© 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 x 8 was found in Petrosimonia and x 9 in other genera. The polyploidy was detected in Kaviria tomentosa (Moq.) Akhani (n 18), Climacoptera crassa (M. Bieb.) Botsch. (n 27) and C. turcomanica (Litw.) Botsch. (n 18, 27). Chromosome numbers of the following species are reported for the first time: Halimocnemis azarbaijanensis Assadi (n 9), H. gamocarpa Moq. (n 9), H. pilosa (Pall.) Akhani (n 9), H. mamamensis (Bunge) Assadi, H. rarifolia (C. Koch) Akhani (n 9), H. mollissima Bunge (n 9), H. pilifera Moq. (n 9), Halocharis sulphurea (Moq.) Moq. (n 9), Petrosimonia glauca (Pall.) Bunge (n 8), and Salsola florida (M. Bieb.) Poir (n 9). 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 x 9 is the basic chromosome number for the Chenopodiaceae family and the other numbers (e.g. x 8 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 n 27 (Fig. 1A). We found 27 bivalents in metaphase I (Fig. 1B) suggesting its probable allohexaploidy. The previous report for this species was 2n 18 (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 (2n 2x 18) (Fig. 1C), tetraploid (2n 4x 36) and hexaploid (2n 2x 54) levels in studied populations (Table 1). In hexaploid population only 27 bivalents in metaphase I was observed (Fig. 1C). But in tetraploid 2009 Fourteen Halophytic Species of Tribes Caroxyloneae and Salsoleae 81 Miotic Dehghani & Chromosome no. (n) Iran. D & 2007). et al. ehran: 35 7 km before Eshtehard, km W Mardabad (Rudeshur), 20.6.2006. D & 282 9 ehran: 80 4.8.2005. D & 342 km before Qom from Tehran, 9 Ardebil: opposite of Eskanlo gas station, 23.7.2005. D & 324Ardebil: opposite of Eskanlo gas Qom: 84 22.6.2005. D & 370 (Houze Soltan salt lake), km before Tehran Esfahan: 18 Esfahan, 4.8.2005. D & 347 km after Delijan towards Azerbaijan: 26 from Jolfa, 24.7.2005. D & 333 km before Poldashat Ardebil: 35 23.7.2005. 326 km before Jolfa from Parsabad, Semnan: 8 km before Semnan from Sorkhe, 20.8.2005. 340T 9 9 9 9 9 9 Ardebil: 55 from Ardebil , 22.7.2005. D & 321 km before Parsabad Azerbaijan: 6 Khalkhal, 21.7.2005. D & 307 km after Khalkhal-Mianeh junction towards 9 9 T Arak: 7 km before Arak from Delijan, 5.8.2005. D & 356 9 Seidlitzia Assadi Ardebil: 150 from Ardebi (33 km before Parsabad lo village), 22.7.2005. D & 320 km before Alahyar 9 Bunge) (Moq.) Bunge] K. Koch) (Bunge) Assadi Zanjan: 75 km before Mianeh from Zanjan, 21.7.2004. D & 304 9 (Litv.) Botsch.(Litv.) Botsch.(Litv.) 35km Tehran: 7 before Eshtehard, km W Mardabad (Rudeshur), 22.6.2005. D & 367 Arak: 26 5.8.2005. D & 360 Meyghan), Kavire km NE Arak (towards 27 18 Moq. 80km 4.8.2005. D & 343 Tehran: before Qom from Tehran, 9 Bunge 35km Tehran: 7 before Eshtehard, km W Mardabad (Rudeshur), 22.6.2005. D & 365 9 (Pall.) Bunge] (Pall.) Moq.) (K. Koch) Akhani (K. Koch) Azerbaijan: 34 km before Khalkhal-Mianeh junction from Zanjan, 21.7.2005. D & 305 9 Moq. 35km Tehran: 7 before Eshtehard, km W Mardabad (RudeShur), 22.6.2005. D & 368 9 (Pall.) Akhani (Pall.) Ardebil: 40 km before Eskanlu from Aslandoz, 23.7.2005. D & 323 9 (M. Bieb.) Botsch.(M. Bieb.) Azerbaijan: 5 (5 Pazi towards km from Poldashat km NW poldasht), 22.7.2005. D & 335 27 Moq. Semnan: 2 km before sangsar from Semnan, 20.8.2005. D & 339 9 Bunge 35km Tehran: 7 before Eshtehard, km W Mardabad (Rudeshur), 20.6.2006. D & 281 8 (Moq.) Akhani 35km Tehran: 7 before Eshtehard, km W Mardabad (Rudeshur), 20.6.2006. D & 283 18 Species Location: (M. Bieb.) Poir ( Poir (M. Bieb.) spec. Azerbaijan: Khalkhal-Mianeh-Maman junction 21.7.2004. D & 308 9 List of studied species/accessions, herbarium vouchers and chromosome numbers of 14 species of Caroxyloneae and Salsoleae from and chromosome numbers of 14 species Caroxyloneae List of studied species/accessions, herbarium vouchers changed (Akhani status are recently for species which are given Doulatyari. The synonyms (M. Bieb.) Bunge ex Boiss. Bunge ex (M. Bieb.) Salsola tomentosa Halanthium rarifolium Gamanthus pilosus Halanthium mamamense Gamanthus gamocarpus ( florida ( ( ( ( Table 1. etrosimonia glauca etrosimonia Salsola florida Halocharis sulphura Halocharis P Kaviria tomentosa Halimocnemis Halimocnemis rarifolia Halimocnemis mollissima Halimocnemis pilifera Halimocnemis pilosa Halimocnemis mamamensis Climacoptera crassa Climacoptera turcomanica Climacoptera turcomanica Climacoptera Halimocnemis azarbaijanensis Halimocnemis gamocarpa 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 2n 2x 18 (Federov, 1974).
Halimocnemis azarbaijanesis Assadi This taxon is described from E Azerbaijan province in NW Iran and probably is conspecific with the closely related species H. mamamensis. Meiosis in this taxon was regular and showed 9 bi- valents at diakinesis (Fig. 1E). Also, 9 diads at each pole of metaphase II were observed (Fig. 1F). This is the first chromosome number report for this species.
Halimocnemis gamocarpa Moq. In the meiosis of this species nine bivalents at diakinesis and metaphase I was observed (Figs. 1G–J). According to available literature, this seems to be the first chromosome number report for this species.
H. mamamensis (Bunge) Assadi (see notes under H. azerbaijanensis) This species is endemic to NW Iran. The gametic chromosome number of three studied popu- lations resulted n 9. Meiosis was regular and nine bivalents were observed in diakinensis and metaphase I (Figs. 1K–L). The mean number of chiasma per each bivalent was 1.11 in Azerbaijan population. This is the first chromosome number report for this species.
H. mollissima Bunge Meiosis in this taxon was regular and showed nine bivalents at diakinesis (Fig. 1M). The mean number of chiasma per each bivalent was 1.2. Chromosome number for this species is reported here for the first time.
H. pilifera Moq In three studied populations a diploidy level of n 9 has been known. Nine bivalents in diaki- nesis were observed (Fig. 1N). The present of lag chromosomes at metaphase II in Tehran popula- tion reported as an aberrant in meiosis of this species (Fig. 1O). This is the first chromosome count for this species.
H. pilosa (Pall.) Akhani This is an endemic species distributing in Transcaucasia and adjacent parts of Iran (Azerbaijan province). Nine diads at metaphase II were observed (Fig. 2A). According to our data, this is the first chromosome count for this species.
H. rarifolia (C. Koch) Akhani Seven populations of different provinces of Iran were studied (Table 1). In all populations sim- ilar ploidy level of n 9 were distinguished. Nine bivalents at diakinesis and first metaphase I was observed (Figs. 2B, C). The mean number of chiasma per each bivalent was 1.45 in Ardebil popula-
Fig. 1. Selected photos of meiotic chromosome compliments of some Iranian Caroxyloneae. Numbers after each name refer to the herbarium number listed in Table 1. A: Diakinesis in Climacoptera crassa (M. Bieb.) Botsch., 335 (n 27); B: Metaphase I in C. crassa 335 (n 27); C: Metaphase I in C. turcomanica (Litv.) Botsch., 367 (n 27); D: Diakinesis in C. turcomanica, 360 (n 18); E: Diakinesis in Halimocnemis azarbai- janensis Assadi, 320 (n 9); F: Metaphase II in H. azarbaijanensis, 320 (n 9); G: Early diakinesis in H. gamocarpa Moq, 343 (n 9); H: Middle diakinesis in H. gamocarpa, 343 (n 9); I: Late diakinesis in H. gamocarpa, 343 (n 9); J: Metaphase I in H. gamocarpa, 343 (n 9); K: Diakenesis in H. mamamensis (Bunge) Assadi, 304 (n 9); L: Metaphase I in H. mamamensis, 307 (n 9). M: Diakinesis in H. mollissima Bunge, 365 (n 9); N: Diakinesis in H. pilifera 368 (n 9); O: Metaphase II. H. pilifera (368). Scale bars 10 mm. 84 A. Doulatyari, S. M. Ghaffari and H. Akhani Cytologia 74(1)
Fig. 2. Selected photos of meiotic chromosome compliments of some Iranian Caroxyloneae and Salsoleae. A: Metaphase II in Helimocnemis pilosa (Pall.) Akhani, 323 (n 9); B: Diakinesis in H. rarifolia (K. Koch) Akhani 324 (n 9); C: Metaphase I in H. rarifolia, 333 (n 9); D: Diakinesis in Halimocnemis sp., 308 (n 9); E: Anaphase I in H. sp 308 (n 9). F: Diakinesis in Halocharis sulphurea Moq., 339 (n 9); G: Metaphase I in H. sulphurea Moq., 339 (n 9); H: Anaphase I in H. sulphurea, 339 (n 9); I: Diakinesis in Kaviria tomentosa (Moq.) Akhani, 283 (n 18); J: Metaphase I in P. glauca 281, (n 8); K: Diakinesis in Sal- sola florida (M. Bieb.) Poir 282 (n 9); L: Metaphase II in S. florida 281 (n 9). Scale bars 10 mm. 2009 Fourteen Halophytic Species of Tribes Caroxyloneae and Salsoleae 85 tion. Based on literature data, the chromosome count in this species is reported for the first time.
Halimocnemis sp. This plant has been collected from a population in the road between Mianeh and Khalkhal. It differs from H. rarifolia and H. mamamense by its erect habit and acute anther appendages. The at- tempts of the last author to recollect this plant and decide on its taxonomic status were not success- ful. Meiosis in this taxon was regular and showed nine bivalents at diakinesis (Fig. 2D), and nine diads at metaphase II were observed (Fig. 2E).
Halocharis sulphura Moq. About seven species of Halocharis distributed in C and SW Asia (Iljin 1949). Our sample showed nine bivalents at diakinesis and metaphase I (Figs. 2F, G). Also chromosome segregation at anaphase I was 9–9 (Fig. 2H). The only others two counts have been reported for H. hispida are 2n 18 (Federov 1974, Podlech & Bader 1974).
Kaviria tomentosa (Moq.) Akhani Meiosis in this species was regular with 18 bivalents at diakinesis which are mostly ring- shaped (Fig. 2I). The mean number of chiasma per each bivalent was 1.67. Bakhshi Khaniki and Maroof (2006) reported the same ploidy level for this taxon based on somatic chromosome compli- ments (2n 36).
Petrosimonia glauca Bunge In contrast to all studied taxa, n 8 was found in this species. Meiosis in this species was regu- lar with 8 bivalents at metaphase I (Fig. 2J). The previous reports for P. crassifoloia (Pall.) Bunge was 2n 16 according to Titova (Fedorov 1974) and 2n 14 for P. triandra (Pall.) Simonk (Fedorov 1974). Our present report confirms the former and rejects the later.
Salsola florida (Bunge) Poir This is a diploid species with n 9. Meiosis in this species was regular and showed 9 bivalents at diakinesis (Fig. 2K), also nine diads at each poles of metaphase II were observed (Fig. 2L). This is the first chromosome count for this species.
Discussion The results obtained from the chromosome studies on pollen mother cells, showed an equal basic chromosome number, x 9 in studied genera except the genus Petrosimonia which has a basic chromosome number of x 8. Our results correspond with previous findings of the rather homoge- nous basic chromosome number in Chenopodiaceae (Turner 1994). The basic chromosome number in the most closely related family Amarathaceae is mostly x 8. In the APG II (2003) classification both families are merged into the broadly defined Amaranthaceae. Chromosome coiling in the species of the genus of Halimocnemis s.l. was considerably long at substages of diakinesis, thus morphology of bivalents at early, middle and late diakinesis were clearly different. The morphology of bivalents at late diakiness and first metaphase was so similar that only by presence of nucleolus, the difference of these two substages could be distinguished (Figs. 1G–F). Two univalents of each bivalent in late diakinesis were connected by a thin layer which false impression that the species is a tetraploid with n 18 (Fig. 1I). This unique meiotic be- havior adds a cytological evidence confirming integration of the genera Gamanthus, Halanthium and Halimocnemis which recently have based on molecular and morphological analysis (Akhani et al. 2007). 86 A. Doulatyari, S. M. Ghaffari and H. Akhani Cytologia 74(1)
Confirming of a new chromosome base in the genus Petrosimonia (x 8) is of great interest. Based on ITS and PsbB-PsbH sequence analysis three species of Petrosimonia and the Ofaiston forms a monophyletic group which is sister to a clade comprising Pyankovia, Climacoptera and Halimocnemis s.l. (including Gamanthus, Halimocnemis s.str., Halanthium and Halotis) (Akhani et al. 2007). Morphologically the species of Petrosimonia and Ofaiston with their unique connate and cage-like anther appendages and medifixed hairs are so distinctive (Iljin 1936). As long as all other species of this calde have not been cytologically investigated we hesitate to explain it in an evolu- tionary context. However, the presence of x 9 chromosome base in other derivatives of Climacoptera clade and more basal clades such as Caroxylon clade suggest that might have been evolved through dysploidy. Deviation of x 9 chromosome base in Chenopodiaceae is very rare. Previously 2n 12 have been reported in Spinacia oleracea and 2n 32 in Chenopodium am- brosoides (Goldblatt & Johnson 1979). The two genera of Climacoptera and Kaviria are diversified in the Irano-Turanian arid and saline habitats. Both genera are extremely variable and there is no agreement on the species circum- scription (Pratov 1986, Hedge et al. 1997, Botschantzev 1980). Apparently the frequency of tetra and hexaploidy level in these genera indicate the role of polyploidy in the evolution and diversifica- tion of these genera. Therefore it is mandatory to extend our chromosome studies as much as possi- ble, given the potential of chromosome studies in the systematics of these genera.
Acknowledgments This paper is based on M. Sc. thesis of the first author (A.D) which is supported by a stu- dentship grant from research council University of Tehran, and additionally by the research project Geobotanical Studies in Different Parts of Iran for H.A. and the grant number 6401011/1/03 for S.M.G.
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