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Chenopodiaceae) from Iran © 2009 The Japan Mendel Society Cytologia 74(1): 79–87, 2009 A Cytological Study of Fourteen Halophytic Species of Tribes Caroxyloneae and Salsoleae (Chenopodiaceae) from Iran Alireza Doulatyari1, Seyed Mahmood Ghaffari2 and Hossein Akhani1,* 1 Department of Plant Sciences, School of Biology, University of Tehran, P. O.Box 14155-6455, Tehran, Iran 2 Institute of Biochemistry and Biophysics, University of Tehran, P. O.Box 13145-1384, Tehran, Iran Received February 11, 2009; accepted February 28, 2009 Summary Original chromosome numbers and meiotic behavior are presented for 25 populations belonging to 14 halophytic species of the genera Climacoptera Botsch., Halimocnemis C. A. Mey. s.l., Petrosimonia Bunge, Halocharis Moq. and Kaviria Akhani & E. H. Roalson (Caroxyloneae) and Salsola florida (M. Bieb.) Poir (Salsoleae). The basic chromosome number of xϭ8 was found in Petrosimonia and xϭ9 in other genera. The polyploidy was detected in Kaviria tomentosa (Moq.) Akhani (nϭ18), Climacoptera crassa (M. Bieb.) Botsch. (nϭ27) and C. turcomanica (Litw.) Botsch. (nϭ18, 27). Chromosome numbers of the following species are reported for the first time: Halimocnemis azarbaijanensis Assadi (nϭ9), H. gamocarpa Moq. (nϭ9), H. pilosa (Pall.) Akhani (nϭ9), H. mamamensis (Bunge) Assadi, H. rarifolia (C. Koch) Akhani (nϭ9), H. mollissima Bunge (nϭ9), H. pilifera Moq. (nϭ9), Halocharis sulphurea (Moq.) Moq. (nϭ9), Petrosimonia glauca (Pall.) Bunge (nϭ8), and Salsola florida (M. Bieb.) Poir (nϭ9). New ploidy levels are also reported for the first time in Climacoptera crassa and C. turcomanica. Meiotic behavior was generally regular in all cases but some irregularities such as laggard chromosomes in Metaphase II were observed in Halimocnemis pilifera. Our results together with available literature data suggest that xϭ9 is the basic chromosome number for the Chenopodiaceae family and the other numbers (e.g. xϭ8 as in Petrosimonia glauca) are derived from dysploidy series. The lowest frequency of chiasma is reported for Halimocnemis mamamensis (1.1 per each bivalent) and the highest for Kaviria tomentosa (1.6 per each bivalent). Key words Chenopodiaceae, Iran, Halophytes, Meiosis, Climacoptera, Halimocnemis, Petrosimo- nia, Salsola, Kaviria Iran is a typical country of large deserts and salty ecosystems. Many desert, salt and drought resistant plant groups have been diversified in this country with diverse climatic condition and interesting geological history. The Chenopodiaceae family with 40 genera and 190 known species in Iran plays a major role in the halophytic and xerophytic vegetation of Iran (Hedge et al. 1997, Ghaffari et al. 2006, Akhani 2008). More than one third of the world known genera of Chenopodiaceae have been known from Iran (Kühn et al. 1993, Hedge et al. 1997). The highest diversity was found in subfamily Salsoloideae s.l. (Hedge et al. 1997, Akhani et al. 2007 and references therein). According to latest phylogenetic analysis the Salsoloideae s.l. includes three tribes Camphorosmeae, Caroxyloneae and Salsoleae s. str. (Akhani et al. 2007). The two tribes Caroxyloneae and Salsoleae are characterized by spiral embryo and Camphoromeae have circular embryo. Based on phylogenetic studies Caroxyloneae include three clades: Caroxylon, Kaviria and Climacoptera. The two latter clades composed of 8 genera which have mostly Central and SW * Corresponding author, e-mail: [email protected] 80 A. Doulatyari, S. M. Ghaffari and H. Akhani Cytologia 74(1) Asian distribution. The Climacoptera clade includes only Irano-Turanian annual species belonging to Petrosimonia, Ofaiston, Pyankovia, Halimocnemis s.l. and Climacoptera which are diversified in the Irano-Turanian deserts. The Kaviria clade includes the newly segregated genus Kaviria from Salsola s.l., Nanophyton and Halocharis. The cytological studies in Chenopodiaceae have a long-standing history. The earliest reports back to 1910 and 1911 when the chromosome number of spinach has been reported by Strasburger and Stomps and followed by several reports on other species by Winge and Dahlgren in 1916 and 1917 (Federov 1974). Based on available literature there are chromosome reports for ca. 16 genera of Salsoleae s.l. and 6 genera of Camphorosmeae (Fedorov 1974, Moore 1982, Goldblatt and Johnson 1979). Several authors reported the karyotype variation in these groups (Semiotroczeva 1974, 1983, Podlech and Bader 1974, Bakhshi Khaniki and Maroof 2006, Zakharyeva (1985, in Goldblatt and Johnson 1979). The chromosome number and karyotype variation in Iranian Chenopodiaceae have been studied in several publications (Uotila 1973, Hekmat-Shoar 1978, Ghaffari 1986, Ebrahimzadeh et al. 1994, Mirzaie Nodoushan and Asadi Corom 2002, Akhani et al. 2005, Ghaffari et al. 2006, Bakhshi Khaniki and Maroof 2006). In this paper we follow our studies on the meiotic behavior and chromosome numbers of one species of Petrosimonia, seven species of Halimocnemis, two species of Climacoptera and one species of Halocharis and one species of Kaviria (all belonging to Caorxyloneae) and one species of Salsola (Salsoleae). Materials and methods Chromosome counts were made from pollen mother cells squashes. The young floral buds were prefixed in the field either in a solution of 6 parts alcohol: 3 parts chloroform: 2 parts propi- onic acid or Carnoy’s I solution (3 parts alcohol: 1 part acetic acid), and after one or two days trans- ferred into 70% alcohol and refrigerated till laboratory work. Anthers were squashed in aceto- carmine 2% or orcein-acetic (La Cour). Microsporocytes in various stages of meiosis studied by Olympus microscope (model BH-2). Suitable slides were made permanent by the Venetian turpen- tine methods (Wilson 1945). Voucher specimens from all studied populations are deposited in the Botanical Biodiversity Research Laboratory, School of Biology, University of Tehran. The nomen- clature and generic circumscription follow the recent morpho-molecular classification (Akhani et al. 2007). Results A list of studied material, number of chromosomes and their vouchers are given in Table 1. The picture of chromosomes compliments of selected populations are given in Figs. 1, 2. The re- sults of genera and individual species are given here. Climacoptera crassa (M. Bieb.) Botsch. This is a hexaploid species with nϭ27 (Fig. 1A). We found 27 bivalents in metaphase I (Fig. 1B) suggesting its probable allohexaploidy. The previous report for this species was 2nϭ18 (Semiotroczeva 1983). C. crassa distinguishes from C. turcomanica in the distribution area and shorter styles and stigma and glabrous teplas (Hedge et al. 1997, Pratov 1986). Climacoptera turcomanica (Litv.) Botsch. Three populations of this species have been studied. We found diploid (2nϭ2xϭ18) (Fig. 1C), tetraploid (2nϭ4xϭ36) and hexaploid (2nϭ2xϭ54) levels in studied populations (Table 1). In hexaploid population only 27 bivalents in metaphase I was observed (Fig. 1C). But in tetraploid 2009 Table 1. List of studied species/accessions, herbarium vouchers and chromosome numbers of 14 species of Caroxyloneae and Salsoleae from Iran. D & DϭDehghani & Doulatyari. The synonyms are given for species which status are recently changed (Akhani et al. 2007). Miotic Species Location: Chromosome no. (n) Climacoptera crassa (M. Bieb.) Botsch. Azerbaijan: 5 km from Poldashat towards Pazi (5 km NW poldasht), 22.7.2005. D & D 335 27 Climacoptera turcomanica (Litv.) Botsch. Tehran: 35km before Eshtehard, 7 km W Mardabad (Rudeshur), 22.6.2005. D & D 367 27 F Climacoptera turcomanica (Litv.) Botsch. Arak: 26 km NE Arak (towards Kavire Meyghan), 5.8.2005. D & D 360 18 SpeciesofTribes ourteen Halophytic Halimocnemis azarbaijanensis Assadi Ardebil: 150 km before Parsabad from Ardebi (33 km before Alahyar lo village), 22.7.2005. D & D 320 9 Halimocnemis gamocarpa Moq. Tehran: 80km before Qom from Tehran, 4.8.2005. D & D 343 9 (ϭGamanthus gamocarpus (Moq.) Bunge] Halimocnemis mamamensis (Bunge) Assadi Zanjan: 75 km before Mianeh from Zanjan, 21.7.2004. D & D 304 9 (ϭHalanthium mamamense Bunge) Ardebil: 55 km before Parsabad from Ardebil , 22.7.2005. D & D 321 9 Azerbaijan: 6 km after Khalkhal-Mianeh junction towards Khalkhal, 21.7.2005. D & D 307 9 Halimocnemis mollissima Bunge Tehran: 35km before Eshtehard, 7 km W Mardabad (Rudeshur), 22.6.2005. D & D 365 9 Halimocnemis pilifera Moq. Tehran: 35km before Eshtehard, 7 km W Mardabad (RudeShur), 22.6.2005. D & D 368 9 Tehran: 80 km before Qom from Tehran, 4.8.2005. D & D 342 9 Arak: 7 km before Arak from Delijan, 5.8.2005. D & D 356 9 Halimocnemis pilosa (Pall.) Akhani Ardebil: 40 km before Eskanlu from Aslandoz, 23.7.2005. D & D 323 9 Caroxyloneae (ϭGamanthus pilosus (Pall.) Bunge] Halimocnemis spec. Azerbaijan: Khalkhal-Mianeh-Maman junction 21.7.2004. D & D 308 9 Halimocnemis rarifolia (K. Koch) Akhani Azerbaijan: 34 km before Khalkhal-Mianeh junction from Zanjan, 21.7.2005. D & D 305 9 (ϭHalanthium rarifolium K. Koch) and Ardebil: opposite of Eskanlo gas station, 23.7.2005. D & D 324 9 Qom: 84 km before Tehran (Houze Soltan salt lake), 22.6.2005. D & D 370 9 Salsoleae Esfahan: 18 km after Delijan towards Esfahan, 4.8.2005. D & D 347 9 Azerbaijan: 26 km before Poldashat from Jolfa, 24.7.2005. D & D 333 9 Ardebil: 35 km before Jolfa from Parsabad, 23.7.2005. 326 9 Semnan: 8 km before Semnan from Sorkhe, 20.8.2005. 340 9 Halocharis sulphura Moq. Semnan: 2 km before sangsar from Semnan, 20.8.2005. D & D 339 9 Petrosimonia glauca Bunge Tehran: 35km before Eshtehard, 7 km W Mardabad (Rudeshur), 20.6.2006. D & D 281 8 Kaviria tomentosa (Moq.) Akhani Tehran: 35km before Eshtehard, 7 km W Mardabad (Rudeshur), 20.6.2006. D & D 283 18 (ϭSalsola tomentosa Moq.) Salsola florida (M. Bieb.) Poir (ϭSeidlitzia Tehran: 35 km before Eshtehard, 7 km W Mardabad (Rudeshur), 20.6.2006. D & D 282 9 florida (M. Bieb.) Bunge ex Boiss. 81 82 A. Doulatyari, S. M. Ghaffari and H. Akhani Cytologia 74(1) 2009 Fourteen Halophytic Species of Tribes Caroxyloneae and Salsoleae 83 population 2 tetravalents and 14 bivalents were observed (Fig. 1D). Previous somatic chromosome count for this species from Central Asia is 2nϭ2xϭ18 (Federov, 1974).
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