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Pollen Morphology of Centaurea L. (Asteraceae) in Iran

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Pollen Morphology of Centaurea L. (Asteraceae) in Iran Acta Botanica Brasilica 27(4): 669-679. 2013. Pollen morphology of Centaurea L. (Asteraceae) in Iran Elaheh Sadat Bagheri Shabestari1,3, Farideh Attar2, Hosein Riahi1 and Masoud Sheidai1 Received: 10 December, 2012. Accepted: 6 June, 2013 ABSTRACT The genus Centaurea L. is one of the largest genera in the family Asteraceae. The number of taxa included in this genus ranges from 500 to 600 worldwide. Iran is one of the major centers of diversity for this genus. Pollen morphology has proved useful in the systematics of Asteraceae, as well as in that of some of its genera and species. The pollen morphology of 19 taxa of the genus Centaurea was investigated in detail by light and scanning electron microscopy. Examination under light microscopy showed pollen grains to be tricolporate, isopolar, radially symmetrical, oblate spheroidal, prolate spheroidal, subprolate, tectum perforate and scabrate. Based on the exine ornamentation observed under scanning electron microscopy, we identified two types of pollen grains, designated type I (dense acute spinules) and type II (sparse spinules). All of the taxa evaluated resemble Wagenitz’s Jacea type. We present micrographs of pollen grains, together with descriptions of the characteristics of their structure. We found that the pollen characters (shape, exine structure, sculpturing) were useful for classification. Key words: Centaurea, palynology, exine ornamentation, light microscopy, scanning electron microscopy Introduction & Ghanbarian 2007), none of which were conducted in Iran. Therefore, the present survey aims to present detailed The genus Centaurea L. (Asteraceae) is represented by quantitative and qualitative data on the pollen morphology 500-600 taxa worldwide (Rendle 1976; Heywood 1979; of the genus, as well as to evaluate the taxonomic value of Hickey & King 1981) and occurs primarily in Asia, Eu- those data. rope, North Africa and North America (Davis et al. 1988). In Iran, the genus is highly (42.35%) endemic (Wagenitz 1980). Of the 86 Centaurea taxa in Iran, 36 are endemic to Material and methods the country (Ghahreman & Attar 1999). The high level of We employed LM and SEM in order to conduct pa- endemism of Centaurea in Iran supports the assertion that lynological studies of 19 species of the genus Centaurea the country is one of the major centers of diversity for the belonging to four sections (Wagenitz 1980): genus (Wagenitz 1986). a) Phaeopappus - C. spectabilis (Fisch. & C.A. Mey) Pollen morphology has been proved to be useful in the Schultz-Bip., C. aucheri subsp. aucheri, C. aucheri subsp. systematics of the Asteraceae family, as well as that of some szowitsii (Boiss.) Wagenitz, C. aucheri subsp. indistincta of its genera and species (Moore et al. 1991). Wagenitz Wagenitz, C. aucheri subsp. elbursensis Wagenitz. C. aucheri (1955) divided the genus Centaurea into eight subgenera, subsp. farsistanica Wagenitz and C. albonitens Turill. on the basis of exine structure and sculpturing: Serratula, b) Cynaroides - C. regia subsp. regia, Boiss. C. imperialis Centaurium, Scabiosa, Crupina, Jacea, Dealbata, Montana Hausskn. ex Bornm., C. phlomoides Boiss. & Hausskn., and Cyanus. Villodre & Garcia-Jacas (2000) stated that C. geluensis Boiss. & Hausskn. and C. gabrielae (Bornm.) pollen type is one of the most reliable characteristics for Wagenitz. establishing phylogeny in the Jacea group. There have c) Paraphysis - including C. amadanensis var. amada- been only a few studies employing light microscopy (LM) nensis, C. amadanensis var. gymnoclada (Jaub. & Spach.) or scanning electron microscopy (SEM) to evaluate the Wagenitz and C. nemecii Nab. pollen morphology of Centaurea species (Uysal et al. 2005; d) Microlophus - C. polypodiifolia var. polypodiifolia, Celik et al. 2005; Celik et al. 2008; Özler et al. 2009; Jafari C. behen L., C. pabotii Wagenitz and C. koieana Bornm. 1 Shahid Beheshti University, GC, Faculty of Biological Sciences, Tehran, Iran 2 University of Tehran, School of Biology and Center of Excellence in Phylogeny of Living Organisms, College of Science, Tehran, Iran 3 Author for correspondence: [email protected] Elaheh Sadat Bagheri Shabestari, Farideh Attar, Hosein Riahi and Masoud Sheidai Voucher specimens are listed in Table 1. Voucher specimens Results are deposited in the Central Herbarium of Tehran University (code, TUH), the Herbarium of the Iranian Research Institute All of the morphological parameters investigated are of Plant Protection (code, IRAN) and the Herbarium of Sha- shown in table 2-4 and in fig. 1-75. Under LM, the pollen hid Beheshti University (code, HSBU). We studied a total of grains are isopolar, radially symmetrical, oblate spheroidal, 15 quantitative and qualitative pollen characters (Table 2-4). prolate spheroidal and subprolate. Their apertures are tri- Characters were selected on the basis of those reported in the colporate. In the polar view, pollen grains are circular and literature (Özler et al. 2009; Zamani et al. 2010) and our own the colpi are acute at both ends (Fig. 58-75). The colpus observations of Centaurea species. Pollen obtained from flower membrane is granulate-perforate. In all taxa examined, the buds at anthesis was prepared for LM according to Wodehouse ectoaperture is bigger than the endoaperture (6.96-18.66 (1935) and Erdtman (1960). The LM observations of acetolyzed μm vs. 2.93-7.73 μm) (Table 3). pollen grains were made with an Olympus binocular micro- The distance between colpi ranged from 22.35±1.31 μm scope at 1000× magnification. Acetolyzed pollen grains were (Table 3), in C. koieana, to 37.11±2.09 μm, in C. amadanensis observed and measured with ImageTool software, version 3 var. amadanensis. The distance between a colpus and the (UTHSCSA, Texas, USA). Measurements are scored for at pole was longest for C. aucheri subsp. aucheri (22.27±1.49 least 20-25 mature pollen grains per specimen. μm) and shortest for C. pabotii (13.37±1.02 μm). For SEM, after acetolysis, the specimens were mounted The polar diameter ranges from 39.98±1.09 (Table 2), on 12.5 mm diameter aluminum stubs and sputter-coated in Centaurea behen, to 55.37±2.05 μm, in C. aucheri subsp. with approximately 25 nm of gold-palladium. The speci- aucheri; the equatorial diameter ranges from 34.69±1.26, in mens were examined and photographed with a scanning C. koieana, to 49.03±1.63 μm, in C. aucheri subsp. aucheri. electron microscope (XL30; Philips, Eindhoven, The Neth- Under LM and SEM, the exine sculpture is scabrate erlands) at an accelerating voltage of 20 kV. The terminology and the tectum is perforate, respectively (Fig. 1-57). Exine follows mainly Ertdman (1952) and Halbitter et al. (2007). thickness ranges from 4.52 μm, in Centaurea koieana, to We analyzed a total of 19 taxa belonging to the genus Cen- 7.81 μm, in C. gabrielae. Under SEM, the number of perfora- taurea. Among such taxa, the exine sculpture is reportedly tions counted ranged from 19.71/μm2, in C. aucheri subsp. scabrate when the sculpturing elements are less than 1 μm indistincta, to 5.04/μm2, in C. polypodiifolia var. polypodii- and microechinate when they are 1-2 μm (Wagenitz 1955). folia. The number of microspines ranged from 5.5/μm2, in Table 1. List of taxa of Centaurea L. (Asteraceae) examined, collectors and herbarium number. Taxa Date Collector, herbarium-accession number C. albonitens 20 June, 1982 Ghahreman et al., TUH-11447 C. amadanensis var. amadanensis 7 June, 2009 Bagheri, HSBU-1007 C. amadanensis var. gymnoclada 8 July, 1995 Ghahreman & Attar, TUH-29650 C. aucheri subsp. aucheri 11 June, 2009 Bagheri, HSBU-1002 C. aucheri subsp. elbursensis 28 May, 2009 Mehrabian, HSBU-1004 C. aucheri subsp. farsistanica 25 June, 1987 Kashkuli, IRAN-7913 C. aucheri subsp. indistincta 28 May, 1968 Termeh, IRAN-7932 C. aucheri subsp. szowitsii 14 June, 2009 Bagheri, HSBU-1003 C. behen 11 July, 2009 Bagheri, HSBU-1005 C. gabrielae 27 April, 1983 Termeh et al., IRAN-842 C. geluensis 23 June, 1999 Veiskarami, TUH-23715 C. imperialis 9 July, 1995 Ghahreman & Attar, TUH-19667 C. koieana 3 June, 1999 Ghahreman et al., TUH-22380 C. nemecii 13 June, 1988 Ghaffari, TUH-5867 C. pabotii 7 June, 2009 Bagheri, HSBU-1006 C. phlomoides 3 July, 1996 Ghahreman & Attar, TUH-19857 C. polypodiifolia var. polypodiifolia 27 June, 1994 Ghahreman & Mozaffarian, TUH-17438 C. regia subsp. regia 3 July, 1996 Ghahreman & Attar, TUH-20565 C. spectabilis 9 June, 2010 Bagheri, HSBU-1001 HSBU – Herbarium of Shahid Beheshti University; TUH – Central Herbarium of Tehran University; IRAN – Herbarium of the Iranian Research Institute of Plant Protection. 670 Acta bot. bras. 27(4): 669-679. 2013. Pollen morphology of Centaurea L. (Asteraceae) in Iran Figures 1-15. Pollen grains (SEM) of species of Centaurea L. (Asteraceae). (1-3) C. albonitens; (4-6) C. aucheri subsp. aucheri; (7-9) C. aucheri subsp. elbursensis; (10-12) C. aucheri subsp. farsistanica; (13-15) C. aucheri subsp. szowitsii. Acta bot. bras. 27(4): 669-679. 2013. 671 Elaheh Sadat Bagheri Shabestari, Farideh Attar, Hosein Riahi and Masoud Sheidai Figures 16-30. Pollen grains (SEM) of species of Centaurea L. (Asteraceae). (16-18) C. aucheri subsp. indistincta; (19-21) C. spectabilis; (22-24) C. amadanensis var. amadanensis; (25-27) C. amadanensis var. gymnoclada; (28-30) C. nemecii. 672 Acta bot. bras. 27(4): 669-679. 2013. Pollen morphology of Centaurea L. (Asteraceae) in Iran Table 2. Size and shape of pollen grains of taxa of Centaurea L. (Asteraceae) examined. (PD – polar diameter; ED – equatorial diameter). Taxa PD μm ED μm PD/ED Shape Sect. Cynaroides C. gabrielae 39.50 (48.23±2.88) 53.02 37.74 (42.75±1.62) 45.12 1.12 prolate spheroidal C. geluensis 39.34 (45.21±3.96) 50.66 37.40 (42.48±3.01) 47.34 1.06 prolate spheroidal C. imperialis 47.08 (48.32±1.04) 49.78 44.06 (45.77±1.24) 47.20 1.05 prolate spheroidal C.
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