia tournefortii. Three chromosome types were also seen in that study. However, according to our results, type B chromosome was not observed in Gundelia tournefortii chromosomes. However, earlier works accepted only one species in the genus, and this could be the cause of this difference. According to Trautvetter (1876), f lower number within the partial synflorescences differs between differ- ent populations of Gundelia. Moreover, the partial syn- florescences in the middle part of the synflorescence are formed by 3 to 8 flowers according to the articles pub- lished in recent years. These flowers have been defined as “cephaloid flowers” in the following sections of the article. The taxa examined in this article were evaluat- ed according to cephaloid flowers, and two groups have been recognised. Group I consists of mainly 3(-5) cepha- loid flowers; Group II consists of mainly 6 or more ceph- aloid flowers (Table 3). In addition to this, according to chromosome types (A-C), taxa are divided into two main groups too. In agreement with our results, these groups are quite overlapping. Namely, type B chromosomes were present in all the taxa of Group I and no type B chro- mosomes was observed in the taxa of Group II with the exception of G. mesopotamica and G. anatolica. These species differ from others by having 6-8 cephaloid flow- ered and type B chromosomes. Also, G. anatolica is the only species in which there are no type C chromosomes. According to Tarıkahya Hacıoğlu and Fırat (2017), G. Fig. 5. PCoA analysis based on six quantitative karyological parameters of investigated taxa. 52 İlker Genç, Mehmet Firat anatolica is a derived species. This difference in chromo- somal morphology supports this argument. According to PCoA analysis, the species belonging to the same group tend to cluster together substantial- ly (Figure 5). The presence of type A and type B chro- mosomes does not show variation at intraspecific level, these chromosome types were observed in all metaphase stages. However, type A chromosomes showed some infraspecific variation. 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