Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 76(1): 27-34, 2023 Firenze University Press www.fupress.com/caryologiaCaryologia International Journal of Cytology, Cytosystematics and Cytogenetics ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-1982 Citation: Sevgi Unal-Karakus, Muhammet Gaffaroglu, Muradiye Kar- asu-Ayata (2023). Comparative cytoge- netics of four endemic Capoeta (Tel- eostei: Cyprinidae) species from Ana- tolia, Türkiye. Caryologia 76(1): 27-34. doi: 10.36253/caryologia-1982 Received: January 17, 2023 Accepted: July 3, 2023 Published: September, 19, 2023 Copyright: © 2023 Sevgi Unal-Karakus, Muhammet Gaffaroglu, Muradiye Kar- asu-Ayata. This is an open access, peer-reviewed article published by Firenze University Press (http://www. fupress.com/caryologia) and distrib- uted under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, pro- vided 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. ORCID SU-K: 0000-0002-6409-7783 MG: 0000-0001-7436-5828 MK-A: 0000-0001-8890-8547 Comparative cytogenetics of four endemic Capoeta (Teleostei: Cyprinidae) species from Anatolia, Türkiye Sevgi Unal-Karakus1,*, Muhammet Gaffaroglu2, Muradiye Karasu- Ayata3 1 Department of Molecular Biology and Genetics, Faculty of Science, Bartin University Bartin, Türkiye 2 Department of Molecular Biology and Genetics, Faculty of Science and Arts, Kirsehir Ahi Evran University, Kirsehir, Türkiye 3 Department of Nutrition and Dietetics, Faculty of Health Sciences, Kirsehir Ahi Evran University, Kirsehir, Türkiye *Corresponding author. E-mail: sunal@bartin.edu.tr Abstract. The genus Capoeta is an important taxon covering a wide distribution in Türkiye. However, only a few genetic studies on Capoeta species reported from Tür- kiye. There is no cytogenetical study in Capoeta aydinensis Turan, Küçük, Kaya, Güçlü & Bektaş, 2017, Capoeta bergamae Karaman, 1969, Capoeta erhani Turan, Kottelat & Ekmekçi, 2008 and Capoeta pestai (Pietschmann, 1933). Thus, in this study, we kar- yotyped through classical cytogenetic techniques (Giemsa staining, Ag-NORs, and C-banding) the four endemic Capoeta species. The diploid chromosome number invar- iably was 150 in the four species. However, chromosome morphologies in the karyo- types had some differences between them. The number of biarmed chromosomes in the karyotypes was higher in all studied species. Their karyotypes contained respec- tively: 54 metacentric, 42 submetacentric and 54 subtelo-acrocentric in C. aydinensis, 56 metacentric, 30 submetacentric and 64 subtelo-acrocentric in C. bergamae, 50 meta- centric, 42 submetacentric and 58 subtelo-acrocentric in C. erhani and 44 metacentric, 40 submetacentric and 66 subtelo-acrocentric chromosomes in C. pestai. C-bands were on the pericentromeres of most chromosomes in the four species. Three chromosome pairs carry rDNA genes in all studied species. The chromosomal locations of these sites were varied between the species. This study provides new insights into the chromo- somal data of the hexaploid cyprinids. Moreover, obtained cytogenetic results should be conclude the cytotaxonomy of the genus Capoeta that distributed in Türkiye. Keywords: Ag-NOR, C-banding, chromosome morphology, chromosome number, scraper. INTRODUCTION Türkiye has one of the most diverse and species-rich freshwater ich- thyofaunas according to the different eco-regions that formed in Anatolian freshwaters (Küçük et al., 2009; Bektaş et al., 2017). The endemic species are https://doi.org/10.36253/caryologia-1982 https://doi.org/10.36253/caryologia-1982 https://orcid.org/0000-0002-6409-7783 https://orcid.org/0000-0001-7436-5828 https://orcid.org/0000-0001-8890-8547 28 Sevgi Unal-Karakus, Muhammet Gaffaroglu, Muradiye Karasu-Ayata much higher than in Western Asia or Europe (Küçük et al., 2009). The members of the genus Capoeta (Valenci- ennes, 1842) (Cyprinidae, Barbinae) distribute from East Europe to West Asia, including Anatolia (Bektaş et al., 2017). Seventeen species named Capoeta antalyensis, C. aydinensis, C. baliki, C. banarescui, C. barroisi, C. berga- mae, C. caelestis, C. capoeta, C. damascina, C. ekmekci- ae, C. erhani, C. oguzelii, C. pestai, C. sieboldii, C. tinca, C. trutta and C. umbla of this genus are presently recog- nized in the inland waters of Türkiye. Except for six spe- cies (C. barroisi, C. capoeta, C. damascina, C. ekmekciae, C. trutta and C. umbla) the other Capoeta members are endemic to Anatolia (Bektaş et al., 2019). Taxonomic problems still exist in Anatolian Capo- eta species and the species diversity of this genus has not been resolved (Turan et al., 2017). Özuluğ and Frey- hof (2008) collected an additional species of C. trutta from Seyhan River in Türkiye. C. turani was described as a new species from this drainage according to the dif- ferent morphological characters (Özuluğ and Freyhof, 2008). C. erhani was described in Ceyhan River of Tür- kiye by Turan et al. (2008). It was distinguished from the other members of C. trutta in the scope of morpho- logical characters (Turan et al., 2008). Otherwise, C. pestai was described from Eğirdir Lake and it was also recorded from Lake Beyşehir. In fact, Beyşehir popula- tion of C. pestai was described as a new species called C. mauricii by Küçük et al. (2009) according to the dif- ferent morphological characters. However, according to the molecular phylogeny study (cyt b gene sequences) in the genus Capoeta by Bektaş et al. (2017), C. turani was synonymized to C. erhani. Also, C. mauricii was syn- onymized to C. pestai (Bektaş et al., 2017). Otherwise, C. bergamae distributes in the western basins of Türkiye, as well as C. aydinensis was described as a new species in the recent years and is presently known from the Büyük Menderes River drainages (Turan et al., 2017). The cytogenetic studies have played an important role in describing the main features in cytotaxonomy and for understanding chromosome evolution in fish species (Gaffaroğlu et al., 2020). However, the karyo- types of fishes are poorly studied compared to the other vertebrates in response to the richness of this group. The karyotype of many fish species is still undescribed due to the difficulty of sampling the individuals, the necessi- ty of having alive individuals, in troubling to obtain kar- yotypes from cell-culture and unsuccessful in obtaining good metaphase spreads (Rossi, 2021). In this context, having too many chromosomes is another reason for this problem . Knowledge of karyotype is necessary for fish cytoge- netics. Detailed investigations of the chromosomes with Giemsa stained karyotypes have only been performed on only seven species namely, C. trutta, C. umbla (Kılıç-Demirok and Ünlü, 2001), C. capoeta, C. bar- roisi (Kaya, 2003), C. damascina (Unal and Gaffaroğlu, 2016), C. antalyensis and C. baliki (Karasu-Ayata et al., 2017) from Türkiye. The chromosomal banding prop- erties have been reported only in C. damascina (Unal and Gaffaroğlu, 2016) and C. antalyensis (Gaffaroğlu et al., 2012). Due to the lack of chromosomal reports, this study aimed to investigate karyotypes with Giemsa staining, C-banding and Ag-NOR staining in four Ana- tolian endemic Capoeta species. MATERIAL AND METHODS Cytogenetic analyses were performed on four Capo- eta species from Türkiye (Table 1, Figure 1). The alive samples were carried to the laboratory. The individuals were treated in vivo for mitotic chromosome preparation by Bertollo et al. (2015). Chromosome preparations were obtained from the cephalic kidney cells after injection of 0.1% colchicine. After hypnotization with 0.075 M KCl, fixation steps (methanol: acetic acid, 3:1) were repeated at least three times in cell suspension. At least 10 meta- phase slide was prepared from each individual. All the experiments followed ethical protocols and after sacri- ficing, the individuals were deposited in 70% ethanol in the laboratory. The process was approved by the Local Animal Ethics Committee of Türkiye (Protocol Number: 68429034/05/17). The Ag-NORs and C-banding were analysed according to the methods reported by Howell and Black (1980) and Sumner (1972). At least 100 metaphase spreads per individual were analysed to confirm the diploid chromosome number. Images were photographed using Leica DM 3000 micro- scope (Leica Microsystems GmbH, Germany) with AKAS software (Argenit Mikrosistem, Türkiye). Chromosomes were measured by digital calliper and classified as meta- centric, submetacentric and subtelo-acrocentric accord- ing to the arm ratios (Levan et al., 1964). Karyotypes were arranged manually. To count the fundamental arm number (FN) meta- and submetacentrics were considered as biarmed whereas subtelo-acrocentrics as uniarmed. RESULTS All studied Capoeta species have diploid chromo- some number 2n = 150 (Figs. 2A, 3A, 4A, 5A) with kar- yotypes composed of mainly biarmed chromosomes. Karyotype formulas were as follows: 54 metacentric, 42 29Comparative cytogenetics of four endemic Capoeta (Teleostei: Cyprinidae) species from Anatolia, Türkiye submetacentric and 54 subtelo-acrocentric in C. aydin- ensis (Fig. 2B); 56 metacentric, 30 submetacentric and 64 subtelo-acrocentric in C. bergamae (Fig. 3B); 50 metacen- tric, 42 submetacentric and 58 subtelo-acrocentric in C. erhani (Fig. 4B) and 44 metacentric, 40 submetacentric and 66 subtelo-acrocentric chromosomes in C. pestai (Fig. 5B). FN was calculated as 234 in C. pestai, 236 in C. bergamae, 242 in C. erhani and 246 in C. aydinensis. Morphologically differentiated sex chromosomes were not detected in all studied species. In terms of C-bands, C. aydinensis contains very few C-bands (Fig. 2C). These C-bands were located on the pericentromeres of chromosome pairs 18, 21, 64 and 69 (Fig. 2D). Thirteen chromosome pairs of C. bergamae has intense pericentromeric C-bands of chromosome pairs 2, 8, 37, 44, 48, 53, 56, 59, 61, 65, 66, 69, 71 (Fig. 3D). C. erhani has slightly pericentromeric C-bands of chromosome pairs 1, 2, 3, 9, 16, 17, 26, 28, 31, 42, 48, 56, 58, 60, 66 and 73 (Figs. 4C, D). Intense pericentromeric C-bands of chromosome pairs 1, 3, 7, 19, 24, 25, 30, 34, 40, 43, 45, 46, 49, 51, 55, 56, 62, 64 and 73 were found in C. pestai (Fig. 5D). Some of the other chromosomes also have less intense pericentromeric C-bands in C. pestai (Fig. 5C) and C. bergamae (Fig. 3C). Multiple Ag-NORs were found in the studied spe- cies. The common Ag-NOR number was six in four Capoeta species (Figs. 2E, 3E, 4E, 5E). These Ag-NORs were located on the terminal regions of metacentric chromosomes 1 and 5 as a strong signal and addition- ally weaker signals of chromosomes 12, 25 and 72 in C. aydinensis (Fig. 2F). Ag-NORs were detected on the terminal regions of the short arms of three submeta- centric chromosome pairs 31, 33 and 37 in C. bergamae (Fig. 3F). Ag-NORs were located on the terminal regions of the short arms of 7th metacentric, 34th and 37th sub- metacentrics in C. erhani (Fig. 4F). Ag-NORs were found on the terminal regions of the short arms of three sub- metacentric chromosome pairs 26, 28 and 30 in C. pestai (Fig. 5F). Also, Ag-NOR number polymorphisms were detected in C. bergamae (Figs. 6A, B), C. erhani (Figs. 7A, B, C, D) and C. pestai (Figs. 8A, B, C, D) in some silver stained metaphases. DISCUSSION In the subfamily Barbinae a large number of spe- cies are polyploid. This subfamily may represent a more complicated polyploid system than other vertebrates. Polyploidy (whole genome duplication), has played an important role in the evolution of cyprinids (Yang et al., 2022). From the subfamily Barbinae (which includes Figure 1. Map of the collected specimens of Capoeta species. 30 Sevgi Unal-Karakus, Muhammet Gaffaroglu, Muradiye Karasu-Ayata only four genera), Barbus and Luciobarbus are tetraploid (2n = 4x) genera (Gaffaroğlu et al., 2013; Karasu-Ayata and Gaffaroğlu, 2019) where the genus Capoeta (2n = 6x) is hexaploid (Unal and Gaffaroğlu, 2016). Only the genus Cyprinion is diploid (2n = 2x) from this subfamily (Gaffaroğlu and Yüksel, 2004). Yang et al. (2022) reported that according to the mitochondrial and nuclear trees the polyploidy was allopolyploid in the subfamily Barbinae. Cytogenetic analyses may provide a useful tool for understanding the karyotype changes in the evolution of the species (Gaffaroğlu et al., 2020). Especially according to the high chromosome number (2n = 150) cytogenetic studies are very limited in the genus Capoeta from Tür- kiye (Table 2) and also from the other countries (Arai, 2011). Cytogenetic data are available for only seven Ana- tolian Capoeta species (Kılıç-Demirok and Ünlü, 2001; Kaya, 2003; Unal and Gaffaroğlu, 2016; Karasu-Ayata et al., 2017). The diploid chromosome number has been conserved in the species of the genus Capoeta in the pre- vious studies (Table 2). The chromosome number 2n = 6x = 150 in this study is consistent with previous reports (Table 2). However, karyotypes showed a pattern consid- ered basal for the genus, or with small variations due to the pericentric inversions and/or translocations in Ana- tolian Capoeta species (Table 2). The number of biarmed chromosomes is higher than uniarmed chromosomes in the Anatolian Capoeta species (Table 2) except C. trutta (Kılıç-Demirok and Ünlü, 2001). This feature is detected in this study as well. We conclude that this karyotype structure with mainly biarmed chromosomes is typical for the genus Capoeta. In detail, C. aydinensis, C. bergamae, C. erhani and C. pestai show very similar karyotype morphologies with some differences. The number of biarmed chromo- somes is as follows 96 in C. aydinensis, 92 in C. erhani, 86 in C. bergamae and, 84 in C. pestai. Otherwise, the Figure 2. Metaphase plates of Capoeta aydinensis by Giemsa stained (A), C-banded (C) and Ag-stained techniques (E) and arranged karyotypes (B, D, F). Arrows indicate the Ag-NORs bearing chro- mosomes. Scale bars = 5 μm. Figure 3. Metaphase plates of Capoeta bergamae by Giemsa stained (A), C-banded (C) and Ag-stained techniques (E) and arranged karyotypes (B, D, F). Arrows indicate the Ag-NORs bearing chro- mosomes. Scale bars = 5 μm. 31Comparative cytogenetics of four endemic Capoeta (Teleostei: Cyprinidae) species from Anatolia, Türkiye number of uniarmed chromosomes is as follows 54 in C. aydinensis, 58 in C. erhani, 64 in C. bergamae and, 66 in C. pestai. The FN ranges from 234 to 246 in this study. Pereira et al. (2012) suggested that distinct FNs with the same chromosome numbers in the species of the genus may be the result of pericentromeric inversions and/ or translocations involving centromeres. Karyotypes of four Capoeta species in this study showed minor varia- tions in their structures and depending on this having distinct FNs, apparently due to above mentioned chro- mosomal rearrangements. In addition, karyotypes with higher FNs are regarded to represent a derived condi- tion (Ganai et al., 2011). According to this hypothesis, C. pestai should be a more primitive scraper whereas C. aydinensis should be the most derived scraper among the four species. From the other countries C. capoeta (Safar, 2000), C. damascina (Gorshkova et al., 2002) and C. sevangi (Kry- sanov, 1999) were reported hexaploidy as detected in four studied species. C. sevangi differs from C. aydinensis, C. bergamae, C. erhani and C. pestai by having 110 uni- armed chromosomes (with FN = 190) (Krysanov, 1999). Moreover, C. antalyensis, C. baliki (Karasu-Ayata et al., 2017) and C. damascina (Unal and Gaffaroğlu, 2016) showed no sex chromosome differentiation like C. aydin- ensis, C. bergamae, C. erhani and C. pestai. Cytogenetic studies were mainly limited to detect chromosome number and morphology in the genus Capoeta (Table 2). Notably, chromosomal banding data (C-banding and Ag-NORs) revealed in only two Capo- eta species to date (Gaffaroğlu et al., 2012; Unal and Gaffaroğlu, 2016). C-bands were located mainly on the pericentromeres and terminal regions of some chromo- somes in four studied Capoeta species. C. aydinensis has the least C-bands compared to the other three species. C. bergamae and C. pestai have more C-banded chro- Figure 4. Metaphase plates of Capoeta erhani by Giemsa stained (A), C-banded (C) and Ag-stained techniques (E) and arranged karyotypes (B, D, F). Arrows indicate the Ag-NORs bearing chro- mosomes. Scale bars = 5 μm. Figure 5. Metaphase plates of Capoeta pestai by Giemsa stained (A), C-banded (C) and Ag-stained techniques (E) and arranged karyotypes (B, D, F). Arrows indicate the Ag-NORs bearing chro- mosomes. Scale bars = 5 μm. 32 Sevgi Unal-Karakus, Muhammet Gaffaroglu, Muradiye Karasu-Ayata mosomes than C. aydinensis and C. erhani. Similarly, C. damascina (Unal and Gaffaroğlu, 2016) and C. antaly- ensis (Gaffaroğlu et al., 2012) had centromeric C-bands as this study. Heterochromatic blocks that were report- ed in C. damascina (Unal and Gaffaroğlu, 2016) are not observed in this study. Due to the lack of the chromo- somal banding data for most of the species of the genus Capoeta from different countries, no comparison should be made. However, our results show the basal chromo- somal banding information for the genus Capoeta. Ag-NOR numbers have a stable distribution pattern among the four species newly analysed. It is assumed that two Ag-NORs in diploid barbins (Yüksel and Gaffaroğlu, 2006), four Ag-NORs in tetraploid barbins (Karasu-Ayata and Gaffaroğlu, 2019) and six Ag-NORs in hexaploid barbins (Unal and Gaffaroğlu, 2016) are common features. The Ag-NORs observed in the spe- cies studied here followed the similar feature observed in the other Capoeta species. C. aydinensis, C. bergamae, C. erhani and C. pestai are similar to C. damascina (Unal and Gaffaroğlu, 2016) and C. antalyensis (Gaffaroğlu et al., 2012) in terms of Ag-NOR numbers. Otherwise, C. bergamae, C. erhani and C. pestai are similar to C. damascina (Unal and Gaffaroğlu, 2016) in terms of loca- tions of Ag-NORs on the submetacentric chromosomes. C. antalyensis (Gaffaroğlu et al., 2012) has Ag-NORs on submeta-subtelocentric chromosomes like C. aydinensis. Moreover, Ag-NOR number polymorphism has not been reported in C. damascina and C. antalyensis (Gaffaroğlu et al., 2012; Unal and Gaffaroğlu, 2016) as observed in C. bergamae, C. erhani and C. pestai. Ribosomal DNA sites are considered as hot spots for chromosomal rearrange- ments such as duplications, fusions, fissions and inver- sions. Also, these sites should be correlated with trans- posable elements or repetitive DNAs (Araya-Jaime et al., 2022). In this context, Ag-NOR number polymorphisms that were detected in the three species in this study should be derived after the above mentioned chromo- somal rearrangements. Figure 6. Ag-NOR polymorphisms of Capoeta bergamae. Four Ag- NORs (A) and, five Ag-NORs (B). Arrows indicate the Ag-NORs bearing chromosomes. Scale bars = 5 μm. Figure 7. Ag-NOR polymorphisms of Capoeta erhani. One Ag- NOR (A), two Ag-NORs (B), three Ag-NORs (C) and, four Ag- NORs (D). Arrows indicate the Ag-NORs bearing chromosomes. Scale bars = 5 μm. Figure 8. Ag-NOR polymorphisms of Capoeta pestai. One Ag-NOR (A), two Ag-NORs (B), three Ag-NORs (C) and, four Ag-NORs (D). Arrows indicate the Ag-NORs bearing chromosomes. Scale bars = 5 μm. 33Comparative cytogenetics of four endemic Capoeta (Teleostei: Cyprinidae) species from Anatolia, Türkiye In conclusion, our results provide new data on the cytogenetic features of four Capoeta species. The endem- ic C. aydinensis, C. bergamae, C. erhani and C. pestai were analysed for the first time. Karyotype differences that were observed in this study highlight cytogenetics as an important tool for cytotaxonomy. The chromosom- al features with classical and molecular cytogenetic tech- niques of the other Capoeta species need to be studied to reveal detailed cytotaxonomy of the genus. REFERENCES Arai R. 2011. Fish karyotypes. A check list. – Springer, Japan. Araya-Jaime CA, Mazzoni Zerbinato de Andrade Silva D, Ribeiro da Silva LR, Neves do Nascimento C, Olivei- ra C, Foresti F. 2022. 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Species Locality Coordinate C. aydinensis (2 individuals) Suçıkan Spring, Dinar, Afyon (Büyük Menderes River) 38°04’N, 30°10’N C. bergamae (8 individuals) Dibekdere Stream, Ahmetli, Manisa (Gediz River) 38°33’N, 27°57’E C. erhani (11 individuals) Çakıt Stream, Şekerpınarı, Pozantı, Adana (Seyhan River) 37°27’N, 34°52’E C. pestai (2 individuals) Kayabaşı Stream, Beyşehir, Konya (South of Beyşehir Lake) 37°29’N, 31°30’E Table 2. Karyological data for the genus Capoeta from Türkiye. 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Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Volume 76, Issue 1 - 2023 Firenze University Press Cytogenetic studies in the Andropogon gayanus-Andropogon tectorum Complex in Southwestern Nigeria Funmilola Mabel Ojo1,*, Chinyere Constance Nwokeocha2, Julius Olaoye Faluyi3 Incidence and frequency of desynapsis in Eremurus persicus (Jaub. & Spach) Boiss.(Asphodelaceae) – A native and important medicinal plant species of Western Himalaya Shivali Verma¹, Irfan I. Sofi², Aijaz H. Ganie², Manzoor A. Shah², Namrata Sharma¹ Comparative cytogenetics of four endemic Capoeta (Teleostei: Cyprinidae) species from Anatolia, Türkiye Sevgi Unal-Karakus1,*, Muhammet Gaffaroglu2, Muradiye Karasu-Ayata3 A karyomorphological comparison of seven species of Achillea L. from Kurdistan of Iran Fatemeh Nezhadi1, Farzad Fayaz2,*, Ezzat Karami2, Hooshmand Safari3, Abdol Rahman Rahimi2 Genotoxicity of a synthetic plant growth regulator, Forchlorfenuron (CPPU), on human lymphocytes using chromosome aberration assay Ayşe Yavuz Kocaman1,*, Berna Yakar2 Phenthoate toxicity evaluation in root meristem of Pisum sativum L. Sazada Siddiqui Karyomorphology of two subspecies of Anthemis maritima (Asteraceae) from Algeria Meryem Nassar1,4,*, Nora Sakhraoui2,4, Gianniantonio Domina3 Cytogenetic effects of Tribulus terrestris L. on meristematic cells of Allium cepa L. and Vicia faba L. Ali Bouzekri1,2,*, Meryem Nassar1,2, Souheila Slimani1,2, Zohra Chekroud1,2 New chromosomal data, karyotype asymmetry and polyploid variations of some Gundelia (Asteraceae) species from Turkey Esra Martin1, Metin Armağan2, Halil Erhan Eroğlu3*, Aslı Doğru-Koca4, Osman Tugay5, Golshan Zare6, Osman Kola7, Mahmut Miski8, Nur Tan9, Ernst Vitek10 Allelopathic and toxicological effects of Origanum vulgare L. essential oil Lejla Husić, Adisa Parić, Aner Mesic* Cytogenetic analysis in Tetragonopterus franciscoensis (Characiformes): another piece to the karyoevolutionary puzzle of tetra fishes Mauricio Barros Fernandes, Jamille de Araújo Bitencourt, Joandson Calixto dos Santos, José Henrique Galdino*, Paulo Roberto Antunes de Mello Affonso