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Intergeneric Classification of Genus Bulbophyllum from Peninsular Malaysia Based on Combined Morphological and Rbcl Sequence Data

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Pak. J. Bot., 48(4): 1619-1627, 2016. INTERGENERIC CLASSIFICATION OF GENUS BULBOPHYLLUM FROM PENINSULAR MALAYSIA BASED ON COMBINED MORPHOLOGICAL AND RBCL SEQUENCE DATA SH. HOSSEINI1* AND K. DADKHAH2 1Department of Biological Science and Biotechnology University of Kurdistan, Sanandaj, Iran 2Department of Statistics, Faculty of Science, University of Kurdistan, Sanandaj, Iran *Corresponding author’s e-mail: [email protected] Abstract Bulbophyllum Thou. is largest genus in Orchidaceae family and a well-known plant of tropical area. The present study provides a comparative morphological study of 38 Bulbophyllum spp. as well as molecular sequence analysis of large subunit of rubisco (rbcL), to infer the intergeneric classification for studied taxa of genus Bulbophyllum. Thirty morphological characters were coded in a data matrix, and used in phenetic analysis. Morphological result was strongly consistent with earlier classification, with exception of B. auratum, B. gracillimum, B. mutabile and B. limbatum status. Furthermore Molecular data analysis of rbcL was congruent with morphological data in some aspects. Species interrelationships specified using combination of rbcL sequence data with morphological data. The results revealed close affiliation in 11 sections of Bulbophyllum from Peninsular Malaysia. Consequently, based on this study generic status of sections Cirrhopetalum and Epicrianthes cannot longer be supported, as they are deeply embedded within the genus Bulbophyllum. Key words: Bulbophyllum, Orchidaceae, Peninsular Malaysia, rbcL Introduction Jamil et al., 2014). The structural portion of this study was performed using qualitative characters of rhizome, The largest collection of subtribe Bulbophyllinae pseudobulb, leaf and flower for 38 different species. (Orchidaceae) have been defined as Bulbophyllum, a DNA sequence data from the large subunit of rubisco genus which form a large, pantropical, and poorly studied (rbcL) which produces the carbon dioxide fixing enzyme group of orchids in Peninsular Malaysia. Bulbophyllum of chloroplast and subsequently combination of species are mostly epiphytic and they are found in morphological binary characters with rbcL sequence different habitats ranging from (sub) tropical dry forests data were analyzed to infer the intergeneric to wet montane cloud forests and most of them are fly classification of Bulbophyllum. The main objectives of pollinated (Tan et al., 2002; Nishida et al., 2004; Teixeira the present study was; 1) to determine and tracing the et al., 2004). Latest traditional taxonomy (Seidenfaden & close relationships among various section belonging to Wood, 1992) was described more than 100 Bulbophyllum the genus Bulbophyllum using rbcL sequence data and species of Peninsular Malaysia into 17 sections. However, structural data and 2) to verify the status of sections based on this study, the members of this large genus have Cirrhopetalum and Epicrianthes. undergone extreme reduction in number of species in the natural habitat and have acquired an efficient adaptation Materials and Methods to the canopy environment. Botanical treatises by such eminent botanists as Sample collection: For this study, 38 species were Meisner (1842), Endlicher (1837), Bentham & Hooker collected from a variety of locations in Peninsular (1883), Hooker (1890), Pfitzer (1888) and Schlechter Malaysia (Table 1). Species are representing 11 sections (1914) recognize Cirrhopetalum as a genus. Garay as described by Holttum (1953), Seidenfaden & Wood (1994) was not concurred with Seidenfaden & Wood (1992). Morphological characters for both vegetative (1992) opinion which had transfered all Cirrhopetalum (included the size of the plant, the characters of rhizome, to Bulbophyllum. He was believed that section leaves and pseudobulb) and flower structures (included Cirrhopetalum is a unique genus in the orchid family the size and structure of the petals and sepals, shape of the because it can be defined unmistakeably through a single lip, column, structure of the inflorescence and flowers and character, i.e., the way the lateral sepals are formed at so on) were studied for identification and description of the base of the column foot. Bulbophyllum species. Voucher specimens for all Garay & Kittredge (1985) have proposed a return to accessions have been deposited in herbarium of biology separate generic status for section Epicrianthes but department, Universiti Putra Malaysia (UPM). Vermeulen (1991) was pointed at affinities with section Leptopus. Systematic study of combined molecular and Morphological study: To construct the species morphological data on this genus will gain us a better interrelationship by numerical taxonomy, thirty insight. Most systematists agree that morphological data qualitative characters were selected based on those can be potentially useful in phylogenetic analyses, reported by Fischer et al., 2007 and our own field especially in combination with molecular data (Kron et observations (Table 2). Qualitative characters were coded al., 2002; Shinwari et al., 1994; Nadeem et al., 2013; as binary/multistate characters (Table 3). 1620 SH. HOSSEINI & K. DADKHAH Table 1. Plant materials used in this study. Gen bank Code Section Taxon Location Herbarium/Voucher accession No. 1. Hirtula B. dayanum Gunung Jerai, Malaysia UPM/ B0014 JF428000 2. Hirtula B. limbatum Johor, Malaysia UPM / B0054 JF428036 3. Hirtula B. hirtulum Fraser s Hill, Malaysia UPM / FAN. FH- 314 JF428030 4. Cirrhopetalum B. flabellum Genting highland, Malaysia UPM / RG 1945 JF427996 5. Cirrhopetalum B. purpurascens Cameron Highland, Malaysia UPM/ B0027 JF428010 6. Cirrhopetalum B. vaginatum Melacca, Malaysia UPM/ FAN. FH- 503 JF428005 7. Cirrhopetalum B. corolliferum Gunung Belumut, Malaysia UPM / B0026 JF428009 8. Cirrhopetalum B. acuminatum Gunung Belumut, Malaysia UPM / RG 2291 JF428021 9. Cirrhopetalum B. auratum Cameron Highland, Malaysia UPM / B0060 JF428040 10. Cirrhopetalum B. gracillimum Genting Highland, Malaysia UPM / B0053 JF428037 11. Aphanobulbon B. flavescens Fraser s Hill, Malaysia UPM / FAN. FH- 062 JF427998 12. Aphanobulbon B. mutabile Fraser s Hill, Malaysia UPM / FAN. FH- 105 JF427997 13. Aphanobulbon B. linearifolium Fraser s Hill, Malaysia UPM / FAN. FH- 258 JF427999 14. Aphanobulbon B. apodum Cameron Highlands, Malaysia UPM / FAN. FH- 276 JF428039 15. Aphanobulbon B. odoratum Pahang, Malaysia UPM / B0056 JF428034 16. Aphanobulbon B. armeniacum Fraser s Hill, Malaysia UPM / SH.K-105 JF428015 17. Desmosanthes B. concinnum Genting highland, Malaysia UPM / RG 2207 JF428006 18. Desmosanthes B. sulcatum Gunung Jerai, Malaysia UPM / FAN. FH- 304 JF427995 19. Desmosanthes B. angustifolium Fraser s Hill, Malaysia UPM / RG 2313 JF427993 20. Desmosanthes B. medusae Johor, Malaysia UPM / B0052 JF428038 21. Desmosanthes B. bakhuizenii Gunung Jerai, Malaysia UPM / SH.K-107 JF428019 22. Desmosanthes B. obtusum Fraser s Hill, Malaysia UPM / FAN. FH- 172 JF427994 23. Sestochilus B. macranthum Cameron Highland, Malaysia UPM / FAN. FH- 153 JF427988 24. Sestochilus B. inunctum Gunung Jerai, Malaysia UPM / SH.K-109 JF427989 25. Sestochilus B. lobbii Cameron Highland, Malaysia UPM / FAN. FH- 426 JF427991 26. Sestochilus B. uniflorum Fraser s Hill, Malaysia UPM / FAN. FH- 107 JF427990 27. Sestochilus B. patens Gunung Jerai, Malaysia UPM / B005 JF427992 28. Sestochilus B. pileatum Gunung Belumut, Malaysia UPM / RG 2281 JF428007 29. Sestochilus B. lasianthum Fraser s Hill, Malaysia UPM / RG 1922 JF428026 30. Careyana B. lilacinum Gunung Jerai, Malaysia UPM / B0029 JF428012 31. Careyana B. sichyobulbon Gunung Jerai, Malaysia UPM / SH.K-111 JF428002 32. Monilibulbus B. stormii Cameron Highlands, Malaysia UPM / B0058 JF428042 33. Monilibulbus B. ovalifolium Cameron Highland, Malaysia UPM / RG 2167 JF428003 34. Globiceps B. coniferum Cameron Highland, Malaysia UPM / RG1757 JF428041 35. Leptopus B. tenuifolium Cameron Highland, Malaysia UPM / B0061 JF428043 36. Polyblepharon B. membranaceum Fraser s Hill, Malaysia UPM / B0024 JF428028 37. Epicrianthes B. cheiropetalum Gunung Jerai, Malaysia Gunung Jerai, UPM / B0018 JF428004 Malaysia Gunung Jerai, Malaysia 38. Epicrianthes B. haniffii Penang, Malaysia UPM / B0031 JF428013 39. Distichorchis D. pahangensis Fraser s Hill, Malaysia UPM / FAN. FH- 180 JF428045 CLASSIFICATION OF THE GENUS BULBOPHYLLUM FROM PENINSULAR MALAYSIA 1621 Table 2. List of characters scored for cluster analysis of studied taxa. No. Characters description 1. Size of the plant: 0 = small (≤3 cm) / 1 = intermediate (4-10cm) / 2 = large (≥10cm) 2. Fresh sheaths or remnants of fresh sheaths covering the rhizome and part of the pseudobulbs: 0 = present/1 = absent Pseudobulbs: 0 = crowded (distance between bulbs is less than the diameter of a bulb)/ 1 = moderately spaced 3. (distance between bulbs is 1–10 times the diameter of a bulb 4. Pseudobulb shape: 0= angled/1= ovoid or oblong 5. Pseudobulb colour: 0 = green& yellow/ 1= brown 6. Pseudobulbs: 0 =flattened or somewhat flattened /1= no flattened 7. Pseudobulb: 0= present the hair around the pseudobulb/1= absent the hair 8. Stem or pseudobulb length: 0= stem/pseudobulb absent or short, 1= elongated 9. Number of leafs per pseudobulbs: 0 = 1/1 = 2 10. Leaf apex: 0= blunt/ 1= cleft/ 2= acute or bilobed acute 11. Leaf thickness: 0 = thick/ 1= thin 12. Leaf size: 0= small (less than 3 cm long)/ 1= intermediate (between 3-12 cm)/ 2= large (more than 12cm) 13. Leaf color: 0= green/ 1= otherwise 14. Peduncle setaceous (bristle like): 0 = yes/1 = no 15. Inflorescence:
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