Thai Forest Bulletin 31

THAI FOREST BULLETIN
(BOTANY) NO. 31
ISSN 04953843
THE FOREST HERBARIUM NATIONAL PARK, WILDLIFE AND PLANT CONSERVATION DEPARTMENT BANGKOK, THAILAND DECEMBER 2003
CONTENTS
Ceropegia hirsuta (Asclepiadaceae), a new record for Thailand.. Tanucha Boonjaras & Obchan Thaithong Notes on two Ixora species, new records for Thailand Voradol Chamchumroon The Rubiaceae of Ko Chang, south-eastern Thailand.... Voradol Chamchumroon & Christian Puff Two new species of Neohouzeoua (Gramineae-Bambusoideae) from Thailand and Myanmar. Soejatmi Dransfield, Rungnapar Pattanavibool & Sarawood Sungkaew A new description of Diospyros coaetanea (Ebenaceae)... Sutee Duangjai & Chamlong Phengklai A new species of Spatholirion (Commelinaceae) from Thailand and further notes on S. ornatum..Kai Larsen & Supee Saksuwan Larsen Notes on Clerodendrum (Lamiaceae). Charan Leeratiwong & Pranom Chantaranothai Matoniaceae (Pteridophyta) - a new family record for Thailand Stuart Lindsay, Somran Suddee, David J. Middleton & Rachun Pooma An acount of the Plantaginaceae of ThailandJohn Parnell Thai Rubiaceae with hooks and thorns... Christian Puff & Voradol Chamchumroon Nonindigenous Rubiaceae grown in Thailand.. Christian Puff & Voradol Chamchumroon Notes on the genus Alpinia (Zingiberaceae) in Thailand... Surapon Saensouk, Pranom Chantaranothai & Kai Larsen Karyology of Jatropha (Euphorbiaceae) in Thailand. Puangpaka Soontornchainaksaeng & Thaya Jenjittikul Cytogenetic studies and taxonomic considerations in some taxa of Mallotus (Euphorbiaceae) in Thailand.. Puangpaka Soontornchainaksaeng, Pranom Chantaranothai & Chadapron Senakun Capparis sikkimensis Kurz subsp. yunnanensis (Capparaceae), a new record for Thailand. Prachaya Srisanga, Chusie Trisonthi & Kongkanda Chayamarit Radermachera eberhardtii (Bignoniaceae), a new record for Thailand. Prachaya Srisanga, Chusie Trisonthi & Thawatchai Santisuk Rhodoleia (Hamamelidaceae), a new generic record for Thailand.... Somran Suddee & David J. Middleton A preliminary study of Gonystylaceae in Thailand.... Chawalit Niyomdham & Metinee Tarumatsawat The Genus Porandra (Commelinaceae) in Thailand . Thaweesak Thitimetharoch, Pranom Chantaranothai, & Robert B. Faden 1 7 13
27 34 39 44 47 53 67 77 97 107
115
125 131 134 136 143
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THAI FOR. BULL. (BOT.) 31: 16. 2003.
Ceropegia hirsuta (Asclepiadaceae), a new record for Thailand

TANUCHA BOONJARAS *& OBCHANT THAITHONG **
ABSTRACT. Ceropegia hirsuta, a new record for Thailand, is described and illustrated.
There are approximately 170 species of the genus Ceropegia distributed in Southeast Asia, India, Madagascar, Tropical Arabia, the Canary Islands, Africa, New Guinea and Northern Australia. Four species were previously reported in Thailand (Kerr, 1951), namely C. arnottiana Wight, C. jucunda Kerr (= C. lucida Wall.), C. siamensis Kerr (= C. monticola W.W. Sm.) and C. sootepensis Craib. As there are no further reports of this genus in Thailand, the discovery of Ceropegia hirsuta Wight & Arn. at Pha Taem National Park in June 2001 represents a new record.
Ceropegia hirsuta Wight & Arn., in Wight, Contrib. 30. 1834; Hook.f., Fl. Brit. India 4: 71. 1883; Gamble, Fl. Pres. Madras 5: 859. 1923; H. Huber, Mem. Soc. Broter. 12: 63. 1957; Stevens in Saldanha & Nicolson, Fl. Hassan Distr.: 444. 1976; Ansari, Fasc. Fl. India 16: 16. 1984. Type: India, Wight Cat. no. 1510 (holotype K). Fig. 1, 2. Herbs or twining herbs; subterranean tuber not seen. Stem ca. 3 mm in diam., pubescent. Leaves opposite, decussate, elliptic to ovate ca. 6.5 by 3 cm, pubescent on both sides, apex acute, base rounded, margin entire, ciliate; petiole ca. 2 cm long, pubescent. Cymes axillary, few-flowered; peduncle up to 1 cm long, pubescent. Flowers large, pedicel ca. 1 cm long, pubescent. Sepals 5, free, linear, 56 mm long with gland at base, glabrous inside, pubescent outside. Corolla urceolate, 45 cm long, corolla tube ca. 3 cm long, curved, inflated at base, funnel-shaped above, sparsely hairy inside, puberulent outside, creamy yellow with purplish red spots; corolla lobes, broadly ovate, 78 by 44.5 mm, hairy inside, margin hairy, folded back, forming plate-like hairy keels inside; apex fused, forming an almost globose dome. Corona in 2 rows; outer corona 5, bifid, deltoid, hairy, inner corona 5, linear, erect, tip hooked, sparsely hairy. Pollinaria 5, pollinium ascending 0.4 mm long, yellow with deltoid pellucid tip, corpusculum spathulate, reddish-brown. Ovaries 2, ca. 2 mm long, marginal placentation, ovules numerous, style very short. Fruits and seeds not seen. Thailand. EASTERN: Ubon Ratchathani [Pha Taem National Park, T. Boonjaras 10 (BCU)]. Distribution. India.
THAI FOREST BULLETIN (BOTANY) 31
Department of Pharmaceutical Botany, Faculty of Pharmacy, Mahidol University, Bangkok 10400, Thailand. Department of Botany, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand.
Ecology. On sandy soil in deciduous dipterocarp forest, 250300 m alt. Flowering JuneAugust. Vernacular. Khreua I Thao () (Ubon Ratchathani). Uses. The whole plant is used as an analgesic. Note. Ceropegia hirsuta is distinguished by the hooked tip of the inner corona, and is the only Asian species with this character. ACKNOWLEDGEMENTS This work was supported by National Research Council of Thailand. Thanks are due to Dr Somran Suddee and Dr Chumpol Khunwasi for valuable suggestions. We also thank Mr Sahut Chantana-orrapint for the photographs. REFERENCE Kerr, A.F.G. 1951. Asclepiadaceae. In: Pendleton, R.L. (ed.), Fl. Siam. Enum. 3(1): 151.
CEROPEGIA HIRSUTA (ASCLEPIADACEAE), A NEW RECORD FOR THAILAND (T. BOONJARAS & O. THAITHONG)
1 mm 5 mm
0.1 mm
1 mm
Figure 1. Ceropegia hirsuta Wight & Arn. A. habit; B. calyx and ovary; C. flower; D. pollinaria, 1. proximal 1 cm view, 2. distal view; E. corona. All from T. Boonjaras 10 (BCU). Drawn by T. Boonjaras. 1 mm
CEROPEGIA HIRSUTA (ASCLEPIADACEAE), A NEW RECORD FOR THAILAND (T. BOONJARAS & O. THAITHONG)
B
Figure 2. Ceropegia hirsuta Wight & Arn. A. habit; B. flower.
THAI FOR. BULL. (BOT.) 31: 712. 2003.
Notes on two Ixora species, new records for Thailand

VORADOL CHAMCHUMROON*
ABTRACT. During a revision of Ixora for the Flora of Thailand, two new records were found for Thailand. Ixora henryi, previously known only from China, is reported from northern Thailand. Ixora cambodiana is recorded from near the border between Thailand and Cambodia. Descriptions and illustrations of the species are provided.
Ixora L. comprises ca. 300 species from Africa and Asia (Mabberley, 1987). There are 20 species in Peninsular Malaysia, occurring in both lowland and upland areas (Corner, 1941). Craib (1934) listed ca. 38 species and seven varieties in Florae Siamensis Enumeratio. Boonbundral (1978) carried out a preliminary study of the genus in Thailand, describing 23 species and three varieties. Whilst revising the genus for the Flora of Thailand the following new records have come to light. Ixora cambodiana Pit., Fl. Indo-Chine 3: 320. 1923. Types: Cambodia, without locality, Jullien s.n. (syntype P); Vietnam, Co-phah, between Hanoi and Bac-ninh, Balansa s.n. (syntype P). Figs. 1, 3. Shrub 3 m high; branchlets glabrous. Leaves elliptic, oblong or oblong-obovate; petiole 0.40.9 cm long, glabrous; blades elliptic-oblong, base cuneate, apex broadly acuminate, 10.512 by 33.1 cm, herbaceous, with 68 pairs of lateral nerves; midrib prominent underneath. Stipules with sheaths 0.40.6 mm long, cuspidate, glabrous outside; awn 0.10.2 cm long. Inflorescence erect, lax, corymbiform, articulate, 89 cm wide and 68 cm long, the inflorescence-supporting leaf pairs with smaller, elliptic blades, (0.7)1 by 0.20.4 cm, peduncle 14 cm long; axes, pedicels and calyces densely covered with short spreading hairs; central first order axes 4 cm long, lateral first order axes 56 cm long; first order bracts without the stipular parts, the latter rather loose, shortly-hairy. Ultimate flower triads with flowers pedicellate; pedicels 0.10.2 cm long, the pedicel of the central flower shorter than the pedicels of the lateral ones; bracteoles present on most pedicels, opposite at the base of the ovary, narrowly triangular to filiform, covered with spreading hairs outside and with colleters inside, 0.51.2 by 0.2 cm. Flowers fragrant. Calyx usually densely hairy, the hairs short; tube 0.20.4 cm long; lobes triangular with acute tips, 0.50.8 cm long. Corolla white or pale pinkish, tube 2.53.5 cm long; lobes 0.60.8 by 0.2 cm, ovate, obtuse or rounded, glabrous or sparely ciliate at base. Anthers pale orange, opening by means of longitudinal slits. Style exserted, 3 mm long; stigma 0.40.5 mm long, pale orange. Ovary 0.10.3 cm long. Fruits globose, 12 cm in diam. Seeds usually 0.70.8 by 0.81 cm, in a 12-seeded drupe, semi-globose with one face flattened-convex.
*The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
Thailand. NORTH-EASTERN: Mukdahan [Dong Bang-I, 16 May 1930, Kerr 21489 (AAU, K); EASTERN: Buri Ram [Khao Phanom Dong Rak, 5 May 2002 Chamchumroon 1472 (BKF)]. Distribution. Thailand, Cambodia, Vietnam. Ecology. Dry evergreen forest. Phenology. Flowering AprilMay. Note. Chamchumroon 1472 is in cultivation at the Rubiaceae garden in Dong Fa Haun Botanic Gardens, Ubon Ratchatani Province. Ixora henryi H. Lv., Fedde, Repert. 13: 178. 1914; Pitard, Fl. Indo-Chine 3: 324. 1923. Types: China, Yunnan, A. Henry 11637 (syntype K!); China, Guizhou, Lou-fou, March 1909, Cavalerie 3496 (syntype K!). Figs. 2, 3. Shrub 23 m high; branchlets glabrous. Leaves opposite, simple, entire, petiolate; petiole 0.40.6 cm long, glabrous; blades elliptic-oblong, base cuneate, apex broadly acuminate, 8.512(18) by 23.1(5) cm, herbaceous, with 68 pairs of lateral nerves; midrib prominent underneath; nerves in 810 pairs. Stipules with sheaths 0.81 cm, cuspidate, glabrous outside; awn 12.5 cm long. Inflorescence sessile, articulate, 2.53.5 cm wide and 23.5 cm long; axes, pedicels and calyces glabrous; central first order axis 0.51 cm long, lateral first order axes 23 cm long; first order bracts with the stipular parts the latter rather loose. Ultimate flower triads with flowers pedicellate; pedicels 0.20.3 cm long or absent, the pedicel of the central flower shorter than the pedicels of the lateral ones; bracteoles present on most pedicels, opposite at the base of the ovary, narrowly triangular to filiform, glabrous outside and with colleters inside, 0.50.9 by 0.10.2 cm. Flowers solitary, fragrant. Calyx tube 24 mm long, lobes triangular with ovate tips, 23 mm long. Corolla white or pale pinkish, tube 2.53.5 cm long; lobes 11.75 by 0.6 cm, ovate, obtuse or rounded, glabrous. Anthers pale orange, opening by means of longitudinal slits. Style exserted, 35 mm long; stigma 0.30.4 mm long, pale orange. Ovary 1.2 mm long. Fruits globose, 78 mm in diam., greenish to red to black. Seeds usually 56 by 5 mm, in 12-seeded drupe; seeds semi-globose with one face flattened-convex. Thailand. NORTHERN: Chiang Rai [Doi Langka, 27 Dec. 1965, Iwatsuki & Fukuoka T-3642 (BKF)]; Lampang [Jae Sawn, 2 March 1997, Maxwell 97167 (BKF)]; Nan [Doi Phu Kha, 27 Feb. 1921, Kerr 4943 (AAU, BK, K); Doi Phu Kha, 21 Nov. 1993, Larsen, S. S. Larsen, Norgaard, Pharsen, Puudjaa & Uerchirakan 44693 (AAU); Doi Phu Kha, 1 Nov. 1995, Pooma 1258 (BKF); Doi Phu Kha, 7 Dec. 1998, Srisanga 392, (QBG); Doi Phu Kha, 12 Feb. 1999, Srisanga 498 (QBG); Doi Phu Kha, 3 Dec. 1999, Srisanga 1206 (QBG); Doi Phu Kha, 12 Jan. 2000, Srisanga 1269 (QBG); Doi Phu Kha, 11 Nov. 2000, Srisanga 1745, (QBG); Doi Phu Kha, 22 Aug. 2001, Srisanga 2040 (QBG); Doi Phu Kha, 22 Aug. 2001, Srisanga & Maknoi 2040 (QBG); Doi Phu Kha, 13 Dec. 1990, Tirvengadum, K. & S. Larsen, Puangpen & Uerchirakan 2010, (AAU)]; NORTH-EASTERN: Loei [Phu Rua, 5 March 1993, Chantaranothai, Middleton, Parnell & Simpson 1101 (K)]; SOUTH-WESTERN: Kanchanaburi [Khao Lieuw Long, 1 April 1968, Beusekom & Phengkhlai 271 (AAU, BKF, C)]. Fig. 3.
NOTES ON TWO IXORA SPECIES, NEW RECORDS FOR THAILAND (V. CHAMCHUMROON)
Distribution. China (Guizhou, Yunnan). Ecology. Along streams in hill evergreen forest. Phenology. Flowering NovemberApril, fruiting AprilJuly. ACKNOWLEDGEMENTS I wish to thank DANIDA for a Scholarship which supported my Ph.D. studies at Kasetsart University and visits to Aarhus Herbarium and Copenhagen Herbarium and the AD (Austrian Academic Exchange) for a scholarship which made it possible to carry out part of the work at the Institute of Botany, University of Vienna, Austria. I would like to thank my supervisors Dr Srunya Vajarothaya, Dr Kongkanda Chayamarit, Dr Lily Kaveeta, and Dr Christian Puff for their valuable suggestions. Thanks are also due to my colleagues at BKF, especially Dr Rachun Pooma, Kanlaya Phattarahirankanok and Narong Koonkhunthod for their help in various ways, especially during fieldwork. REFERENCES Boonbundral, S. 1978. A primary on taxonomy of the genus Ixora in Thailand. M.S. thesis, Chulalongkorn University, Bangkok. 92p. (in Thai). Corner, E.J.H. 1941. Notes on the systematy and distribution of Malayan Phanerogams, IV: Ixora. Gard. Bull. Str. Settlem. 11: 177235. Craib, W.G. 1934. Flora Siamensis Enumeratio 2(2): 147234. Mabberley, D.J. 1987. The Plant Book. Cambridge University Press, Cambridge. 858 p.
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A C D E
Figure 1. Ixora cambodiana Pit.: A. a twig with terminal articulated inflorescence (x); B. stipule with branch (x1); C. petal (x1.5); D. a portion of the inflorescence (x3); E. flower (x1.5). All from Chamchumroon 1472 (BKF). Drawn by V. Chamchumroon.
NOTES ON TWO IXORA SPECIES, NEW RECORDS FOR THAILAND (V. CHAMCHUMROON)
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A E B C
Figure 2. Ixora henryi H. Lv.: A. a twig with terminal articulated inflorescence (x1); B. flower (x1); C. petal (x1); D. a portion of the inflorescence (x2); E. stipule with branch (x1). All from Maxwell 97167 (BKF). Drawn by V. Chamchumroon.
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Ixora cambodiana Pit.
Ixora henryi H. Lv.
Figure 3. Distribution of Ixora cambodiana Pit. and I. henryi H. Lv. in Thailand.
THAI FOR. BULL (BOT.) 31: 1326. 2003.
The Rubiaceae of Ko Chang, south-eastern Thailand

VORADOL CHAMCHUMROON* & CHRISTIAN PUFF**
ABSTRACT. Ko Chang is the second largest island of Thailand. Its Rubiaceae were first treated by K. Schumann in Schmidts Flora of Koh Chang. He recognised 55 species in 27 genera. Based on herbarium and field work, 64 taxa belonging to 33 genera are currently recorded. Although the numbers of taxa in Schumann's treatment and ours do not widely differ, there are major changes in status of the taxa. Eight of the 27 generic names in Schumanns treatment are no longer applicable; only 20 of the 57 taxa names are still in use. Of the 10 species newly described in the Flora of Koh Chang, only one is a good species; none of the new combinations have been maintained. Fifteen taxa have been newly recorded since Schumanns treatment. The relatively large proportion of rubiaceous taxa showing a Ko Chang (or general south-eastern Thailand)/ peninsular Thailand disjunction is remarkable.
INTRODUCTION Ko Chang, the second largest island of Thailand, is situated in Trat Province, in the south-eastern region of Thailand, close to the border with Cambodia (see map, Fig. 1). It is the core part of Mu Ko Chang National Park which became Thailands 45th National Park in 1982. The Rubiaceae, with around 100 genera and over 500 species and infrasepcific taxa, is one of the largest angiosperm families in the Flora of Thailand (Puff, unpublished). They are well represented on Ko Chang as trees, shrubs, climbers or herbs, occurring in a variety of habitats. The first treatment of the family for the island was written by Schumann (1902), at the request of J. Schmidt, editor of Flora of Koh Chang. Duplicates of Schmidts collections were sent to Berlin for Schumann to work on. Although Schumann had a profound knowledge of Rubiaceae, his treatment, which recognised 55 species in 27 genera, had several shortcomings. The main reason was the fact that the Berlin Herbarium (B) lacked sufficient material from Thailand and its surroundings for comparison with the Ko Chang specimens sent to Schumann. Thus he was unable to correctly match numerous collections and subsequently (in part, unjustifiably) described several new taxa and came up with some unusual identifications (see below for details). Work on the family subsequent to Schumanns treatment resulted in major taxonomic realignments of certain taxa. An example is the tribe Gardenieae, where all Asiatic species of Randia are now placed into various other genera (see below). Moreover, collecting activities during recent decades have revealed the presence of a considerable number of rubiaceous taxa not seen by Schumann. For these reasons, an
*The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand. **Institute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna, Austria.
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updated treatment of the Rubiaceae of Ko Chang is presented here, in which 64 taxa in 33 genera are recognised. The present study is based both on field work in Ko Chang and on a study of collections in the following herbaria: AAU, BK, BKF, C, K, W, WU. GEOGRAPHY AND VEGETATION OF KO CHANG Apart from Ko Chang, Mu Ko Chang National Park (NP) includes several small to very small islands (see map, Fig. 1). They are primarily concentrated around the southern part of the main island. Ko Chang itself, ca. 30 x 14 km, runs more or less parallel to the coast and appears very mountainous from the mainland due to the mountain ridge which runs down most of the length of the entire island. The ridge produces several summits, the tallest of which are Khao Lan, Khao Chom Prasat, Khao Khlong Mayom, Khao Salak Phet and Khao Yai. The latter, 743 m asl, is the highest peak. The rocks are mainly granitic and formed ca. 230195 mya. The mountains tend to have rounded slopes rather than sheer cliffs. Rainfall is very high (Trat Province has a mean annual rainfall of > 3000 mm, which is amongst the highest in Thailand; Thai Land Development Department web site, undated), and many very clear streams supply water all year round. The principal rivers of the island are Khlong Son, Khlong Mayom, Khlong Khangkhao, Khlong Bang Bao, Khlong Phrao and Khlong Nonsi. Settlements, tourist facilities, roads, etc. are mostly concentrated in the western coastal areas, while the reminder of the island is covered by largely undisturbed, dense tropical evergreen frest. Dominant tree species include the Dipterocarpaceae namely Dipterocarpus spp., Anisoptera costata and Hopea odorata; Podocarpus neriifolius and Croton spp.; species of the tree-like monocots Pandanus and Caryota; various woody climbers such as Bauhinia spp. Numerous Zingiberaceae such as Amomum spp. are common in the herb layer. Around the coastal villages of Salak Phet, Salak Khok, Khlong Son and Kh1ong Phrao Bay there are some large areas of Beach and Mangrove Forest. Dominant trees of the former include Terminalia catappa, Melaleuca cajuputi, Syzygium grande, S. gratum and Pandanus odoratissimus. Typical mangrove taxa include Rhizophora mucronata, R. apiculata, Ceriops decandra, C. tagal, Bruguiera gymnorrhiza, B. parviflora, Avicennia alba, Xylocarpus granatum, X. moluccensis, Hibiscus tiliaceus and Cerbera odollam. RESULTS Overview Sixty-four taxa of Rubiaceae belonging to 33 genera are known from Ko Chang (Table 1), as compared to 57/27 in Schumann (1902). Although the numbers of taxa in Schumann's treatment and ours do not widely differ, there are major changes in status of the taxa.
THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)
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Figure 1. Map of Ko Chang and surrounding islands (Mu Ko Chang National Park) (from Microsoft Interactive World Atlas 2001); inset: location of Ko Chang (arrow).
Eight of the 27 generic names in Schumanns Rubiaceae treatment in Flora of Koh Chang (RFKC) are no longer applicable (see below for details). Only 20 of the 57 names of taxa in RFKC are still in use. However, several of them are somewhat problematic and, at present, of more or less uncertain status. Of the 10 new species described in RFKC, only one is a good species; four others are doubtful or incompletely known; the remaining are synonyms of other taxa. None of the new combinations published in RFKC has been maintained. A detailed account is given below. Taxonomic changes since the Flora of Koh Chang Genera that are not currently recognised are Borreria, Coffea, Damnacanthus, Nauclea, Petunga, Plectronia, Randia and Stylocoryne. The most significant changes concern Randia. After it had been realized that Randia Houst. ex L. represented an exclusively American genus, differing from African and Asiatic Randia in having unisexual flowers with one-celled ovaries and pollen grains in tetrads, paleotropical taxa were re-investigated both in Africa (Keay, 1958, and many others), and tropical Asia (Tirvengadum, 1983; Wong, 1984; and others). The five taxa of Randia recorded from Ko Chang are now placed in four different genera (Aidia, Fagerlindia, Oxyceros and Rothmannia; see Table 1). Randia armigera has been transferred to Fagerlindia and Randia eucodon to Rothmannia. Randia oppositiflora (the epithet is mis-spelt as the basionym is Webera oppositifolia) is now considered a synonym of Aidia densiflora (cf. Ridsdale, 1996). The two remaining Randia spp. have been transferred to the genus Oxyceros. Whilst it is clear from the collections that R. fasciculata sensu Schum. (= R. siamensis Craib) is O. horridus, the correct identity of R. longiflora is not certain because the cited specimen could not be traced. If transferred to
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Oxyceros, the latter becomes O. longiflorus, but Tirvengadum (1982) cautioned that R. longiflora has often been confused with O. horridus. Plectronia, as used in RFKC, is not accepted here. The four Plectronia species in RFKC are now placed in three different genera. Whilst the transfer of three of the Plectronias to Canthium and Psydrax is merely the result of differing generic concepts within tribe Vanguerieae, Plectronia schmidtii, as first described in RFKC, was a mistake. As already discovered by Craib (1932), it is a member of tribe Octotropideae (formerly Hypobathreae; cf. Robbrecht, 1980, Robbrecht et al., 1993). Craib assigned it to Petunga, but that genus is now considered a synonym of Hypobathrum. Petunga schmidtii has not been formally transferred to Hypobathrum, but we refrain from making the new combination here because the genus is in need of general revision which may render the new combination unnecessary. Canthium in Thailand is not yet revised, and the names currently used for the RFKC Plectronia spp. (adopted from Craib, 1932) may not hold. The status of Plectronia siamensis, only known from Schmidts Ko Chang type collection, remains uncertain. It was transferred to Canthium siamense by Pitard (1924) but it is, as already noted by Craib (1932), very close to C. glabrum (see also under Damnacanthus, below). Also not yet resolved (for Thailand, and for Asiatic material in general) is the generic delimitiation of Canthium. According to Bridson (1992) Canthium s. str. is largely confined to Africa and only presented in Asia by a few taxa. Several Canthium spp. may belong to Psydrax, a genus re-instated by Bridson (1985). A formal transfer to Psydrax has only been published for Canthium nitidum (Psydrax nitidum (Craib) Wong; Wong, 1989). The genus Coffea does not occur naturally in Asia (see Bridson, 1987 for a survey of its complex nomenclatural history); both Coffea spp. in RFKC are thus referable to other genera. Coffea schmidtii, first published in RFKC, is not a Coffea segregate but belongs to Diplospora (D. schmidtii; Craib, 1932). The latter is identical to two other wrongly identified Diplospora spp. (D. malaccensis and D. pubescens sensu K. Schum.). The other species, Coffea fragrans sensu Schumann, is referable to Coffea merguensis Ridl. var. orientalis Craib (Craib, 1934). This, in turn, was transferred to Paracoffea merguensis (Ridl.) Leroy var. orientalis (Craib) Leroy (Leroy, 1971). This name, however, is invalid (see Bridson, 1985 for details). Later on, Leroy (in LobreauCallen & Leroy, 1980) validly transferred C. merguensis to Psilanthus merguensis (Ridl.) Leroy, but var. orientalis was forgotten so that, at present, P. merguensis var. orientalis is not validly published. Leroys (1971) understanding of var. orientalis was somewhat odd. While Craib (1934) distinguished var. merguensis and var. orientalis by their leaf sizes (smaller vs. larger) and corollas (sparsely pubescent vs. glabrous on the outside), and by their geographic range (the former in southern Burma and peninsular Thailand, the latter in south-eastern Thailand), Leroy used leaf consistency, an altered set of leaf length and width differences, and an obscure difference in flowers per inflorescence (pluri- vs. pauci- or uniflorous) to distinguish the two entities. This resulted in an odd distribution pattern: Leroys var. merguensis would be restricted to Burma (Tenasserim), and his var. orientalis would occur in peninsular and in southeastern Thailand (according to him, it also occurs in Burma, but none of the specimens
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listed originated from that country). We cannot follow Leroys distinction of the two varieties, and at present we refrain from formally publishing infraspecific taxa within Psilanthus merguensis because the genus should be revised first for the Flora of Thailand area (currently, there is only a revision for India; cf. Sivarajan et al., 1992). Nauclea has undergone marked changes in its generic delimitation, and both Nauclea spp. in RFKC need to be transferred to Neonauclea. As already noted by Craib (1932), Schumanns species identifications were erroneous, and Craib placed them under Neonauclea calycina which now is considered a synonym of N. pallida (cf. Ridsdale, 1978, 1989). Stylocoryne (also spelled Stylocoryna) is now generally considered synonymous with Tarenna, but the status of S. wallichii in RFCK is not entirely certain because the specimen could not be traced by Craib (1932) or by us. The correct name for this should be Tarenna wallichii, but so far it has only been recorded from peninsular Thailand and the Malay Peninsula (Wong, 1989). As disjunctions between peninsular Thailand and Ko Chang (and south-eastern Thailand in general) are relatively common (see below), it is possible that the species does occur on the island. Borreria is not generally considered a synonym of Spermacoce, and the weedy species recorded in RKCH becomes S. hispida. Damnacanthus is a widely distributed genus, occurring from Japan westwards to China and northern India (Assam, eastern Himalaya); its geographic range passes north of Thailand. Damnacanthus indicus, the most widely distributed species in the genus, covers the entire area (Kanjilal et al., 1939; Lo, 1979; Yamazaki, 1987). Thus, the record of Damnacanthus indicus from Ko Chang, a hot and wet, strictly tropical island much outside the species range, seems very odd. Neither Craib (1934) nor us were able to trace the specimen cited in RFKC (Schmidt 282), or any other collection of Damnacanthus from Thailand. The comment that the species is frequently cultivated also seems strange, as the spiny plant hardly makes an attractive ornamental. We are certain that it is one of the worse misnomers in RFKC. In the absence of the cited specimen, we cannot offer a definitive solution, but we have a strong suspicion that Schumanns Damnacanthus indicus might be one of the thorny Canthium species that was encountered in Ko Chang. The two taxa superficially resemble each other in having paired shoot thorns and clustered white flowers, and both genera have ovaries with a solitary, pendulous ovule per locule (Robbrecht et al., 1991; Li, 1963, Fig. 372). As Canthium s.l. (Canthium and Psydrax) is not yet revised for the Flora of Thailand, we are only able to identify the species as a member of the Canthium horridum group, characterized by paired short-shoot thorns. The new combination Oldenlandia costata is to be considered a synonym of Hedyotis venosa (Craib, 1932). Apparently Schumann had problems with the identification of Ko Chang Oldenlandia. Two others were wrongly identified, as already noted by Craib (1932); see Table 1. The not entirely resolved generic delimitation between Oldenlandia and Hedyotis, has further complicated the matter. Schumanns new species Mussaenda lanceolata, was, according to Craib (1932), a variety of M. cambodiana, differing chiefly in the denser indumentum of branchlets and leaves. Although the genus has not yet been revised for the Flora of Thailand, it appears that M. cambodiana var. lanceolata (only known from Ko Chang) is not worth
18
retaining but should be included in the variable M. cambodiana (recorded mainly from south-eastern Thailand, Cambodia and Laos). The status of the two Lasianthus species newly described by Schumann (L. oligoneurus and L. schmidtii) is not fully resolved. In his revision of the genus, Zhu (2001) noted that both are only known from the type collections. In particular L. oligoneurus is problematic as, according to Zhu, the type is incomplete, lacking fruits. Of the remaining four Lasianthus species in RFKC, only one, L. lucidus, has retained its name. Lasianthus caloneurus became a synonym of L. maingayi, and L. cyanocarpus is now L. hirsutus. L. strigosus Wight, for which Schumann noted that it is supposed to be a Ceylon (species), and of which he claims he has also seen specimens from New Guinea, is actually L. chevalieri, a species restricted to south-eastern Thailand, Vietnam and Hainan Island (China) (Zhu, l.c.). Three species of Ophiorrhiza are listed in RFKC, of which O. brachycarpa was newly described. Craib (1932) placed both other species in RFKC (O. mungos and O. harrisiana) with the former, indicating they were wrongly determined. The name O. mungos certainly cannot be correct; the species is restricted to southern India and Sri Lanka. O. harrisiana is a species widely distributed from India to Indochina and to the Malay peninsula (according to Schanzer (pers. comm), who has revised the genus for Flora of Thailand, the name O. harrisiana is predated by O. rugosa). This and O. brachycarpa are part of a polymorphic group of widely distributed taxa which is not yet fully resolved (Schanzer, pers. comm.). Thus the specific status of O. brachycarpa, is not yet certain. The only Urophyllum known from Ko Chang was described in RFKC as the new species U. schmidtii. The species is hardly distinct from U. blumeanum, a species common in the forests of peninsular Thailand. A revision of Thai Urophyllum might well show that its status as a distinct species cannot be upheld. The only new species described in RFKC that can be maintained appears to be Ixora dolichophylla, a distinct species only known from Ko Chang (the genus is currently under revision by one of us (V.C.)). New Records Since the publication of RFKC, a total of 15 taxa have been newly recorded from the island (see Table 1). Eleven of these are mentioned for the first time in Craibs Enumeratio (Craib 1932, 1934); the others are added by us. New records for the island include three categories of taxa are as follows: Taxa largely restricted to the south-eastern region of Thailand (and sometimes also extending to the neighbouring parts of Cambodia). Good examples are Argostemma pulchellum (Sridith & Puff 2000, Fig. 7), or Duperrea scabrida (replaces the northern D. pavettaefolia in the Southeast; Puff, unpublished). Taxa widely distributed in Thailand (and sometimes also in the surrounding countries). Examples are Psychotria adenophylla and some of the weedy Hedyotis species.
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Taxa with a distinctly disjunct distribution range of south-eastern/peninsular Thailand such as Timonius corneri or Uncaria cordata (see also below). Taxa or species pairs occurring both in Ko Chang (and the south-eastern region in general) and peninsular Thailand (cf. map, Fig. 2) The relatively large number of Rubiaceae showing this kind of disjunction seems noteworthy. At present nine taxa known from Ko Chang fall into this category (Gardenia tubifera, Hedyotis philippensis, Lasianthus constrictus var. latifolius, L. maingayi, Psychotria angulata, Psydrax nitida, Rennellia speciosa: Fig. 2, Timonius corneri, and Uncaria cordata). This represents 14% of all Ko Chang Rubiaceae. In the peninsula, these taxa often reach their northern limit in Ranong Province, and mostly they extend further South to the Malay peninsula (and sometimes also to Sumatra).
Figure 2. Distribution of Rennellia speciosa (based on collections in Thai herbaria).
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Table 1. List of Rubiaceae in Ko Chang, Trat. Column Name in Schumann: to retain the name as given by Schumann (1902), wrongly applied names are indicated by the suffix [sensu K. Schum.] rather than by the normally used sensu K. Schum., non . Column Notes and comments: X, XX = newly recorded after Schumanns treatment (XX = newly recorded by us). *, ** mark taxa newly described in Schumann (1902), or new combinations, respectively.
Current name Aidia densiflora (Wall.) Masam. Argostemma pulchellum Geddes Name in Schumann (1902) Randia oppositiflora (Roxb.) K. Schum. ** Argostemma sp. Notes and comments **; see text X; identified to species by Craib (1932), confirmed by Sridith (1999) according to Craib (1932) *; possibly not specifically distinct from C. glabrum; see text XX; see text
Damnacanthus indicus Gaertn. Plectronia didyma (Roxb.) Kurz [sensu K. Schum.] (p.p.) Chassalia curviflora (Wall.) Thw. same Coptosapelta flavescens Korth. same Dentella repens Forst. same Diplospora schmidtii (K. Schum.) Coffea schmidtii K. Schum. *; Craib Diplospora malaccensis Hook. f. [sensu K. Schum.] D. pubescens Hook. f. [sensu K. Schum.] Duperrea scabrida Craib Fagerlindia armigera (K.Schum.) Randia armigera K. Schum. * Tirveng. Gardenia tubifera Wall. same Guettarda speciosa L. same Hedyotis nodiflora Wall. ex. G. Don H. philippensis (Willd.) Merr. ex C.B. Robinson H. pinifolia Wall. ex G. Don Oldenlandia pinifolia (Wall. ex G. Don) O. Ktze. H. pitardiana Craib Oldenlandia mollis Wall. H. subcarnosa Wall. Oldenlandia lineata Roxb. (pp.) H. venosa Korth. Oldenlandia costata (Roxb.) K. Schum. ** H. wallichii Kurz Hydnophytum formicarum Jack same var. siamensis Becc. Hypobathrum schmidtii [Petunga Plectronia schmidtii C. B. Clarke schmidtii (C.B.Clarke) Craib] *; Petunga racemosa (Roxb.) K. Schum. [sensu K. Schum. 1902] Ixora coccinea L. same I. cibdela Craib I. dolichophylla K. Schum. * same
Canthium glabrum Blume Canthium siamense (K. Schum.) Pitard ? Canthium sp. (with thorns; C. horrridum group; Chamchumroon & Puff 1157, BKF) Canthium horridum group ? Canthium umbellatum Wight
Plectronia glabra (Blume) Kurz Plectronia siamensis K. Schum. *
see text according to Craib (1932); see text
*; see text
XX *; see text
X (Craib, 1932) X (Craib, 1932; as H. congesta)
see Craib (1932) see Craib (1932) **; see Craib (1932) X (Craib, 1932)
*; Petunga schmidtii, an older name for Petunga brevispica; is not formally transferred to Hypobathrum; see text X (Craib 1934) *
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Table 1. (continued).
Current name Ixora javanica (Blume) DC. Name in Schumann (1902) I. amoena Wall.; I. stricta Roxb.; I. stricta Roxb. var. blumeana Hook.f. same Notes and comments
I. kratensis Craib I. nigricans R. Br. ex Wight & Arn. Lasianthus chevalieri Pitard
XX
Lasianthus strigosus Wight [sensu K. Schum.] L. chinensis (Champ.) Benth. L. constrictus Wight var. latifolius Craib L. hirsutus (Robx.) Merr. Lasianthus cyanocarpus Jack L. lucidus Blume same L. maingayi Hook.f. Lasianthus caloneurus K. Schum. * L. oligoneurus K. Schum. * same L. schmidtii K. Schum. * same Morinda citrifolia L. same Mussaenda cambodiana Pierre ex Mussaenda lanceolata K. Schum. Pitard [var. lanceolata (K. Schum.) * Craib] Mycetia squamulosopilosa Pitard Neonauclea pallida (Reinw. ex Havil.) Bakh.f. Nauclea purpurea (Roxb.) Merr. [sensu K. Schum.]; Nauclea synkorynes Korthals [sensu K. Schum.] Oldenlandia diffusa (Willd.) Roxb. same Ophiorrhiza brachycarpa K. same; Schum. ? * O. harrisiana Heyne [sensu K. Schum.] O. mungos L. [sensu K. Schum.] Oxyceros horridus Lour. Randia fasciculata (Roxb.) DC. [sensu K. Schum.] O. longiflorus (Lam.)Yamazaki ? Randia longiflora Lam. Psilanthus merguensis (Ridl.) Leroy
see text XX X (Craib, 1934)
* *; see text *; see text *; see text
X (Craib, 1932); close to M. malayana; see text as Neonauclea calycina in Craib (1932); see text
*; synonomy according to Craib (1932); see text
Psychotria adenophylla Wall. P. angulata Korth. Psychotria sp. ?
P. rubra Poir. P. sarmentosoides Val. P. serpens L. Psydrax nitidum (Craib) Wong
as Randia siamensis in Craib (1934) no specimen seen; possibly O. horridus; see text Coffea fragrans Wall. [sensu K. as Coffea merguensis Ridl. var. Schum.] orientalis Craib in Craib (1934); see text X (Craib, 1934) X (Craib, 1934) Psychotria fulva Ham.ex Hook.f. Uncertain; no specimens seen (also [sensu K. Schum.] not by Craib, 1932: see his note under P. monticola, p. 194); P. fulva does not occur in Thailand Psychotria jackii Hook.f. [sensu K. see Craib (1934) Schum.] Psychotria sarmentosa Blume see Craib (1934) [sensu K. Schum.] same Plectronia didyma (Roxb.) Kurz as Canthium nitidum var. [sensu K. Schum.] (p.p.) suboblique Craib in Craib (1932); new combination in Wong (1989)
22
Table 1. (continued).
Current name Rennellia speciosa Hook.f. Rothmannia eucodon (K. Schum.) Bremek. Scyphiphora hydrophyllacea Gaertn. Spermacoce hispida L. Tarenna hoaensis Pitard T. wallichii (Hook. f.) Ridl. ? Timonius corneri Wong Uncaria cordata (Lour.) Merr. Urophyllum longifolium Hook.f. U. schmidtii C. B. Clarke *
Name in Schumann (1902) same Randia eucodon K. Schum. * same *
Notes and comments
as Borreria articularis in Craib (1934) Tarenna asiatica O. Ktze. [sensu see Craib (1932) K. Schum.] Stylocoryne wallichii (Hook. f.) K. specimen not seen; see text Schum. X (Craib 1932, as T. wallichianus); see Wong (1988) X (Craib 1932, as U. sclerophylla) same same *; possibly not distinct from U. blumeanum; see text
Borreria hispida (L.) K. Schum.
In addition, there are several closely related entities showing the same pattern, e.g. Psilanthus merguensis var. merguenis (peninsula) and var. orientalis (Ko Chang), or Urophyllum blumeanum (peninsula) and the scarcely distinct U. schmidtii (Ko Chang). Other examples are Mycetia malayana (peninsula) and the closely allied M. squamulosopilosa (south-eastern Thailand and Cambodia; Fukuoka, 1989), and Ixora brunonis (peninsula) and I. kratensis (south-eastern Thailand) which, according to Chamchumroon (in prep.), may need to be combined and considered geographical subspecies. If these are added to the statistics, the link between the peninsula and southeastern Thailand is represented by ca. 20 % of all Rubiaceae taxa recorded from Ko Chang. Further notes Prior to the establishment of Mu Ko Chang National Park at the end of 1982 (see above), there was negative human impact on the vegetation of Ko Chang. Settlements, farms, plantations, etc., were established in the more accessible ( flat) coastal areas. For these reasons, some of the classic collecting localities visited by, for example, Schmidt and Kerr (cf. Jacobs, 1962) in the late 19th and early 20th centuries have disappeared. For instance, one of the frequently cited localities, Klong Nonsi (equals Klong Muns), is now a populated area near the Ko Chang District Office with no traces of natural vegetation left. Examples are the type collections of Ixora dolichophylla, Lasianthus oligoneuron and L. schmidtii, all of which come from the Klong Nonsi area. None of these species have ever been recollected anywhere else. ACKNOWLEDGEMENTS Voradol Chamchumroon wishes to thank the AD (Austrian Academic Exchange) for a scholarship which made it possible to carry out part of the present studies at the Institute of Botany, University of Vienna and also the Danish Scholarship
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(DANCED) for support allowing Voradol Chamchumroon to visit the University of Aarhus and the Botanical Museum, Copenhagen. Mr Pramuk Thichakorn, Head of Mu Ko Chang National Park, is acknowledged for his assistance and for allowing us to work in the National Park. REFERENCES Bridson, D.M. 1985. The reinstatement of Psydrax (Rubiaceae subfam. Cinchonoideae, tribe Vanguerieae) and a revision of the African species. Kew Bull. 40: 687725. ________. 1987. Nomenclatural notes on Psilanthus, including Coffea sect. Paracoffea (Rubiaceae tribe Coffeeae). Kew Bull. 42: 453460. ________. 1992. The genus Canthium (Rubiaceae-Vanguerieae) in tropical Africa. Kew Bull. 47: 353401. Craib, W.G. 1932. Florae Siamensis enumeratio. A List of the plants known from Siam with notes of their occurrence. Vol. 2, part 1: Caprifoliaceae & Rubiaceae (in part). 1145. Siam Soc., Bangkok. ________. 1934. Florae Siamensis enumeratio. A List of the plants known from Siam with notes of their occurrence. Vol. 2, part 2: Rubiaceae (concluded) to Dipsacaceae: 147 233. Siam Soc., Bangkok. Fukuoka, N. 1989. Notes on the Rubiaceae from Thailand: 3. Acta Phytotax. Geobot. 40: 107118. Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. 1990. Index Herbariorum. Part I: The herbaria of the world. Ed. 8 [Regnum Vegetabile vol. 120]. New York Botanical Garden, Bronx, New York. Jacobs, M. 1962. Reliquiae Kerrianae. Blumea 11: 427493. Kanjilal, U.N., Das, A., Kanjilal, P.C. & De, R.N. 1939. Flora of Assam. Vol. III. Caprifoliaceae to Plantaginaceae: 1578. Allied Book Center, Dehradun. Keay, R.W.J. 1958. Randia and Gardenia in West Africa. Bull. Jard. Bot. Etat Bruxelles 28: 1572. Leroy, J.-F. 1971. Note sur un Cafier du Sud-Est Asiatique: Paracoffea merguensus (Ridl.) J.-F. Leroy. Adansonia, sr. 2, 11: 603614. Lobreau-Callen, D. & Leroy, J.-F. 1980. Quelques donnes palynologiques sur le genre Coffea et autre genres du cercle de Cafiers. Ass. Sc. Intern. Caf. 9e Colloque, Londres: 250251. Li, H.-L. 1963. Woody flora of Taiwan. Narberth, Livingston. 974 p. Lo, H.-S. 1979. A revision of the genus Damnacanthus Gaertn.f. from China [in Chinese]. Acta Phytotax. Sinica 17: 104109. Pitard, J. 1924. Rubiaceae (suite). In: Lecomte, M.H. (eds), Flore gnrale de l'IndoChine, vol. 3, fasc. 3: 289432. Masson, Paris.
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Ridsdale, C.E. 1978. A revision of the tribe Naucleeae s.s. Blumea 24: 307366. ________. 1989. A revision of Neonauclea. Blumea 34: 177275. ________. 1996. A review of Aidia s.l. (Rubiaceae) in Southeast Asia and Malesia. Blumea 41: 135179. Robbrecht, E. 1980. The Hypobathreae (Rubiaceae-Ixoroideae). 1. Delimitation and division of a new tribe. Bull. Jard. Bot. Nat. Belg. 50: 6977. Robbrecht, E., Puff, C. & Igersheim, A. 1991. The genera Mitchella and Damnacanthus. Evidence for their close alliance; comments on the campylotropy in the Rubiaceae and the circumscription of the Morindeae. Blumea 35: 307345. Robbrecht, E., Bridson, D.M. & Deb, D.B. 1993. The South Indian genus Octotropis (Rubiaceae). An investigation of its characters and reinstatement of the tribal name Octotropideae. In: Robbrecht, E. (ed.), Advances in Rubiaceae Macrosystematics. Op. Bot. Belg. 6: 8192. Schumann, K. 1902. Rubiaceae. In: Schmidt, J. (ed.), Flora of Koh Chang. Contributions to the knowledge of the vegetation in the Gulf of Siam. Bot. Tidsskr. 24: 329341. Sivarajan, V.V., Biju, S.D. & Mathew, P. 1992. Revision of the genus Psilanthus Hook. f. (Rubiaceae tribe Coffeeae) in India. Bot. Bull. Acad. Sinica (Taiwan) 33: 209224. Sridith, K. 1999. A synopsis of the genus Argostemma Wall. (Rubiaceae) in Thailand. Thai For. Bull. (Bot. ) 27: 86137. Sridith, K. & Puff, C. 2000. Distribution of Argostemma Wall. (Rubiaceae), with special reference to Thailand and surrounding areas. Thai For. Bull. (Bot. ) 28: 123138. Tirvengadum, D.D. 1982. A study of the tribe Gardenieae (Rubiaceae) of South and South East Asia. Generic delimitation and revision with enumeration of species. 1176. Thesis Licent. Sci (unpubl.), Aarhus. ________. 1983. New taxa and name changes in tropical Asiatic Rubiaceae. Nordic J. Bot. 3: 455469. Wong, K.M. 1984. The genera of Peninsular Malaysian Rubiaceae formerly confused with Randia. Malay. Nature J. 38: 157. ________. 1988. The Antirheoideae (Rubiaceae) of the Malay Peninsula. Kew Bull. 43: 491518. ________. 1989. Rubiaceae. In: Ng, F.S.P. (ed.), Tree flora of Malaya 4: 324425. Longman Malaysia, Petaling Jaya. Yamazaki, T. 1987. The genus Damnacanthus in Japan, Korea and Taiwan [in Japanese]. J. Phytogeogr. Taxon. 35: 6974. Zhu, H. 2001. A taxonomic revision of the genus Lasianthus Jack (Rubiaceae) from Thailand. Acta Phytotax. Sinica: 116150.
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Figure 3. A. Ko Chang, near Than Mayom waterfall. B-F. Rubiaceae recorded from Ko Chang; B. Psychotria angulata; C. Uncaria cordata; D. Rennellia speciosa; E. Gardenia tubifera; F. Duperrea scabrida (Photographed by C. Puff).
THAI FOR. BULL. (BOT.) 31: 2733. 2003.
Two new species of Neohouzeoua (Gramineae-Bambusoideae) from Thailand and Myanmar

SOEJATMI DRANSFIELD*, RUNGNAPAR PATTANAVIBOOL** & SARAWOOD SUNGKAEW***
ABSTRACT. Two new species of Neohouzeoua A. Camus, N. kerriana and N. fimbriata from southern Thailand and Myanmar, are described and illustrated.
Neohouzeoua A. Camus is found in tropical mainland Asia. The genus was described in 1922 with two species, N. mekongensis A. Camus and N. dulloa (Munro) A. Camus (transferred from the genus Teinostachyum Munro). A. Camus differentiated it from Teinostachyum Munro, because the six stamens have connate filaments, glumes or reduced glumes are absent, the palea lacks keels, and from Schizostachyum Nees again because the filaments are connate, the palea lacks keels but has two long tips and because the rachilla extension is much reduced or absent. When A. Camus described the genus, she did not include descriptions of the habit. Here extended descriptions are added, mainly based on the description of material of N. dulloa and recent collections of N. mekongensis (Dransfield, 1997). The genus possesses erect culms with thin walls, many branches of the same size at each node, indeterminate inflorescences with slender spikelets each containing one floret with the absence of a rachilla extension or with much reduced and slender rachilla extension. The spikelets are usually arranged in a group of four along the main and branch axis. In most bamboo genera possessing three stigmas, the top of the style is level where the stigmas are borne with spreading bases. In Neohouzeoua the style apex or top is level on one side, dipping down on the other side and filled with minute hairs. The stigmas are borne on the level part (Fig. 1, K1 & K2). This feature can be observed especially in mature spikelets and can be used to recognise Neohouzeoua. The inflorescences in N. mekongensis and N. dulloa are much branched, with slender branches, but are also found terminating leafy branches; subtending sheaths/bracts are usually glabrous and the slender spikelets are arranged in a group of four (usually), with glabrous glumes. In the new taxa, the inflorescences are usually borne terminating the leafy branches, with short branches, similar to those of Schizostachyum; the subtending sheaths/bracts are covered with stiff hairs; the spikelets are not clearly arranged in four, are less slender, and the lemmas and paleas are covered
*Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK. **Silviculture and Botany Division, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand. ***Department of Forest Biology, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand.
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with hairs. Superficially these new taxa do not resemble either N. mekongensis or N. dulloa, but we have included them in the genus, because the filaments are connate, the paleas have no keels on the back but bifid apices and two long tips, the rachilla extensions are absent and the stigma bases are erect or straight (Table 1). It is interesting to note that in these two new species of Neohouzeoua the sheaths or subtending bracts in the inflorescences remain intact on the nodes of the main axis, and bear modified blades and auricles, similar to the culm-leaves. It is also remarkable that these two common bamboos, found growing along roadsides, have never been described before. Neohouzeoua kerriana S. Dransf., R. Pattanavibool & S. Sungkaew, sp. nov. auriculis vaginae culmi magnis erectis, setis longis praeditis, setis 12 mm longis, laminis foliorum grandibus, auriculis grandibus erectis stramineis, setis brevibus, lemmatis dense hirsutis, apice lemmatis apicem paleae longiore/superanti distincta. Typus: Thailand, Kerr 12070 (holotypus K!). Fig. 1. Sympodial bamboo. Culms straight and erect, tips arching almost to the ground or leaning on nearby vegetation, about 4(8) m long, 1.1 cm in diam., with relatively thick walls, about 2 mm thick, internodes 2025 cm long, covered with appressed pale hairs, the upper parts covered with dark brown hairs, becoming glabrous and smooth, the hollow internodes layered/lined with a white membrane (easily removed), girdle present. Branches several at each node, the same size, no dominant primary branch. Culm-leaves rigid, 812 by 45 cm, top horizontal or slightly curved in the middle, covered with light caducous brown hairs, becoming glabrous; blades lanceolate, tapering to long tips, spreading or deflexed, 1215 cm long, 56 mm wide near the base, junction with sheath 4 mm wide, usually glabrous, pubescent near the base abaxially; auricles large, erect, 6 mm tall/wide, 21 mm long, glabrous, fringed with long bristles, bristles 12 mm long; ligule very short, minutely serrate. Leaf-blades 2230 by 47 cm, glabrous, tapering to long tips, up to 7 cm long, base attenuate or slightly rounded, petiole 1015 mm long; auricles large, thin or stramineous, glabrous, 10 by 45 mm, fringed with short curly hairs, hairs up to 5 mm long; sheaths glabrous; ligule very short, entire. Inflorescences borne terminally on leafy branches, or on leafless branches, up to 36 cm long, axis glabrous, internodes up to 9 cm long; pseudospikelets 23 cm long, cylindrical, arranged in a group (usually of four) at nodes of the main or/and branch axis, subtending bracts/sheaths up to 25 mm long, glabrous or glabresent, bearing modified blades and erect auricles, auricles with short bristles. Spikelets ca. 25 mm long, cylindrical, with light brown hairs; lemma 1618 by 8 mm, densely hairy towards the apex, otherwise glabrous, 13-nerved, acuminate, with long slender tips of ca. 6 mm long (resembling an awn), often exceeding palea apex; paleas not-keeled, 15 20 by 46 mm, glabrous, densely hairy near the apex, apex bifid, two tips of ca. 2 mm long, 12-nerved, connected with transverse nerves; lodicules not present; stamens 6, filaments joint, anthers with two short apiculate tips; ovary short, with long slender style, stigmas 3, erect/straight at the bases, tops plumose, spreading.
TWO NEW SPECIES OF NEOHOUZEOUA (GRAMINEAE-BAMBUSOIDEAE) FROM THAILAND AND MYANMAR
29
Thailand. PENINSULAR Chumphon [Langsuan, Khao Nam Sao, alt. 8001000 m, fl., 21 Feb. 1927, Kerr 12070 (holotype K!)]; Ranong [Muang, along the route Ranong-Chum Phon Date, alt. 58 m, sterile, 24 Feb. 2001, Sungkaew & Pattanavibool 13 (BKF!, K!)]. Distribution. Southern Thailand. Ecology. Primary or secondary/disturbed forest, on slopes or river banks, up to 1000 m. alt. Vernacular. Phai Racha wang (from Kerr 12070). Etymology. This species is named in honour of Dr A.F.G. Kerr. Notes. In the field this species can be recognised by its bushy habit with arching slender young culms emerging over the clump, and large leaf-blades; the hollow internodes are lined with white membrane, and the auricles of culm leaves are large, erect, and fringed with long bristles. The sheath of the leaf-blade possesses a large auricle fringed with bristles. N. kerriana can be found growing abundantly near streams along the road in Ranong. The type was collected from higher altitude. We believe this bamboo is widespread from Langsuan to Ranong. Neohouzeoua fimbriata S. Dransf., R. Pattanavibool & S. Sungkaew, sp. nov. N. kerrianae S. Dransf. et al. laminis grandibus affinis, sed auriculis vaginae culmi magnis deflexis, setis longis praeditis, setis 25 mm longis, auriculis laminorum foliorum grandis chartaceis deflexis, lemmate minus dense hirsute, apice lemmatis apicem paleae breviore differt. Typus: Myanmar, Parker 2733 (holotypus K!). Fig. 2. Scrambling, sympodial bamboo, up to 13 m long. Culms up to 3 cm in diam., internodes about 40 cm long, with relatively thin walls, light green with white wax and dark brown scattered hairs below the nodes, otherwise glabrous and smooth, a thin white membrane found inside the internode. Branches many at each node, of the same size, hairy on sheaths and auricles. Culm leaves 1316 cm by 6.59 cm near the base, covered densely with appressed dark brown hairs, hairs caducous; blades ovatelanceolate, deflexed, 1321 cm long, about 12 mm wide near the base, 6 mm at the junction with the sheath, tapering to long tips, glabrous adaxially, hairy abaxially especially near the base; auricles large, deflexed/flared, 21 mm long, 9 mm tall/wide, caducous, with long bristles, bristles 25 mm long; ligule 4 mm long, laciniate. Leafblades 2652 by 612 cm, tapering to very long fine tips, up to 3 cm long, base rounded, margins smooth or ciliolate, glabrous, often slightly scabrid towards the apex adaxially, with light brown hairs towards the petiole adaxially, petiole (pseudopetiole) 1 cm long, glabrous or with scattered pale hairs adaxially, base slightly swollen (above the junction with the sheath); sheaths covered with pale hairs, hairs caducous, margins ciliolate, callus or outer ligules prominent, 3 mm long/tall; auricles thick or chartaceus, large, flared, 14 mm long, 4 mm high, margins with long curly bristles, bristles about 20 mm long; ligule (inner ligule) very short but with long fimbriae (6 mm long). Inflorescences borne terminating leafy branches, 1330 cm long, main and branch axis glabrous, subtending bracts/sheaths up to 3.5 cm long, glabrous or pubescent,
30
bearing modified blades, blades spreading or deflexed, 13 cm long, and large auricles, auricles deflexed/flared, with long bristles, bristles up to 12 mm long; pseudospikelets arranged in group at each node of the inflorescence axis, 2530 mm long, cylindrical, bracts/sheaths subtending buds with long light brown hairs near the apex, apex with long tips or with much reduced modified blades. Spikelets 2025 mm long; lemmas 15 18 by 8 mm, acuminate with long tips, 6 mm long, not exceeding the palea, light brown hairs near the apex and along the middle vein and the margins, otherwise glabrous, 21-nerved; paleas 25 by 8 mm, light brown hairs near the apex, otherwise glabrous, apex bifid, tips 3 mm long, 22-nerved; lodicules absent; stamens 6, filaments fused; ovary small with long glabrous style, stigmas 3, base erect/straight, plumose. Young fruit obovoid, 12 mm long, 2.5 mm in diam., remains of the style 30 mm long. Thailand. PENINSULAR: Ranong [Between Kapong and Ranong, alt. 40 m, sterile, 26 Feb. 2001, Sungkaew & Pattanavibool 12 (BKF!, K!)]; Phangnga [Kopah, fl., 8 Dec. 1917, Haniff & Noor SNF2066 (K!, SING!)]. Myanmar. Mergui, Yangwa Khlong, fl., 1 March 1927, Parker 2733 (holotype K!); l.c., sterile, 1 March 1927, Parker 2736 (K!); Mergui, Karathuri-Bokpyin Range, alt. 3 m, fl., 17 Feb. 1925, Sukos 7659 (K!); l.c., fl., 19 March 1925, Scott 951 (K!); Mergui, Maliwun, fl., 23 March 1930, Su Koe 10980 (K!); Tavoy, Maungpok Nwalabo ridge, alt. 1200 m, sterile, 6 Jan. 1919, Rogers 322 (K!). Distribution. Southern Thailand and southern Myanmar. Ecology. On slopes of secondary growth or forest. Vernacular. Yat-wan (Shan) (from Parker 2733). Etymology. The specific name is based on the long bristles found along the margins of the large auricles of culm-leaves, leaf-blades, and sheaths/subtending bracts in the inflorescences. Notes. This species resembles Neohouzeoua kerriana in having a bushy habit with young slender culms projecting above it, and large leaf-blades. It differs from the latter in having large deflexed or flared auricles of the culm leaves, fringed with long bristles, the ligule with long fimbriae, leaf-blades with large deflexed chartaceous auricles fringed with long bristles (25 mm long), and the tip of the lemma not exceeding the tip of the palea. It seems that this bamboo is very common around Tavoy and Mergui in the southern part of Myanmar, and flowers frequently. In a letter to Mr Fisher (attached to the Parker specimens), R. N. Parker wrote that it is a gregarious scrambling bamboo, occasionally climbing to a height of 40 ft, very common on the Yangwa range of hills where it flowered gregariously, the ground in the flowered areas being covered with seedlings. In the letter it says that this bamboo is called Dendrocalamus longifimbriatus Gamble. In fact Dendrocalamus longifimbriatus was described based on mix specimens, the leafy branches match those of Neohouzeoua fimbriata, and the flowering branches, which is chosen as the type of Dendrocalamus longifimbriatus, are Dendrocalamus inflorescences (Dransfield & Wong, in prep.).
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Table 1.
Comparative morphology of Neohouzeoua mekongensis A. Camus, N. kerriana and N. fimbriata

mekongensis kerriana arching or scrambling erect, then drooping to the ground, relatively thick walled large, erect, with short bristles lanceolate, deflexed very short, minutely serrate large, glabrous large, stramineous, erect, with short bristles racemose hairy, with or without modified blades and erect auricles 25 mm long densely hairy towards the apex densely hairy near the apex 2 mm long fimbriata arching or scrambling erect, then drooping to the ground, relatively thick walled large, flared, with long bristles ovate-lanceolate, deflexed 4 mm long, laciniate large, hairy abaxially large, thick, flared with long bristles racemose hairy, with modified blades and deflexed auricles 20 25 mm long pubescent near the apex hairy near the apex 3 mm long
Habit Culms
erect erect, thin walled
Auricle of culmleaves Blades of culmleaves Ligules of culmleaves Leaf-blades Auricles of leafblades Inflorescence Subtending bracts or sheaths Spikelet Lemma Palea Palea tip
small, erect, with long bristles lanceolate, deflexed 3 mm, laciniate medium, glabrous small with long bristles much branched racemose glabrous, rarely with modified blades 19 mm long glabrous glabrous 2 mm long
ACKNOWLEDGEMENTS We would like to thank Dr Kongkanda Chayamarit of the Forest Herbarium, Bangkok (BKF), and her staff, for their help in the field. Soejatmi Dransfield received a travel grant from TOBU through Royal Botanic Gardens, Kew, UK. REFERENCES Camus, A. 1922. Neohouzeoua A. Camus, nov. gen. Bull. Mus. Paris 1922: 100102. Dransfield, S. 1997. Report on fieldwork collecting bamboos in Thailand. Thai For. Bull. (Bot.) 26: 3539.
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Figure 1. Neohouzeoua kerriana: A. inflorescence terminating leafy branch; B. part of inflorescence borne on leafless branch; C. tip of leaf-blade; D. auricle of leaf-blade; E. spikelet; F. lemma; G. palea; H. tip of palea; J. ovary and stamens; Ka. apex of style, front; Kb. apex of style, back. A, GK from the type, CD from Sungkaew & Pattanavibool 13. Drawn by Soejatmi Dransfield.
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Figure 2. Neohouzeoua fimbriata: A. inflorescence; B. apex of leaf-blade; C. spikelet; D. lemma; E. palea; F. apex of plea; G. ovary and stamens; H. apex of style; J. fruit. A, CJ from the type, B from Sungkaew & Pattanavibool 12. Drawn by Soejatmi Dransfield.
THAI FOR. BULL. (BOT.) 31: 3438. 2003.
A new description of Diospyros coaetanea (Ebenaceae)

SUTEE DUANGJAI* & CHAMLONG PHENGKLAI**
ABSTRACT. Following the discovery of female and fruiting specimens of Diospyros coaetanea, the description and keys to this species are emended.
The genus Diospyros L. (Ebenaceae) was treated for the Flora of Thailand by Phengklai (1981). The description of Diospyros coaetanea H.R. Fletcher was based on specimens with male flowers only. During recent fieldwork by Sutee Duangjai in Kanchanaburi, Nakhon Ratchasima and Saraburi, female and fruiting specimens were discovered. Using this material an emended description of D. coaetanea is presented here. Diospyros coaetanea H.R. Fletcher, Kew Bull. 1937: 383; in Fl. Siam. En. 2: 366. 1938; C. Phengklai in Fl. Thailand 2 (4): 385, Fig. 56. 1981. Type: Thailand, Mae Hong Son, Kerr 5466 (holotype BM). Figs. 12. Deciduous tree, up to 20 m high. Bark scaly, grey or black, inner bark reddish. Leaves elliptic or oblong, 20 by 7 cm, base acute, apex acute, subcoriaceous to coriaceous, glabrous on upper surface, pubescent on lower surface; lateral veins in 20 pairs, arched and anastomosing well away from the margin, inconspicuous on upper surface, prominent on lower surface; scalariform veins inconspicuous on lower surface; petiole 710 mm long, glabrous. Male flowers cymose or paniculate, 4(5)-merous; pedicel 23 mm long, glabrous. Calyx campanulate, 610 mm long; lobes as long as calyx-tube, blackish-tomentose on both sides. Corolla tubular, 1517 mm long, 5 mm in diam., divided to one-fifth, tomentose outside, glabrous inside. Stamens 1016, glabrous. Rudimentary ovary pilose at apex. Female flowers solitary, 45-merous; pedicel 23 mm long. Calyx campanulate, 610 mm long; lobes as long as calyx-tube, blackish-tomentose on both sides. Corolla urceolate, 517 mm long, 8 mm in diam., divided to one-quarter, tomentose outside, glabrous inside. Staminodes 610, glabrous. Ovary ovoid, tomentose on top, 8-locular; style solitary; stigmas 4. Fruit a berry, 5.57.5 by 3.55 cm, succulent, glabrous, rounded or depressed at both ends, fruiting calyx divided to middle, tomentose on both sides, apex of calyx lobes acuminate; lobes not reflexed or plicate or undulate, without nerves; fruit-stalk 915 by 79 mm; endosperm smooth. Thailand. NORTHERN: Mae Hong Son, Lampang, Phitsanulok; EASTERN: Nakhon Ratchasima [Sakaerat, S. Duangjai 9 (BKF)]; SOUTH-WESTERN: Kanchaburi [Thong Pha Phum, S. Duangjai 10 (BKF)]; CENTRAL: Saraburi [Phu Khae, S. Duangjai 11 (BKF)]; SOUTH-EASTERN: Chachoengsao.
*Department of Forest Biology, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. **The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
A NEW DESCRIPTION OF DIOSPYROS COAETANEA (EBENACEAE) (S. DUANGJAI & C. PHENGKLAI)
35
Distribution. Endemic to Thailand. Ecology. In mixed deciduous forest. Flowering MarchMay, fruiting March August. Vernacular. Lam ta khwai () (Northern). Uses. The fruits are edible. The key to Diospyros based on female specimens in the Flora of Thailand is emended as follows:
23. Stigmata 4 24. Calyx divided to one fifth or a half; corolla-lobes villous or tomentose outside; leaves glabrous 24a. Corolla campanulate; corolla-lobes villous outside; calyx divided to one-fifth 26. D. areolata 24b. Corolla urceolate; corolla-lobes tomentose outside; calyx divided to a half 56. D. coaetanea 24. Calyx divided to base or nearly so; corolla-lobes glabrous or only hairy along mid-line outside; leaves hairy 29. D. glandulosa
We have also emended the key to fruiting specimens as follows:

15. Fruit up to 1 cm across 16. Fruiting calyx 3-merous 3. D. ferrea 16. Fruiting calyx 4-merous 10. D. buxifolia 15. Fruit 1.57.5 cm across 17. Fruiting calyx not dilated 17a. Apex of fruiting calyx-lobes acute; fruit up to 5 cm across; fruit-stalk up to 4 mm across 54. D. pendula 17b. Apex of fruiting calyx-lobes acuminate; fruit 5.57.5 cm across; fruit-stalk 79 mm across 56. D. coaetanea 17. Fruiting calyx dilated between lobes 55. D. wallichii
ACKNOWLEDGEMENT The authors are grateful to Mrs Jintana Boonchern for preparing the line drawings. REFERENCE Phengklai, C. 1981. Ebenaceae. In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 2(4): 281392.
36
D E F
Figure 1. Diospyros coaetanea H.R. Fletcher: A. habit; B. female flower; C. male flower; D. fruit; E. young fruit; F. seed. Scale bars = 1 cm. A from Duangjai 11 (BKF), B-E, F from Duangjai 10 (BKF), C from Duangjai 9 (BKF). Drawn by J. Boonchern.
A NEW DESCRIPTION OF DIOSPYROS COAETANEA (EBENACEAE) (S. DUANGJAI & C. PHENGKLAI)
37
Figure 2. Diospyros coaetanea H.R. Fletcher: A. leaves; B. male flowers; C. female flower; D. fruit.
THAI FOR. BULL. (BOT.) 31: 3943. 2003.
A new species of Spatholirion (Commelinaceae) from Thailand and further notes on S. ornatum
KAI LARSEN & SUPEE SAKSUWAN LARSEN*
ABSTRACT. Spatholirion calcicola a new species from northern Thailand is described and illustrated. Spatholirion decumbens Fukuoka & N. Kurosaki is shown to be synonymous with S. ornatum Ridl. A key to Spatholirion in Thailand and adjacent areas is provided.
A NEW SPECIES OF SPATHOLIRION In an earlier paper (Larsen & Larsen, 1994), the distribution and variation of Spatholirion ornatum Ridl. in Thailand was discussed. There it was stated that, besides an old collection from Tomoh (= To Mo) in southern Thailand, all the other collections of the species were from the northern part of the country. It was also stated that the collections from northern Thailand, studied in the wild, did not match Ridleys description in every detail, and that it would be interesting to refind the species in southern Thailand and study the life form under the more humid climatic conditions here. In 1995 the Thai-Danish Botanical Expedition visited Hala-Bala Wildlife Sanctuary in Narathiwat province. Hala-Bala is situated on the Thai-Malaysian border adjacent to the Belum Nature Reserve in Malaysia. The locality, named Sirindhorn Falls, near Waeng is close to the old mining village of To Mo visited by Ridley and from where he described S. ornatum. The surroundings consist of primary evergreen forest with acid soil, dark and rich in humus. In this habitat several populations of S. ornatum were found. In BKF there is also a collection C.S.S. 212 from Waeng. These specimens fully match Ridleys original description (Ridley, 1896): and therefore, what has previously been treated as S. ornatum in northern Thailand is a different species. Besides the morphological differences, there is also a striking difference in the ecological requirements of the two taxa. Therefore, the material from northern Thailand is described here as a new species. Spatholirion calcicola K. & S.S. Larsen sp. nov. A S. ornato Ridl. differt caulibus non radicantibus, radicibus crassis tuberiferis, caule ad basin cum innovationibus numerosis, foliis ellipticis utrinque pallide viridibus, axe inflorescentiae floribusque masculinis violaceis; in terra calcarea sylvarum deciduarum crescenti. Typus: Thailand, Nan, Larsen et al. 46420 (holotypus AAU!; isotypi BKF!, K!, L!, P!). Fig. 1.
*Department of Systematic Botany, University of Aarhus, Denmark.
40
Acaulescent herb with innovation shoots at base; roots thick with spherical or fusiform storage tubers; stems not rooting. Leaves: petiole up 25 cm long, 45 mm broad, canaliculate at base, pubescent; lamina sparsely pubescent on upper side, particularly along the midvein in the lower part, 920 by 712 cm, green on both sides, elliptic, attenuate or truncate to cordate at base, apex acute to acuminate. Inflorescence on a ca. 56 cm long peduncle with one basal bract enclosing the bisexual flowers, remaining inflorescence a panicle of male flowers. Peduncle densely woolly. Bract spathaceous, when opened subrotund to lanceolate, 1520 mm broad with pointed apex, margin densely woolly, otherwise sparsely pubescent on the upper surface. Rachis of inflorescence pubescent. Flower buds with dense yellowish pubescence of patent hairs. Bisexual inflorescence cream-coloured with 12 flowers, perianth subequal in length, the outer whorl somewhat broader, 710 mm long, 24 mm broad, stamens with a ca. 5 mm long filament, anther ca. 1 mm long; pistil ca. 1 cm long; male flowers violet, similar to the bisexual ones but smaller in all parts, perianth 34 mm long, stamens ca. 4 mm. Ovary minute, not developing. Capsule puberulous, 3-valved, angulate, 20 by 78 mm. Seeds browish, reniform with an orange aril, ca. 5 by 3 mm. Thailand. Without locality, Kerr 20573 (K); NORTHERN: Chiang Mai [along the road Fang-Chiang Mai, in shade among limestone boulders, alt. 400 m, Aug. 1963, Larsen 2762 (AAU)]; Nan [Tham Pa Toob, 10 km N of Nan, rugged limestone hills covered with secondary, mixed deciduous forest and bamboo thicket, alt. 250 m, 13 Sept. 1995, K. Larsen et al. 46420 (holotype AAU!; isotypes BKF!, K!, L!, P!); Tham Pa Toob, K. Larsen et al. 43586 (AAU, BKF, PSU) ]; Lampang [Pang Puai, alt. 420 m, 24 May 1915, Kerr 3617 (K); Ngao, Sept. 1931, Put 4010 (K); Ngao, Mae Huat, on rocks, alt. 350 m, Sangkhachand 92 (BKF); Jae Sawn Nat. Park, at base of limestone cliffs, 500 m alt. 25 Aug. 1995, Maxwell 95585 (BKF); Pratu Pha, limestone, 9 July 1965. Smitinand & Phengklai 8871 (BKF)]. Distribution. Endemic to northern Thailand. Ecology. Growing in crevices in limestone rocks or in scree; alt. 350500 m. Cytology. All three species have been studied with regard to chromosome number. Larsen & Larsen (1994, Fig. 31) gave the chromosome number 2n=20 for S. ornatum. This number refers to S. calcicola. S. ornatum has now been counted for the first time, from living material brought back and grown at AAU, and has been found to have the same number. All three species have 2n=20. Note. Larsen et al. 46420 was chosen as the holotype as the chromosome number was determined from this plant. Its flowers are preserved in alcohol and colour slides are deposited in AAU. FURTHER NOTES ON S. ORNATUM Spatholirion ornatum Ridl., J. Bot. 1896: 329. 1896. Type: Thailand, Narathiwat, Legeh at Tomoh, Machado s.n. (lectotype SING!; isolectotype K!, selected here). S. decumbens Fukuoka & N. Kurosaki, Acta Phytotax. Geobot. 48: 147. 1997.
A NEW SPECIES OF SPATHOLIRION (COMMELINACEAE) FROM THAILAND AND FURTHER NOTES ON S. ORNATUM
41
We have studied the type material of S. ornatum deposited in K and SING. The sheet at Kew consists of one leaf and a packet containing a few flowers. It seems clear to us that this is a duplicate taken from the better material in SING. Consequently, the sheet in SING has been selected here as the lectotype. In 1994 a Japanese expedition to Thailand visited the area at Waeng in Narathiwat province and collected material of Spatholirion. During our expedition to that area in 1995 we were told by the Head of a local forest station that members of the Japanese expedition were very interested in the plant, which is now grown as an ornamental at the station. Fukuoka & Kurasaki (1997) described their collection as a new species, S. decumbens. According to their paper they saw a photo of the type collection in Kew. However, they did not see the better type material in SING, nor the material cited in Larsen & Larsen (1994), and they also misinterpreted the collections from northern Thailand in BKF. There is no doubt that the locality where S. ornatum and S. decumbens has recently been collected in Narathiwat province is very close to the type locality; it may even be the same place. There are no significant differences between these new collections and the material of Ridley. There is, however, a clear difference between the southern material and the material from northern Thailand, as Fukuoka & Kurasaki correctly point out. They also give a very good description of the main differences between S. ornatum and our new species, but mistook the material from northern Thailand to be S. ornatum. Spatholirion decumbens thus falls within the circumscription of Sphatholirion ornatum when the northern material (now assigned to S. calcicola) is excluded. Thus it is reduced here to a synonym of S. ornatum. The following key will help to identify Spatholirion in Thailand and adjacent regions.
KEY TO THE SPECES OF SPATHOLIRION 1. Scandent herb, leaves lanceolate to lanceolate-elliptic, long acuminate (China) S. longifolium 1. Erect or decumbent herbs, leaves elliptic or ovate (Southern Thailand, Malaysia) 2. Inflorescence creamy white, male flowers yellow-white. Leaves ovate with cordate base, upper sides dark green, almost glabrous, undersides purple; petioles densely sericiously hairy with white hairs, stems rooting at the lower nodes, roots thin. On acid soil in evergreen forest (West Malaysia: Kelantan, Thailand: Narathiwat) S. ornatum 2. Inflorescence pale violet, male flowers violet. Leaves elliptic with attenuate or truncate to cordate base, both sides pale green, upper sides patently hairy; petioles less hairy; stems not rooting; roots thick with spherical storage tubers. In crevices of calcareous rocks or at the base of calcareous boulders in deciduous forest (northern and north-eastern Thailand) S. calcicola
ACKNOWLEDGEMENTS The authors are greatly indebted to Dr Christian Tange who forded a flooded valley in the limestone area North of Nan in order to bring back living material of S. calcicola for growing in the greenhouses in Aarhus. Dr B. llgaard provided a latin traslation of the diagnosis. Dr R.B. Faden kindly read the manuscript. We are also indebted to Prof. Pranom Chantaranothai and Mr Thaweesak Thitimetharoch for further information on the occurrence of S. calcicola in Thailand. The plate was skilfully drawn by Piyakaset Suksathan. The field work was supported by the Danish Natural Science Research Council grant no 11-07-11 and the Carlsberg Foundation supported several visits to herbaria in Europe.
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REFERENCES Faden, R. B. 1998. Commelianaceae. In: Kubitzki, K. et al. (eds), The Families and Genera of Vascular Plants IV: Flowering Plants: Monocotyledons, pp. 109128. Springer-Verlag, Berlin. Fukuoka, N. & Kurosaki, N. 1997. A new species of Spatholirion (Commelinaceae) from Peninsular Thailand. Acta Phytotax. Geobot. 48: 147150. Jacobs, M. 1962. Reliquiae Kerrianae. Blumea 11: 427493. Larsen, K. & Larsen, S.S. 1994. Spatholirion ornatum (Commelinaceae) in Thailand. Thai For. Bull. (Bot.) 22: 8991. Ridley, H.N. 1896. A new genus of Commelinaceae. J. Bot. Lond. 34: 329330.
A NEW SPECIES OF SPATHOLIRION (COMMELINACEAE) FROM THAILAND AND FURTHER NOTES ON S. ORNATUM
43
Figure 1. Spatholirion calcicola: A. habit; B. inflorescence; C. bisexual flower; D. male flower; E. stamen; F. lower part of infructescence with capsule; G. open capsule; HI. seed seen in front and back view. All from Larsen et al. 46420 (AAU). Drawn by P. Suksathan.
THAI FOR. BULL. (BOT.) 31: 44 46. 2003.
Notes on Clerodendrum (Lamiaceae)

CHARAN LEERATIWONG* & PRANOM CHANTARANOTHAI**
ABSTRACT. A new combination, Clerodendrum farinosum var. pubescens is made. Clerodendrum lasiocephalum and C. schmidtii var. macrophyllum are reduced to synonymy of C. chinense var. simplex and C. schmidtii, respectively. Clerodendrum smitinandii is recorded from Cambodia and Vietnam for the first time. Six lectotypes are selected.
The following nomenclatural and typification changes have arisen from work being carried out for the Flora of Thailand Clerodendrum account. Clerodendrum farinosum (Roxb.) Walp. var. pubescens (H.R. Fletcher) C. Leeratiwong & P. Chantaranothai comb. nov. Clerodendrum venosum Wall. ex C.B. Clarke var. pubescens H.R. Fletcher, Bull. Misc. Inform., Kew 1938: 205. 1938; Fletcher, Bull. Misc. Inform., Kew 1938: 428. 1938. Type: Thailand. Chiang Mai, Me Tun, mixed forest, alt. ca. 600 m., A.F.G. Kerr 6190 (lectotype BK!, selected here; isolectotype K!). Kerr 6190 had been named as C. venosum var. pubescens by Fletcher (1938). However, Daniel & Rajendran (1993) proposed C. farinosum as the correct name for V. venosum. Therefore, it is necessary to make a new varietal combination. This variety can be easily separated from the typical variety by the dense hairs on the twigs and the lower surface of the leaves. It is known only from the type collection. Clerodendrum chinense (Osbeck) Mabb. var. simplex (Moldenke) Chen, Novon 1: 58. 1991. C. philippinum Schauer var. simplex Moldenke, Phytologia 20: 338. 1970. Type: Thailand, Ban Musseo, between Tak and Mae Sot, alt. 400 m, F. Floto 7634 (holotype herb. Moldenke). C. lasiocephalum C.B. Clarke in Hook.f., Fl. Brit. India 4: 594. 1885, synon. nov. Type: India, Mishmee, Griffith 6055 (holotype K!). Although closely related to the typical variety, var. simplex differs consistently in the length of the corolla tube, with a single layer of five lobes. Clerodendrum lasiocephalum is similar to C. chinense var. simplex. Therefore the former is reduced to the synonymy of the latter. The species is widespread from India, Myanmar and Thailand (except the peninsular part) to China.
*Department of Biology, Faculty of Science, Prince of Songkla University, Songkhla 90112, Thailand. **Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand.
NOTES ON CLERODENDRUM (LAMIACEAE) (C. LEERATIWONG & P. CHANTARANOTHAI)
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Clerodendrum garrettianum Craib, Bull. Misc. Inform., Kew 1911: 443. 1911. Type: Thailand, Chiang Mai, Doi Suthep, A.F.G. Kerr 1435 (lectotype K!, selected here; isolectotypes BM!, K!). Craib (1911) described C. garrettianum based on two specimens, Kerr 1309 (BM!, K!) and Kerr 1435, from Doi Suthep, Chiang Mai province. The latter collection is selected as the lectotype, as it is a comprehensive well preserved specimen. The species is widespread from north and north-eastern Thailand to Indo-China. Clerodendrum lloydianum Craib, Bull. Misc. Inform., Kew 1914: 284. 1914. Type: Thailand, Phrae, Vanpruk 499 (lectotype K!, selected here; isolectotype K!). Clerodendrum lloydianum is widespread in Thailand (except the peninsular part) and Indo-China. Clerodendrum schmidtii C.B. Clarke, Bot. Tidsskr. 26: 173. 1904. Type: Thailand. Trat, Chang Island, Klawng Nonsi, Schmidt 740 (lectotype K!, selected here). C. hastato-oblongum C.B. Clarke, Bot. Tidsskr. 26: 173. 1904. Type: Thailand. Trat, Chang Island, Klawng Son, Schmidt 692a (holotype C). C. schmidtii C.B. Clarke var. macrophyllum Moldenke, Phytologia 23: 180.1972, syn. nov. Type: Thailand. Trat, Koh Rangyai, 20 km south of Koh Chang, Charoenphol, Larsen & Warncke 5022 (holotype AAU; isotype BKF!). Variety macrophyllum was based on Charoenphol, Larsen & Warncke 5022, which has ovate, obovate or cordate leaves. However, intermediates exist between this variety and the typical plant. We conclude that Moldenke's variety (1972) does not warrant separation at that rank. The species is widespread in north-eastern to southeastern Thailand and Indo-China. Clerodendrum smitinandii Moldenke, Phytologia 7: 79. 1959. Type: Thailand: Chanthaburi, Pong Namrawn, Smitinand 3197 (holotype herb. Moldenke). Smitinand & Abbe 6519 (BKF, K) which was collected from Mt Bokor, Kampot, Cambodia and J. & M.S. Clemens 4261 (K), collected from Mt Basi, Vietnam, May July 1927, are the first records for Cambodia and Vietnam, respectively. Clerodendrum umbratile King & Gamble, Bull. Misc. Inform., Kew 1908: 110. 1908. Type: Malaysia: Perak, Dr Kings collector 722 (lectotype K!, selected here). Eleven syntypes were cited in the protologue of C. umbratile by King and Gamble (1908). The syntypes were collected by botanists from Malaysia and Indonesia. Dr King's collector 722 is selected here as the lectotype because it is in better condition than the other syntypes. The species is widespread from southern Thailand and peninsular Malaysia to Sumatra.
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THAI FOREST BULLEIN (BOTANY) 31
Clerodendrum vanprukii Craib, Bull. Misc. Inform., Kew 1911: 444. 1911. Type: Thailand. Phrae: Huai Kamin, Vanpruk 207 (lectotype K!, selected here; isolectotype K!). Clerodendrum vanprukii is endemic to northern Thailand. ACKNOWLEDGEMENTS We sincerely thank curators and staff of the following herbaria for the loan of material and, in some cases, for assistance during our visits to the institutions: BCU, BK, BKF, BM, K, LINN, PSU, QBG and US. We also thank Dr David Simpson (K) for providing some information. This study is supported by the Biodiversity Research Program, Thailand, BRT 541075. REFERENCES Craib, W.G. 1911. Contributions to the Flora of Siam. II. List of Siamese plants with descriptions of new species - continued. Bull. Misc. Inform., Kew 1911: 443. Daniel, P. & Rajendran, A. 1993. Clerodendrum farinosum, the correct name for C. venosum (Verbenaceae). Taxon 42: 669670. Fletcher, H.R. 1938. Contributions to the Flora of Siam. Additamentum XLIX. Bull. Misc. Inform., Kew 1938: 199209. Anon. 1908. Decades kewenses. Plantarum novarum in herbario Horti Regii conservatarum. Decades XLVIIXLVIII. Bull. Misc. Inform., Kew 1908: 110. Moldenke, H.N. 1972. Novelties from Brazil, Peru, Venezuela, and Thailand. Phytologia 23: 180181.
THAI FOR. BULL. (BOT.) 31: 47 52. 2003.
Matoniaceae (Pteridophyta) - a new family record for Thailand

STUART LINDSAY*, SOMRAN SUDDEE**, DAVID J. MIDDLETON* & RACHUN POOMA**
ABSTRACT. The fern species Matonia pectinata R.Br. has recently been collected in two provinces in Peninsular Thailand (Trang and Yala). These collections represent the first records for this species, the genus Matonia and the family Matoniaceae in Thailand. Trang is also the new northern limit of the known distributional range of M. pectinata.
During a plant collecting trip to the Khao Banthat Mountain Range in Trang province during 2003 a very unusual fern species, clearly absent from the pteridological literature for Thailand (see Tagawa & Iwatsuki, 1979, 1985, 1988, 1989; Boonkerd & Pollawatn, 2000), was found near the summit of Phu Pha Mek. Subsequent research revealed that this species was Matonia pectinata R.Br. and that another plant (specimen at BKF but its existence unpublished) had been collected in Yala province in February 2000. These two collections represent not only the first records for M. pectinata in Thailand but also the first records for the genus Matonia and the family Matoniaceae in Thailand. Matoniaceae is a small family of primitive leptosporangiate ferns. It has a widespread and diverse fossil record (interpreted as 11 or more genera; see Holttum, 1954, 1968, Kramer, 1990, Tryon & Lugardon, 1990, Nishida et al., 1998) but, today, the family is represented by only four species in two genera that are restricted to Southeast Asia. The two genera (which differ most noticeably in their ecological preferences and frond architecture) are Matonia and Phanerosorus. Matonia has two species that are very similar (Kato, 1993): M. pectinata R.Br. known (until now) from Peninsular Malaysia, Riau Archipelago, Lingga Archipelago and Sumatra, and M. foxworthyi Copel., known from Borneo, the Philippines, Moluccas and New Guinea. Kedah Peak, a mountain in the Kedah province of Peninsular Malaysia has, for many years, been accepted as the northern limit of the distributional range of M. pectinata. However, as a result of the discoveries reported here, the Khao Banthat Mountain Range in Trang province of Peninsular Thailand has become the new northern limit of the known distributional range of M. pectinata. Its discovery there and in Yala raises the interesting possibility that more Malaysian species may yet be discovered on some of the underexplored mountains of Peninsular Thailand.
*Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, USA. **The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
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MATONIACEAE Matoniaceae C. Presl, Gefssbndel Farrn 32, t.6. 1847 [Gefssbndel Farrn 32, t.6. 1847 was a pre-print of Abh. Knigl. Bhm. Ges. Wiss. ser.5, 5(2): 340, t.6. 1848]; Diels in Engler & Prantl (eds.) Nat. Pflanzenfam. 4(194): 343350. 1900; Bower, Ferns 2: 220227. 1926; Copel., Gen. Fil. 172173. 1947; Holttum, Revis. Fl. Malaya 2: 5860. 1954 (also 2: 5860, 2nd ed., 1968); Parris, Jermy, Camus, & Paul in Jermy, Studies on the flora of Gunung Mulu National Park, Sarawak 189. 1984; Kramer in Kubitzki, Fam. Gen. Vasc. Pl. 1: 183185. 1990; Kato, Fl. Malesiana ser. 2, 3: 289294. 1998. Terrestrial or lithophytic. Rhizome: creeping, dorsiventral, dicyclic-solenostelic or tricyclic-solenstelic, and densely covered with shiny, pale-brown or orange-brown, multicellular, uniseriate hairs. Fronds: alternate on the dorsal side of the rhizome; erect or pendant; stipes slightly or very hairy at base (hairs identical to those on rhizome) but glabrous and shiny above; lamina pedate with pectinate pinnae, or alternately pinnate with pinnae consisting of resting buds or bud-derived leaflets and linear, simple or forked pinnules; veins free or only weakly anastomosing in sterile portions, usually anastomosing in soriferous portions. Sori: on the underside of fronds, round or elliptic, in one row on each side of costule or midrib; indusia, peltate, deciduous, thick in central portion, membranous and inrolled in marginal portion; sporangia few, large, simultaneously maturing, in 13 layers in a circle around receptacle; each consisting of a short thick stalk and a subglobose capsule with an incomplete oblique annulus. Spores: tetrahedral-globose, trilete, pale, surface granulate. Two genera, each with very distinctive frond architecture: Matonia R.Br. and Phanerosorus Copel. Only Matonia is known in Thailand. MATONIA Matonia R.Br. in Wall., Pl. Asiat. Rar. 1: 16, t.16. 1829; Hook. & Bauer, Gen. Fil. t.43. 1840; Hook., Sp. Fil. 5: 285286. 1864; Copel., Gen. Fil. 172. 1947; Holttum, Revis. Fl. Malaya 2: 5960. 1954 (also 2: 5960, 2nd ed. 1968); Parris, Jermy, Camus, & Paul in Jermy (ed.) Studies on the flora of Gunung Mulu National Park, Sarawak 190. 1984; Kramer in Kubitzki, Fam. Gen. Vasc. Pl. 1: 183185. 1990; Kato, Blumea 38(1): 167172. 1993; Kato, Fl. Malesiana ser. 2, 3: 290292. 1998. Type species: Matonia pectinata R.Br. Terrestrial. Rhizome: creeping, dorsiventral, tricyclic-solenostelic, densely covered with shiny, orange-brown, multicellular, uniseriate hairs. Fronds: alternate in two rows on the dorsal side of the rhizome; erect, stipe brown or chestnut-brown, hairy at base but glabrous and shiny above, much longer than lamina; lamina perpendicular to stipe (umbrella-like), pedate, symmetrical, consisting of a central pinna and, on each side of that, an equal number (513) of progressively shorter pinnae; pinnae pectinate, deeply lobed, linear, coriaceous; costae glabrous on top, glabrous or hairy underneath; pinna segments linear-oblong and obtuse or narrowly deltoid and acute, entire, with revolute margins, shiny on top, often glaucous and papillate underneath; veins forming costal areoles, forked, free or
MATONIACEAE (PTERIDOPHYTA) -A NEW FAMILY RECORD FOR THAILAND
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only weakly anastomosing in sterile portions of segments, always anastomosing in soriferous portions (sori centred on small circular veins fed by 59 radial veinlets). Sori: on the underside of fronds, round, usually 03 per segment (but occasionally 4 or 5), often distributed with a bias towards the basiscopic sides of segments; indusia hemispherical, peltate, deciduous, thick in central portion and membranous and inrolled in marginal portion; sporangia 510 in one layer arranged in a tight circle around a prominent, persistent, receptacle; capsules subglobose-polygonal with incomplete oblique annuli. Spores: tetrahedral-globose, trilete, pale, surface granulate. Two very similar species: M. pectinata R.Br. and M. foxworthyi Copel. Only M. pectinata is known in Thailand. Matonia pectinata R.Br. in Wall., Pl. Asiat. Rar. 1: 16, t.16 (illustrations of the holotype). 1829; Hook. & Bauer, Gen. Fil. t.43 (illustrations of the holotype). 1840. Hook., Sp. Fil. 5: 285286. 1864; Bedd., Ferns Brit. India 2: 186, pl. 186. 1866; Bedd., Handb. Ferns. Brit. India 19 1883; Tansley & Lulham, Ann. Bot. (Oxford) 19(76): 475519, pl. 3133. 1905; Copel., Sarawak Mus. J. 2(3): 388. 1917; Holttum, Revis. Fl. Malaya 2: 5960. 1954 (also 2: 5960, 2nd ed. 1968); Kato, Blumea 38: 167172. 1993; Kato, Fl. Malesiana ser. 2, 3: 289292. 1998. Type: Peninsular Malaysia, Malacca, Mt Ophir (present day Gunung Ledang, Johor), near the summit at ca.1200 m, 1815, Farquhar s.n. (holotype K-W, Wallich List no.184, seen on microfiche). Fig. 1. Rhizome: 46 mm in diam. and densely covered with shiny orange-brown hairs, 33.5 mm long. Fronds: 14.5 cm apart, stipes 17 mm in diam. and 40135 cm long; lamina approximately 2050 cm diam., central pinna 1538 cm long, 1026 shorter lateral pinnae (i.e. an equal number (513), on each side of the central pinna); pinna segments up to 2 cm long and 47 mm broad at base, narrowly deltoid and acute and usually oblique and falcate. Sori and spores as in genus description above except that Thai specimens of M. pectinata have 58 sporangia per sorus (and most commonly 6). Thailand. PENINSULAR: Trang [Yan Ta Khao District, Khao Banthat Mountain Range, near summit of Phu Pha Mek, 1200 m, 7 April 2003, D.J. Middleton, V. Chamchumroon, S. Lindsay, R. Pooma & S. Suwanachat 1992 (A, BCU, BKF, L)]; Yala [Betong District, Sangala Khiri Mountain Range, 1520 m, 23 Feb. 2000, C. Niyomdham, P. Puudjaa & S. Chonkunjana 6082 (BKF)]. Distribution. Peninsular Malaysia, Riau Archipelago, Lingga Archipelago and Sumatra. Ecology. In Thailand and Peninsular Malaysia M. pectinata grows only in exposed areas or in relatively open scrub on or near the tops of isolated mountains at altitudes of 7502000 m. In the Riau and Lingga Archipelagos M. pectinata has also been found at sea level. M. pectinata is a thicket-forming fern adapted to poor sandy soils. Throughout its range, M. pectinata often grows with other thicket-forming ferns such as Dipteris conjugata Reinw., Pteridium aquilinum (L.) Kuhn and various Gleichenia and Dicranopteris species
50
(Tansley & Lulham, 1905; Parris et al., 1984). Its main associates in Thailand are Dipteris conjugata (see photo on page 68 of Niyomdham, 2000) and Gleichenia microphylla R.Br. The gametophytes of M. pectinata are relatively large and long-lived and, like many primitive fern species, can produce multiple embryos (Stokey & Atkinson, 1952). Note. The family and genus descriptions given above are based on specimens of Matoniaceae from throughout the familys range but the description of Matonia pectinata is based solely on the new Thai material. It is important to remember that this new Thai material represents only two plants and to be aware that the one in Trang province has laminae that are particularly small (2025 cm in diam., consisting of a central pinna 1518 cm long and 1014 shorter lateral pinnae) and stipes that are particularly thin (13 mm diam.). The description of M. pectinata in Flora Malesiana (Kato, 1998) reports rhizomes up to 8.5 mm in diam., stipes up to 180 cm long and laminae up to approximately 75 cm in diameter. It remains to be seen whether plants this large will also be found in Thailand. ACKNOWLEDGEMENTS We would like to thank the National Geographic Society for financial support for this work. We would also like to thank Orathai Kerdkaew for the illustrations, and Voradol Chamchumroon and Suwat Suwanachat for their assistance in the field. REFERENCES Boonkerd, T. & Pollawatn, R. 2000. Pteridophytes in Thailand. Office of Environmental Policy and Planning, Bangkok, Thailand. Holttum, R.E. 1954 (2nd ed. 1968). A revised Flora of Malaya, vol. 2, Ferns of Malaya, Government Printing Office, Singapore. Kato, M. 1993. A taxonomic study of the genus Matonia (Matoniaceae). Blumea 38: 167172. ________. 1998. Matoniaceae. Flora Malesiana, Series II, 3: 289294. Kramer, K.U. 1990. Matoniaceae. In: Kramer, K.U. & Green, P.S. (eds), The Families and Genera of Vascular Plants I: Pteridophytes and Gymnosperms, pp. 183185. SpringerVerlag, Berlin, Heidelberg, New York. Niyomdham, C. 2000. Plants of Hala-Bala (title in Thai). Amarin Printing and Publishing Public Co., Ltd. Bangkok. Nishida, H., Yoshida, A. & Nishida, M. 1998. Permineralized matoniaceous fossils from the Cretaceous of Japan. J. Jap. Bot. 73: 2634. Parris, B.S., Jermy, A.C., Camus, J.M., & Paul, A.M. 1984. The Pteridophyta of Mulu National Park. In: Jermy, A.C. (ed.), Studies on the flora of Gunung Mulu National Park, Sarawak, pp. 145233. Kuching Forest Department, Kuching.
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Stokey, A.G. & Atkinson, L.R. 1952. The gametophyte and young sporophyte of Matonia pectinata R.Br. Phytomorphology 2(2 & 3): 138150. Tagawa, M. & Iwatsuki, K. 1979. Flora of Thailand vol. 3, part. 1. Royal Forest Department, Bangkok. ________. 1985. Flora of Thailand vol. 3, part. 2. Royal Forest Department, Bangkok. ________. 1988. Flora of Thailand vol. 3, part. 3. Royal Forest Department, Bangkok. ________. 1989. Flora of Thailand vol. 3, part. 4. Royal Forest Department, Bangkok. Tansley, M.A. & Lulham, R.B.J. 1905. A study of the vascular system of Matonia pectinata. Ann. Bot. (Oxford), 19(76): 475519. Tryon, A.F. & Lugardon, B. 1990. Spores of the Pteridophyta. Springer-Verlag, New York, Berlin, Heidelberg.
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Figure 1. Matonia pectinata R.Br.: A. frond; B. rhizome; C. part of pinna with 4 intact sori; D. part of pinna showing the venation around one sorus (top segment) and under another (middle segment; sorus removed). A from C. Niyomdham et al. 6082 (BKF), BD from D.J. Middleton et al. 1992 (BKF). Drawn by O. Kerdkaew.
THAI FOR. BULL. (BOT.) 31: 5364. 2003.
An account of the Plantaginaceae of Thailand

JOHN PARNELL*
ABSTRACT. One genus, with a single native species, is recognised in the Plantaginaceae following revision of Thai and associated materials. Plantago comprises the widespread P. major which is taken to include P. asiatica (lectotypified here) and P. erosa (lectotypified here). Investigation of stomatal density, stomatal length, seed size, seed number, leaf-lobing, pseudo-pedicel presence, inter-flower spacing and corolla-lobe length revealed that only the latter showed significant difference between material usually assigned to P. asiatica and P. major; with the latter the larger. The detected difference is both small and the reverse of that expected from previous studies. Though field experience suggests that P. major is fairly frequent throughout Thailand the number of collections housed in herbaria is very few and the distributional data given are, necessarily, incomplete.
Plantaginaceae comprises three genera and 250260 species worldwide: Brougiera and Littorella contain three and one species respectively, the rest are all in Plantago. The family is cosmopolitan, being absent only from the Arctic and Antarctic. It is closely allied to the Scrophulariaceae and in recent years, on the basis of rather limited molecular, anatomical and more substantial chemical evidence, the two families have been considered as one (Judd et al., 1999). However, the most recent molecular evidence suggests that there is only weak support for the formation of this new clade (Soltis et al., 2000). Most recently, on the basis of DNA sequence data from the plastid genes rbcL, ndhF and rps2, Olmstead et al. (2001) propose the submergence of the Plantaginaceae within a weakly defined clade which they have called the Veronicaceae. As Olmstead et al. (2001) indicate the name Plantaginaceae postdates the name Veronicaceae but has priority as it is conserved: therefore, the correct name for this clade at family level is Plantaginaceae. At one point in their paper, Olmstead et al. (2001) accept but later they implicitly reject the concept that this clade (Veronicaeae) should be recognised at family level. It seems that Olmstead et al. (2001) prefer to retain the Plantaginaceae as a family, in the narrow conventional sense. Certainly, their analysis shows that the branch lengths linking Plantago and Veronica are very long (209 and 108 respectively) (Olmstead et al., 2001). Therefore, the fact that these genera cluster together may be a false indication of a close relationship due to the Felsenstein effect or long-branch attraction (Felsenstein, 1978, Judd et al., 1999). In these circumstances, it would appear sensible to retain the family Plantaginaceae as a separate entity until more data from more taxa becomes available. This is congruent with the opinions of Pilger (1937) & Rahn (1996) who clearly indicate that they consider the family monophyletic with the latter suggesting that the consistent presence of hairs in the leaf axils is an important autapomorphy.
*Herbarium, School of Botany, Trinity College, Dublin, Dublin 2, Ireland.
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A survey of herbarium material in A, AAU, ABD, BK, BKF, C, E, K and TCD yielded only 59 collections from Thailand, 55 of which are of P. major s.l. Though many collections were duplicated in more than one herbarium, the family is clearly severely under-collected. Therefore, the distributional data given herein are even more than usually tentative. Species descriptions are based on original observations, largely of herbarium material. As measurements have been taken from dried specimens they may differ slightly from fresh material. It is known that a number of species, particularly Plantago indica L., Plantago arenaria Waldst. & Kit., Plantago afra L. (listed as Plantago psillium L.) and Plantago ovata Forsk. are grown for pharmaceutical use as bulk laxatives (Leung, 1980) and that the mucilage from the husk is used as a thickener in some food products (Chua et al., 1994). P. afra and P. arenaria have escaped from cultivation, being found as weeds amongst a crop of cumin and P. ovata is cultivated in Thailand: therefore, all these species are included in this account and appear in the key. Surprisingly, Plantago lanceolata L. has not yet been recorded from Thailand nor any of the immediate surrounding countries and, therefore, is currently excluded. PLANTAGINACEAE Juss., Gen. Pl. 89. 1789 (Plantagines); Decne. in A.DC., Prod. 13: 693737. 1852; Pilger in Engl. & Diels, Pflanzenr. 4. 269: 1937. Upright to spreading annual or perennial herbs with short stems; Thai material never subshrubby. Leaves simple, spirally arranged, with parallel venation, margin entire or lobed to toothed. Petiole forming a sheath at the base; stipules absent, hairs always present in leaf axils. Bracts small, persistent. Flowers (3)4(5)-merous, in long or short spikes on long peduncles, usually hermaphrodite. Sepals fused, lobed. Corolla largely fused, scarious; lobes triangular, spreading and usually reflexed at maturity. Stamens usually 4 (solitary in Bougeria), equal, with long filaments, alternating with corolla-lobes, anthers 2celled, exserted, versatile, opening by longitudinal slits; connective prominent. Ovary superior of 2 carpels; ovules 1-many on an axile placenta. Style 1, long, stigma bifid, exserted. Fruit in all Thai species always a circumscissile capsule, the top segment falling off as a lid. Three genera (only Plantago in Thailand) and ca. 250260 species. Cosmopolitan. PLANTAGO L., Sp. Pl. 112. 1753; Decne. in A.DC., Prod. 13: 694. 1852. Inflorescences 37, spicate or, in some introduced species, capitate. Leaves mainly radicle, usually hairy, entire or lobed. (All native Thai material has a distinct blade and petiole, the latter is sometimes as long as the blade; introduced species may have linear
AN ACCOUNT OF THE PLANTAGINACEAE OF THAILAND (J. PARNELL)
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to narrowly ensiform leaves with the petiole not distinguishable from the blade.) Bracts triangular to ovate or obovate; glabrous or hairy, membranous to herbaceous, keeled, sometimes winged, usually green or brown. Flowers hermaphrodite. Corolla lobes reflexed at maturity, inconspicuous. Stamens 4, exserted; anthers often cauducuous. Fruit enclosed in the remnants of the perianth segments at maturity. Seeds angular, peltate. One native species in Thailand; three species potentially cultivated.
1. Inflorescence a spike, leaves with a distinct, broad blade and obvious petiole 1. P. major 1. Inflorescence capitate, leaves lacking a distinct broad blade and obvious petiole 2 2. Involucral bracts with long, silky, ciliate hairs 2. P. ovata 2. Involucral bracts lacking long, silky, ciliate hairs, though often with a long acumen 3 3. Inflorescence branches sparsely hirsute and minutely, but evidently, papillose, (i.e. long and very short hairs evident). Bracts of two types, the basal suddenly narrowing to a long acumen 34x as long as the base of the bract, the upper lacking a long acumen 3. P. arenaria 3. Inflorescence branches hirsute and not evidently papillose (i.e. hairs apparently the same length). Bracts ovate to lanceolate, all similar, the base never suddenly narrowing to a long acumen 4. P. afra
1. Plantago major L., Sp. Pl. ed. 1. 1:112. 1753; Decne. in A.DC., Prod. 13: 694. 1852; Hook.f., Fl. Brit. India 4: 705. 1885; Gagnep. in Lecomte, Fl. Indo-Chine 4: 1047. 1936; Pilger in Engl. & Diels, Pflanzenr. 4. 269. 4156: 1937; Matthew & Rani in Matthew, Flora of Tamilnadu Carnatic 2: 1285. 1983; H, Cyco Vitnam 2: 1109. 1993; Type: Herb. Linn. 144.1 (lectotype LINN!). P. asiatica L., Sp. Pl. ed. 1 1: 113. 1753; Pilger in Engl. & Diels, Pflanzenr. 4. 269: 5659: 1937; Ridl., Fl. Mal Pen. 2: 225. 1923; Alston in Trimen, Hand. Fl. Ceylon 6: 237. 1931; H, Cyco Vitnam 2: 1109. 1993. Type: Herb Linn. 144.4 (lectotype LINN!, selected here). P. major var. asiatica (L.) Decne. in A.DC., Prod. 13: 694. 1852; Plantago major var. asiatica (L.) Trimen, Hand. Fl. Ceylon 3: 389. 1895. P. erosa Wall. in Roxb., Fl. Ind. Ed. 1 (ed. Carey) 423. 1820, Decne. in A.DC., Prod. 13: 695. 1852; Pilger in Engl. & Diels, Pflanzenr. 4. 269. 6061: 1937; Dassan. in Dassan. & Clayton, Rev. Hndbk. Fl. Ceylon 10: 328330. 1996; Springate, in Grierson & Long, Flora of Bhutan 2: 1342. 2002. Type: Wallich 6412A, lower specimen (lectotype K-W!, selected here). Perennial herb, 520 cm tall, with a single basal rosette of leaves, rosette present on flowering. Bracts ovate to triangular, much smaller than the flowers, sometimes largely green other times mostly membranous with a green midrib. Flowers in long spikes 5 15(20) cm long, the lower often well separated from each other the upper usually congested. Calyx 22.5 mm long, oval, obtuse or acute, scarious, keeled, keel green when fresh, brown when dry. Corolla-lobes spreading, 11.2 mm, recurved; lilac to whitish. Stamens exserted, with caducous, cream-coloured anthers and fragile white filaments, connective 0.10.2 mm long, apiculate, prominent. Stigma ca. 2 mm long, white. Capsule 34 mm, the top 5060% forming a circumscissile capsule. Seeds black, somewhat angular (Fig. 1). Thailand. NORTHERN: Mae Hong Son (Kieo Lom), Chiang Mai (Mae RimSamoeng, Mae Chaem, Chiang Mai, Doi Chiang Dao, Doi Inthanon, Doi Khun Huai Pong, Doi Pa Hom Pok, Doi Suthep, Doi Pui, Khun Yam, Om Koi), Chiang Rai (Doi
56
Tung, Mae Chan), Nan (Muang District, Phu Kha), Lamphun (Ban Muang Nga, Doi Khun Tan), Lampang (Jae Son, Mae Mo), Phrae (Ban Nam Klai, Mae Krae), Phitsanulok (Phu Hin Rong Kla, Phu Miang); (Bannang Sata).
SOUTHWESTERN:
Kanchanaburi (Sai Yok);
PENINSULAR:
Yala
Distribution. Cosmopolitan. Ecology. Roadsides, open and disturbed habitats; largely recorded from elevations above 500 m and from damp or wet habitats. Vernacular. Ya en yuet (), En yuet () (Northern). Uses. Whole plant is powdered and applied to cuts to stop bleeding and infection and to treat strains. In Peninsular Malaysia a decoction of the plant is used to alleviate coughs. Notes. P. erosa Wall. in Roxb., P asiatica L. and P. major L. have all been reported from Southeast Asia. Rahn (1996) indicated that P. erosa was most likely to be a subspecies of P. asiatica and that P. asiatica and P. major differed in their chromosome number (2n=12 and 2n=24 respectively). Linneaus (1753) & Barnoud (1845) distinguished the latter two taxa on the basis that the scape of P. major was rounded and the flowers imbricate whilst the scape of P. asiatica was angled and the flowers were separated from each other along the scape. Taiken et al. (1993) suggested that these taxa differed in various features shown in Table 1. Table 1. Putative differences between P. asiatica and P. major (Taiken et al., 1993).
P. asiatica Stomatal number per mm2 Stomatal index Seed length (mm) (125)165(200) (20.1)27.2(36.4) 2 P. major (225)285(375) (20.5)26(30) 2
Rahn (pers. comm.) also indicated that he believed these species differed as follows: P. major: flowers sessile. Corolla lobes 1 mm. Seeds 634 per capsule. 2n=12. P. asiatica: flowers borne on a short pedicel-like extension of the calyx. Corolla lobes >1.2 mm. Seeds 59 per capsule. 2n=24. Examination of all available Thai material and material from surrounding regions and elsewhere has, with a solitary exception, not confirmed the existence of the putative differences between these species suggested by Linneaus (op. cit.) or Taiken (op. cit.) or Rahn (op. cit.) (Table 2). Table 2. Characteristics of measured material of P. asiatica and P. major; (n=100).
P. asiatica (Thailand) Mean Standard Error Corolla-lobe length Stomatal number per mm2 Stomatal length (mm) 0.81 0.02 143 8.74 0.256 0.007 P. major s.s. (all non-Thai material) Standard Error 0.9 0.03 154 8.7 0.247 0.006 F-value; p 4.25; 0.04 0.81; 0.38 0.84; 0.36
57
0.29; 0.59 0.13; 0.71
Seed length (mm) Seed number / Capsule
1.23 0.05 8.81 0.82
1.20 0.05 8.56 0.74
There appears to be no consistent or statistically significant difference between P. major s.s. and material normally assigned to P. asiatica in respect of stomatal density, stomatal length, seed length or seed number / capsule (all p>0.05; Table 2). Corolla lobe length is statistically different between the two species with the lobes of P. major 0.1 mm longer than P. asiatica on average (F=4.25; p0.5); this finding appears to contradict the suggestion of Rahn (op. cit.) that the corolla-lobes of P. asiatica are longer. Examination of Southeast Asian material attributed to P. asiatica, also revealed that the pedicel-like extension (pseudo-pedicel) of the calyx is not consistently present in duplicate collections of material. Also, whilst some specimens have flowers well separated along the base of the scape, others have the flowers congested and all intermediate conditions may be found. Leaf lobing is similarly inconsistently present and the angularity of the scape variable. Comparison of Southeast Asian material with undoubted material of P. major from Europe and elsewhere showed that material assigned to P. major subsp. intermedia (Gilib.) Lange has identical leaf-lobing to material of P. asiatica with leaf-lobes. Furthermore, material labelled as P. erosa by Rahn (e.g. Sorensen, Larsen & Hansen 1777 (C)) tends to have high stomatal densities ( 250 per mm2), often has lobed leaves and the basal flowers in the inflorescence well separated out from the rest. However, as stated before all intermediate conditions between this form and typical P. major exist. On the basis of the available evidence, I do not believe that P. asiatica warrants recognition at the species level: indeed at present, in my view, even subspecific rank appears difficult to justify - at best, varietal rank might be warranted for this taxon. If varietal rank is accepted the correct name for such material is P. major var. asiatica (L.) Decne. This conclusion is in concord with those of Hooker (1885), Gagnepain (1936), Backer & Bakhuizen van der Brink (1965) & Chua et al. (1994). Though Trimen (1895) was inclined to recognise this taxon at specific level, he did not do so, preferring varietal status. As I do not recognise P. asiatica as a separate species I also cannot follow Rahns suggestion that P. erosa be treated as a subspecies of P. asiatica. My view of the pattern of variation P. major is well summarized by Backer & Bakhuizen van der Brink Jr (1965) who state that the extreme forms differ so much that they might be taken for different species, were it not that they are connected by a series of intergrades. The type of P. asiatica has not been chosen. There are two relevant specimens of P. asiatica (144.4 & 144.3) in LINN and one of P. major (144.1). Of these, 144.4 is the more complete, with many mature and younger flowers - it shows all the features Linnaeus thought distinctive and I therefore have typified by reference to that specimen. 144.1 clearly corresponds to Linnaeuss description of P. major and matches current thinking on this species. It has been already selected as the type. The type of P. erosa is indicated as Wallich 6412 by Dassanayake in Dassanayake & Clayton (1996). However, no formal lectotypification has been made and 6412 is not an unique reference. The sheet bearing the label 6412A in K-W is labelled P. erosa and consists of two specimens. The upper has only one mature leaf and is somewhat small and immature.
58
The lower is well preserved, mature and with the features of P. erosa that Wallich thought distinctive. Presumably it should be designated as 6412B (there is, apparently, no sheet so labelled in K-W). Therefore I have nominated this specimen as the lectotype. All the material (four specimens) on sheet 6412C, D also corresponds to the concept of P. erosa of Wallich. However, 6412E and 6412Suppl. correspond to P. major L. 2. Plantago ovata Forsk., Fl. Aegypt. Arab. 31. 1775; Decne. in A.DC., Prod. 13: 706. 1852. Type: Egypt, Forsskal 249, partly (syntype C, not seen), 250 (syntype C, not seen) & 253 (syntype C, not seen). Annual or biennial, erect herb 415 cm tall with a rosette of leaves which are upward-pointing when flowering. Leaves linear, narrow, almost ensiform, very densely hirsute, sometimes woolly, about as long as flower heads. Flowers in ovate to oblong heads of knobbly appearance. Bracts with long, silky, ciliate hairs which are normally numerous. Thailand. Cultivated. Distribution. Europe to the Middle East and India. Ecology. N/A. Uses. Used in the pharmaceutical industry and widely cultivated. Notes. Used as a source of laxatives and as a source of food thickeners; it may be sold as Psyllium husk. 3. Plantago arenaria Waldst. & Kit., Pl. Rari. Hung. 1: t. 51. 1801; Decne. in A.DC., Prod. 8: 735. 1852; Rahn, Bot. J. Linn. Soc. 120: 184. 1996. Type: not located. P. indica L., Sys. Nat. ed. 10. 896. 1759. nom illegit.; Pilger in Engl. & Diels, Pflanzenr. 4. 269. 419. 1937. Plantago psyllium L., Sp. Pl. ed. 1. 1: 115. 1753, nom ambig. Annual, usually erect herb, 30(50) cm tall, lacking a basal rosette of leaves when flowering. Upper parts minutely papillose and sparsely and shortly hirsute. Leaves linear, opposite, with fascicles in the axils and appearing as a whorl of equally sized leaves or as a whorl of leaves with two members of the whorl a little larger than the rest, shortly hirsute. Flowers in numerous ovate to oblong heads of smooth appearance, with the flowers closely packed. Bracts winged, of two types; the basal with a very long acumen 34 x longer than the rest of the bract; the upper bracts lacking a long acumen. Thailand. Cultivated. Distribution. Europe to the Middle East and India. Ecology. N/A.
59
Uses. Used in the pharmaceutical industry and cultivated. An escape from cultivation in Chiang Mai (Fang). Notes. Used as a source of bulk laxatives and of food thickeners. The nomenclature of this species is confused and probably worth rehearsing as the incorrect name continues to be applied in herbaria in the region and elsewhere despite Rahn (1996), whose work on the phylogeny of the group remains the most up-to-date summary. Panigrahi (1975) showed that the name P. indica was coined by Linneaus in 1759 as a new name for a species he had already described in 1753 as P. psillium: P. indica is therefore a superfluous and invalid new name. Unfortunately, Verdcourt (1969, 1971) has also shown that the name P. psillium is a nomen ambiguum and therefore cannot be used: a view also ascribed to by Rahn (1996). The first available, valid epithet is therefore that coined by Waldstein & Kitaibel. Under the current ICBN rules formal application should, probably, be made to reject P. psillium, as until this is done it remains the correct name for this taxon. However, psillium is certainly a taxonomically confusing epithet irrespective of its nomenclatural priority. The epithet psillium is widely applied to Plantago material used in Asian pharmaceutical preparations. Its usage in these preparations adds to the confusion as manufacturers report that the material is P. ovata. 4. Plantago afra L. Sp. Pl. ed. 2. 168. 1762.; Rahn, Bot. J. Linn. Soc. 120: 184. 1996. Type: not located. P. psyllium L. Sp. Pl. ed. 2. 1: 167. 1762 et auct. mult. non L., Sp. Pl. ed. 1. 1: 115. 1753, Decne. in A.DC., Prod. 13: 734. 1852; Hook.f., Fl. Brit. India 4: 707. 1885; Pilger in Engl. & Diels, Pflanzenr. 4. 269. 424. 1937. Annual herb lacking a basal rosette of leaves when flowering, 820(40) cm tall. Upper parts covered in short hairs and sparsely or densely glandular. Leaves linear, opposite, with small fascicles in the leaf-axils and so appearing as a whorl of leaves with two members of the whorl larger than the rest or a whorl of equally sized leaves. Flowers in ovoid spikes; of somewhat knobbly, somewhat loose appearance. Bracts oval to lanceolate, hairy, all similar and lacking a long acumen. Thailand. Cultivated. Distribution. Europe to the Middle East and India. Ecology. N/A. Uses. Used in the pharmaceutical industry and cultivated. Notes. Used as a source of bulk laxatives and as a source of food thickeners. The nomenclature of this species is complex, ties in with that of the previous species, and is probably worth restating as the incorrect name continues to be applied in herbaria in the region and elsewhere. Verdcourt (1969, 1971) showed that Linnaeus used the name P. psillium in two different senses. Unfortunately, the name which is widely used is based on the later concept of Linnaeus which appeared in the second (1762) edition of Species Plantarum. The name P. psillium as used in the first edition of 1753 is, in fact, an earlier name for what has
60
been called P. indica (see above). Verdcourt (op. cit.) therefore rejected P. psillium as a nomen ambiguum. This leaves P. afra as the first available, valid name (Verdcourt, 1969, 1971). This species is often confused with P. arenaria. REFERENCES Backer, C.A. & Bakhuizen van der Brink, Jr. R.C. 1965. Flora of Java (Spermatophytes only) 2: 446. N.V.P. Noordhoff, Groningen, The Netherlands. Barnoud, F.M. 1845. Monographie Gnrale de la Famille des Plantagines. Forten, Paris. Chua, K.S., Tan, H.T.W. & Turner, I.M. 1994. The Angiosperm Flora of Singapore: Part 3 Plantaginaceae. Gard. Bull. Singapore 46: 103107. Felsenstein, J. 1978. Cases in which parsimony and compatibility methods will be positively misleading. Syst. Zool. 27: 401410. Gagnepain, F. 1936. Plantaginaceae. In: Lecomte, H. & Humbert, H. (eds), Flore Gnrale de lIndo-Chine 4: 1046. Masson et Co., Paris. Hooker, J.D. 1885. Flora of British India 4: 705706. Reeve & Co., London. Judd, W.S., Campbell, C.S., Kellogg, E.A. & Stevens, P.F. 1999. Plant systematics. A phylogenetic approach. Sinauer Associates Inc., Massachusetts. 464 p. Leung, A.Y. 1980. Encyclopedia of common natural ingredients used in food, drugs, and cosmetics. John Wiley & Sons, New York. Linnaeus, C. 1753. Species Plantarum. A Facsimile of the First Edition. Ray Society, 1957. Olmstead, R.G. dePamphilis, C.W., Wolfe, A.D., Young, N.D., Elisons, W.J. & Reeves, P.A. 2001. Disintegration of the Scrophulariaceae. Amer. J. Bot. 88: 348361. Panigrahi, G. 1975. A note on Plantago exigua (Plantaginaceae) and related taxa. Kew Bull. 30: 669673. Pilger, R. 1937. Plantaginaceae. In: Engler, A. & Diels F. (eds), Das Pflanzenreich 4: 269 Neidriech. 466 p. Rahn, K. 1996. A phylogenetic study of the Plantaginaceae. Bot. J. Linn. Soc. 120: 145198. Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H., Hoot, S.B., Fay, M.F., Axtell, M., Swensen, S.M., Prince, L.M., Kress, W.J., Nixon, K.C. & Farris, J.S. 2000. Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Bot. J. Linn. Soc. 133: 381461. Taiken, Z., Tanaka, T. & Tingyuo, K. 1993. Studies on the Plantago plants in China. Leaoning, China. Trimen, H. 1895. A Handbook to the Flora of Ceylon. Vol. 3. Dulua & Co., London.
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Verdcourt, B. 1969. A new plantain from East Africa. Kew Bull. 23: 509.
Verdcourt, B. 1971. Plantaginaceae. In: Milne Redhead, E. & Polhill, R.M. (eds), Flora of Tropical East Africa. Crown Agents for Overseas Governments and Administrations, London. 7 p. THAI SPECIMENS EXAMINED Plantago afra L. Sutheesoen, S., 3354 (BK); Umpai, 566 (BK). Plantago arenaria L. Umpai, 569 (BK). Plantago major L. Anderson, E.F. 5281 (BKF); Barnid, s.n. (TCD) Java; Bragg, K. 16 (A,L); Brun, Bjornland, Schumacher, 122 (C), 440 (C); CC. & CH. 353 (BKF); Chermsirivathana, C., 375 (BK); Chantaranothai, P., Parnell, J., Simpson, D. & Pooma, R. 1014 (TCD); Hansen, B. & Smitinand, T. 12761 (C), 12841 (C), 12891 (AAU, BKF, E, K, L); Iwatsuki, K., Fukuoka, N. & Chintayungkun, A. T-9569 (BKF, L); Kermode, C.W.D. 16656 (K); Kerr, A.F.G. 2912 (E, TCD); Khantchai, 370 (BKF); Koyama, H., Nagamusu, H. T-40140 (A, BKF); Koyama, H., Terao, H. & Wongprasert, T. T-32590 (BKF); Larsen, K., Larsen, S.S., Nielsen, I. & Santisuk, T. 41843 (AAU); Larsen, K., Larsen, S.S., Norgaard, C.T., Pharsen, K., Puudjaa, P. & Ueachirakan, W. 44378 (AAU), 44720 (AAU); Larsen, K., Santisuk, T. & Warncke, E. 1873 (AAU, BKF, E), 2931 (AAU, BKF, E); Larsen, K., Smitinand, T. & Warncke, E. 1020 (AAU, BKF, C); Maxwell, J.F. 87-1477 (L), 94-680 (A, BKF), 96-1245 (A, BKF), 96-1277 (A, BKF); Murata, G., Iwatsuki, K., Phengklai, C. & Charamphol, C. T-15371 (BKF); Niyomdham, C. 75 (AAU, BKF, C, L); Paisooksantivatana, y-1695-85, y-1720-86, y-2304-89 (BK); Phengkhlai, C. T-49886 (BKF), T-49891 (BKF); Samrong, S.P.N.S. 17 (BKF); Shimizu, T., Toyokuni, H., Koyama, H., Yahara, T. & Santisuk, T. 19175 (L); Smitinand, T. 2646 (BKF); Sorensen, T., Larsen, K. & Hansen, B. 1777 (C), 2859 (A, C), 4837 (C); Suvarnakoses, P. 1702 (BKF); Suvatoa, L.A., 39 (BK); Tagawa, M., Iwatsuki, K., Koyama, H., Fukuoka, N., Nalampoon, A. & Chinayungkun, A. T-9292 (BKF); Takahasi, H. T-32590 (BKF), T-62552 (A, BKF), T-61863 (A, AAU, BKF); Tsugaru, S. T-61863 (BKF); Umpai, 463 (BK); Vacharee, 478 (BK); van Beusekom, C.F., Geesink, R., Phengklai, C. & Wongwan, B. 4721 (BKF, C, K, L); Winit, K., 1808 (A), 1873 (BK); s.n. (BK35192) (BK). Plantago ovata L. Umpai, 508 (BK)
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REPRESENTATIVE, NON-THAI SPECIMENS EXAMINED Plantago afra L. Adomovic, s.n. (K) Greece; Al Ani, H.A., 9823 (K) Iraq; Ash, J., 2041 (K) Ethiopia; Asplund, E., 447 (K) Canary Islands; Ball, J. s.n. (TCD) Algeria; Balls, E.K. & Gowerlay, W.B., B713 (K) Turkey; Bally, P.R. & Melville, R., 157671 (K) Somalia; Bang, A.M., 1965 Bolivia; Bourgeau, E., 526 (K) Canary Islands; Bove, s.n. (TCD) Algeria; Bramwell, D. 787 (TCD) Canary Islands; Bramwell, D., Richardson, I.B.K. & Murray, B.G., 186 (K) Morocco; Casey, E.C., 243 (K) Cyprus; Chevallier, L. 474 (TCD) Algeria; Davis, P.F., 41426 (K) Turkey; Deane, H., s.n. (K) India; Ellman, E. & Hubbard, C., 398 (K) Spain; Evans, I., 44 (K) Ethiopia; Gaillardot, 2176 (TCD) Syria; Gillett, J.B., 15257 (K) Jordan; Gregor, A.G., s.n. (K) U.K.; Houck, D.F. 363 (BKF) U.S.A.; Joad, G.C., s.n., s.n., s.n., s.n. (K) France; Kotschy, T. 50 (TCD) Iran; Lehmann, F.C., 6317 (K) Ecuador; Mandon, G., s.n. (K) Madeira; Mosseray, R., s.n. (K) France; Metcalf, s.n. (K) Italy; Mooney, H.F., 2836 (K) Ethiopia; Morton, L., 3699 (K) Iran; Muschler, R., s.n. (K) Egypt; Newbould, J.B., 5687 (K) Kenya; Qazilbash, N.A., 2418 (K) India; Sine nom. s.n. (K) Britain; Trethewy, A.W., 186 (K) Algeria; Whiting, M.H. & Richmond, K., 56 (K) Morocco; Wright, E.P. s.n. (TCD) Algeria; Yip, H.G. 186 (BKF) China. Plantago arenaria Waldst. & Kit. Ball, J. s.n. (TCD) Italy; Bemrose, C.J.V., 32 (K) Britain; Bleak, A.A., s.n. (K); Black, A.A. s.n. (K) Channel Islands; Blom, C., s.n. (K) Sweden; Bogdan, V., 4169 (K) Russia; Boros, A., 100 (K) Hungary; Cadet, T., 4545 (K) Reunion; de Wilde, W.J.J.O. & de Wilde-Duyfjes, B.E.E., 9080 (K) Ethiopia; Favrat, L. & Barbey, W., s.n. (K) Switzerland; Fishwick, R.W., 18 (K) Iran; Gogina, E.E. 178 (TCD) Russia; Gregor, A.G., s.n. (K) Britain; Heginbotham, C., s.n. (K) Britain; Horwood, E.K. 61/42 (TCD) France; Hukseaeb, A., s.n. (K) Russia; Johnston, T.S., s.n. (K) Britain; Lambert, V., 1926 (K) Belgium; Mcel, L., s.n. (K) Slovakia; Melvill,e R., s.n. (K) Cult.; Mennell, H., s.n. (K) Channel Islands; Milne-Redhead, G. & Shaw, H.K.A. 1832 (K) Britain; Pickler, s.n. (K) Austria; Porta, s.n. (K) Italy; Scheppig, C., s.n. (K) Germany; Schwarz, Z., 472 (K) Poland; Sennen, F., 3483 (K) Spain; Shaw, H.K.A., 1832 (K); Sine nom. s.n. (K) Cult.; Tauscher, J.A. s.n. (TCD) Hungary; Tomassini, M. s.n. (TCD) Italy; Turrill, W.B., K557 (K) Bulgaria; Ussher, C.A., s.n. (K) Ireland; Verdcourt, B., 4498 (K) France; Wilkinson, J.W., H2229 (K) Britain; Wisniewski, T., s.n. (K) Romania. Plantago major L. Akeroyd, J.R. & Doogue, D., 7709 (TCD) Ireland; Akeroyd, J.R. & Preston, C.D., 881 (TCD) Greece; Averyanov, L, Ban, N.T., Binh, N.Q., Budantzev, A., Budantzev, L., Hiep, N.T., Loe, P.K., Tam, N.X. & Yakovlev, G., VH411 (AAU) Vietnam; Ball, P.W. & Chater, A.O., 108 (TCD) Serbia; Bonnet, s.n. (TCD); Bor, N.L., 229 (TCD) Ireland; Boufford, D.E., Bartholomew, B., Li, G. & Zhu, G.H., 24481 (AAU) China; Chao, J.M & Kao, M.T., 6233 (AAU) China; Chater, A.O. & Moore, D.M., 439 (TCD) Spain; Chow, K.S., 35 (AAU)
63
China, 70 (TCD) China, 727 (AAU) China; Cirtu, D. & Cirtu, M., 724 (TCD) Romania; Coulter, T.C., 571 (TCD) U.S.A.; Dessanayake, M.D., 590 (TCD) Ceylon; Dixon, H.H., s.n. (TCD) Ireland; D.N.F.C., s.n. (TCD) Ireland; Florence, J., 10788 (TCD) Pitcairn; Gage, A.T., 63 (ABD) India; Ghosh, S.K., s.n. (AAU) India; Gupta, N.C., 36 (TCD) India; Inamasu, Y., 423 (AAU) Japan; Jeppesen, S. & Ljtant, B., 706 (TCD) Denmark; Johnson, E., s.n. (TCD) India; Kermode, C.W.D., 16656 (K) Burma; Kerr, A.F.G., 20968 (K) Laos; Kings Collector 155 (ABD) India; Larsen, K., s.n. (AAU) Ceylon; Lau, Y.E., 210 (AAU) China; Loc, P.K., Hoang, P.H. & Averyanov, L., CBL1325 (AAU) Vietnam, 1597 (AAU) Vietnam; Ling, P.P. & Yao, K., 39 (AAU) China; Ljtnant, B. Nielsen, I. & Esbensen, W., 145 (TCD) Denmark; McKee, H.S., 6141 (K) Burma; MKen, M.J., 110 (TCD) South Africa; Matthew, K.M., 40550 (AAU) India; Maxwell, J.F., 78-202 (AAU) Malaysia; 78236 (TDC) Singapore; Moore, A.G., s.n. (TCD) Ireland; Nakano, T., s.n. (AAU) China; Olney, S.T., s.n. (TCD) U.S.A.; Pearson, C., s.n. (TCD) Ireland; Pakuism, C.E., 1161 (K) Burma; Proskuriakova, G.M., 179 (TCD) Russia; Rahl, A., s.n. (ABD) Burma; Ravenel, H.W., s.n. (TCD) U.S.A.; Reporter on Economic Products, 10355 (TCD) India; Robinson, C.B., 1360 (K) Cambodia; Santisuk, T., s.n., (BKF) U.S.A.; 383 (BKF) China.; Strachey, R. & Winterbottom, J.E., s.n. (TCD) India; Thomson, T., s.n., s.n. (TCD) India; Thwaites, G.H.K., 2246 (TCD) Sri Lanka; Watts, W.A., s.n. (TCD) Ireland; Yao, K., 10441 (AAU) China; Yip, H.G., 186 (AAU, BKF) China. Plantago ovata L. Bornmller, J., 589, 590 (K) Iran; Bourgeau, E., 1662 (K) Spain; Dinsmore, J.E., 18456 (K) Syria; Epling, E., Ellison, L. & Anderson, H., s.n. (K) U.S.A.; Hoover, R.F., 4289 (K) U.S.A.; Hudson, R., 2936 (K) Kenya; Kotschy, T. 836, 30.61 (TCD) Iran; Nelson, A. & Nelson, R.A., 1235 (K) U.S.A.; Nuttall, L.W., 600 (K) U.S.A.; Purpus, C.A., 120 (K) Mexico; Schweinfurth, G., 331 (K) Egypt; Sine nom. s.n., (K) Spain; Syngrasside, O., 313 (K) Cyprus; Townsend, C.C., 65/140a (K) Jordan; Tribe, A.R., T27 (K) Somalia; Wiggins, I.L. & Wiggins, D.B., 15792 (K) Mexico.
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Figure 1. Seed of Plantago major L. (Chantaranothai, P., Parnell, J., Simpson, D. & Pooma, R. 1014 (TCD)).
THAI FOR. BULL. (BOT.) 31: 6574. 2003.
Thai Rubiaceae with hooks and thorns

CHRISTIAN PUFF* & VORADOL CHAMCHUMROON**
ABSTRACT. Several woody Thai Rubiaceae (ca. 30 taxa) are distinguished by the presence of thorns or hooks (curved or recurved thorns). Most of these taxa belong to the tribe Gardenieae (Catunaregam, Ceriscoides, Dioecrecis, Fagerlindia, Oxyceros, Tamilnadia), two to the Vanguerieae (Canthium, Meyna), and one to the Coptosapelteae (Uncaria). The climbing hooks of Uncaria spp. morphologically represent modified inflorescence stalks, while thorns are vegetative lateral shoots with limited growth in all other taxa. In Ceriscoides, thorns are short lateral branches which, in turn, bear very abbreviated leafy and/or flowering shoots. In the monotypic Tamilnadia, short lateral branches terminally bear one or two pairs of short thorns. In the monotypic Dioecrescis, paired thorns are primarily on lateral shoots which occur in clusters in the lower part of the trunk. In the remaining taxa, thorns are short shoots produced in the leaf axils of long shoots, often together with leafy and/or flowering short shoots (accessory bud/shoot formation; thorns then always above the leafy and/or flowering brachyblasts; one or the other may not be developed). Thorns are either straight or curved (sometimes within a genus, e.g. Canthium spp.). The presence of curved thorns or hooks is associated with a climbing or straggling habit (Uncaria; Thai Oxyceros spp.; amongst the thorny Thai Canthium spp., only those with such a habit have curved thorns). Noteworthy is the presence of two different types of hooks (and shoot morphology) within Oxyceros. A key to the taxa is provided.
INTRODUCTION Thorns, either solitary and straight or sometimes branched, occur in a large number of angiosperm families. Their occurrence is not necessarily a generic character but may be confined to a number of species of a genus, often providing a good field character for identifying them. Unfortunately, their presence is sometimes omitted in flora descriptions, which are often based on herbarium specimens, because the thorns are sometimes only found on older, non-collectable stem parts. A small selection of examples from the Thai flora include families such as Apocynaceae (Carissa; Middleton, 1999: fig. 4A), Euphorbiaceae [Shirakiopsis indicum (syn. Sapium indicum), with straight thorns on older stems], Flacourtiaceae (Flacourtia and Scolopia spp.), Guttiferae (Cratoxylum spp.), Leguminosae (e.g. in the introduced and cultivated thorn of Madras, Pithecellobium dulcis), or Rutaceae (Citrus and Feroniella spp.). Hooked thorns (curved or recurved thorns), too, have independently evolved in woody taxa of a number of families and typically occur in climbing or straggling shrubs. Examples from the Thai flora include Artabotrys spp. (Annonaceae), Ancistrocladus tectorius (Ancistrocladaceae), or Indorouchera griffithiana (Hugoniaceae, or Linaceae s.l.). In the Rubiaceae, nine of the approximately 100 genera occurring in the flora of Thailand area have, in total, ca. 30 woody taxa which develop either straight or curved or recurved or hook-like thorns. The majority of these belong to the tribe Gardenieae (Catunaregam, Ceriscoides, Dioecrecis, Fagerlindia, Oxyceros, Tamilnadia), one of
*Institute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna, Austria. **The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Bangkok 10900, Thailand.
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the largest in the flora area; the others to the tribes Vanguerieae (Canthium, Meyna), and Coptosapelteae (Uncaria) (see Table 1). The main aim of this paper is to describe and compare the thorny structures in these taxa, and to provide information on how these thorn-bearing taxa differ from each other. The article is especially intended to aid field botanists in identifying the genera. A key to the thorn-bearing taxa is also provided. Table 1. Survey of Thai Rubiaceae with thorns and hooks. Column Tribe: GAR. Gardenieae; VAN. Vanguerieae; COP. Coptosapelteae
Taxa Catunaregam (all 4 Thai taxa) Ceriscoides (all 3 Thai taxa) Dioecrecis erythroclada (monotypic genus) Fagerlindia (all Thai taxa) Oxyceros (all 5 Thai taxa) Tamilnadia uliginosa (monotypic genus) Canthium (ca. 10 taxa; the remaining without thorns) Meyna (all 3 Thai taxa) Uncaria (all Thai taxa) Tribe GAR. GAR. GAR. GAR. GAR. GAR. VAN. VAN. COP. X X X X X straight thorns X X X X X curved or recurved thorns, hooks
A note on terminology In botanical literature, the terms thorn and spine are often used interchangeably, not taking account the origin of the organs in question. In the present context, the following definition, based on traditional morphology, is applied: Thorns (either straight or hooked, solitary or branched) are modifications of shoots, typically abbreviated shoots with limited growth. Being shoots, they have a normal shoot structure, i.e. have phloem and xylem (this definition largely corresponds to that in Tropical Woody Rubiaceae, the standard work describing Rubiaceae character states, cf. Robbrecht, 1988: 41). Spines, in contrast, are sharp-pointed outgrowths of primarily epidermal and subepidermal tissues, without contributions from the woody body. Examples of the latter are the vicious spines on the leaves of the palm Elaiodoxa, or the sharp-tipped outgrowths on the stems of Bombax spp. (Bombacaceae).
THAI RUBIACEAE WITH HOOKS AND THORNS (C. PUFF & V. CHAMCHUMROON)
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OBSERVATIONS In Thai Rubiaceae, thorns are either straight, curved or recurved (sometimes within a genus, e.g. Canthium spp.). The presence of curved thorns or hooks is associated with a climbing or straggling habit (Uncaria; Thai Oxyceros spp.; amongst the thorny Thai Canthium spp., only those with such a habit have curved thorns; see below). Noteworthy is the presence of two different types of hooks (and shoot morphology) in Oxyceros (see below). Uncaria Schreb. Recurved or climbing hooks occur in all species of the genus. They are normally only found on the paired lateral branches of the woody lianas (i.e. not on the main axes), and the hooks themselves are typically paired at the nodes (cf. fig 1A). Morphologically, they represent modified fertile shoots, i.e. they are modified inflorescence stalks. Observations on living plants sometimes reveal that the production of stalked, axillary, head-like inflorescences eventually stops and that the stalks are transformed into hooks (cf. fig. 1B, arrow). A relevant ontogenetic study confirming this has been published for a Madagascan species of the genus (Guillaumin, 1931). The modification of inflorescence stalks and axes into climbing hooks also occurs in several other plant families. A well known example is the Annonaceae genus Artabotrys. As opposed to the axillary hooks of Uncaria, the inflorescence hook is terminal in Artabotrys, and growth continues via a lateral shoot (sympodial-monochasial growth). In view of the ontogenetic evidence and documentation provided here, Ridsdales (1998) description of vegetative lateral branches modified into hooks cannot be upheld. In all other thorny rubiaceous taxa, thorns are lateral shoots with limited growth. As thorn morphology, and the arrangement and number of thorns are not uniform, genera or groups of genera can be distinguished quite easily: Ceriscoides (Hook.f.) Tirveng. All species of this genus of small trees produce thorns (cf. Reza 2001). Thorns tend to be of equal length and regularly arrangement (i.e. in decussate pairs) on younger branches, but older stems often have a very untidy appearance because some lateral shoots ending in a thorn have become relatively long (to ca. 10 cm) while other thorns remain short (only to ca. 34 cm). Characteristic of this genus is that thorns or thorn-tipped lateral branches, in turn, bear leafy or fertile short shoots. Fig. 1D illustrates this for C. sessiliflora, where a lateral branch ending in a thorn is seen which bears an abbreviated shoot with an inflorescence of which only one flower is left. Fruits often appear to sit on the thorny branches, but a closer examination reveals that they are produced on very short and much abbreviated lateral shoots. Dioecrescis Tirveng. In this monotypic tree genus (D. erythroclada; cf. Tirvengadum, 1983), paired, straight thorns are normally found on most lateral branches of young individuals (Fig. 1E, small tree on the right). On older trees, thorns are typically seen on clustered lateral branches (presumably having arisen due to serial bud/shoot formation)
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which come from the basal parts of the main trunk. Often, several tiers of such thornbearing branch clusters can be observed (Fig. 1E, arrows). The life-span of these branches is limited, so that very old individuals may lack thorns altogether. The presence of these thorn-bearing branches along the lower part of the trunks could be interpreted as a protection against browsing animals. Thorn-bearing branches normally have a regular appearance as all thorns are of roughly the same length (often ca. 2 cm), paired and arranged in a decussate fashion. Tamilnadia Tirveng. & Sastre. Branches of this monotypic tree genus (T. uliginosa; cf. Tirvengaum & Sastre, 1979) bear decussately arranged, short lateral shoots, each of which terminally produces one or two pairs of short thorns (Fig. 1C). Leaves and flowers and fruits are typically produced on very abbreviated shoots immediately below the thorns. Fig. 1C, inset, shows these knob-like brachyblasts, from which the fruit depicted below was removed. Fagerlindia Tirveng. and Oxyceros Lour. These two genera are closely allied and, as noted by Ridsdale (1985), the architecture of the two is basically very similar. Fagerlindia typically has erect main axes (trunks), while in Oxyceros the main axes bend over (straggling or climbing habit). In both, the main shoots bear regularly arranged, decussate lateral branches which, in turn, bear thorns (no thorns on the main trunks!). Fagerlindia invariably has straight thorns, while the Thai species of Oxyceros always have curved or recurved thorns (Oxyceros species with straight thorns are known from Indochina and Southwest China, species with no thorns from Vietnam and Sri Lanka; cf. Tirvengadum, 1982; Ridsdale, 1985, 1998). Fagerlindia shows accessory bud/shoot formation, which is defined as follows: two, buds, rather than one, are produced in the axil of a long shoot leaf, the two being arranged on top of each other (serial bud/shoot formation, as opposed to collateral accessory buds, such as in the banana, Musa spp.). The upper develops into the short-shoot thorn, the lower (i.e., the one immediately above the leaf axil) into a very abbreviated leaf- and/or flower-bearing short shoot. The association between these two kinds of shorts is visible in Fig. 1G (although the leaves in whose axils they have developed have already fallen off). This feature is absent in Oxyceros. Only one bud - the one developing into hooked thorn-is developed in the axil of a foliage leaf (cf. Fig. 1H). Moreover, the Thai species of Oxyceros, show a dimorphic pattern. Whilst in three of the species, the hooked thorns are regularly arranged, i.e. found in pairs at each node of a lateral branch, and are of uniform size (cf. Fig. 1H), this is not so in O. bispinosus and O. scandens. In these two species, the hooks at the first (i.e. basal) node of a lateral axis are recurved and fang-like (see illustration on the right), and both the basal lateral branch internode and this thorn pair are thick and massive (Fig. 1I). The next one or two nodes of the lateral branch usually bear the normal, much smaller, thinner hooked thorns, and the following nodes are always hook less. This undoubtedly is a much more sophisticated climbing aid than in the other taxa. Strictly, the fang-like hooks face downward, but as the plants habit is scandent, shoots with these hook-bearing branches hang downward, as depicted in Fig. 1I.
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Like Fagerlindia (above), all remaining taxa (Catunaregam, Canthium, Meyna) show accessory bud/shoot formation, i.e. thorns and abbreviated leafy and/or flowering shoots are in immediate vicinity to each other. Catunaregam Wolf. A genus of shrubs or small trees with thorns on erect and plagiotropic shoots. Thorns are typically arranged in decussate pairs, but in species such as C. spinosa, where leaf arrangement is variable (decussate and in whorls of three on one plant), they may be both paired and in threes. In the mentioned species, erect shoots often show ternate leaf and thorn arrangements, while plagiotropic shoots bear decussate leaves and thorns. The thorn pairs develop before the leafy and/or fertile brachyblast pairs, so that on younger shoots only thorn pairs are visible. As the buds from which the thorns develop are some distance away from the subtending foliage leaf, thorns are in a somewhat supra-axillary position. The ontogenetically younger leafy or fertile brachyblast does not always develop. While thorns and brachyblasts are typically produced in pairs, development of either one thorn or one brachyblast of a pair may be suppressed, fail or be disturbed by external influences. The result is a rather irregular appearance, with solitary thorns intermingled with paired thorns along a long shoot. This seems to happen quite frequently in species of Catunaregam. Figure 1F, showing four nodes of a branch of C. tomentosa, illustrates this: at the node on the left side, only one thorn is developed but a pair of leafy brachyblasts (although not equally well developed) is present; at the node on the right side, thorns are not developed at all; on the second node from the right, the thorn pair is developed, but one of the short shoots of a brachyblast pair has failed to develop. Canthium Lam. With regard to Thai Canthium, there are two major problems: (1) the generic limits are not resolved, and (2) there is no recent species-level revision (see also Chamchumroon & Puff, 2003 for further comment). Data given here are preliminary. Of the numerous Canthium species listed in Craib (1932: 135145), only ca. 10 are species with thorns, whereas the remainder are unarmed species. At least in some of the thorny taxa, paired, rather massive thorns are present on orthotropic shoots and smaller thorns on lateral branches; in other species thorns are confined to lateral branches. In the group of taxa with thorns on lateral branches, there are species with straight thorns (plants characterized by an erect habit), and species with recurved, hook-like thorns, all of which have a straggling, scrambling or climbing habit. Straight thorns on lateral branches are often short to very short (less than 1 cm long) to hardly discernible. The usually paired thorns are, as in, for example, Fagerlindia and Catunaregam, produced immediately above the leaf and/or fertile much contracted short shoots. Meyna Link. A small Afro-Asiatic genus of ca. 10 poorly delimited species of shrubs and small trees. All (including the three ill-defined species occurring in Thailand) possess paired or, less commonly, ternately arranged thorns.
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The genus is reputed to be very close to Canthium s. str. (Verdcourt & Bridson 1991: 859), and shoot structure is very similar. Paired thorns are borne immediately above leafy and/or fertile short shoots on long shoots (lateral branches), but development of the leafy/fertile brachyblasts may occasionally be suppressed (as seen in Fig. 1K). Thorns are also found on old, erect shoots (e.g. M. velutina, Fig. 1J), and these are normally very long and massive. At least in M. velutina, leaf arrangement within a plant can be variable, i.e. decussate on some shoots and in whorls of three on others (also cf. Catunaregam, above). Consequently, thorn arrangement, too, varies (cf. Fig. 1K, compare with Fig. 1J, both from the same individual). Dichotomous key to Thai Rubiaceae with hooks and thorns It is important to note that this key is valid only for taxa occurring in Thailand. Character states for taxa outside the Flora area may deviate. An example is Oxyceros, which in Thailand consists only of species with curved thorns but elsewhere contains some species with straight thorns.
1. Thorns or hooks curved; plants straggling or climbing, never erect trees or shrubs (2) 2. Inflorescences many-flowered, globose, pedunculate; stipules typically bifid at apex Uncaria 2. Inflorescences not as above; stipules never bifid (3) 3. Branches (long shoots) with abbreviated leafy and/or flowering shoots and thorns immediately above them (arising at the same node; accessory bud/shoot formation); flowers relatively small, always less than 1 cm in diam., mostly greenish, greenish-white or greenish-yellowish, in fascicles on very abbreviated shoots; fruits drupaceous Canthium pro parte 3. Branches (long shoots) with paired hooked thorns (leafy and/or flowering brachyblasts not present in the vicinity of thorns); inflorescences terminal, flowers relatively large, usually > 1 cm in diam., white; fruits berry-like (4) 4. All hooked thorns on a branch of the same kind (Fig. 1H) Oxyceros pro parte 4. Thorn pairs basal on lateral branches fang-like; larger and more robust than hooked thorns at the following nodes (cf. Fig. 1I) Oxyceros pro parte 1. Thorns straight; plants with erect main stems (5) 5. Trees with branches bearing short lateral shoots which terminally produce one or two pairs of short, straight thorns Tamilnadia 5. Arrangement of thorns not as above (6) 6. Thorns or thorn-tipped short lateral branches bearing very abbreviated leafy and/or flowering shoots (cf. Fig. 1D) Ceriscoides 6. Thorns neither with leafy nor with flowering abbreviated shoots (7) 7. Numerous pairs of thorns on often fascicled branches found in the lower part of trunk; branches of the crown mostly thorn-less; trunk and older branches typically rusty- to red-brown; flowers relatively large (to ca. 2 cm in diam.), green, unisexual Dioecrescis 7. Not as above; branches (long shoots) with abbreviated leafy and/or flowering shoots and thorns immediately above them (arising at the same node; accessory bud/shoot formation); flowers either considerably smaller than above or, if relatively large, not green and not unisexual (8) 8. Flowers relatively large, typically more than 1 cm in diam., mostly white (but turning yellow to orange with age in Catunaregam), in many- to few-flowered terminal inflorescences, or solitary flowers terminal on abbreviated shoots; fruits berry-like, without stones (9) 9. Thorns often long, to 4 cm; flowers 510-merous, large (2 cm or more in diam.), solitary (or in groups of few), older corollas turning yellow to orange; fruits large, at least 2 cm long or in diam. Catunaregam 9. Thorns smaller, usually not more than 1 cm long; flowers 5-merous, smaller, usually in many- to fewflowered inflorescences, old flowers not turning conspicuously yellowish or orange; fruits small, often less than 1 cm in diam. Fagerlindia
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8. Flowers relatively small, always less than 1 cm in diam., mostly greenish, greenish-white or greenishyellowish, in fascicles on very abbreviated shoots; fruits drupaceous (10) 10. Thorns often long, to over 5 cm, sometimes both in whorls of 3 or opposite on the same individual; ovary 5-carpellate, drupes typically with 5 stones (sometimes fewer due to abortion) relatively large (to ca. 3 cm in diam.) Meyna 10. Thorns typically much shorter (sometimes thin and inconspicuous and less than 1 cm long), opposite only; ovary 2(3)-carpellate, drupes with 2(3) stones (or, due to arbortion, a solitary stone), normally smaller, mostly not more than 1 cm in diam. Canthium pro parte
ACKNOWLEDGEMENTS Voradol Chamchumroon wishes to thank the AD (Austrian Academic Exchange) for a scholarship which made it possible to carry out part of the present studies at the Institute of Botany, University of Vienna, Austria. REFERENCES Chamchumroon, V. & Puff, C. 2003. The Rubiaceae of Ko Chang, south-eastern Thailand. Thai For. Bull. (Bot.) 31: 1326. Craib, W.G. 1932. Florae Siamensis enumeratio. A List of the plants known from Siam with notes of their occurrence. Vol. 2, part 1: Caprifoliaceae & Rubiaceae (in part), pp. 1145. Siam Soc., Bangkok. Guillaumin, A. 1931. Transformation de la partie infrieure de l'axe de l'inflorescence en crochet chez les Uncaria. C. R. Acad. Sci. Paris 192: 12641265. Middleton, D.J. 1999. Apocynaceae. In: Santisuk, T. & Larsen, K. (eds), Flora of Thailand 17(1): 1153. Reza, M. bin Mohd Azmi. 2001. Systematic studies on Ceriscoides (Rubiaceae Gardenieae) with emphasis on the morphology and taxonomic position of recently documented Malesian taxa. Unpubl. Ph.D. thesis, Univ. of Malaya, Kuala Lumpur. Ridsdale, C.E. 1985. The genus Fagerlindia (Rubiaceae) in the Philippines. Blumea 31: 239244. ________. 1998. Rubiaceae. In: Dassanayake, M. D. (ed.), A revised handbook to the Flora of Ceylon 12: 141343. Robbrecht, E. 1988. Tropical woody Rubiaceae. Opera Bot. Belg. 1: 1271. Tirvengadum, D. D. 1982. A study of the tribe Gardenieae (Rubiaceae) of South and South East Asia. Generic delimitation and revision with enumeration of species. Thesis Licent. Sci (unpubl.), Aarhus. ________. 1983. New taxa and name changes in tropical Asiatic Rubiaceae. Nordic J. Bot. 3: 455469. Tirvengadum, D. D. & Sastre, C. 1979. La signification taxonomique des modes des ramification de Randia et genres affines. Mauritius Inst. Bull. 8: 7798. Verdcourt, B. & Bridson, D. 1991. Rubiaceae (part 3). In: Polhill, R. M. (ed.), Flora of Tropical East Africa: 749956. A.A. Balkema, Rotterdam, Brookfield.
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Figure 1. AB, Uncaria spp.: A. shoot with three pairs of axillary climbing hooks; B. shoot with paired axillary pedunculate inflorescences, the arrow points to a developing inflorescence with clearly hook-like peduncle; C. Tamilnadia uliginosa, branch with shot lateral branches terminally bearing one to two pairs of thorns (inset: detail of short lateral branch showing two terminal pairs of thorns and very contracted short shoots immediately below with fruits); D. Ceriscoides sessiliflora, thorntipped lateral branch bearing short shoot with inflorescence; E. Dioecrescis erythroclada, small tree with thorn-bearing lateral branches on the right; trunk of old tree with clustered thorn-bearing branches (arrows) on the left; FG. Catunaregam tomentosa (F) and Fagerlindia sp., branches with thorns and associated leafy and fruit-bearing short shoots; HI. Oxyceros spp.; H. O. horridus, branch with paired, axillary hooked thorns; I. O. bispinosus, branch with two pairs of lateral branches, each with fang-like hooks near their base; JK. Meyna velutina, pictures from the same tree, J. old stems with large, paired thorns; K. younger branches with thorns in whorls of 3. Further explanations in the text (Photographed by C. Puff).
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THAI FOR. BULL. (BOT.) 31: 7594. 2003.
Non-indigenous Rubiaceae grown in Thailand

CHRISTIAN PUFF* & VORADOL CHAMCHUMROON **
ABSTRACT. A survey of non-indigenous Rubiaceae that are either commercially important or cultivated as ornamentals is given. It covers the genera Coffea, Cinchona, Carphalea, Gardenia, Hamelia, Mussaenda and Pseudomussaenda, Pentas, Psilanthus, Rondeletia and Serissa.
INTRODUCTION The Rubiaceae are an essentially tropical family, fairly evenly distributed in the Old and New World. Various Rubiaceae which originate from other parts of Asia, from Africa, Madagascar, or from Central and South America are cultivated in Thailand. These non-indigenous Rubiaceae found in Thailand fall in two categories: (a) commercially important plants, and (b) ornamentals. The present paper presents a survey of these taxa and provides brief descriptions for them. It explicitly excludes Ixoras, probably the most commonly grown rubiaceous ornamentals, as the genus, including ornamentals, is currently being studied by one of us (VC). Also excluded are taxa native to Thailand that are cultivated; only nonindigenous species are dealt with. COMMERCIALLY IMPORTANT PLANTS Coffea L. The genus Coffea does not occur naturally in Thailand or the rest of Asia. Only planted coffees, namely C. arabica and C. canephora (and possibly C. liberica; see below), are found in the country. All other taxa listed under Coffea in older literature on the Thai flora (e.g. Craib, 1934: 171174) are referable to other genera, primarily Psilanthus (see Chamchumroon & Puff, 2003). Coffea arabica L. (ARABICA Coffee). Fig. 2AD. Shrubs or small trees to ca. 4 m tall. Leaves opposite, with petioles to 10 mm long, elliptic to ovate-lanceolate, 615(18) by 38 cm, acuminate at apex, narrowed to base, with 710 pairs of lateral veins, glabrous, glossy above. Flowers in axillary clusters of few to ca. 20; calyx ring-like, minute, often with very small indistinct teeth; corolla (4)
*Institute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna, Austria. **The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
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5(6)-merous, tube 711 mm long, lobes 818 by 36 mm; anthers to 10 mm long, exserted; style with bifid stigma slightly exserted; ovary 2-locular, each locule with a solitary ovule. Fruits drupaceous, red, oblong-ellipsoid to globose, greatest length ca. 1020 mm when fresh. Seeds 2, ca. 916 mm long, enclosed in a thin endocarp. The natural distributional range of the species is confined to Southwest Ethiopia and to a few localities in neighbouring south-eastern Sudan and northern Kenya (cf. map, Fig. 1). It grows in evergreen forest in Ethiopia at altitudes ranging from 1500 1900 m (Puff, 2003). Coffea canephora Pierre ex Frhner. C. robusta Linden. (ROBUSTA coffee). Fig. 2EF. Shrubs or small trees ca. 2-4 m tall. Leaves with petioles to 20 mm long, oblongelliptic to broadly elliptic, 1235(40) by 511 cm, acuminate at apex, obtuse to cuneate at base, with (8)1115(17) pairs of lateral veins, glabrous, glossy above. Flowers in axillary clusters of ca. 1030(50); calyx ring-like, minute; corolla 56(7)merous, tube ca. 515 mm long, lobes ca. 1020 by 25 mm; anthers to 10 mm long, exserted; style with bifid stigma slightly exserted; ovary 2-locular, each locule with a solitary ovule. Fruits drupaceous, red, oblong-ellipsoid, greatest length ca. 912 mm when fresh. Seeds 2, ca. 711 mm long, enclosed in a thin endocarp. The species occurs naturally from Sudan, Uganda and Northwest Tanzania westward to Sierra Leone, and southwestwards to Angola. It is a component of the African lowland rain forest (cf. Fig. 1, which, however, only shows the core range of the species).
Figure 1. Distribution routes for the cultivated coffee crop in the tropics; the solid lines refer to Coffea arabica (ARABICA coffee), the broken lines to C. canephora (ROBUSTA coffee); numbers are the approximate years of introduction (Ferwerda 1976). The natural distribution range of C. canephora (vertically hatched) is shown incompletely; its stretches further westward as far as Sierra Leone, and southwestward to Angola.
NON-INDIGENOUS RUBIACEAE GROWN IN THAILAND (C. PUFF & V. CHAMCHUMROON)
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Table 1. Differences between Coffea arabica and C. canephora compiled from various sources. See text for further comments.
Character Leaf length (mm) Leaf width (mm) Pairs of lateral veins Flowers per axillary inflorescence Flowers: corolla lobes Fruit size (greatest length; mm) Fruits at maturity Seed (coffee bean) length (mm) Breeding Chromosomes (2n) Flowering time Flower mature fruit "coffee beans": caffeine content "coffee beans": yield (in tons per hectare) Root system Optimal temperature (annual average) Optimal annual rainfall Altitudinal range Coffea arabica 60150(180) 3080 710 (2)620 (4)5(6) 1020 falling off 916 autogamous 44 after rainy season 9 months ca. 0.51.5% 1.53 deep 1524 C 15002000 mm Typically 1000 to 2500 m ("mountain coffee") [8001500 m in Thailand] susceptible susceptible susceptible susceptible resistant primarily America "ca. 74%" "8090%" Coffea canephora (syn. C. robusta) 120350(400) 50110 (8)1115(17) (8)1230(50) 56(7) 912 remaining on plant 711 allogamous 22 irregular 1011 months ca. 24% 2.34 shallow 1830 C 20003000 mm Sea level to ca. 1000 m ("lowland coffee") [usually below 1000 m in Thailand] resistant resistant resistant +/- resistant susceptible especially Old World "ca. 25%" "ca. 10%"
Diseases: coffee rust (Hemileia vastatrix) Diseases: coffee bean disease (beetle) Diseases: nematodes Diseases: spider web disease Diseases: tracheomycosis (Giberella xylarioides) Major production areas World production
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Morphological as well as other diagnostic characters between the two species are summarized in Table 1. It should be noted that, due to the large numbers of cultivars in each of the two species, and due to the continuous development of new strains, individual data may not always fully hold. Data on world production (Table 1, last row) vary greatly from source to source. In recent years, ROBUSTA coffee production in Asia has drastically increased, so that it is likely to the world production ratio is shifting in favour of ROBUSTA (see also below). The remaining few percent of world coffee production is contributed by LIBERICA and EXCELSA coffee (C. liberica), and some other species such as C. stenophylla (West African Highland coffee or Highland coffee of Sierra Leone) and others. ARABICA coffee beans are generally considered to be of higher quality than ROBUSTA beans and always fetch higher prices. Coffee made from pure ARABICA beans is usually considered very palatable, while coffee made from pure ROBUSTA beans is not well liked because it is bitter. Moreover, the beans of ARABICA, being larger and more perfectly shaped than those of ROBUSTA, are more pleasing to the customers eye. Consequently, pure roasted ARABICA beans sell better - and at a higher price. Both in ARABICA and in ROBUSTA, there are many slight variations in flavour. The reasons for these differences are many-fold: different strains produce different flavours; growth of the same strain under different soil and climatic conditions also results in variation in flavour; roasting techniques (primarily the length of roasting and temperature) influences the taste. Many of the coffees (either roasted, packed beans or ground roasted beans) are blends of ARABICA of different origin, or blends of ARABICA and ROBUSTA. A general guidline is that the cheaper the coffee blend the more of the less expensive ROBUSTA will be in it. Instant coffees are primarily made from the lower-priced ROBUSTA beans. A note on coffee fruits and coffee beans, using Coffea arabica as an example (Fig. 2D) The fruits are drupes with a skin-like exocarp and a fleshy mesocarp and typically contain two 1-seeded pyrenes (Fig. 2Da) (in some cultivars, one of the two seed-containing pyrenes is always aborted, and the remaining pyrene, containing the only well-developed seed, becomes globose; for this reason, such coffees are called Pearl Coffee). The two pyrenes are flat on their ventral surface, where they face each other (see Fig. 2Da). Compared to other Rubiaceae with drupaceous fruits, Coffea is somewhat unusual in having very thin, horn-like endocarps (rather than the more common, thick endocarps made up of numerous layers of sclerenchymatic tissue). In coffee processing plants, the seed-containing pyrenes (cf. Fig. 2Db) are first removed from the fruits in various ways. Then the endocarps, being thin, are easily cracked open mechanically, and the empty endocarp cases and seeds (Fig. 2Dc) are separated. The seeds (coffee beans in commercial terms) show a characteristic, slightly curved fold on the ventral side which extends to the inside (compare seed section and
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ventral view, Fig. 2Da and Dc), which is an unusual feature within the family. Well over 95% (by volume) of the seed is occupied by endosperm; the minute embryo (Fig. 2De) is located in the lower part of the seed, in an oblique position (Fig. 2Dd). Yet another unusual feature of a coffee seed is the structure of the exotesta (seed coat). Whereas, in the majority of Rubiaceae, the exotesta is a continuous layer of cells covering the entire seed, the seed coat of Coffea is discontinuous: during growth of a coffee seed the seed coat cannot keep up with the resultant increase in size and diameter and eventually breaks up into isolated islands of exotesta cell groups on the surface of the endosperm. Coffee-growers call this silver-skin (visible on the surface of the seeds in Fig. 2Dcd). This silver-skin can be easily washed off, and the green coffee beans commercially sold are primarily endosperm (remnants of the seed coat are normally only left in the ventral fold of a coffee bean). For anatomical information of both C. arabica and C. canephora seeds see Dentan (1985). History and economic importance for Thailand Figure 1 shows the distribution routes for cultivated ARABICA and ROBUSTA coffee (C. arabica and C. canephora). Interesting and important from an Asiatic point of view is the introduction of ROBUSTA coffee to Java in 1900. This was the starting point which has led to the current lead in ROBUSTA production in Asian countries (whereas ARABICA, although introduced to Asia earlier than to the Americas, lags behind in terms of production; primary ARABICA producers are Brazil and Colombia). Larger scale coffee production in Thailand only started in the 1960s. However, since then, there have been dramatic increases. By 1976, Thailand officially became a coffee exporting nation. By the late 1990s, the country was ranked seventh in world coffee production (i.e. production of ARABICA, ROBUSTA, and other minor coffees put together). Amongst the Southeast Asiatic ROBUSTA coffee producers, Thailand ranks third (behind Vietnam and Indonesia). At present, Thailand produces ca. 80,000 tons of ROBUSTA coffee per year, of which about 60% (ca. 50,000 tons) is exported, whereas the rest is processed locally (see below). ROBUSTA coffee is grown in large-scale plantations primarily in peninsular Thailand, from Chumphon southwards (Ranong, Surat Thani, Phang-Nga, Krabi and Nakhon Si Thammarat). ROBUSTA is easier to cultivate, more disease-resistant, can tolerate wider temperature and moisture extremes, produces more beans and the fruits mature more quickly than ARABICA. Although ROBUSTA beans are generally considered to be of lesser quality than ARABICA beans, Thai ROBUSTA has the reputation of being of good quality and is exported primarily to the U.S.A., Europe, Japan and Singapore. Locally used and processed ROBUSTA is mainly used for canned coffee drinks as well as instant coffee (which, too, is partly exported). In contrast, ARABICA coffee, is primarily grown on a small scale, often by hill tribe families and villages, in Thailands northern provinces (Chiang Mai, Chiang Rai, Lampang, Mae Hong Son, Tak). Only a few hundred tons of beans are produced annually, almost all of which are processed and consumed locally. ARABICA growing in the North started in the 1970s, as part of the opium crop replacement campaign, and
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was supported by the Royal Development Projects, the United Nations and many other governmental and non-governmental organisations. ARABICA growing in Thailand is still somewhat experimental but it is to be excepted that, due to increasing local demands, this might change quickly. Many coffee shop chains have opened in recent years, and nowadays numerous service stations and shopping centres have highly popular coffee outlets. Most of these serve ARABICAS of local production. The Boncaf Website (self-termed The Thai Coffee Website, http://www. boncafe.co.th/thai-coffee/thai-coffee.htm, and links) gives excellent information in a Thai context and is recommended for further reading. The website of the International Coffee Organization (ICO) (http://www.ico.org/) provides extensive general information on various aspects, ranging from botanical data, plant breeding and ecology to coffee prices, trade statistics and economic-political trends concerning coffee. Generalized statistical data are also available from the website of the Food and Agriculture Organization of the UN (FAO) (htto://www.apps.fao.org). Recommended standard references on cultivated coffee are Clarke & Macrae (19851988), Clifford & Willson (1985) and Wrigley (1988). Is Coffea liberica Hiern in Thailand? Craib (1934: 172) mentioned two Kerr collections of cultivated plants of this species, one from Chanthaburi province, the other from Nakhon Si Thammarat, and commented that the species is frequent in cultivation. As these specimens could not be traced, it remains uncertain whether their identity is correct. The situation is rather complex: Hierns original description of C. liberica contained elements of two species, namely C. liberica [sensu stricto] and C. canephora (see Bridson & Vercourt, 1988: 706, 711). It is, therefore, possible that the specimens referred to by Craib are C. canephora. Craibs comment that it is frequently cultivated would support this proposition. Coffea liberica Hiern. C. excelsa A. Chev. (LIBERICA Coffee, EXCELSA Coffee). This species occurs naturally in West Africa, having a range that more or less corresponds to that of C. canephora. It is fairly widely cultivated in tropical Africa, although not in large-scale plantations. The species is also grown elsewhere in the tropics, but is of minor importance (less than 5% of the world coffee production). Beans yield a very bitter coffee of low quality (coffee from EXCELSA strains is said to be somewhat less bitter than coffee from LIBERICA strains). It is the most robust Coffea species: plants can become trees to 20 m tall (with trunks 30 cm in diam.); leaves can be to 55 cm long and over 20 cm wide; flowers and fruits, too, are larger than in the other species. Cinchona calisaya Wedd. C. ledgeriana (Howard) Trimen. Fig. 2G. Trees to ca. 5 m tall. Leaves oblong to ovate, ca. 815 by 28 cm, rounded at apex, cuneate at base, with 79 pairs of lateral veins. Inflorescences terminal, many-flowered.
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Flowers heterodistylous, 5-merous; calyx with small teeth; corolla white to pinkish, tube ca. 812 mm long, lobes to ca 5 mm long, conspicuously villous along the margins; tips of stamens exserted in short-styled flowers (cf. Fig. 2G), included in longstyled flowers; styles and stigmas included. Fruits capsular, with septicidal dehiscence, to ca. 2 cm long, ellipsoidal. Seeds flattened, to ca. 7 by 2 mm, with strongly dentate to more or less fimbriate wing. A neotropical species spontaneously occurring on the eastern slopes of the Andes from central Peru to central Bolivia (Andersson, 1998: map, Fig. 14D). Cinchona trees have been grown throughout the tropics for the extraction of quinine. Northern and southwestern India and Java are major areas of production. Within the genus (ca. 23 species confined to north-western South America) only strains from two species are of any commercial importance, i.e. C. calisaya (often grown as C. ledgeriana), and C. pubescens (often grown under the name C. succirubra). Bark from C. calisava was commercially known as yellow, that of C. pubescens as brown or (the succirubra strain) red (Andersson, 1998). These two species contain the highest amount of quinine alkaloids within the genus. Another species often cited as being of commercial importance, C. officinalis, has been redelimited by the revisor of the genus (Andersson, 1998) to a form that has no medicinal value at all. The name Cinchona came from the Countess of Chinchn, the wife of the fourth Count of Chinchn and Viceroy of Peru, who, in 1638, was cured of a malarial type of fever by using a decoction made from the bark of Cinchona. After her return to Spain, she introduced the drug to European medicine. In 1820 two scientists, Pelletier and Caventou, isolated an alkaloid chemical in the bark which provided the highest antimalarial effect and named it quinine. In the middle of the 19th century, seeds of Cinchona were smuggled out of South America by the British and the Dutch and planted and cultivated in Java by the Dutch and in India and Sri Lanka by the British. They quickly dominated the world production of quinine, whereas Bolivia and Peru, where the trees originated from, played no commercial role as quinine producers. The occupation of Java by the Japanese in 1942, during World War II, resulted in major shortages of natural quinine. For this reason, research to develop a synthetic quinine alkaloid was intensified, and by 1944 the goal was successfully achieved. After World War II, synthetic products largely replaced natural quinine, and the importance of Cinchona as a commercial crop dropped dramatically, but this may change. Because of increasing problems with malaria strains which have developed a resistance to synthesised quinine drugs, natural quinine extracted from quinine bark and the use of natural bark tea and/or bark extracts are making a comeback in the management and treatment of malaria. New strains of drug-resistant malaria can be treated effectively with natural quinine and/or quinine bark extracts (Taylor, 2002). Some production of Cinchona bark is also supported by the soft-drink industry. Quinine is very bitter tasting and commercially sold tonic waters often use quinine as its bitter ingredient/component. Commercially produced tonic water usually contains around 100 to 300 ppm quinine and up to a maximum allowable concentration of 70 mg/l-1(Taylor, l.c.).
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In Thailand, Cinchona was probably never planted on a large scale. A well known locality for planted Cinchona calisaya (as C. ledgeriana) is the medicinal garden (Forestry Experimental Station) on Doi Suthep (Chiang Mai province). There are also some isolated trees near the Doi Inthanon National Park guesthouse. ORNAMENTALS Taxa are presented here in alphabetic sequence. The brief descriptions are primarily based on cultivated material. Descriptions may therefore deviate from those given for the taxa in their natural state (e.g. growth form, leaf sizes and shapes, or flower colour may differ in cultivars). Carphalea kirondron Baillon. Fig. 3AB. Shrubs to 3 m tall (much taller and becoming tree-like if allowed to grow). Leaves mostly opposite but occasionally in whorls of 3, with petioles to 2 cm long, lanceolate to ovate-lanceolate, ca. 410 by 25 cm, acute at apex, narrowed to base, glabrous to variously hairy. Inflorescences terminal, very many-flowered, corymb-like, to 20 cm in diam. Flowers mostly 4-merous, heterodistylous; calyx lobes enlarged, unequal (1 more enlarged than the others and to ca. 2 cm long), bright red; corolla tube narrowly filiform, reddish, ca. 1.52.5 cm long, lobes (creamy-) white, to 5 mm long; filaments and anthers exserted and styles and stigmas included in short-styled forms, upper part of style and 2 filiform stigmas exserted in long-styled form. Fruits indehiscent, nut-like, obconic, to ca. 5 mm long, crowned by the enlarged calyx lobes; typically 1 fertile seed in each of the two locules, but not infrequently one or both seeds are aborted. A species native and endemic to Madagascar (Puff, 1988: map, Fig. 14B). The entire genus is characterised by the presence of coloured, enlarged calyces. It is a member of the tribe Hedyotideae, along with Pentas (see below). The species is infrequently seen in Thailand (several localities in Chiang Mai and Kanchanaburi provinces). Outside Thailand, it has been observed by one of us (C.P.) in the Philippines and in Malaysia (in recent years, plants have been regularly seen on sale in Kuala Lumpur markets). The plant may not be easy to propagate. Seed germination rates at the Singapore Botanic Gardens was extremely low (only ca. 2%; Ng Siew Yin, 1981). Propagation by cuttings also appears to be difficult (also cf. Puff, 1988: 304). Gardenia Ellis. Fig. 3K. Several species, including both native and non-indigenous ones, are cultivated in Thailand. The most commonly seen cultivated native species is probably G. carinata (not treated here). G. augusta (L.) Merr. G. jasminoides Ellis, a name often used in horticultural literature. G. florida L.
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Shrubs ca. 12 m tall. Leaves opposite or occasionally in whorls of 3, shortly petiolate, elliptic-obovate or oblong-lanceolate, ca. 512 by 2.55 cm, acute to acuminate at apex, narrowed to base, coriaceous, glabrous, glossy; stipules sheath-like above insertion of petioles. Flowers solitary and terminal on abbreviated lateral shoots (thus seemingly axillary), fragrant, very often double; calyx made up of 6 linear (-lanceolate) lobes, ca. 23 cm long; corolla white (turning creamy-yellow with age), tube and lobes to ca. 5 cm long; double flowers: the outer (true) corolla lobes 6(9), elliptic-ovate, the inner more numerous, smaller; stamens poorly developed or petaloid; ovary often sterile in double flowers. Fruits or sterile pseudo-fruits longitudinally ribbed, these and the persistent calyx often turning bright orange-reddish. A species native to south-eastern China, Taiwan, Japan and nearby regions of the subtropical eastern hemisphere. Several species, including both native and non-indigenous ones, are cultivated in Thailand. The most commonly cultivated native species is probably G. carinata (not treated here). The most prominent non-indigenous ornamental is G. augusta (L.) Merr. (syn. G. jasminoides Ellis, a name often used in horticultural literature; syn. G. florida L.). The species is widely planted in all parts of Thailand. Cultivated plants are mostly of the normal shrubby type. Prostrate forms (cv. Prostrata), used elsewhere in the tropics as ground cover in protected, partly shady areas, do not seem to be grown in Thailand. Also forms with variegated leaves (cv. Variegata) appear to be absent. Hamelia patens Jacq. (syn. H. erecta Jacq., H. coccinea Swartz). Fig. 3FG. Shrubs or, if allowed to grow, small trees to 7 m. Leaves opposite or in whorls of 3 or 4, petiolate, ovate-elliptic, ca. 612 by 38 cm, (sub)acuminate at apex, narrowed to the base, membranous, glabrous to villous. Inflorescences terminal, many-flowered, with distinct double-helicoid cymes as partial inflorescences; inflorescence axes red. Flowers 5-merous; corollas rather thick and robust, yellow to yellow-orange or orangered, tube ca. 1.52 cm long, lobes erect, very small; anther, styles and stigmas mostly included; ovary red, 5-locular. Fruits berry-like, globose, to ca. 1 cm in diam., yellow to red at first, dark blue-black when fully mature. Seeds minute, numerous. A widespread tropical American species, naturally occurring from tropical Florida and Mexico southwards through Central America, the West Indies, and South America to Chile and Argentina (Elias, 1976: map, Fig. 5). In its natural distribution range, the species is known to be rather aggressive, quickly occupying newly disturbed sites (Elias 1976). Seeds (both in its natural range and elsewhere) are dispersed by birds and germinate readily. The species apparently also reproduces vegetatively by sending out runners and rhizomatous branches (Elias, l.c.). Flowers are typical bird flowers, being pollinated by hummingbirds in the New World; sun birds (and also butterflies) were observed as pollinators in Thailand. As is typical for ornithophilous flowers, they lack any scent. This fast-growing species is widely planted in all parts of Thailand. It is often grown as a living fence or hedge and soon flowers again after pruning. It is often also grown as a solitary bush and can be trimmed into various shapes without negative effects. Individuals left untouched for several years become tree-like but are less attractive in such a state.
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Mussaenda L. and Pseudomussaenda Wernham Mussaenda, Pseudomussaenda and Schizomussaenda are three genera characterised by enlarged, coloured (petaloid) calyx lobes. The taxonomic position and relationship of the tropical African Pseudomussaenda has been much disputed, and several authors (e.g. Bakhuizen van den Brink & Koster, 1963, Baker & Bakhuizen van den Brink, 1965) suggested it should be merged with Mussaenda. Jayaweera (1963) suggested that Pseudomussaenda and Schizomussaenda are not sufficiently different to warrant the segregation as two genera. Based on a detailed investigation of the character states of the three genera, Puff et al. (1993), came to the conclusion that the three genera should be upheld. This was subsequently confirmed by DNA studies (Bremer & Thulin, 1998). In the present context, Schizomussaenda (S. dehiscens; monotypic), a tree-like plant occurring in northern Thailand, is not treated herein as it is not used as an ornamental. Indigenous species of Mussaenda are well represented in the Thai flora, but not all taxa grown as ornamentals are native. The most obvious character distinguishing Mussaenda from Pseudomussaenda is the fruit: it is indehiscent and berry-like in the former and a dehiscent capsule in the latter. Mussaenda erythrophylla Schumach. & Thonn. Fig. 3I. Shrubs to 3 m (scandent and climbing, with stems to 8 m if allowed to grow; usually not seen in cultivation). Leaves with petioles to 4 cm long, elliptic to broadly ovate, ca. 712 by 58 cm, acute to acuminate at apex, rounded, subcordate to cuneate at base, hairy; stipules bifid. Inflorescences terminal, many-flowered, inflorescence axes densely covered with red hairs. Flowers 5-merous, heterodistylous; calyx lobes of most flowers small, ca. 11.5 mm long, lanceolate, with red hairs, but 26 flowers per inflorescence with a single calyx lobe enlarged, foliaceous and coloured bright red (elliptic to round, to ca. 10 by 8 cm); corolla tube with red hairs outside, to ca. 3 mm long, lobes white to creamy-yellow, round, to ca. 1 cm long, a ring of dark red hairs surrounding the throat; anthers and style and stigmas included in the corolla tube both in long- and short-styled forms. Fruits berry-like, ellipsoid, to ca. 2 cm long, with numerous minute seeds. A species native to Western tropical Africa. The species is occasionally seen in gardens and parks, but is not as commonly planted as the M. philippica cultivars described below. It appears to be cultivated much more frequently in the peninsula than elsewhere in Thailand. Mussaenda philippica L.C. Rich. Doa and Queen Sirikit. Fig. 3H. Shrubs, sometimes rather tree-like, to ca. 3 m tall. Leaves with petioles to ca 3 cm long, variable, ovate to lanceolate, ca. 515 by 37 cm, shortly acuminate at apex, rounded to cuneate at base, glabrous to hairy; stipules bifid. Inflorescences terminal, many-flowered. Flowers 5-merous, often completely sterile; all calyx lobes of all flowers of an inflorescence enlarged, foliaceous and white or in various shades of pink, lanceolate to rounded, ca. 610 by 48 cm; corolla tube and lobes to ca. 1 cm long, corolla yellow, yellow-orange to orange, with a darker orange eye. Fruit not developed.
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Normal M. philippica shows the typical Mussaenda characteristics, i.e. it has only relatively few enlarged white calyx lobes per inflorescence. Usually, only one to four flowers of a partial inflorescence have one enlarged, coloured calyx lobe each, whereas the remaining calyx lobes remain small and inconspicuous. It is not grown as an ornamental. The origin of cvs. Doa and Queen Sirikit is partly documented in a University of the Philippines Newsletter published for the 1998 opening of a Mussaenda cultivar garden (http://www.up.edu.ph/newsletter/1998/01/donas.html). An unusual mutation of M. philippica was discovered in the 1930s on Mt Maquilin, in the vicinity of the Los Baos Campus of the University of the Philippines (Luzon). It differed from the normal M. philippica in having all five calyx lobes of all flowers of an inflorescence enlarged and coloured pure white. The aberrant form was eventually formally described as M. philippica var. aurorae Sulit. This sport was taken into cultivation by horticulturists at Los Baos and reproduced vegetatively. It became known as M. philippica Doa Aurora, named after the wife of former President Manuel L. Quezon. This set the precedent for naming subsequently developed cultivars after First Ladies (Doas). Pink Doas (cultivars with all enlarged calyx lobes in various shades of pink): all of these originated by basically crossing cv. Doa Aurora (white enlarged calyx lobes) with M. erythrophylla (bright red enlarged calyx lobes; see above): white + red = pink. Further backcrossing of F1 hybrids with other cultivars resulted in slight colour variations, e.g. cv. Doa Luz (mottled pink with whitish veins), cv. Doa Hilaria (pure pink), cv. Doa Evangelina (dark pinkish-red), and many others. Mussaenda philippica Queen Sirikit also belongs to the group of 'pinks'. It was named after the Queen of Thailand during her visit to the Los Baos campus of the University of the Philippines in 1963. According to the Plant Science Department at the University of Connecticut (http://florawww.eeb.uconn.edu/acc_num/199300381.html), cv. Queen Sirikit resulted from the backcross of the F1 hybrid, M. erythrophylla x M. philippica Doa Aurora, to M. philippica Doa Aurora. Both white and pink M. philippica cultivars are widely grown in Thailand as ornamentals, not infrequently both are planted side by side (Fig. 3H). They appear to require continuously high humidity and undoubtedly are most commonly seen in the Peninsula, but are also frequent in Bangkok and can be observed as far north as Chiang Mai province, where they are grown in the low-lying areas. Whilst the white-calyx cultivars invariably are cv. Doa Aurora, it is not certain whether the pink-calyx cultivars seen in Thailand are always cv. Queen Sirikit. Cultivars in various shades of pink occur which, except for the colour tone of the enlarged calyces, are indistinguishable. It is likely that many of the pink cultivars are rather cv. Doa Luz (a much older cultivar than cv. Queen Sirikit) which has been grown in many parts of Southeast Asia for a long time. Pseudomussaenda flava Verdc. Mussaenda luteola Delile. Mussaenda flava (Verdc.) Bakh. f. Fig. 3J.
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Shrubs to ca. 3 m tall. Leaves opposite, shortly petiolate, elliptic to oblongelliptic, ca. 48 by 24 cm, acute to acuminate at apex, narrowed to the base, softly pubescent; stipules with 2 filiform fimbriae. Inflorescences terminal, severalflowered. Flowers 5-merous, heterodistylous; calyx lobes of most flowers small, ca. 25 mm long, filiform, hairy, but 12 flowers per inflorescence with a single calyx lobe enlarged, foliaceous and coloured white to creamy-yellow (oblong, ovate or elliptic, ca. 25 by 1.54 cm); corolla tube 23 cm long, greenish, lobes ca. 58 mm long, ovate, yellow, throat often with a ring of orange-yellow hairs; anthers and style and stigmas included in the corolla tube both in long- and short-styles forms. Fruits dry, capsular, oblong, to ca. 5 mm long, loculicidally splitting into two valves. The species seems to be cultivated primarily in peninsular Thailand. It usually is planted in groups and is sometimes seen as a living fence. Pentas lanceolata (Forssk.) Defl. Fig. 3C. Perennial herbs, sometimes a little woody at base, to ca. 0.5 m tall, or, in some cultivars, low and cushion-like. Leaves with petioles to ca. 1 cm long, ovate to lanceolate, 46 by 2.53 cm, acute to acuminate at apex, narrowed to the base, thinnish, pubescent to puberulous. Inflorescences terminal, many-flowered, of umbel-like appearance. Flowers 5-merous, mostly heterodistylous; corolla, depending on cultivar, white, in shades of pink, reddish-purplish or lilac or mauve, sometimes 2-toned (dark pink with white stripes, etc.), tube narrowly cylindrical, to ca. 2 cm long, lobes ovate to elliptic, ca. 0.51 cm long; filaments and anthers exserted and styles and stigmas included in short-styled forms, upper part of style and 2 filiform stigmas exserted in long-styled form. Fruits capsular, dehiscing into 2 valves, to ca. 5 mm long; seeds numerous, minute. A variable species, divided into several infraspecific taxa, occurring from Tropical East Africa to Ethiopia to Sudan and also extending into south-western Arabia (Verdcourt, 1976; Puff, 2003). A species sporadically seen cultivated (Bangkok, Chiang Mai and Chiang Rai provinces) and presumably a relatively recent addition to Thailands horticultural scene. Low, cushion-like forms make attractive flowering borders. More erect growing forms are occasionally seen as pot plants. Psilanthus bengalensis (Heyne ex Schult.) Le Roy. Coffea bengalensis Heyne ex Schult. Illustrations: Sivarajan et al. (1992: Fig. 2AE); Pinratana (1980: 37). Shrubs to 23 m tall (if allowed to grow; plants usually pruned and kept low, and often less than 0.5 m). Leaves elliptic-oblong to broadly ovate, ca. 59 by 45 cm, shortly acuminate at apex, narrowed to the base, subcoriaceous, glabrous (except for veins below); stipules small, triangular-aristate. Flowers 15 together, terminal (on main shoots or on abbreviated axillary shoots), fragrant, 5-merous; calyx rim-like, with minute teeth; corolla white, tube to 2 cm long, lobes ovate to elliptic, ca. 1 cm long; tips of anthers exserted, style and stigmas included. Fruits subsessile, drupaceous, ellipsoidal, crowned by persistent calyx, ca. 1 cm long, blue-black when ripe, with 2 seeds.
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A species naturally occurring from Bangladesh to India and Nepal (Sivarajan et al., 1992). An occasionally seen ornamental (Bangkok; northern provinces), probably grown for its sweetly scented flowers. Pinratana (1980) notes that the species is propagated by seed. Rondeletia odorata Jacq. Fig. 3E. Shrubs ca. 12 m tall. Leaves oblong (-obovate), 410 by 24 cm, rounded to acute at apex, obtuse or subcordate at the base, thick, leathery. Inflorescences terminal, several- to many-flowered. Flowers 5-merous; corollas red or red-orange, mostly with a distinct yellow-orange eye, tube ca. 1 cm long, lobes rounded, to ca. 5 mm long; anthers included; stigmas barely exserted from throat; ovary 2-locular. Fruits globose, ca. 4 mm in diam., capsular, splitting into 2 valves, releasing numerous very small, wingless seeds. A species naturally occurring from northern Central America (southern Mexico) southward to north-western South America, and also in the Caribbean. Only seen cultivated in a few places in peninsular Thailand, and in Bangkok. Serissa japonica (Thunb.) Thunb. S. foetida (L.f.) Willd. Fig. 3D. Shrubs to ca. 0.5 m tall. Leaves shortly petiolate, oblong, ca. 1.52.5 by 0.5 cm, acute at apex, narrowed to base, thin-coriaceous, glabrous. Leaves and young shoots foetid when bruised. Flowers solitary or few together terminal on abbreviated short shoots, (4)56-merous, heterodistylous; corolla white to pale pink, tube ca. 5 mm long, lobes ca. 2 mm, with acuminate tips; anthers exserted and filiform stigmas included in short-styled form, reversed in long-style flowers; ovary 2-locular. Fruits small, less than 5 mm long, opening by means of an operculum, with 2 pyrenes. A species native to south-eastern China and possibly an early introduction to Japan (see note in Puff, 1990). In Craib (1934: 230), and often also in horticultural literature, the species is wrongly called Serissa foetida. The species was originally associated with the genus Lycium and was described as L. japonicum in 1780 and as L. foetidum in 1781; hence japonicum is the oldest name. The characteristic foetid odour that is emitted when Serissa foetida plant tissue is damaged is due to paederoside, an iridoid glycoside (Inouye et al., 1988). Tissue damage leads to the release of an enzyme which splits the bad-smelling sulphur component off the iridoid glycoside. The same chemical compound is known from other rubiaceous genera and taxa occurring in Thailand, e.g. Paederia and Leptodermis (in the same tribe, Paederieae, as Serissa), Saprosma and species of Lasianthus. The species is fairly frequently planted in all parts of Thailand but is easily overlooked because it is not particularly showy. It is often used as in the borders of flower
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beds or as a low hedge. Cultivars with variegated leaves (yellowish-white leaf aged; cv. Variegata) are common (cf. Fig. 3D). Cultivars with double flowers are less often seen (formally described as forma duplex and forma pleniflora, cf. Hara, 1952). Propagation by cuttings is easy. Serissa japonica is much used in Bonsai, although not particularly so in Thailand. The plants are fast-growing so that attractive Bonsai trees can be formed by pruning and wiring in a relatively short time. ORNAMENTALS EXPECTED TO BE GROWN IN THAILAND At least two spectacular rubiaceous ornamentals of extra-Asiatic origin, observed in Malayan and Indonesian gardens, have not yet been recorded for Thailand although they should do well, at least in the peninsula. They are only briefly mentioned below: Warszewiczia coccinea (Vahl) Klotzsch Shrubs with striking terminal, spike-like inflorescences to over 50 cm long, composed of numerous congested partial inflorescence arranged along the elongated inflorescence axis. Each several-flowered partial inflorescence has two flowers with a single much enlarged calyx lobe, which is leaf-like in shape and coloured bright red (to ca. 10 by 5 cm; compare with Mussaenda and Pseudomussaenda, above). A species naturally occurring from southern Central America to Brazil. The plant is often called Wild Poinsettia because the enlarged, foliaceous, redcoloured calyx lobes are, at least in colour, superficially similar to the bracts subtending the inflorescences of Poinsettia (Euphorbia pulcherrima). Euclinia longiflora Salisb. Randia macrantha (Schultes) DC. Gardenia macrantha Schultes. Illustrations: Bridson & Verdcourt (1988: Fig. 79), Chin (1977). Bushy shrubs to ca. 4 m tall with spectacular, fragrant, white 5-merous flowers mostly solitary at the end of branches. Corolla tube to over 20 cm long, lobes to 5 cm long, recurved; flowers turn creamy-white with age. An African rain forest species occurring from Uganda westwards to Guinea, and south-west to Angola. Corner (1988: 647) mentions that the species [as Randia macrantha] is frequently cultivated in Malay gardens and known as Angels Trumpets. This is not to be confused with Rothmannia longiflora Salisb. (syn. Randia maculata DC.), a species that has been seen cultivated in Indonesia. It is also a tropical African plant, with a natural distribution range similar to Euclinia longiflora. The bushes, too, produce very large flowers (corolla tubes to ca. 20 cm long) which are erect and solitary, but the corolla tubes are greenish or reddish outside, and the inside of the lobes is white with purplish or reddish markings. Illustration: Hall (1970: Fig. 56, 110).
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ACKNOWLEDGEMENTS Voradol Chamchumroon wishes to thank the AD (Austrian Academic Exchange) for a scholarship which made it possible to carry out part of the present studies at the Institute of Botany, University of Vienna, Austria. REFERENCES Andersson, L. 1998. A revision of the genus Cinchona (Rubiaceae-Cinchoneae). Mem. New York Bot. Gard. 80: 175. Backer, C. A. & Bakhuizen van den Brink, R. C., Jr. 1965. Flora of Java (Spermatophytes only), vol. 2.-N.V.P. Noordhoff, Groningen. Bakhuizen van den Brink, R. C., Jr. & Koster, J. T. 1963. Notes on the Flora of Java VIII. Blumea 12: 6169. Bremer, B. & Thulin, M. 1998. Collapse of Isertieae, re-establishment of Mussaendeae, and a new genus of Sabiceeae (Rubiaceae); phylogenetic relationships based on rbcL data. Pl. Syst. Evol. 211: 7192. Bridson, D. & Verdcourt, B. 1988. Rubiaceae (part 2). In: Polhill, R.M. (ed.), Flora of Tropical East Africa: 415747. Brookfield, A.A. Balkema, Rotterdam. Chamchumroon, V. & Puff, C. 2003. The Rubiaceae of Koh Chang, south-eastern Thailand. Thai For. Bull. (Bot.) 31: 1326. Chin, H.F. 1977. Malaysian flowers in colour. Tropical Press, Kuala Lumpur. Clarke, R.J. & Macrae, R. (ed.). 1985. Coffee. Vol. 1: Chemistry. Elsevier Applied Science Publ., London, New York. ________. 1987. Coffee. Vol. 2: Technology. Elsevier Applied Science Publ., London, New York. ________. 1988. Coffee. Vol. 3: Physiology. Elsevier Applied Science Publ., London, New York. ________. 1988. Coffee. Vol. 4: Agronomy. Elsevier Applied Science Publ., London, New York. ________. 1988. Coffee. Vol.5: Related beverages. Elsevier Applied Science Publ., London, New York. _______. 1988. Coffee. Vol. 6: Commercial and technico-legal aspects. Elsevier Applied Science Publ., London, New York. Clifford, M. N. & Willson, K. C. (eds). 1985. Coffee. Botany, biochemistry and production of beans and beverage. Croom Helm, London, New York, Sydney. Corner, E. J. H. 1988. Wayside Trees of Malaya, 3rd. ed. vol. 2. Malayan Nature Soc., Kuala Lumpur. Craib, W. G. 1934. Florae Siamensis enumeratio. A List of the plants known from Siam with notes of their occurrence. Vol. 2, part 2: Rubiaceae (concluded) to Dipsacaceae. 147233. Siam Society, Bangkok.
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Dentan, E. 1985. The microscopic structure of the coffee bean. In: Clifford, M. N. & Willson, K. C. (eds), Coffee. Botany, biochemistry and production of beans and beverage, pp. 284304. Croom Helm, London, New York, Sydney. Elias, T.S. 1976. A monograph of the genus Hamelia (Rubiaceae). Mem. New York Bot. Gard. 26: 81144. Ferwerda, F.P. 1976. Coffees. Coffea spp. (Rubiaceae). In: Simmonds, N.W. (ed.), Evolution of crop plants, pp. 257260. Longman, London. Hall, N. 1970. Flore du Gabon 17. Famille des Rubiaces (2e partie). Muse Nationale dHistoire Naturelle, Paris. Hara, H. 1952. Enumeratio Spermatophytarum Japonicarum ... Pars secunda. Iwanami Shoten, Tokyo. Inouye, H., Takeda, Y., Nishimura, H., Kanomi, A., Okuda, T. & Puff, C. 1988. Chemotaxonomic studies of rubiaceous plants containing iridoid glycosides. Phytochemistry 27: 25912598. Jayaweera, D.M.A. 1963. The rubiaceous genus Mussaenda: the morphology of the Asiatic species. J. Arnold Arb. 63: 111126. Ng Siew Yin. 1981. Carphalea kirondron Baill. (Rubiaceae). A newcomer to the horticultural scene. IPR News (Parks & Recreation Dept., Singapore) Jan. 1981: 5. Pinratana, Brother Amnuay. 1980. Flowers in Thailand. Nature Series no. 02G. Viratham Press, Bangkok. Puff, C. 1988. Observations on Carphalea Juss. (Rubiaceae-Hedyotideae), with particular reference to the Madagascan taxa and its taxonomic position. Bull. Jard. Bot. Belg. 58: 271323. _______. 1990 [1989]. The affinities and relationships of the Japanese endemic Pseudopyxis (Rubiaceae-Paederieae). Pl. Spec. Biol. 4: 145155. _______. 2003, in press. Rubiaceae. In: Hedberg, I., Sileshi Nemomissa & Edwards, S. (eds), Flora of Ethiopia and Eritrea, vol. 4,1 Nat. Herbarium, Addis Ababa Univ. & Dept. Syst. Botany, Uppsala Univ., Addis Ababa & Uppsala. Puff, C., Igersheim, A. & Rohrhofer, U. 1993. Pseudomussaenda and Schizomussaenda (Rubiaceae): close allies of Mussaenda. Bull. Jard. Bot. Belg. 62: 3568. Sivarajan, V. V., Biju, S.D. & Mathew, P. 1992. Revision of the genus Psilanthus Hook.f. (Rubiaceae tribe Coffeeae) in India. Bot. Bull. Acad. Sinica 33: 209224. Stoffelen, P., & Robbrecht, E. 1995. De plantekunde bril opgezet. Een botanische kijk op koffie. In: Robbrecht, E. (ed.), Kawa, koffie ontsluierd: von koffiestruk tot kopje koffie. Nat. Bot. Gard. Belgium, Meise. Taylor, L. 2002. Herbal Secrets of the Rainforest, 2nd ed. Prima Publishing, Roseville, Califonia. Verdcourt, B. 1976. Rubiaceae (part 1). In: Polhill, R. M. (ed.), Flora of Tropical East Africa. Agents for Overseas Governments and Administration, London. Wrigley, G. (1988). Coffee. Tropical Agriculture Series, Longman, London.
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Figure 2. AD Coffea arabica; EF C. canephora: A, E. flowers; BC, F. fruits; D. a, sectioned fruit showing two seeds, b, seed enclosed in endocarp, c, two seeds (endosperm covered by silver skin), d, seed, portion cut off to show position of embryo, e, embryo; G. Cinchona calisaya. AC, EF photographed by C. Puff; D. paintings by O. Van de Kerckhove, from Robbrecht & Stoffelen (1995: Fig. 20), modified, reproduced with permission of the National Botanic Garden of Belgium.
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Figure 3. A. Carphalea kirondron; B. Carphalea kirondron inflorescence showing red enlarged calyx lobes and some corollas; C. Pentas lanceolata; D. Serissa japonica; E. Rondeletia odorata; F. Hamelia patens; G. Hamelia patens inflorescence, note coloured inflorescence axes; H. Mussaenda philippica cultivars (white: cv. Dona Auror; pink: cv. Queen Sirikit or possibly Dona Luz); I. M. erythrophylla; J. Pseudomussaenda flava; K. Gardenia augusta, double flower. All photographed by C. Puff.
THAI FOR. BULL. (BOT.) 31: 95104. 2003.
Notes on the genus Alpinia (Zingiberaceae) in Thailand

SURAPON SAENSOUK*, PRANOM CHANTARANOTHAI* & KAI LARSEN**
ABSTRACT. Two new records from Thailand, Alpinia scabra (Blume) Baker and A. blepharocalyx K. Schum. with two varieties are described and illustrated.
The genus Alpinia is the largest in the Zingiberaceae with more than 200 species. It occurs throughout tropical Asia to New Guinea, Australia, the Solomon Islands, New Hebrides, New Caledonia, Fiji and Samoa (Larsen et al., 1998). In Thailand 15 species were enumerated by Larsen (1996). During revision of this genus for the Flora of Thailand, Alpinia scabra, A. blepharocalyx var. blepharocalyx and A. blepharocalyx var. glabrior (Hand.-Mazz.) T.L. Wu were found in Thailand. Furthermore one new species was discovered (Saensouk et al., in prep.). At present, 17 species and two varieties of Alpinia are recognized in Thailand. Alpinia scabra (Blume) Baker in Hook.f., Fl. Brit. India 6: 256. 1892; Ridl., J. Str. Br. Roy. Asiat. Soc. 32: 164. 1899; K. Schum. in Engl., Bot. Jahrb. 26: 275 t.2, fig. C. 1899; K. Schum. in Engl., Pflanzenr. 20 (IV, 46): 334. 1904; Boerlage, Icon. Bogor. 4: 223. 1914; Ridl., Fl. Malay Penins. 4: 279. 1924; Smith, Edinb. J. Bot. 47: 57. 1990. Hellina scabra Blume, Enum. Pl. Javae 1: 60. 1827. Languas scabra (Blume) Burk., Gard. Bull. Singapore 6: 260. 1930; Holttum, Gard. Bull. Singapore 13: 158. 1950. Type: Indonesia, Blume s.n. (holotype ?L, not seen). Figs. 2,3. Leafy shoots 23 m tall. Leaves: sheaths with short stiff hairs near base of petiole and ligule; ligule ovate, ca. 1 by 0.5 cm, apex rounded, shortly pubescent; petioles up to 1.5 cm long, shortly pubescent; lamina oblong, 2550 by 46 cm, with stiff hairs on both surfaces, scattered stiff hairs along margin, base cuneate to attenuate, apex acuminate to caudate, to 3.5 cm long. Inflorescence paniculate, 3040 cm long, usually with 23 large branches in the lower part, each subtended by long sheaths; sheaths 23, elliptic, 1520 by 1.5 cm, apex acuminate to caudate to 1.5 cm; peduncles shortly pubescent; rachis rather stout, shortly pubescent, bearing many cincinni; peduncles of cincinni 12.5 cm long with up to 6 flowers on each. Bracts towards base of inflorescence very small, ovate, 28 by 2 mm, apex acute, shortly pubescent. Bracteoles ovate, ca. 1 by 1 mm, apex acute, shortly pubescent. Pedicels slender, ca. 5 mm long, green, glabrous. Flowers white; calyx tubular, 57 by 3 mm, white, apex equally 3lobed, tip of lobes shortly pointed, pubescent; corolla tube slender, 68 by 3 mm, white; dorsal corolla lobe oblong, ca. 10 by 3 mm, apex hooded, up to 3 mm long, white, glabrous
*Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. **Department of Systematic Botany, University of Aarhus, Denmark.
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with crimson spots; lateral lobes oblong, ca. 6 by 3 mm, apex hooded, white, glabrous with crimson spots; labellum shorter than the corolla lobes, obovate, 79 by 45 mm, white, cleft almost to the base, the 2 halves with wavy margins; lateral staminodes elliptic to 1 mm long, glabrous. Filaments ca. 10 mm long, glabrous; anthers ca. 5 by 3 mm; anther-crest ca. 2 mm long, triangular, membranous, glabrous. Ovary ovoid, ca. 1 by 1 mm, glabrous; style slender, glabrous; stigma cup-shaped, glabrous except for cilia along margin. Epigynous glands 2, flat, ca. 3 mm long. Fruit unknown. Thailand. PENINSULAR: Yala [Betong district, Hala Bala Wildlife Sanctuary, 8 Feb. 1997, P. Puudjaa 329 (AAU, BKF)]. Distribution. Peninsular Malaysia and Indonesia (Java). Ecology. In tropical rain forest. Phenology. Flowering February. Note. A. scabra is similar to A. galanga but differs in having larger and richer inflorescences branches, the tip of calyx lobes shortly pointed, the labellum shorter than the corolla lobes and the apex of labellum cleft to the base. The leaf surface is scabrate. Alpinia blepharocalyx K. Schum. in Engl., Pflanzenr. 20 (IV, 46): 334. 1904; T.L. Wu, H.T. Tsai, S.Q. Tong, P.S. Chen, S.W. Zhao & H.W. Li, Fl. Reip. Pop. Sin. 16(2): 89. 1981; Smith, Edinb. J. Bot. 47: 57. 1990; Larsen, Thai For. Bull. (Bot.) 24: 35. 1996; J.H. Shan in Z. Wu & P.H. Raven, Fl. China 24: 339. 2000 Type: China, Henry 11962 (holotype K!; isotype E). A. bracteata Roxb., Hort. Bengal.: 2. 1814; Fl. Ind. 1: 61. 1820; Baker in Hook.f., Fl. Brit. India 6: 256. 1892; Gagnep. in Lecompte, Fl. IndoChine 1: 95. 1908., non A. bracteata Rosc., Trans. Linn. Soc. 11: 281. 1814. Figs. 1, 4.
KEY TO THE VARIETIES 1. Lower surface of lamina densely villous var. blepharocalyx 1. Lower surface of lamina sparsely pubescent along the margin and midrib, otherwise glabrous var. glabrior
var. blepharocalyx Leafy shoots 23 m tall. Leaves: sheaths shortly pubescent along margin near base of ligule and on petiole; ligules oblong-ovate or ovate, 0.51 by 0.51 cm, apex acuterounded or truncate, densely pubescent; petioles 1.54 cm long, glabrous or pubescent; lamina oblong to lanceolate, 5560 by 712 cm, upper surface glabrous, lower surface densely villous, base attenuate, apex acuminate to caudate or mucronate. Inflorescence a drooping raceme, 1020 cm long, when young covered by 1 broad sheath; peduncles 1.510 cm long, densely pubescent; rachis stout, 1520 cm long, densely brown pubescent, bearing many cincinni, peduncles of 1-flowered cincinni 0.51 cm long, densely brown pubescent. Bract absent. Bracteoles green to dark red when young, dry and brittle when mature, broadly obovate when flattened, 2.74.5 by 22.5 cm, apex equally bilobed, pubescent. Pedicels 0.51 cm long, shortly brown pubescent. Calyx tubular, 2.42.6 by 1 cm, apex unequally 3-lobed, deeply split to 1 cm on one side, pubescent; corolla tube 22.5 cm long, shorter than calyx, pubescent; dorsal corolla lobe
NOTES ON THE GENUS ALPINIA (ZINGIBERACEAE) IN THAILAND (S. SAENSOUK, P. CHANTARANOTHAI & K. LARSEN)
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oblong, 2.54 by 1.72.5 cm, apex bluntly hooded, shortly pubescent; lateral lobes smaller, oblong, 1.73.5 by 11.5 cm, apex bluntly hooded, shortly pubescent; labellum strongly concave, red, yellow along margin, obovate to suborbicular, 3.55 by 3.54 cm, margin crinkled, apex 3-lobed, 2 lobes nearly equal, mid-lobe small, apex 2- to 3dentate, pubescent at base of labellum; lateral staminodes small, curved, oblong to linear, 78 mm long, apex acuminate or unequally 2-dentate, sparsely pubescent. Filaments 1520 by 4 mm, glabrous or sparsely shortly pubescent; anthers 1215 by 5 6 mm, shortly pubescent; anther-crest 1 mm long, apex acute, shortly pubescent along margin or wanting. Ovary ovoid, 45 by 34 mm, densely pubescent; style slender, glabrous; stigma cup-shaped, glabrous. Epigynous glands 2, ca. 5 mm long. Fruits unknown. Thailand. NORTHERN: Chiang Mai, Lampang; NORTH-EASTERN: Loei. Distribution. India, Bangladesh, Myanmar, South China, Laos and Vietnam. Ecology. Along margins of evergreen forest and open grassy pine forest, 800 2000 m. Phenology. Flowering DecemberMay. Specimens examined. Nanakorn QBG 5468 (QBG); Nanakorn QBG 8777 (QBG); Nanakorn QBG 9420 (QBG); Maxwell 0109 (CMU); Kerr 4747 (BK, C); Larsen & Larsen 47310 (AAU); Ljtnant & Niyomdham 124 (AAU), Ljtnant & Niyomdham 162 (AAU); Sadakorn 474 (BK), Sutheesorn 2263 (BK); Smitinand 10260 (BK). var. glabrior (Hand.-Mazz.) T. L. Wu in T. L. Wu & S. J. Chen, Acta Phytotax. Sin. 16(3): 35. 1978; Wu, Tsai, Tong, Chen, Zhao & Li, Fl. Reip. Pop. Sin. 16(2): 90. 1981; Shan in Wu & Raven, Fl. China 24: 339. 2000. Languas blepharocalyx var. glabrior Hand.-Mazz., Symb. Sin. 7: 1322. 1936. Type: China, Handel-Mazzeti 5750 (holotype ?W, not seen). Figs. 34. Lower surface of lamina sparsely pubescent along margin and midrib, otherwise glabrous. Thailand. NORTHERN: Chiang Mai; NORTH-EASTERN: Loei. Distribution. India, Bangladesh, Myanmar, South China, Laos and Vietnam. Ecology. Along margins of evergreen forest and open grassy pine forest, 1000 1400 m. Phenology. Flowering FebruaryApril. Specimens examined. Nanakorn s.n. (QBG); Kerr 8662 (BK, C); Kerr 20144 (BK); Suwatabhan 113 (BK); Srensen et al. 2326 (AAU). ACKNOWLEDGEMENTS We are deeply indebted to Department of Biology, Faculty of Science, Khon Kaen University and the AAU Herbarium for their facilities during this study. The first author is very grateful to Mrs Supee S. Larsen, who kindly advised and helped in many
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ways while the first author was in Aarhus. Many thanks to Mr Piyakaset Suksathan and Mr Charun Maknoi for valuable advice and also to Ms Kamolhathai Phulphong for the photographs. Warm thanks also to the Curators and staff of the herbaria visited (AAU, BK, BKF, C, CMU, KL, PSU, QBG and S). The Golden Jubilee Ph.D. Program (Thailand) is acknowledged for financial support. REFERENCES Larsen, K. 1996. A preliminary checklist of the Zingiberaceae in Thailand. Thai For. Bull. (Bot.) 24: 3549. Larsen, K., Lock, J.M., Maas, H. & Maas, P.J.M. 1998. Zingiberaceae. In: Kubitzki, K. et al. (eds), The Families and Genera of Vascular plants IV: Flowering Plants: Monocotyledons, pp. 474495. Springer-Verlag, Berlin.
Figure 1. Lower surface of leaves in varieties of Alpinia blepharocalyx: A. var. blepharocalyx; B. var. glabrior. Scale bars = 5 cm.
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Figure 2. Alpinia scabra: Specimen collected from the Hala Bala Wildlife Sanctuary, Betong, Yala by P. Puudjaa, 8 Feb. 1997 (AAU, BKF).
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Figure 3. Alpinia scabra: A. flower; B. bract; C. bracteole; D. ligule; E. calyx; F. dorsal corolla lobe; G: lateral lobe; H: labellum and lateral staminodes; I: stamen; J: ovary, style and epigynous glands; K: style and stigma. Scale bars = 1 cm.
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Figure 4. Flowering specimens of Alpinia blepharocalyx: A. A. blepharocalyx var. blepharocalyx; B. A. blepharocalyx var. glabrior. Photographed by K. Phulphong (A) and S. Saensouk (B).
THAI FOR. BULL. (BOT.) 31: 105112. 2003.
Karyology of Jatropha (Euphorbiaceae) in Thailand

PUANGPAKA SOONTORNCHAINAKSAENG* & THAYA JENJITTIKUL **
ABSTRACT. The karyology of five species of Jatropha (Euphorbiaceae) in Thailand was investigated. Chromosomes of the microsporocytes of Jatropha were well stained with propionocarmine. Most of them were paired as bivalents at first metaphase and separated to 11:11 at first anaphase. Chromosomes of the taxa studied are of very small size. Bivalent length ranges from 13.67 m. Most species have chromosome numbers of 2n = 22 and a base number of x = 11. J. curcas is chromosomally similar to J. multifida. Both of these taxa have a meiotic configuration of 7ringII + 4rodII. J. integerrima Red flower and Pink flower have the same meiotic configuration of 6ringII +5rodII. J. podagrica has a meiotic configuration of 8ringII + 3rodII. The karyology of J. gossypifolia was determined from first anaphase cells: the chromosomes separated to 11:11. J. curcas and J. multifida and, perhaps, J. gossypifolia appear closely related to each other based on their meiotic configuration and morphological similarity.
INTRODUCTION Jatropha L. is a diverse and widespread genus of 175 species (Airy Shaw, 1972). It is particularly common in Tropical America and Africa, with a few species native to Southwest Asia. Asia and one species in Madacascar. None are native to Southeast Asia, Australasia or Oceania. J. curcas L., the Physic Nut, yields a powerful purgative. The bark of J. dioica Sesse is used for tanning and dyeing. Several species are cultivated, mainly in the tropics (Smith, 2001). There are five species of Jatropha cultivated in Thailand. J. curcas L. and J. gossypifolia L. are used as medicinal plants, whilst J. integerrima Jacq., J. multifida L. and J. podagrica Hook. are attractive ornamentals (Chayamarit et al., 2001). Jatropha curcas has potential as a source of a combustible organic oil. There are many previous records of chromosome numbers in Jatropha as shown in Table 1. Here we report the karyology of the five species present in Thailand. MATERIALS AND METHODS Jatropha curcas, J. gossypifolia, J. integerrima Red flower and Pink flower, J. multifida and J. podagrica (Fig. 1) were collected from naturalised populations. Flower buds were fixed in Carnoys solution and stored in 70 % ethyl alcohol under refrigeration. Slide preparations were made using the propionocarmine squash technique.
*,** Department of Plant Science, Faculty of Science, Mahidol University, Rama VI Rd., Payathai, Bangkok 10400, Thailand. *Corresponding author.
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Karyological observations were made from well spread chromosomes at stages of pollen mother cell development. Chromosome numbers were determined approximately from 25 cells viewed under a light microscope (Olympus model BHA). The best cells were photographed at 100x using an oil immersion objective. Voucher specimens were made for all samples and placed in the Department of Plant Science Herbarium at Mahidol University, the Department of Biology Herbarium at Khon Kaen University and the Royal Garden Suanluang, Rama IX, Bangkok. RESULTS AND DISCUSSION The chromosomes of the microsporocytes of Jatropha were well stained with propionocarmine. Most of them were paired in bivalents at first metaphase and separated to 11:11 at first anaphase (Table 2, Fig. 2). The chromosomes of the Jatropha taxa studied are of very small size. Bivalent length ranges from 13.67 m. The results suggest that Jatropha is fully fertile and stable in its naturalised habitat. Most of the taxa have chromosomes number of 2n = 22 and a base number of x = 11 respectively. These results agree with previous studies (Table 1). The meiotic configuration of Jatropha is as follows: J. curcas is similar to J. multifida. Both of them have meiotic configurations of 7ringII + 4rodII. J. integerrima Red flower and Pink flower show the same meiotic configuration of 6ringII + 5rodII. J. podagrica has a meiotic configuration that is different from the others of 8ringII + 3rodII. J. gossypifolia was determined in first anaphase cells: the chromosomes separated to 11:11 respectively. J. curcas and J. multifida and, perhaps, J. gossypifolia appear closely related to each other based on their meiotic configuration and morphological similarity. J. curcas is the only species which was found to be both diploid and tetraploid. As with many genera, such as Bauhinia, Cassia, Crotalaria, Desmodium and Vigna (Leguminosae), all members of Jatropha have the same chromosome numbers within a ploidy level. J. integerrima Red flower and Pink flower have the same chromosome number and meiotic configuration. Chromosome numbers of most of the studied species of Jatropha are in accordance with those of previous studies and are stable over a wide geographical range. ACKNOWLEDGEMENTS This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training grant BRT 140002.
KARYOLOGY OF JATROPHA (EUPHORBIACEAE) IN THAILAND (P. SOONTORNCHAINAKSAENG & T. JENJITTIKUL)
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Table 1. Previous records of chromosome numbers of Jatropha.

Chromosome number G 1. Jatropha capensis Sond. 2. J. cardiophylla Arg. 3. J. cathartica Teran & Berland 4. J. cordata Arg. 5. J. curcas L. 11,S 22 22 22 22 22;44 Dehgan (1984) Missouri Botanical Garden (2003) Missouri Botanical Garden (2003) Missouri Botanical Garden (2003) Darlington & Wylie (1965); Missouri Botanical Garden (2003); Moore (1973); Somboonsarn (1983); Soontornchainaksaeng & Chaiyasut (1999) Sinpatananon & Hongthongdaeng (1993) Darlington & Wylie (1965); Missouri Botanical Garden (2003); Moore (1973); Somboonsarn (1983); Soontornchainaksaeng & Chaiyasut (1999) Missouri Botanical Garden (2003) Missouri Botanical Garden (2003) Soontornchainaksaeng & Chaiyasut (1999) Soontornchainaksaeng & Chaiyasut (1999) Missouri Botanical Garden (2003) Darlington & Wylie (1965); Missouri Botanical Garden (2003); Soontornchainaksaeng & Chaiyasut (1999) Missouri Botanical Garden (2003) Moore (1973); Soontornchainaksaeng & Chaiyasut (1999)
Species
References
6. J. glandulifera Roxb. 7. J. gossypifolia L.
11
22 22
8. J. hetrophylla Heyne 9. J. integerrima Jacq. Red flower Pink flower 10. J. macrorhiza Benth. 11. J. multifida L.
11 11 11
44 22 22 22 22 22
12. J. pandurifolia Andrews 13. J. podagrica Hook. G = gametophytic count S = sporophytic count
11 -
22 22
Table 2. Chromosome number and meiotic configurations of Jatropha in Thailand.

Species J. curcas L. J. gossypifolia L. J. integerrima Jacq. Red flower Pink flower J. multifida L. J. podagrica Hook. Chromosome number 2n 22 22 22 22 22 22 22 n 11 11 11 11 11 11 11 x 11 11 11 11 11 11 11 Meiotic configuration s 11II 11:11 11II 11II 11II 11II 11II Locality Nakhon Pathom Nakhon Pathom Bangkok Bangkok Nakhon Pathom Nakhon Pathom Coll. no. 041 042 194 195 196 197
Coll. No. = P. Soontornchainaksaeng et al. collection number II = bivalent
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REFERENCES Airy Shaw, H. K. 1972. The Euphorbiaceae of Siam. Kew Bull. 26(2): 191363. Chayamarit, C., Santisuk, T., Larsen, K., Welzen, P. van., Esser, H. J., Nanakorn, W., Chantaranothai, P., Boonthavikoon, T., Pooma, R., Phuphathanaphong, L., Chantharaprasong, C. & Larsen, S. 2001. Systematic study of the family Euphorbiaceae in Thailand. In: Baimai, V. & Kumhom, R. (eds), BRT Research Report 2001, pp. 7888. Biodiversity Research and Training Program, Bangkok, Thailand. Darlington, C.D. & Wylie, A.P. 1965. Chromosome atlas of flowering plants. George Allen and Unvin, London. Dehgan, B. 1984. Phylogenetic significance of interspecific hybridization in Jatropha (Euphorbiaceae) Syst. Bot. 9: 467478. Missouri Botanical Garden. 2003. Index of plant chromosome numbers (ICPN) database. URL: http://www.mobot.mobot.org. Moore, D.M. 1973. Index to plant chromosome numbers for 19671971. Oosthoeks Uitgeversmaatschappij B. V., Domstraat 513, Utrecht, Netherlands. Sinpatananon, A. & Hongthongdaeng, P. 1993. Chromosome number of local mulberry varieties. In: Modern genetics. 8th Seminar on genetics, 29 March1 April 1993, pp. 7679. Somboonsarn, N. 1983. Morphological, anatomical and cytological investigation of some hydrocarbon plants. M.Sc. Thesis, Graduate school, Kasetsart University, Bangkok. Soontornchainaksaeng, P. & Chaiyasut, K. 1999. Cytogenetic investigation of some Euphorbiaceae in Thailand. Cytologia 64: 229234.
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Figure 1. Morphological variation of Jatropha (Euphorbiaceae) in Thailand: A. J. curcas L.; B. J. gossypifolia L.; C. J. integerrima Jacq. Red flower; D. J. integerrima Jacq. Pink flower; E. J. multifida L.; F. J. podagrica Hook.
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Figure 2. Chromosomes in the pollen mother cells of Jatropha. A. First metaphase chromosomes of J. curcas (2n = 22; 7ringII + 4rodII). B. First anaphase chromosomes of J. gossypifolia (n = 11; 11:11). C. First metaphase chromosomes of J. integerrima (cv. Red flower) (2n = 22; 6ringII + 5rodII). D. First metaphase chromosomes of J. integerrima (cv. pink flower) (2n = 22; 6ringII + 5rodII). E. First metaphase chromosomes of J. multifida (2n = 22; 7ringII + 4rodII). F. First metaphase chromosomes of J. podagrica (2n = 22; 8ringII + 3rodII). Scale bar = 10 m.
THAI FOR. BULL. (BOT.) 31: 113122. 2003.
Cytogenetic studies and taxonomic considerations in some taxa of Mallotus (Euphorbiaceae) in Thailand
PUANGPAKA SOONTORNCHAINAKSAENG*, PRANOM CHANTARANOTHAI** & CHADAPRON SENAKUN***
ABSTRACT. An investigation into the cytogenetics of Thai Mallotus (Euphorbiaceae) indicated that all plants have highly distinctive chromosomes at diakinesis and first anaphase. Bivalent length is 1.336 m in pollen mother cells of 160286.67 m. The chromosome numbers of Mallotus can be divided into three groups, viz: 2n = 20 (10II), 22 (11II) and 24 (12II). Chromosome numbers of 11 species are recorded for the first time. The basic numbers of Mallotus are x = 10, 11 and 12. These dysploid numbers may have evolved through a decrease from 2n = 22 to 2n = 20. M. oblongifolius is taxonomically conspecific with M. peltatus but their chromosome numbers are different with 2n = 20 and 2n = 22 respectively. Therefore, these two species should be maintained as distinct taxa.
INTRODUCTION Mallotus Lour. is a paleotropical genus and comprises 150 species ranging from Africa and Madagascar through Southeast Asia and Malesia to Australia and the western Pacific. There are 35 species in Thailand of which two are new records and three newly described. They are used as medicinal plants (M. barbatus, M. floribundus, M. macrostachyus, M. paniculatus and M. philippensis), for wood (M. miquelianus), for dye (M. philippensis) and as ornamental plants (M. floribundus) (Chayamarit et al., 2001). Cytogenetics have played an active part in the understanding of hereditary mechanisms of plant diversity. They also represent one of the numerous disciplines that plant improvement relies upon. The data from this study will be useful for future taxonomic work on Mallotus. MATERIALS AND METHODS Chromosome counts were made from 13 species of Mallotus collected at various locations in Thailand (Table 1). The chromosomes were isolated from young flowers which were fixed in Carnoy's solution for 2448 hours. The anthers were squashed and stained with propionocarmine and the chromosomes then examined using 100x magnification through an Olympus-BHA light microscope. Counts were made from 20 well-spread cells from each species at various stages of meiois. Voucher specimens are deposited at the herbaria of Department of Plant Science, Mahidol University, Department of Biology, Khon Kaen University and the Royal Garden Suanluang, Rama IX, Bangkok.
*Department of Plant Science, Faculty of Science, Mahidol University, Rama VI Rd, Payathai, Bangkok 10400, Thailand. Corresponding author. **Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. ***Walairukhavaj Botanical Research Institute, Mahasarakam University, Mahasarakam, Thailand.
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RESULTS AND DISCUSSION Chromosomes of the microsporocytes of Mallotus were very well stained with propionocarmine. Most of them were identified at diakinesis with a bivalent length 1.336 m in cells of 160286 m. Chromosome numbers and meiotic figures could be divided into three groups. Group I had 2n = 20 (10II) and comprised M. oblongifolius and M. pallidus. Group II had 2n = 22 (11II), and comprised M. barbatus, M. macrostachyus, M. miquelianus, M. montanus, M. peltatus, M. repandus and Mallotus sp. Group III had 2n = 24 (12II) and comprised M. paniculatus, M. resinosus and M. spodocarpus (Table 1; Figs. 1933). Most species studied showed balanced first anaphase cells. Chromosome numbers of 11 species were recorded for the first time. The results revealed that Mallotus has basic numbers of x = 10, 11 and 12. These agree with the findings of other workers. Perry (1943) reported that M. japonicus has 2n = 36, Soontornchainaksaeng & Chaiyasut (1999) and Missouri Botanical Garden (2003) reported that M. babatus has 2n = 22 and M. apelta and M. philippensis have 2n = 11, respectively. These dysploid numbers may reflect ploidy level variation combined with hybridisation in species or cytotypes during evolution. Dysploidy is of relatively frequent occurrence among seed plants within populations or genera. Airy Shaw (1972) accepted both M. oblongifolius and M. peltatus as distinct species. However, van Welzen et al. (2000) reduced M. oblongifolius to a synonym of M. peltatus, suggesting that the only difference between them was the absence or presence of a peltate leaf base. However, specimens (referred to M. peltatus) with peltate leaves often have non-peltate ones and specimens (always referred to M. oblongifolius) with non-peltate leaves often have peltate ones. Our cytological investigation showed that the chromosome numbers of these two species are different, suggesting that the two taxa should be maintained as distinct. Further study is needed to confirm their status. ACKNOWLEDGEMENTS The authors would like to thank Dr Kongkanda Chayamarit, Assistant Professor Chirayupin Chantharaprasong, Mrs Leena Phuphathanaphong, Mr Phongsak Phonsena and the staff of the Euphorbiaceae Project in Thailand for kindly providing valuable information and some materials. This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training-BRT 140002. REFERENCES Airy Shaw, H.K. 1972. The Euphorbiaceae of Siam. Kew Bull. 26: 191363. Chayamarit, K., Santisuk, T., Larsen, K., Welzen, P.V., Esser, H. J., Nanakorn, W., Chantaranothai, P., Boonthavikoon, T., Pooma, R., Phuphathanaphong, L., Chantharaprasong, C. & Larsen, S. 2001. Systematic study of the family Euphorbiaceae in Thailand. In: Baimai, V. & Kumhom, R. (eds) BRT Research Report 2001, pp. 7888. Biodiversity Research and Training Program (BRT), Bangkok, Thailand.
CYTOGENETIC STUDIES AND TAXONOMIC CONSIDERATIONS IN SOME TAXA OF MALLOTUS (EUPHORBIACEAE) IN THAILAND
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Missouri Botanical Garden. 2003. Index of plant chromosome numbers (ICPN) database. URL: http://www.mobot.mobot.org. Perry, B. A. 1943. Chromosome number and phylogenetic relationships in the Euphorbiaceae. Amer. J. Bot. 30: 527543. Soontornchainaksaeng, P. & Chaiyasut, K. 1999. Cytogenetic investigation of some Euphorbiaceae in Thailand. Cytologia 64: 229234. Welzen, P. C. van. 2000. Checklist of the genera of Thai Euphorbiaceae I. Thai For. Bull. (Bot.) 28: 59112. Table 1. Meiotic chromosome counts of Thai Mallotus II = bivalent, Ref. = reference number; prev. = Soontornchainaksaeng, P. & Chaiyasut K. (1999); Coll. no. = P. Soontornchainaksaeng et al.; *Coll. No. = C. Senakun & W. Tongpubal.
Species 1. M. barbatus Mll. Arg. 2. M. macrostachyus Mll. Arg. 3. M. miquelianus Boerl. 4. M. montanus Airy Shaw 5. M. oblongifolius Mll. Arg. 6. M. pallidus Airy Shaw 7. M. paniculatus Mll. Arg. 8. M. peltatus Mll. Arg. 9. M. repandus Mll. Arg. 10. M. resinosus Merr. 11. M. spodocarpus Airy Shaw 12. M. tiliifolius Mll. Arg. 13. Mallotus sp. Chromosome Number 2n n x 22 22 22 22 20 20 24 22 22 24 24 24 22 11 11 11 11 10 10 12 11 11 12 12 12 11 11 11 11 11 10 10 12 11 11 12 12 12 11 Meiotic figure 11II 11II 11II 11II 10II 10II 12II 11II 11II 12II 12II 12II 11II Record /(Ref.) prev 1st 1st 1st 1st 1st 1st 1st 1st 1st 1st 1st Locality Prachuap Khiri Khan Trang Songkhla Songkhla Songkhla Prachuap Khiri Khan Songkhla Chanthaburi Sa Kaeo Prachuap Khiri Khan Lop Buri Chumphon Chachoengsao Coll. no. 148 375 276 281, 291 278 261 296 411 386 *70 203 150 138
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117
7 8
10
Figures 110. Mallotus spp.: 1. M. barbatus; 23. M. macrostachyus; 45. M. miquelianus; 6. M. montanus; 78. M. paniculatus; 910. M. pallidus.
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119
13
11
12
14
15
16
17
18
Figures 1118. Mallotus spp.: 1112. M. paniculatus; 1314. M. peltatus; 15. M. resinosus; 16. M. spodocarpus; 17. M. tiliifolius; 18. Mallotus sp.
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121
19
20
21
22
23
24
25
26
27
28
29
30
33 31 32
Figures 1933. Diakinesis cells of 13 species of Mallotus: Figs. 1921. 2n = 20; 19. & 20. M. oblongifolius, 21. M. pallidus Figs. 2229. 2n = 22; 22. M. barbatus, 23. M. macrostachyus, 24. M. miquelianus, 25. M. montanus, 26. M. peltatus, 27. M. rependus, 28. & 29. Mallotus sp. Figs. 3033. 2n = 24; 30. M. paniculatus, 31. M. resinosus, 32. M. spodocarpus, 33. M. tiliifolius Arrows indicate nucleolus. Scale bar = 10 m.
THAI FOR. BULL. (BOT.) 31: 123128. 2003.
Capparis sikkimensis Kurz subsp. yunnanensis (Capparaceae), a new record for Thailand
PRACHAYA SRISANGA*, CHUSIE TRISONTHI** & KONGKANDA CHAYAMARIT*** ABSTRACT. Capparis sikkimensis Kurz subsp. yunnanensis (Craib & W.W. Sm.) Jacobs is newly recorded for Thailand from Doi Phu Kha National Park in Nan province. A description and illustration of the taxon are provided.
In the account of Capparaceae for the Flora of Thailand (Chayamarit, 1991), 25 indigenous and one introduced species of Capparis L. were recorded. During an expedition by the first author to Doi Phu Kha National Park, Nan province in March 2000, specimens belonging to this genus were collected. They were later identified as C. sikkimensis Kurz subsp. yunnanensis (Craib & W.W. Sm.) Jacobs, a new record for Thailand. The description below is based on the Thai material. Capparis sikkimensis Kurz subsp. yunnanensis (Craib & W.W. Sm.) Jacobs, Blumea 12 (3): 496. 1965. C. yunnanensis Craib & W.W. Sm., Notes Bot. Gard. Edin. 9: 91. 1916; Merr., J. Arn. Arb. 23: 167. 1942. Type: China, Yunnan, A. Henry 12986 (holotype E; isotype A, K, US). C. roxburghii (non DC.) Dunn, J. Linn. Soc. Lond. Bot. 39: 426. 1911. C. bhamoensis Raizada, Ind. For. Rec. n.s. 3: 127, fig. 4. 1941. C. fohaiensis B.S. Sun, Acta Phytotax. Sin. 9: 114. 1964. Figs. 12. Scandent shrub or large climber, 46 m high; branches terete, shallowly furrowed, straight, brown pubescent, later glabrescent; young shoots and inflorescences not surrounded by cataphylls; thorns slightly recurved, in pairs, 34 mm long. Leaves chartaceous, elliptic or ovate, (4.5)1113 by (3)56.5 cm; apex acute or cuspidate; base rounded or broadly cuneate; margin entire; pubescent on both surfaces, later glabrescent; midrib impressed above, raised and prominent below; secondary veins in 68 pairs, prominent on lower surface; petioles 710 mm long, brown-pubescent. Inflorescences solitary in leaf axils or a lax corymb (24-flowered) axillary or on the terminal part of the twig; peduncle 24.5 cm long, brown-pubescent. Flowers fragrant; pedicels up to 4.5 cm long, brown pubescent. Sepals 4, biseriate, free, orbicular concave or shallowly saccate, shortly clawed, 1319 by 1118 mm, inner pair slightly smaller, brown pubescent on both sides. Petals 4, yellow, free, obovate, 2025 by 812 mm, densely tomentose inside, glabrous except for tomentose basal parts outside. Torus 45 mm wide, pubescent. Stamens 6080; filaments ca. 3.5 cm long, hairy at base; anthers ca. 2 mm long, glabrous, dehiscing by lateral slits. Gynophore 2.54 cm long, hairy at base. Ovary ellipsoid, 34 by 23 mm, glabrous; placentas 4. Fruits ellipsoid, 45 by 2.53 cm; pericarp 25 mm thick; stipe 78.5 cm long. Seeds 34, 1520 by 1015 mm.
*Herbarium, Queen Sirikit Botanic Garden, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand. **Biology Department, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand. ***The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
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Thailand. NORTHERN: Nan [Doi Phu Kha National Park 19 13 N 101 06 E, Srisanga 1359 (BKF, QBG); Srisanga 1692 (QBG); Srisanga 2337 (QBG)]. Distribution. China, Myanmar and Vietnam. Ecology. Lower montane forest, 15001600 m altitude. Flowering: March April. Fruiting: SeptemberJanuary. Note. C. sikkimensis subsp. yunnanensis was originally described as C. yunnanensis Craib & W.W. Sm. (1916), and later reduced to subspecific rank under C. sikkimensis Kurz by Jacobs (1965). C. sikkimensis is divided into four subspecies: C. sikkimensis subsp. sikkimensis distributed in north-eastern India and western Myanmar; C. sikkimensis subsp. yunnanensis (Craib & W.W. Sm.) Jacobs in northern Myanmar, China (Yunnan) and Vietnam (Tonkin); C. sikkimensis subsp. masaikai (Lvl.) Jacobs in southern and south-eastern China; and C. sikkimensis subsp. formosana (Hemsl.) Jacobs distributed in Riu Kiu Islands, Taiwan and Hainan. As noted by Jacobs, C. sikkimensis is highly polymorphic, and for his division into subspecies, no more than 30 collections were available. More material, especially from Myanmar-Yunnan, is needed to re-examine the status of the subspecies. For now we have followed Jacobss treatment. C. sikkimensis subsp. yunnanensis is distinguished by its brownish pubescent sepals on both sides, large petals (2025 by 812 mm), long filaments (ca. 3.5 cm) and long gynophore (2.54 cm). ACKNOWLEDGEMENTS Much appreciation is due to the curators and staff of BKF, CMU, KUN, P and QBG for their kind permission to study specimens, and to the staff of Doi Phu Kha National Park for assistance in the field. We are grateful to Ms Soraya Klankoom for the line drawing and Mr Pasakorn Kaewket for technical assistance. This work was supported by a grant from the Thailand Research Fund. REFERENCES Chayamarit, K. 1991. Capparaceae. In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 5(3): 241271. Craib, W. G. & Smith, W. W. 1916. Diagnoses Specierum Novarum. Notes Bot. Gard. Edinb. 9: 9192. Jacobs, M. 1965. The genus Capparis (Capparaceae) from the Indus to the Pacific. Blumea 12 (3): 385541.
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Figure 1. Capparis sikkimensis Kurz subsp. yunnanensis (Criab & W.W. Sm.) Jacobs: A. flowering branch; B. flower. Scale bars = 1 cm. Drawn from Srisanga 1359 (QBG). Drawn by S. Klankoom.
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CAPPARIS SIKKIMENSIS KURZ SUBSP. YUNNANENSIS (CAPPARACEAE), A NEW RECORD FOR THAILAND
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Capparis sikkimensis Kurz subsp. yunnanensis (Capparaceae), a new record for Thailand
A
A B
Figure 2. Capparis sikkimensis Kurz subsp. yunnensis (Craib & W.W. Sm.) Jacobs: A. flowers; B. fruit.
Radermachera eberhardtii (Bignoniaceae), a new record for Thailand
Figure 2. Radermachera eberhardtii Dop: A. inflorescence; B. infructescences.
THAI FOR. BULL. (BOT.) 31: 129131. 2003.
Radermachera eberhardtii (Bignoniaceae), a new record for Thailand

PRACHAYA SRISANGA*, CHUSIE TRISONTHI** & THAWATCHAI SANTISUK***
ABSTRACT. Radermachera eberhardtii Dop, a new record for Thailand, is described and illustrated. The key to Radermachera in the Flora of Thailand is emended. Radermachera microcalyx C.Y. Wu & W.C. Yin, previously known from China (Guangxi and Yunnan), is placed in the synonymy of R. eberhardtii.
Radermachera Zoll. & Moritzi consists of ca. 15 species. The distribution is in tropical Asia: India (Assam), Myanmar, South China, Thailand, Cambodia, Laos, Vietnam, Malaysia, Taiwan, Ryu Kyu Is., Philippines, Borneo and Indonesia (Steenis, 1976; Santisuk, 1985). In the account of Bignoniaceae for the Flora of Thailand (Santisuk, 1987), five indigenous species in the genus Radermachera Zoll. & Moritzi are recorded. During an expedition by the first author to Doi Phu Kha National Park, Nan province, specimens belonging to this genus were collected. They were later identified as R. eberhardtii Dop, a new record for Thailand. C.Y. Wu and W.C.Yin (1979) established R. microcalyx for a plant from southern China (Guangxi and Yunnan). They recognised this species as showing affinity with R. glandulosa (Blume) Miq., but differing in some quantitative characters, i.e. smaller calyx (35 mm long and wide). After examining some of the paratype specimens cited in the protologue (C.W. Wang 75298, 76948 & 81097, all in KUN), those collections of R. microcalyx matched well with R. eberhardti (i.e. yellow flower, more or less straight corolla tube, tubular calyx with five irregular lobes, and leaflets with some scattered immersed minute glands on the lower surface. Hence, the authors feel that R. microcalyx should be treated as conspecific with R. eberhardtii. The following emendment is made to the Flora of Thailand Radermachera key:
1. Leaves 1-pinnately compound 1.*Calyx cupular, evenly truncate or subtruncate. Corolla tube slightly curved, white tinged pink or purplish. Capsules straight, crowded, hanging in bunch 1. R. glandulosa 1.*Calyx tubular with 5 irregular lobes. Corolla tube more or less straight, yellow. Capsules tortuous, widely spaced 6. R. eberhardtii 1. Leaves 2- or 3-pinnately compound or biternate
Radermachera eberhardtii Dop, Bull. Mus. Hist. Nat. (Paris) 32: 233. 1926, incl. var. acuta Dop, l.c.: 234; in Fl. Indo-Chine 4: 587. 1930; Steenis, Blumea 23 (1): 125. 1976; Santisuk & Vidal in Fl. C.L.V. 22: 17. 1985; Santisuk, Nat. Hist. Bull. Siam Soc. 33(2):
*Herbarium, Queen Sirikit Botanic Garden, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand. **Biology Department, Faculty of Science, Chiang Mai University, Chiang Mai 50200, Thailand. ***The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok. 10900, Thailand.
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71. 1985. Type: Vietnam, Eberhardt 1594 (P!, 2 sheets). R. microcalyx C.Y. Wu & W.C. Yin in Wu, Fl. Yunnan. 2: 711, fig. 197, 14. 1979; Zhang & Santisuk in Wu & Raven, Fl. China 18: 218. 1998. synon. nov. Type: China, Yunna, Xishuangbanna, Yan Hui Li 380 (not located). Figs. 1, 2 (p. 127). Evergreen tree 715 m high. Branchlets terete, lenticellate, minutely glandular. Leaves 1 pinnate, 2530 cm long; rachis minutely glandular; leaflets 57; lamina chartaceous to subcoriaceous, elliptic to ovate, (10)1420(25) by (3.5)4.56(11) cm; apex acute, acuminate or caudate; base cuneate, oblique; margin entire; both surfaces glabrous, minutely glandular throughout and with sparse medium glands around base on lower surface; midrib raised and prominent below; secondary veins in 610 pairs; petioles 12 cm or up to 6 cm in terminal leaflet, minutely glandular. Flowers in terminal panicle, ca. 20 cm long, minutely glandular; pedicels 510 mm long, minutely glandular. Calyx green, 67 mm long, minutely glandular, medium glands in groups at middle; tube 45 mm long; lobes 5, irregular, ca. 2 mm long, apex acute. Corolla yellow, 33.5 cm long, minutely glandular; tube 2.53 cm long; lobes ovate to orbicular, 0.61 cm long, hairy especially outside, margin fimbriate. Stamens 4, didynamous, hairy at insertion; filaments glabrous, shorter pair ca. 4 mm long, longer pair 56 mm long; anthers glabrous, 1.52 mm long, dehiscing by longitudinal slits. Staminode 1, ca. 2 mm long. Ovary cylindric, 2 2.5 mm long, minutely glandular; style 710 mm long, glabrous; stigma bilobed. Disc ca. 0.5 mm high, glabrous. Capsule terete, curved, 1025 cm long, minutely glandular, dehiscing by 2 valves; valves 46 mm wide. Seeds flat, 815 by 12 mm included wing. Thailand. NORTHERN: Nan [Doi Phu Kha National Park, 19 13 N 101 05 E, Srisanga 1324 (BKF, QBG); Srisanga 1450 (QBG)]; Chiang Mai [Doi Suthep, Maxwell 88-276 (AAU)]. Distribution. China (Guangxi and Yunnan) and Vietnam. Ecology. By shady stream in evergreen forest, 300950 m altitude. Flowering MarchApril. Fruiting AprilDecember. ACKNOWLEDGEMENTS Much appreciation is due to the curators and staff of AAU, BKF, CMU, KUN, P and QBG for their kind permission to study specimens and staff of Doi Phu Kha National Park for assistance in the field. We are grateful to Ms Soraya Klankoom for the line drawing and Mr Pasakorn Kaewket for technical assistance. This work was supported by a grant from the Thailand Research Fund. REFERENCES Santisuk, T. 1985. Bignoniaceae of mainland South-east Asia. Nat. Hist. Bull. Siam Soc. 33(2): 61101. ________. 1987. Bignoniaceae. In: Smitinand, T. & Larsen, K. (eds), Flora of Thailand 5(1): 3266. Steenis, C.G.G.J. van. 1976. Conspectus of the genera Radermachera and Fernandoa in Indo-Malesia (Bignoniaceae). Blumea 23: 121138. Wu, C. Y. and W. C. Yin. 1979. Bignoniaceae. In: Wu, C. Y. (ed.), Flora Yunnanica 2: 711. Zhang, Z. and T. Santisuk. 1998. Bignoniaceae. In: Wu, C. Y. and Raven, P.H. (eds), Flora of China 18: 218.
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Figure 1. Radermachera eberhardtii Dop: A. leaves; B. inflorescence; C. corolla; D. opening corolla; E. calyx; F. stamen; G. fruit; H. seed. AF from Srisanga 1324 (BKF, QBG); GH from Srisanga 1450 (QBG). Drawn by Soraya Klankoom.
THAI FOR. BULL. (BOT.) 31: 132135. 2003.
Rhodoleia (Hamamelidaceae), a new generic record for Thailand

SOMRAN SUDDEE* & DAVID J. MIDDLETON**
ABSTRACT. The genus Rhodoleia with one species, Rhodoleia championii Hook.f., is newly recorded for Thailand. The genus and species are described and a new key to the Hamamelidaceae is presented.
The Hamamelidaceae has recently been published in the Flora of Thailand and included five genera with a total of six species (Phengklai 2001). Curiously, the genus Rhodoleia was not known from Thailand even though the single species, Rhodoleia championii Hook.f., has been recorded from Southern China, Burma, Vietnam, Sumatra and Peninsular Malaysia (Vink, 1957). As part of a joint collecting initiative between the Forest Herbarium in Bangkok and Harvard University Herbarium the authors found several trees of Rhodoleia championii in Kaeng Krachan National Park, Phetchaburi province. In addition a number of collections of this species have also recently been made from Sukhirin District, Narathiwat by P. Puudjaa and C. Niyomdham. Exell (1933) distinguished seven species in Rhodoleia. Vink (1957), however, reduced these to one species and provided a discussion of the variation patterns in this polymorphous species. Endress (1993) notes that a revision is needed of the genus but in the meantime we will follow Vinks species concept. RHODOLEIA Champion ex Hook.f., Bot. Mag. III, 6: t. 4509. 1850; Exell, Sunyatsenia 1: 95. 1933; Vink, Fl. Malesiana ser. 1, 5: 371. 1957; Endress in Kubitzki, Fam. Gen. Vasc. Pl. 2: 329. 1993. Bisexual shrubs or trees. Leaves simple, entire, secondary venation pinnate. Stipules mostly absent. Flowers bisexual, connate, in peduncled heads, surrounded by an involucre of bracts which are caducous after anthesis. Sepals rudimentary. Petals 04, only those along the outer margin of the head well developed, spathulate to obovate. Stamens 611, each with 4 pollen sacs, dehiscing by 2 septicidal slits to produce 4 narrow valves. Ovary half-inferior, 2-locular. Ovules 68; styles 2, long and slender. Capsules basally united, woody. Seeds narrowly winged.
*The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900. Thailand **Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, USA.
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One species (but see note above) in Southern China, Indochina, Sumatra and Peninsular Malaysia. One species in Thailand. Rhodoleia championii Hook.f., Bot. Mag. III, 6: t.4509. 1850; Exell, Sunyatsenia 1: 96. 1933; Vink, Fl. Malesiana ser. 1, 5: 372. 1957. Type: Hong Kong, Champion s.n. (holotype K; isotypes BM, E). For synonymy see Vink (1957). Fig. 1. Evergreen tree to 40 m tall, dbh to 1 m. Bark rusty-brown, with large paler lenticels, inner bark pale salmon pink, wood white. Leaves: petioles red, 0.84 cm long; blades dark green above, glaucous beneath, ovate, 4.412.2 by 25.3 cm, apex acute to shortly acuminate, base acute to obtuse, midrib flat to slightly sunken above, prominent beneath, secondary veins flat above and beneath, in 79 pairs at a steep angle to the midrib, anastomosing before margin, tertiary venation laxly reticulate. Flowers recurved. Bracts shortly ovate to obovate-oblong, 315 by 410 mm, upper bracts largest, brownishtomentose on abaxial side, glabrous on adaxial side. Sepals rudimentary, apically ciliate. Petals bright red, 1218 by 13 mm, glabrous. Filaments 11.5 cm long, glabrous; anthers elliptic-oblong, 12 mm long. Ovary glabrous. Fruit heads green, ripening brown, 1.92.6 cm across. Seeds 34.5 by 2.53 mm, sterile seeds wingless, fertile seeds narrowly winged. Thailand. SOUTH-WESTERN: Phetchaburi [Kaeng Krachan District, Kaeng Krachan National Park, trail from Khao Phanoen Thung Ranger Substation to Than Thip Waterfall, 1000 m alt., 28 March 2003, David J. Middleton, Chandee Hemrat, Stuart Lindsay, Somran Suddee & Suwat Suwanachat 1780 (A, AAU, BKF, K, L, MO)]; PENINSULAR: Narathiwat [Sukhirin District, Khao Nakharach, 665 m alt., 20 Oct. 1996, Chawalit Niyomdham 4834 (BKF); idem., 8 August 1996, Pachok Puudjaa 254 (BKF); idem., 17 Feb. 1997, Pachok Puudjaa 385 (BKF)]. Distribution. Southern China, Burma, Vietnam, Sumatra, Peninsular Malaysia. Ecology. Primary evergreen forest to 1000 m altitude. Note. With the addition of another genus to the Hamamelidaceae of Thailand a new key, adapted from Phengklai (2001), is presented.
1. Sepals and petals absent. 2. Ovary semi-inferior. Capsules united into a globular or obpyramidal head. Seeds winged 2. Ovary superior. Capsules free, ellipsoid. Seeds not winged 1. Either sepals (sometimes minute) or petals or both present 3. Leaves palmately nerved 3. Leaves pinnately nerved 4. Stipules mostly absent. Sepals rudimentary 4. Stipules present or, if caducous, leaving a scar. Sepals not rudimentary, sometimes caducous 5. Sepals 6. Petals absent. Lower half of capsule surrounded by a receptacle 5. Sepals 4. Petals present. Capsule not surrounded by receptacle
Altingia Distylium Symingtonia Rhodoleia Sycopsis Loropetalum
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ACKNOWLEDGEMENTS We would like to thank the National Geographic Society for financial support for this work. We would also like to thank Orathai Kerdkaew for the illustrations, Chandee Hemrat, Stuart Lindsay and Suwat Suwanachat for their assistance in the field, and the staff of Kaeng Krachan National Park. REFERENCES Endress, P.K. 1993. Hamamelidaceae. In: Kubitzki, K. et al. (eds), The Families and Genera of Vascular Plants II: Flowering Plants: Dicotyledons, pp. 322331. Springer-Verlag, Berlin. Exell, A.W. 1933. A revision of the genus Rhodoleia. Sunyatsenia 1: 95102. Phengklai, C. 2001. Hamamelidaceae. In: Santisuk, T. et al. (eds), Flora of Thailand 7: 400411. Vink, W. 1957. Hamamelidaceae. Flora Malesiana ser. 1, 5: 363379.
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Figure 1. Rhodoleia championii Hook.f.: A. habit; B. inflorescence; C. capsules; D. seeds. A. from D.J. Middleton et al. 1780 (BKF), BD from P. Puudjaa 385 (BKF). Drawn by O. Kerdkaew.
THAI FOR. BULL. (BOT.) 31: 136140. 2003.
A preliminary study of Gonystylaceae in Thailand

CHAWALIT NIYOMDHAM*& METINEE TARUMATSAWAT*
ABSTRACT. Gonystylaceae is considered to be a rare plant family. One species, Gonystylus confusus Airy Shaw, is indigenous to Thailand. The species is found only in peat swamp forest and lowland dipterocarp forest: a few specimens have been collected in Narathiwat province.
Gonystylaceae is considered to be a rare plant family. One species, Gonystylus confusus Airy Shaw, is indigenous to Thailand and is found only in peat swamp forest and lowland dipterocarp forest. A few specimens have been collected in Narathiwat province and are described herein. GONYSTYLACEAE Trees or shrubs. Leaves simple, alternate, exstipulate. Inflorescence usually paniculate. Flowers hermaphrodite, actinomorphic. Calyx 5-lobed, imbricate; petals small, numerous; anthers basifixed, 2-loculed, opening by longitudinal slits. Ovary 35loculed. Style filiform. Stigma small, capitate. Ovules solitary in each locule, pendulous from near the apex. Fruit a capsule; seed large, without endosperm. A monotypic family of about 30 species, from the Nicobar Is., through Malesia to the Solomon Is. and Fiji. GONYSTYLUS Teysmann & Binnendijk, Bot. Zeit. 20 : 265. 1862; Ridl., Fl. Mal. Pen. 1: 321323. 1922; Airy Shaw, Hook. Ic. Pl. 35. t.34743475. 1947; Kew Bull. 916. 1947; Kew Bull. 138147. 1950; Fl. Mal. 4: 349. 1953; Whitmore, Tree Fl. Mal. 2: 387. 1973. Trees or shrubs. Leaves alternate, chartaceous to coriaceous with translucent gland dots. Inflorescence paniculate, the main branches few, more or less elongate, the lateral branches short, consisting of extremely condensed, irregular, nodulose racemes, often reduced to fascicles. Bracts small, falling very early. Flowers longpedicelled. Calyx thickly coriaceous, more or less cupular, imbricate or sub-valvate, calyx-teeth 5 slightly unequal (3 larger and 2 smaller), always hairy inside. Petals often joined at base, 740, deltoid or subulate. Stamens about equal in number to the petals, rarely twice as many; filaments very short and slender, free; anthers basifixed, broadly or
*The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Chatuchak, Bangkok 10900, Thailand.
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narrowly oblong to obovate, 2-locular. Ovary sessile, globose, (2)35(8)-locular; style filiform, elongate, wiry, glabrous or pubescent; stigma small, capitate. Ovules solitary in each cell, pendulous from the apex, anatropous. Fruit a capsule, with thick woody wall, splitting into 25 parts. Seeds large, arillate. One species in Thailand. Gonystylus confusus Airy Shaw, Kew Bull. 2: 10. 1947; Fl. Malesiana 4: 357. 1953; Whitmore, Tree Fl. Mal. 2: 191. 1973; Phengklai & Niyomdham, Fl. peat swamp areas Narathiwat: 242. 1991. Type: Malaysia, Perak, Larut, Kings Collector 3778 (holotype K, not seen). Figs. 13. Tree up to 30 m tall, 220 cm girth; bark dark brown with elongate, adherent scales; inner bark fibrous, reddish to yellowish brown; sapwood white, soft; young part and inflorescence hairy. Leaves oblong-elliptic to oblanceolate-oblong, coriaceous, 6.523.5 cm long, 3.59 cm wide; apex cuspidate to abruptly short acuminate; base subcuneate to rounded, glabrous on both surfaces, sometimes with few, adpressed hairs near the midrib beneath; venation slender, parallel, conspicuous on both surfaces; petiole 12 cm. Inflorescence terminal, and in upper leaf axils, 520 cm long. Flowers small, ca. 6 mm across. Calyx 45-lobed, 56 mm long, 34 mm wide, tomentose outside. Petals ca. 30, subulate-filiform 34 mm long, joined at the very base. Ovary hairy, (2)3(4)- loculed. Fruit globose-ellipsoid, ellipsoid to ovoid, sitting on the remaining calyx, scurfy, rusty brown, splitting into 23(4) parts, 46 cm long, 2.53.5 cm across, (1)23-seeded, globose-ellipsoid to oblong, 2.84 cm long, 2.33.5 across; aril creamy white. Thailand. PENINSULAR: Narathiwat [Su Ngai Pa Di, Pa Wai, 11 Sept. 1985, C. Niyomdham et al. 960 (BKF); Waeng, Sirindhorn Waterfall, 20 Sept. 2001, C. Niyomdham et al. 6569 (BKF); 18 June 2003, M. Tarumatsawat 03-3 (BKF); Su Ngai Ko Lok, Toh Daeng canal, 25 July 2000, T. Noo-Yim et al. 00-36 (BKF); 22 June 2002, T. Noo-Yim et al. 02-2 (BKF); 17 June 2003, M. Tarumatsawat 03-1 (BKF); Su Ngai Ko Lok, Ban Pa Ye, rubber tree plantation, 17 June 2003, M. Tarumatsawat 032 (BKF)]. Distribution. Malay Peninsula and Indonesia (North Sumatra). Ecology. Peat swamp and evergreen low land dipterocarp forests, up to 600 m altitude. Flowering AugustNovember. Fruiting JuneSeptember. Vernacular. Tan yong phru (), Tam yao () Note. The authors have not seen type specimens of this species. However the materials available match the description in Airy Shaw (1947) very well. ACKNOWLEDGEMENTS Special thanks are due to Mr Tanit Noo-Yim, Mr Apichat Rattanavirakul, Mr Sunthorn Toh Dam and Mr Pachok Poodjaa, for their help and hospitality during field work in Narathiwat province.
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Finally, my appreciation is extended to Dr Kongkanda Chayamarit, Curator and all the scientific staff of The Forest Herbarium, who kindly helped to bring this publication out on time. REFERENCES Airy Shaw, A.K. 1947. Notes on the genus Gonostylis Teijsm. & Binnend. (Thymelaeaceae). Kew Bull. 2: 916. ________. 1953. Thymelaeaceae-Gonostyloideseae. In Steenis, C.G.G.J. van (ed.). Fl. Malesiana Ser. 1, 4: 349361.
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Figure 1. Gonystylus confusus Airy Shaw. A. Trunk and leaves. Slash, showing fibrous, reddish to yellowish brown inner bark (Photographed by M. Tarumatsawat); B. Flowering twig; C. Fruiting twigs; D. Fruits and seeds (Photographed by C. Niyomdham).
THAI FOR. BULL. (BOT.) 31: 141 148. 2003.
The Genus Porandra (Commelinaceae) in Thailand

THAWEESAK THITIMETHAROCH*, PRANOM CHANTARANOTHAI* & ROBERT B. FADEN** ABSTRACT. A revision of the genus Porandra D.Y. Hong in Thailand is presented. Three species are recognised, P. microphylla Y. Wan, P. ramosa D.Y. Hong and P. scandens D.Y. Hong. The first two species are newly recorded from Thailand. A key to the species is provided together with illustrations, distributions and ecological data.
INTRODUCTION The genus Porandra was established by Hong (1974) based on two species, namely P. ramosa D.Y. Hong and P. scandens D.Y. Hong. Since his work a third species, P. microphylla Y. Wan has been described from China (Wan, 1986). Faden (1998) considered Porandra to be probably conspecific with Amischotolype but Hong & DeFilipps (2000) maintained the genus. All species except P. scandens were considered endemic to China. The genus was separated from Amischotolype by its climbing habit and anther cells opening by apical pores. In the course of preparing Commelinaceae treatments for the Flora of Thailand, P. ramosa and P. microphylla are newly recorded for the country. In addition, although Hong (1974) and Hong & DeFillips (2000) stated that rhizomes were absent, although we found long horizontal underground rhizomes in P. microphylla and P. scandens and they may also be present in P. ramosa. TAXONOMIC ACCOUNT PORANDRA D.Y. Hong in Acta Phytotax. Sin. 12(4): 462. 1974; Faden, Fam. Gen. Vas. Pl. 4: 120. 1998. Type species: Porandra ramosa D.Y. Hong. Perennial rhizomatous scandent herbs. Leaf blades alternate; petiole unwinged. Inflorescence a very compact globose head, subsessile. Flowers bisexual, subsessile. Sepals equal, sepaline. Petals free, equal, not clawed. Stamens 6, free, equal, all fertile; filaments bearded; anthers dehiscing by apical pores. Capsules trilocular, trivalved, seeds 2 per locule. Seeds arillate. Three species mainly from southern China and adjacent areas. All the species are found in Thailand.
*Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. **Department of Botany, Smithsonian Institution, P.O. Box 37012 Washington DC 20013-7012, USA.
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KEY TO SPECIES 1. Anthers drip-shaped. Leaves often densely pubescent beneath, rarely glabrous with dense pubescence at the apex. Petioles, sepals and capsule densely pubescent 2. P. ramosa 1. Anthers subglobose-oblong or lanceolate-oblong. Leaves usually glabrous, rarely sparsely pubescent beneath. Petioles glabrous. Sepals and capsule sparsely pubescent or glabrous 2. Leaves (15)1825 by 35.5 cm, glabrous, densely ciliate along margins 3. P. scandens 2. Leaves 515(16) by 1.54.5 cm, glabrous or with a line of dense puberulence near margin above, margin glabrous or ciliate at apex 1. P. microphylla
1. Porandra microphylla Y. Wan in Bull. Bot. Res. North-east. Forest. Inst. 6(4): 153. 1986; D.Y. Hong & DeFilipps in Fl. China 24: 24. 2000. Type: China, Guangxi, Long an, Longhushan Conservation Area, Wan Fu 85011 (holotype IBG, not seen; isotype GXSP, not seen). Figs. 1AF & 2A. Woody-like scandent herb, stems to 3 m long, ascending to 1.5 m tall. Leaf blade narrowly elliptic-lanceolate, 515(16) by 1.54.5 cm, glabrous or with a line of dense puberulence above near the margin, base cuneate to obtuse, apex abruptly acuminate to caudate, margins slightly undulate, glabrous or ciliate at the apex; petioles pale green or dark purple, 58 mm long, glabrous; leaf sheaths 26.5 by 0.31 cm, densely puberulent on one side, mouth long-ciliate. Inflorescence a head ca. 1.5 cm in diam., usually with (2)46 flowers; bracteoles ovate, 2.53 by 34 mm, sparsely pubescent on the outside and margins. Flowers white. Sepals green, ellipsoid, 67 by 2.54 mm, pubescent on the outside. Petals white, broadly elliptic or oblanceolate, 67 by 2.53 mm. Stamens: filaments 68 mm long, bearded; anthers subglobose-oblong, ca. 11.2 by 1 mm. Ovary subglobose, ca. 1.2 mm in diam., sparsely puberulent; style ca. 10 mm long. Capsules green, 13 in a head, ovoid, 810 by 68 mm, pubescent throughout. Seeds ellipsoidreniform, ca. 4 by 3 mm. Thailand. NORTHERN: Chiang Mai (Doi Chiang Dao, Doi Inthanon). Distribution. China. Ecology. Common in hill-evergreen forest, alt. 8002000 m; flowering and fruiting from August to February. Vernacular. Phak plap doi () (Doi Chiang Dao-Chiang Mai). Notes. P. microphylla is distinct because of its leaves which are smaller and rather glabrous and its anthers which are subglobose-oblong and often smaller than in the other species. It was first recorded from China, but now it seems to have a much wider range. Specimens examined: G. Murata et al. T-15173 (BKF), T-15658 (BKF); C. Phengklai et al. 7170 (BKF, K); T. Smitinand & I. Alsterlund 6656 (K); T. Smitinand & J.A.R. Anderson 7267 (BKF); T. Thitimetharoch 458 (Herb. Khon Kaen University). 2. Porandra ramosa D.Y. Hong in Acta Phytotax. Sin. 12(4): 462. 1974; D.Y. Hong & DeFilipps in Fl. China 24: 24. 2000. Type: China, Yunnan, Feng Qing, T.T. Y 16255 (holotype PE, not seen). Figs. 1GL & 2B.
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Woody-like scandent herb. Leaf blade elliptic-lanceolate, 1016 by 34 cm, densely brownish or rusty pubescent beneath or rarely densely pubescent at the apex, base acute to cuneate, apex acuminate to caudate, margins ciliate; petioles dark purple, 36 mm long, densely pubescent beneath or on both sides; leaf sheaths 35 by 0.31 cm, pubescent throughout and/or on one side, mouth long-ciliate. Inflorescence a head 1.52 cm in diam., usually with (4)68 flowers; bracteoles broadly ovate, 2.54 by 35 mm, densely pubescent especially on midvein outside, margins often sparsely ciliate. Flowers pinkish purple. Sepals pale green or pinkish purple, ellipsoid, 78 by 34 mm, densely pubescent along midvein outside, margins often sparsely ciliate. Petals pinkish, elliptic-oblanceolate, 78.5 by 2.53 mm. Stamens: filaments 710 mm long, pinkishbearded; anthers purple, drip-shaped, 22.5 by 1.21.5 mm. Ovary subglobose, ca. 1 mm in diam., pubescent, style 1013 mm long. Capsules purple, 13(6) in a head, ovoid, 711 by 58 mm, pubescent apically. Seeds ellipsoid-reniform, 47 by 33.5 mm.
EASTERN:
Thailand. NORTHERN: Chiang Mai (Doi Inthanon, Mae Ka Pak); Loei (Khawk Moei-Phu Kradung). Distribution. China.
NORTH-
Ecology. In hill-evergreen and gallery forests, alt. 10001500 m; flowering and fruiting from April to December. Notes. P. ramosa is readily distinguished from all other members of its genus by its unique drip-shaped anthers and densely pubescent petioles, sepals, capsules and leaves. Formerly known only from southern China but now known to occur in northern and north-eastern Thailand. Specimens examined: H.B.G. Garrett 561 (K), 1132 (K, US); T. Smitinand 2526 (BKF, K). 3. Porandra scandens D.Y. Hong in Acta Phytotax. Sin. 12(4): 462. 1974; D.Y. Hong & DeFilipps in Fl. China 24: 24. 2000. Type: China, Yunna, Xishuangbanna, C.W. Wang 77909 (holotype PE). Figs. 1MR & 2C. Woody-like scandent herb, stems to 6 m long. Leaf blade elliptic-lanceolate, 18 25 by 35.5 cm, base acute-cuneate, apex acuminate-caudate, margins ciliate; petioles pale green or dark purple, 310 mm long, sometimes sparsely pubescent above; leaf sheaths ca. 4.5 by 0.5 cm, densely pubescent on one side, mouth ciliate. Inflorescence a small compact globose head ca. 1.5 cm in diam., usually with (4)68(10) flowers; bracteoles broadly ovate, 3.54 by 45 mm, glabrous, margins sometimes sparsely ciliate. Flowers white. Sepals pale green or pinkish purple, ellipsoid, 68 by 45 mm, boat-shaped, glabrous or sparsely pubescent on outside. Petals greenish white, ellipticoblanceolate, 68 by 2.53.5 mm. Stamens: filaments 68 mm long, white- or purplishbearded; anthers purple, oblong-lanceolate, 1.52.5 by 11.5 mm. Ovary subglobose, ca. 1 mm in diam., glabrous or sparsely pubescent, style 1013 mm long. Capsules 1 3(6) in a head, broadly ovoid or subglobose, 811 by 59 mm, glabrous or sparsely pubescent on upper part. Seeds ellipsoid-reniform, 47.5 by 35 mm. Thailand. NORTHERN : Chiang Mai (Doi Chiang Dao, Doi Suthep, Fang), Chiang Rai (Mae Sariang, Mae Sruai), Kamphaeng Phet (Klong Lan, Mae Wong),
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Lampang (Jae Sawn), Tak (Um Phang); NORTH-EASTERN: Loei (Phu Kradung, Phu Luang); EASTERN: Chaiyaphum (Ban Nam Phrom, Phu Kheio); SOUTH-EASTERN: Chantaburi (Khao Soi Dao); SOUTH-WESTERN: Kanchanaburi (Kwae Noi River Basin, Si Sawat), Ratchaburi (Suan Pueng). Distribution. China, Laos, Vietnam. Ecology. In evergreen forest, alt. 2501800 m; flowering and fruiting from April to December, flowers opening in late afternoon. Vernacular. Yaa ton (), Phak plap doi () (Chiang Dao-Chiang Mai); Phak kap plee () (Kamphaeng Phet); Sang phai () (Phu Kradung-Loei). Uses. Cooked young leaves are edible (Suan Phueng-Ratchaburi). Notes. P. scandens is widespread from southern China to Thailand; Y. Paisooksantivatana & P. Sangkachand ,from Ratchaburi province ,(BK) 872083-Y .xtent of its rangee most-southern represents the Specimens examined: K. Bunchuai 1684 (K), 1170 (K); H.B.G. Garrett 1409 (K); R. Geesink, T. Hattink & C. Phengklai 6700 (K), 6850 (K); R. Geesink & P. Hiepko 7890 (BKF, K); A.F.G. Kerr 3234 (K); A. Kostermans 834 (BKF, K); K. Larsen 43579 (US); K. Larsen, T. Santisuk & E. Warncke 2396 (BKF, K), 2546 (K), 2695 (BKF, K); J.F. Maxwell 73187 (BK), 95728 (BKF); C. Niyomdham et al. 4377 (BKF); Y. Paisooksantivatana & P. Sangkhachand Y-2083-87 (BK); Prayad 939 (BK); Put 3271 (BK, BM, K), 4545 (BK, BM, K); J.F. Rock 176 (US), 241 (US); S. Sutheesorn 2598 (BK); T. Thitimetharoch 410 (Herb. Khon Kaen University), 425 (Herb. Khon Kaen University), 445 (Herb. Khon Kaen University), 472 (Herb. Khon Kaen University); P. Triboun 1300 (Herb. Khon Kaen University). ACKNOWLEDGEMENTS We would like to thank the Royal Golden Jubilee Program (RGJ: grant No. 4.B.KK/42/B.1) for financial support. We are very grateful to the directors and curators of BCU, BK, BM, BKF, CMU, K, LINN, QBG, PSU, US herbaria and also Khon Kaen University Herbarium for permission to study specimens. We also thank the Department of Biology, Faculty of Science, Khon Kaen University and the Smithsonian Institution, Washington for their facilities. REFERENCES Faden, R. B. 1998. Commelinaceae. In: Kubitzki, K. et al. (eds), The Families and Genera of Vascular Plants IV: Flowering Plants: Monocotyledons, pp. 109128. Springer-Verlag, Berlin. Hong, D. Y. 1974. Revisio Commelinacearum Sinicarum. Acta Phytotax. Sinica 12(4): 459 488.
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Hong, D. Y. & DeFilipps, R. A. 2000. Commelinaceae. In: Wu, Z. Y. & Raven, P. H. (eds), Flora of China (Flagellariaceae through Marantaceae) Vol. 24, pp. 1939. Science Press, Beijing.
Figure 1. AF: Porandra microphylla Y. Wan: A. leaves; B. sepal; C. petal; D. stamen; E. pistil; F. capsule and persistent calyx. GL: P. ramosa D.Y. Hong: G. leaves; H. sepal; I. petal; J. stamen; K. pistil; L. capsule and persistent calyx. MR: P. scandens D.Y. Hong: M. leaf; N. sepal; O. petal; P. stamen; Q. pistil; R. capsule and persistent calyx. Drawn by T. Thitimetharoch.
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Figure 2. A. Porandra microphylla Y. Wan; B. P. ramosa D.Y. Hong, from T. Smitinand 2526 (K); C. P. scandens D.Y. Hong. Photographed by T. Thitimetharoch.

Thai Forest Bulletin 31

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THAI FOREST BULLETIN

125 131 134 136 143

THAI FOR. BULL. (BOT.) 31: 16. 2003.

Ceropegia hirsuta (Asclepiadaceae), a new record for Thailand

THAI FOREST BULLETIN (BOTANY) 31

THAI FOREST BULLETIN (BOTANY) 31

THAI FOR. BULL. (BOT.) 31: 712. 2003.

Notes on two Ixora species, new records for Thailand

THAI FOREST BULLETIN (BOTANY) 31

THAI FOREST BULLETIN (BOTANY) 31

THAI FOREST BULLETIN (BOTANY) 31

Ixora cambodiana Pit.

Ixora henryi H. Lv.

Figure 3. Distribution of Ixora cambodiana Pit. and I. henryi H. Lv. in Thailand.

THAI FOR. BULL (BOT.) 31: 1326. 2003.

The Rubiaceae of Ko Chang, south-eastern Thailand

THAI FOREST BULLETIN (BOTANY) 31

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

THAI FOREST BULLETIN (BOTANY) 31

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

THAI FOREST BULLETIN (BOTANY) 31

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

Figure 2. Distribution of Rennellia speciosa (based on collections in Thai herbaria).

THAI FOREST BULLETIN (BOTANY) 31

Plectronia glabra (Blume) Kurz Plectronia siamensis K. Schum. *

see text according to Craib (1932); see text

X (Craib, 1932) X (Craib, 1932; as H. congesta)

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

see text XX X (Craib, 1934)

* *; see text *; see text *; see text

*; synonomy according to Craib (1932); see text

Psychotria adenophylla Wall. P. angulata Korth. Psychotria sp. ?

P. rubra Poir. P. sarmentosoides Val. P. serpens L. Psydrax nitidum (Craib) Wong

THAI FOREST BULLETIN (BOTANY) 31

Name in Schumann (1902) same Randia eucodon K. Schum. * same *

Notes and comments

Borreria hispida (L.) K. Schum.

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

THAI FOREST BULLETIN (BOTANY) 31

THE RUBIACEAE OF KO CHANG, SOUTHEASTERN THAILAND (V. CHAMCHUMROON & C. PUFF)

THAI FOR. BULL. (BOT.) 31: 2733. 2003.

Two new species of Neohouzeoua (Gramineae-Bambusoideae) from Thailand and Myanmar

THAI FOREST BULLETIN (BOTANY) 31

TWO NEW SPECIES OF NEOHOUZEOUA (GRAMINEAE-BAMBUSOIDEAE) FROM THAILAND AND MYANMAR

THAI FOREST BULLETIN (BOTANY) 31

TWO NEW SPECIES OF NEOHOUZEOUA (GRAMINEAE-BAMBUSOIDEAE) FROM THAILAND AND MYANMAR

Comparative morphology of Neohouzeoua mekongensis A. Camus, N. kerriana and N. fimbriata

erect erect, thin walled

THAI FOREST BULLETIN (BOTANY) 31

TWO NEW SPECIES OF NEOHOUZEOUA (GRAMINEAE-BAMBUSOIDEAE) FROM THAILAND AND MYANMAR

THAI FOR. BULL. (BOT.) 31: 3438. 2003.

A new description of Diospyros coaetanea (Ebenaceae)

A NEW DESCRIPTION OF DIOSPYROS COAETANEA (EBENACEAE) (S. DUANGJAI & C. PHENGKLAI)

We have also emended the key to fruiting specimens as follows:

THAI FOREST BULLETIN (BOTANY) 31

A NEW DESCRIPTION OF DIOSPYROS COAETANEA (EBENACEAE) (S. DUANGJAI & C. PHENGKLAI)

THAI FOR. BULL. (BOT.) 31: 3943. 2003.

*Department of Systematic Botany, University of Aarhus, Denmark.

THAI FOREST BULLETIN (BOTANY) 31

THAI FOREST BULLETIN (BOTANY) 31

THAI FOR. BULL. (BOT.) 31: 44 46. 2003.

Notes on Clerodendrum (Lamiaceae)

NOTES ON CLERODENDRUM (LAMIACEAE) (C. LEERATIWONG & P. CHANTARANOTHAI)

* ; see text ; see text *; see text