Caryologia. International Journal of Cytology, Cytosystematics and Cytogenetics 75(4): 67-76, 2022 Firenze University Press www.fupress.com/caryologia ISSN 0008-7114 (print) | ISSN 2165-5391 (online) | DOI: 10.36253/caryologia-1674 Caryologia International Journal of Cytology, Cytosystematics and Cytogenetics Citation: Saeedeh Sadat Mirzadeh Vaghefi, Adel Jalili (2022). Chromosome counts of some species of wetland plants from Northwest Iran. Caryolo- gia 75(4): 67-76. doi: 10.36253/caryolo- gia-1674 Received: May 29, 2022 Accepted: December 06, 2022 Published: April 28, 2023 Copyright: © 2022 Saeedeh Sadat Mir- zadeh Vaghefi, Adel Jalili. This is an open access, peer-reviewed article published by Firenze University Press (http://www.fupress.com/caryologia) 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. Chromosome counts of some species of wetland plants from Northwest Iran Saeedeh Sadat Mirzadeh Vaghefi*, Adel Jalili Research Institute of Forests and Rangelands, Agricultural Research, Education and Extension Organization (AREEO), P. O. Box 13185-116, Tehran, Iran *Corresponding author. E-mail: myalyssum94@gmail.com Abstract. Wetlands scatter as microclimates in mountain areas of Iran. In this investi- gation, the chromosome number of species and populations from Azerbaijan provinces were studied. After seed germination and root fixation, meristem cells were stained and photographs were taken for cells in the metaphase stage by light microscope. Data were analyzed by Micromeasure and excel softwares. As many as 28 populations of 24 species were studied. The chromosome numbers of two species (viz. Ranunculus kotschyi) were reported for the first time. 11 populations in 9 species were reported for the first time from Iranian populations (viz. Alisma plantago-aquatica, Prunella vulgar- is, Scrophularia umbrosa). The range of haploid chromosome numbers is between n=6 and n=21. Ideograms were depicted for each species. Keywords: chromosome number, wetland, ideogram, Azerbaijan. INTRODUCTION Wetlands are considerate as microclimates in alpine areas with special species in Iran. In the steppic mountain areas of Iran are a wide range of wetland vegetation. The wetland habitats are sharply embedded within veg- etation of the IranoTuranian steppes that are more characteristic of this region and are of interest both in themselves and for wider comparison with EuroSiberian wetlands (Naqinezhad, 2012). Most of the wetlands in the northwest part of Iran develop when snow- melt and rain fill the pockmarks left on the landscape by glaciers. This kind of wetlands was categorized in Prairie potholes class (EPA 2001). In angiosperms, the haploid chromosome number varies between n = 2 and n = 132, but the majority of them show a range between n = 7 and n = 12(Sharma, 2009). Variation or constancy in the chromosome number within taxa of different categories has been proven to be important characters for taxonomic groupings (Sharma, 2009). We studied on chromosome number of 28 accessions of 24 species belonging to 21 genera of 12 families wetland plants of Azerbaijan Moun- tains of Iran. The aim of this study was to investigate on chromosome num- ber of Wetland plants of Azerbaijan provinces. 68 Saeedeh Sadat Mirzadeh Vaghefi, Adel Jalili MATERIALS AND METHODS Plant materials Specimens and seeds were collected from natural habitats of Azerbaijan provinces (West, East Azerbai- jan and Ardebil) in 2013 and 2014. Vouchers are depos- ited in TARI (Table 1). The plant samples were identified by Flora of Iran (Assadi et al., 2018) and Flora Iranica (Rechinger, 2005). Chromosome counts The seeds were used for chromosome counts. Seeds were germinated at 20-25°C on moist filter paper in Petri dishes. Root tips of seeds had grown 1–1.5 cm in length were stopped from the germinating by pretreat- ing with alpha-bromonaphthalene for two hours. Then fixed with acetic alcohol (1:3) for 4 h and stored in 70% alcohol at 2–4°C. Then the root tips were rinsed in water and hydrolyzed with 1 N HCl for 10–18 min at 60°C and rinsed in running water for a minimum of 3–5 min. Staining of root tips was carried out for 1–2 h and root tips were put in hematoxylin for 5–10 min to improve staining. Finally, squash preparations were made. Chro- mosome nomenclature follows Levan et al. (1964) and Stebbins (1971). The photos were taken by light micro- scope (BH2 Olympus × 1000). We measured the chromosomes using MicroMeas- ure (version 3.3 (Reeves and Tear, 2000). RESULTS Our goal in this project was to create a chromosome index for the wetland plants of Azerbaijan. It is so dif- ficult to discussion about several species from several families that collected for common goal. Table 1. List of species examined, Family, Locality, Altitude, Voucher number, and Habitat. Voucher no. Alt. Locality Family Species 102715 2392 East Azerbaijan, Tabriz, Arshad chamani Alismataceae Alisma plantago- aquatica L. 102769 2281 West Azerbaijan, Salmas, Jam Valley Poaceae Alopecurus arundinaceus Poir. 102752 1900 Aligo Village Brasicaceae Barbarea plantaginea DC. 8228 ---- East Azerbaijan, Bostanabad Poaceae Catabrosa aquatica (L.) P.Beauv. 102750 2266 Ardebil province, Salmas, Jam Valley Asteraceae Erigeron acris L. subsp. pycnotrichus (Vierh.) Rech. f. 101699 2266 West Azerbaijan, Dalamper Village Geraniaceae Geranium sylvaticum L. 102751 2266 West Azerbaijan, Dalamper Village Hypericaceae Hypericum perforatum L. 102807 2266 West Azerbaijan, Dalamper Village Asteraceae Inula aucheriana DC. 102762 1688 East Azerbaijan, Hashtroud, Zalbil Asteraceae Mulgedium tataricum (L.) DC. 8239 1819 East Azerbaijan, Hashtroud, Baboneh village Poaceae Phragmites australis Trin. ex Steud. 101691 2281 West Azerbaijan, Salmas, Jam Valley Plantaginaceae Plantago atrata Hoppe. 101601 2266 West Azerbaijan, Dalamper Village Lamiaceae Prunella vulgaris L. (Dalamper) 8242 ---- East Azerbaijan, Bostanabad Lamiaceae Prunella vulgaris L. (Bostanabad) 101668 2296 Ardebil, Agh Ghabali Rannunculaceae Ranunculus aquatilis var. diffusus With. 101670 East Azerbaijan, Varzaghan Rannunculaceae Ranunculus dolusus Fisch . & C. A. Mey. 101674 1648 Ardebil province, Ardebil to Bolaghvar Rannunculaceae Ranunculus kotschyi Boiss. 102738 1900 East Azerbaijan, Bostanabad Scrophulariaceae Scrophularia umbrosa Dumort. 102805 1688 East Azerbaijan, Hashtroud, Zalbil Asteraceae Senecio pseudoorientalis Schischk. (Hashtroud) 102761 1648 Ardebil province, Ardebil to Bolaghvar Asteraceae Senecio pseudoorientalis Schischk. )Bolaghvar) 101688 2266 West Azerbaijan, Dalamper Village Fabaceae Trifolium pratense L. 102724 2281 West Azerbaijan, Salmas, Jam Valley Juncaginaceae Triglochin maritima L. (Jam valley( 102728 1650 Ardebil province, Ardebil to Bolaghvar Juncaginaceae Triglochin maritima L. (Bolaghvar) 102753 1900 East Azerbaijan, Bostanabad Asteraceae Tripleurospermum disciforme Sch.Bip. (Bostanabad) 102784 1900 East Azerbaijan, Tabriz, Arshad chamani Asteraceae Tripleurospermum disciforme Sch.Bip. (Jam valley) 102713 2392 East Azerbaijan, Tabriz, Arshad chamani Scrophulariaceae Veronica orientalis Miller (Arshad Chamani) 102716 2281 East Azerbaijan, Bostanabad Scrophulariaceae Veronica orientalis Miller (Bostanabad) 102729 2281 East Azerbaijan, Bostanabad Scrophulariaceae Veronica filiformis Sm. 101677 1650 Ardebil province, Ardebil to Bolaghvar Fabaceae Vicia variabilis Freyn & Sint. ex Freyn 69Chromosome counts of some species of wetland plants from Northwest Iran Of 24 examined species (Table 2), 5 species have two populations (Prunella vulgaris, Tripleurospermum dis- ciforme, Pedicularis sibthorpii, Senecio pseudoorientalis and Veronica orientalis). Another species has just one population. Asteraceae (5 species), Scrophulariaceae (3 species), Poaceae and Ranunculaceae (3 species) families have the most species respectively. The chromosomes counts of the taxa previously reported are in Table 2. Most of them confirm our results. Chromosome numbers of Ranunculus kotschyi (Fig. 2P) and Erigeron acris subsp. pycnotrichus (Fig. 1F) were reported for the first time. Alisma plantago-aquatica (Fig. 1A), Prunella vulgaris (Fig. 2M & N), Scrophular- ia umbrosa (Fig. 2Q), Mulgedium tataricum (Fig. 1J), Ranunculus aquatilis var. diffusus (Fig.1D), Erigeron acris subsp. pycnotrichus (Fig.1F), Geranium sylvaticum (Fig.1G), Triglochin maritima (Fig.3U&V), Veronica fili- formis (Fig. 3Y) were reported for the first time of Irani- an populations. Sporophytic count of Barbarea plantag- inea (Fig. 1C) was reported for the first time from Iran. The details of each taxon reported as in Table 3. Table 2. Chromosome counts of studied species previously described in the literature. References (viz.) 2n References N Species (Dobes et al., 1997)(Wulff, 1939)(Love and Love., 1942) 10, 12, 14 (Kaur et al., 2011) 14 Alisma plantago- aquatica (Kuzmanov, 1993)(Amosova et al., 2019)(Sheidai et al., 2009) 28, 42 (Koull and Gohil 1991) 14 Alopecurus arundinaceus (Astanova, 1999)(Ørgaard and Linde-laursen, 2014) 16 (Aryavand, 1977) (Ghaffari, 2007) 8 Barbarea plantaginea (Sawicka, 1991)(Lövkvist and Hultgård., 1999)(Sheidai et al., 2009) 20 Catabrosa aquatica. (Kaur et al., 2011)(Bala and Gupta, 2013)(Paule et al., 2017) 18 Erigeron acris (other subspecies) (Petrova and Stanimirova, 2001) (Lövkvist and Hultgård 1999)(Dmitrieva, 1986) 28, 24 (Clifford Odets, 1954) 12 Geranium sylvaticum (Ciccarelli et. al. 2001)(Lövkvist and Hultgård, 1999)(Krasnikov and Schaulo 1990)(Kalinka et al., 2014((Baltisberger and Widmer, 2009((Brutovska et al., 2000) 32 (Ghaffari, 2006) 16 Hypericum perforatum (Chehregani and Hajisadeghian, 2009) 9 Inula aucheriana (Probatova, 2004) 36 Mulgedium tataricum (Lövkvist and Hultgård 1999)(Panahi., 1979)(Gervais et al., 1993) 42,48, 72,96 Phragmites australis (Petrova and Stanimirova, 2001)(Lessani and Chariat- panahi., 1979) 12, 24 Plantago atrata (Lövkvist and Hultgård 1999((Krasnikov and Schaulo., 1990) 28 Prunella vulgaris (Dahlgren and Cronberg, 1996) 32,48 Ranunculus aquatilis var. diffusus (Baltisberger, 1991) (Agapova, 1981)(Assadi, 1989) (Ghasemi et al., 2015) 28, 32 Ranunculus dolusus Not reported Ranunculus kotschyi (Javurkova, 1979)(Grau, 1979)(Vitek et al. 1992) 26,52 (Grau, 1979) 13, 26 Scrophularia umbrosa (Ghaffari, 1999) 20 Senecio pseudoorientalis (Probatova, 2000) (Krasnikov and Schaulo, 1990)(Zhang et al., 1993)(Sheidai et al., 1998) 14, 16 Trifolium pratense (Lövkvist and Hultgård. 1999)(Krasnikov, 1991)) Rotreklova, 2004( (Uchiyama, 1989)(Iwatsubo et al., 1998) 48,120 Triglochin maritima (Ghaffari, 1999) (Hayirlioglu-Ayaz, 2011) 18 (Ghaffari, 1999) (Razaq, et al., 1994) 9 Tripleurospermum disciforme (Ghaffari, 1986) 32 (Ghaffari, 1987) 32 Veronica orientalis (Pogan et al., 1990)(Dzhus and Dmitrieva, 2001)(Albach et al., 2009) 14 (Dobes and Vitek, 2000) 7 Veronica filiformis (Hesamzadeh Hejazi and Rasuli, 2006)(Hesamzadeh Hajazi and Ziaei Nasab, 2009) 14 Vicia variabilis 70 Saeedeh Sadat Mirzadeh Vaghefi, Adel Jalili Figure 1. Somatic metaphase chromosome, Ideogram of each population is on the right side of the image: A: Alisma plantago- aquatica (2n = 14); B: Alopecurus arundinaceus (2n = 28); C: Barbarea plantaginea (2n = 16); D: Ranunculus aquatilis var. diffusus (2n = 32); E: Catabrosa aquatica (2n = 20 ); F: Erigeron acris subsp. pycnotrichus (2n = 18); G: Geranium sylvaticum (2n = 28); H: Hypericum perforatum (2n = 32); I: Inula aucheriana (2n = 18); J: Mulgedium tataricum (2n = 36). Scale bar: 1 µm. 71Chromosome counts of some species of wetland plants from Northwest Iran Figure 2. Somatic metaphase chromosome, Ideogram of each population is on the right side of the image: K: Phragmites australis (2n = 42); L: Plantago atrata (2n = 12); M: Prunella vulgaris (Dalamper) (2n = 28); N: Prunella vulgaris (Bostanabad) (2n = 28); O: Ranunculus dolusus (2n = 32); P: Ranunculus kotschyi (2n = 16); Q: Scrophularia umbrosa (2n = 52); R: Senecio pseudoorientalis (Hashtroud) (2n = 40); S: Sene- cio pseudoorientalis )Bolaghvar) (2n = 40); T: Trifolium pratense (2n = 16). Scale bar: 1 µm. 72 Saeedeh Sadat Mirzadeh Vaghefi, Adel Jalili Somatic metaphase and Ideograms of taxa were shown (Figs 1-3). DISCUSSION Only some interesting results are discussed here. The chromosome number of the most of the species support by former studies (Table 2). The variation ranges of haploid chromosome numbers in studied species are between n=6 and n=21. Populations in Triglochin maritima (Fig. 3U & V) comprise different ploidy levels (2n = 24, 12). Some spe- cies like Prunella vulgaris (Fig. 2M & N), Veronica ori- Figure 3. Somatic metaphase chromosome, Ideogram of each population is on the right side of the image: U: Triglochin maritima (Jam val- ley) (2n = 24); V: Triglochin maritima (Bolaghvar) (2n = 12); W: Tripleurospermum disciforme (Bostanabad). (2n = 18); X: Tripleurospermum disciforme (Jam valley( (2n = 18); Y: Veronica filiformis (2n = 14); Y1: Veronica orientalis (Arshad Chamani) (2n = 32); Y2: Veronica orienta- lis (Bostanabad) (2n = 32); Z: Vicia variabilis (2n = 14). Scale bar: 1 µm. 73Chromosome counts of some species of wetland plants from Northwest Iran entalis (Fig. 3Y1 & Y2), Senecio pseudoorientalis (Fig. 2R & S), Tripleurospermum disciforme (Fig. 3W & X) have invariant ploidy level in this study. Alisma plantago-aquatica n = 7 (Fig. 1A), is not par- ticularly variable; nevertheless three different chromo- some numbers have been found for it viz. n = 5 (Wulff, 1939), 6 ( Liehr, 1916; Love and Love, 1942) and 7 ( Love and Love, 1942). Combining the available chromosome studies, the Erigeron has relatively consistent chromosome diversifi- cation with a basic number of 9, and most species con- tain diploid individuals, which suggested Erigeron at the initial phase of polyploid diversification (Baldwin and Speese, 1955). Phragmatis australis is represented by many poly- ploids, cuploids from 3x = 36 to 8x = 96 (without 5 x) and aneuploidies (2n = 42, 44, 46, 49, 50, 51, 52, 54). Tetraploidy and octoploidy are in majority (Gorenflot, 1979). Chromosome numbers of Phragmatis australis showed a high degree of aneuploidy and varied between 2n=42 and 2n=59 (Gervais et al., 1993). The degree of polyploidy is not in direct relation with the individu- al’s habitus. Meiosis study shows that this complex has already passed the maturity, the diploid forms having disappeared (Gorenflot, 1979). Base on this study our case is aneuploidy. Hypericum perforatum has the smallest chromo- somes than other species of this genus. In other studies it had 2n=32 with median centromeric chromosomes as our result (Brutovska et al., 2000). Most of the subgenera of Veronica exhibit only one single basic number, i.e., x = 6, 7, 8, 9, 12, 17, or 20/21. Table 3. Somatic chromosome number (2n), Ploidy level, karyotype formula, ranges of chromosome length and degree of Asymmetry according to STEBBINS (1971) for the studied taxa, Figure numbers are related images in Fig 1 Figure number (Figs.1-3) Stebbins Chromosome range length (µm) Karyotype formula Ploidy level x 2n Taxa A B1 5.3-7.83 2sm+5m 2 7 14 Alisma plantago- aquatica B A2 4.1-5.95 8sm+6m 4 7 28 Alopecurus arundinaceus C A1 1.06-1.29 2sm+6m 2 8 16 Barbarea plantaginea E A1 2.25-3.09 10m 4 5 20 Catabrosa aquatica F A1 3.44-5.10 3sm+6m 2 9 18 Erigeron acris subsp. pycnotrichus G A1 1.51-2.19 2sm+12m 2 14 28 Geranium sylvaticum H A1 0.92-1.37 16m 4 8 32 Hypericum perforatum I A1 2.91-3.8 9m 2 9 18 Inula aucheriana J A2 2.14-3.30 5sm+13m 4 9 36 Mulgedium tataricum K A2 2.8-4.89 10sm+11m 4 An. 42 Phragmites australis L A1 6m 2 6 12 Plantago atrata M B1 0.95-2.28 14m 2 14 28 Prunella vulgaris (Dalamper) N A1 0.7-1.11 14m 2 14 28 Prunella vulgaris (Bostanabad) D A2 2.01-3.31 9sm+7m 4 8 32 Ranunculus aquatilis var. diffusus O A2 1.89-2.87a 4sm+12m 4 8 32 Ranunculus dolusus P A3 3.67-5.38 5sm+3st 2 8 16 Ranunculus kotschyi Q A1 1.51-3.15 4sm+22m 4 13 52 Scrophularia umbrosa R A1 2.41-4.31 4sm+16m 4 10 40 Senecio pseudoorientalis (Hashtroud) S A2 2.6-3.66 2sm+18m 4 10 40 Senecio pseudoorientalis )Bolaghvar) T A1 3.41-4.80 2sm+6m 2 8 16 Trifolium pratense U A1 1.88-2.46 3sm+9m 4 6 24 Triglochin maritima (Jam valley) V A3 4.23-5.16 3sm+3st 2 6 12 Triglochin maritima (Bolaghvar) W A1 2.14-2.7 9m 2 8 18 Tripleurospermum disciforme (Bostanabad). X B1 1.81-2.86 9m 2 9 18 Tripleurospermum disciforme (Jam valley( Y1 A1 0.73-1.31 16m 4 8 32 Veronica orientalis (Arshad Chamani) Y2 A1 1.38-2.19 16m 4 8 32 Veronica orientalis (Bostanabad) Y A1 0.28-0.48 1sm+6m 2 7 14 Veronica filiformis Z A3 1.66-2.37 3sm+4m 2 7 14 Vicia variabilis An.= Aneuploidy. 74 Saeedeh Sadat Mirzadeh Vaghefi, Adel Jalili In this genus, the putative ancestral base number of 9 has been reduced several times to 8 and 7, respectively (aneuploidy/dysploidy), often associated with transi- tion to annual life history. In contrast, no unambiguous increase of chromosome base number has been inferred (Albach et al., 2008). A base chromosome number reduction to x = 8 (aneuploidy), seems to have occurred in Veronica orientalis (tetraploid, 2n=4x=32). Previous results confirm our result (Ghaffari, 1986). Basic chromosome number(x) The frequency of 2x (53.57%) and 4x (46.43%) in this study are almost identical. 6x and more another nx not found. 2x and 4x are most common in flowering plants (Bala and Gupta, 2013). The present result is in agree- ment with the reports of earlier investigators. Polyploidy and habit The overall chromosome numbers during pre- sent study lie on two different levels of ploidy i.e. 2x, 4x. Among these, the diploids are the most common in terms of frequency (53.57 %), followed by tetraploids (46.43 %) (Table 3). As reported at least 47% of species have undergone a recent polyploidy event (Wood et al., 2009)polyploidy has been recognized as an important phenomenon in vascular plants, and several lines of evidence indicate that most, if not all, plant species ultimately have a poly- ploid ancestry. However, previous estimates of the fre- quency of polyploid speciation suggest that the forma- tion and establishment of neopolyploid species is rare. By combining information from the botanical com- munity’s vast cytogenetic and phylogenetic databases, we establish that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase. These frequency estimates are higher by a factor of four than earlier estimates and lead to a standing incidence of polyploid species within genera of 35% (n = 1,506. As Ramsey and Ramsey,(2014) indicated that; Polyploids are able to colonize larger geographic ranges and/or occur in more habitats than related diploids, similar picture occurred in our study: those species with higher ploidy level have wide spread geographical distribution than their related diploid populations or species. For example in Iran, some species such as; Alopecurus arundinaceus, Phragmites australis, Catabrosa aquatica, Hypericum perforatum, Veronica orientalis with higher polyploidy level showed same attitudes (Table 3) (Alinejad et al., 2017; Ghahremaninejad et al.; 2012, Khanhasani et al., 2021; Safikhani et al., 2018; Aref Tabad et al., 2016; Jalili et al., 2014) Karyotype Metacentric and submetacentric are commonly observed. Karyotype of ten species (35.7%) consists of chromosomes with the centromere in median regions (m). 14 species (50%) have metacentric and submeta- centric chromosomes in their formula. Only 2 species have  karyotypes with submetacentric and  subtelocentric chromosomes. Karyotypes of most species (78.58%) were classified in the 1A and 2A Stebbins classes (Table 3). 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