REINWARDTIA A JOURNAL ON TAXONOMIC BOTANY, PLANT SOCIOLOGY AND ECOLOGY Vol. 14(1): 1 - 2 4 8 , December 23, 2014 Chief Editor Kartini Kramadibrata (Mycologist, Herbarium Bogoriense, Indonesia) Editors Dedy Darnaedi (Taxonomist, Herbarium Bogoriense, Indonesia) Tukirin Partomihardjo (Ecologist, Herbarium Bogoriense, Indonesia) Joeni Setijo Rahajoe (Ecologist, Herbarium Bogoriense, Indonesia) Marlina Ardiyani (Taxonomist, Herbarium Bogoriense, Indonesia) Topik Hidayat (Taxonomist, Indonesia University of Education, Indonesia) Eizi Suzuki (Ecologist, Kagoshima University, Japan) Jun Wen (Taxonomist, Smithsonian Natural History Museum, USA) Managing Editor Himmah Rustiami (Taxonomist, Herbarium Bogoriense, Indonesia) Lulut Dwi Sulistyaningsih (Taxonomist, Herbarium Bogoriense, Indonesia) Secretary Endang Tri Utami Layout Editor Deden Sumirat Hidayat Medi Sutiyatno Illustrators Subari Wahyudi Santoso Anne Kusumawaty Correspondence on editorial matters and subscriptions for Reinwardtia should be addressed to: HERBARIUM BOGORIENSE, BOTANY DIVISION, RESEARCH CENTER FOR BIOLOGY- INDONESIAN INSTITUTE OF SCIENCES CIBINONG SCIENCE CENTER, JLN. RAYA JAKARTA - BOGOR KM 46, CIBINONG 16911, P.O. Box 25 Cibinong INDONESIA PHONE (+62) 21 8765066; Fax (+62) 21 8765062 E-MAIL: reinwardtia@mail.lipi.go.id 1 2 3 4 1 3 4 4 Cover images: 1. Begonia holosericeoides (female flower and habit) (Begoniaceae; Ardi et al.); 2. Abaxial cuticles of Alseodaphne rhododendropsis (Lauraceae; Nishida & van der Werff); 3. Dipo- dium puspitae, Dipodium purpureum (Orchidaceae; O'Byrne); 4. Agalmyla exannulata, Cyrtandra coccinea var. celebica, Codonoboea kjellbergii (Gesneriaceae; Kartonegoro & Potter). The Editors would like to thanks all reviewers of volume 14(1): Abdulrokhman Kartonegoro - Herbarium Bogoriense, Bogor, Indonesia Altafhusain B. Nadaf - University of Pune, Pune, India Amy Y. Rossman - Systematic Mycology & Microbiology Laboratory USDA-ARS, Beltsville, USA Andre Schuiteman - Royal Botanic Gardens, Kew, UK Ary P. Keim - Herbarium Bogoriense, Bogor, Indonesia Barry Conn - Royal Botanic Gardens National Herbarium of New South Wales, Sydney, Australia Dato' Abdul Latiff Mohamad - Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia Daniel Potter - Department of Plant Sciences, University of California, Davis, California, USA Deby Arifiani - Herbarium Bogoriense, Bogor, Indonesia Ferry J. W. Slik - University of Brunei Darussalam, Brunei Henti H. Rachmat - Conservation and Rehabilitation Research and Development Center, Bogor, Indonesia Ian M. Turner - Royal Botanic Gardens, Kew, UK Iskandar Z. Siregar - Bogor Agricultural University, Bogor, Indonesia Jay H. Bernstein - Kingsborough Community College, Brooklyn, New York, USA Jens G. Rohwer - University of Hamburg, Hamburg, Germany Joan Pereira - SAN Herbarium, Sabah Forestry Department, Sabah, Malaysia Kuswata Kartawinata - Herbarium Bogoriense, Bogor, Indonesia Lars H. Schmidt - University of Copenhagen, Copenhagen, Denmark Mark Hughes - Royal Botanic Gardens, Edinburgh, UK Masahiro Kato - Kyoto University, Kyoto, Japan Nuril Hidayati - Herbarium Bogoriense, Bogor, Indonesia Ong Poh Teck - Forest Research Institute Malaysia, Kepong, Malaysia Peter C. van Welzen - National Herbarium Netherlands, Leiden University Branch, Leiden, Netherlands Reuben Nilus - Sabah Forestry Department, Sabah, Malaysia Rugayah - Herbarium Bogoriense, Bogor, Indonesia Ruth Kiew - Forest Research Institute of Malaysia, Kepong, Malaysia Uwe Braun - Institut fur Biologie Bereich Geobotanik und Botanischer Garten, Halle (Saale), Germany Yasuaki Sato - Osaka-Sangyo University, Osaka, Japan REINWARDTIA Vol 14, No 1, pp: 13 − 17 13 CODONOBOEA (GESNERIACEAE) SECTIONS IN PENINSULAR MALAYSIA Received February 6, 2014; accepted April 7, 2014 LIM CHUNG LU Herbarium, Forest Research Institute Malaysia (FRIM), 52109 Kepong, Selangor, Malaysia. E-mail: limchunglu@frim.gov.my RUTH KIEW Herbarium, Forest Research Institute Malaysia (FRIM), 52109 Kepong, Selangor, Malaysia. ABSTRACT LIM, C. L. & KIEW, R. 2014. Codonoboea (Gesneriaceae) Sections in Peninsular Malaysia. Reinwardtia 14(1): 13 – 17. — Codonoboea is the largest genus of Gesneriaceae in Peninsular Malaysia with 92 species. Nine sections, Boeopsis, Codonoboea, Didymanthus, Glossadenia, Heteroboea, Pectinati, Reptantes, Salicini and Venusti, have been erected based on morphological characters, such as habit, leaf arrangement and shape, type of inflorescence, number of flowers, corolla type and nectary type. Recent molecular phylogenetic study using ITS and trnL-F sequences show that only section Heteroboea is monophyletic. While the sections of Codonoboea can no longer be maintained as formal taxonomic taxa, as informal groupings they are useful in identification and in constructing keys. Key words: Codonoboea, Gesneriaceae, Peninsular Malaysia, section. ABSTRAK LIM, C. L. & KIEW, R. 2014. Seksi Codonoboea (Gesneriaceae) di Semenanjung Malaysia. Reinwardtia 14(1): 13 – 17. — Codonoboea merupakan marga terbesar dalam Gesneriaceae di Semenanjung Malaysia dengan 92 jenis. Sembilan seksi yaitu Boeopsis, Codonoboea, Didymanthus, Glossadenia, Heteroboea, Pectinati, Reptantes, Salicini dan Venusti telah dibuat berdasarkan karakter morfologi, seperti habitus, bentuk dan susunan daun, tipe perbungaan, jumlah bunga, tipe daun mahkota dan tipe nektar. Penelitian filogenetik molekuler terkini dengan menggunakan sekuen ITS dan trnL-F menunjukkan bahwa hanya seksi Heteroboea yang bersifat monofiletik. Sementara itu seksi Codonoboea tidak bisa dipertahankan sebagai taksa taksonomi secara formal, pengelompokan informal seksi ini dapat dimanfaatkan dalam membangun kunci dan dapat pula dimanfaatkan dalam proses identifikasi. Kata kunci: Codonoboea, Gesneriaceae, Semenanjung Malaysia, seksi. INTRODUCTION Gesneriaceae Rich. & Juss. ex DC. (1816), the African violet family, is a diverse family with 140–150 genera and more than 3500 species worldwide (Weber, 2004) widely distributed in the subtropical and tropical regions in the Old and New World. Peninsular Malaysia is the centre of diversity of Codonoboea Ridl. (Table 1) and in Peninsular Malaysia it is the largest and most diverse genus of the Gesneriaceae with 92 species (Kiew & Lim, 2011; Kiew, 2011; Kiew & Sam, 2012; Lim et al., 2013) with still more awaiting new species description followed by Paraboea (24 species) and Ridleyandra (19 species). In the past, the genus was confused with Didymocarpus Wall., Didissandra C. B. Clarke, Paraboea (C. B. Clarke) Ridl., Loxocarpus R. Br., Henckelia Spreng. and Chirita Buch.-Ham. ex D. Don. However, with evidence from the molecular phylogenetic method, Weber et al. (2011) showed conclusively that Codonoboea is a monophyletic group distinct from the genera mentioned above. Table 1. Distribution of Codonoboea species. In Peninsular Malaysia, its species are morpho- logically very diverse ranging in habit (creeping, Area No. species Peninsular Thailand 4 Peninsular Malaysia 92 Sumatra 9 Borneo 27 Batam & Lingga Is. 2 Philippines 2 Sulawesi 2 Moluccas 1 New Guinea 1 mailto:limchunglu@frim.gov.my REINWARDTIA 14 [VOL.14 rosette or erect herbs), in leaf arrangement (alternate or opposite), branching of inflorescence (from much branched panicles to single-flowered cymes) and in flower size, colour and type (pollen or nectar flower). To facilitate the classification of this large and diverse genus, nine sections have been proposed based on one to several morpho- logical characters (Table 2), but even with nine sections there are still a few odd species, such as C. longipes (C. B. Clarke) Kiew, that cannot be assigned to existing sections. In a large and unwieldy genus, sections can serve several functions for example in (a) grouping similar species together can facilitate identification and the construction of keys by dividing a large number of species into smaller units of similar taxa, (b) making it easier to discern phyto- geographic patterns, and (c) it may reflect evolu- tionary trends. However, the problem with similar morphology is that it may also reflect parallel or convergent evolution in adaptation, for example, similar habitat, pollinator or dispersal mechanism. To assess whether the Codonoboea sections are monophyletic, the molecular phylogenetic method was used on a sample of species from Peninsular Malaysia. MATERIALS Twenty seven taxa of Codonoboea were included in the study with representatives from five of the nine sections from Peninsular Malaysia (Table 3). Another four species from Borneo, the Philippines and Sumatra have not been assigned to a section (Middleton et al., 2013). Species from Microchirita (C. B. Clarke) Y. Z. Wang [M. caliginosa (C. B. Clarke) Y. Z. Wang, M. involucrata (Craib) Y. Z. Wang, M. ruthiae Rafidah and M. viola (Ridl.) A. Weber & Rafidah) and Boea [Boea hygrometrica (Bunge) R. Br. and B. philippensis C. B. Clarke) were selected as out- group taxa. Using molecular phylogenetic technique, two regions, i.e. trnL-F intron-spacer region, and the nuclear ITS1-5.8S-ITS2 region were analysed (Lim, 2014). Mapping of sections character-state transitions was carried out in MacClade 4.08 (Maddison & Maddison, 2003). RESULTS AND DISCUSSION Mapping of sections onto a molecular phylo- genetic tree is shown in Fig. 1. From the tree, only section Heteroboea is monophyletic. Section Heteroboea is well-supported with all species sampled forming a clade together with the undescribed Codonoboea sp. nov. 2. Species of this section are caulescent with spirally arranged leaves and large, solitary, trumpet-shaped flowers with a large annular nectary. Phylograms of ITS1- 5.8S-ITS2 region, trnL-F intron-spacer region as well as the combined results of the two regions show that the species in the clade do not differ Table 2. Sections in Codonoboea. Section Publication Salient characters Boeopsis Ridley (1907) J. Roy. As. Soc. Str. Br. 49: 22; Kiew (1992) Gard. Bull. Sing. 44: 39. Rosette plants, small campanulate corolla Codonoboea Kiew (1990) Blumea 35: 167. Epiphyllous flowers Didymanthus Clarke (1883) Monogr. Phan. 5 (1): 87, tab. 10. Erect stem, opposite and well-spaced petiolate leaves Glossadenia Weber & Burtt (1998) Beitr. Biol. Pflanzen 70: 335. Tongue-like nectary Heteroboea Bentham (1876) Genera Plantarum 2: 1022. Spirally arranged leaves, winged peti- ole Pectinati Ridley (1923) Flora of Malay Peninsula 2: 508. Serrate to deeply toothed lamina Reptantes Ridley (1905) J. Roy. As. Soc. Str. Br. 44: 29. Creeping stem, opposite and well- spaced leaves Salicini Ridley (1896) J. Linn. Soc. Bot. 32: 514; Kiew (1992) Gard. Bull. Sing. 44: 41. Small campanulate corolla, narrow lamina Venusti Kiew (1995) Malay. Nat. J. 48: 203. Foliaceous, conspicuous bracts 2014] 15 LIM & KIEW: Codonoboea (Gesneriaceae) sections in Peninsular Malaysia Table 3. Codonoboea species according to the section. Section Species Boeopsis C. anthonyi, C. codonion, C. floribunda, C. heterophylla, C. leiophylla, C. oreophila, C. pumila, C. puncticu- lata, C. rubiginosa Didymanthus C. albomarginata, C. glabrata, C. malayana, Codonoboea sp. nov. 1 Heteroboea C. bombycina, C. crinita, C. curtisii, C. platypus, C. fasciata, Codonoboea sp. nov. 2 Salicini C. salicina, C. salicinoides, C. tiumanica Venusti C. calcarea Not assigned Borneo: C. bakoensis, C. crenata Philippines: C. corrugata Sumatra: C. racemosa Peninsular Malaysia: C. longipes, Codonoboea sp. nov. 3, Codonoboea sp. nov. 4, Codonoboea sp. nov. 5 Fig 1. Mapping of Codonoboea sections on maximum parsimony majority-rule consensus tree based on combined ITS and trnL-F sequences. Number above branches are majority- rule frequencies. REINWARDTIA 16 [VOL.14 much genetically (Lim, 2013). In fact, it is possi- ble that the species concept in this section is too narrow because several species groups, such as the C. crinita (Jack) C. L. Lim, C. platypus (C. B. Clarke) C. L. Lim and C. bombycina (Ridl.) C. L. Lim complex, are difficult to identify, because while the extreme forms are distinct there are many specimens showing intermediate character states. In contrast, section Boeopsis, Didymanthus and Salicini are not monophyletic in the tree. Their morphological similarity is likely therefore to be the result of convergent evolution due to adaption to pollinator or environmental factors. For example, the salient characters of section Salicini are (a) narrow lamina that can be explained as adaptive to their rheophytic habitat and (b) small campanulate corolla as adaptation to the same pollinator (which is as yet unknown). Similarly, the rosette habit of section Boeopsis is an adaptation to their growing on vertical earth banks or rock faces and the trend from the nectar flowers (large trumpet-shaped corolla and large annular nectar) to the flat-faced pollen flower with a minute or no nectary is a trend that appears to have occurred more than once in Codonoboea. CONCLUSIONS The results from the molecular study call into question the taxonomic value of the sections and indeed Kiew & Lim (2011) did not assign species to section for precisely this reason. Although, section Heteroboea is monophyletic, it is firmly nested within the other Codonoboea species sampled, so it makes no sense to maintain giving it formal taxonomic status. Codonoboea is very diverse morphologically and informal subgeneric groups are still useful for a speciose genus in formulating user-friendly keys for identification. Indeed, the only key for all species in Peninsular Malaysia is that of Ridley (1923) and the ease of using it (always relative in the case of Ridley’s keys) is because it is based on these groups that enable rapid identification. Therefore, it is advocated here that while informal groups are useful that they not be given formal taxonomic status and that the nine sections should therefore lapse into synonym. ACKNOWLEDGEMENTS This research was carried out as part of the Flora of Peninsular Malaysia Project (project no. 01-04-01-0000 Khas) at the Forest Research Inst. Malaysia (FRIM), Kepong, funded by the Ministry of Science, Tech- nology and Innovation. In addition, the first author is indebted to the following for financial support: the Systematic Studies of Selected Taxa of Plants Project [Erycibe (Convolvulaceae), Chirita, Loxocarpus, Henckelia sections Bo eopsis and Salicini (Gesneriaceae) and Utricularia (Lentibulariaceae)] in Peninsular Malaysia (GPP-TFBC-1208-001) at FRIM and funded by Malaysian Forestry Research and Development Board (MFRDB), and to the Postgraduate Research Fund (project no. PS170/2008B) at University of Malaya, Kuala Lumpur. We are much indebted to Royal Botanic Garden, Edinburgh (RBGE) as a collaborating institute for the molecular phylogenetic study, especially to M. Möller for guidance and advice in molecular phylogenetic study. We are grateful to the staff of the FRIM Flora Biodiversity Programme for their assistance in the field trips. To curators, keepers and staff of BM, E, K, KEP, KLU, L, SAR, SING and UKMB, we feel so much indebted for the specimen loans and type specimen photographs provided. 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Malay Peninsula 2: 495–547. WEBER, A. 2004. Gesneriaceae. In: KUBITZKI, K. (Ed.) The families and genera of vascular plants 7: 63–158. WEBER, A. & BURTT, B. L. 1998. Remodelling of D i d y m o c a r p u s a n d a s s o c i a t e d g e n e r a (Gesneriaceae). Beitrage zur Biologie der Pflanzen 70: 293–363. WEBER, A., MIDDLETON, D. J., DAVID, J. M., FORREST, A., KIEW, R., LIM, C. L., RAFIDAH, A. R., SONTAG, S., TRIBOUN, P., WEI, Y. G., YAO, T. L. & MÖLLER, M. 2011. Molecular systematics and remodelling of Chirita and associated genera (Gesneriaceae). Taxon 60(3): 767 –790. REINWARDTIA 18 [VOL.14 INSTRUCTION TO AUTHORS Scope. Reinwardtia is a scientific irregular journal on plant taxonomy, plant ecology and ethnobotany published in December. Manuscript intended for a publication should be written in English. Titles. Titles should be brief, informative and followed by author's name and mailing address in one- paragraphed. Abstract. English abstract followed by Indonesian abstract of not more than 250 words. Keywords should be given below each abstract. Manuscript. Manuscript is original paper and represent an article which has not been published in any other journal or proceedings. The manuscript of no more than 200 pages by using Times New Roman 11, MS Word for Windows of A4 with double spacing, submitted to the editor through . New paragraph should be indented in by 5 characters. For the style of presentation, authors should follow the latest issue of Reinwardtia very closely. Author(s) should send the preferred running title of the article submitted. Every manuscript will be sent to two blind reviewers. Identification key. Taxonomic identification key should be prepared using the aligned couplet type. Nomenclature. Strict adherence to the International Code of Botanical Nomenclature is observed, so that taxonomic and nomenclatural novelties should be clearly shown. English description for new taxon proposed should be provided and the herbaria where the type specimens area deposited should be presented. Name of taxon in taxonomic treatment should be presented in the long form that is name of taxon, author's name, year of publication, abbreviated journal or book title, volume, number and page. Map/line drawing illustration/photograph. Map, line drawing illustration, or photograph preferably should be prepared in landscape presentation to occupy two columns. Illustration must be submitted as original art accompanying, but separated from the manuscript. The illustration should be saved in JPG or GIF format at least 350 pixels. Legends or illustration must be submitted separately at the end of the manuscript. References. Bibliography, list of literature cited or references follow the Harvard system as the following examples. Journal : KRAENZLIN, F. 1913. Cyrtandraceae novae Philippinenses I. Philipp. J. Sci. 8: 163-179. MAYER, V., MOLLER, ML, PERRET, M. & WEBER, A. 2003. Phylogenetic position and generic differentiation of Epithemateae (Gesneriaceae) inferred from plastid DNA sequence data. American J. Bot. 90: 321-329. Proceedings :TEMU, S. T. 1995. Peranan tumbuhan dan ternak dalam upacara adat "Djoka Dju" pada suku Lio, Ende, Flores, Nusa Tenggara Timur. In: NASUTION, E. (Ed.). Presiding Seminar dan Lokakarya Nasional Etnobotani II. LIP1 & Perpustakaan Nasional: 263-268. (In Indonesian). SIMBOLON, H. & MIRMANTO, E. 2000. Checklist of plant species in the peat swamp forests of Central Kalimantan, Indonesia. In: IWAKUMA et al. (Eds.) Proceedings of the International Symposium on: Tropical Peatlands. Pp. 179-190. Book : RIDLEY, H. N. 1923. Flora of the Malay Peninsula 2. L. Reeve & Co. Ltd, London. Part of Book : BENTHAM, G. 1876. Gesneriaceae. In: BENTHAM, G. & HOOKER, J. D. Genera plantarum 2. Lovell Reeve & Co., London. Pp. 990-1025. Thesis : BAIRD, L. 2002. A Grammar of Keo: An Austronesian language of East Nusantara. Australian National University, Canberra. [PhD. Thesis]. Website : http://www.nationaalherbarium.n1/fmcollectors/k/Kostermans AJGH.htm). Accessed 15 February 2012. Reinwardtia Published by Herbarium Bogoriense, Botany Division, Research Center for Biology, Indonesian Institute of Sciences Address: Jin. Raya Jakarta-Bogor Km. 46 Cibinong 16911, P.O. Box 25 Cibinong Telp. (+ 62) 21 8765066; Fax (+62) 21 8765062 E-mail: reinwardtia@mail.lipi.go.id REINWARDTIA Author Agreement Form Title of article Name of Author(s) : I/We hereby declare that: • My/Our manuscript was based on my/our original work. • It was not published or submitted to other journal for publication. • I/we agree to publish my/our manuscript and the copyright of this article is owned by Reinwardtia. • We have obtained written permission from copyright owners for any excerpts from copyrighted works that are included and have credited the sources in our article. 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