2008-Chromosome numbers of five species of the Marattiaceae in Taiwan

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335 Taiwan J For Sci 23(4): 335-9, 2008 Research note Chromosome Numbers of Five Species of the Marattiaceae in Taiwan Tsung-Hsin Hsieh, 1) Yi-Han Chang, 2) Wen-Liang Chiou, 2) Yao-Moan Huang 2,3) SummaryThe chromosome numbers of root tips of all 5 species of the Marattiaceae in Taiwan were determined. Results showed that Angiopteris lygodiifolia (2n = 80), Angiopteris palmiformis (2n = 80), and Marattia pellucida (2n = 78) are diploid; Archangiopteris itoi is triploid (2n = 120), and Archangiopteris somai is tetraploid (2n = 160). These chromosome numbers together with previ- ous evidence support the hypothesis that Archangiopteris itoi is a hybrid between Angiopteris ly- godiifolia and Archangiopteris somai. Key words: Chromosome number, ploidy, Marattiaceae, Taiwan. Hsieh TH, Chang YH, Chiou WL, Huang YM. 2008. Chromosome numbers of five species of the Marattiaceae in Taiwan. Taiwan J For Sci 23(4):335-9. 研究簡報 台灣產五種觀音座蓮舅科植物之染色體數目 謝宗欣 1) 張藝翰 2) 邱文良 2) 黃曜謀 2,3) 本文觀察台灣產5種觀音座蓮舅科植物之染色體數目。觀音座蓮(2n = 80) 、蘭嶼觀音座蓮(2n = 80) 及觀音座蓮舅(2n =78) 為二倍體,伊藤氏原始觀音座蓮為三倍體(2n = 120) ,台灣原始觀音座蓮為四倍 (2n = 160) 。這些染色體數目與已往之研究證據均支持伊藤氏原始觀音座蓮為觀音座蓮與台灣原始觀 音座蓮之雜交後代的假說。 關鍵詞:染色體數、倍體數、觀音座蓮舅科、台灣。 謝宗欣、張藝翰、邱文良、黃曜謀2008。台灣產五種觀音座蓮舅科植物之染色體數目。台灣林業科 23(4):335-91) Department of Biological Sciences and Technology, National University of Tainan, 33 Su-Lin St., Sec. 2, Tainan 70005, Taiwan. 國立台南大學生物科技學系,70005 台南市樹林街二段33 號。 2) Division of Forest Biology, Taiwan Forestry Research Institute, 53 Nanhai Rd., Taipei 10066, Taiwan. 林業試驗所森林生物組,10066 臺北市南海路53 號。 3) Corresponding author, e-mail:[email protected] 通訊作者。 Received July 2008, Accepted September 2008. 2008 7 月送審 2008 9 月通過。

Transcript of 2008-Chromosome numbers of five species of the Marattiaceae in Taiwan

335Taiwan J For Sci 23(4): 335-9, 2008

Research note

Chromosome Numbers of Five Species of the Marattiaceae in Taiwan

Tsung-Hsin Hsieh,1) Yi-Han Chang,2) Wen-Liang Chiou,2) Yao-Moan Huang2,3)

【Summary】

The chromosome numbers of root tips of all 5 species of the Marattiaceae in Taiwan were determined. Results showed that Angiopteris lygodiifolia (2n = 80), Angiopteris palmiformis (2n = 80), and Marattia pellucida (2n = 78) are diploid; Archangiopteris itoi is triploid (2n = 120), and Archangiopteris somai is tetraploid (2n = 160). These chromosome numbers together with previ-ous evidence support the hypothesis that Archangiopteris itoi is a hybrid between Angiopteris ly-godiifolia and Archangiopteris somai.Key words: Chromosome number, ploidy, Marattiaceae, Taiwan.Hsieh TH, Chang YH, Chiou WL, Huang YM. 2008. Chromosome numbers of five species of the

Marattiaceae in Taiwan. Taiwan J For Sci 23(4):335-9.

研究簡報

台灣產五種觀音座蓮舅科植物之染色體數目

謝宗欣1) 張藝翰2) 邱文良2) 黃曜謀2,3)

摘 要

本文觀察台灣產5種觀音座蓮舅科植物之染色體數目。觀音座蓮(2n = 80)、蘭嶼觀音座蓮(2n = 80)及觀音座蓮舅(2n =78)為二倍體,伊藤氏原始觀音座蓮為三倍體(2n = 120),台灣原始觀音座蓮為四倍體(2n = 160)。這些染色體數目與已往之研究證據均支持伊藤氏原始觀音座蓮為觀音座蓮與台灣原始觀音座蓮之雜交後代的假說。

關鍵詞:染色體數、倍體數、觀音座蓮舅科、台灣。

謝宗欣、張藝翰、邱文良、黃曜謀。2008。台灣產五種觀音座蓮舅科植物之染色體數目。台灣林業科學23(4):335-9。

1) Department of Biological Sciences and Technology, National Universi ty of Tainan, 33 Su-Lin St . ,

Sec. 2, Tainan 70005, Taiwan. 國立台南大學生物科技學系,70005台南市樹林街二段33號。2) Div i s ion of Fores t Bio logy, Ta iwan Fores t ry Research Ins t i tu te , 53 Nanha i Rd . , Ta ipe i 10066 ,

Taiwan. 林業試驗所森林生物組,10066臺北市南海路53號。3) Corresponding author, e-mail:huangym@tfri .gov.tw 通訊作者。

Received July 2008, Accepted September 2008. 2008年7月送審 2008年9月通過。

336 Hsieh et al.―Chromosome numbers of Marattiaceae

There are 5 species of the family Marat-tiaceae native to Taiwan: Angiopteris lygo-diifolia Rosenst., Angiopteris palmiformis (Cav.) C. Chr., Archangiopteris itoi Shieh, Archangiopteris somai Hayata, and Marat-tia pellucida Presl (DeVol and Shieh 1994). In Taiwan, Angiopteris lygodiifolia is a very common fern. Angiopteris palmiformis is common on Lanyu (Orchid Island). Marattia pellucida is also only distributed on Lanyu but is rare. Archangiopteris itoi and Arch-angiopteris somai are endemic species and sympatrically distributed in Wulai, northern Taiwan (DeVol 1975, DeVol and Shieh 1994).

Two basic chromosome numbers of Marattiaceae (x = 40, 39) have been docu-mented. The basic chromosome number is 39 for Marattia and 40 for other genera (Braith-waite 1977, Walker 1985, Manickam and Irudayaraj 1988, Nakato 1988, Singh and Roy 1988, Weng and Qiu 1988, Li 1989, Camus 1990, Takamiya 1995). Of these, Angiopteris lygodiifolia was previously reported to be n = 40 (Tsai and Shieh 1983), and Archangiop-teris somai was reported to be 2n = 4x = 160 in Taiwan (Chou et al. 2007). Chromosome numbers of the root tips of all 5 species of the Marattiaceae in Taiwan were determined herein.

Sixteen plants of 5 species were collect-ed from Taiwan and a nearby island, Lanyu. Those plants were transplanted to the Taipei Botanical Garden (Table 1). After about 1 month of cultivation, the root tips were pre-treated for 3~4 h with a mixture of 70 ppm cycloheximide and 250 ppm 8-hydroxy-quinoline (1:1) at about 18~20℃. The root tips were fixed for 1~3 h in a mixture of 45% acetic acid and absolute ethanol (1:3) at about 20℃, and then preserved in 70% ethanol at 4℃. Then they were macerated for 1~3 min in 1 N HCl at 60℃, washed for 10 s and di-gested for 1~2 h in 4% pectinase. Finally, the root tips were squashed in modified Sharma’s solution (Sharma 1982). The chromosome number of root tip cells was observed with a light microscope (Leitz, Dialux 20) and photographed with a digital camera (Nikon, Coolpix 995).

The basic chromosome number of more than 22 species of Angiopteris has been documented to be 40 (Lin et al. 1996). Table 2 and Figures 1~5 show that Angiopteris ly-godiifolia and Angiopteris palmiformis are diploid (2n = 2x = 80). Archangiopteris itoi is triploid (2n = 3x = 120), assuming that the basic number is 40. Archangiopteris somai is tetraploid (2n = 4x = 160), as previous

Table 1. Materials of marattoid species in this study Species Locality Sample size Voucher1)

Angiopteris lygodiifolia Wulai, Taipei Co. 2 CM Chen s.n. Toucheng, Ilan Co. 1 MH Huang 387Angiopteris palmiformis Lanyu, Taitung Co. 3 CW Chen s.n. (2 plants) SW Chung s.n.Archangiopteris itoi Wulai, Taipei Co. 3 YM Huang 1314, 1315, 1316Archangiopteris somai Wulai, Taipei Co. 2 YM Huang 1317, 1318 Lienhuachih, Nantou Co. 3 YM Huang 1319, 1320, 1321Marattia pellucida Lanyu, Taitung Co. 2 CW Chun s.n. PF Lu s.n.1) Living plants cultured at Taipei Botanical Garden.

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Figs. 1~5. Chromosomes of the root tips of members of the Marattiaceae in Taiwan. 1. Angiopteris lygodiifolia (2n = 80); 2. Angiopteris palmiformis (2n = 80); 3. Marattia pellucida (2n = 78); 4. Archangiopteris itoi (2n = 120); 5. Archangiopteris somai (2n = 160). Scale bars = 10 µm.

Table 2. Chromosome numbers of Marattiaceae species in Taiwan Chromosome number Species Present count Previous count Ploidy level Reference 2n n 2n Angiopteris lygodiifolia 80 40 2x Tsai and Shieh (1983) 80 2x Nakato (1988)Angiopteris palmiformis 80 2x Archangiopteris itoi 120 3x Archangiopteris somai 160 160 4x Chou et al. (2007)Marattia pellucida 78 2x

338 Hsieh et al.―Chromosome numbers of Marattiaceae

determination (Chou et al. 2007). Marattia pellucida is diploid (2n = 2x = 78), based on the basic chromosome number of 39, as pre-viously documented in Marattia fraxinea and Marattia smithii (Manickam and Irudayaraj 1988, Takamiya 1995).

The morphology of Archangiopteris itoi is intermediate between Angiopteris ly-godiifolia and Archangiopteris somai. These 3 species are sympatrically distributed in Wulai. Archangiopteris itoi shares common patterns of RAPD markers with Angiopteris lygodiifolia and Archangiopteris somai and was hypothesized to be a hybrid of the lat-ter 2 species (Wu 2002). The chromosome numbers shown in this study also support this hypothesis.

The family Marattiaceae has previously been considered to have 6 genera, Archangi-opteris, Angiopteris, Christensenia, Danaea, Marattia, and Macroglossum (Holttum 1954, Pichi Sermolli 1977). However, the Marattia-ceae was recently treated as having 4 genera: Danaea, Christensenia, Marattia, and Angi-opteris (Camus 1990, Smith et al. 2006, Mur-dock 2008). The genera Archangiopteris and Macroglossum were nested within the Angi-opteris clade based on molecular phylogeny. Angiopteris is monophyletic only if Archangi-opteris and Macroglossum are included (Mur-dock 2008). The same basic chromosome numbers of Angiopteris and Archangiopteris, and the possibility of hybridization between species of these 2 genera indicate no signifi-cant boundary between these 2 genera.

ACKNOWLEDGEMENTS

The authors wish to thank Mei-Hsueh Huang, Chien-Wen Chen, Pi-Fong Lu, and Shih-Wen Chung for collecting live materials.

LITERATURE CITED

Braithwaite AF. 1977. A chromosome count and range extension for Christensenia (Marat-tiaceae). Am Fern J 67:49-50. Camus JM. 1990. Marattiaceae. In: Kubitzki K, editor. The families and genera of vascular plants. Volume I. Berlin, Germany: Springer-Verlag. p 174-80.Chou HM, Huang YM, Wong SL, Hsieh TH, Hsu SY, Chiou WL. 2007. Observations on gametophytes and juvenile sporophytes of Archangiopteris somai Hayata (Marattia-ceae), an endangered fern in Taiwan. Bot Stud 48:205-13.DeVol CE. 1975. Marattiaceae. In: Li HL, Liu TS, Huang TC, Koyama T, DeVol CE, editors. Flora of Taiwan. Volume I. Taipei, Taiwan: Ep-och Publishing Co. p 71-8.DeVol CE, Shieh WC. 1994. Marattiaceae. In: Editorial Committee of The Flora of Taiwan, editors. Flora of Taiwan. Volume I. Taipei, Taiwan: Editorial Committee of The Flora of Taiwan. p 74-9.Holttum RE. 1954. A revised flora of Malaya. Vol. 2, Ferns of Malaya. Singapore: Govt Print Off. 643 p.Li JW. 1989. Chromosome numbers of some species in the genus Angiopteris Hoffm. from Yunnan, China. In: Shing KH, Kramer KU, editors. Proc Int Symp Systematic Pteridology. Beijing, China: Science and Technology Press. p 109-10.Lin SJ, Iwatsuki K, Kato M. 1996. Cytotaxo-nomic study of ferns from China I. Species of Yunnan. J Jpn Bot 71:214-22.Manickam VS, Irudayaraj V. 1988. Cytology of ferns of the western Ghats (South India). New Delhi, India: Today and Tomorrow Print-ers and Publishers. 81 p. Murdock A. 2008. Phylogeny of marattoid ferns (Marattiaceae): inferring a root in the ab-sence of a closely related outgroup. Am J Bot 95:626-41.Nakato N. 1988. Notes on chromosomes of

339Taiwan J For Sci 23(4): 335-9, 2008

Japanese pteridophytes (2). J Jpn Bot 63:214-8.Pichi Sermolli REG. 1977. Tentamen pteri-dophytorum genera in taxonomicum ordinem redigendi. Webbia 31:313-512.Sharma HC. 1982. A technique for somatic counts from root tips of cereal seedlings raised by embryo culture. Curr Sci 51:143-4.Singh VP, Roy SK. 1988. Cytology of forty four species from Sikkim, Himalaya. Indian Fern J 5:162-9.Smith AR, Rryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG. 2006. A classifica-tion for extant ferns. Taxon 55:705-31.Takamiya M. 1995. Chromosomal studies of ferns and fern-allies in the Republics of Fiji and Vanuatu, South Pacific. I. Psilotaceae,

Ophioglossaceae, Marattiaceae and Schizaea-ceae. Acta Phytotax Geobot 46:137-45. Tsai JL, Shieh WC. 1983. A cytotaxonomic survey of the Pteridophytes in Taiwan. J Sci Engin 20:137-58.Walker TG. 1985. Cytotaxonomic studies of the ferns of Trinidad 2. The cytology and taxo-nomic implications. Bull Br Mus (Nat Hist) Bot 13:149-249. Weng RF, Qiu SP. 1988. Chromosome counts of some ferns from Zhejiang. Invest Stud Nat 8:43-52. Wu WH. 2002. A study on the natural fern hybrid – Archangiopteris itoi Shieh [Master thesis]. Taipei, Taiwan: Department of Botany, National Taiwan University.

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