Thai Fores Thai Forest Bulletin

t Bulletin (Botany) Vol. 46 No. 2, 2018 (Botany)

Vol. 46 No. 2, 2018 ISSN 0495-3843 (print) ISSN 2465-423X (electronic)

Forest Herbarium Department of National Parks, Wildlife and Conservation Chatuchak, Bangkok 10900 http://www.dnp.go.th/botany ISSN 0495-3843 (print) ISSN 2465-423X (electronic)

Fores t Herbarium Department of National Parks, Wildlife and Plant Conservation Bangkok, THAILAND THAI FOREST BULLETIN (BOTANY) Thai Forest Bulletin (Botany) Vol. 46 No. 2, 2018

Published by the Forest Herbarium (BKF) CONTENTS Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900, Thailand Page Advisors Wipawan Kiaosanthie, Wanwipha Chaisongkram & Kamolhathai Wangwasit. Chamlong Phengklai & Kongkanda Chayamarit A new of Scleria P.J.Bergius (Cyperaceae) from North-Eastern Thailand 113–122 Editors Willem J.J.O. de Wilde & Brigitta E.E. Duyfjes. Miscellaneous Cucurbit News V 123–128 Rachun Pooma & Tim Utteridge Hans-Joachim Esser. A new species of () from Thailand, and lectotypifications of names for related taxa 129–133 Managing Editor Assistant Managing Editor Orporn Phueakkhlai, Somran Suddee, Trevor R. Hodkinson, Henrik Æ. Pedersen, Nannapat Pattharahirantricin Sawita Yooprasert Priwan Srisom & Sarawood Sungkaew. Dendrobium chrysocrepis (Orchidaceae), a new record for Thailand 134–137 Editorial Board Rachun Pooma (Forest Herbarium, Thailand), Tim Utteridge (Royal Botanic Gardens, Kew, UK), Jiratthi Satthaphorn, Peerapat Roongsattham, Pranom Chantaranothai & Charan David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin, Leeratiwong. The Campylotropis (Leguminosae) in Thailand 138–150 Ireland), David J. Middleton ( Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis Alan Paton, Somran Suddee & Bhanubong Bongcheewin. Chelonopsis thailandica, Biodiversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München, a new species and new record of Chelonopsis () from Thailand 151–154 Germany), Bob Harwood (Northern Territory Herbarium, Darwin, Australia), André Schuiteman Nutdanai Putthisawong & Sahut Chantanaorrapint. A revision of the genus Tapeinidium (Royal Botanic Gardens, Kew, UK), Anders S. Barfod (Aarhus University, Denmark), (Lindsaeaceae) in Thailand 155–161 Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Pimwadee Pornpongrungrueng (Khon Kaen University, Thailand), Stuart Lindsay (National Parks Board, Singapore) Caroline Byrne, John Adrian Naicker Parnell & Kongkanda Chayamarit. Systematics of the Thai and with comments on the Kielmeyeroidae Thai Forest Bulletin (Botany) (TFB) publishes papers on plant (especially of vascular ), (Clusiaceae) 162–216 nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology, cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines. The journal now uses Thai Journal Online (ThaiJO) for online submission and peer review at www.tci-thaijo.org/index.php/ThaiForestBulletin. Manuscripts are considered on the understanding that their contents have not appeared, or will not appear, elsewhere in the same or abbreviated form. Before submitting a manuscript please read the Guidelines for authors. These guidelines must be followed precisely otherwise publication of the manuscript will be delayed. In addition, papers published online will be distributed simultaneously in printed form to several libraries, and bound hard copy volumes will appear later. Exchange with botanical journals or periodicals pertaining to plant taxonomy would be appreciated.

FOREST HERBARIUM Director: Phongsak Phonsena Curator: Nannapat Pattharahirantricin BKF Staff: Somran Suddee, Piyachart Trisarasri, Preecha Karaket, Thanongsak Jonganurak, Pachok Puudjaa, Voradol Chamchumroon, Nanthawan Suphuntee, Narong Koonkhunthod, Montri Saengsawasti, Naiyana Tetsana, Sukontip Sirimongkol, Manop Poopath, Sommanussa Saengrit, Sukid Rueangruea, Baramee Sakolrak, Sawita Yooprasert, Saksan Kaitongsuk, Orathai Kerdkaew.

Front Cover: Brassaiopsis spinosissima Esser

Printed at: Prachachon Co., Ltd. 35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand Tel : 0 2636 6550 THAI FOREST BULL., BOT. 46(2): 113–122. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.01

A new species of Scleria P.J.Bergius (Cyperaceae) from North-Eastern Thailand

WIPAWAN KIAOSANTHIE1, WANWIPHA CHAISONGKRAM2 & KAMOLHATHAI WANGWASIT1,*

ABSTRACT Scleria aureovillosa Kiaosanthie & K.Wangwasit, a new species of Cyperaceae from North-Eastern Thailand, is described and illustrated. It is closely related to S. benthamii C.B.Clarke but diff ers in the and culm surfaces, culm shape, the absence of wings at the leaf sheath, contraligule features, nutlet morphology and micromorphology, and leaf and culm anatomy. An emended section of the key to the species in the Flora of Thailand account of Scleria is provided.

KEYWORDS: anatomy, nutlet, Scleria aureovillosa, taxonomy, Thailand. Published online: 21 September 2018

INTRODUCTION Hypoporum (Nees) C.B.Clarke, Scleria and Trachylomia (Nees) Bauters). The genus Scleria P.J.Bergius (Cyperaceae) comprises ca 250 species, mainly distributed in the During fi eldwork undertaken as part of a Ph.D. tropics and subtropics but also extending into warm study by the fi rst author, specimens were collected temperate regions (Goetghebeur, 1986 & 1998; in Phu Rua District, Loei Province, which could not Haines & Lye, 1983; Zhang et al., 2010). In the Thai be identified using existing identification keys. fl ora, the genus is the fourth largest in the Cyperaceae Moreover, the combination of characters from the with 22 species (Simpson & Koyama, 1998), and in morphology, nutlet micromorphology and anatomy surrounding regions, 29 species in , 34 species of these specimens differed from any previously in , 22 species in Indo-, 25 species known species from Thailand and elsewhere. in , 11 species in and 24 species in China, have been recorded (Clarke, 1894; Camus, MATERIALS AND METHODS 1912; Kern, 1961 & 1974; Khoi, 2002; Newman et al., 2007; Zhang et al., 2010). Scleria can be easily Materials were examined from fi eld collections recognized by the bony nutlets, which often have a and herbarium specimens at ABD, BK, BKF, BM, lobed hypogynium at the base and are not covered BO, CMUB, E, K, KEP, KKU, P, PSU, QBG and by the spikelet glumes. In the past, the genus has SING. Identifi cation was attempted using keys in been variously classifi ed into seven subgenera and the Flora of Thailand (Simpson & Koyama, 1998), up to 13 sections (e.g. Bentham & Hooker, 1883; as well as other publications such as Flora of Clarke, 1894 & 1908; Haines & Lye, 1983; Koyama, Générale de I’ Indo-Chine, the Flora of the Malay 1961; Kern, 1974). Recently, a new infrageneric Peninsula, Flora Malesiana, Flora of Vietnam, a classification by Bauters et al. (2016), based on Checklist of the Vascular Plants of Lao PDR and molecular evidence and supported by morphology, Flora of China etc. Images of the infl orescences, confi rmed Scleria as monophyletic and sister to tribe habit and habitat were taken with a Ricoh CX4 Bisboeckelereae, with four, strongly supported digital camera. Morphological observations were subgenera (Browniae (C.B.Clarke) C.B.Clarke, made using an Olympus SZ-PT stereo microscope.

1 Department of Biology, Faculty of Science, Mahasarakham University, Maha Sarakham 44150, Thailand. 2 Program of Biology, Faculty of Science, Udon Thani Rajabhat University, Udon Thani 41000, Thailand. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p113-122-G8.indd 113 1/16/62 BE 4:30 PM 114 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Nutlets were taken from dry specimens and placed awned, margins hispid. Spikelets in clusters with on a stub with carbon tape, coated with gold by 1‒2 subandrogynous spikelets and 1‒2 staminate SPI-MODULE and examined using a JEOL spikelets; prophyll usually borne at the base of spikelet, JSM6460LV scanning electron microscope (SEM) obovate, 1.5‒2 mm long, 2-keeled, sides membranous, at the Laboratory Equipment Center, Mahasarakham glabrous, reddish-brown, apex obtuse.Subandrogynous University. The outer walls of nutlets were removed spikelets ovate-oblong, 4‒5 x 1‒2 mm; with 3 by soaking in 70% sulphuric acid for 60‒90 minutes. glumes, ovate to broadly-ovate, 2‒4 x 1‒2 mm, sides The samples were then sonicated in an ultrasonic membranous, glabrous, reddish-brown, apex awned, cleaner for 30 minutes and washed with distilled lowest glume empty, shortest; lateral staminate water in the ultrasonic cleaner for a further 10 minutes. fl ower remnant present. Staminate spikelets sessile The samples were dried overnight in silica gel and to pedunculate, ovate-lanceolate, 4‒6 x 1 mm; with were examined with the SEM. Confi rmation of the many glumes, ovate to broad-ovate, 2‒4 x 1‒2 mm, siliceous on nutlet were obtained by analysis using sides membranous or hyaline, glabrous, reddish- a SEM fi tted with an energy dispersive X-ray analyser brown, apex acute, obtuse or emarginate, the two (EDX). The comparative leaf and culm anatomy lowest glumes empty, shortest, apex awned. were observed. Samples were prepared using epidermal 3, linear, 1.5 mm long, yellow to brown. Stigmas 3. peeling and paraffi n methods and stained with safranin Nutlets subglobose to globose, terete, 2.1‒2.5 x and fast greenn (Johansen, 1940; Thamathaworn, 1996). 2.1‒2.2 mm, white, apex black apiculate, surface Observations were made using an Olympus CH30 reticulate, densely irregular silica deposits, single, light microscope. 2‒4 x 2‒3.5 μm, and clusters, 5‒9.5 x 5‒10 μm, hairs 116‒173 μm long, golden villous in 3‒6 rows TAXONOMIC TREATMENT on ridges; disk well developed, 3-lobed, deeply sinus at the base, lobes broadly ovate or deltoid, apex Scleria aureovillosa Kiaosanthie & K.Wangwasit, acuminate. sp. nov. Thailand.― NORTH-EASTERN: Loei [Phu Rua, Similar to Scleria benthamii C.B.Clarke but 23 Aug. 2014, Kiaosanthie WK 1122014 (QBG!, diff ers in having trigonous culms (vs triquetrous in KKU!)]; [Phu Wua Wildlife Sanctuary, S. benthamii), an obtuse contraligule (vs rounded to Summit plateau, 200 m, 14 Oct. 1998, Muasya et al. truncate) and nutlets 2.1‒2.5 mm long (vs 2.6‒2.9 mm 1340 (T71) (K!)]; [Bung Khla, 306 m, long), subglobose to globose, terete, with a black, 28 July 2014, Kiaosanthie WK 1032014 (QBG!, apiculate apex (vs ovoid, subterete to trigonous and KKU!)]. obtuse apex) (Fig. 3 & Table 1). Type: Thailand, Distribution.― [Kampot, 27 June Loei, Phu Rua, 1,155 m, 12 Nov. 2012, Kiaosanthie 1938, Poilane 273499 (P!); Preah Sihanouk, Kampong WK 0152012 (holotype BKF [194620!]; isotypes Seila, 100 m, 13 Sept. 2013, Maxwell 13-178 KKU!, QBG!) (Figs. 1, 2 & 3A‒C). (CMUB!)]. Erect perennial herb, rhizome brown. Culms Ecology.― Growing in seasonally wet, open loosely tufted (37‒)50‒100 cm x 1‒2(‒4) mm, grassy places on hillsides and on sandy soil; trigonous, smooth and glabrous. cauline; 100‒1155 m alt. blade linear, (12‒)21‒40 cm x 2‒3 mm, acute, fl at, Phenology.― Flowering and fruiting June‒ smooth and glabrous; sheath closed, 2.5‒7.2 cm November. long, wingless, purplish to brown, slightly scabrid to smooth; contraligule obtuse, ciliate. Involucral Vernacular.― Kok luk khon thong (กกลกขนทอู ง). leaf-like, (3.2‒)7‒9(‒19.5) cm x 1‒2 mm. The Thai name translates as ‘sedge with golden hairs Infl orescences truncate, linear, loose, 14.5‒34 cm x on the nutlet surface’. (5‒)10‒15 mm; 1-noded, with 1 lateral panicle, the Etymology.― The specifi c epithet of this new panicle sometimes absent; lateral panicle linear- species is taken from the Latin aureus and villus, oblong or spike-like, loose, 4.6‒11 cm x 5‒6 mm, which refers to the distinctive feature of the species peduncle 2‒6 cm long, somewhat short branches; having golden villous hairs on the mature nutlet bracteole glume-like, (3‒)10‒15 mm long, apex surface.

SW 11611-p113-122-G8.indd 114 1/16/62 BE 4:31 PM A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND 115 (W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)

Figure 1. Scleria aureovillosa Kiaosanthie & K.Wangwasit. A. habit; B. part of infl orescence; C. contraligule; D. prophyll; E‒G. glumes; H. nutlet; I. disk 3-lobed. All from the holotype. Drawn by Wipawan Kiaosanthie.

SW 11611-p113-122-G8.indd 115 1/16/62 BE 4:31 PM 116 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 2. Scleria aureovillosa Kiaosanthie & K.Wangwasit. A‒B. part of infl orescence; C. nutlet; D. disk 3-lobed; E. rhizome; F. habitat.

SW 11611-p113-122-G8.indd 116 1/16/62 BE 4:31 PM A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND 117 (W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)

Figure 3. Scanning electron micrographs of nutlets of Scleria. A‒C. Scleria aureovillosa Kiaosanthie & K.Wangwasit, all from the Kiaosanthie WK 0152012 (BKF); D‒F. S. benthamii C.B.Clarke, all from the Kiaosanthie WK 0682013 (BKF).

SW 11611-p113-122-G8.indd 117 1/16/62 BE 4:31 PM 118 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Conservation Status.― Following the IUCN (Bauters et al., 2016). The section is mainly distributed Red List Criteria and Categories (IUCN 2012), in Africa but with eight species occurring in Thailand: Scleria aureovillosa Kiaosanthie & K.Wangwasit six species, S. rugosa R.Br., S. thwaitesiana Boeckeler, should be considered as “Least Concern” (LC). The S. tessellata Willd., S. mikawana Makino, S. parvula species is widespread in north-eastern Thailand and Steud., and S. bifl ora Roxb., are annuals, while the was recorded in Cambodia. remaining two species, S. benthamii C.B.Clarke and Notes.― Scleria aureovillosa Kiaosanthie & S. aureovillosa Kiaosanthie & K.Wangwasit, are K.Wangwasit is classifi ed in sect. Foveolidia Raf. perennials. Kern (1961, 1974) described this section based on the truncated infl orescences, subandrogy- as comprising of annuals only, while Bauters et al. nous and staminate spikelets and 3-lobed disk (2016) confirmed that the section comprises both

Table 1. Comparisons of diagnostic morphological, nutlet micromorphological and anatomical characteristics of Scleria aureovillosa Kiaosanthie & K.Wangwasit and S. benthamii C.B.Clarke.

Characters S. aureovillosa S. benthamii

Morphology Culm shape trigonous triquetrous Culm surface glabrous glabrous to pubescent Leaf surface glabrous glabrous to pubescent Leaf sheath wingless winged or wingless Contraligule obtuse rounded to truncate Infl orescence node 1 node, with 1 lateral panicle 1‒3 nodes, with 1‒2 lateral panicles (sometimes 0) Nutlet shape subglobose to globose, terete ovoid, subterete to trigonous Nutlet length 2.1‒2.5 mm 2.6‒2.9 mm Nutlet apex apiculate, black obtuse, not blac

Nutlet micromorphology Silica deposits single and clustered single Silica shape irregular rounded

Leaf anatomy Leaf types hypostomatic amphistomatic Unicellular prickles on leaf absent present margins Papillae on adaxial surface absent present Keel shape obtuse acute Leaf margin shape obtuse acute Adaxial parenchymatous present at mid rib absent hypodermis Radiated palisade cells absent present Lateral rib sclerenchyma adaxial girders adaxial strands

Culm anatomy Papillae absent present Angular sclerenchyma girders strands

SW 11611-p113-122-G8.indd 118 1/16/62 BE 4:31 PM A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND 119 (W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)

annuals and perennials. Perennial species, with C.B.Clarke. Scleria aureovillosa Kiaosanthie & golden hairs on the reticulated nutlet surface, are K.Wangwasit has hypostomatic leaves with an obtuse present in Africa (S. achtenii De Wild., S. nyasensis keel and margins, bifacial mesophyll with palisade C.B.Clarke, and S. unguiculata E.A.Rob.) and and spongy parenchyma, lateral ribs with adaxial America (S. reticularis Michx.). The American species sclerenchymatous girders, and leaf and culm surfaces is separated from the others by the without trichomes and papillae. Scleria benthamii having lateral clusters, while the three African species C.B.Clarke has amphistomatic leaves with an acute and S. aureovillosa Kiaosanthie & K.Wangwasit keel and acute margins, mesophyll with radiated have loose lateral panicles. Scleria aureovillosa palisade cells surrounding the vascular bundles, Kiaosanthie & K.Wangwasit diff ers from the three lateral ribs with adaxial sclerenchymatous strands, African species by having a single node with 1 lateral and leaf and culm surfaces with unicellular hairs and panicle (sometimes absent), the African species papillae (Figs. 4 & 5). Micromorphology also provides having at least two nodes with 1‒6 lateral panicles additional detail of the silica deposits on nutlet at each node. surfaces. Scleria aureovillosa Kiaosanthie & K.Wangwasit has both single and clusters of silica Leaf and culm anatomical investigations reveal deposits, whereas S. benthamii C.B.Clarke has only diagnostic features between Scleria aureovillosa single silica deposits (Figs. 3C & F). Kiaosanthie & K.Wangwasit and S. benthamii

The following key is emended from the key to Scleria in the Flora of Thailand Cyperaceae account (Simpson & Koyama, 1998).

21. Nutlets not patterned, smoothish to pubescent; disk lobe acute often bidentate at apex S. levis 21. Nutlets reticulate, pubescent or villous; disk lobe acuminate at apex 21a. Leaves and culms glabrous to pubescent; contraligule rounded to truncate; nutlets ovoid, subterete to trigonous, apex obtuse, not black S. benthamii 21b. Leaves and culms glabrous; contraligule obtuse; nutlets subglobose to globose, terete, apex distinctly apiculate, black S. aureovillosa

ACKNOWLEDGEMENTS Bentham, G. & Hooker, J.D. (1883). Genera This work wassupportedd by ScienceAchievement Plantarum. Cyperacaeae. V. 3. part 2. London, Scholarship of Thailand. The authors would like to Henrietta Street, Covent Garden. pp. 1037‒1073. thank Department of Biology, Faculty of Science, Camus, E.G. (1912). Cyperaceae. In: M.H. Lecomte Mahasarakham University for generous places to work (ed.), Flora of Générale de I’ Indo-Chine 1: and Ph.D. study of fi rst author, Prof. Dr David A. 157‒170. Messon et Cie Editeurs, Paris. Simpsonn and Dr Khanit Wangwasitt for the suggestions, Clarke, C.B. (1894). Cyperaceae. In: J.D. Hooker Mrs Nualanong Wichaikul for her assisted in a SEM (ed.), Flora of British India 7: 685‒694. London. examinations, the curators and staff s of the herbarium . (1908). New genera and species of for allowing access to herbarium materials, Asst Cyperaceae. Kew Bulletin of Miscellaneous Prof. Pasakorn Bunchalee, Asst Prof. Worachat Information. Additional ser 8: 58. Tokeaw, Mrs Wilailux Zumstein, Miss Nittiya Goetghebeur, P. (1986). Genera Cyperacearum. Chueawangkham, and Miss Primprapa Poosongsee Eenbijdrage tot de kennis van de morfologie, for their helping during the fi eld studies. systematiekenfylogenese van de Cyperacaeae- genera. Doctoral thesis, Rijksuniversiteit Gent. REFERENCES ______. (1998). Cyperaceae. In: K. Kubitzki (ed.), Bauters, K., Asselman, P., Simpson, D.A., Muasya, The families and genera of vascular plants, A.M., Goetghebeur, P. & Larridon, I. (2016). fl owering plants, monocotyledons 4: 141‒190. Phylogenetics, ancestral state reconstruction, Springer-Verlag, Berlin, Germany. and a new infrageneric classifi cation of Scleria Haines, R.W. & Lye, K.A. (1983). The sedges and (Cyperaceae) based on three DNA markers. rushes of east Africa. African Natural History Taxon 65 (3): 444‒466. Society, Nairobi.

SW 11611-p113-122-G8.indd 119 1/16/62 BE 4:32 PM 120 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 4. Leaf and culm anatomical characteristics of Scleria aureovillosa Kiaosanthie & K.Wangwasit. A‒C. leaf in transverse section: A. keel; B. leaf margin; C. lateral rib; D. adaxial leaf surface; E. culm surface; F. culm in transverse section. All from Kiaosanthie WK 0152012 (BKF).

SW 11611-p113-122-G8.indd 120 1/16/62 BE 4:32 PM A NEW SPECIES OF SCLERIA P.J.BERGIUS (CYPERACEAE) FROM NORTH-EASTERN THAILAND 121 (W. KIAOSANTHIE, W. CHAISONGKRAM & K. WANGWASIT)

Figure 5. Leaf and culm anatomical characteristics of Scleria benthamii C.B.Clarke. A‒C. leaf in transverse section: A. keel; B. leaf margin; C. lateral rib; D. adaxial leaf surface; E. culm surface; F. culm in transverse section. All from Kiaosanthie WK 0682013 (BKF).

SW 11611-p113-122-G8.indd 121 1/16/62 BE 4:32 PM 122 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

IUCN (2012). IUCN red list categories and criteria: Newman, M., Ketphanh, S., Svengsuksa, B., Thomas, version 3.1. second edition. Gland, Switzerland P., Sengdala, K., Lamxay, V. & Armstrong, K. and Cambridge, UK: IUCN. (2007). A Checklist of the Vascular Plants of Lao Johansen, D.A. (1940). Plant microtechnique. The PDR. Royal Botanic GardenEdinburgh,Edinburgh. Maple Company, U.S.A. Ridley, H.N. (1925). The Flora of the Malay Peninsula. Kern, J.H. (1961). The genus Scleria in Malaysia. V. 5: Monocotyledones Gymnospermeae general Flora Malesiana Precursores. Blumea 11: indices. L. Reeve & Co., LTD., London. 140–218. Simpson, D.A. & Koyama, T. (1998). Cyperaceae. In: ______. (1974). Cyperaceae. In: C.G.G.J. van T. Santisuk & K. Larsen (eds), Flora of Thailand. Steenis (ed.), Flora Malesiana ser. 1, 7 (3): 6 (4), pp. 426‒447. Diamond Printing, Bangkok. 435‒753. Thammathaworn, A. (1996). Handbookk for permanent Khoi, N.K. (2002). Cyperaceae. In: N.T. Ban (ed.), slide of plant tissue by paraffin method. Flora of Vietnam 3: 381‒414. Science & Technics Department of Biology, Faculty of Science, Publishing House, Ha Noi. Khon Kaen University, Thailand. Koyama, T. (1961). Classification of the family Zhang, S., Tucker, G.C. & Simpson, D.A. (2010). Cyperaceae 1. Journal of the Faculty of Science, Scleria. In: W. Zhengyi, P.H. Raven & H. Deyaun University of Tokyo 8: 34‒148. (eds), Flora of China 23: 260‒268. Science Press, Beijing & Missouri Botanical Garden Press, St Louis.

SW 11611-p113-122-G8.indd 122 1/16/62 BE 4:32 PM THAI FOREST BULL., BOT. 46(2): 123–128. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.02

Miscellaneous Cucurbit News V

WILLEM J.J.O. DE WILDE & BRIGITTA E.E. DUYFJES1

ABSTRACT Two recently described species are transferred from Zehneria to Pilogyne, namely Pilogyne subcoriacea (Y.D.Zhou & Q.F.Wang) W.J.de Wilde & Duyfjes and Pilogyne longifl orum (G.W.Hu & Q.F.Wang) W.J.de Wilde & Duyfjes. A new species from Thailand is described, Sinobaijiania frondosa W.J.de Wilde & Duyfjes. For Thladiantha angustisepala W.J.de Wilde & Duyfjes a range extension into China is noted.

KEYWORDS: Pilogyne, Sinobijiania, Thladiantha, Cucurbitaceae, South-East Asia, Kenya. Published online: 1 October 2018

INTRODUCTION (I) TWO NEW COMBINATIONS IN PILOGYNE FROM AFRICA This fi fth instalment of Miscellaneous Cucurbit News (for publication details of previous instalments Recent cooperation between China and Kenya see Thai Forest Bulletin (Botany) 39: 1. 2011) addresses for the preparation of a new Flora of Kenya has given the issue of the acceptance (or morphological validity) an impetus to botanical fi eld collecting in Kenya. of the species-rich genus Pilogyne Schrad. as separate Two new species of Cucurbitaceae, both endemics from Zehneria Endl. Similar controversies occur with from upland Kenya, were described in the genus the distinction of Mukia Arn. (against Cucumis L.) Zehneria. Both these species appear to belong in the and Gymnopetalum Arn. (against Trichosanthes L.), genus Pilogyne, and the new combinations are made largely due to modern molecular research. below. Furthermore, a new species of Sinobaijiania C.Jeff rey The rationale of accepting the genus Pilogyne & W.J.de Wilde is described, the specimen turning beside Zehneria (de Wilde & Duyfjes, 2009) has been up unexpectedly hidden as a ‘stowaway’ among controversial, and in the latest papers by De Boer plants collected for ecological research. Remarkably, et al. (2016) and Dwivedi et al. (2018), the merging its female fl owers and fruits remain unknown. Such of Pilogyne with the older Zehneria was urged, based plants are really curious, reminiscent of e.g. the on molecular evidence. However, in the Pacifi c area wide-spread cucurb species Siraitia siamensis a number of still poorly known species cannot be (Craib) C.Jeff rey ex S.Q.Zhong & D.Fang, occurring placed with certainty in either of the two available not rarely all over Thailand, but never seen fertile by genera, and most of the species concerned are suspect the authors in Thailand. Finally, the fi nd of a specimen of being of hybrid origin. These include Zehneria of Thladiantha angustisepala W.J.de Wilde & Duyfjes baueriana Endl., the type of the genus Zehneria. in southern China was a noteworthy extension of the Because this latter species diff ers considerably in known range of that species, thanks to our Russian detail from all other species casually assigned to and Chinese colleagues. Zehneria (see de Wilde & Duyfjes, 2009, fi g. 4), we prefer provisionally to keep the genus Zehneria as

1 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p123-128-G8.indd 123 1/16/62 BE 4:37 PM 124 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

monotypic, restricted to its type species, occurring isotypes EA, HIB), Kenya, Nandi County, S Nandy in lowland Norfolk Island and New Caledonia in the Forest, Kobujoi area. Pacifi c. Zehneria baueriana diff ers in its stout general Pilogyne longifl orum is remarkable by its long habit and in the stamens with short fi laments (fi lament receptacle tube. Its fruit is not yet known. The species, as long as or shorter than the anther), inserted near known from 6 collections in a restricted area of the middle of the receptacle tube. In Pilogyne, wide- Kenya, occurs at 1900–2000 m altitude. spread with many species in Africa and South-East Asia, the general facies of the plants is more delicate, with and fruits variable, but the stamens (2) A NEW SPECIES OF SINOBAIJIANIA always with long filaments, the filaments much FROM THAILAND longer than the anthers. This new species was discovered during iden- In the recent publication by Dwivedi et al. tifi cation of a collection of mainly sterile specimens (2018) concerning the phylogeny of Zehneria based made in 2013 during an ecological and fl oristic study on molecular data, it is urged to accept this genus in of a forest plot in North-Eastern Thailand. The a broad sense, because separating the genera as Sinobijiania collection appeared entangled in a proposed by de Wilde & Duyfjes (2006a) cannot be sterile leafy twig of an unidentifi ed treelet or , proved phylogenetically and the proposed morpho- and consisted of a fertile but leafl ess portion of a logical characters would be too weak. Dwivedi et al. branch. It appeared foliose because of unexpectedly (2018) further argue that this lumping together would large and dense male fl ower bracts, and possibly the be benefi cial for reaching taxonomic stability (in whole fl owering portion of the plant was destitute fl oras and fi eld guides) and avoid confusion among of leaves. Two years later, in the same ecological botanists trying to identify these plants in the fi eld plot, in about the same locality, a sterile apex of a and herbarium. In contrast, the present authors think leafy growing shoot was collected. Female elements, that the problems should not be evaded and that fl owers or fruit, are as yet not found. inevitably the considerable and obvious diff erences Apart from its possibly largely leafl ess habit in morphological traits, especially within the fl owers, when in , the present new Sinobaijiania is will remain a persistent source of error for botanists readily distinct within the genus, and also from most as long as these differences are not clarified and species of the closely related genus Thladiantha: defi ned at the genus level. Therefore, the recognition (1) in its compound male infl orescence, with 3–6 of several genera, including Pilogyne, possibly better bunches of congested fl owers dispersed along the defi ned morphologically, should be maintained. rachis, the fl owers each axillary to broad fl abelliform, toothed bracts; (2) in its fl owers in which the calyx is Pilogyne subcoriacea (Y.D.Zhou & Q.F.Wang) much longer than the corolla. For further information W.J.de Wilde & Duyfjes, comb. nov.— Basionym: compare de Wilde & Duyfjes (2008) and Lu & Zehneria subcoriacea Y.D.Zhou & Q.F.Wang, Jeff rey (2011). Phytotaxa 277(3): 282, f. 1, 2. 2016.— Type: Zhou & Mbuni 16/3 (holotype HIB; isotypes EA, PE), Kenya, Sinobaijiania frondosa W.J.de Wilde & Duyfjes, Mt Kenya, 00° 10’ 12.28’’ S; 37° 13’ 09.63’’ E. sp. nov. — Zehneria spec. A (see Jeff rey, 1967 and Agnew, Distinct within the genus Sinobaijiania in 1974, 2013). fl owers with calyx ca twice as long as corolla, not This species is known from several localities shorter than corolla.— Type: Visser, Chamchumroon, in Kenya, made at 2000–3000 m altitude. Saengrit & Suphuntee 372A, (holotype L!), Thailand, North-Eastern, , Phu Langka National Pilogyne longifl orum (G.W.Hu & Q.F.Wang) W.J.de Park, 17° 58’’ 77’ N; 104° 08’ 35’’ E., 28 June 2013, Wilde & Duyfjes, comb. nov.— Basionym: Zehneria Fig. 1. longifl orum G.W.Hu & Q.F.Wang, Phytotaxa Climber, possibly ca 5 m long, possibly tuberous, 324(1): 89, f. 1, 2. 2017.— Type: Sino-Africa Joint shoots 2–2.5 mm diam. (dry), when fl owering possibly Investigation Team (SAJIT) 006679 (holotype HIB; destitute of leaves. Plant sparsely hairy in most parts

SW 11611-p123-128-G8.indd 124 1/16/62 BE 4:37 PM MISCELLANEOUS CUCURBIT NEWS V (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 125

Figure 1. Sinobaijiania frondosa W.J.de Wilde & Duyfjes. A. portion of male fl owering twig, note large fl ower bracts; B detail of male infl orescence; C. male fl ower seen from outside; D. idem, one removed; E. idem (largely) and two removed; F. stamens; G. detail of inner surface of , the minute papillae apparently are oil glands (all: Visser et al. 372A). Drawn by Jan van Os.

SW 11611-p123-128-G8.indd 125 1/16/62 BE 4:37 PM 126 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

(early glabrescent and appearing as glabrous); pro- Note.— The papillose inner surface of the petals absent; tendrils ca 15 cm long, 2-branched and (fi g. 1G), apparently concerns the presence of oil spiralling below and above the point of branching at glands (Renner & Schaefer 2010). about one fifth from the apex. Leaves (on sterile shoots): petiole 1–2 cm long; lamina narrowly ovate, (3) THLADIANTHA ANGUSTISEPALA 8–10 × 3–4 cm, base cordate, apex acute, margin W.J.DE WILDE & DUYFJES, (remotely) dentate, teeth 1–2(–3) mm long, lower A NEW RECORD FOR CHINA surface sparsely whitish hairy. Male infl orescence compound, paniculate, in axils of fallen (reduced) (with Maxim S. Nuraliev et al. leaves, growing from the nodes together with a strong, – see Acknowledgements) well-developed tendril; panicle (5–)15–17 cm long The range-extension into China of Thladiantha including peduncle to 5 cm long with 2 or 3 scattered, angustisepala, hitherto known from Thailand, Laos, fi nely hairy linear bracts 3–5 mm long; the rachis and Northern Vietnam (Tonkin), became known from higher up bearing 5 or 6 scattered side-branches a collection by Nuraliev et al. G35, from Guangxi (0.5–)1–1.5 cm long, each ending in a condensed (China), made not far from the northernmost known raceme of 5–10 (or more) fl owers (one opening each collections in Vietnam of T. angustisepala, i.e. Cuc day), axillary to broadly wedge-shaped persistent Phuong National Park and Xuan Son National Park, imbricate bracts, 1.5–2.5 cm long and wide, irregu- ca 240 km NNE of the latter. The collection Nuraliev larly sharply incised or lobed at apex, bract-lobes et al. G35 slightly deviates in a less pronounced sharp, to 5 mm long. Male fl owers: pedicel slender, indumentum on all parts. The duplicate specimen 10–15 mm long, sparsely hairy, jointed at about the seen (in L) is a leafy young side-shoot from an older middle; fl owers fi nely hairy, part of pedicel above perennial ground-lying leafl ess shoot rooting at the joint 5–6 mm long, receptacle tube narrow, short- nodes. It bears only one single long-pedicelled campanulate, (1.5–)2 × 2–2.5 mm, details of thicken- fl ower, as is usually present at the base of a still to ing inside (disc) not known; calyx corolla-like, sepals be developed male raceme. free, more-or-less imbricate in bud, spreading, narrowly ovate, at apex narrowed into a long acute- In Gagnepain (Fl. Indo-Chine, 1921), acuminate apex, ca 11 × 4 mm, much longer than Keraudren (Fl. Cambodge, Laos & Vietnam, 1975), petals; corolla minutely hairy, in bud sub-globose, and Lu & Jeffrey (Fl. China, 2011) the name ca 5 mm diam.; petals imbricate, very shortly free Thladiantha angustisepala was not mentioned, as at apex, each 5–5.5 × 3 mm, faintly nerved, inside this latter species was only described in 2006 by de papillose; stamens in total ca 3 mm long, two in pairs Wilde & Duyfjes (2006b) and noted as not occurring and one solitary, fi laments ca 1.5 mm long, with hairs in China. However, on closer study, material of less than 0.1 mm long, anthers with scattered minute T. angustisepala most likely was included in the hairs, dorsifi xed, somewhat curved, ca 2.5 mm long; above mentioned fl oras under the name T. calcarata basal scales 2, each ca 0.8 mm long. Female fl owers, C.B.Clarke by Gagnepain (1921), or T. cordifolia fruits and seeds not known. (Blume) Cogn. (1881), the latter with T. calcarata as a synonym. In those fl oras Thladiantha cordifolia Thailand.— NORTH-EASTERN: Nakhon Phanom obviously was accepted in a wider sense as compared [Phu Langka National Park, 17° 58’’ 77’ N, 104° 08’ to the notion expressed in de Wilde & Duyfjes 35’’ E, 210 m, 28 June 2013, Visser et al. 372A (L)]. (2006b). Thladiantha angustisepala is quite distinct Distribution.— Endemic to Thailand, so far from T. cordifolia, as is evident from its description known only from the type locality. in e.g., the much narrower 1-veined sepals, and Ecology.— Dry dipterocarp forest on clay loam diff erent fruit, readily seen in the photographs and soil with much undergrowth, at ca 210 m altitude. fi gure then presented (plates 1, 2 and in fi g. 6). Phenology.— Flowering in June. The three species in the area with large fl abellate bracts in the male raceme and entire calyx lobes, Etymology.— The species epithet refers to viz. Thladiantha angustisepala, T. cordifolia, and frondose appearance of the large and densely packed T. tonkinensis Gagnep. superfi cially resemble each male fl ower bracts.

SW 11611-p123-128-G8.indd 126 1/16/62 BE 4:37 PM MISCELLANEOUS CUCURBIT NEWS V (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 127

other, especially when only one solitary male fl ower and Lu & Jeffrey (2011). The three species are is present. The latter, T. tonkinensis, was accom- distinct as shown in the following key: modated in T. cordifolia both by Keraudren (1975)

1. Male sepals linear, 1 mm wide or less, 1-veined. Fruit verrucose or ribbed 2. Male petals 15–25 mm long. Fruit ca 3 cm long, striate-verrucose T. tonkinensis 2. Male petals 15–20 mm long. Fruit 4–5 cm long, ± ribbed T. angustisepala 1. Male sepals linear-oblong, 2–3 mm wide, 3-veined. Fruit fenestrately sculptured T. cordifolia

Specimens of Thladiantha angustisepala REFERENCES studied: China, Guangxi Zhuang Autonomous Agnew, A.D.G. (1974). 1st ed. Upland Kenya wild Region, Baise City, Napo County, Baishen town, flowers: A flora of the ferns and herbaceous Nonglong village, foot of limestone hill, near village, fl owering plants of upland Kenya. 177. Oxford N 23° 14’ 25’’ E 105° 33’ 35’’, 1150 m, 22 Nov. University Press. 2016, Nuraliev et al. G35 (L!, MW: MW0754572). ______. (2013). Upland Kenya wild fl owers and Vietnam, Phu Tho Province, Thanh Son District, ferns: A fl ora of the fl owers, ferns, grasses and Xuan Son National Park, around Du Village, N 21° sedges of highland Kenya. East Africa Natural 07,877’ E 104° 56,533’, 361 m, 28 May 2015, History Society, Nairobi. Vislobokov 1,5013 (MW: MW0754573); same location, N 21° 07,998’ E 104° 55,880’, 277 m, 2 Cogniaux, C.A. (1881). Cucurbitaceae. In: A. & C. June 2015, Vislobokov 1,50166 (MW: MW0754574); de Candolle, Monographiae Phanerogamarum same location, N 21° 07,148’ E 104° 55,875’, 547 m, Prodomi 3: 325–951. Masson, Paris. 4 June 2015,Vislobokov 1,50288 (MW: MW0754575). De Boer, H.J., Cross, H.B., de Wilde, W.J.J.O., Duyfjes, B.E.E. & Gravendeel, B. (2016). Molecular phylogenetic analyses of Cucurbitaceae ACKNOWLEDGEMENTS tribe Benincaseae urge for merging of Pilogyne Two ecology students of Leiden University, with Zehneria. Phytotaxa 236(2): 173–183. Nina Soetens and Fenna Westveer, are thanked for De Wilde, W.J.J.O. & Duyfjes, B.E.E. (2006a). their keen eff orts to procure a leafy twig of the new Redefi nition of Zehneria and four new related Sinobaijinia. Jan van Os (Leiden) prepared the genera (Cucurbitaceae), with an enumeration of drawing. the Australasian and pacifi c species. Blumea 51: 1. The following are gratefully acknowledged ______. (2006b). The subtribe Thladianthinae for their contribution to the knowledge of Thladiantha: (Cucurbitaceae) in Indochina and . Maxim S. Nuraliev & Nikolay A. Vislobokov (both: Blumea 51: 493. Faculty of Biology, M.V. Lomonosov Moscow State ______. (2008). Cucurbitaceae. In: T. Santisuk & University, 1, 12, Leninskie Gory, 119234 Moscow, K. Larsen (eds), Flora of Thailand 9(4): 507. Russia & Joint Russian-Vietnamese Tropical Scientifi c Prachachon Co. Ltd., by Niran Hetrakul. and Technological Center, Cau Giay, Hanoi, Vietnam), ______. ( 2009). Miscellaneous Cucurbit news and Fang Wen, Bo Pan, Long-Fei Fu & Yi-Gang Wei III. Garden’s Bulletin Singapore 61(1): (all: Guangxi Key Laboratory of Plant Conservation 205–216. and Restoration Ecology in Karst Terrain, Guangxi Institute of Botany, Guangxi Zhuang Autonomous Dwivedi, A.M., Barfield, S., Pandey, A.K. & Region and Chinese Academy of Sciences, Guilin, Schaefer, H. (2018). Phylogeny of Zehneria 541006, China). (Cucurbitaceae) with special focus on Asia. Taxon 67(1): 55–65. The work of MSN and NAV was carried out in Gagnepain, F. (1921). Thladiantha. Flore Indo- accordance to Government order for the Lomonosov Chine 2: 1075. Masson et Cie., Paris. Moscow State University (project No. AAAA-A16- 116021660105-3).

SW 11611-p123-128-G8.indd 127 1/16/62 BE 4:38 PM 128 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Jeffrey, C. (1967). Cucurbitaceae. In: E. Milne- Lu, A.M. & Jeff rey, C. (2011). Cucurbitaceae. In: Redhead & R.M. Polhill (eds), Flora of Tropical W. Zhengyi & P. Raven (eds), Flora of China East Africa 17: 1. Crown Agents for Oversea 29. Missouri Botanical Garden Press, St. Louis. Governments and Administrations. Renner, S.S. & Schaefer, H. (2009). The evolution Keraudren-Aymonin, M. (1975). Thladiantha. In: and loss of oil-off ering fl owers: new insights A. Aubréville & J.-F. Leroy (eds), Flore du from dated phylogenies for angiosperms and Cambodge du Laos et du Viêtnam 15: 29. bees. Philos. Trans., Ser. B 365: 423–435. doi: Muséum National d’Histoire Naturelle, Paris. 10.1098/rstb.2009.0229.

SW 11611-p123-128-G8.indd 128 1/16/62 BE 4:38 PM THAI FOREST BULL., BOT. 46(2): 129–133. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.03

A new species of Brassaiopsis (Araliaceae) from Thailand, and lectotypifi cations of names for related taxa

HANS-JOACHIM ESSER1

ABSTRACT Brassaiopsis spinosissima Esser is described as a new species from Peninsular Thailand. It is compared with similar species from Thailand (Brassaiopsis ciliata Dunn and B. griffi thii C.B.Clarke) and Peninsular Malaysia (B. sumatrana Ridl.). They are primarily distinguished by diff erences in infl orescence size and position and in details of the indumentum. Two names sometimes confused with the new species, Panax palmatus Roxb. and Hedera polyacantha Wall., are lectotypifi ed and discussed in connection with (Buch.-Ham. ex D.Don) Seem. in order to distinguish them from the new species.

KEYWORDS: Hedera, Malay Peninsula, Panax, typifi cation Published online: 8 October 2018

INTRODUCTION TAXONOMY Brassaiopsis Decne. & Planch. is a small genus Brassaiopsis hainla (Buch.-Ham. ex D.Don) Seem., of Asian Araliaceae with ca 25 species distributed J. Bot. 2: 291. 1864.— Hedera hainla Buch.-Ham. from India and China to Java and . They are ex D.Don, Prodr. Fl. .: 187. 1825. Type: Nepal, characterized by spiny stems and branches, dendritic, Naramhetty, Buchanan Hamilton s.n. (holotype BM!). yellowish-brown to reddish-brown trichomes, fl owers — Hedera polyacantha Wall., Pl. Asiat. Rar. 2: 82, with five united sepals and five free petals, and t. 190. 1831.— Brassaiopsis polyacantha (Wall.) 2–3-locular ovaries and fruits. Most species bear R.N.Banerjee, Indian Forester 93: 341. 1967. Type: palmately-lobed leaves. As in many Araliaceae, leaf Nepal, 1821, Wallich Cat. 4907 A (Wallich s.n.) dimorphism may be present with young leaves quite (lectotype K-W!, designated here). diff erent from older ones. Species sometimes look superfi cially similar and there has been quite some — Panax palmatus Roxb. [Hort. Bengal.: 21. 1814, confusion in the past about their names. Brassaiopsis nom. nud.] Fl. Ind. ed. 1832, 2: 74. 1832. — hainla (Buch.-Ham. ex D.Don) Seem., as circumscribed Brassaiopsis palmata (Roxb.) Kurz, J. Asiat. Soc. here, can be distinguished by leaves that are lobed Bengal, Pt. 2, Nat. Hist. 39: 77, pl. 2. 1870, pro parte for less than half of their diameter (i.e., short lobes). as to type. Type: India, cultivated in Calcutta Botanic Among the species with deeply-lobed leaves (lobed Gardens and indigenous ‘in the moist valleys for more than half of their diameter), one species between the hills over the province of Chittagong’, from Peninsular Thailand is described as new here. Roxburgh s.n. (lectotype BM!, designated here). Notes.— 1. Wallich Cat. 4907, the type of The names Panax palmatus Roxb. [basionym Hedera polyacantha, is a mixture of two elements of Brassaiopsis palmata (Roxb.) Kurz] and Hedera (4907 A and B), both collected by Wallich himself, polyacantha Wall. [basionym of Brassaiopsis as can be seen in the East India Company Herbarium polyacantha (Wall.) R.N.Banerjee] have both been at Kew and as it was also cited in Wallich’s catalogue repeatedly misapplied. This confusion also concerns (Wallich, 1831–1832). Wallich Cat. 4907 A, collected this new species. It is therefore necessary to lectotypify in Nepal, is a fl owering specimen with shallowly these two earlier names and clarify their 5-lobed leaves undoubtedly referable to Brassaiopsis circumscriptions.

1 Botanische Staatssammlung München, Menzinger Strasse 67, 80638 München, Germany; email: [email protected]

© 2018 The Forest Herbarium

SW 11611-p129-133-G8.indd 129 1/16/62 BE 4:45 PM 130 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

hainla, whereas Wallich Cat. 4907 B, likewise from cited the correct synonymy, but erroneously included Nepal, is a sterile specimen with deeply lobed leaves the Malay Peninsula for the distribution off Brassaiopsis with seven very narrow, nearly glabrous lobes. In hainla. his Plantae Asiaticae Rariores, Wallich (1831) illustrated 4907 A in his plate, but with seven leaf Brassaiopsis spinosissima Esser, sp. nov. lobes; he cited no collection but indicated Nepal as the collecting locality. The presence of the quite This species is characterized by deeply-lobed diff erent parts of Wallich Cat. 4907 B explains why leaves with (7–)9 lobes scarcely constricted at base Hedera polyacantha Wall. with deeply divided, that are sparsely pubescent on both sides, with distinct 7-lobed leaves, based on Wallich Cat. 4907 B, was reticulate venation below, and by large infl orescences sometimes distinguished from H. polyacantha Wall. up to 60 cm in length. It is similar to two species ex G.Don with shallowly divided, 5-lobed leaves, with deeply-lobed leaves that are found in Northern referring to the illustration. Seemann (1864) cited Thailand, namely Brassaiopsis ciliata Dunn, which Hedera polyacantha as a synonym of Brassaiopsis is distinguished by its (7–)9–11-lobed leaves that hainla, based on Wallich Cat. 49077 (Nepal). Clarke have characteristic, setose ciliate trichomes above (1879), however, listed Hedera polyacantha as a and are usually constricted at base, and with infl ores- synonym of Brassaiopsis palmata sensu Kurz, refer- cences ca 15–30 cm long, and B. griffi thiii C.B.Clarke, ring only to Wallich’s plate. King (1898) discussed which has (5–)7–9-lobed leaves with the lobes never this confusion in some detail, and he cited only constricted at base and glabrous at least above if not Wallich Cat. 4907 B under Brassaiopsis palmata. on both sides, and with infl orescences ca 20 cm long. The name was not mentioned again until Indian Type: Thailand. Surat Thani, Phanom District, botanists (e.g. Balakrishnan, 1970, under Euaraliopsis Khlong Phanom National Park, ‘Giant Bamboo’ Hutch., a synonym of Brassaiopsis) re-discovered nature trail, 8°48’N, 98°44’E, 300 m, 28 Nov. 2005 it, followed by Philipson (1979). The leaf comprising (fl ), Gardner, Sidisunthorn & Tippayasri ST 1705a Wallich Cat. 4907 B is diffi cult to name. Because (holotype BKF!; isotype M!). Fig.1. fl owers and shallowly lobed leaves were illustrated — Brassaiopsis polyacantha auct. non (Wall.) by Wallich, Wallich Cat. 4907 A is chosen here as R.N.Banerjee: Gardner et al., Forest S. Thailand the lectotype, despite the fact that the number of 1: 231, pl. 323. 2015. lobes shown in the illustration diff ers. In this way Brassaiopsis polyacantha is unambiguously a Deciduous single-stemmed or sparsely branched synonym of B. hainla, as it had been treated by to 14 m, dbh to 30 cm, fl owering when in leaf but Seemann (1864) and Grierson (1991). often leafl ess when fruiting; bark pale grey to creamy, middle bark green, inner bark white; stem and 2. The type of Panax palmatus at BM is branches very spiny. Indumentum of pale-brownish undoubtedly referable to Brassaiopsis hainla, as can to yellowish-brown trichomes ca 0.2–0.3 mm long. also be seen in Icones Roxburghianae 2208 at Kew. Leaves: stipules ca 1.2 mm long; petiole 18–37 cm Seemann (1864) was correct when he placed Panax long, nearly glabrous; blade chartaceous, to ca palmatus in synonymy. However, Kurz (1870, 1877) 30 × 40 cm, palmately (7–)9-lobed, base rounded to applied Roxburgh’s basionym to plants of the decurrent, the basal 7–12 cm undivided, the lobes to Andaman Islands, which differ in having deeply 14–18 × 4–9 cm, longer than the undividedd basal part, lobed leaves. This was an error, and because he was elliptic and scarcely constrictedd at base, margin serrate, followed by most other authors for over a century, apex acuminate, paler below, both surfaces with the combination Brassaiopsis palmata was very scattered trichomes, veins of each lobe ca 7(–9) pairs, often misapplied to various species with deeply lobed complete venation including reticulate veinlets leaves (e.g., Ridley, 1922). As far as could be traced, distinctly visible below, indistinct above. Infl orescences Seemann’s correct synonymy was re-estabished only terminal, at least 30 × 50 cm on specimens (described recently by Grierson (1991). as 60 cm long by Gardner et al. 2015), sometimes 3. Another consequence of these typifi cations is with spines near base, with numerous (often 10–14) that Brassaiopsis hainla is now a Himalayan species, side branches, each subtendedd by a scaly, persistent just reaching Thailand but not Malesia. Esser (2004) bract ca 3 × 1.5 mm, main axis with scattered trichomes;

SW 11611-p129-133-G8.indd 130 1/16/62 BE 4:45 PM A NEW SPECIES OF BRASSAIOPSIS (ARALIACEAE) FROM THAILAND, 131 AND LECTOTYPIFICATIONS OF NAMES FOR RELATED TAXA (H.-J. ESSER)

Figure 1. Brassaiopsis spinosissima. A. habit; B. leaf; C. fl owering umbel; D. umbel in bud; E. fl ower in bud; F. fl ower bud, petals removed showing stamens; G. fl ower bud, petals and removed, disc and style; H. mature fl owers; I. umbel of immature fruits. After Gardner et al. ST 1705a (BKF).

SW 11611-p129-133-G8.indd 131 1/16/62 BE 4:45 PM 132 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

side branches to 25 cm long, with a larger terminal however has digitately compound leaves of a very umbel and numerous smaller lateral umbels each on diff erent shape. a peduncle of ca 5 cm; terminal umbel of each branch In Peninsular Malaysia, plants from Pahang 3 cm diam. [4(–5) cm in fruit], with 15–25 fl owers, and Kedah which are very similar have been named bracts ca 1 mm long. Flowers brownish-cream; as Brassaiopsis palmata or Brassaiopsis polyacantha densely tomentose in bud, soon glabrescent; pedicels by Ridley (1922), Stone (1977) and Philipson (1979). 9–12 mm long, with scattered trichomes; sepals 5, They appear to diff er mainly in minor characters such mostly united, 2 mm long, free apical lobes ca 0.5 mm as subentire leaf lobes, being in leaf when in fruit, long, glabrous; petals 5, greenish-white to dull cream, and are found on streambanks and along rivers. They free, 2.5 × 1 mm; stamens 5, not seen in mature may be conspecifi c with Brassaiopsis spinosissima fl owers; 2–3-locular; style short, undivided. but further studies are needed to clarify their status. Fruits grey-green when immature; pedicels 8–12 mm The only other Malesian species with a similar leaf long, with scattered tichomes; drupes subspherical, shape is Brassaiopsis sumatrana Ridl., from Sumatra not sulcate when dry, 7–8 × 7–8 mm, glabrous, style and the Malay Peninsula, which is however clearly 1.5 mm long. distinct by its much smaller infl orescences (less than Thailand.——PENINSULAR: Surat Thani [Phanom 20 cm long) which appear laterally (pseudolateral, District, Khlong Phanom National Park, ‘Giant overtopped by leaves) with a dense, reddish-brown Bamboo’ nature trail, 8°48’N, 98°44’E, 300 m, 20 indumentum, and leaves with very indistinct Mar. 2005 (fr), Gardner et al. ST 1705 (BKF, M); venation. same locality, 28 Nov. 2005 (fl ), Gardner et al. ST 1705a (BKF, M)]; Nakhon Si Thammarat [Lan Saka ACKNOWLEDGEMENTS District, National Park, SE side of Khao Luang Mountain, between Khiriwong village This study was initially supported by an summut, 8°28′N, 99°45′E, 650 m, 7 Mar. 2006 (fr), InstitutionalPostdoctoralgrantt (ERBCHBGCT930297) Gardner ST 2454 (BKF, M)]. of the Human Capital and Mobility Programme (Commission of the European Communities), Etymology.—The name refers to the particularly organized by John Parnell (TCD). I thank the staff of dense spines of the species. All species off Brassaiopsis the Forest Herbarium (BKF), Bangkok for support, have spiny stems and branches. In this species, and Simon Gardner for providing specimens. I am however, the spines are denser and more pronounced indebted to Mahsarahka Rungkrajang for the beautiful than in most other species, and notably even the illustration. basal part of the infl orescence is spiny. Distribution and Ecology.—Found at the edge of relatively undisturbed evergreen forest, on rugged REFERENCES limestone terrain or over granitic bedrock, between Balakrishnan, N.P. (1970). Nomenclatural notes on 300 and 650 m elevation. some flowering plants—II. Journal of the Bombay Natural History Society 67: 67–76. Notes.— Gardner et al. (2015) provide an excellent illustration under Brassaiopsis polyacantha. Clarke, C.B. (1879). Araliaceae. In: J.D. Hooker (ed.), The present author is partly responsible for their The Flora of British India 2: 720–740. Reeve & mis-identifi cation. Co., London. In Thailand, other species with a similar leaf Esser, H.-J. (2004 [‘2003’]). Brassaiopsis. In: D.G. shape are only known from as Frodin & R. Govaerts, World checklist and mentioned above, Brassaiopsis ciliata Dunn (only bibliography of Araliaceae: 98–105. The Royal known from Nan) and B. griffi thii C.B.Clarke (only Botanic Gardens, Kew. known from Chiang Mai). They diff er, among other Gardner, S., Sidisunthorn, P. & Chayamarit, K. characters, by their smaller infl orescences and the (2015). Forest trees of Southern Thailand, volume diff erent indumentum on their leaves. Large infl ores- 1 (A–Es). The Forest Herbarium, Bangkok & cences of comparable sizes occur in Thailand only The Royal Botanic Gardens, Kew, 749 pp. in (Blume) Regel, which

SW 11611-p129-133-G8.indd 132 1/16/62 BE 4:45 PM A NEW SPECIES OF BRASSAIOPSIS (ARALIACEAE) FROM THAILAND, 133 AND LECTOTYPIFICATIONS OF NAMES FOR RELATED TAXA (H.-J. ESSER)

Grierson, A.J.C. (1991). Araliaceae. In: A.J.C. Ridley, H.N. (1922). The Flora of the Malay Grierson & D.G. Long (eds), Flora of Peninsula 2. Reeve & Co., London, 672 pp. 2.1: 333–350. Royal Botanic Garden, Edinburgh. Seemann, B. (1864). Revision of the natural order King, G. (1898). Materials for a fl ora of the Malay Hederaceae. II: On the genera with a single style. Peninsula 10. Journal of the Asiatic Society Journal of Botany 2: 289–309. Bengal, Part 2, Natural History 67: 1–63. Stone, B.C. (1977). Notes on the systematy of Kurz, W.S. (1870). Report on the vegetation of the Malayan phanerogams: 25. Araliaceae. Garden Andaman Islands. Offi ce of the Superintendent Bulletin Singapore 30: 275–291. of Government Printing, Calcutta, 75 pp. Wallich, N. (1831). Plantae Asiaticae Rariores 2(8): ______. (1877). Forest Flora of British Burma 1. 53–86, t. 176–200. Treuttel and Würtz, London Office of the Superintendent of Government etc. Printing, Calcutta, 549 pp. ______. (1831–1832). A numerical list of dried Philipson, W.R. (1979). Araliaceae I. In: C.G.G.J. specimens: 172–211. Lithographed manuscript, van Steenis (ed.), Flora Malesiana, Series 1 Calcutta and London. Spermatophyta, 9: 1–105. Martinus Nijhoff Publishers, The Hague.

SW 11611-p129-133-G8.indd 133 1/16/62 BE 4:46 PM THAI FOREST BULL., BOT. 46(2): 134–137. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.04

Dendrobium chrysocrepis (Orchidaceae), a new record for Thailand

ORPORN PHUEAKKHLAI1, SOMRAN SUDDEE2, TREVOR R. HODKINSON3, HENRIK Æ. PEDERSEN4, PRIWAN SRISOM5 & SARAWOOD SUNGKAEW1,6,*

ABSTRACT In this paper we report the fi rst confi rmed fi nd of Dendrobium chrysocrepis in Thailand. A morphological description, line drawing and colour plate are presented (all based on Thai material), and some background information on the globally rare species is provided.

KEYWORDS: Dendrobium moschatum, Epidendroideae, northern Thailand. Published online: 11 October 2018

INTRODUCTION ex Hook.f. The original description was based on material from , provided by Charles Parish Dendrobium Sw., one of the largest orchid genera – a chaplain who made numerous plant collections (Wood, 2006), belongs to subtribe Dendrobiinae in in the wider surroundings of Mawlamyine from 1852 tribe Dendrobieae of the large subfamily Epidendroideae. and the following 25 years (Clayton, 2017). In contrast It contains about 1,450 species distributed among to many other orchids imported to Europe in the 28 sections (Pridgeon et al., 2014), 10 of which are 1800s, D. chrysocrepis has remained exceedingly represented in Thailand. Section Dendrobium is rare in cultivation (Wood, 2006: 489), and very few relatively large, comprising of approximately 100 observations of the species in the wild have been species distributed from India in the west to New made since its discovery. Recent records from Guinea and Australia in the south-east and to Japan Yunnan (Jin & Li, 2006 [sub syn. D. menglaensis and Korea in the north-east (Lavarack et al., 2000; X.H.Jin & H.Li]; Li et al., 2009; Xu et al., 2010: 321) Pridgeon et al., 2014). In the latest critical revision have demonstrated that D. chrysocrepis has a much of the Thai material, 53 species were accepted in wider distribution than previously assumed – and groups currently referred to sect. Dendrobium. Thus, thus raised the hope that the rare species might also Seidenfaden (1985) accepted 36 species in sect. occur in Thailand. Against this background, it was Dendrobium s.s., nine species in sect. Callista and gratifying, but not totally unexpected, when the fi rst eight in sect. Brevifl ores; the two latter infrageneric unequivocal fi nd of D. chrysocrepis in Thailand was taxa, however, are now included in sect. Dendrobium made during fi eld work targeting Dendrobium sect. (Pridgeon et al., 2014). Dendrobium in May 2017. One of the least known species in sect. Dendrobium is D. chrysocrepis C.S.P.Parish & Rchb.f.

1 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Chatuchak, Bangkok 10900, Thailand. 2 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 3 Department of Botany School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 4 Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5–7, DK-1350 Copenhagen K, Denmark. 5 146/1 Krungthep-Nonthaburi 42, Krungthep-Nonthaburi Rd. Wongsawang, Bang Sue, Bangkok 10800, Thailand. 6 Center for Advanced Studies in Tropical Natural Resources, Kasetsart University, Bangkhen, Bangkok 10900, Thailand. * Corresponding author: ff [email protected]

© 2018 The Forest Herbarium

SW 11611-p134-137-G8.indd 134 1/16/62 BE 4:51 PM DENDROBIUM CHRYSOCREPIS (ORCHIDACEAE), A NEW RECORD FOR THAILAND 135 (O. PHUEAKKHLAI, S. SUDDEE, T.R. HODKINSON, H.Æ. PEDERSEN, P. SRISOM & S. SUNGKAEW)

DESCRIPTION In the 1990s, the appearance of a few cultivated Dendrobium chrysocrepis C.S.P.Parish & Rchb.f. plants, presumably of wild-collected origin and in ex Hook.f., Bot. Mag. 98: t. 6007. 1872; Hook.f., some cases obtained from Thai dealers, was taken by Fl. Brit. India 5: 744. 1890; B. Grant, Orchids Burma: some orchidologists as evidence of D. chrysocrepis 77. 1895.— Callista chrysocrepis (C.S.P.Parish & occurring in Thailand. However, for each of the Rchb.f. ex Hook.f.) Kunte, Revis. Gen. Pl. 2: 654. plants concerned, Seidenfaden (1996, 1997), in spite 1891. Type: Mawlamyine, Myanmar, March 1871, of thorough investigations, was unable to verify Parish 309 (holotype K! [K000943862]). either the identifi cationn or a wild-collected Thai origin. Consequently, the record reported in this paper — D. menglaense X.H.Jin & H.Li, Ann. Bot. Fenn. represents the fi rst confi rmed fi nd of D. chrysocrepis 43: 296. 2006; G.G.Zhu, Z.H.Ji, J.J.Wood & in Thailand. The only known occurrence in Thailand H.P.Wood in Z.Y.Wu et al. (eds), Fl. China 25: 385. apparently comprises only ca 10 individuals, making 2009. Figs. 1–2. D. chrysocrepis an obvious candidate for the national Lithophytic herb. Stem subclavate, bilaterally Thai Red List. fl attened from a thin base, 10–28 cm long, ca 1 cm in diam., enclosed by persistent leaf sheaths. Leaves ACKNOWLEDGEMENTS lanceolate, up to 7 × 1.5 cm, apex unequally acuminate. produced on leafless stems, one- The first author would like to thank Bob flowered; floral bract ovate, ca 2 mm long, apex Harwood for helpful suggestions. Financial support acute. Flowers 2.5–3 cm long, yellow; pedicel and from the Thailand Research Fund through the Royal ovary ca 2 cm long. Sepals glabrous on both sides, Golden Jubilee Ph.D. Program (Grant No. PHD/ 5–6 veined; dorsal sepal obovate, rounded, 1.8–2 × 0215/2558) to student’s initials and advisor’s initials 0.8–1 cm, 5-veined; lateral sepals at base adnate to is acknowledged. the column, obliquely elliptic-oblong, acute to obtuse, 2.2–2.5 × 0.8–1 cm, 6-veined. Petals obliquely REFERENCES obovate, obtuse to rounded, 1.8–2 × 0.8–1 cm, Clayton, D. (2017). Charles Parish – plant hunter 7-veined. Labellum yellowish-orange, slipper- and botanical artist in Burma. The Ray Society, shaped, densely pubescent on the dorsal side, 1.8–2 × London & Royal Botanic Gardens, Kew. 1.2–1.5 cm, with incurved margin; disc rufously villous in a broad median band. Column yellow to Jin, X.-H. and Li, H. (2006). Coelogyne tsii and light green, ca 1.2 cm long, hairy at apex; mentum Dendrobium menglaensis (Orchidaceae), two short. new species from Yunnan, China. Annales Botanici Fennici 43: 295–297. Thailand.— NORTHERN: Chiang Mai [Angkhang, Lavarack, B., Harris, W. & Stocker, G. (2000). 19 May 2017, Srisom 51 (BKF!, spirit collection)]. Dendrobium and its relatives. Timber Press, Distribution.— China (Yunnan), Myanmar Portland, Oregon. Ecology.— In Thailand, plants are found in Li, L., Ye, D., Li, J. & Xing, F. (2009). A newly lower montane rain forest on a limestone hill growing recorded species and a new synonym of on mossy, humus-rich cliffs, sometimes together Orchidaceae from China. Journal of Tropical with Dendrobium dantaniense Guillaumin; 1700 m and Subtropical Botany 17: 295–297. alt. Flowering: recorded in May. Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Vernacular.— Ueang thung thong (เอื้องถุงทอง) Rasmussen, F.N. (2014). Genera Orchidacearum (here proposed). 6. Epidendroideae (part three). Oxford University Press, Oxford. Note.— Dendrobium chrysocrepis is similar to D. moschatum (Buch.-Ham.) Sw. but diff ers in Seidenfaden, G. (1985). Orchid Genera in Thailand having smaller, bright yellow, solitary fl owers on XII. Dendrobium Sw. Opera Botanica 83: 1–295. bilaterally fl attened stems. Seidenfaden, G. (1996). The reappearance of Phalaenopsis lowii (Orchidaceae) in Thailand. Nordic Journal of Botany 16: 283–286.

SW 11611-p134-137-G8.indd 135 1/16/62 BE 4:51 PM 136 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 1. Dendrobium chrysocrepis C.S.P.Parish & Rchb.f. ex Hook.f.: A. habit; B–C. fl ower (front view and top view); D. dorsal sepal; E. lateral sepals (inside and outside); F. petals (inside and outside); G–H. lip (upper surface and side view); J. lip and column; Drawn by O. Phueakkhlai.

SW 11611-p134-137-G8.indd 136 1/16/62 BE 4:51 PM DENDROBIUM CHRYSOCREPIS (ORCHIDACEAE), A NEW RECORD FOR THAILAND 137 (O. PHUEAKKHLAI, S. SUDDEE, T.R. HODKINSON, H.Æ. PEDERSEN, P. SRISOM & S. SUNGKAEW)

A B

C D E

Figure 2. Dendrobium chrysocrepis C.S.P. Parish & Rchb. f. ex Hook. f.: A–B. habit; C. fl ower (top view); D. lip (side view); E. lip (upper and lower surfaces); Photographed by P. Srisom (A–C) and O. Phueakkhlai (D–E).

Seidenfaden, G. (1997). Contributions to the orchid Xu, Z., Jiang, H., Ye, D. & Liu, E. (2010). The wild fl ora of Thailand XIII. Olsen & Olsen, Fredensborg. orchids of Yunnan [in Chinese]. Yunnan Publishing Wood, H.P. (2006). The Dendrobiums. A.R.G. Group Corporation / Yunnan Science & Technology Gantner Verlag K.G., Liechtenstein. Press, Kunming.

SW 11611-p134-137-G8.indd 137 1/16/62 BE 4:51 PM THAI FOREST BULL., BOT. 46(2): 138–150. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.05

The genus Campylotropis (Leguminosae) in Thailand

JIRATTHI SATTHAPHORN1, PEERAPAT ROONGSATTHAM1, PRANOM CHANTARANOTHAI2 & CHARAN LEERATIWONG1,*

ABSTRACT Seven taxa of Thai Campylotropis are recognized: C. bonii Schindl., C. capillipes (Franch.) Schindl. subsp. prainii (Collett & Hemsl.) Iokawa & H.Ohashi, C. decora (Kurz) Schindl., C. harmsii Schindl., C. parvifl ora (Kurz) Schindl., C. pinetorum (Kurz) Schindl. and C. sulcata Schindl. The reinstatement of C. parvifl ora is proposed here and C. cytisoides f. parvifl ora is reduced to a synonym of C. parvifl ora. A key to the species and descriptions with notes on their distribution, ecological data, vernacular names and photographs are also provided.

KEYWORDS: Campylotropis, , Leguminosae, Thailand. Published online: 29 October 2018

INTRODUCTION BM, C, CAL, CMU, CMUB, E, K, KKU, L, P, PSU, QBG, US and WAG (herbarium acronyms follow The genus Campylotropis Bunge (Desmodieae, Thiers, 2016) were examined. Most type specimens Leguminosae) comprises about 37 species distributed cited have been seen, indicated in the text by !, n.v. in temperate and tropical Asia (Iokawaa & Ohashi, 2002, (non vide) indicates when the type was not seen. 2008; Lewis et al., 2005; Huang et al., 2010). The genus was fi rst described by Bunge (1835) based on C. chinensis Bunge, now a synonym of C. macrocarpa TAXONOMIC TREATMENT (Bunge) Rehder. This genus is similar to genus CAMPYLOTROPIS Michx., but the character to distinguish Campylotropis from Lespedeza is the presence of Bunge, Uchen. Zap. Imp. Kazansk. Univ. 4: 157. one fl ower per bract while Lespedeza has two fl owers 1835; Miq., Fl. Ned. Ind. 1: 229. 1855; Schindl. in per bract (Iokawa & Ohashi, 2002). The name Fedde, Repert. Spec. Nov. Regni Veg. 11: 338, 424. ‘Campylotropis’ is derived from two Greek words, 1912; Rehder, Man. Cult. Trees & 1: 517. ‘campylo’ meaning curved and ‘tropis’ meaning keel 1927; Hutch., Gen. Fl. Pl. 1: 488. 1964; H.Ohashi, which is a distinct character of the genus (Lewis J. Jap. Bot. 49(2): 40. 1974; H.Ohashi et al. in Polhill et al., 2005). Craib (1928) listed seven species of & P.H.Raven, Advances Legume Syst. 1: 300. 1981; Campylotropis (as Lespedeza), and we recognize Thuan in Thuan et al., Fl. Cambodge, Laos & six of those (except C. henryi non Schindl. treated Vietnam 23: 143. 1987; P.Y.Fu, Bull. Bot. Res., as a synonym of C. decora (Kurz) Schindl.) in the Harbin 7(4): 22. 1987; P.Y.Fu, Fl. Reipubl. Popularis seven species treated here. Sin. 41: 92. 1995; Iokawa & H.Ohashi, J. Jap. Bot. 77: 191. 2002; X.F.Gao, in Inst. Bot. Kunming. Acad. Sin., Fl. Yunnan. 10: 550. 2006; P.Huang et al. in MATERIALS AND METHODS Z.Y.Wu et al., Fl. China 10: 292. 2010. Type species: Fresh fi eld collections in Thailand and specimens C. chinensis Bunge (= C. macrocarpa (Bunge) from the following herbaria: AAU, BCU, BK, BKF, Rehder).

1 Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand. 2 Department of Biology and Center of Excellence on Biodiversity (BDC), Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p138-150-G8.indd 138 1/16/62 BE 4:57 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 139 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

— Phlebosprium Jungh., Reisen Durch Java: 346. of pedicels, mostly narrowly triangular, persistent 1847; Hassk., Flora 30: 508. 1847. Type species: or caducous, outside hairy, inside glabrous; bracte- P. cytisoides Jungh. oles occuring at base of calyx, triangular, ovate- — Phlebosporium Benth. & Hook.f., Gen. Pl. 1: triangular or setaceous, persistent or caducous, 524. 1865. Type species: P. cytisoides Benth. outside hairy, inside glabrous. Calyx of 4 sepals, campanulate, outside hairy, inside glabrous; dorsal — Lespdeza sect. Campylotropis (Bunge) Benth., 1-lobed, entire or 2-toothed at apex; lateral 2-lobed, Hook’s J. Bot. Kew Gard. Misc. 4: 48. 1852; Taub. slightly shorter than dorsal one; lowest 1-lobed, in Engl. & Prant, Nat. Pfl anzenfam. 3(3): 332. 1894; longest. Corolla of 5 petals, light to dark violet, dark Nakai, Lespedeza of Japan & Korea: 3. 1927. Type blue or pinkish-white to creamy-white, glabrous; species: L. cytisoides Benth. standard variable in shape, mostly broadly obovate — Lespedeza subgen. Campylotropis (Bunge) or elliptic, clawed, auricle present or absent; wings Maxim, Trudy Imp. S.-Peterburgsk. Bot. Sada 2: mostly oblong, clawed, auricle 1; keel boat-shaped, 345, 347. 1873. Type species: L. cytisoides Benth. narrowly oblong, falcate, clawed, auricle 1, ventral sutures joined. Stamens 10, diadelphous (9+1), base — Lespedeza Michx. subgen. Oxyramphis (Wall. connate into staminal tubes about ¾ of its length; ex Meisn.) Baker in Hook.f., Fl. Brit. India 2: 143. anthers uniform, ovate-oblong or oblong, yellow. 1879.— Oxyramphis Wall., Numer. List nr. 5348. Pistils 1-carpellate; ovary superior, oblong, glabrous 1831, nom. nud. Type species: O. macrostyla to hairy, 1-locular, 1-ovuled; styles fi liform, incurved (D.Don) Wall. ex Meisn. at right angle about ¼ of its length; stigmas terminal, Shrubs, 1–4 m tall. Stems erect or ascending, capitate. Pods a legume, 1-articulated, indehiscent, much branched, terete or multi-angular, hairy. laterally fl attened, apex acuminate or rounded and Leaves pinnately 3-foliolate, alternate, subcoria- mucronate, obvious reticulate veins, hairy; fruiting ceous to chartaceous; petioles hairy; stipules trian- stalks elongated. Seeds 1, reniform, obliquely elliptic gular or triangular-lanceolate or triangular-linear, or oblong, brown, reddish-brown or blackish-brown; striate, persistent, outside hairy, inside glabrous; hilum with annulus aril. stipels absent. Inflorescences axillary raceme or A genus of about 37 species, widely distributed terminal panicle with reduced subtending leaves, in India to Indo-China and East Asia. Six species peduncles hairy. Flowers papilionaceous, 1-fl owered and 1 subspecies in Thailand. per subtending bract, fragile; bracts present at base

KEY TO THE SPECIES 1. Corolla pinkish-white to creamy-white; standard with green blotches on both sides; wings less than or equal to 6 mm long 2. Lower surface of leafl ets covered with ascending velutinous hairs; upper surface of leafl ets with black spots; pods less than or equal to 6 mm long 6. C pinetorum 2. Lower surface of leafl ets covered with appressed hairs; upper surface of leafl ets without black spot; pods more than 6 mm long 5. C. parvifl ora 1. Corolla dark blue or light to dark violet; standard without green blotches; wings more than 6 mm long 3. Upper surface of leafl ets pilose; stems with shortly zigzag lateral branches 4. C. harmsii 3. Upper surface of leafl ets glabrous; stems without shortly zigzag lateral branches 4. Peduncle covered with appressed hairs; pods glabrous or covered with appressed hairs 5. Standard with white and yellow blotches, 7.5–9 mm long and without auricles; peduncle sparsely cover of appressed-hairy; wings less than or equal to 8.5 mm long 1. C. bonii 5. Standard without white and yellow blotches, 10–11 mm long and with auricles; peduncle densely appressed-hairy; wings more than 8.5 mm long 2. C. capillipes subsp. prainii 4. Peduncle covered with patent hairs and glandular hairs; pods patent-hairy 6. Calyx lobes longer than 3 mm long, overlapping, dorsal lobe ovate-triangular; lower surface of leafl ets shortly appressed- hairy; pedicels more than 2 mm long 3. C. decora 6. Calyx lobes shorter or equal to 3 mm long, not overlapping, dorsal lobe triangular; lower surface of leafl ets appressed sericeous; pedicels less than or equal to 2 mm long 7. C. sulcata

SW 11611-p138-150-G8.indd 139 1/21/62 BE 2:56 PM 140 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1. Campylotropis bonii Schindl. in Engl., Bot. long. Seeds reniform to oblong, 6–8 × 3–3.5 mm, Jahrb. Syst. 54 (1): 64. 1916; Thuan in Thuan et al., brown. Fl. Cambodge, Laos & Vietnam 23: 143. 1987; Thailand.— PENINSULAR: Krabi [Sra Morakot Iokawa & H.Ohashi, J. Jap. Bot. 77(4): 201. 2002. Wildlife Sanctuary, 10 Apr. 2003, Middleton et al. — Lespedeza bonii (Schindl.) Gagnep. in Lecomte & 2100 (BKF); Tham Suea Temple, 5 Jan. 2006, Gagnep., Fl. Indo-Chine 2: 520. 1920.Type: Vietnam, Gardner & Sidisunthron ST2161 (QBG, L), ibid., Tuan-Du, Bon 4320 (holotype P P00758118!, iso- 18 Apr. 2007, Gardner ST2895 (BKF, QBG), ibid., types P P00758120!, P P00758119!). 24 Oct. 1991, Larsen et al. 42555 (BKF, P), ibid., — Campylotropis bonii var. anguticarpa Schindl. 8 May 2002, Pooma et al. 3609 (BKF), ibid., 10 in Fedde Repert. Nov. Regni Veg. 22: 271. 1926; May 2017, Satthaphorn & Leeratiwong 80 (PSU), Craib, Fl. Siam. 1(3): 432. 1928. Type: Thailand, ibid., 22 Aug. 2017, Satthaphorn & Leeratiwong 81 Phang Nga, small island near Panyi island, Herb. (PSU)]; Phang Nga [Khao Ping Kan, 25 Oct. 1974, Hort. Singapore no. 4079 (holotype BM!, isotype S.N. (BK); Small Island near Panyi Island, 17 Dec. K!). Fig. 1A–B. 1928, Herb. Hort. Singapore no. 4079 (BM, K)]; Phuket [28 Feb. 1994, Schmidt s.n. (P)]. Shrub, 1–3 m tall. Stems erect or ascending, terete, dark brown, glabrous or sparsely covered Distribution.— Vietnam. with appressed hairs. Leaves: petioles 8–17 mm long, Ecology.— Fissures on limestone outcrops, glabrous or sparsely appressed-hairy; rachis 2–5 mm alt. 150–330 m. Flowering: April to October. long; stipules triangular or triangular-lanceolate, Fruiting: April to October. 2–3 × 0.8–1 mm, outside appressed-hairy; leafl ets obcordate-obdeltoid, subcoriaceous; terminal leafl ets Vernacular.— Lueat nai tai (เลือดในใตต). 1.2–2.5 × 0.9–2 cm, lateral leafl ets 1–2 × 0.5–1.5 cm, Note.— Campylotropis bonii has a light to apex emarginate, base cuneate, margins entire; upper dark purple corolla with white and yellow blotches surface green, glabrous; lower surface light green, on the standard petal, sparsely appressed hairs on appressed-hairy; lateral veins 9–12 pairs; petiolules the peduncle, and distinctly obcordate-obdeltoid 2.5–4 mm long, densely appressed-hairy.Infl orescences leafl ets. axillary raceme, 2.5–3.5 cm long, laxly fl owered; peduncles 8–10 mm long, sparsely appressed-hairy; 2. Campylotropis capillipes (Franch.) Schindl. in bracts triangular, 0.5–1 × 0.2–0.5 mm, glabrous; Fedde, Repert. Spec. Nov. Regni Veg. 11: 341. 1912; bracteoles triangular, 0.7–1 × 0.3–0.5 mm, glabrous; Hand.-Mazz., Symb. Sin. 7: 573. 1933; C.Y.Wu et al., pedicels 7–11 mm long, sparsely appressed-hairy. Index Fl. Yunnan. 1: 573. 1984; P.Y.Fu, Bull. Bot. Calyx green, appressed-hairy; tube 1.8–2 mm long; Res., Harbin 7(4): 28. 1987; P.Y.Fu, Fl. Reipubl. dorsal lobe triangular, 1.5–2 × 1–1.5 mm; lateral lobes Popularis Sin. 41: 98. 1995.— Lespedeza capillipes narrowly triangular, 1.3–1.5 × 0.7–0.8 mm; lowest Franch., Pl. Delavay. 165. 1890; Iokawa & H.Ohashi, lobe triangular to narrowly triangular, 1.5–1.8 × J. Jap. Bot. 77(4): 206. 2002; X.F.Gao, in Inst. Bot. 0.6–1 mm. Corolla light to dark purple with white Kunming. Acad. Sin., Fl. Yunnan. 10: 557. 2006; and yellow blotches at base; standard elliptic-ovate, P.Huang et al. in Y.Z.Wu et al., Fl. China 10: 294. 7.5–9 × 5–7 mm, apex acute, claw 1.5–2 mm long, 2010. Type: China, Yunnan, monte Hee-chan-men, auricles absent; wings oblong, 8–8.5 × 2.5–3 mm, Delavay 2733 (lectotype K K03089370! selected slightly upward, apex rounded, claw 1.5–2.5 mm by Iokawa and Ohashi (2008), isolectotypes K long, auricle 0.5–0.6 mm long; keel 7.5–8 × 2.5–3 mm, K000894863!, P P00758122!). claw 1.5–2.5 mm long, auricle 0.2–0.3 mm long. Stamens 7–9 mm long, staminal tubes 6–7 mm long, subsp. prainii (Collett & Hemsl.) Iokawa & free part of fi laments 2–2.5 mm long; anthers oblong H.Ohashi, J. Jap. Bot. 77(4): 209. 2002; Iokawa & 0.3–0.4 × 0.2–0.3 mm. Pistils 9–9.5 mm long; ovary H.Ohashi, J. Jap. Bot. 83(1): 41. 2008; P.Huang 2–2.5 × 0.3–0.5 mm, appressed-hairy; styles 6–6.5 mm et al. in Y.Z.Wu et al., Fl. China 10: 295. 2010.— long, appressed-hairy at base. Pods obliquely oblong, Lespedeza prainii Collett & Hemsl., J. Linn. Soc., 11–15 × 4–5 mm, brown, apex roundedd and mucronate, Bot. 28: 46. 1890; H.Lév., Cat. Pl. Yun-Nan: 158. sparsely appressed-hairy; fruiting stalks 8–10 mm 1916.— Campylotropis prainii (Collett & Hemsl.)

SW 11611-p138-150-G8.indd 140 1/16/62 BE 4:57 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 141 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

Schindl. in Fedde, Repert. Spec. Nov. Regni Veg. (QBG); Doi Chiang Dao, 12 Nov. 1963, Adisai 648 11: 341. 1912; C.Y.Wu et al., Index Fl. Yunnan. 1: (BK), ibid., 4 Nov. 1963, Bunchuai 1305 (K, L), 576. 1984; P.Y.Fu, Bull. Bot. Res., Harbin 7(4): 28. ibid., 7 Dec. 1965, Hannipman 3278 (C, L), ibid., 1987; P.Y.Fu, Fl. Reipubl. Popularis Sin. 41: 98. 8 Feb. 1983, Koyama et al. T-33268 (BKF), ibid., 1995. Type: Myanmar (Burma), Shan, Collett 951 11 Nov. 1995, Maxwell 95-1174 (BKF, CMUB, L), (holotype K K000894915!). Fig. 1C–D. ibid., 30 Jan. 1996, Maxwell 96-136 (BKF, CMUB), ibid., 27 Sept. 1971, Murata et al. T-15201 (BKF, P), Shrub, 1–3 m tall. Stems erect, multi-angular, ibid., 28 Nov. 2005, Norsaengsri 276 (QBG), ibid., blackish-brown, densely appressed-hairy. Leaves: 23 Oct. 1992, Pooma 690 (BKF, CMUB), ibid., 11 petioles 5–33 mm, densely appressed-hairy; rachis Nov. 2016, Satthaphorn 766 (PSU), ibid., 12 Nov. 1–3 mm long; stipules triangular-lanceolate or 2016, Satthaphorn 777 (PSU), ibid., 27 Oct. 2017, setaceous, 4–10 × 1–2 mm, outside appressed-hairy; Satthaphorn 82 (PSU), ibid., 28 Oct. 2017, leafl ets obovate-oblong or obcordate, chartaceous; Satthaphorn 83 (PSU), ibid., 15 Jan. 2018, terminal leafl ets 0.8–2.8 × 0.5–1.5 cm, lateral leafl ets Satthaphorn & Leeratiwong 90 (PSU); ibid., 18 Dec. 0.5–2.2 × 0.5–2 cm, apex retuse or rounded and 2003, Sawai 492 (KKU), ibid., 19 Dec. 2003, Sawai mucronate, base cuneate or rounded, margins entire; 481 (KKU), ibid., 27 Oct. 1979, Shimizu et al. upper surface dark green, glabrous; lower surface T-210377 (BKF), ibid., 27 Oct. 1979, Shimizu et al. light green, densely appressed-hairy; lateral veins T-21090 (BKF), ibid., Dec. 1959, Smitinand & Abbe 7–12 pairs; petiolules 1–2 mm long, densely appressed- 6244 (K), ibid., 3 Dec. 1961, Smitinand & Anderson hairy. Infl orescences axillary raceme, 2.5–6 cm long, 7347 (BK, K), ibid., 10 Nov. 1962, Smitinand 7762 denselyy fl owered, fl owers usually pendulous; peduncles (BKF), ibid., 21 Nov. 1999, Suksathan 2170 (QBG), 1–2.6 cm long, appressed-hairy; bracts triangular, ibid., 9 Nov. 1997, Triboun 649 (BK); Doi Nom, 2 3–7 × 1–2 mm, appressed-hairy; bracteoles triangular, Nov. 1963, Phusomeseang 47 (K, L); Hui Nam Dang 1–2.5 × 1–2 mm, glabrous; pedicels 3–10 mm long, National Park, 17 Jan. 2002, Chayamarit et al. 3131 densely appressed-hairy. Calyx reddish-brown to (BKF)]; Uttaradit [Phu Soi Dao National Park, 8 red, appressed-hairy; tube 2–2.5 mm long; dorsal Feb. 2010, Norsaengsri 6484 (QBG)]. lobe triangular, 1.2–2.5 × 1–2.5 mm, with or without 2 teeth; lateral lobes triangular 1–2 × 0.5–1.2 mm; Distribution.— Myanmar, China. lowest lobe narrowly triangular, 1.5–2.2 × 0.5–1.2 mm. Ecology.— On exposed rugged limestone, Corolla dark blue to purple; standard obovate- lower montane pine oak forest, alt. 1300–2100 m. oblong, 10–11 × 7–8 mm, apex obtuse or emarginate, Flowering: September to November. Fruiting: claw 1.7–2 mm long, auricles 0.3–0.5 mm long, October to February. refl exed; wings oblong to oblong-obovate, 9.5–10 × 4.5–5.5 mm, apex rounded, claw 2.5–3.5 mm long, Vernacular.— Thua khao dok khram (ถั่วเขา auricle 0.5–1 mm long; keel 9.5–10 × 2.5–3 mm, ดอกคราม). claw 2.5–3.5 mm long, auricle 0.4–0.5 mm long. Note.— Campylotropis capillipes is similar to Stamens 9.5–11.5 mm long, staminal tubes 5.5–9 mm C. bonii in having appressed hairs on the peduncle long, free part of fi laments 1.5–2.5 mm long; anthers and purple corolla, but the former species is easily ovate-oblong, 0.4–0.6 × 0.3–0.4 mm. Pistils separated by having dense appressed hairs (vs. 10.5–13 mm long; ovary 5–7 × 0.6–1 mm, glabrous sparsely appressed hairs) on the peduncle and the or appressed-hairy; styles 5.5–8 mm long, glabrous absence of white and yellow blotches on the standard or appressed-hairy. Pods obliquely elliptic to oblong, petal. The subspecies is diff erent from the typical 10–14 × 3–7 mm, greenish-red to brownish-red, subspecies by the leafl ets being glabrous on the upper apex rounded and mucronate, glabrous or appressed- surface (vs. appressed hairs). hairy; fruiting stalks 5–11 mm long. Seeds reniform or obliquely elliptic, 1.5–2 × 0.7–1.2 mm, reddish-brown. 3. Campylotropis decora (Kurz) Schindl. in Fedde, Repert. Spec. Nov. Regni Veg. 11: 428. 1912; Iokawa Thailand.— NORTHERN: Chiang Mai [Ban Sa & H.Ohashi, J. Jap. Bot. 77(4): 213. 2002; Iokawa Ngin Nhua, 27 Nov. 2014, Pongamornkul 4453 & H.Ohashi, J. Jap. Bot. 83(1): 44. 2008; P.Huang (QBG), ibid., 27 Jan. 2016, Pongamornkul 5639 et al. in Y.Z.Wu et al., Fl. China 10: 295. 2010.

SW 11611-p138-150-G8.indd 141 1/16/62 BE 4:57 PM 142 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

— Lespedeza decora Kurz, J. Asiat. Soc. Bengal, lobes; lowest lobe triangular, 3.5–5 × 1.5–1.8 mm. Pt. 2, Nat. Hist. 42: 231. 1873; Kurz, Forest Fl. Corolla light purple; standard obovate, 9.2–14 × Burma 1: 381. 1877; Collett & Hemsl., J. Linn. 8–10 mm, apex acute or obtuse, claw 1–2 mm long, Soc., Bot. 28: 45. 1890; Prain in King, J. Asiat. Soc. auricle 0.1–0.3 mm long; wings oblong, 9–12 × Bengal, Pt. 2, Nat. Hist. 66: 377. 1897; Craib, Fl. 4–5 mm long, claw 2.4–3 mm long, auricle 0.4–1 mm Siam. 1(3): 433. 1928. Type: Myanmar (Burma), long, apex rounded; keel boat-shaped, 9–13 × Martaban,Kurz 1665 (holotype CAL 0000012094!). 3–3.8 mm, apex acute, claw 2.4–3.8 mm long, auricle — Lespedeza sericophylla Collett & Hemsl., J. Linn. 0.3–0.7 mm long. Stamens 10–16 mm long, staminal Soc., Bot. 28: 45. 1890; Prain in King, J. Asiat. Soc. tubes 7.5–12 mm long, free partt of fi laments 1.5–4 mm Bengal, Pt. 2, Nat. Hist. 66: 374. 1897.— long; anthers oblong, 0.5–0.6 × 0.2–0.3 mm. Pistils Campylotropis sericophylla (Collett & Hemsl.) 11–17 mm long, ovary 2.6–3.5 × 0.7–1 mm long, Schindl. in Fedde, Repert. Spec. Nov. Regni Veg. patent-hairy; style 7.4–14 mm long, glabrous. Pods 11: 428. 1912. Type: Myanmar (Burma), Shan hills, obliquely ovate, 5–10 × 4–5 mm, dark brown, apex Toungyi, Collett 13 (holotype K K000894914!). acuminate, densely patent-hairy and glandular-hairy; fruiting stalks up to 14 mm long. Seeds reniform, — Campylotropis sessilifolia Schindl. in Fedde, 1.5–4.6 × 1–3 mm, blackish-brown. Repert. Spec. Nov. Regni Veg. 11: 427. 1912. Type: Myanmar (Burma), South Shan hills, Toungyi, Thailand.— NORTHERN: Mae Hong Son [Doi Robertson 54 (holotype K K000894917!). Pha Daeng, 26 Dec. 2012, Norsaengsri 10025 (QBG); Huai San, 5 Feb. 2014, Norsaengsri 10815 — Lespedeza henryi sensu Gagnep. in Lecomte & (QBG)]; Chiang Mai [Doi Chiang Dao, 26 Mar. Gagnep., Fl. Indo-Chine 2: 520. 1920; Craib, Fl. 1991, Banziger 918 (CMU, L), ibid., 29 Feb. 1940, Siam. 1(3): 433. 1928, non Schindl. 1912.— Garret 1158 (E, K, L, P), ibid., 28 Nov. 1913, Kerr Campylotropis henryi sensu Thuan in Thuan et al., 2862 (K), ibid., 3 Mar. 1979, Koyama et al. s.n. Fl. Cambodge, Laos & Vietnam 23: 144. 1987, non (BKF), ibid., 28 Jan. 1996, Maxwell 96-116 (BKF, Schindler, 1912. Fig. 1E.–F. CMUB, L), ibid., 21 Dec. 1931, Put 4484 (E, K), Shrub, 0.8–2 m tall. Stems erect, multi-angular, ibid., 14 Jan. 2018, Satthaphorn & Leeratiwong 89 greenish-brownn to blackish-brown, densely appressed- (PSU), ibid., 15 Jan. 2018, Satthaphorn & Leeratiwong hairy. Leaves: petioles 0.2–3.5 cm long, sulcate with 91 (PSU), ibid., 27 Oct. 1979, Shimizu et al. T-21090 adaxial wings, appressed-hairy, angled; rachis (BKF), ibid., 17 Feb. 1958, Smitinand 4252 (L), 1–13 mm long; stipules triangular, 4–7 × 0.5–1.2 mm, ibid., 6 Dec. 1959, Smitinand 6243 (BKF), ibid., 16 outside appressed-hairy, persistent; leafl et obovate Feb. 1958, Sørensen et al. 1243 (C)]; Chiang Rai or obovate-oblong, subcoriaceous; terminal leafl ets [Doi Tung, 2 April 2011, Bult 1161 (BKF, CMUB), 0.6–6.2 × 0.4–3 cm, lateral leafl ets 0.5–5.7 × 0.4–3 cm, ibid., 21 Apr. 2008, Maxwell 08-95 (CMUB, QBG, apex emarginate, rounded or truncate and mucronate, L), ibid., 14 Feb. 2012, Norsaengsri & Tathana 8966 base rounded or slightly cordate, margins entire; (QBG), ibid., 28 Mar. 2012, Norsaengsri & Tathana upper surface green, glabrous; lower surface, light 9268 (QBG), ibid., 17 Feb. 1992, Pooma 649 (BKF, green, densely appressed-hairy; lateral veins 7–12 CMUB); , 16 Feb. 1993, Banziger 1079 pairs; petiolules 1–3 mm long, densely appressed- (CMUB, L)]; Tak [Doi Hua Mot, 22 Dec. 1993, hairy. Inflorescences axillary raceme or terminal Herb. trip. 778 (BCU), ibid., 14 Dec. 2003, Mattapha panicle, 2–10 cm long, densely fl owered; peduncle 508 (KKU), ibid., 14 Jan. 2004, Mattapha 573 0.2–3 cm long, patent-hairy and densely glandular- (KKU), ibid., 11 Feb. 1987, Paisooksantivatana hairy; bract triangular 1–4.5 × 0.5–1.2 mm, patent- y2049-87 (BK), ibid., 23 Jan. 2017, Satthaphorn 78 hairy; bracteoles narrowly triangular, 1–3 × (PSU); Kaeng Soi, 20 Nov. 1920, Kerr 4635 (BK, K)]. 0.1–0.5 mm, patent-hairy; pedicels 3–11 mm long, Distribution.— Myanmar, China, Laos. patent-hairy and densely glandular-hairy. Calyx Ecology.— In open, or on partly shaded rugged reddish-brown, patent-hairy and glandular-hairy; limestone, or deciduous forest, alt. 800–2250 m. tube 2.5–3.5 mm long; dorsal lobe ovate-triangular, Flowering: December to March. Fruiting: January 4–4.8 × 2–3.5 mm, bifi d or not; lateral lobes ovate- to April. triangular, 4–5 × 2–2.2 mm long, overlapping other

SW 11611-p138-150-G8.indd 142 1/16/62 BE 4:57 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 143 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

Vernacular.— Thua khao hin pun (ถั่วเขาหินปูน) auricle 0.5–1 mm long; keel 9–10 × 2.8–3 mm, claw Note.— Campylotropis decora is distinguished 2.5–3.5 mm long, auricle 0.3–0.5 mm long. Stamens by having densely patent hairs mixed with glandular 9.5–13 mm long, staminal tubes 7–11 mm long, free hairs throughout the peduncle, and ovate-triangular part of filaments 2–3 mm long; anthers oblong, overlapping calyx lobes which are distinctly longer 0.2–0.5 × 0.1–0.3 mm. Pistils 11–14 mm long; ovary than the calyx tube. From fi eld observations, specimens 3–3.5 × 0.8–1 mm, densely pilose; styles 7–10 mm at Doi Chiang Dao, Chiang Mai province, had thicker long, pilose. Pods ovate-oblong, 8–20 × 2.3–6 mm, leafl ets than other collections. brown, densely pilose, apex acuminate; fruiting stalks 7–13 mm. Seeds reniform, 4.5–8 × 3–3.5 mm, brown.

4. Campylotropis harmsii Schindl. in Fedde, Repert. Thailand.— NORTHERN: Chiang Mai [Doi Spec. Nov. Regni Veg. 11: 342. 1912; P.Y.Fu, Bull. Chiang Dao, 4 Dec. 2012, Chamchumroon et al. Bot. Res., Harbin 7(4): 28. 1987; P.Y.Fu, Fl. Reipubl. V.C. 1685 (BKF), ibid., 7 Jan. 1975, Geesink et al. Popularis Sin. 41: 97. 1995; Iokawa & H.Ohashi, J. 81377 (BKF, C, K, L, P), ibid., 6 Nov. 1922, Kerr Jap. Bot. 77(5): 255. 2002; X.F.Gao, in Inst. Bot. 6592 (BK, K), ibid., 1 Dec. 1984, Koyama et al. Kunming. Acad. Sin., Fl. Yunnan. 10: 556. 2006; T-39775, ibid., 28 Nov. 2005, Norsaengsri 274 Iokawa & H.Ohashi, J. Jap. Bot. 83(1): 44. 2008; (QBG), ibid., 30 Nov. 2005, Norsaengsri 277 P.Huang et al. in Z.Y.Wu et al., Fl. China 10: 296. (QBG), ibid., 19 Dec. 2014, Saisorn 347 (KKU), 2010.— Lespedeza harmsii (Schindl.) H.Lév., Cat. ibid., 14 Jan. 2018, Satthaphorn & Leeratiwong 88 Pl. Yun-Nan: 158. 1916; Craib, Fl. Siam. 1(3): 433. (PSU), ibid., 30 Nov. 2005, Sawai 985 (KKU)]; Nan 1928. Type: China, Yunnan, Szemao, Henry 9803D [Doi Phu Kha, 27 Feb. 1921, Kerr 4960 (BK, K)]. (lectotype K K000894861! designated by Iokawa & Distribution.— China. Ohashi (2008); isolectotype E E00025802!). Fig. 1G–H. Ecology.— In open, limestone ridge, evergreen forest, alt. 1200–2250 m. Flowering: December to Shrub, 0.5–4 m tall. Stems terete, shortly zigzag January. Fruiting: December to January. lateral branches, blackish-brown, patent-hairy. Leaves: petioles 4–14 mm long, densely appressed- Vernacular.— Thua khao bai khon (ถัวเขาใบขน่ ). hairy; rachis 1–9 mm long; stipules ovate-triangular, Note.— Campylotropis harmsii is different 4–7 × 1–2 mm, outside appressed-hairy; leaflets from other Thai Campylotropis species by having transversely elliptic, chartaceous; terminal leafl ets shortly zigzag lateral branches and pilose hairs on 0.8–3.5 × 0.5–2 cm, lateral leaflets 0.6–2.8 × both leaf surfaces. 0.4–1.5 cm, apex acute and mucronate, base cuneate, margins entire; upper surface dark green, pilose; lower 5. Campylotropis parvifl ora (Kurz) Schindl. in surface light green, densely pilose, midrib and lateral Fedde, Repert. Spec. Nov. Regni Veg. 11: 342. 1912; vein 5–8 pairs; petiolules 1.2–1.7 mm long, densely C.Y.Wu et al., Index Fl. Yunnan. 1: 575. 1984; appressed-hairy. Inflorescences axillary raceme, P.Y.Fu, Bull. Bot. Res., Harbin 7(4): 50. 1987; 2–8 cm long, laxly flowered; peduncles 3–9 mm Thuan in Thuan et al., Fl. Cambodge, Laos & long, densely patent-hairy and sparsely glandular- Vietnam 23: 146. 1987; P.Y.Fu, Fl. Reipubl. hairy; bracts ovate triangular, 1–2.2 × 0.3–1 mm, Popularis Sin. 41: 123. 1995; X.F.Gao, in Inst. Bot. patent-hairy; bracteoles ovate-triangular, 1–1.2 × Kunming. Acad. Sin., Fl. Yunnan. 10: 558. 2006.— 0.2–0.3 mm, appressed-hairy; pedicels 4–13 mm Lespedeza parvifl ora Kurz, J. Asiat. Soc. Bengal, long, densely patent-hairy. Calyx reddish-brown, Pt. 2, Nat. Hist. 42: 231. 1873; Kurz, Forest Fl. densely patent-hairy; tube 2–3 mm long; dorsal lobe Burma 1: 380. 1877; Collett & Hemsl., J. Linn. triangular, 3–3.5 × 3–4 mm; lateral lobes triangular, Soc., Bot.28: 45. 1891; Prain in King, J. Asiat. Soc. 4–4.2 × 1.2–1.5 mm; lowest lobe narrow triangular, Bengal, Pt. 2, Nat. Hist. 66: 377. 1897; H.Lév., Cat. 4.5–5 × 1–1.2 mm. Corolla pinkish-purple to white; Pl. Yun-Nan: 158. 1916; Gagnep. in Lecomte & standard elliptic-oblong, 10–11 × 7–7.5 mm, apex Gagnep., Fl. Indo-Chine 2: 522. 1920; Craib, Fl. acute, claw 1.2–2 mm long, auricle 0.2–0.3 mm long, Siam. 1(3): 433. 1928.— Campylotropis cytisoides infl exed; wings oblong, 10–11 × 5–5.5 mm, apex Miq. f. parvifl ora (Kurz) Iokawa & H.Ohashi, J. rounded, slightly upward, claw 2–2.5 mm long,

SW 11611-p138-150-G8.indd 143 1/16/62 BE 4:58 PM 144 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 1. A–B: Campylotropis bonii Schindl.; C–D: C. capillipes (Franch.) Schindl. subsp. prainii (Collett & Hemsl.) Iokawa & Ohashi; E–F: C. decora (Kurz) Schindl.; G–H: C. harmsii Schindl.

SW 11611-p138-150-G8.indd 144 1/16/62 BE 4:58 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 145 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

Jap. Bot. 77(4): 212. 2002; Iokawa & H.Ohashi, J. (QBG); Kiew Lom, 12 Dec. 2007, Tanaka et al. Jap. Bot. 83(1): 44. 2008, syn. nov. Type: Myanmar HN8196 (QBG); Mae Sariang, 8 July 1968, Larsen (Burma), Martaban, Mason 34 (holotype CAL et al. 2237 (K); Mountain pass between Pai and Pang 0000012096!). Fig. 2A–B. Mapha along road no. 1095, 13 Dec. 1998, Konta et al. 4470 (BKF); Muang Sroi Waterfall, 17 Jan. Shrub, 0.3–2 m tall. Stems erect, multi-angular, 1983, Koyama et al. T-32675 (QBG), ibid., 8 Feb. brown to dark brown, appressed-hairy. Leaves: 2013, Norsaengsri 10245 (QBG)]; Chiang Mai [12 petioles 0.4–5.5 cm long, sparsely appressed-hairy; Nov. 1911, Kerr 1587B (E, P); Ban Au Tam, 21 Jan. rachis 1–10 mm long; stipules triangular-linear, 2015, Pongkamornkul 4552 (QBG); Ban Gong Por 5–10 × 0.5–1 mm, outside appressed-hairy; leafl ets Nue, 26 Jan. 2016, Pongkamornkul 5590 (QBG), narrowly ovate to elliptic, subcoriaceous; terminal 27 Jan. 2016, Pongkamornkul 5649 (QBG); Ban leafl ets 1.3–5 × 0.5–1.6 cm, lateral leafl ets 0.9–3.8 × Huay Bon, 13 Jan. 1988, Paisooksantivatana et al. 0.3–1.6 cm, apex acute or rounded and mucronate, y 2217-88 (BK); Ban Mae Sa Ngin, 17 Nov. 2015, base rounded or cuneate, margins entire; upper Pongkamornkul 5155 (QBG); Ban Pak Tang Muzer, surface dark green, glabrous or slightly appressed- 28 Mar. 2015, Pongkamornkul 4628 (QBG); Ban hairy mainly on midrib; lower surface pale green to Pong Khrai, 9 Feb. 2006, Glamwaewwong 2253 grey, shortly appressed-hairy; lateral veins 7–12 (QBG); Doi Ang Khang, 30 Jan. 1999, Larsen & pairs; petiolules 1.5–2 mm long, densely appressed- Larsen KL47264 (AAU); Doi Chiang Dao, 20 Nov. hairy. Inflorescences axillary raceme or terminal 1990, Banziger 768 (CMU), ibid., 27 Feb. 2003, panicle, 8.5–11 cm long, densely fl owered; peduncles Chamchumroon & Sup V.C.1944 (BKF), ibid., 19 5.2–6 cm long, densely patent-hairy and sparsely Jan. 1991, Maxwell 91-80 (CMU, L), ibid., 27 Jan. glandular-hairy; bracts triangular, 1–1.2 × 1996, Maxwell 96-111 (BKF, CMUB), ibid., 29 Dec. 0.2–0.3 mm, appressed-hairy, reddish-brown, usually 1955, Pleunchit 1046 (BKF), ibid., 19 Dec. 1931, caducous; bracteoles setaceous, 0.5–1 mm long, Put 4398 (BCU, K), ibid., 14 Jan. 2018, Satthaphorn densely appressed-hairy; pedicels 1.5–4 mm long, & Leeratiwong 86 (PSU), ibid., 15 Jan. 2018, appressed-hairy and sparsely glandular-hairy. Calyx Satthaphorn & Leeratiwong 92 (PSU); Doi Inthanon green, densely appressed-hairy; tube 1–2 mm long; National Park, 11 Jan. 1994, Fukuoka & Koyama dorsal lobe narrowly triangular to setaceous, 1.5–2 × T-62083 (BKF), ibid., 3 Jan. 1975, Geesink et al. 0.5–0.6 mm, with or without 2 teeth; lateral lobes 8049 (K, L), ibid., 6 Feb. 1998, Konta & Phengklai narrowly triangularr to setaceous 1.5–1.8 × 0.3–0.4 mm; 4033 (BKF), ibid., 17 Dec. 1998, Konta et al. 4651 lowest lobe narrowly triangular to setaceous, 1.5–2 × (BKF), ibid., 3 Jan. 1975, Koyama et al. T-39663 0.5–0.6 mm. Corolla pinkish-white to creamy-white (BKF), ibid., Nov. 1968, Phengklai & Smitinand with green blotches on both surface of standard; 6047 (BKF), ibid., 26 July 1988, Phengklai et al. standard elliptic-oblong, 5.5–6 × 2.5–3 mm, apex 6981 (BKF), ibid., 17 Dec. 1965, Takawa et al. acute, claw 1–1.2 mm long, auricle absent; wings T2463 (BKF, P), ibid., 15 Apr. 1970, Worawoat 93 oblong, 5–6 × 2–2.2 mm, apex rounded, slightly (BKF); Doi Pha Hom Pok, 25 Feb. 1958, Sørensen upward, claw 1.4–1.5 mm, auricle 0.5–0.6 mm long; et al. 1630 (C); Doi Pui, 8 Jan. 1969, Nooteboom 636 keel 5.5–6 × 1.4–1.5 mm, claw 1–1.2 mm long, (BKF, K, L), ibid., 11 Jan. 1979, Paisooksantiwatana auricle 0.3–0.5 mm long. Stamens 6–7 mm long, 94-79 (BK, CMUB), ibid., 2 Mar. 1974, Sadakorn staminal tubes 4.5–5 mm long, free part of fi laments 314 (BK), ibid., 13 Jan. 2018, Satthaphorn & 2–3 mm long; anthers oblong, 0.4–0.5 × 0.3–0.4 mm. Leeratiwong 85 (PSU); Doi Sutep, 5 Mar. 1966, Pistils 6–7 mm long; ovary 2–3 × 0.5–0.7 mm, densely Chermsirivathana 4577 (BK), ibid., 10 Feb. 1926, appressed-hairy; styles 4–5 mm long, appressed-hairy Collins 1223 (BK, K, US), ibid., 1 Jan. 1905, at base. Pods obliquely obovate, 6.5–9 × 4–5 mm, Hosseus 202 (C, E, K, L, P), ibid., 25 Jan. 1910, dark brown, arranging mainly one side of rachis, Kerr 951 (K, L, P), ibid., 12 Nov. 1911, Kerr 1507 apex acuminate, patent-hairy; fruiting stalks 3–5 mm. (K), ibid., 22 Dec. 1920, Kerr s.n. (BK, P), ibid., Seeds reniform, 2–4.5 × 1.5–2.5 mm, brown. 26 Dec. 1987, Maxwell 87-1643 (CMU, L), ibid., Thailand.— NORTHERN: Mae Hong Son [Ban 10 Mar. 1988, Maxwell 88-321 (BKF, CMU, L), Huai Hi, 1 May 2014, Norsaengsri 11116 (QBG); ibid., 10 Jan. 1969, Nooteboom 710 (L), ibid., Doi Pui, 16 Dec. 2007, Tanaka et al. HN8480 Schultze & Pattanavibul 053/037-05 (WAG), ibid.,

SW 11611-p138-150-G8.indd 145 1/16/62 BE 4:58 PM 146 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

5 Mar. 1966, Sukku 70 (BKF), ibid., 8 Feb. 1958, Puudjaa 7062 (BKF); Thung Yai Naresuan East Sørensen et al. 876 (C, E), ibid., 15 Apr. 1958, Wildlife Sanctuary, 22 Dec. 2011, Watthana & Sørensen et al. 2762 (C, L), ibid., 16 Apr. 1958, La-ongsri 4095 (BKF)]; NORTH-EASTERN: Sørensen et al. 2775 (C), ibid., 19 Jan. 1967, Umpai Phetchabun [, 25 Dec. 1982, 349 (BK); Doi Kio Lom, 18 Jan. 2002, Chayamarit Koyama et al. T-31666 (BKF), ibid., Sawai 446 et al. 3140 (BKF); Doi Sahng Liang, 16 Dec. 1997, (KKU)]; Loei [Na Haew, Nhu Tong Kao, 25 Apr. Maxwell 97-1494 (BKF, CMUB); Huay Mae Mae, 1994, BGO staff s.n. (QBG); Phu Kradueng, 21 Mar. 24 Jan. 1996, Na Nakorn et al. 5657 (QBG); Khun 1958, Sørensen et al. 2383 (C, BKF); Phu Suan Sai, Wang Highland, 11 Jan. 1986, Paisooksirivathana 10 Dec. 1996, BGO Staff 74 (QBG); Phu Ruea y1746-866 (BK); Mae Chaem, 7 Dec. 1998, Maxwell National Park, 5 Dec. 2004, Sawai & Rob 832 98-1438 (CMUB, L); Mae Dad Noi, 25 Jan. 2010, (KKU)]; EASTERN: Chaiyaphum [Between Nam Norsaengsri & Intamusik 6345 (QBG); Mae Rim, Phrom and Thunkamang, 13 Dec. 1971, Beusekom 19 Dec. 2000, Glamwaewwong 25 (QBG), ibid., 18 et al. 4208 (BKF, C, K, L, P); Khonsan, Dec. 1994, Dec. 2001, Glamwaewwong 121 (QBG), ibid., 8 Wangwasit 64 (BK), ibid., Dec. 1994, Wangwasit Jan. 1983, Koyama et al. T-32097 (BKF), ibid., 9 65 (BK); Phu Khiao, 8 Nov. 1984, Murata et al. Jan. 1990, Maxwell 90-47 (CMU, E), ibid., 18 Jan. T-50138 (BKF)]; SOUTH-WESTERN: Kanchanaburi 1994, Na Nakorn et al. 406 (QBG), ibid., 21 Dec. [Ban Pilok, 19 Feb. 1967, Chermsirivathana 640 1995, Na Nakorn et al. 5511 (QBG), ibid., 20 Jan. (BK), ibid., 10 Dec. 1969, Chermsirivathana 1687 1996, Na Nakorn et al. 5559 (QBG), ibid., Jan. 1992, (BK); Huai Ban Kau, 11 Nov. 1971, Beusekom et al. Phengklai et al. 90077 (BKF), ibid., 21 Dec. 1978, 3670 (BKF, K, L)]. Pradit & Munpanid 539 (AAU), ibid., 22 Mar. 2017, Distribution.— Myanmar, China, Laos, Satthaphorn 79 (PSU); ibid., 26 Nov. 1951, Vietnam. Smitinand 1500 (BKF, P), ibid., 1 Jan. 1998, Srisanga & Puff 6 (QBG), ibid., 19 Jan. 2015, Sudjit et al. 3 Ecology.— In open, partly shaded areas on (QBG); Mae Sae, 27 Jan. 1977, S.N. (BKF); Mae limestone mountains and dry evergreen forest, alt. Sanam, 26 Dec. 1978, Niyomdham et al. 143 (BKF, 600–1800 m. Flowering: November to March. K, L); Mae Soi Valley, 4 Jan. 1991, Maxwell 91-22 Fruiting: November to March. (CMU, L); Mae Taeng, 3 Dec. 1977, Santisuk 1448 Vernacular.— Lueat nai (เลอื ดใน), hing men (หิ่งเมน ). (C, K); Mae Taman Reforest Unit, 28 Nov. 1984, Note.— According to Iokawa & Ohashi (2002), Koyama et al. T-39663 (BKF); Mae Wang, 18 Dec. Campylotropis cytisoides f. parvifl ora was established 1998, Konta et al. 4715 (BKF), ibid., 20 Feb. 2004, as distinct from the typical form, C. cytisoides f. Maxwell 04-91 (CMUB, L); Mae Ya Mae Kang, 29 cytisoides, by using the density of glandular hairs, Dec. 1913, Garret 103 (BK, C, E, K, L, P); Mai the length and the texture of leafl ets and the length Muang Nao Arboretum, 25 Dec. 2001,Sankamethawee of infl orescences. From specimen investigation of 384 (BKF, CMUB, L); Om Koi, 30 Nov. 2013, both forms, we have found additional characters to Pongkamornkul 3791 (QBG), ibid., 12 Jan. 1988, distinguish between the two forms. Campylotropis Santisuk 6664 (BKF); Pake Same Village, 28 Dec. cytisoides f. parvifl ora also diff ers from the typical 2000, Pongkamornkul 587 (QBG); Pha Dok Siao form by having a pinkish-white to creamy-white Waterfall, 25 Dec. 2015, Pisuttimarn 413-1 (KKU)]; corolla (vs. purplish-blue), the 1–2 mm long calyx Chiang Rai [Doi Luang National Park, 28 Oct. 1997, tube (vs. 2–3 mm), the 5.5–6 mm long standard (vs. Maxwell 97-1252 (CMUB, L); Khun Jae National 7.5–9 mm) and being sparsely short appressed-hairy Park, 1 Jan. 1998, Maxwell 98-12 (BKF, CMUB, on the lower surface of leafl ets (vs. densely long L); Khun Korn Waterfall, 12 Jan. 1997, KK 26 appressed-hairy). Therefore, we here reinstate this (BCU); Mae Fah Luang, 1 Feb. 2006, Maxwell 06-95 taxon to species level, C. parvifl ora. (CMUB, L); Mae Tameo Village, 2 Mar. 1989, Bragg 63 (CMU)]; Lamphun [Doi Khun Tan National Park, 27 Dec. 1993, Maxwell 93-1564 6. Campylotropis pinetorum (Kurz) Schindl. in (CMUB, L)]; Lampang [Jae Sawn National Park, 3 Fedde, Repert. Spec. Nov. Regni Veg. 11: 429. 1912; Dec. 1995, Maxwell 95-1256 (BKF, CMUB, L)]; Schindl. in Fedde, Repert. Spec. Nov. Regni Veg. Tak [Mae Ra Mard, 24 Dec. 2002, Niyomdham & 20: 285. 1924; Thuan in Thuan et al., Fl. Cambodge,

SW 11611-p138-150-G8.indd 146 1/16/62 BE 4:58 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 147 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

Laos & Vietnam 23: 147. 1987; Iokawa & H.Ohashi, Thailand.— NORTHERN: Chiang Mai [Doi Ang J. Jap. Bot. 77(5): 277. 2002; X.F.Gao, in Inst. Bot. Khang, 2 Feb. 2007, Srisanga et al. 2956 (CMUB, Kunming. Acad. Sin., Fl. Yunnan. 10: 555. 2006; QBG); Doi Inthanon National Park, 14 Feb. 2001, P.Huang et al. in Z.Y.Wu et al., Fl. China 10: 298. Chayamarit et al. 23977 (BKF), ibid., 12 Feb. 1998, 2010.— Lespedeza pinetorum Kurz, J. Asiat. Soc. Konta et al. 4245 (BKF), ibid., 18 Feb. 1998, Konta Bengal, Pt. 2, Nat. Hist. 42: 230. 1873; Kurz, et al. 4371 (BKF), ibid., 20 Dec. 1998, Konta et al. Forest Fl. Brit. Burma 1: 381. 1877; Prain in King, 4915 (BKF), ibid., 31 Dec. 1989, Maxwell 89-1614 J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 375. (CMU, L), ibid., 18 Feb. 1998, Niyomdham 5322 1897; Gagnep. in Lecomte & Gagnep., Fl. Indo- (BKF), ibid., 11 Feb. 1998, Phengklai et al. 11002 Chine 2: 524. 1920; Craib, Fl. Siam. 1(3): 433. (BKF), ibid., 16 Mar. 2004, Pornpongrungrueng 1928. Type: Myanmar (Burma), Martaban, Kurz 459 (KKU), ibid., 13 Jan. 2018, Satthaphorn & 16377 (holotype CAL 0000012101!). Fig. 2C–D. Leeratiwong 84 (PSU), ibid., 7 Jan. 1998, Srisanga et al. 94 (AAU, BKF, QBG); Doi Mon Chong, 1 Shrub, 0.4–1.5 m tall. Stems erect, multi-angular, Jan. 1997, %*26WDৼ (QBG); Doi Pui, Mar. brown, densely yellowish-velutinous. Leaves: petioles 1997, &KD\DPDULW 3KDWKDQDFKDURHQ (BKF), 1.8–2.8 cm long, velutinous; rachis 3–6 mm long or ibid., Mar. 1997, &KD\DPDULW 3KDWKDQDFKDURHQ indistinct; stipules triangular, 7–9 × 2.5–4 mm, 669 (BKF), ibid., 8 Feb. 1979, Pradit 681 (C), ibid., RXWVLGHYHOXWLQRXVSHUVLVWHQWOHDÀHWVQDUURZO\HOOLSWLF 24 Feb. 1975, Sadakorn 513 (BK), ibid., 5 Feb. 2015, VXEFRULDFHRXVWHUPLQDOOHDÀHWV±î±FP Srisanga et al. 3870 (QBG); Doi Sutep, &RFNHUHOO ODWHUDOOHDÀHWV±î±FPDSH[DFXWH s.n. (US), ibid., 8 Jan. 1911, Kerr 1645 (BK, C, K, base cuneate, margins entire; upper surface light green, L, P), ibid., 23 Jan. 1988, Maxwell 88-677 (AAU, ¿QHO\DSSUHVVHGKDLU\HVSHFLDOO\RQYHLQVORZHU BKF, CMU, L), ibid., 10 Jan. 1969, Nooteboom 711 surface pale green, densely ascending velutinous, (BKF, C, K, L), ibid., 9 Feb. 1958, Sørensen et al. PDWXUHOHDÀHWVZLWKVFDWWHUHGEODFNVSRWVODWHUDO 891 (C, BKF), ibid., 8 Jan. 1959, Sørensen et al. veins 7–12 pairs; petiolules 1.5–2 mm long, velutinous. 6589 (C, E), ibid., 19 Jan. 1967, Umpai 361 (BK); ,QÀRUHVFHQFHVD[LOODU\UDFHPH±FPORQJGHQVHO\ Kawng Sang, 22 Jan. 1964, Hansen et al. 10877 (C, ÀRZHUHGSHGXQFOHV±FPORQJYHOXWLQRXV and K, L)]; Chiang Rai [Doi Chang, 17 Feb 1968, sparsely glandular-hairy; bracts triangular, 5–8 × Hansen & Smitinand 12620 (AAU, BKF, C, E, K, 2–3 mm, patent-hairy; bracteoles setaceous, ca 1 mm L, P)]; Phayao [Doi Luang National Park, 10 Feb. long, patent-hairy, early caducous; pedicels 1.5–2 mm 2016, Muangyen 702 (QBG)]; Nan [Khun Sathan long, patent-hairy. Calyx reddish-brown, patent- National Park, 27 Jan. 2013, Norsaeangsri 10110 hairy; tube 2–2.5 mm long; dorsal lobes triangular, (QBG)]; Uttaradit [Phu Miang-Phu Thong Wildlife 2.5–3 × 1.8–2 mm, with or without 2 long teeth, Sanctuary, 27 Jan. 2011, 5RP.ODR%RWDQLFDO*DUGHQ lateral lobes triangular, 2.5–3 × 1–1.1 mm, lowest 0159/2554 (QBG); Phu Soi Dao National Park, 8 lobe narrowly triangular, 2.8–3 × 0.7–1 mm. Corolla Feb. 2010, Norsaeangsri 6482 (QBG)]; Phitsanulok pinkish-white to creamy-white; standard obovate- [Phu Soi Dao National Park, 9 Feb. 2000, Suksathan REORQJ±î±PPDSH[URXQGHGFODZ 2313 (QBG)]; Kamphaeng Phet [Mae Wong 1.5–2 mm long, auricle absent; wings spathulate, National Park, 3 Jan. 1999, Bult 207 (BKF, QBG)]; ±î±PPDSH[URXQGHGFODZ±PP NORTH-EASTERN: Loei [Phu Kradueng National long, auricle 0.8–1 mm long; keel 6–6.5 × 2.5–3 mm, Park, 10 Jan. 1960, Abbe & Smitinand 9427 (BKF), claw 2.8–3 mm long, auricle 0.3–0.5 mm long. ibid., 10 Feb. 1931, Kerr 20063 (K, L); Phu Luang Stamens 8–8.5 mm long, staminal tubes 4–6 mm Wildlife Sanctuary, 27 Nov. 1987, Dee 995 (BKF); ORQJIUHHSDUWRI¿ODPHQWV±PPORQJDQWKHUV Phu Ruea National Park, 16 Jan. 1967, Smitinand oblong, 0.3–0.4 × 0.1–0.2 mm. Pistils 8.5–9 mm 10153 (BKF), ibid., 13 Dec. 1966, Umpai 333 (BK)]. long; ovary 1.5–2 × 0.7–0.8 mm, densely patent- hairy; styles 6.5–7 mm long, densely patent-hairy. Distribution.— Myanmar, Laos, Vietnam. PodsREOLTXHO\RYDWH±î±PPEURZQDSH[ Ecology.— In open, grassy areas, hill evergreen rounded and mucronate, densely patent-hairy; fruit- forest, lower montane forest, alt. 1000–2250 m. ing stalks 2–3 mm long. Seeds oblong or reniform, Flowering: November to March. Fruiting: January swollen, 2.6–2.8 × 1.3–1.5 mm, reddish-brown. to March.

SW 11611-p138-150-G8.indd 147 1/21/62 BE 3:20 PM 148 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Vernacular.— Thua khao (ถั่วเขา), thua doi sunken, 7–12 pairs; petiolules 1–3 mm long, densely (ถั่วดอย), thua pa (ถั่วปา), ka sam pik (กาสามปก). sericeous. Infl orescences axillary raceme or terminal panicle, 2.5–16 cm long, densely fl owered; peduncles Note.— Campylotropis pinetorum resembles 1.2–2 cm long, densely patent-hairy and sparsely C. parvifl ora in having a pinkish-white to creamy- glandular-hairy; bracts triangular, 3.5–4.5 × white corolla. However, the former has black spots 0.4–0.5 mm, appressed-hairy, reddish-brown usually on the upper leaf surface, velutinous hairs on the caducous before anthesis; bracteoles triangular, lower leaf surface and pods less than or equal to 0.5–1 × 0.2–0.3 mm, appressed-hairy; pedicels 6 mm long, while the latter has no black spots on 1–2 mm long, densely patent-hairy, young fl owers the upper leaf surface, appressed hairs on the lower subsessile. Calyx reddish-brown, appressed-hairy; leaf surface and pods more than 6 mm long. tube 1.5–1.8 mm long; dorsal lobe triangular, 2.2–3 × 1.8–2 mm, with or without 2 teeth; lateral lobes 7. Campylotropis sulcata Schindl. in Engl., Bot. triangular, 2–2.2 × 0.8–1 mm, lowest lobe triangular Jahrb. Syst. 54 (1): 65. 1916; C.Y.Wu et al., Index 2.2–3 × 0.7–1 mm. Corolla light to dark purple; Fl. Yunnan. 1: 576. 1984; P.Y.Fu, Bull. Bot. Res., standard elliptic, 8.5–11 × 5–6.5 mm, apex acute, Harbin 7(4): 51. 1987; P.Y.Fu, Fl. Reipubl. Popularis claw 0.8–1 mm long, auricle absent; wings oblong, Sin. 41: 126. 1995; Iokawa & H.Ohashi, J. Jap. Bot. 8–9.5 × 3–4 mm, slightly falcate, apex rounded, claw 77(6): 326. 2002; X.F.Gao, in Inst. Bot. Kunming. 1.5–2 mm long, auricle 0.5–1 mm long; keel 9.5–11 × Acad. Sin., Fl. Yunnan. 10: 554. 2006; Iokawa & 2.5–3 mm, claw 1.5–2.5 mm long, auricle 0.5–1 mm H.Ohashi, J. Jap. Bot. 83(1): 54. 2008; P.Huang long. Stamens 8–10 mm long, staminal tubes 6–8 mm et al. in Z.Y.Wu et al., Fl. China 10: 300. 2010.— long, free part of fi laments 2.5–3 mm long, upward; Lespedeza sulcata (Schindl.) Craib. Fl. Siam. 1(3): anthers ovate-oblong, 0.3–0.4 × 0.2–0.3 mm. Pistils 434. 1928. Type: China, Yunnan, Bon s.n. (holotype 8–12 mm long; ovary 2–2.5 × 0.4–0.5 mm, densely P P03089818!). patent-hairy; styles 6.5–9.5 mm long, densely patent- — Campylotropis rockii Schindl. in Fedde, Repert. hairy. Pods obliquely ovate, 4–7 × 2–4 mm, dark Spec. Nov. Regni Veg. 22: 270. 1926; C.Y.Wu et al., brown, apex acuminate, densely short patent-hairy, Index Fl. Yunnan. 1: 576. 1984; P.Y.Fu, Bull. Bot. calyx and style persistent; fruiting stalks 1–5 mm Res., Harbin 7(4): 29. 1987; P.Y.Fu, Fl. Reipubl. long. Seeds reniform, 1.2–1.5 × 0.5–1 mm, blackish- Popularis Sin. 41: 100. 1995; X.F.Gao, in Inst. Bot. brown. Kunming. Acad. Sin., Fl. Yunnan. 10: 553. 2006. Thailand.— NORTHERN: Mae Hong Son [Doi Type: China, Yunnan, Simao, Ganlanba as west of Laem Rang, 27 Dec. 2012, Norsaengsri 10061 Mekong, Salween water divide, upper Kan-lan-chai, (QBG); Nong Khao Klang (Karen) Village, 29 Oct. Rock 7059 (holotype US, n.v.). 2007, Maxwell 07-680 (QBG)]; Chiang Mai [Doi — Campylotropis purpurascens Ricker in Rehder Chiang Dao, 12 Nov. 2011, Clark 235 (QBG), ibid., et al., J. Wash. Acad. Sci. 36: 39. 1946. Type: China, 2 Nov. 1922, Kerr 6509 (BK, E, K), ibid., 4 Nov. Yunnan, Hila, Shunning, Yü 17571 (holotype A, n.v.; 1995, Maxwell 95-1067 (BKF, CMUB, L), ibid., isotypes E E00025717!, US 00288564!). Fig. 2E–F. 27 Jan. 1996, Maxwell 96-101 (BKF, CMUB, L), ibid., 19 Dec. 2015, Pisuttimarn 410-1 (KKU), ibid., Shrub, 1–2 m tall. Stems erect, multi-angular, 17 Oct. 1994, Pooma 873 (BKF, CMUB), ibid., 11 dark brown, densely patent-hairy. Leaves: petioles Nov 2016, Satthaphorn 75 (PSU), ibid., 14 Jan. 0.4–2.9 cm long, sulcate, densely sericeous; rachis 2018, Satthaphorn & Leeratiwong 877 (PSU), ibid., 1–6 mm long; stipules triangular-linear, 8–10 × 17 Dec. 2003, Sawai 508 (KKU), ibid., 11 Dec. 0.6–1 mm, outside appressed-hairy, persistent; leaf- 1987, Smitinand s.n. (BKF), ibid., 7 Nov. 1997, lets elliptic-oblong or obovate-elliptic, subcoriaceous; Triboun 698 (BK)]; Chiang Rai [Khun Jae National terminal leafl ets 2–6.5 × 0.4–2.8 cm, lateral leafl ets Park, 23 Nov. 1997, Maxwell 97-1416 (BKF, 1.5–4.5 × 0.3–2.3 cm, apex acute or retuse and CMUB, L), ibid., 1 Jan. 1998, Maxwell 98-4 (BKF, mucronate, base cuneate, margins entire; upper CMUB, L)]; Phitsanulok [Phu Hin Rong Kla surface dark green, appressed-hairy; lower surface National Park, 1 Oct. 1990, Chantaranothai et al. light green, densely sericeous, midrib adaxially 90/515 (KKU), ibid., 9 Oct. 1987, Sridith 53 (BCU),

SW 11611-p138-150-G8.indd 148 1/16/62 BE 4:58 PM THE GENUS CAMPYLOTROPIS (LEGUMINOSAE) IN THAILAND 149 (J. SATTHAPHORN, P. ROONGSATTHAM, P. CHANTARANOTHAI & C. LEERATIWONG)

ibid., 30 Oct. 1987, Sridith 76 (BCU), ibid., 29 Oct. Note.— Campylotropis sulcata is distinct in 2001, Watthana & Suksathan 1543 (AAU, CMUB, having a multi-angular stem, patent hairs on the QBG)]; EASTERN: Chaiyaphum [Phu Khiao, 8 Nov. peduncle and light to dark purple petals. The species 1984, Murata et al. T-41784 (BKF)]. may resemble C. decora in some aspects, e.g. multi- Distribution.— China. angular stems, obovate-elliptic leaflets and the peduncle with patent hairs, however it diff ers from Ecology.— In open, rugged limestone, mixed the latter in having sericeous hairs on the lower deciduous forest, scrub forest, alt. 850–2000 m. surface of leaflets (vs. appressed hairs), without Flowering and fruiting: October to January. Fruiting: glandular hairs on the peduncle (vs. with patent hairs October to January. mixed with glandular hairs), bracts which are more Vernacular.— Thua doi khon yao (ถั่วดอยขนยาว). than 2 mm long (vs. less than or equal to 2 mm long) and standard petal without auricles.

Figure 2. A–B: Campylotropis parvifl ora (Kurz) Schindl.; C–D: C. pinetorum (Kurz) Schindl.; E–F: C. sulcata Schindl.

SW 11611-p138-150-G8.indd 149 1/16/62 BE 4:59 PM 150 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

ACKNOWLEDGEMENTS Iokawa, Y. & Ohashi, H. (2002). A taxonomic study of Campylotropis (Leguminosae) I. Journal of The authors are grateful to the curators and Japanese Botany 77(4): 179–222. staff s of the herbaria cited. We would like to thank Prof. Dr Henrik Balslev and Asst Prof. Dr Sahut ______. (2008). Campylotropis (Leguminosae) Chantanaorrapint for their valuable helps. This of China, an Enumeration and Distribution. research was financially supported by Carlsberg Journal of Japanese Botany 83: 36–59. Foundation, Science Achievement Scholarship of Huang, P., Ohashi, H. & Iowaka, Y. (2010). Thailand, Graduate School, Dissertation Funding Camyplotropis. In: Z.Y. Wu, P.H. Raven & D.Y. for Thesis, Prince of Songkla University and Center Hong (eds), Flora of China 10: 292–302. of Excellence on Biodiversity (BDC-PG3-160013). Lewis, G.P., Schrire, B., Mackinder, B. & Lock, M. (2005). Legumes of the World. The Bath Press, REFERENCES UK, 434 pp. Bunge, A.A. (1835). Plantarum mongholico-chinen- Thiers, B. (2016, continuously updated). Index sium decas prima. Uchenya zapiski Imperatorskogo Herbariorum: A global directory of public herbaria Kazanskogo Universiteta. Kazan 4(2): 154–180. and associated staff . New York Botanic Garden’s Craib, W.G. (1928). Florae Siamensis enumertio. Virtual Herbarium. http://sweetgum.nybg.org./ Vol 1. part 3. The Siam Society, Bangkok. ih/. November 25, 2017.

SW 11611-p138-150-G8.indd 150 1/16/62 BE 4:58 PM THAI FOREST BULL., BOT. 46(2): 151–154. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.06

Chelonopsis thailandica, a new species and new record of Chelonopsis (Lamiaceae) from Thailand

ALAN J. PATON1,*, SOMRAN SUDDEE2 & BHANUBONG BONGCHEEWIN3

ABSTRACT Chelonopsis thailandica A.J.Paton, Suddee & Bongch. is described and illustrated here. The new species belongs in Chelonopsis Sect. Aequidens being a small shrub with an equally lobed calyx, lanceolate leaves and eglandular hairs. The new species diff ers from the other species in the section in having a remotely toothed leaf margin and denser indumentum. This is the fi rst species of Chelonopsis to be recorded from Thailand. Only one specimen has been seen and it is possible that the habitat where the collection was made in Chiang Mai Province no longer exists.

KEYWORDS: Chiang Mai, endemic, Gomphostemmateae, Mae Rim district, new species. Published online: 1 November 2018

INTRODUCTION (2008, 2013) are similar to that of Li & Hedge (1994) in Flora of China, the main diff erence relevant to The genus Chelonopsis Miq. is mainly found the description of the new species is that C. albifl ora in south-west China and Japan with one species Pax & K.Hoff m. recognized by Li & Hedge (1994), found in Kashmir (Xiang et al., 2008), in addition is placed in synonymy of C. souliei by Xiang et al. to the new species from Thailand described here. (2008). The genus is placed in the Gomphostemmateae along with Gomphostemma Wall. ex Benth. with another The new species described here was discovered genus, Bostrychanthera Benth., having been merged from studying a specimen on two sheets in the into Chelonopsis (Xiang et al., 2013). Xiang et al. herbarium of Queen Sirikit Botanic Garden. The (2008, 2013) divided the genus into two subgenera: specimen was collected on a road in the Pong Yaeng subgen. Chelonopsis are herbs and have an unequally Subdistrict between Samoeng and Mae Rim Districts lobed calyx, whereas subgen. Aequidens C.Y.Wu & in 1996, which was upgraded to a tarmac road in H.W.Li are shrubs with equally lobed calyces. Wu around 2004. Despite several targeted fi eld work & Li (1965, 1977) and Xiang et al. (2013) further attempts to fi nd more specimens of this plant by the divided subgen. Aequidens into two sections: sect. authors and staff of QBG, and searches in major Microphyllum C.Y.Wu & H.W.Li with ovate leaves European and Thai herbaria in preparation of the and clavate glandular hairs, and sect. Aequidens Lamiaceae account for Flora of Thailand by the C.Y.Wu & H.W.Li has lanceolate leaves and lacks authors, no further specimen has been found. It is clavate glandular hairs, which is similar to the new possible that this plant is now extinct due to destruction species described here. In all, Xiang et al. (2013) of its habitat. The specimen is incomplete and only recognized 14 species, but only two, Chelonopsis has one open corolla. Despite the lack of fruiting souliei (Bonati) Merr. and C. forrestii J.Anthony material and limited fl owering material, this specimen belong in sect. Aequidens. The accounts of Xiang et al. clearly belongs in Chelonopsis having a dilated

1 Science Directorate, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K. 2 Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Chatuchak, Bangkok 10900, Thailand. 3 Department of Pharmaceutical Botany, Faculty of Pharmacy, Mahidol University, Bangkok 10400, Thailand. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p151-154-G8.indd 151 1/16/62 BE 5:02 PM 152 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

corolla tube with short lobes, campanulate calyces opposed to 1–2.5 m tall. The habitat of C. thailandica and few-flowered cymes in the axils of leaf-like is slightly diff erent from other species in the section bracts. It shares the characters of sect. Aequidens by as they are usually found in hot dry river valleys having eglandular hairs and lanceolate leaves, but rather than dry dipterocarp woodland. Unfortunately, diff ers from related Chinese species, C. forrestii and the type specimen of C. thailandica only has one C. souliei as described below. open fl ower and it was not possible to dissect this.

DESCRIPTION ACKNOWLEDGEMENTS Chleonopsis thailandica A.J.Paton, Suddee & Thanks are due to the staff of QBG for assistance Bongch. sp. nov. Type: Thailand, Chiang Mai, Mae during fi eld work and herbarium visits. We would Rim to Samoeng, 25 Sept. 1996, Nanakorn et al. like to thank Dr Prachaya Srisanga of QBG for 7634 (holotype QBG!-2 sheets). Figs. 1 & 2. preparing the high-resolution image for Figs. 1 & 2. Shrub to 0.4 m tall. Stems quadrangular, white We would also like to thank the staff of Aarhus pubescent with simple hairs. Leaves opposite, University Herbarium and the Carlsberg Foundation petiolate, lanceolate, 40–60 × 10–20 mm, apex for hosting the authors of this paper and fi nancial obtuse to acute, base rounded to broadly cuneate, support to the authors to assist with the compilation margin with few remote serrate teeth, upper leaves of the Lamiaceae account for the Flora of Thailand. entire or repand, white pubescent; petioles 5–15 mm long. Inflorescence terminal, lax with 1-flowered REFERENCES pendulous cymes in the axils of leaf-like bracts; Li, H.W. & Hedge, I.C. (1994). Lamiaceae. In: C.Y. pedicels ca 1 mm long. Calyx campanulate, 5-lobed, Wu & P.H. Raven (eds), Flora of China 17: weakly-2 lipped with posterior lobes slightly longer; 135–139. Science Press, Beijing, and Missouri lobes narrowly lanceolate. Corolla white, 12–15 mm Botanical Press, St. Louis. long; tube dilating from base; posterior lip 1-lobed, Wu, C.Y. & Li, X.W. (1965). Materiae ad fl oram very short; anterior lip 3-lobed, 3–4 mm long. Nutlets labiatarum sinensium (1). Acta Phytotaxonomica not seen. Sinica 10: 150–154. Thailand.— NORTHERN: Chiang Mai ______. (1977). Labiatae. In: C.Y. Wu (ed.), Flora Distribution.— Endemic to Thailand, only Reipublicae Popularis Sinica 65(2). Science known from the type locality. Press, Beijing. Ecology.— Open dry deciduous dipterocarp Xiang, C.L., Liu, E.D. & Peng, H. (2008). A key to forest. the genus Chelonopsis (Lamiaceae) and two new Phenology.— Flowering: September. combinations: C. rosea var. siccanea and C. souliei var. cashmerica comb. nov. Nordic Conservation status.— Likely to be Critically Journal of Botany 26(1–2): 31–34. https://doi. Endangered or possibly Extinct. org/10.1111/j.0107-055X.2008.00209.x Note.— This new species diff ers from both Xiang, C.L., Zhang, Q., Scheen, A.C., Cantino, P.D., species of Chelonopsis sect. Aequidens by having a Funamoto, T. & Peng, H. (2013). Molecular leaf margin with serrations irregular and distant in the phylogenetics of Chelonopsis (Lamiaceae: lower leaves and repand in upper leaves, rather than Gomphostemmateae) as inferred from nuclear regularly serrate, and a whitish densely pubescent and plastid DNA and morphology. Taxon 62(2): indumentum, rather than being pubescent to almost 375–386. https://doi.org/10.12705/622.11 glabrous. The new species further differs from C. souliei by having strictly opposite leaves, rather than in whorls of three, and one- rather than three- flowered cymes. Additional differences from C. forrestiii include having a white rather than yellow corolla, and in being a smaller plant, ca 0.4 m tall as

SW 11611-p151-154-G8.indd 152 1/16/62 BE 5:02 PM CHELONOPSIS THAILANDICA, A NEW SPECIES AND NEW RECORD OF CHELONOPSIS (LAMIACEAE) FROM THAILAND 153 (A.J. PATON, S. SUDDEE, B. BONGCHEEWIN)

Figure 1. Chelonopsis thailandica: Sheet I (from Nannakorn 7634 (QBG)). Photos by P. Srisanga.

SW 11611-p151-154-G8.indd 153 1/16/62 BE 5:02 PM 154 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 2. Chelonopsis thailandica: Sheet II (from Nannakorn 7634 (QBG)). Photos by P. Srisanga.

SW 11611-p151-154-G8.indd 154 1/16/62 BE 5:02 PM THAI FOREST BULL., BOT. 46(2): 155–161. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.07

A revision of the genus Tapeinidium (Lindsaeaceae) in Thailand

NUTDANAI PUTTHISAWONG1 & SAHUT CHANTANAORRAPINT1,*

ABSTRACT A revision of the genus Tapeinidium (C.Presl) C.Chr. in Thailand is presented based on herbarium specimens and fi eld surveys. Two species are recognized, namely Tapeinidium luzonicum (Hook.) K.U.Kramer and T. pinnatum (Cav.) C.Chr. A key to species, descriptions, photographs and conservation assessments are provided. In addition, issues with the previous lectotypifi cations of Davallia luzonicum Hook. and D. pinnata Cav. are resolved.

KEYWORDS: conservation assessment, Davallia, fern, lectotypifi cation, pteridophyte. Published online: 22 December 2018

INTRODUCTION T. luzonicum from Khao Luang (Nakhon Si Thammarat) and T. pinnatum from Khao Luang With about 18 currently accepted species, (Nakhon Si Thammarat), Ban Mae Lao (Yala) and Tapeinidium (C.Presl) C.Chr. is the third largest Ko Chang (Trat). Although the genus concept re- genus of the family Lindsaeaceae (Lehtonen et al., mains unchanged from that of Tagawa & Iwatsuki 2010; PPG I, 2016). Although it is primarily a (1985), the realization that many more specimens Malesian genus its members occur from southern have been collected since the publication of the Flora India to Samoa and as far north as the Ryukyu Islands of Thailand account (now also from Chanthaburi, of Japan but are absent from northern India, the Krabi, Phatthalung, Phangnga and Trang) prompted northern parts of mainland South-East Asia (including us to reassess whether the earlier species concepts northern Thailand), China and Australia (Holttum also hold. Our study confi rmed that there are only 1955 [‘1954’]; Kramer, 1967, 1971). The genus was two Tapeinidium species in Thailand, T. luzonicum fi rst described as a subgenus of Microlepia C.Presl and T. pinnatum. But, more importantly, it resulted (Presl, 1851 [‘1849’]) and was subsequently raised in an improved key to these species, more detailed to the generic rank by Christensen (1906), based on descriptions, conservation assessments and the reso- Tapeinidium pinnatum (Cav.) C.Chr. lution of some typifi cation issues. The earliest report of Tapeinidium in Thailand was by Holttum (1955 [‘1954’]) who reported that MATERIALS AND METHODS T. pinnatum “occurs...throughout Malaysia [but] northwards of Malaya it extends only into Lower This study was largely based on fresh specimens Siam”. More than a decade later, Tagawa & Iwatsuki from fi eld surveys and herbarium specimens housed (1967) reported Tapeinidium biserratum (Blume) in BCU, BK, BKF, PSU and QBG. Most morphological Alderw. from Khao Luang, Nakhon Si Thammarat and anatomical characters were examined using province but this was quickly redetermined as stereo and light microscopes. Mature spores were T. luzonicum (Hook.) K.U.Kramer by Kramer (1968 also examined with an electron microscope. [‘1967’]). Tagawa & Iwatsuki (1985) subsequently Ecological and geographical data were compiled produced an account of Tapeinidium for the Flora from field observations, specimen labels and of Thailand that recognized two species based on a publications. handful of specimens from only three localities:

1 Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p155-161-G8.indd 155 1/16/62 BE 5:07 PM 156 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

TAXONOMIC TREATMENT pinnate to tripinnate, coriaceous (or subcoriaceous), Tapeinidium (C.Presl) C.Chr., Index Filic. 631. bearing sparse hairs on lower surface; hairs transparent, 1906; Holttum, Rev. Fl. Malaya ed. 1, 2: 338. 1955 uniseriate, 2–3 cells long. Rachises with U or V shape [‘1954’]; Kramer, Blumea 15(2): 545. 1968 [‘1967’]; vascular bundle in cross section. Veins free, pinnately Kramer in Van Steenis & Holttum (eds), Fl. Males., or dichotomously branched, not reaching the margin Ser. 2, Pterid. 1(3): 184. 1971; Tagawa & Iwatsuki of the lamina. Sori submarginal, terminal on veins, in Smitinand & Larsen (eds), Fl. Thailand 3(2): 145. uninerval or very rarely binerval (exceptionally 1985; Kramer in Kramer & Green (eds), Fam. & trinerval, but not in Thai species), paraphyses present, Gen. Vasc. Pl. 1: 91. 1990; Lethonen et al., Bot. J. uniseriate, 5–11 cells long. Indusia firm, pouch- Linn. Soc. 163: 337. 2010; Shiyong in Zhengyi & shaped, attached at the base and at least the greater Raven, Fl. China. 2−3: 141. 2013.— Microlepia part of the sides, opening towards the margin of the subg. Tapeinidium C.Presl, Epimel. Bot.: 968. 1851 lamina but not reaching it. Spores bilateral, [‘1849’].— Protolindsaya Copel., Philipp. J. Sci., monolete. C. 5: 283. 1910. Type species: Tapeinidium pinnatum A tropical Asian and Pacifi c genus of ca 18 (Cav.) C.Chr. species distributed from southern India to Samoa Plants usually terrestrial, sometimes lithophytic. and as far north as the Ryukyu Islands of Japan but Rhizome very short to moderately long creeping, absent from northern India, the northern parts of solenostelic with an internal sclerifi ed pith, scaly. mainland SE Asia (including northern Thailand), Scales non-clathrate, basally attached, narrowly China and Australia (Holttum 1955 [‘1954’]; triangular, apex long attenuate, margin minutely to Kramer, 1967, 1971); Two species in Thailand, coarsely toothed. Fronds monomorphic. Stipes pale mostly in Peninsular Thailand. to dark brown. Laminae pinnately compound, simply

KEY TO THE SPECIES OF TAPEINIDIUMM IN THAILAND 1. Lamina simply pinnate; pinnae linear-lanceolate, up to 0.7 cm wide; margins crenate, serrate or bi-serrate; basal pinnae smaller than or as large as middle pinnae 1. T. pinnatum 1. Lamina bipinnatisect to tripinnatifi d; pinnae not linear-lanceolate; largest pinnae 1.5–8(–13.5) cm wide; basal pinnae larger than other pinnae 2. T. luzonicum

1. Tapeinidium pinnatum (Cav.) C.Chr., Index Rhizome short creeping, 2–3.5 mm diam., Filic.: 631. 1906; Christensen, Bot. Tidsskr. 32: internodes less than 8 mm long, densely scaly 345. 1916; Holttum, Rev. Fl. Malaya ed. 1, 2: 339, throughout. Scales light brown to brown, narrowly f. 196. 1955 [‘1954’]; Ching, Fl. Reipubl. Popularis triangular, 2–4 mm long, up to 12 cells wide at base, Sin. 2: 8. 1959; Kramer, Blumea 15(2): 553. 1968 apex long attenuate ending with 3–8 uniseriate cells, [‘1967’]; Kramer in Van Steenis & Holttum (eds), base obtuse, margin minutely toothed. Stipes Fl. Males., Ser. 2, Pterid. 1(3): 193. 1971; Kramer, stramineous or pale brown to dark brown, up to 20 cm Gard. Bull. Singapore 26: 8. 1972; Tagawa & long, up to 2.5 mm diam., scaly at base, grooved on Iwatsuki in Smitinand & Larsen (eds), Fl. Thailand adaxial side. Laminae simply pinnate, oblong or 3(2): 146. 1985; Boonkerd & Pollawatn, Pterid. oblong-lanceolate in outline, 20–46 × 14–23 cm, Thailand: 96. 2000.— Davallia pinnata Cav., Descr. coriaceous, terminal part pinnatisect and acute to Pl. (Cavanilles): 277. 1802.— Microlepia pinnata attenuate. Rachises green or pale brown, 0.5–1.5 mm (Cav.) Bedd., Handb. Ferns Brit. India: 64. 1883; diam. at base of lamina, grooved on adaxial side, Christ, Bot. Tidsskr. 24: 111. 1901.— Type: sharply keeled (carinate) on abaxial side. Pinnae up [but incorrectly labelled as being from to 27 pairs, consisting (from the base upwards) of “Chile & the Philippines”], Née s.n. (lectotype MA 1–3 pairs of lower pinnae and 5–24 pairs of upper [MA475617], fi rst step designated by Kramer (1968 pinnae. Lower pinnae slightly stalked (petiolules [‘1967’]), second step designated here; isolecto- less than 1 mm long), opposite or sub-opposite, type MPU [MPU017606] photo seen). For further linear-lanceolate, 3–14 × 0.2–0.6 cm, apex acute or synonymy see Kramer (1971). Fig. 1. acuminate, base cuneate, margin crenate, serrate or

SW 11611-p155-161-G8.indd 156 1/16/62 BE 5:07 PM A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT) 157

bi-serrate; basal pinnae smaller than or as large as Ecology.— In Thailand, Tapeinidium pinnatum other ones. Upper pinnae sessile, alternate, linear- grows on soil or rocks near streams in shady areas lanceolate, 4–15 × 0.3–0.7 cm, apex acute or in lowland evergreen, dry evergreen, and lower acuminate, base cuneate, margin crenate, serrate or montane forests, 80–1150 m altitude. Kramer (1971) bi-serrate; the main pinnae all more-or-less the same suggests that it is a facultative rheophyte. size, the upper ones gradually becoming smaller Proposed IUCN Conservation Assessment.— towards the frond apex. Costae distinct, sharply Least Concern (LC). Tapeinidium pinnatum is the keeled below, slightly grooved above. Veins distinct most common and widespread species of Tapeinidium below, once-forked. Sori submarginal, terminal on (Kramer, 1968 [‘1967’]) and several of the Thai 1(–2) veins, 0.5–0.75 × 0.5–1.5 mm; 1–3 in each localities are in protected areas. lobe, paraphyses 7-11 cells long. Indusia brown, pouch-shaped, free margin subentire. Sporangia The original description of Davallia pinnata spheroidal, 0.2–0.25 × 0.15–0.2 mm; annulus with Cav. (Cavanilles, 1802) was based on specimens 15–16 cells; sporangial stalk triseriate, 4–5 cells collected “in Chile and the Philippines” by the long. Spores light brown, ellipsoid, 35–45 × 20–30 μm, French-born Spanish botanist and explorer, Luis psilate. Née, during his 5-year expedition (1789–1794) to nearly all the Spanish possessions in the Americas Thailand.— SOUTH-EASTERN: Chanthaburi and Asia. While he is known to have collected large [Khitchakut district, Khao Khitchakut National Park, numbers of specimens in both Chile and the Krating waterfall area, 425 m, 10 Jan. 2009, Philippines there seems to be some confusion over Middleton 4671 (BKF, E [E00727425], P what was collected where as Davallia pinnata, like [P02433068])]; Trat [Ko Chang, Khlong Mayom, all members of the genus Tapeinidium, only occurs 100 m, 2 June 1923, Kerr 6503 (BK); 16 Feb. 1955, naturally in the Old World. Aware that Chile was an Smitinand 2164 (BKF); Ko Chang, Lem Dan Kao, incorrect locality, Kramer (1968 [‘1967’]) chose a 1150 m, 3 Oct. 1924, Kerr 9244 (BK); Ko Chang Née specimen (Née s.n.) from the Philippines at MA district, Khlong Phlu Ranger station, 80 m, 6 Jan. to serve as the “Type” (i.e. lectotype). Kramer cited 2009, Middleton et al. 4622 (BKF, E [E00736559])]; this “Type” again in his Flora Malesiana revision PENINSULAR: Krabi [Khao Phanom Bencha National (Kramer, 1971) but we cannot locate any Née Park, Khao Penom, 1000 m, 18 June 2006, Williams specimen at MA that is solely labelled as being from et al. 1910 (BKF); Pha Nam Yot, Khao Ngon Nak, the Philippines. Instead, there are two specimens of Noppharatthara Beach-Phi Phi Islands National Davallia pinnata collected by Née at MA that, due Park, 480 m, 12 Nov. 2018, Putthisawong 853 to the confusion over the collecting locality, are each (PSU)]; Nakhon Si Thammarat [Khao Luang area, still labelled as being from “Chile and the Philippines”. near Khiri Wong village, 16 May 1968, van Beusekom Although these labels are misleading it is reasonable & Phengkhlai 808 (BKF); Khao Nan National Park, to assume that both specimens were collected by Khao Khom, 10 May 2006, Boonkerd et al. 177 Née in the Philippines and, therefore, that either can (BCU)]; Phatthalung [Pa Bon district, Khao Sam Phu, serve as the lectotype of Davallia pinnata. We have 200 m, 29 June 2016, Putthisawong 175 (PSU)]; chosen Née s.n. with barcode MA475617 for the Phangnga [Klong Hin Poeng to Toong Rha, 700 m, second-step lectotypifi cation (see Art. 9.17 of the 27 Mar. 2000, Suksathan 2516, 36666 (QBG)]; Yala Shenzhen Code (Turland et al., 2017)). [Betong district, Ban Malao, 700 m, 25 Aug. 1923, Smith 1913 (BKF); Betong district, Hala-Bala Wildlife Sanctuary, trial up unnamed ‘1490’ mountain 2. Tapeinidium luzonicum (Hook.) K.U.Kramer, reached from shores of Bang Lang Reservoir, 1050 m, Blumea 15(2): 552. 1968 [‘1967’]; Kramer in Van 23 May 2005, Middleton et al. 3581 (BKF, E Steenis & Holttum (eds), Fl. Males., Ser. 2, Pterid. [E00246597])]. 1(3): 191, f. 11. 1971; Kramer, Gard. Bull. Singapore 26: 8. 1972; Tagawa & Iwatsuki in Smitinand & Distribution.— Southern India, Peninsular Larsen (eds), Fl. Thailand 3(2): 146, f. 10.5. 1985.— Malaysia, Singapore, (Sumatra to Davallia luzonica Hook., Sp. Fil. 1: 174, t. 60B, f. ), Sabah, Sarawak, Philippines, Taiwan and 2, 3 & 5. 1846.— Type: Philippines, Luzon, Cuming Japan (Ryukyu Islands). 139, p.p. (lectotype E [E00782194], designated

SW 11611-p155-161-G8.indd 157 1/16/62 BE 5:07 PM 158 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

here; isolectotypes B?, not seen but reported by Thailand.— PENINSULAR: Nakhon Si Thammarat Kramer (1971), GH [GH00020954] photo seen [Khao Nan National Park, Khao Khom, 10 May (mixed with T. pinnatum), GOET [GOET009148] 2006, Boonkerd et al. 173, 1766 (BCU); Khao Luang, photo seen, L [L0052135] photo seen). 1100–1768 m, 21 Jan. 1966, Tagawa et al. T4830 — Tapeinidium biserratum auct. non (Blume) (BKF, L); 1400 m, 23 Aug. 1967, Iwatsuki et al. Alderw.: Holttum, Rev. Fl. Malaya ed. 1, 2: 339, f. s.n. (BKF); 1100 m, 23 May 1968, van Beusekom 197. 1955 [‘1954’]; Tagawa & Iwatsuki, S.E. Asian & Phengkhlai 941 (BKF); 1400 m, 4 July 2017, Studies [Jap. J. S.E. Asian Studies] 5: 75. 1967. For Putthisawong 613 (PSU); 1400 m, 1 Jan. 2018, further synonymy see Kramer (1971). Fig. 2 Putthisawong 710 (PSU)]; Phatthalung [Pa Bon district, Khao Sam Phu, near summit, ca 950 m, 1 Rhizome moderately long creeping, 3–5 mm July 2016, Putthisawong 214 (PSU)]; Kongra district, diam., internodes up to 12 mm long, densely scaly Khao Lon Nom Sao, 900 m, 27 May 2017, Putthisawong throughout. Scales light brown to brown, narrowly 455 (PSU); 1100 m, 28 May 2017, Putthisawong triangular, 2–5 mm long, 6–15 cells wide at base, 467 (PSU)]; Trang [Yan Ta Khao district, Khao apex long attenuate ending with 2–6 uniseriate cells, Banthat mountains, near summit of Phu Pha Mek, base obtuse, margin irregularly toothed. Stipe dull 1200 m, 7 Apr. 2003, Middleton et al. 1993 (A, BKF, brownish yellow or brown to dark brown, up to 40 cm E [E00736795], L, MICH)]. long, up to 2.5 mm diam., scaly at base, grooved on Distribution.— Peninsular Malaysia, Indonesia adaxial side. Laminae bipinnatisect to tripinnatifi d, (Sumatra to Sulawesi), Sabah, Sarawak, and deltoid or oblong-ovate in outline, 16.5–40 × the Philippines. 7.5–30 cm, coriaceous, terminal part pinnatisect and acute to attenuate. Rachises dull brownish yellow, Ecology.— In Thailand, Tapeinidium luzonicum 0.5–1.5 mm diam. at base of lamina, grooved on grows on soil on slopes and stream banks in evergreen adaxial side, sharply keeled (carinate) on abaxial and lower montane forests, 900–1770 m altitude. side. Pinnae up to 24 pairs; consisting (from the base Proposed IUCN Conservation Assessment.— upwards) of 1–3 pairs of lower pinnae, 3–6 pairs of Least Concern (LC). This species is not under any middle pinnae, and up to 15 pairs of upper pinnae. immediate threat and several of the Thai localities Lower pinnae slightly stalked (petiolules less than are in protected areas. 3 mm long), opposite or subopposite, pinnatisect to bipinnatifid (see Fig 2B), lanceolate-deltoid in The original description of Davallia luzonica outline, 4–22 × 1.5–8(–13.5) cm, the basal pinnae Hook. (Hooker, 1846) was based on elements of the largest; pinnules (if present) subsessile, 1–3 pairs, Cuming 139, a mixed collection of at least three subopposite to alternate, shallowly lobed to pin- Tapeinidium species and, reputedly, one Athyriaceae natifi d (see Fig 2B), ovate-lanceolate to linear-lan- species (K000398343, not seen) collected in Luzon, ceolate in outline, 0.5–3.7 × 0.2–0.7 cm, apex obtuse the Philippines. As was common practice at this time to acuminate, base cuneate, basal basiscopic pinnules Hooker did not designate a holotype nor did he cite often the largest. Middle pinnae sessile, alternate, any specimen that could serve as the type of Davallia pinnatisect, oblong-lanceolate in outline, 4–14 × luzonica. Kramer (1968 [‘1967’]) suggested, with 1.2–3 cm. Upper pinnae sessile, alternate, shallowly some reservation [“Type: Cuming 139 (K ?; isotypes lobed to pinnatifid, oblong-lanceolate in outline, B, L).”], that Cuming 139 at K could be the 0.9–3.8 × 0.4–1.1 cm, base cuneate. Costae distinct, “holotype” but for reasons that were not explained sharply keeled below, fl attened or slightly grooved he did not cite any K specimen or propose an above. Veins in pinnules and ultimate lobes pinnate, alternative “holotype” (strictly speaking, a lectotype) veinlets simple or once-forked. Sori submarginal, in his Flora Malesian revision (Kramer, 1971). We terminal on 1(–2) veins, 0.25–0.5 × 0.5–1.5 mm, think it’s possible that by 1971 he had concluded, paraphyses 5-7 cells long. Indusia brown, pouch- as we have done, that the only specimen of Cuming shaped, free margin subentire. Sporangia spheroidal, 139 at K that is Tapeinidium (K000360561) is not 0.2–0.25 × 0.15–0.2 mm; annulus with 11–16 cells; T. luzonicum (so is not Davallia luzonica). Kramer’s sporangial stalk triseriate, 4–5 cells long. Spores light earlier attempt to lectotypify the name Davallia brown, ellipsoid, (30–)35–47.5 × (20–)22.5–32.5 μm, luzonica Hook. with the only Tapeinidium specimen psilate.

SW 11611-p155-161-G8.indd 158 1/16/62 BE 5:07 PM A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT) 159

Figure 1. Tapeinidium pinnatum (Cav.) C.Chr.: A–B. plants in natural habitats (A on soil, B on rock); C. section of fertile pinna showing serrate margins and submarginal sori terminal on veins; D–E. SEM micrographs of spores, D. equatorial view, E. close-up of surface showing lack of spore ornamentation. D & E. from Putthisawong 175 (PSU).

SW 11611-p155-161-G8.indd 159 1/16/62 BE 5:07 PM 160 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Figure 2. Tapeinidium luzonicum (Hook.) K.U.Kramer: A. plant in natural habitat (on soil slope); B. basal pinnae of a tripinnatifi d frond (but note that the division of basal pinnae can vary from pinnatisect (resulting in a bipinnatisect frond) to bipinnatifi d (resulting in a tripinnatifi d frond); C. section of fertile middle pinna showing typical pinnatisect division and submarginal sori terminal on veins; D–E. SEM micrographs of spores, D. equatorial view, E. close-up of surface showing lack of spore ornamentation. B. from Putthisawong 4677 (PSU) and D & E. from Putthisawong 455 (PSU).

SW 11611-p155-161-G8.indd 160 1/16/62 BE 5:08 PM A REVISION OF THE GENUS TAPEINIDIUM (LINDSAEACEAE) IN THAILAND (N. PUTTHISAWONG & S. CHANTANAORRAPINT) 161

of Cuming 139 at K must be disregarded since that Hooker, W.J. (1846). Species Filicum 1. W. Pamplin, specimen is in serious confl ict with the protologue London. 245 pp. (see Art. 9.19 of the Shenzhen Code (Turland et al., Kramer, K.U. (1968 [‘1967’]). The Lindsaeoid ferns 2017)). Since there are no appropriate specimens of of the Old World II. A revision of Tapeinidium. Cuming 139 at K we have chosen a specimen at E to Blumea 15(2): 545–556. serve as the lectotype of Davallia luzonica. This ______. (1971). Lindsaea Group. Flora Malesiana, specimenn of Cuming 1399 (E00782194) was previously Series 2, Pteridophyta 1(3): 177–254. housed in Glasgow University herbarium where Hooker once worked. Lehtonen, S., Tuomisto, H., Rouhan, G. & Christenhusz, M.J.M. (2010). Phylogenetics and classification of the pantropical fern family ACKNOWLEDGMENTS Lindsaeaceae. Botanical Journal of the Linnean We would like to thank the curators of BCU, Society 163: 305–359. BK, BKF, E, GH, GOET, K, L, MA, MPU, P, PSU, PPG I. (2016). A community-derived classifi cation QBG and JSTOR Global Plants for access to for extant lycopods and ferns. Journal of specimens and/or specimen images. We also thank Systematics and Evolution 54: 563–603. S. Lindsay (National Parks Board, Singapore) and Presl, C.B. (1851 [‘1849’]). Epimeliae Botanicae. an anonymous reviewer for useful suggestions. This A. Haase, Prague. 264 pp. work was supported by Prince of Songkla University Tagawa, M. & Iwatsuki, K. (1967). Enumeration of Research Fund (SCI610429S). The first author Thai pteridophytes collected during 1965–66. would like to express his sincere appreciation to the [Japanese Journal of] Southeast Asian Studies. Science Achievement Scholarship of Thailand 5: 23–120. (SAST) for fi nancial support. ______. (1985). Lindsaeaceae. In: T. Smitinand & K. Larsen (eds), Flora of Thailand. 3(2): 129–149. REFERENCES Royal Forest Department, Bangkok. Cavanilles, A.J. (1802). Descripción de las Plantas. Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, Imprenta Real, Madrid. 625 pp. W., Hawksworth, D.L., Herendeen, P.S., Knapp, Christensen, C.F.A. (1906). Index Filicum: sive S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, enumeratio omnium generum specierumque T.W., McNeill, J., Monro, A.M., Prado, J., Price, fi licum et hydropteridum ab anno 1753 ad fi nem M.J. & Smith, G.F. (eds) 2018: International anni 1905 descriptorum adjectis synonymis Code of Nomenclature for algae, fungi, and principalibus, area geographica etc. Hafniae plants (Shenzhen Code) adopted by the [Copenhagen] H. Hagerup. 744 pp. Nineteenth International Botanical Congress Holttum (1955 [‘1954’]). A revised Flora of Malaya, Shenzhen, China, July 2017. Regnum Vegetabile an illustrated systematic account of the Malayan 159. Glashütten: Koeltz Botanical Books. 254 fl ora, including commonly cultivated plants. Vol. pp. DOI https://doi.org/10.12705/Code.2018 2. Ferns of Malaya. Edition 1. Government Printing Offi ce, Singapore. 643 pp.

SW 11611-p155-161-G8.indd 161 1/16/62 BE 5:07 PM THAI FOREST BULL., BOT. 46(2): 162–216. 2018. DOI https://doi.org/10.20531/tfb.2018.46.2.08

Systematics of the Thai Calophyllaceae and Hypericaceae with comments on the Kielmeyeroidae (Clusiaceae)

CAROLINE BYRNE1, JOHN ADRIAN NAICKER PARNELL1,2,* & KONGKANDA CHAYAMARIT3

ABSTRACT The Calophyllaceae and Hypericaceae are revised for Thailand and their relationships to the Clusiaceae and Guttiferae are briefl y discussed. Thirty-two species are defi nitively recognised in six genera, namely: L., Kayea Wall., L. and Mesua L. in the Calophyllaceae and Blume. and L. in the Hypericaceae. A further four species of Calophyllum are tentatively noted as likely to occur in Thailand. Descriptions, full synonyms relevant to the Thai taxa, distribution maps, ecology, phenology, vernacular names, specimens examined and provisional keys are given. All species previously classifi ed in the genus Mesua have been moved to the genus Kayea, except L. Two taxa were found to be endemic to Thailand: Mammea harmandii (Pierre) Kosterm. and Hypericum siamense N.Robson. The distribution for the families in Thailand was found to vary with the Thai Calophyllaceae being found mainly in Central and Peninsular Thailand whilst the Thai Hypericaceae were found mainly in the North and the North-East of Thailand. Overall the numbers of collections housed in herbaria are few and more collections are necessary in order to give a comprehensive account of their distributions in Thailand.

KEYWORDS: Guttiferae, Flora of Thailand. Published online: 24 December 2018

INTRODUCTION from herbarium notes or directly from dried material. Ecological information was taken from specimens, The present work forms the basis of an account from field observations and from the literature. being prepared for the Flora of Thailand. It is largely Distribution maps were constructed using Arc-GIS based on the examination of 2,345 specimens from 9, Version 9.1. These maps show the distribution of worldwide herbaria including: A, AAU, ABD, B, all species in Thailand. The co-ordinates of localities BK, BKF, BM, C, CAL, CMUB, E, G, K, KKU, L, used have been taken from label data, and in the cases LINN, NY, P, PSU, QBG, SING, TCD, U and UPS. where only province names have been provided, a All descriptions are based on observations made co-ordinate point was mapped in the centre of that from herbarium material and living plants observed province to show the presence of the specimen. All while in the fi eld. Dimensions used are based on types that have been seen are indicated by (!) after herbarium specimens and living plants measured in the herbarium abbreviation. Where a new lectotype the fi eld. For dry specimens, fl owers were softened has been chosen, this is indicated after the designated in water before measurements were taken. Vernacular specimen. All lectotypes have been chosen on the names and geographical information were taken same basis: that is that they are the best available of from specimens and the literature. Flower, and bark all potential lectotype specimens or, in rare cases, the exudate colours in Hypericum and Cratoxylum always sole available specimen. In the synonymy, homotypic refer to fresh material: all other colours are drawn

1 Herbarium, Botany Department, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 2 Trinity Centre for the Environment, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 3 Forest Herbarium, National Park, Wildlife and Plant Conservation Department, 61 Phahonyothin Road, Chatuchak, Bangkok 10900, Thailand. * Corresponding author: [email protected]

© 2018 The Forest Herbarium

SW 11611-p162-216-G8.indd 162 1/16/62 BE 5:21 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 163 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

synonyms are indicated by the identity sign (≡). Bar Clusioideae was sister to the Bonnetiaceae (Wurdack codes are given where possible; but, for example, & Davis, 2009). So as to ensure monophyletic family neither ABD or LINN use bar codes and in many circumscriptions Wurdack & Davis (2009) reinstated herbaria not all specimens are bar-coded. It is hoped the Calophyllaceae, a decision repeatedly and recently that this account will enable the keys, descriptions re-confirmed (APG IV, 2016). Sun et al. (2016) and distributional information on taxa to be refi ned undertook a mega-analysis of a super-matrix of 9,300 before they are published in the Flora of Thailand. species and confi rmed that the Podostemaceae and Furthermore, it is clear that a number of taxa are Hypericaceae form a clade (100% Bootstrap support known from very few collections indeed, sometimes [BS]) with the Calophyllaceae being that clade’s only a single collection (e.g., Kayea elegans King): a sister (93% BS), this latter clade being sister to a situation repeatedly drawn attention to as a general more weakly supported clade (72% BS) comprising problem for many taxa in Thailand (Parnell et al., the Clusiaceae and Bonnetiaceae and with the whole 2003; van Welzen et al., 2011). lot forming a clade with 100% BS within the Malphigiales. Sun et al. (2016) also indicate that The Calophyllaceae and Hypericaceae and the Mesua in the Calophyllaceae is not monophyletic Clusiaceae and the Guttiferae necessitating the transfer of a number of species The families Calophyllaceae and Hypericaceae away from this genus. Some of these relationships are dealt with together in this paper as their status appear difficult to accept from a morphological as separate families has varied over time. Indeed, in perspective. In particular, the Podostemaceae are an the past, they have been dealt with as one family aquatic family of angiosperms that occur on rocks under the name Guttiferae or Clusiaceae, and that in waterfalls and rapids (Kato, 2004) and appear is how they were fi rst proposed to be treated for the quite diff erent to the other families in the Malpigiales Flora of Thailand when this work commenced. in terms of habit, indeed diff erent to any other family of Angiosperm. Koi (2005) suggested that saltational The Clusiaceae, Calophyllaceae and evolution might have occurred to give rise to the Hypericaceae belong to the order Podostemaceae, as there is a lack of intermediate (Chase et al., 1993; Davis et al., 2004; Stevens, 2006) species between this family and the Clusiaceae s.l. which contains 700 genera and over 16,000 species and Hypericaceae. But there are morphological in 30 families (Tokuoka &Tobe, 2006; Wurdack & similarities linking them to each other, including Davis, 2009). The phylogenetic placement of this various types of secretory structure and secondary order is poorly understood; it may lie either in the products (Kubitzki et al., 1978; Stevens, 2006; fabid clade of the (rosid I) or the malvid clade Wurdack & Davis, 2009). (rosid II) (APG IV, 2016). Phylogenetic relationships within the Malpighiales are also not fully resolved The Guttiferae (Clusiaceae, nom. alt.) sensu (Stevens, 2001 onwards; Davis et al., 2004; APG lato, comprises 27 genera and 1,050 species. Stevens II, 2003; Tokuoka & Tobe, 2006; APG III, 2009, (2006) considered there to be two subfamilies: Wurdack & Davis, 2009; Sun et al. 2016) though the Kielmeyeroideae and Clusioideae. interfamily relationships are now better understood The Kielmeyeroideae, now consideredd a separate (APG IV, 2016). Most recent phylogenetic studies family Calophyllaceae (Stevens, 2001 onwards; have been based on molecular analyses, using plastid APG III, 2009; APG IV, 2016; Mabberley, 2017) and nuclear gene regions (Savolainen et al., 2000a, contain two tribes, namely tribe Calophylleae with 2000b; Soltis et al., 2000; Stevens, 2001 onwards; 11 genera and tribe Endodesmieae with two genera APG II, 2003; Davis et al. 2004, 2005; Tokuoka & (Stevens, 2001 onwards). They are characterised by Tobe, 2006; Wurdack & Davis, 2009; APG III, 2009; spiral leaves with pellucid dots, moderate-sized APG IV, 2016; Sun et al. 2016). Wurdack & Davis cotyledons and perfect fl owers and as Stevens (2006) (2009) showed that the Clusiaceae are polyphyletic indicates (as Clusiaceae) are found mainly in moist, with the two commonly recognised subfamilies of tropical, lowland or lower montane forests. The the Clusiaceae s.l. not forming a clade: subfamily majority of genera are found in primary forests, some Kielmeroideae was shown to be sister to the species grow in peat swamp forests (Calophyllum Hypericaceae and Podostemaceae, while subfamily

SW 11611-p162-216-G8.indd 163 1/16/62 BE 5:21 PM 164 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

pisiferum Planch. & Triana, C. rupicola Ridl., in the genus Mammea (Clusiaceae): Mammea C. sclerophyllum Vesque, C. teysmannii Miq.), and harmandiii (Pierre) Kosterm. and M. siamensis (Miq.) some grow in black-water fl oodplains (species of T.Anderson. Three species were reported in the Caraipa Aublet, Haploclathra Benth.) (Stevens, genus Mesua (Clusiaceae), namely Mesua ferrea L., 2006). M. ferruginea (Pierre) Kosterm. and M. nervosa Planch. & Triana. The Clusioideae, now considered a separate family Clusiaceae (Stevens, 2001 onwards; APG III, 2009; APG IV, 2016; Mabberley, 2017), contain 14 TAXONOMIC ACCOUNT genera and 500 species (Mabberley, 2017) in three The following is a brief description of the four tribes (Stevens, 2001 onwards), namely Clusieae genera and 21(–25) species of Calophyllaceae and with fi ve genera, Garcinieae with two generan and the two genera and 12 species of Hypericaceae that Symphonieae with seven genera, and further details we have recognised in Thailand respectively: are beyond the scope of this publication. Calophyllum (Calophyllaceae).— Approximately Hypericaceae comprise 8 genera and 560 186 species, 14 (probably 18) of which occur in species (Mabberley, 2017). The family has a world- Thailand. The distribution is worldwide in tropical wide distribution and is divided into three tribes: areas including Africa, North and South America, Vismieae with two genera, Hypericeae with five Tropical and Temperate Asia, Australasia and the genera and Cratoxyleae with two genera. Among the Pacifi c. They are restricted to areas that are humid two Thai genera, Hypericum L. is found in temperate and in lowland or montane rainforests but can grow regions and high montane areas in tropical regions in drier, open areas (Stevens, 1980). Some of the main whereas Cratoxylun is found exclusively in the Old characteristic features of this genus include fi ssured World tropics. bark with exudate, leaves with distinct venation, The Calophyllaceae and Hypericaceae of Thailand terminal or axillary inflorescences, four to eight tepals and drupaceous fruits. Calophyllum inophyllum In his revision, Sangkaew (1999) reported 17 is one species in this genus that is of economic species of Calophyllum L. (under Clusiaceae) in importance as it is used in many countries for diff erent Thailand (Calophyllum calaba L., C. canum Hook.f., purposes. In Java, the seed oil and latex is used to C. depressinervosum M.R.Hend. & Wyatt-Sm., dye batik cloth. The timber is strong and durable C. dryobalanoides Pierre, C. inophyllum L., and used as a general-purpose timber for masts, C.macrocarpum Hook.f.,C. mollee King, C.pisiferum, bridgework, boat building and cabinet making. The C. polyanthum Wall. ex Choisy, C. rupicola, C. sclero- oils produced from the fruits are used to treat ulcers, phyllum, C. soulattri Burm.f., C. symingtonianum rheumatism and skin diseases such as eczema, and a M.R.Hend. & Wyatt-Sm., C. tetrapterum Miq., decoction of the bark and latex is used both internally C. teysmannii, C. thorelii Pierre and C. touranense and externally as a remedy for many infections and Gagnep. ex P.F.Stevens). Forest Herbarium (2001) also for skin and eye diseases and rheumatism. In reported 29 taxa occurring in Thailand in the local medicine, the fl owers, leaves and seeds are Clusiaceae and Hypericaceae together under the also used. Although this tree grows slowly, it is also label Guttiferae. This latter publication also included used for reforestation and aff orestation projects and all of the 17 Calophyllum species recorded by at shorelines to protect the coast (ICRAF, 2008). Sangkaew (1999), six species in the genus Cratoxylum The 14 Thai species we so far recognise and describe (Hypericaceae), namely are: Calophyllum calaba, C. dryobalanoides, (Vahl) Blume, C. cochinchinense (Lour.) Blume, C. inophyllum, C. macrocarpum, C. pisiferum, C. formosum (Jack) Dyer, C. formosum subsp. C. polyanthum, C. rupicola, C. sclerophyllum, prunifl orum (Kurz) Gogelein, C. maingayi Dyer, C. soulattri, C. symingtonianum, C. tetrapterum, C. sumatranum (Jack) Blume subsp. neriifolium C. teysmannii, C. thorelii and C. touranense. Four (Kurz) Gogelein, and one species in the genus other species have been reported by other workers Hypericum (Hypericaceae), namely Hypericum (see discussion below). hookerianum Wight & Arn. Two species were recorded

SW 11611-p162-216-G8.indd 164 1/16/62 BE 5:21 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 165 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Kayea (Calophyllaceae).— Approximately 75 Hypericum (Hypericaceae).— Approximately species occur in tropical Asia namely , 420 species found worldwide but mainly in temperate Myanmar, Cambodia, India, Laos, Malaysia, Nepal, regions of Africa, North America, Asia, Europe and Singapore, Sri Lanka, Sumatra, Thailand and the former Soviet Union. This genus consists of trees Vietnam. This genus comprises trees with opposite, or herbs that show a lot of morphological variation. glabrous and coriaceous leaves. The infl orescences The leaves are opposite and can be sessile or subses- are large with solitary flowers and there are four sile. There are four to fi ve sepals and petals present sepals and petals present. The fruit is an indehiscent and the fl owers are usually yellow and showy. The fl eshy fruit with one to four seeds. Three species occur petals are sometimes spotted with black glands. in Thailand, namely Kayea elegans, K. ferruginea Hypericum species are cultivated as ornamentals as Pierre and K. kunstleri King. they have showy flowers. In Britain and Ireland, there are over 80 varieties available as garden plants Mammea (Calophyllaceae).— Approximately (Lord et al., 2004). Many members of this genus 75 species, occurring throughout the tropics in also yield compounds important to the pharmaceutical Africa, Central America, Madagascar and Tropical industry such as fl avonoids, tannins, phenol-carbonic Asia. Madagascar is thought to be the centre of origin acids and xanthones (Frohne & Pfänder, 2005) and for this genus (Stevens, 2006). The species are all L. is known for its’ medicinal trees with clear or coloured resinous sap and are properties, as it is used as a herbal remedy to help found in areas of low to medium altitude. The leaves mild depression (Frohne & Pfänder, 2005). In are simple and opposite and some have black glandular Thailand six species occur mainly in the Northern dots present. The inflorescences are quite often Region. These are Hypericum henryi H.Lév. & caulifl orous and the fruit is fl eshy and coriaceous Vaniot. subsp. hancockiii N.Robson, H. hookerianum, when dry. There are three species found in Thailand, H. japonicum Thunb., H. napaulense Choisy, namely Mammea brevipes (Craib) Kosterm., H. patulum Thunb. and H. siamense. M. harmandii and M. siamensis. Cratoxylum (Hypericaceae).— Contains eight Mesua (Calophyllaceae).— One species in species in three sections (Kochummen, 1973) found Tropical Asia, namely Mesua ferrea, which has long in northeastern India and southern China to western been cultivated in the area and is found growing Malesia at low altitudes or in low montane areas. naturally at low altitudes. This genus comprises trees Five species occur in Thailand. Species vary from with a sticky exudate which dries black. The leaves deciduous to evergreen trees or shrubs that are usually are opposite, often with dots on the upper surface. glabrous and exude a yellow resinous sap. The leaves The infl orescences are solitary, axillary or terminal are papyraceous and usually have pellucid dots and there are four or fi ve sepals and petals present. present. The infl orescences are terminal panicles or The fruits of this genus are quite variable and can short terminal cymes or sometimes axillary cymes. be drupes, berries or capsules. Mesua ferrea is often There are fi ve sepals and petals present and the corolla used in folk medicine for a variety of skin disorders. is usually pink (red) or white. The genus Cratoxylum It is found naturally growing and it is also cultivated is used as a source of timber (Heywood, 1993) and throughout Asia and has many other uses also: the the wood is used for joinery and for light construction seed is used in meal for poultry as it is a good source and furniture making (MTC, 2017). The Thai taxa of protein and energy, it is used for fi rewood, the are Cratoxylum arborescens, C. chinense, C. formosum timber is used in heavy construction and furniture subsp. formosum, C. formosum subsp. prunifl orum, making, the fl owers are used for dyeing fabrics, to C. maingayi and C. neriifolium Kurz. stuff pillows and in cosmetic products and an oil called nahor is extracted from the seeds and is used to treat dandruff and rheumatism (ICRAF, 2017).

KEY TO THE FAMILIES 1. Fruit indehiscent, with few, often large seeds 1. Calophyllaceae 1. Fruit dehiscent, with many small seeds 2. Hypericaceae

SW 11611-p162-216-G8.indd 165 1/16/62 BE 5:21 PM 166 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1. CALOPHYLLACEAE Tepals 4–8 or Calyx 2–5, free sepals. Corolla petals 4–5, free. Stamens 4–∞, aggregated into fascicles. Evergreen trees or shrubs with canals or glands Ovary superior, carpels 1–many, ovules 2–∞. Fruits present in all parts of the plant; usually glabrous, uni-/ berries, drupes or capsules, size varies immensely. multicellular hairs sometimes present. Leaves simple, Seeds ± winged, ± arillate; cotyledons often huge. petiolate, opposite, alternate or whorled. Infl orescences axillary or terminal, sometimes comprising single Pantropical with 14 genera and 750 species; fl owers, bisexual or unisexual, radially symmetric. 4 genera and 21(–25) species in Thailand.

KEY TO THE GENERA 1. Ovary 1-celled 2. peltate; ovule 1 1. Calophyllum 2. Stigma 4-fi d; ovules 4(–8) 2. Kayea 1. Ovary 2(–many)-celled 3. Sepals 2; ovary 2(–many)-celled 3. Mammea 3. Sepals 4–5; ovary 2-celled 4. Mesua

1. CALOPHYLLUM — Apoterium Blume, Bijdr. Fl. Ned. Ind. 5: 218. L., Sp. Pl.: 513. 1753; L., Gen. Pl. ed. 5: 229. 1754; 1825. Type species: Apoterium sulatri Blume. Choisy, Mém. Soc. Hist. Nat. Paris 1: 228. 1823; Trees with fi ssured bark usually yellow-brown Choisy in DC., Prodr. 1: 562. 1824; Endl., Gen. Pl.: to grey-brown, with two diff erent kinds of exudate: 1028. 1840; Choisy, Descr. Guttif. Inde: 41. 1849; a clear honey exudate or an opaque white exudate. Miq., Fl. Ned. Ind. 1(2): 509. 1859; Planch. & Triana, Leaves simple, petiolate, opposite, coriaceous; close, Ann. Sci. Nat. Bot. Sér. 4, 15: 241. 1862; T.Anderson fi ne parallel veins from midrib towards margin, some in Hook.f., Fl. Brit. India 1: 271. 1874; Vesque, having distinct intramarginal veins, veins are usually Epharmosis 2: 6. 1889; King, J. Asiat. Soc. Bengal, distinct; the midrib is depressed on upper surface. Pt. 2, Nat. Hist. 59: 172. 1890; Vesque in DC. Monogr. Infl orescences terminal or axillary. Floral parts tepals Phan. 8: 529. 1893; Trimen, Handb. Fl. Ceylon 1: 98. 4 or 8, rarely more; fl owers are similar among species; 1893; Engl. in Engl. & Prantl, Nat. Pfl anzenfam. 3(6): white; bracts and bracteoles ovate and caducous. 220. 1895; Pit. in Lecomte, Fl. Indo-Chine 1(4): 316. Stamens 30–600; usually glabrous; basifi xed; vertically 1910; Ridl., Fl. Malay Penins. 1: 181. 1922; Engl. in dehiscent; fi laments are slender and joined at base Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 21: 192. into 4–6 bundles. Ovary 1-celled; 1 ovule; glabrous; 1925; Gagnep. in Humbert, Suppl. Fl. Indo-Chine stigma peltate. Fruit drupe; ovoid to globose; varying 1(3): 268. 1943; Perrier de la Bathie in Humbert, Fl. from yellow/green to orange/brown to black. Seed Madagasc. 136: 3. 1951; M.R.Hend. & Wyatt-Sm., with large cotyledons. Gard. Bull. Singapore 15: 285. 1956; Maheshw., Bull. Distribution.— 186 species in Africa (Tropical, Bot. Surv. India 2: 139. 1960; Backer & Bakh.f., Fl. Madagascar and the Mascarenes), Asia Temperate Java 1: 384. 1963; A.C.Sm. & S.P.Darwin, J. Arnold (East and ), Asia Tropical (Indian Arbor. 55: 216. 1974; P.F.Stevens, Austral. J. Bot. 22: Subcontinent, Malesia), Pacifi c, Australasia, North, 349. 1974; P.F.Stevens, J. Arnold Arbor. 61: 167. Central and South America (Mexico to Argentina 1980; H.W.Li et al., Flora of China 13: 38–40. 2007. including Caribbean); 14 (probably 18) species in ≡ Ponna Rheede ex. Boehm., Ludw. Def. Gen. Pl. Thailand. 239. 1760, nom. illeg. superfl . ≡ Balsamaria Lour., Fl. Cochinch. 2: 469. 1790. nom. illeg. superfl .?. Subsequent to the preparation of this account Type species: Calophyllum inophyllum L. Gardner et al. (2015) confirmed the report of — Calaba Mill., Gard. Dict. Abr., ed. 4. 1754. Sangkaew (1999) concerning the occurence of Callophyllum molle in Thailand. This species is — Augia Lour., Fl. Cochinch.: 337. 1790, nom. rej., known as a rare tree of lowland evergreen forest in pro minore parte. Type species: Augia sinensis Lour. Narathiwat which represents an extension of its [see Stevens, 1980].

SW 11611-p162-216-G8.indd 166 1/16/62 BE 5:22 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 167 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

known range in the Malay Peninsula. Gardner et al. (2015) and Sangkaew (1999) that of C. canum from (2015) also report two specimens of the otherwise southernmost Thailand. Calophyllum depressiner- Malesian Calophyllum wallichianum Planch. & vosum obviously has acuminate leaves with the latex Triana from lowland evergreen or evergreen bamboo canals clearly impressed, and C. canum has a forest in Phangnga, Ranong. The latter species is distinctive swollen terminal bud, fl owers with very similar to Calophyllum macrocarpum and would key large numbers of stamens and leaves with a wavy out along with it in the key below; whilst the former margin. At the time of writing of this account we species is the only species of Thai Calophyllum that had not seen specimens of these species. Apart from is obviously hairy on its leaves, especially on the their inclusion in the key below and its general undersurface of the young leaves. Gardner et al. (2015), improvement, we require further information to Sangkaew (1999) and Stevens (1980) also report the confi rm the colour of their bark exudates; therefore, occurrence of Calophyllum depressinervosum from we would welome further relevant information on south and southeast Thailand, and Gardner et al. all of the above species.

KEY TO THE SPECIES 1. Bark exudate a clear honey colour 2. Outer pair of tepals dorsally pubescent 3. Tepals more than 8; outer pair of tepals equalling the next pair in shape and size 14. C. touranense 3. Tepals 8 or fewer; outer pair of tepals not equal to the next pair in shape and size 4. Stamens less than 100; fruit ≤ 1cm in diameter 1. C. calaba 4. Stamens more than 100; fruit > 1 cm in diameter 5. Leaf apex acuminate to rarely acute; branchlets 4-angled 4. C. macrocarpum 5. Leaf apex acuminate, retuse, or cuspidate; branchlets terete 6. Branchlets fl attened or rarely terete; leaf apex acuminate to acute 6. C. polyanthum 6. Branchlets terete; leaf apex cuspidate to retuse 7. Stamens less than 200; infl orescences terminal and from upper leaf axils; fruit spherical to ellipsoid, drying light green to purple 13. C. thorelii 7. Stamens more than 200; infl orescences axillary racemes; fruit ovoid to rotund, drying reddish-brown to almost black 8. C. sclerophyllum 2. Outer pair of tepals dorsally glabrous 8. Leaves slightly coriaceous; fruit ellipsoid to obovoid; stamens less than 70 10. C. symingtonianum 8. Leaves coriaceous; fruit ellipsoid, globose, obovoid or subglobose; stamens usually more than (65–)70, if 70 or less then other characters not as above 9. Leaf apex retuse to round 3. C. inophyllum 9. Leaf apex acute to acuminate 10. Stamens 35–90; tepals (4–)8; intramarginal veins absent 11. C. tetrapterum 10. Stamens 65–220; tepals 8; intramarginal veins present 11. Tree without buttresses; infl orescences terminal and from upper leaf axils, usually covered with brown pubescence; fruit ovoid, apex acute to acuminate 2. C. dryobalanoides 11. Tree with/without buttresses, if present, small (to <1 m); infl orescences in racemes, usually glabrous, but occasionally covered with brown pubescence; fruit ellipsoid to subglobose, apex acute to round 12. C. teysmannii 1. Bark exudate not a clear honey colour, somewhat opaque, usually milky white/yellow 12. Youngest twigs glabrous 7. C. rupicola 12. Youngest twigs pubescent 13. Fruit yellow-green or pale brown 1. C. calaba 13. Fruit orange or purple or black 14. Youngest twigs and base of infl orescences orange pubescent; fl owers in axillary racemes; leaves ovate or elliptic 5. C. pisiferum 14. Youngest twigs and base of infl orescences red-brown pubescent; fl owers axillary, in cymes; leaves oblong-ovate to elliptic-ovate 9. C. soulattri

SW 11611-p162-216-G8.indd 167 1/16/62 BE 5:22 PM 168 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1. Calophyllum calaba L., Sp. Pl.: 732. 1753; intramarginal vein inconspicuous but present. P.F.Stevens, J. Arnold Arbor. 61: 256. 1980; Infl orescence axillary, covered in brown pubescence, P.F.Stevens in Jarvis et al., Regnum Veg. 127: 28. 3–15 fl owers per infl orescence, pedicels 0.3–2.8 cm 1993. Type: Sri Lanka, Herb. Hermann 3: 3, No. 202 long. Floral parts tepals 4, sometimes 6, the outer (lectotype BM [BM000621800!], designated by pair elliptic-obovate to ovate, 3–5 × 2–3 mm, Stevens [1993]). sometimes dorsally brown pubescent, inner one — Calophyllum saigonense Pierre, Fl. Forest. elliptic-oblong, 4.5–7 × 1.5–3 mm, rarely dorsally Cochinch. 1: t. 105. 1885; Vesque in DC. Monogr. pubescent. Stamens in fascicles, approximately Phan. 8: 602. 1893; Pit. in Lecomte, Fl. Indo-Chine 20–95 per fl ower; fi laments 2–6 mm long; anthers 1(4): 318. 1910; Craib in Schmidt, Bot. Tidsskr. 32: 1–1.2 mm long, orange. Ovary 1.5–2 mm long; style 328. 1915; Craib, Fl. Siam. 1: 121. 1925; Gagnep. 2–4 mm long. Fruit ellipsoid to ovoid, apex rounded, in Humbert, Suppl. Fl. Indo-Chine 1: 274. 1943; 0.5–1 × 0.5–0.8 cm, yellow-green to pale brown Pham & Nguyên, Cây-Có Mièn Nam Viet-Nam 179. when ripe. 1960; Pham, Cây-Có Mièn Nam Viet-Nam 2(2): Thailand.— (Map 1.1.1): NORTH-EASTERN: 303. 1970; H.Keng, Gard. Bull. Singapore 28: 245. Nong Khai [Chaiyaburi, 21 Feb. 1924, Kerr 8526 1976. Type: Cochinchine, Beucar, Pierre 3649 (ABD, BK, BM, E, K, TCD)]; EASTERN: Buri Ram (lectotype P! designated by Stevens [1980]; isolec- [21 Nov. 1976, Phengklai et al. 3331 (A, BKF, totypes E [E00209519!], K [K000380003!], P!). PSU)]; Surin [4 Dec. 1976, Phengklai et al. 3624 — Calophyllum curtisii Ridl., J. Asiatic Soc. Bengal (A, BKF, PSU); Sangkha, 13 Jan. 1924, Kerr 8271 Pt. 2, Nat. Hist. 59: 176. 1890; Curtis, J. Straits (ABD, BK, BM, C, K, P, TCD)]; Roi Et [5 Feb. Branch Roy. Asiatic Soc. 25: 78. 1894; Ridl., Fl. 1925, Wanarale 62 (BK, K); T. Banna, Suwanaphum, Malay Penins. 1: 185. 1922; M.R.Hend. & Wyatt-Sm., 13 Aug. 1982, Sutheesoen 5342 (BK); ibid. 17 Aug. Gard. Bull. Singapore 15: 321. 1956; Kochummen, 1982, Sutheesoen 5358 (BK)]; Yasothon [Mar. 1982, Malayan Forest Rec. 2(17): 221. 1965; Whitmore, Smitinand s.n. (BKF)]; Si Sa Ket [Kantharalak Tree Flora of Malaya 2: 177. 1973; Corner, Gard. District, 23 Mar. 1966, Sangkhachand 201 (BK); 20 Bull. Singapore Suppl. 1: 104. 1978. Type: Malaysia, Jan. 1965, Phengklai 900 (C, K, L, P); 20 Jan. 1965, Malacca, Penang, Curtis 523 (lectotype K Phengklai 38105 (E); Chong Bat Lak, Kantaralak, [K000199886!], designated here; isolectotypes BM 11 Apr. 1976, Maxwell 76-205 (AAU, BK, L); [BM000611470!], BO not seen). Kantharalak District, Dongkrak Range, Chong Bat Lak, 20 Aug. 1976, Maxwell 76-592 (AAU, BK, Trees to 30 m tall; dbh 20 cm; bole straight L)]; Ubon Ratchathani [Lam Don Hoi, 4 Dec. 1968, without buttresses; outer bark deeply fi ssured yellow Smitinand & Turbang 10504 (BKF); Khong Chiam to grey-yellow, inner bark red-pink; exudate clear Falls, 16 Oct. 1990, Smitinand 90-267 (BKF)]; honey coloured or yellow or white, sticky. Branchlets SOUTH-EASTERN: Chon Buri [Nong Yai Bu, 1914, fl attened to terete, some smooth-waxy, some showing Collins 645 (K); Sri Racha, Na Phrow, Apr. 1922, tiny minute hairs on stems, tomentose; internodes Collins 804 (ABD, BM, K); Hup Bon, Sri Racha 1–3 cm long. Leaves petioled, petiole 0.3–1.9 cm Forest, 25 Oct. 1927, Collins 1661 (BM, K); Sri long; young leaves reddish pubescent, blade 3.5–11.2 Racha Forest, 23 Dec. 1927, Collins 1779 (BK, K)]; × 0.9–4.4 cm, elliptic to ovate, concave above and Chanthaburi [Ban Chao Lao, 8 Feb. 1991, Chayamarit convex below, base cuneate to attenuate to slightly 281 (BKF); Khao Sabap, 7 Jan. 1930, Kerr 17993 acute, apex acute to round, sometimes cuspidate; (A, B, BK, BM, C, K, L)]; Trat [Khao Saming, 26 venation very obvious on both surfaces, midrib Jan. 1927, Put 5666 (BK, K, L); Ko Chang, Khlong depressed on upper surface and pronounced on lower Son, 1 Mar. 1900, Schmidt 668 (C, K); Ko Kut, Ao surface, orange/brown on lower surface and red/ Salad, 9 Apr. 2002, Phengklai et al. 13582 (BKF)]; light tan on upper surface, midrib wide (2 mm) on PENINSULAR: Chumphon [Bang Son, 9 Jan. 1927, lower surface, narrowing towards apex to approxi- Kerr 11318 (A, BK, C, K, L, P)]; Ranong [Ban mately 1 mm wide, upper and lower surface smooth, Nakha, 9 Dec. 1976, Santisukk 797 (A, BKF); Khlong coriaceous, green/orange/yellow/brown; secondary Nakha, 24 Feb. 1974, Geesink et al. 7561 (AAU, veins present, dense, arising at same points on both BKF, C, K, L, P); Ban Laem Lieng, 1 Feb. 1927, sides of the midrib, approximately 0.5 mm apart, Kerr 11733 (A, BK, BM, C, K, L, P); 14 Jan. 1929,

SW 11611-p162-216-G8.indd 168 1/16/62 BE 5:22 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 169 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Kerr s.n. (BK)]; Surat Thani [Khao Tham Nong, Na Uses.— The wood is used in house construction. San, 19 June 1966, Sakol 1070 (BK); Ko Pa-ngan, The fruits are edible (Sangkaew, 1999). 4 Nov. 1927, Put 1155 (BK, BM, K); ibid., 4 Nov. Conservation.— IUCN Regional (Thailand) 1927, Put 11566 (BK, BM, K, L); Chaiya, 10 Jan. Status: Least Concern (LC). 1935, Seidenfaden 25766 (C); Ko Tao, 23 Sept. 1928, Kerr 16065 (A, BK, BM, C, K, L, P); Than Khanon, Pierre s.n. (BKF)]; Phangnga [Ko Kho Khao, 4 Dec. 2. Calophyllum dryobalanoides Pierre, Fl. Forest. 1965, Sangkhachand 1185 (BKF, K, L); between Cochinch. 1: t. 106. 1885; Vesque in DC., Monogr. Thai Muang & Thung Maphrao, 29 Jan. 1958, Phan. 8: 601. 1893; Pit. in Lecomte, Fl. Indo-Chine Smitinand 41577 (BKF); Laem Lieng to La-un, 7 Mar. 1(4): 319. 1910; Gagnep. in Humbert, Suppl. Fl. 1930, Kerr 18419 (K, L); Ko Yao Yai, 2 Mar. 1929, Indo-Chine 1: 274. 1943, pro minore parte; Pham Kerr 17291 (A, BK, BM, C, K, L, P); ]; Phuket & Nguyên, Cây-Có Mièn Nam Viet-Nam, 179. [Tha-chai/-chat, Dec. 1986, Phengklai & Smitinand 1960; Pham, Cây-Có Mièn Nam Viet-Nam, 2(2): 6105 (AAU, BKF, L, P)]; Krabi [Ao Luek,15 Mar. 302. 1970; P.F.Stevens, J. Arnold Arbor. 61: 232. 1930, Kerr 185677 (L)]; Phatthalung [20 Oct. 1938, 1980. Type: Cochinchine (Vietnam), Pierre 83 Siwanna s.n. (BKF)]; Trang [Hat Chao Mai NP, (lectotype P [P00526184!], designated by Stevens Sikao District, 14 Dec. 1995, Mauric 15 (BKF); [1980]; isolectotypes A [A000672326!], BM without locality, 27 May 1934, Poilane 23626 [BM000611356!], C( [C10009442!, fruits only], K (BKF)]; Satun [Tarutao, Son Bay, 29 Feb. 1980, [K000677426!], L [L0011725!, L0011726?], P Congdon 400 (A, AAU, PSU); Ky Island, Tarutao, [P00526185!, P00526186!],SING [SING0054598!). 22 Mar. 1980, Congdon 493 (A, AAU); Tarutao Trees to 30 m tall; dbh 20 cm; bole straight Island, 26 Apr. 1981, Congdon 12600 (A); Kanchanadit, without buttresses; outer bark fi ssured, brown-yellow 3 Jan. 1928, Kerr 13653 (BK, BM, K)]; Songkhla or grey-yellow; exudate clear honey coloured. [Kobe Island, Sathingpra District, 26 May 1984, Branchlets quadrangular, slender, some with slightly Srirugsa 835 (A, BKF, PSU)]; Khlong Hoi Khong, brown pubescence, or waxy or glabrous; internode Hat Yai District, 11 Jan. 1985, Maxwell 85-65 (A, 1–2.5 cm long. Leaves petioled, petiole 0.6–1.5 cm BKF, PSU)]; Kho Hong Hill/Prince of Songkla long; blade 4–8.9 × 1.5–3.5 cm, lanceolate to elliptic- University Campus, 29 Dec. 1985, Maxwell 85-1181 entire, coriaceous, very smooth, dark green adaxially, (A, AAU, BKF, L, PSU); Kho Hong Hill/Hat Yai, paler green abaxially, base cuneate to slightly acute, 27 Apr. 1986, Maxwell 86-261 (A, BKF, L, PSU)]; apex acuminate; venation very pronounced, midrib Pattani [21 July 1990, Santisuk s.n. (BKF)]; Yala not obvious on either surface, more pronounced on [30 Jan 1931, Put 3668 (BK, BM, C, K, L); ibid., lower surface, yellow, distance between veins 30 Jan 1931, Put 3670 (A, BK, BM, C, K, L, P)]; 0.2–0.8 mm, veins evenly spaced and meeting at the Narathiwat [Khok Mai Ruea, , 12 Jan. 1986, same points on opposite sides of the midrib, intra- Niyomdham et al. 11366 (A, AAU, BKF, C, K, L, P); marginal veins present. Infl orescence terminal and ibid., 25 Apr. 1986, Niyomdham et al. 1225 (A, BKF, from upper leaf axils, 7–37 fl owers per infl orescence, C, K, L, P)]; Unknown locations [Unknown collector covered with brown pubescence, pedicels 0.6–1 cm 1777 (BKF); 10 Jan. 1974, Conran 291 (PSU); Hewitt long. Floral parts tepals 8, the outer pair oval to 2063 (BM, K)]. suborbicular, 3.5–5 × 3–4 mm, dorsally slightly Distribution.— Andaman Islands, Vietnam to brown pubescent on outer side near base or glabrous, Borneo, Sumatra, Java and the Lesser Sunda Islands. inner one obovate to suborbicular, 4–6 × 5–6 mm, glabrous, the next two the same shape and size, Ecology and phenology.— Savannah, evergreen elliptic-obovate, 4–5 × 3.5–4 mm, glabrous, white. forest, secondary forest; from near sea level to 650 m Stamens in fascicles, 140–170 stamens per fl ower; elevation. Flowering: August–December Fruiting: fi lamentt 2–3 mm long; anther 0.6–1 mm long, orange/ October–April. yellow. Ovary 1.5–2 mm long, style 1.5–3 mm long. Vernacular.— Pha uung (พะอูง); pa-ong (ปะอง); Fruit ovoid, apex acute to acuminate, 1–1.5 × pa-ung (ปะอุง); phanghan klet raet (พังหันเกล็ดแรด) 0.7–1 cm, green. (Sangkaew, 1999).

SW 11611-p162-216-G8.indd 169 1/16/62 BE 5:22 PM 170 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Thailand.— (Map 1.1.2): SOUTH-WESTERN: 76. 1961; H.L.Li, Woody Fl. Taiwan: 601. fi g. 235. Phetchaburi [ Kaeng Krachan, Kaeng 1963; Baker & Bakh.f., Fl. Java 1: 386. 1965; Krachan NP, on road from Ban Krang Camp to Khao Whitmore, Guide Forests Brit. Solomon Is.: 77. Phanoen Thung Ranger Substation, 12 Aug. 2002, 1966; Whitmore, Tree Flora of Malaya 2: 186. Middleton et al. 999 (A)]; SOUTH-EASTERN: Trat [Ko 1973; P.F.Stevens, Austral. J. Bot. 22: 374. 1974; Chang, Khlong Mayom, Salak Khok, 6 Apr. 1923, A.C.Sm. & Darwin, J. Arnold Arbor. 55: 223. fi gs. Kerr 6390 (ABD, BM); Ko Chang, Khlong Non Si, 6–8. 1974; P.F.Stevens, J. Arnold Arbor. 57: 170. 24 Sept. 1924, Kerr 9175 (BM); Bo Rai, 28 Nov. 1976; Bamps et al. in Polhill, Fl. Trop. E. Africa, 1924, Kerr 9462 (BK, K, L)]. Guttiferae: 3. 1978; Perry, Med. Pl. E. & SE. Asia, Distribution.— Cambodia, Vietnam. 173. 1980; P.F.Stevens, J. Arnold Arbor. 61: 324. 1980; S.Gardner et al., Field Guide Forest Trees N. Ecology and phenology.— Evergreen forest, Thailand: 53. 2000; H.W.Li et al., Fl. China 13: 53. secondary forest, near freshwater; from 50 to 1200 2007. Type: Sri Lanka, Herb. Hermann 2: 82, No. m elevation. Flowering: September–December. 201 (lectotype BM [BM000621780!], designated Fruiting: January–March. (August). by Stevens [1980]). Vernacular.— Pha-ong (พะอง) (Sangkaew, — Balsamaria inophyllum Lour., Fl. Cochinch. 2: 1999). 470. 1790. Type: Unknown collector s.n. possibly Uses.— The wood is used in construction. Loureiro s.n. (holotype BM [BM000611357!]). Fragrant oil emitted from the fl ower is used in hair — Calophyllum ovatifolium Noronha, Verh. Batav. dressing (Sangkaew, 1999). Genootsch. Kunsten 5(4): 13. 1790 Type: not Conservation.— IUCN Regional (Thailand) located. Status: Data Defi cient (DD). — Calophyllum bingatorr Roxb., Hortus Bengal. 41. 1814, nom. nud.; Don, Gen. Syst. 1: 622. 1831; 3. Calophyllum inophyllum L., Sp. Pl.: 513. 1753; Roxb., Fl. Ind. 2: 607. 1832. Type: Moluccas, Burm.f., Fl. Ind. 120. 1768; G.Forster, Fl. Ins. Ambon, Rumphius, Herb. Amboin. 2: t. 71. 1741. Austral.: 41. 1768; Choisy in DC. Prodr. 1:562. — Calophyllum blumei Wight, Illus. Ind. Bot. 1: 1824; Blume, Bijdr. Fl. Ned. Ind. 5: 217. 1825; 128. 1840; Walp., Rep. Bot. Syst. 1: 397. 1842. Type: Blanco, Fl. Filip.: 612. 1837; Wight, Icon. Pl. Ind. not located. Orient.: t. 77. 1839; Thwaites, Enum. Pl. Zeyl.: 51. Trees 7–30 m tall; dbh 11–12 cm; bole ± straight, 1858; Miq., Fl. Ned. Ind. 1(2): 510. 1859; Planch. without buttresses; outer bark slightly fissured, & Triana, Ann. Sci. Nat. Bot. Sér. 4, 15: 282. 1862; rough, grey to brown; exudate the colour of clear Benth., Fl. Austral. 1: 183. 1863; T.Anderson in honey. Branchlets striate, slightly brown pubescent Hook.f., Fl. Brit. India 1: 273. 1874; F.Muell., or glabrous, yellow-greenish sap in centre of branch- Descr. Ecology and Phenology Papuan Pl.: 36. lets; internode 2–3 cm long. Leaves petioled, petiole 1875; Vesque, Epharmosis 2: t. 1. 1889; Vesque in 1.5–3 cm long; blade 8–15 × 4–8 cm, elliptic-obovate, DC. Monogr. Phan. 8: 544. 1893; Brandis, Indian sometimes oblong, obtuse, rounded, coriaceous, dark Trees: 54. fi g. 43. 1907; Pit. in Lecomte, Fl. Indo- green and concave adaxially, dull green and convex Chine 1(4): 324. 1910; Heckel, Ann. Mus. Hist. Nat. abaxially, rarely glaucous, base cuneate to attenuate, Marseille 2(10): 262, pl. 25. 1912; Koord-Schum., apex retuse to round, rarely acute or emarginate; Syst. Verz. 1: 4. 1912; Merr., Enum. Philipp. Fl. Pl. midrib depressed on upper surface in channel about 3:79. 1923; Craib, Fl. Siam. 1: 120. 1925; Kaneh., ¾ of lamina length, underside raised, secondary Fl. Micron. 234, fi g. 106 1933; Merr., Trans. Amer. veins present, dense and distinct on both surfaces, Philos. Soc. 24(2): 269. 1935; Kaneh., Formosan 6–8 veins per 0.5 cm. Infl orescence axillary panicles Trees. Rev. ed.: 473. fi g. 433. 1936; Sastri et al., 5–10 cm long, pubescent or glabrous, up to 12 fl owers Wealth India 2; 18. 1950; Heyne, Nutt. Pl. Ned.-Ind. per infl orescence, pedicels 0.6–4 cm long. Floral ed. 3, 1: 1083. 1950; M.R.Hend. & Wyatt-Sm., parts tepals 8, refl exed when fl ower is fully open, Gard. Bull. Singapore 15: 314. pl. 1C. 1956; the outer pair oval to suborbicular, glabrous, imbricate, N.Robson in Exell & Wild, Fl. Zambes. 1: 394. t. the next pair elliptic and the 2 inner pairs elliptic to

SW 11611-p162-216-G8.indd 170 1/16/62 BE 5:22 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 171 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

obovate. Stamens in fascicles but not distinct fascicles, Kasem 1453 (BK)]; Songkhla [Prince of Songkla 40–250 stamens per fl ower; fi laments 4–5 mm long, University Campus, Hat Yai, 16 June 1983, Warapohn white to yellow-green; anthers 1–1.5 mm long, bright 16 (A, PSU); Kho Hong Hill, Hat Yai, 3 Oct. 1991, yellow/orange. Ovary subglobose, 1.5–3.5 mm long; Songsiri 17 (PSU); PSU Campus, Hat Yai, 25 Dec. style 5–7 mm long with fl at stigma, almost white, 1979, Congdon & Hamilton 4899 (A, PSU); Ko Kham 8–9 mm long; 1 ovule, red. Fruit globose, apex round, Island, Chana, Sirirugsa PS425 (PSU)]; Unknown 2.5–3.2 × 3 cm. location [Kerr 24866 (BKF)]. Thailand — (Map 1.1.3): NORTHERN: Chiang Distribution.— widespread from East Africa Mai [27 June 1914, Kerr 3265 (ABD, BM, E, K)]; to Australia. Phayao [Kwan Phayao, 3 July 1994, BGO Staff 1108 Ecology and phenology.— Evergreen and (QBG)]; Phrae [1 July 1911, Vanpruk 268 (BKF, deciduous forests, marshy ground, sandy and rocky K)]; NORTH-EASTERN: [15 June 1932, shorelines; from near sea level to 300 m. Flowering: Lakshnakara 946 (ABD, BK, BM, K, TCD)]; October–December. Fruiting: all year. Visited by SOUTH-WESTERN: Kanchanaburi [19 Jan. 1907, B.S. bees. 3900 (BK)]; CENTRAL: Saraburi [3 Mar. 1987, Pukhae, 14 km from North of Saraburi, in direction of Lopburi, Vernacular.— Kra thing (กระทิง); saraphi thale Lambinon 87/088 (AAU)]; Krung Thep Maha Nakhon (สารภีทะเล); thing (ทิง) (Sangkaew, 1999). [beside the library, Kasetsart University, Bang Khen, Uses.— This species is used for various purposes 26 Nov. 1970, Watdahnahsahp 16 (A, L, PSU); 28 due to its high durability. The wood is used in May 1922, Marcan 849 (ABD, BM); 3 Apr. 1922, underwater construction and for furniture making. Paknam, Marcan 987 (BM); 13 July 1920, Kerr The seed oil is used in soap making and as medicine 4344 (ABD, BM, K, TCD); Kasetsart University to treat conditions such as rheumatism and skin Campus, Bang Khen, 22 Apr. 1975, Maxwell 75-430 infections. The plant also has many other medicinal (AAU, BK); Paknam, July 1924, Kerr s.n. (BM); uses due the presence of saponins and hydrocyanic Royal Palace, 21 Oct. 1960, Smitinand s.n. (BKF)]; acid (Sangkaew, 1999; Stevens, 1980). SOUTH-EASTERN: Chon Buri [14 Aug. 1981, Apiom et al. 9 (KKU); seaside, 22 Jan. 1955, Conservation.— IUCN Global Status: Least Chotmaiwee 28 (BK): Ko Kram Island, Pattaya, East Concern (LC); Regional (Thailand) Status: Least Coast, 22 Aug. 1923, Collins 942 (ABD, K, TCD); Concern (LC). Kasetsart University, Sri Racha Campus, 22 Dec. 2004, Maxwell 04-827 (CMUB)]; Rayong [Chak 4. Calophyllum macrocarpum Hook.f., Fl. Brit. Phong, Klaeng, 29 June 2004, Kertsawang 319 Ind. 1: 273. 1874; King, J. Asiat. Soc. Bengal, Pt. 2, (QBG); Phae Arboretum, Phae District, 23 Nov. Nat. Hist. 59: 179. 1890; Vesque in DC., Monogr. 1979, Shimizu et al. T-232977 (L)]; Trat [Ko Kut SW, Phan. 8: 603. 1893; Ridl., J. Straits Branch Roy. 21 Nov. 1970, Charoenphol et al. 5111 (AAU, E)]; Asiat. Soc. 33: 48. 1900; Ridl., Fl. Malay Penins. Ko Kradat, 15 Feb. 1900, Schmidt 5466 (C); 1928; 1: 187. 1922; M.R.Hend., Gard. Bull. Straits PENINSULAR: Chumphon [Hat Sai Ri, 18 Feb. 1968, Settlm. 4: 224. 1928; M.R.Hend. & Wyatt-Sm., Vachanapong 64 (BK); Saek Island, Parlenam, 9 Gard. Bull. Singapore 15: 317. 1956; Kochummen, June 1969, Jaram 1677 (BM)]; Surat Thani [Ko Tao, Malayan Forest Rec. 2(17): 218. 1965; Burkill, 1 Jan. 1927, Kerr 11217 (ABD, BK, BM, K)]; Dict. Econ. Prod. Malay Penin. 2(1): 416. 1966; Phangnga [North of Thung Maphrao, 19 July 1972, Meijer, Bot. Bull. Herb. Forest Dept., Sabah 7:16. Larsen et al. 31115 (AAU, K, L, P)]; Phuket [23 1967; Whitmore, Tree Flora of Malaya 2: 187. May 1982, Ubolchalaket 111 (P)]; Phatthalung [Oct. 1973; H.Keng, Gard. Bull. Singapore 28: 244. 1915, Vanpruk 772 (BKF, K)]; Satun [Tarutao, near 1976; Corner, Gard. Bull. Singapore Suppl. 1: 104. Malaeca Creek, 20 Oct. 1979, Congdon 115 (A, PSU); 1978; P.F.Stevens, J. Arnold Arbor. 61: 452. fi g. 28, west side of Pulau Adang, 20 Oct. 1979, Congdon g, h. 1980. Type: Malaysia, Malacca, Maingay 115A (P); Tarutao NP, Ao Son, 10 Nov. 1979, 1728 (lectotype K [K000199888! & K000199889! Congdon 1477 (A, AAU, PSU); La-ngu, Ko Pulon Mai – a single specimen on two sheets], designated by Phai, 10 Apr. 2003, Phengklai et al. 14760 (BKF); Stevens [1980]). Tarutao Island, 20 Apr. 1969, Chermsirivathana &

SW 11611-p162-216-G8.indd 171 1/16/62 BE 5:22 PM 172 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Trees 25–35 m tall; dbh up to 110 cm; bole Conservation.— IUCN Regional (Thailand) straight, usually without buttresses; outer bark Status: Data Defi cient (DD). deeply fi ssured, fi ssures boat-shaped, yellow-brown to dark-brown to black, some areas of bark emitting 5. Calophyllum pisiferum Planch. & Triana, Ann. a reddish hue, inner bark brown to black; exudate Sci. Nat., Bot. Sér. 4, 15: 294. 1862; Ridl., Fl. Malay the colour of clear honey. Branchlets flattened, Penins. 1: 184. 1922; M.R.Hend. & Wyatt-Sm., 4-angled, slight brown pubescence at node, inter- Gard. Bull. Singapore 15: 345. 1956; Whitmore, nodes 0.5–4 cm long. Leaves petioled, petiole Tree Flora of Malaya 2; 172. 1973; P.F.Stevens, J. 1.5–3.2 cm long; blade 8–16.5 × 3–4.6 cm, elliptic Arnold Arbor. 61: 518. fi g. 32, k, l. 1980. Type: to oblanceolate to lanceolate, concave adaxially, Malaysia, Malacca, Gaudichaud 866 (lectotype G convex abaxially, coriaceous, very smooth, light [G00032033!], designated by Stevens [1980]; green to khaki green, base cuneate to acute, apex isolectotypes P [P01900978!, P01900979!]). acuminate to rarely acute; venation pronounced on both surfaces, midrib on upper surface depressed in — C. retusum Wall. ex Choisy var. cochinchinense channel approximately ½ of lamina length, very Pit. in Lecomte, Fl. Indo-Chine 1(4): 321. 1910; pronounced on lower surface, arising at same point Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 275. on both sides of midrib, approximately 0.5 mm apart, 1943; Pham & Nguyên, Cây-Có Mièn Nam Viet-Nam, 5–10 veins per 0.5 cm. Infl orescence axillary; 5–15 179. 1960; Pham, Cây-Có Mièn Nam Viet-Nam. fl owers per infl orescence; covered with red-brown 2(2): 303. 1970. Type: Cochinchine (Vietnam), pubescence; pedicels 0.7–3.3 cm long. Floral parts Thorel 1395 (lectotype P [P01900966!], designated tepals 8 (rarely 10), the outer pair ovate to subor- by Stevens [1980]; isolectotypes P [P01900967!, bicular, 6–7 × 5–6 mm, dorsally densely pubescent; P01900968!], B not seen, K [K000380005!], G inner ones ovate to elliptic to oblong-elliptic,7–17.5 [G00032033!]) × 3–8 mm, dorsally pubescent; the next 2 with the — Calophyllum retusum Wall. ex Choisy var. cam- same shape and size, oblong-elliptic, 6–13 × 2–3 mm, bodgense Pit. in Lecomte, Fl. Indo-Chine 1(4): 321. dorsally slightly pubescent. Stamens in fascicles, 1910; Gagnep. in Humbert, Suppl. Fl Indo-Chine 160–200 per fl ower; fi lament 3–6 mm long; anther 1:275. 1943; Pham, Cây-Có Mièn Nam Viet-Nam. 1–1.3 mm long. Ovary 2–3 mm long; style 3–5 mm 2(2): 303. 1970. Type: Cambodia, Hahn 866 (holotype: long, stigma peltate. Fruit ellipsoid; 6.5–15 × 4.5–8 P! [P01900969!]). cm; acute at apex; yellow-green when ripe, drying — Calophyllum sangkae Craib, Bull. Misc. Inform. wrinkled with longitudinal striations and dark brown. Kew 1925: 18. 1925; Craib, Fl. Siam. 1: 122. 1925. Thailand — (Map 1.1.4): PENINSULAR: Nakhon Type: Thailand, Surin, Kerr 8283 (lectotype K Si Thammarat [Krung Ching Waterfall, 21 Feb. [K000380004!], designated here; isolectotypes: 1991, Niyomdham et al. 2250 (BKF)]; Trang [Khao ABD [ABDUH:2/518!], BM [BM000611358!], Chong, 30 Apr. 1969, Phusomsaeng & Smitinand BK!, P [P00526175!]). 240 (BK, K, L, P); 15 June 1973, Phusomsaeng et — Calophyllum motleyi Ridl., Bull. Misc. Inform. al. 1617 (BKF, L)]; Narathiwat [Khao Sam Sip, 25 Kew 1938:122. 1938. Type: Indonesia, Borneo, Sept. 1996, Niyomdham 4792 (BKF)]. Motley 865 (holotype K [K000380013!]). Distribution — Malay Peninsula to Borneo — Calophyllum retusum auct. non Wall. ex Choisy: excluding Java. T.Anderson in Hook.f., Fl. Brit. India 1:272. 1874, Ecologyy and phenology.— Evergreen rainforest; pro parte; Pierre, Fl. Forest. Cochinch. 1: pl. 102. from 100 to 220 m elevation. Flowering: May–July. 1865; Vesque, Epharmosis 2: t. 25. 1889; King, J. Fruiting: August–June. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 50: 176. 1890, pro majore parte. Vernacular.— Chuat (ชวด) (Phusomsaeng & Smitinand 240); tang hon (ตังหน) (Sangkaew, 1999). Shrubs to 5 m or trees to 35 m tall; dbh 60 cm; bole straight without buttresses; outer bark fi ssured, Uses.— The wood is used in construction and brown-yellow to grey-brown, inner bark dark red; furniture making (Sangkaew, 1999). exudate clear to slightly opaque. Branchlets slightly

SW 11611-p162-216-G8.indd 172 1/16/62 BE 5:22 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 173 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Map 1.1.1 Distribution of Calophyllum calaba in Thailand. Map 1.1.2 Distribution of Calophyllum dryobalanoides in Thailand.

Map 1.1.3 Distribution of Calophyllum inophyllum in Thailand. Map 1.1.4 Distribution of Calophyllum macrocarpum in Thailand.

SW 11611-p162-216-G8.indd 173 1/16/62 BE 5:22 PM 174 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

fl attened to rectangular, covered in minute orange (Leeratiwong 2002-32); pa-ong (ปะอง) (Sangkaew, pubescence; internodes 1–2.5 cm long. Leaves 1999). petioled, petiole 0.3–0.7 cm long; blade 3.5–10.8 × 1–3.3 cm, elliptic or ovate, coriaceous, smooth, Uses.— Branches are used for constructing orange/yellow to brown concave adaxially, convex houses and boat poles (Sangkaew, 1999). abaxially, base round to cuneate, apex acute to round; Conservation.— IUCN Global Status: Least venation distinct on both surfaces, midrib depressed Concern (LC); Regional (Thailand) Status: Near on upper surface and pronounced on lower surface, Threatened (NT). secondary veins not arising at same points of midrib and some splitting again forming intersecondary veins, 6. Calophyllum polyanthum Wall., Numer. List 7–13 veins per 0.5 cm. Infl orescences axillary, race- 4844. 1821, nom. nud.; Wall. ex Choisy, Descr. mose, covered with brown pubescence, 7–15 fl owers Guttif. Inde: 43. 1829; Choisy, Mém. Soc. Phys. per infl orescence, pedicels 0.5–1.5 cm long. Floral Genève 12: 423. 1851; Planch. & Triana, Ann. Sci. parts tepals 4, the outer pair ovate, 4–5 × 2–3 mm, Nat., Bot. Sér. 4, 15: 278. 1862; Bedd., Fl. Sylv. S. dorsally brown pubescent, inner pair elliptic to India 3: 22. 1871; T.Anderson in Hook.f., Fl. Brit. obovate-elliptic, 5–7 × 3–5 mm, slightly dorsally India 1: 274. 1874; Kurz, J. Asiat. Soc. BengaL, Pt. pubescent to glabrous. Stamens in fascicles, 2, Nat. Hist. 43: 88. 1874; Kurz, Forest Fl. Burma approximately 30–70 stamens per fl ower; fi laments 1: 95. 1877; Gamble, List Trees Darjeeling Dist.: 7. 3–3.5 mm long; anthers 0.4–0.6 mm long, oblong- 1878; W.Theob., Burma 2: 636. 1883; Vesque, elliptic. Ovary 1.5–3 mm long; style 4–5 mm long. Epharmosis 2: t. 6. 1889; Vesque in DC., Monogr. Fruit oval to ellipsoid with an acute apex, 0.9–1.3 × Phan. 8: 555. 1893; Prain, Bengal Pl. 1: 246. 1903; 0.7–1 cm, rough ridges on surface, orange when ripe. Brandis, Indian Trees: 54. 1907; A.M.Cowan & Thailand.— (Map 1.1.5): EASTERN: Surin Cowan, Trees N. Bengal: 17. 1929; Craib, Fl. Siam. [Sangka, 14 Jan. 1924, Kerr 8283 (ABD, BK, BM, 1: 121. 1925; Kanjilal et al., Fl. 1:14. 1934; P)]; SOUTH-EASTERN: Prachin Buri [Huai Kasian, Sastri et al., Wealth India 2: 19. 1950; Maheshw., 14 Nov. 1964, Sakol 109 (BK)]; Chanthaburi [East Bull. Bot. Surv. India 2: 144. 1960; Dutt et al., of Makam, 18 Jan. 1958, Sørensen et al. 464 (C)]; Indian Forester 100: 65. 1974; P.F.Stevens, Trat [Ko Kut SW, 21 Nov. 1970, Charoenphol et al. J. Arnold Arbor. 61: 220. 1980; S.Gardner et al., 50977(AAU,BKF, P); ibid., 21 Nov. 1970,Charoenphol Field Guide Forest Trees N. Thailand: 53. 2000; et al. 51066 (AAU, BKF, C, K); Ko Kut, 6 Apr. 1959, H.W.Li et al., Fl. China 13: 39. 2007. Type: India, Smitinand 5722 (BKF); Khao Saming, 26 Nov. Assam, Wallich 4844 (holotype G not seen (fi de 1924, Kerr 9435 (BK, BM, C, K, L, P); Khao Kuap Stevens, 1980); isotypes BM [BM000611322!], FI (or Panom Tom), 25 Dec. 1929, Kerr 17774 (A, BK, [FI010322!],G [G00355193!],GH [GH00067241!], BM, C, K, L, P); Waterfall, Kudi Island, 9 Apr. 2002, K [K001104055!], P [P00526192!]). Leeratiwong 2002-32 (PSU)]; PENINSULAR: — Calophyllum smilesianum Craib, Bull. Misc. Narathiwat [Pawai, Su-ngai Padi, 25 Aug. 1988, Inform. Kew 1924: 85. 1924; Craib, Fl. Siam. 1: Niyomdham & Ueachirakan 1916 (AAU, BKF, C, K, 122. 1925; Gagnep. in Humbert, Suppl. Fl. Indo- L, P); ibid., 5 Sept. 1988, Niyomdham & Ueachirakan Chine 1:270. 1943. Type: Thailand, Kerr 5792 1935 (AAU, BKF, E, K, L, P); Su-ngai Padi, 10 (lectotype K [K000677427!] second step designated Oct. 1988, Niyomdham 1952 (AAU, BKF, K, L)]. here – first step designated by Stevens [1980]; Distribution.— Laos, Cambodia, southern isolectotypes BM [BM000611353!], E!, P Vietnam, Malaysia and Indonesia. [P00526163!]).— Calophyllum smilesianum Craib Ecology and phenology.— Evergreen forest, var. luteum Craib, Bull. Misc. Inform. Kew 1924: peat swamp forest, by water, in rocky areas from 86. 1924; Craib, Fl. Siam. 1: 122. 1925; Gagnep. in near sea level to 1700 m elevation. Flowering: Humbert, Suppl. Fl. Indo-Chine 1: 271. 1943. Type: October–January. Fruiting: November–April. Thailand, Doi Pahom Pok, Kerr 5180 (lectotype K not seen, designated by Stevens [1980]; isolecto- Vernacular.— Katanghan nam (กะตังหันนํ้า) types ABD [ABDUH:2/519!], BK!, BM (Kerr 9435); ka thang han bai lek (กะทังหันใบเล็ก) [BM000611354!]).

SW 11611-p162-216-G8.indd 174 1/16/62 BE 5:23 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 175 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

— Calophyllum williamsianum Craib, Bull. Misc. Feb. 1998, Konta & Phengklai 4003 (BKF); Chom Inform. Kew 1924: 86. 1924; Craib, Fl. Siam.1: 122. Thong District, Doi Hua Sua, 17 Dec. 1998, Konta 1931; Gagnep. in Humbert, Suppl. Fl. Indo-Chine et al. 4673 (BKF); Doi Tiu, 8 Mar. 1921, Kerr 5038 Suppl. 1: 270. 1943. Type: Thailand, Doi Tiu (Nan), (ABD, BM, E, K, P); Doi Pahom Pok, Muang Fang, Kerr 5038 (lectotype K! [bar code not available], 1 Apr. 1921, Kerr 5180 (ABD, BK, BM, K); Doi designated by Stevens [1980]; isolectotypes BK Inthanon, Mae Cham, 19 Oct. 1988, Santisuk 6821 [BK255126!], E [E00209520!], P [P00526164!]). (BKF); Doi Inthanon NP, 27 July 1988, Phengklai et al. 7049 (BKF, C, P); ibid., 28 July 1988, Trees 6–40 m tall; dbh 150–400 cm; bole Phengklai et al. 7089 (BKF); ibid., 28 July 1988, straight without buttresses; outer bark slightly Phengklai et al. 7105 (BKF); ibid., 28 July 1988, fi ssured, brown-grey/yellow; exudate the colour of Phengklai et al. 7155 (AAU, C, K, P); ibid., 2 Nov. clear honey. Branchlets fl attened, sometimes terete, 1962, Smitinand et al. 7708 (BKF); ibid., 10 Nov, brown, slightly pubescent. Leaves petioled, petiole 1962, Smitinand et al. 77877 (BKF); Doi Inthanon, 1.2–2.5 cm long, slightly pubescent to glabrous; Khum Mae Wang, 7 Feb. 1998, Phengklai et al. blade 3–19 × 1.8–6.5 cm, round to obovate-oblong, 10881 (BKF); Doi Inthanon, Doi Hua Suea, 17 Dec. yellow to dark brown, base attenuate to cuneate, 1998, Phengklai et al. 112977 (BKF); west side of apex acuminate to acute, lateral veins distinct on Doi Inthanon NP, 6 Aug. 1988, Fukuoka T-62568 both surfaces, very slender, radiating at same points (BKF); Doi Inthanon NP, 8 Dec. 1998, Kanzaki & from both sides of the midrib and fading near the Kwanchai C654 (CMUB); Doi Suthep-Pui, 30 Mar. edges of the leaves, midrib pronounced on lower 2004, Maxwell 04-175 (CMUB); Doi Suthep NP, surface and faint on upper surface, depressed in near Chang Khian Village, 13 June 1996, Gardner channel on upper surface about ⅔ of the lamina & Kopachan s245b1 (BM, CMUB); Mae Taeng, length, brown/black on upper surface and orange/ Doi Hua Chang, 19 Mar. 1990, Smitinand 90-38 (E, yellow on lower surface. Infl orescence terminal and K, P); Forest station to the top, Doi Chang, Mae from upper leaf axils, covered in brown pubescence, Taeng District, 23 Oct. 1979, Shimizu et al. T-20574 pedicels 1–1.8 cm long. Floral parts tepals 8, the outer (A, AAU, BKF, L)]; Nan [Doi Phu Kha, Pua, 2 July pair oval to suborbicular, 3–4 × 3–3.5 mm, dorsally 1999, Srisanga et al. 816 (QBG); ibid., 26 Feb. grey-brown pubescent, inner ones ovate, 5–8 × 1997, Pooma et al. 14466 (A, BKF, CMUB); ibid., 4.5–5.5 mm, dorsally slightly pubescent and densely 26 Sept. 2000, Srisanga 1725 (QBG); ibid., 22 Aug. so towards base, the next 2 with the same shape and 2001, Srisanga 2021 (QBG); ibid., 15 May 1992, size, ovate, 6–6.5 × 5–5.5 mm, slightly pubescent Santisuk s.n. (BKF)]; Phitsanulok [Foothill of Phu along margin towards base, yellowish-white. Stamens Mieng Mountain, 27 July 1966, Larsen 938 (AAU)]; in fascicles, approximately 200–277 per flower; NORTH-EASTERN: Phetchabun [Thung Salaeng fi laments 2.5–4.5 mm long; anthers 1–1.7 mm long, Luang, 16 Feb. 1968, Phengklai s.n. (BKF)]; Loei oblong, basally connate, orange/brown, centre of [Phu Kradung, Bunpheng 706 (BKF); Kao Keo anther darker. Ovary 2–2.2 mm long; style 3.5–4 mm Kang, 10 Apr. 1922, Kerr 5792 (ABD, BM, E, K, long; stigma peltate. Fruit ovoid to globose, 3–4 × P); Khao Krading, 14 Mar. 1924, Kerr 8752 (ABD, 2–3 cm, acute to round at apex, smooth or fl aking at BK, BM, E, K, P, TCD); Phu Kradung NP, 28 Nov. surface, green or black when ripe. Seeds 35 × 30 mm. 1972, Smitinand 11780 (BKF)]; PENINSULAR: Thailand —(Map 1.1.6): NORTHERN: Mae Chumphon [Kao Tong, 18 Jan. 1927, Kerr 11523 Hong Son [North Mae Hong Son, en route to Doi (A, BK, C, K, L, P)]; Trang [Khao Chong Botanical Chang, 31 May 1977, Santisuk 1148 (A, BKF)]; Garden, 10 Aug. 1993, Smitinand et al. 5 (BKF); Chiang Mai [Doi Ka, 8 Apr. 1925, Winit 1375 (BK, Chong, 25 Jan. 1957, Smitinand 41066 (BKF)]. BKF, K); Mae Taeng, 5 Dec. 1977, Santisuk 1491 Distribution.— India to Southwestern China, (A, C, K, L, P); Doi Pa Kao, Apr. 1912, Kerr 2608 Cambodia and Indonesia. (BM, E, K); Mae Chaem, 9 Sept. 1912, Kerr 2691 (K); Mae Sanga Watershed Management Unit, Mae Ecology and phenology.— Evergreen forest Chaem District, 6 May 2000, Sookchaloem et al. and lower montane forest from 60 to 1950 m elevation. 3340 (BKF); Doi Inthanon, 26 Apr. 1931, Put 3765 Flowering: January–July. Fruiting: May–February. (BK, BM, K, L); eastern side of Doi Inthanon, 6

SW 11611-p162-216-G8.indd 175 1/16/62 BE 5:23 PM 176 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Vernacular.— Pa-ong (พะอง) (Kerr 8752); kho- pair ovate and the inner pair oblong-elliptic. Stamens approximately 50 stamens per flower; anthers mai-do ( ); sa-chum-mun ( ); ma haen คอไหมดอ ซาจุมมุน 0.4–1 mm long. Ovary style 2–2.7 mm long. Fruit doi (มะแหนดอย) (Sangkaew, 1999). ovoid-globose, 8 mm diam., wrinkled, pale-yellow Uses.— Wood is used in construction and to green. furniture making (Sangkaew, 1999). Thailand.— (Map 1.1.7): PENINSULAR: Conservation.— IUCN Regional (Thailand) Narathiwat [Waeng, 17 Apr. 1972, Sangkhachand Status: Least Concern (LC). et al. 10677 (BKF, C, L); Pawai, Su-ngai Padi, 17 Sept. 1987, Niyomdham & Sriboonma 1623 (BKF)]; Unknown location [Unknown collector 57479 (L)]. 7. Calophyllum rupicola Ridl., Trans. Linn. Soc. London, Bot. 3(9): 278. 1893; Ridl., Fl. Malay Distribution.— Northeast Malaya to Sumatra. Penins. 1: 182. 1922; M.R.Hend. & Wyatt-Sm., Ecology and phenology.— Peat swamp forest, Gard. Bull. Singapore 15: 346. 1956; Whitmore, evergreen forest from near sea level to 270 m Tree Flora of Malaya 2: 168. 1973; P.F.Stevens, J. elevation. Flowering: October–November. Fruiting: Arnold Arbor. 61: 515. 1980. Type: Malaysia, December–April. Malacca, Pahang, Ridley 26366 (holotype SING!; isotypes BM [BM00946495!], K!). Vernacular.— Tang hon nam (ตงหั นนํ้า) (Sangkaew, 1999); tang hon bai lek (ตังหนใบเล็ก) (Niyomdham & — Calophyllum rupicola Ridl. variety?: M.R.Hend. Sriboonma 1623) & Wyatt-Sm., Gard. Bull. Singapore 15: 347, pl. 27. 1956, pro parte [not validly published as it is not Conservation.— IUCN: Regional (Thailand) accepted by the authors and is not given a name; see Status: Critically Endangered (CR). McNeill et al. 2012,- Art. 36.1] — Calophyllum rupicola Ridl. var. elatum Whitmore, 8. Calophyllum sclerophyllum Vesque, Epharmosis Gard. Bull. Singapore 26: 270. 1973, pro parte, typo 2: t. 33. 1889; Vesque in DC., Monogr. Phan. 8: 587. haud excluso; Whitmore, Tree Flora of Malaya 2: 1893; Merr., Bibliogr. Enum. Born. Pl.: 394. 1921; 168. 1973, pro parte. Type: ? Malaysia, FRIM 2538 Masam., Enum. Phan. Born.: 476. 1942; Heyne, [K000677416, L0012089, L0012090, SING0055128] Nutt. Pl. Ned.-Ind., ed. 3. 1: 1085. 1950; M.R.Hend. is Calophyllum tetrapterum var tetrapterum. & Wyatt-Sm., Gard. Bull. Singapore 15: 324. 1956; J.A.R. Anderson, Gard. Bull. Singapore 20: 154. Trees 10–35 m tall; dbh 30 cm; without 1963; Pukol & Ashton, Checkl. Brunei Trees: 93. buttresses; outer bark thin, rough with small fi ssures 1964; Ashton, Oxford Forest Mem. 25: pl. 32. 1964; greyish-brown, inner bark pink; exudate milky- Smythies, Common Sarawak Trees 64: pl. 22. yellow (opaque-whitish), sticky. Branchlets terete 1965; Kochummen, Malayan Forest Rec. 2(17): to quadrangular, smooth though striations present 214. 1965; Whitmore, Tree Flora of Malaya 2: 191. on some specimens, glabrous, brown to red/brown, 1973; J.A.R. Anderson, Trees Peat Swamp scars absent. Leaves petioled, petiole 0.4–0.9 cm Sarawak: 88. 1973; Corner, Gard. Bull. Singapore long, blade 5.2–15 × 2.2–3.6 cm, elliptic to elliptic- Suppl. 1: 105. 1978; P.F.Stevens, J. Arnold Arbor. lanceolate to obovate-oval, approximately half of 61: 447. 1980. Type: Borneo, Sarawak, Beccari PB leaves glaucous and other half coriaceous, shiny, 2705 (holotypeO not seen; isotypes FI [FI007648!], base narrowly tapering, apex bluntly acute to K! [K000380015!], M not seen, P not seen). rounded to slightly cuspidate; midrib prominent on — Calophyllum rhizophorum Boerl. & Koord. in both surfaces though fades and disappears near apex Koord.-Schum., Syst. Verz. 2: 40. 1911; J.A.R (approximately 10–20 cm from apex) on some Anderson, Trees Peat Swamp Sarawak, fi g. 26C. specimens, varies from light yellow/green to red/ 1973. Type: Indonesia, Sumatra, Koorders 10333 brown on both surfaces, secondary veins present on (holotype BO not seen). both surfaces, on upper surface some leaves show fading of venation at/near edges of leaves. — Calophyllum subluridum Wyatt-Sm., Malayan Inflorescence axillary, pedicels 0.3–0.8 cm long, Forest Rec. 17: 113. 1952; nom. nud. slender, glabrous. Floral parts tepals 4, the outer

SW 11611-p162-216-G8.indd 176 1/16/62 BE 5:23 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 177 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

— Calophyllum teysmannii Miq. var. inophylloide 9. Calophyllum soulattri Burm.f. ,Fl. Ind.: 121. auct. non (King) P.F.Stevens: Phengklai & 1768; F.Muell., J. Roy. Soc. New S. Wales 24: 174. C. Niyomdham, Flora in Peat Swamp Areas of 1890; F.Muell., Bot. Centralbl. 44: 29. 1891; Merr., Narathiwat, 196. fi g. 108. 1991. Interpr. Herb. Amboin. : 371. 1917; Merr., Philipp. J. Sci. 19: 366. 1921; Merr., Enum. Philip. Fl. Pl. 3: Trees 20–45 m tall; dbh up to 300 cm; outer 81. 1923; Holthius, Blumea 5: 214. 1942; Heyne, bark shallowly fi ssured, scaly pale green to light Nutt. Pl. Ind.-Ned. ed. 3. 1: 1085. 1950; Sastri et al., brown, inner bark fi ssured, soft, pink to red or dark Wealth India 2: 20. 1950, pro parte; M.R.Hend. & orange; exudate the colour of clear honey. Branchlets Wyatt-Sm., Gard. Bull. Singapore 15: 319. 1956; sturdy, thick, terete, dark brown to black, pubescent, Maheshw., Bull. Bot. Surv. India 2: 142. 1960; waxy or glabrous, striations and scars present. Backer & Bakh.f., Fl. Java 1: 386. 1963; Whitmore, Leaves petioled, petiole 1.9–2.8 cm long; blade Guide Forests Brit. Solomon Is.: 77. 1966; Burkill, 6.2–16.7 × 3.1–7.9 cm, obovate, oval or rotund, Dict. Econ. Prod. Malay Penin. 2(1): 416. 1966; thickly coriaceous, base cuneate to slightly attenuate, Whitmore, Gard. Bull. Singapore 22: 15. 1967; apex blunt or mainly retuse; midrib prominent on Meijer, Bot. News Bull. Forest Dept., Sabah 7: 16. both surfaces, fading towards apex on upper surface 1967; Pham, Cây-Có Mièn Nam Viet-Nam. 2(2): for approximately 2/3 of lamina length, dark brown 300. fi g. 1970; Foreman, Check List Vasc. Pl. to black abaxially and light tan to dark brown Bougainville: 42. 1971; Whitmore, Tree Flora of adaxially, both secondary veins and intramarginal Malaya 2; 192. 1973; T.G.Hartley et al., Lloydia veins present and very prominent on both surfaces. 36: 276. 1973; P.F.Stevens, Austral. J. Bot. 22: 399. Infl orescence axillary racemes, pedicels up to 3 cm 1974; P.F.Stevens, J. Arnold Arbor. 57: 175. 1976; long, 9–11 fl owers per infl orescence. Floral parts H.Keng, Gard. Bull. Singapore 28: 245. 1976; tepals 8, the outer pair orbicular to suborbicular, Corner, Gard. Bull. Singapore Suppl. 1:105. 1978; dorsally brown pubescent, the next pair orbicular to Perry, Med. Pl. E. SE. Asia, 174. 1980; P.F.Stevens, suborbicular, the next two pairs obovate. Stamens J. Arnold Arbor. 61: 277. 1980. Type: Indonesia, 200–360 per fl ower; anthers 1.3–1.8 mm long. Ovary Java, Burman s.n. (holotype: G [G00032034!]). style 4.5–6.5 mm long. Fruit ovoid to rotund with acute or round apex, 2.4 × 1.6 cm;, surface very — Calophyllum lanceolatum Warb., Bot. Jahrb. Syst. fi nely wrinkled, drying dull reddish brown to almost 13: 381. 1891; nom. illeg, ≡ Calophyllum warburgii black. Engl. & Prantl, Nat. Pflanzenfamilien 3(6): 221. 1893; Lauterb., Nova Guinea Bot. 8: 843. 1912; Thailand.— (Map 1.1.8): PENINSULAR: Lauterb., Bot. Jahrb. Syst. 58: 13. 1922; A.C.Sm., Narathiwat [Bang Kuntang, Tak Bai, 7 July 1983, J. Arnold Arbor. 22: 345. 1941, pro majore parte. Niyomdham 654 (BKF); Pa Ye, Su-ngai Padi, 14 Type: Indonesia, Kei Inseln, Warburg 20048 (lecto- Apr. 1988, Niyomdham & Ueachirakan 1815 (BKF); type: US [US00114158!], designated here; isolecto- Bang Nara, 22 Aug. 1973, Smitinand 11960 (BKF)] types: A [A00067341!, A00067342!], MICH Distribution.— Malay Peninsula to Borneo [MICH1115518!], LAE not seen). excluding Java. — Calophyllum hibbardii Elmer, Leafl . Philipp. Bot. Ecology and phenology.— Peat Swamp Forest 2: 503. 1908. Type: Philippine Islands, Elmer 9837 at near Sea Level. Flowering: July–August. Fruiting: (lectotype L [L0011778!], designated here; isolecto- (April) August–December. types A [A00067335!], BM [BM000611478!], BO Vernacular.— Ya-kang ( ); kra thing phru not seen, E [E00346383!], F [F0057186F!], FI ยะกัง [FI010315!], G not seen, HBG [HBG517738!], K (กระทิงพรุ) (Sangkaew, 1999). [K000677476!], LY not seen, MO not seen, NY Uses.— A good timber species used in general [NY00072325!], US not seen, W not seen). construction (Sangkaew, 1999). — Calophyllum paludosum C.T.White, J. Arnold Conservation.— IUCN Regional (Thailand) Arbor. 31: 98. 1950; Whitmore, Guide Forests Brit. Status: Critically Endangered (CR). Solomon Is. 77. 1966; Whitmore, Gard. Bull. Singapore 22: 14. 1967. Type: Solomon Islands,

SW 11611-p162-216-G8.indd 177 1/16/62 BE 5:23 PM 178 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Map 1.1.5 Distribution of Calophyllum pisiferum in Thailand. Map 1.1.6 Distribution of Calophyllum polyanthum in Thailand.

Map 1.1.7 Distribution of Calophyllum rupicola in Thailand. Map 1.1.8 Distribution of Calophyllum sclerophyllum in Thailand.

SW 11611-p162-216-G8.indd 178 1/16/62 BE 5:23 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 179 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

BSIP (Walker & White) 192 (holotype BRI [BRI- [Tarutao Island, Talo-Oo-Dang, 27 Apr. 1981, AQ0340217!]; isotypes A [A00067323!], CANB Congdon 1261 (A, AAU, PSU)]; Songkhla [Ton [CANB189036!], K [K00038002!], LAE not seen, Nga Chang Reserve, Hat Yai, 27 Aug. 1981, Sirirugsa MEL [MEL75509!]). 442 (PSU); ibid., 2 Aug. 1985, Maxwell 85-7666 (A, — Calophyllum wallichianum auct. non Planch. & AAU, BKF, L, PSU)] Triana: Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. Distribution.— Cambodia, Vietnam, Malaysia 45:119. 1876; Theobald, Burma 2: 637. 1883; to Australia. Maheshw., Bull. Bot. Surv. India 2: 146. 1960. Ecology and phenology.— Primary evergreen Trees 10–20 m tall; dbh up to 30 cm; bole forest, by water, peat swamp area, found on granitic straight, with/without buttresses, if present, small; bedrock from 50 to 500 m elevation. Flowering: outer bark shallowly fi ssured, almost smooth, light March–October. Fruiting: all year. brown, heartwood dark brown; exudate white. Vernacular.— Tang hon loko ( ) (Kerr Branchlets fl attened, with reddish-brown pubescence, ตังหนโละโกะ internodes 3–10cm long. Leaves petioled, petiole 19035); tang hon bai yai (ตังหนใบใหญ) (Sangkaew, 1.1–2.3 cm long; blade 6–30 × 3–7.5 cm, oblong-ovate 1999). to elliptic-ovate, coriaceous, yellow-brown, sparsely Uses.— Used for masts and spars and in house hairy to almost glabrous, base cuneate to slightly construction (Stevens ,1980). acute, apex acute to acuminate; venationn very obvious, Conservation.— IUCN Global Status: Least arising at same point from both sides of the midrib, Concern (LC); Regional (Thailand) Status: Least midrib wider and more apparent on lower surface, Concern (LC). depressed in channel on upper surface for about ½ of lamina length, dark green above and pale green underneath, immature leaves with light brown midribs. 10. Calophyllum symingtonianum M.R.Hend. & Infl orescence axillary, in cymes, sometimes appearing Wyatt-Sm., Gard. Bull. Singapore 15: 338. pl. 18. paniculate, 11–29 fl owers per infl orescence, covered 1956; Kochummen, Malayan Forest Rec. 2(17): with brown pubescence, pedicels 1–3 cm long. Floral 222. 1965; Whitmore, Tree Flora of Malaya 2: 192. parts tepals 4, the outer pair oval to suborbicular, 1973; P.F.Stevens, J. Arnold Arbor. 61: 229. 1980. 3–4.5 × 4.5–5 mm, glabrous or slightly pubescent Type: Malaysia, Malacca, Pahang, Nur SFN 32633 at margin, inner ones obovate, 5–6.5 × 5–6 mm, (holotype SING [SING0055134!]; isotypes A glabrous or slightly pubescent at margin, tepals, [A00067289!], K [K000677413!], KEP not seen, fi laments and style white. Stamens in fascicles, 59–97 L [L0012087!], MO not seen, P!, S [S11-34387!], stamens per fl ower; fi laments 3–5 mm long; anthers UC not seen, US [US00114173!]). 1–2.6 mm long, white or yellow. Ovary 2–2.5 mm Trees to 40 m tall; up to dbh 300 cm; without long; style 3–5 mm long. Fruit ovoid to spherical, buttresses; outer bark fi ssured, rough, greyish-brown 0.8–1 × 0.7–1 cm, round at apex, stalk 2–2.9 cm to yellowish-brown, inner bark pink; exudate the long, purple to black. colour of clear honey, sticky. Branchlets terete, Thailand — (Map 1.1.9): SOUTH-WESTERN: glabrous, smooth, thin, dark brown to greyish. Kanchanaburi [Thung Phra Ruesi, 5 May 1992, Leaves petioled, petiole 1–1.3 cm long; blade, Smitinand 255 (BKF)]; PENINSULAR: Chumphon 6.3–9.4 × 2.4–3 cm, elliptic to lanceolate, slightly [Bang Son, 13 Mar. 1928, Put 1562 (BK, BM, K); coriaceous, smooth, green to red/brown, base tapering Tongtapai Sara, 13 Apr. 1967, Sutheesoen 2204 to acute, apex acuminate; midrib discrete on both (BKF)]; Krabi [Khao Pra-Bang Khram Wildlife surfaces, slightly more prominent on lower surface, Sanctuary, 14 Jan. 2006, Maxwell s.n. (CMUB)]; light green to brown on upper surface and pale green Nakhon Si Thammarat [Phrom Lok Falls, Khao to yellow on lower surface, secondary veins present, Luang NP, Phrom Khiri District, 24 Feb. 1987, arising from same points on both sides of the midrib, Maxwell 87-194 (A, BKF, L, P, PSU)]; Phatthalung Infl orescence terminal and from upper leaf axils, [Tamot Falls NP, Tamot, 9 Aug. 1986, Maxwell 86- some with pale brown pubescence. Floral parts 5466 (A, AAU, BKF, L, PSU)]; Trang [Bangsak, 19 tepals 8; outer pair of tepals dorsally glabrous or Apr. 1930, Kerr 19035 (A, BK, BM, K, L, P)]; Satun (occasionally) dorsally slightly pubescent towards

SW 11611-p162-216-G8.indd 179 1/16/62 BE 5:23 PM 180 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

base. Stamens 46–70 per fl ower. Fruit ellipsoid to Prod. Malay Penin. Ed. 2. 1: 412. 1966; Whitmore, slightly obovoid, apex acute to round, 2.5 cm long, Tree Flora of Malaya 2: 180. 1973; Corner, Gard. glabrous and wrinkled, dull reddish brown to dark Bull. Singapore, Suppl. 1: 104. 1878. Type: Malaysia, purple when dry. Malacca, Maingay 1660 (lectotype K [K000199894!], Thailand.— (Map 1.1.10): Trang, Songkhla, designated by Stevens [1980]). Yala (locations taken from Sangkaew, 1999), — Calophyllum prainianum King, J. Asiat. Soc. Unknown location [No collector name or number Bengal, Pt. 2, Nat. Hist. 59: 175. 1890; Vesque in (BKF)]. DC Monogr. Phan. 8: 550. 1893; Ridl., Fl. Malay Distribution.— Malay Peninsula. Penins. 1: 183. 1922. Types: Malaya, Perak, Larut, King’s Collector [Kunstler] 5366 (syntypes BM Ecology and phenology.— Evergreen forest [BM000946498!], FI not seen, G not seen, K!, P from 460 m elevation. Flowering: unrecorded. [P01900949!], UC not seen), King’s Collector Fruiting: February–April. [Kunstler] 7243 (syntypes BM [BM000946496!], Vernacular.— Tang hon (ตังหน) (Sangkaew, FI not seen, K [K000677415!], L [L0012091!], P!). 1999). — Calophyllum venustum King, J. Asiat. Soc. Uses.— Hard wood used to build houses Bengal, Pt. 2, Nat. Hist. 59: 180. 1890; Vesque in DC. (Sangkaew, 1999). Monogr. Phan. 8: 549. 1893; Ridl., Fl. Malay Penins. 1: 186. 1922. Type: Malaysia, Malacca, Perak, Larut, Conservation.— IUCN Regional (Thailand) King’s Collector [Kunstler] 7763 (lectotype L Status: Data Defi cient (DD). [L0012092!], designated here; isolectotypes FI not seen, G not seen, K [K000677414!]). 11. Calophyllum tetrapterum Miq., Pl. Jungh.: — Calophyllum foetidum Ridl., J. Straits Branch 291. 1854; P.F.Stevens, J. Arnold Arbor. 61: 505. Roy. Asiat. Soc. 54: 18. 1910; Fl. Malay Penins. 1: 1980. Type: Indonesia, Sumatra, Junghuhn s.n. 186. 1922. Type: Singapore, Ridley 11958 (lectotype (holotype U!; isotypes BO!, L!). SING [SING0055098!], designated by Stevens — Calophyllum bancanum Miq., Fl. Ned. Ind., [1980]: isolectotypes BM!, K!). Eerste Bijv. 1(3): 499. 1861; Kurz, Natuurk. Tijdschr. — Calophyllum lanceola Ridl., J. Straits Branch Ned.-Indië 27: 192. 1864; F.Muell. in Walp. Ann. Roy. Asiat. Soc. 82:170. 1920; Ridl., Fl. Malay Syst. Bot. 7: 357. 1868; Scheff er, Natuurk. Tijdschr. Penins. 1: 82. 1922. Type: Malaysia, Malacca, Ridley Ned.-Indië 31: 354. 1870, 32: 405. 1873. Type: 5751 (holotype SING!). Indoneaia, Bangka, Teysmann HB 3214 (holotype U [U0002361!]; isotypesBO not seen, K [K0006774557!], — Calophyllum rupicola Ridl. var. elatum Whitmore, L [L0012098!]). Gard. Bull. Singapore 26: 270. 1973; Whitmore, Tree Flora of Malaya 2: 169. 1973. Type: Malaya, — Calophyllum gracile Miq., Fl. Ned. Ind., Eerste Malacca, Kelantan, FRI (Kochummen) 2538 (holo- Bijv. 1(3): 498. 1861; F.Muell. in Walp. Ann. Syst. type KEP not seen, isotypes K [K000677416!], L Bot. 7: 357. 1868.— Calophyllum pulcherrimum [L0012089!, L0012090!], SING [SING0055114!]). Wall ex. Choisy var. gracile (Miq.) Boerl., Cat. Hort. Bogor. 2:82. 1901. Type: Indonesia, Sumatra, — Calophyllum globuliferum Ridl., Bull. Misc. Teysmann HB 649 (holotype U; isotypes BO not Inform. Kew 1938: 121. 1938. Type: Indonesia, seen, K [K000677459!], L!, MEL not seen). Borneo, Kalimantan, Bangarmassing, Motley 618 (holotype K [K000380016!]). — Calophyllum fl oribundum Hook.f., Fl. Brit. India 1: 272. 1874; King, J. Asiat. Soc. Bengal, Pt. 2, Nat. — Calophyllum dryobalanoides auct. non Pierre: Hist. 59: 175. 1890; Ridl., Fl. Malay Penins. 1:184. Craib, Fl. Siam. 1: 120. 1925; Gagnep. in Humbert, 1922; M.R.Hend., Gard. Bull. Straits Settlem. 4:224. Suppl. Fl. Indo-Chine, 1: 274. 1943, pro parte. 1928; M.R.Hend. & Wyatt-Sm., Gard. Bull. — Calophyllum pulcherrimum auct. non Wall. ex Singapore 15: 332. pl.16. 1956; Smythies, Common Choisy: T.Anderson in Hook.f., Fl. Brit. India 1: Sarawak Trees 61. 1965; Kochummen, Malayan 271. 1874; Pierre, Fl. Forest. Cochinch. 1: pl. 104A. Forest Rec. 2(17): 215. 1965; Burkill, Dict. Econ. 1885, pro parte; Vesque, Epharmosis 2: t. 21. 1889;

SW 11611-p162-216-G8.indd 180 1/16/62 BE 5:23 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 181 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Vesque in DC. Monogr. Phan. 8: 570. 1893, pro Kaeom, Sawi, 14 Apr. 1967, Sutheesoen 22277 (BK)]; parte; Curtis, J. Straits Branch Roy. Asiat. Soc. 25: Ranong [Khuraburi, Ko Phra Thong, 4 Apr. 2003, 78. 1894; Pit. in Lecomte, Fl. Indo-Chine 1(4): 321. Phengklai et al. 13729 (BKF)]; Surat Thani [Khao 1910; Ridl., Fl. Malay Penins. 1: 182. 1922, pro Kaup, 25 Dec. 1929, Kerr 17772 (BK, BM, K, L)]; parte; Burkill & M.R.Hend., Gard. Bull. Straits Krabi [Khao Pra-Bang Khram Wildlife Sanctuary, Settlem. 3: 347. 1925; Craib, Fl. Siam. 1: 121. 1925; 17 Jan. 2006, Maxwell 06-655 (CMUB); ibid., 25 Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 274. Mar. 2006, Maxwell 06-201 (CMUB)]; Ao Luek, 1943; Pham & Nguyên, Cây-Có Mièn Nam Viet- 15 Mar. 1930, Kerr 185677 (A, BK, BM, C, K, P); Nam, 179. 1960; Pham, Cây-Có Mièn Nam Viet- Trang [Khao Chong, 30 Apr. 1969, Phusomsaeng Nam. 2(2): 301. fi g. 1970. & Smitinand 240 (AAU); Khlong Mayom, Salak Khok, Ko Chang, 6 Apr. 1923, Kerr 6930 (BKF, Trees to 25 m tall; dbh 30 cm; bole straight, K)]; Satun [Rawi Island, 15 June 1980, Congdon with or without small buttresses; outer bark coarsely 631 (A); Tarutao NP at waterfall above Choe-lae but shallowly fi ssured or smooth, grey-brown to dark Village east side of Adang, 19 Oct. 1980, Congdon grey-black, inner bark soft, pink; exudate the colour 951 (AAU, PSU); Tarutao, 20 Jan. 1928, Kerr 14213 of clear honey. Branchlets terete to fl attened, slender, (BK, BM, C, K, L, P)]; Songkhla [Ton Nga Chang glabrous, internode, 1–3 cm long, brown/red/grey. Reserve, Hat Yai, 30 July 1983, Srirugsa 592 (PSU); Leaves petioled, petiole 0.5–1.5 cm long; blade ibid., 10 Jan. 1984, Srirugsa 7377 (PSU); ibid., 18 3.5–13.2 × 1.4–4 cm, obovate to elliptic, slightly Apr. 1985, Maxwell 85-4177 (A, AAU, BKF, E, L, glaucous, red-brown, some green/yellow and concave PSU)]; Khlong Hoi Khong, Hat Yai, Maxwell 85-65 adaxially, convex abaxially, coriaceous, base cuneate (P); Narathiwat [Waeng, 30 June 1972, Nitrasirirak to attenuate, apex acute to acuminate, midrib not 206 (BKF); Bala-Hala, Waeng, 3 May 1997, very pronounced on either surface, depressed in Puudjaa 573 (BKF); Waeng, Sangkhachand et al. channel about ½ of lamina length on upper surface, 1067 (C)]; Unknown location [Hui Tonpong, 25 underside raised, secondary veins present, prominent Mar. 1920, Punyabukkana 874 (ABD, BKF, K)]. on both surfaces, dense, arising roughly from the same place on both sides of the midrib, spacing of Distribution.— Cambodia, Vietnam, Malay veins varies from leaf to leaf, approximately 6–17 Peninsula to Indonesia excluding Java. veins per 0.5 cm, intramarginal veins absent. Ecology and phenology.— Evergreen forest, Infl orescence axillary, 3–11 fl owers per infl orescence, tropical rainforest, water areas, peat areas, savannah covered with brown pubescence, pedicels 0.5–2.5 cm from near Sea Level to 700 m elevation. Flowering: long. Floral parts tepals (4–)8, sometimes 5, 6, 7 or all year. Fruiting: January–May (October). 8, outer pair ovate, oval to suborbicular, 3–5 × 3–4 mm, glabrous to slightly pubescent along margin Vernacular.— Tang hon (ตังหน) (Sangkaew, and apex, next pair obovate or ligulate, 4–8 × 3–5 mm, 1999). glabrous, the next 2 pairs obovate to spathulate, 5–7 Uses.— Young leaves are eaten as a vegetable × 2–3 mm, or the innermostt ones oblong-oblanceolate, (Sangkaew, 1999). 5–6 × 1–1.5 mm, glabrous. Stamens in fascicles, Conservation.— IUCN Global & Regional 36–86 stamens per fl ower, fi laments, 2–5 mm long, (Thailand) Status: Least Concern (LC). anthers, 1–1.3 mm long, oblong or elliptic-oblong. Ovary 1–2 mm long; style 2–4.5 mm long. Fruit ovoid-globose, 0.6–1 × 0.5–0.8 cm, acute to round 12. Calophyllum teysmannii Miq., Fl. Ned. Ind. at apex, dark green when ripe. Suppl. 1(3): 499. 1861; P.F.Stevens, J. Arnold Arbor. 61: 431. 1980. Type: Indonesia, Sumatra, Teysmann Thailand.— (Map 1.1.11): SOUTH-EASTERN: HB 650 (holotype U [U0002364!]; isotypes BO Trat [Ko Chang, Jungle at Khlong Non Si, 14 Feb. not seen, L [L0012088!], P [P01900948!]). 1900, Schmidt 528 (C, K); Khao Saming, 26 Jan. 1927, Put 567 (BK, K, L); Ko Chang, Khlong — Calophyllum intramarginale M.R.Hend. & Mayom, 22 Feb. 1900, Schmidt 603a (C, K); Khlong Wyatt-Sm., Gard. Bull. Singapore 15: 342, pl. 22. Non Si, Ko Chang, 26 Sept. 1924, Kerr 9175 (A, 1956. Type: Malaysia, Malacca, Moysey & Kiah BK, C, E, K, L, P)]; PENINSULAR: Chumphon [Khao SFN 31900 (holotype SING [SING0055110!]; isotypes A [A00067279!], K!, KEP not seen).

SW 11611-p162-216-G8.indd 181 1/16/62 BE 5:24 PM 182 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

— Calophyllum inophylloide King var. singapurense Distribution.—Cambodia, Indonesia, Malaysia. M.R.Hend. & Wyatt-Sm., Gard. Bull. Singapore 15: Ecology and phenology.— Peat Swamp Forest 316, pl. 9. 1956; Kochummen, Malayan Forest Rec. from near sea level to 250 m elevation. Flowering: 2(17): 215. 1965; Smythies, Common Sarawak April–August. (October). Fruiting: (April) July– Trees, 61. 1965; Whitmore, Gard. Bull. Singapore August. 26: 270. 1973; Whitmore, Tree Flora of Malaya 2: 186. 1973; Corner, Gard. Bull. Singapore Suppl. 1: Vernacular.— Tang hon bai neep (ตังหนใบหนีบ) 104. 1978. Type: Singapore, Corner SFN 32518 (Sangkaew, 1999). (holotype SING [SING0055108!]; isotypes K Conservation.— IUCN Regional (Thailand) [K000199910!], KEP not seen). Status: Critically Endangered (CR). Trees 5–35 m tall; dbh 95 cm; bole straight, small buttresses sometimes present, if present up to 13. Calophyllum thorelii Pierre, Fl. Forest. 70 cm tall; outer bark shallowly fi ssured and scaly, Cochinch. 1: t. 103. 1885; Vesque in DC., Monogr. greyish-brown to dark brown, inner bark orange- Phan. 8: 601. 1893; Pit. in Lecomte, Fl. Indo-Chine brown to black; exudate the colour of clear honey. 1(4): 322. 1910; Craib, Fl. Siam. 1: 122. 1925; Branchlets fl attened, slender; with brown or black Gagnep. in Humbert, Fl. Suppl. Indo-Chine 1: 269. pubescence, waxy or glabrous, internodes 0.5–3.5 cm 1943; P.F.Stevens, J. Arnold Arbor. 61: 218. 1980. long. Leaves petioled, petioles 3–12 mm long; blade Type: Cochinchine (Vietnam), Pierre 34 (lectotype 3.5–9 × 1.5–4.5 cm, elliptic-oblong or obovate, P [P01900945!], second step designated here; fi rst otherwise oblong-lanceolate to obovate, concave step designated by Stevens [1980]; isolectotypes adaxially, convex abaxially, coriaceous, red-brown A [A00067243!], BM [BM000611352!], K to khaki green, base acute to cuneate, apex acute to [K000677428!], P [P01900946 !, P01900947 !]). retuse, midrib depressed in channel on upper surface — Calophyllum thorelii Pierre var. oxycarpum about ⅓ of lamina length, underside raised, lateral Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1:270. veins on both surfaces distinct but sometimes obscure, 1943; nom. inval. 6–10 veins per 0.5 cm, intramarginal veins present. Infl orescence racemes, 3–11 fl owers per infl ores- Trees to 30 m tall; dbh up to 35 cm; outer bark cence, usually glabrous but sometimes covered with deeply fi ssured, brownish-yellow, inner bark (slash- brown pubescence, pedicels 0.5–4.3 cm long. Floral mark) pink to reddish-brown; exudate the colour of parts tepals 8, the outer pair oval to suborbicular, clear honey. Branchlets terete, smooth, light to dark 4–5 × 4–4.5 mm, glabrous or slightly pubescent at red-brown, longitudinally ridged, scars present. margin, next pair elliptic, next ones obovate, inner Leaves petioled, petioles 2.2–3 cm long, blade elliptic ones ovate to suborbicular, all 5–8 × 5.5–7 mm, to oblanceolate, occasionally oblong, 8.1–15 × glabrous or slightly pubescentt at margin, white to pale 2.9–4.8 cm, strongly coriaceous, green to light dull yellow, green in bud. Stamens in fascicles, 65–212 brown adaxially, yellow to tan abaxially, base cuneate stamens per fl ower; fi laments, 3–4.5 mm long, white; to gradually tapering, apex cuspidate, occasionally anthers, 1–1.5 mm long, yellow. Ovary 1–2.5 mm acute, leaf edges curling when dry, midrib depressed long; style 2–5 mm long, white, stigma peltate. Fruit and glaucous adaxially, very prominent and yellow- ellipsoid to subglobose, 2–2.5 × 1.8–2 cm, apex tan abaxially, secondary and intramarginal veins acute to round, striate to smooth, yellow-green to present, prominent on both surfaces, 0.8 mm apart, pale brown when ripe. arising from same points on opposite sides of the Thailand.— (Map 1.1.12): PENINSULAR: midrib. Infl orescence terminal and from upper leaf Narathiwat [Pa Ye, Su-ngai Padi, 14 Apr. 1988, axils. Floral parts tepals 8, outer pair ovate to oval, Niyomdham & Ueachirakan 1815 (AAU, C, E, K, L, dorsally pubescent, next pair ovate to obovate-oblong, P); Pha Khluai, Sukhirin, 17 Apr. 1996, Niyomdham next 2 pairs obovate to elliptic-oblong. Stamens & Puudjaa 47177 (BKF); Khao Nakharat, Sukhirin, 124–178 per flower. Fruit spherical to ellipsoid, 20 Oct. 1996, Niyomdham 48366 (BKF); Phru Khok 1.8–2.8 × 1.2–2 cm, light green to purple, stringy Ka La, 18 Aug. 1997, Niyomdham et al. 51277 (AAU, fi bres present inside fruit. BKF); To Daeng, Aug. 1993, Niyomdham s.n. (BKF)].

SW 11611-p162-216-G8.indd 182 1/16/62 BE 5:24 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 183 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Map 1.1.9 Distribution of Calophyllum soulattri in Thailand. Map 1.1.10 Distribution of Calophyllum symingtonianum in Thailand.

Map 1.1.11 Distribution of Calophyllum tetrapterum in Thailand. Map 1.1.12 Distribution of Calophyllum teysmannii in Thailand.

SW 11611-p162-216-G8.indd 183 1/16/62 BE 5:24 PM 184 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Thailand.— (Map 1.1.13): NORTHERN: Leaves petioled, petiole 0.9–1.9 cm long, light tan Lampang [Mt. Salor, 29 Oct. 1925, Winit 1492 to brown on lower surface and brown to red on upper (BK)]; Nakhon Sawan [Nakawn Tai, 11 Apr. 1922, surface; blade 5.4–16 × 2.5–5.5 cm, lanceolate, elliptic, Kerr 5816 (BK)]; EASTERN: Nakhon Ratchasima ovate to oval, glabrous, brown to red to olivaceous [Kao Lam, Kanat, 14 Jan. 1925, Kerr 9978 (BK); coriaceous, some slightly glaucous, base aequilateral ibid., 14 Jan. 1925, Kerr 9979 (BK)]; CENTRAL: to cuneate, apex acute to acuminate, midrib incon- Saraburi [Sam Lan, no date, Geesink et al. 6800 spicuous on both surfaces but more prominent (BKF); SOUTH-EASTERN: Chon Buri [Nong Ta Yu, abaxially, wider at base, gradually narrowing towards Sri Racha Forest, 12 Jan. 1926, Collins 11277 (C, K, apex, abaxially almost rectangular and raised L); Phra Chedi Mountains, Sri Racha District, 6 Mar. approximately 0.1–0.3 mm above surface, secondary 1975, Maxwell 75-2100 (AAU, BK, L)]; Chanthaburi veins present and weakly prominent on both surfaces, [at Chanthaburi River on path leading to Prachadu intramarginal veins present. Infl orescence terminal, Hill, Feb. 1916, Collins 563 (ABD, BM, K); Laem from upper leaf axils and from adjacent foliate axils, Sing, 23 Nov. 1924, Kerr 9368 (BK, C, K, L, P); pedicels 0.3–0.6 cm long. Floral parts tepals Khao Sabap, Chanthabun, 5 Dec. 1924, Kerr 9556 (8)9–13, outer pair suborbicular, dorsally pubescent, (A, BK, C, K, L, P)]; Trat [Ko Chang Island, Ao the next pair suborbicular, the inner pair elliptic. Ong Kang, 7 May 1974, Geesink et al. 6579 (AAU, Stamens approximately 400; fi laments up to 4 mm BKF, C, K, L, P); Ko Chang, 7 May 1974, Maxwell long; anthers oblong. Ovary approximately 1.5(–3) 74-404 (AAU, BK)]; Unknown locations [Winit mm long; style (3(–5) mm long. Fruit spherical to 1751 (BK)]. globose, 2.5–4 × 1.5–2.2 cm, round at apex; wrinkled Distribution.— Vietnam. with longitudinal striations, purple/red to black when dry; greenish brown when ripe. Ecology and phenology.— Rocky deciduous forest, evergreen forest, bamboo jungle from 30 to Thailand.— (Map 1.1.14): NORTH-EASTERN: 400 m elevation. Flowering: October–January. Phetchabun [Thung Salaeng Luang, 12 Dec. 1993, Fruiting: December–May. Pooma 728 (BKF)]; Nakhon Phanom [Phu Lankha, 16 Dec. 1968, T.P. 1951 (BKF)]; EASTERN: Nakhon Vernacular.— Kang han (กังหัน); ka thang han Ratchasima [Khao Yai NP, Khao Khieo, 18 Oct. (กะทังหัน) (Sangkaew, 1999). 1969, van Beusekom & Charoenphol 1724 (A, BKF, Uses.— Wood is used for construction and the C, K, L, P); Khao Yai NP, Khao Laem, 19 Oct. 1969, fruits are edible. The flowers are very fragrant van Beusekom & Charoenphol 1765 (BKF, L); Khao (Sangkaew, 1999). Yai NP, 19 Nov. 1962, Smitinand & Robbins 7913 (BKF); Khao Rom, Khao Yai, 3 Dec. 1983, Fukuoka Conservation.— IUCN Regional (Thailand) & Ito T-34637 (BKF); Khao Yai NP, 8 Sept. 1979, Status: Near Threatened (NT). Santisuk s.n. (BKF)]; Unknown locations [17 Dec. 1968, C.P. 1957 (BKF); 11 Sept. 1938, Poilane 14. Calophyllum touranense Gagnep. ex P.F.Stevens 277/9 (27719?) (BKF); Santisuk s.n. (BKF)]. in J. Arnold Arbor. 61: 226. 1980. Type: Indochina Distribution.— Vietnam. (Vietnam), Clemens & Clemens 4162 (holotype A [A00067245!]; isotypes K [K000677429!], NY Ecology and phenology.— Evergreen forest, [NY00066984!], P [P01900942!]).— C. touranense montane forest, rocky or stony ground from 650 to Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1: 271. 1500 m elevation. Flowering: January–March 1943, nom. inval. (description in French only); (October). Fruiting: February–June and September– Phạm-hoàng Hộ, Câycỏ Việtnam, 1(1): 573 1588, October. 1991. Vernacular.— Tang hon khao (ตังหนเขา) (Sangkaew, 1999). Trees 7–25 m tall; dbh 100 cm; outer bark whitish-grey or brownish-yellow, deeply fi ssured, Uses.— Used in house construction (Sangkaew, inner bark (slash-mark) red; exudate colour of clear 1999). honey. Branchlets terete fl aking, rough to smooth Conservation.— IUCN Regional (Thailand) to pubescent, scars present, grey-brown to red-brown. Status: Least Concern (LC).

SW 11611-p162-216-G8.indd 184 1/16/62 BE 5:25 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 185 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

2. KAYEA prominent. Infl orescences solitary and large, or in Wall., Pl. Asiat. Rar. 3: 5. 1832; Wight, Ill. Ind. Bot.: terminal or axillary panicles and small. Calyx sepals 129. 1840; Thwaites, Enum. Pl. Zeyl. 51. 1858; 4, imbricate. Corolla petals 4, imbricate, white or Planch. & Triana, Ann. Sci. Nat. Sér. 4, 15: 296. pink. Stamens numerous; fi laments basally connate 1861; Benth. & Hook.f., Gen. Pl. 1: 176. 1862; or free; anthers small; dehiscence vertical. Ovary T.Anderson in Hook.f., Fl. Brit. Ind. 2: 276. 1874; 1-celled; ovules 4 (rarely 8); style slender; stigmas Kurz, Forest Fl. Burma 1: 96. 1877; Engl. & Prantl, narrow, 4-fi d. Fruit fl eshy, indehiscent, 1–4 seeded, Nat. Pfl anzenfam. 3(6): 222. 1895; Brandis, Indian enclosed in enlarged calyx (in majority of species). Trees: 56. 1907; Pit., Fl. Indo-Chine 1(4): 325. 1910; Seeds thick, testa thin, woody. Ridl., Fl. Malay Penins. 1: 188. 1922. Type species: Distribution.— 75 species in tropical Asia: Kayea fl oribunda Wall. Borneo, Myanmar, Cambodia, Ceylon, India, Trees or shrubs; glabrous. Leaves opposite, Indonesia, Malaysia, Philippines; 3 species in blade lanceolate, coriaceous; veins distant, not Thailand.

KEY TO THE SPECIES 1. Infl orescence not solitary, axillary or terminal, usually paniculate; fruit globose 2. K. ferruginea 1. Infl orescence solitary, axillary or terminal; fruit not globose 2. Fruit ovoid to ellipsoid, <5 cm long, with persistent calyx; leaf apex acuminate to cuspidate 3. K. kunstleri 2. Fruit ellipsoid, >7 cm long, with or without persistent calyx; leaf apex long caudate 1. K. elegans

1. Kayea elegans King, J. Asiat. Soc. Beng., Pt. 2, axillary or terminal, ca 10 mm diam.,pedicel up to Nat. Hist. 59(2): 183. 1890; Vesque in DC., Monogr. 10 cm long with several ovate-acute bracts at its Phan. 8: 622. 1893. ≡ Mesua elegans (King) Kosterm., base. Calyx sepals equal, coriaceous. Corolla petals Reinwardtia 6: 427. 1969; Whitmore, Tree Flora of oblong, acute, smallerr than sepals, white. Ovaryy ovoid, Malaya 2: 162–236. 1973; Turner, Gard. Bull. attenuate. Fruit ellipsoid and tapering into a stout Singapore 47: 264. 1995. Type: Malaysia, Malacca, beak, surface smooth with ridges at 4 corners, Larut, Perak, King’s Collector 73466 (lectotype K 7.5–8.2 × 2.7–3.8 cm, ochrish grey, endocarp thin. [K000677315!], designated here; isolectotypes BM Seeds 3–6, oblong to round to almost square, covered [BM000611304!], BO not seen, GH!). in a thin paper-like layer inside fruit, black. — Kayea caudata King in J. Asiat. Soc. Beng., Pt. 2, Thailand.— (Map 1.2.1): PENINSULAR: Ranong Nat. Hist. 59(2): 183; 1890; Vesque in DC., Monogr. [Khlong Kam Phuan, 26 Apr. 1973, Geesink & Phan.7: 621. 1893. Type: Malaya, Malacca, Perak, Santisuk 4950 (AAU, BKF, L, P)]). King’s Collector 79377 (holotype K [K000677312!]). Distribution.— Malay Peninsula, Singapore. Trees 12–18 m tall; outer bark and inner bark Ecologyy and phenology.— Evergreen forest from red. Branchlets drooping, slender, thin, terete, scarring 450 to 600 m elevation. Flowering: approximately present, yellow/brown/grey; exudate not recorded. March. Fruiting: approximately April. Leaves petioled, petioles 0.4–0.7 cm long; blade Conservation.— IUCN: Global Status: Least 5.6–9.9 × 1.6–2.6 cm, oblong to elliptic to lanceolate Concern (LC); Regional (Thailand) Status: Critically in shape, base cuneate, some almost angustate, apex Endangered (CR). long caudate, apex length 1.3–2 cm long, upper and lower surfaces glaucous, some slightly coriaceous, rigid, some undulate, midrib pale yellow and 2. Kayea ferruginea Pierre, Fl. Cochinch. t. 99. depressed on upper surface, yellow-tan and slightly 1889; Ridl., J. Straits Branch Roy. Asiat. Soc. 54, 21. prominent on lower surface (almost depressed); 1897; Pit. in Lecomte, Fl. Indo-Chine 1(4): 326. veins visible, no intramarginal veins present, very 1910; Ridl., Fl. Malay Penins. 1: 189–190. 1922; faint on upper surface, veins not arising at same point Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3): on opposite side of midrib. Infl orescence solitary, 276. 1943. ≡ Mesua ferruginea (Pierre) Kosterm.,

SW 11611-p162-216-G8.indd 185 1/16/62 BE 5:26 PM 186 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Reinwardtia 6: 427.1969; Whitmore, Tree Flora of 3. Kayea kunstleri King, J. Asiat. Soc. Bengal, Pt. 2, Malaya 2: 230. 1973; Turner, Gard. Bull. Singapore Nat. Hist. 59(2): 182. 1890; Vesque in DC. Monogr. 47: 264. 1995. Type: Vietnam, Pierre 1050 (lecto- Phan. 8: 620. 1893; Ridl., Fl. Malay Penins. 1: type K [K000677333!] , designated here; isolecto- 190–191. 1922. ≡ Mesua kunstleri (King) Kosterm., typesA [A00067758!],GH!, K!, NY [NY00075970!], Reinwardtia 6: 427. 1969; Whitmore, Tree Flora of P!). Syntype: Vietnam, Pierre 4564 (P!). Malaya 2: 228. 1973. Type: Malaysia, Malacca, Larut, Perak, King’s Collector 6850 (lectotype K Trees or shrubs 8–12 m tall; dbh 25 cm; outer [K000677324!], designated here; isolectotypes bark grey-brown, smooth to scaly, inner bark pink BM [BM000611308!], A [A00067764!], BO not to red, exudate not recorded. Branchlets green/grey/ seen, FI [FI010334!], SING [SING0194784!], GH!). brown/black, smooth to slightly fl aking, scars present, Syntypes: Malaysia, Malacca, Perak, Malaysia, King’s round. Leaves petioled, petiole 0.7–1.1 cm long, Collectorr3301 (BO n ot seen, SING [SING0194782!]); bronze to green/yellow; midrib very pronounced on Penang, Curtis 1419 (SING [SING0194783!]); lower surface, very depressed in channel on upper Malacca, Maingay 1766 (not located). surface; blade 5–14.7 × 2.1–4.6 cm, elliptic to lanceolate, coriaceous to smooth, with a slight shine — Kayea curtisii King, Ann. Roy. Bot. Gard. to glaucous and dull, base cuneate, apex acute, some Calcutta 5: 144, t. 174B. 1896. Type: Malaysia, acuminate, secondary veins approximately 16 on Malacca, Penang, Curtis 805 (lectotype K each side of blade, 10–14 mm apart, intersecondary [K000677313!], designated by Whitmore [1973]; veins present, tertiary veins dense and intramarginal isolectotype BO not seen, SING [SING0194787!]. veins present, all very faint and fi ne. Infl orescence — Kayea rivulorum Ridl., J. Straits Branch Roy. paniculate, pedicel up to 3 cm long, forming a Asiat. Soc. 54: 22. 1910. Syntypes: Malacca, rounded ball of small spheres, approximately 15 Malaysia, Goodenough 19766 (SINGG [SING0194794!]); clustered together at terminal point, each ca 1.5 cm Selangor, Malaysia, Ridley 7349 (GH!, SING long, usually 3–5 flowers clustered in leaf-axils. [SING0194793!]). Calyx sepals persistent, in 2 opposite pairs. Corolla petals acute, oblong, imbricate, white. Stamens in Shrubs to small trees, 0.5 to 8 m tall; dbh fascicles, numerous, anther & fi lament approximately 10–30 cm; outer bark brown, inner bark pink; 2.5 mm long, anthers orange/yellow. Ovary with exudate whitish-cream. Branchlets brown/red to long single style protruding. Fruit globose, with light brown, thin, striate, scars present. Leaves acute tip, 3–3.5 × 2.5–3.5 cm, texture rough, brown, petioled, petiole 0.4–1.6 cm long, blade 6.7–11.6 × pedicel 3 mm long, enlarged calyx surrounding the 1.4–3.6 cm, elliptic to oblanceolate, dull green to fruits. Seeds 1. brown to orange/yellow, some glaucous, some dots/ glands present, base cuneate to acute, apex cuspidate Thailand.— (Map 1.2.2): PENINSULAR: Surat (often with an obvious mucro), a few acuminate, Thani [Bang Yai River Street, 4 June 1966, Sakol coriaceous to thin and delicate, venation extremely 1018 (BK); Ban Duara, Sept. 1924, Bourke 2501 faint on some leaves, varying from leaf to leaf, veins (BK, K)]; Narathiwat [Tak Bai, 29 Apr. 1984, appear to fade on older leaves, arising at diff erent Niyomdham 778 (BKF, P); Tak Bai, Ban Khunthong, points on either side of the midrib, midrib pale cream 29 Apr. 1984, Niyomdham 810 (BKF, K, P)]. to green on upper surface and light brown to tan on Distribution.— Cambodia, Malay Peninsula, lower surface, not very obvious on either surface, Singapore. slightly more prominent on lower surface. Ecology and phenology.— Common along Infl orescence solitary, axillary or terminal, pedicel rivers and streams, near sea level. Flowering: length 1.5–3 cm long, smooth, bud smooth. Calyx March–June. sepals outer 2 smaller, round-oval, green, inner 2 larger, elliptic, white, persistent. Corolla petals Vernacular.— Bunnak nam (บุนนาคนํ้า), la-do- oblong-acuminate, caducous, white, up to 1 cm long. pa-he (ลาดอปาเฮะ) (Niyomdham 778). Stamens numerous, in fascicles, anther & fi lament Conservation.— IUCN Regional (Thailand) 1.5–2 cm long, filament cream, anthers round to Status: Data Defi cient (DD). almost oblong, yellow. Ovary yellow, style 1. Fruit ovoid to ellipsoid, 3.5–4.5 cm long texture wood-like

SW 11611-p162-216-G8.indd 186 1/16/62 BE 5:26 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 187 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Map 1.1.13 Distribution of Calophyllum thorelii in Thailand. Map 1.1.14 Distribution of Calophyllum touranense in Thailand.

Map 1.2.1 Distribution of Kayea elegans in Thailand. Map 1.2.2 Distribution of Kayea ferruginea in Thailand.

SW 11611-p162-216-G8.indd 187 1/16/62 BE 5:24 PM 188 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

with scales, green outside, cream inside, fruiting Distribution.— Malay Peninsula, Singapore. calyx yellow-greenish, persistent. Seeds 1–4. Ecology and phenology.— Evergreen forest; Thailand.—(Map 1.2.3): PENINSULAR: Krabi open disturbed growth from 50 to 600 m elevation. [4 Apr. 1930, Kerr 18842 (BK, BM, C, K, L, P)]; Flowering: March–June. Fruiting: July–September. Nakhon Si Thammarat [Khao Luang, near Khiri Vernacular.— Nakaboot (Maxwell 84-179). Wong Village, 16 May 1968, van Beusekom & Phengklai 778 (BKF)]; Phatthalung [10 km East of Conservation.— IUCN Global & Regional Na Thawi, 24 Nov. 1990, Larsen et al. 41691 (AAU)]; (Thailand) Status: Least Concern (LC). Satun [Thale Ban NP, 12 July 2000, Middleton et al. 391 (A, BKF)]; Songkhla [Kho Hong Hill, Hat Yai, 3. MAMMEA 17 May 1960, Chirayupin 2 (BK); Kho Hong Hill, L., Sp. Pl.: 512. 1753; Juss., Gen. Pl.: 257. 1789; Hat Yai, 1997, Sritungnant 11 (PSU); ibid., 21 Sept. Benth. & Hook.f., Gen. Pl. 1: 176. 1862; Engl. in 1976, Cheuntai 18 (PSU); ibid., 4 Jan. 1974, Cannon Engl. & Prantl, Nat. Pfl anzenfam. 3(6): 219. 1895; JRC 243 (PSU); ibid., 30 Oct. 1990, Larsen et al. Kosterm., Pengum. Lemb. Pusat Penjel. Kehut 72: 409566 (AAU); ibid., 4 June 1992, Larsen et al. 42652 15. 1961; Backer & Bakh.f., Fl. Java 1: 384. 1963; (AAU, P); ibid., Hat Yai, 1 Sept. 1984, Maxwell Whitmore, Tree Flora of Malaya 2: 226. 1973; 84-1799 (PSU); ibid., 31 Dec. 1985, Maxwell 85-1186 P.F.Stevens, Aust. J. Bot. 22: 413–423. 1974. Type (PSU); Hat Yai, 16 Apr. 1963, Pradit 190 (BK); species: L. (lectotype designated PSU, behind Biology Building, 2 Feb. 1979, by Britton & P.Wilson, Scient. Surv. Porto Rico 5: Congdon & Hamilton 249 (PSU); Khuan Mit, south 583. 1924). of Hat Yai, 29 Apr. 1979, Congdon & Hamilton 403 (PSU); Sadao, 12 May 1970, Sutheesoen 1683 (BK); — Calysaccion Wight, Ill. Ind. Bot. 1:130. 1840. Utapao, 14 May 1919, Kerr 3684 (BM, K); PSU Type species: Calysaccion longifolium Wight. Campus, 11 Apr. 1976, Sutheesoen 3692 (BK); Ban Trees; exudate clear or coloured and resinous. Thepha, 24 Mar. 1928, Kerr 14742 (BK, BM, C, K, Leaves simple, opposite, blade entire, often coriaceous L, P); Prakawp, 10 July 1928, Kerr 158466 (BK, and persistent, stipules absent, some with black BM, K, L); Khao Mot Daeng, Kho Hong, Hat Yai, glandular dots. Infl orescences terminal or axillary, 6 Oct. 1991, Larsen et al. 421866 (AAU, BKF, P, some often caulifl orous, solitary or cymose, bisexual PSU); Yo Island, Muang District, 27 Aug. 1986, or unisexual, sometimes in clusters in axils of fallen Maxwell 86-624 (BKF, L, PSU)]; Yala [Banang leaves. Calyx sepals 2, entirely closed and united in Station, 23 July 1923, Kerr 73766 (BK, BM, K)]; bud, later splitting into 2 reflexed sepals during Narathiwat [Waeng, Phusomsaeng 440 (L); Khao fl owering. Corolla petals 4–6, free or nearly entirely Re Chan, To Mo, 20 Apr. 1931, Lakshnakara 728 connate, usually white. Stamens numerous, free or (BK, BM, C, K, L, P); Khao Cha Bor, To Mo, 27 basally connate, dehiscence vertical. Ovary 2 (–many) Apr. 1931, Lakshnakara 816 (BM, C, K, L, P); celled; ovules 4; style short or not present, if present, Waeng, 13 Aug. 1972, Phusomsaeng et al. 1027 stigma peltate. Fruit fleshy, coriaceous when dry. (BKF); Nikhom Waeng, 14 Apr. 1968, Prayad 1296 Seeds 1–4, embryo with 2 fused cotyledons. (BK); ibid., 28 Apr. 1968, Prayad 1331 (BK); Sungei Padi, Chatwarin Falls, 5 Mar. 1974, Larsen & Larsen Distribution.— 75 species in Borneo, Myanmar, 32998 (AAU, BKF, K, L, P); Waeng, Phusomsaeng China, Indonesia (Flores, Java), Philippines, Samoa, BKF 41118 (K)]; Unknown locations [Ban Tu Gor, North and South America; 3 species in Thailand. Tan Yong Mas, 30 Apr. 1931, Lakshnakara 843 (BM, C, K, L, P); Sirirugsa s.n. (PSU)].

KEY TO THE SPECIES 1. Emarginate leaves absent 2. M. harmandii 1. At least some emarginate leaves present 2. Leaves obovate or elliptic; 11–29.5 cm long; pedicels up to 0.5 cm long 1. M. brevipes 2. Leaves obovate, oblanceolate or oblong; 7.5–25 cm long; pedicels longer than 0.5 cm 3. M. siamensis

SW 11611-p162-216-G8.indd 188 1/16/62 BE 5:26 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 189 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

1. Mammea brevipes (Craib) Kosterm. in Pengum. 2. Mammea harmandii (Pierre) Kosterm., Addit. Lemb. Pusat Penjel. Kehut 72: 13. 1961; Whitmore, Notes Mimos.; the Genera Mammea L. and Tree Flora of Malaya 2: 226. 1973. ≡ Ochrocarpos Ochrocarpos Thou.: 13. 1956; Kosterm., Commun. harmandii Pierre var. brevipes Craib, Fl. Siam. 1: Forest Res. Inst. Bogor 72: 22. 1961. ≡ Ochrocarpus 119. 1925. Type: Thailand, Phetchabun, Kerr 5706 harmandii Pierre, Fl. Cochinch. 3: t. 93. 1883; Pit. (lectotype K [K000677516!], designated here; in Lecomte, Fl. Indo-Chine 1(4): 295. 1910; Petelot, isolectotype BM [BM000611511!]). Pl. medic. Cambodge, Laos, Vietnam (Recherches — Ochrocarpus [sic] siamensis auct. non T.Anderson: agron. C.L.V. 14) 1: 59. 1952. ≡ Calysaccion Ridl., Fl. Malay Penins. 1: 180. 1922. based on harmandii (Pierre) Pierre, Bull. Soc. Linn. Paris 2: Robinson 6304 (K!). 1226. 1896–1897.Type: Laos, Harmand 1072 (lecto- type P [P00642270!], designated here; isolectotype Trees 8–12m tall; outer bark smooth to rough, P [P00642271!]). dark brown to grey; exudate not recorded. Branchlets terete. Leaves petioled, 0.6–2 cm long; blade 11–29.5 Trees 6–10 m tall, dbh 25–30 cm; outer bark × 5–11 cm, elliptic to obovate, coriaceous, leaf edges reddish-brown, inner bark dark red to pale fl esh-like; slightly curling downwards especially near the leaf exudate yellow, sticky. Branchlets terete with longi- base, base cuneate some almost angustate, apex tudinal ridges, smooth. Leaves petioled, 1–1.7 cm rounded or cuspidate, if cuspidate, tips 3–4 mm long, long, blade 13.1–17 × 4.4–5.6 cm, obovate, lanceolate, at least some emarginate, some glaucous, khaki elliptic, coriaceous, undulate, base cuneate, apex green to dark green adaxially, pale yellow abaxially, mostly cuspidate, few round or acute, midrib raised midrib raised and yellow abaxially, dark brown- and light green-brown abaxially, green-brown green adaxially, secondary, intersecondary, tertiary adaxially, netted veining visible, obscure, no intra- and intramarginal veins visible, secondary veins marginal veins present. Inflorescence clusters of 10–15 per side, 7–10 mm apart. Infl orescence many fl owers borne on the branches, fl owers arising on fl owers clustered together, shortly petioled, pedicels branches where small branches join larger branches, terete, up to 0.5 cm long, borne on inconspicuous arising from separate points but close together on irregularly rounded protuberances 3–5 mm wide, the branches. Calyx sepals elliptic, persistent. each subtended by 4–8 closely imbricate bracts. Corolla petals 2, elliptic, longitudinal veins present Calyx sepals elliptic, 7–9 × 5–7 mm, persistent, on petals. Stamens oblong, anther and fi lament 3 mm papyraceous, obtuse or slightly acute at apex with long. Ovary globose; stigma peltate; style up to 4 mm 15–18 parallel nerves. Corolla petals 4–6, obovate long. Fruit ellipsoid, rough, 2.3–2.5 × 1.0–1.4 cm, to round, 10–17 × 4–8 mm. Stamens in fascicles, yellow, vertically dehiscing, tip acute, petiole up to approximately 200, anthers oblong, 1–3 mm long, 1–2.2 cm long. free or weakly connate at the base. Ovary ellipsoid; Thailand.— (Map 1.3.2): NORTHERN: Chiang 2–2.5 mm diam., style terete, approximately 2 mm Rai [Doi Tung, near Temple, 17 May 1992, Santisuk long, stigma peltate; 2–lobed, approximately 2 mm s.n. (BKF)]; EASTERN: Nakhon Ratchasima diam. Fruit oval to elliptic, 2–2.7 × 1–1.3 cm, hard, [Sakaerat, 22 Oct. 1971, van Beusekom et al. 3291 wood-like, purple/red, petiole up to 3 mm long. (BKF, C, K, L, P); Khao Phrik, Sikhieo District, 9 Seeds seed(s) embedded in pulp, when fresh. May 1976, Maxwell 76-3077 (L); CENTRAL: Saraburi [Sam Lan Forest, Muang District, 1 Mar. 1975, Thailand.— (Map 1.3.1 ): NORTH-EASTERN: Maxwell 75-174 (AAU, BK, L); ibid., 17 July 1976, Phetchabun [Muang, 28 Mar. 1922, Kerr 5706 Maxwell 76-434 (AAU, BK, L)]; Unknown location: (ABD, BM, K); Chondaen, 8 Mar. 1931, Kerr 20392 [8 Feb. 1956, Smitinand s.n. (BKF)]. (ABD, BK, BM, E, K)]. Distribution.— Laos, Malay Peninsula Distribution.— Endemic. (Langkawi). Ecology and phenology.— Evergreen forest, Ecology and phenology.— Evergreen forest, on limestone from 400 to 500 m elevation. Flowering: deciduous forest from 150 to 650 m. Flowering: March. February–May. Fruiting: July–October. Conservation.— IUCN Regional (Thailand) Conservation.— IUCN Regional (Thailand) Status: Critically Endangered (CR). Status: Data Defi cient (DD).

SW 11611-p162-216-G8.indd 189 1/16/62 BE 5:26 PM 190 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

3. (Miq.) T.Anderson, J. Linn. Feb. 1960, Kasin 8066 (BK); Doi Suthep, 21 Oct. Soc. 9: 261. 1867; Hook.f., Fl. Brit. Ind. 1: 271. 1958, Sørensen et al. 58100 (C); ibid., 1958, Sørensen 1874; Vesque in DC., Monogr. Phan. 8: 527. 1893; et al. 5829 (C); Muang Tun, 30 June 1922, Kerr Kosterm., Pengum. Lemb. Pusat Penjel. Kehut 72: 6193 (ABD, BK, BM, K); Doi Suthep, between the 26. 1961; Whitmore, Tree Flora of Malaya 2: 226. Agricultural Station and TV-7 Station, 9 June 1988, 1973. ≡ Calysaccion siamense Miq., Mus. Bot. 1: Maxwell 88-739 (AAU, L); East side, 76/1 Soi 5, 209. 1863. ≡ Ochrocarpos siamensis (Miq.) Suthep Road, 8 Mar. 1992, Maxwell, 92-70 (A, T.Anderson in Hook.f. Fl. Brit. Ind. 1: 270–271. CMUB, E, P); Huai Kaeo Arboretum, Muang, 12 1874; Kurz, Forest Fl. Burma 1: 94. 1877; Gamble, Mar. 1986, Phengklai s.n. (BKF); Doi Suthep, Kerr Man. Ind. Timb. 56. 1881; Pierre, Fl. Cochinch. t. s.n. (BM)]; Lampang [Jaehomwittya School, 94–96. 1885; Pit. in Lecomte, Fl. Indo-Chine 1(4): Chaehom District, 16 Feb. 2001, Panatkool 464 293–294. 1910; Ridl., Fl. Malay Penins. 1: 180. (CMUB)]; Phetchabun [Ban Wang Saphung, Wang 1922; Craib, Fl. Siam. 1:119. 1925; Gagnep. in Pong District, 1 Feb. 2001, Wongprasert 012-31 Humbert, Suppl. Fl. Indo-Chine 1(3): 254. 1943. (BKF)]; EASTERN: Nakhon Ratchasima [Sakaerat. Type: Thailand, Bangkok, Teijsman s.n. (holotype? Pak Thong Chai District, Reforestation Station, 21 BO not seen; isotypes: L [L0012276!], U Apr. 1987, Soejarto et al. 6022 (A, L); Khao Phrik, [U0002418!]). Sikieo District, 9 May 1976, Maxwell 76-3077 (BK)]; SOUTH-WESTERN: Phetchaburi [18 Feb. 1923, Trees to 20 m tall, dbh 10–30 cm; outer bark Marcan 1176 (ABD, BM); Khao Wong, 2 Mar. 1963, thick, roughly cracked and flaking, sometimes Sakol 459 (BK)]; CENTRAL: Krung Thep Maha smooth or slightly fi ssured, dark grey, inner bark Nakhon [1899, Zimmermann 105 (BM, L); red; exudate pale yellow. Branchlets smooth, Krungthep, 2 Mar. 1927, Lakshnakara 305 (ABD, yellowish. Leaves shortly petioled, petiole 0.5–1.5 cm BK, K); ibid., 8 June 1931, Lakshnakara 854 (BK, long, blade 7.5–25 × 2.5–7 cm, oblanceolate, obovate BM); Marcan 17077 (ABD, BM); Unknown collector or oblong, coriaceous, young leaves purple, mature 5953 (L); Feb. 1955, Phloenchit s.n. [BKF 11561] dark-green above, yellow-green abaxially, base (BKF)]; SOUTH-EASTERN: Chon Buri [Naphro, Near narrowly cuneate, apex blunt to slightly cuspidate, Sri Racha, 19 Apr. 1923, Collins 789 (ABD, BK, E, some emarginate, midrib dark green adaxially, green K)]; PENINSULAR: Chumphon [Unknown collector abaxially, side veins numerous on both surfaces. 4360 (K)]; Songkhla [Ma Be Ya Village, Foothills Infl orescence clustered on old woody twigs behind of Klong Rhang Hill, Na Mom District, 15 Feb. leaves, pedicels light green to yellow, up to 2 cm 1986, Maxwell 86-70 (A, L, PSU); PSU, Hat Yai long. Male and female fl owers on separate trees, District, 7 June 1986, Maxwell 86-343 (A, BKF, fl owers approximately 1.2–2.5 cm long, white or PSU); Wat-Pako, Sathingpra, 23 Apr. 1974, Vitchu pale yellow. Calyxx sepals elliptic, 2–7 mm long, light V30 (PSU)]. green. Corolla petals 4, oblong, 6–8 mm long. Stamens between 60–90 stamens per fl ower, fi laments Distribution.— Laos, Malay Peninsula white, free, anthers yellow, 1–2 mm long. Ovary (Penang). style and ovary light green maturing darker green, Ecology and phenology.— Dry, mixed evergreen style 1, short with 2-lobed stigma. Fruit oval to forest, deciduous forest, on granite bedrock from 50 ellipsoid with short blunt tip, up to 4 × 2.5 cm but to 460 m elevation. Flowering: February–June. usually smaller, 2-valved, rind with sparse white Fruiting: June. The fl owers contain aromatic oil and latex, juicy and dull yellow inside, orange on outside. the species is widely cultivated. Seeds 1, with thin yellow coating (aril). Vernacular.— Sarapee (สารภี) (Palee 30); Thailand.— (Map 1.3.3): NORTHERN: Chiang Sa ra pi (สารภี) (Lakshnakara 854). Mai [Chiang Mai University, Biology Department, 3 June 1992, Palee 300 (A, CMUB, E, P); Suan Kaeo, Uses.— The pollen is used by Thai people as Mae Rim, 5 Mar. 2002, Glamwaewong 1566 (QBG); a cosmetic. The fl owers are used to make necklaces. Doi Suthep, on lower slopes, 14 Mar. 1909, Kerr Conservation.— IUCN Regional (Thailand) 548 (BM, K, P); Doi Suthep, Ban Salienue, Hot, 18 Status: Least Concern (LC).

SW 11611-p162-216-G8.indd 190 1/16/62 BE 5:26 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 191 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Map 1.2.3 Distribution of Kayea kunstleri in Thailand. Map 1.3.1 Distribution of Mammea brevipes in Thailand.

Map 1.3.2 Distribution of Mammea harmandii in Thailand. Map 1.3.3 Distribution of Mammea siamensis in Thailand.

SW 11611-p162-216-G8.indd 191 1/16/62 BE 5:24 PM 192 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

4. MESUA Ridl., Fl. Malay Penins. 1:192. 1922; Kirtikar & L., Sp. Pl.: 515. 1753; Juss., Gen. Pl. : 258. 1789; Basu, Indian Med. Pl. 2(1): 274. 1933; Kamjilal, Wight, Prodr. Fl. Ind. Orient. 102. 1834; Miq., Fl. Fl. Assam 1: 111. 1934; Kitamura, Fauna Fl. Nepal Ned. Ind. 1(1): 509. 1859; Planch. & Triana, Ann. Himalaya 1: 179. 1955; MacMillan, Trop. Pl. & Sci. Nat., Bot. Sér. 4, 15: 298. 1861; Benth. & Gard. 5: 87. 1956; Sastri, Wealth of India 6: 349. Hook.f., Gen. Pl. 1: 176. 1862; T.Anderson in 1962; Maheshwari, Bull. Bot. Surv. India 5: 337. Hook.f., Fl. Brit. Ind. 2: 276. 1874; Kurz, Forest Fl. 1963; Whitmore, Tree Flora of Malaya 2: 227–228. Burma 1: 96. 1877; King, J. Asiat. Soc. Bengal, Pt. 1973; Kosterm. in Dassan., Rev. Handb. Fl. Ceylon 2, Nat. Hist. 59(2): 184–185. 1890; Engl. in Engl. 1: 107. 1980; Gamble, Man. Ind. Tim. (ed. 2a) 59. & Prantl, Nat. Pfl anzenfam. 3(6): 218–219. 1895; 1992; H.W.Li et al., Fl. China 13: 38. 2007. Type: Prain, Bengal Pl. 1: 245. 1903; Pit. in Lecomte, Fl. Sri Lamka, Herb. Hermann 1: 38, No. 203 (lectotype Indo-Chine 1(4): 328. 1910; Gamble, Fl. Madras BM [BM000621365!], designated by Maheshwari 76. 1915; Haines, Bot. Bihar Orissa 2: 55. 1921; [1963] (see Jarvis, 2007 for further details) Ridl., Fl. Malay Penins. 1: 191. 1922; Whitmore, — Nagassarium Rumph., Herb. Amboin. 7: 3, t. 2. Tree Flora of Malaya 2: 227–228. 1973; Kosterm. in 1750. nom. inval.— Calophyllum nagassarium Dassan., Rev. Handb. Fl. Ceylon 1:107, 1980; Gamble, Burm., Fl. Ind.: 121. 1768. ≡ Mesua nagassarium Man. Ind. Tim. (ed. 2a) 59. 1992; H.W.Li et al., Fl. (Burm.f.) Kosterm., Ceylon J. Sci., Biol. Sci. 12:71, China 13: 38. 2007. Type species: Mesua ferrea L. 1976; Kosterm. in Dassan., Rev. Handb. Fl. Ceylon 1:107, 1980; P.F.Stevens, J. Arnold Arbor. 61:682, Trees; inner bark usually with sticky exudate 1980. Type: [Indonesia, Amboina and Java], Herb. drying black. Leaves opposite, blade coriaceous, Burman s.n. (holotype: G not seen). often with dots, veins numerous, slender. Infl orescences solitary, axillary or terminal. Calyx sepals 4–5, — Mesua speciosa Choisy in DC. Prodr. 1: 562. imbricate. Corolla petals 4–5, imbricate. Stamens 1824. Type: India, Rheed. Hort. Malab. 3: 63, t. 53. numerous, fi laments basally connate. Ovaryy superior, Trees up to 30 m tall, dbh 95 cm; bole straight, 2-celled; ovules 4; style long; stigma peltate, stigma cylindrical, often fl uted at base; outer bark ash-grey lobes broad. Fruit fl eshy or woody, drupe, or turning dark-dull brown, scaly, irregularly fi ssured, capsule, 1–4 seeded. Seeds testa fragile, aril absent. inner bark brownish-red to pink or red; exudate Distribution.— 5 species in tropical Asia; 1 aromatic, clear; sapwood creamy white to pinkish- species in Thailand. brown; heartwood dark red, hard, tough and heavy. Branchlets young twigs slender, grey and terete, end bud inconspicuous. Leaves petioled, 0.4–1.2 cm Mesua ferrea L., Sp. Pl.: 515. 1753; Burm.f., Fl. long, green above and below; blade 1–15 × 1–18 cm, Ind. 121. 1768; Choisy in DC. Prodr. 1:562. 1824; elliptic, oblong or lanceolate, glaucous abaxially, Wight, Prodr. Fl. Ind. Orient.: 102. 1834; Wight, coriaceous or delicate, base acute or obtuse, apex Icon. Pl. Ind. Orient.: t. 118. 1839; Graham, Cat. acute, young leaves pink, midrib faint and depressed Pl. Bombay: 26. 1839; Thwaites, Enum. Pl. Zeyl.: on both surfaces, slightly more pronounced on lower 5. 1858; Miq., Fl. Ind. Bat. 1: 509. 1860; Planch. & surface, side veins very fine, almost invisible, Triana, Ann. Sci. Nat., Bot. Sér 4, 15: 299. 1861; secondary, intersecondary, tertiary and intramarginal Dalzell & Gibson, Bombay Fl.: 31. 1861; Beddome, veins faint especially on young leaves, numerous, Fl. Sylv. S. India: t. 64. 1871; T.Anderson in somewhat patent, invisible on the white waxy lower Hook.f., Fl. Brit. Ind. 2: 277. 1874; Kurz, Forest Fl. surface. Inflorescence solitary, axillary, on short Burma 1: 97. 1877; Gamble, Man. Ind. Timb.: 27. peduncles, 0.8–2.3 cm long, slender, 4–10 cm diam., 1881; Pierre, Fl. Cochinch. 7: t. 97. 1885; Watt, Dict. sweet scented. Calyx sepals 4, 12–15 mm long, Econ. Prod. India 5: 236. 1891; Vesque in DC., imbricate, in 2 rows, small outer pair and larger inner Monogr. Phan. 8: 630. 1893; Talbot, Trees Bombay pair; light green, persistent, densely velvety puberulous Pres:.16. 1894; Woodrow, J. Bombay Nat. Hist. outside, orbicular, imbricate, fl eshy. Corolla petals 4, Soc. 1: 81. 1901; Prain, Bengal Pl. 1: 246. 1903; 16–40 mm long, white, cuneate, obovate or obcordate, Brandis, Indian Trees: 55. 1907; Gamble, Fl. Madras curled and erose at the margins, brown/purple 1: 77. 1915; Haines, Bot. Bihar Orissa 2: 55. 1921;

SW 11611-p162-216-G8.indd 192 1/16/62 BE 5:26 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 193 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

striations present, caducous. Stamens numerous; 720 (BKF, K)]; Nakhon Si Thammarat [28 Mar. anthers large, linear 4–10 mm long, orange-golden- 1955, Thaworm 50 (BKF)]; Trang [Chong, 8 Apr. yellow. Ovary up to 5 mm, ovoid, 2-locular; style 1949, Boongird 62 (A); Khao Chong, 14 Apr. 1969, 1–1.1 cm long; stigma peltate. Fruit ovoid to Phusomsaeng 171 (C, E, K, P); ibid., 1 Mar. 1966, ellipsoid with conical point, 2.5–3.5 cm, striate; Bunnab 376 (BKF, L); ibid., 13 June 1969, sepals enlarged up to 4 cm long, calyx persistent, Sangkhachand 1863 (BK); Chong, 25 Jan. 1958, dark orange or purple/brown, pericarp tough. Seeds Smitinand 4117 (BKF); Kantang, 14 Mar. 1929, 1–4, up to 2.4 cm woody, smooth, glossy brown, oily. Kerr 17485 (ABD, BK, BM, C, K, TCD)]; Yala [Banglang, Thanto, 20 Sept. 1992, Niyomdham 5126 Thailand.— (Map 1.4.1): NORTHERN: Mae Hong Son [Pang Mu, 8 July 1958, Smitinand 4597 (BKF)]; Narathiwat [Nikhom Waeng, 22 Sept. 1966, (BKF)]; Chiang Mai [CMU, in front of Humanities Prayad 4177 (BK); Khao E-dang, Sukhirin, 23 Aug. Faculty, Muang District, 13 Jan. 2001, Intien 15 (A, 1998, Puudjaa 498 (BKF); Waeng, 6 Sept. 1966, CMUB); ibid., 5 Aug.,1995, Tumersnis 32 (CMUB); Sangkhachand 1317 (BKF, L); Su-ngai Padi, Huai Pu, QBG, Mae Rim, 20 Mar. 1998, Watthana Chatwarin Falls, 5 Mar. 1974, Larsen & Larsen 86 (QBG); Huai Kaeo Arboretum, 7 June 2000, 32998 (L)]. Sankamethawee 122 (CMUB); ibid., 15 Feb. 1992, Distribution.— Sri Lanka, India, tropical Nepal, Pooma 632 (BKF); ibid., 15 May 1995, BGO Staff Myanmar, Cambodia, Laos, Vietnam, Malaysia. 33666 (BKF, QBG); Muang, Brun et al. 159 (C); Ecology and phenology.— Evergreen forest, Maesa Botanic Garden, 22 Aug. 1989, Pooma 291 deciduous forest, near the sea, on limestone from 10 (BKF); Botanic Garden, Mae Rim, 25 May 1994, to 1050 m elevation. Flowering: February–May. BGO Staff 710 (QBG); Doi Suthep, 1 July 1998, Fruiting: May–October. Pongamornkul 131 (QBG); ibid., 11 June 1910, Kerr 1211 (BM, K, TCD); ibid., Oct. 1970, Dixen Vernacular.— Mai bun nak (ไมบุนนาค) (Vanpruk s.n. (AAU)]; Doi Suthep, East side, Khonthathan 720); bun nak (บุนนาค) (Kerr 5916). Falls Area, May 1991, Maxwell 89-631 (E); Doi Uses.— The wood from this species is one of Suthep Pui NP, Park HQ, Muang, 13 Sept. 1995, the strongest woods found in Asia and is used for Kopachon s162b1 (BM, CMUB); ibid., 6 Nov. 1997, construction work, for railway sleepers and in boat Sritong, s162b2 (CMUB); Danta Bua, 31 July 1914, building. Oil from the seed is used in soap making Kerr s.n. (BM); Chiang Dao Watershed Station, 4 and the fl owers and fl ower buds are used in cosmetics June 1973, Geesink et al. 56877 (AAU, BKF, C, E, K); and perfumery. In India the leaves and fl owers are Chaiprakan, Sidongyen Subdistrict, Lahn Village, also used as hair decorations and the fruits are 24 May 1991, Maxwell 91-472 (A, E, P); Phayao sometimes eaten. [Doi Luang NP, east side of Jahn Bah Tawng, Muang Conservation.— IUCN Regional (Thailand) District, 6 May 1997, Maxwell 97-451 (A, CMUB); Status: Least Concern (LC). ibid., 9 Aug. 1997, Maxwell 97-844 (A, BKF, CMUB)]; Sukhothai [Mt. Khao Luang, 2 May 1922, Kerr 5916 (AAU, ABD, BK, BM, K)]; CENTRAL: 2. HYPERICACEAE Krung Thep Maha Nakhon [Phrayathai, 10 Feb. Evergreen or deciduous herbs, shrubs or trees 1955, Agooroo 22 (BKF); 1899, Zimmermann 110 with glands or canals present in most parts of the (BM, K, L, P); 20 Feb. 1921, Marcan 581 (BM, K); plant, uni/multicellular hairs usually present. Leaves 7 Mar. 1926, Marcan 1998 (BM, K); Kerr s.n. opposite, alternate or whorled. Inflorescences (BM)]; SOUTH-EASTERN: Chanthaburi [Cultivated terminal or cymose; fl owers single, perfect. Calyx in Arboretum, Makham District, 29 Mar. 1995, (2–)4–5 sepals, free. Corolla (3–)45 petals, free. Santisuk et al. s.n. (BKF)]; PENINSULAR: Ranong Stamens 9–∞, free, connate or fasciculate, nectary [Khao Phota Luang Kaeo, 22 June 1974, Geesink absent. Ovary superior, 1–∞ ovules. Fruit capsule et al. 7434 (BKF, K); Hat Hin Dam, 24 Apr. 1974, or baccate (berry-like), sometimes drupe, ± wings. Larsen & Larsen 33352 (AAU, K, P)]; Phuket [Muang Mai, 9 Mar. 1929, Kerr 17400 (ABD, BK, Distribution.— Worldwide with 8 genera and BM, C, E, K, TCD)]; Krabi [Apr. 1915, Vanpruk 560 species; 2 genera and 11 species in Thailand.

SW 11611-p162-216-G8.indd 193 1/16/62 BE 5:26 PM 194 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

KEY TO THE GENERA 1. Herbs with horizontal runners; fl owers usually yellow 1. Hypericum 1. Deciduous to evergreen trees or shrubs; fl owers pink or white 2. Cratoxylum

1. HYPERICUM spreading or eventually refl exed, aestivation always L., Sp. Pl.: 783. 1753; L., Gen. Pl. 5: 341. 1754; contorted, usually yellow or sometimes white, dotted Juss., Gen. Pl.: 257. 1789; Wight, Prodr. Fl. Ind. with black glands. Stamens numerous, basally connate, Orient.: 99. 1834; Dyer in Hook.f., Fl. Brit. India 1: distinct, sometimes in fi ve free antepetalous fascicles; 253. 1874; Gamble, Man. Ind. Timb.: 48. 1881; Engl. anthers dehisce introrsely by longitudinal slits; usually in Engl. & Prantl, Nat. Pfl anzenfam. 3(6): 215. 1895; same colour as but shorter than petals, stamen length Prain, Bengal Pl. 1: 243. 1903; Gagnep. in Lecomte, varies within fascicles, longest stamens on the outside Fl. Indo-Chine 1(4): 284. 1910; Craib, Bull. Misc. of the fascicle ring. Ovary superior; unilocular; styles Inform. Kew: 66. 1913; Gamble, Fl. Madras: 69. 2–5; stigmas distinct, persistent, varies in size and 1915; Haines, Bot. Bihar Orissa: 2: 51. 1921; Ridl., shape from narrowly elliptic to ovoid to globose. Fl. Malay Penins. 1:151. 1922; Craib, Fl. Siam. Fruit a woody or coriaceous septicidal capsule. Seeds 1:110. 1925; R.Keller in Engl. & Prantl, Nat. small, globose to ovoid or ellipsoid; sometimes very Pfl anzenfam. ed. 2, 25:175. 1925, pro parte excl. numerous (over 1000), winged seeds present on sect. IV Elodea; Gagnep. in Humbert, Suppl. Fl. species with dehiscent fruit. Indo-Chine 1(3): 248. 1943; Backer & Bakh.f., Fl. Distribution.— 488 species worldwide (absent Java 1: 382. 1963; N.Robson, J. Roy. Hort. Soc. 95: from tropical lowlands, southern oceanic islands, 482. 1970; N.Robson in Fl. Males., Ser. 1, Spermat. hot and cold deserts); 6 species in Thailand. 8: 4–6. 1974; N.Robson, Bull. Brit. Mus. (Nat. Hist.), In a number of cases the acknowledged World Bot. 12(4): 163–325. 1985; N.Robson, Taxon 39: expert on Hypericum, Robson, has used the term 134–135. 1990; N.Robson, Syst. Biodivers. 4(1): ‘holotype’: we have almost always followed his lead, 19–98. 2005; H.W.Li & N.Robson, Fl. China 13: but believe in a number of cases that Art. 9.10 2–35. 2007. Type species: Hypericum perforatum L. (Turland et al. 2018) should be applied [Art 9.10 (lectotype designated by Britton & Brown, Ill. Fl. deals with the use of ‘a term as defi ned in the Code N. U.S. ed. 2, 2: 529. 1913). as denoting a type, in a sense other than that in which Herbs to occasionally trees up to 12 m high; it is so defi ned’. The article indicates that this is to herbs spreading by means of horizontal runners be ‘treated as an error to be corrected (for example, before becoming erect or ascending. Stems eventually the use of the term lectotype to denote what is in terete, green to red when young and brown to black fact a neotype)’] We believe that Robson’s designa- when older, fi ssured when older, stems can turn grey tion of a holotype is usually correctable under Art. and the cork layers fl ake off in strips. Leaves sessile 9.11 (Turland et al. 2018) to lectotype. In many cases or subsessile, opposite, entire, venation present, the use of the term ‘holotype’ should be construed sometimes parallel, glands of two types, dark glands as the fi rst stage of a necessary lectotypifi cation and are blackish-red cells containing a wax and pale that, therefore, second a step lectotypifi cation (Art. glands are schizogenous intercellular spaces lined 9.17; Turland et al. 2018) is required. We are, however, by cells which secrete an oil. Infl orescences solitary, in most cases unwilling to undertake this, preferring in three’s, cymose or umbellate. Calyx 5 sepals, it to be undertaken by an expert on this genus in its basally connected, shape very variable with the same worldwide context. glands as the leaves, margins entire. Corolla 5 petals,

KEY TO THE SPECIES 1. Herbs with fl owers under 2.5 cm across 2. Sepals usually toothed with gland-tipped teeth; stems terete; ovary 3-celled 1. H. elodeoides 2. Sepals entire, with/without glands; stems quadrangular; ovary 1-celled 4. H. japonicum 1. Shrubs up to 3m tall, with fl owers 5–8 cm across 3. Sepals 4; petals 4; leaves oblong or ovate-oblong to elliptic with clear intramarginal vein 6. H. siamense 3. Sepals 5; petals 5; leaves oblong, ovate-oblong, elliptic, lanceolate or oblanceolate without intramarginal vein

SW 11611-p162-216-G8.indd 194 1/16/62 BE 5:26 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 195 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

4. Sepals entire or denticulate; ovate, oblong or elliptic; stems spreading, 2-lined 5. H. patulum 4. Sepals entire; stems arching or erect 5. Leaves variable: lanceolate, oblanceolate, elliptic, oblong, obtuse; sepals with obvious longitudinal striations, ± equal, imbricate; stems arching, young branches compressed; stamens 60–80 3. H. hookerianum 5. Leaves narrowly lanceolate to elliptic; sepals without obvious striations, not equal, not imbricate; stems erect, young branches thin; stamens 40–60 2. H. henryi subsp. hancockii

1. Hypericum elodeoides Choisy in DC., Prodr. 1: teeth, apex slightly pointed to round, bracts with 552. 1824. Type: Nepal. Wallich 4812A (?holotype black glandular hairs. Corolla petals 6–13 mm long, G [GDC] [00210401!]; isotypes BM [BM000521690!], elliptic to oblanceolate, caducous, yellow/cream, K!, S [S11-31777!], SING. with longitudinal lines present, apex slightly pointed. — Hypericum napaulense Choisy in DC., Prodr. 1: Stamens in 3 fascicles. Ovary 3-locular; styles 552. 1824; Dyer in Hook.f., Fl. Brit. India 1: 253. 0.9–1.3 mm. Fruit ovoid to elliptic, 3-celled, green. 1874; Gagnep. in Lecomte, Fl. Indo-Chine 1(4): 284. Seeds 0.5–0.6 mm long. 1910; Craib, Fl. Siam. 1:111. 1925; Gagnep. in Thailand.— (Map 2.1.1): NORTHERN: Chiang Humbert, Suppl. Fl. Indo-Chine 1(3): 251. 1943; Mai [Summit of Doi Inthanon, 19 Aug. 1927, Garrett H.W.Li & Li Yan-hui in Li Hsiwen, Fl. Reipubl. 421 (ABD, BM, E, K, L, P, TCD); Doi Chiang Dao, Popularis Sin. 50(2): 1–112. 1990; H.W.Li & 23 Oct. 1992, Pooma 703 (BKF); Doi Pa Kao, 8 May N.Robson, Fl. China 13: 24. 2007. Type: Nepal, 1921, Kerr 5392 (ABD, BK, BM, K); Doi Inthanon, Gossain Than, Wallich s.n. (?holotype: G [GDC] 30 Oct. 1962, Smitinand et al. 7643 (BKF); Doi [G00210678!]; isotypes BM!, K!). Chiang Dao, 11 Dec. 1987, Santisuk s.n. (BKF)]. — Hypericum pallens D.Don, Prodr. Fl. Nepal.: 219 Distribution.— India, Bhutan, Nepal, South (Jan.–Feb.) 1825. Type: West Himalaya, Wallich China, Myanmar. 4814 (holotype K!). Ecology and phenology.— Marshy areas, on — Hypericum setosum Wall., Numer. List. [Wallich] limestone from 1400 to 2500 m elevation. Flowering: n. 4814. 1828, nom. nud. May–August and October–December. — Hypericum wightianum Wall., Numer. List. Conservation.— IUCN Regional (Thailand) [Wallich] n. 4818. 1828, nom. nud; Wall. ex Wight Status: Least Concern (LC). & Arn., Prodr. Fl. Ind. Orient.: 99. 1834; Gamble, Notes.— 1. We have carefully chosen the most Fl. Madras: 69. 1915. Type: India, Neelgherries, complete available Wight collection as the lectotype Wight 3366 (lectotype E [E00174238!], designated of Hypericum wightianum over any Wallich material here; isolectotypes BM [BM000551503!], E as Wight (1834) states clearly that he had not seen [E00174230!], K [K000677235!]). Wallich’s material. His descriptionn can only, therefore, Shrubs or perennial herbs up to 1 m tall; be based on Wight 336. glabrous; decumbent. Branchlets terete, slender, 2. Hypericum elodeoides is a widespread striations present, light golden-brown to dark red- and variable species sometimes separated from brown. Leaves sessile; blade 0.7–2.5 × 0.4–1.5 cm, H. napaulense on the basis of its glandular petal elliptic to oblanceolate, base cuneate to slightly taper- margin, terete stem, acute leaves and ribbed sepals. ing, apex slightly acute to rounded to subacuminate, Robson et al. (2014) unite them. parallelodromous veins faint; round (black) dots sometimes present on upper surface, light green to tan on upper surface and dark brown/black on lower 2. Hypericum henryi H.Lév. & Vaniot subsp. surface, glabrous, delicate, smooth, midrib depressed hancockii N.Robson, Bull. Brit. Mus. (Nat. Hist.), on upper surface and pronounced on lower surface. Bot. 12(4): 261 1985; H.W.Li & N.Robson, Fl. Infl orescence pedicel up to 5 cm long, solitary or China 13: 12. 2007. Type: China, Yunnan, Hancock corymbose cymes with up to 17 fl owers, thin and 116 (holotype K!; isotype K!). delicate, spine-like hairs present on all stalks of — Hypericum garrettii Craib in Bull. Misc. Inform. infl orescences. Calyx sepals, 0.5–0.6 × 0.1–0.2 cm, Kew 1913; 66. 1913, pro parte, quoad Kerr 6300). oblanceolate, persistent, some with gland-tipped

SW 11611-p162-216-G8.indd 195 1/16/62 BE 5:27 PM 196 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

— Hypericum garrettii var. ovatum Craib, Fl. Siam. Distribution.— Myanmar, Vietnam, China, 1:111. 1925. Type: Thailand, Chiang Mai, Kerr 5188 Sumatra. (lectotype ABD [ABDUH:2/516!]; isolectotypes: Ecology and phenology.— Evergreen forest, BK!, BM [BM000617086!], E [E00209541!], K mountain, exposed/open ridges from 280 to 2560 m [K000380029!]: designation of a holotype by elevation. Flowering: All year. Fruiting: July– Robson [1985] is correctable under Art. 9.11 November. [Turland et al. 2018] to lectotype). Conservation.— IUCN Regional (Thailand) Shrubs to 3 m tall. Branchlets terete, erect, Status: Least Concern (LC). slender, smooth, new branches thin, red-brown, internodes 1–2 cm long. Leaves sessile; blade 1.5–4.3 × 0.4–2 cm, lanceolate to elliptic, green to golden 3. Hypericum hookerianum Wight & Arn., Prodr. yellow to red/brown, some 2-toned, sometimes Fl. Ind. Orient. 1: 99. 1834; Dyer in Hook.f., Fl. glaucous, glabrous, delicate, base cuneate to rounded, Brit. India 1: 254. 1874; Gamble, Fl. Madras 69. apex acute to cuspidate to obtuse, midrib depressed 1915; R.Keller in Engl. & Prant, Nat. Pfl anzenfam. on upper surface and pronounced on lower surface, ed. 2, 21: 176. 1925; Y.Kimura in Hara, Fl. E. fading slightly towards apex, but still visible, Himalaya 210. 1966; K.M.Matthew in Rec. Bot. secondary veins visible, intersecondary and tertiary Surv. India 20: 45. 1969; N.Robson, J. Roy. Hort. veins faint, secondary veins not arising from same Soc. 95: 490. 1970; N.Robson in Hara & Williams, point on opposite sides of midrib, some leaves Enum. Fl. Pl. Nepal 2: 61. 1979; N.Robson, Bull. trinerved, black glands present. Inflorescence Brit. Mus. (Nat. Hist.) Bot. 12 (4): 255. 1985; pedicels 0.3–1.5 cm long, branches terete, paniculate, H.W.Li & N.Robson, Fl. China 13: 10. 2007. ≡? 3 flowers per corymb. Calyx sepals, 6 × 4 mm; Brathys nepalensis Blume, Mus. Bot. 2: 19. 1852. unequal, obovate to ovate or elliptic, orange. Corolla ≡ Norysca hookeriana (Wight & Arn.) Wight, ll. petals 8–25 × 6–15 mm; yellow, ovate. Stamens Ind. Bot. 1: 113. 1840; Blume, Mus. Bot. 2: 22. 40–60 per fascicle, anthers longest 5–13 mm, usually 1856; Y.Kimura in Nakai & Honda, Nov. Fl. Jap. half as long as petals, round, yellow. Ovary ovoid, 10: 98. 1951.Type: India, Wight 332 (lectotype K styles 4, 4–6 mm long, protruding. Seeds brown. [K000677209!]; isolectotypes E [E00174236!, E00174237!): designation of a holotype by Robson Thailand.— (Map 2.1.2): NORTHERN: Chiang [1985] is correctable under Art. 9.11 [Turland et al. Mai [Doi Inthanon, trail to the summit, Chom Thong 2018] to lectotype. District, 22 Mar. 2007, Byrne 9 (TCD); Kio Mae Pan Nature Trail, Doi Inthanon, 22 Mar. 2007, Byrne — Hypericum garrettii Craib in Bull. Misc. Inform. 10 (TCD); ibid., 22 Mar. 2007, Byrne 11 (TCD); Kew 66. 1913, (pro parte); Craib, Fl. Siam. 1:110. Huai Mae, Klang Pat, Pha Mon, 23 Feb.1982, 1925; Gagnep. in Humbert, Suppl. Fl. Indo-Chine Suvarmasuddhi 318 (BKF); Doi Inthanon, Pa Ngem 1(3): 248. 1943. Type: Thailand, Chiang Mai, Doi N Peak, 14 July 1922, Kerr 6300 (ABD, BM, K); Inthanon, Garrett 67 (lectotype K [K000380030!], higher elevation of Doi Inthanon,7 Jan. 1983, designated here; isolectotypes BM [BM000617687!], Koyama et al. T-32090 (BKF); Pa Ngem, 6 Apr. E [E00209544!]). 1925, Winit 1350 (ABD, BKF, K); Doi Pha Hom Shrubs 1–3 m tall. Branchlets terete, arching, Pok, Muang Fang, 2 Apr. 1921, Kerr 5188 (ABD, stems brown, smooth, slender, scars present, young BM, E, K, TCD); 12 Feb. 1983, Koyama et al. branches compressed. Leaves sessile-subsessile; T-33411 (BKF); 12 Feb. 1983, Koyama et al. blade 3.3–6.3 × 1.3–2.4 cm, lanceolate, oblanceolate, T-33438 (BKF); 23 Mar. 1965, Phengklai 973 (C, elliptic, oblong or obtuse, base cuneate, apex some- E); Feb. 1978, Løjtnant & Niyomdham 1666 (AAU); what acute to acuminate, delicate, papyraceous, 13 Sept. 1967, Iwatsuki et al. 9690 (AAU, BKF, E, glabrous, few pellucid dots present, vibrant green to K, L)]; Chiang Rai [Doi Tung, NW side of Pa Hung, red to dull red/black, midrib depressed on both 25 Dec. 2006, Maxwell 06-10066 (CMUB)]; Nakhon surfaces, adaxial midrib colour very variable, red, Sawan [Khlong Lan, Mo Ko Chu Mountain, 3 Jan. yellow/white or dark red, abaxial midrib colourr red or 1999, van de Bult 209 (BKF, CMUB)]; NORTH- brown, secondary, intersecondary and tertiary veins EASTERN: Sakhon Nakhon [An Gop, Phu Phan NP, visible, no intramarginal veins. Infl orescence terminal, 26 Feb. 1993, Chantaranothai et al. 973 (TCD)].

SW 11611-p162-216-G8.indd 196 1/16/62 BE 5:27 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 197 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

1–5(–10)- flowered, from apical node, pedicels et al. 90-6266 (K); Doi Inthanon NP, summit of Doi 0.3–1.6 cm long. Calyx sepals 3.5–9 × 2–6.5 mm, Inthanon, 23 Sept. 1991, Maxwell 91-7766 (AAU, E, oblong, elliptic to obovate-spathulate, rounded, entire P); higher elevation of Doi Inthanon, Payap District, ovate to obovate, ± imbricate, green, red/brown, 19 Dec. 1965, Tagawa et al. T-28566 (BKF); near longitudinal veins present. Corolla petals 9–18 × Pagoda, the west side of the main route, Doi 5–12 mm, obovate, entire, caducous bright yellow Inthanon, 4 Aug. 1988, Koyama T-61088 (BKF); to golden yellow, longitudinal veins present. Stamens Kio Mae Pan, Doi Inthanon, 1998, BGO Staff s.n. 60–80 per fascicle, anthers oblong, 4–6(–8) mm (QBG)]; Unknown location [1987, Robson s.n. long, approximately ¼ – ½ as long as petals, anthers (AAU, BKF)]. bright yellow to orange-yellow. Ovary 3–7 × 4–6 mm Distribution.— Bhutan, Myanmar, China, long, styles 5, longer than anthers, ovoid to globose. India (Madras), Nepal, Vietnam. Fruit green, ovoid. Seeds ca 0.5 mm long, elliptic, dark brown to red. Ecology and phenology.— Primary evergreen forest, marshy ground, open savannah, swamp from Distribution.— Thailand (Map 2.1.3): 1850 to 2595 m elevation. Flowering: April–July. NORTHERN: Mae Hong Son [Doi Chang, 19 Feb. Fruiting: September–October. 1968, Hansen & Smitinand 12674 (AAU, BKF, E, K, L, P)]; Chiang Mai [Doi Inthanon NP, 29 Jan. Vernacular.— Bua thong (บัวทอง) (BGO Staff 1996, BGO Staff 24 (QBG); Doi Inthanon, Pa Ngem 7117). N Peak, 12 Oct. 1910, Garrett 67 (BM, E, K); Pa Conservation.— IUCN Regional (Thailand) Ngem N Rocks, Doi Inthanon, Garrett 70 (BM); Status: Least Concern (LC). Kio Mae Pan, 15 Oct. 2003, Mattapha 394 (KKU); Doi Inthanon NP, 3 Aug. 1984, W.N. 548 (BKF); 4. Thunb. in Murray, Syst. Doi Pha Hom Pok, 23 Mar. 1965, Phengklai & Veg., ed. 14: 702. (May–June) 1784; Thunb., Fl. Sangkhachand 973 (BKF); Doi Inthanon NP, 12 Jap.: 295. (Aug.) 1784; Choisy, Prodr. Monogr. Aug. 1995, Pooma 1072 (BKF); Doi Pha Hom Pok, Hyperic.: 48. 1821; Blume, Bijdr. Fl. Ned. Ind. 5: 23 Nov. 1998, Suksathan 1488 (QBG): Western 143. 1825; D. Don, Prodr. Fl. Nepal.: 219. 1825; fl ank of Doi Inthanon, Mae Pau, 7 Dec. 1969, van Wight, Prodr. Fl. Ind. Orient.: 99. 1834; Choisy in Beusekom & Phengklai 23977 (AAU, BKF, C, E, Zoll., Syst. Verz.: 151. 1854; Miq., Ann. Mus. Bot. P); Inthanon NP, 8 Nov. 1992, Balick & Nanakorn Lugduno-Batavi 2: 259. 1866; Dyer in Hook.f., Fl. 3410 (AAU); Doi Inthanon, Chom Thong 16 Sept. Brit. India 1: 263. 1874; Kurz, J. Asiat. Soc. Bengal, 1995, Nanakorn et al. 43966 (QBG); Mae Chaem Pt. 2, Nat. Hist. 43: 84. 1874; Trimen, Fl. Ceylon 1: District: Top area of Doi Inthanon in Doi Inthanon 93. 1893; Engl. in Engl. & Prantl, Nat. Pfl anzenfam. NP, 20 Dec. 1998, Konta et al. 4880 (BKF); Kio 3(6): 215. 1895; Bailey, Queensl. Fl. 1: 101. 1899; Mae Pan, Doi Inthanon, 21 Dec. 1997, Niyomdham Prain, Bengal Pl. 1: 243. 1903; Gagnep. in Lecomte, 5244 (BKF); Doi Inthanon, 2 May 1921, Kerr 5312 Fl. Indo-Chine 1(4): 284. 1910; Koord.-Schum., (ABD, BK, BM, K, TCD); Doi Inthanon, Nov. 1986, Syst. Verz.: 187. 1911–1914; Gamble, Fl. Madras: Phengklai & Smitinand 6072 (BKF); ibid., 21 Aug. 69. 1915; Haines, Bot. Bihar Orissa. 2: 51 1921; 1996, BGO Staff 71177 (QBG); Doi Chiang Dao, 3 Ridl., Fl. Malay Penins. 1: 151. 1922; Lauterb., Bot. Dec. 1961, Smitinand & Anderson 7303 (BKF); Doi Jahrb. Syst. 5: 58. 1922; Craib, Fl. Siam. 1: 111. Inthanon NP, 4 Aug. 1988, Phengklai et al. 7416 1925; Ridl., Bull. Misc. Inform. Kew: 59. 1926; (K); ibid., Phengklai et al. 7484 (BKF); ibid., 30 M.R.Hend., Gard. Bull. Straits Settlem. 4: 222. Oct. 1962, Smitinand et al. 7645 (BKF); Kio Mae 1928; Corner, Wayside Trees of Malaya: 324. 1940; Pan, Doi Inthanon, 28 Nov. 1996, BGO Staff 7928 Y.Kimura, Bot. Mag. (Tokyo) 54: 87. 1940; Gagnep. (QBG); Doi Inthanon, 31 Dec. 1974, Geesink et al. in Humbert, Suppl. Fl. Indo-Chine 1(3): 248. 1943; 7998 (K, L); Kio Mae Pan, Doi Inthanon, 20 Dec. Backer & Bakh.f., Fl. Java 1: 382. 1963; Ohwi, Fl. 1998, Phengklai et al. 11394 (BKF); Doi Chiang Jap.: 631. 1965; N.Robson in Fl. Males., Ser. 1, Dao, 10 Nov. 1962, Smitinand & Poore 30238 Spermat. 8: 4–6. 1974; N.Robson: Taxon 39: 134– (AAU); Doi Inthanon, 28 Nov. 1993, Larsen et al. 135. 1990; H.W.Li & N.Robson, Fl. China 13: 34. 44980 (AAU); Doi Inthanon NP, forest trail at 42 2007. ≡ Sarothra japonica (Thunb.) Y.Kimura et al., km along summit road, 7 Oct. 1990, Chantaranothai

SW 11611-p162-216-G8.indd 197 1/16/62 BE 5:27 PM 198 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Map 1.4.1 Distribution of Mesua ferrea in Thailand. Map 2.1.1 Distribution of Hypericum elodeoides in Thailand.

Map 2.1.2 Distribution of Hypericum henryi subsp. hancockii Map 2.1.3 Distribution of Hypericum hookerianum in Thailand. in Thailand.

SW 11611-p162-216-G8.indd 198 1/16/62 BE 5:24 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 199 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Nova Fl. Jap. 10: 235. 1951. Type: Japan, Honshu, style and ovary light green; styles 3. Fruit 3–5 mm Thunberg s.n. (lectotype UPS!, isolectotype: BM!: long, globose, red. Seeds many, approximatelyy 0.5 mm designation of a holotype by Robson [1985] is long, cylindrical, brown or yellow, longitudinally correctable under Art. 9.11 [Turland et al. 2018] to ribbed. lectotype). Thailand.— (Map 2.1.4): NORTHERN: Mae Hong — Hypericum chinense Osbeck, Dagb. Ostind. Resa Son [Khun Yuam, 4 Sept. 1974, Larsen & Larsen 244. 1757, nom. rejic. prop. Type: China, Osbeck 34138 (AAU, K, P)]; Chiang Mai [Doi Inthanon, 23 s.n. (holotype S!) (see note at end of description Feb. 1983, Unknown collector 28 (K); Mai Muang about rejection of the name). Nao Arboretum, 12 Apr. 2001, Sankamethawee 157 — Hypericum pusillum Choisy, Prod. Monogr. (A); Chiang Dao, 7 July 1998, Pongamornkul 161 Hyperic.: 50. 1821. Type: not located. (QBG); Muang Fang, 6 June 1973, Sadakorm 227 (BK); Doi Suthep, 4 Mar. 1966, Chermsirivathana & — Brathys laxa Blume., Mus. Bot. 2: 19. 1852. ≡ Boonkird 434 (BK); Doi Suthep, 23 Mar. 1905, Sarothra laxa (Blume ) Y.Kimura, Nakai & Honda, Unknown collector 485 (P); Doi Inthanon, 24 Mar. Nova Fl. Jap. 10: 241. 1951.Type: Japan, Pierot 252 1967, Chermsirivathana & Boonkird 730 (BK); ibid. (lectotype: L [L0012247!], in sched.. Possible 12 Aug. 1995, Pooma 1070 (BKF); Ob Luang table- syntype: Japan, Keiske s.n. (L [L0012248!]). land, along road from Bo Luang to Om Koi, 12 June — Brathys japonica Blume, Mus. Bot. 2: 19. 1852. 1968, van Beusekom & Phengklai 1188 (AAU, BKF, Type: Japan, Siebold s.n. (holotype L!). C, E, K, L, P); Doi Suthep, 23 Mar. 1937, Deignan — Hypericum calycatum Jacquem. ex Dyer in 1526 (A); west of Fang, 25 Feb. 1958, Sørensen et al. Hook.f., Fl. Brit. India 1: 256. 1874. Type: Japan, 1663 (C); 10 km south of Bo Luang along the Om Honshu, Thunberg s.n. (holotype? UPS!; isotype Koi trail, 2 July 1968, Larsen et al. 1942 (AAU, BKF, BM!). C, E, L, P); Doi Suthep, 14 Mar. 1958, Sørensen et al. 2701 (C); Mae Sanam, 27 June 1978, Phengklai — Hypericum thunbergii Franch. & Sav., Enum. Pl. et al. 4155 (BKF); Om Koi, 28 June 1978, Phengklai Jap. 2: 300. 1878. Types: Japan, Yokosaka, Savatier et al. 41677 (BKF); Doi Suthep, 23 Mar. 1905, Hosseus 158 (syntypes P [P04637564!, P04637565!, 485 (BM, K); Bo Luang, 7 June 1973, Geesink et al. P04637566!, P04637568!, P04637570!], Japan, 5833 (C, E, L, P); ibid., 12 June 1973, Geesink et al. Sagone 3411 (not located). 58677 (BKF, C, E, K, L, P); Chom Thong, 30 May Annual or perennial herbs, with erect or 1979, Vidal et al. 6198 (P); Ban Tha Ma Kiang, Phrao, decumbent branched stems, 15–45 cm long; stems 14 June 1997, Nanakorn et al. 9204 (QBG); Om Koi, green, approximately 3 mm wide at base, glabrous. 20 Jan. 1966, Hansen et al. 10843 (C, K, L); near Bo Branchlets slender, stems quadrangular. Leaves Luang at 37 km, Hansen et al. 110177 (C); Doi Suthep, sessile, papyraceous, decussate, sometimes glaucous east side, above Khonthathan Falls, Muang, 26 Dec. beneath; blade 6–10 × 2–6 mm, ovate to lanceolate, 1988, Maxwell 88-1408 (L); Ban Pa Pae, Mae Taeng base cordate-amplexicaul some auricled, apex District, 7 Nov. 1990, Maxwell 90-1244 (A); Ban Tha rounded or obtuse, midrib green adaxially, light Ma Kiang, San Sai Subdistrict, Phrao Valley, Phrao, green abaxially, side veins 1–8 each side. 1 Aug. 1992, Maxwell 92-403 (A, E, P); summit of Infl orescence 1–30-fl owered, 6–10 mm across, axes Doi Poa Lohn, Mae Seuk Subdistrict, Mae Jam green, pedicels up to 1 cm long, fl owers terminal, District, 14 Jan. 1997, Maxwell 97-38 (A, BKF); Doi mostly elongated infl orescences, sometimes solitary. Inthanon, 23 July 1988, Tsugaru T-61735 (A, AAU, Calyx sepals ± free, 2–6 mm long, elliptic to oblong- BKF); San Pa Tong, 10 Jan. 1986, Paiseeksantivatana lanceolate, subequal to unequal, apex acute to Y 1742-866 (BK)]; Chiang Rai [Chiang Khien, 3 Mar. rounded, 3–5 longitudinal veins present, entire, light 1912, Kerr 2486 (BM, E, K, TCD); ibid., 3 Mar. green, persistent. Corolla petals, orange-yellow; 1912, Kerr 2486A (BM); Ban Dai, Sidonmun subdis- oblanceolate, obovate or elliptic, 2–5 × 1–1.5 mm trict, group 3, Chiang Saen, 6 Mar. 1989, Maxwell long; persistent. Stamens 7–25 stamens per fascicle, 89-312 (L)]; Nan [Doi Phu Kha NP, Pua, 4 Nov. 1999, ca 3 mm long, persistent, anthers yellow, fi laments Srisanga 1119 (QBG); ibid., 25 May 2000, Srisanga light orange. Ovary ovoid to subglobose; stigma, 1399 (QBG)]; Tak [Huai Khek, Lom Sak 23 Feb.

SW 11611-p162-216-G8.indd 199 1/16/62 BE 5:27 PM 200 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1964, Hansen et al. 11236 (BKF, C)]; NORTH- 5. Thunb. in Murray, Syst. EASTERN: Loei [Phu Kradung, 19 Mar. 1948, Veg., ed. 14: 700. (May–June) 1784; Thunb., Fl. Juntabendlham 149 (BK); ibid., 19 Mar. 1974, Jap.: 295, t. 17. (Aug.) 1784; Curtis, Bot. Mag. 50: Chermsirivathana & Boonkird 1885 (BK); ibid., 19 t. 5693. 1823; Choisy in DC., Prod. 1: 545. 1824; Mar. 1974, Chermsirivathana & Boonkird 1889 (BK); Dyer in Hook.f., Fl. Brit. India 1: 254. 1874; ibid., 16 Jan. 1966, Hennipman 3704 (BKF, C, L); Gamble, Man. Ind. Timb.: 48. 1881; Gamble in ibid., 1970, Charoenphol et al. 4808 (AAU)]; Sakhon Humbert, Suppl. Fl. Indo-Chine 1(3): 248. 1943; Nakhon [Phu Phan NP, ca 100 km west of Park HQ, Zollinger, Syst. Verz. 1–2: 151. 1854; Maxim. in 19 Oct. 1990, Chantaranothai & Parnell 90/799 Bull. Acad. Imp. Sci. Saint-Pétersbourg 27: 429. (TCD)]; EASTERN: Chaiyaphum [Thung Kra Mang, 1882; Hemsl., J. Linn. Soc., Bot. 23: 73. 1886; Phu Khieo, 9 Aug. 1972, Larsen et al. 31604 (AAU, Kuntze, Rev. Gen. Bot. 1: 60. 1891; Koehne, Deut. K, L, P)]; SOUTH-EASTERN: Chanthaburi [Pong Nam Dendrol.: 415. 1893; R.Keller, Bull. Herb. Boissier Ron, 11 Mar. 1956, Smitinand 32866 (BKF, K, L, P); 5: 638. 1897; Diels, Bot. Jahrb. Syst. 29: 476. 1900; Khao Phloi Waen, 10 Jan. 1930, Kerr 18049 (K, H.Lév., Bull. Soc. Bot. France 53: 499. 1906; TCD)]; PENINSULAR: Surat Thani [Thung Luang, 1 R.Keller in Engl. & Prantl, Nat. Pfl anzenfam. ed. 2, Apr. 1927, Kerr 12512 (ABD, BM, K)]; Unknown 21: 176. 1925; Rehder, J. Arnold Arbor. 15: 100. location [Unknown collector 49 (KKU)] 1934; Y.Kimura, Bot. Mag. (Tokyo) 54: 88. 1940; Distribution.— India, Sri Lanka, Bhutan, Y.Kimura in Nakai & Honda, Nov. Fl. Jap. 10; 100, Nepal, Myanmar, Cambodia, Laos, Vietnam, China, f. 41. 1951; Lauener, Notes Roy. Bot. Gard. S Korea, Taiwan, Russia, Japan, Malaysia, Singapore, Edinburgh 27: 4. 1966; N.Robson, J. Roy. Hort. Indonesia (Sumatra to Irian Jaya), Philippines, Soc. 95: 491. 1970; N.Robson, Bull. Brit. Mus. (Nat. South-East and South Australia, New Zealand. Hist.), Bot. 12(4): 261 1985; H.W.Li & N.Robson, Fl. China 13: 13. 2007. ≡ Komana patula (Thunb. Ecology and phenology.— Deciduous forest, in Murray) Y.Kimura ex Honda, Nom. Pl. Japonic. bamboo forest, wet grassland, bog, savannah from 509. 1939. ≡ Norysca patula (Thunb. in Murray) 50 to 2500 m elevation. Flowering: April–June Voigt, Hort. Suburb. Calcutt. 90. 1845. ≡ Eremanthe (October). Fruiting: Apr.–June (November). patula (Thunb. in Murray) K.Koch, Hort. Dendrol. Vernacular.— Bua thong (บวทองั ) (Pongamornkul 65. 1853. ≡ Komana patula (Thunb. in Murray) 161). Y.Kimura ex Hisauti, Kikasyokubutsu, 179. 1950; cf. Y.Kimura, Nakai & Honda, Nova Fl. Jap. 10: 97, 99. Conservation.— IUCN Regional (Thailand) 1951.Type: Japan, Thunberg s.n. (?holotype UPS!). Status: Least Concern (LC). — Hypericum uralum sensu Hance in J. Bot. 16: Notes.— The name Hypericum chinense 104. 1878, non. H. uralum Buch.-Ham. ex D.Don Osbeck has never been used since it was published. in Bot. Mag. 50: t. 2375. 1823. Type: Nepal, Merrill (1916) pointed out its priority over H. japoni- Narainhetty, Hamilton s.n. (lectotype BM!; desig- cum Thunb. in Murray but thought the name nated by Robson in Robson et al. [2014]). H. chinense L. applied to the same species. Hypericum chinense L. is a cultivated shrub and this — Hypericum argyi H.Lév. & Vaniot, Bull. Soc. name has been used almost exclusively in botanical Bot. France 54: 591. 1908; H. Lév., Fl. Kouy- and horticultural literature until 1985, when Robson Tcheou. 198: 1914; H. Lév., Mem. Real. Acad. Ci. (1985) pointed out the above synonymy. Since then Barcelona 3(12): 553. 1916. Type: China, d’Argy H. chinense L. has been replaced by H. monogynum s.n. (?holotype E!; isotype A [A00067732!]). L., and H. japonicum Thunb. in Murray has been Shrubs to 3 m tall; with spreading branches, conserved over H. chinense Osbeck; see Robson greyish-brown. Branchlets terete, becoming 2-lined, (1990) for more information regarding the name. thick, rough, dark brown to red-orange, scarring Robson (1985) uses the author citation ‘Thunb. present. Leaves sessile to petioled, if petioled, up to ex Murray’ as the authority for a number of species 0.5–2 mm long, in whorls at top of branch and sparse of Hypericum, but Bartholomew et al. (1997) argue along stems; blade 1.5–6.0 × 0.5–3.0 cm, lanceolate correctly that the citation should be Thunb. in to oblanceolate or ovate, oblong or elliptic, base Murray. cuneate, apex acuminate to obtuse, light brown to

SW 11611-p162-216-G8.indd 200 1/16/62 BE 5:27 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 201 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

dark red smooth, delicate, margins undulate, midrib Corolla petals 2–2.4 × 1.2–1.7 cm, obovate, yellow, visible and prominent on upper surface, black on some tinged red on dorsal side, longitudinal striations both surfaces, venation ± obscure, without intramar- present. Stamens approximately 45–50 stamens per ginal veins. Infl orescence pedicels 0.2–0.7 cm long, fascicle, anthers 10–11 mm long, oblong, yellow to 1–15-fl owered. Calyx sepals ± free, 5–10 × 3.5–7 mm orange. Ovary ovoid; styles 2.5–6 mm long. Fruit ovate to elliptic or oblong, imbricate, unequal or ovoid. Seeds approximately 1 mm long, orange- subequal, entire to sometimes denticulate, sometimes brown. red. Corolla petals 1.2–1.8 × 1–1.4 cm, oblong to Distribution — Thailand (Map 2.1.6): obovate, golden yellow, longitudinally striated. NORTHERN: Chiang Mai [Doi Chiang Dao, 6 Jan. Stamens 50–70 stamens per fascicle, longest stamens 1922, Kerr s.n. (K); ibid., 10 July 1998, Pongamornkul 7–12 mm long; anthers bright yellow. Ovary ovoid; 171 (QBG); ibid., 8 July 1998, Pongamornkul 392 styles 4–5.5 mm long. Fruit ovoid. Seeds dark brown. (QBG); ibid., 17 Aug. 1963, Smitinand & Sleumer Thailand.— (Map 2.1.5): NORTHERN: Chiang 1055 (L); ibid., 8 July 1998, Suksathan 11177 (QBG); Mai [Doi Chiang Dao, 17 Aug. 1963, Smitinand et ibid., 15 Oct. 2000, Suksathan 2842 (QBG); ibid., al. 1055 (BKF); ibid., 3 Mar. 1979, Koyama et al. 15 Oct. 2000, Suksathan 2845 (QBG); ibid., 6 Jan. 15639 (AAU, BKF); the swampy place and its’ 1995, BGO Staff 2895 (QBG); ibid., 7 Jan. 1995, vicinity of the summit, Doi Inthanon, 4 Aug. 1988, BGO Staff 2919 (QBG); ibid., 10 Nov. 1962, Tsugaru T-61869 (A, AAU, BKF); Doi Chiang Dao, Smitinand et al. 7789 (BKF, E); ibid., 27 Sept. 1971, 6 Mar. 1922, Kerr s.n. (BK)]. Murata et al. T-15211 (AAU, K); ibid., 3 Dec. 1961, Distribution.— Myanmar, China, Japan, New Smitinand & Anderson 25915 (K); ibid., 10 Nov. Zealand, Sumatra, Vietnam. 1962, Smitinand & Poore 30328 (L); below the summit of Doi Chiang Dao, 14 Sept. 1967, Shimizu Ecology and phenology.— Open ridges from et al. T-10109 (BKF); Payap, Doi Chiang Dao, 7 1900 to 2520 m elevation. Flowering: May– Dec. 1965, Hennipman 32777 (L); Doi Chiang Dao, September. Fruiting: July–October. 26 Sept. 1971, Murata et al. T-15262 (BKF, K, P): Conservation.— IUCN Regional (Thailand) higher elevation of Doi Chiang Dao, 26 Sept. 1971, Status: Data Defi cient (DD). Murata et al. T-15280 (BKF); Doi Chiang Dao, 13 Nov. 1922, Kerr s.n. (ABD, BM)]. 6. Hypericum siamense N.Robson, Bull. Brit. Ecology and phenology.— Limestone ridges, Mus. (Nat. Hist.), Bot. 12 (4): 240 1985. Type: exposed cliff edges from 1800 to 2200 m elevation. Thailand, Chiang Mai, Doi Chiang Dao, Kerr s.n. Flowering: July–January. (holotype BM [BM000617688!]; isotype ABD Vernacular.— Bua thong (บัวทอง) (Pongamornkul [ABDUH2/517!]). 392). Bushy shrubs, 0.7–3 m tall or perennial herbs. Conservation.— IUCN Regional (Thailand) Branchlets terete, rigid, scars present, rough, inter- Status: Least Concern (LC). nodes 0.5–2 cm long, brown to red. Leaves subsessile to sessile; blade 2.3–3.6 × 0.6–1.8 cm, lanceolate to 2. CRATOXYLUM elliptic to oblong or ovate-oblong, base cuneate to tapering, apex acute or acuminate, delicate, some Blume, Verh. Batav. Genootsch. Kunsten 9: 172, glaucous, few black round glands present, midrib 174. 1823; Blume, Mus. Bot. 2: 17. 1852; Miq., Fl. depressed in upper surface and pronounced on lower Ned. Ind. 1(2): 515. 1859; Dyer in Hook.f., Fl. Brit. surface, midrib narrowing towards apex but still India 1: 257. 1874; Kurz, Forest Fl. Burma 1: 83. visible, brown/red on both surfaces, secondary veins 1877; Gamble, Man. Ind. Timb.: 48. 1881; King, J. and intramarginal veins visible, intersecondary veins Asiat Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 145. 1890; and tertiary veins obscure. Infl orescence pedicels up Engl. in Engl. & Prantl, Nat. Pfl anzenfam. 3(6): 215. to 1 cm long, solitary and axillary, or 1–5-fl owered 1895; Prain, Bengal Pl. 1: 245. 1903; Brandis, Indian and corymbiform. Calyx sepals free, 0.8–1.2 × Trees: 47. 1907; Gagnep. in Lecomte, Fl. Indo-Chine 0.3–0.6 cm; triangular to ovate; imbricate or refl exed 1(4): 287. 1910; Gamble, Fl. Madras: 70. 1915; Ridl., to spreading, unequal; longitudinal striations present. Fl. Malay Penins. 1: 152. 1922; Craib, Fl. Siam.

SW 11611-p162-216-G8.indd 201 1/16/62 BE 5:27 PM 202 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1: 111. 1925; Gagnep. in Humbert, Suppl. Fl. Indo- eucamptodromous. ,QÀRUHVFHQFHs terminal panicles, Chine 1(3): 251. 1943; Backer & Bakh.f., Fl. Java or short terminal or axillary cymes. Calyx sepals 5, 1: 383. 1963; Gogelein, Blumea 15: 454. 1967; Baas, persistent, coriaceous, usually accrescent. Corolla Blumea 18: 369. 1970; N.Robson in Fl. Males., Ser. petals 5, caducous to subpersistent, obovate, exceed- 1, Spermat. 8: 4–6. 1974; Corner, Wayside Trees of ing sepals, pink or white. Stamens 3 or 5 fascicles, 0DOD\DŁHornschuchia Blume, Cat. DGHOSKRXVSHUVLVWHQWDQWKHUVGRUVL¿[HGVOHQGHU Gew. Buitenzorg: 15. 1823: nom. illeg. Type species: sometimes with a brown dot on the connective. Cratoxylum hornschuchii Blume, nom. illeg.>Ł Ovary 3–celled, ovoid to ellipsoid; styles free; ovules Hornschuchia hypericina Blume] ±’Seeds oblong, winged all round or partly winged; embryo erect. Deciduous to evergreen trees or shrubs; all glabrous except C. formosum subsp. SUXQLÀRUXP; Distribution.— 6 species in Borneo, Myanmar, exudate yellow resinous sap turning black when dry. Cambodia, India, Indonesia, Laos, Malaysia, South Leaves opposite, sessile to petiolate, entire, papyra- and Southwest China, Philippines, Vietnam; 5 species ceous, pellucid dots usually present, base attenuate in Thailand. to subcordate, apex rounded to cuspidate, venation

KEY TO THE SPECIES )ORZHUVLQVPDOOFOXVWHUVRQROGOHDÀHVVWZLJV 2. Flowers white; leaves obovate, elliptic, lanceolate, oblanceolate or round; fruits 4–6 mm diam.; seeds 6–8 per loculus 3. C. formosum 2. Flowers dark red; leaves obovate, elliptic or oblong; fruits 2–4 mm diam.; seeds 5–6 per loculus 4. C. maingayi 1. Flowers on leafy twigs (YHUJUHHQWUHHXSWRPKLJKÀRZHUVLQWHUPLQDOS\UDPLGDOSDQLFOHVOHDYHVRERYDWHREODQFHRODWHREORQJRUHOOLSWLF 1. C. arborescens 'HFLGXRXVWUHHRUVKUXEXSWRPKLJKÀRZHUVLQVKRUWWHUPLQDOSDQLFOHVRUF\PHVOHDYHVREORQJHOOLSWLFRUODQFHRODWH /HDYHVVHVVLOHWRVXEVHVVLOHÀRZHUVQRWLQWKHD[LOVRIPDWXUHOHDYHVSHGLFHOVRIWHQORQJHUWKDQ±PP 5. C. neriifolium /HDYHVSHWLROHGVRPHÀRZHUVLQWKHD[LOVRIPDWXUHOHDYHVSHGLFHOV±PP 2. C. cochinchinense

1. Cratoxylum arborescens (Vahl) Blume, Mus. — Ancistrolobus glaucescens Turcz., Bull. Soc. Imp. Bot. 2: 17. 1852; Dyer in Hook.f., Fl. Brit. India 1: Naturalistes Moscou 31(1): 383. 1858. Type: Lobb 258. 1874; Kurz, Forest Fl. Burma 1: 83. 1877; King, 419 (lectotype: BM [BM000624819!], designated J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 146. here)]; isolectotypes BM [BM000624818], KW 1890; Ridl., Fl. Malay Penins. 1: 152. 1922; Engl. [KW001000762!], L!). LQ(QJO 3UDQWO1DW3ÀDQ]HQIDPHG   Evergreen trees to 45 m tall; outer bark scaly, 184. 1925; Corner, Gard. Bull. Straits Settlem. 10: reddish/brown, red hue obvious, glabrous; exudate 22, 31. 1939; Meijer, Bot. News Bull. Forest Dept., not recorded. Branchlets young shoots with continuous Sabah 7: 64. 1967; Gogelein, Blumea 15: 471. 1967; interpetiolar scars. Leaves petioled, 0.5–1.0 cm long; N.Robson in Fl. Males., Ser. 1, Spermat. 8: 4–6. red to green; blade 5–16 × 2–6 cm obovate-oblong to 1974; Corner, Wayside Trees of Malaya 1: 325. 1988; obovate-oblanceolate or elliptic, coriaceous, base Phengklai & Niyomdham, Flora in peat swamp areas cuneate to attenuate apex acute to cuspidate, numerous RI1DUDWKLZDWŁ Hypericum arborescens faint secondary veins present. ,QÀRUHVFHQFH a many- 9DKO6\PE%RWWŁ Vismia arbo- ÀRZHUHGWHUPLQDOS\UDPLGDOSDQLFOHXVXDOO\ERUQH rescens (Vahl) Choisy, Prodr. Monogr. Hyperic. 36. on leafy twigs; pedicels 0.2–0.4 cm long, slender, 1821. Type: India, 1778, König s.n. (holotype: C homostylous. Calyxx sepals 3.5–6 × 2–4.5 mm, obovate [10009464 !]. to oblong, concave. Corolla petals 4.5–7 × 2–5 mm, — Hypericum coccineum Wall., Numer. List. obdeltoid, usually caducous, deep red or pink or very [Wallich] n. 4823. 1828. nom. nud. rarely orange or white. Stamens 30–40 stamens per — Cratoxylum blancoii Blume, Mus. Bot. 2:17. 1852. fascicle, 4–5 mm long, green-yellow. Ovaryy 1.5–2 mm Type: Philippines, Luzon, Antipolo, Ramos s.n. long, pistil 3–5 mm. Fruit round to oblong; 7–9 × (Illustrative specimen/ neotype: Merrill 851, [US!]. 4 mm. Seeds 10–18 per loculus; ca 5 × 0.8 mm, narrowly oblong.

SW 11611-p162-216-G8.indd 202 1/21/62 BE 3:24 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 203 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Thailand.— (Map 2.2.1): PENINSULAR: — Hypericum biflorum Lam., Encycl. 4(1): 170. Narathiwat [Tak Bai, 22 July 1984, Niyomdham 23 1797; Hook & Arnott, Bot. Beechey Voy. 173. 1833. (AAU, K); To Mo, Ban Bacho, 22 Apr. 1931, Type: China. collector unknown (P-LAM? not Lakshnakara 758 (BK, BM, K); Tak Bai, 25 May located. ?Neotype: ?Gogelin 1967, publication not 1984, Niyomdham 829 (A, AAU, BKF, K); Tak Bai, located, China, Bladh s.n. LD not seen, S! Ku Chum, 15 Sept. 1987, Niyomdham & Sriboonma [S-11-34218]). 1601 (AAU, BKF, E, K, L, P); Ben Yuan Yahng, — Hypericum pulchellum Wall., Numer. List. Su-ngai Padi, 7 June 1987, Maxwell 87-532 (A, [Wallich] n. 4821. 1828. nom. nud. BKF, L, PSU)]. — Hypericum carneum Wall., Numer. List. [Wallich] Distribution.— India, Myanmar, Malaysia, n. 4820. 1828. nom. nud. Singapore, Borneo, Java, Sumatra. — Hypericum horridum Wall., Numer. List. Ecology and phenology.— Evergreen forest, [Wallich] n. 4822. 1828; nom. nud. peat swamp forest from near sea level to 460 m elevation. Flowering: May–September. — Ancistrolobus ligustrinus Spach., Ann. Sci. Nat., Bot. Sér.2, 5: 352, t. 6. 1836. Type: China, circa Uses.— The wood is used for indoor Macao herb. Decaisne s.n. (not traced). ≡ Cratoxylum construction. ligustrinum Blume, Mus. Bot. 2: 16. 1852; Merr., Vernacular.— Ngong ngang (โงงงัง) (Niyomdham Trans. Amer. Philos. Soc. 24, 2: 268. 1935; Corner, 23); gawng ngahng (Maxwell 87-532). Gard. Bull. Straits Settlem. 10: 34. 1939; Meijer, Conservation.— IUCN Global Status: Least Bot. News Bull. Forest Dept., Sabah 7: 64. 1967; Concern (LC); Regional (Thailand) Status: Critically Corner, Wayside Trees of Malaya 1: 325. 1988. Endangered (CR). — Ancistrolobus sp., Wight, Ill. Ind. Bot. 1: 3. 1840; nom. nud. 2. Cratoxylum cochinchinense (Lour.) Blume, — Cratoxylum polyanthum Korth., Verh. Nat. Gesch. Mus. Bot. 2: 17. 1852; Corner, Gard. Bull. Straits Ned. Bezitt., Bot. 175, t. 36. 1842; Korth., Flora, Settlem. 10: 26–27, 34. 1939; Smythies, Comm. 31:579. 1848; Miq., Fl. Ned. Ind. 1(2): 515. 1859; Sarawak Trees: 69. 1965; Gogelein, Blumea 15: Dyer in Hook.f., Fl. Brit. India 1: 257. 1874; Kurz, 463. 1967; N.Robson in Fl. Males., Ser. 1, Spermat. J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 85. 8: 4–6. 1974; Corner, Wayside Trees of Malaya 1: 1874; Kurz, Forest Fl. Burma 1: 83. 1877; King, J. 325. 1988; P.H.Hô, Ill. Fl. Vietnam 1(1): 579, f. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59(2): 145. 1890; 1604 1991; Thin, Sida 17: 742. 1997; S.Gardner Brandis, Indian Trees: 47. 1907; Gagnep. in Lecomte, et al., Field Guide Forest Trees N. Thailand: 49. Fl. Indo-Chine 1(4): 287. 1910; Ridl., Fl. Malay 2000; H.W.Li et al., Fl. China 13: 36–37. 2007. ≡ Penins. 1: 152. 1922; Craib, Fl. Siam. 1:112. 1925; Hypericum cochinchinense Lour., Fl. Cochinch. 2: Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3): 472. 1790. ≡ Vismia cochinchinensis (Lour.) 253. 1943. Type: Indonesia. Borneo, Korthals s.n. Spreng., Syst. Veg. (ed. 16) 3: 350. 1826. Type: “In (lectotype L [L0012110!], designated by Gogelein sylvis Cochinchinae”, Loureiro s.n. (not located). [1967]; isolectotypes: BM [BM000624802!], NY [NY00084789!], L [L0012111!, L0012112!, — Hypericum chinense Retz., Observ. Bot. (Retzius) L0012113!, L0012114!, L0012115!, L0012116!, 5: 27. 1789; nom. illeg. ≡ Elodes [as Elodea] chi- L0012118!, L0012119!, L0012120!, L0012121!, nensis (?Retz.) Hance, London J. Bot. 7: 472. 1848. L0012122!, L0012124!). Type: China, “East China” (Hong Kong) possibly Bladh s.n. (isotype LD not seen). — Cratoxylum myrtifolium Blume, Mus. Bot. 2: 17. 1852. Type: Indonesia, Borneo, Korthals & Müller — Hypericum petiolatum auct. non L.: Lour., Fl. s.n. (syntype L [L0012125!]). Cochinch. 2: 479. 1790. ≡ Cratoxylum petiolatum (Lour) Blume., Mus. Bot. 2: 17. 1852. Type: China, — Cratoxylum wightiii Blume, Mus. Bot. 2: 18. 1852. “incultum propre Cantonem Sinarum”, Loureiro s.n. Type: Griffi th 1104 (holotype CGE not seen; isotype (not located). E [E00438025!]). ≡ Cratoxylum polyanthum var.

SW 11611-p162-216-G8.indd 203 1/16/62 BE 5:27 PM 204 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Map 2.1.4 Distribution of Hypericum japonicum in Thailand. Map 2.1.5 Distribution of Hypericum patulum in Thailand.

Map 2.1.6 Distribution of Hypericum siamense in Thailand. Map 2.2.1 Distribution of Cratoxylum arborescens in Thailand.

SW 11611-p162-216-G8.indd 204 1/16/62 BE 5:25 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 205 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

wightii Dyer in Hook.f., Fl. Brit. India 1: 25. 1874; stamens present per fascicle, sometimes congested. Corner, Gard. Bull. Straits Settlem. 10: 21–36. 1939. Ovary 2–3 mm long; style 1–3 mm long. Seeds — Elodes sp., Griff ., Notul. 4: 569. 1854; nom. nud. (5–)6–8 per loculus, 6–8 × 2–3 mm, oblanceolate, elliptic or oblong. — Ancistrolobus brevipes Turcz., Bull. Soc. Imp. Naturalistes Moscou. 31(1): 383. 1858. Type: Thailand.— (Map 2.2.2): NORTHERN: Chiang Singapore, Lobb 485 (isotypes BM [BM000624803!, Mai [Botanic Garden, Mae Rim, 30 Mar. 1998, BM000624804!], KWW [KW001000859!], W not seen). Sucheera 6 (QBG); ibid., 25 Oct. 1993, BGO Staff 63 (QBG); ibid., 6 June 1994, BGO Staff 600 (QBG); — Cratoxylum lanceolatum Miq., Fl. Ind. Bat. ibid., 24 Mar. 1996, BGO Staff 6109 (QBG); ibid., Suppl. 1: 500. 1861. Type: Indonesia, Sumatra, 30 Mar. 1998, Watthana 966 (QBG); ibid., 5 Nov, Teysmann HB 3813 (holotype U [U0002393!]; 1997, Watthana & Siriphum 22 (QBG); ibid., 5 July isotypes K!, L!).— Cratoxylum polyanthum var. 1989, Pooma 2077 (BKF); Huai Pan, no date, Serm genuinum Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. 115 (QBG); San Pan Si, QBG, Mae Rim, 9 July Hist. 43(2): 85. 1874; nom. inval. 2002, Glamwaewong 241 (QBG); Mae Ta, 23 Apr. — Cratoxylum polyanthum var. macrocarpum 1922, Winit 714 (ABD, BKF, K); Doi Suthep, 4 Apr. Boerl., Cat. Hort. Bog. 62. 1901. Type: Indonesia, 1910, Kerr 1080 (BM, C, K, L, TCD); Doi Suthep Boerlage s.n. (isotypes K!, L [L0012127!],). to Ban Neh, 30 Dec. 1921, Rock 1553 (A, K); Nov. 1960, Unknown collector 1938 (BK); Doi Suthep, — Cratoxylum hypoleuca Elm., Leafl . Philip. Bot. 13 Apr. 1958, Sørensen et al. 2682 (BKF, C, K); 5: 1787. 1913. Type: Philippines, Elmer 12913 ibid., 12 May 1958, Sørensen et al. 3369 (C, K); (isotypes A [A00067463!], BM [BM000624805!], Doi Inthanon NP, 22 July 1988, Phengklai et al. 6746 E[E00438024!],GH [GH00067464!], L [L0012126!], (BKF, E, K); Mae Kahn, 11 Oct. 2001, Maxwell K!,P [P0476972!], U [U002394!],US[US00901797!, 01-514 (A, CMUB); Doi Lahn, 13 June 2003, US00114120!], W not seen). Maxwell 03-130 (A, CMUB); Doi Suthep, 28 Apr. Trees or shrubs 2–30 m tall; dbh 35 cm; outer 1988, Maxwell 88-544 (L); Mae Kahn (Karen bark deeply fi ssured, yellow at cambium, sapwood Village), Doi Luang NP, east side, Mae Dtawn Nai white, heartwoodd brown, hard; exudate reddish-brown. Village Area, 19 June 1997, Maxwell 97-679 Branchlets terete, smooth, light brown to dark brown, (CMUB, BKF); along Mae Klang River, 30 July slender, delicate, scars present. Leaves petioled, 1988, Fukuoka T-623766 (PSU); Ban Mae-Hoi, Chom 0.2–0.5 cm long; blade 2.9–12.1 × 1.0–4.7 cm, Thong, Cholsuk s.n. (BKF)]; Chiang Rai [Payapri oblong to elliptic, base cuneate to attenuate, apex Village (Akha), 19 May 1997, Gardner & round, acuminate or acute, smooth and delicate to Sidisunthorn 2093 (A, CMUB); above Payapri Lao papyraceous to coriaceous and shiny, dark tan/red to Mae Village, 25 Nov. 2005, Maxwell 05-646 light yellow/green, some glaucous, black dots present (CMUB); Doi Giah, 10 Apr. 2006, Maxwell 06-295 on upper surface, density of black dots varying with (CMUB)]; Lamphun [Doi Khuntan NP, trail from each leaf, midrib faint on both surfaces, a little more Mah Meun Station to Pha Tup Falls, Mae Tha, 30 pronounced on lower surface but faint, narrowing Apr. 1994, Maxwell 94-572 (A, CMUB)]; Phrae approximately ½ mm towards apex; very faint on [Huai Khamin, RHS of Hui, 23 Aug. 1940, Somkid upper surface, light green to red/brown, secondary, 4677 (BKF)]; Sukhothai [Soke Phra Ruang Waterfall, intersecondary, tertiary and intramarginal veins visible 8 June 1972, Maxwell 72-280 (AAU, BK)]; on both surfaces, especially obvious on lower surface, Phitsanulok [Nong Hin, 21 May 1967, Phusomsaeng veins not arising at same points on either side of 218 (C, L); Thung Salaeng Luang, 19 July 1966, midrib. Inflorescence flowering stems flattened, Larsen et al. 465 (AAU, BKF, L, P); Thong Nang fl ower on leafy twigs in axils of mature leaves (usually Tow, 3 Dec. 1966, Prayad 561 (BK); Kaeng Sophi 2 per leaf pair), some also in terminal clusters, 1–5 Waterfall, 10 Dec. 1966, Sangkhachand 585 (BK); fl owered, pedicels 0.1–0.2 cm long. Calyx sepals Thung Salaeng Luang NP, ca 80 km east of persistent, 5–7 × 2–5 mm, oval, obovate, obtuse, Phitsanulok, 20 July 1973, Murata et al. T-16507 dull green to dark purple. Corolla petals 5–10 × (BKF, L, P); ibid., 25 July 1973, Murata et al. 2.5–5 mm, red, crimson, pink or orange, longitudinal T-17058 (C, K, L, P)]; Kamphaeng Phet [20 June veins present on petals. Stamens 4–8 mm long, 45–55

SW 11611-p162-216-G8.indd 205 1/16/62 BE 5:28 PM 206 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

1930, Kiah 24350 (ABD)]; NORTH-EASTERN: Loei Charoenchai & Poompuang 269 (CMUB); Khlong [Si Than Forest, Wang Saphung, 8 Apr. 1926, Din Sai Area, 9 Nov. 1997, Charoenchai 450 (CMUB); 204 (ABD, BK, BKF, K); Chung Pae, 6 May 1959, ibid., 14 Mar. 1998, Charoenchai 527 (BKF, Khantchai 1045 (BKF, K, L)]; Sakon Nakhon [26 CMUB); Tu Rean Forest, 1960, Bunpheng s.n. (L)]; June 1932, Lakshnakara 1003 (AAU, ABD, BK, SOUTH-EASTERN: Sa Kaeo [Aran Pratet, 18 Oct. BM, E, K)]; Nakhon Phanom [Tha-Uthen, 12 Aug. 1928, Put 2023 (AAU, ABD, BK, K); Muang, 1963, Pradit 422 (BK); ibid., 12 Aug. 1963, Pradit Srisanit s.n. (BKF)]; Prachin Buri [5 Aug. 1920, 4477 (BK)]; Khon Kaen [Kanchanapisaele Temple, Phengklai et al. 3711 (A, BKF, PSU)]; Chon Buri Phu Wieng District, 25 Mar. 2003, Kantachote 168 [Khao Nang Nom, Phanus Nikhom, 9 Apr. 1976, (KKU)]; EASTERN: Nakhon Ratchasima [Khao Yai Vachance 60 (BK); Sri Racha, no date, Collins 126 NP, 18 July 1973, Murata et al. T-16272 (BKF, L, (BM, K); ibid., June–Aug. 1913, Collins 199 (BM, P)]; Surin [Rattana, 19 May 1965, Sakol 2866 (BK); C, E, K, L, TCD); low hill, back of Sri Racha, 16 Vao Yai, 9 July 1963, Uarem? 408 (BK); between Nov. 1926, Collins 1300 (AAU, ABD, K); Khao Loong & Kong Bung Pru Villages, 16 Oct. 2003, Chalak, near Sri Racha, 29 Aug. 1926, Collins 1453 Maxwell 03-329 (CMUB)]; Ubon Ratchathani [UR (ABD, K); Khao Khieo Opened Zoo, 10 Dec. 2000, Gene Conservation Station, Ban Bahai Village, Huai Phengklai et al. 12888 (BKF); Kasetsart University Yang Subdistrict, Khong Chiam, 18 Sept. 2001, at Ao Udom, west side of Kow Hill, 23 Aug. 2003, Greijmans 183 (CMUB)]; SOUTH-WESTERN: Uthai Maxwell 03-2366 (CMUB); Khao Khieo, Sri Racha Thani [Nong/Prue Forest, Ban Rai, 6 July 1963, District, 16 Feb. 1975, Maxwell 75-131 (BK); ibid., Bunnak 9577 (BKF); Kanchanaburi [Saneh Pawng 11 Apr. 1975, Maxwell 75-420 (BK)]; Rayong [Chak Village, Sangkhlaburi District, 9 May 2003, Phong, Klaeng, 17 May 2004, Kertsawang 293 Kansuntisukmongkol 83 (CMUB); near Neeckey, (QBG)]; Chanthaburi [Khao Sabap, Phlieo Waterfall, near Wangka, 29 Apr. 1946, Kwae Noi River Basin 16 June 1970, Vachanapong 100 (BK); no date, B.S. Exp. 274 (BK); Huai Aek, 24 Nov. 1969, 170 (BKF); Ma Kuam, 22 Nov. 1930, Lakshnakara Sangkhachand 1598 (BKF, C, K, L, P); Sadong, 482 (AAU, ABD, BK, E, K); outside entrance to Salag Prah Wildlife Sanctuary, 20 Nov. 1971, van river on the way to the lighthouse, Collins 5566 (K); Beusekom et al. 40377 (BKF, C, K, L, P)]; Ratchaburi Khao Sabap, Chanthabun, 5 July 1927, Put 890 [Hua Hin–Pak Ta Wan, Ladell 204 (ABD, K); (AAU, ABD, BK, BM, C, E, K, L, P); Laem Ling, Nawng Ke, Apr. 1926, Collins 1571 (AAU, ABD, 8 Apr. 1923, Marcan 13477 (ABD); Phlieo Falls, 4 BK, K)]; Phetchaburi [Ang Num Yen, 16 Jan. 1969, Apr. 1971, Maxwell 71-245 (AAU, BK, L)]; Trat Vachanapong 320 (BK); T. Nong Ya Pong, 14 Mar. [23 Aug. 1954, Sangkhachand 179A (L); Ko Chang, 1965, Sakol 545 (BK); Huai Sai, Cha-am, 16 Mar. Khlong Son, 15 Apr. 1955, Bunnak 418 (A, BKF); 2001, Puudjaa 935/1 (BKF)]; Prachuap Khiri Khan Makham, 17 Mar. 1956, Sangkhachandd 629 (C, E, [Pranburi, Phengnaren 141 (BKF); Hua Hin, 5 Aug. K, L, P); West slopes of Khao Sabap, 25 Feb. 1935, 1920, Marcan 344 (BM, K); Sam Roi Yot, 25 Mar. Seidenfaden 2683 (C); near Ban Saphan Hin, ca 60 1987, Soejarto et al. 5785 (BKF, L); Nawng Kang, km SE from Trat, 4 Aug. 1973, Murata et al. T-17628 8 July 1926, Kerr 10911 (AAU, ABD, BK, K)]; (BKF)]; PENINSULAR: Chumphon [Pathio, 24 May CENTRAL: Saraburi [Pukhae, Muang District, 1947, 1969, Jaray 29 (BK); Tha Sae, 30 May 1969, Jaray Dee 81 (BKF); Forestry Station, Pukhae, Thananan 877 (BK); Bang Son, 9 Mar. 1928, Put 1469 (AAU, 24966 (BKF); Sahm Lan, 1 Dec. 1973, Maxwell 73- ABD, BK, C, E, K, L)]; Ranong [Kapoe, 18 Nov. 6977 (AAU, BK)]; Nakhon Nayok [Khao Yai NP, 1965, Sangkhachand 1139 (C)]; Surat Thani [18 Muang District, along the road to Heo Suwat Falls Aug. 1975, Prapat 477 (BKF, C, L); Khao Thapet, near Lam Takhong Camping site at 43 km, 18 Aug. 5 June 1966, Sakol 10266 (BK); Ko Prob, 17 July 2002, Aramwith 7 (A, CMUB); Pakphli District, 26 1966, Sakol 13877 (BK); Pha Phet, no date, Smitinand June 1991, Parinya et al. 366 (BK); Pakhli District, 5592 (BKF)]; Phangnga [Thung Chali, Nang Yo, 20 June 1991, Somprasong & Sangkhachand 49 (BK); July 1979, Niyomdham et al. 356 (BKF, C, K, L, P); ibid., 27 June 1991, Parinya et al. 60 (BK); Khao low hill along road, south of Takua Pa, 12 May 1968, Yai NP, Muang District, 25 June 2001,Boonkongchart van Beusekom & Phengklai 732 (AAU, BKF, C, E, 69 (A, BKF, CMUB); ibid., 4 June 2002, K, L, P); Kopah, 14 Dec. 1907, Haniff 2948 (BM,

SW 11611-p162-216-G8.indd 206 1/16/62 BE 5:28 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 207 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

K)]; Phuket [on way from Ton Sai Wildlife Reserve K)]; Yala [Muang, 28 July 1988, Saephu 8 (PSU)]; to Pa Klok, 7 July 1979, Niyomdham et al. 217 Narathiwat [Yi-ngo, 18 May 1961, Sangkhachand (BKF, C); Foothills of Khao Phra Mountain, 8 July 155 (C, L, P); Tak Bai, 12 July 1983, Niyomdham 1972, Robson 30761 (AAU, E, K, P); Khao Phra 692 (BKF, C)]; Unknown locations [5 Dec. 1921, Mountain, 8 July 1972, Larsen et al. 30764 (AAU, Vanpruk 298 (BKF, K); Jan.–May 1927, Squires E, K, P)]; Krabi [Nai Chong, Amphoe Muang, 14 333 (E); 1862–1866, Thorel 1062 (E); 22 Apr. 1958, Apr. 1969, Chermsirivathana & Kasem 13277 (BK); Sørensen et al. 3015 (C); 1862, Pierre 32366 (E); Kho Pra-Bang Khram Wildlife Sanctuary, 15 Aug. %LWVHQXORNH *URৼ (K); Vanpruk s.n. (BKF, 2006, Maxwell 06-590 (CMUB)]; Nakhon Si K); Godefroy-Lebeuf s.n. (K). Thammarat [Chawang, 10 Aug. 1956, Sanan 767 Distribution.— Myanmar, China, Indochina, (AAU, BKF, L); Chong Khao, Thung Song District, Malay Peninsula,Borneo, Philippines, Sumatra. 31 Aug. 1982, Shimizu et al. T-28951 (BKF)]; Phatthalung [Khaun-kha-nun, 6 Sept. 1996, Ecologyy and phenology.— Evergreen, deciduous Purintavasagul 223 (PSU)]; Trang [Vanpruk 602 or tropical rainforest, savannah, scrub from near sea (K); Chum Het, 15 Apr. 1928, Kerr 15212 (ABD, level to 1100 m elevation. Flowering: March–June. BK, K): Si Kao, 18 Apr. 1930, Kerr 19017 (AAU, Fruiting: August. ABD, BK, K); near Med Plant Exp. Plot, Trang Vernacular.— Teo (êĉĚü))( (Ladell 204); taew (Ēêüš ) Hort. Exp. Sta., Amper Sikao, 28 July 1987, (Bunnak 418);) tiu kliang (êĉĚüđÖúĊĚ÷Ü) (%*26WDৼ); Paisooksantivatana & Sangkhachand Y-2135-87 tarat ki tai (êćĒøé×ĊĚĕêš) (Sangkhachand 179A); seu (BK)]; Satun [Khuan Kalong, 10 May 1967, gwae joh (Maxwell 01-514). Unknown collector 365 (BKF); ibid., 9 May 1967, Conservation.— IUCN Global & Regional Phengnaren 496 (BKF, K, L, P)]; Songkhla [12 (Thailand) Status: Least Concern (LC). Sept. 1979, Udomsak 166 (PSU); Ton Nga Chang Wildlife Sanctuary, 20 km west of Hat Yai, 4 Aug. 1980, Students 22 (PSU); Hat Yai Municipal Park, 3. (Jack) Dyer in Hook.f., Wat Kawn Chong, Hat Yai, no date, Congdon & Fl. Brit. Ind. 1: 258. 1874; Kurz, Forest Fl. Burma Hamilton 22 (A, PSU); Ban Tun, Muang District, 1: 83. 1877; King,n J. Asiat. Soc. Bengal, Pt. 2, 16 Sept. 1980, Suwit 32 (PSU); Park, Hat Yai, 11 Nat. Hist. 59(2): 145. 1890; Koord. & Val., Bijdr. Sept.,1980, Students 366 (PSU); PSU Campus, 18 Boomsoort. Java 5: 137. 1900; Gagnep. in Lecomte, Oct. 1982, Peechate 377 (PSU); Tumbol Kho Sa Ba, Fl. Indo-Chine 1(4): 287. 1910; Ridl., Fl. Malay The-Pha, 24 Nov. 1943, K.K. et al. 54 (PSU); Hat Penins. 1:152. 1922; Craib, Fl. Siam. 1: 111. 1925; Yai, 16 Apr. 1960, Pradit 202 (BK); Khuan Mit, 29 Corner, Gard. Bull. Straits Settlem. 10: 23, 28. 1939; Apr. 1979, Congdon & Hamilton 404 (PSU); Kho Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3): Hong Hill, Hat Yai, 23 Aug. 1979, Srirugsa 404 251. 1943; Backer & Bakh.f., Fl. Java 1: 383. 1963; (PSU); Khlong Hae, Hat Yai, 24 Apr. 1985, Meijer, Bot. News Bull. Forest Dept., Sabah 7: 64. Vachanapong 654 (BK); Rataphum, 25 May 1970, 1967; Gogelein, Blumea 15: 467. 1967; N.Robson Sutheesoen 1733 (BK); Tepa, 23 Mar. 1928, Kerr in Fl. Males., Ser. 1, Spermat. 8: 4–6. 1974; Corner, 14709 (ABD, BK, BM); Kho Hong Hill, Hat Yai, Wayside Trees of Malaya 1: 325. 1988; P.H. Hô, 30 Oct. 1990, Larsen et al. 40938 (PSU); Ton Nga Ill. Fl. Vietnam 1(1): 578, f. 1602. 1991; Thin, Sida Chang Wildlife Sanctuary, 1 Nov. 1990, Larsen et 17(4): 742. 1997; H.W.Li et al., Fl. China 13: 37–38. al. 41030 (AAU, BKF, P, PSU); Khao Chum Sak, ŁElodes formosa Jack, Malay. Misc. 2(7): Hat Yai, 12 June 1992, Larsen et al. 42818 (AAU, ŁTridesmis formosa Korth., Verh. Nat. P); Kho Hong Hill, Hat Yai 25 May 1985, Maxwell *HVFK %RW  W   Ł Tridesmis jackii 85-5377 (A, AAU, BKF, E, L, P, PSU); Khlong Hoi Spach, Ann. Sci. Nat., Bot., Sér. 2, 5: 352. 1836. Type: Khong, Hat Yai, 2 Oct. 1985, Maxwell 85-934 (AAU, Indonesia, Sumatra, Jack s.n. (holotype CAL!) BKF, L, PSU); near Boripath Waterfall, 12 Jan. Trees to 8 m tall; dbh 10 cm; outer bark thick, 1978, Srirugsa PS21 (PSU); Tung Lung, 19 Apr. 1974, VFDO\DQGÀDNLQJLQQHUEDUNZLWKZDWHU\H[XGDWH Vitchu V9 (PSU)]; Pattani [Banang Station, 2 July Branchlets larger branches armed with long stout 1923, Kerr 7316 (AAU, ABD, BK, BM, E, K), 8 thorns, stems spiny, brown, interpetiolar scars Sept. 1923, Kerr 7745 (AAU, ABD, BK, BM, E,

SW 11611-p162-216-G8.indd 207 1/21/62 BE 3:25 PM 208 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

interrupted. Leaves petioled, petiole 0.2–0.8 cm long; FOXVWHUVRIÀRZHUHGF\PHVRI±RQROGOHDÀHVV blade 0.2–10.4 × 1.3–4 cm, oval, elliptic, lanceolate, twigs, pedicels 0.3–1(–1.5) cm long. Calyx sepals round, oblanceolate or obovate, light green, some 5–7 × 2–3 mm, oblong to elliptic, persistent,green- yellow/green or brown/red, base cuneate, few being grey. Corolla petals, 5; obovate to oblong; persistent?, almost angustate, apex acute, round, obtuse, acumi- ÀRZHUVZKLWH Stamens fused into 3 slender fascicles nate or cuspidate, midrib faint adaxially, obvious of 20–30 stamens, anthers round, slightly shorter abaxially, narrowing towards apex, veination netted, than the styles. Ovary 2–4.5 mm long, styles 3, obvious on some leaves, faint on others, secondary, 2–8 mm long, green, glabrous. Fruit 4–6 mm, oval, intersecondary, tertiary and intramarginal veins oblong, obovate with pointed tip purple/deep red; present, veins not arising from same points on both pedicel 6–11 mm long, hairy; covered by persistent sides of the midrib, 7–12 pairs of very obvious side sepals at base. Seeds 6–8 per loculus, 6–7.5 × 2–4 mm, veins joined in loops near margins. ,QÀRUHVFHQFH in obovate, oblong or oblanceolate.

KEY TO THE SUBSPECIES OF C. FORMOSUM 1. All parts glabrous 1. subsp. formosum 1. Young twigs, pedicels, calyx and leaves pubescentt 2. subsp. SUXQLÀRUXP

1. subsp. formosum C, E, K)]; Roi Et [Kasetwisai, Ban Nam Om, 9 June — Hypericum biflorum (non Lamark) Choisy in 1982, Paisooksantivatana & Sutheesoem Y1027-82 DC., Prod. 1:546. 1824. Type: China, ?König. (BK)]; Si Sa Ket [Kantharalak District, 24 Mar. 1966, Sangkhachand 203 (BK); ibid., 24 Mar. 1966, — Tridesmis ochnoides Spach, Hist. Veg. Phan. 5: Sangkhachand 208 (BK); ibid., 28 Mar. 1966, 359. 1836; Ann. Sci. Nat. Bot. 2(5): 351, t. 4a. 1836; Sangkhachand 214 (BK); ibid., 12 Apr. 1966, Blume, Mus. Bot. 2:18. 1852. Type: ?holotype: FI Sangkhachand 245 (BK)]; Ubon Ratchathani [Nam [FI1005999!], possible first stage lectotype, see Tok Sae, NE, 26 Feb. 1967, Phusomsaeng 33 (BKF, Gogelein, Blumea 15468-469. 1967. C, E, K, L, P); Wormchumsak, 25 Feb. 1961, — Hypericum aegiptium Blanco, Fl. Filip. 615. Chirayupin 187 (BK); Dong Phahuan, no date, 1837. Type: Philippines, Palawan, Taytay, Merrill Niyomdham 4598 (AAU)]; SOUTH-WESTERN: Species Blancoanae 632 (Illustrative specimen/ Kanchanaburi [Dongyai, 14 Aug. 1971, Phengklai neotype)([S!]). et al. 2928 (C, E, K, L); Between Kritee & Meung — Cratoxylum pentadelphum Turcz., Bull. Soc. Chah, 7 July 1973, Geesink & Phengklai 6169 (C, Naturalistes Moscou 36: 580. 1863. Type: Indonesia, E, K, L, P)]; Prachuap Khiri Khan [Khlong Lah Met, 17 July 1921, Winit 628 (ABD, BK, BKF); Java,+RUV¿HOGVQ (lectotype KWW [KW001000481!], designated here; isolectotypes BM!, CGE not seen). Huai Yang Forest Reserve, 21 Apr. 1997, Lamkay 40-112 (KKU); no date, Godefroy-Lebeuf s.n. (K)]; All parts glabrous; connective without glands. CENTRAL: Saraburi [Sam Lan, 30 Mar. 1974, Thailand.— (Map 2.2.3a): NORTHERN: Chiang Maxwell 74-2199 (AAU, BK, BKF); Sam Lan Forest, Mai [Doi Suthep, 15 Apr. 1909, Garrett 32 (K)]; Muang District, 15 June 1974, Maxwell 74-590 Sukhothai [29 Apr. 1964, Pradit 899 (BK)]; NORTH- (AAU, BK, L)]; Nakhon Nayok [Pakphli District, EASTERN: Kalasin [Kalasin Morning Market, A. June 1991, Somprasong & Sangkhachand 53 (BK); Muang Kalasin, 28 Jan. 1990, Widmer 30 (BKF)]; Kao Sila, 25 Mar. 1975, Sutheesoen 3242 (BK)]; Maha Sarakham [Pa Khok Dang Khaeng, Maha SOUTH-EASTERN: Sa Kaeo [Aran Pratet, 18 Oct. Sarakham University, 10 May 1996, %*26WDৼ 1928, Put 2020 (AAU, ABD, BK, BM, E, K); (QBG)]; Khon Kaen [Nam Phong, 22 June 2003, Prachin Buri [Taphraya, 4 Jan. 1966, S. & J. 2093 Chantaranothai et al. s.n. (KKU)]; EASTERN: (BK)]; Chon Buri [Sri Racha Forest, 16 miles inland Chaiyaphum [Pa-Hin-Ngam Forest Park, 17 Mar. from sea, Aug. 1913, Collins 159 (AAU, BK. E, K); 1993, Suddee 266 (BKF); East Chaiyaphum, 18 Apr. Sri Racha Forest, Collins 578 (K); Sri Racha Forest, 1971,Chaloenphol 85 (L); Pak Pang via Chaiyaphum, Khong Yai Bu, 29 Mar. 1921, Collins 723 (ABD, E, 30 Jan. 1931, Kerr 19970 (AAU, ABD, BK, BM, K, TCD); Sri Racha Forest, 4 Mar. 1920, Kerr 4033

SW 11611-p162-216-G8.indd 208 1/21/62 BE 3:25 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 209 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

(ABD, BM, K, TCD); Chantalen Falls, 18 Dec. 1974, Ecology and phenology.— Evergreen forest, Chermsirivathana & Sangkhachand 1969 (BK); Ko deciduous forest, beach, scrub area from 20 to 900 Khram, Sattahip, 14 Aug. 1999, Phengklai et al. m elevation. Flowering: March–June. Fruiting: 119300 (BKF); Ang Chong Nam, District, April–October. 29 Feb. 1976, Maxwell 76-129 (BK, P)]; Rayong Vernacular.— Tao ( ) (Kerr 14431); teo ( ) [Chak Phong, Klaeng, 3 Apr. 2004, Kertsawang 252 แตว ติ้ว (Kerr 9748);) tao-som (แตวสม) (Sangkhachand 355); (QBG)]; Chanthaburi [Laem Sing, 8 Apr. 1923, tiu khaao ( ) (BGO Staff 3). Marcan 1339 (ABD, BM); Makam, 8 Apr. 1959, ติ้วขาว Smitinand 5751 (AAU, BKF, L); Plain of Makam, Uses.— The wood is used in construction. 9 Apr. 1959, Sørensen et al. 7232 (C)]; Trat [Leam Conservation.— IUCN Global & Regional Koh, 28 Mar. 1955, Sangkhachand 355 (BKF, L); (Thailand) Status: Least Concern (LC). Ko Chang Noi, Schmidt 6966 (C, K); Amphoe Khao 2. subsp. pruniflorum (Kurz) Gogelein, Saming to Khlung, 2 Jan. 1930, Kerr 17923 (AAU, Blumea 15: 469. 1967; S.Gardner et al., Field Guide ABD, BK, BM, C, K, L)]; PENINSULAR: Chumphon Forest Trees N. Thailand: 48. 2000. ≡ Tridesmis [Phon, 16 Mar. 1958, Sørensen et al. 2142 (C, K)]; prunifl ora Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat Surat Thani [Na Muang Falls, Ko Samui, 2 June Hist. 41(2): 293. 1872. ≡ Cratoxylum prunifl orum 1960, Chirayupin 152 (BK); Ko Samui, 8 Apr. 1927, (Kurz) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat Hist. Kerr 12539 (ABD, BK, BM, E, K); Ban Na San, 43(2): 84. 1874; Kurz, Forest Fl. Burma 1:84. 1877; 13 Aug. 1927, Kerr 13324 (AAU, ABD, BK, E, K)]; Craib, Fl. Siam. 1: 113. 1925. ≡ Cratoxylum Phangnga [Khao Nanghong, 9 May 1967, Sutheesoen prunifolium (Kurz) Dyer in Hook.f., Fl. Brit. India 2563 (BK)]; Krabi [Sakeo, 23 Dec. 1924, Kerr 9748 1: 258. 1874; Brandis, Indian Trees, 47. 1907; (AAU, ABD, BK, BM, C, E, K, L)]; Trang [Angtong Gagnep. in Lecomte, Fl. Indo-Chine 1(4): 287. 1910; Falls, Sikao District, 16 Oct. 1978, Taworn 78 (PSU); Gagnep. in Humbert, Suppl. Fl. Indo-Chine 1(3): Amphoe Nayong, Khao Chong NP, trail from 251. 1943.Type: Myanmar, Kurz s.n. (not located, Botanic Gardens HQ to Ton Yai, 9 July 2000, probably CAL). Middleton et al. 345 (A, E, TCD); Khao Chong, 10 Mar. 1966, Bunnab 415 (BKF); ibid., 16 June 1973, — Hypericum prunifolium Wall., Numer. List. Phusomsaeng et al. 1594 (BKF); ibid., 18 Mar. 1969, [Wallich] 7276. 1828; nom. nud. Sangkhachand 1798 (BKF); ibid. 25 June 1969, — Cratoxylum dasyphyllum Hand.-Mazz., Sinensia Sangkhachand 1913 (BKF); ibid., 8 Mar. 1976, 2: 4. 1931. Type: China, Ching 7980 (isotypes L Chermsirivathana 2203 (BK); Trang Forest near [L0012128!], NY [NY00084786!], W not seen). Chong Waterfalls, 18 Sept. 1933, Collins 2357 (AAU, BM, K, P); Chong, 20 Aug. 1935, Smitinand Young twigs, pedicels, calyx and leaves 2960 (BKF, L); Ko Libong, 21 Apr. 1930, Kerr pubescent; connective with glands. 190566 (AAU, ABD, BK, BM, E, K); Khao Chong, Thailand.— (Map 2.2.3b): NORTHERN: Mae 12 Mar. 1974, Larsen & Larsen 33255 (AAU, C, Hong Son [Khun Yuam, 4 Apr. 1977, Nimanong et K, P); ibid., 13 Aug. 1975, Maxwell 75-831 (BK, al. 17877 (PSU); Di Bao-hae, Mae Sariang, 23 Apr. L)]; Satun [Khlong Ton, 10 Mar. 1928, Kerr 14431 1973, Sutheesoen 2379 (BK)]; Chiang Mai [Botanic (AAU, ABD, BK, BM, K)]; Songkhla [near Suanisan Garden, Mae Rim, 18 Oct. 1993, BGO Staff 6 Restaurant, Hat Yai, 16 Sept. 1991, Pollawat 6 (QBG); ibid., 21 Mar. 1996, Morakot 8 (QBG); (PSU); Hat Yai. Khuan Mit, 29 Apr. 1979, Congdon ibid., 30 Mar. 1998, Watthana 98 (QBG); ibid., 6 404 (AAU, BKF); Hat Yai, Kho Hong Hill, summit Apr. 1994, BGO Staff 587 (QBG); Mae Sanam ridge, 31 Mar. 1985, Maxwell 85-348 (A, BKF, E, Arboretum, 17 Apr. 2000, Sankamethawee 110 L, PSU); ibid., 10 Oct. 1985, Maxwell 85-9466 (A, (CMUB); Doi Suthep, 1 Apr. 1949, Soradetch 426 AAU, L, PSU)]; Narathiwat [Bacho, 18 Apr. 1961, (BKF); Chom Thong, Fang, Pong Namron, 17 Aug. Sangkhachand 65 (C)]; Unknown location [Apr. 1968, Phengnaren 579 (BKF, L); Doi Suthep, 27 1870, Pierre 1466 (E)]. Apr. 1909, Kerr 611 (BM, K, P, TCD); Doi Pha Distribution.— Cambodia, Laos, Vietnam, Dam, between Hang Dong & Bo Luang, 5 July 1968, Malaysia, Indonesia, Philippines Larsen et al. 2152 (AAU, BKF, E, L, P); Doi Suthep, Sørensen et al. 2910 (C): Mae Sa Mai, 30 Mar. 1996,

SW 11611-p162-216-G8.indd 209 1/16/62 BE 5:28 PM 210 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

BGO Staff 62166 (QBG); Doi Inthanon NP, 4 May in southerly direction, 12 Aug.,1998, van de Bult 79 1996, BGO Staff 6389 (QBG); ibid., 18 July 1988, (BKF, CMUB)]; NORTH-EASTERN: Phetchabun [ Phengklai et al. 6551 (C, K); Doi Suthep, east side, Sangkhachand 3112 (C, L)]; Loei [Namtok Tad Huai Kaeo Falls Area, 4 July 1987, Maxwell 87-604 Hueang, Na Haeo, 9 Dec. 1996, BGO Staff 27 (BKF, L); Muang District, Doi Suthep, east side, (QBG); ibid., 7 Nov. 1995, BGO Staff 120 (QBG); East side of Khonthathan Falls Area, 6 Apr. 1988, Si Tang, Wang Saphung District, 19 Mar. 1952, Maxwell 88-430 (AAU, BKF, L); Jawn Tong, 19 Smitinand 12166 (BKF); Na Haeo, 28 July 1995, July 1991, Maxwell 91-661 (A, AAU, E, P); Jawn BGO Staff 39766 (QBG); HQ-Phu Son, Phu Ruea Tong, Mae Soi Valley, Mae Soi Subdistrict, without NP, 12 Dec. 1996, BGO Staff 81844 (QBG)]; EASTERN: date, Maxwell 90-525 (L); Muang District, east side Chaiyaphum [18 Apr. 1971, Chaloenphol 85 (BK); of the Central University Library, Chiang Mai Nam Phrom, 10 Dec. 1971, van Beusekom et al. University, 11 May 1992, Maxwell 92-213 (A, 4100 (C)]; Buri Ram [Lam Nong Rong, 23 Mar. CMUB, E); Mae Pah Boo (Karen Village), 23 May 1920, Suksakorn 892 (K)]; Surin [3 Dec. 1976, 1995, Maxwell 95-428 (A, CMUB); Doi Lahn, 18 Phengklai et al. 3590 (PSU)]; Si Sa Ket [Kantharalak Sept. 1996, Maxwell 96-1204 (A, BKF, CMUB); District, 21 Apr. 1966, Sangkhachand 2966 (BK)]; Doi Bah Gluay, 3 May 1997, Maxwell 97-434 (A, SOUTH-WESTERN: Kanchanaburi [Saneh Rawng BKF, CMUB); Jahm Luan Subdistrict, Huay Yah Village, Sangklaburi, 19 May 2003, Dtai (Karen) Village, slopes along Mae Jon River, Kansuntisukmongkol 1866 (A, CMUB); Thung Yai 11 June 1998, Maxwell 98-649 (A, CMUB, L); Doi Naresuan West Wildlife, Thong Pha Phum, 14 Apr. Suthep-Pui NP, 22 Mar. 2000, Maxwell 00-148 (A, 2006, van de Bult 885 (CMUB); near Kin Sayok CMUB); Doi Inthanon NP, 2 Dec. 1998, Hara C533 about 120 km north west of Kanburi, 12 July 1946, (CMUB); Mae Ya Waterfall, Doi Inthanon, 29 Nov. Kostermans 10900 (A, C, L); Dongyai, 14 Aug. 1971, 1999, Hara E98 (CMUB); Tad Noi, Somthong, Phengklai et al. 2928 (AAU); Ban Kao, 17 Nov. 14 Feb. 1938, Taengsuwan s.n. (BKF); Me Tun, 1961, Larsen 82977 (C); 12 Mar. 1926, Kerr 10604 28 Apr. 1915, Winit 315 (BM, K)]; Chiang Rai [Khun (ABD, BK, BM, E, K, TCD); Muang Cha Area, Chae NP, 5 Apr. 1998, Maxwell 98-397 (BKF, Sangkhla Buri District, 8 July 1973, Maxwell 73-208 CMUB); Doi Luang NP, 29 May 1998, Maxwell (BK)]; CENTRAL: Saraburi [Sam Lan Forest, Muang 98-604 (A, BKF, CMUB)]; Lamphun [Ban Pha District, 19 May 1974, Maxwell 74-528 (AAU, BK); Mon (Chom Thong), 31 May 1979, Vidal et al. 6205 ibid., 8 Feb. 1975, Maxwell 75-108 (BK, L)]; (BKF, E, K, L, P); Doi Khun Tan NP, 1 May 1994, Nakhon Nayok [Muang District, 3 May 2002, Maxwell 94-579 (A, CMUB); ibid., 28 Sept. 1994, Charoenchai & Poompuang 2466 (CMUB)]; SOUTH- Maxwell 94-1050 (A, CMUB)]; Forest at EASTERN: Chon Buri [Sri Racha, 4 Mar. 1920, Kerr Jaehomwitthaya School, 19 Apr. 2000, Panatkool 4034 (ABD, BM, K); Khao Khieo, Sri Racha District, 2577 (A, CMUB); ibid., 21 May 2000, Panatkool 2 Feb. 1975, Maxwell 75-102 (BK, L); Phra Chedi 309 (A, CMUB); ibid., 25 Mar. 2001, Panatkool mountain, Ko Kieo, Sri Racha, 11 Apr. 1975, Maxwell 481 (A, CMUB); Me Kang, 29 Apr. 1925, Winit 75-419 (BK, L)]; Chanthaburi [28 Mar. 1956, 1403 (ABD, BK, BKF, K); Muang Ngao, 16 July Sangkhachand 640 (L); Soi Dao, Larsen 10015 (C); 1931, Put 4013 (ABD, BK, BM, E, K); Ngao, Pra- Khao Phra Bat, north of Chanthaburi, 27 Aug. 1972, tu-pha, 2 Sept. 1996, Phengklai & Fukuoka 10127 Larsen et al. 32141 (AAU, E, K, P)]; PENINSULAR: (BKF); Huai Tak, 28 May 1954, Tharawattananon Krabi [Kow Pra-Bahng Krahm Wildlife Sanctuary, BKF 40372 (BKF); Doi Luang NP, 22 Apr. 1997, 30 Mar. 2006, Maxwell 06-2400 (CMUB)]; Unknown Maxwell 97-3866 (A, CMUB)]; Phrae [Huai Mae locations [1914, Vanpruk 479 (P); Truck Lek, 25 Sai, Padaeng District, 21 May 1949, Sukket 3 (BKF); Apr. 1922, Marcan 7377 (BM); Muan Jawn, 3 Apr. Wang Mon, 19 Feb. 1912, Vanpruk 285 (BKF, K); 1922, Kerr s.n. (BK, P); Kerr s.n. (BM)]. Huai Khamin, 23 Aug. 1940, Somkid 468 (BKF)]; Distribution.— Myanmar, Laos, Cambodia Tak [Lan Sang, 31 May 1973, Geesink et al. 5596 Vietnam, southern China. (C); Lan Sang NP, 22 July 1973, Murata et al. T-16683 (BKF, P)]; Phitsanulok [Thung Salaeng Ecology and phenology.— Evergreen forest, Luang, 19 July 1966, Larsen et al. 468 (AAU, deciduous forest, rocky dipterocarp savannah, on BKF)]; Nakhon Sawan [Khlong Lan, 2 km from Tr. limestone from 20 to 950 m elevation. Flowering: March–May. Fruiting: April–November.

SW 11611-p162-216-G8.indd 210 1/16/62 BE 5:28 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 211 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Vernacular.— Teaw (ติ้ว) (BGO Staff 27);) ra- — Cratoxylum parvifolium Merr., J. Arnold Arbor. gein (ราเงง) (Sangkhachand 640); tiu khon (ติ้วขน) 19: 56. 1938. Type: Vietnam, Clemens & Clemens (BGO Staff 6); tiu daeng (ติ้วแดง) (Watthana 98); tiu 3454 (lectotype A [A00067445!], designated here; (ติ้ว) (BGO Staff 8184); sah gwe jo (Maxwell 95-428); isolectotypes BM [BM000624784!], L!, MICH sih weh jo (Maxwell 98-604). [MICH1115535!], P [P01901058!], U [U226859!], US!, W not seen). Uses.— The bark is used as a medicine to treat diarrhoea in domestic animals and the young leaves — Cratoxylum thorelii Pierre ex Gagnep., Suppl. are used as a substitute for tea. (Li et al., 2007). Fl. Indo-Chine 1: 252. 1943. Type: Cambodia, Compon Luong, Thorel 2065 (?holotype P Conservation.— IUCN Regional (Thailand) [P01901059!]; isotypes L [L0012141!],P [P01901060!, Status: Least Concern (LC). P019001061!]). Deciduous shrubs or trees to 20 m tall, outer 4. Cratoxylum maingayi Dyer in Hook.f., Fl. Brit. bark of many, thin fl aking layers, brown, inner bark India 1: 258. 1874; King, J. Asiat. Soc. Bengal, Pt. 2, brown; exudate yellow. Leaves petioled, 0.5–1 cm Nat. Hist. 59(2): 145. 1890; Gagnep., Notul. Syst. long; blade 2–9 × 1–4.5 cm, obovate to elliptic to (Paris) 1: 19. 1909; Ridl., Fl. Malay Penins. 1:152. oblong, coriaceous, base round to cuneate, apex acute 1922; Gogelein, Blumea 15: 470. 1967; N.Robson to round, midrib dark green adaxially, greyish-pale in Fl. Males., Ser. 1, Spermat. 8: 4–6. 1974; Corner, to light green adaxially, secondary veins present, Wayside Trees of Malaya 1: 325. 1988; Phengklai raised on both surfaces. Infl orescence pedicels light & Niyomdham, Flora in peat swamp areas of green, 0.2–0.6 cm long, axillary cymes of 2–4 fl owers Narathiwat: 197. 1991; S.Gardner et al., Field Guide on leafl ess twigs, fl owers pink-white. Calyx sepals Forest Trees N. Thailand: 49. 2000. Type: Malaya, 2.5–5 × 1–3 mm, oblong, light green. Corolla petals Malacca, Penang, Maingay 145 (isotypes BM 6–12 × 2–5 mm, elliptic, dark red, lateral veins present. [BM000624807!], CAL not seen, CGE not seen, Stamens numerous, anthers cream; fi laments white. K!, L [L0012144!, L0012145!]). Ovary 2–3 mm long, styles as long as petals. Fruit — Cratoxylum harmandii Pierre, Fl. Forest. 2–4 mm diam., oblong to acute. Seeds winged, 5–6 Cochinch. 1: t. 53. 1882; Gagnep., Notul. Syst. per loculus, oblong to ovate-oblong. (Paris) 1: 18. 1909. Type: Vietnam, Harmand 3235 Thailand.— (Map 2.2.4): NORTHERN: Chiang (?holotypeP [P001901062!]; isotypeL [L0012143!]). Mai [Maesa Botanic Garden, 5 July 1989, Pooma — Cratoxylum formosum var. thorelii Pierre ex 2077 (CMUB); Muang Fang, 3 June 1992, Maxwell Gagnep., Notul. Syst. 1; 19. 1909. Type: Cambodia, 92-255 (A, CMUB, E, P); Doi Suthep-Pui NP, SW Grand-Lac, Godefroy 295 (?holotype P [P01901063!, side off road to Sisangwen Falls, Hang Dong District, P01901064!]; isotypes L [L0012142!], K!. 5 Nov. 1992, Maxwell 92-6666 (CMUB, P); Mae Pah — Cratoxylum cochinchinense var. calcareum Ridl., Boo (Karen Village), 23 May 1995, Maxwell 95-426 Bull. Misc. Inform. Kew 1938: 115. 1938. Type: (A, CMUB)]; NORTH-EASTERN: Nakhon Phanom Indonesia, Borneo, Haviland 1463 (lectotype SING [Phu Phan NP, 14 Dec. 1982, Koyama et al. T-31004 [SING0062298!], Designated here; isolectotypes (A, BKF)]; PENINSULAR: Ranong [Kapoe, 18 K!, L [L0012146!]). Nov.1965, Sangkhachand 1139 (BKF, K, L)]; Trang [Khao Chong, 26 Oct. 1965, Boonnab 92 (BKF)]; — Cratoxylum acuminatum Merr., Papers Mich. Narathiwat [Pawai, Su-ngai Padi, 9 Sept. 1985, Acad. Sc. 23: 185. 1938. Type: Indonesia, Sumatra, Niyomdham et al. 928 (A, AAU, BKF, C, K, L)]. Rahmat Si Boeea 7687 (lectotype SING [SING0062299!]), designated here; isolectotypes A Distribution.— Indochina, Malay Peninsula, [A000677446!], L [L0043274!], MICH Singapore, Sarawak, Sumatra. [MICH1115534!], S [S11-34223!]. Ecology and phenology.— Lowland forest, — Cratoxylum subglaucum Merr., Papers Mich. peat swamp forest from near sea level to 850 m Acad. Sc. 23: 186. 1938. Type: Indonesia, Sumatra, elevation. Flowering: May–June. Fruiting: November– Rahmat Si Boeea 79077 (isotypes A [A00067448!], December. L!, MICH [MICH1115536!], S [S-G-1700!], SING Vernacular.— Taeo (แตว)(Sangkhachand [SING0062300!]). 1139); sah gwee jo (Maxwell 95-4266).

SW 11611-p162-216-G8.indd 211 1/21/62 BE 3:20 PM 212 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Map 2.2.2 Distribution of Cratoxylum cochinchinense in Map 2.2.3a Distribution of Cratoxylum formosum subsp. Thailand. formosum in Thailand.

Map 2.2.3b Distribution of Cratoxylum formosum subsp. Map 2.2.1 Distribution of Cratoxylum maingayi in Thailand. prunifl orum in Thailand.

SW 11611-p162-216-G8.indd 212 1/16/62 BE 5:25 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 213 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Conservation.— IUCN Global & Regional Vanpruk 2700 (BKF, K); Huai Khamin, 24 Aug. 1940, (Thailand) Status: Least Concern (LC). Somkid 469 (BKF); Song, Mae Yom NP, Tao Pun Subdistrict, 10 Oct. 1991, Maxwell 91-872 (AAU, 5. Cratoxylum neriifolium Kurz, J. Asiat. Soc. CMUB, E, P)]; Phitsanulok [in strip of cut out forest Bengal, Pt. 2, Nat. Hist. 41(2): 293. 1872; Dyer in near rivulet (Ran Ti River), Apr.–May 1946, Hook.f., Fl. Brit. India 1: 257. 1874; Kurz, Forest Kostermans 274 (A, K, L)]; NORTH-EASTERN: Udon Fl. Burma 1: 85. 1877; Gamble, Man. Ind. Timb.: Thani [Nong Bua, 7 Mar. 1924, Kerr 8619 (ABD, BK, 48. 1881; Prain, Bengal Pl. 1: 244. 1903; Brandis, BM,C, E,K,P,TCD)]; EASTERN:Nakhon Ratchasima Indian Trees, 47. 1907; Gagnep. in Lecomte, Fl. [Pak Thong Chai, 27 May 1968, Phengnaren 529 Indo-Chine 1(4): 291. 1910; Craib, Fl. Siam. 1: (BKF)]; SOUTH-WESTERN: Uthai Thani [Kao Pa-sha, 112. 1925; Gagnep. in Humbert, Suppl. Fl. Indo- Nong Chang, 29 May 1974, Sutheesoren 3008 (BK)]; Chine 1(3): 254. 1943. ≡ Kanchanaburi [Kwae Noi Basin, Unknown collector Blume subsp. neriifolium (Kurz) Gogelein, Blumea 143 (L); Tham Phe, 25 Dec. 1961, Phengklai 322 15: 463. 1967; S.Gardner et al., Field Guide Forest (L); Brangkasi, about 100 km south of Wangka, 19 Trees N. Thailand: 49. 2000. Syntypes: Myanmar, June 1946, den Hoed & Kostermans 474 (BK, K, L); Chittagong, Pegu Myanmar, Kurz (not located). south of Ka Tha Lai in Pan Puang River Valley, about 40 km south east of Wangka, 13–16 June 1946, — Hypericum neriifolium Wall., Numer. List. 4824 Kostermans 869 (A, BK, K, L); Tapoh, 27 Dec. (pro parte). 1828, nom. nud. 1961, Danish Expedition 8991 (A, C, E); Thung Yai Deciduous trees 7–20 m tall; dbh 40 cm; outer Naresuan Wildlife Reserve, Lai Wa Subdistrict, Ban bark thick, roughly cracked and flaking, black, Saneh Pawng (Karen Village), 15 Aug. 1993, fissured, very rough; exudate not recorded. Maxwell 93-929 (A, CMUB)]; PENINSULAR: Trang Branchlets terete to fl attened, smooth, scars absent, [near Medicinal Plant Exp. Plot, Trang Hort. Exp. black to black-red to green. Leaves sessile; blade Station, Amper Sikao, 28 July 1987,Paisooksantivatana 5.7–10.3 × 1.5–3.1 cm, lanceolate to elliptic to oblong & Sangkhachand Y2133-877 (BK)]; Unknown locations coriaceous and tough, occasionally glaucous, base [Somkid 103 (BKF)]. rounded to slightly cordate, apex acute to rounded to acuminate, midrib obviously depressed on upper surface and pronounced on lower surface, light yellow to red/brown on upper surface, yellow to dark red on lower surface, secondary, intersecondary and intramarginal veins present, all visible. Infl orescence fl owering branch fl attened, then spreading into many- branched terete delicate pedicels, fl owers usually in groups of 3, forming an elongated infl orescence not in the axils of mature leaves but on leaf twigs, length of pedicel variable. Calyx sepals persistent, 6 × 3 mm, obovate; greenish-brown. Corolla petals twice as long as sepals, bright scarlet. Stamens fi laments free. Ovaryy 1.5–3 mm long. Fruit 8–9 × 4–5 mm, covered by persistent sepals. Seeds 6–8 per loculus.

Thailand.— (Map 2.2.5): NORTHERN: Chiang Mai [Muang Payao, 11 July 1931, Put 3970 (AAU, ABD, BK, BM, E, K)]; Chiang Rai [Payao, 23 July 1966, Prayad 441 (BK); Huai Dong, at Tew Dam, 13 Aug. 1926, Winit 1794 (ABD, BK, BKF, K, TCD)]; Lampang [Mae Huat, Ngao, 21 May 1954, Sabhasi 2 (BKF); Chae Hom, 4 Feb. 1921, Kerr 47766 (AAU, ABD, BK, BM, K)]; Phrae [Pai Ton, 1 July 1911, Map 2.2.5 Distribution of Cratoxylum neriifolium in Thailand.

SW 11611-p162-216-G8.indd 213 1/16/62 BE 5:25 PM 214 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Distribution.— India, Myanmar, Laos, B.S., Jansen, R.K., Kim, K.J., Wimpee, C.F., Cambodia. Smith, J.F., Furnier, G.R., Strauss, S.H., Xiang, Ecology and phenology.— Deciduous and Q.Y., Plunkett, G.M., Soltis, P.S., Swensen, evergreen forests, on limestone from 150 to 800 m S.M., Williams, S.E., Gadek, P.A., Quinn, C.J., elevation. Flowering: April–August. Fruiting: Eguiarte, L.E., Golenberg, E., Learn, G.H., (August) September–March. Graham, S.W. Jr., Barrett, S.C.H., Dayananden, S. & Albert, V.A. (1993). Phylogenetics of seed Vernacular.— Tao (แตว))( (Kerr 8619); khi chiu plants: an analysis of nucleotide sequences from (ขี้ติ้ว) (Kerr 47766); tio dam (ติ้วดํา) (Winit 1794). the plastid gene rbcL. Annals of the Missouri Uses.— Used in construction, and for wooden Botanical Garden 80: 528–580. utensils (e.g. handles of chisels and hammers). D’Arcy, W.G. (1980). Family 123. Guttiferae. In: Conservation.— IUCN Regional (Thailand) R.E. Woodson & R.W. Schery and collaborators, Status: Least Concern (LC). Flora of Panama. Annals of the Missouri Botanical Gardens 67: 969–1043. ACKNOWLEDGEMENTS Davis, C.C. & Chase, M.W. (2004). Elatinaceae are sister to Malpighiaceae: Peridiscaceae belong We would like to thank the Irish Research to Saxifragales. American Journal of Botany 91: Council for Science, Engineering and Technology 262–273. (IRCSET) for funding this research project, an Davis, C.C., Webb, C.O., Wurdack, K.J., Jaramillo, anonymous referee and especially Dr Hajo Esser for C.A. & Donoghue, M.J. (2005). Explosive radia- helpful comments on, corrections to and detailed tion of the Malpighiales supports a Mid- reading of an earlier versions of this paper. Cretaceous origin of Modern Tropical Rain Forests. American Naturalist 165: E36–E65. REFERENCES Frohne, D. & Pfänder, H.J. (2005). Poisonous Plants: APG II (2003). An update of the Angiosperm A Handbook for Doctors, Pharmacists, Phylogeny Group classifi cation for the orders and Toxicologists, Biologists & Veterinarians. families of fl owering plants. APG II. Botanical Timber Press, Portland, 488 pp. Journal of the Linnean Society 141: 399–436. Gardner, S., Sidisunthorn, P. & Chayamarit, K. APG III (2009). An update of the Angiosperm (2015). Forest Trees of Southern Thailand. Vol. Phylogeny Group classifi cation for the orders 1. The Forest Herbarium, Bangkok and the and families of flowering plants. APG III. Royal Botanic Gardens, Kew, 748+1 pp. Botanical Journal of the Linnean Society 161: Gogelein, A.J.F. (1967). A revision of the genus 105–121. Cratoxylum. Blumea 15: 453–475. APG IV (2016). An update of the Angiosperm Heywood, V.H., Brummit, R.K., Culham, A. & Phylogeny Group classifi cation for the orders Seberg, O. (2007). Families of and families of flowering plants: APG IV. the World. Royal Botanic Gardens, Kew, 424 pp. Botanical Journal of the Linnean Society 181: ICRAF. World Agroforestry Centre. (2008). http:// 1–20. www.worldagroforestrycentre.org/SEA/ Bartholomew, B., Nicolson, D.H. & Nordenstam, Products/AFDbases/AF/index.asp. Accessed 8th B. (1997). Author citation of Thunberg’s new April 2008. species in Murray’s “Systema vegetabilium,” ______.(2017). The Agroforestree Database. Mesua ed. 14. Taxon 46: 311–314. ferrea. http://www.worldagroforestry.org/treedb2/ Chase, M.W., Soltis, D.E., Olmstead, R.G., Morgan, speciesprofi le.php?Spid=1770. Accessed July D., Les, D.H., Mishler, B.D., Duvall, M.R., 2017. Price, R.A., Hills, H.G., Qiu, Y.L., Kron, K.A., Jarvis, C.V. (2007). Order out of Chaos. Linnean Rettig, J.H., Conti, E., Palmer, J.D., Manhart, plant names and their types. Linnean Society of J.R., Sytsma, K.J., Michael, H.J., Kress, W.J., London and the Natural History Museum, Karol, K.A., Clark, W.D., Hedrén, M., Gaut, London, 1017 pp.

SW 11611-p162-216-G8.indd 214 1/21/62 BE 3:21 PM SYSTEMATICS OF THE THAI CALOPHYLLACEAE AND HYPERICACEAE WITH COMMENTS ON THE KIELMEYEROIDAE (CLUSIACEAE) 215 (C. BYRNE, J.A.N. PARNELL & K. CHAYAMARIT)

Kato, M. (2004). Taxonomic studies of Podostemaceae Robson, N.K.B. (1985). Studies in the genus of Thailand. 1. Hydrobryum and related genera Hypericum L. (Guttiferae) 3. Sections 1. with crustaceous roots (Subfamily Podostemoi- Campylosporus to 6a. Umbraculoides. Bulletin deae). Acta Phytotaxonomica et Geobotanica of the British Museum Natural History (Botany) 55: 133–165. 12: 163–235. Kochummen, K.M. (1973). Hypericaceae. In: T.C. ______. (1990). Studies in the genus Hypericum L. Whitmore, Tree Flora of Malaya, pp. 248–252. (Guttiferae) 8. Sections 29. Brathys (part 2) and Longman, Kuala Lumpur & Singapore. 30. Trigynobrathys. Bulletin of the British Koi, S., Imaichi, R. & Kato, M. (2005). Endogenous Museum Natural History (Botany) 20: 1–151. leaf initiation in the apical-meristemless shoot Robson, H.B.K., Carine, M., Pattinson, D. Nurk, N., of Cladopus queenslandicus (Podostemaceae) Crockett, S. & Wajer, J. (2014). Hypericum online. and implications for evolution of shoot http://hypericum.myspecies.info/taxonomy/ Morphology. International Journal of Plant term/723/descriptions accessed 21 May 2017. Science 166: 199–206. Sangkaew, S. (1999). Taxonomic studies of the genus Kubitzki, K., Mesquita, A.A.L. & Gottlieb, O.R. Calophyllum L. (Guttiferae) in Thailand. Masters (1978). Chemosystematic implications of xan- thesis. Kasetsart University, Bangkok, 144 pp. thones in Bonnetia and Archytaea. Biochemical Savolainen, V., Fay, M.F., Albach, D.C., Backlund, Systematics and Ecology 6: 185–187. A., van der Bank, M., Cameron, K.M., Johnson, Li, Xiwen, Li. Jie & Stevens, P.F. (2007). Clusiaceae S.A, Lledo, M.D., Pintaud, J.C., Powell, M., (Guttiferae). In Flora of China 13: 37–38. Sheahan, M.C., Soltis, D.E., Soltis, P.S., Weston, Science Press, Beijing & Missouri Botanical P., Whitten, W.M., Wurdack, K.J. & Chase, M.W. Garden, St. Louis. (2000a). Phylogeny of the : a nearly Lord, T., Armitage, J., Cubey, J., Grant, M. & complete familial analysis based on rbcL gene Whitehouse, C. (2004). RHS Plant Finder sequences. Kew Bulletin 55: 257–309. 2004–2005. Dorling Kindersley, London, 956 pp. Savolainen, V., Chase, M.W., Hoot, S.B., Morton, Mabberley, D.J, (2017). Mabberley’s Plant Book. C.M., Soltis, D.E., Bayer, C., Fay, M.F., Bruijn, Cambridge University Press, Cambridtge, 1102 A.Y. de, Sullivan, S. & Qiu, Y.L. (2000b). pp. Phylogenetics of flowering plants based on combined analysis of plastid atpB and rbcL gene Maheswari, J.K. (1963). Taxonomic studies on sequences. Systematic Biology 49: 306–362. Indian Guttiferae II. The genus Mesua Linn. Bulletin of the Botanical Survey of India 5: Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., 335–343. Albach, D.C., Zanis, M., Savolainen, V., Hahn, W.H., Hoot, S.B., Fay, M.F., Axtell, M., Swensen Merrill, E.D. (1916). Osbeck’s Dagbok öfwer en S.M., Prince, L.M., Kress, J.W., Nixon, K.C. & Ostindsk Resa. American Journal of Botany 3: Farris, J.S. (2000). Angiosperm phylogeny inferred 580–588. from 18s rDNA, rbcL, and atpB sequences. MTC. Malaysian Timber Council (2017). Resources Botanical Journal of the Linnean Society 133: forestry and sustainability. Derum. http://www. 381–461. mtc.com.my/resources-WoodWizard.php. Stevens, P.F. (1980). A revision of the Old World Accessed 9 July 2017. species of Calophyllum (Guttiferae). Journal of Parnell, J. A. N., Simpson, D. A., Moat, J., Kirkup, the Arnold Arboretum 61: 117–699. D. W., Chantaranothai, P., Boyce, P. C., Bygrave, ______. (2001 onwards). Angiosperm Phylogeny P., Dransfi eld, S., Jebb, M. H. P., Macklin, J., Website. [and more or less continuously updated Meade, C., Middleton, D. J., Muasya, A. M., since] http://www.mobot.org/MOBOT/research/ Prajaksood, A., Pendry, C. A., Pooma, R., Apweb. Version 7, May 2006; accessed 8 Apr. Suddee, S. and Wilkin, P. (2003). Plant collecting 2008. Version 12, July 2012; accessed March spread and densities: their potential impact on 2012. Version 14, accessed 14th May 2018. biogeographical studies in Thailand. Journal of Biogeography 30: 193–209.

SW 11611-p162-216-G8.indd 215 1/16/62 BE 5:29 PM 216 THAI FOREST BULLETIN (BOTANY) VOL. 46 NO. 2

Stevens, P.F. (2006). Clusiaceae and Hypericaceae. van Welzen, P.C., Madern, A., Raes, N., Parnell, In: K. Kubitzki (ed.), The Families and Genera J.A.N., Simpson, D.A., Byrne, C., Curtis, T., of Vascular Plants 9, pp. 48–66, 194–201. Macklin, J., Trias-Blasi, A., Prajaksood, A., Springer-Verlag, Berlin. Bygrave, P., Dransfi eld, S., Kirkup, D.W., Moat, Sun, M., Naaem, R., Su, J.-X., Cao, Z.-Y., Burleigh, J., Wilkin, P., Couch, C., Boyce. P.C., Chayamarit, J.G., Soltis, P.S., Soltis, D.E. & Chen, Z.-D. K., Chantaranothai, P., Esser, H.-J., Jebb, M.H.P., (2016). Phylogeny of the Rosidae: A dense taxon Larsen, K., Larsen, S.S., Nielsen, I., Meade, C., sampling analysis. Journal of Systematics and Middleton, D.J., Pendry, C.A., Musaya, A.M., Evolution 54: 363–391. Pattharahirantricin, N., Pooma, R., Suddee, S., Staples, G., Sungkaew, S. & Teerawatananon, A. Tokuoka, T. & Tobe, H. (2006). Phylogenetic analyses (2011). The current and future status of Floristic of Malpighiales using plastid and nuclear DNA Provinces in Thailand. In: Y. Trisurat, R.P. sequences, with particular reference to the Shrestha, & R. Alkemade (eds), Land use, climate embryology of Euphorbiaceae sensu stricto. change and biodiversity modeling. Perspectives Journal of Plant Research 119: 599–616. and applications, pp. 219–247. Information Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, Science Reference, Hershey. W., Hawksworth, D.L., Herendeen, P.S., Knapp, Whitmore, T.C. (1973). XVIII. Guttiferae. In: Notes S., Kusber, W-H., Li, De-Zhu, Marhold, K., May, on the systematy of Malayan Phanaerogams T.W., McNeill, J., Monro, A.M., Prado, J., Price, XVIII–XXII*. Gardens Bulletin Singapore 26: M.J. & Smith, G.F. (2018). International Code 269–284. of Nomenclature for algae, fungi and plants (Shenzen Code). Regnum Vegetabile 159. Wurdack, K.J. & Davis, C.C. (2009). Malpighiales Koeltz Botanical Books, Glashütten, 254 pp. Phylogenetics: gaining ground on one of the most recalcitrant clades in the Angiosperm Tree of Life. American Journal of Botany 96: 1551–1570.

SW 11611-p162-216-G8.indd 216 1/16/62 BE 5:29 PM Reviewers of manuscripts for Thai Forest Bulletin (Botany) Vol. 46(2), 2018

Julie Barcelona University of Canterbury, Christchurch, New Zealand Kenneth Bauters Ghent University, Ghent, Belgium Hans-Joachim Esser Botanische Staatssammlung München, Germany Bob Harwood Northern Territory Herbarium, Darwin, Australia Matthew Jebb National Botanic Gardens Glasnevin, Dublin 9, Ireland Xiaohua Jin Institute of Botany, Chinese Academy of Sciences, Beijing, China Charan Leeratiwong Prince of Songkla University, Songkhla, Thailand Stuart Lindsay National Parks Board, Singapore Pete Lowry Missouri Botanical Garden, St. Louis, Missouri, USA David Middleton Singapore Botanic Gardens, Singapore Pimwadee Pornpongrungrueng Khon Kaen University, Khon Kaen, Thailand Andre Schuiteman Royal Botanic Gardens, Kew, UK David Simpson Royal Botanic Gardens, Kew, UK Tim Utteridge Royal Botanic Gardens, Kew, UK Chun-Lei Xiang Chinese Academy of Sciences, Beijing, China

SW 11611-A-G8.indd 1 1/16/62 BE 5:34 PM THAI FOREST BULLETIN (BOTANY) Thai Forest Bulletin (Botany) Vol. 46 No. 2, 2018

Published by the Forest Herbarium (BKF) CONTENTS Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900, Thailand Page Advisors Wipawan Kiaosanthie, Wanwipha Chaisongkram & Kamolhathai Wangwasit. Chamlong Phengklai & Kongkanda Chayamarit A new species of Scleria P.J.Bergius (Cyperaceae) from North-Eastern Thailand 113–122 Editors Willem J.J.O. de Wilde & Brigitta E.E. Duyfjes. Miscellaneous Cucurbit News V 123–128 Rachun Pooma & Tim Utteridge Hans-Joachim Esser. A new species of Brassaiopsis (Araliaceae) from Thailand, and lectotypifications of names for related taxa 129–133 Managing Editor Assistant Managing Editor Orporn Phueakkhlai, Somran Suddee, Trevor R. Hodkinson, Henrik Æ. Pedersen, Nannapat Pattharahirantricin Sawita Yooprasert Priwan Srisom & Sarawood Sungkaew. Dendrobium chrysocrepis (Orchidaceae), a new record for Thailand 134–137 Editorial Board Rachun Pooma (Forest Herbarium, Thailand), Tim Utteridge (Royal Botanic Gardens, Kew, UK), Jiratthi Satthaphorn, Peerapat Roongsattham, Pranom Chantaranothai & Charan David A. Simpson (Royal Botanic Gardens, Kew, UK), John A.N. Parnell (Trinity College Dublin, Leeratiwong. The genus Campylotropis (Leguminosae) in Thailand 138–150 Ireland), David J. Middleton (Singapore Botanic Gardens, Singapore), Peter C. van Welzen (Naturalis Alan Paton, Somran Suddee & Bhanubong Bongcheewin. Chelonopsis thailandica, Biodiversity Center, The Netherlands), Hans-Joachim Esser (Botanische Staatssammlung München, a new species and new record of Chelonopsis (Lamiaceae) from Thailand 151–154 Germany), Bob Harwood (Northern Territory Herbarium, Darwin, Australia), André Schuiteman Nutdanai Putthisawong & Sahut Chantanaorrapint. A revision of the genus Tapeinidium (Royal Botanic Gardens, Kew, UK), Anders S. Barfod (Aarhus University, Denmark), (Lindsaeaceae) in Thailand 155–161 Piyakaset Suksathan (Queen Sirikit Botanic Garden, Thailand), Pimwadee Pornpongrungrueng (Khon Kaen University, Thailand), Stuart Lindsay (National Parks Board, Singapore) Caroline Byrne, John Adrian Naicker Parnell & Kongkanda Chayamarit. Systematics of the Thai Calophyllaceae and Hypericaceae with comments on the Kielmeyeroidae Thai Forest Bulletin (Botany) (TFB) publishes papers on plant taxonomy (especially of vascular plants), (Clusiaceae) 162–216 nomenclature, phylogeny, systematics, plant geography, and floristics, and in morphology, palynology, cytotaxonomy, chemotaxonomy, anatomy and other relevant disciplines. The journal now uses Thai Journal Online (ThaiJO) for online submission and peer review at www.tci-thaijo.org/index.php/ThaiForestBulletin. Manuscripts are considered on the understanding that their contents have not appeared, or will not appear, elsewhere in the same or abbreviated form. Before submitting a manuscript please read the Guidelines for authors. These guidelines must be followed precisely otherwise publication of the manuscript will be delayed. In addition, papers published online will be distributed simultaneously in printed form to several libraries, and bound hard copy volumes will appear later. Exchange with botanical journals or periodicals pertaining to plant taxonomy would be appreciated.

FOREST HERBARIUM Director: Phongsak Phonsena Curator: Nannapat Pattharahirantricin BKF Staff: Somran Suddee, Piyachart Trisarasri, Preecha Karaket, Thanongsak Jonganurak, Pachok Puudjaa, Voradol Chamchumroon, Nanthawan Suphuntee, Narong Koonkhunthod, Montri Saengsawasti, Naiyana Tetsana, Sukontip Sirimongkol, Manop Poopath, Sommanussa Saengrit, Sukid Rueangruea, Baramee Sakolrak, Sawita Yooprasert, Saksan Kaitongsuk, Orathai Kerdkaew.

Front Cover: Brassaiopsis spinosissima Esser

Printed at: Prachachon Co., Ltd. 35 Soi Pipat, Silom Road, Bangrak, Bangkok 10500, Thailand Tel : 0 2636 6550 Thai Fores Thai Forest Bulletin

t Bulletin (Botany) Vol. 46 No. 2, 2018 (Botany)

Vol. 46 No. 2, 2018 ISSN 0495-3843 (print) ISSN 2465-423X (electronic)

Forest Herbarium Department of National Parks, Wildlife and Plant Conservation Chatuchak, Bangkok 10900 THAILAND http://www.dnp.go.th/botany ISSN 0495-3843 (print) ISSN 2465-423X (electronic)

Fores t Herbarium Department of National Parks, Wildlife and Plant Conservation Bangkok, THAILAND