Studies in Malesian Gentianaceae, V. The Fagraea complex in Borneo: New generic assignments and recombinations
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1 Gardens Bulletin Singapore 65(2): Studies in Malesian Gentianaceae, V. The Fagraea complex in Borneo: New generic assignments and recombinations K.M. Wong 1, M. Sugumaran 2 and J.B. Sugau 3 1 Herbarium, Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore wkm2000@gmail.com (corresponding author) 2 Rimba Ilmu Botanic Garden, Institute of Biological Sciences, University of Malaya, Kuala Lumpur, Malaysia maran@um.edu.my 3 Forest Research Centre, Forestry Department, P.O. Box 1407, Sandakan, Sabah, Malaysia John.Sugau@sabah.gov.my ABSTRACT. A new classification of the Fagraea complex (Gentianaceae) based on recent molecular studies and taxonomic considerations now recognises the genera Cyrtophyllum, Fagraea in the strict sense, Limahlania, Picrophloeus, and Utania. The Bornean species of this complex are listed following the new perspective and the remaining necessary new combinations (seven, in Utania) are made. Keywords. Borneo, Cyrtophyllum, Fagraea, Gentianaceae, Limahlania, Picrophloeus, Utania Introduction Wong & Sugau (1996) revised the Bornean species of the Fagraea Thunb. complex. In their work, they reviewed a number of species limits that had previously been masked by the broader concepts adopted by Leenhouts (1962b), and revived a number of names that had been enumerated or described by Cammerloher (1923) and others. In addition, a number of species were also newly recognised. Subsequently, Sugumaran & Wong (2012) demonstrated good support from molecular evidence for reasonably clear lineages that could be identified with genera within the Fagraea Complex. Their new classification recognised Cyrtophyllum Reinw., Fagraea (in the strict sense), Limahlania K.M.Wong & M.Sugumaran, Picrophloeus Blume, and Utania G.Don. Genera and sections Of the three sections of Fagraea recognised by Leenhouts (1962b), F. section Cyrtophyllum (Reinw.) Blume (Blume 1838) is considered to have included two distinct groups, which Wong & Sugau (1996) simply referred to as the F. fragrans complex (axillary inflorescences) and F. elliptica complex (terminal inflorescences). There are further differences (e.g., plant architecture) and these groups are now recognised as
2 236 Gard. Bull. Singapore 65(2) 2013 Cyrtophyllum (five species, revised by Wong & Sugumaran 2012b) and Picrophloeus (four species: Wong 2012). All species of F. section Fagraea (sensu Leenhouts 1962b) remain in Fagraea sensu stricto, except for F. crenulata Maingay ex Clarke which was transferred to the new monospecific genus, Limahlania K.M.Wong & M.Sugumaran (Wong & Sugumaran 2012a). The species of F. section Racemosae Benth. (Bentham 1856), with a rather distinct inflorescence structure in which the elongate panicle bears condensed branches that are much shorter than the rachis (in the other genera of this complex, the basal cyme branches are similar to the rachis in length), are transferred to the genus Utania G.Don. Taxonomic assignment of Bornean species in the Fagraea Complex Here we outline the generic designation of 43 known Bornean species from the Fagraea Complex according to this new classification. Species distributions have been given in the Bornean enumeration of Wong & Sugau (1996) except that of the newly described Cyrtophyllum minutiflorum K.M.Wong, in Wong & Sugumaran (2012b). CYRTOPHYLLUM (4 spp.) Fagraea caudata Ridl. = Cyrtophyllum caudatum (Ridl.) K.M.Wong, Gard. Bull. Singapore 64: 499 (2012). Fagraea fragrans Roxb. = Cyrtophyllum fragrans (Roxb.) DC., Prodromus 9: 31 (1845). Fagraea gigantea Ridl. = Cyrtophyllum giganteum (Ridl.) Ridl., Fl. Malay Penins. 2: 421 (1923). Cyrtophyllum minutiflorum K.M.Wong, Gard. Bull. Singapore 64: 507 (2012). FAGRAEA (27 spp.) Fagraea acutibracteata K.M.Wong & Sugau, Sandakania 8: 49 (1996). Fagraea auriculata Jack, Malayan Misc. 2, 7: 82 (1822). Fagraea blumei G.Don, Gen. Hist. 4: 69 (1837) (in Don 1838). Fagraea borneensis Scheff., Flora 52: 309 (1869). Fagraea carnosa Jack, Malayan Misc. 2, 7: 81 (1822). Fagraea dulitensis K.M.Wong & Sugau, Sandakania 8: 59 (1996). Fagraea floribunda K.M.Wong & Sugau, Sandakania 8: 62 (1996). Fagraea havilandii K.M.Wong & Sugau, Sandakania 8: 64 (1996). Fagraea iliasii K.M.Wong & Sugau, Sandakania 8: 66 (1996). Fagraea involucrata Merrill, J. Straits Branch Roy. Asiat. Soc. 77: 233 (1917). Fagraea kalimantanensis (Leenh.) K.M.Wong & Sugau, Sandakania 8: 69 (1996). Fagraea kinabaluensis K.M.Wong & Sugau, Sandakania 8: 69 (1996). Fagraea kuminii K.M.Wong & Sugau, Sandakania 8: 71 (1996).
3 Fagraea complex in Borneo: name changes, new combinations 237 Fagraea littoralis Blume var. borneensis K.M.Wong & Sugau, Sandakania 8: 73 (1996). Fagraea longipetiolata K.M.Wong & Sugau, Sandakania 8: 75 (1996). Fagraea macroscypha Baker, Kew Bull.: 25 (1896). Fagraea megalantha K.M.Wong & Sugau, Sandakania 8: 77 (1996). Fagraea oblonga King & Gamble, J. Asiat. Soc. Bengal 74, 2: 612 (1908). Fagraea oreophila K.M.Wong & Sugau, Sandakania 8: 78 (1996). Fagraea rarissima K.M.Wong & Sugau, Sandakania 8: 82 (1996). Fagraea renae K.M.Wong & Sugau, Sandakania 8: 82 (1996). Fagraea resinosa Leenh., Bull. Jard. Bot. État Bruxelles 32: 429 (1962a). Fagraea ridleyi King & Gamble, J. Asiat. Soc. Bengal 74, 2: 612 (1908). Fagraea splendens Blume, Mus. Bot. 1: 168 (1850). Fagraea stonei K.M.Wong & Sugau, Sandakania 8: 88 (1996). Fagraea tuyukii K.M.Wong & Sugau, Sandakania 8: 90 (1996). Fagraea sp. A., K.M.Wong & Sugau, Sandakania 8: 92 (1996). LIMAHLANIA (1 sp.) Fagraea crenulata Maingay ex Clarke in Hooker f., Fl. Brit. India 4 (1883) 83 = Limahlania crenulata (Maingay ex Clarke) K.M.Wong & M.Sugumaran, Gard. Bull. Singapore 64: 492 (2012). PICROPHLOEUS (4 spp.) Fagraea belukar K.M.Wong & Sugau = Picrophloeus belukar (K.M.Wong & Sugau) K.M.Wong, Gard. Bull. Singapore 64: 513 (2012). Fagraea collina K.M.Wong & Sugau = Picrophloeus collinus (K.M.Wong & Sugau) K.M.Wong, Gard. Bull. Singapore 64: 514 (2012). Fagraea elliptica auctt. incl. Wong & Sugau, Sandakania 8 (1996) 21 = Picrophloeus javanensis Blume, Bijdr. Fl. Ned. Ind. 16: 1020 (1826). Fagraea rugulosa K.M.Wong & Sugau = Picrophloeus rugulosus (K.M.Wong & Sugau) K.M.Wong, Gard. Bull. Singapore 64: 520 (2012). UTANIA (7 spp.) Fagraea cuspidata Blume, Mus. Bot. 1: 170 (1850); Wong & Sugau, Sandakania 8: 28, f. 9 (1996) = Utania cuspidata (Blume) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Blume, s.n., Borneo, Tanjong Java (iso L). Fagraea montana K.M.Wong & Sugau, Sandakania 8: 31, f. 10 (1996) = Utania montana (K.M.Wong & Sugau) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Aban SAN 50722, Sabah, Ranau, Mamut, copper mining area (holo SAN; iso K, L). Fagraea philippinensis K.M.Wong & Sugau, Sandakania 8: 35, f. 12 (1996) = Utania philippinensis (K.M.Wong & Sugau) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Ahern s Coll. For. Bur. 3270, Philippines, Luzon, Rizal (holo K).
4 238 Gard. Bull. Singapore 65(2) 2013 Fagraea spicata Baker, Kew Bull. (1896) 25; Wong & Sugau, Sandakania 8: 38 (1996) = Utania spicata (Baker) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Creagh, s.n., British North Borneo, east coast (holo K). Fagraea stenophylla Becc. ex Merrill, J. Straits Branch Roy. Asiat. Soc. 77: 236 (1917); Wong & Sugau, Sandakania 8: 38, f. 13 (1996) = Utania stenophylla (Becc. ex Merrill) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Native Coll. Bur. Sci. 2828, Sarawak, Upper Baram, Selungo (iso A, UC). Fagraea teysmannii Cammerloher, Bull. Jard. Bot. Buitenzorg, ser. 3, 5: 314, f. 3 (1923); Wong & Sugau, Sandakania 8 (1996) 40 = Utania teysmannii (Cammerloher) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Teysmann, s.n., Karimata, Soengai Tajan (lecto L; chosen by Wong & Sugau 1996). Fagraea volubilis Wall. in Roxb., Fl. Ind. 2: 36 (1824); Wong & Sugau, Sandakania 8: 40 (1996) = Utania volubilis (Wall.) M.Sugumaran, Gard. Bull. Singapore 64: 493 (2012). TYPE: Jack, E. Bencoolen (sheet marked E. Bencoolen on bottom left) (holo K-W). Fagraea volubilis var. microcalyx K.M.Wong & Sugau, Sandakania 8: 43, f. 14 (1996) = Utania volubilis var. microcalyx (K.M.Wong & Sugau) K.M.Wong, M.Sugumaran & Sugau, comb. nov. TYPE: Burley, Tukirin et al. 372, Kalimantan, 5 km NE of Haruwu village (holo KEP; iso A, BO). ACKNOWLEDGEMENTS. We thank the Keepers and Curators of the A, BO, K, KEP, KLU, L, NY, US, SAN, SAR and SING herbaria for permission to consult reference materials and loans of specimens. References Baker, J.G. (1896) Decades Kewensis. Plantarum novarum in Herbario Horti Regii Conservatarum. Decas XXIII XXV. Kew Bull. (1896): Bentham, G. (1856) Notes on Loganiaceae. J. Linn. Soc., Bot. 1: Blume, C.L. (1826) Bijdragen tot de Flora van Nederlandsch Indië. Part 14, pp ; Part 16, pp Batavia: Lands Drukkerij. Blume, C.L. (1838) Rumphia. Vol pp., 105 pl. Amstelodam: C.G. Sulpke. Blume, C.L. (1850) Museum Botanicum Lugduno-Batavum. Vol pp., 59 fig. Lugduni- Batavorum: E.J. Brill. Cammerloher, H. (1923) Die Loganiaceen und Buddleiaceen Niederländisch-Indiens. Bull. Jard. Bot. Buitenzorg, ser. III 5(4): Candolle, A.L.P.P. de (1845) Prodromus Systematis Naturalis Regni Vegetabilis. Vol pp. Paris: Sumptibus Sociorum Treuttel et Würtz. Don, G. (1838) A General System of Gardening and Botany. Vol. 4. viii pp. London: J.G. & F. Rivington, etc.
5 Fagraea complex in Borneo: name changes, new combinations 239 Hooker, J.D. (1883) The Flora of British India. Vol pp. London: L. Reeve & Co. Jack, W. (1822) Descriptions of Malayan Plants. Malayan Miscellanies 2(7). iii + 96 pp. Bencoolen: Sumatran Mission Press. King, G. & Gamble, J.S. (1908) Materials for a Flora of the Malayan Peninsula. J. Asiat. Soc. Bengal 74(2): Leenhouts, P.W. (1962a) Notes on some species of Fagraea Thunb. Bull. Jard. Bot. État Bruxelles 32: Leenhouts, P.W. (1962b) Loganiaceae. In: Steenis, C.G.G.J. van (ed) Flora Malesiana. Series 1, vol. 6. Pp Groningen: Wolters-Noordhoff Publishing. Merrill, E.D. (1917) Alabastra Borneensia. J. Straits Branch Roy. Asiat. Soc. 77: Ridley, H.N. (1923) The Flora of the Malay Peninsula. Vol. 2. vi pp. London L. Reeve & Co. Roxburgh, W. (1824) Flora indica; or, Descriptions of Indian plants. Vol pp. Serampore: Mission Press. Scheffer, R.H.C.C. (1869) Observationes phytographicae. Flora; oder, (allgemeine) botanische Zeitung 52: Sugumaran, M. & Wong, K.M. (2012) Studies in Malesian Gentianaceae, I. Fagraea sensu lato complex genus or several genera? A molecular phylogenetic study. Gard. Bull. Singapore 64(2): Wong, K.M. (2012) Studies in Malesian Gentianaceae, IV. A revision of Picrophloeus. Gard. Bull. Singapore 64(2): Wong, K.M. & Sugau, J.B. (1996) A revision of Fagraea (Loganiaceae) in Borneo, with notes on related Malesian species and 21 new species. Sandakania 8: Wong, K.M. & Sugumaran, M. (2012a) Studies in Malesian Gentianaceae, II. A taxonomic framework for the Fagraea complex, including the new genus Limahlania. Gard. Bull. Singapore 64(2): Wong, K.M. & Sugumaran, M. (2012b) Studies in Malesian Gentianaceae, III. Cyrtophyllum reapplied to the Fagraea fragrans alliance. Gard. Bull. Singapore 64(2):
Studies in Malesian Gentianaceae II: A taxonomic framework for the Fagraea complex, including the new genus Limahlania
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